How do plants produce ethylene




















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A dominant mutant receptor from Arabidopsis confers ethylene insensitivity in heterologous plants. Nat Biotechnol. Ethylene signaling: simple ligand, complex regulation. Curr Opin Plant Biol. Download references. The author thanks Dr. Bram Van de Poel and members of the Chang lab for comments on the manuscript. You can also search for this author in PubMed Google Scholar. Correspondence to Caren Chang. Reprints and Permissions. Chang, C. BMC Biol 14, 7 Download citation.

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Skip to main content. Search all BMC articles Search. Download PDF. Abstract Ethylene gas is a major plant hormone that influences diverse processes in plant growth, development and stress responses throughout the plant life cycle. What is the plant hormone ethylene?

Why is ethylene important to agriculture? How was the ethylene hormone discovered? Is there anything different about a gaseous hormone? How do plants synthesize ethylene? Full size image. How much ethylene is needed to trigger a response in plants?

Is ethylene harmful to humans? How was the ethylene-signaling pathway determined? Using classical methods of differential screening of ripe fruit cDNA banks, several genes whose expression is regulated by ethylene or induced during ripening were isolated and cDNA clones corresponding to genes strongly expressed during tomato ripening were isolated Mansson et al.

To study ethylene-associated events during ripening, cDNAs corresponding to the mRNAs of ethylene-regulated genes have been isolated and characterized Mansson et al.

Subsequently, other important genes expressed during ripening were isolated, identified and characterized, such as those encoding polygalacturonase PG and pectin methyltransferase Smith et al. Many natural tomato mutants affected at ripening have been used to study gene expression and regulation.

Two principal groups of mutants were used, i. Treatment of the rin mutant with ethylene allowed transcription of genes corresponding to some of these mRNAs, indicating that the rin -type fruits possess the capacity to respond to ethylene Lincoln and Fischer, ; Knapp et al.

This suggests either the absence of a specific receptor to ripening in the transduction pathway or a deficiency in the ethylene signal transduction cascade, linking perception of this phytohormone to the ripening responses.

Dellapenna et al. To identify the function of genes and their role in the ripening process, an antisense RNA strategy has been used by several research groups and several transgenic plants showing reduced expression of ripening related genes have been obtained Gray et al. Transgenic tomato plants expressing an antisense polygalacturonase gene showed a reduction in PG transcripts as well in enzymatic activity during ripening and it was has been shown that in fruits with antisense PG the degradation of cellular wall pectins was inhibited but other aspects of maturation, such as ethylene production and lycopene accumulation were not affected Smith et al.

Other transgenic plants, genetically modified to get alter ethylene production have been obtained by various authors. Fruits from plants transformed with an antisense ACC oxidase clone showed a strong reduction in ethylene production, delayed ripening and a considerable increase in conservation period Hamilton et al. Melon, genetically transformed using a melon antisense ACCO clone resulted in a considerable reduction in fruit ethylene synthesis and, unlike wild type fruits, they showed no increase in ethylene production during peak respiration climacteric crisis either when attached to the plants or post-harvest Ayub et al.

These melons showed an inhibition of maturation as indicated by the absence of yellowish of peel and much reduced softening coupled with a high sugar concentration produced as a result of the prolonged ripening time Ayub et al.

In all cases, ripening of transgenic fruits can be restored by the application of exogenous ethylene Zegzouti, To block ethylene biosynthesis an antisense ACCS clone which completely blocked ripening was introduced into tomato Oeller et al.

Other tomato models involved a modification of the ethylene biosynthetic pathway by the over-expression of a bacterial ACC deaminase Klee et al. The genetic manipulation of alcohol dehydrogenase levels in ripening tomato fruit have been shown to affect the balance of some flavor aldehydes and alcohols Speirs et al.

Much information concerning the biochemical components of the ethylene perception and signal transduction pathways of ethylene have been obtained during the past decade through the development of molecular and genetic strategies using A.

The components of ethylene signal transduction cascade are described below. The process of ethylene perception starts when this molecule interacts with a receptor linked to the endoplasmic reticulum ER membrane Giovannoni, ; Stepanova and Alonso, The pleiotropic effects of the etr1 mutation in A.

