DeVries &toenniessen - Securing The Harvest PDF

Acknowledgements
This book grew out of a 2-year exploration conducted by the Food Security theme of
The Rockefeller Foundation focusing on the potential for crop genetic improvement to
contribute to food security among rural populations in Africa. The exploration carried
the authors to ten countries of sub-Saharan Africa and a number of related national,
regional, and international meetings on several continents. Along the way, innumerable
individuals farmers, researchers, seed merchants, policy experts and others contributed their views, comments and experiences related to crop improvement in African
agriculture, and the authors are very grateful for their assistance.
In particular, we would like to thank Foundation colleagues Gordon Conway,
Bob Herdt, John Lynam, John OToole and Ruben Puentes for reading through the
manuscript and sharing their views. David Jewell and Gebisa Ejeta also read early drafts
and gave useful comments. In addition, we would like to express our gratitude to the
following individuals who assisted with the study by sharing their views and providing
information: Marianne Banziger, Jeffrey Bennetzen, Malcolm Blackie, Ronnie
Coffman, Joel Cohen, Ken Dashiell, Alfred Dixon, Peter Ewell, John Hartmann, Tom
Hash, Dave Hoisington, Lee House, Justice Imanyowa, Jane Ininda, Saleem Ismael,
Monty Jones, Richard Jones, Bill Kiezzer, Laurie Kitch, Jenny Kling, Dennis Kyetere,
Isaac Minde, Larry Murdoch, Patricia Ngwira, Hannington Obiero, Joseph Ochieng,
Moses Onim, James Otieno, Yvonne Pinto, Kevin Pixley, Fred Rattunde, Darrell
Rosenow, B.B. Singh, B.N. Singh, Elizabeth Sibale, Ida Sithole, Margaret Smith,
Aboubacar Toure, Lamine Traore, Wilberforce Tushemereirwe, Eva Weltzien and John
Whyte. Finally, we would like to express our sincere appreciation to Sarah Dioguardi
and Mulemia Maina, who provided excellent care and technical assistance in preparing
the manuscript.
Inevitably, when attempting to address as broad a range of issues as biotechnology
to seed production in a number of important crops, mistakes and discrepancies will
occur, both in terms of the facts gathered and the assertions made. Although the authors
have tried to avoid these, they apologize in advance for those that remain, and take full
responsibility for them.
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Executive Summary
Food crops grown under low-input, rain-fed conditions in sub-Saharan Africa are
affected by a wide range of biological and environmental constraints, but remain the
best, if not the only, means of improving food security among the rural poor. To reach
maturity and yield well, crop varieties must be able to resist or tolerate these stress
factors. Due to wide variation in environmental conditions over space and time, the
particular set of constraints which operate in any given area is continually changing.
Moreover, local processing and consumption needs often exert additional quality
requirements in order for improved crop varieties to be adopted by small-scale farmers.
To be successful, breeding programmes for Africa must take into consideration this
variation and relevant varietal preferences of farmers. By analysing these requirements,
selecting appropriate parental materials, and making selections under relevant local
conditions with regular farmer input, new varieties with the right combination of
genetic resistances and tolerances can be produced.
This kind of approach differs significantly from the methodology of selecting for
high yield potential and broad adaptation which continues to give good results in more
stable and more highly modified agricultural environments such as those in developed
countries and the irrigated regions of developing countries. The major implication is a
need for more localized, agro-ecology-based breeding programmes, where the principal
objective is to assemble a set of traits that reduce yield losses and thereby confer greater
yield stability. Over time, yield-enhancing genes may still be added and make a significant contribution to overall performance, but the emphasis during the current phase of
breeding programmes should be placed on critical resistance factors.
The need to develop a range of improved varieties for Africa, each well adapted to
local conditions, argues strongly for giving priority to well-funded and staffed crop
breeding programmes at the national level. Country-level programmes have lower costs
and are able to deploy larger numbers of teams which can operate in close proximity to
the various agro-ecologies that need to be covered by any given programme.
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Executive Summary
International agricultural research centres (IARCs) have a major role to play in

facilitating the development of fully capable national agricultural research systems
(NARSs) able to produce the steady flow of new offerings required by farmers. In
addition, international centres and advanced research institutes should devote
significant resources and attention to the more difficult, intractable constraints of crop
production which affect the important crop species of Africa. Such intractable
constraints are numerous and have not been solved despite considerable effort using
conventional techniques. By combining their talent and resources, and drawing on the
strengths of biotechnology, international centres and advanced research institutes may
now be able to overcome many, if not most, of these difficult problems.
Biotechnology remains a highly underdeveloped resource for improved food
production in Africa, largely due to underinvestment by governments and international
donors. Africa already has a number of scientists trained in biotechnology who are
unable to utilize their knowledge owing to lack of facilities and operating funds. Since
this situation may continue for some time to come, development of fully functional
biotechnology capacity in all NARSs is not likely. However, those countries that can
adequately staff both conventional and molecular breeding units should be encouraged
to do so. Tissue culture of clonally propagated crops has already proved its value
to agriculture in Africa. A second application of biotechnology which could prove
cost-effective in the short to medium term is marker-aided selection for a range of traits,
with the primary objective being to combine as many resistance traits as are required to
maximize crop performance under low-input conditions. Finally, as national biosafety
regulations and systems become operational, it will become more logical to invest in
national expertise and facilities for crop genetic engineering, whereby critical resistance
traits may be transferred directly into otherwise well-adapted varieties.
Localized, agro-ecology-based crop improvement schemes need to be supported
by similarly oriented seed enterprises. In Africa, investment in the seed sector has
historically been very low, in part influenced by the poor success record of large seed
companies on the continent. Large, monopolistic seed companies have perceived little
advantage in pursuing the locally directed breeding programmes needed to develop
a range of varieties adapted to the various niches created by environmental variation.
Multinational seed companies that rely solely on their own offshore breeders and gene
banks find it difficult to overcome the adaptation barriers of Africa; and their historical
reluctance to commercialize germplasm under licensing agreements with the public
sector further diminishes the attraction of operating in Africa.
Conversely, smaller entities operating in a competitive environment that ally
themselves to NARS breeding programmes for access to new varieties may perform well
with respect to small farmers interests. Their most obvious limitations size and lack of
capital can serve as an effective entry point for governments, private investors and
donor agencies. Limited production of foundation seed is one bottleneck to the growth
of this and related models for development of the seed industry. More harmonious
regulatory structures across the region are also needed.
Taken together, the combination of new science, new ways of working with
farmers, new opportunities for private sector seed supply, and a greater appreciation of
Africas diverse agro-ecologies represent a new era in crop genetic improvement for
Africa. Old arguments for products already being on the shelf lose their meaning in
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Executive Summary
view of what scientists and farmers can achieve today, if the needed effort and resources
are put forward.
In this context, the importance of responsive, relevant public policy in the furtherance of a healthy, functional germplasm sector in Africa can hardly be overestimated.
Public policy makers need to be committed to solving the problem of food insecurity on
the continent, and to employing relevant, up-to-date policy that can strengthen the
breeding and seed sectors. The worldwide biotechnology debate has provided the latest
opportunity to put African agriculture in the spotlight and emphasize the need to move
policy structures forward rapidly and responsibly. A major tenet of these changes must
be to encourage private investment of all kinds in the seed sector. Another is the
reinforcement of public sector capacity in crop improvement, using both conventional
and molecular techniques. The establishment of an effective set of biosafety regulations
is also critical to taking advantage of recent advancements in crop improvement.
In spite of its potential, genetic improvement of crops will always face limitations
with regard to what it can offer to farmers in regards to their levels of productivity. No
matter what efficiencies genetic enhancement is able to build into crop plants, they will
always draw their nutrition from external sources, and this places enormous importance
on the investments that can be made in the soils of Africa. Overall improvements in agricultural productivity are likely to move in tandem with improvements made in the management of soil nutrients by African farmers. Shortfalls in the level of nutrient supply
that are possible through the uses of organic methods must be complemented by making
fertilizer broadly more accessible to small-scale farmers. Because of the need to demonstrate the potential of the combined effects of genetic improvement and improved soil
fertility, crop improvement initiatives and soil fertility management programmes should
operate in similar environments and test their results on the same or similar sites.
These policy and technology innovations can combine well with the revolution in
farmer participation in agricultural research. One very critical entry point for farmer
participation is the need to understand better the various agro-ecologies that can be
targeted by public breeding programmes. Farmers are the best source of information
regarding the number and prioritization of production constraints, as well as the spatial
distribution of differing agro-ecologies. In view of the importance and complexity of
their preferences for processing, taste, growth habit, and multiple uses of crop plants,
farmers also need to be made part of the process of varietal selection. While there is no
set procedure for farmer participation in breeding schemes in Africa, it seems obvious
that breeding programmes which operate in close proximity to farmers and their base of
knowledge will have definite advantages over those which do not involve farmers.
Within this rapidly evolving professional context, crop genetic improvement can be
viewed as a highly underexploited resource for improving food security among Africas
majority, rural populations. Indeed, with late-maturing, low-yielding crop varieties
dominating the farming systems of much of Africa, crop genetic improvement still
has the potential to play an important role in the development of more productive agricultural systems throughout the continent.
A new paradigm for germplasm improvement in Africa, and indeed in other regions
of the developing world, can be envisioned. It is a paradigm driven first and foremost by
the urgent need for food security in Africa among a growing population of very poor,
rural people who have been left behind by globalization and the interests of the private
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Executive Summary
sector. The impetus for understanding the details of their needs in terms of better, more
resilient crops leads directly to the application of an enriched set of technologies for
crop improvement, including conventional, field-based selection and laboratory-based
modification and enhancement of the germplasm. Public sector technology development in this paradigm is linked directly to a broad interface of private initiative through
non-governmental organizations (NGOs), farmers associations and private business.
And, it is backed up by the commitment to serving the peoples needs through seed
distribution by an efficient seed sector.
Providing the crop varieties needed to improve food security across the vast continent of Africa is an enormous challenge. No donor or national government acting alone
would be able to mobilize the commitment and resources necessary to make a major
change in this area. Novertheless, there is very little likelihood that Africa will be food
secure without an intensive, long-term programme of investment in crop improvement
which takes advantage of the full range of approaches now available.
While it would be going too far to declare that improved food security through
higher-performing crops throughout Africa is readily do-able, it is at least possible to
break down the process into conceivable steps as follows:
1. Constructing the breeding teams within NARSs supported by IARCs.
2. Delineating and classifying the agro-ecologies which merit targeting.
3. Determining farmer preferences for new varieties.
4. Employing appropriate parental materials and breeding methods aimed to produce
new varieties within an acceptable time frame.
5. Getting seed to farmers via public and private means.
Running concurrently with the process above, biotechnology studies aimed at
developing solutions to intractable problems can be initiated at any time. Products of
those studies feed into step 4. In view of the recent, negative trend in food availability
and child nutrition in sub-Saharan Africa, food security in Africa is one of the most
critical challenges facing humankind today. The record of private investment during the
post-structural adjustment era does not present a convincing argument that growth in
the private sector alone will lead Africa out of poverty and food insecurity. The public
sector still plays an enormously important role in offering hope to the poor and excluded
throughout the continent. And within this grouping, public sector capacity in the
genetic improvement of food crops presents an exciting opportunity to make real
progress.
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Foreword
One of the many outcomes of the global media debate on biotechnology has been a
heightened level of awareness, and one wants to believe interest in how the food crops
that provide our nutrition are developed, grown, and eventually end up on our tables.
This is a positive outcome for several reasons. First, because agriculture in the developed
world, although playing a huge role in the way we live, tends to remain out of sight and
out of mind for nearly all of us except the 2 or 3% who are farmers and farm workers.
Second, because it has reminded us that food security is never a resolved issue. One way
or another, we have to keep on producing enough food for 6 billion people today and 8
billion by 2025, or there could be mass starvation. Finally, the debate on biotechnology
has provided spokespersons of the agricultural community with the opportunity of
explaining to the rest of the world just how dependent we already are, with or without
biotechnology, on genetic improvement of food crops and on inputs such as fertilizer.
As the one remaining major world region where agriculture has yet to be transformed from subsistence, low-yield systems dependent on shifting cultivation to
efficient, modern systems capable of producing regular surpluses, the question of crop
improvement is especially important to Africa. Africa is also the sole world region where
many indices of food security have shown a serious decline in recent years. In the context
of continued high population growth and an increased emphasis on keeping Africas
unique natural environment intact, it is clear that crop yields must be substantially and
sustainably increased. As they have in all other parts of the world, more efficient,
better-performing crop varieties can play an important role in achieving this goal.
This book came about as part of a major restructuring of The Rockefeller Foundation which has resulted in a renewed commitment to the poor and excluded of the
world, who have largely been left behind by globalization and economic growth. The
studys Africa focus reflects a greater emphasis being placed by our Food Security
Programme on that part of the world bypassed by the Green Revolution. Its attention
to issues ranging from frontier research in biotechnology to participatory methods of
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Foreword
seed dissemination via farmers groups reflects a greater concern with the application of
science to the needs of the poor that result in real, positive changes in their lives and
livelihoods.
The title of Part I of this book, Biotechnology, Breeding, and Seed Systems for African
Crops: Re-thinking a 10,000-year-old Challenge, reflects what is an ambitious attempt by
the authors to encapsulate in a brief format our current understanding of the nature of
the task of extending better-performing crop varieties to Africas farmers. While it is
clear that any one group can only focus on selected portions of this process, it is hoped
that the opportunities identified can mobilize additional resources and generate new
partnerships which cover the full scope of the challenge ahead.
It has been of particular interest to me to note the important roles the authors foresee for gaining a greater understanding of agro-ecologies in Africa and for the application
of participatory methods as well as biotechnology. By generating new crop varieties with
greater yield stability, greater productivity and greater local acceptability, and by getting
the new genetic resources into farmers hands through more responsive seed systems,
they believe increased food security can be attained.
Gordon Conway
New York
January 2001
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Biotechnology, Breeding,
and Seed Systems for
African Crops: Re-thinking
a 10,000-year-old Challenge
Chapter I
Introduction and Summary
Introduction
The final decades of the 20th century achieved the fastest growth of the global economy
ever recorded over a similar period. Rapid scientific and technological innovation,
coupled with the opening up of economies throughout the world, permitted more
people to improve their well-being than had ever been possible before over such a
relatively brief period of time. Globalization, as the phenomenon of capital mobility
and global distribution of technology came to be called, brought employment and
opportunity to innumerable groups of people who only one generation previously were
unable to imagine such changes.
For many of the worlds poor, the most immediate effect of global economic
growth meant simply eating better and enjoying the many subtle but influential benefits
of adequate nutrition and improved health. World food prices fell dramatically, while
life expectancy rose sharply. As living standards rose, greater numbers of children were
also able to attend school.
At the centurys close, however, it became apparent that not all the worlds
population was equally swept along in the positive trend. A significant portion of
the worlds population had failed to benefit from globalization. Furthermore, the
widespread rolling back of social services within the public sector, coupled with the
widespread belief that in the new world order everyones needs would be adequately met
through the marketplace, meant that many members of this group had less chances of
escaping poverty than ever before.
The inability of a large segment of the worlds population to benefit from the
current socio-economic advances presents the world with intellectual and moral
challenges that cannot be ignored. Africa, it is widely accepted, lies at the core of this
challenge. While home to fewer total numbers of this left-behind group than Asia, it
also embodies fewer of the factors necessary for inclusion in the growth-led process of
globalization. In Africa, large portions of the population do not have access to sufficient
food. Economies are not growing at rates required to generate new opportunities for
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Chapter 1
growing populations. The ensuing widespread frustration felt by local populations is

proving a fertile ground for civil conflict and regional wars.
With upwards of 70% of Africas workforce engaged in farming, agriculture
represents an important channel for extending new opportunities for improving the
well-being of hundreds of millions of people throughout the continent. Unfortunately,
continued reliance on technologies and practices designed for a previous era means that
agriculture has largely become a trap for Africas rural population, guaranteeing a life of
poverty and isolation.
It does not have to be this way. The arrival of the information age, combined with
new biological technologies, and new ways of linking people to them, means that
peoples lives can be improved in a relatively brief period of time. This book explores the
ways to take advantage of the new capacity for global knowledge sharing and increased
public and private capital gains from the past decades, and direct them at one important
group of technologies crop genetic resources to broadly improve access to adequate
food and enhance the well-being of Africas rural poor.
This book is not intended as an analysis of all the activities ongoing in crop genetic
improvement in Africa. Rather, it is intended as a collection of observations obtained
from a wide range of geographical locations, as well as institutions involved in making
crop genetic resources more valuable and more useful to African farmers. In making
those observations, the authors also attempt to understand what has worked in Africa,
what has not, and how the lessons learned might be grouped together to provide some
guiding principles for crop genetic improvement (the term is used to imply all methods
available, including biotechnology, conventional breeding, and seed dissemination)
work in the future. Nevertheless, they are the first to recognize that much needs to be
learned in Africa, and that the views of many qualified people must yet be sought.
Improved food security, led by increased productivity among Africas many smallscale farmers, has been the aim of significant national and international effort in recent
decades. However, the relatively underdeveloped, non-globalized state of African
agriculture presents scientists, farmers and development agents with challenges at a
number of levels. African agriculture at the close of the 20th century remains by and
large an organic, living system, where biophysical signals within and between cropping
systems still pulse and exert checks and balances on the levels of success that can be
enjoyed by any single organism within the system. Strategies for increasing crop
productivity which operate within this context must be significantly different from those
applied in modernized, highly manipulated agricultural systems of developed regions of
the world. They must also differ substantially from strategies applied in Asia, previously
cited as a potential model for the challenge in Africa. This book attempts to show why
this is the case, and take a new look at the potential role of crop genetic improvement in
making sustainable improvements in the food security status of poor rural people in
Africa.
Improved varieties of African crops are destined for cultivation in soils that are very
low in fertility and where attacks by pests and diseases and periodic drought often
further reduce yields. These factors have led some to conclude that genetic improvement
of African crops cannot result in major social benefits. Indeed, some popular arguments
contend that increasing food production can do little to stave off hunger.
But crop improvement within this context is not just about raising yield
thresholds, just as efforts aimed at making food more abundant in chronically food
Chapter 1
insecure regions may have little impact on national food balances. Increasing the
amounts of food produced among poor farmers is aimed at improving nutrition
and maximizing options among people whose options are few. Better crop varieties
for African farmers also involve increasing yield stability and safeguarding the
meagre investments of some of the worlds most vulnerable people. This book argues
that real gains in food availability for the poor and excluded of Africa are possible
through publicly based, multi-tiered crop improvement strategies which are informed
by farmers needs and attuned to the agro-ecologies in which the new varieties will be
used.
The argument put forward in this text does not contend that better varieties alone
are the answer to food insecurity in Africa. Rather, more resilient, higher yielding
varieties are viewed as an important component of a broadly improved and bettersupported African farming environment. Improved varieties will inevitably perform
better on more fertile soils, just as farmers efforts at securing the harvest will go much
further within national and international policy environments that support and value
their livelihood.
Nevertheless, this study does recognize and seek to capitalize on the advantages of
seed and other planting materials in situations where, at present, little other assistance
can be offered to farmers. Seeds are animate technologies that can be easily transported
and transferred from one hand to another. Seed is also often the cheapest input available.
Improved varieties, therefore, are frequently the only modernized input used by African
farmers. They are the first step in securing the harvest.
The frame of reference for this study is one which will be very familiar to many field
researchers and development agents working in Africa. It is that of a single mother of
several children whose primary means of income is a 1 ha plot of unimproved land on an
eroded hillside. Depending on which part of the continent she is from, her principal
crop may be maize, sorghum, cassava, millet, rice, or banana. Inevitably, her farm will
contain other crops as well, such as cowpea, common bean, finger millet, groundnut,
and if she is lucky, a few cash crops such as vegetables. From each harvest, she must
provide for virtually all the needs of her family throughout the year, including clothing,
health care, education costs and housing. Because she can afford few purchased inputs,
the yield potential of her farm at the outset of the season is low she can expect to
harvest a maximum of perhaps 2000 kg of produce. Meagre though it may be, in most
years, through a wise combination of sales, barter and home consumption, she may be
able to cope at this low level of productivity.
Figure 1.1ad depicts hypothetically her farms productivity potential under different levels of intervention with adapted, accessible technologies, including better crop
varieties and more fertile soils. During the course of any given season, innumerable
threats to the crops appear on the scene (Fig. 1.1a). In the case of maize, the threats
might be drought, maize streak virus, stem borers, and the parasitic weed, Striga. If she
relies on cassava, the threats to her harvest may include African cassava mosaic virus,
bacterial blight and green mites. Periodic drought during the season has a further,
negative effect on yield. The impact of drought plus whatever combination of pests and
diseases attacks the crop in a given year can often reduce the average harvest on her farm
by perhaps 5060%, to 1000 kg of harvestable produce. At this level of productivity, the
family is on the edge of survival. If the losses are greater, or if disease enters the home,
some members may not survive.
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A4138
Strategies for securing the harvest in marginal farming zones of Africa.
Chapter 1
Fig. 1.1.
The effect of crop varieties which resist and/or tolerate these constraints (Fig. 1.1b)
can reduce such losses and raise her harvest above the theoretical survival line of
1000 kg. Meanwhile, improved soil fertility, shown separately in Fig. 1.1c, could allow
her to raise her initial potential harvest to perhaps 3000 kg, but without better varieties
the harvest is still reduced to some point at or near the survival line by the end of the
season. Improved soil fertility and resilient crops combined (Fig. 1.1d), could provide her
with the kind of productive potential and yield stability necessary to raise her harvest to
perhaps 2000 kg, a major improvement.
As uncomfortable as it may make us feel to contemplate the situation of this woman
and her children, this is the reality of millions of farm families throughout Africa today.
Certainly, they stand in need of development in the broadest sense. They need better
roads, better schools, better health care, and more employment opportunities. But they
also need better crop varieties, and in particular varieties which are resilient to drought,
low nutrient soils, insect pests and the myriad of diseases which attack crops in Africa.
Improving productivity securing the harvest in low-input systems where
farmers cannot afford purchased inputs means delivering as many useful traits as
possible within the seed. The end product of these efforts, moreover, must be usable and
acceptable by rural households. On a continent where upwards of 70% of the total
population are engaged in farming, better and more resilient crops which produce a
larger and more dependable harvest can be an effective strategy for delivering more food
and earning potential to those who need it most.
Directing science and technology at the ground-level needs of poor farmers may
not be the most effective way to increase food production on a national level. Better-off
farmers in more favourable farming environments may be quicker to adopt new technologies and produce higher yields with them once they are in place. Nor are resilient
crop varieties a new idea. But unlike in Asia, rain-fed, marginal farming conditions are
not a secondary focus in Africa, to be targeted once the more favourable areas have
been tapped. The difficult conditions and household scenarios like that faced by our
woman on the hill and her children are the only target whose solution will bring about
meaningful change in the vast, rural areas of the continent. Recent observations of the
revolution being brought about by globalization indicate they will not be delivered
unless very practical, results-oriented programmes are implemented by agricultural
research and development agencies within the public sector (Flavel, 1999; Persley and
Doyle, 1999). It is an enormous challenge, made more difficult by the very limited
resources currently being put forward to address it. Understanding how the challenge
can be approached putting together the pieces, if you will of some very promising
recent advances in the science and methods of working with the poor would seem a
useful subject to explore.
Summary
Recent years have seen vast improvements in our understanding of the genetic make-up
of crop plants and the techniques available for enhancing them. It is now possible to do
more for the woman on the hill than ever before. Indeed, the failure of the Green
Revolution to take root previously in Africa means that, in one form or another, most of
this potential is yet to be realized. Nevertheless, accessing these advances and directing
Chapter 1
Chapter 1
them toward the needs of the poor in an increasingly private sector-driven development
agenda is a major challenge which requires support from many sides.
Crop genetic resources are assets that the poor and excluded can own and
further modify to meet their needs. Experience and observation have shown that African
farmers are intensely interested in questions of crop variety performance. Most are
already engaged in informal variety trials of their own design. Their expertise can be
tapped in the search for resistance genes, in making selections, in growing out progeny
and in adapting varieties to local conditions. Many of them can help deploy and
disseminate the new varieties which result.
African farmers are supported by a group of committed, well-trained scientists and
technicians who understand well the tasks they face. Nevertheless, their numbers are as
yet insufficient to ensure full success. Moreover, their lack of access to operating funds
regularly reduces the rate and extent to which their knowledge can be applied.
Additional training is needed, especially in the newer techniques for genetic improvement and in understanding better Africas diverse agro-ecologies; and additional
financial support is required to allow them to put their strategies into action.
Some of the solutions are close at hand. For example, introgressing resistance to
maize streak virus in African maize populations should be a relatively simple task. If
known resistance genes were transferred into locally well adapted genotypes, maize
production across Africa might be increased by several million tonnes (see Plate 1).
Achieving solutions to other constraints will require more complex, high-risk ventures.
Downy mildew disease of millet is controlled by up to 17 genes (Hash et al., 1996), and
the fungal pathogen can rapidly evolve new pathotypes. Resistance in cereal crops to the
parasitic weed, Striga, is so ephemeral a trait that researchers are working at transferring
in more durable resistance genes from wild relatives (Ejeta et al., 2000; Kling et al.,
2000).
This book attempts to consider both the broad context of the role of crop genetic
improvement in improving food security in Africa and the more specific, scientific
challenges inherent to improvement strategies within important crop species. As such, it
is divided into two parts. This first part of the book looks at a range of human and
environmental factors which condition efforts aimed at benefiting farmers through
improved crop varieties, and then focuses on the discrete but interlinked roles of crop
breeding, biotechnology, and seed systems in developing and delivering new products to
farmers. The second part focuses on the challenges of genetic improvement and seed
dissemination for seven crop species of broad importance to African agriculture.
Chapter 1
2.1
The Challenge
Overview
The exploration that gave rise to this book tried, to the extent possible, to take a clean
slate approach to understanding the role of crop genetic improvement in African agriculture, and recognize those factors which seemed most influential in how small scale
farmers take advantage (or fail to take advantage) of improved crop varieties. An important sub-theme of this study was to understand why the Green Revolution of Asia and
Latin America did not have a greater level of impact in Africa.
Inevitably, the complexity inherent in the range of factors (farmer income, profitability, infrastructure, education, environmental factors, institutional factors, etc.) which
affect crop improvement in Africa obliged the authors to group some of those factors
which were perceived as less important tinto those which were believed to be of major,
continent-wide importance. The result is a short list of interacting factors which
includes:
the range and intensity of biophysical constraints to crop growth;

large agro-ecological variation;
the under-developed state of seed sectors in most countries;
the absence of policies which encourage crop improvement; and,
very low and declining soil fertility in much of Africa.
While depicted here as constraints, the chapter largely tries to communicate a message of optimism that previous barriers to raising agricultural productivity in Africa can
be overcome through new knowledge, new science and better methods of working with
farmers.
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Chapter 2
2.2 A Myriad of Production Constraints

The African continent south of the Sahara is dominated by agriculture. Approximately
70% of Africans live in rural areas and an estimated 50 million families derive their
livelihood from farming. The vast majority of these farms cover an area of less than 5 ha
and are hand-tilled. Crops are grown using a minimal amount of purchased inputs
(i.e. seed, fertilizer, etc.) (Wiggins, 2000).
Under these conditions, African crops are threatened by a daunting array of
debilitating production constraints which farmers can do little to change. In this book,
these constraints are loosely categorized as either routine or intractable. Routine
constraints are those which may be more or less effectively controlled through plant
breeding aimed at raising genetic resistance or tolerance levels through conventional
crossing and selection methods. Intractable constraints are those which are difficult or
impossible to control through conventional crop improvement (see Plate 2).
Categorization of a constraint as either intractable or routine is of course dependent
upon the ability of the farmer to alter the growing environment. The very limited investment capacity of small-scale farmers in Africa means that many potentially routine
production problems are, in fact, intractable. This increases the significance of genetic
crop improvement as a strategy in their potential control. Likewise, the dominance
of production constraints shifts the breeding strategy from one aimed at maximum
yield potential under high input use to one aimed at limiting losses from identified
constraints under low input use.
The potential to manage production constraints through crop genetic improvement
has increased steadily throughout the history of plant breeding, but has been greatly
expanded through the emergence of biotechnology. Although the diverse applications of
biotechnology may eventually make it a useful approach to the control of routine constraints to food production in Africa as well, for the purposes of this book, biotechnology is considered primarily applicable in the case of intractable constraints. An
incomplete, but illustrative short list of intractable constraints to production for seven
crops in significant portions of Africa is given below (Table 2.1).
While constraints to crop production exist throughout the world, they are more
intense in the tropics. Wellman (1968) studied the incidence of diseases on a number of
important food crops and noted far more in the tropics than in temperate areas. Dover
and Talbot (1987) reported that preharvest losses due to pests and diseases are approximately 3550% in some tropical areas (Table 2.2).
Biophysical constraints in Africa pose a greater threat to increasing agricultural
productivity than in other developing regions of the world. African farmers use vastly
fewer off-farm inputs and largely continue to apply traditional methods of cultivation
(Wiggins, 2000). In contrast, Latin American and Asian farmers have broadly modernized their cultivation methods over the past three decades (Table 2.3).
Table 2.3 indicates the vastly contrasting pace of development in different regions
in the developing world over the previous three decades. The agricultural sectors of Asia
and Latin America, which began the period with higher levels of development in all
categories (irrigation, fertilizer use and mechanization), have developed more rapidly
than Africas.
Latin America maintained a fourfold advantage in the percentage of irrigated
agricultural land over Africa. Asia, which began the period with a massive, 24-fold
11
The Challenge
advantage in the percentage of irrigated land, continued to add irrigated land at a rapid
pace, while Africa, continued to grow from a miniscule base.
Significant variations in rates of growth are also noted in fertilizer use and mechanization. Asia and Latin America finished the three-decade period with more than
fivefold and threefold increases in fertilizer application rates, respectively, while African
Table 2.1. Examples of intractable constraints to production among small-scale
farmers for seven important African food crops.
Focus crop
Intractable traits
Maize
Sorghum
Millet
Rice
Cowpea
Cassava
Banana
Striga, stem borers, phosphorus uptake

Striga, anthracnose, phosphorus uptake
Striga, head miner, downy mildew
Gall midge, rice yellow mottle virus
Maruca pod borers, bruchids, thrips
Root rots, green mite
Banana weevil, nematodes, black sigatoka
Table 2.2.
Crop disease incidence in tropical compared with temperate zones.

Number of diseases
Crop species
Temperate areas
Tropics
Sweet potato
Rice
Beans
Potato
Maize
15
54
52
91
85
187
500600
253280
175
125
Source: Dover and Talbot (1987) after Wellman (1968).

Table 2.3. Rates of usage of irrigation, fertilizers and mechanical land preparation
in Africa, Asia and Latin America.
Irrigated area
(% of total
agricultural land)
Sub-Saharan Africa
Asian developing
countries
Latin America and
Caribbean
Fertilizer
application
(kg ha1)a
No. of tractors
in use
( 103)
1970
1997
1970
1997
1970
1997
0.4
0.6
1.2
2.9
84
159
9.6
13.4
9.0
52.5
488
4610
1.5
2.4
4.4
14.6
637
1589
Source: FAO (2000).

aFigures vary. These were calculated by dividing FAO total fertilizer consumption by
total cultivated area.
12
Chapter 2
farmers managed only an increase of between two- and threefold. African farmers now
apply fertilizer at lower rates than Asian and Latin American farmers did three decades
ago. Lower input use in Africa is probably substituted in part by added labour input,
without which, yield levels would be lower. This translates to lower labour productivity,
and an accompanying drag on management capacity within the household, with
resultant negative impacts on factors such as sanitation, education and infant health.
Even more striking differences were noted for mechanical land preparation. In 1998,
Africa had one-third and one-quarter, respectively, the number of tractors in use as Asia
and Latin America in 1970.
All these factors irrigation, fertilizer and mechanization exert a homogenizing
force on crop growth conditions when they are present. Irrigation (and drainage) makes
water more uniformly available to the plant throughout the season, allowing for the
plant leaf canopy to remain fully extended over the full growing cycle. Irrigation also
significantly reduces risks associated with other forms of investment, such as fertilizers,
which fail to provide a cost-effective response in the absence of water. Fertilizer application, in addition to supplying basic nutrition for the development of vegetative and
reproductive structures, reduces variability of nutrient supply within the field, generally
increasing the value of genetically uniform crop varieties. Tillage performed by tractors
reaches deeper into the soil profile and, over time, reduces localized variation in the
fields topography.
The effect of input use is both a whole-farm environment that is more favourable
to crop growth than the surrounding, natural environment, and reduced within-farm
variation. It is in part the reduction of this within-farm variation that makes possible the
cultivation of highly uniform varieties of a single crop species possible throughout large
areas of North America, Asia and Latin America. As we will attempt to demonstrate, the
same is not true in Africa.
The preponderance of production constraints among African staple crops calls
for increased funding for research on crop genetic improvement to overcome those
constraints. While biotechnology is not an automatic solution to these constraints,
it should be viewed as a useful tool for improved food security in Africa. Tissue culture
can assist in the rapid multiplication of pathogen-free and true-breeding lines. Genotypic analysis through marker-assisted breeding can be used to identify favourable
individual plants with valuable, difficult-to-measure traits. Gene transfer through
genetic engineering can overcome limited genetic variation within a given species.
However, as emphasized throughout this book, biotechnology research should be linked
from the beginning to viable field-based breeding programmes, and, ultimately, to seed
dissemination strategies to prevent their results from remaining on-the-shelf.
2.3 Africas Diverse Cropping Landscape

Africas cultivated area is of immense size and has great environmental variation. African
farmers have developed complex cropping systems to fit environments ranging from the
slopes of Mt Kenya to the fringes of the Sahara, each with its unique mix of biotic and
abiotic constraints. For this reason, cropping patterns and dietary staples vary widely
from one end of the continent to another. Moreover, observations of small- compared
with medium- and large-scale farmers in Africa show that small-scale farmers tend to
The Challenge
13
cultivate a wider range of crop species, most likely as a strategy for maintaining household food security during a maximum portion of the year independent of household
purchasing power. The need for diversification may drive farmers to cultivate a very
wide range of crop and animal species (see Conway (1997) for an example from western
Kenya). In isolated regions of the continent where species diversity is limited, intraspecies (or, varietal) variation may be substituted. Farmers in Sudans Bar el Gazahl
region cultivate up to four varieties of sorghum whose morphological and growth habit
differences rival those found among different crop species.
Farmer crop deployment strategies extend to the species and subspecies level
according to complex environmental and social norms. Rice farmers in northern Mali
grow large plots of relatively high-yielding Asiatic (Oryza sativa) rice on upper-level
terraces of their farms on the Niger flood plain for normal consumption during the year.
In lower-lying parts of their farms, they grow preferred, African (O. glaberrima) varieties
for use mainly during special occasions such as religious holidays, weddings, and
baptisms. Farmers in the Bugusera region of Rwanda and Burundi grow different
varieties of sorghum and bananas in the same fields for use in either beer-making or as
weaning foods for infants. Farmers in a wide range of agro-ecologies of eastern and
southern Africa grow small plots of sesame as a source of cooking oil, which otherwise
represents a major household cash expenditure.
The interaction of opportunities and constraints that farmers manage creates the
resultant farming systems that embody the use of all available resources human,
ecological, genetic, and other for achieving food security. Researchers have at various
junctures attempted to understand this full picture of the farming system through
extensive interaction with farmers prior to intervening through research initiatives
(Hildebrand, 1981; Merrill-Sands, 1986).
The complexity of farmers decision-making environments can be startling. Table
2.4 shows the agricultural calendar prepared by farmers and extension agents in Tete
Province, Mozambique (Buhr, 1990). A tremendous amount of information can be
inferred from the chart, which depicts, for example, what some agricultural economists
have long asserted about African farming systems, namely, that labour is often a
constraint in modifying existing cultivation practices (Barker and Cordova, 1978;
Hildebrand and Poey, 1985). While labour shortages are obviously apt to exist during
the September to November planting period, additional shortages can occur during
much of the rest of the year, as well, including during the time of weeding and harvest,
when certain niche crops and second seasons (or relay crops) of main crops must
be planted. These labour shortages often lead to late planting in large parts of Africa.
This, combined with the existence of a recurrent hunger period prior to the main
harvest season, gives rise to the intense interest farmers in Mozambique and
elsewhere have shown in earlier-maturing maize. Early-maturing varieties can also lead
to the introduction of a second relay crop, which can be grown on residual moisture,
thus permitting a broad intensification of farming systems (Haugerud and Collinson,
1990).
While such complexity can appear overwhelming to researchers attempting to make
contributions through the transfer of improved technologies, experience suggests that it
is just this level and type of information that is needed in targeting different agroecologies from a limited number of research sites, as in the case of plant breeding. These
principals are explored in greater detail in Chapter 4.
14
pea
Chapter 2
The Challenge
pea
Okra
15
16
Chapter 2
Because small-scale farmers in Africa cultivate largely unimproved (i.e. unterraced,

non-irrigated, and undrained) fields, they have deployed a rich mix of crops, each
bearing an adaptive advantage within some niche on the farm. The implication is that
farmers of a given agro-ecology seeking to improve their overall productivity may be in
need of improved, adapted cultivars of several crop species. Likewise, significantly
improving any one crop may be of great benefit to one group of African farmers but of
little importance to another (see Plate 3).
While household economies and farm-level variability create crop deployment
patterns at one level, large-scale environmental variation on the African continent
creates crop deployment trends on a far wider scale. Understanding these trends can be
of use in devising strategies for agricultural research at national and regional levels.
Figure 2.1 shows the geographical distribution of production of staple crops within the
major climatic zones of Africa.
As would be expected, the distribution of crop species across differing geographical
regions varies considerably. Within a region, however, priorities can be relatively easily
identified. In order to significantly affect household food security status of rural
populations, efforts focusing on a given region obviously must focus on a range of crops
used extensively on farms within that region.
Fig. 2.1. Per capita production (in kg) of primary crops in sub-Saharan Africa. Ma,
maize; R, rice; S, sorghum; Mi, millet; Ca, cassava; B, banana; Co, cowpea. Source:
FAO (2000).
17
The Challenge
A major impact on food security in Africas semi-arid sahelian zone will require
inclusion of drought-tolerant crops such as sorghum, millet and cowpea.
For humid, lowland regions such as coastal West Africa and the Congo Basin,
strategies should include a focus on cassava, with important, secondary efforts on
maize and rice.
Sorghum and millets, formerly major crops in eastern and southern Africa, are
losing acreage and would appear to have decreasing importance in this region.
Today, strategies for most rural populations of eastern and southern Africa need to
focus primarily on maize, with important, secondary efforts on cassava, common
bean and banana.
Grain legumes remain important in the diets of people in all areas except the Congo
Basin, although their inherently lower yield prevents them from competing with
productivity levels of starchy crops. This translates to a focus on cowpea in lowland
areas and common beans in mid-altitude and highland areas.
90
80
70
60
50
40
30
20
10
0
Fig. 2.2.
M
ille
t
rg
hu
ic
So
at
he
M
ai
ze
Asia
Africa
Crop consumption
(kg person1 year1)
Dealing with environmental variation requires a strategy which encompasses the genetic
challenge (namely, introgressing target traits useful to a range of crop species into a range
of crop genetic backgrounds) (Buddenhagen and de Ponti, 1983) and the institutional
challenge (working through structures which link differing national programmes facing
similar crop improvement tasks). However, through better understanding of the
prioritization given by farmers to different species and traits within those species, it
should be possible to develop coherent, national strategies for improving the genetic
basis for crop production for a range of species.
Species selection by farmers across various subregions is reflected in consumption
patterns for Africa as a whole. Figures 2.2 and 2.3 show figures for per capita consumption of the principal food crops in Asia and Africa during 1997. The data reveal the use
of a wider range of food crops in Africa. Consumption of only two crops rice and
wheat accounts for 70% of non-animal food consumption in Asia. Meanwhile, the
consumption of Africas four most important crop-based food products wheat, maize,
banana/plantain and cassava accounts for only 67% of its total, with much of the
wheat being imported.
The broad implications for crop improvement strategies in the two regions are
obvious. Whereas Asias struggle largely hinged on the ability of researchers and farmers
Cereal crop consumption trends in Asia and Africa, 1997.
18
Chapter 2
80
Asia
Africa
Crop consumption
(kg person1 year1)
70
60
50
40
30
20
10
es
ls
n
ai
nt
la
/p
Pu
s
m
Ya
na
na
Ba
Po
Sw
ta
to
ee
tp
ot
at
o
C
as
sa
va
Fig. 2.3.
Consumption trends of selected non-cereal crops in Asia and Africa, 1997.
to devise more productive rice and wheat-based farming systems, in Africa, broad-based
food security will require sustainable productivity increases within its respective agroecological systems based on maize, sorghum, cassava, millet, rice, pulses and bananas,
among other crops. Consequently, this book focuses on seven priority food crops of
sub-Saharan Africa. In Asia, the initial success of modern varieties of irrigated rice and
wheat was followed by success in developing varieties for many rain-fed areas devoted
to these and other crops. Most of Africa, however, is characterized by farming conditions that have reduced or delayed the impact of genetic improvement found in
other parts of the world. While all the major regions of the world include areas
with these sets of conditions, in Africa, they dominate. As indicated by Byerlee (1996),
these include marginal crop production areas, areas with very poor infrastructure, and
areas where quality traits outweigh the yield advantages of improved varieties, among
others.
An observational accounting of land use patterns among small-scale farmers in
Africas little-modified agricultural landscape indicates that crops are employed largely
according to their ecological niche. Issues such as temperature, natural drainage, rainfall
patterns, soil fertility, and pest and disease occurrence, to a large extent, govern which
crops can be used where. In the tropics, the temperature regime is mainly influenced by
elevation. Most attempts at classifying cropping systems have focused on this factor. In
the following, an attempt is made to offer descriptions of the trends in agricultural land
use in differing environments in Africa.
Lowlands. Valley bottoms of lowland humid environments are widely sown with rice. In
off-seasons, raised beds often produce sweet potato. In sloping areas, cassava and upland
rice are grown on highly leached soils. As rainfall decreases, soil phosphorus levels
increase and maize can be grown. Semi-arid lowland zones are dominated by sorghum
and millet cultivation. Cowpea is the most important pulse crop grown in all welldrained lowland environments. Few pulses grow well in poorly drained lowland
environments, and diets often lack this element.
Mid-altitude zones. Mid-altitude zones of Africa are dominated by maize. In higher
rainfall areas, however, maize productivity is reduced by foliar and storage pests and
diseases and reduced sunlight, and cassava and/or bananas are commonly grown.
The Challenge
19
Cassava is also substituted for maize in very high population zones and areas with very
poor soils. In lower rainfall areas, moisture stress reduces yields and sorghum becomes
the dominant crop. Poorly drained mid-altitude environments are planted to rice. Beans
and pigeon pea are the most popular pulses in mid-altitude zones.
Highlands. Areas above 2000 m except Ethiopia are planted to English potato and
highland bananas, with interspersed plantings of maize and wheat. In Ethiopia,
highland areas are planted to teff. The dominant pulse of the African highlands is beans.
In the Great Lakes regions, beans also supply a large percentage of carbohydrates.
The result is a rich and resourceful utilization of crop genetic resources which contribute economic advantages, nutrition, and cultural significance to rural households
across the continent. Moreover, this diversity, while presenting its own challenges to
crop improvement, holds the promise that improvements made on crop species can be
used in ecologically sustainable ways. As explored in greater depth later in this book,
both the variety of crops and the importance of their adaptation, in turn, highlight
the need for decentralized breeding operations and the continual involvement of farmers in identifying traits and selecting improved crop varieties for multiplication and
commercialization.
Regional crop improvement programmes have made some attempts at understanding the complexity of agro-ecologies in Africa in order to target better their breeding efforts and varietal testing programmes (see Plate 4). The International Center for
Maize and Wheat Improvement (CIMMYT), for example, has recognized nine maize
production mega-environments in sub-Saharan Africa based on three different altitude
ranges in three different ecologies: lowland tropics, subtropical, and highland tropics
(CIMMYT, 1990). The International Center for Tropical Agriculture (CIAT) has
recognized 14 bean production environments in sub-Saharan Africa based on similar
criteria (Wortmann, 1998). Such groupings of environmental variation into aggregated
geographical units is critical for the targeting of crop genetic resources aimed at
achieving continent-wide coverage. However, the level of resolution achieved by such
efforts to date remains imprecise in comparison with its importance in hitting the target
consistently, throughout Africa. Thus, crop-specific agro-ecological analysis remains a
critical area of untapped potential for broadly improving the impact of crop genetic
improvement.
In most cases, this will be a task taken on by the national agricultural research
systems (NARSs), at times reinforced by ecological and geographic information
systems (GIS) studies performed by international agricultural research centres (IARCs)
outreach programmes. Progress has been made in several countries. As an example, the
Kenya Agricultural Research Institute (KARI) recognizes five maize breeding agroecologies in Kenya which are used to focus breeding efforts, depicted in Fig. 2.4.
Researchers in western Kenya have recognized additional subclassifications within the
moist mid-altitude zone, which can be used to add further precision to crop improvement efforts (Amadou Niang, personal communication). Thus, even cursory inventory
of maize agro-ecologies in Kenya may result in six or more broad families of maize
varieties. Kenya represents one of the most intensively studied countries in Africa, and,
perhaps not coincidentally, one where crop genetic improvement has made significant
impact (Gerhart, 1975). Wider and more intensive analysis of agroecologies needs to be
conducted by crop improvement teams in all African countries.
20
Chapter 2
Fig. 2.4. Constraints to maize production in major agro-ecologies (Highlands, Mid-Altitude

Moist, Mid-Altitude Intermediate Moist, Mid-Altitude Dry, Mid-Altitude Intermediate Dry, and
Tropical Lowlands) of Kenya identified by KARI maize breeding teams. DTM, days to maturity.
Trying to make accurate determinations of varietal needs for numerous groups of

African farmers living in widely varied agricultural agro-ecologies is a real challenge.
Without it, however, the chances of success are slim. Experienced crop improvement
specialists in Africa can cite the many new varieties which have been developed for
The Challenge
21
African farmers, but which have never been adopted. Former US Secretary of Agriculture, Clayton Yeutter, writing recently on the biotechnology revolution (ISAAA,
2000), stated:
Newer genetic modifications, impressive as they may be in the laboratory and in the pages
of professional journals, are of little real world relevance unless those desirable traits are
transmitted through seeds with good yield characteristics. Otherwise, farmers in the U.S.,
Africa, or anywhere else, simply will not plant those crops.
Unlike Asia, new crop varieties for Africa cannot be developed based on the assumption
that fertilizer will be subsidized and made available through government programmes.
To be adopted in Africa, new varieties need to be well adapted to local conditions and
provide yield advantages with few external inputs. Recent approaches to breeding
focused on selection under low-input African conditions (Bnziger et al., 1997;
Bahia and Lopes, 1998) have proved effective in identifying varieties with superior
performance under drought and low soil nutrient status. Such adaptation to environmental stress needs to be combined with good levels of resistance to foliar diseases, insect
pests and, in some cases, the ability to grow vigorously during early stages of development to shade out weeds. While landraces will in most cases have reasonable levels of
resistance to all these constraints, for reasons related to co-evolution and the relatively
slow rate of genetic change via mass selection methods performed by farmers, it is
unlikely these levels will match those possible through scientific breeding programmes.
To develop varieties that poor farmers find useful, it is necessary to understand
environmental variation in Africa and listen to farmers advice on issues of growing
environments and household utilization, a topic explored more extensively in Chapter
4. In Africa, perhaps more than in any other part of the world, the science of genetic
improvement must be paired with the art of understanding people and the environment. This will require additional investment in areas which serve to consolidate the
presently diffuse, dispersed base of knowledge on agro-ecologies and crop user systems
in Africa. Recent initiatives such as the atlases on bean, cassava and maize are a good start
in this direction (Carter et al., 1992; CIAT, 1998).
Household preferences, as well, cannot be overlooked in breeding programmes and
consideration should be given to the overall crop usage environment in which the
adoption must take place. Some crop/user system combinations in developing countries
constitute situations where yield advantages of improved varieties can easily be outweighed by the importance of quality traits (Herdt and Capule, 1983). Very poor
farmers often cannot afford to pay for industrial milling services, and must carry out all
processing tasks in the home. Thus, farmer preference for flint-textured maize varieties
among resource-poor farmers in Malawi was key to identifying flint hybrids which
achieved high levels of adoption in the early 1990s (Nhlane, 1990; Smale et al., 1993).
Likewise, food scientists who have analysed sorghum quality characteristics have become
increasingly capable of predicting the acceptability of improved varieties based on the
quality of food products they produce. Studies conducted using sorghum flours from
West, southern and East Africa revealed significant differences in flour texture and total
water content of porridges consumed. Households preferred varieties with high amylose
starch content and low flour lipids and proteins (Fliedel and Aboubacar, 1998). Few
improved varieties have scored high in such tests. Nevertheless, breeders have often
22
Chapter 2
failed to take full advantage of the ability of food scientists or consumers to inform them
of the probable success of their offerings at the household level.
The need to link as much field-based information as possible to crop improvement
programmes argues for a high degree of integration of disciplines and connectivity
between breeders working at international, regional and national levels. Decisionmaking matrices that may seem very complicated to breeders may be a relatively simple
matter for farmers who use crops in various forms everyday. Since many aspects of
adaptation and farmer preference do not relate to expertise commonly embodied within
crop research institutes, adequate linkages need to be established with agencies or
individuals who do embody this expertise, including farmers and NGOs.
2.4 A Seed Sector Dominated by Market Failure

While the trend toward privatization and globalization of the germplasm sector has
undoubtedly resulted in the distribution of better seed-based technologies to farmers in
developed regions of the world, these policy changes will not function to the same extent
in Africa in the short or medium terms. Private seed companies are constrained to
operating in environments where they can make acceptable profits. In Africa, multinational seed companies may be motivated to popularize one or even several highyielding maize hybrids among better-off farmers in favourable areas, but it is less likely
that they will find it profitable to devote significant resources to developing varieties
with the very specific adaptation advantages required by small-scale, low-input farmers.
Even if such varieties enabled resource-poor farmers to double their yields, this would
often mean an increased harvest of only 1 t ha1 or less. The share of increased profits a
seed company might capture from such a modest increase is small in comparison with
profits available in developed regions of the world (Tripp, 2000). Additionally, the
degree of complexity involved in designing a full range of varieties required by different
categories of farmers cultivating farms in very different agro-ecologies further limits
potential profitability.
Low effective demand and relatively small profits available from seed in much of
Africa, in comparison with the rest of the world, have delayed the commercialization of
the seed market. Low rates of economic growth forecast for much of Africa are not likely
to attract large-scale investment from outside the continent of the type needed to achieve
broad coverage of farmers needs for seed. Rather, indigenous seed companies that operate closer to local markets and on lower margins should be considered as a solution with
wider potential. To date, however, little international or national assistance has been
directed at this type of company. The strategy put forward in this book places high
importance on the development of Africas private seed companies.
Regardless of the strategy employed, given the economic realities in Africa and the
difficulties seed companies face in attracting clientele, growth of the seed sector is likely
to be slow and sporadic. The implication is that public sector-based strategies for seed
dissemination will be critical to realizing the benefits of crop genetic improvement in
Africa for some time to come. In fact, the absence of a sufficient effort by either private
or public sector breeding interests has left an enormous gap in the seed supply offered to
African farmers. While things can be done to encourage such investment, alternative
strategies and continued experimentation are needed (see Chapter 6).
The Challenge
23
Finally, in the absence of investment by private seed companies, the rapid-fire,

signalresponse, product-refining process that is the great advantage of distribution
systems conducted via private enterprise will not operate for seed in Africa. Feedback on
performance and preference issues must be gathered through other means. This fact
drastically increases the need for continual participation by farmers in variety refinement
and seed dissemination. It also points to a critical role for research managers who oversee
crop improvement initiatives and are ultimately responsible for ensuring that useful
varieties emerge from such efforts. This challenge is discussed in greater detail below.
2.5 Policies and Institutions are Critical to the Success of Crop

Improvement
Policies that favour food security like those which favour education and health
provide a foundation for development and the passing on of these basic human needs
to successive generations (Sen, 1981). Of these three commonly cited priorities for
development, however, the most basic and immediate is security against hunger. In
Africa, where none of the three needs has as yet been broadly secured for society, equity
arguments can be advanced that food security ranks as the most essential priority. Yet
public and donor funding for agriculture has lagged far behind other priority sectors in
Africa. A recent review of public spending in Uganda showed that agriculture accounted
for just 4% of total expenditures from national and donor agencies sources, compared
with 11 and 20%, respectively, for health and education (Uganda Ministry of Finance,
2000).
Few African countries have prioritized food security through development of the
agricultural sector. While speeches made by African leaders are invariably peppered with
references to freedom from hunger and development of the economy through technological advance, public sector spending on agricultural research and extension in
Africa declined from 1981 to 1991 (Pardey et al., 1997). African governments have
received little real encouragement to develop their agricultural sectors from Western
governments, several of which have de-emphasized the agricultural portfolios of their aid
packages to Africa during the 1990s. The US government made payments to farmers of
$7.3 billion in 1995 and 1996 (USDA, 1998). Meanwhile, the prevailing belief is that
the agricultural sector in Africa should develop itself.
Thus, a fifth challenge is situated within the institutional framework of crop genetic
improvement: how individuals, groups, and institutions are organized to achieve results
in relation to goals which require collaborative arrangements, resulting in a physical
product which is usable by farmers. Overall institutional or departmental performance
influences significantly the output of breeding teams and is generally unrelated to the
academic preparation of the individuals involved. The area of public policies and
institution performance attains greater importance in relation to the many regulatory
issues and intellectual property rights attached to techniques and products of
biotechnology.
At a national level, there is a need for crop-based strategies for genetic improvement
that make use of the full range of scientific capacity which can be applied. National
breeding programmes are the front lines of public sector breeding in Africa. For many of
the self-pollinated crops, and for open-pollinated varieties (OPVs) of cross-pollinated
24
Chapter 2
crops, national programme varieties are likely to continue to be the dominant means by
which genetic improvements move out to small-scale farmers in Africa. The efficiency of
this process is highly sensitive to the science policy of each institution and the regulatory
framework governing plant varieties.
National and international policy on intellectual property and plant variety
protection is being debated in various corners of the globe, and outcomes are difficult to
predict (Barton, 1998; Erbisch and Maredia, 1998; Koo and Wright, 1999). Indeed,
confusion exists in many cases regarding what forms of biological property can be
protected where and for what purpose. By some interpretations, the trend would seem to
threaten access by developing countries to genetically engineered crops and, quite possibly, to other biotechnology applications as well, as donor agencies reduce support for all
but conventional science applications. The widespread patenting of breeding materials
by both private companies and public universities in the USA and Europe has inevitably
reduced the flow of germplasm from those regions to Africa. However, even at this early
stage, progress has been made towards sharing critical intellectual property that would
seem to counter that argument (Mugo, 2000). What seems clear is that developing
countries require advocates who work in the interest of broadening access by them to
emerging technologies, and not simply the products of those technologies.
2.6 The Soil Fertility Problem

Better varieties, of course, are only part of the challenge. Of equal or greater importance
to realizing the full productive potential of crop plants is the supply of plant nutrition
through healthy, fertile soil. Traditionally, major increases in productivity have mainly
come about as a result of a combination of improved production systems (irrigation,
drainage, improved cultural practices, introduction of fertilizers) and the introduction of
more efficient varieties (Matlon and Adesina, 1991). The introduction of improved
wheat and rice varieties in Asia, for example, coincided with wider availability of
inorganic fertilizers and irrigation (Herdt and Capule, 1983). Cheap and widely
available inorganic fertilizer in Nigeria facilitated rapid expansion and intensification of
maize production following the introduction of improved, adapted varieties in the
1970s (IITA, 1995). More recently, improved maize varieties and increased fertilizer
applications (encouraged by credit facilities) in Ethiopia during the period 1994 to 1996
produced a dramatic, 31% increase in average yield (Quinones et al., 1997) (see Plate 5).
Fertilizer consumption in Africa has stagnated, even while land brought under
cultivation has increased dramatically. From 1970 to 1982, there was a slight but steady
increase in fertilizer consumption in Africa, but there has been no increase in total
consumption since then (Fig. 2.5). Meanwhile, the continent has added 270 million
persons. The reduced fertilizer use per capita has been made up through bringing
new land into cultivation and by mining soils of their nutrients through continual
cultivation without replenishment of nutrients. As a result, net nutrient outflows per
year in Africa are estimated at 63 kg ha1 year1 on average (Debrah, 2000). Lower input
use in Africa is probably substituted in part by added labour input, without which, yield
levels would most likely be even lower.
Herein lies an enormous problem. Farmers in sub-Saharan Africa use by far the least
amount of fertilizer in the world. In 1993, average application of total nutrients in Africa
The Challenge
25
Fig. 2.5. Total fertilizer consumption in Africa (excluding Egypt and Libya). Source:
IFA (2000).
was 10 kg ha1, compared with 83 kg ha1 in other developing regions (Heisey and
Mwangi, 1996). Moreover, subsidies which formerly encouraged the use of fertilizers
were largely removed starting in the early 1980s, and have not been re-applied. The
halt in increase of fertilizer applications in Africa shown in Fig. 2.5 coincides nearly
exactly with the implementation of those policies. Studies conducted by Holden and
Shanmugarathan (1994) and Bumb and Baanante (1996) both showed that higher
fertilizer prices following the removal of subsidies led to reduced application of
inorganic fertilizers in several African countries.
Today, domestic fertilizer prices in Africa are far above world prices. While world
urea prices in 2000 ranged between $80 and 100 t1, urea prices in several African
countries range from $400 to $842 t1 (Debrah, 2000). At prevailing fertilizer costs and
farm-gate prices for commodities, the economics of fertilizer use are not favourable.
Extensive analysis of fertilizer responses, fertilizer prices and producer prices for maize in
Malawi from 1994 to 1997 resulted in researchers recommending zero application of
ferilizer on maize produced for market in 34 out of 41 agro-ecologies (Benson, 1997). In
summary, therefore, at current fertilizer prices in Africa, there is little perspective for
fertilizer application among African farmers to increase (Sanchez et al., 1997).
Privatization of agricultural input markets has removed much of the flexibility governments formerly availed themselves of in encouraging the use of agricultural inputs,
including fertilizers (Cromwell, 1996). To an increasing degree, therefore, poor farmers
are left with fewer options.
The response among soil fertility researchers to reduced applications of inorganic
fertilizers has been to focus on the cycling of nutrients in low-input systems and the use
of lower-cost methods of adding nutrients, such as legume rotations, green manures,
and improved fallows (Sanchez et al., 1997). While significant amounts of nitrogen can
be added to soils through the cultivation of green manure crops, the limitations of
this method for maintaining soil fertility must not be overlooked. Crop recovery of
nitrogen contributed by the leaves of leguminous plants is generally lower (1030%
recovered) than that contributed by inorganic fertilizers (2050%) (Palm, 1995).
More importantly, legume biomass contributes little of the phosphorus required to
complement the nitrogen and potassium contributed by such additions. For example,
26
Chapter 2
cover crops of velvet bean (Mucuna pruriens) and Crotolaria ochroleuca contribute on
average 3542 kg nitrogen and 79 kg potassium per tonne of biomass, but only
1.62 kg of phosphorus (Palm et al., 1997). A substantial crop of either of these green
manures of, say, 5 t ha1 would thus yield a maximum of 10 kg P ha1 far below the
minimum required for a 2 t ha1 crop of maize.
Humankinds dependence on the HaberBosch process of synthesizing nitrogen for
use in producing its food requirements has been extensively analysed by Smil (1991),
who concluded that no alternatives to the use of inorganic nitrogen currently exist
for densely populated developing countries. While many African countries have low
absolute population:land ratios, Binswanger and Pingali (1989) revealed that in reality,
due to highly uneven resource endowment in Africa, many countries have high effective
population densities. Heisey and Mwangi (1996) have also described many African
countries as land-scarce. These considerations, coupled with the recognized labour
constraints many small-scale farmers operate under in Africa, argue for continued efforts
aimed at making fertilizers broadly more accessible to small-scale farmers. In an analysis
of the factors leading to low fertilizer usage among small-scale African farmers, Debrah
(2000) identified as critical factors: high fertilizer cost, low farmer profitability, unequal
access, low credit availability, and socio-economic factors related to farmers attitudes
toward fertilizer use.
The Sustainable Community-oriented Development Programme (SCODP) is an
NGO aimed at improving access to fertilizer and seed among farmers in Siaya District of
western Kenya. The organization promotes the use of these inputs and improved crop
management practices through 11 small farm input shops located in rural market
centres. By breaking large (50 kg) bags of fertilizer into units ranging from 100 g to 2 kg,
the organization became the largest supplier of fertilizer in the district in only 4 years.
Moreover, the organization was able to operate at a profit, and projected growth in sales
of over 400% by 2002 (SCODP, 2000). Such initiatives SCODP is supported by several donor agencies may reveal how fertilizers can be supplied at more affordable prices
than at present. Nevertheless, the scale of such activity remains very small. Only 10% of
farmers in Siaya currently use fertilizer, and fertilizer application rates in the SCODP
project area remain low. Even if SCODP were to achieve projected growth in sales, average fertilizer application rates in the District would remain at 6 kg ha1.
Finally, the case of broad provision of small quantities of fertilizer to African
farmers deserves attention. Between 1999 and 2000 in Malawi, the Starter-Pack
programme distributed 2.86 million packages to 2.4 million farm households (indicating full coverage but some errors in distribution as well). Packs contained a range of crop
species, depending on the agro-ecology in which they were targeted, but generally
consisted of 2 kg of cereal crop seed, 2 kg of legume seed, and 10 kg of fertilizer. Average
household harvest increased from 1087 kg ha1 to 1904 kg ha1, resulting in an average
increase of 96 kg per household (Mann, 1999). Distribution of the packages boosted
national maize production by 25% and extended household food sufficiency from 6.1 to
8.7 months (Levy et al., 2000). However, the sustainability of this programme is already
being challenged.
Projected low rates of economic growth coupled with population increases in
sub-Saharan Africa during the coming decade (Conway, 1997) carry important implications for technology development and application among small-scale farmers. If, as projected, fertilizer applications in Africa are likely to remain low for the foreseeable future,
27
The Challenge
the primary contribution from plant breeding will come from efforts aimed at making
crop plants more productive and dependable within low-input, limited-infrastructure
production environments. Improved nutrient management based on accessible, practicable methods of returning nutrients to the soil can make land more productive, and
more resilient crop plants can help produce better harvests; however, it is likely that
Africas crop production environment will generally remain low in fertility.
Low soil fertility and low fertilizer use, combined with a lack of irrigation, very low
pesticide use, and low rates of tractor use in Africa, effectively mean that crop genetics
and improved planting material remain as one of the most effective means by which
African farmers can be assisted. The absence of these productivity-enhancing, agroecology-homogenizing factors of production makes the task of crop improvement more
complex and more difficult in Africa than in other parts of the world. But low input use
does not reduce the importance of crop genetic improvement in the lives of Africas rural
poor, rather, it raises it.
2.7 What, Then, is Needed?

