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Crop Improvement

Gale Encyclopedia of Food & Culture:

Crop Improvement

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Crop improvement refers to the genetic alteration of plants to satisfy human

needs. In prehistory, human forebears in various parts of the world brought
into cultivation a few hundred species from the hundreds of thousands
available. In the process they transformed elements of these species into
crops though genetic alterations that involved conscious and unconscious
selection, the differential reproduction of variants. Through a long history of
trial and error, a relatively few plant species have become the mainstay of
agriculture and thus the world's food supply. This process of domestication
involved the identification of certain useful wild species combined with a
process of selection that brought about changes in appearance, quality, and
productivity. The exact details of the process that altered the major crops is
not fully understood, but it is clear that the genetic changes were enormous
in many cases. In fact some crop plants have been so changed that for many
of them, maize, for example, their origins are obscure, with no extant close
wild relatives.

The selection process was unconscious in many cases. For example, in wild
wheats, the grains scatter by disarticulation, separation of the seed from the
seed head. When these grains were harvested by cutting the heads with a
sickle, an unconscious selection occurred for "nonshattering" types that
would then be continually replanted. For some crops a clear conscious
selection occurred, especially when the variant was obvious and would be
maintained by vegetative propagation. Something so clearly useful as a
seedless banana must have been immediately seized upon and maintained
("fixed") by planting offshoots of the plant. The changes wrought in
domestication included alteration in organ size and shape; loss of many
survival characters, such as bitter or toxic substances; disarticulation of
seeds in grains; protective structures, such as spines and thorns; seed
dormancy; and change in life span—increased in crops grown for roots or
tubers and decreased in crops grown for seed or fruit. Selection by bulking
desirable types (mass selection) is a powerful technique for making rapid
changes easily while maintaining genetic diversity in the population.

The selection of naturally occurring variants is the basis of crop improvement.

Over thousands of years this technique resulted in the development of
modern basic crops. The discovery of techniques for asexual (vegetative)
propagation, such as by using natural offshoots, rooting stem cuttings, or
various grafting techniques, made it possible to "fix" genetic variants. This
was the technique used for many tree fruits, enabling identical plants to be
cultivated in orchards. Naturally produced seedlings derived from
intercrosses of these selected plants were then available for selection again.
Many present-day fruit crops are similar to those cultivated in antiquity, and
some ancient selections are still cultivated—dates, for example. As humans
carried improved crops to new locations, opportunities opened to increase
genetic variation from natural intercrosses with new wild populations.

The changes that occur can be dramatic over time, as seen in the
proliferation of breeds of animals and especially the wide range of changes
brought about by fanciers of dogs, chickens, and pigeons. The observation of
these changes influenced the thinking of Charles Darwin to suggest that
natural selection, the survival of the fittest, could lead to enormous genetic
changes if carried out over a long enough time, and could lead to the origin
of new species.

In the eighteenth and nineteenth centuries a conscious attempt was made to

predict the performance of plants that could be expected from one seed
generation to the next. The concept that ancestry was important in crop
improvement led to refinement in the selection process, brought about by
keeping records and the assessment of lineage. Furthermore it became
obvious that variation could be managed by controlling the mating process,
an extension of what had long been known in animal breeding. This new type
of selection, termed pedigree selection, was found to increase the efficiency
of the process. Progeny testing (evaluating the genetic worth by progeny
performance) increased efficiency of this process. The origins of commercial
plant breeding began in the second half of the nineteenth century among
seed producers. It involved controlled crosses (hybridization) between
selections to control genetic recombination, followed by selection of
improved types. This is still the basis of traditional plant breeding.
Interestingly much of this early type of plant breeding was carried out
without a clear understanding of the genetic mechanism involved in
inheritance.

Until the famous experiments with the garden pea by Gregor Mendel (1822–
1884), a Catholic priest in Brünn, a Moravian town then in the Austro-
Hungarian Empire, the basic theory of inheritance involved the concept of
blending. Mendel unraveled the basic concept of inheritance and clearly
showed that characters in the pea were due to elements, later called genes,
that remained unaltered as they were inherited. Many characters in peas,
such as tallness and dwarfness, were shown to be controlled by a pair of
genes, of which one member was not always expressed (the concept of
dominance and recessiveness). Mendel demonstrated that the gametes of
the pea contained one member of the gene pairs that controlled characters
and that recombined randomly at fertilization. Mendel's paper was published
in 1866, but it had no impact until the paper was "rediscovered" in 1900,
when it created a sensation. It was soon obvious that the differences in
appearance among plants (phenotypes) could be explained by the interaction
of various genes (genotypes) as well as interaction with the environment.

Twentieth-Century Developments

In the twentieth century plant breeding developed a scientific basis, and crop
improvement was understood to be brought about by achieving favorable
accumulations and combinations of genes. Taking advantage of known
genetic diversity could facilitate this, and appropriate combinations were
achieved through recombinations brought about by the sexual process
(hybridization). Furthermore it was possible to move useful genes by special
breeding strategies. Thus a gene discovered in a wild plant could be
transferred to a suitable adapted type by a technique known as the backcross
method. A sexual hybrid was made, followed by a series of backcrosses to
the desirable (recurrent) parent, while selecting for the new gene in each
generation. After about five or six back-crosses, the offspring resembled the
recurrent parent but contained the selected gene.

