However, the mechanism of plant perception of free cutin monomers is currently unknown (Boller and Felix, 2009). Markus Riederer, Julius-von-Sachs-Institut für Biowissenschaften, Universität Würzburg, Würzburg, Germany. 3A), implying that the predominant structural motif must be a copolymer with an unknown polyhydroxy compound, presumably glycerol (Pollard et al., 2008). A, Schematic diagram highlighting the major structural features of the cuticle and underlying epidermal cell layer (not drawn to scale). Free primary alcohols can occur in the wax mixture, or they can be esterified to a fatty acid in order to form wax esters. 12 Plants. Needle browning and death in the flagged crown of Abies mariesii in the timberline ecotone of the alpine region in central Japan. Confocal laser scanning microscopy elucidation of the micromorphology of the leaf cuticle and analysis of its chemical composition. Studies assessing the impact of UV radiation on plant life have emphasized the role of the cuticle and underlying epidermis as optical filters for solar radiation. Plant cuticle structure. In contrast to the lack of association with cutin, extensive removal of wax from tomato fruit, accomplished by brief immersion of the fruit in an organic solvent, indicates that waxes contribute approximately 95% of the cuticle-mediated resistance to water diffusion, at least in tomato fruit (Leide et al., 2007). Crossref Annual Plant Reviews Volume 23: Biology of the Plant Cuticle The past decade has seen considerable progress in assembling models for the biosynthesis of its two major components, the polymer cutin and cuticular waxes. The relative order of these steps is not known, although it has been shown that the ω-hydroxylation precedes the midchain hydroxylation and that the final product of these steps is most likely a dihydroxyhexadecanoic acid-CoA ester (Li-Beisson et al., 2009). The elongation cycles can be terminated by a thioesterase to form free VLCFAs, or the VLCFA-CoA esters can undergo further modifications. In each of three Arabidopsis mutants exhibiting organ fusions, lacerata, bodyguard, and fiddlehead, ectopic organ fusions and cuticular permeability defects could be partially suppressed by a second mutation in SERRATE (Voisin et al., 2009). Mining the surface proteome of tomato (Solanum lycopersicum) fruit for proteins associated with cuticle biogenesis. edition, in English A Synthetic Community Approach Reveals Plant Genotypes Affecting the Phyllosphere Microbiota. Despite its similarity to lipolytic enzymes, the recombinant protein acts as an acyltransferase in vitro, forming polyester oligomers from 2-MHG (Yeats et al., 2012b). Furthermore, heterologous expression of the combination of CER1, CER3, a cytochrome b5, and LACS1 in yeast resulted in the formation of very-long-chain alkanes (Bernard et al., 2012). Written in English. Later studies indicated that cutin levels are also increased in WIN1/SHN1-overexpressing plants and that the up-regulation of genes encoding cutin biosynthetic enzymes precedes the induction of wax biosynthetic genes (Kannangara et al., 2007). 10.1146/annurev.arplant.59.103006.093219. Annual Plant Reviews, Volume 23 A much clearer picture is now emerging of the fine structure of the plant cuticle and its surface, the composition of cuticular waxes and the biosynthetic pathways leading to them. Nevertheless, based on the discovery of this effect, surfaces with high degrees of hydrophobicity and microscopic texture have been employed as effective biomimetic technical materials (Bhushan, 2012), and improved self-cleaning surfaces in agricultural crops may be a productive avenue of research. Although the specific sequence of all intracellular biosynthetic steps will require additional characterization of the substrate specificity of each enzyme, biochemical characterization of Arabidopsis bifunctional GPATs indicates that they have a strong preference for ω-hydroxylated acyl-CoA, suggesting that hydroxylation precedes the transfer to glycerol (Yang et al., 2012). Thus, cutin plays an important role as a physical barrier to many pathogens, yet extreme deficiencies in Arabidopsis can result in increased resistance to some pathogens by way of a secondary, but not well understood, mechanism that involves the induction of plant defenses. The regulation of cuticle biosynthesis is complex and involves interacting signaling networks associated with environmental stress responses, pathogen responses, and feedback regulation based on the structure and integrity of the cuticle itself. Joseph T. Wells, CPA, CFE, is the founder and Chairman of … After the synthesis of wax and cutin precursors, they are exported from the ER, across the plasma membrane, through the polysaccharide cell wall, and to the nascent cuticular membrane. The past decade has seen considerable progress in assembling models for the biosynthesis of its two major components, the polymer cutin and cuticular waxes. The cutin monomers that are released during polymeric cutin hydrolysis can act as elicitors of plant defense responses and are thus classified as damage-associated molecular patterns. Lastly, a defect in the formation of floral nanoridges was also identified in the Arabidopsis mutant defective in cuticular ridges (dcr), which showed a substantial deficiency in floral cutin but a less drastic alteration of leaf and stem cutin (Panikashvili et al., 2009). 