Quite obviously, improving all crops for all possible traits in all agro-ecologies in Africa
is not feasible in the short or medium terms. One obvious difficulty with agro-ecological
approaches is in achieving the scientific and technical acuity at all the levels necessary
broadly across the continent. Without making a detailed analysis of the number of
breeders and other crop scientists currently employed at the national and international
levels in Africa, it is safe to say that there are currently insufficient numbers to apply
systematically agro-ecology-based strategies for crop improvement. However, if one
accepts that the biophysical challenges in Africa are great, and that new science
combined with new methods for understanding and deploying the varieties that farmers
desire does create a new realm of the possible in Africa, then a concerted, worldwide
effort at improving overall agricultural productivity in Africa, including the genetic
performance of Africas food crops, is justified. Moreover, simple calculations of
the financial requirements for funding the national component of the challenge
(shown in Chapter 4) reveal that making this effort is not necessarily prohibitively
expensive.
Making use of lessons learned over the past decade of working closer with farmers
in Africa, and combining those lessons with breakthroughs made in genetics, the
following steps can and should be taken.
Identify traits that genuinely reduce productivity at the small-scale farmer level in
a wide range of environments in Africa and for which a (full or partial) genetic
solution may be possible.
Understand and conceptualize the various agro-ecologies within which new crop
varieties need to be deployed, and determine the priority constraints, adaptation
advantages, and user preferences which need to be targeted in developing new
varieties.
Identify individuals and institutions capable of making meaningful contributions
to a genetic solution to each trait.
28
Chapter 2
Provide opportunities for interaction between groups of scientists, aimed at

identifying strategies for developing the available genetic resources into a farmerusable product.
Implement strategies via vertically integrated crop improvement initiatives which
incorporate biotechnology, breeding, and seed systems.
Disseminate new technologies via farmer-focused, agro-ecologically informed
initiatives which consider the full range of agronomic and genetic technologies
which can benefit farmers.
Due to the complexity and size of the challenge implied, a long-term commitment is
required. Only through successive research steps, each carried to the next stage, can
progress remain on track and capable of delivering needed products.
Because biotechnology, breeding, and seed systems form relatively discrete
functions within the crop improvement process, they can, to a certain extent, be
considered as separate activities, each with its own sectoral strategy. They are considered
as such in Chapters 4, 5 and 6 of this book. First, however, it is worthwhile to document
the level and the kind of food that Africa needs and to understand how improved farmer
productivity might contribute to a solution. Chapter 3 presents a brief look at the nature
of food scarcity in Africa.
The Roots of Hunger
Rural communities in Africa are under pressure on several fronts. Profits from farming
at the current, low levels of productivity, are too small to allow farmers to reinvest in
their land and maintain sustainable production systems (Eicher, 1990; Blackie, 1994).
Meanwhile, continual increases in population have depleted the available resource base
and eroded many social entitlements which hitherto provided for a state of equilibrium
in rural areas of Africa (Lele, 1989). Finally, steady increases in agricultural productivity
in developed regions of the world (increasingly facilitated by biotechnology), combined
with persistent payments of massive subsidies to North American and European
farmers, have continued to push world grain prices downward, making it increasingly
difficult for marginal land farmers in developing countries around the world to operate
profitably (FAO, 2000). Rural areas by definition offer a limited set of economic
alternatives to agriculture, and Africa has attracted very little direct foreign investment
to create new jobs, even in urban areas. As a result, economic growth in rural areas has
been insufficient to offer alternative means of employment for the rural poor (Eicher,
1990; Oyejide, 1993), and agriculture remains their only real option for survival and
income.
Africa has the highest percentage agricultural population and the second highest
cultivated area in the world (Table 3.1). However, average cereal yields are the worlds
lowest, and less than half of Asia and Latin America.
Another reason for rising hunger in Africa during the past few decades is high rates
of population growth. In 1970, Asia, Latin America and Africa had similar rates of
population growth (Fig. 3.1). Yet population growth in Asia began decreasing rapidly
thereafter and by 1978 had decreased by 25%. Latin American population growth rates
declined more slowly, but by 1995 fell by nearly 20%. In Africa, the rate of population
growth rose sharply between 1970 and 1995. At present, the population growth rate in
Africa is nearly double that of Asia.
A high population growth rate is especially injurious in countries with low or
negative economic growth, where the number of lives to support simply outstrips the
rate of appearance of new opportunities for adding value within the household. In
29
29
Chapter 3
30
Chapter 3
Table 3.1. Key agricultural indicators from agricultural sectors of selected regions
of the world.
Africa
Asia
Latin America
North America
Cultivated area
(million ha)
Agricultural
population
(millions)
Percentage
agricultural
population
Cereal yields
(kg ha1)
1070
1604
747
494
416
1922
111
8
57.8
55.9
23.3
2.6
1107
2886
2545
4189
Source: FAO (1998).
Fig. 3.1. Comparative population growth rates in Asia, Latin America and Africa,
19701995. Source: FAO (2000).
sub-Saharan Africa between 1987 and 1997, the average annual percentage increase in
gross national product per capita was 0.7% (World Bank, 1998). Meanwhile, per
capita foreign direct investment in Latin America in 1998 was $126, while in Asia it was
$35. Africa lagged far behind in attracting investment, at only $6.85 per capita (see
Table 3.2).
High levels of population growth, combined with low or negative economic
growth, have led to reduced access to food among Africas inhabitants. Although Asia,
with 70% of the developing worlds total population, has far greater numbers of people
who are undernourished, sub-Saharan Africa has almost double the percentage (33%
compared with 17% in Asia) of hungry people (FAO, 2000). Within Africa, eastern and
southern Africa account for the greatest number of undernourished people. Per capita
food consumption in these regions has decreased during the period 1980 to 1995
(Fig. 3.2). In eastern Africa there was a 5.5% decrease in consumption and in southern
Africa a 9.3% decrease.
Child nutrition and growth rate is an especially relevant indicator of socioeconomic development. A recent study by the United Nations and the International
Food Policy Research Institute (IFPRI) revealed that 35.2% of children in sub-Saharan
30
Chapter 3
31
The Roots of Hunger
Table 3.2.
Key economic indicators, developing regions and the world, 1998.

Population
(millions)
Region
Africa
East Asia and
Pacific
Latin America
and Caribbean
World
GDPa
GDP per capita
(billions of $)
($)
Foreign direct
investment
(billions of $)
6,642.3
36,332.7
,518
4.4
1,800.3
1,900.3
1,056
64.2
6,509.2
6,000.3
2,100.3
30,200.3
4,124
5,033
64.3
619
Source: The World Bank (2000).

aGross domestic product.
Fig. 3.2. Africa: per capita consumption of cereals plus roots and tubers plus pulses,
19801995. Source: FAO (2000).
Africa suffered from stunted growth. The figure when considering all developing
countries was 32.5% (UN/IFPRI, 2000). In East Africa, 48.1% of children exhibit
stunted rates of growth, the highest in the world. High birth rates in Africa mean that
the total number of children with stunted growth has increased rapidly over the past two
decades. Africa is the only developing region in the world where the percentage of
underweight children is increasing (Fig. 3.3).
Global statistics do not adequately explain causes of hunger. Regional or subregional trends in production and imports often mask severe food shortfalls within a
given country or areas of a given country. Table 3.3 shows the trend in production and
imports of the primary staple, maize, in selected countries of eastern and southern
Africa. Percentage consumption made up of imported grain tripled in Kenya between
the 15-year period up to 1990 and the 5-year period following 1990. Malawi has since
recorded surplus production of maize, thanks in large measure to a countrywide
initiative for free distribution of agricultural inputs, but such subsidies are unpopular
with donors and are likely to be terminated.
31
Chapter 3
32
Chapter 3
Fig. 3.3. Prevalence of underweight children in Africa and Asia, 19802005. Source:
UN/IFPRI (2000).
Table 3.3. Average annual maize production and import trends in Kenya, Malawi
and Zimbabwe during 19761990 and 19911995
Imports
(1000 t year1)
Production
(1000 t year1)
Import % of
production
2107
1331
1613
3.2
2.7
1.7
2524
1387
1425
9.4
24.0
24.6
19761990
Kenya
Malawi
Zimbabwe
68
36
28
19911995
Kenya
Malawi
Zimbabwe
238
333
351
Source: FAO (2000).
Among the vast number of isolated, agriculturally based villages throughout Africa,
hunger is often directly tied to subsistence farming systems and the attendant cycles of
production, harvest and off-season activity. Hunger periods in Africa periods during
which there is very little food left in the granaries or in the ground (in the case of tuber
crops) normally occur during the months prior to harvest of the main crop. During
these periods, hunger can be abated by growing short-cycle annuals (often cowpea or
common beans) or even earlier-maturing varieties of the main staple, such as maize
(Chapman et al., 1997). Identifying the most appropriate alternative, however, requires
plant introductions and extensive on-farm research.
The impact of low technology adoption on productivity and land use has been
immense. While agricultural productivity in Africa as measured by cereal yields has
32
Chapter 3
33
The Roots of Hunger
Fig. 3.4. Cereal yield trends, Asia compared with Africa, 19701997. Source: FAO
Internet Database.
increased by approximately 18%, area under cultivation (FAO, 2000, data not shown)
has increased by 32%. During the same period, cereal yields in Asia increased by 45%,
while area cultivated increased by only 11% (Fig. 3.4).
Food aid is often cited as a logical, if partial, solution to food insecurity in Africa
and other food-insecure regions of the world. In 1999, the USA provided approximately
58% of total food aid shipments worldwide. In that year, the USA shipped a total
of 9.84 million Mt of food aid to other countries. Of this, 1.14 million Mt,
approximately 11%, went to Africa (USAID, 1999). The total value of this food was
approximately $467 million. Food aid shipments to Africa from the European Union
during the same period were approximately 140,000 t (Walter Middleton, personal
communication).
Some food aid (especially emergency food aid, which accounts for a third of Africas
total from US sources) represents a key component of food security because it is
channelled into areas and population groups with crucial needs. Nevertheless, with an
estimated, total cereal harvest in Africa during 1999 of 75.54 million Mt (FAO, 2000),
and additional, estimated total harvest of cassava and pulses totalling 27 million Mt,
total food aid shipments to Africa from the USA in 1999 accounted for only 1.1%
of food consumption. Thus, food aid represents only a very partial solution to food
insecurity in Africa, and cannot be depended on over the long term.
Commercial importation of cereals plus pulses and root crops into sub-Saharan
Africa in 1998 totalled approximately 16.1 Mt (FAO, 2000), equivalent to 15.7% of
total consumption of these commodities. Taken together, grain imports and food aid in
1999 thus supplied approximately 16.7% of total food consumption. While both food
aid shipments and commercial imports are subject to fluctuations, the likelihood that
either will begin to supply the major portion of the food consumed in Africa is low.
With harvests currently accounting for roughly 83% of consumption, overall food
security in Africa remains solidly dependent upon local agricultural production.
33
Chapter 3
34
Chapter 3
Several writers have chronicled Africas declining food security (Pingali et al., 1987;
Conway, 1997; Ravaillion and Chen, 1997), and it is not the purpose of this study to
present an extensive analysis of the current status of Africas food security. Moreover,
while this book focuses on ways to increase food production, it is understood that
achieving this goal will not solve the problems of hunger related to lack of access to land
or to poverty in urban areas. Rather, the book focuses on one, perhaps significant
component of a solution to low productivity among small-scale farmers in Africa, that of
more productive and more resilient crops.
Recently, several authors have argued that hunger among the poor is not a food production problem, but rather one resulting from a complex set of interrelated factors such
as markets, roads, prices and information, among others (Moore Lapp et al.,
1998; Altieri and Rosset, 1999). While all of these factors undoubtedly play a part
in improving the earning potential and food security of the rural poor in Africa,
observations and discussions with farmers reveal that there are also many millions of
rural poor whose food intake and farm income is limited by the size of their harvest.
Agriculture accounts for upwards of 80% of total employment in many countries of
sub-Saharan Africa. Higher farm yields among these people, made possible in part by
more productive and resilient crops, will inevitably mean healthier, more productive
lives and greater hope for the future. However, if farmers are to improve their levels of
productivity for the benefit of themselves and the rest of the continent, it is critical that
improved seeds be provided.
34
Chapter 3
Breeding Between an Art and

a Science
4.1 Overview
Crop improvement through the continual selection of better food-producing plants
is an activity as ancient as agriculture itself. Through several stages beginning with
the mass selection practices begun by the worlds first farmers, through the breeding
revolution made possible by the discovery of Mendelian genetics, to today, as the world
begins to make use of the first products of biotechnology, the improvement of crop
plants has remained one of humankinds most engrossing vocations.
One of the best-known results of the crop improvement process has been an
increase in the upper threshold of productivity in harvested portion per unit of land area
planted. Increasing the yield potential of food crops has been one of the most important
factors in the steady reduction of food costs throughout the world and the freeing of
human and financial capital for other endeavours.
The dramatically increased yield made possible by re-structured, Green Revolution crop varieties has benefited literally billions but has also proved to be a controversial outcome. Agricultural productivity increases in most regions of the world have
resulted in lower food prices, making food more accessible to the poor and contributing
to increased life expectancy and providing a platform for broader, economic
development (Lipton and Longhurst, 1989; Renkow, 1993; David and Otsuka, 1994).
The opportunity to profit from higher-yielding varieties initially prompted wealthier
farmers on better lands to invest in more purchased inputs and irrigation, often
leading to changes in the welfare of poorer farmers (Frankel, 1971; Griffin, 1974). It has
also led to continuous cropping and the build-up of pests and diseases (Khush, 1990).
People farming poorer lands faced a greater risk of crop failure, obtained smaller yield
increases, and were more reluctant to make such investments (Lipton and Longhurst,
1989). Over time, however, higher-yielding varieties with greater resistance to pests,
pathogens, and other stresses were developed and spread throughout much of Asia,
benefiting farmers in marginal and prime areas as well as consumers, through lower
35
35
Chapter 4
36
Chapter 4
prices. In Africa, there is essentially no irrigation and inputs are expensive, so more
resilient crops will be needed on both good and marginal lands.
Small-scale farmers who depend in part on their own produce for nutrition and
livelihoods may profit more from crop improvement techniques which enhance and
stabilize yields by limiting losses than from higher yield thresholds (Buddenhagen, 1983;
Widawsky and OToole, 1990; Herdt, 1991). Yield stabilizing traits come in many
types, but usually translate to an increased ability of plants to resist or tolerate biological
and environmental stress factors such as pests, diseases, drought and low soil fertility.
Because the farmers most in need of such technologies are those least able to pay for
them, making crops more resilient to marginal conditions remains a neglected area, both
for science and the crop improvement sector, broadly speaking. In part because of weak
public breeding programmes and in part because of the transfer of priority for crop
genetic improvement to the private sector, the potential of crop genetic improvement in
mitigating against low productivity in Africa, where the private sector has not responded
as expected, remains under-exploited.
In this chapter, we attempt to build an argument for increased efforts at
developing crop varieties for African farmers which perform better under the marginal,
low-input farming conditions that prevail across much of the continent. Differing both
in scope and methodology from breeding initiatives which resulted in the Green
Revolution, the proposed approach would make use of advances both in biotechnology
and in farmer-driven, participatory methods of plant breeding described below. The
central theme, however, is that the first, critical step in making use of either of those
and other innovations is the development of plant breeding capacity across the
continent. As noted elsewhere in this study, improved, adapted crop varieties for African
farmers are not being proposed as a substitute for the numerous, other components of
improved, agricultural systems, but are rather intended to serve as one component of
those systems.
4.2 Aims and Contributions of Plant Breeding

Plant breeding is a process of identifying and managing the full range of plant traits
(plant growth characters which are controlled by their genes) in order to produce better
combinations of those traits in new and more valuable crop varieties. Breeders accomplish this mission by controlling pollination events and making specific combinations of
cross-pollinations and self-pollinations. Although the selection strategies employed by
breeding programmes are numerous and complex, most plant breeding is aimed at
improving food crops in four basic ways.
1. Restructuring of the plant, often in conjunction with time to maturity, whereby an
increased portion of assimilate is directed to usable plant structures such as seeds or
fruits.
2. Altering the time to maturity, through management of crop responses to photoperiod.
36
Chapter 4
37
Breeding Between an Art and a Science
3. Introgression of resistance and/or tolerance genes which allow the plant to perform
better in the presence of biotic (i.e. pests and diseases) or environmental (i.e. temperature, moisture, or nutrient availability) constraints.
4. Heterosis, which occurs when genetically diverse parent plants are combined to
form more vigorous hybrids.
Breeding programmes develop both open-pollinated varieties (OPVs) and hybrid
varieties. The distinction between OPVs and hybrid varieties is an important one. OPVs
are varieties resulting solely from the recombination and selection of plant traits within
segregating populations (following cross-pollinations, plant genes segregate in random
fashion during successive generations, making the final outcome of any given crosspollination difficult to predict, and forcing breeders to follow large numbers of segregating plant families). OPVs are the end-products of these segregating populations once the
variety has stabilized. Hybrid varieties are very different. Hybrids are the generation of
seed produced by crossing non-segregating (fixed through repeated self-pollinations)
parental lines. For reasons that are not completely understood, the crossing of contrasting, fixed parental lines gives rise to a generation of seed that is more vigorous and higher
yielding. Once a hybrid has been grown in the field for one season, however, the genes in
the harvested seed are beginning to segregate, and the yield advantage is progressively
lost. Therefore, while OPVs can be saved from year to year without loss of performance
of the variety, the same is not true for hybrids.
Examples of each of these aspects of crop improvement are evident in different
periods of agricultural advancement in various parts of the world, beginning with
the worlds first breeders, the first farmers (and most likely, women). As each plant
cultivated by the early farmers contained a set of genes controlling a range of plant traits,
certain traits linked to survival and productivity such as plant vigour, disease resistance,
seed number and seed viability would naturally have been enhanced through successive
generations of sowing and harvest. This process, sometimes referred to as mass
selection, continues today wherever seeds from previous harvests are saved and
replanted the following season.
It is known that farmers of Assyria and Babylonia artificially pollinated date palms
as early as 700 BC (Poehlman, 1979). The first recorded observation of sexual reproduction in plants, however, was made by Camerarius, a German, in 1694, and the first
systematic studies of artificial plant hybridization were done by a fellow German, Joseph
Kolreuter, from 1760 to 1766. During the 1800s, plant breeding evolved gradually
through studies performed by such individuals as Thomas Andrew Knight, president of
the Horticultural Society of London from 1811 to 1838, by Le Couteur and Shirreff in
England, and by Louis Leveque de Vilmorin and his son in France in the middle and late
1800s. Gregor Mendels classic studies on inheritance of genetic traits in garden pea,
first reported in 1866 but rediscovered by the scientific community in 1900, led to the
establishment of the science of genetics.
In the USA, commercial maize hybrids were first made available to American
farmers in the 1930s (Crow, 1998). The development of high-yielding hard red winter
wheat varieties in the USA is traced to the introduction into Kansas of a Turkish variety,
Turkey Red, by Russian Mennonites in 1873. The hard red spring wheat variety
Marquis originated from a cross made by Dr C.E. Saunders in 1892. Much higher
wheat yields were made possible by the introduction into the USA from Japan of the
37
Chapter 4
38
Chapter 4
short, semi-dwarf variety of wheat, Noren 10, in 1948. Norman Borlaug combined the
Noren 10 germplasm with Mexican wheat varieties that led to breakthroughs in
Mexican wheat yields in the 1950s. Early phases of Asias Green Revolution of the 1960s
and 1970s were based largely on the restructuring of rice and wheat plants through the
use of dwarfing genes (House, 1996). Rapid increases in maize production and yields in
West Africa during the 1980s came about as a result of the release of earlier-maturing
varieties with higher levels of disease resistance (IITA, 1995).
Thus, the history of plant breeding is highly international, marked by both chance,
fortuitous introductions made by farmers and more deliberate transfers of traits
involving teams of scientists. The history of crop improvement in Africa is much the
same, with numerous examples of both types of advance. A durra variety of sorghum
cultivated by Touareg nomads in northern Mali is commonly believed to have been
introduced by pilgrims returning from Mecca. Meanwhile, rice varieties recently developed by the West African Rice Development Association (WARDA) combine traits
from two distinct species and required several applications of biotechnology.
Neither of these types of advancement, however, can serve as an effective substitute
in Africa for consistent, informed, crossing, selection, and recombination carried out by
plant breeding teams that is the mainstay of crop improvement systems throughout the
world. At present, with notable exceptions in a number of countries, it is this central, key
component that is most lacking in Africa, and most in need of public sector support.
While in the USA, Europe and parts of Asia and Latin America a major portion of this
activity has transitioned to the private sector, in Africa the same is not true. Just as food
security was first attained through publicly funded initiatives (including the Green
Revolution) in other parts of the world, African governments and their collaborating
institutions must make this commitment, whose goal will be the establishment of
adequately supported, NARS crop improvement teams for all essential food crops
throughout the continent.
Of equal or greater importance to the productivity of crop plants is the stability and
long-term sustainability of cropping systems, as a whole (Buddenhagen, 1996). The
development of improved varieties has been described as essential to the creation of a
sustainable farming system under situations of intensification (Fischer, 1993, cited in
Byerlee, 1996; Hoffman et al., 1993). Although plant breeding contributions to the
sustainability of cropping systems are less widely recognized, evidence has been
presented for sustainability improvements emanating from releases of pest- and diseaseresistant rice varieties in Asia in the 1980s (Conway, 1997) and early-maturing flint
maize varieties released in Malawi and Mozambique in the 1990s (Smale et al., 1993;
Chapman et al., 1997). More recently, maize breeding in Kenya has resulted in maize
varieties with greater resistance to foliar diseases. After only 2 years of breeding,
experimental varieties tested during a season of high disease incidence yielded higher
than commercial hybrids (Ininda and Ochieng, 2000).
As applications of biotechnology become more routine, they are being increasingly
integrated with breeding programmes to result in a very different sort of crop improvement programme. In Africa, however, the integration of such techniques across most of
the continent is still some years away. Therefore, it is still necessary to consider the
breeding subsector as a stand-alone pursuit, mainly conducted by teams of public sector
field researchers and in critical need of support from national governments and international donor agencies, alike. This section deals with the major movements afoot in
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breeding programmes in Africa and attempts to identify priorities within and across
these institutional boundaries.
As noted above, farmers in Africa have been making plant selections for thousands
of years. These selections have resulted in popular landrace varieties of indigenous
crops such as sorghum, millet and cowpea, which have recognized names within and
even across different agro-ecological zones. Examples of these include the popular
Striga-tolerant variety of sorghum in Mali, Seguetana; the high-yielding and pest- and
disease-resistant cowpea in Mozambique, Namuesse; and the widely adapted millet
landrace from Togo, Iniadi. Selections made from introduced crops such as maize
and cassava have also resulted in landraces of these crops being distributed among
large numbers of farmers. Catete is a popular, early-maturing landrace of maize
in Angola. Fumo de Comboio is a disease-resistant, high-yielding variety of cassava in
Mozambique. Reep is a late-maturing, yellow maize variety used widely in southern
Sudan.
Useful demonstrations of the value of these selection methods occur when varieties
of cereal crops are introduced directly, without modification, from North America or
other regions of the world. In addition to photoperiod differences which frequently
result in altered times of flowering and harvest, such introductions normally suffer from
very high incidence of crop pests and diseases. Added, major differences in grain quality
and plant type preferences preclude most temperate germplasm from being of direct use
in Africa, although certain useful crop varietal traits have frequently been transferred
to African backgrounds via back-crossing and population improvement methods of
breeding.
Modern plant breeding began in Africa in the early 20th century, following its
emergence in Europe and North America, via the influence of colonialist governments.
French scientists began breeding rice in West Africa as early as the 1930s and millet
in the 1950s. Research on hybrid maize was initiated in Rhodesia in 1932. The first
hybrid variety to be released in Africa was the maize variety SR-1, released in 1949
(Mashingaidze, 1994). Breeding programmes aimed at developing varieties for smallscale farmers were generally neglected, however, until after the Green Revolution in
Asia. Such programmes received a significant boost with the establishment of the
International Institute for Tropical Agriculture (IITA) in Ibadan, Nigeria, in 1967.
Today, IITA conducts breeding on cassava, yam, banana, cowpea, soybean and
maize. Since then, several other centres of the Consultative Group on International
Agricultural Research (CGIAR) have established breeding programmes in Africa.
Several cases of yield increases in Africa led by the introduction of improved
varieties clearly indicate the potential of crop improvement to alleviate hunger.
Cassava yields increased dramatically in Nigeria following the introduction of

improved varieties in the mid-1980s (Nweke et al., 1994).
Maize production in West Africa over the same period increased by an estimated
average of 4.1% annually following the development of early-maturing, droughtresistant varieties (IITA, 1995; Smith et al., 1997).
Rapid adoption of hybrid maize in western Kenya during the 1960s and 1970s led
to dramatic increases in productivity (Gerhart, 1975).
White and Sitch (1994) reported yield increases in sorghum, sweet potato, cowpea,
and maize in Mozambique when improved, adapted varieties were introduced.
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Chapter 4
Table 4.1.
Yield increases observed from the introduction of improved varieties.
Country
Crop species
% Increase
in yield
Angola
Mali
Mozambique
Mozambique
Mozambique
Senegal
Sudan
Congo (Dem. Rep.)
Congo (Dem. Rep.)
Maize
Sorghum
Sweet potato
Maize
Sorghum
Cowpea
Maize
Maize
Cowpea
46
24
61
71
133
100
53
18
108
Reference
Nankam et al., 1996
Dembele et al., 1997
White and Sitch, 1994
UC-Riverside, 1994
Janson and Kapukha, 1995
Asanzi and DeVries, 1995
Asanzi and DeVries, 1995
DeVries and Olufowote (1997) analysed results from intensive, NGO-managed

campaigns to increase farmer productivity through testing and dissemination of
improved, adapted varieties of staple food crops in six African countries. The results
of on-farm measurements of yield increases are shown in Table 4.1.
Given the absence of extensive irrigated land and continued low use of external
inputs, agro-ecologically based breeding, combined with full exploitation of heterosis in
the formation of adapted hybrid varieties must substitute for a Green Revolution-style
approach to breeding in Africa. These aims can be significantly assisted by breakthroughs in biotechnology. The results from this effort will not be as quick to appear or
as dramatic as during the Green Revolution in Asia, but, taken together, they represent
an effective, rational strategy against food scarcity throughout the continent.
4.3 Crop Improvements Counter Arguments

In spite of diligent efforts by national and international scientists alike, it must be
recognized that plant breeding for many important food crops in Africa has been
plagued by low adoption rates among the majority, small-scale farmers. While in some
cases this is simply a matter of input- (fertilizer- and pesticide-) responsive varieties not
being adopted due to the absence of those inputs on African farms, the causes are often
more complex. While there is some indication of greater gains in this area over the past
decade (Maredia et al., 2000), in general, adoption rates continue to present a strong
challenge to breeding as a strategy for improving food security among the rural poor.
This phenomenon deserves careful analysis before the proposition can be made that
increased efforts at crop improvement in Africa are justified, and efforts have been made
to build such an analysis into all the major sections of this book.
In spite of relatively large expenditures of funds and human resources on maize
breeding in most of Africa over the past three decades, only an estimated 37% of farmers
regularly plant improved varieties (Morris, 1998). Similarly for sorghum and millet,
Ahmed et al. (2000), in citing adoption rates in seven countries where the highest level
of impact from breeding had been achieved, found that the highest level of adoption of
improved varieties outside of South Africa was 35%, while the average adoption rate for
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these countries was 29%. Surveys on adoption of improved cassava in Nigeria, the
African country where cassava breeding has arguably had its greatest success to date,
show that approximately 55% of farmers who have had access to improved varieties have
adopted them (Nweke et al., 1993).
A system-wide review of CGIAR impact on crop genetic improvement by CGIAR
(2000) revealed that in Africa:
Traditional varieties represented 70% or higher usage for sorghum, millet, beans
and cassava.
Wheat ranked highest in adoption of modern varieties, with approximately 63%,
followed by maize with roughly 48% adoption, and rice with 40% adoption.
The balance of farmers cultivating these crops continue to plant local landraces,
often altered by chance cross-pollinations with improved varieties grown in the vicinity.
The aim of crop improvement is not to replace these landraces systematically with
improved varieties. In most cases, they embody traits which must be conserved if new
offerings are to be successfully introduced (and, as this section points out, the final decision must be left up to farmers). However, the collected observations made by scientists
and farmers over the years are that in many cases landraces do not represent the best that
can be achieved today. The rate of evolution of these varieties through mass selections by
farmers has not kept pace with the numerous, rapid changes that have taken place in the
rural settings where they are grown. Chief among these changes, of course, has been
population growth, constantly creating more pressure on the land and the crops grown
on it. But other changes have resulted in new conditions, as well, including the
introduction of new pests and diseases resulting from greater movement of people and
goods, the more frequent incidence of drought, potentially arising from global warming,
and the reduction of soil fertility and soil water-holding capacity (Ngwira, 1989). As a
result, many landraces suffer from susceptibility to pests and diseases and environmental
stresses (Rajaram et al., 1988; Kyetere et al., 1997; Ininda and Ochieng, 2000). Among
the traits farmers commonly cite as advantages in improved varieties is reduced time to
maturity. But significantly reducing the time to maturity, or increasing resistance to
pests and diseases, cannot be achieved in the short or medium terms by mass selection. It
requires the intervention of plant breeders.
4.4 Farmer Participation in Crop Improvement

The factors that influence adoption rates of improved varieties are many and varied.
Certainly, access to improved varieties, given the low coverage of Africa by seed
companies, is an important one which is explored later in this book. But in a context
where genes for desirable traits must be introgressed into genetic backgrounds which are
also desirable, ecological adaptation and farmer preferences play a major role in
adoption. For these reasons, one of the most important changes in breeding programmes for developing countries in recent times has not been based on genetics at all,
but on the increased emphasis placed on the participation of farmers in the variety
development and selection process. This innovation has proved most critical in areas
where seed markets often do not operate efficiently and farmers are therefore less able to
communicate their varietal preferences through the marketplace.
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Chapter 4
Aspects of farmer varietal preference are, in fact, intricately intertwined with factors
which confer varietal adaptation. However, whereas varietal adaptation can be evaluated
by breeders on the basis of performance factors, aspects related to farmer preference can
best be evaluated by end users of the varieties (Adesina and Baidu-Forson, 1995;
DeVries and Fumo, 1995; Kitch et al., 1998). Such aspects of farmer preference
commonly include time to harvest (with a widespread bias toward early maturity),
quality of secondary harvested components (stalks, fodder production, edible leaves),
grain quality (texture, taste), and processing qualities (ease of de-hulling, pounding or
grinding). Taste characteristics, in particular, tend to be overlooked by relatively affluent
outsiders who consume a wide variety of food products and have access to a variety of
condiments. Simpler, monotonous diets rely primarily on the flavour and texture of the
food product itself (Fliedel and Aboubacar, 1998).
In fact, the rationale for involving farmers directly in breeding for marginal areas is
simple: diverse agro-ecologies and farmer preferences common to small-scale farming
contexts tend to complicate the decision-making tasks faced by breeders. Farmers, on
the other hand, understand almost intuitively which offerings among a range of choices
are best for them, given their various priorities for use of the crop. Moreover, because
crop varieties are usually developed by non-users (few researchers are farmers), they
require regular input from farmers to be able to structure their selection indices
accurately. Participatory plant breeding methods have been described by De Boef et al.
(1993), Okali et al. (1994), Sperling and Loevinsohn (1996), Witcombe et al. (2000),
and Thro and Spillane (2000). The most commonly cited range of reasons for involving
farmers in the selection process includes the following.
1. Gaining a better understanding of farmer preferences. Farmers who consume a portion
of their crop within the household may insist on taste, texture, and processing requirements which are difficult to screen for by breeding teams. Simply allowing farmers to
observe their growth and taste under local conditions can avoid years of time and effort
trying to push a variety that is not acceptable for these reasons.
2. Permitting more precise selection for individual environments. Creating testing
stations in every agro-ecology is often cost-prohibitive. Likewise, counting on breeders
to be able to line up all the agro-ecologies and resistance and tolerance traits and decide
which variety is best in each area is often not a reasonable proposition. Testing varieties
on-farm and letting farmers decide which perform best can simplify the process of
agro-ecology-based breeding.
3. Empowering farmers vis--vis the decision-making process (Jones et al., 1999). Farmers
who have not found improved varieties useful in the past can become highly interested
in contributing to crop improvement when consulted. In fact, permitting their voices to
be heard on these issues can be viewed as part of the broader process of democratization,
worldwide.
The crop improvement process embodies several opportunities for meaningful interaction between farmers and researchers, beginning with a needed, intensive learning
phase when researchers become aware of agro-ecological variation and the interactions of
crops and user systems. Early inbred generations (three generations of controlled breeding or later) are stages when farmers can be consulted on issues of plant type, maturity,
and grain quality (Butler et al., 1995). Nevertheless, a review of participatory plant
breeding programmes worldwide performed by the CGIAR (Jones et al., 1999) found
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that in very few cases were farmers consulted prior to the genetic fixing of traits in candidate varieties. More common was the involvement of farmers in priority setting, for
example via surveys conducted prior to setting selection indices. Important conditioning
factors for success of farmer participation in such schemes were: (i) the willingness and
interest of farmers to set aside time for the work; and (ii) the presence of clear points of
view among farmers consulted regarding the traits required in the crop species.
Generally speaking, participatory breeding to date in Africa has been mainly
confined to priority setting and variety selection. Participatory variety selection usually
involves exposing farmers (through scheduled visits) to a wide range, or, a basket of
candidate varieties grown by researchers in a common planting (sometimes referred to as
mother trials) followed by farmer-conducted evaluation of a smaller number of selected
varieties on a large number of farms (sometimes referred to as baby trials). Box 4.1
below offers an example of an IARC breeding programme which makes extensive use of
farmer expertise. As participatory methods become more established, it is hoped that
farmers will be consulted more extensively prior to the fixing of traits.
A second aspect of farmer participation in crop improvement aims at greater
tapping of biodiversity and the large variation that exists within landraces of crops
grown in Africa. For some time it has been known that resistance genes existed in low,
but useful, frequencies in a number of African crops. What has been missing was a
means of isolating them, in order to feed resistance sources back into breeding programmes. Recent proposals for using rural training facilities in teaching farmers how to
Box 4.1.
Participatory rice variety selection in West Africa
When the West Africa Rice Development Association (WARDA) experienced a

breakthrough in the breeding of interspecific crosses between African rice (Oryza
glaberimma) and Asian rice (O. sativa), it decided to involve farmers in making
selections of varieties for release. Interspecific rice varieties represented an entirely
new plant type with various combinations of traits contributed by each species. The
African rice genome contributed vigorous early growth for reduced competition
from weeds and resistance to a number of important pests and diseases. Asian rice
characters that were expressed included branching tillers, which supported more
grain. In order to determine which combinations of traits were of most importance
to farmers, WARDA employed a 3-year, participatory process, gradually moving
from a large number of varieties to a limited number which could be presented for
release and multiplication through national research programmes.
In year 1 of the WARDA process, 60 lines are introduced to farmers through
trials grown in farmers fields. WARDA scientists make three visits during the growing season to discuss with farmers the performance of each variety at critical stages
of growth. In year 2, the list is narrowed down to seven varieties. Farmers evaluate
each variety for various characteristics, and evluations are recorded by the WARDA
Economics Unit. In the final year of participatory selection, WARDA multiplies
those varieties that have been selected by farmers and offers them for sale.
Interspecific varieties have consistently been among those selected by farmers
in tests that included both interspecifics and normal rice varieties. Breeders at
WARDA are continuing to search through screening trials of interspecific progeny
for varieties that may offer new, valuable plant types and resistance to intractable
problems of rice production in Africa.
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Chapter 4
identify insects and diseases represent a new means of linking farmers to breeding
programmes which may lead to a new form of participatory gene discovery (FAO/
Zimbabwe MOA, 1998).
The potential impact of increasing the participation of farmers in the crop improvement process in Africa should not be underestimated. Gradually, a methodology is
emerging for ensuring that the crucial ingredient of farmer preferences is included in
breeding improved crops for poor farmers. Nevertheless, the complexities in terms of
taking timely decisions and maintaining the rhythm and steady progress necessary to get
improved lines moved through a programme, likewise, should not be ignored. And the
increasing dogma over farmer participation should not be allowed to interfere with those
decisions that still need to be taken by breeders.
The number and range of farmers that can be included in breeding and selection
programmes carried out by a small number of scientists is very limited. Transport and
other costs associated with including large numbers of farmers in selection processes in
each agro-ecology could limit the overall effectiveness of breeding programmes. Neither
should farmer input on breeding be viewed as a panacea. There is evidence that farmers
generally underestimate the importance of disease resistance in increasing and stabilizing
yields (Trutmann, 1996). Only one farmer out of 243 interviewed in Cameroon
identified nematodes as a cause of lodging of bananas in an area where the problem is
considered by researchers to be widespread (Hauser et al., 1998). This can be especially
important in areas where disease incidence is sporadic, but subject to epidemics. In light
of the lessons already learned, farmer participation and other means of obtaining
information from the farmer level can be viewed as catalysts to the central responsibility,
which should remain with breeders.
However, the most salient feature of farmer participation in crop improvement
remains simple: a critical knowledge base exists among farmers that needs to be accessed
in order for crop improvement to be effective in developing varieties which perform
better under local farming conditions. Gaining access to this knowledge base can be
achieved through a wide range of means, but inevitably requires that breeders take the
time to listen to farmers and understand the ways in which they use crop species and
varieties to provide food security in their households.
4.5 Crop Improvements Ground Zero: National Breeding

Programmes
National breeding programmes are the front lines of public sector breeding in Africa.
National programmes continue to be the primary place of employment of Africas
best-trained scientists. For many self-pollinated food crops, national programme varieties are likely to continue to be the sole source of new varieties. Regional breeding
networks, though often coordinated by international or regional entities, still depend
on national programmes to propose and promote the release of promising materials.
National programmes which maintain a strong focus on breeding of commercial crops
such as maize can also serve as an important source of new varieties marketed by private
seed companies, through licensing agreements. For these and other reasons, understanding the promise and limitations of plant breeding in Africa requires an analysis of factors
influencing breeding capacity at the national level.
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The promise of public sector breeding programmes today lies essentially in their
ability to develop improved varieties for farmers who are not targeted by private sector
seed companies1. In Africa, this includes the vast majority of farmers of all crops.
Therefore, public sector breeding programmes play a far more important role in developing countries than developed ones. While NARSs today embody a greater number of
better-trained staff than at any time previously in Africa (Pardey et al., 1997), they also
suffer from a number of critical weaknesses, including strategic and financial gaps.
Strategy gaps
The organization of crop improvement programmes is driven by factors related to the
size of programme to be implemented, the clients to be targeted, and the kind of products to be produced (House, 1985; Fehr, 1987). In the past in Africa, many such factors
were influenced by the monopolistic control maintained by public sector entities over
varietal development and seed distribution. National seed companies which faced no
competition had little interest in marketing a broad array of varieties finely tuned to different agro-ecosystems. Emphasis was therefore placed on broadly adapted materials,
and selections were based on performance across widely differing farming conditions,
even though the limitations of such strategies for developing countries were called into
question by several authors (Ceccarelli, 1989; Simmonds, 1991). At least part of this
emphasis was reinforced by breeding strategies in the USA, which, at least in public
institutions, was driven by the search for broad adaptation, as illustrated by the following excerpt from a widely used plant breeding text:
Breeding populations for wider adaptation may result in fewer varieties or hybrids. The
desirability of this is obvious. Most certainly, such development could be more deliberately
planned and productive if factors associated with wide adaptation were better defined.
(Smith, 1966).
Deregulation of the seed sector, with its attendant diversification of products and
purveyors of new varieties, coupled with more participatory approaches which attempt
to offer a basket of new varieties, as opposed to a very limited number, are both having
a major impact on the ways in which public breeding programmes in Africa need to be
organized. Among NARSs, several trends have emerged which point to needs for
strategy adjustments. For example, increased activity by private seed companies which
focus on supplying maize farmers in high-potential areas with hybrid seed can be viewed
as an opportunity for national maize breeding teams to concentrate on the development
of OPVs with higher levels of adaptation to different agro-ecologies. Yet, in many cases,
NARs continue to devote the majority of their breeding resources to hybrid maize
development.
As emphasized elsewhere in this book, better targeting of small-scale farmers needs
will require combining numerous resistance and tolerance traits. This, in turn, will
The opportunities which exist for public breeding programmes to serve the needs of private sector seed
companies are recognized and explored in some depth in the section on Seed Systems in this book.
However, it is their role as an agency focusing on non-commercial uses of breeding which forms the
primary concern of this section.
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Chapter 4
require greater collaboration among scientists trained in various disciplines, including

pathology, entomology, and physiology, who, due to continued, low-input (such as
fungicides, insectides and herbicides) use in Africa, are generally only able to make an
impact on farmers lives if their focus traits are embodied in the seed. In many cases,
these scientists continue to operate on the periphery of crop improvement programmes,
often to the detriment of the products eventually developed. Targeting small-scale
farmers calls for a much greater level of interaction of scientists and technicians from a
range of plant scientists central to crop performance at the farmer level. Indeed, in some
circumstances, their greater attention to factors related to crop performance at the
farmer level has led to their assuming leading roles on integrated breeding efforts in
Africa and other developing regions.
Chapters 8 to 14 of this book focus on traits of specific significance for seven important species of food crops in Africa. While numerous traits are identified for each crop,
only a subset of these is critical to crop performance in each agro-ecology.
Understanding the distribution and boundaries of the various agro-ecologies within
each country, defining the set of traits important within each agro-ecology, and then
selecting the best-adapted parents with the best combining abilities would appear to be a
logical and straightforward approach to meeting the challenge of implementing a
national breeding strategy. Because of the large land area to be covered and Africas
constant ecological variation, national programmes will be the key to implementing
these strategies. They can, however, receive critical support from IARCs, both in
defining agro-ecologies and in the supply of parental breeding materials.
In Kenya, Uganda and Malawi recent development of new national breeding
strategies has helped to revitalize programmes and focus the attention of breeding teams
on producing new products. Key components of such exercises were consultations with
farmers and consideration of the various agro-ecologies to be covered by the strategy.
Strategy building at the national level has also been furthered through sponsoring of
regional meetings wherein such subjects as strategy development and management of
public breeding programmes in the context of a deregulated seed industry form the
primary focus of attention.
Financial gaps
Strategy development at a national level will not result in major changes in output if
trained personnel and operating funds are not available for implementing breeding
strategies. Studies on the structural capacity of NARSs to conduct agricultural research
indicate that good progress has been made in terms of staffing up, with the number of
national scientists growing at an average annual rate of 5% between 1961 and 1991
(Pardey et al., 1997). Funding to carry out research, however, has become scarcer over
this period (Byerlee, 1996). In many national programmes of sub-Saharan Africa,
governments provide sufficient funding to cover staff salaries (albeit at very low levels),
but not the necessary operating costs. Although national research programmes were at
one time a popular area of expenditure for African governments, support to agricultural
research began to decline in the 1980s. By 1991, expenditure per scientist was only
66% of the 1961 level (Pardey et al., 1997). The result has been both a reduction
in overall activity and output and an increasing dependence upon donor agencies for
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operational costs. In recent studies, funds from international donor agencies were found
to account for 45% of crop improvement research expenditure in Africa (Pardey et al.,
1991). For biotechnology research, the figure rises to 65% (Cohen, 1998).
As access to operating funds has decreased, the result has been gradually decreasing
levels of activity among breeding teams, reflected in smaller breeding programme
nurseries covering fewer breeding environments. At a certain point, minimal population
sizes fail to embody sufficient genetic variation to warrant continued activity (Falconer,
1989). Appropriately targeted infusions of financial support to African NARSs, therefore, will be critical to taking advantage of recent breakthroughs in crop genetic
improvement. Although increased levels of funding for agricultural research from
African governments would be a welcome policy development, it is questionable
whether this is likely, given the current financial crisis many African governments are
facing. In view of competing priorities for national budget expenditure in the areas of
education and health systems, the current formula, whereby governments generally
cover researchers salaries and donor agencies cover a sizeable portion of operating costs,
may not be the worst option.
In order for such a formula to function effectively, however, some key principles
must be observed. First, NARSs policy-makers must work from effective master
plans in overseeing and approving the application of donor funds. This argues for
more intensive coordination of donor resources by NARS headquarters. Second, donor
institutions need to become more transparent and more cognisant of the application of
each others resources, and plan accordingly. This is particularly important in the use of
resources aimed at developing end-products, as in the case of breeding. At present, there
are the beginnings of a loose-knit consortium of donors (including the Swedish-funded
Bio-Earn initiative, the Gatsby Charitable Trust, and The Rockefeller Foundation)
which contribute to breeding and biotechnology in Africa. Informal information shared
among the managers of these programmes is increasing the complementarity of funding
for crop genetic improvement. However, greater levels of coordination among a wider
group of donor agencies are still needed.
Several authors have questioned the rationale for increased funding to NARS,
especially those focusing on marginal areas, sometimes even stating there was
overinvestment in such institutions (Winkelmann, 1994; Byerlee, 1996; Evenson,
2000), and advocating for a reduction in the number of crop improvement programmes. Much of this analysis has focused on the case of rice and wheat in Asia,
however, where yield thresholds have been challenged and NARSs have remained
relatively well-staffed and financed over the past few decades. Observations from Africa
point to a rather different picture. The impact of the AIDS epidemic has certainly
created a need for added training of researchers to replace those who have died or
become ill (Marianne Banziger, personal communication). Second, the greater overall
complexity of Africas crop improvement challenge described in Chapter 2, combined
with dilapidated or non-existent infrastructure, may mean that crop improvement gains
come at a higher financial cost in Africa than Asia.
Calculating the cost of Africas crop improvement budget with any precision is a
difficult task. In particular, researchers salaries and costs of their administrative support,
as well as infrastructure costs, are highly variable from country to country and such
information is difficult to obtain. However, calculating the operating costs of such
programmes is somewhat more feasible. Setting aside for the moment the cost of formal
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training, and using information from The Rockefeller Foundations annual grantmaking reports, an average annual cost per crop per country for breeding operations can
be estimated at approximately $70,000. Based on the assumption that an average of
seven food crops warrant such investments on an annual basis in each country of
sub-Saharan Africa, and counting a total of 41 countries, $20 million could potentially
cover the operating costs of breeding operations in sub-Saharan Africa. If operating costs
account for half of total costs, then a total of $40 million could potentially cover the full
budget, with the exception of the cost of formal training.
Adding in the costs of formal training is an equally subjective exercise, but one that
is still useful for purposes of gaining an order-of-magnitude estimate of the costs
involved. Again, based on seven crops in 41 countries and an average of four crop
scientists per crop (but assuming that all of these would work on at least two crops), and
using an estimated cost of $150,000 for a PhD programme, the cost of a formal,
postgraduate training programme for crop improvement in Africa is in the order of
$4.35.7 million annually, depending on whether one assumes a 15- or 20-year average
length of career. Even making generous concessions for the costs of administering such
training programmes, the total cost of formal, postgraduate training for crop improvement in Africa should not reach beyond $10 million, annually.
Therefore, while recognizing the error-prone nature of such calculations, and while
not wishing to attach an overly great importance on identifying a discrete cost for what is
inevitably and necessarily a very disjointed and variable effort, it can nevertheless be suggested that the overall, annual cost of a serious initiative on broadly improving the
genetic performance of Africas food crops through conventionally based, national crop
improvement programmes might be estimated at $50 million, $20 million of which
(based on the current model) can be covered by African governments. By comparison, in
2000, the total value of the African food aid programme administered by one US-based
NGO alone was over $100 million (Walter Middleton, personal communication).
4.6 Applied Science Powerhouses: International Agricultural Research

Centres
Crop mandates
NARSs in Africa receive critical reinforcement in crop improvement efforts from
IARCs, in particular from the member centres of the CGIAR. The work of the CGIAR
is carried out by 16 research centres headquartered in various countries throughout the
world. Six centres CIAT, CIMMYT, CIP (International Potato Center), ICRISAT
(International Crops Research Institute for the Semi-Arid Tropics), IITA and WARDA
(West Africa Rice Development Association) currently have scientists based in Africa
working on crop genetic improvement.
CGIAR centres have traditionally managed gene banks and developed broadly
adapted source populations and breeding lines for use by NARSs and seed companies
in selecting adapted, improved varieties. They have also carried out both general and
highly focused training programmes for crop improvement specialists working at the
national level. More recently, IARCs have become active in the formulation and
management of commodity-based, regional crop improvement networks. The CGIAR
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centre contribution to NARS productivity in breeding in developing countries, worldwide, has been estimated at 30% (Evenson, 2000). They also increasingly focus on crop
management issues.
International agricultural research centres have considerable presence in Africa. In
1997, Africa accounted for 40% of allocations for research expenditure by the CGIAR,
representing the largest single region in the world (CGIAR, 1998). Table 4.2 lists the
number of full-time internationally recruited scientists currently employed by the
CGIAR in Africa.
Table 4.3 lists the centres breeding activity within the context of regional crop
improvement networks. Most IARCs have actively promoted the development of
networks focused on improvement of their mandate crops. These networks have based
their membership on the three main regional agricultural research coordination bodies,
CORAF (West Africa), ASARECA (East Africa) and SACCAR (Southern Africa).
International agricultural research centres continue to serve an important purpose
in crop genetic improvement in Africa. They have broadly improved the genetic
potential of germplasm adapted to Africa through the introgression of novel traits and
increased yield potential. In the case of maize and, to a lesser extent, sorghum, they have
also developed advanced, inbred lines for use in the formation of commercial, hybrid
varieties adapted to all the major subregions. Finally, they have built up and maintained
extensive African germplasm collections of all the major food crops, a task which, due to
insecurity in a large number of countries, could not have been assured by NARSs or any
other group.
IARC breeding programmes have had a major positive impact on agricultural productivity in Africa. Their continued support is vital to achieving greater gains in the
future. Still, adoption rates of improved varieties lag behind that which might be
expected given IARCs extensive effort on crop improvement in Africa, and more effective approaches must be sought. There appears to be a gap between IARCs and NARSs,
where NARSs have not employed IARC-bred source materials in developing finished
varieties or been able to use breeding methodologies promoted by IARCs. In many
cases, NARSs have continued to rely primarily on their own, relatively narrow germplasm base. In other cases, they have failed to mount breeding programmes at all, in
spite of possessing scientists trained to the same levels as IARCs. In the absence of this
critical breeding linkage, source materials have sometimes been presented as adapted,
ready-for-release varieties, and national breeding programmes relegated to the role of
testing IARC-bred material.
Table 4.2.
IARC crop improvement scientists in Africa.
Institute
Scientists in crop improvement
CIAT
CIMMYT
CIP
ICRISAT
IITA
WARDA
Total
6
7
4
15
35
9
76
49
Chapter 4
50
Chapter 4
Table 4.3.
Centre
Scope of IARC crop improvement responsibilities in Africa.

Mandate crops
Primary IARC breeding sites

/regional focus
Banana/plantain On, Nigeria/West Africa

Kampala, Uganda/East Africa
Cassava
Ibadan, Nigeria/West Africa
Bvumbwe, Malawi
Cowpea
Kano, Nigeria/Africa
Maize
Ibadan, Nigeria/West, Central Africa
Bouak, Cte dIvoire/West, Central
Africa
CIAT
Beans
Lilongwe, Malawi
CIMMYT Maize
Harare, Zimbabwe/Southern Africa
Addis Ababa, Ethiopia/
African Highlands
Nairobi, Kenya/East Africa
Wheat
Testing only/Southern Africa
Addis Ababa, Ethiopia
CIP
Potato
Testing only/Africa
Sweet potato
Nairobi, Kenya/Africa
ICRISAT Sorghum
Bamako, Mali/West Africa
Nairobi, Kenya/East Africa
Bulawayo, Zimbabwe/Southern
Africa
Millets
Bulawayo, Zimbabwe/Southern
Africa
Pigeon pea
Nairobi, Kenya/Africa
Groundnut
Lilongwe, Malawi/East and Southern
Africa
WARDA Rice
Bouak, Cte dIvoire/West Africa
(non-irrigated)
St Louis, Senegal/West Africa
(irrigated)
Entebbe, Uganda
IITA
Networks
MUSACO
BARNESA
CEWARRNET
EARRNET
SARRNET
RENACO
WECAMAN
ECABRN
SABRN
MWIRNET,
SADLF
AHI
ECAMAW
MWIRNET
ECAMAW
PRAPACE
PRAPACE
WCASRN/ROC
ARS
ECARSAM
SMINET
SMINET
WCAMRN/
ROCAFREMI
INGER
ECSARRN
Lingering low yields among African farmers for crops such as maize and rice, where
adoption of improved varieties has been appreciable, call into question the overall value
of the improved germplasm to local farmers, and whether the public crop improvement
system, including the combined efforts of both IARCs and NARSs, cannot be
improved upon. While, for lack of better options, offered materials (both unfinished,
IARC breeding lines and unenhanced, NARS varieties) may represent the best improved
varieties available, they do not represent the full extent of what modern breeding
could normally do for farmers. Their lack of key identifiable traits (drought tolerance,
maturity, grain type, disease incidence, etc.) may reduce their acceptance by farmers and
50
Chapter 4
51
diminish the value of the crop improvement system, overall (Simmonds, 1991). An
observation from the recent, system-wide review of the IARC contribution to varietal
development (Evenson, 2000) perhaps captures the challenge most succinctly:
Not all germplasm produced in an IARC program is of equal value to all NARS programs.
The proportion of germplasm relevant to a given NARS program depends on the
differences in soil and climate conditions in the NARS region and in the IARC location
and on the efforts of IARC program to actually target germplasm for the NARS program.
(Evenson, 2000)
Increasing the interface between IARC and NARS breeding programmes is

necessary to realize the full potential of plant breeding in Africa. There is no doubt that
the provision of source breeding materials and technical backstopping by IARCs can
improve the success of the national programme. The relationship between them needs
to be broadened and strengthened so that NARSs recognize the value of IARC breeding
materials and IARCs understand the agro-ecological and institutional constraints
NARSs face. Weaknesses in NARS breeding capacity need to be eliminated so that
investments in IARCs can pay off. Likewise, IARC perception and understanding of
agro-ecologies and farmer preferences needs to be strengthened so that source materials
reflect more closely the priorities identified by farmers and breeding programmes.
Finally, IARCs serve a major need in mounting genetic improvement programmes
aimed at overcoming intractable constraints to production which may be beyond the
financial and scientific reach of national programmes. Thus, while IARCs account for a
small portion of the total work force, they are a key component for crop improvement
throughout the continent. This argues for an increase in the number of full-time IARC
scientists engaged in breeding in Africa.
Capacity building
In 1961, there were 2000 full-time agricultural researchers in Africa. By 1991 the
number had risen to 9000. Approximately 65% of agricultural researchers in 1991 had
attained postgraduate degrees, compared with 45% in 1961 (Pardey et al., 1997). As
NARS scientific staff have gained higher levels of training, the central task of IARCs
engaged in breeding in Africa has evolved. NARS scientists returning from training
overseas have been equipped with strong theoretical backgrounds which can be put to
use in developing new products for farmers. However, PhD programmes in the fastmoving field of genetics provide little background in methods of practical breeding.
Moreover, many returning scientists have never managed full-scale breeding operations.
Building practical breeding capacity including the science policy tasks discussed above
is therefore an increasingly important mission for IARCs.
4.7 Breeding Linkages Within a Continuum of Crop Improvement

Activities
The possible integration of biotechnology research, breeding, and seed systems in
developing and delivering new genetic technologies for small-scale farmers in Africa is
51
Chapter 4
52
Chapter 4
Fig. 4.1.
Collaborative model for overcoming genetic constraints in African crops.
shown in Fig. 4.1. It proposes a process by which identified production problems can be
addressed using a range of research methods. Examples of problems deemed routine
might include the appearance of a new disease, for example, the appearance of grey leaf
spot disease of maize in eastern and southern Africa in the past few years, for which
sources of resistance have been identified.
Intractable problems arising in marginal environments most likely will require
specialized research, such as that conducted by a different category of institution, herein
referred to as advanced research institutes (ARIs). Intractable problems might include
drought tolerance, parasitism by Striga, or attack by insect pests. These are problems for
which progress via conventional breeding techniques has proved difficult or very slow,
and for which no reliable means of selection are available. Here, NARS work must be
linked with that of IARCs and ARIs.
4.8 Managing the Complexity of Adaptation

Chapter 2 explored some of the complexities of designing crop varieties with specific
adaptation advantages in low-input farming systems. Managing this complexity will
require a systematic approach to understanding the nature and boundaries of agroecologies. In most cases, this will be done in stages, and at varying levels of resolution.
Table 4.4 lists some of the mega-environment characteristics for selected crops.
4.9 An Emerging Paradigm for Breeding in Africa

During the 20-odd years following independence in most African countries, monopolistic, public seed companies were managed and mandated by government to serve the
needs of all farmers. With a single outlet for seed of improved varieties, national
breeding programme strategies were more or less set by the marketing interests of the
52
Chapter 4
53
Production ecologies of Africa for selected crops.
Table 4.4.
Predominant
region
Crop
Agro-ecology
Maize
(CIMMYT,
1990)
Lowland tropical, Northern Guinea Maize streak virus

early
savannah, Congo (MSV), southern
Basin, southern leaf blight, stalk
Mozambique
rot, ear rot
Lowland tropical, Southern Guinea MSV, southern
intermediate
savannah, Congo leaf blight, ear rot,
Basin
stalk rot
Lowland tropical, West African
MSV, polysora
late
forest zones
rust, southern leaf
blight, ear rot,
stalk rot
Mid-altitude, early Northern
Turcicum leaf
Mozambique
blight, grey leaf
spot (GLS), MSV,
ear rot
Mid-altitude,
Zimbabwe,
MSV, ear rot,
intermediate
Malawi,
GLS, turcicum
Mozambique
leaf blight,
common rust
Mid-altitude, late Nigeria,
Turcicum leaf
Cameroon,
blight, GLS,
Zambia, eastern common rust,
Angola, Tanzania, MSV, ear rot
Uganda, Kenya,
Ethiopia
Highland, early/ Western Kenya, Turcicum leaf
intermediate
Great Lakes
blight, GLS, common rust, MSV,
ear rot
Highland, late
Ethiopia, Kenya, Turcicum leaf
Great Lakes,
blight, GLS,
Tanzania
common rust, ear
rust, MSV, stalk
rot
Humid/
Cte dIvoire,
Weeds, acidity,
sub-humid zone Guinea, Sierra
blast, drought,
Leone
nitrogen
deficiency
Rain-fed lowland Nigeria, Benin,
Weeds, water
Liberia,
control, rice
Mozambique,
yellow mottle
Tanzania
virus (RYMV),
nitrogen
deficiency,
drought
Rice
(WARDA,
1998)
Constraints
53
Chapter 4
% of
total area
13
10
23
15
27
10
40
38
54
Chapter 4
Table 4.4.
Crop
continued.
Agro-ecology
Predominant
region
Constraints
Irrigated
Cameroon,
Nitrogen
Nigeria,
deficiency,
Tanzania, Kenya weeds, RYMV,
iron toxicity,
nematodes,
gall midge
Sahel irrigated
Senegal, Mali,
Nitrogen
Niger, Burkina
deficiency, cold,
Faso
salinity, weeds,
alkalinity
Mangrove
Coastal West
Sulphate acidity,
swamp
Africa
salinity, crabs
Deep
Northern Sahel Water control, low
water/floating
yielding varieties,
low fertilizer use
efficiency
Sorghum
Southern Guinea Nigeria, Ghana, Anthracnose,
(ICRISAT, 1992; savannah
Chad, Cameroon, sooty stripe,
Fred Rattunde, (> 1000 mm)
Sudan
smut, Striga
personal
communication)
Northern Guinea Northern Nigeria, Shoot fly, stem
savannah
northern Ghana, borers
Chad, southern
Burkina Faso
Striga, downy
Sahel zones
Mali, Burkina
mildew
(< 1000 mm)
Faso, Niger,
Nigeria
Eastern and
Kenya, Tanzania, Stem borer, grain
mould, midge,
southern Africa Zambia,
shoot fly, drought
Zimbabwe,
Mozambique,
Botswana,
Namibia
Heat and drought,
Millet (ICRISAT, Sahel zones
Chad, Niger,
head miners,
1992, and Fred
Mali, Senegal
Striga
Rattunde,
personal
communication)
Northern Guinea Nigeria, Burkina Downy mildew,
savannah
Faso, Chad, Mali, stem borers,
drought, Striga
semi-arid East
Africa
Downy mildew,
Southern Guinea Ghana, Togo,
drought
savannah
Nigeria
54
Chapter 4
% of
total area
5
4
6
30
30
20
20
40
20
15
55
Table 4.4.
continued.
Crop
Agro-ecology
Cassava
(Henry and
Gottret, 1996)
Southern Africa,
East African
Highlands
Lowland humid
tropics
Predominant
region
Constraints
Botswana, Namibia, Downy mildew,

Zimbabwe, Ethiopia drought
West African forest Mosaic virus

zones, Congo
(ACMV),
Basin, Mozambique bacterial blight,
anthracnose
Lowland
ACMV, bacterial
sub-humid
blight, spider
Tropics
mite, drought,
mealy bug
Semi-arid
Sahelian zones,
Drought,
tropics
southern Africa
anthracnose,
bacterial blight,
burrowing bug
Mid-altitude
East and southern ACMV, spider
zones
Africa, Cameroon
mite, bacterial
blight, burrowing
bug, termites
Sub-tropic
South Africa
Drought, bactezones
rial blight, mealy
Banana
bug, spider mite
(INIBAP, 1995) West African
Guinea savannah, Black sigatoka,
lowland
transitional zones, weevils, bunchy
Congo basin
top virus
East African
Uganda, western
Black sigatoka,
Highland
Kenya, Great Lakes nematodes,
weevils,
fusarium, banana
streak virus
Lowland
Coastal east Africa
Mid-altitude
South Africa,
Fusarium
Cowpea
commercial
Zimbabwe
(Laurie Kitch,
Forest Zone
Coastal West Africa Weeds, maruca
personal
and Guinea
pod-borers,
communication) savannah
thrips, foliar
disease
Northern Guinea Coastal West Africa Bruchids, pod
savannah
bugs, thrips
Sudan
Nigeria, Ghana,
Aphids, bacterial
savannah
Benin, Togo
blight, mosaic
viruses, bruchids
Sahelian zones Senegal, Mali,
Bacterial blight,
Striga, aphids,
Niger, Burkina
Macrophomena,
Faso, Chad
bruchids
55
Chapter 4
% of
total area
15
35
36
10
10
36
57
6
1
5
10
40
30
56
Chapter 4
Table 4.4.
continued.
Crop
Agro-ecology
Cowpea
(contd)
Mid-altitude,
humid
Predominant
region
Kenya, Tanzania,
Uganda, northern
Mozambique
Mid-altitude, dry Kenya, Zimbabwe,
Botswana
Constraints
% of
total area
Thrips, pod bugs,

viruses, bruchids
Thrips, pod bugs,

viruses, bruchids
10
seed company. In the absence of competition from other suppliers, most seed companies
did the logical thing: in order to keep marketing and operational costs at a minimum,
they marketed a minimal number of improved varieties to as broad a grouping of
farmers as possible.
This restricted outlet for new varieties did little to encourage breeders to create the
steady flow of increasingly well adapted varieties which is the norm in deregulated seed
markets. Rate of release of new varieties stagnated, accounting for the fact that many
food crop varieties in use in many African countries are well over 20 years old. Figure 4.2
shows the rate of release of new maize varieties in selected eastern and southern African
countries during the period 1960 to 1995. Following deregulation of many seed sectors
during the late 1980s and early 1990s, increasing numbers of seed companies were
allowed to compete for the same market, with the result that the rate of release of new
maize varieties increased sharply and has maintained itself during the late 1990s (data
not shown).
Increased competition among seed companies will continue to fuel greater levels of
attention to farmers needs in Africa, with the net result being higher levels of activity in
the area of breeding. Thus, the old emphasis on varieties with broad adaptation
acceptable to a maximum number of farmers can be replaced with an approach that
emphasizes increased adaptation within a given agro-ecology. In this kind of market, one
plausible hypothesis is that breeding programmes (both those attached to seed
companies and those attached to NARSs which license their varieties with seed
Fig. 4.2. Rate of maize variety releases in Kenya, Malawi, Tanzania and Zambia,
19611995. Source: Zambezi (1997).
56
Chapter 4
57
companies) which pay closer attention to environmental variation stand to experience

increased demand for their products. If so, this would follow a trend among privatized
seed markets such as that of the USA, where seed companies often test experimental
varieties in over 1000 sites (Jensen, 1994).
Multi-year, multi-location testing of varieties can generate considerable understanding among breeding teams related to the differences among agro-ecologies (Jensen,
1994). As agro-ecologies become better known, breeding strategies emerge which
anticipate the needs of farmers, with information feeding back to the selection of
parents. For a commercial crop with considerable potential seed sales such as maize, this
may come about naturally as a result of competition among companies. For other,
non-commercial crops, such strategies will need to be constructed within the public
sector. The emerging new paradigm, then, is characterized by one wherein multidisciplinary crop improvement teams systematically begin to work backward from the
farmers needs dictated in turn by agro-ecological and end-use characteristics to the
formation of selection strategies and choice of parents for the formation of new breeding
populations most likely to bear positive results in as brief a period as possible. Within
such an approach, farmer participation is critical throughout the process.
4.10 Africa Breeding Challenges Summary

