Animal husbandry is the management and care of farm animals by humans for profit, in which genetic

qualities and behaviour, considered to be advantageous to humans, are further developed. The term can
refer to the practice of selectively breeding and raising livestock to promote desirable traits in animals for
utility, sport, pleasure, or research,
[1]
but also refers to the efficient exploitation of a species in agriculture
advantageous to humans.
[2]

Contents
[show]
History of breeding[edit]
See also: Selective breeding
Animal husbandry has been practiced for thousands of years, since the first domestication of animals.
Selective breeding for desired traits was first established as a scientific practice by Robert
Bakewell during the British Agricultural Revolution in the 18th century. One of his most important breeding
programs was with sheep. Using native stock, he was able to quickly select for large, yet fine-boned
sheep, with long, lustrous wool. TheLincoln 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.
[3]
These sheep were exported widely and have
contributed to numerous modern breeds.
Under his influence, English farmers began to breed cattle for use primarily as beef for consumption -
(previously, cattle were first and foremost bred for pulling ploughs as oxen
[citation needed]
). Long-horned
heifers were crossed with the Westmoreland bull to eventually create the Dishley Longhorn. Over the
following decades, farm animals increased dramatically in size and quality. In 1700, the average weight of
a bull sold for slaughter was 370pounds (168 kg). By 1786, that weight had more than doubled to 840
pounds (381 kg).
Animal herding professions specialized in the 19th century to include the cowboys of the United States
and Canada, charros and vaqueros of Mexico,gauchos and huasos of South America, and the farmers
and stockmen of Australia.
In more modern times herds are tended on horses, all-terrain vehicles, motorbikes, four-wheel
drive vehicles, and helicopters, depending on the terrain and livestock concerned. Today, herd managers
often oversee thousands of animals and many staff. Farms, stations and ranches may employ breeders,
herd health specialists, feeders, and milkers to help care for the animals.
Breeding techniques[edit]
Techniques such as artificial insemination and embryo transfer are frequently used today, not only as
methods to guarantee that females breed regularly but also to help improve herd genetics. This may be
done by transplanting embryos from high-quality females into lower-quality surrogate mothers - freeing up
the higher-quality mother to be reimpregnated. This practice vastly increases the number of offspring
which may be produced by a small selection of the best quality parent animals. On the one hand, this
improves the ability of the animals to convert feed to meat, milk, or fiber more efficiently, and improve the
quality of the final product. On the other, it decreases genetic diversity, increasing the severity of disease
outbreaks among other risks.
History in Europe[edit]
The semi-natural, unfertilized pastures formed by traditional agricultural methods in Europe, were
managed and maintained by the grazing and mowing of livestock.
[4]
Because the ecological impact of this
land management strategy is similar to the impact of a natural disturbance, the agricultural system will
share many beneficial characteristics with a natural habitat including the promotion of biodiversity.
[4]
This
strategy is declining in the European context due to the intensification of agriculture,
[4]
and the
mechanized chemical based methods that became popular during and following the industrial revolution.
Sustainability[edit]
In organic and biodynamic farming methods, animals play an important role in achieving closed or
sustainable system by providing multiple functions to the agricultural system. In the Balinese context, for
example, ducks are herded through rice paddies at particular stages of the growing season. The ducks
eat insects and weeds that inhibit the growth of seedlings, and prevent the farmer from using herbicides
or pesticides.
[5]
Their droppings contribute nitrogen to the soil as a natural source of fertilizer, and the
movement of their feet and bills in the water of the terrace ecosystem increases the oxygen content of the
soil thus increasing nutrient availability.
[5]

Climate change[edit]
Due to the significant contribution of agriculture to the emissions of non-CO2 greenhouse gases methane
and nitrous oxide, the relationship between humans and livestock is being analyzed for its potential to
help mitigate climate change. Strategies for the mitigation include optimizing the use of gas produced
from manure for energy production (biogas).
[6]





















Selective breeding (also called artificial selection) is the process by which humans breed
other animals and plants for particular traits. Typically, strains that are selectively bred are domesticated,
and the breeding is normally done by a professional breeder. Bred animals are known as breeds, while
bred plants are known as varieties, cultigens, or cultivars. The cross of animals results in what is called
a crossbreed, and crossbred plants are calledhybrids.
There are two approaches or types of artificial selection, or selective breeding. First is the traditional
breeders approach in which the breeder or experimenter applies a known amount of selection to a
single phenotypic trait by examining the chosen trait and choosing to breed only those that exhibit higher
or extreme values of that trait. The second is called controlled natural selection, which is
essentially natural selection in a controlled environment. In this, the breeder does not choose which
individuals being tested survive or reproduce, as he or she could in the traditional approach. There are
also selection experiments, which is a third approach and these are conducted in order to determine the
strength of natural selection in the wild. However, this is more often an observational approach as
opposed to an experimental approach.
[1]

In animal breeding, techniques such as inbreeding, linebreeding, and outcrossing are utilized. In plant
breeding, similar methods are used. Charles Darwindiscussed how selective breeding had been
successful in producing change over time in his book, On the Origin of Species. The first chapter of the
book discusses selective breeding and domestication of such animals as pigeons, cats, cattle, and dogs.
Selective breeding was used by Darwin as a springboard to introduce the theory of natural selection, and
to support it.
[2]

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.
[3]

Contents
[show]
History[edit]
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.
[4]
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.
[5]
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, and gave various examples.
[6]

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.
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.
[7]

These sheep were exported widely, including to Australia and North America, and have contributed to
numerous modern breeds, despite that 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 theEnglish
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 asoxen
[citation needed]
, 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 breedingfrom
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.
[8]

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.
[9]

Animal breeding[edit]
Main article: Animal breeding
Animals with homogeneous appearance, behavior, and other characteristics are known as particular
breeds, and they are bred through culling particular traits and selecting for others. 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
typical breeder 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.
[10]
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
reproducing with them.
[10]

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.
[11][12][13]

Plant breeding[edit]
Main article: 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-yieldingvarieties 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 deletedgenes.
[citation needed]

Selective breeding in aquaculture[edit]
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.
[14]

Control of the reproduction cycle was one of the main reasons as it is a requisite for selective breeding
programmes. Artificial reproduction was not achieved because of the difficulties in hatching or feeding
some farmed species such as eel and yellowtail farming.
[15]
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.
[16]

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.
[14]

Quality traits in aquaculture[edit]
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. (Gjedrem
1985). This trait is of great economic importance for all aquaculture species as faster growth rate
speeds up the turnover of production (Gjedrem 1983). Improved growth rates show that farmed
animals utilize their feed more efficiently through a correlated response (Gjedrem 1985).
Survival rate survival rate may take into account the degrees of resistance to diseases (Gjedrem
1985). This may also see the stress response as fish under stress are highly vulnerable to diseases
(Gjedrem 1983). 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 (Gjedrem 1985).
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 (Gjedrem 1985).
Finfish response to selection[edit]
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 (Thodeson et al.1999). 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 (Storset et al. 2007).
Rainbow trout (S. gairdneri) was reported to show large improvements in growth rate after 7-10
generations of selection (Donaldson and Olson 1957). 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
(Okamoto et al. 1993). Coho salmon (Oncorhynchus kisutch) increase in weight was found to be more
than 60% after four generations of selective breeding (Hershberger et al. 1990). 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 (Tran and Nguyen
1993). 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 (Bondari, 1983). More
recently, the overall response of Channel Catfish response to selection for improved growth rate was
found to be approximately 80%, i.e., an average of 13% per generation (Dunham 2006).
Shellfish response to selection[edit]
Oysters
Selection for live weight of Pacific oysters showed improvements ranging from 0.4% to 25.6% compared
to the wild stock (Langdon et al. 2003). Sydney-rock oysters (Saccostrea commercialis) showed a 4%
increase after one generation and a 15% increase after two generations (Nell et al. 1996, 1999). 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. (2001) 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 (Culloty et al. 2004). Over the years Rossmore oysters have shown to
develop lower prevalence to B. ostreae infection and percentage mortality. Ragone Calvo et al. (2003)
selectively bred the eastern oyster, Crassostrea virginica, for resistance against co-occurring
parasitesHaplosporidium 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 (Goyard et al. 1999). Marsupenaeus japonicas showed a 10.7% increase in
growth after the first generation (Hetzel et al. 2000). 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 form 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 (Cock et al. 2009). 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). Tang et al. (2000) confirmed this by showing no mortalities in IHHNV-
challenged Super Shrimp post larvae and juveniles.
Aquatic species versus terrestrial livestock[edit]
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
(Gjedrem and Baranski 2009). Applying such genetic improvement program to aquaculture species will
increase productivity to meet the increasing demands of growing populations.

