Cultivar

From Wikipedia, the free encyclopedia

Osteospermum 'Pink Whirls'
A cultivar selected for its intriguing and colourful flowers

The term cultivar most commonly refers to an assemblage of plants selected for desirable characters that are maintained during propagation. More generally, cultivar refers to the most basic classification category of cultivated plants in the International Code of Nomenclature for Cultivated Plants (ICNCP). Most cultivars arose in cultivation, but a few are special selections from the wild.

Popular ornamental garden plants like roses, camellias, daffodils, rhododendrons, and azaleas are cultivars produced by careful breeding and selection for floral colour and form. Similarly, the world's agricultural food crops are almost exclusively cultivars that have been selected for characters such as improved yield, flavour, and resistance to disease, and very few wild plants are now used as food sources. Trees used in forestry are also special selections grown for their enhanced quality and yield of timber.

Cultivars form a major part of Liberty Hyde Bailey's broader group, the cultigen, which is defined as a plant whose origin or selection is primarily due to intentional human activity. A cultivar is not the same as a botanical variety, which is a taxonomic rank below subspecies, and there are differences in the rules for creating and using the names of botanical varieties and cultivars. In recent times, the naming of cultivars has been complicated by the use of statutory patents for plants and recognition of plant breeders' rights.

The International Union for the Protection of New Varieties of Plants (UPOV – French: Union internationale pour la protection des obtentions végétales) offers legal protection of plant cultivars to persons or organisations that introduce new cultivars to commerce. UPOV requires that a cultivar be "distinct, uniform", and "stable". To be "distinct", it must have characters that easily distinguish it from any other known cultivar. To be "uniform" and "stable", the cultivar must retain these characters in repeated propagation.

The naming of cultivars is an important aspect of cultivated plant taxonomy, and the correct naming of a cultivar is prescribed by the Rules and Recommendations of the International Code of Nomenclature for Cultivated Plants (ICNCP, commonly denominated the Cultivated Plant Code). A cultivar is given a cultivar name, which consists of the scientific Latin botanical name followed by a cultivar epithet. The cultivar epithet is usually in a vernacular language. For example, the full cultivar name of the King Edward potato is Solanum tuberosum 'King Edward'. 'King Edward' is the cultivar epithet, which, according to the Rules of the Cultivated Plant Code, is bounded by single quotation marks.

Etymology

Liberty Hyde Bailey (1858–1954) coined the words cultigen in 1918 and cultivar in 1923.

The word cultivar originated from the need to distinguish between wild plants and those with characteristics that arose in cultivation, presently denominated cultigens. This distinction dates to the Greek philosopher Theophrastus (370–285 BC), the "Father of Botany", who was keenly aware of this difference. Botanical historian Alan Morton noted that Theophrastus in his Historia Plantarum (Enquiry into Plants) "had an inkling of the limits of culturally induced (phenotypic) changes and of the importance of genetic constitution" (Historia Plantarum, Book 3, 2, 2 and Causa Plantarum, Book 1, 9, 3).

The International Code of Nomenclature for algae, fungi, and plants uses as its starting point for modern botanical nomenclature the Latin names in Linnaeus' (1707–1778) Species Plantarum (tenth edition) and Genera Plantarum (fifth edition). In Species Plantarum, Linnaeus enumerated all plants known to him, either directly or from his extensive reading. He recognised the rank of varietas (botanical "variety", a rank below that of species and subspecies) and he indicated these varieties with letters of the Greek alphabet, such as α, β, and λ, before the varietal name, rather than using the abbreviation "var." as is the present convention. Most of the varieties that Linnaeus enumerated were of "garden" origin rather than being wild plants.

In time the need to distinguish between wild plants and those with variations that had been cultivated increased. In the nineteenth century many "garden-derived" plants were given horticultural names, sometimes in Latin and sometimes in a vernacular language. From circa the 1900s, cultivated plants in Europe were recognised in the Scandinavian, Germanic, and Slavic literature as stamm or sorte, but these words could not be used internationally because, by international agreement, any new denominations had to be in Latin. In the twentieth century an improved international nomenclature was proposed for cultivated plants.

Liberty Hyde Bailey of Cornell University in New York, United States created the word cultivar in 1923 when he wrote that:

The cultigen is a species, or its equivalent, that has appeared under domestication – the plant is cultigenous. I now propose another name, cultivar, for a botanical variety, or for a race subordinate to species, that has originated under cultivation; it is not necessarily, however, referable to a recognized botanical species. It is essentially the equivalent of the botanical variety except in respect to its origin.

In that essay, Bailey used only the rank of species for the cultigen, but it was obvious to him that many domesticated plants were more like botanical varieties than species, and that realization appears to have motivated the suggestion of the new category of cultivar.

Bailey created the word cultivar, which is generally assumed to be a portmanteau of cultivated and variety. Bailey never explicitly stated the etymology of cultivar, and it has been suggested that it is instead a contraction of cultigen and variety, which seems correct. The neologism cultivar was promoted as "euphonious" and "free from ambiguity". The first Cultivated Plant Code of 1953 subsequently commended its use, and by 1960 it had achieved common international acceptance.

Cultigens

Bread wheat, Triticum aestivum, is considered a cultigen, and is a distinct species from other wheats according to the biological species concept. Many different cultivars have been created within this cultigen. Many other cultigens are not considered to be distinct species, and can be denominated otherwise.

The words cultigen and cultivar may be confused with each other. A cultigen is any plant that is deliberately selected for or altered in cultivation, as opposed to an indigen; the Cultivated Plant Code states that cultigens are "maintained as recognisable entities solely by continued propagation". Cultigens can have names at any of many taxonomic ranks, including those of grex, species, cultivar group, variety, form, and cultivar; and they may be plants that have been altered in cultivation, including by genetic modification, but have not been formally denominated. A cultigen or a component of a cultigen can be accepted as a cultivar if it is recognisable and has stable characters. Therefore, all cultivars are cultigens, because they originate in cultivation, but not all cultigens are cultivars, because some cultigens have not been formally distinguished and denominated as cultivars.