With notable exceptions in very poor countries and in countries where protracted
periods of instability have depleted public sector ranks, NARSs in Africa have achieved
the needed capacity to perform most of the routine breeding work entailed in
developing new varieties. Participatory techniques of breeding are progressively being
incorporated into national breeding programmes, offering a potential solution to the
perennial problem of incorporating farmer preferences into new offerings. The next
phase of their challenge is in securing resources and instituting programme strategies
that produce a steady stream of new varieties. At another level, the challenge is in
developing functional relationships with private seed companies that result in mutually
beneficial licensing agreements for commercializing new, public varieties. If breeding
programmes can undertake these kinds of reforms and pursue them vigorously, there is
genuine hope that public sector plant breeding can fulfill its potential in sparking
increases in productivity among Africas millions of small-scale farmers.
The challenges perceived by IARCs should closely resemble those being addressed
by NARSs, namely, creating a steady stream of improved, adapted breeding materials.
However, they should aim at playing a facilitative role rather than a lead role in
achieving those aims. With few exceptions, national programmes in Africa have demonstrated their ability to develop new varieties using conventional breeding methods.
Increasingly, these programmes are staffed by scientists with identical training to those
employed by international research centres. Breeding well-adapted source materials for
use in combination with local materials in boosting yield potential or resistance to pests,
diseases, or environmental stress is a critical role for the IARCs. However, an equally
important role exists in assisting the development and implementation of nationally
based breeding strategies.
57
Chapter 4
Biotechnology: Expanded
Possibilities
5.1 Overview
Plant biotechnology spans a broad and rapidly expanding range of research techniques
aimed at direct control over the genetic make-up of crops through the manipulation of
plant cell cultures and through the analysis and isolation of DNA. The most common
applications of biotechnology to plant breeding can be separated into five broad
categories: tissue culture, DNA marker technology, genetic engineering, genomics and
bioinformatics. Tissue culture involves in vitro regeneration of whole, functioning
plants from single cells or small portions of parent plants. Marker technology allows
many of the loci, genes, and alleles that are important in crop improvement and already
present in the gene pool available to breeders to be identified, located on the
chromosomes, and more effectively transferred via conventional crossing. It enables
interactions between genes to be determined and facilitates the identification and use of
new favourable alleles from wild relatives. Its most useful form, marker-assisted selection
(MAS) involves whole-plant selection based on DNA markers closely linked to genes of
interest. Genetic engineering refers to the in vitro transfer of genes into plant cells
followed by regeneration of whole plants containing these genes in the germline
(Hoisington et al., 1998). As laboratories become more fluent with biotechnology
methods of research, these three areas are increasingly being used in synergistic combination with each other.
More recently, a new application of biotechnology, termed genomics, has evolved
out of work in molecular genetics. Plant genomics can be described as identifying the
function of all of a plants genes and how they work together to determine when, where,
and why traits are expressed. Using gene chip (the plotting of thousands of gene
segments on plates or micro-arrays) technology the interrelationships and interactions
between genes and whole pathways can now be studied, and should help breeders to
create varieties with more exact combinations of desired traits (Wang et al., 2000). Such
aims have been facilitated in part by the development of methods for isolating expressed
59
59
Chapter 5
60
Chapter 5
sequence tags short segments of gene transcripts, which have been sequenced and can
be used to help identify the level of expression and function of the genes that generated
them. Because gene-encoding sequences represent only 10% of most genomes, this
method allows researchers to concentrate on the more informative portions of genomes.
The international effort currently under way to sequence the entire rice genome as a
model cereal genome should be completed in 2 to 3 years. It is already generating full
sequence data and is providing a wealth of useful new information for genomics
research. Because the genes that code for numerous plant traits and processes are quite
similar across species, this knowledge can be applied to genetic research on other crops.
It is widely believed that genomics will eventually replace the comparatively imprecise
method of identifying genes through markers, which usually do not identify the gene
itself.
Stemming from the explosion of information on plant genomes, yet another new
application, that of bioinformatics, has become of primary importance. Bioinformatics,
as the term implies, is essentially the management of information on gene structure,
position and function in ways that allow the data to be used to make broader interpretations related to the behaviour of whole organisms. Since cereal genomes are very
similar in gene content and gene order, bioinformatics should facilitate comparisons and
sharing of information across crop species. This, combined with the fact that bioinformatics requires powerful computational capabilities, sophisticated software, networking, and specialized human resources, argues in favour of having bioinformatics
centres that work on several crops. Together, genomics and bioinformatics are aimed
at eventually allowing researchers with access to the information to understand the
functioning of whole genomes.
While various methods of directly manipulating DNA and transferring it to plants
have been in use for nearly two decades, the power of biotechnology to transform
agriculture only became apparent following the release of several transgenic varieties of
staple food and fibre crops grains, legumes and cotton varieties resistant to insects and
herbicides. The response of farmers was dramatic, and plantings of transgenic varieties
increased rapidly in countries where they have been commercialized. While yields of
some transgenic crops were higher, the major advantages to farmers were significant
overall reductions in the cost of production and greater flexibility in rotating production
with other crops. In fact, transgenic varieties looked set to cover most of developed
country agriculture until controversy over their importation and testing broke out in
Europe.
Biotechnology is already contributing to smallholder agriculture in several ways.
Tissue culture is facilitating the rapid propagation and dissemination of clean planting
material of vegetatively propagated crops (e.g. banana, cassava, potato, sweet potato and
yams). Anther culture and the creation of doubled haploid plants are speeding the
process of producing genetically stable breeding lines (e.g. rice, wheat and barley).
Anther culture also helps overcome sterility problems in progeny resulting from wide
crosses made within species (e.g. Indica rice Japonica rice) and between related species
(e.g. Oryza sativa Oryza glaberrima). Marker-assisted selection (MAS) is speeding the
process of back-crossing desired genes into locally well adapted cultivars (e.g. resistance
to streak virus in maize, yellow mottle virus in rice, and African mosaic virus in cassava).
MAS is also leading to more durably resistant varieties by facilitating the pyramiding
of different resistance genes in the same variety (e.g. more durable blast resistance and
60
Chapter 5
61
Biotechnology: Expanded Possibilities
more durable gall midge resistance in rice). To date, however, the only transgenic crop
being grown by numerous smallholder farmers is insect-resistant (Bacillus thuringiensis)
Bt cotton in South Africa, Mexico and China. It has led to significant reductions in
pesticide use and costs, and is much appreciated by these farmers.
The technological gap which separates African agriculture and that of much of the
rest of the world today is perhaps most starkly apparent in the status of biotechnology
research. Nearly one-third of cropland in the USA in 1999 was planted to transgenic
crops (James, 1999). A survey of seed companies in Europe in 1999 showed that
one-third of all companies already employ genetic engineering in their crop improvement programmes and by 2002 80% of all seed companies will employ biotechnology
(Arundel et al., 2000). In India, there are over 30 agricultural research teams making
routine use of plant biotechnology (Dill, 1997), and in the rest of Asia, even relatively
underdeveloped countries such as Vietnam and Indonesia have burgeoning biotechnology laboratories making routine use of molecular mapping techniques, transformation
technologies and other applications. Yet when this book was drafted, there were only
three national-level research laboratories in sub-Saharan Africa (outside of South Africa)
using molecular applications of biotechnology. The implications, in terms of Africas
capacity to innovate and advance its food production systems in a way similar to the rest
of the world, must be recognized.
However, in spite of the delayed uptake of biotechnology in Africa, such
comparisons give a rather distorted image of the current potential for biotechnology to
affect the lives of the rural poor in Africa, for several reasons:
Many African scientists have received training in biotechnology research in

advanced research facilities and are ready to apply the techniques on the crops they
know;
Several biotechnology products (including transgenic banana, cassava, maize and
rice) aimed at African agriculture have been developed in outside laboratories and
await only the appropriate regulatory approvals for importation;
Available applications of plant biotechnology are highly suited to managing genetic
traits of importance to tropical agriculture, such as resistance to insect pests,
diseases, and environmental stress;
Crop yields in Africa are so low, in part due to lack of technology, that even
relatively modest improvements in technology can have significant local impact.
In view of Africas lagging status in the development and adoption among farmers
of conventionally produced, modern, improved varieties, adding molecular methods
of crop improvement may appear of lower priority at this time. However, it would be
a mistake to delay the installation and application of biotechnology for plant breeding
in Africa. Judging by the very rapid rates of adoption of transgenic varieties in countries where they have been introduced (James, 2000) and the increasingly routine use
made of other biotechnology applications in breeding programmes in the developed
world, farmers around the world can benefit from the use of biotechnology in crop
improvement.
To deny African farmers the benefit of these kinds of products would only put the
continent further behind in terms of the access its farmers have to modern agricultural
technology. The result would probably be continued low yields and higher food costs
than those enjoyed by the rest of the world. Therefore, while research planners in Africa
61
Chapter 5
62
Chapter 5
as elsewhere must strive to create a balance between investment in lower-cost,

easier-to-use, conventional methods and more complex methods of crop improvement,
there is significant value in gradually introducing biotechnology to African crop
improvement programmes. One way or another, African governments will need to make
decisions concerning agricultural biotechnology. To do so wisely there will need to be
African scientists who understand and can use the technology when appropriate.
5.2 Areas of Plant Biotechnology Research

While five areas of plant biotechnology are described above (tissue culture, markerassisted selection, genetic engineering, genomics and bioinformatics), the application of
only three of these will be reviewed in relation to biotechnology in Africa. Although the
latter two are of immense potential importance and undoubtedly can make significant
contributions to African crop varieties, the authors believe that they fall into a category
of upstream research which cannot be justified within the context of NARSs aimed at
reducing hunger in Africa.
Tissue culture
Tissue culture is based on the ability of some organisms to regenerate themselves from a
single cell or small clumps of cells. This is possible because the information necessary for
whole-organism growth, differentiation and regulation is present in each cell of the
organism. Tissue culture has been most widely used in Africa as a means of generating
disease-free propagules of vegetatively reproduced crops such as cassava, banana, potato,
sweet potato and yam. Such micropropagation is the most commonly applied form of
biotechnology in Africa (Cohen, 1998). Cost is usually the key factor in determining the
utility of tissue culture in commercial agriculture. Methods of meristem culture can be
adapted to local conditions such that facility expenses are low, yet field (post-flask)
survival rates are high. Tissue culture is in routine usage in numerous laboratories in
Africa, particularly in those countries where agriculture is based on cultivation of clonal
crops, such as the humid regions of west and central Africa. In addition, Kenya has used
tissue culture for rapid propagation of improved varieties of potato, pyrethrum and
sugarcane since the mid-1970s (Odame and Kameri-Mbote, 2000). Advances in tissue
culture techniques over the past decade have resulted in most crops being regenerated
from undifferentiated cells in vitro. Tissue culture is an important tool in transformation
methodologies discussed below. Tissue culture has been coupled with breeding programmes (as in the case of anther culture of rice) to speed up the regeneration, fixation
and duplication of genotypes of interest.
Examples of the use of tissue culture in crop improvement in Africa include:
A new rice plant type for West Africa resulting from embryo rescue of wide crosses
made between Asian rice (Oryza sativa) and African rice (Oryza glaberrima)
followed by anther culture of the hybrids to stabilize breeding lines (WARDA,
1998).
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Bananas propagated from apical meristem in Kenya have been shown to have
increased vigour and suffer lower yield loss from weevils, nematodes and fungal
diseases (ISAAA, 1997).
Non-governmental organizations involved in supply of planting material of
potatoes, sweet potatoes, cassava and banana to farmers affected by conflict in
Burundi multiplied planting stock through contracts with public laboratories that
used tissue culture to increase rapidly the number of clean seedlings available
(DeVries, 1999) (see Plate 6).
Marker-assisted selection (MAS)

The most common application of molecular genetics in plant breeding is markerassisted selection, which allows detection and localization within the plant genome of
genes controlling traits of interest to researchers. MAS applications are based on two
principal methods: (i) comparison of the differing products of reactions from DNAcleaving enzymes on DNA of alternative genotypes having differences in their base pair
compositions that alter cleaving sites (RFLP technology); and (ii) comparison of
patterns from the synthesis of repetitive sequences of DNA in alternative genotypes. The
latter uses relatively quick and inexpensive polymerase chain reaction (PCR) technology
and is preferable as a practical breeding tool for scoring numerous plants. Markerassisted selection was cited by CGIAR centres as the biotechnology application they
expect to be most useful in the future (CGIAR, 2000). It is also currently the application
on which the CGIAR is concentrating the greatest amount of resources (MAS accounts
for 28% of CGIAR biotechnology expenditures, the largest single item, followed by
genetic transformation, at 22%) (CGIAR, 2000).
Positive identification of genes within single plants allows for more precise selection
of the most favourable genotypes. Localization of genes along the chromosomes of
plants permits the construction of gene maps. Isolation of genes via molecular genetics
can permit their cloning in preparation for transfer using genetic engineering. However,
it is important to note that all methods of genetic marker selection rely on accurate
correlation being made between a given genotypes laboratory results and field level
performance. Thus, marker-assisted selection cannot succeed without an attached,
highly functional plant breeding capability.
MAS has a multitude of applications to crop improvement, but has proved
most useful as a tool to speed back-crossing of qualitative traits (those controlled by
a single gene or few genes) (Young, 1999). With marker-assisted back-crossing a
desired introgression can be achieved in four to six generations rather than ten or
more because the markers allow for more rapid elimination of undesired portions of
the donor genome while facilitating retention of the desired segment. In order to
maximize genetic variation for a given, quantitative trait (one controlled by multiple
genes), individual genes controlling the trait, termed quantitative trait loci, or QTLs
must be present in their most favourable format. By mapping these loci and tracking
their occurrence in large numbers of genotypes, it is possible to identify those
individuals with the most favourable make-up. As several traits can often be traced
using the same DNA, MAS holds the promise of more effective combining of
valuable traits within a given crop variety. At present in sub-Saharan Africa, MAS
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laboratories are under development in Zimbabwe, Kenya, Cte dIvoire, Nigeria and
South Africa.
Marker-assisted selection should be considered as a breeding tool when the following criteria are met with regard to the trait of interest.
The trait is important.

Phenotypic screening is reliable and accurate but difficult and/or expensive.
There already exists a large plant population segregating for the trait, preferably
with a simple pedigree.
Numerous mapped markers covering the whole genome are readily available.
Mapping of QTLs governing quantitative (multi-gene) traits, followed by their

selection based on detection of tightly linked molecular markers has been posited as a
means of improving the management of quantitative traits in breeding programmes
(Knapp, 1991; Dudley, 1993; Tanksley, 1993). While in principle this method could
become of great use in transferring complex traits (such as Striga resistance) from source
materials to improved varieties, in practice, this use of molecular markers has not as yet
proved feasible (Young, 1999). In addition to the cost of developing the genetic maps,
different QTLs have been detected for the same trait measured in different sites or in
progeny from different parentage (CIMMYT, 2000). This is an area that deserves review
and analysis, so that developing-country laboratories can decide whether QTL mapping
should be a priority for them. One approach that has been suggested by researchers at
CIMMYT (2000) is to analyse QTLs for complex traits in up to five different segregating populations and up to 20 different environments leading to the identification of
consensus regions for all genotypes. Marker-assisted selection strategies could then be
proposed which eliminated the need to construct new linkage maps for each new cross.
If successful, this will represent a major step forward in the employment of MAS
strategies for crop improvement.
Examples of the ways in which MAS may be applied to assist poor farmers in Africa
include:
Back-crossing the gene for resistance to maize streak virus (MSV) into well adapted
local varieties. Excellent genetic resistance to MSV has been known for over
20 years but not widely deployed. The locus of the resistance has now been located
on chromosome 1 and closely linked markers identified. Using markers it should be
possible to introgress this resistance into numerous well adapted local varieties.
This would be an excellent first use of molecular markers in national breeding
programmes.
Mapping of QTLs for resistance to weevils. Weevils are storage pests, which destroy
large portions of the harvest among Africas poorest farmers who lack modern grain
storage facilities. Natural resistance exists in several crops, however screening for
weevil damage among a large number of varieties is very laborious. Being able to
identify varieties with the genes for weevil resistance in the laboratory would
represent a major advantage for breeders.
Likewise, drought tolerance is a complex trait, controlled by many QTLs, that is
difficult to measure in the field in many seasons, simply because rains occur and all
stress symptoms disappear. Identifying molecular markers for this trait in a range of
cereal crops could permit the development of a wide range of materials with the
drought tolerance trait.
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Genetic engineering
Genetic engineering (also called genetic transformation) involves the direct transfer of
DNA between different varieties, species, genera and even between plants and animals.
It is the more invasive of the three primary applications of plant biotechnology and
therefore the one subject to the highest level of biosafety regulation. The two most
common methods of gene transfer are the intentional infection of recipient plant tissue
by Agrobacterium, natures own genetic engineer, and via bombardment of plant tissue
with particles coated with foreign DNA. Agrobacterium cells carry plasmids and can
cause a portion of the plasmid, called the transfer or tDNA, to become incorporated
into the plant genome the Agrobacterium infects. Through recombinant DNA technology, scientists can subtract deleterious genes from and add beneficial genes to the
tDNA prior to infection. Agrobacterium-mediated transfer was originally developed as a
technique for transforming dicotyledonous plants, but successful use of the technique
has now been reported for most cereal crops. Its major advantage over particle
bombardment is that it usually introduces just a single copy of the new genes to the
plant genome.
Genetic engineering is most commonly employed as a means of introducing a new
trait or variation for a given trait when naturally occurring variation is absent or
insufficient within the target crop species. A useful example is given by a number of
crops, including cotton, maize, potatoes and rice, which lack effective host plant
resistance to chewing insects. These crops have been transformed with gene constructs
of a protein produced by the bacterium Bacillus thuringiensis (Bt) that interferes with the
digestive systems of several genera of insect pests that chew and burrow inside the plant
and are therefore difficult to control with pesticides. Progeny of transformants have
shown significantly increased resistance to chewing insects (Barton et al., 1987;
Ghareyazie et al., 1997).
To date in sub-Saharan Africa genetic transformation of plants has been achieved
only in South Africa and Nigeria. In South Africa smallholder farmers are reported
to have adopted Bt cotton with great success (J.F. Kirsten, personal communication).
Also, for some outbreeding species like banana and cassava, where back-crossing is
problematic due to strong in-breeding depression, genetic transformation may be more
effective than conventional crossing as a means of moving desired genes into numerous
well adapted local varieties.
Examples of the ways in which genetic engineering might be applied to crop
improvement in Africa include:
Providing resistance to insect pests of cowpea. Cowpea is a highly nutritious crop

that grows well in marginal areas of Africa providing protein and vitamins to some
of the worlds poorest rural people. Its one serious constraint is susceptibility to
chewing and sucking insects. No sources of resistance are known to exist within
cultivated or wild cowpea genomes (Fatokun et al., 1997). Transferring a gene into
cowpea which confers broad-based resistance against such insects would reduce
losses to these pests for a large group of Africas poorest farmers.
Making rice a new source of dietary pro-vitamin A. Rice is an important staple food
in West Africa and its production is growing in other regions of the continent.
European scientists have recently added genes to rice that enable it to synthesize
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nutritionally significant levels of pro-vitamin A (-carotene) in the grain (Ye,

2000). Scientists at the West Africa Rice Development Association have produced a
new rice plant type from crosses between African rice and Asian rice which combine
desirable characteristics of both. These new rices are highly desired by farmers and
are spreading rapidly. If genes for -carotene production could be crossed into the
new plant type, a powerful driving force would exist for disseminating a new source
of pro-vitamin A to populations that need it.
5.3 The Interface Between Biotechnology and Breeding

The descriptions above make apparent the wide differences between alteration of crops
through plant breeding and the direct management of traits through biotechnology.
What is less obvious is the myriad ways in which biotechnology and plant breeding
interact. In fact, crop biotechnology is directly dependent upon plant breeding for its
impact. To appreciate the extent of the relationship between the two it is necessary to
explore the concept of the phenotype.
A varietys value to a given agricultural system is based on its phenotype. An
organisms phentoype is generally understood to arise from the combined forces of its
genetic make-up (its genotype), its environment, and the interaction of the two. Because
crop varieties are grown in different, relatively uncontrolled environments, knowing
their genetic make-up is critical, but insufficient in determining whether any given
individual will be of use in local farming systems. To make this determination, the
variety must be evaluated in numerous growing environments. Often, this evaluation
process involves making final selections, or fine-tuning, of the crop for final release as
a commercial product.
It is perhaps an unfair generalization to state that plant breeders manage the
phenotype while biotechnologists manage the genotype when, in fact, plant breeders
have referred to candidate genotypes based on carefully maintained pedigrees,
knowledge of gene expression and quantitative genetics, and the use of morphological
markers for nearly a century. However, plant breeders are primarily focused on the ways
plants reconstruct themselves phenotypically following a cross between two known
genotypes. Biotechnologists, on the other hand, are more focused on assembling genetic
constructs which have the basis for a given type of performance.
Molecular markers represent a virtually limitless set of loci for qualitative and
quantitative genetic analysis in any organism and are neutral with respect to both
phenotype and environment. However, initially to establish genetic linkage between
DNA markers and the loci for important traits requires the ability to observe and
measure reliably and accurately the phenotype in segregating populations under
different environmental conditions.
While identifying genetic markers and performing transformations are strictly
laboratory based activities, identifying valuable phenotypes can only be done under field
conditions. The interpretation of what is taking place, therefore, is almost always a
combined effort, and offers ample justification for systematically creating linkages
between programmes, without which much effort and expenditure can go to waste. It is
for this reason that making use of biotechnology in Africa is initially dependent on establishing functional, active, field breeding programmes.
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5.4 Biotechnology in Africa

While few are currently using advanced biotechnology methods, African institutions are
making progress toward becoming active practitioners of the techniques. In surveys
conducted by the International Service for National Agricultural Research (ISNAR)
in 1998 (Cohen, 1998), 37 countries of Africa reported an average of six full-time
scientific staff devoted to biotechnology research. The average annual expenditure
on plant biotechnology research in African countries was approximately $121,000.
Approximately 65% of this money came from international donor agencies. The
most important area of biotechnology techniques applied was micropropagation of
vegetatively propagated crops via tissue culture, which accounted for 52% of the total
number of activities. In most cases, African biotechnology laboratories are an add-on to
an existing crop improvement programme. This is a good strategy and helps to lower
cost, but they are still primarily donor funded. In only three countries of sub-Saharan
Africa (South Africa, Kenya and Zimbabwe) have national agencies been established to
promote agricultural biotechnology as a strategy for national economic development.
This is in sharp contrast to Asia and Latin America where governments have established
new agencies, such as the China National Center for Biotechnology Development
and the Indian Department of Biotechnology, that provide significant funding, over
and above their traditional agricultural R&D budgets, to build national capacity and
competitiveness in agricultural biotechnology. Even in Europe where opposition to
genetically modified organisms (GMOs) is most vocal, governments and industry
continue to invest significant new funds in development of agricultural biotechnology
research centres.
Most African countries are far from making this level of commitment and do
need to strengthen their seed distribution and conventional breeding programmes
before shifting significant resources to biotechnology. However, unless some greater
investment is made in training Africans who can understand the benefits and risks of
biotechnology and collaborate with researchers elsewhere, there is a real possibility the
biotechnology revolution will pass Africa by much as the Green Revolution previously
did. A necessary first step is the training of additional Africans in modern plant
breeding, including the application of new molecular and bioinformatic tools for crop
genetic improvement. Then, as they return from training, facilities would be needed
where they could use their new skills at their home institutions. The Rockefeller
Foundations experience in Asia suggests that an effective biotechnology capacity can
be built into an existing breeding programme by adding three or four well trained
scientists, about $100,000 worth of new facilities and equipment, plus $30,00050,000
per year in operating funds.
Expenditure on biotechnology applications by centres of the CGIAR involved in
crop improvement in Africa totalled $10.33 million (CGIAR, 2000). The exact portion
of that work directed at the needs of African farmers is unknown; however, if one
assumes 100% of that performed by the Africa-based organizations, IITA and WARDA,
and 40% (the CGIAR-wide portion of resources directed toward Africa) of that
performed by other centres, the approximate figure would be $5.7 million. The
Africa-related CGIAR centre with the largest biotechnology programme is CIMMYT,
with $3.2 million, followed by IITA, with $2.0 million (CGIAR, 2000).
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According to the ISNAR survey, within IARCs carrying out biotechnology research
in Africa, use of molecular markers was the leading technique applied, accounting for
75% of all activities. It should also be noted that several IARCs and several laboratories
in South Africa have reported successful transformation of important crop species,
including maize, cassava and cowpea. Viruses, insect pests and plant diseases ranked
as the three most important production constraints being addressed, each with
approximately 19% of the 94 total activities, followed by crop quality, with 12% of the
activity.
With regard to the crops reviewed in the ISNAR study, tissue culture methods have
been applied at numerous sites managed by IARCs. IITA maintains large tissue culture
facilities at its headquarters in Ibadan focusing on cassava, banana and yam. IITA is also
engaged in identification of markers for genes controlling resistance to Striga and is
actively pursuing genetic transformation of cowpea. WARDA employs anther culture in
the development of interspecific varieties of rice and has recently established a laboratory
for the use of molecular markers at its headquarters in Bouake, Cte dIvoire (see
Plate 7). CIMMYT has applied MAS for several important traits of African maize,
and is assisting in the creation of national biotechnology laboratories in Kenya and
Zimbabwe.
Several biotechnology projects are nearing implementation stages within African
NARSs. However, the distribution of these projects is highly skewed in terms of the
institutions and crops involved.
1. Marker-assisted selection (MAS) for maize streak virus resistance is scheduled for
implementation in Kenya during 20012002 (KARI, 1998).
2. MAS will also be applied to drought and stem borer resistance in Kenya and
Zimbabwe during the same period (CIMMYT, 1998).
3. KARI and CIMMYT have initiated a 5-year programme aimed at developing
insect-resistant maize varieties using transgenic B.t. technology.
4. A six-country programme funded by the Swedish International Development
Agency (SIDA) is aimed at enhancing and broadening capacity in biosafety and
biotechnology in eastern Africa.
5. KARI has imported and begun multiplying transgenic sweet potato varieties
resistant to feathery mottle virus.
6. The Ugandan government has agreed to fund a 5-year initiative implemented by
INIBAP, the National Agricultural Research Organization of Uganda, and others to
develop transgenic East African highland bananas with resistance to black sigatoka
disease, nematodes and weevils (INIBAP, 2000).
7. Researchers at the University of Zimbabwe have used a gene construct for resistance
to aphid-borne mosaic virus to achieve resistance in transgenic tobacco (Mlotshwa,
2000) and are now collaborating with Michigan State University to transform cowpea
with the gene (Sithole-Niang, 2000).
In conjunction with the projects listed above, and others, some countries have
invested in biotechnology infrastructure. In addition to the many facilities in operation,
which make use of tissue culture methods, the following are either already assembled or
nearing completion.
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1. In Zimbabwe, a large research campus has been developed at the Scientific and
Industrial Research and Development Center. One of the SIRDC institutes is dedicated
to biotechnology research. Five full-time SIRDC scientists are currently working on a
variety of topics, which range from wine making to fingerprinting of sweet potatoes, to
marker-assisted selection of drought tolerance in maize.
2. Also in Zimbabwe, at the University of Zimbabwe, a small laboratory has been set
up in the Crop Science Department for identifying QTLs for resistance to Striga in
sorghum.
3. At the Centre Nationale de Recherche Agricole, in Abidjan, a dedicated biotechnology
laboratory has been set up for performing mapping studies and genetic transformation
of cassava and yams.
4. In Kenya, at the National Agricultural Research Laboratory, the Kenya Agricultural
Research Institute is building a biotechnology laboratory suitable for a range of
biotechnology applications.
Also worthy of mention are: genetic transformation of tobacco has been achieved
at the Tobacco Research Institute in Harare, Zimbabwe; and several laboratories in
South Africa have confirmed routine capacity for genetic transformation of agronomic
crops.
However, the level of activity in biotechnology is a poor indication of the overall
human capacity on the continent. In fact, a much larger number of scientists
have acquired skills and knowledge, but they are often unable to use them due to lack
of facilities and funding for research and they are seldom brought together to form
the critical mass of talent necessary to make real research progress (Kezire et al.,
2000; Odame and Kameri-Mbote, 2000). Unless the level of activity in biotechnology
increases dramatically in the coming years, this capacity could begin to erode for lack
of practice. One way of helping these scientists to make progress and keep up to date
is to establish collaborative research linkages which enable them to spend time
(at least 23 months per year) conducting research relevant to their home country
at an advanced laboratory in Europe or North America where equipment, supplies,
information, new methods, mentors and peers are all more readily available. The
Rockefeller Foundation calls such arrangements career fellowships and experience
indicates that the fellows work long and hard, accomplish much, and make important
contributions to the host laboratory as well as their home institution.
Another threat to the application of existing capacity at present is the anti-GMO
campaign being waged by a number of individuals and agencies primarily based outside
Africa. Concerns raised by these groups have slowed the deployment of transgenic crops
throughout the world (Paarlberg, 2000). Several IARCs are also unable to field-test
transgenic varieties due to recently initiated national regulations on transgenics
(CGIAR, 2000; Johanson and Ives, 2000). One positive sign that progress toward
assessing such new technologies would not be completely stopped was the official
importation in Kenya in March 2000 of transgenic sweet potato cuttings with resistance
to feathery mottle virus (Daily Nation, 19 August 2000).
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5.5 Biotechnology for African Crop Challenges Summary

This chapter has argued that because of limitations in the downstream areas of breeding
and seed systems, programmes concerned with biotechnology applications for Africa
should not be limited to biotechnology. In order for investments in biotechnology to
have an impact in the lives of the poor, they must be made in tandem with targeted
support to downstream, field-based activities.
Yet there are numerous crop improvement challenges in Africa which will not
respond to conventional efforts. Fortunately, a core group of African scientists have
already received training and others will soon return who are ready to apply their new
knowledge to priority constraints of the poor. Given the promise of the technology
itself, and given the size of the challenge in attempting to make major differences in crop
performance under marginal conditions in Africa, it seems clear that now is the time to
begin developing and applying relevant biotechnology applications in Africa.
Commercial biotechnology firms have shown limited interest in applying their
capacity to resolving food security needs in developing countries (Persley, 1999). In the
case of Africa, this trend is not expected to change in the near future. Therefore, if
biotechnology is to have the anticipated impact on the lives of Africans, it will be due to
investment in public sector capacity in biotechnology applications.
Building on existing national commitment to biotechnology capacity, continued
development and use of tissue culture techniques, where applicable, should pay early
dividends to crop improvement programmes. Marker-assisted selection aimed at
combining numerous resistance traits in a single crop variety is another example of a
potential biotechnology application that is relevant and accessible for IARCs and
national programmes to adopt. Its use in Africa should be expanded as NARSs gain
capacity in molecular methods and the technology itself is improved. IARCs can play a
critical role in backstopping the setting up of molecular laboratories and the integration
of molecular selection techniques within African NARSs.
While the development of transgenic crop varieties by NARSs in Africa is as yet
some way off, preparations can proceed to assure biosafety measures are in place to
permit their evaluation, introduction, and eventual development. According to Ndiritu
(1999), South Africa, Kenya, Zimbabwe, Botswana, Malawi, Mauritius, Cameroon and
Zambia either have or are in the process of adopting explicit biosafety regulations and
guidelines, and are participating in the negotiations for an international biosafety
protocol. The ability of national programmes to assess safely the value of transgenic crop
varieties is dependent on containment facilities, which are currently lacking in most
African countries. Increased testing and verification capacity of transgenic materials
within NARSs is also essential to increasing access to products of biotechnology research
based elsewhere.
5.6 The Potential of Apomixis

With regard to crop varieties, what Africa really needs for all major food staples is a large
number of locally well-adapted, improved cultivars that can increase both the quantity
and stability of production, plus more effective mechanisms of distributing these
cultivars to all farmers. Due to Africas many unique ecological and socio-economic
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niches and its limited markets and infrastructure, farmer participation in cultivar
development, selection and dissemination will be essential in meeting these objectives.
If apomixis could be introduced as a flexible new tool for breeding African crops, it
would greatly facilitate farmer participatory breeding, lower the cost of producing new
varieties, improve their performance under all growing conditions, and speed the
process of seed delivery, including farmer-to-farmer trade.
Apomixis (agamospermy) is a form of asexual reproduction through seed that
occurs naturally in some plants. It is a genetically controlled process that bypasses female
meiosis and fertilization to produce seeds genetically identical to the maternal parent.
The seeds formed and the progeny plants they produce are true breeding genetic clones
ready for immediate performance evaluation. In recent years there has been progress in
identifying the genes controlling the components of apomictic seed development and
in moving these genes into crop plants (Jefferson and Bicknell, 1996; Veille-Calzada
et al., 1996; Grossniklaus et al., 1998; Ozias-Akins et al., 1998; Luo et al., 1999). While
no new apomictic crop varieties have been released, several relevant patents have been
issued, primarily to public sector research organizations. For use in Africa, apomixis
would have the following advantages.
For essentially all crops, numerous genotypes that performed well under local
conditions could be genetically fixed early in the selection cycle and developed
directly into cultivars desired by farmers. Under such a scheme, variability would be
generated through traditional hybridization with the population of resulting plants
grown and evaluated by farmers under local conditions. The plants that performed
best would be selected and crossed with an apomictic male parent. The resulting
progeny plants would be apomictic and the best could be selected by farmers as true
breeding, superior cultivars. Farmers would thus become key actors in the breeding
of diverse cultivars for diverse environments. The number of true breeding, locally
well-adapted, superior cultivars would be large enough to encourage the use of
mixtures of cultivars, thus enhancing the genetic diversity of the crop both locally
and regionally.
Many of the cultivars selected by farmers and genetically fixed by apomixis would
have hybrid vigour. Such hybrid cultivars not only have greater productivity under
good conditions, but also have increased stability of production under adverse
conditions. Apomixis would bring the benefits of hybrid vigour to numerous crops
and to many smallholder farmers who never previously benefited from hybrid
technology.
Important African crops such as cassava, sweet potatoes, yams and potatoes which
are traditionally propagated vegetatively could be converted to true-seed propagation. These are polyploid crops that are difficult to breed. Their seeds either segregate genetically or suffer from severe inbreeding depression when plants are selfed
to make them true breeding. Consequently, elite cultivars are usually propagated
via tissue segments that are genetic clones of the donor plant. However, the rate of
vegetative multiplication is slow (roughly six offspring per parent plant), pathogens
are often transmitted along with the tissue segments, and the costs of storage,
shipping and planting are high. With apomixis, elite cultivars would produce seeds
that are genetic clones of the parent plant and that are, for the most part, pathogen
free due to the normal pathogen elimination mechanisms associated with
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seed development. African farmers could save, trade and disseminate seeds of
their favourite elite cultivars much as Asian farmers saved and traded the seeds
responsible for the Green Revolution.
Apomixis as a flexible breeding tool has the potential to be one of the most important
innovations in the history of agriculture, benefiting all farmers, including those who
have benefited little from previous innovations. While considerable research is still
needed to make this a reality, the power of recent advances in plant molecular biology
and progress to date in understanding the biological mechanisms of apomixis make
this a challenging but achievable goal. A mission-oriented international research
collaboration designed to share results with everyone would enable all crop breeding
programmes, including those serving farmers in Africa, to benefit fully from the
technology.
5.7 Intellectual Property Rights

The free exchange of materials and information has been a hallmark of the international
agricultural research system. Such exchange has been key to past successes of the system
and it will be key to future successes in Africa. The genetic improvement of plants
is a derivative process, in which each enhancement is based directly on preceding
generations, and the process of adding value requires access to the plant material itself.
Most important food crops originated in developing countries, and much of the value in
todays seeds has been added over the centuries, as farmers have selected their best plants
as a source of seed for their next crop. Traditionally, these landraces and the indigenous
farmer knowledge associated with them were provided free of charge to the world
community. In exchange, public sector research and breeding programmes added value
and returned scientific knowledge and improved breeding lines as global public goods
to developing and developed countries. Now, however, the rules of the game are
changing, even before Africa has had the opportunity to benefit much from such global
public goods.
Over the past decade, in industrial countries, applied crop-biotechnology research
and the production of improved seeds have increasingly become functions of the
for-profit private sector. This has led to a significant increase in the total research effort
committed to the plant sciences and crop improvement, but the results of such research,
in both the public and private sectors, are now often protected as various forms of
intellectual property, including patents, material transfer agreements, plant breeders
rights and trade secrets. Furthermore, intellectual property rights (IPR) are globalizing.
Industrial countries have made IPR an important component of international
trade negotiations where they use IPR to exploit their competitive advantage in
research and development. Larger developing countries seeking to join the World Trade
Organization have been required by the Trade Related Aspects of Intellectual Property
Rights (TRIPS) provisions to put in place IPR systems that include protection of crop
varieties. The least developed countries, including most in Africa, have until 1 January
2006 to implement such IPR systems. However, since most African farmers cannot
afford to purchase new seed for each planting, it is important that Africas new plant
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variety protection laws include provisions allowing farmer-saved seed and use of varieties
as a resource for further breeding. This is in contrast to granting utility patents on plants
which extend protection to the progeny and its seeds such that breeders cannot legally
use protected varieties as breeding material. The International Undertaking on Plant
Genetic Resources for Food and Agriculture needs to be approved and implemented
as a means of ensuring the conservation, sustainable use, and free flow of seeds for the
benefit of people in Africa and elsewhere.
Protection of intellectual property is to be expected when dealing with for-profit
companies. The major IPR change that is threatening the operations of the international
agricultural research system has occurred within public sector plant research institutions.
In the USA, the 1980 BayhDole Act gave universities and other public-funded research
institutions the right to obtain patents on and commercialize inventions made under
government research grants. Similar arrangements are emerging in Europe, Japan,
Australia, and most other industrialized countries. The result is that while the majority
of the significant discoveries (e.g. pathogen-derived plant resistance to virus infection)
and enabling technologies (e.g. biolistic transformation methods) are still generated with
public funding in research institutions and agricultural universities, these discoveries are
no longer being treated as public goods. Rather, they are being patented and licensed,
often exclusively, to the for-profit sector. Such discoveries now primarily flow from the
public sector to the for-profit sector and if they flow back out usually come under
material transfer agreements (MTA) which significantly restrict their use, usually for
research purposes only. Crop genetic improvement is a derivative process and each
incremental improvement that involves biotechnology now comes with a number of IP
constraints which accumulate with each transfer or further improvement. To deal with
this predicament, the private sector is becoming greatly centralized into a global
oligopoly dominated by five leading firms. They are the product mergers made in part to
accumulate the IP portfolios necessary to produce biotechnology-derived finished
crop varieties with freedom to operate. Such consolidation could lead to a loss of
competition among purveyors of agricultural technology and make it excessively
difficult for new firms to enter the industry, worldwide, and may constitute arguments
for restricting the exercise of intellectual property rights (Barton, 1999).
The publicly funded agricultural research community, however, has not followed
suit. Leading academic researchers are primarily interested in research competitiveness.
They readily sign research MTAs to keep competitive but are then restricted from
further transferring their research products. Their universities now have technology
transfer offices where the incentives are to maximize IP royalty income, often by
granting exclusive licences. The net result is that improved plant materials produced by
academic scientist-inventors are highly IP-encumbered and commercially useful only to
a big company having an IP portfolio large enough to cover most of the IP constraints.
The international agricultural research system does not have such an IP portfolio and as
a consequence the traditional flow of materials through the system is breaking down,
particularly at the point where useful new technologies and improved plant materials
flow from public sector researchers in developed countries to international centres and
national crop improvement programmes in developing countries. Africa, in particular, is
being short changed of the benefits of biotechnology because, unlike Asia and Latin
America, it has little capacity to replicate research results patented elsewhere, for the
73
Chapter 5
74
Chapter 5
benefit of poor farmers in countries where the IP is not protected. Africa is much more
dependent on partnering with others, but publicly funded researchers in industrial
countries are no longer partners who can freely share their most important discoveries.
A new mechanism is needed, such as universities retaining the right to grant
charitable licences, and then pooling such licences into an IP portfolio designed to
facilitate use of research results to help food-insecure subsistence farmers in places like
Africa. Such an IP portfolio could help reinvigorate the international agricultural
research systems by re-establishing the flow of advanced scientific knowledge and
research materials to and through the system for the benefit of smallholder farmers in
Africa and other poor countries.
74
Chapter 5
Seed Systems: Reaching the

Poor in Numbers
6.1 Overview
Seeds form the foundation of all agriculture. Without seed there is no next seasons crop.
The genetic traits embodied within seeds reflect and determine, to a large extent, the
nature of farming systems, themselves. In turn, the nature of seed systems (the term is
used herein to describe the prevailing methods for accessing, storing, and exchanging
seeds and other planting material) determine in large measure who benefits from the
advances made in plant breeding and biotechnology. Perhaps most important within the
context of this book, seeds serve as a tangible representation of technologies developed
for use and long-term ownership by poor farmers. Once seed is obtained, farmers can
use it at will in directing their own advancement.
Seed is consistently recognized as the most important and least expensive of cash
inputs for farming (Venkatesan, 1994). The importance attributed to seed is most often
described in terms of its role as the factor which sets the upper limits on productivity and
yield stability (Morris, 1998; Srivastava and Jaffee, 1993), although at a far more basic
level, the sheer availability of seed has often affected African farmers ability to sow a
crop (Cromwell, 1996).
The seed demandsupply relationship in a large portion of Africas smallholder
farming systems appears to represent a situation of market failure: farmers in need of
seed of particular qualities cannot afford to pay for them at rates which would make it
attractive for suppliers to enter the market. The resulting, low effective demand for seed
of varieties designed for the growing conditions of low-input farms effectively prevents
their development. In agricultures sliding scale of efficiencies, moreover, the lack of this
key, initial input may prevent farmers from moving upward to productivity levels where
private sector exchanges operate more efficiently. As an example, yields of maize in the
USA during the early 20th century had reached a plateau at approximately 1.5 t ha1
prior to the introduction of hybrids in the mid-1930s (Chrispeels and Sadava, 1994).
Subsequent phases of rapid yield increase (and rapid expansion of the seed industry)
75
76
Chapter 6
were brought on by the successive introductions of double-cross, three-way and single-cross hybrids to produce current average yields of over 7 t ha1 (Norskog, 1995;
Crow, 1998). Sustainable increases in yields in Africa are likely to be less than in the
USA, however, there is little doubt that the current, unmet demand for improved,
seed-based technologies represents a key factor in Africas declining per capita agricultural productivity.
Several authors have pointed out that seed systems in Africa and other developing
regions have received less attention in proportion to their importance than other
sub-sectors, such as agricultural research and extension systems (Venkatesan, 1994). Perhaps because of perceptions that the potential of improved varieties of basic food crops
in Africa did not duplicate or was less stable than results obtained in Asia, seed systems in
Africa have received less attention than merited. Poor access to seed among small-scale
farmers in Africa has been recognized as a major constraint to crop improvement by several authors, yet facilitating consistent, broad-based access to seed of improved crop varieties remains a complex issue with no simple solutions. In a recent review of seed systems
in Kenya, Malawi, Zambia and Zimbabwe by Tripp (2000), several different types of
seed demand are identified, including demand for improved varieties with higher yield,
demand for re-supply of seed lost following disasters, demand resulting from poverty
and chronically low yields, and demand for fresh seed of known varieties. This chapter
will endeavour to explore these four types of demand and offer some practical suggestions toward meeting the different needs expressed by them.
Beginning in the late 1980s and early 1990s, the seed sectors of most African
countries were progressively liberalized. Many formerly monopolistic, parastatal seed
companies were privatized and regulations were relaxed to allow the entry of wholly
private firms (Cromwell, 1996). While over the long term this policy is likely to pay
important benefits for farmers through gains in efficiency and competition, in the short
term the change in policy has meant that government and donor agencies which
previously supported seed supply via public institutions have lost much of their ability to
do so, as public seed companies have been privatized and overall responsibility for seed
distribution has been shifted to the private sector. As hybrid maize seed markets
represent the primary incentive for private seed sales, countries where the primary food
staple is other than maize (including all of West Africa) have suffered particularly low
levels of seed sector investment.
Responding to demand for seed of hybrid and commercial crops grown in Africa
such as cotton and maize, appears to be a relatively straightforward matter of applying
sound business and technical strategies common to private seed sector development
anywhere, albeit one compounded in terms of complexity by Africas vast size,
underdeveloped infrastructure, and very low farmer incomes. Responding to the
low-level or intermittent demand for open-pollinated and non-commercial crops such as
cowpea and cassava, however, has yet to be performed successfully by purely private
companies, and thus implies continued involvement of public agencies, NGOs, and
small, grass roots farmer associations (Sperling, 1994; David et al., 1997). Regardless
of the type of agency involved, however, the size of the seed problem in Africa and
its overwhelming importance in delivering the hope and promise of crop genetic
advancements to farmers, merits close examination and broader adherence to strategies
that work.
77
Seed Systems
6.2 Issues in Seed Supply

Overview
Total seed consumption worldwide is estimated at 120 million tonnes per year (Kelly
and George, 1998). In developing countries, over 80% of seed of staple food crops is
farmer-saved seed (Jaffee, 1991). As the vast majority of seed used in Africa is farmersaved seed, estimates of total consumption must be calculated using areas planted and
common seeding rates, coupled with observed rates of variety replacement or renewal of
seed stocks. Estimates of annual seed consumption for a number of species in Africa are
presented in Table 6.1. These calculations show that total annual seed consumption of
major crops in Africa is approximately 2 million tonnes. Using restocking rates for
self-pollinated crops and non-hybrid seed of open-pollinated crops of once every 3 years,
and restocking rates for hybrid maize of once per year point to a potential seed market of
approximately 700,000 t per year.
Farmer-saved seed
The vast majority of seed used on farms throughout the world is seed saved from season
to season using a wide variety of techniques. The first step involved is selection of the
portion of harvest to be kept as seed. Several reports suggest that farmers make their
selections on the basis of plant quality characters in the field (Wright et al., 1995;
Scowcroft and Polak Scowcroft, 1997). However, recent studies conducted in Ghana
and Zambia found that less than 25% of farmers actually select seed in the field (Walker
and Tripp, 1997).
On-farm seed storage practices vary from region to region and crop to crop.
Farmers in central Mozambique construct special silos of woven grass hoisted on poles.
Farmers in southern Sudan store seed in racks placed above cooking fires where smoke
acts as an insecticide. In Somalia, farmers store sorghum seed in underground pits
covered with woven mats and soil. In general, such techniques appear adequate for
Table 6.1. Estimates of annual seed consumption for principal cereal and pulse
crops in Africa.
Groundnut
Maize
Rice (upland)
Rice (lowland)
Beans
Cowpea
Sorghum
Millet
Total
Area planted
(million ha)
Seeding rate
(kg ha1)
Total
(1000 t)
9.0
25.2
3.1
4.7
3.2
7.2
23.0
20.2
85.5
80
20
60
20
40
10
5
4
720
504
186
94
128
72
115
81
1900
78
Chapter 6
preserving cereal seed viability. Legume seed, which is more difficult to produce and
store, presents added difficulties (Desai et al., 1997). Farmers in Mali, for example, often
lack access to viable groundnut seed at planting time. With the exception of cowpea, germination rates of farm-stored seed of four grain crops exceeded the nationally accepted
minimum germination rates in Ghana (Walker and Tripp, 1997). Cowpea also ranked
lowest in germination rate of saved seed among six crops in a survey conducted in
Mozambique (Sitch, 1996).
With farmer-saved seed being such a common practice in Africa, the question must
be asked whether introduced seed does, indeed, represent an advantage to small-scale
farmers. The answer to this question is bound up in the complex, multipurpose
decision-making environment of seed users who are at the same time homesteaders,
commercial producers and members of small communities. One principle of varietal
adoption (and thereby, out-sourcing of seed, at least once) which is becoming
increasingly accepted is that farmers must first observe the new variety growing in their
own farming environment for an entire cycle, during which they will evaluate its overall
usefulness in the context of the various purposes they hold for that crop. Nevertheless,
when benefits are perceived, in general, it can be stated that sentimental attachment to
local varieties, while often present, will generally give way to more practical concerns
over increasing the harvest (Muhhuku, 2000). In most cases, however, farmers will
continue to cultivate old varieties alongside new ones for some time.
Therefore, on-farm, farmer-managed trials of experimental varieties represent one
of the most critical stages of crop improvement. Unfortunately, the rather unglamorous
(and, relatively speaking, expensive) work of multiplying sufficient seed, transporting it
to the field, and the laying out and planting of plots has often been neglected by breeding teams and donor agencies alike. The logistical complexities of multi-location varietal
testing (often compounded by lack of efficient means of communication, bad roads, fuel
shortages, etc.) are likely to hinder the effectiveness of crop improvement work,
including biotechnology, for some time to come. Given the reality of the ways in which
small-scale farmers in Africa make decisions on new varieties, however, it is logical to ask
whether funding of upstream work on biotechnology and breeding is worthwhile if no
one is willing to support and conduct the multi-location, participatory testing phase.
Farmer-saved seed stock stored using local methods is intermittently subject to
ruptures caused by drought, pest and disease outbreaks, and civil unrest. Renewal of
depleted seed stocks caused by at least the former two occurrences would, in many rural
economies, represent opportunities for seed companies. In Africas highly depressed
economy, such demand has so far largely remained ineffective, that is, insufficiently
backed by purchasing power to stimulate the creation of commercial supply networks.
Nevertheless, recent studies have shown that even very poor farmers will purchase seed
when it is sold via appropriate channels (David and Otsuka, 1994; David and Sperling,
1999; Rohrbach and Malusalila, 2000).
Over the past decades, large amounts of donor and government funding in Africa
have been dedicated to establishing seed supply systems serving rural areas. However,
much of this funding was applied during a period when seed supply was controlled by
the public sector. The experience gained under this regime is of little use in determining
how best to apply new funding under liberalized market conditions, making seed
systems one of the most experimental and open-ended working environments of the
entire crop improvement process.
Seed Systems
79
Seed organizations
When seed is not saved on-farm, African farmers procure new seed from a variety of
sources. In the Mozambique survey cited above, 53% of farmers reported redistribution
of their own seed stocks to other farmers. Sixty per cent of the recipients of this seed
were family members, while 28% were neighbours. Seventy-four per cent of these
transfers were done for free (Sitch, 1996). In Zambia, studies also revealed that most
off-farm seed was procured from other farmers as a gift, while in Ghana, the majority
was purchased (Walker and Tripp, 1997), either in the marketplace or from other
farmers.
While a large portion of seed used on African farms is likely to continue to be
farmer-saved seed, seed dissemination capacity continues to be an important factor to
the overall impact of crop improvement programmes. In order for advances in crop
improvement to reach the farmer, someone must first supply the seed. Aside from farmers themselves, a number of entities are involved in seed supply in Africa. These
are identified below, along with brief characterizations of their respective modes of
operation.
Multinational companies
These are largely publicly traded companies represented in more than one country, often
involving worldwide operations. Recent developments in biotechnology and intellectual
property regimes have caused a rapid consolidation of the international seed industry,
recently chronicled by James (1998) and James (2000), with DuPont (incorporating
Pioneer Hi-Bred International), Syngenta (Novartis and Astra-Zeneca), Monsanto
(Agracetus, Asgrow, Calgene, Dekalb, Holdens, among others), AgrEvo (Hoechst,
PGS, Rhone-Poulenc, Sun Seeds) and Dow Elanco (Dow Chemical, Eli Lily and Co.)
emerging as the largest actors.
Private sector investment in research in developing countries has remained
relatively small compared with their total budgets (Byerlee, 1996). While all of the
companies listed above have at least some presence on the African continent, their
presence in the seed markets of sub-Saharan Africa is relatively limited. In Africa, the
group is represented by several joint ventures with the main focus on hybrid maize and
cotton seed sales. To date, most of the investments made by multinational companies in
sub-Saharan Africa have been concentrated in a few countries of eastern and southern
Africa, including Kenya, Malawi, Zimbabwe and South Africa. Extensive earlier
investments made by Pioneer Hi-Bred International in West Africa (Cameroon, Cte
dIvoire, Nigeria) were written off in 1993 (Rusike and Eicher, 1997).
Multinational seed companies often enter into multiple country seed markets via
breeding and seed production operations based in one or two. Several companies have
breeding operations in South Africa and Zimbabwe which are used to develop hybrids
for additional markets outside those countries. They rely on the knowledge of their
breeders plus extensive participation in national variety trials to produce varieties
developed from proprietary parental materials already within their seed banks for release
in distant ecologies.
The principal advantage of multinational companies in gaining access to local
markets is their size, which allows them to manage large collections of broadly adapted
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Chapter 6
germplasm and make investments in seed markets, which may take several years to show
a profit. A major disadvantage of the current mode of operation of multinational
companies in Africa is their reluctance to set up individual breeding operations in each
country, in part due to their high overheads in research and promotion of new varieties
(Byerlee and Lopez-Pereira, 1994). Minimum investment levels were recently estimated
by one large seed company at approximately $5 million. As such, their releases, although
often bearing useful traits (such as early maturity or, in the case of maize, resistance to
maize streak virus), lack genuine adaptation. Market share of multinational companies
in the major hybrid maize markets of Africa has remained small (Tripp, 2000).
Former national seed companies
Former national seed companies are the surviving entities of privatized (partially or
fully), parastatal companies. Few such companies still enjoy monopoly status, although
low or ineffective competition in many countries means that they continue to function
as oligopolies, with very large market shares (Tripp, 2000). In countries where there
has been appreciable competition, they are operating alongside private companies, competing for the same market. Such companies have historically enjoyed direct, formalized
relations with NARSs, with significant carry-over benefits during the liberalization
process. As a result, they continue to embody significant advantages over newcomers via
their ownership of adapted, improved varieties developed prior to market liberalization.
This fact has allowed public companies to carry seed of a wide range of crop species at
relatively little added cost. With deregulation, however, most NARSs are now free to
supply new releases to the highest bidder, and relations with public seed companies have
changed.
Private, national companies
Outside of a few countries (South Africa and Zimbabwe, and more recently Kenya and
Malawi), seed markets in Africa have not experienced the proliferation of private,
independent companies which deregulation of the seed industry should permit.
However, in Kenya, where seed industry deregulation has been more or less complete,
a significant number of small companies focusing on niche markets have been registered
within the past few years (John Kedera, personal communication).
Private, national companies may yet play an important role in seed distribution in
Africa, as they have in India and Brazil (see Lopez-Pereira and Filippello, 1995). By
sub-contracting most tasks, operating in areas they know well, and taking advantage of
close contacts with national breeding programmes, private national companies can
potentially attain the sort of efficiencies which would allow them to undersell multinationals. However, these arrangements have been cited as a constraint to growth by
Tripp (2000), in part due to the lack of medium- and large-scale growers who can
produce large quantities of seed on contract.
Farmer associations
Farmers organizations have often become involved with seed supply as a means of
guaranteeing timely access to seed of acceptable quality for producing a crop. The Maize
Seed Association of Zimbabwe and the Kenya Farmers Association are two large-scale
Seed Systems
81
examples of this type of group. Such groups have usually been dominated by the
interests of large-scale farmers, and often have involved official state sponsorship or
substantial support from donor agencies. Moreover, their focus on issues more related to
marketing of produce than with the supply of seed will tend to reduce their competitiveness in the future, and large farmer associations are not projected to be a significant force
in seed supply over the medium and longer terms. More recently, smaller associations
which operate locally have been involved in seed multiplication and distribution, and
proved a certain level of effectiveness (Mduruma, 1999). (See Box 6.1.)
NGOs
As regulations controlling use and distribution of seed have been relaxed, NGOs have
become increasingly involved in issues of seed supply, especially in the context of
emergency relief operations directed at rural populations whose farming activities have
been disrupted by conflict, drought, or other environmental disasters (Cromwell, 1996;
Osborn, 1996; Chapman et al., 1997; Tripp, 2000). Quantities of seed distributed by
NGOs in the context of emergency relief programmes can be measured in the tens of
thousand of tonnes, and are growing (Osborn, 1996). The seed-related activities of
NGOs and donor agencies in the context of relief programmes have been well
documented in separate studies conducted by the Overseas Development Institute
(1996) and Osborn (1996), which estimated that $10 million were being spent annually
in nine countries of eastern Africa. In 1994, a single organization (World Vision)
distributed seed to over 300,000 farm families in Mozambique alone (see Box 6.2).
Other major theatres for emergency seed distribution campaigns in Africa have been
Kenya, Somalia, Liberia, Sierra Leone, Angola, Rwanda, Burundi and the former Zaire.
Uganda was recently the focus of a major campaign aimed at distribution of cassava
varieties resistant to African cassava mosaic virus (Otim-Nape et al., 1997).
In such contexts, the dire need of farmers, coupled with the opportunity of
obtaining finance from donor agencies, has led many NGOs with inadequate knowledge and experience in agriculture into the distribution of seed. This is problematic for
a number of reasons. In many cases, opportunistic traders have sold such agencies seed
which was, in fact, grain. In other cases, NGOs have imported seed of varieties which
were not adapted and which produced very poor yields. Nevertheless, a small number of
more technically oriented NGOs have distinguished themselves in the area of seed relief
and agricultural recovery programmes through effective responses to seed shortages.
In view of the large quantities of seed being distributed through relief programmes
in Africa, it is now essential that these schemes become integrated with national and
international crop variety development efforts. Distribution of inappropriate or poor
quality seed by organizations who have little understanding of adaptation and variety
performance issues needs to be curtailed. Donor agencies (who fund such activities) and
NARSs (who should regulate them) are perhaps the most important levels of control in
this area.
Seed distributed through long-term development projects operated by NGOs is
rarer and poorly documented. Because seed is a valued commodity, seed distribution
under non-emergency conditions often blurs the lines between humanitarian and
commercial operations, and distribution of free seed by NGOs has at times been
criticized by private companies as a deterrent to market development. Meanwhile, the
ease of supplying donor-funded, emergency seed programmes has often caused private
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Chapter 6
Box 6.1.
Proving it can be done: Seed Co Ltd.
In searching for solutions to the problem of seed access among small-scale farmers
of Africa, the debate often turns to the lack of investment in private seed networks
and, eventually, questions over whether the sale of seed to small-scale farmers in
Africa can be done profitably. Africas dispersed and underdeveloped markets,
poor infrastructure, and low farmer incomes present obvious challenges to the
expansion of private seed markets. Nevertheless, a response to the question may
already be available in Zimbabwes Seed Co.
The origins of Seed Co date to 1940, when the Seed Maize Association
was formed at the request of the government to multiply and market popular openpollinated maize varieties (McCarter, 2000). In 1957 a second association, the
Crop Seeds Association, was formed to concentrate on seed production for selfpollinated crop species. In 1983 the two associations merged to form the Seed
Co-operative Company of Zimbabwe Ltd. In 1996 the company was renamed Seed
Co Ltd and was listed on the Zimbabwe Stock Exchange. Today, Seed Co markets
37 varieties covering seven crop species (Seed Co, 2000). The company is present
in Mozambique, Zambia, Zimbabwe and Malawi and has annual regional sales of
approximately 50,000 t. Equally important, an estimated 80% of its customers are
small-scale farmers, who purchase certified seed of improved varieties in packs
ranging in size from 0.5 to 50 kg. According to Seed Co:
An important key in the development of the Zimbabwean seed industry was
the signing of legal agreements between the Ministry of Agriculture and Seed
Co that entitled the company to the exclusive right to multiply and market a
range of Government bred products. In exchange, the company had to undertake to produce agreed volumes of seed, including 2030% carryover, and to
sell seed at agreed prices. These agreements have resulted in large volumes of
quality seed being made available to Zimbabwean farmers at prices three
times lower than those of South Africa and approximately one-ninth of the
United States . . . Seed Co now employs nine breeders and owns two research
stations
(McCarter, 2000).
In todays environment of open competition among multiple companies,
such collusion between government and a single company would not be permitted,
and prior to becoming reorganized as a private company, the relationship between
government and Seed Co often came under criticism. To date, for example, there
are no sales of open-pollinated maize seed in Zimbabwe, a policy which was
maintained in part with the support of Seed Co. Mozambican farmers, on the
other hand, have been able to purchase good quality OPVs for nearly a decade.
Moreover, it is possible that competition between Seed Co and more recent entries
into the region (Pioneer, Monsanto and Pannar (South Africa) are now active in
Zimbabwe and several other countries) will result in even lower prices and higher
quality products. Nevertheless, in terms of creating a formula to deliver public
goods effectively to needy users, Seed Co and its predecessors can be cited as one
which has paid off.
83
Seed Systems
Box 6.2.
Seed sector development in Mozambique
In 1987, World Vision International initiated a programme of support to

Mozambican farmers affected by the long-running civil war. Hundreds of
thousands of rural families had been displaced by fighting. World Visions
programme focused on supplying farming households with seed of maize,
sorghum, millet, cowpea and groundnut to re-commence farming. Following
several seasons of poor results using commercial offerings from Zimbabwean and
South African suppliers, World Vision began a series of multilocation trials that
included experimental varieties from IARCs, regional seed companies, and local
landraces. Beginning in 1990, results of trials were passed on to the national
research institute and Semoc, the national seed company.
The figure below summarizes the results of 4 years of NGO-supervised
on-farm trials (as all research stations were located in war-affected areas, all
research was conducted on-farm). World Vision backed up trial information with
extensive data sets constructed from farmer interviews, observations from field
visits, and surveys. The programme developed collaborative relations with the
national university by hosting senior thesis projects and collaborated with
CIMMYT, ICRISAT and CIP in facilitating in-country testing and training sessions
focused on specific crops.
Due to the war, Semoc breeders and seedsmen were unable to travel outside
the capital city, and were therefore eager to receive information from others regarding the performance of candidate materials, especially data from farmers fields.
The programmes first breakthrough came in the form of an early maturing, openpollinated, flint maize variety (DMR-EMSRW-1) developed at IITA which carried
resistance to maize streak virus and downy mildew. The variety was renamed
Matuba (a later selection was released as Semoc-1) and multiplied by Semoc.
Between 1993 and 1997, 15,000 tonnes of Matuba and Semoc-1 were sold by
Semoc. Another early outcome was the identification of Namuesse cowpea.
Although many improved cowpea varieties were tested, none out-performed
Namuesse, a local variety with good yield, intermediate growth habit, and
84
Chapter 6
Box 6.2.
Continued
resistance to thrips. Macia sorghum is a pure line variety developed by ICRISAT

with good yield, intermediate height, and excellent palatability and preparation
characteristics. Both Namuesse and Macia have since become popular
commercial varieties for Semoc.
As vegetatively propagated crops, distribution of stock of the best-performing
sweet potato and cassava varieties presented greater challenges. The CIPdeveloped sweet potato variety, 15 Dias became highly sought after. Numerous
womens associations began multiplication and local vending of vines. The most
popular cassava variety, Fernando Po, was also a local variety. It has been multiplied in numerous field stations run by World Vision and provincial services of
the Mozambican Ministry of Agriculture. Through international seed tenders,
Semoc has gone on to supply seed of Semoc-1, Macia, and Namuesse cowpea
to Angola and Malawi. In July 1998, Semoc was purchased by Seed Coop of
Zimbabwe. Seed Coop cited its interest in gaining rights to lowland tropical
germplasm as one of its principal reasons for purchasing Semoc.
companies to eschew working on increasing their market share in non-emergency

markets. Several Ugandan agencies which have distributed seed while operating as
NGOs are converting to private, for-profit seed companies (Laker-Ojok, 2000).
Community-based seed production schemes
As seed multiplication initiatives have moved downstream and more development
groups have become involved, community-based seed production has gained increasing popularity. In these projects, improved seed and technical assistance is focused
on targeted, pilot villages in order to train farmers in seed production, storage, and
distribution. While the concept may be attractive, this model is faced with several
serious challenges, mostly related to the sustainability of such initiatives (Shumba and
Mwale, 1999; Tripp, 2000). Scaling-up from or even replicating the village level is often
very difficult. So-termed pilot schemes in a very limited number of villages often
consume a sizeable portion of the resources available for the whole country.
Seed products
Transgenic varieties
Because of the high intellectual property (IP) density of transgenic varieties, it is likely
that most (though not all) offerings of these products will be undertaken by multinational companies seeking to establish new markets in Africa. An early example is
insect-resistant Bt cotton, which has already been released in South Africa and fieldtested in Zimbabwe. Bt cotton is a likely candidate because of the presence of several
large companies that can unite the seed demands of large numbers of producers and
because the technology significantly reduces production costs in a crop where African
production is competitive.
Seed Systems
85
An alternative scenario for the distribution of transgenic varieties in Africa is

partnerships between private corporations and public sector breeding programmes,
which would also help to strengthen the capacity of NARSs to oversee and utilize new
molecular technologies. Several examples of such partnerships already exist in Africa,
with the potential for others to be initiated.
Hybrid varieties
Because of their cost an average of $40 per season for most small-scale farmers, which
may represent as much as 10% of total yearly income hybrid varieties are likely to
continue to be marketed primarily to semi-commercial farmers, who farm upwards of
5 ha or more of land. Moreover, the production and marketing of hybrid seed is not well
suited to community-based operations. The start-up costs involved in establishing seed
operations in this sector, including long lead times for registration of new varieties, seed
cleaning equipment, storage and transport costs, coupled with the small number of
clients, indicate that activity in this area is likely to depend on access to capital in the
form of loans, venture capital and grants.
The provision of such capital would significantly increase the possibility for
development of a start-up company sector in areas of currently ineffective demand
for seed. This is explored in more detail, below.
Open-pollinated varieties
Open-pollinated varieties are often considered more suitable for small-scale farmers
because they can be recycled without a loss of yield potential. For this same reason, they
have proved of little interest to private seed companies throughout the world1. Because
OPVs offer a low-risk entry point for developing seed markets among non-commercial
farmers, their distribution should not be limited to public sector agencies. Public
agencies which have developed productive, adaptated OPVs should be encouraged
to engage with private seed companies regarding test marketing of such products.
Improved OPVs could provide a stepping stone towards the use of other yieldenhancing technologies.
Seed prices
Demand for seed among small-scale farmers embodies most of the components of
demand in other markets. Quantities demanded tend to increase with decreasing prices.
Demand elasticities at both high and low price levels, however, are influenced by
consumer knowledge and product quality (Morris, 1998). Several impinging factors
prevent the comparison of pricequantity relations across Africa; however, in two
countries with similar seed industries, price does appear to influence quantities sold.
Table 6.2 shows examples of the price of maize seed in African countries.
One notable exception is the case of EMBRAPAs collaboration with private, start-up seed companies in
the cerrados region of Brazil. Based on careful tagging and continual release of new cycles of selection of
the popular maize OPV, B-201, private companies have been able to create a continuous source of
demand.
86
Chapter 6
Table 6.2.
1999.
Country
Ghana
Zimbabwe
Uganda
Kenya
Malawi
Tanzania
Nigeria
Ethiopia
Zambia
Mean maize seed prices in nine countries of sub-Saharan Africa in

