Role of ethylene on various ripening pathways and on the development of sensory quality of Charentais cantaloupe melons

Charentais melons (Cucumis melo L., var cantalupensis Naud.) in which ethylene biosynthesis has been suppressed by an antisense ACC oxidase gene have been used to better understand the role of ethylene in the regulation of the ripening process of climacteric fruit and on the development of sensory qualities. We have shown that a number of biochemical and molecular processes associated with the ripening of climacteric fruit are ethylene-independent. In some cases, such as softening of the flesh, the same pathway comprises both ethylene-dependent and -independent components. The various ethylene-dependent events exhibit differential sensitivity to ethylene. The threshold level for degreening of the rind is 1 ppm, while 2.5 ppm are required to trigger the ethylene-dependent component of the softening process. The saturating level of ethylene for all these events is less than 5 ppm, which is by far lower than the internal ethylene concentrations found in the fruit at the climacteric peak (around 100 ppm). Detachment of the fruit influences the development of respiratory climacteric. Fruit remaining attached to the vine, although producing higher levels of ethylene, exhibit a reduced climacteric rise in respiration as compared to detached fruit. The response of antisense ACO fruit to exogenous ethylene in terms of respiration is higher in detached than in attached fruit. Ethylene-suppressed melons show a severe reduction of aroma volatiles production, particularly in ester production. In the biosynthetic pathway of aliphatic esters, the dehydrogenation of fatty acids and aldehydes appears to be ethylene-dependent. In contrast, alcohol acetylation comprises ethylene-dependent and ethylene-independent components, probably corresponding to differentially regulated alcohol acetyl transferases. In terms of sensory quality, these data show that the extension of shelf-life through the inhibition of ethylene production has some beneficial effects on texture and sugar accumulation but is detrimental for the generation of aroma

The pivotal role of ethylene in plant growth

Being continuously exposed to variable environmental conditions, plants produce phytohormones to react quickly and specifically to these changes. The phytohormone ethylene is produced in response to multiple stresses. While the role of ethylene in defense responses to pathogens is widely recognized, recent studies in arabidopsis and crop species highlight an emerging key role for ethylene in the regulation of organ growth and yield under abiotic stress. Molecular connections between ethylene and growth-regulatory pathways have been uncovered, and altering the expression of ethylene response factors (ERFs) provides a new strategy for targeted ethylene-response engineering. Crops with optimized ethylene responses show improved growth in the field, opening new windows for future crop improvement. This review focuses on how ethylene regulates shoot growth, with an emphasis on leaves

Recent Developments on the Role of Ethylene in the Ripening of Climacteric Fruit

It has long been recognised that ethylene plays a major role in the ripening process of climacteric fruit. A more thorough analysis, however, has revealed that a number of biochemical and molecular processes associated with climacteric fruit ripening are ethylene-independent. One of the crucial steps of the onset of ripening is the induction of autocatalytic ethylene production. In ethylene-suppressed melons, ACC synthase activity is induced at the same time as in control melons, indicating that ACC biosynthesis during the early stages of ripening seems to be a developmentally-regulated (ethylene-independent) process. The various ripening events exhibit differential sensitivity to ethylene. For instance, the threshold level for degreening of the rind is 1ppm, while 2.5 ppm are required to trigger some components of the softening process. The saturating level of ethylene producing maximum effects is less than 5 ppm, which is by far lower than the internal ethylene concentrations found in the fruit at the climacteric peak (over 100 ppm). In many fruit chilling temperatures hasten ethylene production and ripening and in some late season pear varieties, exposure to chilling temperatures is even absolutely required for the attainment of the capacity to synthesize autocatalytic ethylene. This is correlated with the stimulation of expression of ACC oxidase and of members of the ACC synthase gene family. Ethylene operates via a perception and transduction pathway to induce the expression of genes responsible for the biochemical and physiological changes observed during ripening. However, only a few genes induced via the ethylene transduction pathway have been described so far. We have used a differential display method to isolate novel ethylene-reponsive (ER) cDNA clones of tomato that potentially play a role in propagating the ethylene response and in regulating fruit ripening. Collectively, these data permit a general scheme of the molecular mechanisms of fruit ripening to be proposed