Advantages and disadvantages[edit]
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. The former tests differences between line means while the
latter is dependent upon variance and covariance components. Essentially, 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 and covariances, 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.
[1]













Plant breeding is the art and science of changing the traits of plants in order to produce desired
characteristics.
[1]
Plant breeding can be accomplished through many different techniques ranging from
simply selecting plants with desirable characteristics for propagation, to more complex molecular
techniques (see cultigen and cultivar).
Plant breeding has been practiced for thousands of years, since near the beginning of human civilization.
It is now practiced worldwide by individuals such as gardeners and farmers, or by professional plant
breeders employed by organizations such as government institutions, universities, crop-specific industry
associations or research centers.
International development agencies believe that breeding new crops is important for ensuring food
security by developing new varieties that are higher-yielding, resistant to pests and diseases, drought-
resistant or regionally adapted to different environments and growing conditions.
Contents
[show]
History[edit]
Main article: History of plant breeding
Plant breeding started with sedentary agriculture and particularly the domestication of the
first agricultural plants, a practice which is estimated to date back 9,000 to 11,000 years.
[2]
Initially early
farmers simply selected food plants with particular desirable characteristics, and employed these as
progenitors for subsequent generations, resulting in an accumulation of valuable traits over time.
Gregor Mendel's experiments with plant hybridization led to his establishing laws of inheritance. Once this
work became well known, it formed the basis of the new science of genetics, which stimulated research
by many plant scientists dedicated to improving crop production through plant breeding.
Modern plant breeding is applied genetics, but its scientific basis is broader, covering molecular
biology, cytology, systematics, physiology, pathology, entomology, chemistry, and statistics(biometrics). It
has also developed its own technology.
Classical plant breeding[edit]

This section needs additional citations
for verification. Please help improve this
article by adding citations to reliable
sources. Unsourced material may be
challenged and removed. (December 2011)
For the role of crossing and plant breeding in viticulture, see Propagation of grapevines.
Classical plant breeding uses 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 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
vitrotechniques 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 in the last 100 years include:
1. Increased quality and yield of the crop
2. Increased tolerance of environmental pressures (salinity, extreme temperature, drought)
3. Resistance to viruses, fungi and bacteria
4. Increased tolerance to insect pests
5. Increased tolerance of herbicides
Before World War II[edit]


Garton's catalogue from 1902
Intraspecific hybridization within a plant species was the first process to be developed.
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.
[3]
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.
[4][5]

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. Wheat hybrids were bred to increase the crop production of Italy during
the so-called "Battle for Grain" (19251940). 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
reproducewith 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
divisioninhibitor 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 hybridembryo 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 calledsomaclonal
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:
1. Collection of variation
2. Selection
3. Evaluation
4. release
5. Multiplication
6. 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 orDNA 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 vegetativepropagation 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 calledcisgenic 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 ormicroinjection. 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 and related species.
Another limitation of viral vectors is that the virus is not usually passed on the progeny, so
every plant has to be inoculated.
The majority of commercially released transgenic plants 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, the cotton bollworm, a common cotton pest, feeds on Bt
cotton it will ingest the toxin and die. Herbicides usually work by binding to certain plant
enzymes and inhibiting their action. The enzymes that the herbicide inhibits are known as
the herbicides target site. Herbicide resistance can be engineered into crops by expressing a
version of target site protein that is not inhibited by the herbicide. This is the method used to
produce glyphosate resistant crop plants (See Glyphosate)
Genetic modification of plants that can produce pharmaceuticals (and industrial chemicals),
sometimes called pharmacrops, is a rather radical new area of plant breeding.
Issues and concerns[edit]
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 at Austin, 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.
[6]
"
The debate surrounding genetically modified food during the 1990s peaked in 1999 in terms
of media coverage and risk perception,
[7]
and continues today - for example, "Germany has
thrown its weight behind a growing European mutiny over genetically modified crops by
banning the planting of a widely grown pest-resistant corn variety.".
[8]
The debate
encompasses the ecological impact of genetically modified plants, the safety of genetically
modified food and concepts used for safety evaluation like substantial equivalence. Such
concerns are not new to plant breeding. Most countries have regulatory processes in place
to help ensure that new crop varieties entering the marketplace are both safe and meet
farmers' needs. Examples include variety registration, seed schemes, regulatory
authorizations for GM plants, etc.
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
[citation needed]
. Efforts to strengthen breeders' rights, for example, by lengthening periods of
variety protection
[citation needed]
, are ongoing.
When new plant breeds or cultivars are bred, they must be maintained and propagated.
Some plants are propagated by asexual means while others are propagated by seeds. Seed
propagated cultivars require specific control over seed source and production procedures to
maintain the integrity of the plant breeds results. Isolation is necessary to prevent cross
contamination with related plants or the mixing of seeds after harvesting. Isolation is
normally accomplished by planting distance but in certain crops, plants are enclosed in
greenhouses or cages (most commonly used when producing F1 hybrids.)
Role of plant breeding in organic agriculture[edit]
Critics of organic agriculture claim it is too low-yielding to be a viable alternative to
conventional agriculture. However, part of that poor performance may be the result of
growing poorly adapted varieties.
[9][10]
It is estimated that over 95% of organic agriculture is
based on conventionally adapted varieties, even though the production environments found
in organic vs. conventional farming systems are vastly different due to their distinctive
management practices.
[10]
Most notably, organic farmers have fewer inputs available than
conventional growers to control their production environments. Breeding varieties specifically
adapted to the unique conditions of organic agriculture is critical for this sector to realize its
full potential. This requires selection for traits such as:
[10]

Water use efficiency
Nutrient use efficiency (particularly nitrogen and phosphorus)
Weed competitiveness
Tolerance of mechanical weed control
Pest/disease resistance
Early maturity (as a mechanism for avoidance of particular stresses)
Abiotic stress tolerance (i.e. drought, salinity, etc...)
Currently, few breeding programs are directed at organic agriculture and until recently those
that did address this sector have generally relied on indirect selection (i.e. selection in
conventional environments for traits considered important for organic agriculture). However,
because the difference between organic and conventional environments is large, a
given genotype may perform very differently in each environment due to an interaction
between genes and the environment (see gene-environment interaction). If this interaction is
severe enough, an important trait required for the organic environment may not be revealed
in the conventional environment, which can result in the selection of poorly adapted
individuals.
[9]
To ensure the most adapted varieties are identified, advocates of organic
breeding now promote the use of direct selection (i.e. selection in the target environment) for
many agronomic traits.
There are many classical and modern breeding techniques that can be utilized for crop
improvement in organic agriculture despite the ban on genetically modified organisms. For
instance, controlled crosses between individuals allow desirable genetic variation to be
recombined and transferred to seed progeny via natural processes. Marker assisted
selection can also be employed as a diagnostics tool to facilitate selection of progeny who
possess the desired trait(s), greatly speeding up the breeding process.
[11]
This technique has
proven particularly useful for the introgression of resistance genes into new backgrounds, as
well as the efficient selection of many resistance genes pyramided into a single individual.
Unfortunately, molecular markers are not currently available for many important traits,
especially complex ones controlled by many genes.
Addressing global food security through plant breeding[edit]
For future agriculture to thrive there are necessary changes which must be made in
accordance to arising global issues. These issues are arable land, harsh cropping conditions
and food security which involves, being able to provide the world population with food
containing sufficient nutrients. These crops need to be able to mature in several
environments allowing for worldwide access, this is involves issues such as drought
tolerance. These global issues are achievable through the process of plant breeding, as it
offers the ability to select specific genes allowing the crop to perform at a level which yields
the desired results.
Minimal land degradation[edit]
Land degradation is a major issue, as it can negatively impact the capability of the land to be
productive. Poor agricultural management has a huge impact on the degradation of soil
worldwide and it is Africa and Asia that are most affected. Through education and
development of modified plants, these statistics can be reduced and agricultural land can
become more productive. Plant breeding allows for an increase in yield with out the extra
strain on the land. The genetically modified, Bt white maize, was introduced to South Africa
and was surveyed in 33 large commercial farms and 368 small landholders properties and in
both cases a higher yield was recorded.
[12]

Increased yield without expansion[edit]
With an increasing population the production of food needs to increase with it, it is estimated
that a 70% increase in food production is needed by 2050 in order to meet the Declaration of
the World Summit on Food Security. But with the natural degradation of agricultural land,
simply planting more crops is no longer a viable option. There for new varieties of plants
need to be developed through plant breeding that generates an increase of yield without
relying on an increase in land area. An example of this can be seen in Asia, where food
production per capita has increased twofold, has been achieved through not only the use of
fertilisers but through the use of better crops that have been specifically designed for the
area.
[13][14]

Breeding for increased nutritional value[edit]
Plant breeding can contribute to global food security as it is a cost-effective tool for
increasing nutritional value of forage and crops. Improvements in nutritional value for forage
crops from the use of analytical chemistry and rumen fermentation technology have been
recorded since 1960; this science and technology gave breeders the ability to screen
thousands of samples within a small amount of time, meaning breeders could identify a high
performing hybrid quicker. The main area genetic increases were made was in vitro dry
matter digestibility (IVDMD) resulting in 0.7-2.5% increase, at just 1% increase in IVDMD a
single Bos Taurus also known as beef cattle reported 3.2% increase in daily gains. This
improvement indicates plant breeding is an essential tool in gearing future agriculture to
perform at a more advanced level.
[15]

Breeding for tolerance[edit]
Plant breeding of hybrid crops has become extremely popular worldwide in an effort to
combat the harsh environment. With long periods of drought and lack of water or nitrogen
stress tolerance has become a significant part of agriculture. Plant breeders have focused
on identifying crops which will ensure crops perform under these conditions; a way to
achieve this is finding strains of the crop that is resistance to drought conditions with low
nitrogen. It is evident from this that plant breeding is vital for future agriculture to survive as it
enables farmers to produce stress resistant crops hence improving food security.
[16]

Participatory plant breeding[edit]
The development of agricultural science, with phenomenon like the Green
Revolution arising, have left millions of farmers in developing countries, most of whom
operate small farms under unstable and difficult growing conditions, in a precarious situation.
The adoption of new plant varieties by this group has been hampered by the constraints of
poverty and the international policies promoting an industrialized model of agriculture. Their
response has been the creation of a novel and promising set of research methods
collectively known as participatory plant breeding. Participatory means that farmers are more
involved in the breeding process and breeding goals are defined by farmers instead of
international seed companies with their large-scale breeding programs. Farmers' groups and
NGOs,
[17]
for example, may wish to affirm local people rights over genetic resources,
produce seeds themselves, build farmers' technical expertise, or develop new products for
niche markets, like organically grown food.
[18][19][20]















