Table 1. Transgenic crops with altered phenotype, senescence and ripening due to overexpression and suppression of ethylene receptor and downstream signaling molecules.
Crop | Gene | Function of the Gene | Expression | Effect/Phenotype | Conclusion | References | |
---|---|---|---|---|---|---|---|
Tomato |
NR (Receptor gene) Semi Dominant |
Functionally redundant ethylene insensitive in all tissues. |
Antisense suppression |
Normal ripening as in wild type fruit. No effect on ethylene sensitivity, increased levels of LeETR4 mRNA. No effect on ethylene sensitivity of seedlings |
Not essential for ripening. Ethylene perception and signaling via the NR receptor is consistent with the receptor inhibition model. Functional compensation of LeETR4 for reduced NR expression |
Hackett et al., 2000172 Lanahan et al., 1994163 Tieman et al., 2000167 |
|
|
NR (Never Ripe) Semi Dominant |
Functionally redundant ethylene insensitive in all tissues. |
Overexpression in LeETR4 suppressed lines |
Eliminates ethylene sensitivity, lack of epinasty, flower senescence and normal fruit set, rescued the ethylene response phenotype. |
Proves the functional redundancy of the receptors. Negative Regulators of ethylene response. Only associated with ripening. |
Tieman et al., 2000167 |
|
|
Nr (Mutant NR gene) |
Unable to bind ethylene, delay senescence, imparts ethylene insensitivity. |
Antisense suppression |
Activated ethylene responses and onset of normal ripening, plant with two homozygous copies of transgene produced wild type fruit. Single gene in a homozygous state gave intermediate phenotype. |
Threshold level of mutant receptor is needed to suppress the ethylene response pathway. Support receptor inhibition model. |
Hackett et al., 2000172 Lanahan et al., 1994163 |
|
|
LeEIL(1-3) Transcription Factor |
Functionally redundant, positive regulator of ethylene signaling. |
Antisense suppression |
Single LeEIL suppression results no change. Multiple LeEIL gene suppression delays leaf epinasty, flower abscission, flower senescence. |
Normal levels of expression of the other two LeEILs can compensate for the loss of single gene. |
Tieman et al., 200123 |
|
|
LeEIL1 in tomato Nr mutant |
Functionally redundant, positive regulators of multiple ethylene receptors. |
Overexpresession in ethylene insensitive non ripening Nr mutant of tomato |
Partial restoration of ripening. Seedling triple response, upregulation of all ripening related genes were not restored. |
LeEIL1 may not have expressed at sufficiently high levels to restore these functions,LeEIL2, LeEIL3 are also required. |
Chen et al., 2004175 |
|
|
LeEIN2 |
Positive mediator of ethylene signaling and fruit development |
Suppression (VGIS) |
Delayed fruit development and ripening with fewer seeds and locules, suppressed ethylene inducible and ripening related genes, partially blocked ripening with large green parthenocarpic sector on fruit |
Probable involvement of LeEIN2 in a crosstalk between ethylene and auxin during fruit development |
Zhu et al., 2006174 |
|
|
LeETR4 |
Imparts ethylene in sensitivity, delay the onset of ripening until the seeds have matured. |
Suppression |
Increased ethylene sensitivity, fruits ripen prematurely flower senesce early without an increase in ethylene level. Severe epinastic curvature of petioles, no change in the levels of NR. |
Negative regulator of the ethylene signaling pathway, lowering the receptor levels increases sensitivity to ethylene. An important regulator of ripening |
Tieman et al., 2000167 |
|
|
LeETR4 using immature fruit specific Tfm7 promoter |
Delays onset of ripening in immature fruits |
RNAi mediated suppression |
Early ripening, total yield, fruit size and flavour related chemical composition had no negative effect. Leaves and other tissues were unaffected. |
Tissue specific approach to hasten fruit development without unwanted effects and enhanced ethylene response by ethylene receptor depletion. |
Kevany et al., 2008171 |
|
|
LeETR1 |
Functionally redundant |
Antisense expression of receiver domain and the 3′ UTR of LeETR1 |
