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Genetically modified crops

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… … …
"GM Crops" redirects here. For the journal formerly printed
under that name, see GM Crops & Food.

Genetically modified crops (GM crops) are plants used

in agriculture, the DNA of which has been modified using
genetic engineering methods. Plant genomes can be
engineered by physical methods or by use of
Agrobacterium for the delivery of sequences hosted in T-
DNA binary vectors. 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, environmental conditions,
reduction of spoilage, 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 of pharmaceutical agents, biofuels, and other
industrially useful goods, as well as for bioremediation.[1]

Farmers have widely adopted GM technology. Acreage

increased from 1.7 million hectares in 1996 to 185.1 million
hectares in 2016, some 12% of global cropland. As of
2016, major crop (soybean, maize, canola and cotton)
traits consist of herbicide tolerance (95.9 million
hectares) insect resistance (25.2 million hectares), or
both (58.5 million hectares). In 2015, 53.6 million ha of
GM maize were under cultivation (almost 1/3 of the maize
crop). GM maize outperformed its predecessors: yield
was 5.6 to 24.5% higher with less mycotoxins (−28.8%),
fumonisin (−30.6%) and thricotecens (−36.5%). Non-
target organisms were unaffected, except for Braconidae,
represented by a parasitoid of European corn borer, the
target of Lepidoptera active Bt maize. Biogeochemical
parameters such as lignin content did not vary, while
biomass decomposition was higher.[2]

A 2014 meta-analysis concluded that GM technology

adoption had reduced chemical pesticide use by 37%,
increased crop yields by 22%, and increased farmer
profits by 68%.[3] This reduction in pesticide use has
been ecologically beneficial, but benefits may be reduced
by overuse.[4] Yield gains and pesticide reductions are
larger for insect-resistant crops than for herbicide-
tolerant crops.[5] Yield and profit gains are higher in
developing countries than in developed countries.[3]
Pesticide poisonings were reduced by 2.4 to 9 million
cases per year in India alone and widespread introduction
of Bt cotton led to 25% decline in farmer suicides in India.
[6]
A 2011 review of the relationship between Bt cotton
adoption and farmer suicides in India found that "Available
data show no evidence of a 'resurgence' of farmer
suicides" and that "Bt cotton technology has been very
effective overall in India."[7]

There is a scientific consensus[8][9][10][11] that currently

available food derived from GM crops poses no greater
risk to human health than conventional food,[12][13][14][15]
[16]
but that each GM food needs to be tested on a case-
by-case basis before introduction.[17][18][19] Nonetheless,
members of the public are much less likely than scientists
to perceive GM foods as safe.[20][21][22][23] The legal and
regulatory status of GM foods varies by country, with
some nations banning or restricting them, and others
permitting them with widely differing degrees of
regulation.[24][25][26][27]

However, opponents have objected to GM crops on

grounds including environmental impacts, food safety,
whether GM crops are needed to address food needs,
whether they are sufficiently accessible to farmers in
developing countries[28] and concerns over subjecting
crops to intellectual property law. Safety concerns led 38
countries, including 19 in Europe, to officially prohibit their
cultivation.[2]

History
…

Main article: History of genetic engineering

Humans have directly influenced the genetic makeup of

plants to increase their value as a crop through
domestication. The first evidence of plant domestication
comes from emmer and einkorn wheat found in pre-
Pottery Neolithic A villages in Southwest Asia dated about
10,500 to 10,100 BC.[29] The Fertile Crescent of Western
Asia, Egypt, and India were sites of the earliest planned
sowing and harvesting of plants that had previously been
gathered in the wild. Independent development of
agriculture occurred in northern and southern China,
Africa's Sahel, New Guinea and several regions of the
Americas.[30] The eight Neolithic founder crops (emmer
wheat, einkorn wheat, barley, peas, lentils, bitter vetch,
chick peas and flax) had all appeared by about 7,000 BC.
[31]
Traditional crop breeders have long introduced foreign
germplasm into crops by creating novel crosses. A hybrid
cereal grain was created in 1875, by crossing wheat and
rye.[32] Since then traits including dwarfing genes and
rust resistance have been introduced in that manner.[33]
Plant tissue culture and deliberate mutations have
enabled humans to alter the makeup of plant genomes.[34]
[35]

