Tomatoes originated in the Andean region of South America under extremely
variable climatic conditions. Wild relatives of tomato grow from sea level to subalpine
elevations with some ecotypes adapted to flooded conditions and others to
extreme drought. Domestication of tomato led to its cultivation as a crop on all
continents and traits have been selected to promote abundant production of fruit.
Selective breeding from the narrow genetic base of domesticated tomato as well
as the introduction of exotic germplasm from the numerous wild relatives of
tomato have developed tomato plants producing high-quality fruit for fresh
consumption as well as for processed, prepared and stored products valued at
approximately US$5 billion annually. Advances in agricultural biotechnology
recently have provided the opportunity to expand the genetic resources available
for tomato improvement. The goals of tomato genetic engineering have been to
protect the tomato crop from environmental and biological assaults, and to
improve the quality of tomato fruit in order to deliver greater value in processed
tomato products or more healthful and attractive fresh fruit.
Tomato fruit are a significant source of nutrition for substantial portions of
the world’s human population because this vegetable crop is widely cultivated
and consumed extensively as both a fresh vegetable and concentrated processed
products. Tomatoes are rich sources of vitamins, especially ascorbic acid and β-carotene, and antioxidants such as lycopene. A single small tomato is sufficient
to supply about a quarter of the vitamins A and C recommended for humans to
consume daily (Hamner and Maynard, 1942; Beecher, 1998). Most of the
nutritional components in tomato fruit are stabilized by the acid pH of the fruit
tissue and many of the human nutrients are conserved during the relatively short and mild processing used in the preparation of most tomato food products.
Tomatoes are grown in industrial quantities in many temperate locations, but the
stability of the concentrated processed product has made it possible to transport
tomato products widely and to prolong the storage of tomato products.
Table 8.1 Transgenic tomato modifications
Trait
Gene
Regulation
Expression
Fruit ripening
Ethylene reduction
Bacterial ACC deaminase
Constitutive
Expression
Ethylene reduction
Phage SAMase
Fruit specific
Expression
Ethylene reduction
Tomato ACC synthase
Constitutive
Antisense suppression
Ethylene reduction
Tomato ACC synthase
Constitutive
Sense suppression
Ethylene reduction
Tomato ACC oxidase
Constitutive
Antisense suppression
Fruit softening
Tomato fruit PG
Constitutive
Antisense suppression
Fruit softening
Tomato fruit PG
Constitutive
Sense suppression
Fruit softening
Tomato fruit PME
Constitutive
Antisense suppression
Fruit softening
Tomato fruit PG and PE
Constitutive
Antisense suppression
Fruit softening
Tomato fruit expansin
Constitutive
Sense suppression
Fruit abscission
Tomato fruit Cel1 and Cel2
Constitutive
Antisense suppression
Fruit composition
Sucrose accumulation,
Tomato fruit invertase
Constitutive
Antisense suppression
Solids content
Constitutive
Expression
Starch accumulation
Arabidopsis sucrose synthase
Constitutive
Expression
Fatty acid and flavor content
Constitutive
Expression
Color
Tomato phytoene synthase
Constitutive
Antisense suppression
Parthenocarpic
Bacterial tryptophan monoxygenase
Constitutive
Expression
Seeds
Increased dormancy
Tomato NCED
Constitutive
Expression
Decreased dormancy
Arabidopsis abi-1
Constitutive
Expression
Pathogen and pest resistance and tolerance
TMV
TMV N
Constitutive
Expression
CMV
Cucumber mosaic virus CP
Constitutive
Expression
TSWV
Tomato spotted wilt virus N
Constitutive
Expression
PhMV
Physalis mottle tymovirus CP
Constitutive
Expression
Pseudomonas syringae pv
tomato
Tomato Pto
Constitutive
Expression
Xanthomonas campestris pv. Vesicatoria
Pepper Bs2
Constitutive
Expression
Cladosporium fulvum
Tomato Cf 9
Constitutive
Expression
Verticillium dahliae
Tomato chitinase
Constitutive
Expression
Fusarium oxysporum f.sp. lycopersici
Tobacco chitinase and Β 1,3
- glucanase
Constitutive
Expression
Trichoderma hamatum Xanthomonas campestris
