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A genetically modified tomato, or transgenic tomato, is a
tomato The tomato is the edible berry of the plant ''Solanum lycopersicum'', commonly known as the tomato plant. The species originated in western South America, Mexico, and Central America. The Mexican Nahuatl word gave rise to the Spanish word ...
that has had its
gene In biology, the word gene (from , ; "... Wilhelm Johannsen coined the word gene to describe the Mendelian units of heredity..." meaning ''generation'' or ''birth'' or ''gender'') can have several different meanings. The Mendelian gene is a b ...
s modified, using genetic engineering. The first trial
genetically modified food Genetically modified foods (GM foods), also known as genetically engineered foods (GE foods), or bioengineered foods are foods produced from organisms that have had changes introduced into their DNA using the methods of genetic engineering. Gene ...
was a tomato engineered to have a longer
shelf life Shelf life is the length of time that a commodity may be stored without becoming unfit for use, consumption, or sale. In other words, it might refer to whether a commodity should no longer be on a pantry shelf (unfit for use), or no longer on a ...
(the
Flavr Savr Flavr Savr (also known as CGN-89564-2; pronounced "flavor saver"), a genetically modified tomato, was the first commercially grown genetically engineered food to be granted a license for human consumption. It was developed by the Californian company ...
), which was on the market briefly beginning on May 21, 1994. The first direct consumption tomato was approved in Japan in 2021. Primary work is focused on developing tomatoes with new traits like increased resistance to pests or
environmental stresses Stress, either physiological, biological or psychological, is an organism's response to a stressor such as an environmental condition. Stress is the body's method of reacting to a condition such as a threat, challenge or physical and psycholog ...
. Other projects aim to enrich tomatoes with substances that may offer health benefits or be more
nutritious Nutrition is the biochemical and physiological process by which an organism uses food to support its life. It provides organisms with nutrients, which can be metabolized to create energy and chemical structures. Failure to obtain sufficient nu ...
. As well as aiming to produce novel crops, scientists produce genetically modified tomatoes to understand the function of genes naturally present in tomatoes. Wild tomatoes are small, green and largely unappetizing, but after centuries of breeding there are now thousands of varieties grown worldwide. Agrobacterium-mediated genetic engineering techniques were developed in the late 1980s that could successfully transfer genetic material into the nuclear
genome In the fields of molecular biology and genetics, a genome is all the genetic information of an organism. It consists of nucleotide sequences of DNA (or RNA in RNA viruses). The nuclear genome includes protein-coding genes and non-coding g ...
of tomatoes. Genetic material can also be inserted into a tomato cell's chloroplast and
chromoplast Chromoplasts are plastids, heterogeneous organelles responsible for pigment synthesis and storage in specific photosynthetic eukaryotes. It is thought that like all other plastids including chloroplasts and leucoplasts they are descended from sy ...
plastome Chloroplast DNA (cpDNA) is the DNA located in chloroplasts, which are photosynthetic organelles located within the cells of some eukaryotic organisms. Chloroplasts, like other types of plastid, contain a genome separate from that in the cell n ...
s using biolistics. Tomatoes were the first food crop with an edible fruit where this was possible.