Modern advances in genetics have allowed humans to

more directly alter plants genetics. In 1970 Hamilton
Smith's lab discovered restriction enzymes that allowed
DNA to be cut at specific places, enabling scientists to
isolate genes from an organism's genome.[36] DNA ligases
that join broken DNA together had been discovered earlier
in 1967,[37] and by combining the two technologies, it was
possible to "cut and paste" DNA sequences and create
recombinant DNA. Plasmids, discovered in 1952,[38]
became important tools for transferring information
between cells and replicating DNA sequences. In 1907 a
bacterium that caused plant tumors, Agrobacterium
tumefaciens, was discovered and in the early 1970s the
tumor inducing agent was found to be a DNA plasmid
called the Ti plasmid.[39] By removing the genes in the
plasmid that caused the tumor and adding in novel genes
researchers were able to infect plants with A. tumefaciens
and let the bacteria insert their chosen DNA sequence
into the genomes of the plants.[40] As not all plant cells
were susceptible to infection by A. tumefaciens other
methods were developed, including electroporation,
micro-injection[41] and particle bombardment with a gene
gun (invented in 1987).[42][43] In the 1980s techniques
were developed to introduce isolated chloroplasts back
into a plant cell that had its cell wall removed. With the
introduction of the gene gun in 1987 it became possible to
integrate foreign genes into a chloroplast.[44] Genetic
transformation has become very efficient in some model
organisms. In 2008 genetically modified seeds were
produced in Arabidopsis thaliana by dipping the flowers in
an Agrobacterium solution.[45] In 2013 CRISPR was first
used to target modification of plant genomes.[46]

The first genetically engineered crop plant was tobacco,

reported in 1983.[47] It was developed creating a chimeric
gene that joined an antibiotic resistant gene to the T1
plasmid from Agrobacterium. The tobacco was infected
with Agrobacterium transformed with this plasmid
resulting in the chimeric gene being inserted into the
plant. Through tissue culture techniques a single tobacco
cell was selected that contained the gene and a new plant
grown from it.[48] The first field trials of genetically
engineered plants occurred in France and the US in 1986,
tobacco plants were engineered to be resistant to
herbicides.[49] In 1987 Plant Genetic Systems, founded by
Marc Van Montagu and Jeff Schell, was the first company
to genetically engineer insect-resistant plants by
incorporating genes that produced insecticidal proteins
from Bacillus thuringiensis (Bt) into tobacco.[50] The
People's Republic of China was the first country to
commercialise transgenic plants, introducing a virus-
resistant tobacco in 1992.[51] In 1994 Calgene attained
approval to commercially release the Flavr Savr tomato, a
tomato engineered to have a longer shelf life.[52] Also in
1994, the European Union approved tobacco engineered
to be resistant to the herbicide bromoxynil, making it the
first genetically engineered crop commercialised in
Europe.[53] In 1995 Bt Potato was approved safe by the
Environmental Protection Agency, after having been
approved by the FDA, making it the first pesticide
producing crop to be approved in the US.[54] In 1996 a
total of 35 approvals had been granted to commercially
grow 8 transgenic crops and one flower crop (carnation),
with 8 different traits in 6 countries plus the EU.[49] By
2010, 29 countries had planted commercialised
genetically modified crops and a further 31 countries had
granted regulatory approval for transgenic crops to be
imported.[55]

The first genetically modified animal to be commercialised

was the GloFish, a Zebra fish with a fluorescent gene
added that allows it to glow in the dark under ultraviolet
light.[56] The first genetically modified animal to be
approved for food use was AquAdvantage salmon in 2015.
[57]
The salmon were transformed with a growth hormone-
regulating gene from a Pacific Chinook salmon and a
promoter from an ocean pout enabling it to grow year-
round instead of only during spring and summer.[58]

Methods
…

Main article: Genetic engineering techniques

Plants (Solanum chacoense) being

transformed using agrobacterium

Genetically engineered crops have genes added or

removed using genetic engineering techniques,[59]
originally including gene guns, electroporation,
microinjection and agrobacterium. More recently, CRISPR
and TALEN offered much more precise and convenient
editing techniques.

