'''Plant breeding''' has been practiced for thousands of years.
Domestication, classical
plant breeding and
genetic engineering are all processes that alter the
genome of a plant to enhance its qualities as a
crop.
Plant breeding is practiced worldwide by
government institutions and commercial enterprises. International development agencies believe that breeding new crops is important for ensuring
food security and developing practices of
sustainable agriculture through the development of crops suitable for their environment .
Domestication
This map shows the sites of domestication for a number of crops. Places where crops were initially domesticated are called centres of origin
Domestication of plants is a
selection process conducted by
humans to produce plants that meet the needs of the farmer and the consumer. The practice is estimated to date back 9,000-11,000 years. Many crops in present day cultivation are the result of domestication in ancient times, about 5,000 years ago in the
Old World and 3,000 years ago in the
New World. In the
Neolithic period, domestication took a minimum of 1,000 years and a maximum of 7,000 years. Today, all of our principal
food crops come from domesticated varieties.
A cultivated crop
species that has evolved from
wild populations due to selective pressures from traditional
farmers is called a
landrace. Landraces, which can be the result of natural forces or domestication, are plants (or
animals) that are ideally suited to a particular region or environment. An example are the landraces of
rice,
Oryza sativa subspecies indica, which was developed in
South Asia, and
Oryza sativa subspecies
japonica, which was developed in
China.
Classical plant breeding
Classical plant breeding uses deliberate
interbreeding ('''crossing''') of closely or distantly related species to produce new crops with desirable properties. Plants are crossed to introduce
traits/
genes from one species into a new genetic background. For example, a
mildew resistant
pea may be crossed with a high-yielding but susceptible pea, the goal of the cross being to introduce mildew resistance without losing the high-yield characteristics. Progeny from the cross would then be crossed with the high-yielding parent to ensure that the progeny were most like the high-yielding parent, ('''backcrossing'''), the progeny from that cross would be tested for yield and mildew resistance and high-yielding resistant plants would be further developed. Plants may also be crossed with themselves to produce
inbreed varieties for breeding.
Classical breeding relies on
homologous recombination of two genomes to generate
genetic diversity. It also makes use of a number of
molecular techniques to generate diversity and produce plants that would not exist in
nature.
The Yecoro [[wheat (right)
cultivar is sensitive to salinity, plants resulting from a hybrid cross with cultivar W4910 (left) show greater tolerance to high salinity]]
Traits that breeders' have tried to incorporate into crop plants in the last 100 years include:
# Increased
quality and
yield of the crop
# Increased
tolerance of environmental pressures (
salinity, extreme
temperature,
drought)
# Resistance to
viruses,
fungi and
bacteria
# Increased tolerance to
insect pests
# Increased tolerance of
herbicides
Before World War II
Intraspecific hybridization within a plant species was demonstrated by
Charles Darwin and
Gregor Mendel, and was further developed by geneticists and plant breeders. In the early
20th century, plant breeders realized that Mendel's findings on the non-random nature of
inheritance could be applied to seedling populations produced through deliberate
pollinations to predict the frequencies of different types.
In 1908,
George Harrison Shull described
heterosis, also known as hybrid vigor. Hetrosis describes the tendency of the progeny of a specific cross to outperform both parents. The detection of the usefulness of heterosis for plant breeding has lead to the development of inbred lines that reveal a heterotic yield advantage when they are crossed.
Maize was the first species where hetrosis was widely used to produce hybrids.
By the
1920s,
statistical methods were developed to analyze gene action and distinguish heritable variation from variation caused by environment. In 1933, another important breeding technique, cytoplasmic male sterility (CMS), developed in maize, was described by
Marcus Morton Rhoades. CMS is a maternally inherited trait that makes the plant produce
sterile pollen, enabling the production of hybrids and removing the need for
detasseling.
These early breeding techniques resulted in large yield increase in the
United States in the early 20th century. Similar yield increases were not produced elsewhere until after
World War II, the
Green Revolution increased crop production in the developing world in the 1960s.
After World War II
Following
World War II a number of techniques were developed that allowed plant breeders to hybridize distantly related species, and artificially induce genetic diversity.
When distantly related species are crossed, plant breeders make use of a number of
plant tissue culture techniques to produce progeny from other wise fruitless mating. Interspecific and intergeneric hybrids are produced from a cross of related species or genera that do not normally
sexually reproduce with each other. The
cereal triticale is a
wheat and
rye hybrid. The first generation created from the cross was sterile, so the
cell division inhibitor
colchicine was used to double the number of
chromosomes in the
cell. Cells with an uneven number of chromosomes are sterile.
Failure to produce a hybrid may be due to pre- or post-fertilization incompatibility. If fertilization is possible between two species or genera, the hybrid
embryo aborts before maturation. When the cross is incompatible after fertilization, the embryo resulting from an interspecific or intergeneric cross can be rescued and cultured to produce a whole plant. This technique has been used to produce
new rice for Africa, an interspecific cross of Asian rice
(Otyza sativa) and African rice
(Otyza glaberrima).
Hybrids may also be produced by a technique called
protoplast fusion. In this case protoplasts are fused, usually in an electric field. Viable recombinants can be regenerated in culture.
Chemical
mutagens like EMS and
DMSO,
radiation and
transposons are used to generate
mutants with desirable traits to be bred with other cultivars. Classical plant breeders also generate genetic diversity within a species by exploiting a process called
somaclonal variation, which occurs in plants produced from tissue culture, particularly plants derived from
callus. Induced
polyploidy, and the addition or removal of chromosomes using a technique called chromosome engineering may also be used.