For related content, see genetic engineering, genetically modified organism, genetically
modified food, and genetically modified food controversies.
Genetically modified crops (GMCs, GM crops, or biotech crops) are plants used in agriculture,
the DNA of which has been modified using genetic engineering techniques. In most cases the aim is to
introduce a new trait to the plant which does not occur naturally in the species. Examples in food crops
include resistance to certain pests, diseases, or environmental conditions, reduction of spoilage, or
resistance to chemical treatments (e.g. resistance to a herbicide), or improving the nutrient profile of the
crop. Examples in non-food crops include production ofpharmaceutical agents, biofuels, and other
industrially useful goods, as well as for bioremediation.
Farmers have widely adopted GM technology. Between 1996 and 2011, the total surface area of land
cultivated with GM crops had increased by a factor of 94, from 17,000 square kilometers (4,200,000
acres) to 1,600,000 km
2
(395 million acres). 10% of the world's crop lands were planted with GM crops in
2010. As of 2011, 11 different transgenic crops were grown commercially on 395 million acres (160
million hectares) in 29 countries.
There is broad scientific consensus that food on the market derived from GM crops poses no greater risk
to human health than conventional food.
[1][2][3][4][5]
GM crops also provide a number of ecological
benefits.
[6]
However, opponents have objected to GM crops per se on several grounds, including
environmental concerns, whether food produced from GM crops is safe, whether GM crops are needed to
address the world's food needs, and economic concerns raised by the fact these organisms are subject to
intellectual property law.
Contents
[show]
Gene transfer in nature and traditional agriculture[edit]
Scientists first discovered that DNA naturally transfers between organisms in 1946.
[7]
It is now known that
there are several natural mechanisms for flow of genes, or (horizontal gene transfer), and that these
occur in nature on a large scale for example, it is a major mechanism for antibiotic resistance in
pathogenic bacteria, and it occurs between plant species.
[8]
This is facilitated
bytransposons, retrotransposons, proviruses and other mobile genetic elements that naturally translocate
to new sites in a genome.
[9][10]
They often move to new species over an evolutionary time scale
[11]
and
play a major role in dynamic changes to chromosomes during evolution.
[12][13]

The introduction of foreign germplasm into crops has been achieved by traditional crop breeders by
artificially overcoming fertility barriers. A hybrid cereal was created in 1875, by
crossing wheatand rye.
[14]
Since then important traits have been introduced into wheat, including dwarfing
genes and rust resistance.
[15]
Plant tissue culture and the induction of mutations have also enabled
humans to artificially alter the makeup of plant genomes.
[16][17]

History[edit]
Main article: History of genetic engineering
The first genetically modified plant was produced in 1982, using an antibiotic-resistant tobacco
plant.
[18]
The first field trials of genetically engineered plants occurred in France and the USA in 1986,
when tobacco plants were engineered to be resistant to herbicides.
[19]
In 1987, Plant Genetic
Systems (Ghent, Belgium), founded by Marc Van Montagu and Jeff Schell, was the first company to
develop genetically engineered (tobacco) plants with insect tolerance by expressing genes encoding for
insecticidal proteins from Bacillus thuringiensis (Bt).
[20]
The Peoples Republic of China was the first
country to allow commercialized transgenic plants, introducing a virus-resistant tobacco in 1992,
[21]
which
was withdrawn from the market in China in 1997.
[22]:3
The first genetically modified crop approved for sale
in the U.S., in 1994, was the FlavrSavr tomato, which had a longer shelf life.
[23]
In 1994, the European
Union approved tobacco engineered to be resistant to the herbicide bromoxynil, making it the first
commercially genetically engineered crop marketed in Europe.
[24]
In 1995, Bt Potato was approved safe
by the Environmental Protection Agency, making it the first pesticide producing crop to be approved in the
USA.
[25]
The following transgenic crops also received marketing approval in the US in 1995: canola with
modified oil composition (Calgene), Bacillus thuringiensis (Bt) corn/maize (Ciba-Geigy), cotton resistant to
the herbicide bromoxynil (Calgene), Bt cotton (Monsanto), soybeans resistant to the
herbicideglyphosate (Monsanto), virus-resistant squash (Asgrow), and additional delayed ripening
tomatoes (DNAP, Zeneca/Peto, and Monsanto).
[19]
As of mid-1996, a total of 35 approvals had been
granted to commercially grow 8 transgenic crops and one flower crop of carnations, with 8 different traits
in 6 countries plus the EU.
[19]
In 2000, with the production of golden rice, scientists genetically modified
food to increase its nutrient value for the first time.
Methods[edit]
Main article: Techniques of genetic engineering


Plants (Solanum chacoense) being transformed using agrobacterium
Genetically engineered plants are generated in a laboratory by altering their genetic makeup. This is
usually done by adding one or more genes to a plant's genome using genetic
engineering techniques.
[26]
Most genetically modified plants are generated by the biolistic method (particle
gun) or byAgrobacterium tumefaciens mediated transformation. Plant scientists, backed by results of
modern comprehensive profiling of crop composition, point out that crops modified using GM techniques
are less likely to have unintended changes than are conventionally bred crops.
[27][28]

In research tobacco and Arabidopsis thaliana are the most genetically modified plants, due to well
developed transformation methods, easy propagation and well studied genomes.
[29][30]
They serve as
model organisms for other plant species.
In the biolistic method, DNA is bound to tiny particles of gold or tungsten which are subsequently shot into
plant tissue or single plant cells under high pressure. The accelerated particles penetrate both the cell
wall and membranes. The DNA separates from the metal and is integrated into plant genomeinside
the nucleus. This method has been applied successfully for many cultivated crops,
especially monocots like wheat or maize, for whichtransformation using Agrobacterium tumefaciens has
been less successful.
[31]
The major disadvantage of this procedure is that serious damage can be done to
the cellular tissue.
Agrobacteria are natural plant parasites, and their natural ability to transfer genes provides another
method for the development of genetically engineered plants. To create a suitable environment for
themselves, these Agrobacteria insert their genes into plant hosts, resulting in a proliferation of plant cells
near the soil level (crown gall). The genetic information for tumour growth is encoded on a mobile, circular
DNA fragment (plasmid). When Agrobacterium infects a plant, it transfers thisT-DNA to a random site in
the plant genome. When used in genetic engineering the bacterial T-DNA is removed from the bacterial
plasmid and replaced with the desired foreign gene. The bacterium is a vector, enabling transportation of
foreign genes into plants. This method works especially well for dicotyledonous plants like potatoes,
tomatoes, and tobacco. Agrobacteria infection is less successful in crops like wheat and maize.
Introducing new genes into plants requires a promoter specific to the area where the gene is to be
expressed. For instance, if we want the gene to be expressed only in rice grains and not in leaves, then
an endosperm-specific promoter would be used. The codons of the gene must also be optimized for the
organism due to codon usage bias. The transgenic gene products should also be able to be denatured by
heat so that they are destroyed during cooking.
Glyphosate resistance[edit]
One of the most famous kinds of GM crops are "Roundup Ready", or glyphosate-resistant trait.
Glyphosate, (the active ingredient in Roundup) kills plants by interfering with the shikimate pathway in
plants, which is essential for the synthesis of the aromatic amino
acids phenylalanine, tyrosine and tryptophan. More specifically, glyphosate inhibits the enzyme 5-
enolpyruvylshikimate-3-phosphate synthase (EPSPS).
The rationale behind developing this trait, was that the selective herbicides used on grain and grass crops
at the time were highly toxic, and not effective against narrow leaved weeds. Thus, developing crops that
could withstand spraying with glyphosate would both reduce environmental and health risks, and give an
agricultural edge to the farmer.
[32]

The shikimate pathway is not present in animals, which instead obtain aromatic amino acids from their
diet.
Some micro-organisms have a version of EPSPS that is resistant to glyphosate inhibition. One of these
was isolated from an Agrobacterium strain CP4 (CP4 EPSPS) that was resistant to glyphosate.
[33][34]
This
CP4 EPSPS gene was cloned and transfected into soybeans. The CP4 EPSPS gene was engineered for
plant expression by fusing the 5' end of the gene to a chloroplasttransit peptide derived from
the petunia EPSPS. This transit peptide was used because it had shown previously an ability to deliver
bacterial EPSPS to the chloroplasts of other plants. Theplasmid used to move the gene into soybeans
was PV-GMGTO4. It contained three bacterial genes, two CP4 EPSPS genes, and a
gene encoding beta-glucuronidase (GUS) from Escherichia coli as a marker. The DNA was injected into
the soybeans using the particle acceleration method. Soybean cultivar A54O3 was used for
the transformation. The expression of the GUS gene was used as the initial evidence of transformation.
GUS expression was detected by a staining method in which the GUS enzyme converts a substrate into a
blue precipitate. Those plants that showed GUS expression were then taken and sprayed with
glyphosate, and their tolerance was tested over many generations.
Types of genetic engineering[edit]


Transgenic maize containing a gene from the bacteria Bacillus thuringiensis
Transgenic plants have genes inserted into them that are derived from another species. The inserted
genes can come from species within the samekingdom (plant to plant) or between kingdoms (for
example, bacteria to plant). In many cases the inserted DNA has to be modified slightly in order to
correctly and efficiently express in the host organism. Transgenic plants are used to express proteins like
the cry toxins from Bacillus thuringiensis,herbicide resistant genes
and antigens for vaccinations
[35]
Transgenic carrots have been used to produce the drug Taliglucerase
alfa which is used to treat Gaucher's disease.
[36]
In the laboratory, transgenic plants have been modified
to increase their photosynthesis (currently about 2% at most plants to the theoretic potential of 9
10%.
[37]
This is possible by changing the rubisco enzyme (i.e. changing C3 plants into C4 plants
[38]
), by
placing therubisco in a carboxysome, by adding CO
2
pumps in the cell wall,
[39][40]
by changing the leaf
form/size.
[41][42][43][44]
Plants have been engineered to exhibit bioluminescence which might one day be a
sustainable alternative to electric lighting.
[45]
Still other transgenic plants have been modified to fix
ambient nitrogen in the plant.
[46]

