INTRODUCTION

1.1

Maize Production and consumption

Maize (Zea mays L.) is the worlds most abundantly grown cereal crop, with an annual
production of over 870 million metric tons (Abdel-Rhman 2015) with the United States,
China, and Brazil being the top three maize-producing countries in the world, producing
approximately 563 million metric tons/year (Ranum et al. 2004). The average yield of
maize in most African countries is about 1.6 tons ha -1, which is much lower compared to
the world average yield of 5 tons ha-1 (Jamil et al. 2012).
Agriculture is the engine for growth in Africa. Cereals like Sorghum, Millets, Wheat,
Maize and Rice are major staple foods of the most population. These cereals are grown
over an area of 98.6 million hectares producing 162 million tons/year. Maize alone,
occupies over 34 million hectares with an average production if over 70 million metric
tons/year (Macauley 2015).
In Sub-Saharan Africa, Maize (Zea mays L.) is a staple food for 50% of the population. It
covers nearly 17% of the estimated 200 million ha cultivated land in SSA, and is
produced in diverse production environments and consumed by people with varying food
preferences and socio-economic backgrounds (Macauley 2015).
In Kenya, maize is a staple food for over 90% of the population (Mbithi and Van
Huylenbroeck 2000). About 1.6 million hectares are under maize annually where
production takes place in both small and large scale farms, however the bulk of it is from
small scale farmers who usually cultivate less than two hectares using limited technology
(Wambugu et al. 2012). In Kenya, the average maize yield is about 1.8 t/ha but a yield
potential of over 6 t/ha is possible (Schroeder et al. 2013), which is far below the crops
genetic potential.
The increase in maize productivity however, has no kept pace with increase in demand
(Midega et al. 2016). This is largely due to a number of biotic and abiotic factors. The
most important abiotic stresses include, drought, soil infertility, heat, and waterlogging.
Drought is one of the factors most frequently limiting maize production, food security,
and economic growth in Sub-Saharan Africa (Wegary et al. 2014). About 40 % of
Africas maize-growing area faces occasional drought stress, resulting in yield losses of

1025 %. Around 25 % of the maize crop suffers frequent drought, with losses of up to
half the harvest (Fisher et al. 2015). Maize is very sensitive to water stress and like other
cereal crops, there are some critical growth stages where soil moisture deficiencies
greatly affect the productivity (akir 2004). Water stress in the vegetative growth period
delays the development of stem and leaf cells, causing short plants and small leaf area
(Lauer 2003). Drought during tasselling could result in 4050% yield reduction, and
drought after pollination decreases single seed weight and causes a significant yield
reduction (Lauer 2003). Supplementary irrigation could potentially improve maize
production in drought prone areas (Nyombayire et al. 2011). However, the majority of
smallholder farmers cannot access irrigation due various barriers (Nyombayire et al.
2011).
Soil infertility is also an important abiotic factor affecting maize production. Soils in the
tropics are generally inherently infertile and most, especially those under subsistence
agriculture have been constantly mined of nutrients over the years without replenishment
with fertilizer (Smaling et al. 1997). In the tropics, drought and low soil fertility (mainly
nitrogen deficiency), frequently occur together (Nyombayire et al. 2011).
Nitrogen is the most limiting nutrient in the lowland and mid-altitude agro- ecologies in
the tropics and worst still a considerable proportion of maize in these areas is grown by
small scale farmer under low nitrogen conditions (Nyombayire et al. 2011).This means
that the nitrogen requirement of the crop must be met by the addition of organic or
inorganic fertilizers (Nyombayire et al. 2011). However, the high cost of inorganic
fertilizers makes it unaffordable to the majority of the subsistence farmers in most
African countries (Bnziger et al. 2004). As a result of these stresses, the International
Maize and Wheat Improvement Center (CIMMYT) initiated programs to improve
tropical maize for stress tolerance under both low N and drought (Edmeades 2013).
Biotic stresses also contribute to yield losses. The most important biotic stresses include;
Pests, diseases and weeds. Major diseases in Sub-Saharan Africa include northern leaf
blight (Exserohilum turcicum), maize streak disease transmitted by hoppers (Cicadulina
spp), gray leaf spot (Cercospora zeae maydis and common leaf rust (Puccinia sorghi).
Major insect pests of economic importance in Sub-Saharan Africa include; stem borers