Formal definition

The Cultivated Plant Code notes that the word cultivar is used in two different senses: first, as a "classification category" the cultivar is defined in Article 2 of the International Code of Nomenclature for Cultivated Plants (2009, 8th edition) as follows: The basic category of cultivated plants whose nomenclature is governed by this Code is the cultivar. There are two other classification categories for cultigens, the grex and the group. The Code then defines a cultivar as a "taxonomic unit within the classification category of cultivar". This is the sense of cultivar that is most generally understood and which is used as a general definition.

A cultivar is an assemblage of plants that (a) has been selected for a particular character or combination of characters, (b) is distinct, uniform and stable in those characters, and (c) when propagated by appropriate means, retains those characters.

Different kinds

A cultivar of the orchid genus Oncidium

Which plants are chosen to be named as cultivars is simply a matter of convenience as the category was created to serve the practical needs of horticulture, agriculture, and forestry.

Members of a particular cultivar are not necessarily genetically identical. The Cultivated Plant Code emphasizes that different cultivated plants may be accepted as different cultivars, even if they have the same genome, while cultivated plants with different genomes may be regarded as the same cultivar. The production of cultivars generally entails considerable human involvement although in a few cases it may be as little as simply selecting variation from plants growing in the wild (whether by collecting growing tissue to propagate from or by gathering seed).

Cultivars generally occur as ornamentals and food crops: Malus 'Granny Smith' and Malus 'Red Delicious' are cultivars of apples propagated by cuttings or grafting, Lactuca 'Red Sails' and Lactuca 'Great Lakes' are lettuce cultivars propagated by seeds. Named cultivars of Hosta and Hemerocallis plants are cultivars produced by micropropagation or division.

Clones

Leucospermum 'Scarlet Ribbon'
A cross performed in Tasmania between L. glabrum and L. tottum

Cultivars that are produced asexually are genetically identical and known as clones; this includes plants propagated by division, layering, cuttings, grafts, and budding. The propagating material may be taken from a particular part of the plant, such as a lateral branch, or from a particular phase of the life cycle, such as a juvenile leaf, or from aberrant growth as occurs with witch's broom. Plants whose distinctive characters are derived from the presence of an intracellular organism may also form a cultivar provided the characters are reproduced reliably from generation to generation. Plants of the same chimera (which have mutant tissues close to normal tissue) or graft-chimeras (which have vegetative tissue from different kinds of plants and which originate by grafting) may also constitute a cultivar.

Seed-produced

Some cultivars "come true from seed", retaining their distinguishing characteristics when grown from seed. Such plants are termed a "variety", "selection" or "strain" but these are ambiguous and confusing words that are best avoided. In general, asexually propagated cultivars grown from seeds produce highly variable seedling plants, and should not be labelled with, or sold under, the parent cultivar's name.

Seed-raised cultivars may be produced by uncontrolled pollination when characteristics that are distinct, uniform and stable are passed from parents to progeny. Some are produced as "lines" that are produced by repeated self-fertilization or inbreeding or "multilines" that are made up of several closely related lines. Sometimes they are F1 hybrids which are the result of a deliberate repeatable single cross between two pure lines. A few F2 hybrid seed cultivars also exist, such as Achillea 'Summer Berries'.

Some cultivars are agamospermous plants, which retain their genetic composition and characteristics under reproduction. Occasionally cultivars are raised from seed of a specially selected provenance – for example the seed may be taken from plants that are resistant to a particular disease.

Genetically modified

Genetically modified plants with characteristics resulting from the deliberate implantation of genetic material from a different germplasm may form a cultivar. However, the International Code of Nomenclature for Cultivated Plants notes, "In practice such an assemblage is often marketed from one or more lines or multilines that have been genetically modified. These lines or multilines often remain in a constant state of development which makes the naming of such an assemblage as a cultivar a futile exercise."  However, retired transgenic varieties such as the Fish tomato, which are no longer being developed, do not run into this obstacle and can be given a cultivar name.

Cultivars may be selected because of a change in the ploidy level of a plant which may produce more desirable characteristics.

Cultivar names

Viola 'Clear Crystals Apricot'
The specific epithet may be omitted from a cultivar name

Every unique cultivar has a unique name within its denomination class (which is almost always the genus). Names of cultivars are regulated by the International Code of Nomenclature for Cultivated Plants, and may be registered with an International Cultivar Registration Authority (ICRA). There are sometimes separate registration authorities for different plant types such as roses and camellias. In addition, cultivars may be associated with commercial marketing names referred to in the Cultivated Plant Code as "trade designations" (see below).

Presenting in text

A cultivar name consists of a botanical name (of a genus, species, infraspecific taxon, interspecific hybrid or intergeneric hybrid) followed by a cultivar epithet. The cultivar epithet is enclosed by single quotes; it should not be italicized if the botanical name is italicized; and each of the words within the epithet is capitalized (with some permitted exceptions such as conjunctions). It is permissible to place a cultivar epithet after a common name provided the common name is botanically unambiguous. Cultivar epithets published before 1 January 1959 were often given a Latin form and can be readily confused with the specific epithets in botanical names; after that date, newly coined cultivar epithets must be in a modern vernacular language to distinguish them from botanical epithets.

Examples of correct text presentation:

Cryptomeria japonica 'Elegans'

Chamaecyparis lawsoniana 'Aureomarginata' (pre-1959 name, Latin in form)

Chamaecyparis lawsoniana 'Golden Wonder' (post-1959 name, English language)

Pinus densiflora 'Akebono' (post-1959 name, Japanese language)

Apple 'Sundown'

Some incorrect text presentation examples:

Cryptomeria japonica "Elegans" (double quotes are unacceptable)

Berberis thunbergii cv. 'Crimson Pygmy' (this once-common usage is now unacceptable, as it is no longer correct to use "cv." in this context; Berberis thunbergii 'Crimson Pygmy' is correct)

Rosa cv. 'Peace' (this is now incorrect for two reasons: firstly, the use of "cv."; secondly, "Peace" is a trade designation or "selling name" for the cultivar R. 'Madame A. Meilland' and should therefore be printed in a different typeface from the rest of the name, without quote marks, for example: Rosa Peace.)