In the early twentieth century it was demonstrated that the extra vigor long
associated with wide crosses (called hybrid vigor or heterosis), particularly in
naturally cross-pollinated crops, could be exploited in plant breeding. For
maize, a new system of hybrid breeding was developed, using a combination
of inbreeding and outbreeding. Inbreeding was accomplished by crossing the
plant with itself. This led to a decline in vigor as the step was repeated over
several generations. Outbreeding was achieved by intercrossing the inbred
lines to restore vigor. The hybrid between inbreds derived from divergent
inbreds (called a single cross or F1 hybrid) was uniform (homogeneous), and
some were superior to the original populations before inbreeding. During the
process of inbreeding, the inbreds became weak, but vigor was restored in
the F1. To increase seed set from weak inbreds, two hybrids were crossed;
this was known as the double cross method.

Hybrid breeding technique in a sense is similar to arranging a Rubic's cube,

where contradictory steps need to be taken to achieve the appropriate
reformulation. In hybrid breeding, the first step produces a series of weak
inbreds, followed by a series of specific combination, to produce a series of
new hybrids. Hybrid maize breeding led to enormous increases in
productivity, which were soon exploited in a wide variety of seed-propagated
crops, including naturally self-pollinated ones, such as tomato and rice.

A number of genetic techniques were developed and refined in twentieth-

century breeding, such as improved techniques to search for and store
increased genetic variability, different techniques to develop variable
populations for selection, and improved methods of testing to separate
genetic from environmental effects. The exact details of the process for crops
necessarily differed among naturally cross-pollinated plants (such as maize)
and naturally self-pollinated plants (such as soybean or tomato) as well as
those plants in which vegetative propagation (usually cross-pollinated)
permitted the fixing of improved types directly.
Conventional plant breeding can be defined as systems for selection of
superior genotypes from genetically variable populations derived from sexual
recombination. The system is powerful because it is evolutionary; progress
can be cumulative, with improved individuals continually serving as parents
for subsequent cycles of breeding. Genetic improvement by conventional
breeding has made substantial changes when the efforts have been long-
term. Characters improved include productivity, quality, and resistance to
diseases, insects, and stress. There are, however, limits to the progress of
conventional breeding. These are due to limitations of the sexual system,
because it is usually not possible to incorporate genes from nonrelated
species or to incorporate small changes without disturbing the particular
combination of genes that make a particular type unique. Thus a useful gene
in cabbage cannot be transferred to wheat. Limitations of conventional
breeding are particularly apparent when a needed character (such as disease
or insect resistance) is unavailable in populations that can be incorporated by
sexual crosses. Mutations may be induced, but they are often deleterious or
connected with undesirable effects.

With conventional breeding, it is also not possible to improve a unique

genotype, such as "Bartlett" pear, by adding a single character, since the
recombination that results from hybridization makes it impossible to recon-
figure this cultivar exactly. Finally, conventional breeding has technical or
economic limitations to detect infrequent or rare recombinants, the lack of
sufficient time to generate cycles of recombination, space to grow necessary
populations to recover superior recombinants, or resources to be able to
select, identify, evaluate, and fix desired recombinants.

Developments Using Dna

It has been suggested that many of the limitations of conventional breeding

can be overcome with advances in molecular biology that rely on DNA, the
genetic material.

Recombinant DNA technology, called transgene technology or genetic

engineering, is the most powerful and revolutionary of the new genetics
developed in the last half of the twentieth century. It is possible to isolate
stretches of DNA from one organism, store it in a bacterial host, select unique
combinations, and then incorporate them into the DNA of another species,
where it can be expressed. This technique, which relies on cell and tissue
culture, is truly a marvelous process. Refinements in the technique make it
possible to concentrate mutations in desired genes, further increasing
variability. Other uses of molecular biology known as genomics involve the
detailed mapping of the DNA and the identification of useful stretches of the
molecule. This makes it possible to improve the efficiency of selection,
because it is based directly on the genes rather than the organism, where the
effects may be confounded by environment and genetic interactions.

The limitations of the new breeding methods include technical problems,

such as the difficulty of transformation, problems of gene expression, or the
lack of knowledge concerning suitable genes to transfer. There are also
nontechnical issues, such as legal problems, since the techniques and the
genes are usually patented. However, in the short run the greatest
impediment has been problems of consumer acceptability and fear of the
unknown. The term "Frankenfood" has been coined to refer to food altered by
the process of using exotic genes incorporated by transgene technology. No
convincing evidence shows that genetic engineering has produced harmful
products, and an abundance of evidence shows that many foods derived from
traditional systems have inherent problems (consider the allergic reactions of
many people to peanuts). Nevertheless, molecular techniques have incited
fear of this new technology in many people. Moreover, the surplus of food in
the West has reduced the imperative to make the case for the need for new
technology to consumers.

The biotechnology industry has sold the technology to farmers (who have
accepted it) and ignored consumers. They have not been sophisticated in
exploiting the environmental virtues inherent in the new technology, such as
reducing pesticides, or in making the case that increased yield could free up
the agricultural use of fragile environments. Because of the benefits that
could accrue from this new technology, especially in the problem areas of the
world, it seems certain that future progress in plant breeding will involve both
conventional and unconventional techniques, but the immediate course of
events is fraught with uncertainty.

Bibliography

Bassett, Mark J., ed. Breeding Vegetable Crops. Westport, Conn.: AVI
Publishing, 1986.

Janick, Jules, and James N. Moore, eds. Fruit Breeding. 3 vols. New York: John
Wiley, 1996.

Poehlman, John Milton. Breeding Field Crops. 3d ed. Westport, Conn.: AVI
Publishing, 1986.

—Jules Janick

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