2. It is important to note that, in this important model species, the cutin of stems and leaves is atypical in that its major component is a dicarboxylic acid (Fig. This strongly suggests that a complex including CER1 and CER3 with cytochrome b5 as an electron donor catalyzes the reduction and decarbonylation of VLCFA-CoA in order to form cuticular alkanes. Studies of the orthologous CFL1 gene in Arabidopsis indicated that it down-regulates cutin biosynthesis by suppressing the function of HDG1, a homeodomain-leucine zipper IV transcription factor (HD-ZIP IV), which has been shown to induce the expression of several cutin biosynthesis genes (Wu et al., 2011). EMBED (for wordpress.com hosted blogs and archive.org item tags) Want more? Life Science Ch. Mangifera indica The fusion zones are often marked by two adjacent polysaccharide cell walls with no visible cuticle separating the two organs, although the fused epidermal layers maintain their identity, as indicated by the differentiation of internal nonfunctional stomata within fusion zones (Sieber et al., 2000). In tomato fruit, severely decreased cutin levels in three cd mutants was associated with increased susceptibility to infection by Botrytis cinerea surface inoculation and also to opportunistic microbes (Isaacson et al., 2009). This edition doesn't have a description yet. Furthermore, as the cuticle is exclusively synthesized by epidermal cells, the regulation of epidermis identity during development can also be considered to play a regulatory role in cuticle development. An additional family of proteins, composed of CER2, CER26, and CER26-like, appears to be required for the elongation of fatty acids to lengths greater than 28C (Haslam et al., 2012; Pascal et al., 2013). . The cutin biopolymer matrix. Biology Test. ... a mature fertilized plant ovule consisting of an embryo and its food source and having a protective coat or testa. © 2013 American Society of Plant Biologists. There has been impressive progress in revealing the molecular biology underlying VLCFA-derived wax biosynthesis, and to this end, Arabidopsis has provided an excellent experimental model (Bernard and Joubès, 2013). Indeed, in recent years, there have been many instances of unexpected associations between the cuticle and diverse aspects of plant biology. It is thought that this self-cleaning surface helps to prevent the buildup of dust that would block sunlight and slow photosynthesis and that this could also play an important role in washing away pathogen spores before they germinate. (Image courtesy of Dr. Lacey Samuels.) This is consistent with a model in which cuticular waxes localize within either crystalline or amorphous domains of the cuticle, with aliphatic compounds forming crystallite “rafts” that are impervious to water, forcing water, and other polar metabolites, to diffuse by a circuitous route through the amorphous domains that are formed by more polar and cyclic waxes (Riederer and Schreiber, 1995). Conversely, the Arabidopsis lacs2 mutant and cutinase overexpressers exhibited no alteration in their susceptibility to a range of other fungal pathogens (Bessire et al., 2007), and the lacs2 mutation also increased susceptibility to a normally avirulent strain of Pseudomonas syringae (Tang et al., 2007). Characteristics of Fruit ( Cuticle architectural organization can be discerned using a number of microscopic techniques. Curiously, these enzymes have sequence homology to BAHD acyltransferases, but conserved catalytic amino acid residues of this family of enzymes are dispensable for the elongation-promoting activity of CER2 (Haslam et al., 2012). Moreover, it is not known how branching or cross linking of cutin affects cuticle function, and the identification of additional cutin synthases will allow this to be investigated using genetic approaches. Mechanical properties of cuticles and their primary determinants. The cuticle has been described as the outermost layer covering all aerial plant organs. One of the most intriguing mechanisms of cuticle regulation resulted from characterization of the cer7 mutant. An additional intracellular acyltransferase required for the synthesis of cutin polyester is a glycerol 3-phosphate acyltransferase (GPAT). In summary, resistance to water loss is primarily attributed to wax and not cutin, but there is