OPV ($)
Hybrids ($)
1.00
0.92
1.62
0.73
0.13
3.00
0.30
1.69
1.62
1.29
0.90
1.75
0.84
1.10
Seed prices vary throughout the world, and are generally analysed by calculating
prevailing seed:grain price ratios in each local system, although comparative, absolute
prices have been listed as well (Lopez-Pereira and Filippello, 1995). Again, using maize
as an indicator, seed tends to be far cheaper in developing countries and Africa in
particular, than in industrial countries. The average price of hybrid maize seed in the
USA in 1994 was $3.72 kg1 compared with $0.75 kg1 in all developing countries
(Lopez-Pereira and Filippello, 1995). Average price of maize seed in Africa is 6.6 times
the average price of grain (CIMMYT, 1994). At these costs, farmers producing between
1.0 and 1.5 t ha1 in Africa would require yield increases of 20% to compensate for the
additional cost of seed.
The data gathered to date suggest that low effective demand for seed in Africa acts
as a hindrance to growth and diversification of the industry. Accordingly, much of the
development assistance applied to seed systems in Africa should be directed toward
reducing the high transaction costs associated with exposing farmers to improved
seed. Seed is a commodity whose quality cannot be judged prior to purchase and whose
overall performance will not be known for several months afterwards (Cromwell, 1996).
The implications for improved seed sectors include the need for extensive on-farm
testing and intensive information campaigns whenever new varieties are disseminated.
6.3 Sustainable Supply of Seed in Africa

Development of sustainable seed supply would serve three needs in Africa: (i) provide a
source of planting material to farmers who for whatever reason have been unable to
conserve seeds from previous seasons; (ii) provide a channel for the deployment of
genetic improvements made possible by conventional plant breeding; and (iii) provide a
means for the future deployment of advances made through biotechnology.
It is generally accepted that private, unrestrained seed markets offer the most
sustainable means of supply of seed to farmers. Yet in Africa, despite an estimated annual
seed market of 700,000 t, coverage by private seed companies is limited. Private seed
companies are constrained to operating in environments where they can make acceptable profits. Seed companies, at best, can expect to capture about one-third of the
increase in profits farmers obtain from using their improved varieties. While a doubling
Seed Systems
87
of yields from 1 to 2 t ha1 may represent significantly increased profits for small-scale
farmers, the opportunity to capture one-third of the profits generated by what is still
only a 1 t ha1 increase may well not be attractive to large commercial seed companies.
Acceptable profits are a function of a companys cost structure, market share, and
opportunity costs, and are difficult to analyse from outside any given firm, making
critical analysis of seed industry trends in Africa a very difficult task. Empirical
observations, however, would suggest that levels of demand and profits in much of
Africa are generally not sufficient to support extensive interest from an exclusively
industrialized seed market, and both public and private seed distribution is needed.
Seed industry development has shown an uneven pattern of development in Africa,
with hybrid maize seed sales accounting for a large portion of the private companies in
operation. Accordingly, as of 1998, West Africa, with the lowest per capita consumption
of maize in Africa, had the lowest ratio of private seed companies per country, with
approximately 0.4 companies per country, or less than one company operating per two
countries. The ratio in East Africa was 1.5:1 and in southern Africa it was 1.2:1. The
country with the highest number of private seed companies in operation was Zimbabwe,
with five.
Although the seed industry is in a phase of evolution which may bring about
changes in this regard, at present the area where large, multinational companies and
large national companies can operate profitably can be assumed to be relatively small.
Meanwhile, problems associated with their cost and issues of performance have greatly
reduced the level of influence exerted by public seed companies (Morris, 1998). Thus,
the question naturally arises as to who is available to supply seed in areas where large
companies cannot operate efficiently.
Yet the alternatives are limited. In view of the recognized limitations of private seed
companies, some authors have called into question the adequacy of the private sector
response in relation to the diversity of farmers seed needs (Cromwell, 1996). Public
companies which formerly handled seed of a wide variety of crop species are in some
cases being replaced by private enterprises interested solely in seed sales of one or a
limited number of highly profitable crops. Lack of coverage by seed companies has led to
a series of community-based seed initiatives, which promote seed production and
distribution by local farmers (see Plate 8). This work should continue, with greater
emphasis placed on lower-cost replication of models in order to reach a larger number of
farmers. Another area of future potential may be represented by small, start-up firms,
which begin by operating locally, and grow with time. Thus far, the growth of this group
has been slow, however, in part due to rigid regulatory structures, which are explored
below.
6.4 Seed Policy and Seed Regulatory Structures

As seed policies have been changed to favour private sector activity, procedures for
registration and release of new varieties have come under increased scrutiny. Tripp and
Louwaars (1997) performed a study of seed regulatory structures in Africa and
concluded that there was a need to reorganize variety registration and performance
testing within systems so that seed regulatory agencies see themselves as allies rather than
opponents of regulatory reform.
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A pilot project on Harmonization of Seed Policies and Regulations in Eastern

Africa was initiated in Kenya, Tanzania and Uganda in 1999. As a result of a series of
meetings, officials have agreed to a range of changes in seed policy and regulations
governing the release of new varieties. Chief among these is the abandonment of the
mandatory, 3 years of testing of new varieties (Anonymous, 2000). Phytosanitary restrictions imposed on ten crops were also reduced from 33 to 3. In Kenya, formal release of
new varieties until recently required 2 years of testing in Area Yield Trials followed by 2
years of testing in National Performance Trials, following which a candidate variety is
submitted for release by two separate Varietal Release Committees. Many observers
have criticized this type of system as being too slow and formalized. More recent guidelines call for two seasons of testing, followed by approval from a single committee (John
Kedera, personal communication).
As seed systems have become more competition-based, it has been necessary to
de-link research institutes from management and control of release mechanisms. In their
place, new structures have been created, often under the auspices of national plant health
inspection services. It is assumed that these structures will be able to deal with seed companies, NARSs, and other purveyors of new varieties on a more equitable basis.
Varietal release structures are key to efficient flow of improved germplasm to
farmers. Assisting these groups in putting together transparent, cost-effective, selfsupporting mechanisms for evaluation and release of new varieties should be viewed as a
potential area of support by donor agencies interested in seed issues. Decreasing the level
of formality will undoubtedly be a key to the success of such reforms, and will probably
grow out of participatory breeding methods, which by their nature break down the
barriers between farmers and the final, varietal offerings, and in some cases put
significant quantities of improved seed into circulation prior to official release by the
government (Tripp and Rohrbach, 2000).
Seed certification is another key area of concern to those whose aim is reform of the
seed sector in Africa (Venkatesan, 1994). Sales of seed to commercial farmers may need
to be subject to relatively strict certification procedures in order to protect consumer
confidence and provide for legal recourse in cases of large transactions. However, formal
seed certification and labelling is viewed as a major constraint to seed market development among non-commercial farmers. In fact, what is sought is an acceptable means of
assessment of candidate varieties that places primary emphasis on the responses of
farmers, and allows the free flow of improved germplasm toward small-scale farmers
through a variety of channels. Achieving this kind of seed regulatory environment
among non-commercial farmers, however, will require the involvement of those who
determine the process which pertains to formal sale of seed to commercial farmers. As
deregulation proceeds, it should be borne in mind that in several active seed sectors, for
example in the USA and India, formalized release of varieties is done on a voluntary
basis, with informal industry standards being relied upon to provide quality control.
Analysis of the constraints to the development of sustainable seed supply among
small-scale farmers in Africa indicates four broad areas which provide points of entry for
a range of actors from both public and private sectors: breeding, farmer access, seed
markets, and regulation.
Seed Systems
89
Breeding
As discussed elsewhere in this report, varietal performance is inevitably the principal
factor in creating sustainable, effective demand for seed of improved varieties.
Participatory variety selection provides a new framework for identifying varieties farmers
value most and provides guidance to breeders in further crop improvement efforts.
In seed sectors which have recently undergone liberalization, the rate of development and release of new varieties may limit the progress new seed entities can make in
establishing a client base. Crop improvement, whether through conventional breeding
or molecular breeding, remains by and large a long-term, expensive undertaking. This
clearly limits the number of seed companies that are likely to establish their own breeding programmes in Africa over the medium term. The programme size and budgets of
public breeding sectors of African countries also limit the speed with which they can
develop new offerings. Both public and private breeding efforts continue to be limited
by availability of qualified personnel.
Farmer access
Overall availability of seed, either through private stockists or publicly based dissemination programmes, continues to limit both the development of viable seed markets and
the overall impact of crop improvement. Private dealers must strive to establish a
maximum number of distribution points in the major farming areas. Likewise, public
breeding efforts need to assist in building farmer awareness of the advantages of new
products through multilocation testing initiatives.
While it has been amply demonstrated that African farmers are willing to pay for
seed when it is of clear benefit to them, seed prices will continue to be a factor in levels of
demand for some time to come. Farm gate (producer) prices will play an important role
in determining what is an affordable seed price. Recent experience seems to indicate that
demand for hybrid seed of grain crops will fall off when seed prices exceed four to five
times the price of grain. Seed:grain price ratios for open-pollinated crops may face a
limit of only 2 or 3:1.
Packaging of seed in small quantities has emerged as an important means of
reaching small-scale farmers. Packages of 25 kg are common. In some cases, for some
grain crops, packages of 1 kg or even less may be appropriate. A recent project aimed at
testing the demand for seed of non-traditional seed products (open-pollinated sorghum,
millet and groundnut) in Zimbabwe found that, contrary to previous belief, farmers
will purchase seed of these crops when it is offered in accessible, sensible ways. They
found that 500 g packages were the most popular among small-scale farmers (Rohrbach
and Malusalila, 2000). Seed suppliers in western Kenya have proposed that gearing
packaging to cash amounts that farmers can reasonably part with (from $0.75 to 1.50) at
any given time may be a means of increasing sales. Packaging of seed in transparent,
plastic bags has also been cited as a means of assuring farmers they are buying a product
of acceptable quality.
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Seed markets
Related to the issue of building farmer awareness, seed market development requires
ample, accurate information on the characteristics of new varieties. Farmers are very
reluctant to pay for new varieties they have never seen before. Participatory multilocation testing and demonstration projects are required to overcome this barrier to
adoption. Moreover, it is almost certain they will not purchase varieties if the seller
cannot provide them with clear information regarding their growth characteristics,
including those which may represent an improvement on what is currently being grown.
Profit sharing among different entities within the seed distribution chain may be
important in establishing seed markets as well. In at least one case, recommendations
have been made for margins to be highest where volume of sales is lowest, to encourage
more vigorous and extensive distribution (Mark Wood, personal communication).
Moreover, because seed sales are seasonal and unpredictable in the early stages of
introducing new varieties, providing stockists with credit guarantees may be an effective
means of building seed markets.
As seed has become an increasingly popular focus of development agencies, free
seed distribution has become a factor in the establishment of private seed markets. While
emergency, (free) seed distribution following a major rupture in crop production cycles
is certainly an effective means of assisting in recovery, it seems clear that free
seed distribution needs to be managed in a way that does not stifle demand among
farmers who would otherwise purchase seed (Tripp and Rohrbach, 2000). Generally
speaking, governmental coordination of seed distribution activities should provide the
means of preventing seed distribution from constraining the establishment of viable
seed markets.
Regulation
Clearly, seed sector liberalization should not be synonymous with allowing seed sectors
to operate in a free-for-all. Quality issues are extremely important in providing farmers
with a fair return for their seed purchases; however, overall product quality can only be
judged several months after the purchase. Seed sector liberalization in several African
countries has been accompanied with increasing cases of seed fraud, where unscrupulous
seed dealers have taken advantage of demand for seed by packaging ordinary grain as
seed and selling it to farmers as certified seed.
As community-based seed production activities have increased, seed certification
has become an increasingly important issue for seed regulators. Following liberalization,
seed regulatory units in Africa have experienced serious difficulties in responding to the
demands for inspection of increasing numbers of formal-sector seed growers and the
many far-flung, farmer-grown seed initiatives. This may involve new categories of seed
certification and new categories of seed certifiers, for example, trained NGO employees
or government extension agents.
Overly bureaucratic variety release procedures have previously been cited as a constraint to seed sector development in Africa. While countries in other regions of the
world have eliminated formal variety release requirements or instituted voluntary release
structures, there appears to be little support for such an approach in Africa, in the short
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Seed Systems
term. Recently, seed regulatory bodies have been pressured to accept as little as a single
seasons data (presumably from more than a single site), which may be submitted
following tests conducted by the varietys developer or other recognized agency,
provided that recognized, accepted commercial check varieties have been included
for comparison.
6.5 African Seed Systems Challenges Summary

The public sector challenge
Table 6.3 summarizes the priority areas of focus for strategies aimed at more sustainable
seed supply among small-scale farmers in Africa.
For many self-pollinated crops rice, beans, cowpea and sorghum dissemination
of improved varieties in Africa will continue to rely at least in part on public sector-based
programmes aimed at targeted groups of farmers and involving NARSs, NGOs,
community-based organizations and, to some extent, small, private companies.
Likewise, distribution of clonally propagated crops (sweet potato, yam, banana and
cassava) which target needs of small-scale farmers will be best approached via the public
sector. Participatory methods of research, including dissemination of small quantities of
seed directly to farmers for testing from research teams, goes some way in circumventing
Table 6.3.
Principal activities related to diffusion of improved varieties.
Activity
Description
On-farm testing
Multilocation testing under farmer conditions serves

to verify that the candidate varieties satisfy adaptation and farmer varietal preferences, and serves to
inform farmers of the pending availability on a wider
scale of a new variety
Crop improvement networks that carry out extensive
on-farm testing are in a position to involve farmers in
the selection of varieties for release and
multiplication
Seed multiplication of both foundation (by national
breeding programmes or private sector) and certified
seed (by NGOs, contract farmers, community-based
organizations and private sector)
National programmes are increasingly entering into
agreements with private companies for licensing of
new releases. Marketing/dissemination by public
agencies is often carried out via distribution
campaigns
Participatory selection
process
Multiplication
Marketing/dissemination
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Chapter 6
the laborious, official release process. Numerous studies have shown that farmers will
share seed with family members and acquaintances, and that some dissemination can be
achieved in this context, although other studies have shown that seed sharing is not automatic, and may be quite limited (David and Sperling, 1999). In these cases, the involvement of researchers or extension agents in the establishment of informal agreements with
farmers may speed dissemination and widen the area of impact (B.B. Singh, personal
communication).
The aim of such initiatives would not be comprehensive distribution of seed to
every farmer in a given area, but achieving sufficient rates of diffusion to ensure that
valuable offerings can move via farmer-to-farmer exchange. Participatory breeding
methods which aim at exposing farmers to crop varieties while they are still under development should encourage seed dissemination initiatives of this type, and may force
major changes in the varietal release process, at least as far as it concerns non-commercial
farmers.
Needless to say, the above-mentioned process is dependent upon public sector
funding provided by national governments and donor agencies. Clearly, in cases where a
useful public good (a new variety) can be effectively distributed to needy consumers via a
public-sector, campaign-type distribution programme, participants should not hesitate
to do so. The underdeveloped state of the economy in Africa, coupled with the
continued occurrence of hunger, is ample evidence that the private sector cannot be
relied upon to fill all gaps. Thus, one inescapable reality is that crop improvement aimed
at benefiting the majority of farmers in Africa will require continued support from the
public sector. Over the long term, however, the establishment of a viable and sustainable
private sector seed exchange needs to be encouraged.
How this is likely to develop in Africa is difficult to predict, in part because the
actors are highly differentiated by make-up, size, and access to emerging genetic technologies. Because the market is segmented, products of rather different origin are likely
to be a key factor, and a diversity of seed distribution entities is indicated.
The private sector challenge

Small, start-up seed companies which focus on niche markets for new varieties and are
able to operate on the basis of smaller markets may offer the best chance of sustainable
seed supply in marginal areas and for crops of less commercial importance. Finding
innovative ways of easing their start-up and entry into operation could ultimately render
important benefits to farmers. The general scarcity of venture capital in Africa almost
always ensures that there are simpler, more immediate opportunities for return on
investment available to local entrepreneurs than the development of a seed company.
Providing capital, in-kind support, technical assistance, and other forms of encouragement is likely to be necessary for the creation of the type of seed industry capable of
responding to small-scale farmers needs over the longer term. There is a growing list of
positive experiences related to supporting small-scale seed commerce in Africa (SCODP,
1999; Laker-Ojok, 2000; Muhhuku, 2000; Rohrbach and Malusalila, 2000).
In addition, support to public sector breeding programmes which take on the
expensive, long-term task of developing new varieties will encourage the operations of
small, private companies by offering varieties which respond to the needs of small-scale
Seed Systems
93
farmers in less favourable growing environments. NARSs in Africa continue to embody

resources which can be used to develop varieties that can be licensed to seed companies.
Finally, it must be recognized that the vast, informal seed market and system of
exchange in Africa is poorly understood. Additional study is required to understand
more fully how to make use of this resource in improving access by poor farmers to
better genetic resources.
Conclusions
The first six chapters of this book have tried to call attention to a set of urgent problems
faced by farmers trying to secure harvests on hills and savannah plains and in highlands
and valleys across Africa. It has tried to identify and describe briefly scientific resources
and methods for using them, some of which the authors believe are likely to produce
useful, sustainable results.
Ultimately, however, achieving food security among Africas rapidly growing
population of rural poor is not a scientific or even an economic goal, but a humanitarian
one. In the end, it is not science that will prevail, but African farmers exerting their will
to succeed and conquer hunger, farm by farm, harvest by harvest. Therefore, perhaps the
most important observation regarding African agriculture is that African farmers do not
wish to be left behind the rest of the world in achieving food security.
Following an analysis of the resources and approaches now available for developing
more productive and more resilient varieties of food crops in Africa, several broad
conclusions and recommendations can be made.
Farmers in Africa cultivate a wide range of crops adapted to a diverse physical

environment. Low initial yield potential on these farms is further reduced by losses
of yield to both routine and intractable production constraints, contributing to
widespread food insecurity and reducing opportunities for economic growth.
Crop-specific assessments of problems encountered by small-scale farmers reveal a
wide range of traits where genetic improvement could significantly reduce losses
among farmers unable to purchase inputs or labour needed to combat these threats
by other means. Higher productive potential available through modern plant types
could make additional contributions. Many of these aims can be realized through
effective plant breeding programmes conducted by national programmes.
Because improved varieties in Africa must fit into highly varied, marginal farming
conditions, and because farmers consume a significant portion of their harvests,
local adaptation and farmer varietal preferences play major roles in determining
the overall adoption rate of improved varieties. The added complexity of breeding
95
95
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96
Chapter 7
for these needs presents a strong case for applying a more participatory, agroecology-based approach to crop improvement which takes account of the particular
mix of constraints to production that exist in any given zone. Engaging farmers in
the decision-making processes related to varietal development makes good sense
and may increase adoption rates.
Development of national breeding programme strategies involving identification
of agro-ecologies, relevant breeding priorities, optimal source germplasm, more efficient use of human resources, and identification of on-going needs for capacity
building will be critical to making any investments in plant breeding and biotechnology pay off. International agricultural research centres can assist national
teams in developing effective strategies for crop improvement and provide key
backstopping roles in the implementation of these strategies.
The existence of both routine and intractable biological constraints to crop
production among small-scale farmers argues for strong roles for both conventional
breeding and biotechnology and for collaboration between national and international research groups. There has been a significant increase in the numbers of
qualified plant scientists within Africas national agricultural research system, and
they are ready to assume an increased role in plant breeding at the national and
regional levels. However, strengthening Africas biotechnology research capacity
will require further investment and assistance from international agricultural
research centres and advanced research institutes.
In the case of crops for which routine methods for application of molecular breeding or genetic transformation have not yet been developed, there are obvious needs
for increased efforts on development of basic biotechnology research tools. In most
cases, this will require the capabilities and commitment of advanced laboratories.
Moving the products of crop improvement beyond the laboratory or field plot and
into farmers hands is a complex undertaking. On the one hand, deregulation of
seed industries in Africa means better prospects for getting new products to farmers
via a more active private and NGO seed sector. However, decreased support to public institutions broadly speaking means they will be in a weaker position to fill gaps
left by private sector activity at the outset. An as-yet unknown factor in this regard is
the likelihood of growth within the small-scale or local seed business sector, and
whether public assistance can help in getting this sector moving. Increased support
for both types of undertaking will be necessary to make investments in crop genetic
improvement pay off.
In order to fulfil their stated aims of improving food security in Africa, crop genetic
improvement initiatives must be tied directly to seed dissemination schemes of one
kind or another. Failure to appreciate the complexity and costs associated with
broad dissemination of improved varieties has left many valuable products on the
shelf and left many farmers without alternatives for improving their production
systems.
Finally, the powerful combination of biotechnology, agro-ecological approaches to
research, and participatory methods of plant breeding represent a new era for crop
genetic improvement in Africa. This opportunity to secure Africas harvest should
not pass without receiving the effort and commitment it merits.
96
Chapter 7
Maize
8.1 Brief History of Maize Cultivation and Utilization in Africa

Maize was introduced into Africa by the Portuguese at the beginning of the 16th century
(Reader, 1998). It has since become Africas second most important food crop, behind
cassava, and is grown in a wide range of environments ranging from Nigers northern
Sahel to Ethiopias highlands to converted forest lands of Sierra Leone.
The popularity of maize among African farmers grew slowly until the early part of
the 20th century. During World War I, the colonial government in Kenya encouraged
(and provided seed for) farmers to plant maize as part of the war effort. Maize
cultivation in southern Africa was initially linked to the spread of commercial mining, as
maize required less labour to grow and process than the traditional grain crops, millet
and sorghum (Byerlee and Heisey, 1997).
Although its palatability is often cited as a reason for maizes continued popularity
among rural populations of eastern and southern Africa, higher productivity and
lower labour demands can probably be assumed to be at least as important. Sorghum
and millet yields in eastern and southern Africa since 1980 have averaged 765 and
729 kg ha1, respectively, compared with yields of maize over the same period of
1.19 t ha1 (FAO, 1998). While a portion of these differences can be attributed to the
different environments in which the crops are grown, even when grown under identical
conditions in semi-arid southern Africa, maize was shown to yield higher (Waddington
and Karigwindi, 1995). Comparatively low labour requirements appear to be a second
factor in the popularity of maize among small-scale farmers in Africa. Increased school
attendance among children who formerly performed bird-scaring chores is cited as an
important factor in the shift of land out of sorghum and millet toward maize in
semi-arid regions of southern Africa (Rohrbach, 1994).
Per capita consumption of maize in Africa is highest in eastern and southern Africa.
Maize consumption in Kenya, Tanzania, Malawi, Zimbabwe, Zambia and Swaziland
averages over 100 kg per year (CIMMYT, 1990), giving maize a similar position in
terms of dietary importance in those countries to rice in Asia.
99
99
Chapter 8
100
Chapter 8
Maize is mainly consumed in African households as a thick porridge, produced by

hand pounding (usually preceded by soaking) or grinding in a hammer mill, followed by
boiling. Households that depend on hand pounding generally prefer harder, flint-type
varieties whose endosperm and embryo can be milled as an integral, whole grain
(Nhlane, 1990; Smale et al., 1994). Households that mill their grain generally prefer
dent varieties. Eastern and southern Africa almost exclusively grow white maize, with
small pockets of yellow landraces in coastal regions and southern Sudan. West and
central African households use both white and yellow maize. Seasonally throughout
Africa, a considerable amount of maize is consumed fresh, both on and off the cob,
either roasted or boiled, as a snack food.
Ninety-five per cent of maize produced in Africa is grown by small- and mediumscale farmers who cultivate 10 ha or less. Yields on these farms are usually low, averaging
1.2 t ha1 (Byerlee and Heisey, 1997). Meanwhile, the productivity range of maize
farmers in Africa is perhaps wider than for any other crop. While subsistence farmers of
coastal West Africa struggle to produce 700800 kg ha1 on farms as small as half of a
hectare, large scale, commercial farmers of Zimbabwe harvest some of the highest cereal
crop yields in the world, regularly topping 10 t ha1 on farms larger than 1000 ha.
8.2 Maize Production Levels and Trends in Africa

Maize production trends in sub-Saharan Africa and its subregions are shown in Fig. 8.1.
The graph depicts eastern and southern Africa as the dominant maize-growing regions
of Africa until approximately 1985. Beginning from a relatively small production base in
the early 1980s, maize production in West Africa rose above eastern and southern Africa
by the early 1990s. This was fuelled by phenomenal increases in maize cultivation in
Africas most populous country, Nigeria, facilitated in large part by the development at
Fig. 8.1.
Maize production trends in Africa, 19781999. Source: FAO (2000).
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101
Maize
IITA of earlier-maturing, disease-resistant maize varieties which could be grown in the

favourable Sudan savannah ecologies (IITA, 1995).
Following generally good rates of production increase during the 1980s, rates of
growth of maize production throughout Africa appear to have declined during the
1990s (see Table 8.1). This was influenced by several drought years in southern Africa
and more generally by increases in the price of fertilizer following removal of subsidies
(Blackie, 1994); in some areas this caused farmers to revert to crops such as cassava
and sorghum, which generally grow better on poor soils. The trend in east Africa is
particularly worrying. Although the population has grown by 20% during the period
1989 to 1998, annual maize harvests for the subregion have actually declined.
Per capita annual maize production in sub-Saharan Africa since 1975 as a whole has
varied between 25 and 40 kg (data not shown). Increasing production in West Africa led
to rapid increases during 1985 to 1989, but poor harvests during the early 1990s
reduced levels considerably. Per capita annual production since 1995 appears to have
stabilized at roughly 35 kg. Over the same period, sorghum production per capita has
ranged between 20 and 28 kg per person per year.
8.3 Maize Production Constraints

Higher maize yields in Africa relative to sorghum and millet are aided in part by farmers
tendencies to cultivate maize on well-watered and more fertile land. Compared with
traditional crops, however, maize is relatively susceptible to moisture and nutrient stress.
Drought and low soil fertility are ubiquitous production constraints on small-scale
farmers fields in Africa (Edmeades et al., 1994). Waddington et al. (1994) estimated
average annual losses of maize production due to moisture stress in eastern and southern
Africa of 13% of total production, or 1.8 million tonnes per year.
Recent estimates of yield reductions due to environmental stresses such as drought
(see Plate 9) and low soil fertility have been aided by satellite imaging and geographical
information system modelling (Hodson et al., 1999). Modelling losses due to biotic
stresses is more difficult. Outbreaks of important pests and diseases of maize are related
to complex egg-laying and sporulation responses which are in turn dependent upon
difficult-to-predict global environmental factors (moisture and temperature regimes)
and human activity (management of crop residues, crop rotations, intercropping) (see
Plate 10). Nevertheless, using estimates of incidence and average yield losses per plant,
estimates of production losses can be made.
Table 8.1. Rate of growth (%) of maize production in Africa and subregions,
19701997.
Region
West Africa
East Africa
Southern Africa
Africa
19701979
19801989
19901997
2.2
5.9
4.1
2.4
15.4
0.0
7.2
7.3
2.3
1.6
3.9
0.5
Source: FAO (1998).
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Chapter 8
Priority biotic constraints of maize in Africa include insect pests (stem borers, grain
borers, and weevils), foliar diseases, ear rots and the parasitic weed, Striga. The ranking
of these pests in terms of economic losses varies significantly across agro-ecosystems.
Table 8.2 shows the estimated area affected by the main pests and diseases of maize in
Africa.
At an estimated 60% of total area affected, maize streak virus (MSV) ranks as the
most widespread biotic constraint to maize production in Africa. However, attempting
to estimate production losses due to MSV provides an example of how difficult such a
task can be, even on a single plant basis. Ampong-Nyarko et al. (1998) found that grain
yield loss in maize due to MSV attack was related to the growth stage at which attack
occurred. Plants attacked at early stages of growth (up to seven-leaf stage) suffered 80%
or higher loss of yield, while plants attacked shortly thereafter (at the nine-leaf stage)
suffered only 20% yield loss.
Table 8.2. Distribution of principal maize production constraints across agroecologies of Africa.
Lowland tropical Sub-tropical
Drought
Striga
Leaf blight
E. turcicum
H. maydis
Rust
P. sorghi
P. polysora
Maize streak virus
Stalk rot
Not specified
Fusarium
Diplodia
Ear rot
Not specified
Fusarium
Diplodia
Stem borers
Not specified
Chilo
Buseola
Pink stem borer
Storage pests
Weevils
Larger grain borer
Termites
Highland
% Area
36
30
21
20
0
1
21
56
35
1
100
0
40
28
2
26
73
42
3
37
58
0
7
28
23
60
37
9
0
1
0
0
0
0
16
18
5
2
29
28
15
25
10
10
0
19
36
33
20
20
35
19
7
15
7
8
69
22
19
0
76
0
33
10
37
33
20
41
38
20
12
15
19
23
21a
Source: adapted from CIMMYT (1988).

aEstimates based on observations in ten maize-producing countries of sub-Saharan
Africa.
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Jeffers and Chapman (1994) found that the most important diseases of maize across
the mid-altitude, transition, and highland zones of eastern and southern Africa are
turcicum leaf blight (Exserohilum turcicum) and common rust (Puccinia sorghi). Average
reduction in yield due to foliar and stem diseases across three ecological zones in three
seasons in Cameroon averaged 1012 g per plant. At a conservative estimate of 25,000
plants ha1 on farmers fields, this would represent a loss of approximately 330 kg ha1
(Cardwell et al., 1997). Applying these losses across 11.7 million ha of maize production
in humid lowland and mid-altitude environments in Africa (CIMMYT, 1990) translates to losses of 3.8 million tonnes per annum. Similar methods can be applied to other
important pests and diseases of maize in Africa as shown in Table 8.3. While such estimates are prone to high levels of error, if accurate, the calculations would indicate that
upwards of 75% of Africas maize harvest is lost to pests and diseases prior to harvest.
In 1989, subsequent to the CIMMYT study cited above, downy mildew disease of
maize (Peronosclerospora sorghi) reached epidemic proportions in parts of Nigeria and
began to spread. Downy mildew continues to be a serious disease of maize in seven states
of West Africa, the Democratic Republic of Congo, and Mozambique. Recent studies
have shown that oospores of downy mildew can transmit the disease through transport
of market grain and crop debris (Adenle et al., 1998). Downy mildew-resistant varieties
(DMRESR-W and DMRESR-Y) have been developed at IITA.
Even more recently, maize yields in eastern and southern Africa have been reduced
by infection from grey leaf spot (Cercospora zea-maydis), which was first reported in
Uganda in 1994 (George Bigirwa, personal communication) and has subsequently
become an important disease of maize throughout most of eastern and southern Africa
(Pixley, 1997). Grey leaf spot disease increased dramatically between 1991 and 1997 in
Malawi (Ngwira, 1998).
Postharvest insect pests, maize weevil (Sitopholus zeamais) and larger grain borer
(Prostephanus truncatus), account for serious losses of harvest in household maize storage
facilities in large areas of Africa. Host plant resistance for both storage pests has been
documented in a range of genotypes (Meikle and Markham, 1998) and especially in the
Tanzanian landrace Kilima (Derera et al., 2000); however, both traits are multigenic in
Table 8.3.
Estimates of production losses due to pests and diseases in Africa.
Pest/disease
Striga
Blights
Rusts
MSV
Stem borers
Total
Area affected
(million ha)
Estimated yield loss

(%)
4.33
14.0
10.5
12.36
16.48
40a
20b
35c
37d
20e
Estimate of total annual

loss of production
(million t)
2.07
3.36
4.41
5.48
3.9
19.22
aAuthors
estimate.
and Chapman (1994).
cKim and Brewbaker (1977).
dAmpong-Nyarko et al. (1998).
eBosque-Perez and Mareck (1991).
bJeffers
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nature and represent difficult challenges for breeding. Molecular methods may prove
useful in screening for resistance for these traits, provided markers can be identified for
their quantitative trait loci (QTL).
8.4 Maize Improvement Through Breeding and Biotechnology

Maize breeding advances in Africa
Maize is one of the most highly bred crops in the world. Maize improvement up to the
early 1900s was limited to recurrent selection methods. Population improvement
through a series of recurrent selection procedures is aimed at maximizing percentages of
favourable alleles at each locus of importance to crop performance in a given environment. Population improvement remains the primary means of improving levels of
performance in base populations from which inbred lines are developed.
In 1908 George Shull, a researcher at Cold Spring Harbor Laboratory in New York,
published a paper which described the basic techniques by which vigour in an experimental variety of maize had been significantly increased through hybridization. This
marked the beginning of the use of heterosis in plant breeding. In 1924 the first sale of
hybrid maize seed occurred. By 1939, over 90% of maize grown in the state of Iowa was
of hybrid varieties (Crow, 1998). These original hybrids sparked a yield gain of approximately 15%, or 300 kg ha1 (Griliches, 1957). Subsequent breeding, focused more on
local adaptation factors, has added approximately 50 kg ha1 year1 (Duvick, 1992).
By contrast, 60 years later in Africa, only 20% of the maize area planted is of hybrid
seed (Morris, 1998). Many African countries where maize is produced still do not have
commercial hybrid varieties. An estimated 63% of maize grown in Africa is of
unimproved, or landrace, varieties (Morris, 1998) (see Plate 11).
Undoubtedly, a major factor in the low usage of hybrid maize in Africa is related to
poorly developed seed industries, which are in turn reflective of relatively poorly
developed economies as a whole. In the absence of seed sectors capable of organizing
large-scale production of hybrid seed, breeders at both international and national levels
have concentrated primarily on population improvement, through recurrent selection
methods of breeding. Although population improvement has proved effective in the
development of open-pollinated varieties (OPVs) which can be cheaply produced and
released, these products have seldom been taken up by private companies (Heisey et al.,
1998), and their distribution through other means has been constrained by a lack of
public sector capacity and funds to promote them.
In response to investments in breeding and seed industry development, farmers in
Kenya and Zimbabwe were early and enthusiastic adopters of hybrid maize (Gerhart,
1975; Rattray, 1969). Gerhart (1975) calculated that rates of adoption of hybrid maize
varieties in western Kenya during the 1960s and 1970s were higher than those of US
farmers during the 1930s and 1940s. Adoption rates among farmers in Zimbabwe were
no less impressive. In 1960, breeders in Zimbabwe released the single-cross hybrid
SR52, the first commercial use of a single-cross hybrid in the world. SR52 was
reported to increase maize yields among its users by an average of 46% (Weinmann,
1975). Within 8 years, SR52 was in use in over two-thirds of the maize area planted by
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commercial farmers (Rattray, 1969). Today in Zimbabwe, seed companies market over
40 different hybrid maize varieties (Zambezi, 1997).
Although hybrids have had the advantage of being promoted by private companies,
recurrent selection methods of breeding have also made important contributions
to maize productivity in Africa. Although the yield components responsible for the
advantages of improved, open-pollinated maize in Africa can be difficult to identify,
increased harvest index, increased vigour, and resistance to foliar diseases all appear to
have made contributions. It is important to note that even in countries with relatively
low adoption rates of identified, improved varieties, the effects of periodic seed
purchases by farmers, free distribution of improved varieties, chance out-crossing
of landraces with neighbouring plots of improved maize, and farmer selection of
seed have inevitably resulted in a broad-based rise in maize yield potential in Africa.
On-station yield differences between improved, open-pollinated maize and local
varieties are remarkable. In trials of 34 early-maturing OPVs grown at 18 sites in southern Africa in 1996, improved varieties averaged 31% higher yield than local varieties
(Pixley, 1996). The improved commercial OPV Manica evaluated in 30 on-farm trials
without fertilizer or other inputs in Mozambique in 1994 yielded 24% higher than the
local (unimproved) variety. A three-way hybrid evaluated in the same study yielded 13%
and 34% higher than Manica and the local variety, respectively (White and Sitch,
1994). Other countries where OPVs have been widely cultivated include Nigeria, where
TZ-B and TZPB achieved high rates of adoption in the late 1970s, and Ghana, where
adoption of the high-lyseine variety, Obatanpa, and other improved maize varieties,
averages 54% (Tripp and Louwaars, 1997). These results point to the kind of broadbased improvements in food security that can be achieved through conventional plant
breeding programmes in Africa.
Advances in maize biotechnology

Molecular genetics
Applications of molecular genetics in maize have progressed rapidly throughout the
1990s. Molecular marker maps have been generated for maize using RFLPs, RAPDs,
AFLPs and microsatellites. Molecular mapping techniques have been applied to a variety
of traits of importance in maize including turcicum blight resistance (Schechert, 1997),
anthesis-silking interval (Ribaut et al., 1996), grey leaf spot disease resistance (Bubeck
et al., 1993) (see Plate 12) and resistance to common rust (Coe et al., 1988).
Studies of the efficiency of marker-assisted selection (MAS) techniques compared
with phenotypic evaluations have been conducted for a variety of traits in maize
(Eathington et al., 1997; Lubberstedt et al., 1998). Quantitative trait loci (QTLs)
identified using RFLP markers produced correlation values with grain yield of 0.86
compared with phenotypic correlation values of 0.36 (Eathington et al., 1997), implying higher gains from selection for markers correlated with field performance than from
selections based on field testing alone. The ability to select the best lines was significantly
improved through the combined use of phenotypic and marker information in both
high-yielding and low-yielding environments, but the usefulness of markers was higher
in the high-yielding environments. However, analysis of four independent maize
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populations for six traits using QTLs identified by RFLP markers showed inconsistent
results. The authors concluded that employing MAS in this case would necessitate
separate QTL mapping for each population (Lubberstedt et al. 1998).
Results from CIMMYTs Applied Biotechnology Center laboratory suggest that the
use of genetic markers for aid in selection schemes is so far applicable in the more simply
inherited traits where QTLs can be identified that described 40% or more of phenotypic
variation. The possibility of using markers in such cases opens up genuine opportunities
for incorporation of multiple resistance traits in a single selection programme, whereby
field-based evaluations for adaptation can be augmented by differing marker analyses for
difficult-to-screen resistance traits.
Studies with QTLs for complex traits where markers describe less than 40% of
phenotypic variation encountered difficulties with respect to consistency of marker
identification (Dave Hoisington, personal communication).
Until 2000, the biotechnology centre at IITA was the sole laboratory in subSaharan Africa (excluding South Africa) engaged in marker identification and markerassisted selection in maize. During 2000, new national molecular laboratories were
launched in Kenya and Zimbabwe, focusing on marker selection for drought tolerance
and resistance to stem borers. MAS schemes which involve collaborations between
laboratories sited outside of Africa and Africa-based testing facilities are on-going in
Kenya, Zimbabwe and Malawi.
Genetic engineering
The first maize plant was regenerated from plant cells in the 1970s (Phillips et al., 1988).
Since then, a variety of regeneration techniques have been developed using callus tissue,
cell suspensions, excised plant parts, and immature embryos. Immature embryos have
proved the most useful tissue in maize transformation techniques (Hoisington et al.,
1998). The first commercially released transgenic maize variety was developed using the
gene gun (Gordon-Kamm et al., 1990). Nevertheless, transformation efficiency to date
using the particle gun remains relatively low, at around 1% (Hoisington et al., 1998).
Agrobacterium-mediated transfer of DNA into maize was first reported in 1996 (Ishida
et al., 1996). Since then, Agrobacterium has been used with increasing frequency due to
its ability to transfer larger DNA segments in lower copy numbers (Hoisington et al.,
1998).
Although transformation of maize for a variety of traits has become routine in
many laboratories, no group has as yet reported genotype-independent transformation
methods, an important benchmark in genetic engineering of specific crop species (Paul
Christou, personal communication). As such, applied genetic engineering work on
maize is often based on initial transformation of one of several genotypes of known
transformability, following which the trait is back-crossed into target germplasm.
Transgenic maize accounted for 11.1 million ha of production in 1999, placing it
second behind soybean (21.6 million ha) in area planted to transgenic crops (James,
2000). All commercial transgenic maize plantings to 1999 carried either insect resistance
(Bt proteins) or herbicide tolerance traits. All transgenic maize in commercial use to date
has been marketed by private companies, however an initiative begun in 2000 between
KARI, CIMMYT and the Novartis Foundation aims to develop insect-resistant maize
for Kenya using Bt genes.
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8.5 Principal Challenges for Maize Improvement in Africa

Investments made in building the capacity of national maize breeding programmes
during the 1980s and 1990s mean that practical strategies for improvement can now
move forward relatively rapidly. However, as maize remains both a commercial crop and
an essential component of household food security, it is essential that all initiatives
begin and end with farmers. Thus, as with other cereal crops, careful attention must
be paid to issues of farmer preferences and utilization. Effective seed dissemination will
be key to achieving high levels of impact from investments made in breeding and
biotechnology.
Faltering maize production and productivity levels in all sub-regions of Africa are
matters of concern. Maize is the dominant supplier of carbohydrate in the diets of most
African households in eastern and southern Africa. However, the very rapid rate of
growth in maize production in West Africa during the 1970s and 1980s is evidence of its
potential to improve food security and opportunities for rural economic growth in that
part of Africa as well.
The principal weaknesses of maize in the African context are its susceptibility to
abiotic stress, especially drought, and a number of damaging pests and diseases. In view
of continued low applications of fertilizer, irrigation, and other inputs, it is likely that
pressures from these constraints will increase during the coming decade. Thus, strategies
which focus on increasing the yield stability of maize through better combinations of
resistance and tolerance traits would appear to be the most appropriate strategy for most
small-scale maize farming systems.
Continual maize plantings in bimodal production systems of eastern Africa are
likely to increase pressures from pests and diseases. The outbreak of grey leaf spot during
the 1998 season has been identified as a major cause of food shortages in Tanzania.
Losses due to maize streak virus and turcicum leaf blight in Kenya during the same
season were especially high. The spread and increased level of infestation by Striga in
areas of low soil fertility has been well documented in both Kenya and Malawi (Frost,
1995; Ngwira et al., 1998). These events may indicate a trend which could intensify in
coming years.
In view of the potential for increased pressure from pests and diseases and drought,
an expanded emphasis on these constraints through a progressive, iterative, and
participatory process is recommended. Guiding principles for such an approach would
include:
Reinforcing the product development strategies and overall capacity of key

national breeding programmes so that a pipeline of resilient, public sector
varieties (both open-pollinated and hybrid) is developed in each sub-region of
Africa.
Promoting closer, fuller partnerships between national programmes and the
Africa-based IARCs involved in maize improvement so that more effective,
product-oriented breeding strategies can be implemented at national level.
Identification of important, intractable pests and diseases and abiotic stresses for
intensified upstream research involving biotechnology and breeding.
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8.6 Maize Seed Systems

Maize forms the primary basis of most commercial seed markets in Africa. While some
commerce has been developed around open-pollinated varieties, over the past decade
there has been a clear move toward hybrid varieties, especially in eastern and southern
Africa. The results of this study have shown that considerations of the cost this move
implies to farmers should be balanced with the urgent need for development of
sustainable seed systems in a competitive, privatized market. In most cases, the benefits
carried to farmers through healthy, adapted, vigorous, hybrid seed do cover the price
paid. Nevertheless, extensive on-farm testing and demonstration is required to persuade
farmers of the benefit of planting improved, purchased seed.
Ensuring African maize seed systems are fully competitive will, however, require
input from donor agencies. The deregulation of seed sectors in Africa combined with
Africas diverse agro-ecologies create opportunities for new private operators in Africa.
Small, private seed companies of African origin could fill a niche in Africas seed markets
by helping to commercialize a larger number of varieties with better adaptation
to specific agro-ecologies. Small companies of this kind can work with national
programmes through licensing agreements that bring in royalty payments to help defray
the cost of public breeding programmes. However, this process may require start-up
technical and financial support of several kinds.
A programme focused on resilient crops will inevitably target some areas where
margins cannot support private supply of seed. In these areas, public seed production
will need to be developed for deployment of improved genetic material embodied in
open-pollinated varieties. Multilocation on-farm testing can bring experimental varieties
to small groups of farmers earlier than via official release mechanisms. Multiplication
initiatives coordinated by researchers and NGOs, combined with seed sharing among
farmers, will distribute seed more widely. Pilot, community-based seed schemes in
various countries are currently underway which will provide needed information
regarding factors correlated with replicability, sustainability, and cost-effectiveness
of such schemes.
8.7 Review of Priority Areas of Research and Development

To produce new maize varieties for Africa, a balanced programme approach is needed
which takes advantage of potential for improvement from both conventional breeding
and biotechnology approaches. Some recent advances in African maize research and
production help to illustrate the type of opportunities which exist.
1. Drought resistance Rain-fed farming systems across Africa are subject to intermittent periods of drought which significantly reduce maize yields. Yield losses due to
drought in non-temperate zones of the world are estimated at 19 million tonnes
annually. Heisey and Edmeades (1999) estimated that 21% of the maize area in Africa is
often affected by drought stress. Drought stress has been exacerbated in recent decades
of declining soil fertility, which is often associated with reduced soil water-holding
capacity. Research at CIMMYT on the effects of drought stress on maize over a 25-year
period resulted in the identification of several screening methods capable of identifying
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Maize
genotypes which tolerate to some degree the effects of moisture stress. Of these, reduced
anthesis-silking interval (ASI) trait has been selected as a relatively straightforward,
easily-measured trait which correlates well with yield under stressed conditions. By
reducing the interval between pollen shed and silking in maize, better synchronization is
achieved within these two plant processes critical to fertilization. Shortened ASI is now
being used as an evaluation criterion in ten countries of southern Africa through the
Southern Africa Drought and Low Fertility Network. Additional upstream research on
drought resistance using molecular markers and genomics is also warranted.
2. Nutrient use efficiency Resistance to low soil nitrogen has been observed to be
associated with drought tolerance in maize. More recent work on phosphorus
acquisition efficiency in maize appears to show improved performance for this trait
through increased resistance to soil acidity, related to better-developed root hairs and
exudation of organic acids by some maize varieties. In view of the general sensitivity of
maize to low nitrogen and phosphorus availability and of the variation expressed in
some genotypes, both these mechanisms appear to merit further research.
3. Multiple resistance to foliar diseases and ear rots Reduced maize grain production
due to pathogen-caused loss of photosynthetic area and phytotoxic effects is a preventable type of loss which affects small-scale farmers most directly. Making maize more
resilient to multiple foliar pathogens and ear rots is becoming increasingly possible by
the identification of molecular markers for resistance genes. Efforts in this area would
include marker identification work taken on primarily by international centres, followed
by marker-assisted selection programmes conducted in Africa by national breeding
programmes.
The identification of a stable molecular marker for MSV resistance on chromosome

1 of maize (Hoisington, personal communication) means that back-crossing of this
gene into well adapted local varieties can be accelerated in national programmes. It
also means that cloning and transfer of this gene can now proceed, facilitating its
movement into a wide variety of otherwise unmodified sources.
Clearly, a concerted emphasis on breeding for grey leaf spot resistance is required
for eastern and southern Africa. Molecular markers have been identified that are
linked to genes for resistance to grey leaf spot.
Ear rots which produce compounds toxic to humans have been a focus of research
conducted in Kenya and parts of West Africa. Improved methods of selection for
resistance to these pathogens may yield added benefits to human health.
Turcicum leaf blight is a major cause of yield loss. Resistance levels to the disease in
improved maize varieties are variable. Efforts to improve levels of resistance through
deployment of the four known resistance genes should continue.
4. Stem borers A considerable amount of work has already been invested in stem
borer resistance, including identification of markers for resistance to multiple species of
borers, screening methodologies and identification of potential donor lines. Further
work can proceed through both conventional and marker-assisted selection methods.
However, in view of the success of Bt maize in controlling stem borers in other regions,
the development of Bt maize for Africa should be given priority. CIMMYT is now
producing transgenic lines of African maize with Bt genes for borer resistance.
5. Postharvest insect pests CIMMYT Harare has initiated research on a limited
number of maize lines adapted to southern Africa which showed resistance to weevils.
Breeding work should now be undertaken to confirm the usefulness of this resistance
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within breeding programmes. Due to the difficulty of screening for resistance, however,
identification of molecular markers is likely to speed introgression of resistance genes
into African-adapted populations. Meanwhile, transgenic maize has been produced with
avidin glycoprotein from chicken egg white, which has been found to prevent development of insects that damage pests in storage (Kramer et al., 2000).
6. Striga Research on Striga-resistant maize has yielded promising avenues for
continued work involving gene doning in Tripsacum, gene cloning via mutator
transposons, and assembly of technology packages utilizing herbicide-resistant maize.
7. Ecosystem development factors In addition, IARCs and NARSs are pursuing
ecosystem development strategies in underexploited environments such as West Africas
southern guinea savannah and humid forest zones and eastern and southern Africas
semi-arid regions. As the agro-ecologies of these areas become better understood, this
should also lead to the identification and prioritization of needed traits and may require
the development of new base populations for breeding purposes.
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Sorghum
9.1 Brief History of Sorghum Cultivation and Utilization in Africa

Sorghum, together with millet, represents Africas most important contribution to the
world food supply (see Plate 13). Sorghum was domesticated in Ethiopia and parts of
Congo, with secondary centres of origin in India, Sudan and Nigeria (Fredricksen,
1986; Mukuru, 1993).
Harlan and de Wet (1972) described five races of cultivated sorghum which have
come into common usage among sorghum breeders. They are: durra, kafir, guinea,
bicolour and caudatum. All five major races of sorghum originated and continue to be
cultivated in Africa, with several races often being used for differing purposes within the
same agro-ecosystem. Guinea sorghum varieties are cultivated primarily in West and
central Africa, with some landraces spreading as far south as Mozambique. Kaffir types
originated in eastern and southern Africa. Durra sorghums developed primarily in
Ethiopia and the Horn, but are also spread across a wide section of Nigeria and savannah
areas of West Africa. Caudatum varieties were developed in Kenya and Ethiopia.
Bicolour, the least important of cultivated races, is sparsely distributed through East
Africa (de Wet et al., 1970).
Although sorghum cultivation has become an important component of agriculture
in a few industrial countries, it remains largely a developing country crop. Ninety per
cent of the worlds area cultivated to sorghum is in developing countries, mainly in
Africa and Asia. In Africa, 74% of sorghum produced is consumed in the home
(FAO/ICRISAT, 1996), primarily as thick or thin porridges, or as traditional beer.
Other African foods prepared from sorghum include green ears, flat breads and rice-like
dishes prepared using boiled sorghum (NAS, 1996). Sorghum stover is an important
source of animal feed in mixed farming situations.
Sorghum has a nutritional profile roughly similar to that of maize. Most varieties
register approximately 9% protein, generally 12% higher than maize; however,
sorghum is generally lower in fat content by a similar amount. Both grains are low in
lysine, and the crude protein digestibility of sorghum is severely reduced by high
111
111
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percentages of prolamine and tannins, necessitating additional processing of grain in

the home. It is probably due to the grains prolamine content that sorghum is often
fermented prior to consumption (NAS, 1996). Tannins (present to discourage bird
damage) are removed in the de-hulling process.
9.2 Sorghum Production Levels and Trends in Africa

In 1995, world production of sorghum was 53 million tonnes, or 4% of total cereal
production, making sorghum the worlds fourth most important grain crop (House,
1996). Due to its excellent adaptation to semi-arid and arid climates, the proportion of
total grain production represented by sorghum in semi-arid countries of Africa is very
high (see Table 9.1).
The rate of growth of sorghum production in Africa during the period 1985
to 1994 was 1.7%, however, the growth rate of area planted to sorghum was 2.9%,
indicating faltering yields of sorghum in some areas. Although maize is generally
considered to be more sensitive to low soil fertility, reductions in soil fertility levels
throughout Africa have affected sorghum yields as well, reducing growth rate of
sorghum yields during 1985 to 1994 to 1.2%. Partially due to the harsh conditions
in which the crop is normally grown and partially due to low harvest indices of most
popular cultivars, sorghum is relatively low-yielding. Average sorghum yields in Africa
are 780 t ha1 (FAO/ICRISAT, 1996).
Figure 9.1 shows sorghum production trends in sub-Saharan Africa as a whole
and in the sub-regions. The graph plainly shows the bulk of African sorghum
production to be centred in West Africa. West Africa is responsible for 60% of total
sorghum production in Africa and roughly 25% of all sorghum grown in developing
countries (FAO/ICRISAT, 1996).
In some farming systems, maize has replaced sorghum as the principal cereal crop
(Rohrbach, 1994). However, the trend over the past 35 years shows sorghum holding a
relatively constant position. In southern Africa, for example, total sorghum production
was 15% that of maize production in 1961. By 1997, that figure had decreased only
slightly, to 12%.
Table 9.1. Importance of sorghum production in selected countries of sub-Saharan Africa.

Country
Sorghum production (% of total cereals)
Burkina Faso
Cameroon
Chad
Mali
Rwanda
Sudan
Africa
53
40
41
38
52
72
18
Source: House et al. (1997) and FAO/ICRISAT (1996).
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Sorghum
Fig. 9.1.
Sorghum production trends in Africa, 19751998. Source: FAO (1998).
9.3 Sorghum Production Constraints

Sorghum crops suffer from attack by 16 major diseases, among which grain mould
(Curvularia sp. and Fusarium moniliforme) (see Plate 14), long smut (Ustilago sp.),
anthracnose (Colletotrichum graminicola), grey leaf spot (Cercospora zeae-maydis), sooty
stripe (Ramulispora sorghi) and head smut (Sphacelotheca reiliana) are of major
importance in Africa (Thakur et al., 1997).
Sorghum is also attacked by a variety of panicle- and stover-feeding insects. The
panicle pest of greatest importance in African sorghum is probably sorghum midge
(Stenodiplosis sorghicola), followed by African sorghum head bug (Eurystylus oldi)
(Henzell et al., 1997), although landraces of sorghum often have resistance to the latter
pest. Insects which attack the foliage and stems of sorghum include green bug
(Schizaphis graminum), shoot fly (Atherigona soccata) and spotted stem borer (Chilo
partellus). Stem borers are an especially important pest of sorghum in eastern and
southern Africa, with few or no sources of host-plant resistance having been discovered
(van den Berg, 2000).
A crop-loss compared with crop-improvement benefit analysis of sorghum
production constraints performed by ICRISAT in 1992 (ICRISAT, 1992) provided
a ranking of pest and disease importance in sorghum for Africa and Asia combined
(Table 9.2).
Among the other major constraints to sorghum production in Africa, day-length
sensitivity has received considerable attention. Large areas of sorghum cultivation in
Africa are devoted to photoperiod-sensitive varieties which commence reproductive
stages of growth when days shorten to a critical period, regardless of their time of
planting. Fixed timing of grain filling regardless of overall plant maturity can reduce
yield significantly. However, the primary function of photoperiod sensitivity is to ensure
that the grain matures under dry conditions, as sorghum grain (and home-stored seed)
deteriorates rapidly when stored wet. As such, elimination of photoperiod sensitivity in
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Table 9.2. Yield loss estimates and return to crop improvement for sorghum in
Africa and Asia.
Identified constraint
Yield loss
($ million)
Potential gain via crop improvement

($ million)
1744
334
129
274
153
102
77
198
21
143
124
121
102
83
67
43
38
15
Drought
Stem borer
Grain mould
Shoot fly
Striga
Anthracnose
Leaf blight
Head bug
Smut
sorghum varieties for use by small-scale farmers is limited by rainfall patterns and
farmers flexibility in postharvest management of the crop (Gomez and Chantereau,
1997).
Sorghum and millet are the most important grain crops of the semi-arid regions of
Africa and Asia. Both crops display impressive abilities to withstand low soil moisture
status and high air and soil temperatures. Drought tolerance in sorghum is still poorly
understood, but has been attributed to several mechanisms, including leaf rolling
under water stress, osmotic adjustment and the stay-green character (for post-flowering
drought) present in some varieties (Rosenow et al., 1997). However, there is disagreement among sorghum breeders regarding the usefulness of the stay-green trait under
severe drought conditions. At high levels of moisture stress, leaf senescence may be
advantageous in reducing transpiration to a sustainable level (Fred Rattunde, personal
communication). Researchers gathered to discuss options for improving drought
tolerance in sorghum in 1999 cited stay-green, lodging resistance, resistance to charcoal
rot, seed filling and stem reserves as potential selection criteria (CIMMYT, 2000).
9.4 Sorghum Improvement Through Breeding and Biotechnology

Sorghum breeding advances in Africa
Breeding methods
Sorghum is considered to have high levels of untapped diversity. The largest collection
of sorghum germplasm is the US sorghum collection (held at the National Seed Storage
Laboratory in Fort Collins, Colorado, and the USDA-ARS Plant Genetic Resources
Conservation Unit in Griffin, Georgia), containing over 40,000 accessions. Less than
3% of these have been used in crop improvement (Dahlberg et al., 1996).
Sorghum is a mostly self-pollinated crop with a variable percentage of out-crossing
existing in most cultivars, ranging from 515%. The most common methods of
selection are based on pedigree techniques developed for self-pollinated crops such as
rice and wheat. However, breeding teams at Purdue University, University of Nebraska,
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Kansas State University, and ICRISAT-Asia Center have applied population improvement techniques (including reciprocal recurrent selection) on sorghum based on
quantitative genetics theories of increasing favourable alleles within base populations
(Rattunde et al., 1997). Gains from selection from such programmes tend to be quite
high, in the order of 150400 kg ha1 (Rattunde et al., 1997). However, a long-term
population improvement programme conducted in Mali during 1970 to 1988 has since
been discontinued for lack of sufficient progress.
Following widespread success of caudatum races of sorghum in India, international
sorghum improvement programmes in Africa have based much of their efforts on
caudatum germplasm. Caudatum releases grown under medium-to-high input levels in
Africa have consistently shown yield advantages over guinea varieties, in part due
to higher planting densities made possible by shorter plant types. Although caudatum
varieties have performed well and achieved a high level of acceptance among local
farmers in southern Africa (Obilana et al., 1997; Rohrbach and Makhwaje, 1999), in
West Africa adoption rates have remained low (Tour et al., 1998). Recently, more
emphasis has been placed on guineacaudatum crosses and guinea crosses with
other races, and several groups have begun development of guinea hybrids and guinea
synthetics (Fred Rattunde, personal communication). At the same time, some farmers
have demonstrated increasing interest in caudatum varieties based on ease of harvest,
higher yield potential and better quality of forage. Guinea sorghum stalks, although
often used in house construction, are indigestible by cattle. As housing materials
improve, therefore, farmers may become more open to use of caudatum varieties.
Recent breeding efforts by several groups have emphasized the use of tan plant
mutants, which are reported to carry higher yield potential and white grains which do
not discolour on contact with pigmented glumes (Rosenow, 1997). However, there
is not complete agreement regarding the potential of tan plant types for African
agriculture, where yield is often reduced by biotic and abiotic stresses and few markets
reward farmers for producing a higher quality product. Studies of isolines of tan and
non-tan plants have shown adaptation advantages in non-tan lines in West Africa
(Rattunde, personal communication).
Problems of low adoption
Improved sorghums have achieved moderate levels of adoption in parts of southern
Africa (Obilana et al., 1997), Cameroon, and Chad (Yapi et al., 1999). In other areas,
adoption rates of improved sorghum among small-scale farmers have been low (Ibrahim
et al., 1995; Ahmed et al., 2000). Low rates of adoption of improved varieties are
probably influenced by poorly developed commercial and/or public seed systems and
low usage of fertilizers and other inputs in areas where sorghum is grown (Ahmed et al.,
2000). However, a review of on-farm trial data showing higher yields but low acceptability among farmers points to other factors, as well (White and Chapman, 1996).
These trends point to the need for more farmer-focused, participatory methods of
improvement which identify constraints and varietal preferences prioritized by local
farmers. These issues are explored in greater detail below.
Persistently low rates of adoption of improved varieties in West and central Africa
point to a potential need for modification in approaches employed in the improvement
of the crop. Some basic principles for a new approach might include:
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As an indigenous crop consumed primarily in the home, acceptable grain quality

for food processing and preparation is a sine qua non to widescale adoption among
small-scale farmers.
Better understanding of traits which confer acceptable grain quality may allow
greater efficiency in transferring both routine and intractable resistance and
tolerance traits to farmers via higher adoption rates.
Because of the complexity of quality-related issues, success in this area will most
likely depend upon direct collaboration among breeders, plant scientists, food
scientists and farmers. As such, deployment of resistance genes for priority pests
and diseases will be limited by breeders ability to deliver these improvements
within a package which is acceptable to local consumers.
The success of any new approach would appear to depend upon farmer participation both in priority setting for breeding programmes and in selection of varieties, and a
greater understanding of the importance of grain quality in marketing of harvests
destined for urban markets. In fact, farmer and consumer participation in sorghum
improvement in West Africa may represent one of the best opportunities yet for
demonstration of the power of wider participation in crop improvement initiatives
(Ejeta, personal communication).
Grain quality issues
Herdt and Capule (1983) previously analysed the persistent use of traditional varieties of
rice in Thailand in spite of the availability of improved varieties. They determined that
in some areas the price discount for improved varieties effectively offset the advantages
they represented in yield. When improved varieties began to be introduced with quality
characteristics preferred by farmers, they were adopted rapidly. Likewise, food scientists
who have analysed sorghum quality characteristics have become increasingly capable of
predicting the acceptability of improved varieties based on the quality of food products
they produce (Fliedel and Aboubacar, 1998). Studies conducted using sorghum flours
from West, southern and east Africa revealed significant differences in flour texture and
total water content of porridges consumed. Households preferred varieties with high
amylose starch content and low flour lipids and proteins. Few improved varieties have
scored high in such tests; nevertheless, breeding teams have failed to take full advantage
of food scientists ability to inform them of the probable success of their offerings at the
household level.
Research conducted in parallel to this by economists at Michigan State University
and ICRISAT (Rohrbach and Makhwaje, 1999) has revealed that sorghum grain market
development suffers from a host of quality-related issues beginning at the farm gate and
extending to the level of confectioners hoping to make usable products from these
grains. This relates to problems of cleaning, grading, and differentiating of sorghum
grain according to colour, grain size and overall preparation characteristics.
Out of this impasse has formed a small but growing contingent of sorghum
specialists calling for a new, more participatory and more integrated approach to
sorghum improvement which would build in marketing and quality aspects. The
integration of breeders with food scientists and economists working on utilization issues
would represent a radical shift from current, production-factor-led initiatives, and, due
to its complexity, would involve certain risks of its own. However, it is important to note
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the growing belief that sorghums integral position within the dietary and household
management traditions of Africa may require a different approach to that employed in
other grain crops such as maize and rice.
Advances in sorghum biotechnology

Molecular genetics
RFLP linkage maps have been developed for sorghum and several are highly saturated
(Chittenden et al., 1994; Pereira et al., 1994; Tao et al., 1996); however, few important
genes have been placed on these maps, and they have thus proved of little practical use
for purposes of genetic improvement. Development of sorghum linkage maps has been
greatly assisted by the high level of conservation of genomes between sorghum and
maize (Binelli et al., 1992; Berhan et al., 1993) and by the relatively small size of its
genome (three times smaller than maize). A sorghum bacterial artificial chromosome
(BAC) library has been constructed at Texas A&M University (Woo et al., 1994), and
the same programme currently aims to develop a high-resolution molecular map of
sorghum in order to facilitate marker-assisted selection (Johanson and Ives, 2000).
Sorghum breeders concluded in mid-2000 that development of a saturated simplesequence repeat (SSR) map of sorghum is still a priority for furthering sorghum
improvement through molecular breeding (CIMMYT, 2000). Meanwhile, Bhattramakki et al. (2000) have published an integrated RFLP and SSR linkage map composed
of 147 SSR and 323 RFLP loci.
Several studies are currently underway aimed at employing MAS for the improvement of sorghum. Researchers at Texas Tech University are involved in identification of
markers for the stay-green trait (Nguyen et al., 1996). A second team at Texas Tech has
initiated a project on marker identification for osmotic adjustment (drought tolerance
trait) in sorghum. Researchers at Texas A&M are working on marker identification for
various disease resistance traits, including downy mildew, anthracnose, leaf blight, head
smut and grain moulds (Magill et al., 1997).
Genetic engineering
Once considered recalcitrant to regeneration in vitro, methods have now been developed
for cell culture regeneration using immature embryos, young leaf bases, shoot apex, and
immature inflorescences (Bhaskaran and Smith, 1990). Successful anther culture has
also been reported from six varieties of sorghum (Kumaravadivel and Rangasamy,
1994). Useful cultivars have also been developed from somaclonal variation methods
(Smith et al., 1997).
Sorghum has been transformed and stable expression of genes obtained using
microprojectile bombardment of immature embryos (Casas et al., 1995; Rathus et al.,
1996) and more recently via Agrobacterium-mediated transfer by scientists at Texas
A&M (Nguyen et al., 1996). Pioneer Hi-Bred of the USA has successfully transformed
sorghum with a high lysine gene, but have not commercialized this genotype (Johanson
and Ives, 2000). Transformation of sorghum via Agrobacterium, however, has so far
reportedly been less efficient than for rice and maize (Nguyen et al., 1996). CSIR
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Bio/Chemtek of South Africa have reported only chimeric (non-stable) transformation

of sorghum (Johanson and Ives, 2000).
Thus, whereas significant academic research has been invested in sorghum biotechnology, the application of this new knowledge to genetic improvement of sorghum
for Africa has lagged considerably. In view of the important role the public sector is
destined to play in this low-value, high-volume African crop, the stage appears set for a
donor or group of donor agencies and African NARSs to enter into collaborations aimed
at extending the benefits of this research to farmers, possibly focusing on resistance to
stem borers, via Bacillus thuringiensis.
9.5 Principal Challenges for Sorghum Improvement in Africa

West Africa focus
Given its predominance as a crop and dietary component in the region, this document
focuses primarily on challenges of sorghum improvement in West Africa. Sorghum
accounts for a far greater percentage of total cereal production (55%) in West Africa
than in either eastern (10%) or southern Africa (less than 5%). Furthermore, the impact
to date of sorghum improvement, as measured by uptake of released varieties, has been
greater in eastern and southern Africa than in West Africa (improved sorghum varieties
currently cover approximately 26% of total area in each of Botswana and Zambia and
40% of total area in Zimbabwe (Obilana et al., 1997), compared with 20% in Mali
(Ahmed et al., 2000)), implying that some of the gains available through sorghum
improvement may have already been achieved in eastern and southern Africa. Moreover,
advances made in developing and disseminating drought tolerant maize in southern
Africa may contribute to past trends of replacing sorghum with maize in that region. It is
unlikely, however, that maize can be made sufficiently drought tolerant to replace
sorghum in extensive areas of West Africa which receive less than 600 mm of rainfall
annually.
Focus on adaptation and accessibility

As an almost purely locally-consumed commodity, the need for an emphasis on end-user
acceptance in West Africa is understood. To the extent possible, therefore, biotechnology and breeding efforts must be applied to locally adapted and accepted germplasm. In
addition, in view of the regions marginal production conditions and the small resource
base of the vast majority of farmers, sorghum improvement focusing on West Africa
should aim at developing products which are within the reach of small-scale producers.
Based on recent economic indicators and analysis from West Africa, it would
appear that to treat the regions sorghum farmers as a homogeneous unit of producers
and consumers would be a mistake (Coulibaly et al., 1998). Therefore, increased analysis
of agro-ecological diversity in sorghum-based farming systems of West Africa may
lead to more precise targeting of breeding efforts. Devaluation of the West African
franc in the early 1990s has led to an upsurge in agricultural production in some
sorghum-producing countries, most notably Mali and Burkina Faso. Meanwhile, major
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increases in cotton production have improved liquidity among small-scale producers.