Examples


Delayed ripening

Tomatoes have been used as a model organism to study the
fruit ripening Ripening is a process in fruits that causes them to become more palatable. In general, fruit becomes sweeter, less green, and softer as it ripens. Even though the acidity of fruit increases as it ripens, the higher acidity level does not make the ...
of climacteric fruit. To understand the mechanisms involved in the process of ripening, scientists have genetically engineered tomatoes. In 1994, the
Flavr Savr Flavr Savr (also known as CGN-89564-2; pronounced "flavor saver"), a genetically modified tomato, was the first commercially grown genetically engineered food to be granted a license for human consumption. It was developed by the Californian company ...
became the first commercially grown genetically engineered food to be granted a license for human consumption. A second copy of the tomato gene ''
polygalacturonase Endo-polygalacturonase (EC 3.2.1.15, pectin depolymerase, pectolase, pectin hydrolase, and poly-α-1,4-galacturonide glycanohydrolase; systematic name (1→4)-α-D-galacturonan glycanohydrolase (endo-cleaving)) is an enzyme that hydrolyzes the α- ...
'' was inserted into the tomato genome in the
antisense In molecular biology and genetics, the sense of a nucleic acid molecule, particularly of a strand of DNA or RNA, refers to the nature of the roles of the strand and its complement in specifying a sequence of amino acids. Depending on the context ...
direction. The polygalacturonase
enzyme Enzymes () are proteins that act as biological catalysts by accelerating chemical reactions. The molecules upon which enzymes may act are called substrates, and the enzyme converts the substrates into different molecules known as products ...
degrades pectin, a component of the tomato cell wall, causing the fruit to soften. When the antisense gene is expressed it interferes with the production of the polygalacturonase enzyme, delaying the ripening process. The Flavr Savr failed to achieve commercial success and was withdrawn from the market in 1997. Similar technology, but using a truncated version of the polygalacturonase gene, was used to make a
tomato paste Tomato paste is a thick paste made by cooking tomatoes for several hours to reduce the water content, straining out the seeds and skins, and cooking the liquid again to reduce the base to a thick, rich concentrate. It is used to impart an inten ...
. DNA Plant Technology (DNAP), Agritope and
Monsanto The Monsanto Company () was an American agrochemical and agricultural biotechnology corporation founded in 1901 and headquartered in Creve Coeur, Missouri. Monsanto's best known product is Roundup, a glyphosate-based herbicide, developed in ...
developed tomatoes that delayed ripening by preventing the production of ethylene, a
hormone A hormone (from the Greek participle , "setting in motion") is a class of signaling molecules in multicellular organisms that are sent to distant organs by complex biological processes to regulate physiology and behavior. Hormones are require ...
that triggers ripening of fruit. All three tomatoes inhibited ethylene production by reducing the amount of
1-aminocyclopropane-1-carboxylic acid 1-Aminocyclopropane-1-carboxylic acid (ACC) is a disubstituted cyclic α-amino acid in which a cyclopropane ring is fused to the C atom of the amino acid. It is a white solid. Many cyclopropane-substituted amino acids are known, but this one oc ...
(ACC), the
precursor Precursor or Precursors may refer to: * Precursor (religion), a forerunner, predecessor ** The Precursor, John the Baptist Science and technology * Precursor (bird), a hypothesized genus of fossil birds that was composed of fossilized parts of u ...
to ethylene. DNAP's tomato, called Endless Summer, inserted a truncated version of the ''ACC synthase'' gene into the tomato that interfered with the endogenous ''ACC synthase''. Monsanto's tomato was engineered with the ''ACC deaminase'' gene from the soil bacterium '' Pseudomonas chlororaphis'' that lowered ethylene levels by breaking down ACC. Agritope introduced an S-adenosylmethionine hydrolase (SAMase) encoding gene derived from the '' E. coli'' bacteriophage T3, which reduced the levels of S-adenosylmethionine, a precursor to ACC. Endless Summer was briefly tested in the marketplace, but
patent A patent is a type of intellectual property that gives its owner the legal right to exclude others from making, using, or selling an invention for a limited period of time in exchange for publishing an enabling disclosure of the invention."A ...
arguments forced its withdrawal. Scientists in India have delayed the ripening of tomatoes by silencing two genes encoding N- glycoprotein modifying enzymes, α-mannosidase and β-D-N-acetylhexosaminidase. The fruits produced were not visibly damaged after being stored at room temperature for 45 days, whereas unmodified tomatoes had gone rotten. In India, where 30% of fruit is wasted before it reaches the market due to a lack of refrigeration and poor road infrastructure, the researchers hope genetic engineering of the tomato may decrease wastage.


