One major technique of plant breeding is selection, the process of selectively propagating plants with desirable characteristics and eliminating or "culling" those with less desirable characteristics. [3]

Another technique is the deliberate interbreeding (crossing) of closely or distantly related individuals to produce new crop varieties or lines with desirable properties. Plants are crossbred to introduce traits/genes from one variety or line 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 then be tested for yield (selection, as described above) and mildew resistance and high-yielding resistant plants would be further developed. Plants may also be crossed with themselves to produce inbred varieties for breeding.

Classical breeding relies largely on homologous recombination between chromosomes to generate genetic diversity. The classical plant breeder may also make use of a number of in vitro techniques such as protoplast fusion, embryo rescue or mutagenesis (see below) to generate diversity and produce hybrid plants that would not exist in nature.

Traits that breeders have tried to incorporate into crop plants include:

Improved quality, such as increased nutrition, improved flavor, or greater beauty
Increased 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
Longer storage period for the harvested crop
Before World War II[edit]

Garton's catalogue from 1902
Successful commercial plant breeding concerns were founded from the late 19th century.[clarification needed] Gartons Agricultural Plant Breeders in England was established in the 1890s by John Garton, who was one of the first to commercialize new varieties of agricultural crops created through cross-pollination.[4] The firm's first introduction was Abundance Oat, one of the first agricultural grain varieties bred from a controlled cross, introduced to commerce in 1892.[5][6]

In the early 20th century, plant breeders realized that Mendel's[who?] 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. Wheat hybrids were bred to increase the crop production of Italy during the so-called "Battle for Grain" (1925–1940). Heterosis was explained by George Harrison Shull. It 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 led to the development of inbred lines that reveal a heterotic yield advantage when they are crossed. Maize was the first species where heterosis was widely used to produce hybrids.

Statistical methods were also 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. This enables the production of hybrids without the need for labor-intensive 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[edit]

In vitro-culture of Vitis (grapevine), Geisenheim Grape Breeding Institute
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 otherwise 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. These crosses are referred to as Wide crosses. For example, the cereal triticale is a wheat and rye hybrid. The cells in the plants derived from the first generation created from the cross contained an uneven number of chromosomes and as result was sterile. The cell division inhibitor colchicine was used to double the number of chromosomes in the cell and thus allow the production of a fertile line.

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 may abort before maturation. If this does occur the embryo resulting from an interspecific or intergeneric cross can sometimes be rescued and cultured to produce a whole plant. Such a method is referred to as Embryo Rescue. This technique has been used to produce new rice for Africa, an interspecific cross of Asian rice (Oryza sativa) and African rice (Oryza 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 DMS, radiation and transposons are used to generate mutants with desirable traits to be bred with other cultivars - a process known as Mutation Breeding. 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 favored parent are made to make the new plant as similar to the favored 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 (See backcrossing ). This process removes most of the genetic contribution of the mildew resistant parent. Classical breeding is therefore a cyclical process.

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 poison solanine was unintentionally increased to unacceptable levels in certain varieties of potato through plant breeding. New potato varieties are often screened for solanine levels before reaching the marketplace.

Modern plant breeding[edit]
Modern plant breeding may use techniques of molecular biology to select, or in the case of genetic modification, to insert, desirable traits into plants. Application of biotechnology or molecular biology is also known as molecular breeding (see: Molecular breeding).

Modern facilities in molecular biology have converted classical plant breeding to molecular plant breeding
Steps of plant breeding[edit]
The following are the major activities of plant breeding:

Collection of variation
Selection
Evaluation
Release
Multiplication
Distribution of the new variety
Marker assisted selection[edit]
See main article on Marker assisted selection.
Sometimes many different genes can influence a desirable trait in plant breeding. The use of tools such as molecular markers or DNA fingerprinting can map thousands of genes. This allows plant breeders to screen large populations of plants for those that possess the trait of interest. The screening is based on the presence or absence of a certain gene as determined by laboratory procedures, rather than on the visual identification of the expressed trait in the plant.

Reverse breeding and doubled haploids (DH)[edit]
See also main article on Doubled haploidy.
A method for efficiently producing homozygous plants from a heterozygous starting plant, which has all desirable traits. This starting plant is induced to produce doubled haploid from haploid cells, and later on creating homozygous/doubled haploid plants from those cells. While in natural offspring genetic recombination occurs and traits can be unlinked from each other, in doubled haploid cells and in the resulting DH plants recombination is no longer an issue. There, a recombination between two corresponding chromosomes does not lead to un-linkage of alleles or traits, since it just leads to recombination with its identical copy. Thus, traits on one chromosome stay linked. Selecting those offspring having the desired set of chromosomes and crossing them will result in a final F1 hybrid plant, having exactly the same set of chromosomes, genes and traits as the starting hybrid plant. The homozygous parental lines can reconstitute the original heterozygous plant by crossing, if desired even in a large quantity. An individual heterozygous plant can be converted into a heterozygous variety (F1 hybrid) without the necessity of vegetative propagation but as the result of the cross of two homozygous/doubled haploid lines derived from the originally selected plant. patent

Genetic modification[edit]
See main article on Transgenic plants.
Genetic modification of plants is achieved by adding a specific gene or genes to a plant, or by knocking down a gene with RNAi, to produce a desirable phenotype. The plants resulting from adding a gene are often referred to as transgenic plants. If for genetic modification genes of the species or of a crossable plant are used under control of their native promoter, then they are called cisgenic plants. Sometimes genetic modification can produce a plant with the desired trait or traits faster than classical breeding because the majority of the plant's genome is not altered.

To genetically modify a plant, a genetic construct must be designed so that the gene to be added or removed 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, and the gene or genes of interest must 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. In some instances markers for selection are removed by backcrossing with the parent plant prior to commercial release.

The construct can be inserted in the plant genome by genetic recombination using the bacteria Agrobacterium tumefaciens or A. rhizogenes, or by direct methods like the gene gun or microinjection. U