Group names

Where several very similar cultivars exist they can be associated into a Group (formerly Cultivar-group). As Group names are used with cultivar names it is necessary to understand their way of presentation. Group names are presented in normal type and the first letter of each word capitalised as for cultivars, but they are not placed in single quotes. When used in a name, the first letter of the word "Group" is itself capitalized.

Presenting in text

Brassica oleracea Capitata Group (the group of cultivars including all typical cabbages)

Brassica oleracea Botrytis Group (the group of cultivars including all typical cauliflowers)

Hydrangea macrophylla Groupe Hortensia (in French) = Hydrangea macrophylla Hortensia Group (in English)

Where cited with a cultivar name the group should be enclosed in parentheses, as follows:

Hydrangea macrophylla (Hortensia Group) 'Ayesha' 

Legal protection of cultivars and their names

Since the 1990s there has been an increasing use of legal protection for newly produced cultivars. Plant breeders expect legal protection for the cultivars they produce. According to proponents of such protections, if other growers can immediately propagate and sell these cultivars as soon as they come on the market, the breeder's benefit is largely lost. Legal protection for cultivars is obtained through the use of Plant breeders’ rights and plant Patents but the specific legislation and procedures needed to take advantage of this protection vary from country to country.

Controversial use of legal protection for cultivars

The use of legal protection for cultivars can be controversial, particularly for food crops that are staples in developing countries, or for plants selected from the wild and propagated for sale without any additional breeding work; some people consider this practice unethical.

Trade designations and selling names

The formal scientific name of a cultivar, like Solanum tuberosum ‘King Edward’, is a way of uniquely designating a particular kind of plant. This scientific name is in the public domain and cannot be legally protected. Plant retailers wish to maximize their share of the market and one way of doing this is to replace the cumbersome Latin scientific names on plant labels in retail outlets with appealing marketing names that are easy to use, pronounce, and remember. Marketing names lie outside the scope of the Cultivated Plant Code which refers to them as "trade designations". If a retailer or wholesaler has the sole legal rights to a marketing name then that may offer a sales advantage. Plants protected by plant breeders' rights (PBR) may have a "true" cultivar name – the recognized scientific name in the public domain – and a "commercial synonym" – an additional marketing name that is legally protected. An example would be Rosa Fascination = 'Poulmax', in which Rosa is the genus, Fascination is the trade designation, and ‘Poulmax’ is scientific cultivar name.

Because a name that is attractive in one language may have less appeal in another country, a plant may be given different selling names from country to country. Quoting the original cultivar name allows the correct identification of cultivars around the world.

The main body coordinating plant breeders' rights is the International Union for the Protection of New Varieties of Plants (Union internationale pour la protection des obtentions végétales, UPOV) and this organization maintains a database of new cultivars protected by PBR in all countries.

International Cultivar Registration Authorities

Dahlia 'Akita'
A cultivar selected for flower form and colour

An International Cultivar Registration Authority (ICRA) is a voluntary, non-statutory organization appointed by the Commission for Nomenclature and Cultivar Registration of the International Society of Horticultural Science. ICRAs are generally formed by societies and institutions specializing in particular plant genera such as Dahlia or Rhododendron and are currently located in Europe, North America, China, India, Singapore, Australia, New Zealand, South Africa and Puerto Rico.

Each ICRA produces an annual report and its reappointment is considered every four years. The main task is to maintain a register of the names within the group of interest and where possible this is published and placed in the public domain. One major aim is to prevent the duplication of cultivar and Group epithets within a genus, as well as ensuring that names are in accord with the latest edition of the Cultivated Plant Code. In this way, over the last 50 years or so, ICRAs have contributed to the stability of cultivated plant nomenclature. In recent times many ICRAs have also recorded trade designations and trademarks used in labelling plant material, to avoid confusion with established names.

New names and other relevant data are collected by and submitted to the ICRA and in most cases there is no cost. The ICRA then checks each new epithet to ensure that it has not been used before and that it conforms with the Cultivated Plant Code. Each ICRA also ensures that new names are formally established (i.e. published in hard copy, with a description in a dated publication). They record details about the plant, such as parentage, the names of those concerned with its development and introduction, and a basic description highlighting its distinctive characters. ICRAs are not responsible for assessing the distinctiveness of the plant in question. Most ICRAs can be contacted electronically and many maintain web sites: for an up-to-date listing.

Mutation breeding

From Wikipedia, the free encyclopedia

Mutation breeding, sometimes referred to as "variation breeding", is the process of exposing seeds to chemicals or radiation in order to generate mutants with desirable traits to be bred with other cultivars. Plants created using mutagenesis are sometimes called mutagenic plants or mutagenic seeds. From 1930 to 2014 more than 3200 mutagenic plant varieties were released that have been derived either as direct mutants (70%) or from their progeny (30%). Crop plants account for 75% of released mutagenic species with the remaining 25% ornamentals or decorative plants. However, although the FAO/IAEA reported in 2014 that over 1,000 mutant varieties of major staple crops were being grown worldwide, it is unclear how many of these varieties are currently used in agriculture or horticulture around the world, as these seeds are not always identified or labeled as being mutagen or having a mutagenic provenance.

Process

There are different kinds of mutagenic breeding such as using chemical mutagens like ethyl methanesulfonate and dimethyl sulfate, radiation and transposons are used to generate mutants. Mutation breeding is commonly used to produce traits in crops such as larger seeds, new colors, or sweeter fruits, that either cannot be found in nature or have been lost during evolution.

Radiation breeding

Exposing plants to radiation is sometimes called radiation breeding and is a sub class of mutagenic breeding. Radiation breeding was discovered in the 1920s when Lewis Stadler of the University of Missouri used X-rays on maize and barley. In the case of barley, the resulting plants were white, yellow, pale yellow and some had white stripes. In 1928, Stadler first published his findings on radiation-induced mutagenesis in plants. During the period 1930–2004, radiation-induced mutant varieties were developed primarily using gamma rays (64%) and X-rays (22%).


Radiation breeding may take place in atomic gardens; and seeds have been sent into orbit in order to expose them to more cosmic radiation.

