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Plant membrane lipids are primarily composed of 16-carbon and 18-carbon fatty acids containing up to three double bonds. By contrast, the seed oils of many plant species contain fatty acids with significantly different structures. These unusual fatty acids sometimes accumulate to >90% of the total fatty acid content in the seed triacylglycerols, but are generally excluded from the membrane lipids of the plant, including those of the seed. The reasons for their exclusion and the mechanisms by which this is achieved are not completely understood. Here we discuss recent research that has given new insights into how plants prevent the accumulation of unusual fatty acids in membrane lipids, and how strict this censorship of membrane composition is. We also describe a transgenic experiment that resulted in an excessive buildup of unusual fatty acids in cellular membranes, and clearly illustrated that the control of membrane lipid composition is essential for normal plant growth and development.

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Land plants secrete a layer of wax onto their aerial surfaces that is essential for survival in a terrestrial environment. This wax is composed of long-chain, aliphatic hydrocarbons derived from very-long-chain fatty acids (VLCFAs). Using the Arabidopsis expressed sequence tag database, we have identified a gene, designated CUT1, that encodes a VLCFA condensing enzyme required for cuticular wax production. Sense suppression of CUT1 in transgenic Arabidopsis plants results in waxless (eceriferum) stems and siliques as well as conditional male sterility. Scanning electron microscopy revealed that this was a severe waxless phenotype, because stems of CUT1-suppressed plants were completely devoid of wax crystals. Furthermore, chemical analyses of waxless plants demonstrated that the stem wax load was reduced to 6 to 7% of wild-type levels. This value is lower than that reported for any of the known eceriferum mutants. The severe waxless phenotype resulted from the downregulation of both the decarbonylation and acyl reduction wax biosynthetic pathways. This result indicates that CUT1 is involved in the production of VLCFA precursors used for the synthesis of all stem wax components in Arabidopsis. In CUT1-suppressed plants, the C24 chain-length wax components predominate, suggesting that CUT1 is required for elongation of C24 VLCFAs. The unique wax composition of CUT1-suppressed plants together with the fact that the location of CUT1 on the genetic map did not coincide with any of the known ECERIFERUM loci suggest that we have identified a novel gene involved in wax biosynthesis. CUT1 is currently the only known gene with a clearly established function in wax production.

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The fatty-acyl composition of the seed oil was determined for 100 ecotypes of Arabidopsis thaliana. Despite coming from diverse geographical locations, seed fatty-acyl profiles of all ecotypes were remarkably similar. They contained identical fatty acids, including the characteristic C20 and C22 very-long-chain fatty acids (VLCFAs). The total proportions of seed VLCFA varied between 22% and 35% w/w of the total seed fatty acid content.

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Transgenic Arabidopsis plants overexpressing the Arabidopsis FATTY ACID ELONGATION1 gene under the control of the 35S promoter from cauliflower mosaic virus accumulated very-long-chain fatty acids (VLCFAs) throughout the plant. In some transformants, C20 and C22 VLCFAs accounted for >30% of the total fatty acids, accumulating at the expense of C16 and C18 fatty acids. These C20 and C22 fatty acids were incorporated into all of the major membrane glycerolipid classes. Plants with a high VLCFA content displayed a dramatically altered morphology, which included the failure of flowering shoots to elongate, a modified spatial pattern of siliques, an altered floral phenotype, and a large accumulation of anthocyanins. In addition, these plants also exhibited a unique alteration of the chloroplast membrane structure. We discuss a possible role for VLCFAs in establishing the shape/curvature of the membranes, which in turn may affect the shape of the cell and ultimately that of the whole plant.