Large increases in local prices of rice and wheat (the two major imported food commodities) have led to substitution of those products in the diets of many urban consumers.
Increased demand for the locally produced cereals, sorghum and millet, have created
incentives for farmers to intensify production of those crops. Of late, farmers have
shown increasing interest in caudatum varieties in Mali such as ICRISAT cultivars
ICSV 901 and ICSV 1079 (Dale Hess, personal communication).
Overall, the scenario created by economic reforms in West Africa appears to favour
moves toward higher-productivity production systems. Major changes to be anticipated
among farmers would include increased rates of usage of inorganic fertilizers and
higher-yielding varieties. Within this context, the potential for adoption of hybrid
sorghums may be higher than in the past. The opportunity also exists, therefore, for
science to support such improvements through the development of new technologies,
including adapted hybrid varieties.
Dual track approach

Promoting food security, equity, and economic growth among sorghum producers in
West Africa during the coming decade may lend itself to a dual track approach, which
pursues differing strategies for resource-poor and better-off farmers. Breeding strategies
and objectives emanating from this approach are explored in greater detail below. A brief
description of the principals guiding science and breeding in each environment is given
by a ranking of the principal thrusts within each improvement initiative, as follows.
Medium/high input environments:
1. Adaptation
2. Yield
3. Quality1
4. Resistance
Low input environments:
1. Quality
2. Resistance
3. Yield
4. Adaptation
9.6 Sorghum Seed Systems

Sorghum seed delivery is underdeveloped in most parts of Africa, and in its current state
should be considered a major constraint to the distribution of the fruits of research. In
spite of its importance to crop improvement at the farmer level, little study has been
dedicated to understanding how sorghum seed delivery in Africa can be made more
sustainable, although studies of traditional seed collection and conservation have been
conducted (World Vision International, 1995; INTSORMIL/INRAN, 1998). Seed
companies in eastern and southern Africa have been reluctant to transfer seed
Including aspects of both plant type and grain quality.
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production and marketing capacity from maize to sorghum, primarily out of concern for
sorghums low profitability and erratic seed demand (Ahmed et al., 2000). In Niger,
farmers who purchase improved varieties do so primarily following a drought year,
because improved varieties are earlier maturing than local varieties (Salifou, 1998).
Farmers who purchased improved seed in Niger were primarily farmers who had some
link to a producers association where credit and other purchasing assistance is available.
In contrast, hybrid sorghum seed sales in Sudan and South Africa have been largely
profitable (Ahmed et al., 2000; Ejeta, 1998). Experience with hybrids in these countries
and India have prompted some researchers to propose a shift toward hybrid sorghum in
West Africa as well, most notably in Niger, where a conference was recently held on the
subject of hybrid sorghum and millet (INTSORMIL/INRAN, 1998).
The perceived potential for hybrid sorghum varieties in West Africa is based largely
on experience from elsewhere in the developing world. Hybrid sorghum varieties were
first released in the USA in 1956 and in India in 1964. Sorghum seed markets in India
and Sudan developed largely on the basis of breeders enhanced ability to package yield
potential and stress tolerance within hybrid varieties and seed companies ability to
realize an acceptable profit margin from dependable sales of hybrid seed. The first hybrid
sorghum in Africa, Hajeen Dura-1 was released in Sudan in 1983. Hybrid sorghum
varieties have been available in Zimbabwe for some time and have recently been introduced in Zambia and Botswana. Until recently, however, no hybrids had been commercialized in West Africa.
Hybrid varieties of sorghum have demonstrated high levels of heterosis. The
average level of heterosis in released hybrids in Africa to date is 45% (House, 1996). The
US figure is only 36%. Unlike in maize, where hybrid varieties have often been
perceived (often mistakenly) as having less advantages in marginal conditions, hybrid
sorghums are generally accepted as having both high yield (hybrids in Niger, for example, generally yield twice as much as local varieties under similar management) and
greater resistance to environmental stress, plus increased seedling vigour and earlier
maturity. The importance of these traits in semi-arid environments lend increased value
to hybrid sorghums in most of the areas where sorghum is an important crop. Breeders
of long experience with sorghum seed systems further state that commercial supply of
sorghum seed has rarely developed without hybrids.
At present in Niger there is no public seed production/distribution entity and only
a single, small, private seed company, which in 1997 produced 200 t of seed of various
crop species. Previous production of seed in Niger (3511 t in 1985 92% of total needs)
focused primarily on millet and was heavily subsidized by the government. The CMDT
(Malian cotton parastatal) is reported to have developed hybrid varieties and is testing
them on-farm this season. In Mali, as well, seed distribution capacity is very low.
Government projects in both Mali and Niger distribute seed of improved cultivars
locally in small quantities. There is no private cereal seed company in Mali. In the
absence of other means, NGOs have increasingly taken up the task of sorghum seed
distribution, with some level of success (Rosenow, 1997).
In view of the lack of responsiveness of the private sector to seed distribution
opportunities, researchers have consistently called for the development of the informal
seed system. To date, however, no clear plan has been advanced for non-commercial
distribution of sorghum seed.
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In prioritizing upstream research for the various constraints to production of sorghum,
several options exist.
1. Striga. The most important biotic constraint to production of sorghum in
sub-Saharan Africa is probably Striga hermonthica. Breeding initiatives at ICRISAT and
Purdue University have developed sorghum varieties with moderate levels of resistance.
More recently, wide crosses to wild sorghum have shown promise (Gebisa Ejeta,
personal communication).
2. Resistance to anthracnose. Anthracnose is perhaps the most damaging disease of
sorghum in a large area of West Africa, and both landraces and introduced varieties are
affected. Developing adequate levels of resistance has proved extremely difficult, due to
low heritability of the trait.
3. Resistance to downy mildew. As adoption of more uniform, improved varieties
increases, downy mildew incidence is likely to rise. Management of downy mildew via
host-plant resistance is difficult, but may yet present a more accessible possibility than
use of fungicides among small-scale farmers.
4. Insect resistance. Resistance to panicle pests in sorghum may form another priority
area of research. Sorghum midge is reportedly the most ubiquitous and damaging insect
species of sorghum, worldwide. Resistance is controlled by an unknown number of
recessive to partially dominant genes (Aggrawal et al., 1988). It is a polygenic trait which
has been suggested as an ideal candidate for improved control through markerassisted selection (Henzell et al., 1996). Stem borer-resistant sorghum would be of
undisputable value to sorghum farmers of southern Africa. The possibility of developing
transgenic Bt sorghum has been mentioned by several authors, but thus far no strategies
have been advanced. Its development should be considered a priority.
5. Sorghum grain mould. Sorghum grain mould is a major problem on improved
cultivars in most of West and central Africa. High levels of resistance to grain mould is
one reason farmers of the region remain attached to traditional, guinea varieties. Guinea
varieties produce a loose, branching head which dries quickly following rains. Most
improved varieties have more compact, non-branching heads. Breeding initiatives have
discovered various sources of resistance, including waxes, glume pigmentation, polyphenols and corneous grain texture (Stenhouse et al., 1996). Marker identification
studies are also underway. Antifungal proteins have also been proposed as a means of
resistance. Their use will require genetic transformation methods.
6. Heterosis in adapted materials. Several researchers have advanced plans for the
development of guinea hybrid sorghum varieties for West Africa. Given high potential
levels of heterosis in such varieties, yields could be significantly increased, provided the
new varieties were well adapted and seed was available at affordable prices.
7. Pest/disease complexes of localized importance. Beyond these intractable problems,
there exist a number of pests and diseases of perhaps lesser individual importance that
combine to form complexes which reduce yields considerably in farmers fields. Like
head bugs and grain moulds, they may create barriers to adoption of improved, higher
yielding cultivars. Resistance to downy mildew, anthracnose and charcoal rot could
form the focus of molecular studies, and also form the focus of downstream support to
national breeding programme efforts.
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8. Phosphorus acquisition efficiency. Sorghum varieties with differing levels of

phosphorus uptake efficiency have been identified in Brazil (Schaffert et al., 1999).
These genotypes, which are adapted to Brazils cerrado region, should be tested for
performance in low phosphorus soils of Africa. In the event a mechanism for increased
uptake efficiency can be identified, breeding strategies could be advanced to enhance
expression of this trait in adapted germplasm.
9. Sorghum seed systems. In view of the above discussion, seed systems must represent a
focus of any crop improvement programme aimed at sorghum. The scope of the present
study has been too brief to gather information needed to make recommendations on the
improvement of sorghum seed supply. In order to gather more information, it will be
necessary to sponsor a study and/or a workshop focused on sustainable supply of pure
line varieties.
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Pearl Millet
10.1 Brief History of Millet Cultivation and Utilization in Africa

Pearl millet (subsequently referred to as millet) is a crop of vital importance to millions
of African families living in semi-arid regions of the continent (see Plate 15). Millet is
one of the worlds most resilient crops. In many areas where millet is the staple food,
nothing else will grow. Several phrases from a recent publication by ICRISAT (1996)
perhaps sum it up best:
We are talking about a crop that is virtually unimprovable a crop that grows where not
even weeds can survive. A crop that has been improved by farmers and through natural
selection for thousands of years. A crop that produces nourishment from the poorest soils
in the driest regions in the hottest climates. A crop that grows straight out of sand dunes.
A crop that survives sand storms and flash floods.
Millet is descended from wild grasses native to the central Saharan plateau region of
Niger. From there it spread to East Africa and India, where millet ranks as the fourth
most important cereal. In West Africa millet is consumed primarily as a thick porridge,
or toh, but it is also milled into flour to prepare breads and cakes. Millet is the mostpreferred cereal grain grown in Sahelian countries, Senegal, Mali, Niger and Burkina
Faso, and is consumed in preference to sorghum. In northern Nigeria, millet flour is
used in making a popular fried cake known as masa. Roasted young ears are a popular
food for children. While sorghum is perhaps a better-known crop in most of the world,
most inhabitants of the Sahel actually prefer to consume millet, a fact which should
encourage greater investments in its improvement.
Feeding trials conducted in India have shown that millet is nutritionally superior
for human growth to maize and rice (NAS, 1966). It has slightly higher protein content
(average of 16%) than maize and roughly twice the fat content (57%) of most maize
varieties, and is particularly high in calcium and iron. It has lower levels of fibre
and most vitamins, although its pro-vitamin A content is relatively high. One important
problem for households which rely on millet as a food staple is its tendency to spoil
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rapidly (as a result of the fat content) following preparation. As constraints to labour
increase in Africa, this constraint is likely to increase in importance, giving rise to the
need for alternatives.
10.2 Millet Production Levels and Trends in Africa

Table 10.1 shows the importance of millet production in selected countries of subSaharan Africa. Five countries in West Africa (Nigeria, Niger, Mali, Burkina Faso and
Senegal) produce 85% of the continents total millet crop. Sudan accounts for 50% of
millet production in eastern and southern Africa. Figure 10.1 shows the relative importance of millet production in West Africa compared with the other two regions. West
Africa is also the only region where millet production has significantly increased over
time. However, all of this growth is due to increased area cultivated, and not increased
Table 10.1.
Africa.
Importance of millet production in selected countries of sub-Saharan
Country
Burkina Faso
Chad
Mali
Niger
Nigeria
Senegal
Sudan
Africa
Millet production
(1000 t)
Millet production
(% of total cereals)
734
228
815
1,769,
4,952,
667
385
11,740,
32
31
38
91
26
75
13
10
Source: House et al. (1997) and FAO/ICRISAT (1996).
Fig. 10.1.
Millet production trends in Africa, 19751998. Source: FAO (1999).
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Pearl Millet
yields. Growth rate of millet yields in sub-Saharan Africa during 1985 to 1994 was
2.3%.
Nevertheless, graphs are of little use in capturing the importance of millet in the
overall food security of Africa. Statistics are reported for countries, and not ecologies.
Along with sorghum, millet is the most important crop of semi-arid zones of Africa, and,
of the two, millet is the better adapted to marginal conditions. Millet is able to grow and
produce reasonable yields on 300 mm of rainfall per season, while sorghum requires
400 mm and the lower limit for maize is 500600 mm (ICRISAT, 1996). As such, like
sorghum, until higher-yielding grain crops such as maize or rice can be made highly
drought-tolerant, millet has a guaranteed place in the farming systems and diets of a
large and widely dispersed range of semi-arid regions in Africa.
10.3 Millet Production Constraints

Even by African standards, millet is a very low yielding crop under most small-scale
farming practices. A detailed survey of farming systems of the Sudan savannah ecology
of West Africa carried out by IITA showed that farmers generally harvest less than
500 kg ha1 of millet grain from cowpea/millet intercrops or cowpea/millet/sorghum
intercrops (Singh and Mohammed, 1998). While millet may be ideally suited to
cultivation in dry areas, it is far from immune to production constraints. Millets
principal defect is its tendency toward low harvest indices. While farmers make use of
stover as well as grain, their selection efforts have had to contend with tight linkages
between panicle size and maturity and reverse-phase linkages between tillering and seed
and panicle size (Rai et al., 1997). As a result, landraces of millet have harvest indices
as low as 14%, compared with 40% in hybrid millets (Kassam and Kowal, 1975).
Incidence of major diseases on landraces appears to be higher than on improved varieties
(Rai et al., 1997).
Surveys of farmers fields conducted to obtain estimates of losses from different
categories of pests in Senegal, Chad, Mali and the Gambia by Dively and Coop (1993)
produced the data summarized in Table 10.2.
The data from West Africa produce a rough ranking of panicle pests of: (1) birds,
(2) head miners and smut, (3) downy mildew and grasshoppers.
In all, millet is reported to suffer attack from some 111 different pathogens,
of which four downy mildew (Sclerospora graminicola), smut (Tolyposporium
penicillariae), ergot (Claviceps fusiformis) and rust (Puccinia substriata) are of major
importance in Asia and Africa (Singh et al., 1993). Of these, downy mildew is by far
the most widespread and damaging. Screening methods, selection techniques, source
germplasm and inheritance patterns have been developed for all four of these diseases
and have been summarized by Hash et al. (1997).
Smut resistance is readily available and inheritance of the trait is additive, making it
a relatively simple disease to breed for (Hash et al., 1997). Ergot resistance is controlled
by multiple recessive genes, and therefore has proved quite difficult to transfer into new
varieties. Downy mildew is able to evolve new virulent pathotypes rapidly in response to
control measures, including host plant resistance. Thus, although many sources of
resistance have been identified, all of these tend to vary in terms of their stability across
sites and years. Moreover, although gene action responsible for resistance to downy
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Table 10.2. Quantifying and ranking of millet pests and diseases in four West
African countries (after Dively and Coop, 1993).
Yield
Loss
Rank 14
Senegal
800
200
25
Mali
556
264
47
1083
131
21
172
97
56
1. Head miners
2. Downy mildew
3. Birds
4. Smut
1. Grasshoppers
2. Meloid beetles
3. Pachnoda beetles
4. Birds
1. Downy mildew
2. Birds
3. Head miners
4. Smut
1. Head miners
2. Grasshoppers
3. Birds
4. Smut
The Gambia
Chad
mildew is dominant, a number of genes are involved and inheritance is primarily

non-additive in nature, making transfer more difficult (Talukdar et al., 1994).
Millet panicles can be heavily damaged by millet head miners (Heliocheilus
albipunctella) (Henzell et al., 1997), a group of insects comprising a complex of
approximately a dozen damaging caterpillar species. Yield losses from head miner have
been described as 1385% in Senegal, 50% in Mali, and 6% in Niger (Toure and
Yehouenou, 1995). They reported no resistance or tolerance had been observed to date
for this pest. Progress in breeding for resistance to head miner was until recently reduced
by difficulties associated with development of a satisfactory screening method. Screening
methods have now been developed which can be broadly applied. Millet does not
normally suffer important losses due to foliar- and stem-feeding insects in West Africa.
The major insect pest of pearl millet in southern Africa is the armoured bush cricket
(Acanthoplus spp.) (Minja, 2000). In large areas of southern Africa, stored millet is
damaged by the rice weevil, Sitophilus oryzea; however, significant differences have been
noted among genotypes tested for resistance (Leuschner et al., 2000). Millet suffers
important losses to the postharvest insects Tribolium castaneum and Cryptolestes
ferrugineus in humid zones of the Sahel (Lale and Yusuf, 2000).
The most important biotic constraint to millet in many parts of West Africa is the
parastic weed, Striga hermonthica. In some seasons, infestation of fields is devastating,
with each host supporting the growth of 50100 Striga plants. To date, no resistance has
been documented in millet, although little research has focused on screening for this
trait.
Bird damage has probably been associated with open-panicle crop species in Africa
since the time of their domestication. Scaring birds from fields has, therefore, been a
traditional chore of children during the time of grain maturity, for just as long. As
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education has been extended to rural areas of Africa, labour to scare birds has become
scarce, increasing the need for bird-resistant varieties.
In the marginal environs in which millet is usually cultivated, water and nutrient
stress is a constant limitation to growth and productivity. Research on improving
drought and low nutrient tolerance in millet is covered in a separate study. Specific
mention is made here only of the priority placed on tolerance to low phosphorus
soils, which has been recommended for selection through marker-assisted selection
techniques (C.T. Hash, personal communication).
10.4 Millet Improvement Through Biotechnology and Breeding

Millet breeding advances in Africa
Millet is a cross-pollinated crop bred using techniques similar to maize, namely, by
hybridizing parents with complementary traits and extracting varieties or lines from
segregating generations. Various methods of recurrent selection are used to improve
populations.
The impact of work focusing on traits found within local varieties presents an
interesting case in millet. Selections from a local landrace originating in Togo, Iniadi,
have become the most widespread millet breeding material in the world (Andrews and
Anand Kumar, 1996). Iniadis success as a breeding material appears to be related to its
high general combining ability, earliness, high yield and grain quality. Fully 50% of
ICRISATs varieties and breeding lines contain Iniadi germplasm in their pedigrees, and
half of all commercial hybrids are produced using male sterile lines from Iniadi. Iniadi
materials also went into the development of Okashana1. Okashani1 is grown on
almost 50% of farms in Namibia (Rohrbach et al., 1999).
Whereas improved millet varieties have been widely accepted by farmers in southern Africa, adoption rates in West Africa, similar to the case of sorghum, have remained
low. Virtually all millet production in Niger is based on cultivation of local landraces,
despite several concerted attempts at distributing seed of improved varieties through
national seed campaigns during the 1980s. One possibility that breeders have not fully
understood what it is farmers desire in their millet varieties could potentially benefit
from the participatory breeding methods described earlier in this book, beginning with a
more full appreciation of the interactions between agro-ecologically based traits and
requirements within the user system.
High levels of heterosis have been demonstrated in millet. Hybrid millets have been
popular in the USA and India since the late 1960s, although hybrid use in India is
confined to areas of relatively high potential. Hybrids in India have 1520% higher
yields than improved OPVs (Rai et al., 1997); however, widespread use of single-cross
hybrid millet varieties in India has led to high levels of infection from downy mildew.
Hybrids have also proved to be more susceptible to smut, ergot and rust.
In view of low millet yields in Africa, researchers have proposed top-cross and
varietal-cross hybrids, which would include sufficient variation to avoid risk of
epidemics. The development of top-cross hybrids has been limited by lack of male sterile
lines within landraces, however limited tests of top-cross hybrids have shown much
promise (John Whitcombe, personal communication). Yields of hybrid millet varieties
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in Africa have shown 4050% yield increase over landraces (ICRISAT, 1992). Interpopulation hybrids using local landraces have also shown significantly increased yields
over improved OPVs (Lambert, 1983). Nevertheless, to date there is no hybrid millet
available for farmers in sub-Saharan Africa, a deficiency that merits concerted attention.
Male-sterile pearl millet populations with dwarfing traits have been developed for use as
seed parents in the development of hybrid millet for Nigeria (Rai et al., 2000), but no
hybrid varieties have as yet been released.
Methods of millet breeding are divided between recurrent selection (within populations formed with the incorporation of various desirable traits in mind) and pedigree
selection techniques aimed at developing fixed lines for use in hybrid formation. Increasingly, combinations of fixed lines are being used to form synthetic varieties. Rai et al.
(1997) recommended the use of composite varieties (mixtures of genotypes maintained
in bulk), as they can be used in or derived from hybrid breeding methods.
Advances in millet biotechnology

Tissue culture has been used to generate somaclonal variants of potential use in breeding
programmes (Vasil and Vasil, 1980; Prasad et al., 1984). Methods have also been
established for the development in vitro of multiple shoots from shoot apical meristems
(Devi et al., 2000). Bui-Dang-Ha and Pernes (1982) and, more recently, Shigemune
and Yoshida (2000) have reported regenerating fertile plants from anthers and microspores. However, use of these techniques is not yet widespread, and transgenic pearl
millet varieties are not at present in use in any breeding programme. In Africa, South
Africas Council for Scientific and Industrial Research (CSIR-Biotek) is carrying out
research aimed at genetic transformation of millet.
An RFLP map has been generated for pearl millet (Liu et al., 1994; Devos et al.,
1995). The map has been employed to identify 16 putative markers for race-specific
resistance to downy mildew (Hash et al., 1997), and appears poised for use in a variety of
other aims, should markers be identified. Markers linked to quantitative trait loci have
been identified for high harvest index, grain filling, and yield under terminal drought
stress (Yadav et al., 1999). Among the candidate traits for selection through markers
are head miner resistance, Striga resistance, drought tolerance and tolerance to low
phosphorus soils (Hash, personal communication). Molecular marker techniques are
also being applied to the task of characterizing pathogens of pearl millet (Sastray et al.,
1995).
10.5 Principal Challenges for Millet Improvement in Africa

As a relatively under-researched crop, little information is available on the millet varietal
preferences of farmers and the factors involved in the adoption of new varieties. Recent
results from farmer participatory breeding work carried out by ICRISAT in Mali suggest
a preference for large seeds, bold grain and earlier maturing varieties (Rai et al., 1997).
Grain quality issues, which have proved so important in the acceptance of sorghum
cultivars, may also be of importance in adoption of improved millets. Farmers also
distinguish between varieties which tolerate heat stress (especially at seedling stage), and
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Pearl Millet
those which do not. Exchange of varieties among farmers is known to occur, but is also
poorly understood. Additional information is needed on farmer varietal preferences and
how these relate to adaptation within different millet agro-ecologies. The status of
understanding of agro-ecologies for both sorghum and millet is at a rudimentary stage,
and requires additional research.
Finally, as a preferred food crop among large populations, and one with distinct
adaptation advantages in large areas of Africa, there are prospects for increasing yields
based on demand-driven strategies. One plausible strategy is the development of hybrid
millet varieties, targeted at countries where economies are relatively strong and millet is
an important crop, such as Senegal, Mali and northern Nigeria.
10.6 Millet Seed Systems

Millet seed systems are essentially the same as for sorghum, discussed in Chapter 9.

Among the traits which have been shown to reduce millet production and productivity,
a number can be identified for priority attention for programmes focused on millet
improvement. Although, as in the case of sorghum, a West Africa focus appears logical
for millet improvement in Africa, most of the traits listed below are of importance in
eastern and southern Africa, as well.
1. Resistance to head miners. Head miner damage is a major problem throughout Africa
and one for which no resistance has been deployed to date. Several reports indicate the
problem may have worsened over recent years. A gap appears to exist between the level
of understanding among laboratory-based scientists and breeders, whose narrowing
could result in rapid progress toward extending solutions to farmers. Moreover, as head
miners preferentially attack early-maturing millet varieties, susceptibility to this pest has
reduced the application of earliness traits in new varieties. As a classic, difficult-toscreen-for trait, marker-assisted selection techniques could probably benefit this area of
research.
2. Striga resistance. This is viewed as a long-term undertaking, but one which could
pay major benefits to farmers of West Africa, where Striga incidence on millet is a
devastating problem. Very likely, development of the trait will require a biotechnology
approach, as resistance has not as yet been documented in cultivated or wild-relative
germplasm. Transformation of millet using genes cloned from species with resistance
(cowpea, rice) may eventually prove a valid approach.
3. Bird resistance. Quelea birds can ravage unprotected crops of millet. Bird-scaring as
a means of protection is becoming more difficult to provide as increasing numbers
of children go to school. Bristled panicles have been shown to offer good levels of
resistance, and the trait can be manipulated through conventional breeding tactics.
Some groups of farmers have indicated they would accept bristled pannicles, provided
they offered adequate protection from this devastating problem. Exploratory research
into the effectiveness of the trait and its acceptance among farmers may prove useful.
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4. Resistance to low-phosphorus soils. One of the critical weaknesses of improved pearl

millet varieties in on-farm evaluations in West Africa over the past 15 years has been
their lack of adaptation to the low levels of available phosphorus in soils outside the
well-managed research stations where they were bred. Marker-assisted back-crossing
would provide the most efficient method for transferring improved ability to take up soil
phosphorus from one or more landraces to more agronomically elite, disease-resistant,
improved cultivars having acceptable grain quality and high grain yield potential.
However, before this can be done, the major loci contributing to improved phosphorus
uptake ability will have to be mapped, and tightly linked flanking markers for them
identified. Strategies for developing improved phosphorus use efficiency in sorghum
should be linked to similar efforts in millet.
5. Study of farmer varietal preferences. As described above, there has been very little
research aimed at elucidating genuine, small-scale farmers varietal preferences for pearl
millet in West Africa. All of the above traits will need to be combined into varieties
which satisfy farmers and consumers needs in terms of grain quality, plant type,
and maturity, etc. This can only be obtained through extensive interviews with local
producers and consumers. Therefore, a series of surveys, to be conducted within each
major agro-ecosystem of West Africa, is recommended.
6. Development of non-traditional hybrids. Given high levels of heterosis in stress
environments exhibited in millet hybrids of varying types, and millets popularity in
areas of West Africa, applied research targeted toward the testing of top-cross and
varietal-cross hybrids needs to be explored. Sustained, commercial seed supply for this
open-pollinated crop is as in the case of maize likely to follow on the development of
some form of hybrid. To date, very little research has been applied to hybrid millet for
Africa.
7. Resistance to downy mildew. High levels of loss to downy mildew are generally
associated with the use of single-cross hybrids, however, extensive observations have also
been made of incidence in landraces grown in West Africa. Durable resistance has
proved a difficult trait to select for, and thus, could benefit from added support via
MAS.
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Rice
11.1 Brief History of Rice Cultivation and Utilization in Africa

Africa is the centre of origin of one cultivated species of rice, Oryza glaberrima, which
was domesticated in the northern Niger valley by Africas first farmers. Oryza sativa was
introduced by European explorers beginning in the 16th century and from Indonesia,
via Madagascar. It has become the dominant species, although pockets of glaberrima
production continue to exist in various parts of West Africa. Rice ranks as Africas fourth
most important grain crop, behind maize, sorghum and millet, and is the primary source
of carbohydrates of farmers in parts of Liberia, Sierra Leone, Guinea, Nigeria and Mali.
For many farmers in East and southern Africa rice is an important secondary crop relied
on as both a source of income, as a niche crop in low-lying areas of small farms, and for
consumption on special occasions.
Because of its wide popularity as a food item, rice is among the most liquid of all
crop assets in Africa. Rice consumption in Africa has a high income elasticity, and
increases in its projected demand in Africa are tightly linked to increased urbanization
and economic growth, in part due to its ease of preparation among smaller, labourlimited households. These patterns are most evident in West Africa, where several
pockets of rapid economic growth have fuelled growth in demand for rice. Demand for
rice has increased at an annual rate of 5.6% since 1962 (WARDA, 1997).
In spite of its status as a cash crop, rice is still very important as a source of income
or food for very poor farmers of West Africa, especially in the inland valley swamp
ecologies of the savannah zones. In high rainfall areas of West Africa, rice and cassava are
relied on as the best extractor of phosphorus on highly leached soils (Sahrawat et al.,
1999).
Among traditional rice farmers of West Africa and Madagascar, rice is consumed in
a wide variety of forms, including porridge and as cakes. Among most consumers,
however, rice is eaten in the conventional way, boiled or parboiled, and served with a
relish containing fish or other animal protein. Highest per capita rice consumption
in Africa is in Guinea Bissau (112 kg per person year1), followed by Sierra Leone
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(88.6 kg per person year1), Guinea (73 kg per person year1), and Gabon (72 kg per
person year1).
11.2 Rice Production Trends in Africa

The annual rice production in selected countries in sub-Saharan Africa is shown in
Table 11.1.
Figure 11.1 reveals the predominance of West Africa in total African rice production. Production levels in southern Africa are highly influenced by Madagascar.
Likewise, those for East Africa are primarily accounted for by Tanzania, which contributes 80% of rice production in the region.
Table 11.1. Annual rice production in selected

countries of sub-Saharan Africa.
Country
tonnes
Cte dIvoire
Guinea
Mali
Nigeria
Sierra Leone
Congo, Democratic Republic
Madagascar
Mozambique
Tanzania
1,222,650
763,955
589,048
3,275,000
411,300
365,000
2,447,000
191,000
810,800
Source: FAO Internet database.
Fig. 11.1.
Rice production trends in Africa, 19751999. Source: FAO (1999).
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11.3 Rice Production Constraints

Management practices, especially water management, are of primary importance in
production limitations in Africa. Total control type of irrigation systems account for
only 12% of the total rice area in Africa, and water availability can be assumed to limit
rice growth at one or several stages of development in all other production systems (see
Plate 16). Nitrogen and phosphorus limit rice production on a wide range of soils in
West Africa and Madagascar.
Biotic factors are also of high importance in rice productivity in Africa. The
particular mix of biological pressures which affects crop production is often grouped
by production ecology. WARDA has described the range of priority constraints by
production ecology (WARDA, 1997), and the information is reproduced in Table 11.2.
Selected constraints cited in Table 11.2 are considered below.
Drought
Rice production is affected by drought stress in a variety of water-limited environments
of Africa. Upland rice, normally grown in high rainfall areas, may still suffer from
moisture stress at various periods during the season. Recent releases of early-maturing
rice offer the chance for farmers to avoid the effects of drought through earlier flowering
Table 11.2.
Priority rice constraints by rice production ecology (WARDA, 1997).
Rice ecology
Share of regional
Yields (t ha1)
rice area
(%)
Current Potential Priority constraints
Humid/
sub-humid zone
40
2.54.5
Rainfed lowland
38
1.4
3.05.5
Irrigated
2.8
5.07.0
Sahel irrigated
3.5
5.08.5
Mangrove swamp
2.56.0
Deep water/
floating
1.2
1.53.0
Weeds, acidity, blast,

drought, nitrogen
deficiency
Weeds, water control,
rice yellow mottle virus
(RYMV), nitrogen
deficiency, drought
Nitrogen deficiency,
weeds, RYMV, iron
toxicity, nematodes,
gall midge
Nitrogen deficiency,
cold, salinity, weeds,
RYMV, alkalinity
Sulphate acidity,
salinity, crabs
Water control, low
yielding varieties, low
fertilizer use efficiency
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and more determinant growth habits of rice varieties. Additional progress may be
possible through selecting for more extensive root systems.
Weeds
Weeds are a major problem in upland rice cultivation, where numerous grass species
of similar structure and adaptation as rice compete with the crop for light, water and
nutrients. Because rice is a relatively small plant (as compared with maize, sorghum and
millet) and slow to establish a full canopy, it suffers greater damage from competition
from medium-sized weeds (see Plate 17).
Insects
African rice gall midge (Orseolia oryzae) is predominantly a problem in irrigated and
lowland rice systems. Resistance to gall midge has been detected both in O. glaberrima
and in interspecific varieties. A moist environment is necessary for egg survival. Stem
borer is a lesser problem, also most serious in irrigated areas. Resistance to stem borers is
low in the rice genome, making rice a potential candidate for transformation using Bt.
Rice yellow mottle virus
RYMV is endemic to Africa, and primarily affects lowland rice ecologies. Although its
incidence is irregular, the effects of the disease can be devastating when it occurs.
Ghesquiere et al. (1997) developed varietal resistance to RYMV in doubled haploid
populations developed from crosses of upland japonica and lowland indica rice varieties.
Pinto et al. (1999) transformed African rice with RYMV transgenes. Resistance to
RYMV has been detected in both O. glaberrima and O. sativa varieties, and has recently
been successfully mapped to chromosome 4 (Ndjioniop et al., 1999).
Blast (Magnaporthe grisea)
Blast fungus is the most serious fungal disease of rice in Africa. It occurs throughout the
continent wherever large areas are cultivated to rice. Blast resistance has been noted in
recently developed interspecific rice varieties. Genetic studies of resistance have been
complicated by variability of the pathogen and lack of rice genotypes with single
resistance genes. One study has identified sites of major gene resistance at four different
loci (Mackill and Bonman, 1992).
Iron toxicity
This constraint occurs in the very high-rainfall zones of West Africa where excessive
leaching of other cations has left toxic concentrations of iron and aluminum in the soil.
It is most pronounced in lowland soils, but can occur in upland environments, as well.
WARDA invested heavily in breeding for iron toxicity tolerance in Liberia during the
1980s, leading to development of Suakoko 8, an iron-tolerant, intermediate variety
which has been adopted by some farmers in the region.
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11.4 Rice Improvement Through Biotechnology and Breeding

Rice breeding advances in Africa
The French research organization, Institut de Recherche de lAgronomie Tropicale (IRAT)
performed much of the early breeding on rice for West African conditions. Several of the
more popular varieties developed by this programme during the 1950s and 1960s are
still cultivated by small-scale farmers. Until the early 1990s IITA also maintained a
programme on rice improvement in Nigeria, which eventually was transferred to Cte
dIvoire and merged with the West Africa Rice Development Association (WARDA),
founded in 1971. WARDA divides its rice improvement programme into three sections:
upland, lowland, and irrigated, with the two former programmes based in Bouake, Cte
dIvoire, and the latter based in St Louis, Senegal.
At present, only the national programmes in Nigeria, Sierra Leone, Senegal,
Burkina Faso and Togo are involved in rice breeding. All others rely on evaluation of
lines developed by WARDA or other national programmes. All 17 WARDA member
countries are linked via a rice breeding task force which meets periodically to discuss
results of trials, set priorities, and obtain new materials.
The WARDA programme has made significant advances during recent years
through a breakthrough in breeding interspecific varieties selected from fertile progeny
of crosses between O. glaberrima and O. sativa lines (Jones et al., 1999). Previous
attempts at crossing the two species had proved fruitless, owing to high levels of sterility
in the progeny. One component of the breakthrough came through the use of anther
culture, which overcomes sterility by fixing lines in homozygous state directly from first
(BC1, F1) and second (BC2, F1) backcross generations.
Interspecific lines have successfully combined some of the most favourable aspects
of each species. Traits contributed by O. glaberrima include increased weed competition
through greater leafiness in lower parts of the plant, resistance to RYMV, African rice
gall midge and blast, drought tolerance and tolerance of iron toxicity. Traits contributed
by O. sativa include better response to increased fertility, higher yield through branched
tillers, and resistance to shattering (WARDA, 1999). Using participatory methods of
rice varietal selection, interspecific varieties have become popular in several countries,
including the major rice producers Cte dIvoire, Guinea and Nigeria.
Advances in rice biotechnology

Due to its high importance as a food crop and its small genome size, rice has become a
model plant for genetic mapping and genome analysis. The rice genome project is an
international effort led by Japan aimed at complete sequencing of the rice genome.
Various countries and laboratories have agreed to take responsibility for sequencing
portions of the genome.
Biotechnology applications for rice improvement were given a major boost at both
international and national levels through implementation of a 15-year, $100 million
International Program on Rice Biotechnology, sponsored by The Rockefeller
Foundation during the period 1985 to 2000. As a result, numerous new varieties with
new traits such as increased synthesis of vitamin A precursor, increased resistance to
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bacterial blight, blast, nematodes, gall midge and tungro virus are at various stages of
testing and release throughout Asia.
Prospects for applications of biotechnology in rice were reviewed by Khush and
Toenniessen (1991). The status of rice biotechnology has more recently been reviewed
by Tyagi et al. (1999). Rice has been reliably transformed via four methods: direct
transfer of DNA into protoplasts, electroporation of intact cells, DNA delivery via
particle gun bombardment, and Agrobacterium-mediated transformation. The first
genotype-independent system for rice transformation was developed by Christou et al.
(1991) using the particle gun. More recently, drawbacks related to transgene integration
patterns have led to greater use of Agrobacterium-mediated methods. Lectin genes from
snow-drop plants for insect resistance via lectin genes (Fujimoto et al., 1993), increased
synthesis of vitamin A precursor (Ye et al., 2000), rice tungro virus resistance (Fauquet
et al., 1997) and rice stripe virus resistance (Hayakawa et al., 1992) have all been
transferred into rice via genetic transformation. Transgenic rice with resistance to rice
yellow mottle virus has been developed via gene silencing, but has so far not been field
tested in Africa (Pinto et al., 1999).
Molecular genetics applications are perhaps more advanced in rice than for any
other food crop. RFLP and PCR-based markers are now available for a wide range of
important genes, including resistance to fungal diseases and viruses. In Asia, a large
number of public research facilities now have the capacity to select for these traits using
molecular techniques, creating broad scope for rice improvement in the region far into
the future. Moreover, new efforts are now being directed at detection of QTLs for
tolerance to moisture stress (John OToole, personal communication).
Saturated rice genetic linkage maps (McCouch et al., 1997; Cho et al., 1998) have
permitted identification of QTLs for many useful agronomic traits. Marker-assisted
breeding can now be applied to enhance the efficiency of conventional breeding
methods. RFLP markers were used to reduce the number of back-cross generations
required to incorporate three genes controlling blast resistance in rice (Hittalmani et al.,
1999). In an attempt at improving efficiency of selection for drought tolerance in
upland rice, Sarail et al. (1999) identified two QTLs associated with root thickness and
root penetration.
11.5 Principal Challenges for Rice Improvement in Africa

The low level of breeding capacity in national programmes of sub-Saharan Africa
represents a major structural challenge to the improvement of rice in the region.
Although WARDA and the breeding task force now operating in West Africa can
facilitate access to improved lines, the likelihood of adaptation of these genotypes to
local environments is inevitably left somewhat to chance. Moreover, integration of
molecular techniques in a manner similar to that achieved in recent years in Asia is
impossible unless there is widespread capacity in conventional breeding. Therefore,
there appears to be ample justification for a broad-based initiative aimed at increasing
breeding capacity.
Once established, this more decentralized breeding strategy will be able to take
ample advantage of the advances made by WARDA through its interspecific breeding
programme, including utilization of the technique for lowland and irrigated
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environments. Once again, trait-based strategies which aim at reducing losses due to
pests and diseases would appear to be a relevant, general thrust. Prioritization of these
traits at a national and sub-national level is a priority for future years. Identification of
such priorities will require the mobilization of breeding teams to interact extensively
with farmers regarding the incidence of biological and environmental constraints and
rice varietal preferences among local farmers. The existence of task forces which give
priority to vigorous, continual participation of a large number of national programmes
represents a major resource in the future implementation of regional strategies
constructed along these lines.
In the interim, rice improvement strategies can be focused on rapid deployment of
new traits and fixed lines made possible by breeding performed by WARDA. This aim
can be advanced by systematic distribution of segregating breeding families and lines
and the provision of support to national programmes for effective, multilocation testing
of these materials.
Drought is the most frequently cited source of yield loss among rice farmers
throughout the world. Likewise, in Africa, both lowland and upland (but most severely,
upland) rice production is regularly constrained by the incidence of periodic droughts in
the region (WARDA, 1997) (see Plate 18).
Strategies for improving drought tolerance in rice have recently been reviewed by
Ito et al. (1999). Molecular tools for detecting QTLs have been developed at Texas Tech
University, Cornell University and at IRRI (Nguyen et al., 1996) and now make
possible the selection for drought tolerance based on specific traits. Additionally, several
landraces of rice grown in the northern Sahel regions of West Africa have been observed
by the author to display tolerance to drought. Prospecting for genes in these landraces
using molecular genetics may be considered a secondary component of strategies on
drought tolerance for rice in Africa.
11.6 Rice Seed Systems

The development of sustainable rice seed systems is challenged by two factors inherent
to the crop itself, namely, its self-pollinating habit and high requirement for seed per
unit of cultivated area. The result of these two factors is a low level of interest among
private seed dealers in stocking rice seed (because of few repeat sales among farmers who
save seed from season to season) and the high cost (mainly transport costs) of rapid
substitution of old varieties for new. For this reason, the broad-based improvement of
rice seed systems is more amenable to methods that rely on gradual diffusion of small
quantities through highly decentralized seed networks.
A logical alternative to this method is that of the seed distribution campaigns which
rely on the public and NGO sectors. This, in turn, is dependent on well-informed
testing and evaluation methods typified by the Participatory Variety Selection (PVS)
methods discussed in Chapter 4. Collective learning from PVS methods in Guinea
recently led to major investments by the government in large-scale multiplication and
dissemination of improved, interspecific rice varieties (Spencer and Edwin, 1999).
Follow-on strategies of this type may be relevant in other rice-growing countries where
interspecific and other improved rice varieties have been widely tested, including Cte
dIvoire, Nigeria and Burkina Faso.
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1. Drought tolerance. Upland rice systems in Africa are subject to frequent periods of
drought stress, especially during vegetative phases of growth. To date, little research has
been aimed at improving the productivity of rice farmers faced with this constraint.
Nevertheless, recent reports of interspecific varieties with higher drought tolerance may
indicate potential for broad improvement for this trait. Researchers who have studied
the effects of moisture stress on rice have increasingly focused on insufficient root
penetration of the soil profile as a cause of the crops susceptibility to drought. Longer
roots which reach deeper into the soil profile are able to maintain growth further into a
drying period and may result in higher yields at the end of the season. Assaying for rooting is a difficult and time-consuming task that may be assisted by the identification of
molecular markers for this trait which can be used to select quickly for longer roots.
2. Increase national breeding capacity. Only five of the 12 West African countries with
major rice-growing areas actively breed new rice varieties. Extended breeding capacity
should be employed in breeding rice for a wider range of agro-ecologies, based on
combining resistance and tolerance traits for the major constraints in each area. A
well-developed network of national programmes is available to test and disseminate
improved varieties which come from such efforts.
3. Increased understanding of rice agro-ecologies. As previously discussed, breeding
teams can increase the efficiency of their targeting of new varieties and selection of
parents for new crosses through a better understanding of the character and boundaries
of rice agro-ecologies in Africa. At present, this type of study is confined to inner-valley
continuum ecologies, and generally does not involve breeding teams. Broadening the
focus of agro-ecological study to other areas and including breeders and other plant
scientists may make it more applicable to varietal development.
4. Rapid deployment of fixed and segregating lines with resistance to major pests and
diseases. This can be facilitated by continued fixing of new lines by the WARDA
interspecific breeding programme, and can be carried out in close conjunction with the
PVS method of selection and multiplication. Participatory variety selection needs to be
intensified to reach larger numbers of testing sites.
5. Increased capacity in molecular breeding at WARDA and selected national programmes.
Molecular reference maps for African rice are still being developed (Marie-Noelle
Ndjiondiop, personal communication), following which localization of the newly-developed marker for RYMV and other important traits can be performed. The point at
which integrating molecular breeding with conventional selection methods becomes
appropriate in NARS breeding programmes will depend on progress made in identifying
useful markers, analysis of the cost-effectiveness of the methods, and the pace of capacity
building at national level.
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II
Exploring New Strategies for

Improving Africas Food
Crops1
The following seven chapters are not intended as presentations of all the challenges facing crop
improvement specialists who work with these seven crops. Indeed, the same agro-ecological diversity
which this document has attempted to emphasize ensures that no single summary can adequately capture
all the issues which are of importance to breeders and farmers throughout Africa. Rather, the chapters
attempt to characterize the major challenges ahead in attempting broadly to improve the performance of
these crops when grown under marginal, low-input conditions common to small-scale farmers in much of
Africa. Therefore, rather than serving as detailed guides for priority setting among breeders, it is hoped that
this part will serve as a stimulus for further discussion and, eventually, increased support and efforts aimed
at addressing the identified challenges.
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12
Cowpea
12.1 Brief History of Cowpea Cultivation and Utilization in Africa

Cowpea is a tropical legume crop of African origin. Most recent speculation on the
crops centre of origin focuses on a band of diversity of wild cowpea stretching across
southern Africa from Namibia to Mozambique, with a centre of speciation in the
Transvaal region of South Africa (Padulosi and Ng, 1997). Nevertheless, the centre of
greatest diversity of cultivated cowpea is in the northern Guinea savannah regions of
West Africa (Ng, 1995). In many fields, almost continuous variation exists between the
more elite, large-seeded varieties of cultivated cowpea, the small-seeded, more weedy
varieties, and true wild species of cowpea (Rawal, 1975). Cultivated cowpea has been
shown to cross regularly with wild cowpea growing on the periphery of fields in East
Africa (Remy Pasquet, personal communication). Cowpea plant remains dating to
1500 BC have been discovered in a cave dwelling in Ghana (Flight, 1976).
Cowpea is an extremely resilient crop, and is cultivated under some of the most
extreme agricultural conditions in the world (see Plate 19). Cowpea varieties grown in
the Sahel and on the fringes of the Sahara are drought and heat tolerant. Other cultivars
are tolerant to acid soils, extremely poor soil fertility, and shading from other crops
(Singh, 1998). Cowpeas highly diverse plant architecture has allowed farmers to
develop varieties which fill a wide range of unique niches: highly determinate cowpea
varieties are grown for grain in monoculture situations, while spreading types are grown
as a dual-purpose (grain/fodder) crop interplanted with cereals, and as a relay crop using
residual moisture.
Cowpea is cultivated for its leaves, green pods, grain, and stover. While all parts of
the plant are used to some degree in each region of the continent, in West Africa cowpea
is primarily grown for its grain and stover (cowpea haulms contain 20% protein and are
highly sought after as cattle feed), while in eastern and southern Africa it is cultivated
primarily for its leaves. Cowpea grain is consumed directly following boiling, as a
component of meals which also include porridge made from cereals or root crops.
Cowpea grain cakes (made from mashed and fried seed) are also sold as a fast food along
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roadsides in Nigeria. In eastern and southern Africa, cowpea leaves are commonly added
to sauces and served with porridge, or boiled and consumed in a manner similar to
spinach.
12.2 Cowpea Production Levels and Trends in Africa

Data in Fig. 12.1 should be used only as an indication of the major cowpea producing
areas of Africa. Figures on cowpea production are imprecise, and for several important
producers (Mozambique, Zimbabwe), no recent data are available.
Current estimates place annual world cowpea grain production at 3 million tonnes
(Singh et al., 1997). Approximately 64% of this is grown in West and central Africa,
which accounts for 80% of total production in Africa. Nigeria, in turn, accounts for
upwards of 75% of production in West and central Africa (FAO, 1999). However, it is
also an important crop in marginal areas of eastern and southern Africa in Sudan,
Somalia, Mozambique and southern Zimbabwe. Most cowpea is grown as an intercrop
with cereals, and little of the harvest reaches regional markets. The sole important export
market for cowpea is believed to be from Niger to Nigeria, which is the worlds largest
consumer of cowpea.
Surveys have shown negative income elasticity for cowpea consumption, indicating
cowpea is a crop of the poor. Cowpea thrives on poor soils and in semi-arid regions,
making it a common intercrop with sorghum and millet-based farming systems of the
Sahelian countries. In many areas, cowpea is a crop cultivated by women, and is often
used as a weaning food.
Fig. 12.1.
Cowpea production trends in Africa, 19751999.
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Cowpea
12.3 Cowpea Production Constraints

Cowpea is attacked by over 35 diseases caused by viruses, bacteria, fungi and nematodes
(Singh et al., 1997), some of which cause significant reduction of yield. These are
grouped by pathogen, below.
Cowpea viral diseases
The most damaging viral diseases of cowpea are the seed-borne diseases (Hampton et al.,
1997). Their symptoms generally appear most strongly on the leaves, where they cause
stunting and deformation. Among these, several are of importance in Africa, including
cowpea yellow mosaic virus, cowpea aphid-borne mosaic virus and cowpea severe
mosaic virus. The genetics of resistance to cowpea viruses has been extensively studied
(Provvidenti, 1993; Scully and Federer, 1993), and numerous sources of resistance to
cowpea viruses have been identified.
Cowpea bacterial diseases
Bacterial blight caused by Xanthomonas campestris and bacterial pustule (Xanthomonas
sp.) are the two most important bacterial diseases of cowpea in Africa (Emechebe and
Florini, 1997). Bacterial blight is the most devastating disease of cowpea in dry regions
of West and central Africa (Wydra and Singh, 1998). Pathogens of both diseases are
transmitted via seed, and spread of the diseases is often caused by planting infested seed;
however, good sources of resistance have been identified for both diseases (Emechebe
and Shoyinka, 1985). Genes for resistance are available for most bacterial diseases, but,
their area specificity and frequent, sudden appearance in new areas illustrate the need for
decentralized breeding schemes for cowpea improvement.
Cowpea fungal diseases
Eleven major fungal diseases of cowpea have been identified among which anthracnose
(Colletotrichum gloeosporioides), Ascochyta blight (Ascochyta phaseolorum), brown blotch
(Colletotrichum capsici) and brown rust (Uromyces sp.) are considered to be of greatest
importance in Africa (Emechebe and Florini, 1997). In addition, Septoria leaf spots and
scab (Elsinoe phaseoli) are important constraints in more humid regions.
Cowpea insect pests
Insect pests represent the most serious constraint to cowpea production throughout
Africa. In many areas, losses due to insect pests are so high that yields seldom rise above
100150 kg ha1 (Kitch et al., 1997). The most important insect pests in cowpea
production systems in Africa are probably aphids (Aphis craccivora), pod borers
(Maruca vitrata), thrips (Megalurothrips sjostedti) and the pod-sucking bug Clavigralla
tomentosicollis. Aphids are most damaging in the Sahel in areas of less than 300 mm
rainfall, while maruca is most important in higher rainfall areas (IITA, 1998).
Unfortunately, cultivated cowpea genome appears to offer few useful sources of durable
resistance to the major insects. Although several wild species are known to be resistant to
maruca pod borers and the pod-sucking complex, and wide crosses made to wild
relatives V. oblongifolia and V. lutea have produced fertile progeny using embryo rescue,
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the results have been mixed (Fatokun et al., 1997). Cowpea bruchids (Callosobruchus
maculatus and Bruchidius atrolineatus) cause extensive damage to cowpea grain in
storage, but actually infest the green pods while still in the field (Murdock et al., 1997).
Much effort has been made to devise and popularize appropriate methods of protecting
against damage to cowpea grain in storage, sometimes with positive results (Murdock
et al., 1997). Nevertheless, bruchids continue to destroy much of the crop throughout
Africa.
Striga
Striga is an important constraint to cowpea production in much of West Africa.
Varieties have been developed which show high levels of resistance to Striga populations
of Nigeria, Burkina Faso, Cameroon and Mali, but which are susceptible to populations
in Benin. Other varieties are resistant to the Benin populations but susceptible to others.
Work is on-going to develop varieties which have broad resistance against all known
Striga populations (IITA, 1998).
12.4 Cowpea Improvement Through Biotechnology and Breeding

Cowpea breeding advances in Africa
The large number of biological constraints encountered in cowpea cultivation in Africa,
coupled with wide natural diversity within the cowpea genome, offer many channels for
improvement of the crop. However, because cowpea is used primarily as a niche crop
by small-scale farmers, successful breeding requires extensive knowledge of local farming
systems. Kitch et al. (1998) analysed varietal preferences among men and women
farmers in Cameroon and identified 26 different criteria used in making selections,
of which less than half were related to yield. The need for this type of detailed
knowledge argues for decentralized breeding initiatives based on increased analysis of
agro-ecological variation and farmer preferences, as the rate of usage of varieties bred
internationally and regionally is very low (Ndiaga Cisse, personal communication).
Nevertheless, national breeding efforts can be effectively reinforced by efforts among
international breeding centres which focus on intractable constraints to production and
longer-term population improvement for yield potential.
IITA has the worldwide mandate for cowpea breeding, and maintains a breeding
centre in Kano, Nigeria. The USAID-funded Bean/Cowpea Collaborative Research
Support Programme is also active in breeding cowpea varieties for northern Sahel and
savannah ecologies focusing on resistance to important pests and diseases (Ehlers and
Hall, 1997). Unfortunately, only a few national programmes (Burkina Faso, Mali,
Ghana, Nigeria and Senegal) have employed cowpea breeders who perform their own
crosses. In some cases, cowpea breeders have been drawn into administrative positions,
and in others, there is a lack of funding channelled to their programmes from either
national or international sources to permit planting a fully functional nursery on a regular basis. This deficiency will need to be corrected if crop improvement in cowpea is to
have serious impact among small-scale farmers. Cowpea varieties are not, as a rule,
broadly adapted and the value of even highly prized plant genetic traits can be masked by
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Cowpea
lack of adaptation or farmer preferences if those traits are not effectively introgressed
into the appropriate background.
The IITA programme has developed varieties for use under two major categories:
time-to-maturity (early, 6070 day; medium, 7590 day; late, 85120 day) and photoperiod sensitivity (non-sensitive and sensitive). In addition, they have developed
speciality varieties with high grain quality, high foliage production and tolerance of
environmental stresses (Singh et al., 1997). These varieties vary with respect to the
farming systems they are adapted to (intercropping compared with mono-cropping).
With increasing time to maturity, plant growth habit also changes from highly
determinate, bush types to indeterminate, vine types. A separate effort has been directed
at transferring useful traits such as early maturity and resistance to thrips, aphids, viruses
and bruchids into popular landraces of West and central Africa (IITA, 1998)
Another major focus has been the development of varieties with multiple virus
resistance and multiple bacterial disease resistance. At present, a series of varieties has
resistance to five major cowpea viruses. Varieties IT96D-660 and IT96D794 are
resistant to cowpea aphid-borne mosaic virus, blackeye cowpea mosaic virus and cowpea
mosaic virus (Hughes and Singh, 1998). Sources of resistance are also available for
fungal disease pathogens, including anthracnose, Cercospora phytophtera, brown blotch
and leaf smut. Varieties are also available with resistance to root knot nematodes.
Extensive efforts have also been made to develop resistance to insects, with
encouraging results for some insect pests (Singh et al., 1997) (see Table 12.1). Resistance
has now been reported to be available against attack by thrips, bruchids and aphids
(Adjadi et al., 1985; Singh, 1993). As has been previously stated, however, adequate
resistance to maruca and pod-sucking bugs has not been achieved, despite exhaustive
screening of the cultivated cowpea genome.
Drought and heat tolerant cowpea varieties, named Mouride and Melakh, have
been developed for water-limited environments of the Sahel (Cisse et al., 1995, 1997)
through collaborative research between US universities and NARS in Africa (Hall et al.,
1997). These extra-early varieties proved highly popular among Senegalese cowpea
farmers who commercialized their harvest.
Table 12.1. Sources of resistance to major pests and diseases of cowpea developed by IITA (after Singh et al., 1997)
Striga
Variety
IT90K-277-2
IT89KD-374-57
IT90K-261-3
IT89KD-457
TVX 3236
IT90K-76
IT90K-59
IT82D-889
IT83S-818
Five virus
diseases
Bacterial
diseases
Bruchids Aphids
R
R
R/MR
MR/R
R/R
R
R
R
R
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MR
R
R
Thrips
R
MR
MR
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Striga and Alectra both attack cowpea throughout Africa. Sources of resistance have
been identified and the genetics of resistance has been studied (Aggarwal, 1984; Singh
and Emechebe, 1990). Varieties with resistance to Striga have been distributed
to numerous African countries. Cowpea could prove to be an important source of
resistance to Striga for other crop species, through genetic transformation.
Some of the most recently developed, improved cowpea varieties combining
resistance to major diseases, insect pests, and Striga gesnerioides have shown over 50%
higher yield potential than existing improved varieties, with 1.5 t ha1 grain and 3 t ha1
of fodder in Sahelian ecologies and 3 t ha1 grain and 5 t ha1 fodder in Sudan savannah
ecologies. Screening for drought tolerance and root characteristics revealed that four
varieties, IT96D-604, IT95K-222-3, IT90K-222-5 and IT95K-1115-10 were most
drought tolerant (IITA, 1998).
Advances in cowpea biotechnology

Biotechnology may hold significant promise for cowpea improvement. Cowpea yields
are so highly affected by insect pests that five- to seven-fold increases are experienced
with one or two applications of insecticide. Maruca pod borers and the pod-sucking
complex of insects are two pests for which no adequate source of genetic resistance has
yet been discovered. Cowpea weevil is another for which known resistance sources offer
only slight improvements.
Several researchers in the USA have worked on transformation systems for cowpea,
using both Agrobacterium and the gene gun methods. IITA, as well, has worked on
cowpea transformation via electroporation of pollen grains, microparticle bombardment
of pollen grains and Agrobacterium transformation of immature flower buds (Thotapilly
et al., 1998). Plants were regenerated from tissue which tested positive for reporter
genes, but which lost expression in the T2 generation. To date, cowpea transformation
has not become a routine procedure. At the time of writing of this book, researchers were
planning to convene a meeting on cowpea biotechnology aimed at establishing a global
strategy for biotechnology applications in Africa. Bt cowpea has been proposed as a
means of achieving resistance to cowpea pod borers, as well as via transformation using
protease inhibitors, -amylase inhibitors and lectins (Monti et al., 1997). At the time of
publication of this book, researchers from a number of institutes in Africa, the USA and
Australia were planning to embark on an ambitious plan to transform African cowpea
using gene constructs encoding Bt and -amylase inhibitor proteins, which are expected
to offer protection against maruca pod borers and cowpea bruchids, respectively. A separate initiative was also underway to transfer a gene construct for resistance to cowpea
aphid-borne mosaic virus into cowpea using novel transformation methods
(Sithole-Niang, 2000).
IITA has also made progress on development of an RFLP map of cowpea using a
cross between cowpea and a wild relative. At the last report, the cowpea map had 92
markers distributed among 85 loci (Fatokun et al., 1997). Average genomic distance
between markers is approximately 10 cM. These markers are distributed into ten linkage
groups; however, cowpea has 11 chromosomes. Since probes used for cowpea mapping
were the same as those used for mapping mung bean, synteny studies have been possible
between cowpea and other legumes. A high degree of conservation was observed, with
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Cowpea
90% of markers hybridizing in similar positions in the two crops. Useful traits for which
putative QTLs have been identified include seed weight, pod number, pod length, plant
height, days to flowering, and days to maturity (Fatokun et al., 1997), while current
efforts are focused on identifying markers for resistance or tolerance to bruchids and
thrips. Timko (2001, unpublished) has since reported identifying 413 RAPD markers
on the cowpea genome, distributed among 11 linkage groups.
12.5 Cowpea Seed Systems

Very little commercial seed of cowpea is marketed in Africa, although seed companies
in Mozambique, Zimbabwe and Ghana do market registered varieties. Most seed is
obtained through informal exchange between farmers, with some seed being purchased
in local markets (Walker and Tripp, 1997). As a self-pollinated, non-commercial
crop, cowpea seed dissemination is suited to public sector-led campaigns which
may focus on increasing access among farmers to a single variety or group of selected
varieties. Significant impact may be possible through NARS/NGO collaborations which
focus on broad testing and dissemination of farmer-selected varieties focusing on the
drought-resistant and pest- and disease-resistant varieties indicated in the above section
on breeding.
Recently, a seed distribution initiative was undertaken in northern Nigeria which
appears to show promise for other areas, as well. Thirty-six experienced cowpea growers
were given 3 kg of breeder seed to grow simultaneously as foundation seed and as a
demonstration plot. They in turn sold seed on to 262 farmers who showed interest in
the varieties. This group produced some 12 t of seed for sale to hundreds more farmers
(Singh and Olufano, 1998).