Cisgenic plants are made using genes found within the same species or a closely related one, where
conventional plant breeding can occur. Some breeders and scientists argue that cisgenic modification is
useful for plants that are difficult to crossbreed by conventional means (such as potatoes), and that plants
in the cisgenic category should not require the same level of legal regulation as other genetically modified
organisms.
[47]

Business of GM Crops[edit]
The global value of biotech seed alone was US$13.2 billion in 2011, with the end product of commercial
grain from biotech maize, soybean grain and cotton valued at approximately US$160 billion or more per
year.
[48]

Players in agriculture business markets include seed companies, agrochemical companies, distributors,
farmers, grain elevators, and universities that develop new crops and whose agricultural extensions
advise farmers on best practices.
The largest share of the GMO crops planted globally are from seed created by the United
States firm Monsanto.
[49]
In 2007, Monsanto's trait technologies were planted on 246 million acres
(1,000,000 km
2
) throughout the world, a growth of 13 percent from 2006. However, patents on the first
Monsanto products to enter the marketplace will begin to expire in 2014, democratizing Monsanto
products. Syngenta, DuPont (especially via its Pioneer Hi-Bred subsidiary), and Bayer CropScience are
also major players in the US and Europe. In addition, a 2007 report from the European Joint Research
Commission predicts that by 2015, more than 40 per cent of new GM plants entering the global
marketplace will have been developed in Asia.
[50]

In the corn market, Monsanto's triple-stack cornwhich combines Roundup Ready 2-weed control
technology with YieldGard (Bt) Corn Borer and YieldGard Rootworm insect controlis the market leader
in the United States. U.S. corn farmers planted more than 32 million acres (130,000 km
2
) of triple-stack
corn in 2008,
[51]
and it is estimated the product could be planted on 56 million acres (230,000 km
2
) in
20142015. In the cotton market, Bollgard II with Roundup Ready Flex was planted on approximately
5 million acres (20,000 km
2
) of U.S. cotton in 2008.
[52]

According to the International Service for the Acquisition of Agri-Biotech Applications (ISAAA), in 2010
approximately 15 million farmers grew biotech crops in 29 countries. Over 90% of the farmers were
resource-poor in developing countries.
[53]
6.5 million farmers in China and 6.3 million small farmers in
India grew biotech crops (mostly Bacillus thuringiensis cotton). The Philippines, South Africa (biotech
cotton, maize, and soybeans often grown by subsistence women farmers) and another twelve developing
countries also grew biotech crops in 2009.
[54]
10 million more small and resource-poor farmers may have
been secondary beneficiaries of Bt cotton in China.
According to a review published in 2012 and based on data from the late 1990s and early 2000s, much of
the GM crop grown each year is used for livestock feed, and increased demand for meat will lead to
increased demand for GM crops with which to feed them.
[55]
Feed grain usage as a percentage of total
crop production is 70% for corn and more than 90% of oil seed meals such as soybeans. About 65 million
metric tons of GM corn grains and about 70 million metric tons of soybean meals derived from GM
soybean are fed to livestock each year.
[55]

Uses, actual and proposed[edit]
GM crops grown today, or under experimental development, have been modified with traits intended to
provide benefit to farmers, consumers, or industry. These traits include improved shelf life,disease
resistance, stress resistance, herbicide resistance, pest resistance, production of useful goods such as
biofuel or drugs, and ability to absorb toxins, for use in bioremediation of pollution. Due to high regulatory
and research costs, the majority of genetically modified crops in agriculture consist of commodity crops,
such as soybean, maize, cotton and rapeseed.
[56][57]
Recently, some research and development has been
targeted to enhancement of crops that are locally important in developing countries, such as insect-
resistant cowpea for Africa
[58]
and insect-resistantbrinjal (eggplant) for India.
[59]

Improved shelf life[edit]
The first genetically modified crop approved for sale in the U.S. was the FlavrSavr tomato, which had a
longer shelf life.
[23]
It is no longer on the market. As of 2013, an apple that has been genetically modified
to resist browning, known as the Nonbrowning Arctic apple produced by Okanagan Specialty Fruits, is
awaiting regulatory approval in the US and Canada. A gene in the fruit has been modified such that the
apple produces less polyphenol oxidase, a chemical that manifests the browning.
[60]
If approved by U.S.
regulators in the coming months, they will be one of the first genetically engineered fruits on store shelves
in America.
[61]

Improved nutrition[edit]
Some GM soybeans on the market today offer improved oil profiles for processing or healthier edible
oils.
[62]

[63]
GM plants are being developed by both private companies and public research institutions such
as CIMMYT, the International Maize and Wheat Improvement Centre.
[64]
Other examples include a
genetically modified cassava with lower cyanogen glucosides and enhanced with protein and other
nutrients,
[65]
while golden rice, developed by the International Rice Research Institute (IRRI), has been
discussed as a possible cure for Vitamin A deficiency.
[66]
An international group of academics has
generated a vitamin-enriched corn derived from South African white corn variety M37W with 169x
increase in beta carotene, 6x the vitamin C and 2x folate it is not in production anywhere, but proves
that this can be done.
[67]
Camelina sativa has been modified in research labs to produce plants that
accumulate high levels of oils similar to fish oils.
[68][69]

Stress resistance[edit]
Plants engineered to tolerate non-biological stresses like drought,
[70][71]
frost,
[72][73]
high Soil
salinity,
[74][75]
and nitrogen starvation
[76]
or with increased nutritional value (e.g. Golden rice
[77]
) were in
development in 2011.
Herbicide resistance[edit]
One of the most prevalent type of GM crops have a "Roundup Ready", or glyphosate-resistant
trait.
[78]
Tobacco plants have been engineered to be resistant to the herbicide bromoxynil.
[24]
Crops have
been commercialized that are resistant to the herbicide glufosinate, as well.
[79]
As weeds have grown
resistant to glyphosate and other herbicides used in concert with resistant GM crops, companies are
developing crops engineered to become resistant to multiple herbicides to allow farmers to use a mixed
group of two, three, or four different chemicals.
[80]

Pathogen resistance insects or viruses[edit]
Tobacco, corn, rice and many other crops, have been generated that express genes encoding for
insecticidal proteins from Bacillus thuringiensis (Bt).
[25][81]
Papaya, potatoes, and squash have been
engineered to resist viral pathogens, such as cucumber mosaic virus which, despite its name, infects a
wide variety of plants.
[82]

Production of biofuels[edit]
Algae, both hybrid and GM, is under development by several companies for the production of
biofuels.
[83]
Jatropha has also been modified to improve its qualities for fuel product. Swiss-
basedSyngenta has received USDA approval to market a maize seed trademarked Enogen, which has
been genetically modified to convert its own starch to sugar to speed the process of making ethanol for
biofuel.
[84]
In 2013, the Flemish Institute for Biotechnology was investigating poplar trees genetically
engineered to contain less lignin so that they would be more suitable for conversion into biofuels.
[85]
Lignin
is the critical limiting factor when using wood to make bio-ethanol because lignin limits the accessibility
of cellulose microfibrils to depolymerization by enzymes.
[86]

Production of useful by-products[edit]
Drugs[edit]
In 2012, the FDA approved the first plant-produced pharmaceutical, a treatment for Gaucher's
Disease.
[87]
Bananas have been developed, but are not in production, that produce
human vaccinesagainst infectious diseases such as Hepatitis B.
[88]
Tobacco plants have been developed
and studied, but are not in production, that can produce therapeutic antibodies.
[89]

Materials[edit]
Several companies and labs are working on engineering plants that can be used to make
bioplastics.
[90]
Potatoes that produce more industrially useful starches have been developed as
well.
[91]
Additionally, oilseed can be modified to produce fatty acids for detergents,
substitute fuels and petrochemicals.
Bioremediation[edit]
Scientists at the University of York developed a weed (Arabidopsis thaliana) that contains genes from
bacteria that can clean up TNT and RDX-explosive contaminants from the soil: It was hoped that this
weed would eliminate this pollution.
[92]
16 million hectares in the USA (1.5% of the total surface) are
estimated to be contaminated with TNT and RDX. However the weed Arabidopsis thaliana was not tough
enough to withstand the environment on military test grounds and research is continuing with the
University of Washington to develop a tougher native grass.
[93]

Genetically modified plants have also been used for bioremediation of contaminated
soils. Mercury, selenium and organic pollutants such as polychlorinated biphenyls (PCBs), TNT
and RDXexplosive contaminants have been removed from soils by transgenic plants containing genes for
bacterial enzymes.
[93][94]

Marine environments are especially vulnerable since oil spills of coastal regions and the open sea are
poorly containable and mitigation is difficult. In addition to pollution through human activities, millions of
tons of petroleum enter the marine environment every year from natural seepages. Despite its toxicity, a
considerable fraction of petroleum oil entering marine systems is eliminated by the hydrocarbon-
degrading activities of microbial communities. Particularly successful is a recently discovered group of
specialists, the so-called hydrocarbonoclastic bacteria (HCCB).
[95]

Extent of worldwide use of GM crops[edit]
C
o
u
n
t
r
y
20
10

pl
an
te
d
ar
ea
(
mi
lli
o
n
he
ct
ar
es
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[9
6]

20
09

A
gr
ic
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tu
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ea
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mi
lli
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ct
ar
es
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[9
7]

P
er
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e
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wi
th
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M
cr
o
p
s
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i
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h

c
r
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A
66.8 403
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So
yb
ea
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aiz
e,
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tto
n,
Ca
no
la,
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ua
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o
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[9
6]

20
09

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[9
7]

P
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of
a
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wi
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G
M
cr
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h

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r
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p
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sh
,
Pa
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ya
,
Alf
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a,
Su
ga
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ee
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o
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r
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20
10

pl
an
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[9
6]

20
09

A
gr
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tu
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ea
(
mi
lli
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ct
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[9
7]