Use of chemical mutagens

High rates of chromosome aberrations resulting from ionizing radiation and the accompanied detrimental effects made researchers look for alternate sources for inducing mutations. As a result, an array of chemical mutagens has been discovered. The most widely used chemical mutagens are alkylating agents. Ethyl methanesulfonate (EMS) is the most popular because of its effectiveness and ease of handling, especially its detoxification through hydrolysis for disposal. Nitroso compounds are the other alkylating agents widely used, but they are light-sensitive and more precautions need to be taken because of their higher volatility. EMS has become a commonly used mutagen for developing large numbers of mutants for screening such as in developing TILLING populations. Although many chemicals are mutagens, only few have been used in practical breeding as the doses need to be optimised and also because the effectiveness is not high in plants for many.

History

According to garden historian Paige Johnson

After WWII, there was a concerted effort to find 'peaceful' uses for atomic energy. One of the ideas was to bombard plants with radiation and produce lots of mutations, some of which, it was hoped, would lead to plants that bore more heavily or were disease or cold-resistant or just had unusual colors. The experiments were mostly conducted in giant gamma gardens on the grounds of national laboratories in the US but also in Europe and countries of the former USSR.

Comparison to other agronomic techniques

In the debate over genetically modified foods, the use of transgenic processes is often compared and contrasted with mutagenic processes. While the abundance and variation of transgenic organisms in human food systems, and their effect on agricultural biodiversity, ecosystem health and human health is somewhat well documented, mutagenic plants and their role on human food systems is less well known, with one journalist writing "Though poorly known, radiation breeding has produced thousands of useful mutants and a sizable fraction of the world's crops...including varieties of rice, wheat, barley, pears, peas, cotton, peppermint, sunflowers, peanuts, grapefruit, sesame, bananas, cassava and sorghum." In Canada crops generated by mutation breeding face the same regulations and testing as crops obtained by genetic engineering. Mutagenic varieties tend to be made freely available for plant breeding, in contrast to many commercial plant varieties or germplasm that increasingly have restrictions on their use such as terms of use, patents and proposed genetic user restriction technologies and other intellectual property regimes and modes of enforcement.

Unlike genetically modified crops, which typically involve the insertion of one or two target genes, plants developed via mutagenic processes with random, multiple and unspecific genetic changes have been discussed as a concern but are not prohibited by any nation's organic standards. Reports from the US National Academy of Sciences state that there is no scientific justification for regulating genetic engineered crops while not doing so for mutation breeding crops.

Several organic food and seed companies promote and sell certified organic products that were developed using both chemical and nuclear mutagenesis. Several certified organic brands, whose companies support strict labeling or outright bans on GMO-crops, market their use of branded wheat and other varietal strains which were derived from mutagenic processes without any reference to this genetic manipulation. These organic products range from mutagenic barley and wheat ingredient used in organic beers to mutagenic varieties of grapefruits sold directly to consumers as organic.

New mutagen techniques

Restriction endonucleases

Interest in the use of bacterial restriction endonucleases (RE) to study double-stranded breaks in plant DNA began in the mid-nineties. These breaks in DNA, otherwise known as DSBs, were found to be the source of much chromosomal damage in eukaryotes, causing mutations in plant varieties. REs induce a result on plant DNA similar to that of ionizing radiation or radiomimetic chemicals. Blunt ended breaks in the DNA, unlike sticky ended breaks, were found to produce more variations in chromosomal damage, making them the more useful type of break for mutation breeding. While the connection of REs to chromosomal aberrations is mostly limited to research on mammalian DNA, success in mammalian studies caused scientists to conduct more studies of RE-induced chromosomal and DNA damaged on barley genomes. Due to restriction endonucleases' ability to facilitate damage in chromosomes and DNA, REs have the capability of being used as a new method of mutagenesis to promote the proliferation of mutated plant varieties.

Space-breeding

The ability of plants to develop and thrive is dependent on conditions such as microgravity and cosmic radiation in space. China has been experimenting with this theory by sending seeds into space, testing to see if space flights will cause genetic mutations. Since 1987, China has cultivated 66 mutant varieties from space through their space-breeding program. Chromosomal aberrations greatly increased when seeds were sent into aerospace compared to their earth-bound counterparts. The effect of space flight on seeds depends on their species and variety. For example, space-bred wheat saw a large growth in seed germination in compared to its Earth-bound control, but space-bred rice had no visible advantage compared to its control. For the varieties that were positively mutated by space flight, their growth potential exceeded that of not only their Earth-grown counterparts, but also their irradiated counterparts on Earth. Compared to traditional mutagenic techniques, space-bred mutations have greater efficacy in that they experience positive effects on their first generation of mutation, whereas irradiated crops often see no advantageous mutations in their first generations. Though multiple experiments have shown the positive effects of space flight on seed mutation, there is no clear connection as to what aspect of aerospace has produced such advantageous mutations. There is much speculation around cosmic radiation being the source of chromosomal aberrations, but so far, there has been no concrete evidence of such connection. Though China's space-breeding program has been shown to be very successful, the program requires a large budget and technological support that many other countries are either unwilling or unable to provide, meaning this program is unfeasible outside of China. Due to such restraints, scientists have been trying to replicate space condition on Earth in order to promote the same expedient space-born mutations on Earth. One such replication is a magnetic field-free space (MF), which produces an area with a weaker magnetic field than that of Earth. MF treatment produced mutagenic results, and has been used to cultivate new mutant varieties of rice and alfalfa. Other replications of space conditions include irradiation of seeds by a heavy 7 Li-ion beam or mixed high-energy particles. These space-bred varieties are already being introduced to the public. In 2011, during the National Lotus Flowers Exhibition in China, a mutant lotus, called the "Outer Space Sun", was shown at the flower show.