Environmental stress tolerance

Abiotic In biology and ecology, abiotic components or abiotic factors are non-living chemical and physical parts of the environment that affect living organisms and the functioning of ecosystems. Abiotic factors and the phenomena associated with them under ...
stresses like frost, drought and increased salinity are a limiting factor to the growth of tomatoes. While no genetically modified stress-tolerant plants are currently commercialised, transgenic approaches have been researched. An early tomato was developed that contained an antifreeze gene (''afa3'') from the
winter flounder The winter flounder (''Pseudopleuronectes americanus''), also known as the black back, is a right-eyed ("dextral") flatfish of the family Pleuronectidae. It is native to coastal waters of the western north Atlantic coast, from Labrador, Canada to ...
with the aim of increasing the tomato's tolerance to frost, which became an icon in the early years of the debate over genetically modified foods, especially in relation to the perceived ethical dilemma of combining genes from different species. This tomato gained the moniker "fish tomato". The antifreeze protein was found to inhibit ice recrystallization in the flounder blood, but had no effect when expressed in transgenic tobacco. The resulting tomato was never commercialized, possibly because the transgenic plant did not perform well in its frost-tolerance or other agronomic characteristics. Another failed cold tolerant is the ''E. coli'' GR transgenic: Others had successfully produced cold tolerant ''
Nicotiana tabacum ''Nicotiana tabacum'', or cultivated tobacco, is an Annual plant, annually grown herbaceous plant of the ''Nicotiana'' genus. The plant is tropical in origin, is commonly grown throughout the world, and is often found in Cultivation of tobacco, ...
'' by inserting various enzymes into the plastids that had already been observed to be more active under cold stress in the donor organism. Brüggemann et al. 1999 thus assumed the same would hold for a transfer of '' E. coli''s
glutathione reductase Glutathione reductase (GR) also known as glutathione-disulfide reductase (GSR) is an enzyme that in humans is encoded by the GSR gene. Glutathione reductase (EC 1.8.1.7) catalyzes the reduction of glutathione disulfide ( GSSG) to the sulfhydryl f ...
→ the chloroplasts of '' S. lycopersicum'' and '' S. peruvianum''. They overexpressed the donated GR and this was supplementing the endogenous GR. Although total GR activity was increased, no improvement in cold tolerance did obtain. Other genes from various species have been inserted into the tomato with the hope of increasing their resistance to various environmental factors. A gene from rice (''Osmyb4''), which codes for a
transcription factor In molecular biology, a transcription factor (TF) (or sequence-specific DNA-binding factor) is a protein that controls the rate of transcription of genetic information from DNA to messenger RNA, by binding to a specific DNA sequence. The f ...
, that was shown to increase cold and drought tolerance in transgenic '' Arabidopsis thaliana'' plants, was inserted into the tomato. This resulted in increased drought tolerance, but did not appear to have any effect on cold tolerance. Overexpressing a
vacuolar A vacuole () is a membrane-bound organelle which is present in plant and fungal cells and some protist, animal, and bacterial cells. Vacuoles are essentially enclosed compartments which are filled with water containing inorganic and organic mo ...
Na+/H+ antiport (''AtNHX1'') from ''A. thaliana'' lead to salt accumulating in the leaves of the plants, but not in the fruit and allowed them to grow more in salt solutions than
wildtype The wild type (WT) is the phenotype of the typical form of a species as it occurs in nature. Originally, the wild type was conceptualized as a product of the standard "normal" allele at a locus, in contrast to that produced by a non-standard, " ...
plants. Tobacco osmotic genes overexpressed in tomatoes produced plants that held a higher water content than wildtype plants increasing tolerance to drought and salt stress.


Pest resistance

The insecticidal toxin from the bacterium '' Bacillus thuringiensis'' has been inserted into a tomato plant. When field tested they showed resistance to the tobacco hornworm ('' Manduca sexta''), tomato fruitworm ('' Heliothis zea''), the tomato pinworm (''
Keiferia lycopersicella ''Keiferia lycopersicella'', the tomato pinworm, is a moth of the family Gelechiidae. It is found in warm areas in Mexico, California, Texas, Georgia, Florida, Hawaii, Cuba, Hispaniola and the Bahamas. It has also been reported from greenhouses ...
'') and the tomato fruit borer ('' Helicoverpa armigera''). A 91-day feeding trial in rats showed no adverse effects, but the Bt tomato has never been commercialised. Tomatoes resistant to a root knot nematode have been created by inserting a cysteine proteinase inhibitor gene from
taro Taro () (''Colocasia esculenta)'' is a root vegetable. It is the most widely cultivated species of several plants in the family Araceae that are used as vegetables for their corms, leaves, and petioles. Taro corms are a food staple in Afri ...
. A chemically synthesised ''cecropin B'' gene, usually found in the giant silk moth (''
Hyalophora cecropia ''Hyalophora cecropia'', the cecropia moth, is North America's largest native moth. It is a member of the family Saturniidae, or giant silk moths. Females have been documented with a wingspan of five to seven inches (160 mm) or more. These m ...
''), has been introduced into tomato plants and
in vivo Studies that are ''in vivo'' (Latin for "within the living"; often not italicized in English) are those in which the effects of various biological entities are tested on whole, living organisms or cells, usually animals, including humans, and ...
studies show significant resistance to
bacterial wilt Bacterial wilt is a complex of diseases that occur in plants such as Cucurbitaceae and Solanaceae (tomato, common bean, etc.) and are caused by the pathogens ''Erwinia tracheiphila'', a gram-negative bacterium, or '' Curtobacterium flaccumfacien ...
and bacterial spot. When the cell wall proteins, polygalacturonase and expansin are prevented from being produced in fruits, they are less susceptible to the fungus ''
Botrytis cinerea ''Botrytis cinerea'' is a necrotrophic fungus that affects many plant species, although its most notable hosts may be wine grapes. In viticulture, it is commonly known as "botrytis bunch rot"; in horticulture, it is usually called "grey mould" o ...
'' than normal tomatoes. Pest resistant tomatoes can reduce the ecological footprint of tomato production while at the same time increase farm income.Groeneveld, Rolf, Erik Ansink, Clemens van de Wiel, and Justus Wesseler (2011) Benefits and costs of biologically contained GM tomatoes and eggplants in Italy and Spain. Sustainability. 2011, 3, 1265-1281