P
er
c
e
nt
a
g
e
of
a
gr
ic
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tu
re
ar
e
a
wi
th
G
M
cr
o
p
s
B
i
o
t
e
c
h

c
r
o
p
s
Br
azi
l
25.4 265
9.60
%
So
yb
ea
n,
M
aiz
e,
Co
tto
n
Ar
ge
nti
22.9 141
16.3
0%
So
yb
ea
C
o
u
n
t
r
y
20
10

pl
an
te
d
ar
ea
(
mi
lli
o
n
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ct
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es
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[9
6]

20
09

A
gr
ic
ul
tu
re
ar
ea
(
mi
lli
o
n
he
ct
ar
es
)
[9
7]

P
er
c
e
nt
a
g
e
of
a
gr
ic
ul
tu
re
ar
e
a
wi
th
G
M
cr
o
p
s
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i
o
t
e
c
h

c
r
o
p
s
na n,
M
aiz
e,
Co
tto
n
Ind
ia
9.4 180
5.22
%
Co
tto
n
Ca
na
8.8 68
13.0
M
aiz
C
o
u
n
t
r
y
20
10

pl
an
te
d
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ea
(
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lli
o
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[9
6]

20
09

A
gr
ic
ul
tu
re
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ea
(
mi
lli
o
n
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ct
ar
es
)
[9
7]

P
er
c
e
nt
a
g
e
of
a
gr
ic
ul
tu
re
ar
e
a
wi
th
G
M
cr
o
p
s
B
i
o
t
e
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h

c
r
o
p
s
da 2% e,
So
yb
ea
n,
Ca
no
la,
Su
ga
rb
ee
t
C
o
u
n
t
r
y
20
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pl
an
te
d
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ea
(
mi
lli
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ct
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es
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[9
6]

20
09

A
gr
ic
ul
tu
re
ar
ea
(
mi
lli
o
n
he
ct
ar
es
)
[9
7]

P
er
c
e
nt
a
g
e
of
a
gr
ic
ul
tu
re
ar
e
a
wi
th
G
M
cr
o
p
s
B
i
o
t
e
c
h

c
r
o
p
s
Re
st
of
the
wo
rld
14.7 3,883
0.38
%
---
-
In the United States, the United States Department of Agriculture (USDA) reports on the total area of
GMO varieties planted.
[98]
According to National Agricultural Statistics Service, the states published in
these tables represent 8186 percent of all corn planted area, 8890 percent of all soybean planted area,
and 8193 percent of all upland cotton planted area (depending on the year).
USDA does not collect data for global area. Estimates are produced by the International Service for the
Acquisition of Agri-biotech Applications (ISAAA) and can be found in the report, "Global Status of
Commercialized Transgenic Crops: 2007".
[99]

Farmers have widely adopted GM technology (see figure). Between 1996 and 2011, the total surface
area of land cultivated with GM crops had increased by a factor of 94, from 17,000 square kilometers
(4,200,000 acres) to 1,600,000 km
2
(395 million acres).
[48]
10% of the world's crop lands were planted
with GM crops in 2010.
[48]
As of 2011, 11 different transgenic crops were grown commercially on 395
million acres (160 million hectares) in 29 countries such as the USA, Brazil, Argentina, India, Canada,
China, Paraguay, Pakistan, South Africa, Uruguay, Bolivia, Australia, Philippines, Myanmar, Burkina
Faso, Mexico and Spain.
[48]
One of the key reasons for this widespread adoption is the perceived
economic benefit the technology brings to farmers. For example, the system of planting glyphosate-
resistant seed and then applying glyphosate once plants emerged provided farmers with the opportunity
to dramatically increase the yield from a given plot of land, since this allowed them to plant rows closer
together.
[100]
Without it, farmers had to plant rows far enough apart to control post-emergent weeds with
mechanical tillage.
[100]
Likewise, using Bt seeds means that farmers do not have to purchase insecticides,
and then invest time, fuel, and equipment in applying them. However critics have disputed whether yields
are higher and whether chemical use is less, with GM crops. See Genetically modified food
controversies article for information.


Land area used for genetically modified crops by country (19962009), in millions of hectares. In 2011,
the land area used was 160 million hectares, or 1.6 million square kilometers.
[48]

In the US, by 2009/10, 93% of the planted area of soybeans, 93% of cotton, 86% of corn and 95% of the
sugar beet were genetically modified varieties.
[101][102][103]
Genetically modified soybeans carried herbicide-
tolerant traits only, but maize and cotton carried both herbicide tolerance and insect protection traits (the
latter largely the Bacillus thuringiensis Bt insecticidal protein).
[104]
These constitute "input-traits" which are
aimed to financially benefit the producers, but may have indirect environmental benefits and marginal cost
benefits to consumers. The Grocery Manufacturers of America estimated in 2003 that 7075% of all
processed foods in the U.S. contained a GM ingredient.
[105]

Europe has relatively few genetically engineered crops
[106]
with the exception of Spain where one fifth of
maize grown is genetically engineered,
[107]
and smaller amounts in five other countries.
[108]
The EU had a
'de facto' ban on the approval of new GM crops, from 1999 until 2004;
[109]
in a controversial move.
[110]
GM
crops are now regulated by the EU.
[111]
Developing countries grew 50 percent of genetically engineered
crops in 2011.
[48]

In recent years there has been rapid growth in the area sown in developing countries. A total of 29
countries worldwide grew GM crops in 2011 by approximately 16.7 million farmers and 50% of GM crops
grown worldwide were grown in developing countries. For example, the largest increase in crop area
planted to GM crops in 2011 was in Brazil (303,000 km
2
versus 254,000 km
2
in 2010). There has also
been rapid and continuing expansion of GM cotton varieties in India since 2002 with 106,000 km
2
of GM
cotton harvested in India in 2011.
[48]
However the use of GM crops in India has been controversial, as
discussed in detail in theGM controversies article.
According to the 2011 ISAAA brief: "While 29 countries planted commercialized biotech crops in 2010, an
additional 31 countries, totaling 60 have granted regulatory approvals for biotech crops for import for food
and feed use and for release into the environment since 1996.... A total of 1,045 approvals have been
granted for 196 events (NB: an "event" is a specific genetic modification in a specific species) for 25
crops. Thus, biotech crops are accepted for import for food and feed use and for release into the
environment in 60 countries, including major food importing countries like Japan, which do not plant
biotech crops. Of the 60 countries that have granted approvals for biotech crops, USA tops the list
followed by Japan, Canada, Mexico, South Korea, Australia, the Philippines, New Zealand, the European
Union, and Taiwan. Maize has the most events approved (65) followed by cotton (39), canola (15), potato
and soybean (14 each). The event that has received regulatory approval in most countries is herbicide
tolerant soybean event GTS-40-3-2 with 25 approvals (EU=27 counted as 1 approval only), followed by
insect resistant maize MON810 with 23 approvals, herbicide tolerant maize NK603 with 22 approvals
each, and insect resistant cotton (MON1445) with 14 approvals worldwide."
[48]

Examples of genetically modified crops[edit]
Currently, there are a number of food species for which a genetically modified version is being
commercially grown (percent modified in the table below are mostly 2009/2010 data).
[102][103][112][113][114][115]

Cro
p
Properties
of the
genetically
modified
variety
Modifi
cation
[specify]

P
e
r
c
e
n
t

m
o
d
i
f
i
e
d

P
e
r
c
e
n
t

m
o
d
i
f
i
e
d

i
n

U
S
i
n

w
o
r
l
d
Alf
alfa
Resistance
to glyphosat
e or glufosin
ate herbicide
s
New
genes
added
/transf
erred
into
plant
geno
me.
P
l
a
n
t
e
d

i
n

t
h
e

U
S

f
r
o
m

2
0
0
5

2
0
0
7
;

2
0

0
7

2
0
1
0

c
o
u
r
t

i
n
j
u
n
c
t
i
o
n
;

2
0
1
1

d
e
r
e
g
u
l
a
t
e
d
Ca
nol
Resistance
to herbicides
New
genes
8
7
2
1
a/
Ra
pes
eed
(glyphosate
or
glufosinate),
see Roundu
p Ready
Canola high
laurate
canola,
[116]
O
leic acid
canola
[117]

added
/transf
erred
into
plant
geno
me
%

(
2
0
0
5

d
a
t
a
[
1
1
5
]
)
%
Cor
n,
fiel
d
(Ma
ize)
Resistance
to glyphosat
e or glufosin
ate herbicide
s. Insect
resistance
via
producing Bt
proteins,
some
previously
used as
pesticides in
organic crop
production.
Added
enzyme,
alpha
amylase,
that converts
starch into
sugar to
facilitate
ethanol
production.
[1
18]

New
genes,
some
from
the
bacteri
um Ba
cillus
thurin
giensi
s,
added
/transf
erred
into
plant
geno
me.
[119
]

8
6
%
[
1
0
2
]

2
6
%
Cot
ton
(co
tto
nse
ed
oil)
Kills
susceptible
insect pests
gene
for
one or
more
Bt
crystal
protei
ns
transf
erred
into
plant
geno
me
9
3
%
4
9
%
Pa
pay
a
(Ha
wai
ian)
Resistance
to
the papaya
ringspot
virus.
[120]

New
gene
added
/transf
erred
into
plant
geno
me
8
0
%

Pot
ato
(fo
od)
NewLeaf: Bt
resistance
against
Colorado
beetle and
resistance
againstPotat
o virus
Y (removed
from market
in 2001
[91]
)
New
Leaf:
Bt
cry3A,
coat
protei
n from
PVY
[12
1]

