Genetically modified tomato

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Plant physiologist Athanasios Theologis with tomatoes that contain the bioengineered ACC synthase gene Tomatoes ARS.jpg
Plant physiologist Athanasios Theologis with tomatoes that contain the bioengineered ACC synthase gene

A genetically modified tomato, or transgenic tomato, is a tomato that has had its genes modified, using genetic engineering. The first trial genetically modified food was a tomato engineered to have a longer shelf life (the Flavr 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 new traits like increased resistance to pests or environmental stresses. [3] Other projects aim to enrich tomatoes with substances that may offer health benefits or be more nutritious. As well as aiming to produce novel crops, scientists produce genetically modified tomatoes to understand the function of genes naturally present in tomatoes.

Contents

Agrobacterium -mediated genetic engineering techniques were developed in the late 1980s that could successfully transfer genetic material into the nuclear genome of tomatoes. [4] Genetic material can also be inserted into a tomato cell's chloroplast and chromoplast plastomes using biolistics. Tomatoes were the first food crop with an edible fruit where this was possible. [5]

Examples

Delayed ripening

Tomatoes have been used as a model organism to study the fruit ripening of climacteric fruit. To understand the mechanisms involved in the process of ripening, scientists have genetically engineered tomatoes. [6]

In 1994, the Flavr Savr became the first commercially grown genetically engineered food to be granted a license for human consumption. A second copy of the tomato gene polygalacturonase was inserted into the tomato genome in the antisense direction. [7] The polygalacturonase enzyme 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 The Flavr Savr failed to achieve commercial success and was withdrawn from the market in 1997.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. [8]

DNA Plant Technology (DNAP), Agritope and Monsanto developed tomatoes that delayed ripening by preventing the production of ethylene, [8] a hormone that triggers ripening of fruit. [9] All three tomatoes inhibited ethylene production by reducing the amount of 1-aminocyclopropane-1-carboxylic acid (ACC), the precursor 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. [8] Monsanto's tomato was engineered with the ACC deaminase gene from the soil bacterium Pseudomonas chlororaphis that lowered ethylene levels by breaking down ACC. [10] 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. [11] Endless Summer was briefly tested in the marketplace, but patent arguments forced its withdrawal. [12]

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. [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

Abiotic stresses like frost, drought and increased salinity are a limiting factor to the growth of tomatoes. [15] While no genetically modified stress-tolerant plants are currently[ when? ] commercialised, transgenic approaches have been researched. An early tomato was developed that contained an antifreeze gene (afa3) from the winter flounder 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". [16] The antifreeze protein was found to inhibit ice recrystallization in 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 the E. coli GR transgenic: Others had successfully produced cold tolerant Nicotiana tabacum 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 → 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. [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 a transcription factor, 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. [19] Overexpressing a vacuolar 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 plants. [20] [21] Tobacco osmotic genes overexpressed in tomatoes produced plants that held a higher water content than wildtype plants increasing tolerance to drought and salt stress. [22]

Pest resistance

The insecticidal toxin from the bacterium Bacillus thuringiensis has 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 knot nematode have been created by inserting a cysteine proteinase inhibitor gene from taro. [27] A chemically synthesised cecropin B gene, usually found in the giant silk moth ( Hyalophora cecropia ), has been introduced into tomato plants and in vivo studies show significant resistance to bacterial wilt and bacterial spot. [28] When the cell wall proteins, polygalacturonase and expansin are prevented from being produced in fruits, they are less susceptible to the fungus Botrytis cinerea than normal tomatoes. [29] [30] Pest resistant tomatoes can reduce the ecological footprint of tomato production while at the same time increase farm income. [31]

Improved nutrition

Tomatoes have been altered in attempts to add nutritional content. In 2000, the concentration of pro-vitamin A was increased by adding a bacterial gene encoding phytoene desaturase, although the total amount of carotenoids remained 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 group Genewatch, told The Independent that she believed, "If you change the basic biochemistry, you could alter the levels of other nutrients very important for health". [33] 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 for the production of anthocyanin from Arabidopsis thaliana [34] whereas another used transcription factors from snapdragon ( Antirrhinum ). [35] When the snapdragon genes were used, the fruits had similar anthocyanin concentrations to blackberries and blueberries. [36] The inventors of the GMO blue tomato using snapdragon genes, Jonathan Jones and Cathie Martin of the John 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 of isoflavone, known for its potential cancer preventive properties, by introducing the soybean isoflavone synthase into tomatoes. [40]