Improved nutrition

Tomatoes have been altered in attempts to add nutritional content. In 2000, the concentration of pro-
vitamin A Vitamin A is a fat-soluble vitamin and an essential nutrient for humans. It is a group of organic compounds that includes retinol, retinal (also known as retinaldehyde), retinoic acid, and several provitamin A carotenoids (most notably ...
was increased by adding a bacterial gene encoding
phytoene Phytoene () is a 40-carbon intermediate in the biosynthesis of carotenoids. The synthesis of phytoene is the first committed step in the synthesis of carotenoids in plants. Phytoene is produced from two molecules of geranylgeranyl pyrophosphate (G ...
desaturase, although the total amount of carotenoids remained equal. The researchers admitted at the time that it had no prospect of being grown commercially due to the anti-GM climate. Sue Meyer of the pressure group Genewatch, told ''
The Independent ''The Independent'' is a British online newspaper. It was established in 1986 as a national morning printed paper. Nicknamed the ''Indy'', it began as a broadsheet and changed to tabloid format in 2003. The last printed edition was publish ...
'' that she believed, "If you change the basic biochemistry, you could alter the levels of other nutrients very important for health". More recently, scientists created blue tomatoes that have increased the production of anthocyanin, an antioxidant in tomatoes in several ways. One group added a
transcription factor In molecular biology, a transcription factor (TF) (or sequence-specific DNA-binding factor) is a protein that controls the rate of transcription of genetic information from DNA to messenger RNA, by binding to a specific DNA sequence. The f ...
for the production of anthocyanin from '' Arabidopsis thaliana'' whereas another used transcription factors from snapdragon (''
Antirrhinum ''Antirrhinum'' is a genus of plants commonly known as dragon flowers, snapdragons and dog flower because of the flowers' fancied resemblance to the face of a dragon that opens and closes its mouth when laterally squeezed. They are native t ...
''). When the snapdragon genes were used, the fruits had similar anthocyanin concentrations to
blackberries The blackberry is an edible fruit produced by many species in the genus ''Rubus'' in the family (biology), family Rosaceae, hybrids among these species within the subgenus ''Rubus'', and hybrids between the subgenera ''Rubus'' and ''Idaeobatus' ...
and blueberries. The inventors of the GMO blue tomato using snapdragon genes, Jonathan Jones and Cathie Martin of the
John Innes Centre The John Innes Centre (JIC), located in Norwich, Norfolk, England, is an independent centre for research and training in plant and microbial science founded in 1910. It is a registered charity (No 223852) grant-aided by the Biotechnology and ...
, founded a company called Norfolk Plant Sciences to commercialize the blue tomato. They partnered with a company in Canada called New Energy Farms to grow a large crop of blue tomatoes, from which to create juice to test in clinical trials on the way to obtaining regulatory approval. Another group has tried to increase the levels of
isoflavone Isoflavones are substituted derivatives of isoflavone, a type of naturally occurring isoflavonoids, many of which act as phytoestrogens in mammals. Isoflavones are produced almost exclusively by the members of the bean family, Fabaceae (Leguminosae) ...
, known for its potential cancer preventive properties, by introducing the soybean ''isoflavone synthase'' into tomatoes. In 2021 Japanese Sanatech Seed issued Sicilian Rouge High GABA tomato variety with increased GABA levels.


