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Genetically modified tomato

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Plant physiologistAthanasios Theologis with tomatoes that contain the bioengineeredACC synthasegene

Agenetically modified tomato,ortransgenic tomato,is atomatothat has had itsgenesmodified, usinggenetic engineering.The first trialgenetically modified foodwas a tomato engineered to have a longershelf life(theFlavr Savr), which was on the market briefly beginning on May 21, 1994.[1]The first direct consumption tomato was approved in Japan in 2021.[2]Primary work is focused on developing tomatoes with newtraitslike increased resistance to pests orenvironmental stresses.[3]Other projects aim to enrich tomatoes with substances that may offer health benefits or be morenutritious.As well as aiming to produce novel crops, scientists produce genetically modified tomatoes to understand the function of genes naturally present in tomatoes.

Agrobacterium-mediatedgenetic engineeringtechniques were developed in the late 1980s that could successfully transfer genetic material into thenucleargenomeof tomatoes.[4]Genetic material can also beinsertedinto a tomato cell'schloroplastandchromoplastplastomesusingbiolistics.Tomatoes were the first food crop with an edible fruit where this was possible.[5]

Examples

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Delayed ripening

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Tomatoes have been used as amodel organismto study thefruit ripeningofclimactericfruit. To understand the mechanisms involved in the process of ripening, scientists have genetically engineered tomatoes.[6]

In 1994, theFlavr Savrbecame the first commercially grown genetically engineered food to be granted a license for human consumption. A second copy of the tomato genepolygalacturonasewasinsertedinto the tomato genome in theantisensedirection.[7]The polygalacturonaseenzymedegradespectin,a component of the tomatocell wall,causing the fruit to soften. When the antisense gene is expressed itinterfereswith 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 The Flavr Savr failed to achieve commercial success and was withdrawn from the market in 1997.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 atomato paste.[8]

DNA Plant Technology(DNAP),AgritopeandMonsantodeveloped tomatoes that delayed ripening by preventing the production ofethylene,[8]ahormonethat triggers ripening of fruit.[9]All three tomatoes inhibited ethylene production by reducing the amount of1-aminocyclopropane-1-carboxylic acid(ACC), theprecursorto ethylene. DNAP's tomato, called Endless Summer, inserted a truncated version of theACCsynthasegene into the tomato that interfered with theendogenousACC synthase.[8]Monsanto's tomato was engineered with theACCdeaminasegene from the soil bacteriumPseudomonas chlororaphisthat lowered ethylene levels by breaking down ACC.[10]Agritope introduced an S-adenosylmethionine hydrolase (SAMase) encoding gene derived from theE. colibacteriophageT3, which reduced the levels of S-adenosylmethionine, a precursor to ACC.[11]Endless Summer was briefly tested in the marketplace, butpatentarguments forced its withdrawal.[12]

Scientists in India have delayed the ripening of tomatoes by silencing two genes encoding N-glycoproteinmodifying 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.[13]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.[14]

Environmental stress tolerance

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Abioticstresses like frost, drought and increasedsalinityare a limiting factor to the growth of tomatoes.[15]While no genetically modified stress-tolerant plants are currently[when?]commercialised,transgenicapproaches have been researched. An early tomato was developed that contained anantifreeze gene(afa3) from thewinter flounderwith the aim of increasing the tomato's tolerance to frost, which became an icon in the early years of thedebate 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".[16]The antifreeze protein was found to inhibit icerecrystallizationin the flounder blood, but had no effect when expressed in transgenic tobacco.[17]The resulting tomato was never commercialized, possibly because the transgenic plant did not perform well in its frost-tolerance or other agronomic characteristics.[17]Another failed cold tolerant is theE. coliGR transgenic: Others had successfully produced cold tolerantNicotiana tabacumby inserting various enzymes into theplastidsthat 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 ofE. coli'sglutathione reductase→ the chloroplasts ofS. lycopersicumandS. 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.[18]