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

Improved taste

When geraniol synthase 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 of lycopene. [41]

Vaccines

Tomatoes (along with potatoes, bananas and other plants) are being investigated as vehicles for delivering edible vaccines. Clinical trials have been conducted on mice using tomatoes expressing antibodies or proteins that stimulate antibody production targeted to norovirus, hepatitis B, rabies, HIV, anthrax and respiratory syncytial virus. [42] Korean scientists are looking at using the tomato to express a vaccine against Alzheimer's disease. [43] Hilary Koprowski, who was involved in the development of the polio vaccine, led a group of researchers in developing a tomato expressing a recombinant vaccine to SARS. [44]

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. [45] The plant peptide hormone, 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. [46] [47]

Related Research Articles

<i>Bacillus thuringiensis</i> Species of bacteria used as an insecticide

Bacillus thuringiensis is a gram-positive, soil-dwelling bacterium, the most commonly used biological pesticide worldwide. B. thuringiensis also occurs naturally in the gut of caterpillars of various types of moths and butterflies, as well on leaf surfaces, aquatic environments, animal feces, insect-rich environments, and flour mills and grain-storage facilities. It has also been observed to parasitize other moths such as Cadra calidella—in laboratory experiments working with C. calidella, many of the moths were diseased due to this parasite.

<span class="mw-page-title-main">Genetically modified maize</span> Genetically modified crop

Genetically modified maize (corn) is a genetically modified crop. Specific maize strains have been genetically engineered to express agriculturally-desirable traits, including resistance to pests and to herbicides. Maize strains with both traits are now in use in multiple countries. GM maize has also caused controversy with respect to possible health effects, impact on other insects and impact on other plants via gene flow. One strain, called Starlink, was approved only for animal feed in the US but was found in food, leading to a series of recalls starting in 2000.

<span class="mw-page-title-main">Genetically modified organism</span> Organisms whose genetic material has been altered using genetic engineering methods

A genetically modified organism (GMO) is any organism whose genetic material has been altered using genetic engineering techniques. The exact definition of a genetically modified organism and what constitutes genetic engineering varies, with the most common being an organism altered in a way that "does not occur naturally by mating and/or natural recombination". A wide variety of organisms have been genetically modified (GM), including animals, plants, and microorganisms.

<span class="mw-page-title-main">Genetic engineering</span> Manipulation of an organisms genome

Genetic engineering, also called genetic modification or genetic manipulation, is the modification and manipulation of an organism's genes using technology. It is a set of technologies used to change the genetic makeup of cells, including the transfer of genes within and across species boundaries to produce improved or novel organisms. New DNA is obtained by either isolating and copying the genetic material of interest using recombinant DNA methods or by artificially synthesising the DNA. A construct is usually created and used to insert this DNA into the host organism. The first recombinant DNA molecule was made by Paul Berg in 1972 by combining DNA from the monkey virus SV40 with the lambda virus. As well as inserting genes, the process can be used to remove, or "knock out", genes. The new DNA can be inserted randomly, or targeted to a specific part of the genome.

<span class="mw-page-title-main">Genetically modified food</span> Foods produced from organisms that have had changes introduced into their DNA

Genetically modified foods, also known as genetically engineered foods, or bioengineered foods are foods produced from organisms that have had changes introduced into their DNA using various methods of genetic engineering. Genetic engineering techniques allow for the introduction of new traits as well as greater control over traits when compared to previous methods, such as selective breeding and mutation breeding.