Gene guns (also known as biolistics) "shoot" (direct high

energy particles or radiations against[60]) target genes
into plant cells. It is the most common 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 DNA inside the
nucleus. This method has been applied successfully for
many cultivated crops, especially monocots like wheat or
maize, for which transformation using Agrobacterium
tumefaciens has been less successful.[61] The major
disadvantage of this procedure is that serious damage
can be done to the cellular tissue.

Agrobacterium tumefaciens-mediated transformation is

another common technique. Agrobacteria are natural plant
parasites.[62] Their natural ability to transfer genes
provides another engineering method. To create a suitable
environment for themselves, these Agrobacteria insert
their genes into plant hosts, resulting in a proliferation of
modified plant cells near the soil level (crown gall). The
genetic information for tumor growth is encoded on a
mobile, circular DNA fragment (plasmid). When
Agrobacterium infects a plant, it transfers this T-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.

Electroporation is used when the plant tissue does not

contain cell walls. In this technique, "DNA enters the plant
cells through miniature pores which are temporarily
caused by electric pulses."

Microinjection is used to directly inject foreign DNA into

cells.[63]

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.[64][65]

In research tobacco and Arabidopsis thaliana are the most

frequently modified plants, due to well-developed
transformation methods, easy propagation and well
studied genomes.[66][67] They serve as model organisms
for other plant species.

Introducing new genes into plants requires a promoter

specific to the area where the gene is to be expressed.
For instance, to express a gene only in rice grains and not
in leaves, an endosperm-specific promoter is used. The
codons of the gene must be optimized for the organism
due to codon usage bias.

Types of modifications
…

Transgenic maize containing a gene from

the bacteria Bacillus thuringiensis

Transgenic
…
Transgenic plants have genes inserted into them that are
derived from another species. The inserted genes can
come from species within the same kingdom (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 be correctly and efficiently expressed
in the host organism. Transgenic plants are used to
express proteins, like the cry toxins from B. thuringiensis,
herbicide-resistant genes, antibodies,[68] and antigens for
vaccinations.[69] A study led by the European Food Safety
Authority (EFSA) also found viral genes in transgenic
plants.[70]

Transgenic carrots have been used to produce the drug

Taliglucerase alfa which is used to treat Gaucher's
disease.[71] In the laboratory, transgenic plants have been
modified to increase photosynthesis (currently about 2%
at most plants versus the theoretic potential of 9–10%).
[72]
This is possible by changing the rubisco enzyme (i.e.
changing C3 plants into C4 plants[73]), by placing the
rubisco in a carboxysome, by adding CO2 pumps in the
cell wall,[74] or by changing the leaf form or size.[75][76][77]
[78]
Plants have been engineered to exhibit
bioluminescence that may become a sustainable
alternative to electric lighting.[79]

Cisgenic
…
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 regulatory scrutiny as transgenics.
[80]

Subgenic
…
Genetically modified plants can also be developed using
gene knockdown or gene knockout to alter the genetic
makeup of a plant without incorporating genes from other
plants. In 2014, Chinese researcher Gao Caixia filed
patents on the creation of a strain of wheat that is
resistant to powdery mildew. The strain lacks genes that
encode proteins that repress defenses against the mildew.
The researchers deleted all three copies of the genes
from wheat's hexaploid genome. Gao used the TALENs
and CRISPR gene editing tools without adding or changing
any other genes. No field trials were immediately planned.
[81][82]
The CRISPR technique has also been used by Penn
State researcher Yinong Yang to modify white button
mushrooms (Agaricus bisporus) to be non-browning,[83]
and by DuPont Pioneer to make a new variety of corn.[84]

Multiple trait integration

…
With multiple trait integration, several new traits may be
integrated into a new crop.[85]

Economics

Yield

Traits

Crops
…

See also: List of genetically modified crops

Herbicide tolerance
…
Countries
First
Crop Use approved No
approved[192]
in

Ap
with
Animal 2007
Alfalfa US 2005
feed[193] th
app
20

Cooking oil Australia 2003

Margarine Canada 1995

Emulsifiers
Canola
in
US 1995
packaged
foods[193]