0
%
0
%
Pot
ato
(sta
rch
)
Amflora:
resistance
gene against
an antibiotic,
used for
selection, in
combination
with
modification
Amflor
a
antibio
tic
resista
nce
gene
from
bacteri
0
%
0
%
s for better
starch
production
[12
2]

a;
modifi
cation
s to
endog
enous
starch
-
produ
cing
enzym
es
Ric
e
Golden
Rice:
genetically
modified to
contain beta
-carotene (a
source
ofvitamin A)
Curre
nt
versio
n of
Golde
n Rice
under
develo
pment
contai
ns
genes
from
maize
and a
comm
on soil
microo
rganis
m.
[123]
Previo
us
protot
ype
versio
n
contai
ned
three
new
genes:
two
F
o
r
e
c
a
s
t

t
o

b
e

o
n

t
h
e

m
a
r
k
e
t

i
n


from d
affodil
s and
the
third
from
a bact
erium
2
0
1
4

o
r

2
0
1
5
[
1
2
4
]

So
ybe
ans
Resistance
to glyphosat
e (see Roun
dup Ready
soybean)
or glufosinat
eherbicides;
make less
saturated
fats;
[125]
Kills
susceptible
insect pests
Herbic
ide
resista
nt
gene
taken
from
bacteri
a
inserte
d into
soybe
an;
knock
ed out
native
genes
that
cataly
ze
satura
tion;
gene
for
one or
more
9
3
%
7
7
%
Bt
crystal
protei
ns
transf
erred
into
plant
geno
me
Sq
uas
h
(Zu
cch
ini/
Co
urg
ette
)
Resistance
to
watermelon,
cucumber
and
zucchini/cou
rgette yellow
mosaic
viruses
[117][12
6][127]

Contai
ns
coat
protei
n
genes
of
viruse
s.
1
3
%

(
f
i
g
u
r
e

i
s

f
r
o
m

2
0
0
5
)
[
1
1
5
]


Su
gar
Resistance
to
New
genes
9
5
9
%
bee
t
glyphosate,
glufosinate
herbicides
added
/transf
erred
into
plant
geno
me
%

(
2
0
1
0
)
;

r
e
g
u
l
a
t
e
d

2
0
1
1
;

d
e
r
e
g
u
l
a
t
e
d

2
0
1
2
Su
gar
Resistance
to certain
New
genes
can
e
pesticides,
high sucrose
content.
added
/transf
erred
into
plant
geno
me
Sw
eet
pep
per
s
Resistance
to cucumber
mosaic
virus
[128][129]

Contai
ns
coat
protei
n
genes
of the
virus.

S
m
a
l
l

q
u
a
n
t
i
t
i
e
s

g
r
o
w
n

i
n

C
h
i
n
a
To
mat
oes
Suppression
of the
enzyme poly
galacturonas
e (PG),
A
revers
e copy
(an an
tisens
T
a
k
e
n
S
m
a
l
l
retarding
fruit
softening
after
harvesting,
[1
30]
while at
the same
time
retaining
both the
natural color
and flavor of
the fruit
e gen
e) of
the
gene
respo
nsible
for the
produ
ction
of PG
enzym
e
added
into
plant
geno
me

o
f
f

t
h
e

m
a
r
k
e
t

d
u
e

t
o

c
o
m
m
e
r
c
i
a
l

f
a
i
l
u
r
e
.

q
u
a
n
t
i
t
i
e
s

g
r
o
w
n

i
n

C
h
i
n
a
Wh
eat
Resistance
to
New
genes
u
n
u
n
glyphosate
herbicide
added
/transf
erred
into
plant
geno
me
k
n
o
w
n
k
n
o
w
n
Effects on farming practices[edit]

This section
requires expansion with:
examples and additional
citations of how
farmers's use of GM
crops changes their
practices. (September
2012)
Managing emergence of resistance[edit]
Constant exposure to a toxin creates evolutionary pressure for pests resistant to that toxin.
One method of reducing resistance is the creation of non-Bt crop refuges to allow some nonresistant
insects to survive and maintain a susceptible population. To reduce the chance an insect would become
resistant to a Bt crop, the commercialization of transgenic cotton and maize in 1996 was accompanied
with a management strategy to prevent insects from becoming resistant to Bt crops, and insect resistance
management plans are mandatory for Bt crops planted in the USA and other countries. The aim is to
encourage a large population of pests so that any resistance genes that are recessive are greatly diluted
within the population.
[131]

This means that with sufficiently high levels of transgene expression, nearly all of the heterozygotes (S/s),
i.e., the largest segment of the pest population carrying a resistance allele, will be killed before they reach
maturity, thus preventing transmission of the resistance gene to their progeny.
[132]
The planting of refuges
(i. e., fields of nontransgenic plants) adjacent to fields of transgenic plants increases the likelihood that
homozygous resistant (s/s) individuals and any surviving heterozygotes will mate with susceptible (S/S)
individuals from the refuge, instead of with other individuals carrying the resistance allele. As a result, the
resistance gene frequency in the population would remain low.
Nevertheless, limitations can affect the success of the high-dose/refuge strategy. For example,
expression of the Bt gene can vary. For instance, if the temperature is not ideal, this stress can lower the
toxin production and make the plant more susceptible. More importantly, reduced late-season expression
of toxin has been documented, possibly resulting from DNA methylation of thepromoter.
[133]
So, while the
high-dose/refuge strategy has been successful at prolonging the durability of Bt crops, this success has
also had much to do with key factors independent of management strategy, including low initial resistance
allele frequencies, fitness costs associated with resistance, and the abundance of non-Bt host plants that
have supplemented the refuges planted as part of the resistance management strategy.
[134]

Companies that produce Bt seed are addressing this as well, by introducing plants with multiple Bt
proteins. Monsanto did this with Bt cotton in India, where the product was rapidly adopted.
[135]

Regulation[edit]
Main articles: Regulation of genetic engineering and Regulation of the release of genetic
modified organisms
The regulation of genetic engineering concerns the approaches taken by governments to assess and
manage the risks associated with the development and release of genetically modified crops. There are
differences in the regulation of GM crops between countries, with some of the most marked differences
occurring between the USA and Europe. Regulation varies in a given country depending on the intended
use of the products of the genetic engineering. For example, a crop not intended for food use is generally
not reviewed by authorities responsible for food safety.
[136][137]

Controversy[edit]
Main article: Genetically modified food controversies
The genetically modified foods controversy is a dispute over the use of food and other goods derived
from genetically modified crops instead of from conventional crops, and other uses ofgenetic
engineering in food production. The dispute involves consumers, biotechnology companies, governmental
regulators, non-governmental organizations, and scientists. The key areas of controversy related to
genetically modified food are: whether GM food should be labeled, the role of government regulators, the
effect of GM crops on health and the environment, the effect on pesticide resistance, the impact of GM
crops for farmers, and the role of GM crops in feeding the world population.
There is broad scientific consensus that food on the market derived from GM crops poses no greater risk
than conventional food.
[1][3][138]
No reports of ill effects have been documented in the human population
from GM food.
[4][139][140]
Although labeling of genetically modified organism (GMO) products in the
marketplace is required in many countries, in the United States, the Food and Drug Administration does
not require labeling of GMO products in the marketplace, nor does it recognize a distinction between
GMO and non-GMO foods.
[141]

Some advocacy groups such as Greenpeace and the World Wildlife Fund have concerns that risks of GM
food have not been adequately identified and managed, and have questioned the objectivity of regulatory
authorities. Other environmental groups, including The Nature Conservancy
[142]
and former anti-GMO
campaigner Mark Lynas support the use of GMOs as beneficial for the environment.
[143]








A large number of farmers in India depend on animal husbandry for their livelihood. In addition to
supplying milk, meat, eggs,wool and hides, animals, mainly bullocks, are the major source of power for
both farmers and drayers. Thus, animal husbandry plays an important role in the rural economy. The
gross value of output from this sector was 358 billion (US$5.7 billion) in FY 1989, an amount that
constituted about 25 percent of the total agricultural output of 1.4 trillion (US$22.4 billion).
[1]

Contents
[show]
Production[edit]


Capra aegagrus hircus goatherd inAndhra Pradesh
In FY 1992, India had approximately 25 percent of the world's cattle, with a collective herd of 193 million
head. India also had 110 million goats, 75 million water buffalo, 44 million sheep, and 10 million pigs. Milk
production in FY 1990 was estimated to have reached 53.5 million tons, and eggproduction had reached
a level of 23.3 billion eggs. Dairy farming provided supplementary employment and an additional source
of income to many small and marginal farmers. The National Dairy Development Board was established
in 1965 under the auspices of Operation Flood at Anand, inGujarat, to promote, plan, and organize dairy
development through cooperatives; to provide consultations; and to set up dairy plants, which were then
turned over to the cooperatives. There were more than 63,000 Anand-style dairy cooperative societies
with some 7.5 million members in the early 1990s. The milk produced and sold by these farmers
brought 320 million (US$5.1 million) a day, or more than 10 trillion (US$160.0 billion) a year. The
increase in milk production permitted India to end imports of powdered milk and milk-related products. In
addition, 30,000 tons of powdered milk were exported annually to neighboring countries. There was about
50000 cows in India.
[2]



Battery cage facilities in Haryana
Today, India has the world's largest dairy herd (composed of cows and buffaloes), at over 304 million
strong,
[3]
and stands first in milk production, with 112.5 million tonnes of milk produced in 2009-
2010.
[3]
India is also the third largest egg-producer in the world, at over 180 million eggs being produced
every day or 65.7 billion eggs for the year 2011-12,
[4]
and the worlds sixth largest producer of poultry
meat.
[5]
While the majority of Indias animal products are consumed domestically, exports are growing.
India is the top global exporter of buffalo meat, and is also the fourth largest exporter of soybean meal, an
important ingredient in commercial feed for farmed animals.
[5]
In addition, Indias leading poultry
producers, including Suguna, Venkys, and the Amrit Group, are increasing sales to countries in other
parts of Asia and the Middle East. International investment is also expanding.
[6]
In 2008, U.S.-based
Tyson Foods acquired a 51 percent stake in Godrej, an Indian conglomerate that is a major producer of
animal feeds and poultry, and the International Finance Corporation (IFC) has an equity stake in
Suguna.
[5]
An estimated 200 million egg-laying hens are kept in battery cages, and more than half of the
2-billion-plus meat chickens produced each year are factory-farmed. Vertical integration and contract
farming for poultry and eggs is also widespread. While the milk and cheese sectors still include many
small-scale producers and cows and buffaloes in extensive systems the number of large, factory-style
dairy operations is increasing.
[7]