Ion beam technology

Ion beams mutate DNA by deleting multiple bases from its code. Compared to traditional sources of radiation, like gamma rays and X-rays, ion beams have been shown to cause more severe breaks in DNA that are more difficult to weave back together, causing the change in DNA to be more drastic than changes caused by traditional irradiation. Ion beams change DNA in a manner that makes it look vastly different than its original makeup, more so than when traditional irradiation techniques are used. Most experimentation, using ion beam technology, has been conducted in Japan. Notable facilities using this technology are TIARA of the Japan Atomic Energy Agency, RIKEN Accelerator Research Facility, and various other Japanese institutions. During the process of ion beam radiation, seeds are wedged between two kapton films and irradiated for roughly two minutes. Mutation frequencies are notably higher for ion beam radiation compared to electron radiation, and the mutation spectrum is broader for ion beam radiation compared to gamma ray radiation. The broader mutation spectrum was revealed through the largely varied amount of flower phenotypes produced by ion beams. Flowers mutated by the ion beams exhibited a variety of colors, patterns, and shapes. Through ion beam radiation, new varieties of plants have been cultivated. These plants had the characteristics of being ultraviolet light-B resistant, disease resistant, and chlorophyll-deficient. Ion beam technology has been used in the discovery of new genes responsible for the creation of more robust plants, but its most prevalent use is commercially for producing new flower phenotypes, like striped chrysanthemums.

Mature pollen treated with gamma radiation

Gamma radiation is used on mature rice pollen to produce parent plants used for crossing. The mutated traits in the parent plants are able to be inherited by their offspring plants. Because rice pollen has a very short lifespan, researchers had to blast gamma rays at cultured spikes from rice plants. Through experimentation, it was revealed that there was a greater variety of mutation in irradiated pollen rather than irradiated dry seeds. Pollen treated with 46Gy of gamma radiation showed an increase in grain size overall and other useful variations. Typically, the length of each grain was longer after the crossing of irradiated parent rice plants. The rice progeny also exhibited a less chalky visage, improving on the appearance of the parent rice plants. This technique was used to develop two new rice cultivars, Jiaohezaozhan and Jiafuzhan, in China. Along with facilitating the creation of these two rice cultivars, the irradiation of mature rice pollen has produced roughly two hundred mutant rice lines. Each of these lines produce rice grains of both a higher quality and larger size. The mutations produced by this technique vary with each generation, meaning further breeding of these mutated plants could produce new mutations. Traditionally, gamma radiation is used on solely adult plants, and not on pollen. The irradiation of mature pollen allows mutant plants to grow without being in direct contact with gamma radiation. This discovery is in contrast to what was previously believed about gamma radiation: that it could only elicit mutations in plants and not pollen.

Notable mutagen varieties

 Argentina

• Colorado Irradiado groundnut (mutant created with X-rays; high fat content and yield, 80% of groundnuts grown in Argentina in the 1980s was Colorado Irradiado)
• Puita INTA-CL rice mutant (herbicide resistance and good yield; also grown in Bolivia, Brazil, Costa Rica and Paraguay)

 Australia

• Amaroo rice mutant variety (60-70% of rice grown in Australia was Amaroo in 2001)

 Bangladesh

• Binasail, Iratom-24 and Binadhan-6 rice mutants 
• Binamoog-5 mung bean mutant variety

 Cuba

• Maybel tomato mutant (excellent drought resistance)
• GINES rice mutant (created using proton radiation; grows well in salty conditions)

 People's Republic of China

• Henong series soybean mutants
• Jiahezazhan and Jiafuzhan rice (mutations obtained by pollen irradiation; high yield and quality, very adaptable, resistant to plant hopper and blast)
• Lumian Number 1 cotton
• Purple Orchard 3 Sweet potato
• Tiefeng 18 soybean
• Yangdao Number 6 rice
• Yangmai 156 wheat
• Zhefu 802 rice mutant (irradiated with gamma rays; resistant to rice blast, good yield even in poor conditions, the most planted rice variety between 1986-1994)
• 26Zhaizao indica rice mutant (created with gamma rays)

 Czech Republic

• Diamant barley (high yield, short height mutant created with X-Rays)

 Egypt

• Giza 176 and Sakha 101 high yield rice mutants

 Finland

• Balder J barley mutant (better drought resistance, yield and sprouting)
• Puhti and Ryhti stiff straw oat mutants

 France

• High oleic sunflowers (covering more than 50 % of the sunflower acreage)

 Germany

• Trumpf barley

 Ghana

• Tek bankye mutant cassava (good poundability and increased dry matter content)

 India

• Co-4, Pant Mung-2, and TAP mung bean mutants
• MA-9 cotton - the world's first mutant cotton, released in 1948 (X-ray radiation; drought tolerance, high yielding)
• PNR-381 Rice
• Pusa 408 (Ajay), Pusa 413 (Atul), Pusa 417 (Girnar), and Pusa 547 chickpea mutants (resistant to Ascochyta blight and wilt diseases, and have high yields)
• Sharbati Sonora wheat
• Tau-1, MUM 2, BM 4, LGG 407, LGG 450, Co4, Dhauli (TT9E) and Pant moong-1 blackgram (YMC, (Yellow mosaic virus) resistance)
• TG24 and TG37 groundnut mutants

 Italy

• Durum wheat (especially Creso mutant, created with thermal neutrons)

 Japan

• Osa Gold Pear (disease resistance) 
• Most rice varieties grown in Japan have the sd1 mutant allele from the Reimei rice variety

 Myanmar

• Shwewartun rice mutant (created by irradiating IR5 rice to give better yield, grain quality and earlier maturity)

 Pakistan

• Basmati 370 short height rice mutant
• NIAB-78 cotton mutant (high yielding, heat tolerant, early maturing)
• CM-72 chickpea mutant (created with 150Gy of gamma rays; high yielding, blight resistant)
• NM-28 mungbean mutant (short height, uniform and early maturing, high seed yield)
• NIAB Masoor 2006 lentil mutant (created with 200Gy of radiation; early maturing, high yield, resistant to disease)

 Peru

• UNA La Molina 95 barley mutant (developed in 1995 for growing above 3,000 m)
• Centenario Amarinth "kiwicha" mutant (high quality grain and exported as a certified organic product)
• Centenario II barley mutant (developed for growing in the Andean highlands with high yield, high quality flour and tolerance to hail)

 Sudan

• Albeely banana mutant (better quality, high yield and better stand)

 Thailand

• RD15 and RD6 aromatic indica rice mutants (created with gamma rays and released in 1977-8; RD 15 is early ripening, RD6 has a valuable glutinous endosperm) Thailand is the biggest exporter of aromatic rice in the world