Delayed abscission and shorter internodes with no effect on ethylene sensitivity, normal triple response, and reduced auxin movement. No effect on NR transcripts. |
Receiver domain of LeETR1 and LeETR2 has a role to play in signaling and can not be compensated by NR receptor |
Whitelaw et al., 2002176 |
|
|
etr1-1 (mutated ethylene receptors from Arabidopsis) |
Imparts dominant ethylene insensitivity |
Overexpression |
Long thin hypocotyls, normal root growth, more open and expended apical regions. Fruits with delayed ripening and senescence similar to Nr mutants |
Confers ethylene insensitivity in heterologous plants |
Wilkinson et al., 1997112 |
|
|
LeETR1 and LeETR2 |
Non redundant, plays role in ethylene perception |
Antisense suppression |
Shortened, thickened hypocotyls, enhanced abscission of petiole explants, faster senescence of flowers both in presence and absence of flowers. |
LeETR1 might have a more specific role to play in signaling, each member has a specific role to play and are not totaly redundant. |
Wang et al., 2003170 |
|
Carnation |
etr1-1 using CaMV 35S promoter |
Imparts global ethylene insensitivity |
overexpression |
Diminished ethylene insensitivity and diseased resistance with delayed senescence. No petal inrolling, firm petals. |
Heterologous expression imparts insensitivity in carnation |
Bovy et al., 199979 |
|
etr1-1 using flower specific fbp1 promoter |
Imparts global ethylene insensitivity |
overexpression |
Delayed flower senescence with increased disease resistance. Ethylene sensitivity increased in other parts of flower apart from petals, better seed germination and reduced leaf abscission. |
Flower specific promoter fixes the compromised disease resistance status |
Bovy et al., 199979 |
||
Petunia |
etr1-1/Nr |
Imparts global ethylene insensitivity |
overexpression |
Ethylene insensitivity with delayed flower senescence. Turgid corollas of flower, delayed abscission. |
Cross species transfer |
Wilkinson et al., 1997112 |
|
etr1-1 using flower binding protein from Petunia (fbp1) or apetala 3 promoter from Arabidopsis |
Imparts global ethylene insensitivity |
overexpression |
Diminished disease resistance and reduced adventitious root hair formation, increased ethylene insensitivity with delayed senescence, seedling root hair formation observed. 73% of fbp1 and 32% of ap3 plants showed more than 100% increase in flower life. |
Flower specific promoter enhances disease resistance in cross species transfer |
Cobb et al., 2002114 |
||
boers (mutated form of BOERS from Brassica oleracea) |
Imparts ethylene insensitivity |
overexpression |
Plants retained turgidity and pigmentation longer, larger flowers, extended flower longevity, diminished disease resistance with higher mortality, increased ethylene evolution and reduced ethylene sensitivity, self incompatible. |
boers is a dominant allele imparting insensitivity in the heterozygous transformed plants |
Shaw et al., 2002177 |
||
PhEIN2 (downstream signaling protein) |
Involved in regulation of accumulation of EIN3 and EIL1 proteins. Positive regulators of ethylene cascade |
Suppression by expressing sense RNA and RNAi mediated silencing |
Transgenic lines using sense RNA expression showed moderate delay in flower senescence and fruit ripening with no premature death reported. RNAi construct suppressed lines showed great delay in flower senescence and fruit ripening with great deal of premature death. Corollas were turgid even after the ovaries were turgid. |
Reduced or inhibited adventitious root formation (sense RNA expression) Reduced ethylene sensitivity (RNAi suppression) |
Shibuya et al. 2004119 |
||
Coriander |
AtERS1 (mutated dominant receptor) |
Confers ethylene insensitivity |
overexpression |
Delayed leaf senescence with prolonged total chlorophyll content in transgenic leaves, delayed flower senescence. |
Delayed senescence is not due to drastic reduction in ethylene biosynthesis rather it is due to the effect of dominant negative ERS1 transgene. Distantly related crop species can have extended shelf life exploiting heterologus gene transfer. |