Operation Flood[edit]
Operation Flood, the world's largest integrated dairy development program, attempted to establish
linkages between rural milk producers and urban consumers by organizing farmer-owned and -managed
dairy cooperative societies. In the early 1990s, the program was in its third phase and was receiving
financial assistance from the World Bank and commodity assistance from the European Economic
Community. At that time, India had more than 64,000 dairy cooperative societies, with close to 7.7 million
members. These cooperatives established a daily processing capacity of 15.5 million liters of whole milk
and 727 tons of milk powder.
[8]



Fishing in India is a major industry in its coastal states, employing over 14 million people.
Fish production in India has increased more than tenfold since its independence in 1947. According to
the Food and Agriculture Organization (FAO) of the United Nations, fish output in India doubled between
1990 and 2010.
[1]

India has 8,118 kilometers of marine coastline, 3,827 fishing villages, and 1,914 traditional fish landing
centers. India's fresh water resources consist of 195,210 kilometers of rivers and canals, 2.9 million
hectares of minor and major reservoirs, 2.4 million hectares of ponds and lakes, and about 0.8 million
hectares of flood plain wetlands and water bodies.
[2]
As of 2010, the marine and freshwater resources
offered a combined sustainable catch fishing potential of over 4 million metric tonnes of fish. In addition,
India's water and natural resources offer a tenfold growth potential in aquaculture (farm fishing) from 2010
harvest levels of 3.9 million metric tonnes of fish, if India were to adopt fishing knowledge, regulatory
reforms, and sustainabilitypolicies adopted by China over the last two decades.
The marine fish harvested in India consist of about 65 commercially important species/groups. Pelagic
and midwater species contributed about 52% of the total marine fish in 2004.
India is a major supplier of fish in the world. In 2006 the country exported over 600,000 metric tonnes of
fish, to some 90 countries, earning over $1.8 billion.
[3]
Shrimps are one of the major varieties exported.
The giant tiger prawn (Penaeus monodon) is the dominant species chosen for aquaculture, followed by
the Indian white prawn (Fenneropenaeus indicus). Shrimp production from coastal aquaculture during
2004 stood at approximately 120,000 tonnes. Farmed shrimp accounted for about 60% of shrimp
exported from the country.
Marine and freshwater catch fishing combined with aquaculture fish farming is a rapidly growing industry
in India. In 2008 India was the sixth largest producer of marine and freshwater capture fisheries, and the
second largest aquaculture farmed fish producer in the world.
[4]
Fish as foodboth from fish farms and
catch fisheriesoffers India one of the easiest and fastest way to address malnutrition and food security.
Despite rapid growth in total fish production, a fish farmers average annual production in India is only 2
metric tonnes per person, compared to 172 tonnes in Norway, 72 tonnes in Chile, and 6 tonnes per
fisherman in China.
[4]
Higher productivity, knowledge transfer for sustainable fishing, continued growth in
fish production with increase in fish exports have the potential for increasing the living standards of Indian
fishermen.
As of 2010, fish harvest distribution was difficult within India because of poor rural road infrastructure, lack
of cold storage and absence of organized retail in most parts of the country.
In 2013, with access to Sri Lankan waters closed after the defeat of the Liberation Tigers of Tamil Eelam,
Indian fishers quickly overfished their own waters and production plummeted.
[5]

Contents
[show]
History[edit]


A fisherman in the backwaters ofKerala.
Fishing and aquaculture in India has a long history. Kautilya's Arthashastra (321300 B.C.) and King
Someswara's Manasottara (1127 A.D.) each refer to fish culture.
[6]
For centuries, India has had a
traditional practice of fish culture in small ponds in Eastern India. Significant advances in productivity
were made in the state of West Bengal in the early nineteenth century with the controlled breeding of carp
in Bundhs (tanks or impoundments where river conditions are simulated). Fish culture received notable
attention in Tamil Nadu (formerly the state of Madras) as early as 1911, subsequently, states such as
West Bengal, Punjab, Uttar Pradesh, Gujarat, Karnataka and Andhra Pradesh initiated fish culture
through the establishment of Fisheries Departments. In 2006, Indian central government initiated a
dedicated organization focussed on fisheries, under its Ministry of Agriculture.
Brackishwater farming in India is also an age-old system confined mainly to the Bheries (manmade
impoundments in coastal wetlands) of West Bengal and pokkali (salt resistant deepwater paddy) fields
along the Kerala coast. With no additional knowledge and technology input, except that of trapping the
naturally bred juvenile fish and shrimp seed, these systems have been sustaining production levels of
between 500 and 750 kg/ha/year with shrimp contributing 20 to 25 percent of the total Indian production.
[6]

Growth[edit]


Fishing boats on Palk Strait in Tamil Nadu


Fish boats of Tamil Nadu
It rose from only 800,000 tons in FY 1950 to 4.1 million tons in the early 1990s. From 1990 through 2010,
Indian fish industry growth has accelerated, reaching a total marine and freshwater fish production to
about 8 million metric tons. Special efforts have been made to promote extensive and intensive inland fish
farming, modernize coastal fisheries, and encourage deep-sea fishing through joint ventures. These
efforts led to a more than fourfold increase in coastal fish production from 520,000 tons in FY 1950 to
2.4 million tons in FY 1990. The increase in inland fish production was even more dramatic, increasing
almost eightfold from 218,000 tons in FY 1950 to 1.7 million tons in FY 1990. The value of fish and
processed fish exports increased from less than 1 percent of the total value of exports in FY 1960 to 3.6
percent in FY 1993.
Between 1990 to 2007, fish production in India has grown at a higher rate than food grains, milk, eggs,
and other food items.
[7]

Economic benefits[edit]
Fishing in India contributed over 1 percent of India's annual gross domestic product in 2008.
Catch fishing in India employs about 14.5 million people. The country's rich marine and inland water
resources, fisheries and aquaculture offer an attractive and promising sector for employment, livelihood,
and food security. Fish products from India are well received by almost half of world's countries, creating
export-driven employment opportunities in India and greater food security for the world. During the past
decades the Indian fisheries and aquaculture has witnessed improvements in craft, tackle and farming
methods. Creation of required harvest and post-harvest infrastructure has been receiving due attention of
the central and state governments. All this has been inducing a steady growth.
[2]

To harvest the economic benefits from fishing, India is adopting exclusive economic zone, stretching 200
nautical miles (370 km) into the Indian Ocean, encompasses more than 2 million square kilometers. In the
mid-1980s, only about 33 percent of that area was being exploited. The potential annual catch from the
area has been estimated at 4.5 million tons. In addition to this marine zone, India has about 14,000 km of
brackish water available for aquaculture, of which only 600 km were being farmed in the early 1990s;
about 16,000 km of freshwater lakes, ponds, and swamps; and nearly 64,000 kilometers
of rivers and streams.
In 1990, there were 1.7 million full-time fishermen, 1.3 million part-time fishermen, and 2.3 million
occasional fishermen, many of whom worked as saltmakers, ferrymen, or seamen, or operated boats for
hire. In the early 1990s, the fishing fleet consisted of 180,000 traditional craft powered by sails or oars,
26,000 motorized traditional craft, and some 34,000 mechanized boats.
Aquaculture[edit]


Carp - a commonly farmed fish in India
[8]

India laid the foundation for scientific carp farming in the country between 1970 and 1980, by
demonstrating high production levels of 8 to 10 tonnes/hectare/year in an incubation center. The late
1980s saw the dawn of aquaculture in India and transformed fish culture into a more modern enterprise.
With economic liberalization of early 1990s, fishing industry got a major investment boost.
India's breeding and culture technologies include primarily different species of carp; other species such
as catfish, murrels and prawns are recent additions.
The culture systems adopted in the country vary greatly depending on the input available in any particular
region as well as on the investment capabilities of the farmer. While extensive aquaculture is carried out
in comparatively large water bodies with stocking of the fish seed as the only input beyond utilising
natural productivity, elements of fertilisation and feeding have been introduced into semi-intensive culture.
The different culture systems in Indian practice include:
[6]

Intensive pond culture with supplementary feeding and aeration (1015 tonnes/ha/yr)
Composite carp culture (46 tonnes/ha/yr)
Weed-based carp polyculture (34 tonnes/ha/yr)
Integrated fish farming with poultry, pigs, ducks, horticulture, etc. (35 tonnes/ha/yr)
Pen culture (35 tonnes/ha/yr)
Cage culture (1015 kg/m/yr)
Running-water fish culture (2050 kg/m/yr)
Aquaculture resources in India include 2.36 million hectares of ponds and tanks, 1.07 million hectares
of beels, jheels and derelict waters plus in addition 0.12 million kilometers of canals, 3.15 million hectares
of reservoirs and 0.72 million hectares of upland lakes that could be utilised for aquaculture
purposes. Ponds and tanks are the prime resources for freshwater aquaculture in India. However, less
than 10 percent of India's natural potential is used for aquaculture currently.
The FAO of the United Nations estimates that about 1.2 million hectares of potential brackishwater area
available in India is suitable for farming, in addition to this, around 8.5 million hectares of salt affected
areas are also available, of which about 2.6 million hectares could be exclusively utilised for aquaculture
due to the unsuitability of these resources for other agriculture based activities. However, just like India's
fresh water resources, the total brackishwater area under cultivation is only just over 13 percent of the
potential water area available. India offers opportunities for highly productive farming of shrimp in its
brackishwater resources.
Carp hatcheries in both the public and private sectors have contributed towards the increase in seed
production from 6321 million fry in 19851986 to over 18500 million fry in 2007. There are 35 freshwater
prawn hatcheries in the coastal states producing over 200 million seed per annum. Furthermore, the 237
shrimp hatcheries with a production capacity of approximately 11.425 billion post larvae per year are
meeting the seed requirement of the brackish water shrimp farming sector.
[6]