(Chilo partellus) and (Busseola fusca), grain weevils (Sitophilus zeamays), and the larger
grain borer (Prostephanus truncantus).
Weeds are one of the major constraints to production, reducing yields by up to one third
or more in some areas. According to FAO, worldwide, 13 % loss of agricultural
production is credited to weeds. In Africa, more than 50% of crop losses are due to weeds
(Sibuga 1997). Weed competition is most serious when the crop is young. Weeds need to
be cleared from a field prior to planting a crop and again during the growing season for
optimal yields to be achieved. The critical period of weed-crop competition is
approximately equal to the first one-third to one-half of the life cycle of the crop
(Gianessi and Williams 2011). Many weed species occur in maize but those causing most
problems are perennial grasses and parasitic weeds (Striga spp.). Weed control is labor
intensive when done with widely used and traditional hand tools. Improved weed
management will help to alleviate labor constraints and improve crop yields. Parasitic
weeds such as Striga spp. can cause up to 100% yield loss. Striga infestation is common
in areas with low soil fertility and affects an estimated 20- 40 million ha in Sub-Saharan
Africa (Lagoke et al., 1991)
1.2

Striga Weed

Striga, commonly known as witchweed, is the most economically important parasitic

weed seed plant in the world (Atera et al. 2013). The genus is classified in the family of
Orobanchaceae (Gethi et al. 2005) with 28 species of parasitic plants that occur naturally
in parts of Africa, Asia and Australia (Atera et al. 2013). The major agricultural Striga
species are Striga hermonthica and S.asiatica, infecting cereals (maize, sorghum, millet
and upland rice). Striga hermonthica (Del.) Benth is the most important parasitic weed of
cereal crops in Western Kenya (Ngesa et al. 2015). It exists as an obligate parasite in its
early life obtaining nourishment from the host plant. It first attaches itself to the roots of
its host then penetrates the phloem and xylem from where it drains off water and
nutrients, hinders photosynthesis and as a result, causes a phytotoxic effect in a matter of
days. This stage of Striga growth is known as below ground growth and results into
damage of the host crop exhibited by stunted growth and a reduction in crop yields
(Gacheru et al. 2002). Later, after emerging from the soil, the weed becomes a facultative
parasite, photosynthesizing approximately 20% of its growth requirements and obtaining

the rest from its host (Watson et al. 2007). The roots of these cereal crops produce
exudates derived from carotenoid biosynthetic pathway (Matusova et al. 2005). These
exudates are required for Striga germination. Striga is particularly a problem in areas with
low moisture and low soil fertility, decreased use of fallow and minimal use of organic or
inorganic fertilizer (De Groote et al. 2008). Striga flowers shed seeds within the life cycle
of its host. Seeds are tiny (< 0.3 mm) and one plant can produce 50,000200,000 seeds
(CIMMYT, 2014), thus, distributed by surface run-off, eroded soil and wind and use of
contaminated farm equipment. This prolific reproductive nature coupled with seed
longevity of the Striga species compounds the detrimental effect of Striga weed in
cereals.
1.3

Effects of Striga

Striga hermonthica problem has been in existence as early as 1936 in the fields of farmers
within Lake Victoria Basin, Western Kenya (Khan et al., 2006). Striga infestation is as a
result of mono-cropping of cereals which host the parasite and declining soil fertility
(Oswald 2005, CIMMYT 2014). In the Lake Victoria Basin of Kenya alone, 0.24 million
hectares of arable land, which is 15% of the total arable land in the region is infested.
This causes yield losses between 10-100% and or monetary losses of up to US$ 41
million annually (Jamil et al. 2011). The yield loss in maize due to Striga effects
represents 12.3% of the 2.4 million metric tons of maize that Kenya produces annually
(Atera et al. 2013).
Initial symptoms in maize occur while the parasite is still subterranean; they are evident
in water soaked leaf lesions, chlorosis, folded leaves and wilting even where there is
sufficient soil moisture, reduced inter nodes and eventual leaf and plant desiccation and
necrosis. Severe stunting and drought-like symptoms such as leaf margin curling also
indicate Striga infection (Berner et al. 1997). In many cases, the damage caused by the
weed is so severe that a farmer can only harvest 0.5tons ha-1 of maize instead of the
potential 5 tons, or zero yields under heavy infestation (Jamil et al. 2012). These losses
largely depend on Striga density, host species and genotype, land use system, soil
nutritional status and rainfall patterns ( (Atera et al. 2013). The most affected are the poor
subsistence farmers, who are not aware of the threat that Striga poses to their land quality

and food security as the weed continues to increase its soil seed bank and spreading to
new areas.
1.4

Striga control strategies

A number of control strategies including crop rotation, intercropping with legumes

(pigeon pea, groundnut, and soybean), late planting, deep planting, transplanting and
push-pull technologies and use of leguminous trap crops, which stimulate suicidal
germination and therefore reduce the seed bank have been put forward to help cub the
weed menace (Henry et al. 2015, IITA n.d, Ellis-Jones et al. 2004). Most of these
techniques require special implements to effect, others like use of fertilizer are not
practical with resource poor farmers. These methods are partially effective and as a result
have not been widely adopted. In view of this, seed-based technologies that include;
germplasm based-Striga resistance, use of herbicide coated seeds, and biological control
methods are some of the methods that scientists are using to help control the striga
menace.
1.4.1