1. National cowpea breeding programmes. Since cowpea varieties must conform to
localized, low-input farming systems in order to be adopted, wide adaptation is not
common. All valuable genetic traits identified and rendered manageable through
biotechnology or breeding will inevitably have to be introgressed into adapted populations at the national level. With only five NARSs currently active in this work, the
prospects for broad improvement of cowpea in Africa are poor.
2. Insect resistance. Insect pests (thrips, bruchids, pod-sucking insects) constitute the
greatest constraint to cowpea production in Africa. Little or no usable resistance has as
yet been detected in cultivated or wild genomes. Strategies have been advanced to
transform African cowpea with gene constructs encoding production of Bt and amylase inhibitor proteins. Recently, research conducted at IITA showed that orchid
and snowdrop lectins were found to be insecticidal to Maruca vitrata and hence may be
used to control this pest through transgenic approaches. In addition, affinity-purified
lectins from African yam beans (Sphenostylis stenocarpa) were tested against pod-sucking
bugs and cowpea weevils using an artificial seed system, and were demonstrated to be
lethal to both pests.
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3. Transformation and gene expression systems. Rates of success with genetic transformation of cowpea to date (if achieved at all) are far below levels of efficiency needed to
plan an improvement strategy. Cowpea transformation is being pursued at IITA, Michigan State University, the University of Arkansas, in China, and India. Researchers at the
University of California at San Diego have discovered a promoter which constitutively
expresses in legume seeds. This could be useful in the enhancing expression of resistance
genes to weevils, thrips and maruca pod borers.
4. Identification of resistance genes. Little usable resistance has been found against
maruca pod-boring or pod-sucking insects. Cowpea bruchid resistance has also been
extremely difficult to isolate and manage. A systematic procedure has been developed at
Purdue University for identification of resistance genes (Larry Murdoch, personal
communication). Genes have already been isolated from common beans for -amylase
inhibitor for bruchid resistance.
5. Improved nutritional character. Although cowpea is already a good source of protein
and carbohydrate, food quality analysis of some 52 cowpea varieties at IITA recently
indicated significant genetic variability for protein, fat, and iron content. The top four
improved varieties had 17% higher protein and 12% higher iron content than the mean
of four popular local varieties (IITA, 1998).
6. Gene flow studies. Because cowpea is an indigenous crop to Africa, there is a high
probability that transgenic resistance genes could flow to wild relatives growing in the
borders of fields. As a measure of biosafety, studies need to be undertaken to measure the
likelihood and degree of risk involved.
7. Virus resistance. Gene constructs have been developed for resistance to a number of
viruses (Mlotshwa, 2000). At present, several of these have been shown to be effective in
model systems, using tobacco (Ida Sithole, personal communication). These could
prove useful in a cowpea transformation system.
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Cassava
13.1 Brief History of Cassava Cultivation and Utilization in Africa

Although still a subject of some debate, the centre of origin of cassava is generally
believed to be the southern border of the Amazon basin (Olsen and Schaal, 1999).
Cassava was introduced in Africa in the Congo River delta by the Portuguese in the 15th
century (Jones, 1959), and spread rapidly to many agro-ecologies (Hahn et al., 1979).
Cassava is today grown in most agro-ecologies of the continent; however, cassava is most
important in farming systems of the humid forest regions, where the productivity of
grain crops is reduced by low sunlight, foliar pests and diseases, and grain storage is more
difficult. Cassava has very high yield potential, making it a viable alternative to grain
crops where population pressures have led to tradeoffs between food quality and
quantity. Commercial cassava yields as high as 20 t ha1 have been registered under
experimental conditions. However, because of high labour requirements at planting and
harvest, cassava production throughout the world continues to be dominated by smallscale, non-mechanized systems.
Cassava is well known for being able to grow and produce food even in very poor
soils. For that reason, it is often grown at the margins of farms while the better land
is reserved for the production of grain crops. In addition, once established, cassava is
relatively drought tolerant, and when mature can survive up to 6 months without rains.
Cassavas ability to produce food under marginal conditions has made it a popular crop
of Africas poor farmers who are unable to invest in fertilizer or pesticides to protect the
crop against environmental stresses and biotic constraints. This fact, coupled with
asexual propagation of the crop, has created a major role for crop improvement not
only are there few other alternatives to building in the performance traits needed by
farmers, but, once finished, there is a very good chance the crop will stably express those
traits (see Plate 20).
Cassava is widely consumed as a porridge, which is prepared from dried and
pounded roots, but is eaten in a very wide range of forms in different parts of the
continent. Cassava is reported to be consumed in 28 different forms in Cameroon, alone
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(Jim Whyte, personal communication). In urban areas of West Africa, widespread development of cassava-processing methods (consisting of pounding, soaking and drying to
produce a fermented flake known as gari) have resulted in cassava becoming an important commercial commodity. Such processing capacity does not exist in east and southern Africa, and cassava has remained a traditional, rural starchy staple in those regions.
Cassava is also consumed as a snack food in various parts of the continent. Varieties
used as snack food are sweet types, low in cyanic acid, which can be boiled and eaten
or even consumed raw. Rapidly increasing cassava cultivation in Sahelian countries
over the past decade has been primarily based on the use of these types (Tshiunza
et al., 1999). Cassava is also widely grown for its leaves, which are used in making
sauces. Once again, leaves from varieties with high cyanic acid content must be properly
processed to remove the toxic compounds. Cassava flour is also sometimes used in making bread for local consumption. Recently, initiatives in West Africa have aimed at
developing the export market potential for production of dried cassava chips used as
animal feed in Europe. This market is currently supplied by Asian production.
Cassavas combined abilities to produce high yields under poor conditions and store
its harvestable portion underground until needed make it a classic food security crop.
In recent years, this has proved of critical importance to many people in Africa caught up
in civil conflicts and unable to cultivate the normal range of annual crops. Displaced
groups of people in Mozambique during that countrys 16-year war often survived on
abandoned cassava fields. Because it is a vegetatively propagated crop, such plantings can
also serve as a ready supply of planting material during rehabilitation following conflict
or drought. It is a notable fact that cassava processing and marketing are often controlled
by women. Thus, resources from cassava production are often targeted toward the needs
of women and children.
As implied above, however, cassavas productive capacity in low-input conditions
comes at a certain cost in terms of carbohydrate quality and protein concentration.
13.2 Cassava Production Levels and Trends in Africa

While aggregate production statistics on cassava are subject to large degrees of error,
the figures in Table 13.1 give a general idea of the trend in cassava production in
Africa. According to figures from the FAO (1999), the rate of increase of cassava
production has been higher than any other crop in Africa over the past 15 years.
Since 1990, this increase has been fuelled by rapid increases in productivity following the release of improved varieties in Nigeria (Nweke et al., 1994), and more recently,
in the Sahel (Tshiunza et al., 1999). Rapid increases in cassava production occurred in
West Africa (primarily Nigeria) following the release of high-yielding, early-bulking
varieties and the establishment of small-scale processing facilities (Fig. 13.1).
13.3 Cassava Production Constraints

At the time of cassavas introduction, few major pests or diseases hindered its
production. With time, however, a number of important biological constraints to production have been introduced. Thus, while cassava may tolerate well extensive farm
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Cassava
Table 13.1. Annual cassava production in major African

cassava-producing countries.a
Production (t year1)
Country
Benin
Cte dIvoire
Ghana
Guinea
Nigeria
Kenya
Madagascar
Uganda
Angola
Congo, Democratic Republic of
Mozambique
2,377,339
1,700,000
7,226,900
811,869
32,695,000
910,000
2,404,000
3,400,000
3,210,570
17,100,000
5,639,000
aIn comparing cassava production figures with those of grain

crops it should be borne in mind that cassava production figures
are reported at 70% moisture content, while most grain crops are
reported at approximately 15% moisture content.
Source: FAO (1999).
Fig. 13.1.
Cassava production trends in Africa, 19751999.
management practices commonly used in its production in Africa, cassava yields are
severely affected by pests and diseases (IITA, 1998). Chief among these are the two
insect pests, cassava green mite (CGM) (Mononychellus tanajoa) and cassava mealy bug
(Phenacoccus manihoti), and two foliar diseases, cassava mosaic disease (also known as
African cassava mosaic virus) (ACMV) and cassava bacterial blight (CBB) (Xanthomonas
campestris). At present, a heavy incidence of cassava brown streak disease has also been
reported in Mozambique (Alfred Dixon, personal communication).
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ACMV and its East African variant EACMV are caused by virus strains which can
be transmitted both mechanically, through handling of planting stock, and via an insect
vector, the white fly (Bemisia tabaci) (Storey and Nichols, 1938). Recently, an especially
virulent form of cassava mosaic disease (now being referred to as Ugandan variant, or
UgV) spread through Uganda, western Kenya, southern Sudan and northern Tanzania
(Legg, 1999). ACMV is endemic in Africa. Cassava bacterial blight was probably
introduced to Africa from South America by accident. It has now been reported
throughout Africa. Root rots, caused by Botryodiplodia theobromae, Fusarium spp. and
Phytophtora sp. have recently been reported to be increasing in Nigeria (Wydra and
Singh, 1998).
Among the insect pests of cassava in Africa, cassava mealy bug and green mite are
considered to be the most important. Both of these were introduced to Africa through
importation of vegetative planting material (Hahn et al., 1979). While cassava mealy
bug infestations have been successfully controlled by distribution of its natural enemy
from South America, the green mite remains a serious constraint to production in parts
of Africa, especially during dry periods, although African landraces have been identified
which carry resistance (see Plate 21). IITA is also pursuing an extensive campaign
on biological control of cassava green mite using an exotic phytoseiid predator,
Typhlodromalus aripo.
13.4 Cassava Improvement Through Biotechnology and Breeding

Cassava breeding advances in Africa
Because of the limited diversity among the introductions made by the Portuguese and
because of infrequent sexual reproduction of the plant in most African environments,
the African cassava gene pool has remained relatively narrow. This lack of genetic
diversity, however, appears to be a constraint within the whole of the cultivated cassava
genome. Olsen et al. (1999) discovered that cultivated cassava contained only 25% of
the diversity of its wild progenitors, compared with 75% for maize.
Cassava is a cross-pollinated crop. However, breeding has historically been limited
by the plants erratic flowering habit. Breeding of cassava in Africa achieved a major
advance with the discovery in 1989 of a site in Ubiaja, Edo State, Nigeria where a broad
base of cassava accessions produced flowers and could be used in making crosses
(Ekanayake, 1996). The development of this facility has led to a drastic increase in
the number and diversity of genetic backgrounds that can be incorporated into new
varieties, both from African landraces and from breeding stock from Latin America (via
CIAT) and other parts of the world. IITA breeders have now developed many new
breeding populations, with their progeny being sent out both as seed and as micropropagated plantlets to many national programmes. Because of long time-to-flower and
severe inbreeding depression in cassava, breeders commonly rely on screening very large
F1 populations, in the hope of finding genotypes which combine a favourable mix
of target traits. More recently, however, inbreeding methods have met with more
success, and fifth-generation inbreds have been developed (Alfred Dixon, personal
communication).
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Resistance to both ACMV and CBB were discovered in a wild relative of cultivated
cassava, Manihot glaziovii, and successfully transferred into cultivated cassava via wide
crosses (Nichols, 1947). This led to the development of a donor parent, clone 58308,
which was later recognized as stably resistant to disease and widely used as a source of
resistance to both diseases in the cassava breeding programme at IITA. Since then,
breakthroughs in breeding have permitted the development of resistant varieties such as
TMS 30372. Indeed, perhaps the most impressive results of cassava improvement,
Africa-wide, have occurred in the rate of development of ACMV-resistant varieties,
whose number increased from 29 in 1989 to 106 in 1993 and 334 in 1997. Similarly,
the number of ACMV-resistant plantlets distributed to collaborating programmes,
Africa-wide, has risen from 3007 in 1993 to 16,864 in 1997. Recently, African landraces
have been identified which carry high levels of resistance to ACMV (Dixon, 1999).
In spite of such advances, cassava breeding remains a highly centralized activity, with
nearly all breeding being conducted at a single site, and other cassava improvement
programmes being relegated to testing. In view of continued low adoption rates of
improved cassava, more decentralized agro-ecology-based breeding activity is viewed as a
priority.
Cassava breeding methodology is based on crossing devised to combine traits of
various parents, followed by clonal selection based on performance for various target
traits. A current priority for breeding in East Africa, for example, is combining high levels of resistance to ACMV and CGM in several adapted clones as these two constraints
seem to result in especially heavy losses when found in combination (Legg et al., 1998).
The IITA methodology begins with crossing performed at Ubiaja, followed by
preliminary evaluations for ACMV, CBB and green mite resistance in Ibadan. Breeding
at IITA has led to the development of a wider range of disease- and insect-resistant
varieties with desirable agronomic (early bulking, high yield, low branching, among
others) and culinary traits (easy processing and desired cyanic acid content) with
adaptation to most of the cassava-producing agro-ecologies in Africa (Dixon, 1999).
This was followed with the massive dissemination of seeds and tissue-cultured plantlets
for further selection for adaptation by NARSs. A total of 38,811 in vitro plantlets were
distributed to 39 African countries between 1994 and 1997. In addition, almost
2.4 million sexual seeds were distributed to various African countries during the same
period (Lynam, 1998). During the 1990s the cassava breeding programme at IITA has
also imported large numbers of accessions from Latin America, aimed at broadening the
gene pool.
Two networks organized around breeding for root crops have been created in East
(EARNET) and southern Africa (SARNET). No such network exists at present for West
and central Africa.
Advances in cassava biotechnology

By far the most commonly employed application of biotechnology in cassava has been
micropropagation via in vitro meristem culture. Perfection of these techniques at IITA
and several national programmes has facilitated wider distribution of disease-free germplasm of a greater range of diversity across Africa than could have been achieved through
distribution of cuttings. Micropropagation has been extensively employed in the
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distribution of cassava cuttings in Burundi following disruption of farming systems

during the conflict there (DeVries, 1999).
Genetic engineering of cassava has been constrained by the lack of a reproducible
system for generating transgenic plants. Recently, however, regeneration of plants from
undifferentiated callus tissue has been achieved using suspension culture techniques
(Taylor et al., 1996). These cultures were then used in regenerating transgenic cassava
plants via microparticle bombardment of the culture medium (Schopke et al., 1996).
Hong-Qing Li et al. (1996) have also reported the development of transgenic plants via
transformation using Agrobacterium. Stable expression of agronomic genes in transformed plants, however, has proved difficult to achieve. Genetic modification of cassava
for modified starch composition has been achieved by researchers at Wageningen,
Netherlands, and field testing is expected to begin soon (Johanson and Ives, 2000).
Researchers at Ohio State University have also developed transgenic cassava with
reduced cyanide toxicity (Johanson and Ives, 2000).
Biotechnology applications for cassava have been delayed in large part by the overall
lack of information on cassava genetics, caused by the low level of investment and the
biological barriers to genetic dissection mentioned above. In addition, cassava is believed
to be an allopolyploid, with 36 somatic chromosomes. For this reason, it has long been
difficult to distinguish between true heterozygosity and duplicated genes in various parts
of the genome (Lefevre and Charrier, 1993).
In spite of these complexities, cassava has a relatively small genome, and this has
facilitated the development of genetic maps using molecular markers. A genetic linkage
map for cassava has been constructed using 132 RFLP markers, 30 RAPDs, three
microsatellite markers, and three isoenzyme markers (Fregene et al., 1996). This map
is based on an F1 population from a cross between elite varieties from Nigeria and
Columbia. In addition, Chavariagga-Aguirre et al. (1998) have identified 32 microsatellite markers, for which 22 primers have been developed.
Since 1997, CIAT has been developing PCR-based markers for the cassava genome.
Over 1000 putative simple sequence repeats (SSRs) have been identified, and primers
have been developed for over 60 of these markers. The aim of the project is to identify
several hundred SSR markers in order to saturate the genome map. The same group has
developed expressed sequence tags (ESTs) from approximately 250 polymorphic cDNA
sequences. QTLs have been identified for useful traits such as early bulking, postharvest
deterioration, starch quality and content, and morphological traits (CIAT, 1998).
Since 1996, researchers at CIAT and IITA have collaborated on the mapping of genes
for resistance to ACMV (CIAT/IITA, 1999). Two F1 mapping populations with different
sources of resistance to the disease were analysed using 186 SSR markers. Using bulk
segregant analysis, a single gene, believed to be from a chromosomal segment of glaziovii
origin, was identified as CSY1. Future phases of the project are focusing on map-based
cloning of the resistance gene, molecular characterization of virus strains, and markerassisted introgression of the resistance gene into Latin American cassava populations.
13.5 Principal Challenges for Cassava Improvement in Africa

One of the most significant challenges associated with cassava improvement is its high
level of genotype environment interactions. Breeding programmes, therefore, must
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continually develop large numbers of new genotypes for testing in a wide range of
environments (Dixon et al., 1996). This constraint also carries obvious implications
favouring the decentralization of breeding programmes. However, although the two
crops are of similar importance in Africa, the number of NARSs with breeding
programmes for cassava is roughly one-fifth the number of those with maize breeding
programmes.
While the difficulties associated with cassava breeding would naturally tend to set
limits on the number of institutes actively engaged in breeding, it seems clear that more
countries should have such programmes, and that decentralization of the regional
breeding programme carried out by IITA is a priority challenge for the future. In fact,
national programme scientist involvement in breeding has been part of the IITA
programme in Uganda since 1991; however, developing improved varieties with specific
adaptation to mid-altitude ecologies will require that a greater number of NARSs
become involved. While population development for West Africa is at far more
advanced stages within the IITA programme, decentralization and greater involvement
of national scientists and farmers in the breeding and selection process is relevant in this
region, as well.
Cassava is an ideal crop for participatory approaches to crop improvement in
that significant variability can be maintained at early evaluation stages and yet lines
are fixed clonally after the cross. Such approaches are best developed at the level
of the national programmes, since local farmer preferences can be introduced into
the evaluation process. However, systematic methods need to be developed and
evaluated before integrating participatory approaches into network and NARSs
activities.
The IITA cassava breeding programme for West and central Africa has been quite
successful in developing populations for the different ecologies of that region. Cassava
varieties for mid-altitude zones of East and central Africa have been drawn largely from
these populations, in spite of important differences which exist between the ecologies.
Meanwhile, it has been postulated that Latin American cassava populations adapted to
higher altitude zones of that continent may have significant adaptation advantages in
East and southern Africa. Therefore, development of new populations in east and
southern Africa is viewed as a major future challenge.
In a more general sense, cassavas status as a standby, food security crop in much of
eastern Africa would appear to have reduced adoption rates of improved varieties with
higher productivity potential (see Plate 22). Areas of Africa where adoption of improved
varieties have been highest (generally thought to be Nigeria and Ghana) are those
where processing technology is also most developed and widespread. Improvement
programmes, therefore, may do well to pay close attention to opportunities for commercialization and processing of the crop.
13.6 Cassava Seed Systems

Cassava germplasm is not disseminated via commercial channels in Africa. This
constraint, coupled with slow and very laborious methods of multiplication via cuttings
have severely constrained diffusion of improved germplasm in many parts of the
continent.
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Cassava dissemination has often depended on campaign approaches which aim at

distributing large quantities of improved planting stock via specially funded initiatives.
This approach has been employed as a response to the disruption of agricultural systems
as a result of conflict in Mozambique, Angola, Liberia, Sudan, Burundi and various
other countries. In Uganda and western Kenya, large-scale multiplication initiatives
have recently been implemented in response to the outbreak of new-variant ACMV in
those areas. In the case of Angola, tissue-cultured plantlets of over 500 accessions of
improved cassava were transferred directly from the IITA laboratory to improvised
hardening-off facilities in production areas before being transferred to multiplication
beds. The impact of these projects is primarily related to the rapid transfer of varieties
with disease resistance in areas which had not previously been reached.
Using rapid multiplication techniques, these projects have increased the rate at
which farmers can now receive improved germplasm. In western Kenya, ratios (in area
terms) of seed field:planting field of 1:10 have been employed, with primary and
secondary seed fields designated for production of initial increases and production of
planting stock for farmers, respectively (Onim, 1998). In this case, primary multiplication sites were located on research stations, while secondary sites were located on
rented land or farmers fields.
Recently, national programmes have been very instrumental in large-scale multiplication and distribution of improved germplasm. Three high-yielding varieties have
been released in Ghana and are being multiplied by the Crops Research Institute, MOA,
and various NGOs. In response to the recent new-variant ACMV epidemic, the national
programme in Uganda has released four IITA-breed varieties which are already being
grown on 80,000 ha. As of late 1998, a total of 1.3 million seedlings were in multiplication in Serere and Namulonge, Uganda and Mtwapa, Kenya. Multiplication
and distribution of ACMV-resistant varieties is underway in Nigeria, Sierra Leone,
Angola, Malawi, Mozambique, Namibia, Tanzania, Zambia, Zimbabwe, Swaziland,
Lesotho, Togo, Benin, Guinea and the Gambia, and in most EARNET countries,
including Kenya, Uganda, Rwanda and Madagascar.
One of the more difficult challenges facing cassava improvement programmes in
Africa is the development of effective methods of multiplication and distribution of
clean planting material. Since cassava planting material is bulky and the rate of multiplication is slow, innovative methods for seed multiplication and distribution need to be
developed. The development of a sustainable seed system will need to be built around a
steady flow of new varieties, a primary multiplication point, and a distributed set of
secondary and tertiary multiplication points.
Significant progress has been made in in vitro multiplication techniques and
post-flask management of plantlets, but there is still a need to improve further plant
establishment under different agro-ecological conditions, especially in more harsh
environments, and to reduce the costs involved. Tissue culture capacity in Africa is both
limited and often inefficient in handling large-scale multiplication. Existing capacity is
primarily devoted to quarantine and research activities, and has limited experience in
larger scales of operation. The role of tissue culture in cassava multiplication in Africa
remains questionable, requiring some evaluation of organization and economic costs.
If tissue culture is not the primary multiplication point for planting material, then
cost-effective alternatives need to be developed.
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Cassava

A review of the overall strategy for development of cassava in Africa reveals two principal
elements, namely, reinforcing cassavas role as a food security crop in existing rural
production and consumption systems and developing cassavas market and income
generation potential in areas where markets exist. Likewise, the review points toward
two key elements to the development of a cassava improvement strategy for Africa. First
is that the target is uniquely smallholder systems. Second, a research strategy to expand
cassava markets and develop the crops commercial potential should at the same time
reinforce cassavas role as a food security crop. Increased productivity, linked to more
efficient processing methods, could increase farmers incomes at the same time as
improving the security of the subsistence food base.
Breeding targets are both a function of genotypic adaptation to biotic and abiotic
stressesamalgamated in the concept of agro-ecology and a function of root and
leaf quality characteristics, determined by end use and product markets. Breeding is
structured in terms of agro-ecologies, but overall targets are an overlay of agro-ecologies
and product markets, as the two are not congruent.
1. Broad introduction of early-bulking, stress-resistant varieties. An estimated 80% of
Africas cassava harvest continues to come from late-bulking, unimproved landraces.
Therefore, a major challenge remains in the area of breeding and participatory variety
selection programmes to increase exposure of farmers to improved varieties with higher
productivity potential and other, yield-stabilizing traits such as resistance to ACMV and
CBB.
2. Improved nutritional characteristics. Given cassavas relatively poor quality carbohydrate and low protein contents, coupled with its increasing popularity among very
poor farmers of marginal zones in Africa, it is clear that an emphasis should be placed on
key aspects of its nutritional profile. Increased iron, zinc, and vitamin A precursor have
been cited as potential targets for micronutrient enhancement (Alfred Dixon, personal
communication).
3. Characterization and targeting of cassava breeding environments. Little has been done
to understand better the factors which contribute to good adaptation. Yet the potential
impact of improved cassava is often hampered by low uptake by farmers. The best
approach to take in regard to priority is to describe common areas where cassava is
or is rapidly becoming an important crop, and study the major constraints and
characteristics of its growth and utilization.
4. Decentralization of cassava breeding programmes. At present, virtually all crossing is
being done at a single site in Nigeria, while cassava improvement at a national level is
relegated to screening large numbers of F1 seed. Enabling national programmes to
perform their own crosses based on information gathered on agro-ecologies and
knowledge of the combining ability of parental clones may unlock new opportunities
for uptake of cassava at a local level. Linked to this, uptake of improved varieties
may also be improved by involving farmers in selection processes at an earlier stage in
their development. Participatory selection methods should therefore accompany this
initiative.
5. Pest and disease resistance. Linked to the above, a determined focus on reducing
losses from important pests and diseases is a clear priority. ACMV, cassava green mite,
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and CBB are observed to cause damage to the harvest in large areas of Africa. Of critical
importance at this time is responding to the serious outbreak of the UgV variant of
ACMV in east Africa (through rapid dissemination of adapted, resistant clones) and the
outbreak of cassava brown streak disease in northern Mozambique.
6. Population improvement for mid-altitude agro-ecologies. Cassava is an increasingly
important crop in mid-altitude environments of eastern and southern Africa. Yet the
range of improved materials adapted to these ecologies is still limited.
7. Improved root quality characteristics. As industrial-level cassava processing becomes
more common, demand for speciality varieties with high starch, longer preservation, or
high soluble sugars is expected to rise. Recent evidence suggests that marker-assisted
selection may be effective in selecting for these traits. Meanwhile, root rot, associated
with poor soil fertility, is reported to be an increasing problem in parts of Nigeria.
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Banana
14.1 Brief History of Banana Cultivation and Utilization in Africa

Plantain and bananas serve as important food crops in much of Africa. Together they
provide more than 25% of carbohydrate needs of over 70 million people on the
continent (IITA, 1998). Cultivated bananas are derived from two species of the genus
Musa, M. acuminata and M. balbisiana (Stover and Simmonds, 1987). M. acuminata
originates in Malaysia, while M. balbisiana originates in India (Simmonds, 1966).
Bananas cultivated in Africa are diploid and triploid genetic combinations of A and B
genomes contributed by one or both of these species.
African bananas are grouped into three categories, including East African (mainly
dessert) bananas (AA, AAA, ABB, and AB), the African plantain bananas (AAB) grown
mainly in central and West Africa, and the East African Highland banana (AAA), used
for cooking and in beer preparation (Karamura, 1998). Although not of African origin,
African bananas have evolved into an important zone of secondary genetic diversity. In
particular, the lowland regions of West Africa contain the worlds largest range of
genetic diversity in plantain, while the highlands of East Africa are an important centre
of diversity of cooking bananas (Ortiz and Vuylsteke, 1994).
Banana is a clonally propagated plant. Triploid genotypes are virtually or
completely sterile and develop their fruit through vegetative parthenocarpy. Diploid
landraces and tetraploid cultivars (mostly artificial hybrids) are also cultivated.
Commercial production of banana and plantain is characterized by the use of a very
limited number of varieties. Cavendish, for example, is currently the most widely
cultivated variety of dessert banana, and is grown throughout the world, while Cuerno
(Horn) is a widely cultivated variety of plantain.
Bananas and plantains are consumed in a wide variety of manners in Africa. Dessert
bananas are consumed raw as snacks and desserts. Plantains are fried in various ways and
eaten as side dishes and fast foods. Cooking bananas and highland bananas are pounded
into thick porridges (fufu and matooke). Beer bananas are fermented and consumed
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as traditional wine in the Great Lakes regions of Uganda, Democratic Republic of

Congo, Rwanda and Burundi.
14.2 Banana Production Trends in Africa

Sub-Saharan Africa produces about 35% of the worlds bananas and plantains (Table
14.1). Banana and plantains have been estimated to supply more than 25% of the
carbohydrates of approximately 70 million people in Africas humid forest and midaltitude regions (Vuylsteke et al., 1992).
Banana production increases have been slow, and generally not kept pace with
population growth (Fig. 14.1). This is due mainly to decreasing yields in the East
African Highlands, caused in part by increasing incidence of weevils, black Sigatoka
disease and nematodes. In West Africa, as well, plantain production has been seriously
affected by high levels of incidence of black Sigatoka.
East Africa (most notably the Great Lakes region covering portions of Rwanda,
Burundi, Tanzania, Kenya and Congo) is the largest producer and consumer region for
bananas in Africa. The Great Lakes region is estimated to produce 15 million tonnes of
bananas per year, and per capita consumption is the highest in the world (INIBAP,
Table 14.1.
Production of bananas and plantains in Africa.

Production
(1000 t year 1)
Region
West and Central Africa
Angola
Cameroon
Dem. Rep. of Congo
Rep. of Congo
Cte dIvoire
Gabon
Ghana
Guinea
Liberia
Nigeria
East Africa
Burundi
Kenya
Madagascar
Malawi
Rwanda
South Africa
Tanzania
Uganda
Others
Total
Per capita consumption

(kg)
318
1274
1831
80
1194
159
637
318
159
1990
24
85
69
46
98
142
64
46
44
25
1506
452
301
151
2108
151
1656
8432
301
23,018
88
34
17
9
5
180
43
222
Source: FAO (1999).
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Banana
2000). However, increases in production have not kept pace with population growth,
indicating stagnating banana production in these principal areas. The decline in productivity of banana plantings in Uganda has been extensively studied and documented
(Frison et al., 1999; Karamura, 1999) (Fig. 14.2). Causes of this trend have been blamed
on increasing incidence of pests and diseases, most notably nematodes, weevils (associated with declining soil fertility), black Sigatoka and Fusarium (associated with the
increased incidence of these diseases, worldwide) (see Plate 23).
The small number of pests and diseases, coupled with the wide exchange of information regarding their causal agents among a relatively small number of researchers,
makes it difficult, if not impossible, to consider Africa-based research on pests and
Fig. 14.1.
(1999).
Banana and plantain production in Africa, 19751999. Source: FAO
Fig. 14.2.
Declining banana yields in Rwanda and Uganda, 19701997.
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pathogens in isolation from the rest of the world. Therefore, in contrast to sections on
pests and pathogens of other crops in this book, those affecting banana and plantain in
Africa will be considered in relation to their incidence, worldwide.
14.3 Banana Production Constraints

Bananas throughout the world suffer from a relatively small number of pests and
diseases, which however can be highly devastating to yield and production. Moreover, a
number of the most important pests and diseases in Africa have been increasing in recent
years (Wilson, 1988), often associated with the widespread phenomenon of reduced
fallow periods (IITA, 1998). Among the most serious pests and diseases are: banana weevil (Cosmopolites sordidus), a complex of nematodes (Pratylenchus goodeyi, Helicotylenchus
multicinctus and Radopholus similis), black streak/black Sigatoka (Mycosphaerella
figiensis), yellow Sigatoka (M. musicola), Fusarium wilt (Fusarium oxysporum), banana
bunchy top virus and banana streak virus.
Of these, black streak/black Sigatoka is considered to be the most serious biotic
constraint to banana and plantain production in Africa (Swennen et al., 1989; Ortiz and
Vuylsteke, 1994). Black streak/black Sigatoka disease was accidentally introduced in
central Africa in the 1980s (Wilson and Buddenhagen, 1986). All plantains and East
African highland bananas are susceptible to black streak/black Sigatoka.
Worldwide, banana bunchy top virus is perhaps the most important virus affecting
Musa (Dale, 1987). In Africa, it has been most widely observed in central Africa
(Diekmann and Putter, 1996), however, it is not believed to cause heavy losses.
Banana streak virus (BSV) was first described on banana plantings in Cte dIvoire
by Lassoudiere in 1974 and first isolated by Lockhart in 1986. BSV is a badnavirus
which occurs throughout banana-producing areas of the world. Whereas numerous
recent studies have shown that badnavirus is present in many plantings, significant loss
of production caused by the disease has been limited to a small number of locations.
In nature, BSV is believed to be vectored by the mealy bug. BSV sequences are
incorporated into the host genome. It is believed that tissue culture triggers transcription
of the virus, leading to symptom expression. De novo generation of BSV infection of
clean germplasm has occurred in a number of progeny reproduced through tissue
culture, especially among the recently produced tetraploid hybrid varieties (Hughes and
Tenkouano, 1998).
Based on detection of integrated BSV sequences in banana chromosomal DNA
isolated from asymptomatic plants, there is speculation that activation of the virus
may occur as a result of tissue culture or other stresses. Episomal BSV appeared
in tissue-cultured plants which shared more than 99% sequence homology with
integrated BSV from parent plants (Lockhart et al., 1998). The disease causes mild
symptoms which include chlorotic and necrotic streaks on leaf tissues, and severe
symptoms which can include distorted bunches, heart rot, and plant death. Yield
reductions from these symptoms range from 7 to 90% (Frison and Sharrock, 1998).
The most important insect pest of banana and plantain in Africa is the banana
weevil (Cosmopolitus sordidus). Banana weevils enter the plant through the soil and bore
through the base of the pseudostem, thus weakening the plant. Both highland bananas
and plantains are susceptible to weevils (Vuylsteke et al., 1993).
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A less well understood but nevertheless well-documented problem of plantain

production in West and central Africa is commonly known as yield decline. Yield
decline is usually associated with medium-sized plantings among smallholder producers
who grow bananas in open fields (Ortiz and Vuysteke, 1994). This disease may be the
result of a complex of production-related constraints such as declining soil fertility,
including micronutrient deficiencies, weevils and nematodes.
Nematodes are recognized as important pests of bananas and plantains in most
producing areas. Average annual losses worldwide are believed to be in the order of 20%
(Sasser and Freckman, 1987). The most damaging species of nematodes on bananas
are the burrowing nematode (Radopholus similis), root-lesion nematodes (Pratylenchus
coffeae and P. goodeyi), and spiral nematodes (Helicotylenchus multicinctus) (Speijer and
De Waele, 1997). Above-ground symptoms of nematode damage include plant lodging,
stunting, chlorosis and reduction of bunch weight.
The threat to banana production posed by Fusarium wilt is greatest in plantation
situations. The pathogen proliferates in the vascular system of plants, causing symptoms
of terminal wilt, yellowing of leaves, and loss of yield. Chemical control measures are not
effective at controlling the disease. Soils which have been infested with the pathogen
cannot be cultivated with susceptible varieties of banana for up to 30 years. Fusarium
wilt was first discovered in Australia by Bancroft in 1876. By the 1960s it had spread
to most of the major banana and plantain producing regions of the world. The
susceptibility of the dessert varieties Gros Michel and Ladyfinger to Fusarium wilt led
to their virtual elimination from use worldwide. They were eventually replaced by the
variety Cavendish, which now accounts for nearly all of the worldwide export market
for dessert bananas. Although Cavendish is immune to Race 1 of Fusarium wilt, it has
proven to be susceptible to Race 4.
In East Africa, Race 1 of Fusarium wilt has been known to occur on introduced
varieties since the 1950s (Jameson, 1953; Ploetz, 1990). It has been reported on East
African highland bananas since the 1980s; however, incidence in infected plantings has
been estimated at less than 5% of plants (Ploetz, 1994). Race 4 of Fusarium wilt is an
important disease on plantations of Cavendish bananas in South Africa.
14.4 Banana Improvement Through Biotechnology and Breeding

Banana breeding advances in Africa
Banana improvement wordwide is characterized by the extremely small number
of teams actively involved in breeding. Important centres for banana and plantain
improvement are located in: Honduras, at the Fundacion Hondurena de Investigacion
Agricola (FHIA); in Onne, Nigeria, at the International Institute for Tropical Agriculture (IITA); in Brazil, at the Empresa Brasileira de Pesquisa Agropecuaria (EMBRAPA);
in Montpelier, France at the Centre International de Rescherche Agricole pour le
Developpement (CIRAD-FLHOR); in Cameroon, at the Centre de Recherche sur la
Banane et Plantain (CRBP); at the Agricultural Research Centre in South Africa; and at
the Banana Board of Jamaica (Frison et al., 1997).
Breeding of banana and plantain has achieved a major breakthrough in recent years
with the development of a hybridization technique advanced by FHIA (Rowe and
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Rosales, 1994). In this method, male fertile diploid bananas are used to pollinate
popular triploid varieties to produce tetraploid hybrids. In most cases, fertile diploids are
sought which can contribute important traits not found in cultivated triploid species.
The result is a hybrid tetraploid which expresses traits of both parents. Following
identification of improved, adapted tetraploids, breeders make additional crosses to
produce sterile triploids.
Following a long-term effort employing this approach, FHIA recently began
distributing a series of tetraploid hybrid bananas with novel traits and good general
agronomic characteristics for various uses. However, adoption rates of these hybrids in
East Africa have remained low, due to poor cooking quality (INIBAP, 2000). These
varieties have been tested and released by farmers in several important banana-growing
regions of Africa. Subsequently, IITA breeders in Uganda began employing similar
methods aimed at improving East African Highland bananas (Ortiz and Vuylsteke,
1994). Adoption of improved varieties developed by this method, however, have also
been reported to be limited by low yield potential and poor cooking quality (INIBAP,
2000).
Unfortunately, progeny of tissue-cultured tetraploid hybrids have been diagnosed
with high percentage infection of banana streak virus, and have been quarantined in
some parts of the world. Because there is no reason to believe that these varieties should
have higher susceptibility to BSV, many questions have been posed, with few definitive
answers. It has been postulated that tissue culture may have led to the expression of BSV,
which is an integrated DNA virus (Frison and Sharrock, 1998).
Breeding for resistance to black streak/black Sigatoka disease and other biological
constraints at IITA represents another recent, significant breakthrough in banana/
plantain breeding in Africa. Beginning in 1987, IITA screened Musa accessions for
resistance to black streak/black Sigatoka and found 30 sources of resistance, most of
which were fertile diploid types (Swennen and Vuylsteke, 1991). Following the breeding
technique developed by Rowe, researchers crossed male-fertile, resistant diploid accessions to female-fertile, tripoid cultivated varieties to obtain resistant, tetraploid hybrids.
Additional diploid accessions produced through this process have since been crossed to
tetraploid hybrids to obtain male-sterile, secondary triploids (Ortiz and Vuylsteke,
1994). IITA has developed five hybrid cultivars (PITA-2, PITA-3, PITA-8,
PITA-14 and PITA-17) with resistance to black Sigatoka. These are being tested
on-farm in Uganda. In addition, IITAs PITA-16 has been shown to be black Sigatoka,
lodging, nematode and (moderately) weevil resistant (Vuylsteke et al., 1998). PITA-3
has been identified for release and distribution in Cte dIvoire.
Nematode resistance has been detected in bananas, with Yangambi Km5 (AAA)
and Pisang Jari Buaya (AA) having been classified as highly and completely resistant
to R. similis, respectively (Speijer and De Waele, 1997). Banana improvement for
resistance to nematodes is constrained by lack of an efficient screen, making the trait a
candidate for eventual selection via molecular marker.
Recently, researchers in Uganda have produced convincing evidence of a genetic
factor in the control of weevils in East African Highland banana (Gold et al., 1998).
Data from liquid chromatograph analyses indicated that several compounds could be
identified in resistant varieties which were absent in susceptible varieties. Ongoing
research is aimed at identifying the compounds responsible and, eventually, developing
an efficient laboratory screen for this trait.
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Advances in banana biotechnology

A review of biotechnology applications for banana was published in 1998 by Crouch
et al.
Tissue culture of bananas via multiple shoot tip culture based on the addition of
cytokinins to standard media has become a widespread method of propagating selected,
clean clones for cultivation in field and plantation (Vuylsteke et al., 1997). In Kenya,
selected farmers have adopted the cultivation of tissue-cultured banana on both large
and small scale (ISAAA, 1997). Tissue-cultured banana plants exhibit significantly
increased vigour and yield, and earlier maturity. However, broader use of tissue-cultured
bananas depends on scaling up both the culturing and dissemination systems. Strategies
aimed at commercial-scale dissemination of tissue-cultured bananas have been advanced
for support of donors and African governments. Propagation of banana via cell suspension has been achieved in various laboratories (Novak et al., 1989; Dheda et al., 1991);
however, the technique has not become routine (see Plate 24).
Molecular genetics studies of banana have focused primarily on identification of
genetic variation and useful genes using PCR-based markers (Crouch et al., 1998). A
molecular linkage map has been developed for banana using a range of marker systems
(Faur et al., 1993), and several hundred SSR markers have been identified by various
research teams (Jarret et al., 1994; Lagoda et al., 1995; Kaemmer et al., 1997). High
levels of polymorphism have been detected, and strategies have been advanced for
utilization of RAPD markers to analyse genetic variation in East African Highland
bananas (Patrick Rubahaiyo, personal communication). More recently, AFLP markers
have been identified and advanced as effective systems for genetic analysis and improvement in banana (Crouch et al., 1998).
Due to the significant barriers inherent in conventional breeding of bananas,
molecular breeding has been viewed as an advancement of significant potential benefit
for the crop. Banana improvement programmes with molecular genetics capacity have
proposed numerous applications of molecular methods, including the cloning of microsatellites, analysis of breeding systems (especially, the enhancement of tetraploid
generation methods), disease diagnostics, and marker-assisted selection. To date,
however, no clear strategy has been formulated for genetic improvement of banana or
plantain in principal areas of production that integrates the two methodologies.
Tragically, an airline crash in Abidjan, Cte dIvoire, in early 2000 robbed the banana
improvement community in Africa of three very dedicated and critical sources of
expertise on this subject area.
Dessert banana and plantain have been successfully transformed using electroporation of protoplasts (Sagi et al., 1994), particle bombardment of embryogenic cells
(Sagi et al., 1995), and Agrobacterium (May et al., 1995). To date, Agrobacteriummediated methods have proved the most successful (Crouch et al., 1998). Enhanced in
vitro resistance to Fusarium wilt and black Sigatoka has been reported in transgenic
bananas developed at the Catholic University of Leuven (Remy et al., 1998).
Researchers at the University of Leeds and the University of Wales, UK, are developing a
gene construct aimed at conferring resistance to weevils (Johanson and Ives, 2000).
However, no transgenic bananas are currently in cultivation, due in part to the lack of
opportunities to field-test them. In addition, East African Highland bananas have not as
yet been transformed. In their review of genetic transformation possibilities, Crouch
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et al. (1998) identified resistance to the viral diseases, banana streak virus and banana
bunchy top virus as the two top priorities. In 2000, the Ugandan government committed $500,000 toward the development of transgenic banana with resistance to black
sigatoka, weevils and nematodes.
14.5 Principal Challenges for Banana Improvement in Africa

A worldwide banana improvement strategy advanced by the World Bank in 1998
proposed the following allocation of priorities for genetic research: black Sigatoka
resistance (especially in Cavendish-type banana) 50%; nematode resistance 25%;
Fusarium wilt resistance 15%; and virus diseases 10%. In the absence of more
agro-ecologically focused prioritization, these can be assumed to be at least partially
relevant for Africa, as well.
Fusarium wilt and black Sigatoka-resistant bananas for highland areas of East Africa
represent a major challenge for banana breeders, not least because farmers in this region
have not taken enthusiastically to previous offerings. East African Highland bananas
have physical and chemical properties that distinguish them from other subgroups
(Hartman et al., 1998). Recent analyses of these characteristics seemed to indicate that
percentage dry matter correlates well with taste preferences.
Two additional areas of focus are more decentralized breeding systems and multilocation testing. Banana breeding remains a technically difficult undertaking. Banana
flowers exhibit low fertility, plants take 1 year to mature and 2 years to produce seed, and
each plant requires upwards of 6 m2 of research land. Methods devised by FHIA and
IITA for banana improvement represent innovations that could be extended to national
level in countries where banana production is important (Hartman and Vuylsteke,
1998). Multilocation testing of the materials which have already been developed
through these techniques could contribute to higher adoption rates as well as more accurate priority setting for future breeding efforts.
Molecular breeding of banana could replace laborious, time-consuming screens in
the field (some which may require up to 1 year) by developing tightly linked markers for
major traits. However, providing sufficiently large segregating populations for screening
is dependent on further improvements in seed production and more systematic methods
for converting hybrid tetraploids into sterile triploids.
14.6 Banana Seed Systems

As relatively long-lived crops, substitution of improved cultivars for traditional ones
should be viewed as a long-term goal. Indeed, few initiatives have aimed at systematic
transfer of improved banana varieties to small-scale farmers, and therefore little
information is available regarding farmers common sources of planting stock. A recent
survey performed by IITA in Cameroon provides some insight. Among 243 banana
farmers interviewed, 89.3% of farmers procured their planting material from preceding
plantings, 13.3% were purchased off-farm, and 5.8% were obtained through trades with
neighbours (Hauser et al., 1998). Recently, dissemination of improved planting material
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Banana
of banana throughout Africa has been seriously constrained by quarantine regulations

associated with the incidence of banana streak virus in tissue-cultured germplasm
(Vuylsteke et al., 1998).
Due to the high costs (transport, labour) related to such substitutions, farmers will
probably only be persuaded to do so if there are significant, easily recognized advantages
in the new varieties. Incremental yield increases are not viewed as a likely motivating
factor for small farmers. Significantly higher resistance to diseases or pests, however,
could well serve as a sufficiently strong persuasion factor, since farmers who lose a crop
to pests and diseases may lose a 612 month investment.
Dissemination of improved dessert banana for commercial, semi-commercial, and
export markets can be a relatively straightforward exercise. Because of the bulkiness of
the planting material, however, the opportunities for small-scale seed enterprises to
engage in broad-based dissemination of improved banana are probably limited.
Commercial growers and small-scale producers of banana are likely to follow very
different decision-making processes regarding improved cultivars.
Dissemination of improved banana for small-scale producers is likely to always
require public resources, even if the major operations are subcontracted to private sector
groups. In the case of tissue-cultured bananas, this could come in the form of culturing
and hardening off of plantlets in soil, prior to sale at subsidized prices to stockists in the
target areas. For non-tissue-cultured bananas, this may come in the form of maintaining
large planting stock nurseries in the project area for direct sale to farmers and
wholesaling to stockists. The latter activity could be taken on by local NGOs supported
by donor agencies and governments.