 United Kingdom

• Golden Promise barley (semi-dwarf, salt tolerant mutant created with gamma rays) Is used to make beer and whisky

 United States

• Calrose 76 Rice (short height rice induced with gamma rays)
• Luther and Pennrad barley (high yield mutant varieties; Pennrad also resistant to winter)
• Murray Mitcham Peppermint (Verticillium wilt tolerance)
• Sanilac bean (X-ray radiation; high yielding mutant - also the Gratiot and Sea-way bean varieties were cross-bred from Sanilac)
• Stadler wheat (high yield mutant with resistance to loose smut and leaf rust and earlier maturity)
• Star Ruby and Rio red varieties of the Rio Star Grapefruit (created using thermal neutron techniques)
• Todd's Mitcham Peppermint (Verticillium wilt tolerance)

 Vietnam

• VND 95-20, VND-99-1 and VN121 rice mutants (increased yield, improved quality, resistance to disease and pests)
• DT84, DT96, DT99 and DT 2008 soybean mutants (developed using gamma rays to grow three crops a year, tolerance to heat and cold and resistance to disease)

In 2014, it was reported that 17 rice mutant varieties, 10 soybean, two maize and one chrysanthemum mutant varieties had been officially released to Vietnamese farmers. 15% of rice and 50% of soybean was produced from mutant varieties.

Release by nation

As of 2011 the percentage of all mutagenic varieties released globally, by country, were:

• (25.2%)  People's Republic of China
• (15.0%)  Japan
• (11.5%)  India
• (6.7%)  Russia
• (5.5%)  Netherlands
• (5.3%)  Germany
• (4.3%)  United States
• (2.4%)  Bulgaria
• (1.7%)  Vietnam
• (1.4%)  Bangladesh

Selective breeding

From Wikipedia, the free encyclopedia

A Belgian Blue cow. The defect in the breed's myostatin gene is maintained through linebreeding and is responsible for its accelerated lean muscle growth.

This Chihuahua mix and Great Dane shows the wide range of dog breed sizes created using selective breeding.

Selective breeding transformed teosinte's few fruitcases (left) into modern maize's rows of exposed kernels (right).

Selective breeding (also called artificial selection) is the process by which humans use animal breeding and plant breeding to selectively develop particular phenotypic traits (characteristics) by choosing which typically animal or plant males and females will sexually reproduce and have offspring together. Domesticated animals are known as breeds, normally bred by a professional breeder, while domesticated plants are known as varieties, cultigens, or cultivars. Two purebred animals of different breeds produce a crossbreed, and crossbred plants are called hybrids. Flowers, vegetables and fruit-trees may be bred by amateurs and commercial or non-commercial professionals: major crops are usually the provenance of the professionals.

In animal breeding, techniques such as inbreeding, linebreeding, and outcrossing are utilized. In plant breeding, similar methods are used. Charles Darwin discussed how selective breeding had been successful in producing change over time in his 1859 book, On the Origin of Species. Its first chapter discusses selective breeding and domestication of such animals as pigeons, cats, cattle, and dogs. Darwin used artificial selection as a springboard to introduce and support the theory of natural selection.

The deliberate exploitation of selective breeding to produce desired results has become very common in agriculture and experimental biology.

Selective breeding can be unintentional, e.g., resulting from the process of human cultivation; and it may also produce unintended – desirable or undesirable – results. For example, in some grains, an increase in seed size may have resulted from certain ploughing practices rather than from the intentional selection of larger seeds. Most likely, there has been an interdependence between natural and artificial factors that have resulted in plant domestication.

History

Selective breeding of both plants and animals has been practiced since early prehistory; key species such as wheat, rice, and dogs have been significantly different from their wild ancestors for millennia, and maize, which required especially large changes from teosinte, its wild form, was selectively bred in Mesoamerica. Selective breeding was practiced by the Romans. Treatises as much as 2,000 years old give advice on selecting animals for different purposes, and these ancient works cite still older authorities, such as Mago the Carthaginian. The notion of selective breeding was later expressed by the Persian Muslim polymath Abu Rayhan Biruni in the 11th century. He noted the idea in his book titled India, which included various examples.

The agriculturist selects his corn, letting grow as much as he requires, and tearing out the remainder. The forester leaves those branches which he perceives to be excellent, whilst he cuts away all others. The bees kill those of their kind who only eat, but do not work in their beehive.

— Abu Rayhan Biruni, India

Selective breeding was established as a scientific practice by Robert Bakewell during the British Agricultural Revolution in the 18th century. Arguably, his most important breeding program was with sheep. Using native stock, he was able to quickly select for large, yet fine-boned sheep, with long, lustrous wool. The Lincoln Longwool was improved by Bakewell, and in turn the Lincoln was used to develop the subsequent breed, named the New (or Dishley) Leicester. It was hornless and had a square, meaty body with straight top lines.

These sheep were exported widely, including to Australia and North America, and have contributed to numerous modern breeds, despite the fact that they fell quickly out of favor as market preferences in meat and textiles changed. Bloodlines of these original New Leicesters survive today as the English Leicester (or Leicester Longwool), which is primarily kept for wool production.

Bakewell was also the first to breed cattle to be used primarily for beef. Previously, cattle were first and foremost kept for pulling ploughs as oxen, but he crossed long-horned heifers and a Westmoreland bull to eventually create the Dishley Longhorn. As more and more farmers followed his lead, farm animals increased dramatically in size and quality. In 1700, the average weight of a bull sold for slaughter was 370 pounds (168 kg). By 1786, that weight had more than doubled to 840 pounds (381 kg). However, after his death, the Dishley Longhorn was replaced with short-horn versions.

He also bred the Improved Black Cart horse, which later became the Shire horse.

Charles Darwin coined the term 'selective breeding'; he was interested in the process as an illustration of his proposed wider process of natural selection. Darwin noted that many domesticated animals and plants had special properties that were developed by intentional animal and plant breeding from individuals that showed desirable characteristics, and discouraging the breeding of individuals with less desirable characteristics.

