Skip to main content
. 2012 Jul 1;7(7):827–846. doi: 10.4161/psb.20321

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