Conventional Resistance

Development of host resistance/tolerant maize, i.e. Breeding maize varieties that are
resistant (prevents or limits Striga attachment or growth) or tolerant (variety still gives
acceptable yields despite Striga attack) is the most widely spread seed-based and highly
effective method. It is the most feasible and environmentally friendly method for smallholder farmers to control Striga. It has been used successfully in sorghum where great
advances have been made in understanding how the tolerance/resistance work through
biotechnology (http://www.iita.org/). The International Institute of Tropical Agriculture
(IITA) and CIMMYT have developed open-pollinated maize varieties, hybrids, and
inbred lines that are resistant to S. hermonthica. IITA has employed recurrent selection
and inbreeding to develop inbred lines that has less S. hemonthica damage symptoms and
yield loss and has less striga emergence thus reducing seed production (Menkir 2011).
IITA developed inbred lines (Reg. no. GP-428 to GP-453 and PI 639925 to PI 639950)
with resistance to Striga and are adapted to the lowlands (TZSTRI101 to TZSTRI117)
and midaltitudes (TZSTRI118 to TZSTRI126) (Menkir 2006). In addition to Striga
resistance, the lowland tropical inbred lines also have resistance to Southern corn leaf
blight, Southern corn rust and maize streak virus while the midaltitude inbred lines have

resistance to Northern leaf blight, corn rust and maize streak virus (Menkir, 2006). Some
of the inbred lines evaluated showed that a decrease in number of emerged Striga was
associated with a decrease in number of Striga attached to the maize roots (Menkir,
2006). Therefore, maize hybrids involving S. hermonthica resistant parental lines can
provide sufficient levels of resistance to Striga.
1.4.2

Herbicide resistant maize

This approach is based upon inherited resistance of maize to a systemic herbicide

(imazapyr), a mechanism widely referred to as imazapyr resistance (IR). When IR maize
seed is coated with the herbicide, germinated Striga seeds attempting to parasitize the
resulting maize plant are destroyed (Woomer and Musa 2005). IR maize or StrigAway
maize technology comprises two main elements a herbicide-resistant maize seed and
imazapyr, a systemic imidazolinone herbicide. The herbicide-resistant maize is coated
with low doses of the herbicide, about 30g imazapyr per hectare, to control Striga
(CIMMYT 2014). As the IR maize germinates, it absorbs some of the herbicide used in
coating it. The germinating maize stimulates Striga to germinate and as it attaches to the
maize root, it is killed before it can cause any damage. The herbicide that is not absorbed
by the maize plant diffuses into the soil and kills Striga seeds that have not
germinated and forms a localized protection zone around growing maize roots
(http://www.iita.org). The StrigAway maize technology, therefore, decreases the level of
Striga in the farm through direct attacks on the Striga plants and seeds. Results by
Kanampiu et al. (2001) indicated that herbicide-resistant maize coated with 30g imazapyr
per hectare gave almost total striga control up to 13 weeks after planting which translates
to a yield increase of up to 3.0 tons ha -1. A study by Makumbi et al. (2015) on Agronomic
performance and genotype x environment interaction of herbicide-resistant maize
varieties in Eastern Africa, showed that the best IR maize variety (STR-VE-216)
outyielded the Striga-tolerant and commercial genotypes by 113 and 89%, respectively,
under Striga-infested conditions. Similarly, IR OPVs supported significantly fewer
emerged Striga plants relative to the check varieties. Under Striga-free conditions, IR
OPVs showed GY advantage over commercial varieties. In Kenya four hybrids were
developed by CIMMYT and allocated to three Kenyan seed companies (Kenya, Western,

and Lagrotech) and the Kenya Agricultural Research Institute for commercialization, two
OPVs were allocated to Western and Freshco Seed companies (Ransom et al. 2012).
1.5