Darwin used the term "artificial selection" twice in the 1859 first edition of his work On the Origin of Species, in Chapter IV: Natural Selection, and in Chapter VI: Difficulties on Theory:

Slow though the process of selection may be, if feeble man can do much by his powers of artificial selection, I can see no limit to the amount of change, to the beauty and infinite complexity of the co-adaptations between all organic beings, one with another and with their physical conditions of life, which may be effected in the long course of time by nature's power of selection.

— Charles Darwin, On the Origin of Species

We are profoundly ignorant of the causes producing slight and unimportant variations; and we are immediately made conscious of this by reflecting on the differences in the breeds of our domesticated animals in different countries,—more especially in the less civilized countries where there has been but little artificial selection.

— Charles Darwin, On the Origin of Species

Animal breeding

Three generations of "Westies" in a village in Fife, Scotland

Animals with homogeneous appearance, behavior, and other characteristics are known as particular breeds, and they are bred through culling animals with particular traits and selecting for further breeding those with other traits. Purebred animals have a single, recognizable breed, and purebreds with recorded lineage are called pedigreed. Crossbreeds are a mix of two purebreds, whereas mixed breeds are a mix of several breeds, often unknown. Animal breeding begins with breeding stock, a group of animals used for the purpose of planned breeding. When individuals are looking to breed animals, they look for certain valuable traits in purebred stock for a certain purpose, or may intend to use some type of crossbreeding to produce a new type of stock with different, and, it is presumed, superior abilities in a given area of endeavor. For example, to breed chickens, a breeder typically intends to receive eggs, meat, and new, young birds for further reproduction. Thus, the breeder has to study different breeds and types of chickens and analyze what can be expected from a certain set of characteristics before he or she starts breeding them. Therefore, when purchasing initial breeding stock, the breeder seeks a group of birds that will most closely fit the purpose intended.

Purebred breeding aims to establish and maintain stable traits, that animals will pass to the next generation. By "breeding the best to the best," employing a certain degree of inbreeding, considerable culling, and selection for "superior" qualities, one could develop a bloodline superior in certain respects to the original base stock. Such animals can be recorded with a breed registry, the organization that maintains pedigrees and/or stud books. However, single-trait breeding, breeding for only one trait over all others, can be problematic. In one case mentioned by animal behaviorist Temple Grandin, roosters bred for fast growth or heavy muscles did not know how to perform typical rooster courtship dances, which alienated the roosters from hens and led the roosters to kill the hens after mating with them.

The observable phenomenon of hybrid vigor stands in contrast to the notion of breed purity. However, on the other hand, indiscriminate breeding of crossbred or hybrid animals may also result in degradation of quality. Studies in evolutionary physiology, behavioral genetics, and other areas of organismal biology have also made use of deliberate selective breeding, though longer generation times and greater difficulty in breeding can make such projects challenging in vertebrates.

Plant breeding

Researchers at the USDA have selectively bred carrots with a variety of colors.

Plant breeding has been used for thousands of years, and began with the domestication of wild plants into uniform and predictable agricultural cultigens. High-yielding varieties have been particularly important in agriculture.

Selective plant breeding is also used in research to produce transgenic animals that breed "true" (i.e., are homozygous) for artificially inserted or deleted genes.

Selective breeding in aquaculture

Selective breeding in aquaculture holds high potential for the genetic improvement of fish and shellfish. Unlike terrestrial livestock, the potential benefits of selective breeding in aquaculture were not realized until recently. This is because high mortality led to the selection of only a few broodstock, causing inbreeding depression, which then forced the use of wild broodstock. This was evident in selective breeding programs for growth rate, which resulted in slow growth and high mortality.

Control of the reproduction cycle was one of the main reasons as it is a requisite for selective breeding programs. Artificial reproduction was not achieved because of the difficulties in hatching or feeding some farmed species such as eel and yellowtail farming. A suspected reason associated with the late realisation of success in selective breeding programs in aquaculture was the education of the concerned people – researchers, advisory personnel and fish farmers. The education of fish biologists paid less attention to quantitative genetics and breeding plans.

Another was the failure of documentation of the genetic gains in successive generations. This in turn led to failure in quantifying economic benefits that successful selective breeding programs produce. Documentation of the genetic changes was considered important as they help in fine tuning further selection schemes.

Quality traits in aquaculture

Aquaculture species are reared for particular traits such as growth rate, survival rate, meat quality, resistance to diseases, age at sexual maturation, fecundity, shell traits like shell size, shell colour, etc.

• Growth rate – growth rate is normally measured as either body weight or body length. This trait is of great economic importance for all aquaculture species as faster growth rate speeds up the turnover of production. Improved growth rates show that farmed animals utilize their feed more efficiently through a correlated response.
• Survival rate – survival rate may take into account the degrees of resistance to diseases. This may also see the stress response as fish under stress are highly vulnerable to diseases. The stress fish experience could be of biological, chemical or environmental influence.
• Meat quality – the quality of fish is of great economic importance in the market. Fish quality usually takes into account size, meatiness, and percentage of fat, colour of flesh, taste, shape of the body, ideal oil and omega-3 content.
• Age at sexual maturation – The age of maturity in aquaculture species is another very important attribute for farmers as during early maturation the species divert all their energy to gonad production affecting growth and meat production and are more susceptible to health problems (Gjerde 1986).
• Fecundity – As the fecundity in fish and shellfish is usually high it is not considered as a major trait for improvement. However, selective breeding practices may consider the size of the egg and correlate it with survival and early growth rate.

Finfish response to selection

Salmonids

Gjedrem (1979) showed that selection of Atlantic salmon (Salmo salar) led to an increase in body weight by 30% per generation. A comparative study on the performance of select Atlantic salmon with wild fish was conducted by AKVAFORSK Genetics Centre in Norway. The traits, for which the selection was done included growth rate, feed consumption, protein retention, energy retention, and feed conversion efficiency. Selected fish had a twice better growth rate, a 40% higher feed intake, and an increased protein and energy retention. This led to an overall 20% better Fed Conversion Efficiency as compared to the wild stock. Atlantic salmon have also been selected for resistance to bacterial and viral diseases. Selection was done to check resistance to Infectious Pancreatic Necrosis Virus (IPNV). The results showed 66.6% mortality for low-resistant species whereas the high-resistant species showed 29.3% mortality compared to wild species.