Wang and Kumar 200491 |
|
Campanula |
etr1-1 with fbp1 |
Imparts global ethylene insensitivity |
Overexpressed |
Enhanced tolerance to exogenous ethylene and reduced ethylene sensitivity, stably passed to next generation of plants, weak expression of etr1-1 in leaves, intact fertility and no compromise on disease resistance. |
Sensitivity to ethylene in plants is independent of the copy number of the T-DNA. |
Srikandarajah et al., 200757 |
|
Nemesia |
CmETR1/H69A, missensed mutated ethylene receptor from Cucumis melo |
Imparts reduced ethylene sensitivity |
Overexpression |
Delayed flower senescence by 1--3 days and lesser root hair formation compared to non transgenic wild type. |
Artificially mutated ethylene receptor gene conferred reduced ethylene sensitivity in heterologus plants |
Cui et al., 2004109 |
|
Lettuce |
CmERS1/H70A |
Imparts stable sterlity, reduced fertility along with ethylene insensitivity |
Overexpressed |
Apart from reduced ethylene sensitivity, induced sterility in vegetatively propagated plants and pollen propagated plants |
This mutated receptor gene checks the pollen dispersal from transgenic plants and prohibit unwanted development of traits in other plants and weeds, can induce sterility in heterologus transgenic plants. |
Takada et al., 2007140 |
|
Chrysanthemum |
mDG-ERS1 (etr1-4), mutated nucleotide substitution corresponding to etr1-1. |
Imparts ethylene insensitivity |
Overexpressed |
Delayed leaf yellowing on exposure to exogenous ethylene and leaves on lower part of stem remain attached to it without yellowing, reduced ethylene sensitivity, detached shoots in darkness without ethylene treatment showed reduced senescence. |
Heterologus gene transformation remained unsuccessful; the mutated receptor could work for compositae family members. |
Narumi et al., 2005b85 Satoh et al., 2006, 2007, 200887,88,89 |
|
Cm-ETR1/H69A (missensed mutated ethylene receptor from Cucumis melo) |
Imparts ethylene insensitivity, male sterility, prevents transgene flow via pollen |
Overexpression |
Drastic reduction in number of pollen grains at temp 20--35 °C but observed at 10-15 C. Due to suppression of Cm-ETR1/H69A expression at low temperature and optimal growth temp for plant at 15--20 °C. Female fertility was also reduced. |
Optimal growth temperature for Chrysanthemum collides with its vegetative, reproductive growth resulting in mature pollens to flourish |
Shinoyama et al., 201290 |
||
Lotus |
Atetr1-1 |
Mutated dominant ethylene insensitive ethylene receptor |
Overexpression |
Delayed abscission and senescence of petals, ethylene insensitive, twisted coiled hypocotyls, no triple response, fewer lateral roots, 7-fold increased nodulation in highly ethylene insensitive transformants, 1.7-fold increases in bacteroid infection. |
Multiple roles of ethylene in nodule initiation by influencing root cell interactions and radial positioning, independent of autoregulation and nitrate inhibition of nodulation |
Lohar et al., 2009105 |
|
Cm-ERS1/H70A |
Inhibits ethylene sensitivity, promotes root nodulation. |
Overexpression |
Confers ethylene insensitivity and fixes the transgene in T3 generation, reduced ethylene sensitivity due to 1-ACC resistance, increased flowering |
Transgenic alteration of ethylene perception alters the rhizobial infection and nodulation phenotype. |
Nukui et al., 2004104 |
||
Cucumber | At-etr1-1 with CaMV 35S, AP3, CRC promoter | Imparts dominant ethylene insensitivity | Overexpression | Prevents carpel development resulting in male flowers with CaMV35S promoter. Increased femaleness and conversion of bisexual to female buds with CRC promoter. Exclusive male flower production with AP3 promoter. |
Heterologus ethylene insensitivity. Ethylene perception is required to promote femaleness in melon. Dual role of ethylene in sex expression and carpel maturation. Site for ethylene perception lies in stamen primordia and not in carpel primordial. | Little et al., 2007145 |