Flavr Savr, 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 Calgene in the 1980s. The tomato has an improved shelf-life, increased fungal resistance and a slightly increased viscosity compared to its non-modified counterpart. It was meant to be harvested ripe for increased flavor for long-distance shipping. The Flavr Savr contains two genes added by Calgene; a reversed antisense polygalacturonase gene which inhibits the production of a rotting enzyme and a gene responsible for the creation of APH(3')II, which confers resistance to certain aminoglycoside antibiotics including kanamycin and neomycin. On May 18, 1994, the FDA completed its evaluation of the Flavr Savr tomato and the use of APH(3')II, concluding that the tomato "is as safe as tomatoes bred by conventional means" and "that the use of aminoglycoside 3'-phosphotransferase II is safe for use as a processing aid in the development of new varieties of tomato, rapeseed oil, and cotton intended for food use." It was first sold in 1994, and was only available for a few years before production ceased in 1997. Calgene made history, but mounting costs prevented the company from becoming profitable, and it was eventually acquired by Monsanto Company.

<span class="mw-page-title-main">Ripening</span> Process in fruits that causes them to become more palatable

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 fruit seem tarter. This effect is attributed to the Brix-Acid Ratio. Climacteric fruits ripen after harvesting and so some fruits for market are picked green.

<span class="mw-page-title-main">Genetically modified crops</span> Plants used in agriculture

Genetically modified 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, 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.

<span class="mw-page-title-main">Plant Genetic Systems</span> Bayer Subsidiary

Plant Genetic Systems (PGS), since 2002 part of Bayer CropScience, is a biotech company located in Ghent, Belgium. The focus of its activities is the genetic engineering of plants. The company is best known for its work in the development of insect-resistant transgenic plants.

<span class="mw-page-title-main">Tomato</span> Edible berry of the tomato plant, Solanum lycopersicum

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 Nahuatl word tomatl gave rise to the Spanish word tomate, from which the English word tomato derives. Its domestication and use as a cultivated food may have originated with the indigenous peoples of Mexico. The Aztecs used tomatoes in their cooking at the time of the Spanish conquest of the Aztec Empire, and after the Spanish encountered the tomato for the first time after their contact with the Aztecs, they brought the plant to Europe, in a widespread transfer of plants known as the Columbian exchange. From there, the tomato was introduced to other parts of the European-colonized world during the 16th century.

<span class="mw-page-title-main">Genetically modified plant</span> Plants with human-introduced genes from other organisms

Genetically modified plants have been engineered for scientific research, to create new colours in plants, deliver vaccines, and to create enhanced crops. Plant genomes can be engineered by physical methods or by use of Agrobacterium for the delivery of sequences hosted in T-DNA binary vectors. Many plant cells are pluripotent, meaning that a single cell from a mature plant can be harvested and then under the right conditions form a new plant. This ability is most often taken advantage by genetic engineers through selecting cells that can successfully be transformed into an adult plant which can then be grown into multiple new plants containing transgene in every cell through a process known as tissue culture.

In botany, drought tolerance is the ability by which a plant maintains its biomass production during arid or drought conditions. Some plants are naturally adapted to dry conditions, surviving with protection mechanisms such as desiccation tolerance, detoxification, or repair of xylem embolism. Other plants, specifically crops like corn, wheat, and rice, have become increasingly tolerant to drought with new varieties created via genetic engineering. From an evolutionary perspective, the type of mycorrhizal associations formed in the roots of plants can determine how fast plants can adapt to drought.

<span class="mw-page-title-main">Anthocyanin</span> Class of chemical compounds

Anthocyanins, also called anthocyans, are water-soluble vacuolar pigments that, depending on their pH, may appear red, purple, blue, or black. In 1835, the German pharmacist Ludwig Clamor Marquart gave the name Anthokyan to a chemical compound that gives flowers a blue color for the first time in his treatise "Die Farben der Blüthen". Food plants rich in anthocyanins include the blueberry, raspberry, black rice, and black soybean, among many others that are red, blue, purple, or black. Some of the colors of autumn leaves are derived from anthocyanins.

<span class="mw-page-title-main">Genetically modified bacteria</span> First organisms to be modified in the laboratory

Genetically modified bacteria were the first organisms to be modified in the laboratory, due to their simple genetics. These organisms are now used for several purposes, and are particularly important in producing large amounts of pure human proteins for use in medicine.

