Plant Breeding and Crop Improvement | Research & Encyclopedia Articles

Plant Breeding and Crop Improvement

Plant breeding began when early humans saved seeds and planted them. The cultural change from living as nomadic hunter-gatherers, to living in more settled communities, depended on the ability to cultivate plants for food. Present knowledge indicates that this transition occurred in several different parts of the world, about 10,000 years ago.

Today, there are literally thousands of different cultivated varieties (cultivars) of individual species of crop plants. As examples, there are more than 4,000 different peas (Pisum sativum), and more than 5,000 grape cultivars, adapted to a wide variety of soils and climates.

The methods by which this diversity of crops was achieved were little changed for many centuries, basically requiring observation, selection, and cultivation. However, for the past three centuries most new varieties have been generated by deliberate cross-pollination, followed by observation and further selection. The science of genetics has provided a great deal of information to guide breeding possibilities and directions. Most recently, the potential for plant breeding has advanced significantly, with the advent of methods for the incorporation of genes from other organisms into plants via recombinant DNA-techniques. This capacity is broadly termed "genetic engineering." These new techniques and their implications have given rise to commercial and ethical controversies about "ownership," which have not yet been resolved.

The genetic discoveries of Gregor Mendel with pea plants, first published in 1866, were revolutionary, although Mendel's work remained obscure until translated from German into English by William Bateson in 1903. Nevertheless, the relationship between pollen lodging on the stigma and subsequent fruit production was realized long before Mendel's work. The first hybrid produced by deliberate pollen transfer is credited to Thomas Fairchild, an eighteenth-century, English gardener. He crossed sweet William with the carnation in 1719, to produce a new horticultural plant.

Towards the end of that century, Thomas Andrew Knight, another Englishman, demonstrated the practical value of cross-pollination on an unprecedented scale. He produced hybrid fruit trees by cross-pollination, and then grafted shoots of their seedlings onto established, compatible root stalks. This had the effect of greatly shortening the time until fruit production, so that the horticultural success of the hybridization could be evaluated. After selecting the best fruit, the hybrid seeds could be planted, and the process of grafting the seedlings and selection could be continued. The best hybrids, which were not necessarily stable through sexual reproduction, could be propagated by grafting. Thomas Knight was also responsible for the first breeding of wrinkled-seeded peas, the kind that provided Mendel with one of his seven key characters (round being dominant, with one allele sufficient for expression; wrinkled being recessive, requiring two copies of the allele for expression).

The concept of "diluting" hybrids by crossing them back to either parent also developed in the latter part of the nineteenth century. This strategy was introduced to ameliorate undesirable characters that were expressed too strongly. In genetic terms, there are two kinds of back-crossing. When one parent of a hybrid has many recessive characters, these are masked in the F₁ (first filial) hybrid generation by dominant alleles from the other parent. However, a cross of the F₁ hybrid with the recessive parent will allow the complete range of genetic variation to be expressed in the F₂ progeny. This is termed a test cross. A cross of the F₁ to the parent with more dominant characters is termed a back cross.

The broad aims of current plant breeding programs have changed little from those of the past. Improvements in yield, quality, plant hardiness, and pest resistance are actively being sought. In addition, the ability of plants to survive increasing intensities of ultraviolet radiation, due to damage in the ozone layer, and to respond favorably to elevated atmospheric concentrations of carbon dioxide are being assessed. To widen the available gene pools, collections of cultivars and wild relatives of major crop species have been organized at an international level. The United Nations' Food and Agriculture Organization (FAO) supported the formation of the International Board for Plant Genetic Resources in 1974. However, many cultivars popular in the nineteenth century have already fallen into disuse and been lost. The need to conserve remaining "heritage" varieties has been taken up by associations of enthusiasts in many countries, such as the Seed Savers' Exchange in the United States

Genetically identical plants, or clones, have been propagated from vegetative cuttings for thousands of years. Modern cloning techniques are used extensively to select for cultivars with particular characteristics, since there are limits to what can be achieved through direct hybridization. Some individual species or groups of cultivars cannot be genetically crossed. Sometimes this is because of natural polyploidy, when plant cells carry extra copies of some or all of the chromosomes, or because of inversions of DNA within chromosomes. In cases where cross-fertilization has occurred, "embryo rescue" may be used to remove hybrid embryos from the ovules and culture them on artificial media.

Pollen mother-cells in the anthers of some species have been treated with colchicine, to generate nuclei with double the haploid chromosome number, thus producing diploid plants that are genetically-identical to the haploid pollen. The use of colchicine to induce polyploidy in dividing vegetative cells first became popular in the 1940s, but tetraploids generated from diploids tend to mask recessive alleles. Generating diploids from haploids doubles all of the existing recessive alleles, and thereby guarantees the expression of the recessive characters of the pollen source.

In other difficult cases, the barriers to sexual crossing can sometimes be overcome by preparing protoplasts from vegetative (somatic) tissues from two sources. This involves treatment with cell-wall degrading enzymes, after which the protoplasts are encouraged to fuse by incubation in an optimal concentration of polyethylene glycol. A successful fusion of protoplasts from the two donors produces a new protoplast that is a somatic hybrid. Using tissue cultures, such cells can, in some cases, be induced to develop into new plants.