When a desirable trait has been bred into a species, a number of crosses to the favoured parent are made to make the new plant as similar as the parent as possible. Returning to the example of the mildew resistant pea being crossed with a high-yielding but susceptible pea, to make the mildew resistant progeny of the cross most like the high-yielding parent, the progeny will be crossed back to that parent for several generations. This process removes most of the genetic contribution of the mildew resistant parent. Classical breeding is therefore a cyclical process.
It should be noted that with classical breeding techniques, the breeder does not know exactly what genes have been introduced to the new cultivars. Some
scientists therefore argue that plants produced by classical breeding methods should undergo the same safety testing regime as genetically modified plants. There have been instances where plants bred using classical techniques have been unsuitable for human consumption, for example the nerve toxin
solanine was accidentally re-introduced into varieties of
potato.
Genetic engineering
:''See main article on
Transgenic plants.''
Genetic engineering of plants is achieved by adding a specific gene or genes to a plant, or by knocking out a gene with RNAi, to produce a desirable
phenotype. The resulting plants are often referred to as
transgenic plants. Genetic engineering can produce a plant with the desired trait or traits faster than classical breeding because the majority of the plant's genome is not being altered.
To genetically engineer a plant, a genetic construct must be designed so that the gene to be added or knocked-out will be expressed by the plant. To do this, a
promoter to drive
transcription and a termination sequence to stop transcription of the new gene must also be introduced to the plant. A marker for the selection of transformed plants is also included. In the
laboratory,
antibiotic resistance is a commonly used marker: plants that have been successfully transformed will grow on media containing antibiotics; plants that have not been transformed will die. Markers for selection are removed by mating (backcrossing) with the parent plant prior to commercial release.
The construct can be inserted in the plant genome by
recombination using the bacteria
Agrobacterium tumefaciens or
A. rhizogenes, or by direct methods like the
gene gun or microinjection. Using plant
viruses to insert genetic constructs into plants is also a possibility, but the technique is limited by the host range of the virus. For example,
Cauliflower Mosaic Virus (CaMV) only infects
cauliflower.
The majority of commercially released transgenic plants, commonly referred to as
genetically modified organisms, are currently limited to plants that have introduced resistance to
insect pests and
herbicides. Insect resistance is achieved through incorporation of a gene from
Bacillus thuringiensis (Bt) that encodes a
protein that is toxic to some insects. For example, if cotton pest the
cotton bollworm feeds on Bt cotton it will ingest the toxin and die. Herbicide resistance, particularly to the herbicide
Roundup, is achieved through
tissue culture. Plants are cultured on media containing the herbicide, and eventually some natural genetic mutation will arise that enables the plant to survive in the presence of the herbicide. The gene is then located (mapped) by crossing with susceptible species, and once identified can be introduced into other species.
Genetic engineering of plants that can produce pharmaceuticals (and industrial chemicals), sometimes called
pharmacrops, is a rather radical new area of plant breeding.
Issues and concerns
Modern plant breeding, whether classical or through genetic engineering, comes with issues of concern, particularly with regard to food crops. The question of whether breeding can have a negative effect on
nutritional value is central in this respect. Although relatively little direct research in this area has been done, there are scientific indications that, by favoring certain aspects of a plant's development, other aspects may be retarded. A study published in the
Journal of the American College of Nutrition in 2004, entitled
Changes in USDA Food Composition Data for 43 Garden Crops, 1950 to 1999, compared nutritional analysis of
vegetables done in 1950 and in 1999, and found substantial decreases in six of 13
nutrients measured, including 6% of
protein and 38% of
riboflavin. Reductions in
calcium,
phosphorus,
iron and
ascorbic acid were also found. The study, conducted at the Biochemical Institute, University of Texas, concluded in summary:
"We suggest that any real declines are generally most easily explained by changes in cultivated varieties between 1950 and 1999, in which there may be trade-offs between yield and nutrient content."
The debate surrounding genetic modification of plants is huge, encompassing the
ecological impact of genetically modified plants and the safety of
genetically modified food.
Plant breeders' rights is also a major and controversial issue. Today, production of new varieties is dominated by commercial plant breeders, who seek to protect their work and collect royalties through national and international agreements based in intellectual property rights. The range of related issues is complex. In the simplest terms, critics of the increasingly restrictive regulations argue that, through a combination of technical and economic pressures, commercial breeders are reducing
biodiversity and significantly constraining individuals (such as farmers) from developing and trading seed on a regional level. Efforts to strengthen breeders' rights, for example, by lengthening periods of variety protection, are ongoing.
See also
Notes
References
- Borojevic, S. 1990. Principles and Methods of Plant Breeding. Elserier, Amsterdam. Order: ISBN 0444988327
- McCouch, S. 2004. Diversifying Selection in Plant Breeding. PLoS Biol 2(10): e347.
- Briggs, F.N. and Knowles, P.F. 1967. Introduction to Plant Breeding. Reinhold Publishing Corporation, New York.
- Gepts, P. (2002). A Comparison between Crop Domestication, Classical Plant Breeding, and Genetic Engineering. Crop Science 42:1780–1790
- Origins of Agriculture and Crop Domestication - The Harlan Symposium
- news@nature.com. 1999 Are non-GM crops safe?
- Sun, C. et al. 1998. From indica and japonica splitting in common wild rice DNA to the origin and evolution of asian cultivated rice. Agricultural Archaeology 1998:21-29
External links
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