<span class="mw-page-title-main">Genetically modified rice</span>

Genetically modified rice are rice strains that have been genetically modified. Rice plants have been modified to increase micronutrients such as vitamin A, accelerate photosynthesis, tolerate herbicides, resist pests, increase grain size, generate nutrients, flavors or produce human proteins.

<span class="mw-page-title-main">Polygalacturonase</span>

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-1,4 glycosidic bonds between galacturonic acid residues:

<span class="mw-page-title-main">History of genetic engineering</span>

Genetic engineering is the science of manipulating genetic material of an organism. The first artificial genetic modification accomplished using biotechnology was transgenesis, the process of transferring genes from one organism to another, first accomplished by Herbert Boyer and Stanley Cohen in 1973. It was the result of a series of advancements in techniques that allowed the direct modification of the genome. Important advances included the discovery of restriction enzymes and DNA ligases, the ability to design plasmids and technologies like polymerase chain reaction and sequencing. Transformation of the DNA into a host organism was accomplished with the invention of biolistics, Agrobacterium-mediated recombination and microinjection. The first genetically modified animal was a mouse created in 1974 by Rudolf Jaenisch. In 1976 the technology was commercialised, with the advent of genetically modified bacteria that produced somatostatin, followed by insulin in 1978. In 1983 an antibiotic resistant gene was inserted into tobacco, leading to the first genetically engineered plant. Advances followed that allowed scientists to manipulate and add genes to a variety of different organisms and induce a range of different effects. Plants were first commercialized with virus resistant tobacco released in China in 1992. The first genetically modified food was the Flavr Savr tomato marketed in 1994. By 2010, 29 countries had planted commercialized biotech crops. In 2000 a paper published in Science introduced golden rice, the first food developed with increased nutrient value.

<span class="mw-page-title-main">Cry1Ac</span> Crystal protein

Cry1Ac protoxin is a crystal protein produced by the gram-positive bacterium, Bacillus thuringiensis (Bt) during sporulation. Cry1Ac is one of the delta endotoxins produced by this bacterium which act as insecticides. Because of this, the genes for these have been introduced into commercially important crops by genetic engineering in order to confer pest resistance on those plants.

<span class="mw-page-title-main">Cry6Aa</span>

Cry6Aa is a toxic crystal protein generated by the bacterial family Bacillus thuringiensis during sporulation. This protein is a member of the alpha pore forming toxins family, which gives it insecticidal qualities advantageous in agricultural pest control. Each Cry protein has some level of target specificity; Cry6Aa has specific toxic action against coleopteran insects and nematodes. The corresponding B. thuringiensis gene, cry6aa, is located on bacterial plasmids. Along with several other Cry protein genes, cry6aa can be genetically recombined in Bt corn and Bt cotton so the plants produce specific toxins. Insects are developing resistance to the most commonly inserted proteins like Cry1Ac. Since Cry6Aa proteins function differently than other Cry proteins, they are combined with other proteins to decrease the development of pest resistance. Recent studies suggest this protein functions better in combination with other virulence factors such as other Cry proteins and metalloproteinases.>

<span class="mw-page-title-main">Ethylene (plant hormone)</span> Alkene gas naturally regulating the plant growth

Ethylene (CH
2=CH
2) is an unsaturated hydrocarbon gas (alkene) acting as a naturally occurring plant hormone. It is the simplest alkene gas and is the first gas known to act as hormone. It acts at trace levels throughout the life of the plant by stimulating or regulating the ripening of fruit, the opening of flowers, the abscission (or shedding) of leaves and, in aquatic and semi-aquatic species, promoting the 'escape' from submergence by means of rapid elongation of stems or leaves. This escape response is particularly important in rice farming. Commercial fruit-ripening rooms use "catalytic generators" to make ethylene gas from a liquid supply of ethanol. Typically, a gassing level of 500 to 2,000 ppm is used, for 24 to 48 hours. Care must be taken to control carbon dioxide levels in ripening rooms when gassing, as high temperature ripening (20 °C; 68 °F) has been seen to produce CO2 levels of 10% in 24 hours.

References

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