1. Development of Sigatoka and Fusarium-resistant East African Highland banana.
This is perhaps the single, highest potential impact objective for Africa, due to
the high dependence on banana among farmers of the Great Lakes region. Major
difficulties stem from issues related to farmer preferences in varieties used for
traditional dishes. Transgenic varieties of plantain with reported high levels of resistance
to Sigatoka disease are currently blocked from entry into areas where they could
have a major impact due to lack of necessary biosafety protocols covering transgenic
crops.
2. Identification of useful markers for breeding applications. This work would be aimed
at eliminating the need for time-consuming screens for resistance traits. Candidate traits
for marker identification would be resistances to nematodes, weevils, viruses, and major
foliar diseases. Putative resistance to weevils has been noted in some varieties of East
African Highland banana.
3. Multilocation testing of improved varieties. Because banana improvement will always
be a task carried out by only a few research groups, broad-based testing is critical to
inform these teams of the potential for adoption of improved varieties. At present,
testing of improved bananas is carried out only on a very limited scale.
4. Further research on banana streak virus. Although BSV is a relatively low-level threat
to production in most of Africa, it can reach epidemic levels in certain cases, as in the
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recent outbreak in Rakai and surrounding districts of Uganda. More critically, poor
understanding of BSV infection and expression has led regulators to operate with
extreme caution in the dissemination of improved, but possibly BSV-infected, varieties.
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References
Abadasi, J.A., Singh, B.B., Ladeinde, T.A.O., Soyinka, S.A. and Emechebe, A.M. (1987)
Inheritance of resistance to brown blotch. Septoria leaf spot and scab in cowpea. India
Journal of Genetics 47, 299303.
Adenle, V., Cardwell, K.F., Ayinde, O., Onukwu, D. and Ogbe, G. (1998) Studies on
the penetration and establishment of downy mildew in maize seeds and correlation
with seed transmission. In: Project 5: Integrated Management of Maize Pests and Diseases.
Annual Report 1998. International Institute of Tropical Agriculture, Ibadan, Nigeria,
pp. 4.
Adesina, A.A. and Baidu-Forson, J. (1995) Farmers Perceptions and Adoption of New Agricultural
Technology: Evidence from Analysis in Burkina Faso and Guinea, West Africa. Elsevier,
1986Oct. 1995. Amsterdam, Netherlands.
Adjadi, O., Singh, B.B. and Singh, S.R. (1985) Inheritance of bruchid resistance in cowpea. Crop
Science 25, 740742.
Aggrawal, B.L. and House, L.R. (1982) Breeding for pest resistance in sorghum. In: Proceedings of
the International Symposium on Sorghum in the Eighties, 27 Nov, 1981. ICRISAT, Andra
Pradesh, India, pp. 435446.
Aggrawal, B.L., Abraham, C.V. and House, L.R. (1988) Inheritance of resistance to midge in
sorghum. Insect Science Applications 9, 4345.
Ahmed, M.M., Sanders, J.H. and Nell, W.T. (2000) New sorghum and millet cultivar introduction in sub-Saharan Africa: impacts and research agenda. Agricultural Systems, 111.
Altieri, M.A. and Rosset, P. (1999) Ten reasons why biotechnology will not ensure food security, protect the environment and reduce poverty in the developing world. AgBioForum
2(3&4), 155162. Retrieved 1 January, 2000 from the World-wide Web: http://www.
agbioforum.org.
Ampong-Nyarko, K., Odindo, M.O., Khan, Z.R. and Overholt, W.A. (1998) Maize Streak Virus in
Eastern and Southern Africa Vector Epidemiology. International Center for Insect Physiology
and Ecology project report, Nairobi, Kenya.
Andrews, D.J. and Anand Kumar, K. (1996) Use of the West African pearl millet land race Iniadi
in cultivar development. Plant Genetic Resources Newsletter 105, 1522.
171
171
Chapter
172
References
Andrews, D.J., King, S.B., Whitcombe, J.R., Singh, S.D., Rai, K.N., Thakur, R.P., Talukdaar, B.S.,
Chavan, S.B. and Singh, P. (1985) Breeding for disease resistance and yield in pearl millet.
Field Crops Research 11, 241258.
Anonymous (2000) Harmonization of Seed Policies and Regulations in Eastern Africa: Agreements,
Recommendations and Way, 16 pp.
Arnanson, J., de Beyssac, B. Conilh, Philogene, B.J.R., Bergvinson, D., Serratos, J.A. and Hamilton, R.I. (1994) Mechanisms of resistance in maize grain to maize weevil and larger grain
borer. In: Mihm, J.A. (ed.) Insect Resistant Maize. Proceedings of an International Symposium, 27
Nov.3 Dec. 1994. CIMMYT, D.F., Mexico, pp. 9195.
Arundel, A., Hocke M. and Tait J. (2000) How important is genetic engineering to European
seed firms? Nature Biotechnology 18(6), 578.
Asanzi, C. and DeVries, J.D. (1995) End of Season Report: Zaire Agricultural Recovery Program. WV
International, Lubumbashi, Zaire, 16 pp.
Atokple, I.D.K., Singh, B.B. and Emechebe, A.M. (1995) Genetics of resistance to Striga and
Alectra in cowpea. Journal of Heredity 86, 4549.
Bahia, A.F.C. and Lopes, M.A. (1998) State of the art developing grain cultivars for acid
savannas of Brazil. In: Schaffert, R.E. (ed.) Proceedings of a Workshop to Develop a Strategy for
Collaborative Research and Dissemination of Technology in Sustainable Crop Production in Acid
Savannas and other Problem Soils of the World. Purdue University and EMBRAPA, pp. 2745.
Bancroft, J. (1876) Report of the board appointed to enquire into the cause of disease affecting livestock
and plant, Queensland. Notes and Proceedings.
Bangarwa, K.S., Lodhi, G.P. and Grewal, R.P.S. (1987) Inheritance of resistance to red leaf spot
disease in sorghum. Indian Journal of Genetics and Plant Breeding 47, 351352.
Bnziger, M., Betrn, F.J. and Lafitte, H.R. (1997) Efficiency of high-nitrogen selection environments for improving maize for low-nitrogen target environments. Crop Science 37,
11031109.
Barker, R. and Cordova, V.G. (1978) Labor utilization in rice production. In: IRRI. Economic
Consequences of the New Rice Technology. International Rice Research Institute, Los Baos,
Philippines.
Barton, J.H. (1998) International intellectual property and genetic resource issues affecting
agricultural biotechnology. In: Ives, C.L. and Bedford, B.M. (eds) Agricultural Biotechnology
in International Development. CAB International, Wallingford, UK, pp. 273283.
Barton, J.H. (1999) Biotechnology for Developing-Country Agriculture: Problems and Opportunities.
Intellectual Property Management. Focus 2. Brief 7 of 10. October 1999. International Food
Policy Research Institute (IFPRI), Washington, DC.
Barton, K.A., Whitely, H. and Yang, N. (1987) Bacillus thuringiensis endotoxin in transgenic
tobacco provides resistance to lepidopteran insects. Plant Physiology 85, 11031109.
Bata, H.D., Singh, B.B., Singh, S.R. and Ladeinde, T.A.O. (1987) Inheritance of resistance to
aphid in cowpea. Crop Science 27, 892894.
Bateman, K.S., Hinch, J.M., Ralton, J.E., Clark, A.E., McKenzie, J.A., Imrie, B.C. and Howlett,
B.J. (1989) Inheritance of resistance in cowpea to Phytophthora vignae in whole plants
cuttings and stem callus cultures. Australian Journal of Botany 37, 511517.
Bennetzen, J.L. (1996) The potential of biotechnology for the improvement of sorghum and
pearl millet. In: INTSORMIL, ICRISAT. Proceedings of the International Conference on Genetic
Improvement of Sorghum and Pearl Millet, held on September 2227, 1996 at Holiday Inn Plaza,
Lubbock, Texas.
Benson, T.D. (1997) Annotated Bibliography of the Work on Area-Specific Fertiliser Recommendations
for Maize in Malawi. Maize Commodity Team Annual Report for the Year 1995/96.
Chetidze Agricultural Research Station, Malawi.
van den Berg, J. (2000) Evaluation of SMIP-developed sorghum cultivars for resistance to
stem borer (Chilo partellus) and the aphid (Melanaphis sacchari). In: Minja, E.M. and van den
172
Chapter
173
References
Berg, J. (eds) Proceedings of the Workshop on Management of Sorghum and Pearl Millet Pests in
the SADC Region, 1013 February 1998, Matopos Research Station, Zimbabwe. ICRISAT,
Bulawayo, Zimbabwe.
Berhan, A.M., Hulbert, S.H., Butler, L.G. and Bennetzen, J.L. (1993) Structural and evolution of
the genomes of sorghum and maize. Theoretical and Applied Genetics 86, 598604.
Bhaskaran, S. and Smith, R.H. (1990) Regeneration in cereal tissue culture: a review. Crop Science
30, 13281336.
Bhattramakki, D., Dong, J., Chhabra, A.K. and Hart, G.E. (2000) An integrated SSR and RFLP
linkage map of Sorghum bicolor (L.) Moench. Genome 43, 9881002.
Binelli, G., Gianfranceschi, L., Pe, M.E., Taramino, G., Busso, C., Stenhouse, J. and Ottaviano,
E. (1992) Similarity of maize and sorghum genomes as revealed by maize RFLP probes.
Theoretical and Applied Genetics 84, 1016.
Binswanger, H.P. and Pingali, P. (1989) Technological priorities for farming in sub-Saharan
Africa. Journal of International Development, 1, 4665.
Blackie, M.J. (1994) Maize productivity for the 21st century: the African challenge. In: Jewell,
D.C., Waddington, S.R., Ransom, J.K. and Pixley, K.V. (eds) Maize Research for Stress
Environments. Proceedings of the Fourth Eastern and Southern Africa Regional Maize Conference,
held at Harare, Zimbabwe, 28 March1 April 1994. CIMMYT, Mexico DF, Mexico, pp.
xixxiii.
Bosque-Perez, N.A. and Mareck, J.H. (1990) Distribution and species composition of lepidopterous maize borers in southern Nigeria. Bulletin of Entomological Research 80, 363368.
Bubeck, D.M., Goodman, M.M., Beavis, W.D. and Grant, D. (1993) Quantitative trait loci
controlling resistance to gray leaf spot in maize. Crop Science 33, 838847.
Buddenhagen, I.W. (1983) Breeding strategies for stress and disease resistance in developing
countries. Annual Review of Phytopathology 21, 385409.
Buddenhagen, I.W. (1996) Modern plant breeding: an overview. In: Persley, G.J. (ed.) Biotechnology and Integrated Pest Management. CAB International, Wallingford, UK.
Buddenhagen, I.W. and de Ponti, O.M.B. (1983) Crop improvement to minimize future losses
to diseases and pests in the tropics. FAO Plant Protection Bulletin 31, 1130.
Bui-Dang-Ha and Pernes, J. (1982) Androgenesis in pearl millet. I. Analysis of plants obtained
from microspore culture. Zeitshrift fr Pflanzenzuchtung 108, 317327.
Bumb, B.L. and Baanante, C.A. (1996) The role of fertilizer in sustaining food security and
protecting the environment to 2020. Food, Agriculture and the Environment Discussion Paper
17. IFPRI, Washington, DC.
Butler, L.M., Myers, J., Nchimbi-Msolla, S., Massangye, E., Mduruma, Z., Mollel, N. and
Dimosa, P. (1995) Farmer evaluation of early generation bean lines in Tanzania: comparisons of farmers and scientists trait preferences. Southern Africa Development Community
(SADC) Regional Bean Research Workshop, 24 Oct. 1995, Oil and protein Seed Center.
Potchefstroom, South Africa, 20pp.
Byerlee, D. (1996) Modern varieties, productivity, and sustainability: recent experience and
emerging challenges. World Development 24 Nos. 14. The World Bank, Washington, DC.
Byerlee, D. and Eicher, C.K. (1997) Africas Emerging Maize Revolution. Lynne Rienner
Publishers, Boulder, Colorado.
Byerlee D. and Heisey, P.W. (1997) Evolution of the African maize economy. In: Byerlee, D. and
Eicher, C.K. (eds) Africas Emerging Maize Revolution. Lynne Rienner Publishers, Boulder,
Colorado.
Byerlee, D. and Lopez-Pereira, M. (1994) Technical Change in Maize: a Global Perspective. Economics
Working Paper No. 9402. CIMMYT, Mexico DF.
Cardwell, K.F., Schulthess, F., Ndemah, R. and Ngoko, Z. (1997) A systems approach to assess
crop health and maize yield losses from pests and diseases in Cameroon. Agriculture,
Ecosystems and Environment.
173
Chapter
174
References
Carson, M.L. (1995) A new gene in maize conferring the chlorotic halo reaction to infection by
Exserohilum turcicum. Plant Disease 79, 717720.
Carter S.E., Fresco, L.O. and Jones, P.G. with Fairbairn, J.N. (1992) An Atlas of Cassava in Africa:
Historical, Agro-ecological and Demographic Aspects of Crop Distribution. CIAT, Cali, Colombia.
Casas, A.M., Kononowicz, A.K., Bressan, R.A. and Hasegawa, P.M. (1995) Cereal transformation
through particle bombardment. Plant Breeding Rev 13, 235264.
Ceccarelli, S. (1989) Wide adaptation: how wide? Euphytica 40, 197205.
CGIAR (1998) CGIAR 1997 Financial Report. CGIAR, Washington, DC, 33 pp.
CGIAR (1998) Report of the 13th Meeting of the CGIAR Finance Committee. CGIAR Secretariat,
Washington, DC, 34 pp.
CGIAR (2000) CGIAR Systemwide Program on Participatory Research and Gender Analysis for
Technology Development and Institutional Innovation. Biotechnology-assisted Participatory Plant
Breeding: Complement or Contradiction? Working Document No. 4. CIAT, Cali, Colombia.
CGIAR (2000) Systemwide Review of Plant Breeding Methodologies in the CGIAR. CGIAR Secretariat, Washington, DC.
CGIAR (2000) Charting the CGIARs Future Reshaping the CGIARs Organization. International
Centers Week 2000. Washington, DC.
CGIAR (2000) Charting the CGIARs Future A New Vision for 2010. IAEG Study: CGIARs Impact
on Germplasm Improvement. Washington, DC.
Chang, R.Y. and Peterson, P.A. (1995) Genetic control of resistance to Bipolaris maydis: one gene
or two genes. Journal of Heredity 86, 9497.
Chapman, J. and White, J.C.N. (1997) World Visions experience with seed supply during
emergency and resettlement programs in Mozambique and Angola; Implications for the
future. Invited paper, presented at: Enhancing research impact through improved seed
supply: Options for strengthening national and regional seed supply systems, 1014 March
1997. Workshop sponsored by ICRISAT, ICARDA, IITA and GTZ. ICRISAT, Bulawayo,
Zimbabwe.
Chapman, J., White, J. and Nankam, C. (1997) World Visions experience with seed supply
during emergency and resettlement programs in Mozambique and Angola: implications
for the future. In: Enhancing research impact through improved seed quality: options for strengthening national and regional seed supply systems. Proceedings of a workshop sponsored by ICRISAT,
ICARDA, IITA and GTZ in Harare, Zimbabwe, March 1014, 1997.
Chavarriaga-Aguirre, P., Maya, M.M., Thoeme, J., Duque, M.C., Iglesias, C.I., Bonierbale, M.W.,
Kresovich, S. and Kochert, G. (1998) Using microsatellites, isozymes and AFLPs to
evaluate genetic diversity and redundancy in the cassava core collection and to assess the
usefulness of DNA-based markers to maintain germplasm collections. Molecular Breeding 5,
263273.
Chittenden, L.M., Schertz, K.F., Lin, Y.R., Wing, R.A. and Paterson, A.H. (1994) Detailed RFLP
map of Sorghum bicolour S. propinquum, suitable for high-density mapping, suggests
ancestral duplication of sorghum chromosomes or chromosomal segments. Theoretical and
Applied Genetics 87, 925933.
Cho, Y.G., McCouch, S.R., Kuiper, M., Kang, M.R., Pot, J., Groenen, J.T.M. and Eun, M.Y.
(1998) Integrated map of AFLP, SSLP and RFLP markers using a recombinant inbred
population of rice (Oryza sativa L.). Theoretical and Applied Genetics 97, 370380.
Chrispeels, M.J. and Sadava, D.E. (1994) Plants, Genes and Agriculture. Jones & Barlett, London,
478 pp.
Christou, P., Ford, T.L. and Kofron, M. (1991) Production of transgenic rice (Oryza sativa L.)
plants from agronomically important indica and japonica varieties via electric discharge
particle acceleration of exogenous DNA into immature zygotic embryos. Bio/technology 9,
957962.
174
Chapter
175
References
CIAT (1998) Saturation of the genetic map of cassava with PCR-based markers and the use of the genetic
map in the improvement of cassava. Progress report to the Rockefeller Foundation. CIAT, Cali,
Colombia.
CIAT/IITA (1999) Mapping of genes for resistance to ACMV in cassava. Progress report to the Rockefeller
Foundation. CIAT/IITA, Cali, Colombia, and Ibadan, Nigeria.
CIMMYT (1988) Maize Production Regions in the Developing Countries. CIMMYT, El Batan,
Mexico.
CIMMYT (1990) 1989/90 CIMMYT World Maize Facts and Trends: Realizing the Potential of Maize
in Sub-Saharan Africa. CIMMYT, Mexico, DF.
CIMMYT (1994) CIMMYT in 1993: Helping the poor through innovative agricultural research.
CIMMYT, Mexico, DF.
CIMMYT (1994) Maize Research for Stress Environments: Proceedings of the Fourth Eastern and
Southern Africa Regional Maize Conference, held at Harare, Zimbabwe, 28 March 1 April 1994,
CIMMYT, Mexico DF.
CIMMYT (1996) Annual Research Report of CIMMYT-Zimbabwe: November 1994 to October 1995.
CIMMYT-Zimbabwe, 100 pp.
CIMMYT (1998) Change for the Better: CIMMYT 1998 Annual Report. CIMMYT, Mexico DF, 28
pp.
CIMMYT (1999) CIMMYT 1997/98 World Maize Facts and Trends; Maize Production in DroughtStressed Environments: Technical Options and Research Resource Allocation, Heisey, P.W. and
Edmeades, G.O. CIMMYT, Mexico D.F., 68 pp.
CIMMYT (2000) Innovative and Integrated Approaches to Improve the Tolerance of Maize to
Water-Limited Environments. Proposal Document. The International Maize and Wheat
Improvement Center (CIMMYT), Mexico DF, 28 pp.
CIMMYT (2000) Proceedings of an international workshop on molecular approaches for the genetic
improvement of cereals for stable production in water-limited environments. CIMMYT, Mexico,
DF. In press.
Cisse, N., Ndiaye, M., Thiaw, S. and Hall, A.E. (1995) Registration of Mouride cowpea. Crop
Science 35, 12151216.
Cisse, N., Ndiaye, M., Thiaw, S. and Hall, A.E. (1997) Registration of Melakh cowpea. Crop
Science 37, 1978.
Coe, E.H., Neuffer, M.G. and Hoisington, D.A. (1988) The genetics of corn. In: Sprague, G.F.
and Dudley, J.A. (eds) Corn and Corn Improvement. Madison, Wisconsin, pp. 81258.
Cohen, J. (1998) Biotechnology for African Crops. Study Commissioned by the Rockefeller
Foundation, ISNAR, The Hague.
Collier, P., Elbadawi, I. and Sambanis, N. (2000) Why are there so many civil wars in Africa?
Prevention of future conflicts and promotion of inter-group cooperation. Paper prepared for
the UNECA Ad Hoc Experts Group Meeting on The Economics of Civil Conflicts in Africa, Addis
Ababa, Ethiopia, April 78, 2000. World Bank.
Conway, G. (1997) The Doubly Green Revolution. Penguin Books, London.
Coulibaly, O., Vitale, J.D. and Sander, J.H. (1998) Expected effects of devaluation on cereal
production in the Sudanian region of Mali. Systems 57, 489503.
Council on Scientific and Industrial Research (1966) The Wealth of India, Vol. 8. Publications
and Information Directorate, CSIR, New Delhi.
Craig, J. and Odvody, G.N. (1992) Current status of sorghum downy mildew control.
In: Milliano, W.A.J. et al. (eds) Sorghum and Millet Diseases. ICRISAT, Andra Pradesh, India,
pp. 213219.
Cromwell, E. (1996) Governments, Farmers and Seed in a Changing Africa. CAB International,
Wallingford, UK.
175
Chapter
176
References
Crouch, J.H., Vuylsteke, D. and Ortiz, R. (1998) Perspectives on the application of biotechnology to assist the genetic enhancement of plantain and banana (Musa spp.). Electronic J.
Biotech. April 15, 1998. http://ejb.ucv.cl/content/vol1/issue1/full/2/index.html#90
Crow, J.F. (1998) Anecdotal, historical and critical commentaries on genetics. Genetics 148,
923928.
Dahlberg, J.A., Hash, C.T., Kresovich, S., Maunder, B. and Gilbert, M. (1996) Sorghum and
pearl millet genetic resources utilization. Proceedings of the International Conference on Genetic
Improvement of Sorghum and Pearl Millet. Meeting held in Lubbock, Texas, September 2227 1997.
INTSORMIL/ICRISAT, pp. 4254.
Daily Nation (2000) KARI Launches first GM Crop. Article by staff reporter Zipporah Mussah,
p. 4, August 19, 2000. Daily Nation, Nairobi, Kenya.
Dale, J.L. (1987) Banana bunchy top: an economically important tropical plant virus disease.
Advances in Virus Research 33, 301325.
David C. and Otsuka, K. (1994) Modern Rice Technology and Income Distribution in Asia. Lynn
Riener, Boulder, Colorado.
David, S. and Sperling, L. (1999) Improving technology delivery mechanisms: lessons from
research in eastern and central Africa. Agricultural and Human Values 16, 381388.
David, S., Kasozi, S. and Wortmann, C. (1997) An Investigation of Alternative Bean Seed
Marketing Channels in Uganda. Occasional Publications Series, No. 19. CIAT, Cali, Colombia,
16 pp.
De Boef, W., Amanor, K., Wellard, K. and Bebbington, A. (1993) Cultivating Knowledge: Genetic
Diversity, Farmer Experimentation and Crop Research. Intermediate Technology Publications,
London, 206 pp.
De Leon, C. and Pandey, S. (1989) Improvement of resistance to ear and stalk rots and
agronomic traits in tropical maize gene pools. Crop Science 29, 1217.
Debrah, S.K. (2000) African fertilizer situation and outlook: challenges, opportunities and
implications. Paper presented at the 6th Annual International Conference of the Arab Fertilizer
Association, 31 January to 2 February 2000, Cairo, Egypt.
Dembele, P., Sogoba, J. and Darra, J. (1997) Rapport sur resultats tests sorgho resistant au striga. WV
International, 7 pp.
Denic, M. (1996) Simultaneous selection for earliness and resistance to downy mildew and
streak virus in maize. pp. 219225. In: Ransom et al. (eds) Maize Productivity Gains through
Research and Technology Dissemination. Proceedings of the 5th Eastern and Southern Africa
Regional Maize Conference 37 June. CIMMYT, Harare, Zimbabwe.
Derera, J., Giga, D.P. and Pixley, K.V. (2000) Inheritance of maize weevil resistance in maize
hybrids among maize lines from southern Africa, Mexico and CIMMYT-Zimbabwe.
In: Maize Production Technologies for the Future: Challenges and Opportunties. Proceedings of the
6th Eastern and Southern Africa Regional Maize Conference held in Addis Ababa, Ethiopia, 2125
September, 1998. CIMMYT, Harare, Zimbabwe.
Desai, B.B., Kotecha, P.M. and Salunkhe, D.K. (1997) Seed Handbook. Marcel Dekker, New York,
627 pp.
Devi, P., Zhong, H. and Sticklen, M.B. (2000) In vitro morphogenesis of pearl millet (Pennisetum
glaucum (L.) R.Br.): efficient production of multiple shoots and inflorescences from shoot
apices. Plant Cell Reports 19, 546550.
Devos, K.M., Tittaway, T.S., Busso, C.S., Gale, M.D., Whitcombe, J.R. and Hash, C.T. (1995)
Molecular tools for the pearl millet nuclear genome. In: Sorghum and Millets Newsletter 36,
6466.
DeVries, J.D. (1997) NGOs, scientists and the poor: competitors, combatants or collaborators.
Australian Development Studies Network. Development Bulletin 44, 6870.
DeVries, J.D. (1999) Evaluation report of the World Vision Burundi Agricultural Recovery Program.
World Vision, Bujumbura, 26 pp.
176
Chapter
177
References
DeVries, J.D. and Chapman, J. (1996) Striga-resistant sorghum initiative. Interim report no. 1 to
USAID. World Vision International, Washington, DC. 15 pp.
DeVries, J.D. and Fumo, E. (1995) Maize varietal preferences and constraints to production in
central Mozambique. In: Jewell, D.C., Waddington, S.R., Ransom, J.K. and Pixley, K.V.
(eds) Maize Research for Stress Environments. Proceedings of the Fourth Eastern and Southern
Africa Regional Maize Conference held at Harare, Zimbabwe, 28 March 1 April, 1994.
CIMMYT, Harare, p. 306.
DeVries, J.D. and Ochieng, J.A.W. (eds) (1998) Advances in Striga Research in Kenya: Proceedings of
a Workshop held in Kisumu on 45 December 1997. KARI, Nairobi, Kenya, 141 pp.
DeVries, J.D. and Olufowote, J. (1997) The role of NGOs in crop improvement and seed multiplication. In: Enhancing Research Impact through Improved Seed Quality: Options of Strengthening
National and Regional Seed Supply Systems. Proceedings of a workshop sponsored by
ICRISAT, ICARDA, IITA and GTZ in Harare, Zimbabwe, March 1014, 1997.
De Wet, J.M.J., Harlan, J.R. and Price, E.G. (1970) Origin of variability in the Spontanea complex
of Sorghum bicolor. American Journal of Botany 57, 704707.
Dheda, D., Dumortier, F., Panis, B., Vuylsteke, D. and Langhe, E. De (1991) Plant regeneration
in cell suspension cultures of the cooking banana cv. Bluggoe (Musa spp, ABB group).
Fruits 46, 125135.
Diekmann, M. and Putter, C.A.J. (1996) FAO/IPGRI Technical Guidelines for the Safe Movement of
Germplasm. No. 15. Musa. 2nd edn. FAO, Rome, Italy; IPGRI, Rome, Italy.
Dill, J. (1997) World Wide List of Rice Biotechnology Projects. A publication of the Rice
Biotechnology Quarterly, Dept. of Biology, Winthrop University, Rock Hill, South
Carolina.
Dively, G.P. and Coop, L. (1993) Millet Loss Assessment Project, Mali. 190. USAID, Washington,
DC, 29 pp.
Dixon, A.G.O. (1999) IITA Working Paper: Cassava Germplasm Development in Africa: Past, Present
and Future. IITA, Ibadan, Nigeria.
Dixon, A., Asiedu, R., Wendt, J. and Mahungu, N. (1996) Development and improvement
of broad-based populations. In: Cassava Productivity in the Lowlands and Mid-altitude
Agroecologies of Sub-Saharan Africa. Annual Report. IITA, Ibadan, Nigeria, pp. 69.
Dover, M.J. and Talbot, L.M. (1987) To Feed the Earth: Agro-ecology for Sustainable Development.
World Resources Institute, Washington, DC.
Dudley, J.W. (1993) Molecular markers in plant improvement: manipulation of genes affecting
quantitative trait. Crop Science 33, 660668.
Duvick, D.N. (1992) Genetic contributions to advances in yield of US maize. Maydica 37, 6979.
Eastwell, K.C., Keifer, M.C. and Bruening, G. (1983) Immunity of cowpeas to cowpea mosaic
virus. In: Goldberg, R.B. (ed.) Plant Mol. Biol. UCLA Symposia on Molecular and Cell Biology
Alan R. Liss, New York, pp. 201211.
Eathington, S.R., Dudley, J.W. and Rufener II, G.K. (1997) Marker effects estimated
from testcrosses of early and late generations of inbreeding in maize. Crop Science 37,
16791685.
Edmeades, G.O., Chapman, S.C., Bolanos, J., Banziger, M. and Lafitte, H.R. (1994) Recent
evaluations of progress in selection for drought tolerance in tropical maize. In: Jewell,
D.C., Waddington, S.R., Ransom, J.K. and Pixley, K.V. (eds) Maize Research for Stress
Environments. Proceedings of the Fourth Eastern and Southern Africa Regional Maize Conference,
Harare, Zimbabwe, 28 March 1 April, 1994, pp. 94100.
Ehlers, J.D. and Hall, A.E. (1997) Cowpea (Vigna unguiculata L. Walp.). Field Crops Research 53,
187204.
Eicher, C.K. (1990) Africas Food Battles. In: Eicher, C.K. and Staatz, J.M. (eds) Agricultural
Development in the Third World. Johns Hopkins University Press, Baltimore, Maryland,
pp. 503530.
177
Chapter
178
References
Ejeta, G. (1998) Hybrid seed experience in Sudan. Paper presented at Regional Hybrid Sorghum and
Pearl Millet Seed Workshop, held in Niamey, Niger, September 28 October 2, 1998.
Ejeta, G., Mohammed, A., Rich, P., Melake-Berhan, A., Housley, T.L. and Hess, D.E. (2000)
Selection for specific mechanisms of resistance to Striga in sorghum. In: Haussman, B.I.G.,
Hess, D.E., Koyama, M.L., Grivet, L., Rattunde, H.F.W. and Geiger, H.H. (eds) Breeding for
Striga Resistance in Cereals. Proceedings of a Workshop held at IITA, Ibadan, Nigeria, from 1820
August 1999. Margraf Verlag, Weikersheim, Germany, pp. 2937.
Ekanayake, I. (1996) Investigations on the induction of flowering in cassava. In: Cassava
productivity in the lowlands and mid-altitude agroecologies of sub-Saharan Africa. Annual Report,
IITA, Ibadan, Nigeria, p. 32.
Elwinger, G.F., Johnson, M.W., Hill, R.R. and Ayers, J.E. (1990) Inheritance of resistance to Gray
Leaf Spot in Corn. Crop Science 30, 350358.
Emechebe, A.M. and Florini, D.A. (1997) Shoot and pod diseases of cowpea induced by fungi
and bacteria. In: Singh, B.B., Mohan Raj, D.R., Dashiell, K.E. and Jackai, L.E.N. (eds)
Advances in Cowpea Research. Copublication of International Institute of Tropical
Agriculture (IITA) and Japan International Research Center for Agricultural Sciences
(JIRCAS). IITA, Ibadan, Nigeria, pp. 176192.
Emechebe, A.M. and Shoyinka, S.A. (1985) Fungal and bacterial diseases of cowpeas in Africa.
In: Singh, S.R. and Rachie, K.O. (eds) Cowpea Research, Production and Utilization. John
Wiley & Sons, Chichester, UK, pp. 173192.
Erbisch, F.H. and Maredia, K.M. (1998) Intellectual Property Rights in Agricultural Biotechnology.
Biotechnology in Agriculture Series, No. 20. CAB International, Wallingford, UK.
Falconer, D.S. (1989) Introduction to Quantitative Genetics. Longman Group, London, UK,
pp. 5461.
FAO (1998) FAO agricultural statistics page on the World-wide Web: www.fao.org
FAO (19982000) www.fao.org. FAOSTAT Database. Internet database. http://apps.fao.org/
lim500
FAO (1999) www.fao.org. Internet Database.
FAO (2000) www.fao.org. Internet Database.
FAO/ICRISAT (1996) The World Sorghum and Millet Economies. FAO/ICRISAT, 68 pp.
FAO/Zimbabwe MOA (1998) African Regional Workshop on Farmer Field Schools for IPM. Meeting
Report. FAO/Zimbabwe MOA, Harare, 35 pp.
Fatokun, C.A., Perrino, P. and Ng, N.Q. (1997) Wide crossing in African Vigna species. In:
Singh, B.B., Mohan Raj, D.R., Dashiell, K.E. and Jackai, L.E.N. (eds) Advances in Cowpea
Research. Copublication of International Institute of Tropical Agriculture (IITA) and Japan
International Research Center for Agricultural Sciences (JIRCAS). IITA, Ibadan, Nigeria.
pp. 5057.
Fauquet, C.M., Huet, H., Ong, C.A., Sivamani, E., Chen, L., Viegas, P., Marmey, V.P., Wang, P.,
Daud, M., de Kochko, A. and Beachy, R.N. (1997) Control of the rice tungro disease by
genetic engineering is now a reality! In: Abstracts, General Meeting of the International
Programme of Rice Biotechnology, Malacca, p. 59.
Faur, S., Noyer, J.L., Horry, J.P., Bakry, F., Lanaud, C. and Gonzlez de Len, D. (1993) A
molecular marker-based linkage map of diploid bananas (Musa acuminata). Theoretical and
Fehr, W.R. (1987) Principles of Cultivar Development. McGraw-Hill, New York, pp. 120134.
Fischer, R.A. (1993) The sustainability debate and wheat science in Australia, developing
countries and CIMMYT. Paper presented to the 8th Regional Wheat Workshop for Eastern,
Central and Southern Africa, Kampala, Uganda, June 610, 1993.
Flavel, R. (1999) Biotechnology for Developing-Country Agriculture: Problems and Opportunities.
Biotechnology and Food and Nutrition Needs. Focus 2. Brief 2 of 10. October 1999. International
Food Policy Research Institute (IFPRI), Washington, DC.
178
Chapter
179
References
Fliedel, G. and Aboubacar, A. (1998) Overview of sorghum and millet food uses. Paper presented
at Regional Hybrid Sorghum and Pearl Millet Seed Workshop, held in Niamey, Niger, September 28
October 2 1998.
Flight, C. (1976) The Kintamp culture and its place in the economic prehistory of West Africa.
In: Harlan, J.R., de Wet, J.M.J. and Stemler, A.B.L. (eds) Origins of African Plant
Domestication. Mouton, The Hague, Netherlands, pp. 212221.
Frankel, F.R. (1971) Indias Green Revolution: Economic Gains and Political Costs. Princeton
University Press, Princeton, New Jersey.
Fredricksen, R.A. (1986) Compendium of Sorghum Diseases. The American Phytopathological
Society, St Paul, Minnesota.
Fregene, M., Akano, A., Gedil, M. and Guitierrez, J. (1996) Final Progress Report to the Rockefeller
Foundation on the Molecular Mapping of Genes Conferring Resistance to the Cassava Mosaic
Disease (CMD) in African Cassava Germplasm. IITA/CIAT, Ibadan, Nigeria.
Frison, E.A. and Sharrock, S.L. (1998) Banana Streak Virus: a Unique VirusMusa Interaction?
Proceedings of a workshop of the PROMusa virology working group held in Montpellier,
France, 1921 January, 1998. IPGRI, Rome, Italy; INIBAP, Montpellier, France.
Frison, E.A., Orjeda, G. and Sharrock, S.L. (eds) (1997) PROMUSA: a Global Programme for
Musa Improvement. Proceedings of a meeting held in Gosier, Guadeloupe, March 5 and 9,
1997. International Network for the Improvement of Banana and Plantain, Montpellier,
France, and The World Bank, Washington, DC, USA.
Frison, E.A., Gold, C.S., Karamura, E.B. and Sikora, R.A. (eds) (1999) Mobilizing IPM for
Sustainable Banana Production in Africa. Proceedings of a workshop on banana IPM held in
Nelspruit, South Africa, 2328 November 1998. INIBAP, Montpellier, France.
Frost, H.M. (1995) Striga Research and Survey in Kenya. National Agricultural Research Project,
KARI/ODA Crop Protection Project. Final report. Nairobi, Kenya, 61 pp.
Fujimoto, H., Itoh, K., Yamamoto, M., Kyozuka, J. and Shimamoto, K. (1993) Insect resistant
rice generated by introduction of a modified -endotoxin gene of Bacillus thuringiensis.
Bio/Technology 11, 11511155.
Galiba, M. (1983) Inheritance of resistance to sooty stripe disease. Sorghum Newsletter 26, 120.
Gerhart, J. (1975) The Diffusion of Hybrid Maize in Western Kenya. CIMMYT, Mexico, DF.
Ghareyazie, B., Alinia, F., Menguito, C.A., Rubia, L.G., De Palma, J.M., Liwanag, E.A., Cohen,
M.B., Khush, G.S. and Bennett, J. (1997) Enhanced resistance to two stem borers in an
aromatic rice containing a synthetic CryIA(b) gene. Molecular Breeding 3, 401404.
Ghesquiere, A., Albar, L., Lorieux, M., Ahmadi, N., Fargette, D., Huang, N., McCouch, S.R. and
Nottenghem, J.L. (1997) A major QTL for RYMV maps to a cluster of blast resistance genes
on chromosome 12. Phytopathology 87, 12431256.
Gold, C., Vuylsteke, D. and Kaggundu, A. (1998) Screening of East African highland banana for
weevil response. In: Project 7: Improving Plantain- and Banana-based Systems. Annual Report
1998. International Institute of Tropical Agriculture, Ibadan, Nigeria, p. 45.
Gomez, F. and Chantereau, J. (1997) Breeding photoperiod sensitive sorghums. In:
INTSORMIL/ICRISAT, 1997. Proceedings of the International Conference on Genetic Improvement of Sorghum and Pearl Millet. Meeting held in Lubbock, Texas, September 2227 1997.
Gordon-Kamm, W.J., Spencer, T.M., Magnano, M.L., Adams, T.R., Daines, R.J., Start, W.G.,
Obrien, J.V., Chambers, S.A., Adams, W.R. and Willets, N.G. (1990) Transformation of
maize cells and regeneration of fertile transgenic plants. Plant Cell 2, 603618.
Griffin, K. (1974) The Political Economy of Agrarian Change: An Essay on the Green Revolution.
Harvard University Press, Cambridge.
Griliches, Z. (1957) Hybrid corn: an exploration in the economics of technological change.
Econometrica 25, 501522.
179
Chapter
180
References
Grimanelli, C. (1998) Ensuring equity in the access to hybrid vigor: introducing apomixis
in maize and other crops. In: Proceedings of the Sixth Eastern and Southern Africa Regional
Maize Conference, held in Addis Ababa, 2125 September, 1998. EARO, Ethiopia and
CIMMYT.
Grossniklaus U., Koltunow, A.M., van Lookeren, M. and Campagne, M.M. (1998) A bright
future for apomixis. Trends in Plant Science 3, 415416.
Hahn, S.K., Terry, E.R., Leuschner, K., Akobundu, I.O., Okali, C. and Lal, R. (1979) Cassava
Improvement in Africa. Reprint from Field Crops Research 2, 193226.
Hahn, S.K., Howland, A.K. and Terry, E.R. (1980) Correlated resistance of cassava to mosaic and
bacterial blight diseases. Euphytica 29, 305311.
Hall, A.E., Singh, B.B. and Ehlers, J.D. (1997) Cowpea breeding. Plant Breeding Reviews 15,
21274.
Hampton, R.O., Thottappily, G. and Rossel, H.W. (1997) Viral diseases of cowpea and their
control by resistance-conferring genes. In: Singh, B.B., Mohan Raj, D.R., Dashiell, K.E. and
Jackai, L.E.N. (eds) Advances in Cowpea Research. Copublication of International Institute of
Tropical Agriculture (IITA) and Japan International Research Center for Agricultural
Sciences (JIRCAS). IITA, Ibadan, Nigeria, pp. 159175.
Hanna, W.W. (1992) Utilization of germplasm from wild species. In: Desertified Grasslands.
Academic Press, London, pp. 251257.
Harlan, J.R. and de Wet, J.M.J. (1972) A simplified classification of cultivated sorghum. Crop
Science 12, 172176.
Hartman, J., Vuylsteke, D., Kangire, A. and Makumbi, D. (1998) Evaluation and genetic
analysis of Fusarium wilt resistance. In: Project 7: Improving Plantain- and Banana-based
Systems. Annual Report 1998. International Institute of Tropical Agriculture, Ibadan,
Nigeria, p. 44.
Hartman, J., Vuylsteke, D., Talengera, D. and Makumbi, D. (1998) Breeding at ESARC
(Uganda). In: Project 7: Improving Plantain- and Banana-based Systems. Annual Report 1998.
International Institute of Tropical Agriculture, Ibadan, Nigeria, p. 48.
Hartman, J., Vuylsteke, D., Ferris, S. and Makumbi, D. (1998) A survey of farmers practices in
preparing plantain planting material in southern Cameroon. In: Project 7: Improving
Plantain- and Banana-based Systems. Annual Report 1998. International Institute of Tropical
Agriculture, Ibadan, Nigeria, p. 42.
Hash, C.T., Witcombe, J.R., Thakur, R.P., Bhatnagar, S.K., Singh, S.D. and Wilson, J.P. (1997)
Breeding for pearl millet disease resistance. In: INTSORMIL/ICRISAT. Proceedings of the
International Conference on Genetic Improvement of Sorghum and Pearl Millet. Meeting held in
Lubbock, Texas, September 2227 1997, pp. 337373.
Hassan, R.M. (1998) Maize Technology Development and Transfer. A GIS Application for Research
Planning in Kenya. CIMMYT/KARI/CAB International, University Press, Cambridge.
Haugerud, A. and Collinson, M.P. (1990) Plants, genes and people: improving the relevance of
plant breeding in Africa. Experimental Agriculture 26, 341362.
Hauser, S., Tschienkoua, M., Madong, B., Nyobe, T. and Dibog, L. (1998) A survey of farmers
practices in preparing plantain planting material in southern Cameroon. In: Project 7:
Improving Plantain- and Banana-based Systems. Annual Report 1998. International Institute of
Tropical Agriculture, Ibadan, Nigeria, p. 4.
Haussmann, B.I.G., Hess, D.E., Gunter Welz, H. and Geiger, H.H. (2000) Improved methodologies for breeding Striga-resistant sorghums. Field Crops Research 66, 195211.
Hayakawa, T., Zhu, Y., Itoh, K., Kimura, Y., Izawa, T., Shimamoto, K. and Toriyama, S. (1992)
Genetically engineered rice resistant to rice stripe virus, an insect-transmitted virus.
Proceedings of the National Academy of Sciences USA 89, 98659869.
Heisey, P.W. and Mwangi, W. (1996) Fertilizer Use and Maize Production in Sub-Saharan Africa.
CIMMYT Economics Working Paper No. 96-01. CIMMYT, Mexico DF, 34 pp.
180
Chapter
181
References
Heisey, P.W., Morris, M.L., Byerlee, D. and Lopez-Pereira, M.A. (1998) Economics of hybrid
maize adoption. In: Morris, M.L. (ed.) Maize Seed Industries in Developing Countries. Lynne
Rienner Publishers, Boulder, Colorado, pp. 143158.
Henry, G. and Gottret, V. (1996) Global Cassava Trends: Reassessing the Crops Future. Working
Document No. 157. CIAT, Cali, Columbia.
Henzell, R.G. and Hare, B.W. (1996) Sorghum breeding in Australia: Public and private
endeavours. In: Proceedings of the Third Australian Conference, 2022 February 1996, Tamworth,
NSW. Australian Institute of Agricultural Science, Melbourne, Occasional Publication
No. 93, pp. 159171.
Henzell, R.G., Peterson, G.C., Teetes, G.L., Franzmann, B.A., Sharma, H.C., Youm, O.,
Ratnadass, A., Toure, A., Raab, J. and Ajayi, O. (1997) Breeding for resistance to panicle
pests of sorghum and pearl millet. In: INTSORMIL/ICRSAT, 1997. Proceedings of the International Conference on Genetic Improvement of Sorghum and Pearl Millet. Meeting held in Lubbock,
Texas, September 2227 1997, pp. 255280.
Herdt, R.W. (1991) Research priorities for rice biotechnology. In: Khush, G.S. and Toenniessen,
G.H. (eds) Rice Biotechnology. CAB International, Wallingford, UK, pp. 1954.
Herdt, R.W. and Capule, C. (1983) Adoption, Spread, and Production Impact of Modern Rice Varieties
in Asia. IRRI, Los Baos, Philippines.
Hildebrand, P.E. (1981) Combining disciplines in rapid appraisal: the sondeo approach.
Agricultural Administration 8, 423432.
Hildebrand, P.E. and Poey, F. (1985) On-farm Agronomic Trials in Farming Systems Research and
Extension. Lynne Rienner Publishers, Boulder, Colorado.
Hittalmani, S., Kumar, G.K., Kulkarni, N. and Shashidhar, H.E. (1999) DNA markers assist in
reducing the number of generations of backcrosses in breeding for blast resistance in rice.
Paper presented at the General meeting of the International Program on Rice Biotechnology, held
on September 2024, 1999 in Phuket, Thailand. Meeting sponsored by the Rockefeller
Foundation, New York.
Hodson, D.P., Rodriguez, A., White, J.W., Corbett, J.D. and Banziger, M. (1999) Africa Maize
Research Atlas (v. 2.0). CIMMYT. Mexico, DF., CD-ROM.
Hoffman, M.P., Thurston, H.D. and Smith, M.E. (1993) Breeding for resistance to insects and
plant nutrients. In: Callaway, M.B. and Francis, C.A. (eds) Crop Improvement for Sustainable
Agriculture. University of Nebraska, Lincoln.
Hoisington, D., Listman, G.M. and Morris, M.L. (1998) Varietal development: applied biotechnology. In: Morris, M.L. (ed.) Maize Seed Industries in Developing Countries. Lynne Rienner,
London, pp. 1335/77102.
Hoisington, D., Khairallah, M., Reeves, T., Ribaut, J.M., Skovmand, B., Taba, S. and Warburton,
M. (1999) Plant genetic resources: what can they contribute toward increased crop
productivity? Proceedings of the National Academy of Sciences USA 96, 59375943.
Holden, S.T. and Shanmugarathan, N. (1994) Structural Adjustment, Production Subsidies and
Sustainable Land Use. Dis. Pap. D-11/1994. Department of Economics and Social Sciences,
Agricultural University of Norway.
House, L.R. (1985) A Guide to Sorghum Breeding. ICRISAT, Patacheru, India.
House, L.R. (1996) Inaugural Address. In: Proceedings of the International Conference on Genetic
Improvement of Sorghum and Pearl Millet. September 2327, 1996. Lubbock, Texas.
House, L.R., Verma, B.N., Ejeta, G., Rana, B.S., Kapran, I., Obilana, A.B. and Reddy, B.V.S.
(1997) Developing countries and potential of hybrid sorghum. In: INTSORMIL/ICRISAT,
1997. Proceedings of the International Conference on Genetic Improvement of Sorghum and Pearl
Millet. Meeting held in Lubbock, Texas, September 2227 1997. INTSORMIL/ICRISAT,
pp. 8496.
Hughes, J.A. and Singh, B.B. (1998) Screening cowpea lines for multiple resistance. In: Integrated
Management of Legume Pests and Diseases. IITA Annual Report, 1998.
181
Chapter
182
References
Hughes, J. and Tenkouano, A. (1998) BSV occurrence, epidemiology, and cultivar interactions.
In: Improving Plantain and Banana-based Systems. IITA Annual Report, 1998.
Ibrahim, O.E., Ahmed, A.T., Omer, M.E., Hamdoun, A.M., Babiker, A.E. and Boreng, P.
(1995) Status of sorghum production, technology generation, transfer and adoption
by farmers in the Sudan. In: Mukuru, S.Z., Ejeta, G. and Ibrahim, N. (eds) Sorghum
and Millets Research in Eastern and Central Africa: Proceedings of a workshop organized to
reestablish a sorghum and millets network in the region, 69 November 1995, Kampala, Uganda,
pp. 157167.
ICRISAT (1992) Medium Term Plan, 1992. Monitoring and Evaluation. Pantancheru, India,
pp. 7580.
ICRISAT (1992) International Crops Research Institute for the Semi-Arid Tropics (ICRISAT) West
African programs annual report 1991. B.P. 12404, Niamey, Niger.
ICRISAT (1996) Improving the Unimprovable: Succeeding with Pearl Millet. ICRISAT, Patancheru,
Andhra Pradesh.
Idachaba, F.S. (1985) Priorities for Nigerian agriculture in the fifth national development plan,
19861990. Nigerian Institute of Social and Economic Research, Ibadan, Nigeria.
IITA (1972) Annual Report of Root and Tuber Improvement Program. IITA, Ibadan, Nigeria.
IITA (1994) Annual Report. IITA, Ibadan, Nigeria, p. 64.
IITA (1995) Annual Report. IITA, Ibadan, Nigeria, p. 55.
IITA (1998) Project 11: CowpeaCereals systems: Improvement in the Dry Savannas. Annual Report
1998. International Institute of Tropical Agriculture, Ibadan, Nigeria.
Imanywoha, J. (1998) Improvement of maize variety 1p 16 in resistance to diseases and tolerance
to low nitrogen. Project proposal, Dec. 1998.
INIBAP (1993) Bananas, Plantains and INIBAP. Annual Report 1993 of The International Network
for the Improvement of Banana and Plantain. Montpellier, France.
INIBAP (1995) Annual Report of The International Network for the Improvement of Banana and
Plantain. Montpellier, France.
INIBAP (2000) Novel approaches to the improvement of banana production in Eastern Africa
the application of biotechnological methodologies. Project Proposal Document. INIBAP,
Rome, Italy, 14 pp.
Ininda, J. and Ochieng, J. (2000) Coordinated Ecosystem Breeding Project. 1st year Report. Kenya Agricultural Research Institute, Nairobi, Kenya.
INTSORMIL/INRAN (1998) Traditional Seed Selection and Conservation Methods of Cereals and
Legumes in Niger: Implications for an Informal Seed System. (B. Ouendeba., T. Abdoulaye, G.
Ibro and K. Anand Kumar) Niamey, Niger.
ISAAA (1997) Annual Report, 1996. Advancing Altruism in Africa. ISAAA, Ithaca, New York.
ISAAA (2000) New Partnerships for Prosperity: Building Public/private Agri-biotech Networks for
Resource Poor Farmers in Southeast Asia and Africa. Biennial Report 19971999. Washington,
DC.
Ishida, Y., Saito, H., Ohta, S., Hieie, Y., Komari, T. and Kumashiro, T. (1996) High efficiency
transformation of maize mediated by Agrobacterium tumefaciens. Nature Biotechnology 14,
745749.
ISNAR (1998) Biotechnology of African Crops. Study Commissioned by the Rockefeller Foundation.
ISNAR, IITA.
Ito, O., OToole, J.C. and Hardy, B. (1999) Genetic improvement of Rice for Water-Limited
Environments. Proceedings of a Workshop on Genetic Improvement of Rice for Water-Limited Environments, 13 December, 1998. International Rice Research Institute, Los Baos, Philippines.
Jaffee, J. (1991) The Balance between Public and Private Sector Activities in Seed Supply Systems. The
World Bank, Washington, DC.
James, C. (1997a) Progressing public-private sector partnerships in international agricultural research and
development. ISAAA Briefs No. 4. ISAAA, Ithaca, New York, 31 pp.
182
Chapter
183
References
James, C. (1997b) Global Status of Transgenic Crops in 1997. ISAAA Briefs No. 5. New York, 30 pp.
James, C. (2000a) Global Status of Commercialized Transgenic Crops: 1999. ISAAA Briefs No. 17.
ISAAA, Ithaca, New York, 65 pp.
James, C. (2000b) Global review of transgenic crops in 1999. ISAAA, Briefs No. 5. ISAAA, Ithaca,
New York, 31 pp.
Jameson, J.D. (1953) Outbreaks and new records. Uganda. FAO Plant Protection Bulletin 1, 62.
Janson, G. and Kapukha, P. (1995) Southern Sudan Agricultural Recovery Program Annual Report.
Jarret, R.L., Bhat, K.V., Cregan, P., Ortiz, R. and Vuylsteke, D. (1994) Isolation of microsatellite
DNA markers in Musa. InfoMusa 3, 34.
Jeffers, D.P. and Chapman, S.C. (1994) Yield losses associated with Exserohilum turcicum and
Puccinia sorghi in high disease incidence environments. In: Jewell, D.C., Waddington, S.R.,
Ransom, J.K. and Pixley, K.V. (eds) Maize Research for Stress Environments. Proceedings of the
Fourth Eastern and Southern Africa Regional Maize Conference, Harare, Zimbabwe, 28 March
1 April, 1994, pp. 157159.
Jefferson, R.A. and Bicknell, R. (1996) The potential impacts of apomixis: a molecular genetic
approach. In: Sobra, B.W.S. (ed.) The Impact of Plant Molecular Genetics. Birkhuser, Boston,
pp. 87101.
Jensen, S. (1994) Genetic improvement of maize for drought tolerance. In: Jewell, D.C.,
Waddington, S.R., Ransom, J.K. and Pixley, K.V. (eds) Maize Research for Stress Environments.
Proceedings of the Fourth Eastern and Southern Africa Regional Maize Conference, Harare,
Zimbabwe, 28 March 1 April, 1994, pp. 6775.
Johanson, A. and Ives, C. (2000) An Inventory of Agricultural Biotechnology for the Eastern and
Central Africa Region. Institute of International Agriculture, Michigan, 59 pp.
Jones. A.L. (ed.), Weltzein, E., Smith, M.E., Meitzner, L.S. and Sperling, L. (1999) Technical and
Institutional Issues in Participatory Plant Breeding from the Perspective of Formal Plant Breeding.
A Global Analysis of Issues, Results, and Current Experience. CGIAR.
Jones, W.O. (1959) Manioc in Africa. Standard University Press, Stanford, California, 315 pp.
Kaemmer, D., Fischer, D., Jarret, R.L., Baurens, F.C., Grapin, A., Dambier, D., Noyer, J.L.,
Lanaud, C., Kahl, G. and Lagoda, P.J.L. (1997) Molecular breeding in the genus Musa: a
strong case for STMS marker technology. Euphytica 96, 4963.
Kanampiu, F.K. (1998) Herbicide-resistant maize to control Striga infestations. In: DeVries, J.D.
and Ochieng, J.A.W. (eds) Advances in Striga Research in Kenya. Proceedings of a workshop held
in Kisumu, Kenya, 45 December 1995, 141 pp.
Karamura, D.A. (1999) Numerical Taxonomic Studies of the East African Highland Bananas
(Musa AAA-East Africa) in Uganda. PhD thesis, Department of Agricultural Botany,
University of Reading. IPGRI, Reading.
KARI (1998) Understanding the mechanisms of maize streak virus resistance of maize lines
from Kenya and from eastern and southern Africa. Project proposal by the Kenya
Agricultural Research Institute. KARI, Nairobi, Kenya, 46 pp.
Kassam, A.H. and Kowal, J.M. (1975) Water use, energy balance and growth of gero millet at
Samaru, Northern Nigeria. Agricultural Methods 15, 333342.
Kelly, A.F. and George, R.A.T. (1998) Encyclopaedia of Seed Production of World Crops. John Wiley
& Sons, Chichester.
Kezire, B.B., Asiimwe, P. and Kyetere, D. (2000) Agricultural biotechnology assessment in
sub-Saharan Africa: Country-specific study Uganda. Paper presented at the Regional Workshop on Building National Biotechnology Innovation Systems held in Mombasa, Kenya, December
68, 2000. Africa Centre for Technology Studies (ACTS), Nairobi, Kenya, 23 pp.
Khush, A.G.S. (1990) Rice breeding: accomplishments and challenges. Plant Breeding Ab. 60,
461469.
Khush, G.S. and Toenniessen, G.H. (1991) Rice Biotechnology. CAB International, Wallingford,
UK, 313 pp.
183
Chapter
184
References
Kim, S.K. (1994) Genetics of maize tolerance to Striga hermonthica. Crop Science 34, 900907.
Kim, S.K. and Brewbaker, J.L. (1976) Effect of Puccinia sorghi on yield and several agronomic
traits of maize in Hawaii. Crop Science 16, 874877.
Kitch, L.W., Shade, R.E. and Murdock, L.L. (1991) Resistance to the cowpea weevil
(Callosobruchus maculatus) larva in pods of cowpea (Vigna unguiculata). Entomologia
Experimentalis Applicata 60, 183192.
Kitch, L.W., Bottenberg, H. and Wolfson, J.L. (1997) Indigenous knowledge and cowpea pest
management in sub-Saharan Africa. In: Singh, B.B., Mohan Raj, D.R., Dashiell, K.E. and
Jackai, L.E.N. (eds) Advances in Cowpea Research. Copublication of International Institute of
Tropical Agriculture (IITA) and Japan International Research Center for Agricultural
Kitch, L.W., Boukar, O., Endondo, C. and Murdock, L.L. (1998) Farmer Acceptability Criteria in
Breeding Cowpea. Cambridge University Press, UK.
Kling, J.G., Fajemisin, J.M., Badu-Apraki, B., Diallo, A., Menkir, A. and Melake-Berhan, A.
(2000) Striga resistance breeding in maize. In: Haussman, B.I.G., Hess, D.E., Koyama, M.L.,
Grivet, L., Rattunde, H.F.W. and Geiger, H.H. (eds) Breeding for Striga Resistance in Cereals.
Proceedings of a Workshop held at IITA, Ibadan, Nigeria, from 1820 August 1999. Margraf
Verlag, Weikersheim, Germany, pp. 103118.
Knapp, S.J. (1991) Using molecular markers to map multiple quantitative trait loci: models for
backcross, recombinant inbred, and doubled haploid progeny. Theoretical Applications of
Genetics 81, 333338.
Koo, B. and Wright, B.D. University of California. (1999) EPTD Discussion Paper No. 51. Dynamic
Implications of Patenting for Crop Genetic Resources. Environment and Production Technology
Division, International Food Policy Research Institute, Washington, DC.
Kramer, K.J., Morgan, T.D., Throne, J.E., Dowell F.E., Bailey, M. and Howard, J.A. (2000)
Transgenic avidin maize is resistant to storage insect pests. Nature Biotechnology 18,
670674.
Kumar, H. (1994) Field resistance in maize cultivars to stem borer Chilo partellus. Annals of
Applied Biology 124, 333339.
Kumaravadivel, N. and Sree Rangasamy, S.R. (1994) Plant regeneration from sorghum anther
cultures and field evaluation of progeny. Plant Cell Reports 13, 286290.
Kyetere, D.T., Kikafunda-Twine, J., Imanywoha, J.B. and Bigirwa, G. (1997) Cereals Programme,
Annual Report, 1997 A & B. NARO, Kampala, Uganda.
Lagoda, P.J.L., Noyer, J.L., Dambier, D., Baurens, F.C. and Lanaud, C. (1995) Abundance and
distribution of SSR (simple sequence repeats) in the Musaceae family: Microsatellite
markers to map the banana genome. In: Proceedings of the FAO/IAEA International
Symposium on Induced Mutations and Molecular Techniques for Crop Improvement, Vienna,
FAO/IAEA, pp. 287295.
Laker-Ojok, R. (2000) AT(Uganda)s role in input distribution. Paper presented at a workshop on
Development of Sustainable Seed Systems for Small-scale Farmers, held at Colline Hotel, Mukono,
Uganda, 2223 May, 2000.
Lale, N.E.S. and Yusuf, B.A. (2000) Insect pests infesting stored pearl millet Pennisetum glaucum
(L.) R.Br. in northeastern Nigeria and their damage potential. Cereal Research Communications 28, 181186.
Lambert, C. (1983) LIRAT et amelioration du mil. Agronomie Tropicale 28, 7888.
Lassoudiere, A. (1974) La mopsaique dite a tirets du bananier Poyo en Cte dIvoire. Fruits 29,
349357.
Lefevre, F. and Charrier, A. (1993) Heredity of seventeen isozyme loci in cassava (Manihot
esculenta Crantz). Euphytica 66, 171178.
Legg, J.P. (1999) Emergence, spread and strategies for controlling the pandemic of cassava
mosaic virus disease in east and central Africa. Crop Protection 18, 627637.
184
Chapter
185
References
Legg, J.P., Whyte, J., Khizzah, B. and Ogwang, J. (1998) CGM/CMD interaction studies. In:
Project 6. Integrated Management of Cassava Pests and Diseases. 1998 Annual Project Report.
International Institute of Tropical Agriculture, Ibadan, Nigeria, pp. 89
Lele, U.J. (1989) Managing agricultural development in Africa. In: Eischer, C.K. and Staatz, J.M.
(eds) Agricultural Development in the Third World, 2nd edn. Johns Hopkins University Press,
London, pp. 531539.
Leuschner, K., Monyo, E.S., Chinhema, E., Tembo, E. and Martin, D. (2000) Pearl millet grain
size and hardness in relation to resistance to Sitophilus oryzea (L.) (Coleoptera Curculionidae).
African Crop Science Journal 8, 7783.
Li, H.Q., Sautter, C., Potrykus, I. and Puonti-Kaerlas, J. (1996) Genetic transformation of cassava
(Manihot esculenta Crantz). Nature Biotechnology 14, 736744.
Lipton, M. and Longhurst, R. (1989) New Seeds and Poor People. Johns Hopkins University Press,
Baltimore, Maryland.
Liu, C.J., Whitcombe, J.R., Pittaway, T.S., Nash, M., Hash, C.T., Busso, C.S. and Gale, M.D.
(1994) An RFLP-based genetic map of pearl millet. Theoret. Applied Genetics 89, 481487.
Lockhart, B.E.L. (1986) Purification and serology for a bacilliform virus associated with banana
streak virus disease. Phytopathology 76, 995999.
Lockhart, B.E.L., Ndowora, T.C., Olszewski, N.E. and Dahal, G. (1998) Studies on integration of
banana streak badnavirus sequences in Musa: Identification of episomally-expressible
badnaviral integrants in genotypes. In: Frison, E.A. and Sharrock, S.L. (eds) Banana Streak
Virus: a Unique Virus-Musa Interaction? Proceedings of a workshop of the PROMUSA Virology
working group held in Montpellier, France, January 1921, 1998. International Plant Genetic
Resources Institute, Rome, Italy; International Network for the Improvement of Banana
and Plantain, Montpellier, France, pp. 4247.
Lopez-Pereira, M.A. and Filippello, M.P. (1995) Emerging Roles of the Public and Private Sectors of
Maize Seed Industries in the Developing World. CIMMYT Economics Program Working Paper
9501. CIMMYT, Mexico, DF., 84 pp.
Lubberstedt, T., Melchinger, A.E., Fahr, S.F., Klein, D., Dally, A. and Westhoff, P. (1998) QTL
mapping in testcrosses of flint lines of Maize: III. Comparison across populations for
forage traits. Crop Science 38, 12781289.
Luo, M., Bilodeau, P., Koltunow, A., Dennis, E.S., Peacock, W.J. and Chaudbury, A.M. (1999)
Genes controlling fertilization-independent seed development in Arabidopsis thaliana.
Proceedings of the National Academy of Sciences USA 96, 296301.
Lynam, J. (1998) Evaluation Findings. Part 1. Crop-based Production System Projects. Cassava
Evaluation Report. IITA, Ibadan, Nigeria.
Mackill, D.J. and Bonman, J.M. (1992) Inheritance of blast resistance in near-isogenic lines of
rice. Philippines.
Magill, C.W., Boora, K., Sunitha Kumari, R., Osorio, J., Oh, B.J., Gowda, B., Chui, Y. and
Fredricksen, R. (1997) Tagging sorghum genes for disease resistance: expectations and
reality. In: INTSORMIL/CRISAT. Proceedings of the International Conference on Genetic
Improvement of Sorghum and Pearl Millet. Meeting held in Lubbock, Texas, September 2227 1997.
Malawi Government (1999) Ministry of Agriculture and Irrigation, Department of Agriculture
Research and Technical Services. Rebuilding the Malawi Maize Pathology Project and
Development of a Research Action Plan. Final Report, June 1999. Lilongwe, 47 pp.
Mann, C. (1999) Starter Pack Scheme Assessment Report. Project evaluation document prepared for
The Rockefeller Foundation, New York, 36pp.
Mann, C.G. (1999) Kenya: biotechnology in Africa: why the controversy? In: Persley, G.J. and
Lantin, M.M. (eds) Agricultural Biotechnology and the Poor. Proceedings of an International
Conference. Washington, DC, 2122 October 1999, pp. 109114.
185
Chapter
186
References
Maredia M.K., Byerlee, D. and Pee, P. (2000) Impacts of food crop improvement research:
evidence from sub-Saharan Africa. Food Policy 25, 531559.
Mashingaidze, K. (1994) Maize research and development. In: Rukini, M. and Eicher, C.K. (eds)
Zimbabwes Agricultural Revolution. University of Zimbabwe Publications, Harare.
Matlon, P. and Adesina, A. (1991) Prospects for sustainable growth in sorghum and millet productivity in West Africa. In: Vosti, S.A., Reardon, T., von Urff, W. and Witcover, J.
(eds) Agricultural Sustainability, Growth and Poverty Alleviation: Issues and Policies. Proceedings
of the Conference held from 23 to 27 September, 1991 in Feldafing, Germany, pp. 363387.
May, G., Afza, R., Mason, H., Wiecko, A., Novak, F. and Arntzen, C. (1995) Generation of
transgenic banana (Musa acuminata) plants via Agrobacterium-mediated transformation.
Bio/Technology 13, 486492.
McCarter, B. (2000) Can Biotechnology Bridge the Gap for Resource Poor Farmers? Occasional paper.
Seed Co Ltd, Harare, Zimbabwe.
McCouch, S.R., Chen XiuLi, Panaud, P., Temnykh, S., Xu YunBi, Cho YongGu, Huang Ning,
Ishii, T. and Blair, M. (1997) Microsatellite marker development, mapping and
applications in rice genetics and breeding. Plant Molecular Biology 35, 8999.
Mduruma, Z.O. (1999) Lessons, experiences and future challenges of community-based
seed production in Tanzania. In: Maize Production Technology for the Future: Challenges and
Opportunities: Proceedings of the sixth eastern and southern Africa regional maize conference, 2125
September, 1998, Addis Ababa, Ethiopia. CIMMYT and EARO, pp. 150154.
Meikle, W.G. and Markham, R.H. (1998) Evaluating postharvest resistance to maize pests. In:
Project 5: Integrated Management of Maize Pests and Diseases. Annual Report 1998. International
Institute of Tropical Agriculture, Ibadan, Nigeria, p. 13.
Melton, A., Ogle, W.L., Barnett, O.W. and Caldwell, J.D. (1987) Inheritance of resistance to
viruses in cowpea. Phytopathology 77, 642.
Merril-Sands, D. (1986) Farming systems research: Clarification of terms and concepts.
Experimental Agriculture 22, 87104.
Miflin, B.J. (2000) Crop biotechnology. Where Now? Meeting report. Plant Physiology 123, 1727.
Minja, E.M. (2000) Management of the armoured bush cricket in Namibia and Zambia:
Farmers methods. In: Minja, E.M. and van den Berg, J. (eds) Proceedings of the Workshop on
Management of Sorghum and Pearl Millet Pests in the SADC Region, 1013 February 1998,
Matopos Research Station, Zimbabwe. ICRISAT, Bulawayo, Zimbabwe.
Mlotshwa, S. (2000) The helper component-proteinase of cowpea aphid-borne mosaic virus. PhD
dissertation, University of Zimbabwe, 111 pp.
Mloza-Banda, H.R., Kapondamgaga, P.H. and Kaotcha, R.M. (1999) Interim Research Report of the
Striga Research Project. Bunda College of Agriculture, Lilongwe, Malawi, 24 pp.
Monti, L.M., Murdock, L.L. and Thottappilly, G. (1997) Opportunities for biotechnology in
cowpea. In: Singh, B.B., Mohan Raj, D.R., Dashiell, K.E. and Jackai, L.E.N. (eds) Advances
in Cowpea Research. Copublication of International Institute of Tropical Agriculture (IITA)
and Japan International Research Center for Agricultural Sciences (JIRCAS). IITA, Ibadan,
Nigeria, pp. 341351.
Moore Lapp, F., Collins, J. and Rosset, P. (1998) World Hunger: 12 Myths, 2nd edn. Grove
Press/Earthscan.
Morris, M.L. (1998a) Overview of the world maize economy. In: Morris, M.L. (ed.) Maize Seed
Industries in Developing Countries. Lynne Rienner, London, pp. 1335.
Morris, M.L. (1998b) Maize in the developing world: waiting for a green revolution. In: Morris,
M.L. (ed.) Maize Seed Industries in Developing Countries. Lynne Rienner, London.
Mugo, S. (2000) Presentation of IRMA Project goals, objectives, and activities. In: Mugo, S.,
Poland, D., DeGroote, H. and Hoisington, D. (eds) Proceedings of the Stakeholders Meeting held
at Panafric Hotel, Nairobi, Kenya. March 3 2000. IRMA Project Document No. 2.
186
Chapter
187
References
Muhhuku, F. (2000) Farmer Demand for Seed of Improved Varieties: Workshop on the development
of sustainable seed systems for small-scale farmers. NARO, Mukono, Uganda, 2223 May,
2000.
Mukuru, S.Z. (1992) Breeding for grain mold resistance. In: de Milliano, W.A.J., Fredricksen,
R.A. and Bengston, G.D. (eds) Sorghum and Millet Diseases: A Second World Review.
ICRISAT, Patancheru, India, pp. 273285
Mukuru, S.Z. (1993) Sorghum and millet in eastern Africa. In: Byth, D.E. (ed.) Sorghum and
Millets Commodity and Research Environments. ICRISAT, Patancheru, Andhra Pradesh,
India.
Murdock, L.L., Shade, R.E., Kitch, L.W., Ntoukam, G., Lowenberg-DeBoer, J., Huesing, J.E.,
Moar, W., Chambliss, O.L., Endondo, C. and Wolfson, J.L. (1997) Postharvest storage of
cowpea in sub-Saharan Africa. In: Singh, B.B., Mohan Rah, D.R., Dashiell, K.E. and Jackai,
L.E.N. (eds) Advances in Cowpea Research. Copublication of International Institute of
Tropical Agriculturee (IITA) and Japan International Research Center for Agricultural
Mwangi, W. (1996) The Economics of Commercial Maize Seed Supply. Module 9 in CIMMYT Seed
Manual. CIMMYT, Mexico, DF.
Nankam, C., Sallah, A. and Nhunga, M. (1996) WV Angola Agricultural Recovery Program
Technical Report of Activities (First Season, 19951996). World Vision International, Luanda,
Angola.
NAS (National Academy of Sciences) (1996) Lost Crops of Africa. Vol. 1, Grains. National
Academy Press, Washington, DC.
Ndiritu, C.G. (1999) Kenya: biotechnology in Africa: why the controversy? In: Persley, G.J. and
Lantin, M.M. (eds) Agricultural Biotechnology and the Poor. Proceedings of an International
Conference. Washington, DC, 2122 October 1999, pp. 109114.
Ndjiondiop, M.N., Albar, L., Fargette, D., Fauquet, C. and Ghesquiere, A. (1999) The genetic
basis of high resistance to rice yellow mottle virus (RYMV) in cultivars of two cultivated
rice species. Plant Disease (in press).
Ng, N.Q. (1995) Cowpea. In: Smartt, J. and Simmonds, N.W. (eds) Evolution of Crop Plants, 2nd
edn. Longman, Harlow, UK, pp. 326332.
Nguyen, H.T., Xu, W., Rosenow, D.T., Mullet, J.E. and McIntyre, L. (1996) Use of biotechnology
in sorghum breeding. In: Proceedings of the International Conference on Genetic Improvement
of Sorghum and Pearl Millet. Meeting held in Lubbock, Texas, September 2227 1997.
Ngwira, P. (1989) The status of maize diseases in Malawi. In: Gebrekidan, B. (ed.) Maize Improvement, Production and Protection in Eastern and Southern Africa. Proceedings of the Third
Eastern and Southern Africa Regional Maize Workshop. Govt. of Kenya and CIMMYT,
Nairobi and Kitale, Kenya, September 1822, 1989, pp. 230238.
Ngwira, P.N., Pixley, K.P., DeVries, J.D. and Kanaventi, C.M. (1998) Major maize disease
problems and farmers varietal preferences in Malawi. In: Proceedings of the Sixth Eastern and
Southern Africa Regional Maize Conference, held in Addis Ababa, 2125 Sept., 1998.
Nhlane, W.G. (1990) Breeding flint maize hybrids (hard endosperm grain) in Malawi in
response to smallholder processing needs. In: Gebrekidan, B. (ed.) Maize improvement, production and protection in Eastern and Southern Africa: Proceedings of the third eastern and southern
Africa regional maize workshop. Nairobi, Kenya.
Nichols, R.F.W. (1947) Breeding cassava for virus resistance. East African Agriculture Journal 12,
184194.
Nkama, I. and Malleshi, N.G. (1998) Production and nutritional quality of traditional Nigerian
masa from mixtures of rice, pearl millet, cowpea and groundnut. Food and Nutrition
Bulletin 19, 366373.
187
Chapter
188
References
Norskog, C. (1995) Hybrid Seed Corn Enterprises: a Brief History, 1st edn. Maracom Corp.,
Willmar, Minnesota.
Novak, F.J., Afza, R., van Duren, M., Perea-Dallos, M., Conger, B.V. and Xiaolang, T. (1989)
Somatic embryogenesis and plant regeneration in suspension cultures of dessert (AA and
AAA) and cooking (ABB) bananas (Musa spp.). Bio/Technology 7, 154159.
Nweke, F.I., Ugwu, B.O. and Dixon, A.G.O. (1992) The Spread and Performance of Improved
Cassava Varieties in Nigeria: An assessment of Adoption. Collaborative Study of Cassava in
Africa. Working paper no. 15. First Draft.
Nweke, F.I., Poulson, R. and Strauss, J. (1994) Cassava production trends in Africa. In: Ofori, F.
and Hahn, S.K. (eds) Tropical Roots Crops in a Developing Economy. Proceedings of the 9th
Symposium of the International Society for Tropical Roots Crops, 2026 October, 1991,
Accra, Ghana. Govt of Ghana, pp. 311321.
Obilana, A.B., Monyo, E.S. and Gupta, S.C. (1996) Impact of genetic improvement in sorghum
and pearl millet: developing country experiences. In: Proceedings of the International
Conference on Genetic Improvement of Sorghum and Pearl Millet. Meeting held in Lubbock,
Texas, September 2227 1997. INTSORMIL/ICRISAT, pp. 119141.
Odame, H. and Kameri-Mbote, P. (2000) Agricultural biotechnology assessment in sub-Saharan
Africa. Country specific study Kenya. Paper prepared for the Regional Workshop on Building
National Biotechnology Innovation Systems: New Forms of Institutional Arrangements and
Financial Mechanisms. African Centre for Technology Studies, Nairobi, Kenya.
ODI (1996) Good Practice Review 4: Seed Provision During and After Emergencies. Overseas
Development Institute, London, 137 pp.
Oh, B.J., Frederiksen, R.A. and Magill, C.W. (1996) Identification of RFLP markers linked to a
gene for downy mildew resistance (Sdm) in sorghum. Canadian Journal of Botany 74, 315.
Okali, C., Sumberg, J. and Farrington, J. (1994) Farmer Participatory Research: Rhetoric and Reality.
Intermediate Technology, London, 159 pp.
Olsen, K.M. and Schaal, B.A. (1999) Evidence on the origin of cassava: phylogeography of
Manihot esculenta. Proceedings of the National Academy of Sciences USA 96, 55865591.
Onim, M. (1998) Multi-location multiplication of ACMV-resistant cassava varieties in western Kenya.
Report to the KARI/IITA Cassava Steering Committee. Lagrotech Consultants, Kisumu, Kenya,
22 pp.
Ortiz, R. and Vuylsteke, D. (1994) Plantain breeding at IITA. In: Jones (ed.) The Improvement and
Testing of Musa: A Global Partnership. Proceedings of the first global conference of the
International Musa Testing Program held at FHIA, Honduras, 2730 April, 1994. INIBAP,
Montpellier, France, pp. 6376.
Osborn, T. (1996) Seeds for Disaster Mitigation and Recovering in the Greater Horn of Africa. USAID
Office of Foreign Disaster Assistance (OFDA), Washington, DC, 137 pp.
Otim-Nape, G.W., Bua, A. and Thresh, J.M. (1997) Progress in cassava technology transfer in
Uganda. In: Proceedings of the National Workshop on Cassava Multiplication, Masindi, Uganda,
January, 1996. NARO/NRI Publication, 139 pp.
Ou, S.H. (1985) Rice Diseases. Kew Institute, London, 368 pp.
Oyejide, T.A. (1993) Effects of trade and macroeconomic policies on African Agriculture. In:
Bautista, R.M. and Valdes, A. (eds) The Bias Against Agriculture: Trade and Macroeconomic
Policies in Developing Countries. International Center for Economic Growth, San Francisco,
California, pp. 241262.
Ozias-Akins, P.E., Lubbers, L., Hanna, W.W. and McNay, J.W. (1993) Transmission of the
apomictic mode of reproduction in Penniserum: co-inheritance of the trait and molecular
markers. Theoretical and Applied Genetics 85, 632638.
Paarlberg, R.L. (2000) 2020 Vision Discussion Paper 33 Governing the GM crop revolution: Policy
choices for developing countries. IFPRI, Washington, DC.
188
Chapter
189
References
Padulosi, S. and Ng, N.Q. (1997) Origin, taxonomy and morphology of Vigna unguiculata (L.)
Walp. In: Singh, B.B., Mohan Raj, D.R., Dashiell, K.E. and Jackai, L.E.N. (eds) Advances in
Cowpea Research. Co-publication of IITA and Japan International Research Center for
Agricultural Sciences. IITA, Ibadan, Nigeria.
Palm, C.A. (1995) Contribution of agroforestry trees to nutrient requirements of intercropped
plants. Agroforestry Systems 30, 105124.
Palm, C.A., Myers, R.J.K. and Nandwa, S.M. (1997) Combined use of organic and inorganic
nutrient sources for soil fertility maintenance and replenishment. In: Replenishing Soil
Fertility in Africa. SSSA Special Publication Number 51. Proceedings of an international
symposium cosponsored by Divisions A-6 (International Agronomy) and S-4 (Soil Fertility
and Plant Nutrition), and the International Center for Research in Agroforestry, held at
the 88th Annual Meetings of the American Society of Agronomy and the Soil Science Society of America, Indianapolis, Indiana, 6 November 1996. Soil Science Society of America.
Madison, Wisconsin, USA, pp. 193217.
Pardey, P.G., Roseboom, J. and Anderson, J.R. (1991) Topical perspectives on national
agricultural research. In: Pardey, P.G., Roseboom, J. and Anderson, J.R. (eds) Agricultural
Research Policy: International Quantitative Perspectives. Cambridge University Press,
Cambridge, pp. 265308.
Pardey, P.G., Roseboom, J. and Beintema, N.M. (1997) Investments in African agricultural
research. World Development 25, 409423.
Patel, P.N. and Hall, A.E. (1988) Inheritance of heat-induced brown discoloration in seed coats
of cowpea. Crop Science 28, 929932.
Pereira, M.G., Lee, M., Bramel-Cox, P., Woodman, W., Doebley, J. and Whitkus, R. (1994)
Construction of an RFLP map in sorghum and comparative mapping in maize. Genome 37,
236243.
Persley, G.P. (1999) Biotechnology for Developing-Country Agriculture: Problems and Opportunities.
Letter to a Minister. Focus 2. Brief 1 of 10. October 1999. International Food Policy Research
Institute (IFPRI), Washington, DC.
Persley, G.P. and Doyle, J.J. (1999) Biotechnology for developing-country agriculture: Problems
and opportunities. IFPRI Focus 2. Brief 1. IFPRI: Washington, DC.
Phillips, R.L., Somers, D.A. and Hibberd, K.A. (1988) Cell/tissue culture and in vitro manipulation. In: Sprague, G.F. and Dudley, J.W. (eds) Corn and Corn Improvement, 3rd edn.
Agronomy Monograph No. 18. American Agronomy Association, Crop Science Society of
America, and Soil Science Society of America, Madison, Wisconsin.
Pingali, P., Bigot, Y. and Binswanger, H.P. (1987) Agricultural Mechanization and the Evolution
of Farming Systems in Sub-Saharan Africa. Johns Hopkins University Press, Baltimore,
Maryland.
Pinto, Y.M., Kok, R.A. and Baulcombe, D.C. (1999) Resistance to rice yellow mottle virus
(RYMV) in cultivated African rice varieties containing RYMV transgenes. Nature Biotech.,
17, 702707.
Pixley, K.V. (1996) CIMMYT mid-altitude maize breeding program Report of activities during
1995/96. CIMMYT-Harare Annual Report, 199596. CIMMYT, Harare, Zimbabwe.
Pixley, K.V. (1997) CIMMYT mid-altitude maize breeding programme. In: Jewell, D.C., Pixley,
K.V., Banziger, M., Waddington, S.R., Varughese, G., Zambezi, B.T. and Mekuria, M. (eds)
CIMMYT-Harare Annual Report, 199697. CIMMYT, Harare, Zimbabwe.
Pixley K.V. and Zambezi, B.T. (1996) Maize Germplasm Available from CIMMYT Zimbabwe.
CIMMYT, Harare, 30 pp.
Ploetz, R.C. (1990) Population biology of Fusarium oxysporum f. sp. Cubense. In: Ploetz, R.C.
(ed.) Fusarium Wilt of Banana. American Pathology Society Press, St Paul, Minnesota,
pp. 6376.
189
Chapter
190
References
Ploetz, R.C. (1994) Fusarium wilt and IMTP Phase II. In: The Improvement and Testing of Musa: a
Global Partnership. Proceedings of the first global conference of the International Musa Testing Program held at FHIA, Honduras, 2730 April, 1994. INIBAP, Montpellier, France, pp. 5769.
Poehlman, J.M. (1979) Breeding Field Crops, 2nd edn. AVI Publishing Company, Inc., Westport,
Connecticut.
Prakash, C.S. and Shivashankar, G. (1984) Inheritance of resistance to bacterial blight in cowpea.
Genetica Agraria 38, 110.
Prasad, B., Prabhu, M.S. and Shantamma, C. (1984) Regeneration of downy mildew resistant
plants from infected tissues of pearl millet cultured in vitro. Current Science 53, 816817.
Provvidenti, R. (1993) Genetics of resistance to viral diseases of bean. In: Kyle, M.M. (ed.)
Resistance to Viral Diseases of Vegetables: Genetics and Breeding. Timber Press, Portland, Oregon,
pp. 112152.
Quinones, M.A., Borlaug, N.E. and Dowswell, C.R. (1997) A fertilizer-based green revolution for
Africa. In: Replenishing Soil Fertility in Africa. Soil Science Society of America, Madison,
Wisconsin, pp. 8195.
Rabobank (1994) The World Seed Market. Report by the Rabobank Nederland, Netherlands.
Rachie, K.O. and Anand Kumar, K. (1994) Pearl millet improvement at ICRISAT An update.
Int. Sorghum Millet Newsletter 35, 129.
Rachie, K.O. and Majmudar, J.V. (1980) Pearl Millet. Pennsylvania State University Press,
University Park.
Rai, K.N., Anand Kumar, K., Andrews, D.J., Gupta, S.C. and Ouendeba, B. (1997) Breeding
pearl millet for grain yield and stability. In: Proceedings of the International Conference on
Genetic Improvement of Sorghum and Pearl Millet. Meeting held in Lubbock, Texas, September
2227 1997. INTSORMIL/ICRISAT, pp. 7183.
Rai, K.N., Andrews, D.J. and Rao, A.S. (2000) Feasibility of breeding male-sterile populations for
use in developing inter-population hybrids of pearl millet. Plant Breeding 119, 335339.
Rajaram, S., Singh, R.P. and Torres, E. (1998) CIMMYT approaches in breeding wheat for rust
resistance. In: Simmonds, N.W. and Rajaram, S. (eds) Breeding Strategies for Resistance to the
Rusts of Wheat. CIMMYT, Mexico DF.
Rathus, C., Adkins, A.L., Henry, R.J., Adkins, S.W. and Godwin, I.D. (1996) Progress towards
transgenic sorghum. In: Foale, M.A., Henzell, R.G. and Kneipp, J. (eds) Proceedings of the
3rd Australian Sorghum Conference, 2022 Feb 1996, Tamworth. Australian Institute of
Agricultural Science, Melbourne, Australia. Occasional Publication No. 93, pp. 409412.
Ratnadass, A. and Ajayi, O. (1995) Panicle insects pests in sorghum in West Africa. In: Nwanze,
N.F. and Youm, O. (eds) Panicle insect pests of sorghum and pearl millet. Proc. Int. Consult.
Workshop, 47 Oct. 1993. Niamey, Niger. ICRISAT, Andra Pradesh, India, pp. 2938.
Rattray, A.G.H. (1969) Advances and achievements in crop research. In: Proceedings of the
Conference on Research and the Farmer, Salisbury, Rhodesia, Sept. 1819, 1969. Dept of Research
and Specialist Services, Harare.
Rattunde, H.F.W., Weltzien, E., Bramel-Cox, P., Kofoid, K., Hash, C.T., Schipprack, W.,
Stenhouse, J.W. and Presterl, T. (1997) Population improvement of pearl millet and
sorghum: current research impact and issues for implementation. In: Proceedings of the
International Conference on Genetic Improvement of Sorghum and Pearl Millet. Meeting held in
Lubbock, Texas, September 2227 1997. INTSORMIL/ICRISAT, pp. 188212.
Ravallion, M. and Chen, S. (1997) What Can New Survey Data Tell Us about Recent Changes in
(Income) Distribution and Poverty. World Bank Economic Review.
Rawal, K.M. (1975) Natural hybridisation among weedy and cultivated Vigna unguiculata (L.)
Walp. Euphytica 24, 699707.
Reader, J. (1997) Africa: a biography of the continent. Hamish Hamilton, London.
190
Chapter
191
References
Redden, R.J., Dobie, P. and Gatehouse, A.M.R. (1983) The inheritance of seed resistance to
bruchids in cowpea. Australian Journal of Agricultural Research 34, 681695.
Remy, S., Francois, I., Schoofs, H., Panis, B., Cammue, B., Swennen, R. and Sagi, L. (1998)
Genetic transformation as a technology to create disease resistance in banana. Acta
Horticulturae, in press.
RenKow, M. (1993) Differential technology adoption and income distribution in Pakistan:
Implications for research resource allocation. American Journal of Agricultural Economics 75,
3343.
Ribaut, J.M., Hoisington, D.A., Deutsch, J.A., Jiang, C. and Gonzalez de Leon, D. (1996)
Identification of quantitative trait loci under drought conditions in tropical maize. 1.
Flowering parameters and the anthesis-silking interval. Theoretical and Applied Genetics 92,
905914.
Rohrbach, D.D. (1994) Improving farmer wellbeing in semi-arid areas. In: Jewell, D.C.,
Waddington, S.R., Ranson, J.R. and Pixley, K.V. (eds) Maize Research for Stress Environments.
Proceedings of the Fourth Eastern and Southern Africa Regional Maize Conference, held
at Harare, Zimbabwe, 28 March1 April 1994, pp. 296303.
Rohrbach, D.D. and Makhwaje, E. (1999) Adoption and impact of new sorghum varieties in
Botswana. Southern African Development Community (SADC)/International Crops
Research Institute for the Semi-Arid Tropics (ICRISAT), Sorghum and Millet Improvement Program (SMIP). Bulawayo, Zimbabwe. (Semiformal publication).
Rohrbach, D. and Malusalila, P. (2000) Developing rural retail trade of seed through small
packs. Paper presented at the Zimbabwe Seed Sector Stakeholders Meeting, 25 June 1999, Matopos
Research Station, Zimbabwe.
Rohrbach, D.D., Lechner, W.R., Ipinge, S.A. and Monyo, E.S. (1999) Impact from Investments in
Crop Breeding: the Case of Okashana 1 in Namibia. Impact Series no. 4. Patancheru 502 324,
Andra Pradesh, India, International Crops Research Institute for the Semi-Arid Tropics,
48 pp.
Rosenow, D.T. (1997) Host country program enhancement in Mali. In: INTSORMIL Annual
Report, 1997.
Rosenow, D.T., Ejeta, G., Clark, L.E., Gilbert, M.L., Henzell, T.G., Borrel, A.K. and Muchow,
R.C. (1997) Breeding for pre- and post-flowering drought stress resistance in sorghum.
In: Proceedings of the International Conference on Genetic Improvement of Sorghum and Pearl
Millet. Meeting held in Lubbock, Texas, September 2227 1997. INTSORMIL/ICRISAT,
pp. 400411.
Rowe, P. and Rosales, F. (1994) Musa breeding at FHIA. In: Jones (ed.) The Improvement and
Testing of Musa: a Global Partnership. Proceedings of the first global conference of the
international Musa testing program held in FHIA, Honduras, 2730 April 1994. INIBAP,
Montpellier, France, pp. 117129.
Rowe, P. and Rosales, F.E. (1996) Bananas and plantains. In: Janick, J. and Moore, J. (eds) Fruit
Breeding, Vol. 1, Tree and Tropical Fruits. John Wiley & Sons, New York, pp. 167211.
Rusike, J. and Eicher, C.K. (1997) Institutional innovations in the maize seed industry. In:
Byerlee, D. and Eicher, C.K. (eds) Africas Emerging Maize Revolution. Lynne Rienner,
London.
Sagi, L., Remy, S., Panis, B., Swennen, R. and Volckaert, G. (1994) Transient gene expression in
electroporated banana (Musa spp., cv. Bluggoe, ABB group) protoplasts isolated from
regenerable embryogenic cell suspensions. Plant Cell Reports 13, 262266.
Sagi, L., Panis, B., Remy, S., Schoofs, H., De Smet, K., Swennen, R. and Cammue, B. (1995)
Genetic transformation of banana (Musa spp.) via particle bombardment. Bio/Technology
13, 481485.
Sahrawat, K.L., Jones, M.P. and Diatta, S. (1999) The role of tolerant genotypes and plant
nutrients in the management of acid soil infertility in upland rice. Paper presented at the
191
Chapter
192
References
consultants meeting on the use of nuclear techniques to develop management practices for increasing
crop production and soil fertility in acid soils, March 13, 1999, FAO, Rome, Italy; IAEA Division
of Nuclear Techniques in Food and Agriculture, International Atomic Energy Agency,
Vienna, Austria.
Salifou, E.M. (1998) Experience and economics of seed production in Niger. Paper presented at
regional hybrid sorghum and pearl millet seed workshop, held in Niamey, Niger, September 28
October 2 1998.
Sanchez, P.A., Shepherd, K.D., Soule, M.J., Place, F.M., Buresh, R.J. and Izac, A.N. (1997) Soil
fertility replenishment in Africa: an investment in natural resource capital. In: Replenishing
Soil Fertility in Africa. Soil Science Society of America, Madison, Wisconsin, pp. 146.
Sarail, A.K. and Singh, V.P. (1999) Seed set potential and seed yield of A R combinations in
rice (Oryza sativa L.). Seed Research 27, 140145.
Sasser, J.N. and Freckman, D.W. (1987) A world perspective on nematology: the role
of the Society. In: Veech, J.A. and Dickson, D.W. (eds) Vistas on Nematology. Society of
Nematologists, Hyattsville, USA, pp. 714.
Sastray, J.G., Ramakrishna, W., Sivaramkrishnan, S., Thakur, R.P., Gupta, V.S. and Ranjekar,
P.K. (1995) DNA fingerprinting detects genetic variability in the pearl millet downy
mildew pathogen. Theoretical and Applied Genetics 91, 856861.
Schaffert, R.E., Alves, V.M.C., Bahia, A.F.C., Pitta, G.V.E., Santos, F.G. and de Oliveira, C.A.
(1999) Sorghum genetic resources with contrasting phosphorus efficiency. In: Workshop on
improving phosphorus acquisition efficiency in marginal soils, held on October 1722, 1999 in Sete
Lagoas, MG, Brazil. EMBRAPA, pp. XI: 113.
Schechert, A.W. (1997) Quantitative genetic and marker-based studies on the importance of
resistance of maize to Setosphaeria turcica in Kenya. PhD Thesis, University of Hohenheim.
Schopke, C., Chavarriaga, P., Fauquet, C.M. and Beachy, R.N. (1995) Cassava tissue culture
and transformation: improvement of culture media and the effect of different antibiotics
on leaf tissues. In: Roca, W.M. and Thro, A.M. (eds) Proceedings of the First International
Scientific Meeting of the Cassava Biotechnology Network. CIAT working document 123, pp.
140145.
Schopke, C., Taylor, N., Carcamo, R., Konan, N.K., Marmey, P., Henshaw, G., Beachy, R.N. and
Fauquet, C. (1996) Regeneration of transgenic cassava plants (Manihot esculenta Crantz)
from microbombarded embryogenic suspension cultures. Nature Biotechnology 14, 731735.
Scowcroft, W.R. (1996) Seeds of Hope Project Completion Report. CIAT, Cali, Colombia, 42 pp.
Scowcroft, W.R. and Polak Scowcroft, C.E. (1997) Seed Security: Disaster Response and Strategic
Planning. Australian Centre for Oil Seed Research, and Agriculture Australia Consultants,
Horsham, Victoria, Australia.
Scully, B.T. and Federer, W.T. (1993) Application of genetic theory in breeding for multiple viral
resistance. In: Kyle, M.M. (ed.) Resistance to Viral Diseases of Vegetables: Genetics and Breeding.
Timber Press, Portland, Oregon, pp. 167195.
Seed Co (2000) Seed Co Seed Manual 20002001. Seed Co Ltd, Harare, Zimbabwe, 28 pp.
Sen, A. (1981) Poverty and Famines. Clarendon Press, Oxford.
Serratos, J.A., Arnason, J.T., Blanco-Labra, A. and Mihm, J.A. (1994) Genetics of maize grain
resistance to maize weevil. In: Mihm, J.A. (ed.) Insect Resistant Maize. Proceedings of an
International Symposium, 27 Nov3 Dec., 1994. CIMMYT, Mexico DF, pp. 132138.
Shigemune, A. and Yoshida, T. (2000) Methods of anther culture of pearl millet and ploidy level
of regenerated plants. Japanese Journal of Crop Science 69, 224228.
Shull, G.H. (1908) The composition of a field of maize. American Breeders Association Reports 4,
296301.
Simmonds, N.W. (1962) The Evolution of the Bananas. Longman, London, 170 pp.
Simmonds, N.W. (1966) Bananas, 2nd edn. Longmans, London.
192
Chapter
193
References
Simmonds, N.W. (1991) Selection for local adaptation in a plant breeding programme.
Theoretical and Applied Genetics 82, 363367.
Singh, B.B. (1993) Cowpea breeding. In: Archival Report (19881992) of Grain Legume
Improvement Program. IITA, Ibadan, Nigeria, pp. 1053.
Singh, B.B. and Mohammed, S.G. (1998) Quantitative characterization of cropping systems in
the Sahel. In: Project 11: CowpeaCereals Systems Improvement in the Dry Savannas. Annual
Report 1998. International Institute of Tropical Agriculture, Ibadan, Nigeria, 8 pp.
Singh, B.B. and Olufano, O. (1998) Farmer for farmer diffusion of improved cowpea seeds. In:
Project 11: CowpeaCereals Systems Improvement in the Dry Savannas. Annual Report 1998.
International Institute of Tropical Agriculture, Ibadan, Nigeria, p. 36.
Singh, B.B., Emechebe, A.M. and Atokple, I.D.K. (1993) Inheritance of Alectra resistance in
cowpea genotype B 301. Crop Science 33, 7072.
Singh, B.B., Chambliss, O.L. and Sharma, B. (1996) Recent advances in cowpea breeding. In:
Singh, B.B., Mohan Raj, D.R., Dashiell, K.E. and Jackai, L.E.N. (eds) Advances in Cowpea
Research. Co-publication of IITA and Japan International Research Center for Agricultural
Sciences, IITA, Ibadan, Nigeria.
Singh, B.B., Mohan Raj, D.R., Dashiell, K.E. and Jackai, L.E.N. (eds) (1997) Advances in Cowpea
Research. Copublication of International Institute of Tropical Agriculture (IITA) and
Japan International Research Center for Agricultural Sciences (JIRCAS). IITA, Ibadan,
Nigeria.
Singh, S.D., Lal, S. and Pande, S. (1993) The changing scenario of maize, sorghum and pearl
millet diseases. In: Sharma, H.C. and Veerabhadra, M. (eds) Pests and Pest Management
in India The Changing Scenario. Plant Protection Association of India, Rajendragar,
Hyderabad, Andhra Pradesh 500030, India, pp. 130139.
Sitch, L. (1996) Fate of seed/planting material of farmer selected varieties distributed through
World Visions extension program. Monograph. World Vision of Mozambique, Maputo,
Mozambique.
Sithole-Niang, I. (2000) Cowpea improvement through genetic engineering. Project Proposal.
University of Zimbabwe, Harare.
Smale, M. and Heisey, P.W. (1997) Maize technology and productivity in Malawi. In: Byerlee, D.
and Eicher, C.K. (eds) Africas Emerging Maize Revolution. Lynne Rienner Publishers,
Boulder, Colorado.
Smale, M., Kaunda, Z.H.W., Makina, H.L., Mkandawire, M.M.M.K., Msowoya, M.N.S., Mwale,
D.J.E.K. and Heisey, P.W. (1991) Chimanga Cha Makolo, in hybrids and composites: An
analysis of farmers adoption of maize technology in Malawi, 19891991. CIMMYT Economics
Working Paper 91/04. International Maize and Wheat Improvement Center (CIMMYT),
Mexico DF.
Smale, M., Kaunda, Z.H.W., Makina, H.L. and Mkandawire, M.M.M.K. (1993) Farmers
evaluation of newly released maize cultivars in Malawi: a comparison of local maize, semi-flint and
dent hybrids. International Maize and Wheat Improvement Center (CIMMYT), Lilongwe
and Harare.
Smale, M., Kaunda, H.W., Makina, H.L. and Mkandawire, M.M.M.K. (1994) Farmers evaluation of newly released maize cultivars in Malawi: a comparison of local maize, semi-flint
and dent hybrids. CIMMYT, Mexico, DF.
Smil, V. (1991) Population growths and nitrogen: an exploration of a critical existential link.
Population and Development Review 17, 569601.
Smith, D.C. (1966) Plant breeding development and success. In: Frey, K.J. (ed.) Plant Breeding.
Iowa State University Press, Ames, Iowa, p. 32.
Smith, J. (1993) Targetting hybrid maize to appropriate agricultural systems in the northern
guinea savannah of West Africa. Unpublished paper. IITA, Ibadan, Nigeria.
193
Chapter
194
References
Smith, J., Weber, G., Manyong, M.V. and Fakorede, M.A.B. (1997) Fostering sustainable
increases in maize productivity in Nigeria. In: Byerlee, D. and Eicher, C.K. (eds) Africas
Emerging Maize Revolution. Lynne Rienner, Boulder, Colorado, pp. 107126.
Song, W.Y., Wang, G.L., Chen, L.L., Kim, H.S., Pi, L.Y., Holsten, T., Gardner, J., Wang, B., Zhai,
W.X., Zhu, L.H., Fauquet, C. and Ronald, P. (1995) A receptor kinase-like protein encoded
by the rice disease resistance gene, Xa21. Science 270, 18041806.
Speijer, P.R. and De Waele, D. (1997) Screening of Musa germplasm for resistance and tolerance to nematodes. INIBAP Technical Guidelines 1. IPGRI, Rome, Italy; INIBAP, Montpellier, France.
Spencer, D. (1986) Agricultural research: lessons of the past, strategies for the future. In: Berg,
R.J. and Whitaker, J. (eds) Strategies for African Development. University of California Press,
Berkeley, pp. 182214.
Spencer, D.S.C. and Edwin, J. (1999) Assessment of the Prospects of the WARDA Interspecific Rice
Varieties in West Africa. Consultancy report compiled for The Rockefeller Foundation, New
York, 32 pp.
Sperling, L. (1994) Analysis of Bean Seed Channels in the Great Lakes Region: South Kivu, Zaire, Southern Rwanda, and Select Bean-Growing Areas of Burundi. Occasional Publications Series, No. 13.
CIAT/RESAPAC, Butare, Rwanda.
Sperling, L. and Loevinsohn, M. (1996) Using Diversity: Enhancing and Maintaining Genetic
Resources On-farm. IDRC, New Delhi, India.
Srivastava, J.P. and Jaffee, S. (1993) Best Practices for Moving Seed Technology: New Approaches
to Doing Business. World Bank Technical Paper No. 176. World Bank, Washington, DC,
36 pp.
Stenhouse, J.W., Bandyopadhyay, R., Singh, S.D. and Subramanian, V. (1996) Breeding for grain
mold resistance in sorghum. In: Proceedings of the International Conference on Genetic Improvement of Sorghum and Pearl Millet. Meeting held in Lubbock, Texas, September 2227 1997.
Stevens, R.D. and Jabara, C.L. (1988) Agricultural Development Principles. Johns Hopkins
University Press, Baltimore, Maryland, 477 pp.
Sthapit, B.R., Joshi, K.D. and Witcombe, J.R. (1995) Farmer participatory high altitude
rice breeding in Nepal: providing choice and utilizing farmers expertise. In: Sperling L.
and Loevinsohn, M. (eds) Using Diversity Enhancing and Maintaining Genetic Resources
On-farm. International Development Research Centre (IDRC), New Delhi, India, pp.
186205.
Storey, H.H. and Nichols, R.F.W. (1938) Studies of the mosaic diseases of cassava. Annals of
Applied Biology 25, 790806.
Stover, R.H. and Simmonds, N.W. (1987) Bananas. 3rd edn. Longmans, London.
Stuber, C.W. (1991) Biochemical and molecular markers in plant breeding. Plant Breeding
Reviews 9, 3761.
Sustainable Community-oriented Development Programme (SCODP) (1999) News Brief, June
1999. SCODPs Seed Mini-pack: a tool to improve small farmers access to improved seeds
in Kenya. In: SCODP, Phase 2. Further development of SCODPs farm input supply
business in Western Kenya: Promotion of the use of fertilizer and seed amongst small farmers in Kenya. Project Proposal. Sega, Kenya.
Swennen, R. and Vuylsteke, D. (1991) Bananas in Africa: diversity, uses and prospects for
improvement. In: Ng, N.Q., Perrino, P., Attere, F. and Zedan, H. (eds) Crop Genetics
Resources of Africa: Proceedings of an International Conference held in Ibadan, Nigeria, 1720 Oct.
1988. IITA/IBPGR/UNEP/CNR, Ibadan, Nigeria.
Swennen R., Vuylsteke, D. and Hahn, S.K. (1989) Combating the black Sigatoka threat to
plantains. IITA Research Briefs 9(2), 24. Ibadan, Nigeria.
194
Chapter
195
References
Talukdar, B.S. and Singh, S.D. (1993) Stability and heritability for downy mildew resistance in
two pearl millet pollinators. In: Annual Report 1992. Cereals Program ICRISAT, Patancheru,
India, pp. 7071.
Talukdar, B.S., Singh, S.D. and Hash, C.T. (1994) Breeding for resistance to diseases in pearl
millet. In: Singh, H.G., Mishra, S.N., Singh, T.B., Hari Har Ram and Singh, D.P. (eds)
Crop Breeding in India. International Book Distributing Company, Lucknow, India,
pp. 176185.
Taneja, S.L. and Leuschner, K. (1984) Resistance screening and mechanisms of resisitance in
sorghum to shoot fly. In: Proceedings of the International Sorghum Entomology Workshop, 1521
July, 1984. Texas A & M University College Station, Texas, pp. 115129.
Tanksley, S.D. (1993) Mapping polygenes. Annual Review of Genetics 27, 205234.
Tanksley, S.D., Young, N.D., Paterson, A.H. and Bonierbale, M.W. (1989) RFLP mapping in
plant breeding: New tools for an old science. Bio/Technology 7, 257264.
Tao, Y.Z., McIntyre, C.L. and Henzell, R.G. (1996) Applications of molecular markers to
Australian sorghum breeding programs. Proceedings of the Third Australian Sorghum Conference.
Taylor, N.J., Edwards, M., Kiernan, R.J., Davey, C.D.M., Blakesly, D. and Hesnshaw, G.G. (1996)
Development of friable embryogenic callus and embryogenic suspension culture systems in
cassava (Manihot esculenta Crantz). Nature Biotechnology 14, 726730.
Tenkouano, A., Miller, F.R., Fredricksen, R.A. and Rosenow, D.T. (1993) A single locus with
multiple alleles as the genetic basis of anthracnose resistance in sorghum. Theoretical and
Thakur, R.P. and Chahal, S.S. (1987) Problems and strategies in the control of ergot and smut in
pearl millet. In: Whitcombe, J.R. and Beckerman, S.R. (eds) Proceedings of the International
Pearl Millet Workshop, 711 April, 1986, ICRISAT, Andra Pradesh, India, pp. 147160.
Thakur, R.P., Talukdar, B.S. and Rao, V.P. (1983) Genetic of ergot resistance. In: Abstracts of
contributed papers of the XV International Congress of Genetics, 1221 Dec., 1983. New Delhi,
p. 737.
Thakur, R.P., Frederiksen, R.A., Murty, D.S., Reddy, B.V.S., Bandyopadhyay, R., Giorda, L.M.,
Odvody, G.N. and Claflin, L.E. (1997) Breeding for disease resistance in sorghum. In:
Proceedings of the International Conference on genetic Improvement of Sorghum and Pearl Millet.
Meeting held in Lubbock, Texas, September 2227 1997. INTSORMIL/ICRSAT, pp. 303336.
Thotapilly, G., Mignouna, J., Ilori, C.O. and Fowale, I. (1998) Pollen electrotransformation in
cowpea (Vigna unguiculata [L]Walp). In: Project 15: Molecular and Cellular Biotechnology for
Crop Improvement. Annual Report 1998. International Institute of Tropical Agriculture,
Ibadan, Nigeria, pp. 89.
Thro, A.M. and Spillane, C. (2000) Biotechnology-assisted Participatory Plant Breeding: Complement
or Contradiction? Working Document No. 4. CGIAR Systemwide Program on Participatory
Research and Gender Analysis for Technology Development and Institutional Innovation.
Timko, M. (2001) The cowpea genetic map and tools for breeders. Paper presented at a symposium
on the genetic improvement of cowpea, January 812, 2001, Dakar, Senegal.
Toenniessen, G.H. (1995) Plant biotechnology and developing countries. Tibtech 13, 404409.
Toenniessen, G.H. (1998) Plant biotechnology: potential for sustainable development. Paper
presented at 4th Asia-Pacific Conference on Agricultural Biotechnology. Darwin, Australia, July
1316, 1998.
Toure, K. and Yehouenou, A. (1995) Les insectes de lepi de mil en Afrique de louest. In:
Nwanze, K.F. and Youm, O. (eds) Panicle Pests of Sorghum and Pearl Millet: Proceedings of an
International Consultative Workshop, 47 Oct., 1993, ICRISAT sahelian Center, Niamey, Niger.
ICRISAT, Patancheru 502 324, Andra Pradesh, India, 320 pp.
Tour, A., Rattunde, H.F.W. and Akintayo, I. (1998) Sorghum hybrids in West Africa.
Conference Paper presented at Regional Hybrid Sorghum and Pearl Millet Seed Workshop,
Niamey, Niger, Sept. 28Oct 2 1998.
195
Chapter
196
References
Tripp, R. (2000) Strategies for Seed System Development in Sub-Saharan Africa: A study of Kenya,
Malawi, Zambia, and Zimbabwe. Working Paper Series no. 2. Socioeconomic and Policy
Program, International Crops Research Institute for the Semi-Arid Tropics. Bulawayo,
Zimbabwe, 50 pp.
Tripp, R. and Louwaars, N. (1997) Seed regulation: choices on the road to reform. Food Policy 22,
433446.
Tripp, R. and Rohrbach, D. (2000) Policies for African Seed Enterprise Development.
(Unpublished).
Trutmann, P. (1996) Participatory diagnosis as an essential part of participatory breeding: A
plant protection perspective. In: Eyzaguirre, P. and Iwanaga, M. (eds) Participatory Plant
Breeding. IPGRI, Rome.
Tshiunza, M., Okechukwu, R.U. and Dixon, A.G.O. (1999) Status of cassava in semiarid zones of
Central and West Africa. COSCASA Working Paper No. 1. Collaborative study of cassava in
the semiarid zone of Africa. International Institute of Tropical Agriculture, Ibadan,
Nigeria.
Tyagi, A.K., Mohanty, A., Bajaj, S., Chaudhury, A. and Maheshwari, S.C. (1999) Transgenic rice: A
valuable monocot system for crop improvement and gene research. Centre for Plant Molecular
Biology and Department of Plant Molecular Biology, University of Delhi, New Delhi,
India.
Uganda, Ministry of Finance, Planning and Economic Development (2000) Ugandas Poverty
Eradication Action Plan: Summary and Main Objectives. Poverty reduction strategy paper.
Kampala, Uganda, 67 pp.
UN/IFPRI (2000) Chapter 1: Nutrition throughout the life cycle. In: Fourth Report on the World
Nutrition Situation. Geneva, Switzerland: United Nations Administrative Committee on
Coordination/Sub-Committee on Nutrition and the International Food Policy Research
Institute, Geneva and Washington, DC.
UPWARD (1996) Into Action Research: Partnerships in Asian Rootcrop Research and Development.
UPWARD, Los Baos, Philippines.
University of California, Riverside (1994) 1993 Executive Summary: A program to develop
improved cowpea cultivars, management methods, and storage practices for semi-arid
zones, 15 pp.
USAID (1996) Seeds for Disaster Mitigation and Recovery in the Greater Horn of Africa. Report
prepared by Chemonics International and USDA Famine Mitigation Activity, 135 pp.
USAID (2000) United States International Food Assistance: Report for 1999. USAID, Washington,
DC, 77 pp.
USDA (1998) Agriculture Fact Book 1998. Washington, DC.
Van Rensburg, J.J. and Gevers, H. (1993) Inheritance of antibiosis to the maize stalk borer,
Buseola fusca. South African Journal of Plant and Soil 10, 3540.
Vasil, V. and Vasil, K. (1980) Isolation and culture of cereal protoplasts. II. Embryogenesis
and plantlet formation form protoplasts of pearl millet. Theoretical and Applied Genetics
56100.
Veille Calzada, J.P., Crane, C.F. and Stelly, D.M. (1996) Apomixis: the asexual revolution. Science
274, 13221323.
Veldhuizen, L. et al. (1997) Farmers Research in Practice: Lessons from the Field. Intermediate
Technology, London.
Venkatesan, V. (1994) Seed Systems in Sub-Saharan Africa: Issues and Options. World Bank Discussion
Paper No 266. World Bank, Washington, DC, 112 pp.
Verdiaer, V., Boher, B., Maraite, H. and Geiger, J.P. (1994) Pathological and molecular characterization of Xanthomonas campestris strains causing diseases of cassava. Applied Environmental
Microbiology 60, 44784486.
196
Chapter
197
References
Vuylsteke, D., Ortiz, R. and Swennen, R. (1992) Plantains and bananas. In: Sustainable Food
Production in Sub-Saharan Africa. IITA, Ibadan, Nigeria, pp. 8691.
Vuylsteke, D., Ortiz, R., Pasberg-Gauhl, C., Gold, C., Ferris, S. and Speijer, P. (1993) Plantation
and banana research at the International Institute of Tropical Agriculture. HortScience 28,
873874.
Vuylsteke, D., Swennen, R. and De Langhe, E. (1996) Field performance of somaclonal variants
of plantain (Musa spp., AAB group). Journal of the American Society of Horticultural Science
121, 4246.
Vuylsteke, D., Ortiz, R., Ferris, S. and Crouch, J. (1997) Plantain improvement. Plant Breeding
Reviews 14, 267320.
Vuylsteke, D., Hartman, J., Tenkouano, A., Van de hauwe, I. and INIBAP (1998) International
dissemination of improved hybrids. In: Project 7: Improving Plantain- and Banana-based
Systems. Annual Report 1998. International Institute of Tropical Agriculture, Ibadan,
Nigeria, pp. 5354.
Waddington, S.R. and Karigwindi, J. (1995) CIMMYT maize agronomy in southern Africa.
In: Annual Research Report. November 1993 to October 1994. CIMMYT-Zimbabwe, Harare,
80 pp.
Waddington, S.R., Edmeades, G.O., Chapman, S.C. and Barreto, H.J. (1994) Where to with
agricultural research for drought-prone environments? In: Jewell, D.C., Waddington, S.R.,
Ransom, J.K. and Pixley, K.V. (eds) Maize Research for Stress Environments. Proceedings of the
Fourth Eastern and Southern Africa Regional Maize Conference, Harare, Zimbabwe, 28 March
1 April, 1994, pp. 129151.
Walker, D.J. and Tripp, R. (1997) Seed management by small-scale farmers in Ghana and
Zambia. In: FAO Seed Conference held in Rome, Italy, 1997. FAO, Rome, Italy.
Wang, R., Guegler, K., LaBrie, S.T. and Crawford, N.M. (2000) Genomic analysis of a nutrient
response in Arabidopsis reveals diverse expression patterns and novel metabolic and
potential regulatory genes induced by nitrate. Plant Cell 12, 14911510.
WARDA (1997) Medium Term Plan 19982000: For Presentation to the Technical Advisory
Committee, Consultative Group on International Agricultural Research, Rome, Italy.
WARDA, Bouak, Cte dIvoire
WARDA (1998) Annual Report. West Africa Rice Development Association, Bouake, Cte
dIvoire.
WARDA (1999) Rice Interspecific Hybridization Project: Research Highlights 1999. WARDA, Bouak,
Cte dIvoire
WARDA (1999) The Spark that Lit the Flame. West African Rice Development Association,
Bouake, Cte dIvoire.
Weinmann, H. (1975) Agricultural Research and Development in Southern Rhodesia: 19241950.
Series in Science, No. 2. University of Rhodesia, Salisbury.
Wellman, F.L. (1968) More diseases on crops in the tropics than in the temperate zone. Ceiba 14,
1728.
Welz, H.G., Schechert, A., Pernet, A., Pixley, K.V. and Geiger, H.H. (1998) A gene for resistance
to the maize streak virus in CIMMYT maize inbred line CML 202. Molecular Breeding 4,
147154.
White, J. and Sitch, L. (1994) 1993/94 Yield Trials: Tete, Zambezi, Sofala and Nampula
Provinces. World Vision International and Instituto Nacional de Investigacao
Agronomica, Maputo, Mozambique, 63 pp.
Widawsky, D.A. and OToole, J.C. (1990) Prioritizing the Rice Biotechnology Research Agenda for
Eastern India. The Rockefeller Foundation, New York, 86 pp.
Wiggins, S. (2000) Interpreting changes from the 1970s to the 1990s in African agriculture
through village studies. In: Streeten, P.P. and Craswell, J. (eds) World Development 28,
631662.
197
Chapter
198
References
Wilson, G.F. (1988) Plantain in Western Africa: report of a mission organized by INIBAP and
sponsored by IFAD, IITA and CIRAD. Montpellier, France: INIBAP, 68 pp.
Wilson, G.F. and Buddenhagen, I.W. (1986) The black Sigatoka threat to plantain and banana in
West Africa. IITA Research Briefs 7, 3.
Winkelmann, D. (1994) Quintessential internationalism in agricultural research. In: Anderson,
J.R. (ed.) Agricultural Technology: Policy Issues for the International Community. CAB International, Wallingford, UK.
Witcombe, J.R. (2000) The impact of participatory plant breeding and other breeding strategies
on the genetic base of crops. In: Cooper, H.D., Hodgkin, T. and Spillane, C. (eds)
Broadening the Genetic Base of Crop Production. CAB International, Wallingford, UK.
Witcombe, J.R. and Hash, C.T. (2000) Resistance gene deployment strategies in cereal hybrids
using marker-assisted selection: gene pyramiding, three-way hybrids, and synthetic parent
populations. Euphytica 112, 175186.
Woo, S.S., Jiang, J., Gill, B.S., Paterson, A.H. and Wing, R.A. (1994) Construction and characterization of a bacterial artificial chromosome library for sorghum. Nucleic Acids Research 22,
49224931.
World Bank (1998) www.worldbank.org
World Vision International (1995) Southern Sudan Agricultural Recovery Program. 1995 Annual
Report, Janson, G. and Kapukha, P. (eds). Nairobi, Kenya.
Wortmann, C.S. (1998) In: Wortmann, C.S., Kirkby, R.A., Eledu, C.A. and Allen, D.J. (eds)
Atlas of Common Bean (Phaseolus vulgaris L.) Production in Africa. CIAT, Cali, Columbia.
Wright, M., Donaldson, T., Cromwell, E. and New, J. (1994) The Retention and Care of Seeds by
Small-scale Farmers. NRI Report No. R2103.
Wright, M., Delimini, L., Luhanga, J., Mushi, C. and Tsini, H. (1995) The Quality of Farmer Saved
Seed in Ghana, Malawi and Tanzania, NRI Research Report.
Wydra K. and Singh, B.B. (1998) Project 11: CowpeaCereals Systems Improvement in the Dry
Savannas. Annual Report 1998. International Institute of Tropical Agriculture, Ibadan,
Nigeria.
Yadav, R.S., Hash, C.T., Bidinger, F.R. and Howarth, C.J. (1999a) QTL analysis and markerassisted breeding of traits associated with drought tolerance in pearl millet. In: Ito, O.,
OToole, J. and Hardy, B. (eds) Genetic Improvement of Rice for Water-limited Environments,
pp. 211223.
Yadav, R.S., Hash, C.T., Bidinger, F.R., Dhanoa, M.S. and Howarth, C.J. (1999) Identification and
Utilization of Quantitative Trait Loci (QTLs) to Improve Drought Tolerance in Pearl Millet
(Pennisetum glaucum (L.) R.Br.) CIMMYT, El Batan, Mexico.
Yadav, O.P., Weltzien-Rattunde, E., Bindinger, F.R. and Mahalakshmi, V. (2000) Heterosis in
landrace-based topcross hybrids of pearl millet across arid environments. Euphytica 112,
285295.
Yapi, A.M., Debrah, S.K., Dehala, G. and Njomaha, C. (1999) Impact of Germplasm Research
Spillovers: the Case of Sorghum Variety S 35 in Chad and Cameroon. Impact Series no. 3.
ICRISAT, Patancheru 502 324, Andra Pradesh, India.
Ye, Xudong (2000) Engineering the provitamin A (-carotene) biosynthetic pathway into
(Carotenoid-free) rice endosperm. Science 287, 303305.
Youm, O. and Kumar, K.A. (1995) Screening and breeding for resistance to millet head miner. P.
201109. In: Nwanze, N.F. and Youm, O. (eds) Panicle Insect Pests of Sorghum and Pearl Millet.
Proc. Int. Consult. Workshop. 47 Oct., 1993. Niamey, Niger. ICRISAT, Andra Pradesh, India,
pp. 20.
Young, N.D. (1999) A cautiously optimistic vision for marker-assisted breeding. Mini-review.
Molecular Breeding 5, 505510.
Zambezi, B.T. (1997) Characteristics of Maize Cultivars Released in Selected Countries of the Southern
African Development Community. International Maize and Wheat Improvement Center
(CIMMYT), Maize Programme, Harare, Zimbabwe, 41 pp.
198
Chapter
Appendix A: Production
(1000 t) of principal food crops
in sub-Saharan Africa, 1997
Population
Maize
(million)
West Africa
Benin
Burkina Faso
Cameroon
Central African
Rep.
Chad
Cte dIvoire
Gambia
Ghana
Guinea
Guinea-Bissau
Mali
Nigeria
Niger
Togo
Senegal
Sierra Leone
East Africa
Burundi
Eritrea
Ethiopia
Kenya
Rwanda
Somalia
Sudan
Tanzania
Uganda
Rice
Sorghum
Millet
Bean/
Cassava cowpea
240.4
5.7
11.1
13.9
9797.7
714.4
366.5
600.0
7237.9 10,864.7 10,146.0 11,527.1