Improved taste

When
geraniol Geraniol is a monoterpenoid and an alcohol. It is the primary component of citronella oil and is a primary component of rose oil, palmarosa oil. It is a colorless oil, although commercial samples can appear yellow. It has low solubility in w ...
synthase from lemon basil (''
Ocimum basilicum Basil (, ; ''Ocimum basilicum'' , also called great basil, is a culinary herb of the family Lamiaceae (mints). It is a tender plant, and is used in cuisines worldwide. In Western cuisine, the generic term "basil" refers to the variety also k ...
'') was expressed in tomato fruits under a fruit-specific promoter, 60% of untrained taste testers preferred the taste and smell of the transgenic tomatoes. The fruits contained around half the amount of
lycopene Lycopene is an organic compound classified as a tetraterpene and a carotene. Lycopene (from the neo-Latin '' Lycopersicum'', the tomato species) is a bright red carotenoid hydrocarbon found in tomatoes and other red fruits and vegetables. Occu ...
.


Vaccines

Tomatoes (along with
potato The potato is a starchy food, a tuber of the plant ''Solanum tuberosum'' and is a root vegetable native to the Americas. The plant is a perennial in the nightshade family Solanaceae. Wild potato species can be found from the southern Unit ...
es, bananas and other plants) are being investigated as vehicles for delivering edible
vaccines A vaccine is a biological preparation that provides active acquired immunity to a particular infectious or malignant disease. The safety and effectiveness of vaccines has been widely studied and verified.< ...
. Clinical trials have been conducted on mice using tomatoes expressing antibodies or proteins that stimulate antibody production targeted to
norovirus Norovirus, sometimes referred to as the winter vomiting disease, is the most common cause of gastroenteritis. Infection is characterized by non-bloody diarrhea, vomiting, and stomach pain. Fever or headaches may also occur. Symptoms usually devel ...
,
hepatitis B Hepatitis B is an infectious disease caused by the '' Hepatitis B virus'' (HBV) that affects the liver; it is a type of viral hepatitis. It can cause both acute and chronic infection. Many people have no symptoms during an initial infection. ...
,
rabies Rabies is a viral disease that causes encephalitis in humans and other mammals. Early symptoms can include fever and tingling at the site of exposure. These symptoms are followed by one or more of the following symptoms: nausea, vomiting, ...
,
HIV The human immunodeficiency viruses (HIV) are two species of ''Lentivirus'' (a subgroup of retrovirus) that infect humans. Over time, they cause acquired immunodeficiency syndrome (AIDS), a condition in which progressive failure of the immune ...
, anthrax and
respiratory syncytial virus Respiratory syncytial virus (RSV), also called human respiratory syncytial virus (hRSV) and human orthopneumovirus, is a common, contagious virus that causes infections of the respiratory tract. It is a negative-sense, single-stranded RNA virus. ...
. Korean scientists are looking at using the tomato to express a vaccine against Alzheimer's disease.
Hilary Koprowski Hilary Koprowski (5 December 191611 April 2013) was a Polish virologist and immunologist active in the United States who demonstrated the world's first effective live polio vaccine. He authored or co-authored over 875 scientific papers and co ...
, who was involved in the development of the
polio vaccine Polio vaccines are vaccines used to prevent poliomyelitis (polio). Two types are used: an inactivated poliovirus given by injection (IPV) and a weakened poliovirus given by mouth (OPV). The World Health Organization (WHO) recommends all chi ...
, led a group of researchers in developing a tomato expressing a
recombinant vaccine A vaccine is a biological preparation that provides active acquired immunity to a particular infectious or malignant disease. The safety and effectiveness of vaccines has been widely studied and verified.< ...
to
SARS Severe acute respiratory syndrome (SARS) is a viral respiratory disease of zoonotic origin caused by the severe acute respiratory syndrome coronavirus (SARS-CoV or SARS-CoV-1), the first identified strain of the SARS coronavirus species, ''seve ...
.


Basic research

Tomatoes are used as a model organism in scientific research and they are frequently genetically modified to further understanding of particular processes. Tomatoes have been used as a model in map-based cloning, where transgenic plants must be created to prove that a gene has been successfully isolated. The
plant peptide hormone Peptide signaling plays a significant role in various aspects of plant growth and development and specific receptors for various peptides have been identified as being membrane-localized receptor kinases, the largest family of receptor-like molecule ...
, systemin was first identified in tomato plants and genetic modification has been used to demonstrate its function, by adding antisense genes to silence the native gene or by adding extra copies of the native gene.


References

{{tomatoes Genetically modified organisms in agriculture Tomatoes