Somatic fusion is of particular interest for characters related to the chloroplast or mitochondrion. These plastids contain some genetic information in their specific, non-nuclear DNA, which is responsible for the synthesis of a number of essential proteins. In about two-thirds of the higher plants, plastids with their DNA are inherited in a "maternal" fashion--the cytoplasm of the male gamete is discarded after fusion of the egg and sperm cells. In contrast, in the minority of plants with biparental inheritance of plastid DNA, or when fusion of somatic protoplasts occurs, there is a mixing of the plastids from both parents. In this way, there is a potential for new plastid-nucleus combinations.

For chloroplasts, one application of plastid fusion is in the breeding of resistance to the effects of triazine herbicides. For mitochondria, an application relevant to plant breeding is in the imposition of male sterility. This is a convenient character when certain plants are to be employed as female parents for a hybrid cross. The transfer of male-sterile cytoplasm in a single step can avoid the need for several years of backcrosses to attain the same condition. Somatic hybridization has been used successfully to transfer male sterility in rice, carrot, rapeseed (canola), sugar beet, and citrus. However, this character can be a disadvantage in maize, where male sterility simultaneously confers sensitivity to the blight fungus, Helminthosporium maydis. This sensitivity can lead to serious losses of maize crops.

Replicate plant cells or protoplasts that are placed under identical conditions of tissue culture do not always grow and differentiate to produce identical progeny (clones). Frequently, the genetic material becomes destabilized and reorganized, so that previously concealed characters are expressed. In this way, the tissue-culture process has been used to develop varieties of sugar cane, maize, rapeseed, alfalfa, and tomato that are resistant to the toxins produced by a range of parasitic fungi. This process can be used repeatedly to generate plants with multiple disease resistance, combined with other desirable characters.

The identification of numerous mutations affecting plant morphology has allowed the construction of genetic linkage maps for all major cultivated species. These maps are constantly being refined. They serve as a guide to the physical location of individual genes on chromosomes.

DNA sequencing of plant genomes has shown that gene expression is controlled by distinct "promoter" regions of DNA. It is now possible to position genes under the control of a desired promoter, to ensure that the genes are expressed in the appropriate tissues. For example, the gene for a bacterial toxin (Bt) (from Bacillus thuringiensis) that kills insect larvae might be placed next to a leaf-development promoter sequence, so that the toxin will be synthesized in any developing leaf. Although the toxin might account for only a small proportion of the total protein produced in a leaf, it is capable of killing larvae that eat the genetically-modified leaves.

Agrobacterium tumefaciens and A. rhizogenes are soil bacteria that infect plant roots, causing crown gall or "hairy roots" diseases. Advantage has been taken of the natural ability of Agrobacterium to transfer plasmid DNA into the nuclei of susceptible plant cells. Agrobacterium cells with a genetically-modified plasmid, containing a gene for the desired trait and a marker gene, usually conferring antibiotic resistance, are incubated with protoplasts or small pieces of plant tissue. Plant cells that have been transformed by the plasmid can be selected on media containing the antibiotic, and then cultured to generate new, transgenic plants.

Many plant species have been transformed by this procedure, which is most useful for dicotyledonous plants. The gene encoding Bt, as well as genes conferring resistance to viral diseases, have been introduced into plants by this method.

Two methods have been developed for direct gene transfer into plant cells—electroporation and biolistics. Electroporation involves the use of high-voltage electric pulses to induce pore formation in the membranes of plant protoplasts. Pieces of DNA may enter through these temporary pores, and sometimes protoplasts will be transformed as the new DNA is stably incorporated (i.e., able to be transmitted in mitotic cell divisions). New plants are then derived from cultured protoplasts. This method has proven valuable for maize, rice, and sugar cane, species that are outside the host range for vector transfer by Agrobacterium.

Biolistics refers to the bombardment of plant tissues with microprojectiles of tungsten coated with the DNA intended for transfer. Surprisingly, this works. The size of the particles and the entry velocity must be optimized for each tissue, but avoiding the need to isolate protoplasts increases the potential for regenerating transformed plants. Species that cannot yet be regenerated from protoplasts are clear candidates for transformation by this method.

In 1992, a tomato with delayed ripening became the first genetically-modified (GM) commercial food crop. More than 40 different GM crops are now being grown commercially. GM corn and cotton contain bacterial genes that kill insects and confer herbicide-resistance on the crops. GM squash contains viral genes that confer resistance to viruses. Potatoes carry the Bt gene to kill the Colorado potato beetle and a viral gene that protects the potato from a virus spread by aphids. Mauve-colored carnations carry a petunia gene required for making blue pigment. In many cases, GM crops result in increased yields and reduced use of pesticides. New research is focused on producing GM foods containing increased vitamins and human or animal vaccines.

GM crops are controversial. There is concern that the widespread dissemination of the Bt gene will cause insects to become resistant. It has been reported that pollen from Bt corn is toxic to the caterpillars of monarch butterflies. It also is possible that GM crops will interbreed with wild plants, resulting in "superweeds" resistant to herbicides. There is also concern that the antibiotic-resistance genes, used as markers for gene transfer, may be passed from the plants to soil microorganisms or bacteria in humans who eat the food. Finally, the possibility of allergic reactions to the new compounds in food exists. Many countries have banned the production and importation of GM crops.

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