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 atranscription factor,that was shown to increase cold and drought tolerance in transgenicArabidopsis thalianaplants, was inserted into the tomato. This resulted in increased drought tolerance, but did not appear to have any effect on cold tolerance.[19]Overexpressing avacuolarNa+/H+antiport(AtNHX1) fromA. thalianalead to salt accumulating in the leaves of the plants, but not in the fruit and allowed them to grow more in salt solutions thanwildtypeplants.[20][21]Tobaccoosmoticgenes overexpressed in tomatoes produced plants that held a higher water content than wildtype plants increasing tolerance to drought and salt stress.[22]

Pest resistance

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The insecticidal toxin from the bacteriumBacillus thuringiensishas been inserted into a tomato plant.[23]When field tested they showed resistance to the tobacco hornworm (Manduca sexta), tomato fruitworm (Heliothis zea), the tomato pinworm (Keiferia lycopersicella) and the tomato fruit borer (Helicoverpa armigera).[24][25]A 91-day feeding trial in rats showed no adverse effects,[26]but the Bt tomato has never been commercialised. Tomatoes resistant to a root knotnematodehave been created by inserting acysteineproteinase inhibitorgene fromtaro.[27]Achemically synthesisedcecropin Bgene, usually found in the giant silk moth (Hyalophora cecropia), has been introduced into tomato plants andin vivostudies show significant resistance tobacterial wiltandbacterial spot.[28]When the cell wall proteins, polygalacturonase andexpansinare prevented from being produced in fruits, they are less susceptible to the fungusBotrytis cinereathan normal tomatoes.[29][30]Pest resistant tomatoes can reduce theecological footprintof tomato production while at the same time increase farm income.[31]

Improved nutrition

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Tomatoes have been altered in attempts to add nutritional content. In 2000, the concentration of pro-vitamin Awas increased by adding a bacterial gene encodingphytoenedesaturase,although the total amount ofcarotenoidsremained equal.[32]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 groupGenewatch,toldThe Independentthat she believed, "If you change the basic biochemistry, you could alter the levels of other nutrients very important for health".[33]More recently, scientists createdblue tomatoesthat have increased the production ofanthocyanin,anantioxidantin tomatoes in several ways. One group added atranscription factorfor the production of anthocyanin fromArabidopsis thaliana[34]whereas another used transcription factors from snapdragon (Antirrhinum).[35]When the snapdragon genes were used, the fruits had similar anthocyanin concentrations toblackberriesandblueberries.[36]The inventors of the GMO blue tomato using snapdragon genes, Jonathan Jones and Cathie Martin of theJohn Innes Centre,founded a company called Norfolk Plant Sciences[37]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.[38][39]

Another group has tried to increase the levels ofisoflavone,known for its potential cancer preventive properties, by introducing the soybeanisoflavone synthaseinto tomatoes.[40]

In 2021 Japanese Sanatech Seed issued Sicilian Rouge High GABA tomato variety with increasedGABAlevels.[2]

Improved taste

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Whengeraniolsynthase from lemon basil (Ocimum basilicum) 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 oflycopene.[41]

Vaccines

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Tomatoes (along withpotatoes,bananasand other plants) are being investigated as vehicles for delivering ediblevaccines.Clinical trialshave been conducted on mice using tomatoes expressingantibodiesor proteins that stimulate antibody production targeted tonorovirus,hepatitis B,rabies,HIV,anthraxandrespiratory syncytial virus.[42]Korean scientists are looking at using the tomato to express a vaccine againstAlzheimer's disease.[43]Hilary Koprowski,who was involved in the development of thepolio vaccine,led a group of researchers in developing a tomato expressing arecombinant vaccinetoSARS.[44]

Basic research

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Tomatoes are used as amodel organismin scientific research and they are frequently genetically modified to further understanding of particular processes. Tomatoes have been used as a model inmap-based cloning,where transgenic plants must be created to prove that a gene has been successfully isolated.[45]Theplant peptide hormone,systeminwas 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.[46][47]

References

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