Argentina 2001

Australia 2002

Brazil 2008
Fiber
Cottonseed Colombia 2004

Cotton oil Costa Rica 2008

Animal Mexico 2000
[193]
feed
Paraguay 2013

South Africa 2000

US 1994

Argentina 1998

Brazil 2007

Canada 1996

Colombia 2007

Cuba 2011
Animal
Gr
feed
Portug
high-
European C
fructose 1998
Maize Union Re
corn syrup
Slov

corn Rom

starch[193] Honduras 2001

Paraguay 2012

Philippines 2002

South Africa 2002

US 1995

Uruguay 2003

Argentina 1996

Bolivia 2005

Brazil 1998

Canada 1995
Animal
Chile 2007
feed
Soybean Costa Rica 2001
Soybean
oil[193] Mexico 1996

Paraguay 2004

South Africa 2001

US 1993

Uruguay 1996

Canada 2001

Comm
20
Sugar [197]
Food pro
Beet US 1998
block
resum

Insect resistance
…
Countries
First
Crop Use approved Note
approved[192]
in

Argentina 1998

Australia 2003

Brazil 2005

Burkina Faso 2009

China 1997

Colombia 2003

Costa Rica 2008

Fiber
Larg
Cottonseed
produ
Cotton oil India 2002
of B
Animal
cotton
feed[193]
Mexico 1996

Myanmar 2006[N 1]

Pakistan 2010[N 1]

Paraguay 2007

South Africa 1997

Sudan 2012

US 1995

12 h
plan
Eggplant Food Bangladesh 2013 on 1
farm
2014

Argentina 1998

Brazil 2005
Animal Colombia 2003
feed
Centr
high- Mexico 1996 origin
fructose maize
Maize
corn syrup
Paraguay 2007
corn Philippines 2002
starch[193]
South Africa 1997

Uruguay 2003

US 1995

543 h
bt po
Poplar Tree China 1998
plante
2014

Other modified traits

…
Countries
Firs
Crop Use Trait approved
approve
in

Cooking oil High Canada 19

Margarine laurate
US 19
canola
Canola Emulsifiers in
packaged Phytase
US 19
foods[193] production

Delayed Australia 19
senescence Norway 19

Australia 19

Colombia 20

Carnation Ornamental Modified

European
flower 19
Union
colour

Japan 20

Malaysia 20

Norway 19

Animal feed Increased Canada 20

high- lysine US 20
fructose corn Canada 20
Maize syrup
Drought
corn tolerance US 20
[193]
starch

China 20
Virus
Papaya Food[193]
resistance US 19

Modified
Petunia Ornamental flower China 1997
colour

Virus Canada 19
[193]
Food
resistance US 19
Potato
Modified
Industrial[205] US 20
starch

Australia 20

Modified
Rose Ornamental flower Colombia 2010
colour
Japan 20

US 20

Increased Argentina 20

Animal feed oleic acid Canada 20

production US 19
Soybean Soybean
oil[193] Stearidonic Canada 20
acid
US 20
production

Virus
Squash Food[193] US 19
resistance

Sugar Drought
Food Indonesia 20
Cane tolerance

Nicotine
Tobacco Cigarettes US 20
reduction

GM Camelina
…
Several modifications of Camelina sativa have been done,
see §Edible oils and §Non-pesticide pest management
products above.

Development
…
The number of USDA-approved field releases for testing
grew from 4 in 1985 to 1,194 in 2002 and averaged around
800 per year thereafter. The number of sites per release
and the number of gene constructs (ways that the gene of
interest is packaged together with other elements) – have
rapidly increased since 2005. Releases with agronomic
properties (such as drought resistance) jumped from
1,043 in 2005 to 5,190 in 2013. As of September 2013,
about 7,800 releases had been approved for corn, more
than 2,200 for soybeans, more than 1,100 for cotton, and
about 900 for potatoes. Releases were approved for
herbicide tolerance (6,772 releases), insect resistance
(4,809), product quality such as flavor or nutrition
(4,896), agronomic properties like drought resistance
(5,190), and virus/fungal resistance (2,616). The
institutions with the most authorized field releases include
Monsanto with 6,782, Pioneer/DuPont with 1,405,
Syngenta with 565, and USDA's Agricultural Research
Service with 370. As of September 2013 USDA had
received proposals for releasing GM rice, squash, plum,
rose, tobacco, flax, and chicory.[206]

Farming practices
…

This section needs expansion with: examples and

additional citations of how farmers' use of GM Learn
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changes their practices. You can help by adding to it.