The plant hormone ethylene restricts Arabidopsis growth via the epidermis

The gaseous hormone ethylene plays a key role in plant growth and development, and it is a major regulator of stress responses. It inhibits vegetative growth by restricting cell elongation, mainly through cross-talk with auxins. However, it remains unknown whether ethylene controls growth throughout all plant tissues or whether its signaling is confined to specific cell types. We employed a targeted expression approach to map the tissue site(s) of ethylene growth regulation. The ubiquitin E3 ligase complex containing Skp1, Cullin1, and the F-box protein EBF1 or EBF2 (SCFEBF1/2) target the degradation of EIN3, the master transcription factor in ethylene signaling. We coupled EBF1 and EBF2 to a number of cell type-specific promoters. Using phenotypic assays for ethylene response and mutant complementation, we revealed that the epidermis is the main site of ethylene action controlling plant growth in both roots and shoots. Suppression of ethylene signaling in the epidermis of the constitutive ethylene signaling mutant ctr1-1 was sufficient to rescue the mutant phenotype, pointing to the epidermis as a key cell type required for ethylene-mediated growth inhibition

Cadmium-induced ethylene production and responses in Arabidopsis thaliana rely on ACS2 and ACS6 gene expression

Background: Anthropogenic activities cause metal pollution worldwide. Plants can absorb and accumulate these metals through their root system, inducing stress as a result of excess metal concentrations inside the plant. Ethylene is a regulator of multiple plant processes, and is affected by many biotic and abiotic stresses. Increased ethylene levels have been observed after exposure to excess metals but it remains unclear how the increased ethylene levels are achieved at the molecular level. In this study, the effects of cadmium (Cd) exposure on the production of ethylene and its precursor 1-aminocyclopropane-1-carboxylic acid (ACC), and on the expression of the ACC Synthase (ACS) and ACC Oxidase (ACO) multigene families were investigated in Arabidopsis thaliana. Results: Increased ethylene release after Cd exposure was directly measurable in a system using rockwool-cultivated plants; enhanced levels of the ethylene precursor ACC together with higher mRNA levels of ethylene responsive genes: ACO2, ETR2 and ERF1 also indicated increased ethylene production in hydroponic culture. Regarding underlying mechanisms, it was found that the transcript levels of ACO2 and ACO4, the most abundantly expressed members of the ACO multigene family, were increased upon Cd exposure. ACC synthesis is the rate-limiting step in ethylene biosynthesis, and transcript levels of both ACS2 and ACS6 showed the highest increase and became the most abundant isoforms after Cd exposure, suggesting their importance in the Cd-induced increase of ethylene production. Conclusions: Cadmium induced the biosynthesis of ACC and ethylene in Arabidopsis thaliana plants mainly via the increased expression of ACS2 and ACS6. This was confirmed in the acs2-1acs6-1 double knockout mutants, which showed a decreased ethylene production, positively affecting leaf biomass and resulting in a delayed induction of ethylene responsive gene expressions without significant differences in Cd contents between wild-type and mutant plants

Climacteric fruit ripening: Ethylene-dependent and independent regulation of ripening pathways in melon fruit

Cantaloupe melons have a typical climacteric behaviour with ethylene playing a major role in the regulation of the ripening process and affecting the ripening rate. Crossing of Cantaloupe Charentais melon with a non-climacteric melon indicated that the climacteric character is genetically dominant and conferred by two duplicated loci only. However, other experiments made by crossing two non-climacteric melons have generated climacteric fruit, indicating that different and complex genetic regulation exists for the climacteric character. Suppression of ethylene production by antisense ACC oxidase RNA in Charentais melon has shown that, while many ripening pathways were regulated by ethylene (synthesis of aroma volatiles, respiratory climacteric and degreening of the rind), some were ethylene-independent (initiation of climacteric, sugar accumulation, loss of acidity and coloration of the pulp). Softening of the flesh comprised both ethylene-dependent and independent components that were correlated with differential regulation of cell wall degrading genes. These results indicate that climacteric (ethylene-dependent) and non-climacteric (ethylene-independent) regulation coexist during climacteric fruit ripening. In addition, ethylenesuppressed melons allowed demonstrating that the various ethylene-dependent events exhibited differential sensitivity to ethylene and that ethylene was promoting sensitivity to chilling injury. Throughout this review, the data generated with melon are compared with those obtained with tomato and other fruit

A dominant repressor version of the tomatoSl-ERF.B3gene confers ethylene hypersensitivity via feedback regulation of ethylene signaling and response components