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The Arabidopsis FATTY ACID ELONGATION1 (FAE1) gene encodes a putative seed-specific condensing enzyme. It is the first of four enzyme activities that comprise the microsomal fatty acid elongase (FAE) involved in the biosynthesis of very-long-chain fatty acids (VLCFAs). FAE1 has been expressed in yeast and in tissues of Arabidopsis and tobacco, where significant quantities of VLCFAs are not found. The introduction of FAE1 alone in these systems is sufficient for the production of VLCFAs, for wherever FAE1 was expressed, VLCFAs accumulated. These results indicate that FAE1 is the rate-limiting enzyme for VLCFA biosynthesis in Arabidopsis seed, because introduction of extra copies of FAE1 resulted in higher levels of the VLCFAs. Furthermore, the condensing enzyme is the activity of the elongase that determines the acyl chain length of the VLCFAs produced. In contrast, it appears that the other three enzyme activities of the elongase are found ubiquitously throughout the plant, are not rate-limiting and play no role in the control of VLCFA synthesis. The ability of yeast containing FAE1 to synthesize VLCFAs suggests that the expression and the acyl chain length specificity of the condensing enzyme, along with the apparent broad specificities of the other three FAE activities, may be a universal eukaryotic mechanism for regulating the amounts and acyl chain length of VLCFAs synthesized.

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We report here the initial characterization of the alcohol dehydrogenase (Adh) gene family of the allotetraploid Gossypium hirsutum, a crop plant that is highly sensitive to waterlogging. Twelve Adh cDNAs were isolated from a library constructed from RNA prepared from anaerobically stressed root tips. Nine of the twelve cDNAs fell into one class, while each of the other three cDNAs fell into a separate class. The 3'-untranslated regions had little or no homology between classes implying that each of these four classes is encoded by a different gene. The most abundant class of Adh cDNA was expressed under the control of the 35S promoter in transgenic cotton and encodes the ADH2 isozyme, the isozyme induced most strongly by low oxygen conditions. Cotton Adh genomic segments were sequenced. The promoter regions of these Adh genes contain the cis-acting motifs which have been shown in other plant species to be necessary for anaerobic induction of transcription. A gene conversion event is likely to have occurred between the 3' ends of two of the genes. Sequence comparison of the Adh genomic and cDNA clones, and Southern analysis of genomic DNA suggest the existence of multiple copies of Adh genes in cotton. Five different members of this Adh gene family of cotton have been identified, of which four are very similar.

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The alcohol dehydrogenase (ADH) system in cotton is characterized, with an emphasis on the cultivated allotetraploid species Gossypium hirsutum cv. Siokra. A high level of ADH activity is present in seed of Siokra but quickly declines during germination. When exposed to anaerobic stress the level of ADH activity can be induced several fold in both roots and shoots of seedlings. Unlike maize and Arabidopsis, ADH activity can be anaerobically induced in mature green leaves. Three major ADH isozymes were resolved in Siokra, and it is proposed that two genes, Adh1 and Adh2, are coding for these three isozymes. The genes are differentially expressed. ADH1 is predominant in seed and aerobically grown roots, while ADH2 is prominent in roots only after anaerobic stress. Biochemical analysis demonstrated that the ADH enzyme has a native molecular weight of approximately 81 kD and a subunit molecular weight of approximately 42 kD, thus establishing that ADH in cotton is able to form and is active as dimers. Comparisons of ADH activity levels and isozyme patterns between Siokra and other allotetraploid cottons showed that the ADH system is highly conserved among these varieties. In contrast, the diploid species of cotton all had unique isozyme patterns.

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Leaf water potential, leaf relative water content, and relative transpiration of barley were determined daily under greenhouse conditions at 3 growth stages: tillering to boot, boot to heading, and heading to maturity. The leaf moisture characteristic curve (relative water content versus leaf water potential) was the same for leaves of the same age growing in the same environment for the first 2 stages of growth, but shifted at the heading to maturity stage to higher leaf relative water content for a given leaf water potential. Growth chamber experiments showed that the leaf moisture characteristic curve was not the same for plants growing in different environments.Relative transpiration data indicated that barley stomates closed at a water potential of about -22 bars at the 3 stages studied.The water potential was measured for all the leaves on barley to determine the variation of water potential with leaf position. Leaf water potential increased basipetally with plant leaf position. In soil with a moisture content near field capacity a difference of about 16.5 bars was observed between the top and bottom leaves on the same plant, while in soil with a moisture content near the permanent wilting point the difference was only 5.6 bars between the same leaf positions.