Rubber tree hevein
Constitutive
Expression
pv. vesicatoria
Tamoto LeETR4
Constitutive
Expression
Sclerotinia sclerotiorum
Collybia velutipes oxalate decarboxylase
Constitutive
Expression
Botrytis cinerea
Pear fruit PGIP
Constitutive
Expression
Phytophthora infestans
Grape reserveratrol
Constitutive
Expression
Manduca Sexta
Tomato prosystemin
Insect induced
Expression
Insect resistance
Bt toxins
Constitutive
Expression
Insect resistance
Bt cry1Ac
Constitutive
Expression
Nematode resistance
Rice cystatinc-I
Constitutive
Expression
Root knot nematode, aphid, viral resistance
TomatoMi
Root-specific
Expression
Plant defense responses
Extracellular responses
Agrobacterium ipt
Constitutive
Expression
Environmental stresses
Salt stress
Yeast HAL2
Constitutive
Expression
Drought
Arabidopsis ABI-1
Constitutive
Expression
Chilling and oxidative stress sensitivity
Tomato catalase
Constitutive
Antisense suppression
Heavy metal tolerance
Bacterial ACC deaminase
Root-specific or stress induced
Expression
Herbicide resistance
Thiazopyr resistance
Rabbit liver esterase
Constitutive
Expression
Quinclorac resistance
Tomato ACC synthase
Constitutive
Antisense suppression
Fenthion (insecticide) sensitivity
Tamoto Prf
Constitutive
Antisense suppression
Metabolic modifications
Increased sucrose unloading
Sucrose phosphate synthase
Root-specific or fruit-specific
Expression
Foliage coloration
Increased anthcyanin
Antirrhinum del
Constitutive
Expression
Floral patterns
Indeterminate flowering
Tomataogamous
Constitutive
Expression
Precocious termination
Tomataogamous
Constitutive
Antisense suppression
Tomato was one of the first plants to be transformed by Agro bacteriumtumefaciens and regenerated into fertile, productive plants (Fillatti et al., 1987).
The success of early work to obtain transgenic plants allowed for the first commercial
release of a transgenic food product, the Flavr Savr tomato, with extended
shelf life of the ripe fruit. The transformation of a large number of tomato varieties
has been reported, suggesting that essentially any variety is amenable to genetic
transformation. For example, fresh market varieties (Moneymaker, Better Boy),
greenhouse varieties (Ailsa Craig), small-fruited fresh varieties (VFNT Cherry)
and processing varieties (UC82b) as well as wild tomato relative (L. schilense (Agharbaouiet al., 1995), L. peruvanium (Rudas et al., 1997) and L. hirsutum (Smith et al., 1996)) have all been transformed in academic and commercial
research laboratories. Most of the successful transformation protocols for tomato
utilize Agro bacteria to deliver transgenes to the hypocotyl sections of newly
germinated seedlings, but biolistic approaches also have been utilized. The
success of the floral dip methods used in Arabidopsis has not been reported for
tomato. Antibiotic resistance of transformed tissues is frequently the preferred
method of selection of transgenic tissues, because of its historical success.
However, public concerns about the contents of genetically modified food
products will undoubtedly lead to the utilization of new selection methods,
including positive selection for growth on selective media (Haldrupet al., 1998).
Because fruit are the economically significant crop from tomato plants, many
transgenic modifications have targeted the fruit ripening processes to develop
products that better withstand harvest, handling, transportation and storage
practices utilized in commercial distribution. To reduce processing costs and
effectively increase processing yield, transgenic fruit have been developed with
increased solids content. To provide novel value-added products, tomato fruit
have also been engineered to produce increased components of nutritional value
and to produce pharmaceutical compounds. To enhance production efficiency
and yield, tomato plants have been engineered for resistance to herbicides,
extreme temperatures and pathogens by the transgenic expression of foreign
genes not accessible by classical breeding methods. A comprehensive listing of
transgenic tomato modifications that have successfully altered aspects of plant
growth, morphology and cultivation are summarized in Table 8.1.