Rainbow trout (S. gairdneri) was reported to show large improvements in growth rate after 7–10 generations of selection. Kincaid et al. (1977) showed that growth gains by 30% could be achieved by selectively breeding rainbow trout for three generations. A 7% increase in growth was recorded per generation for rainbow trout by Kause et al. (2005).

In Japan, high resistance to IPNV in rainbow trout has been achieved by selectively breeding the stock. Resistant strains were found to have an average mortality of 4.3% whereas 96.1% mortality was observed in a highly sensitive strain.

Coho salmon (Oncorhynchus kisutch) increase in weight was found to be more than 60% after four generations of selective breeding. In Chile, Neira et al. (2006) conducted experiments on early spawning dates in coho salmon. After selectively breeding the fish for four generations, spawning dates were 13–15 days earlier.

Cyprinids

Selective breeding programs for the Common carp (Cyprinus carpio) include improvement in growth, shape and resistance to disease. Experiments carried out in the USSR used crossings of broodstocks to increase genetic diversity and then selected the species for traits like growth rate, exterior traits and viability, and/or adaptation to environmental conditions like variations in temperature. Kirpichnikov et al. (1974) and Babouchkine (1987) selected carp for fast growth and tolerance to cold, the Ropsha carp. The results showed a 30–40% to 77.4% improvement of cold tolerance but did not provide any data for growth rate. An increase in growth rate was observed in the second generation in Vietnam. Moav and Wohlfarth (1976) showed positive results when selecting for slower growth for three generations compared to selecting for faster growth. Schaperclaus (1962) showed resistance to the dropsy disease wherein selected lines suffered low mortality (11.5%) compared to unselected (57%).

Channel Catfish

Growth was seen to increase by 12–20% in selectively bred Iictalurus punctatus. More recently, the response of the Channel Catfish to selection for improved growth rate was found to be approximately 80%, i.e., an average of 13% per generation.

Shellfish response to selection

Oysters

Selection for live weight of Pacific oysters showed improvements ranging from 0.4% to 25.6% compared to the wild stock. Sydney-rock oysters (Saccostrea commercialis) showed a 4% increase after one generation and a 15% increase after two generations. Chilean oysters (Ostrea chilensis), selected for improvement in live weight and shell length showed a 10–13% gain in one generation. Bonamia ostrea is a protistan parasite that causes catastrophic losses (nearly 98%) in European flat oyster Ostrea edulis L. This protistan parasite is endemic to three oyster-regions in Europe. Selective breeding programs show that O. edulis susceptibility to the infection differs across oyster strains in Europe. A study carried out by Culloty et al. showed that ‘Rossmore' oysters in Cork harbour, Ireland had better resistance compared to other Irish strains. A selective breeding program at Cork harbour uses broodstock from 3– to 4-year-old survivors and is further controlled until a viable percentage reaches market size. Over the years ‘Rossmore' oysters have shown to develop lower prevalence of B. ostreae infection and percentage mortality. Ragone Calvo et al. (2003) selectively bred the eastern oyster, Crassostrea virginica, for resistance against co-occurring parasites Haplosporidium nelson (MSX) and Perkinsus marinus (Dermo). They achieved dual resistance to the disease in four generations of selective breeding. The oysters showed higher growth and survival rates and low susceptibility to the infections. At the end of the experiment, artificially selected C. virginica showed a 34–48% higher survival rate.

Penaeid shrimps

Selection for growth in Penaeid shrimps yielded successful results. A selective breeding program for Litopenaeus stylirostris saw an 18% increase in growth after the fourth generation and 21% growth after the fifth generation. Marsupenaeus japonicas showed a 10.7% increase in growth after the first generation. Argue et al. (2002) conducted a selective breeding program on the Pacific White Shrimp, Litopenaeus vannamei at The Oceanic Institute, Waimanalo, USA from 1995 to 1998. They reported significant responses to selection compared to the unselected control shrimps. After one generation, a 21% increase was observed in growth and 18.4% increase in survival to TSV. The Taura Syndrome Virus (TSV) causes mortalities of 70% or more in shrimps. C.I. Oceanos S.A. in Colombia selected the survivors of the disease from infected ponds and used them as parents for the next generation. They achieved satisfying results in two or three generations wherein survival rates approached levels before the outbreak of the disease. The resulting heavy losses (up to 90%) caused by Infectious hypodermal and haematopoietic necrosis virus (IHHNV) caused a number of shrimp farming industries started to selectively breed shrimps resistant to this disease. Successful outcomes led to development of Super Shrimp, a selected line of L. stylirostris that is resistant to IHHNV infection. Tang et al. (2000) confirmed this by showing no mortalities in IHHNV- challenged Super Shrimp post larvae and juveniles.

Aquatic species versus terrestrial livestock

Selective breeding programs for aquatic species provide better outcomes compared to terrestrial livestock. This higher response to selection of aquatic farmed species can be attributed to the following:

• High fecundity in both sexes fish and shellfish enabling higher selection intensity.
• Large phenotypic and genetic variation in the selected traits.

Selective breeding in aquaculture provide remarkable economic benefits to the industry, the primary one being that it reduces production costs due to faster turnover rates. This is because of faster growth rates, decreased maintenance rates, increased energy and protein retention, and better feed efficiency. Applying such genetic improvement program to aquaculture species will increase productivity to meet the increasing demands of growing populations.

Advantages and disadvantages

Selective breeding is a direct way to determine if a specific trait can evolve in response to selection. A single-generation method of breeding is not as accurate or direct. The process is also more practical and easier to understand than sibling analysis. Selective breeding is better for traits such as physiology and behavior that are hard to measure because it requires fewer individuals to test than single-generation testing.

However, there are disadvantages to this process. Because a single experiment done in selective breeding cannot be used to assess an entire group of genetic variances, individual experiments must be done for every individual trait. Also, because of the necessity of selective breeding experiments to require maintaining the organisms tested in a lab or greenhouse, it is impractical to use this breeding method on many organisms. Controlled mating instances are difficult to carry out in this case and this is a necessary component of selective breeding.