481.5
26.8
120.2
28.0
0.5
89.5
942.9
603.9
375.0
55.0
400.0
71.0
3.4
82.0
99.1
576.0
8.5
1093.0
80.5
9.0
289.8
5354.0
3.0
452.2
60.3
9.4
7629.0
162.0
11.0
2137.0
2214.0
78.0
128.0
52.0
2107.0
740.0
17.0
112.3
1287.0
16.7
227.0
696.6
135.0
614.0
3268.0
67.0
41.0
173.7
411.3
714.6
38.0
55.3
4.3
2.0
2.0
533.0
80.0
6.7
14.3
1.2
18.3
7.6
1.1
11.5
118.4
9.8
4.3
8.7
4.4
194.6
6.4
3.4
60.1
28.4
5.9
10.2
27.9
31.5
20.8
38.0
426.6
19.4
12.9
316.9
5.0
19.0
540.3
7297.0
435.0
151.8
118.2
21.5
5944.5
85.0
59.0
1226.0
130.0
130.0
153.0
3369.0
498.5
294.0
12.0
144.7
248.4
65.4
66.1
153.8
8.0
29.0
738.9
5902.0
1713.0
58.3
426.5
21.7
1850.0
15.0
19.0
270.0
55.0
1.0
641.0
347.0
502.0
62.5
424.8
1.5
1781.8
173.4
2.5
184.7
7602.3
56.3
148.9
9.3
77.4
2431.8
130.0
225.0
62.5
13.0
2.5
1426.0
572.8
2487.7
73.7
10.0
91.0
23.9
139.3
1650.0
420.0
46.7
33.1
1357.0
270.0
3.0
400.0
120.0
13.0
11.0
271.0
269.0
167
167
Chapter
168
Appendix A: Production of Food Crops
Appendix A.
Continued.
Population
Maize
(million)
Southern Africa
Angola
Botswana
Congo, Dem.
Rep.
Congo, Rep.
Lesotho
Madagascar
Malawi
Mozambique
Namibia
Swaziland
Zambia
Zimbabwe
Total Africa
132.8
11.6
1.5
48.0
2.7
2.1
15.8
10.1
18.3
1.6
0.9
8.5
11.7
567.8
7299.7
369.5
11.6
Rice
3189.5
25.0
Sorghum
544.9
0.0
16.8
Millet
429.6
61.9
2.0
Bean/
Cassava cowpea
6542.5
581.6
2955.0
66.3
1000.0
347.0
50.0
38.0 4200.0
143.0
20.0
0.3
195.1
6.0
142.1
29.1
14.2
178.0 2558.0
1.0
2421.7
1226.5
65.7
39.5
4.1
253.0
1042.0
180.2
262.5
44.2 1334.2
49.4
9.5
107.5
108.2
0.4
0.7
5.8
960.2
12.5
30.8
61.0
187.5
2192.2
0.4
105.0
115.0
40.0
45.0
24,726.4 11,142.0 17,354.1 12,425.6 20,501.4 6,799.7
Source: FAO, FAOSTAT Database, 1999.
168
Chapter
Appendix B: Per capita

production (kg per person)
of principal food crops in
sub-Saharan Africa, 1997
Population
(million) Maize
West Africa
Benin
Burkina Faso
Cameroon
Central African
Rep.
Chad
Cte dIvoire
Gambia
Ghana
Guinea
Guinea-Bissau
Mali
Nigeria
Niger
Togo
Senegal
Sierra Leone
East Africa
Burundi
Eritrea
Ethiopia
Kenya
Rwanda
Somalia
Sudan
Tanzania
Uganda
Rice
Sorghum
Millet
Bean/
Cassava cowpea
240.4
5.7
11.1
13.9
40.9
125.3
33.0
43.2
30.8
4.7
8.1
4.0
45.4
21.1
84.9
28.8
42.2
4.9
54.4
5.1
48.2
84.5
0.0
27.0
11.3
12.9
0.9
6.5
3.4
6.7
14.3
1.2
18.3
7.6
1.1
11.5
118.4
9.8
4.3
8.7
4.4
194.6
6.4
3.4
60.1
28.4
5.9
10.2
27.9
31.5
20.8
24.1
14.8
40.3
7.1
59.7
10.6
8.2
25.2
45.2
0.3
105.2
6.9
2.1
39.3
25.3
3.2
35.6
78.0
13.2
12.5
1.9
66.9
35.6
5.0
16.8
90.0
13.9
12.4
91.7
122.7
53.4
27.6
6.8
9.5
20.0
93.5
3.7
5.9
1.9
0.7
0.2
0.1
16.9
3.8
11.2
63.7
1.4
10.8
17.3
0.7
17.3
47.0
61.6
44.4
35.3
13.6
4.9
30.5
13.3
17.4
20.4
4.6
22.0
15.0
120.8
15.8
14.1
3.5
37.1
4.6
55.1
8.4
1.1
26.4
64.2
49.8
174.8
13.6
49.0
4.9
9.5
2.3
5.6
4.5
1.9
0.2
0.0
23.0
11.0
24.1
42.6
9.3
29.7
1.3
97.4
22.8
2.3
16.1
64.2
5.7
34.6
1.1
17.6
12.5
20.3
7.9
10.6
1.3
0.1
45.3
27.5
3.6
12.1
13.9
42.9
10.9
3.8
7.0
42.2
0.9
6.7
20.3
1.3
0.4
8.6
12.9
169
169
Chapter
170
Appendix B: Per Capita Production of Food Crops
Appendix B.
Continued.
Population
(million) Maize
Southern Africa
Angola
Botswana
Congo, Dem.
Rep.
Congo, Rep.
Lesotho
Madagascar
Malawi
Mozambique
Namibia
Swaziland
Zambia
Zimbabwe
Total Africa
Rice
Sorghum
Millet
Bean/
Cassava cowpea
132.8
11.6
1.5
38.5
31.9
7.7
24.1
2.2
3.3
11.2
2.4
5.3
1.3
49.1
50.1
21.9
5.7
48.0
2.7
2.1
15.8
10.1
18.3
1.6
0.9
8.5
11.7
567.8
20.8
7.4
67.6
11.3
121.4
56.9
30.9
120.2
113.0
187.4
43.7
7.2
0.1
161.9
6.5
9.8
0.5
1.5
19.9
1.0
13.8
0.1
3.9
14.3
5.9
0.8
3.6
9.0
30.6
0.8
2.4
67.2
0.0
7.2
9.8
21.9
87.5
0.4
72.9
22.1
3.4
37.2
3.0
2.2
6.8
153.3
25.0
6.4
3.8
7.5
Source: FAO, FAOSTAT Database, 1999.
170
Chapter
Index
advanced research institutes (ARIs), 52

African agriculture
compared with other regions, 3, 4, 7, 9,
1011, 11, 29, 30
consumption patterns, 1718
and political support, 9
agricultural indicator comparisons, 30, 31
agro-ecologies, 1920, 9596
adaptation by cassava, 155, 156
analysis and crop improvements, 27, 28
analysis for maize improvement, 110
analysis for rice improvement, 138
and crop design, 42, 44, 57
crop diversity, 1222
regional production constraints, 5356
seed company adjustment to, 45
and sorghum adaptation, 119
Agrobacterium-mediated gene transfer, 65
and bananas, 163164
and cassava, 152
and cowpea, 144
and maize, 106
and rice, 136
and sorghum, 117118
AIDS, impact on research, 47
Angola
banana production trends, 158
cassava production, 149, 154
Catete maize, 39
crop yield increases, 40
NGOs seed supply, 81
apomixis, 7072
bananas, 13, 16, 17, 18, 39

breeding research centres, 161
genetic engineering and pest resistance,
65
history and utilization, 157158
hybrid varieties, 161162
improvement challenges, 164
improvement techniques, 161164
intractable traits, 11
pests and diseases, 160161, 162,
163164
production constraints, 55, 160161
production trends, 158160
research and development priorities,
165166
seed systems, 164165
tissue culture propagation, 63
transgenic research, 68
transgenic varieties, 61
Bean/Cowpea Collaborative Research Support
Programme, 142
beans, 19
pests and diseases, 11
Benin cassava production, 149, 154
bioinformatics, 59, 60
biosafety and biotechnology, 68, 70
biotechnology, 5974, 117
banana improvement, 163164
199
199
Index
200
Index
biotechnology continued
cassava improvement, 151152
cowpea breeding, 144145
and farmer involvement, 78
and intellectual property rights, 7274
maize improvement, 104106, 105106
millet breeding programmes, 128
and production constraints, 10
research challenges, 70
research constraints, 96
rice improvement, 135136
transformation and gene expression, 146
bird damage to crops, 112, 125, 126127,
129
Botswana, hybrid sorghum sales, 120
Burkina Faso
millet production levels, 124
rice breeding, 135
sorghum production, 112, 119
Burundi
NGO seed supply, 81
sorghum and banana varieties, 13
Cameroon
sorghum production, 112
cassava, 16, 17, 18, 19, 39
decentralized production, 155
65
genome, 152
nutritional improvement programme,
155
pests and diseases resistance, 155156
production trends, 148, 149
155156
cereal yield trends, 33
CGIAR see Consultative Group on
International Agricultural Research
(CGIAR)
Chad
millet pests and diseases, 126
child nutrition, 3031, 32, 140
CIAT see International Center for Tropical
Agriculture (CIAT)
CIMMYT see International Center for Maize
and Wheat Improvement
(CIMMYT)
CIP see International Potato Center (CIP)
civil unrest, 4
and agricultural disruption, 154
and seed supply, 78, 83
common bean, 17
and hunger periods, 32
community-based seed supply, 87, 90, 91,
108
Congo, banana production trends, 158
Congo (Dem. Rep.)
cassava production, 149
rice production, 132
Consultative Group on International
Agricultural Research (CGIAR), 39,
41, 4243
biotechnology expenditure, 67
using marker-assisted selection, 63
consumption patterns, 1718
and crop improvements, 21
Cte dIvoire
biotechnology research, 69
IARC crop improvement research, 50
cotton
Bt variety, 84
transgenic Bt variety, 61, 65
cowpea, 16, 18
drought-tolerant species, 17
early maturity, 143
65
and hunger periods, 32
landrace varieties, 39
low germination, 78
nutritional improvement goal, 146
200
Index
201
Index
pest-resistant varieties, 8384

production constraints, 5556,
141142
production levels, 140
145146
seed systems, 145
crop diversity, 5
crop improvement, 5, 2728
and farmer acceptance, 2021, 4041,
115116, 155
farmer involvement, 4144
history, 3738
national breeding programmes, 4448
on-farm trials, 78
reducing crop losses, 95
research model, 52
variety improvements during war, 83
and yield potential, 3536
crop research programmes, and local
preference information, 2122
crop varieties
for improved yield, 5, 7
for low-input farming, 52
performance as selection factor, 8
Crops Research Institute, 154
Crotolaria ochroleuca, 2526
DNA marker technology see marker-assisted

selection (MAS)
drought, 5
and maize production, 101, 107
resistance, 64
resistant cassava, 17, 147
resistant cowpea, 17, 143
resistant maize, 108109
resistant rice, 137, 138
and rice production, 133134
and sorghum production, 114, 118,
120
dwarfing genes, 38
East Africa
banana production trends, 158, 159
cassava production trends, 149
cowpea production levels, 140
maize production trends, 100, 101
sorghum production trends, 113
environmental variation and crop choice, 16
Ethiopia
fertilizer availability, 24
fallows improvement, 25
farmers
acceptance of new varieties, 2021, 40
access to seed, 76, 88, 89
associations and seed purchase, 8081
distributing seed to neighbours, 77, 78,
79, 92, 108
involvement in breeding programmes,
4144, 78, 89
low adoption of improved sorghum,
115116
millet varietal preferences, 127
poverty as factor in improvements, 36
seed saving, 7779
and transgenic crops, 60
underestimating diseases, 44
fertilizers
application, 10
availability and crop improvement,
2427
lack of subsidies, 21
Starter Pack programme, 26
usage, 11, 12
food aid, 33
food crops
per capita production tables, 169170
production tables, 167168
food importation, 31, 32, 33
food security, 4, 23, 3334, 95
and cassava, 148, 153
and population growth, 2930
and sorghum production, 119
Gabon banana production trends, 158

Gambia, The
improved cassava distribution, 154
gene banks, 48
genetic engineering, 12, 59, 6566, 117118
maize, 106
genetic improvements and seed market, 96
genetically modified organisms (GMOs),
opposition to, 67, 69
genomics, 5960
geographic information systems (GIS), 19
201
Index
202
Index
Ghana
cassava improved varieties adoption, 153,
154
cowpea seed systems, 145
open-pollinated varieties of maize, 105
green manures, 25, 26
Green Revolution, 7, 9, 35, 39, 40
groundnuts, seed availability, 78
Guinea
governmental improved rice distribution,
137
heat stress in millet, 128129

heat tolerant cowpea, 143
highlands, crop choice, 17, 19
hunger, 29, 30, 34
and crop improvement, 92
period, 13, 15, 32
hybrid crop varieties, 37, 7576, 85
of bananas, 161162
millet, 127128, 130
of sorghum, 120
IARC see International Agricultural Research

Centers (IARC)
ICRISAT see International Crops Research
Institute for the Semi-Arid Tropics
(ICRISAT)
IITA see International Institute for Tropical
Agriculture (IITA)
infrastructure affecting hunger, 34
INIBAP see National Agricultural Research
Organization of Uganda (INIBAP)
intellectual property rights, 24, 7274
and transgenic varieties, 84
International Agricultural Research Centers
(IARC), 19, 46, 4849, 50, 51, 57
and biotechnology, 70
and maize improvement programmes,
107
International Center for Maize and Wheat

Improvement (CIMMYT), 19, 48,
49, 50
and biotechnology research, 68
transgenic millet research, 106
International Center for Tropical Agriculture
(CIAT), 19, 48, 49, 50
cassava genetic research, 152
International Crops Research Institute for the
Semi-Arid Tropics (ICRISAT), 48,
49, 50
International Institute for Tropical
Agriculture (IITA), 39, 48, 49, 50
banana improvement, 161, 162
cassava improvement programme, 150,
151, 153, 154
cowpea breeding programme, 142143,
144145, 146
crop improvements, 83
International Potato Center (CIP), 48, 49, 50
iron toxicity in rice, 134
irrigation, 10, 11, 12, 36
KARI see Kenya, Agricultural Research

Institute (KARI)
Kenya
Agricultural Research Institute (KARI),
19, 68, 106
grain imports, 31, 32
hybrid maize adoption, 39
hybrid maize production, 104
maize breeding, 38
national breeding stategies, 46
NGO seed supply, 81
seed policy and regulation, 88
Sustainable Community-oriented
Development Programme (SCODP),
26
transgenic research, 68, 106
transgenic sweet potato importation, 69
202
Index
203
Index
labour supply, 13, 95, 147

land scarcity, 26
land use
and low technology impact, 3233
patterns, 16, 1819
landrace varieties, 39, 41, 72
of cassava, 150151
of millet, 127, 128
resistance genes, 4344
legume rotation, 25
Lesotho, improved cassava distribution, 154
Liberia
NGO seed supply, 81
low-input farming, 26, 27, 46, 95
crop design, 52
and maize production, 107
and seed availability, 7576
and sorghum production, 119
lowlands, crop choice, 17, 18
Madagascar
maize, 16, 17, 18
Agrobacterium-mediated transgenic, 106
anthesis-silking interval trait, 109
Bacillus thuringiensis and pest resistance,
65, 68
and fertilizer application, 24
flint-textured, 21, 38, 83
hybrid development by NARs, 45
Kenya agro-ecologies, 19
maturation time, 32
new varieties release rate, 56
nutrient use efficiency, 109
open-pollinated varieties, 105
pest- and disease-resistant, 109110
production constraints, 1920, 53,
101104, 107
108110
resistance genes, 8, 64, 68

seed prices, 86
seed systems, 108
surplus production in Malawi, 31
Malawi
early-maturing flint maize, 38
fertilizer prices, 25
flint-textured maize, 21
maize surpluses, 31, 32
Mali
lack of seed companies, 120
rice varieties, 13
seed storage, 78
sorghum production, 112, 119
sorghum varieties, 38
Striga-tolerant sorghum, 39
marginal land improvement strategies, 6
marker-assisted selection (MAS), 12, 59, 60,
6364, 68, 70, 105106
and banana improvement, 165
and cassava, 156
sorghum improvement, 117
material transfer agreements (MTA), 73
maturation time, 36, 38, 41
mechanization, 10, 11, 12
micropropagation, 67
of cassava, 151152
mid-altitude regions, crop choice, 17, 1819
millet, 16, 18
crop improvement, 127128
farmer varietal preferences, 127, 130
hybrid varieties, 127128, 130
129130
203
Index
204
Index
milling, 21
molecular genetics see biotechnology
Mozambique
decision-making environment, 13,
1415
early-maturing flint maize, 38
Fumo de Comboio cassava, 39
NGO seed supply, 81
seed storage, 77
Seguetana cowpea, 39
yield increases with new varieties, 39, 40
multinational seed companies, 83, 87
Namibia, improved cassava distribution, 154

NARS see national agricultural research
systems (NARSs)
National Agricultural Research Organization
of Uganda (INIBAP), 68
national agricultural research systems
(NARSs), 19, 38, 45, 4652, 49, 50,
52
and cassava breeding programmes, 153
interface with IARC, 51
relationship with seed companies, 80
seed dissemination, 5657, 91
staffing and finance, 4648
national breeding programmes, 4448, 96
for cowpea, 145
national seed companies, 87
Niger
sorghum seed purchasing, 120
Nigeria
cassava distribution, 154
cassava improved varieties adoption, 153
cassava production trends, 148, 149
cassava yield increases, 39
fertilizer availability and maize
production, 24
rice breeding, 135

nitrogen, 25, 26
non-governmental organizations (NGOs),
40
and banana variety dissemination, 165
cassava breeding, 154
maize seed distribution, 108
seed dissemination, 81, 84, 91, 96, 120,
137
supplying OPVs, 76
supplying plant material, 63
open-pollinated varieties (OPVs), 23, 37, 82,

83, 85
maize, 45, 104, 105
millet, 128
seed dealers lack of interest, 76
Participatory Variety Selection, 137

pearl millet see millet
of bananas, 160161, 162, 163164
of cassava, 149150, 151
of cowpea, 141142, 143144
of maize, 8, 101104, 107
of millet, 125126
and new crop varieties, 83
as production constraints, 5356
resistance and quantitative trait loci
(QTLs), 64
resistance and tissue culture, 63
resistant bananas, 165166
resistant cassava, 155156
resistant cowpea, 145, 146
resistant millet, 129
resistant rice, 138
resistant sorghum, 121
of rice, 133, 134, 135136, 137
of sorghum, 113
suffered by landrace varieties, 41
underestimated by farmers, 44
phenotypes, 66, 105, 106
phosphorus, 25, 127, 130
phytosanitary restrictions, 88
pigeon pea, 19
plant biotechnology, expenditure, 67
plant breeding, 5157
aims, 3640
204
Index
205
Index
biotechnology training for researchers,

67, 69
challenges, 57
decentralized cassava production, 155
farmer involvement, 4144, 89
history, 3739
maize improvement, 104106
multiplication of cassava, 154
NARSs and seed market, 5657
national, 2324, 4448
on-farm testing, 78, 91, 154
and phenotypes, 66
rice, 135137
sorghum improvement, 114115
plant restructuring, 36, 38
plant variety protection, 24
planting decisions, 13
political strategies and agriculture, 9, 2324
pollination, 2324
hybrid varieties 37
open-pollinated varieties (OPVs) see
open-pollinated varieties (OPVs)
polymerase chain reaction (PCR), 63
banana improvement, 163
polyploid crops, and apomixis, 71
population growth, 26, 2930, 41
and maize production, 101
post-harvest pests and diseases in maize, 102,
103104, 109119
potassium, 25
potatoes, 19
65
research institutes, 48, 49, 50
primary crop production per capita, 16
production constraints, 9, 12, 5356, 95, 96
bananas, 160161
cassava, 148150
maize, 20, 101104, 107
millet, 125127
rice, 133134
routine or intractable, 10
sorghum, 113114
pulses, 18
QTLs see quantitative trait loci (QTLs)

qualitative trait loci, 63, 66
quality importance, 21
quantitative trait loci (QTLs), 63, 64, 66
and cassava improvement, 152
and cowpea improvement, 144145
and maize improvement, 104, 105106
and rice improvement, 136
research programmes
biotechnology, 5974
financial constraints, 45, 4648
resources, 8
scientist training, 51, 67, 69, 70
resistance genes, 8, 37, 4344, 45, 52, 6061,
64, 68, 8384
cassava, 151
cowpea, 143, 146
maize, 105
millet, 125126
rice, 136
sorghum, 119
restriction fragment length polymorphism
(RFLP), 63, 105, 106
cowpea linkage map, 144145
millet linkage map, 128
sorghum linkage maps, 117
rice, 16, 17, 18
65
crop improvement, 135136
crossbreeding programme, 43
genome project, 60, 135
Oryza glaberrima, 13, 43, 131, 134, 135
Oryza sativa, 13, 43, 131, 134, 135
pro-vitamin A incorporation, 6566
research and development priorities, 138
research institutes, 48, 49, 50
seed systems, 137
sterility between crosses, 60
transgenic varieties, 38, 61, 66
Rockefeller Foundation
career fellowships, 69
rice biotechnology, 135136
205
Index
206
Index
Rwanda
NGO seed supply, 81
sorghum and banana varieties, 13
Sahel cassava production trends, 148

SCODP see Sustainable Community-oriented
Development Programme (SCODP)
seed certification, 88, 90
seed consumption estimates, 77
seed market, 9
constraints of small-scale production,
2223, 45, 52, 5657, 75, 87
deregulation, 56, 57, 76, 81, 90, 96
lack of African companies, 8687
multinational companies, 7980
national seed companies, 80
private sector seed companies, 9293,
105
public sector companies, 9293
trade volume, 77
variety development and marketing, 90,
91
seed policy and regulation, 8788, 9091
seed prices, 8586
seed systems
of bananas, 164165
of cassava, 153154
comunity-based production, 84
cowpea, 145
definition, 75
disasters and supplies, 78, 83
farmer-saved seed, 77, 78, 79, 92, 108
government funding, 78
government involvement, 90, 92
maize, 108
millet, 129
multiplication of supply, 91
open-pollinated varieties, 85
purchases by farmer associations,
8081
sorghum, 119120
storage, 7778
supply fraud, 81
sustainable supply, 8687, 91, 120, 122
transgenic crops, 84
self-pollinated varieties dissemination by

public sector, 91
semi-arid land and crop improvement, 17
Senegal
cowpea yield increases, 40
rice breeding, 135
sesame crops for oil, 13
Sierra Leone
NGO seed supply, 81
rice breeding, 135
soil fertility, 4, 2427
decline, 9
Somalia
NGO seed supply, 81
seed storage, 77
sorghum, 16, 18
biotechnology and improvement,
117118
day-length sensitivity, 113114
durra varieties, 38
flour quality preferences, 21
grain quality issues, 116117, 119
hybrid varieties, 120, 121
improved strains adoption by farmers,
115116
pest-resistant varieties, 84, 121
phosphorus acquisition efficiency,
122
production levels and trends, 112
121122
varieties in Bugusera region, 13
varieties in Sudan, 13
South Africa
hybrid sorghum sales, 120
206
Index
207
Index
Southern Africa, 50
cassava production trends, 149
cowpea production trends, 140
sorghum production trends, 112, 113
soybean, 39
sterility in plant breeding, 60, 127, 128, 162
storage pests in maize, 102, 103104
stress tolerance, 36, 37
Striga infestation
of cowpea, 142, 144
of maize, 102, 103, 107, 110
of millet, 126
and sorghum production, 114, 121
Striga resistance, 8, 39, 52, 64, 128, 129
in maize, 110
Sudan
maize yield increases, 40
Reep maize, 39
seed storage, 77
sorghum varieties, 13
Sustainable Community-oriented
Development Programme (SCODP),
26
sustainable farming and intensification, 38
Swaziland, improved cassava distribution, 154
sweet potatoes, 18
Tanzania
teff, 19
temperate vs. tropical zone disease, 11
tissue culture, 12, 59, 60, 6263, 67, 128
of bananas, 163, 165
of cassava, 154
national research centres, 6869
tobacco transgenic research, 68, 69
Togo
Indiana millet, 39
rice breeding, 135
training see research programmes, scientist

training
transgenic crop varieties, 38, 6061, 66
and biosafety, 70
maize, 106
tropical vs. temperate zone disease, 11
Uganda
banana improvement research, 162
NGO seed supply, 81
transgenic banana development, 68,
164
velvet bean, Mucuna pruriens, 2526
WARDA see West Africa Rice Development

Association (WARDA)
West Africa
cassava production trends, 148, 149
cowpea production trends, 140
maize production increases, 39
sorghum production trends, 113,
118119, 120
West Africa Rice Development Association
(WARDA), 38, 43, 48, 49, 50, 50
rice breeding programme, 135, 136137,
138
wheat, 17, 19
woman on a hill concept, 5, 7
yam, 39
yield decline of bananas, 161
yield increases with new varieties, 39, 40
yield potential, 35
yield stabilizing traits, 36
207
Index
208
Index
Zaire NGO seed supply, 81

Zambia
seed gifts, 79
Zimbabwe
biotechnology research, 68, 69
government support for seed supply,

8284
hybrid maize production, 104
maize imports, 32
208
Index

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Acknowledgements

International agricultural research centres (IARCs) have a major role to play in

Introduction and Summary

growing populations. The ensuing widespread frustration felt by local populations is

Introduction and Summary

Strategies for securing the harvest in marginal farming zones of Africa.

Introduction and Summary

the range and intensity of biophysical constraints to crop growth;

2.2 A Myriad of Production Constraints

Striga, stem borers, phosphorus uptake

Crop disease incidence in tropical compared with temperate zones.

Source: Dover and Talbot (1987) after Wellman (1968).

Source: FAO (2000).

2.3 Africas Diverse Cropping Landscape

Because small-scale farmers in Africa cultivate largely unimproved (i.e. unterraced,

Cereal crop consumption trends in Asia and Africa, 1997.

Consumption trends of selected non-cereal crops in Asia and Africa, 1997.

Fig. 2.4. Constraints to maize production in major agro-ecologies (Highlands, Mid-Altitude

Trying to make accurate determinations of varietal needs for numerous groups of

2.4 A Seed Sector Dominated by Market Failure

Finally, in the absence of investment by private seed companies, the rapid-fire,

2.5 Policies and Institutions are Critical to the Success of Crop

2.6 The Soil Fertility Problem

2.7 What, Then, is Needed?

Provide opportunities for interaction between groups of scientists, aimed at

The Roots of Hunger

Source: FAO (1998).

The Roots of Hunger

Key economic indicators, developing regions and the world, 1998.

Source: The World Bank (2000).

Source: FAO (2000).

The Roots of Hunger

Breeding Between an Art and

4.2 Aims and Contributions of Plant Breeding

Breeding Between an Art and a Science

Breeding Between an Art and a Science

Cassava yields increased dramatically in Nigeria following the introduction of

Yield increases observed from the introduction of improved varieties.

DeVries and Olufowote (1997) analysed results from intensive, NGO-managed

4.3 Crop Improvements Counter Arguments

Breeding Between an Art and a Science

4.4 Farmer Participation in Crop Improvement

Breeding Between an Art and a Science

Participatory rice variety selection in West Africa

When the West Africa Rice Development Association (WARDA) experienced a

4.5 Crop Improvements Ground Zero: National Breeding

Breeding Between an Art and a Science

require greater collaboration among scientists trained in various disciplines, including

Breeding Between an Art and a Science

4.6 Applied Science Powerhouses: International Agricultural Research

Breeding Between an Art and a Science

IARC crop improvement scientists in Africa.

Scientists in crop improvement

Scope of IARC crop improvement responsibilities in Africa.

Primary IARC breeding sites

Banana/plantain On, Nigeria/West Africa

Breeding Between an Art and a Science

Increasing the interface between IARC and NARS breeding programmes is

4.7 Breeding Linkages Within a Continuum of Crop Improvement

Collaborative model for overcoming genetic constraints in African crops.

4.8 Managing the Complexity of Adaptation

4.9 An Emerging Paradigm for Breeding in Africa