Freshwater aquaculture activity is prominent in the eastern part of the country, particularly the states of
West Bengal, Orissa and Andhra Pradesh with new areas coming under culture in the states of Punjab,
Haryana, Assam and Tripura. Brackishwater aquaculture is mainly concentrated on the coasts of Andhra
Pradesh, Tamil Nadu, Orissa and West Bengal. With regards to the market, while the main areas of
consumption for freshwater fish are in West Bengal, Bihar, Orissa and northeast India, cultured
brackishwater shrimps supply India's fish export industry.
Rajiv Gandhi Center for Aquaculture(RGCA)[edit]
Rajiv Gandhi Centre for Aquaculture is the Research and Development arm of the Marine Products
Export Development Authority (MPEDA), which, inspired by the late Prime Minister Rajiv Gandhis vision
of making India a technologically advanced nation, founded this Centre of Excellence in Aquaculture and
dedicated it to the development of the Indian Aquaculture Industry. RGCA is actively involved in the
development of various Sustainable Aquaculture Technologies that are bio-secure, eco-friendly, traceable
and with low carbon outputs, for seed production and grow out farming of various aquatic species, those
having export potential in particular. RGCA is also developing a state-of-the-art technology transfer and
training centre for disseminating the technologies developed at the various projects established at
different locations in the country to the aquaculture industry in India.
Distribution of fish industry in Indian states[edit]


A fisherman in Kerala


A Boat(being manufactured) atBheemunipatnam Beach in Andhra Pradesh
Fishing is a diverse industry in India. The table below presents the top ten fish harvesting states in India,
for the 2007-2008 agriculture year.
Leading fish producing states in India, 20072008
Rank
[9]
State Total production (metric tonnes)
1 West Bengal 1,447,260
2 Andhra Pradesh 1,010,830
3 Gujarat 721,910
4 Kerala 667,330
5 Tamil Nadu 559,360
6 Maharashtra 556,450
7 Orissa 349,480
8 Uttar Pradesh 325,950
9 Bihar 319,100
10 Karnataka 297,690
Between 2000 and 2010, the freshwater prawn farming in India has grown rapidly. The state of Andhra
Pradesh dominates the sector with over 86 percent of the total production in India with approximately 60
percent of the total water area dedicated to prawn farming, followed by West Bengal. Mixed farming of
freshwater prawn along with carp is also very much accepted as a technologically sound culture practice
and a viable option for enhancing farm income. Thirty five freshwater prawn hatcheries, at present
producing about 200 million seed per annum, cater for the requirements of the country.
[6]

Law and regulations[edit]
India has a federal structure of government. According to India's constitution, the power of enacting laws
is split between India's central government and the Indian states. The state legislatures of India have the
power to make laws and regulations with respect to a number of subject-matters, including water
(i.e., water supplies, irrigation and canals, drainage and embankments, water storage and water power),
land (i.e., rights in or over land, land tenure, transfer, and alienation of agricultural land), fisheries, as well
as the preservation, protection and improvement of stock and the prevention of animal disease. There are
many laws and regulations that may be relevant to fisheries and aquaculture adopted at state level.
At the central level, several key laws and regulations are relevant to fisheries and aquaculture. These
include the British-era Indian Fisheries Act (1897), which penalizes the killing of fish by poisoning water
and by using explosives; the Environment (Protection) Act (1986), being an umbrella act containing
provisions for all environment related issues affecting fisheries and aquaculture industry in India. India
also has enacted the Water (Prevention and Control of Pollution) Act (1974) and the Wild Life Protection
Act (1972). All these legislations must be read in conjunction with one another, and with the local laws of
a specific state, to gain a full picture of the law and regulations that are applicable to fisheries and
aquaculture in India.
Research and training[edit]
Fisheries research and training institutions are supported by central and state governments that deserve
much of the credit for the expansion and improvements in the Indian fishing industry. The principal
fisheries research institutions, all of which operate under the Indian Council of Agricultural Research, are
the Central Marine Fisheries Research Institute at Kochi (formerly Cochin),Kerala; the Central Inland
Fisheries Institute at Barrackpore, West Bengal; and the Central Institute of Fisheries
Technology at Willingdon Island near Kochi. Most fishery training is provided by the Central Institute for
Fishery Education in Mumbai, which has ancillary institutions in Barrackpore, Agra (Uttar Pradesh),
and Hyderabad (Andhra Pradesh). The Central Fisheries Corporation inCalcutta is instrumental in
bringing about improvements in fishing methods, ice production, processing, storing, marketing, and
constructing and repairing fishing vessels. Operating under a 1972 law, the Marine Products Export
Development Authority(MPEDA), headquartered in Kochi, has made several market surveys abroad and
has been instrumental in introducing and enforcing hygiene standards that have gained for Indian fishery
export products a reputation for cleanliness and quality.
Programmes[edit]


Fishermen in Andhra Pradesh


Kochi, a fishing center in Kerala
The Government of India launched National Fisheries Development Board in 2006. Its headquarters are
in Hyderabad, located in a fish shaped building. Its activity focus areas are:
[10]

Intensive Aquaculture in Ponds and Tanks
Fisheries Development in Reservoirs.
Coastal Aquaculture
Mariculture
Seaweed Cultivation
Infrastructure: Fishing Harbours and Landing Centres
Fish Dressing Centres and Solar Drying of Fish
Domestic Marketing
Technology Upgradation
Deep Sea Fishing and Tuna Processing
The implementation of two programs for inland fisheriesestablishing fish farmers' development
agencies and the National Programme of Fish Seed Developmenthas led to encouragingly increased
production, which reached 1.5 million tons during FY 1990, up from 0.9 million tons in FY 1984. A network
of 313 fish farmers' development agencies was functioning in 1992. Under the National Programme of
Fish Seed Development, forty fish-seed hatcheries were commissioned. Fish-seed production doubled
from 5 billion fry in FY 1983 to 10 billion fry in FY 1989. A new program using organic waste for
aquaculture was started in FY 1986. Inland fish production as a percent of total fish production increased
from 36 percent in FY 1980 to 40 percent by FY 1990.
Major harbours[edit]


Kochi shipyard


A fishing dock in Maharashtra
Apart from four main fishing harbours--Mangalore (Karnataka), Kochi (Kerala), Chennai (Tamil
Nadu), Vishakhapatnam (Andhra Pradesh), andRoychowk in Kolkata (West Bengal)--twenty-three minor
fishing harbors and ninety-five fish-landing centers are designated to provide landing and berthing
facilities to fishing craft. The harbors at Vishakhapatnam, Kochi, and Roychowk were completed by 1980;
the one at Madras was completed in the 1980s. A major fishing harbor was under construction
at Sassoon Dock in Mumbai in the early 1990s, as were thirteen additional minor fishing harbors and
eighteen small landing centers. By early 1990, there were 225 deep-sea fishing vessels operating in
India's exclusive economic zone. Of these, 165 were owned by Indian shipping companies, and the rest
were chartered foreign fishing vessels.
The government provides subsidies to poor fishermen so that they can motorize their traditional craft to
increase the range and frequency of operation, with a consequent increase in the catch and earnings. A
total of about 26,171 traditional craft had been motorized under the program by 1992.
Restrictions[edit]
The banning of trawling by chartered foreign vessels and the speedy motorization of traditional fishing
craft in the 1980s led to a quantum jump in marine fish production in the late 1980s. The export of marine
products rose from 97,179 tons (Rs531 billion) in FY 1987 to 210,800 tons (Rs17.4 trillion) in FY 1992,
making India one of the world's leading seafood exporting nations. This achievement was largely a result
of significant advancements in India's freezing facilities since the 1960s, advancements that enabled
India's seafood products to meet international standards. Frozen shrimp, a high-value item, has become
the dominant seafood export. Other significant export items are frozen frog legs, frozen lobster tails, dried
fish, and shark fins, much of which is exported to seafood-loving Japan. During the eighth plan, marine
products were identified as having major export potential.
Institutes[edit]
There are several specialized institutes that train fishermen. The Central Institute of Fisheries Nautical
and Engineering Training in Juhu instructs operators of deep-sea fishing vessels and technicians for
shore establishments. It has facilities in Madras and Vishakhapatnam for about 500 trainees a year. An
Institute named Fisheries Institute of Technology and Training (FITT) was established with the
participation of TATAs in Tamil Nadu, to improve the socioeconomic condition of fishers.
[11]
The
Integrated Fisheries Project, also headquartered in Kochi, was established for the processing,
popularizing, and marketing of unusual fish. Another training organization, theCentral Institute of Coastal
Engineering for Fisheries in Bangalore, has done techno-economic feasibility studies on locations of
fishing harbor sites and brackish-water fish farms. At present there are 19 Fisheries colleges and one
fisheries university (CIFE: Central Institute of Fisheries Education, Mumbai)functioning in various states of
the country,providing Professional Fisheries education with a view of developing Professionalism in the
field of Fisheries. Among the fisheries colleges, Fisheries college and Research Institute located in
Tuticorin, Tamil Nadu is the more popular college because of the maximum number of intake of MFSc
and PhD candidates every year. Other colleges such as the College of Fisheries, Panangad, College of
Fisheries, and Mangalore are also working well for the professionalism.
To improve returns to fishermen and provide better products for consumers, several states have
organized marketing cooperatives for fishermen. Nevertheless, most traditional fishermen rely on
household members or local fish merchants for the disposal of their catches. In some places, marketing is
carried on entirely by fisherwomen who carry small quantities in containers on their heads to nearby
places. Good wholesale or retail markets are rare.