Ethylene Response Factors (ERFs) are downstream components of the ethylene signal transduction pathway, although their role in ethylene-dependent developmental processes remains poorly understood. As the ethylene-inducible tomato Sl-ERF.B3 has been shown previously to display a strong binding affinity to GCC-box-containing promoters, its physiological significance was addressed here by a reverse genetics approach. However, classical up- and down-regulation strategies failed to give clear clues to its roles in planta, probably due to functional redundancy among ERF family members. Expression of a dominant repressor ERF.B3-SRDX version of Sl-ERF.B3 in the tomato resulted in pleiotropic ethylene responses and vegetative and reproductive growth phenotypes. The dominant repressor etiolated seedlings displayed partial constitutive ethylene response in the absence of ethylene and adult plants exhibited typical ethylene-related alterations such as leaf epinasty, premature flower senescence and accelerated fruit abscission. The multiple symptoms related to enhanced ethylene sensitivity correlated with the altered expression of ethylene biosynthesis and signaling genes and suggested the involvement of Sl-ERF.B3 in a feedback mechanism that regulates components of ethylene production and response. Moreover, Sl-ERF.B3 was shown to modulate the transcription of a set of ERFs and revealed the existence of a complex network interconnecting different ERF genes. Overall, the study indicated that Sl-ERF.B3 had a critical role in the regulation of multiple genes and identified a number of ERFs among its primary targets, consistent with the pleiotropic phenotypes displayed by the dominant repression lines

Effects of Elevated Substrate-Ethylene on Colonization of Leek (\u3cem\u3eAllium porrum\u3c/em\u3e) by the Arbuscular Mycorrhizal Fungus \u3cem\u3eGlomus aggregatum\u3c/em\u3e

There are very few studies of hormonal regulation of arbuscular mycorrhiza formation that include the gaseous hormone ethylene. Ethylene is considered inhibitory to the formation of arbuscular mycorrhizae; however, very low concentrations may promote their formation. We used an improved method of exogenous ethylene application to determine whether ethylene concentration dependent changes in colonization occur in the leek (Allium porrum L. cv. Giant Musselburgh) – Glomus aggregatum Schenck & Smith emend. Koske system. This improved method allowed for a continuous flow of constant concentration of the gas to be applied to a substrate. The 0.6 μL/L substrate–ethylene treatment reduced both root and leaf length and resulted in significantly lower arbuscular colonization compared with controls, whereas the 0.3 μL/L treatment reduced root length only and did not significantly affect colonization levels. Despite continuous application of exogenous ethylene, the amount of ethylene detected in inoculated substrates was reduced to near zero 20 days after inoculation. This decrease may be either due to an increased capacity for ethylene oxidation by arbuscular mycorrhizal roots or because arbuscular mycorrhizal fungi (or other microbes in the pot-cultured inoculum) are capable of metabolizing ethylene. The present study highlights the need for investigations into arbuscular mycorrhizal fungal physiology and the mechanisms by which ethylene regulates arbuscular mycorrhiza formation

Ethylene supports colonization of plant roots by the mutualistic fungus Piriformospora indica

The mutualistic basidiomycete Piriformospora indica colonizes roots of mono- and dicotyledonous plants, and thereby improves plant health and yield. Given the capability of P. indica to colonize a broad range of hosts, it must be anticipated that the fungus has evolved efficient strategies to overcome plant immunity and to establish a proper environment for nutrient acquisition and reproduction. Global gene expression studies in barley identified various ethylene synthesis and signaling components that were differentially regulated in P. indica-colonized roots. Based on these findings we examined the impact of ethylene in the symbiotic association. The data presented here suggest that P. indica induces ethylene synthesis in barley and Arabidopsis roots during colonization. Moreover, impaired ethylene signaling resulted in reduced root colonization, Arabidopsis mutants exhibiting constitutive ethylene signaling, -synthesis or ethylene-related defense were hyper-susceptible to P. indica. Our data suggest that ethylene signaling is required for symbiotic root colonization by P. indica

Ethylene seems required for the berry development and ripening in grape, a non-climacteric fruit

While the grape has been classified as a non-climacteric fruit whose ripening is thought to be ethylene independent, we show here that a transient increase of endogenous ethylene production occurs just before veraison (i.e. inception of ripening). We observed that ethylene perception, at this time, is required for at least the increase of berry diameter, the decrease of berry acidity and anthocyanin accumulation in the ripening berries; these latter experiments were performed with 1-methylcyclopropene, a specific inhibitor of ethylene receptors. The potential roles of ethylene in berry development and ripening are discussed