Resistance
…
Bacillus thuringiensis
…
Constant exposure to a toxin creates evolutionary
pressure for pests resistant to that toxin.[207] Over-
reliance on glyphosate and a reduction in the diversity of
weed management practices allowed the spread of
glyphosate resistance in 14 weed species in the US,[206]
and in soybeans.[5]

To reduce resistance to Bacillus thuringiensis (Bt) crops,

the 1996 commercialization of transgenic cotton and
maize came with a management strategy to prevent
insects from becoming resistant. Insect resistance
management plans are mandatory for Bt crops. The aim is
to encourage a large population of pests so that any
(recessive) resistance genes are diluted within the
population. Resistance lowers evolutionary fitness in the
absence of the stressor, Bt. In refuges, non-resistant
strains outcompete resistant ones.[208]

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 maturation, thus preventing
transmission of the resistance gene to their progeny.[209]
Refuges (i. e., fields of nontransgenic plants) adjacent to
transgenic fields 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 remains lower.

Complicating factors can affect the success of the high-

dose/refuge strategy. For example, if the temperature is
not ideal, thermal stress can lower Bt toxin production and
leave the plant more susceptible. More importantly,
reduced late-season expression has been documented,
possibly resulting from DNA methylation of the promoter.
[210]
The success of the high-dose/refuge strategy has
successfully maintained the value of Bt crops. This
success has depended on factors independent of
management strategy, including low initial resistance
allele frequencies, fitness costs associated with
resistance, and the abundance of non-Bt host plants
outside the refuges.[211]

Companies that produce Bt seed are introducing strains

with multiple Bt proteins. Monsanto did this with Bt cotton
in India, where the product was rapidly adopted.[212]
Monsanto has also; in an attempt to simplify the process
of implementing refuges in fields to comply with Insect
Resistance Management(IRM) policies and prevent
irresponsible planting practices; begun marketing seed
bags with a set proportion of refuge (non-transgenic)
seeds mixed in with the Bt seeds being sold. Coined
"Refuge-In-a-Bag" (RIB), this practice is intended to
increase farmer compliance with refuge requirements and
reduce additional labor needed at planting from having
separate Bt and refuge seed bags on hand.[213] This
strategy is likely to reduce the likelihood of Bt-resistance
occurring for corn rootworm, but may increase the risk of
resistance for lepidopteran corn pests, such as European
corn borer. Increased concerns for resistance with seed
mixtures include partially resistant larvae on a Bt plant
being able to move to a susceptible plant to survive or
cross pollination of refuge pollen on to Bt plants that can
lower the amount of Bt expressed in kernels for ear
feeding insects.[214][215]

Herbicide resistance
…
Best management practices (BMPs) to control weeds may
help delay resistance. BMPs include applying multiple
herbicides with different modes of action, rotating crops,
planting weed-free seed, scouting fields routinely,
cleaning equipment to reduce the transmission of weeds
to other fields, and maintaining field borders.[206] The
most widely planted GM crops are designed to tolerate
herbicides. By 2006 some weed populations had evolved
to tolerate some of the same herbicides. Palmer amaranth
is a weed that competes with cotton. A native of the
southwestern US, it traveled east and was first found
resistant to glyphosate in 2006, less than 10 years after
GM cotton was introduced.[216][217]

Plant protection
…
Farmers generally use less insecticide when they plant Bt-
resistant crops. Insecticide use on corn farms declined
from 0.21 pound per planted acre in 1995 to 0.02 pound in
2010. This is consistent with the decline in European corn
borer populations as a direct result of Bt corn and cotton.
The establishment of minimum refuge requirements
helped delay the evolution of Bt resistance. However,
resistance appears to be developing to some Bt traits in
some areas.[206]

Tillage
…