Combining ability

Combining ability is the ability of parents or cultivars to combine amongst each other
during hybridization so that favorable gene expressions are transmitted to their progenies.
Combining ability is of two types: general and specific combining ability. General
combining ability (GCA) is the average performance of a line in hybrid combinations,
while specific combining ability (SCA) is the performance of a hybrid compared to the
average performance of the parent-inbred lines of the hybrid (Sprague and Tatum 1942).
The performance of a hybrid is related to GCA and SCA of the inbred lines involved in
the cross where diallel analysis is used to estimate the GCA and SCA effects and their
implications in breeding. Diallel mating design is a method whereby all inbred lines or
open-pollinated varieties are crossed in all possible combinations for field evaluation
purposes.
At the gene action level, GCA is associated with additive effects of the genes, while SCA
is related to dominance and epistatic effects (non-additive effects) of the genes. GCA
effect is an indicator of the relative value of the lines and their parental populations in
terms of frequency of favorable genes and of its divergence, as compared to the other
parents in the diallel. Thus, the analysis of GCA allows identification of superior parents,
to be used in intra-population breeding programs. The SCA effect of two populations
expresses the differences of gene frequencies between them and their divergence, as
compared to the diallel parents. A study by Olakojo and Olaoye (2005) evaluated ten
inbred parents of varying tolerance to Striga. Lutea in a half diallel to generate 45F1
hybrids which were evaluated in three Striga endemic regions. The results showed the
GCA effects for Striga syndrome ratings were generally low with a negative values. This
implied that those parent inbreds were good sources of genes for higher grain yield. SCA
effects were also low with negative values for Striga syndrome rating, showing good
Striga tolerance. In another study by Karaya et al. (2014), fourteen female lines were
mated using North Carolina Design II with six males and evaluated under both artificial
Striga infestation and Striga free environments. The ratio of GCA: SCA means squares
exhibited a predominance of additive gene effect in the inheritance of Striga resistance

traits as opposed to dominance gene effects. Inbred lines with good GCA for yield and
Striga resistance traits were also identified. Similar studies were conducted by Olaoye
and Bello (2009) on ten Open Pollinated Varieties (OPVs) and their 45F 1 hybrids in
Striga prone and Striga free zones in Nigeria. Both GCA and SCA effects for Striga
related characters such as Striga shoot counts, syndrome ratings, flowering Striga shoots
and barren maize plants were generally low. This suggested additive and dominant gene
action in tolerance to Striga. Dao et al. (2014) did a study where twenty-four CIMMYT
and INERA white lines were crossed to two CIMMYT testers and twenty six IITA and
INERA yellow inbred lines were crossed to two IITA testers. Two trials composed of 48
white testcrosses and 52 yellow test crosses plus three checks were evaluated in wellwatered and drought stress conditions in the dry season over two years. Significant GCA
effects due to lines and testers for many traits including grain yield were observed. Of
the 24 CIMMYT and INERA white lines, 15 lines could be classified into heterotic
groups based on the SCA effects and testcross mean grain yield in well-watered
environment and, 10 lines were classified under drought stress condition. With this
regard, the information about combining ability of experimental breeding materials is
imperative to a breeding program aiming to develop high yielding hybrids and composite
varieties.

1.8

Heterotic Grouping

Melchinger and Gumber (1998) defined a heterotic group as a group of related or

unrelated genotypes from the same or different populations, which display similar
combining ability and heterotic response when crossed with genotypes from other
genetically distinct germplasm groups.
Heterotic groups are formed by isolation and selection (natural or human) for adaptedness
over time and are critical in maximizing the expression of heterosis in hybrids (Pawali
2000). The expression of heterosis depends on the level of dominance controlling the
trait. For example, grain yield in maize is expected to exhibit heterosis as a consequence
of partial to complete dominance of genes controlling the trait (Hallauer and Miranda
Filho 1988). Heterosis of a character indicates the extent to which a character can be
inherited from one generation to another. This is a valuable tool used to predict the

magnitude of genetic gains that follows selection. The combination of lines of different
heterotic groups results in hybrids with higher chances of genetic expression of the target
effects of hybridization (Ricci et al. 2007). In the study by Dao et al. (2014) eighty five
percent of the IITA and INERA yellow lines were classified into heterotic groups in both
drought and non-drought conditions. Legesse et al. (2009) carried out a study to examine
the combining ability and heterotic grouping of twenty six highland transition maize
inbred lines. The results revealed two best contrasting heterotic groups, Pool 9A-MHM
and 142-1-e, composed of 13 and 10 inbred lines, respectively. This heterotic group may
have a wide applicability in the East African highland maize breeding programs.
Fan et al. (2009) classified 23 maize inbred lines into heterotic groups using four testers.
He reported that using heterotic groups specific and general combining ability (HSGCA)
increased maize breeding efficiency by 16.7% to 23.6% compared with simple sequence
repeat (SSR) and specific combining ability combined with line pedigree and hybrid yield
information (SCA-PY) methods, respectively.

The objectives of these study was,

i.

To estimate the general combining ability (GCA) and the specific combining
ability (SCA) of Sixty four herbicide resistant inbred lines under striga infestation.

ii.

To classify the inbred lines into heterotic groups based on SCA for grain yield
with 2 inbred line testers.

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