As a matter of fact, the ETR1 gene from A. This gene encodes a protein whose N-terminal hydrophobic end forms a dimmer linked by an S-S bond Shaller and Bleecker, responsible for its location in the membrane. The C-terminal end shows high homology with the histidine-kinase family proteins implicated in signal transduction in prokaryotes known as the two-component system Johnson and Ecker, The expression of the coding region of this gene in yeasts showed that the ETR1 protein binds to the plasma membrane as a dimmer and is also capable of binding ethylene and that the ETR1 protein acts as an ethylene receptor Shaller and Bleecker, , while Rodriguez et al.

The five members of the ETR family are related on the basis of the common structural elements in the protein. Additionally, specific substitutions of amino acids in the ethylene binding domain in all members confer dominant insensibility to this hormone in whole plants. There are strong indications that ethylene receptors act as negative regulators of the ethylene signal transduction pathway and that they are activated in the absence of ethylene acting direct or indirectly in the activation of a cascade downstream component, denominated the CTR1 constitutive-triple-response Giovannoni, ; Stepanova and Alonso, Studies shown that transcription factors are part of the ethylene signal transduction pathway, with the cloning of the EIN3 gene encoding a nuclear protein providing the first direct evidence of nuclear regulation in this transduction pathway Chao et al.

In Arabidopsis the expression of the ERF1 gene is rapidly induced by treatment with ethylene Solano, et al. Another relevant fact is that EIN3 homodimmers are capable of having in vitro interactions with a promoter element in the ERF1 gene. This indicates that ERF1 is part of the primary signal transduction cascade and is downstream of the previously identified components, suggesting that a transcriptional cascade operates in ethylene signaling Bleecker and Kende, , Giovannoni, ; Stepanova and Alonso, Emerging genomics tools including expressed sequence tags ESTs and expression arrays are also likely to accelerate the discovery of homologous genes from additional species and the identification of additional novel ripening regulators Adams-Phillips et al.

Several theories on ethylene signal perception and transduction have been proposed to explain the mechanisms by which ethylene receptors could promote signal transduction through a cascade involving several components Zarembinski and Theologis, ; Ecker, ; Bleecker and Kende, The model recently proposed by Bleecker and Kende and subsequently reviewed by Giovannoni and Stepanova and Alonso places the components of the ethylene signal transduction pathway in a linear array and defends the theory that ethylene negatively regulates the joint binding of ETR1 and CTR1 to the receptor, resulting in de-repression of response pathways Figure 2.

The order of the components in this hypothetical linear chain is based on the analysis of epistatic genes, gene expression studies and the study of biochemical interactions.

In this model, ethylene negatively regulates the family of receptors associated with the endoplasmic reticulum membrane and which are related to the two-component catalytic bacterial receptor family. The histidine-kinase transmitter domains of members of this receptor family interact with the CTR1 Raf-like kinase regulator domain. A target for the EIN3 transcription factors is the ERF1 gene promoter , which is a member of a second family of transcription factors and is rapidly induced in response to ethylene and is capable of activating a set of responses to ethylene when expressed.

The large diversity of gene types is representative of the multitude of events affected by ethylene during fruit ripening. Ethylene receptor genes and components of the ethylene transduction pathway have been discovered in the recent years, as described above.

However, the number of genes demonstrated to be induced through this pathway is low in regard to the variety of physiological responses of plants to ethylene. For this reason, novel early ethylene-regulated ER genes from late immature green tomato fruit have been isolated using the differential display technique, in order to obtain a broader understanding of the molecular basis by which ethylene coordinates the ripening process Zegzouti et al.

A large set of clones have been isolated, showing homologous genes involved in transcriptional and post-transcriptional regulation, signal transduction components, stress-related proteins and primary metabolism. However, as yet a number of these ER clones have no assigned function and reverse genetics is currently being used to investigate the function of these genes and address their role in the ripening process Pech et al.

The latest data have indicated that ER50 is a CTR-like clone, potentially involved in the ethylene transduction pathway Leclercq et al. These genes are being extensively studied in order to determine their function and mechanisms of regulation by ethylene.



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