not a direct correlation between the amount of either component and CWP. in Oxford, UK, . In 20 backcrossed families, CWP was inversely correlated with the amount of alkanes in the wax but not the total amount of wax, and the more rapidly desiccating parent had three times the wax coverage as the parent that exhibited low postharvest water loss (Parsons et al., 2012). MYB41 is induced by ABA, drought, and osmotic stress, leading to the down-regulation of cutin biosynthesis genes and the disruption of cuticle structure (Cominelli et al., 2008). This ultimately highlights the fact that the cuticle is a specialized modification of the cell wall, and like other modifications, such as lignification or suberization, it should be considered within the context of polysaccharide cell wall components. The first cutin synthase has been identified (Girard et al., 2012; Yeats et al., 2012b), but there are certainly additional cutin synthases, and whether they are closely related to CD1 or belong to distinct protein families remains to be discovered. Brand new Book. The identification of CD1 as the first known cutin synthase raises several questions about the specificity and generality of the reaction that it catalyzes. Typical cutin monomers and polymeric structure. The polymeric structure of cutin is not well understood. A new paradigm has recently been proposed for the diffusion of polar compounds and water across the cuticle. Christopher E. Jeffree, Science Faculty Electron Microscope Facility, Edinburgh, UK. However, in Arabidopsis, a number of cutin-deficient mutants and plants that ectopically overexpress fungal cutinases exhibit enhanced resistance to B. cinerea (Bessire et al., 2007, 2011; Chassot et al., 2007; Tang et al., 2007). The protein is localized in the outer cell wall of the epidermis below the cuticle, which led the authors to propose that BDG may be involved in cutin polymerization, although the increased amounts of polymeric cutin in the mutant would argue against this (Kurdyukov et al., 2006a). WIN1/SHN1 is part of a three-member gene family in Arabidopsis, and silencing of all three genes led to a reduction in the amount of cutin but not waxes (Shi et al., 2011). www.plantphysiol.org/cgi/doi/10.1104/pp.113.222737. After cutin is polymerized, is modification of the polymeric structure required to accommodate organ expansion? In this case, increased cuticular permeability appears to enhance the diffusion of inoculum-derived elicitors that induce the production of small, polar antifungal compounds, which in turn inhibit B. cinerea growth (Bessire et al., 2007). The IRG1 gene was found to encode a C2H2 zinc finger transcription factor that had previously been identified as a regulator of dissected leaf morphology (Chen et al., 2010). An edition of Biology of the plant cuticle (2006) Biology of the plant cuticle. They both appear to act upstream of, and directly activate, WIN1/SHN1 but also some cuticle biosynthetic genes (Oshima et al., 2013). Number of times cited according to CrossRef: FTIR spectroscopic features of the pteridosperm Ruflorinia orlandoi and host rock (Springhill Formation, Lower Cretaceous, Argentina). Osmophores and floral fragrance in The fine structure of the plant cuticle. In terms of photosynthesis, there is likely a tradeoff between a self-cleaning surface and the increased dispersion of light by epicuticular wax crystals, as discussed below. The plant cuticle is most typically associated with providing a fixed barrier to excessive transpirational water loss, allowing gas exchange and transpiration to be dynamically controlled by stomata. Experiments further addressing this hypothesis will be particularly interesting, given the surprising finding that the cer9 mutant actually exhibits enhanced drought tolerance and water use efficiency (Lü et al., 2012). Table of contents for Biology of the plant cuticle / edited by Markus Riederer & Caroline Muller. Markus Riederer, Julius--von--Sachs--Institut fur Biowissenschaften, Universitat Wurzburg, Wurzburg, Germany. A long unresolved question in wax biosynthesis is the enzymatic basis of alkane synthesis. Most recently, two breakthroughs in the long-sought molecular bases of alkane formation and polyester synthesis have allowed construction of nearly complete biosynthetic pathways for both waxes and cutin. Smooth, glossy “glabrous” cuticles typically reflect only small amounts of light (less than 10%), but glaucous plant surfaces are moderately reflective and generally show approximately 20% to 30% reflectance in the UV and visible spectra (Pfündel et al., 2006). The plant cuticle is an extracellular hydrophobic layer that covers the aerial epidermis of all land plants, providing protection against desiccation and external environmental stresses. In Arabidopsis, both of these oxidations are performed by the cytochrome P450 enzyme midchain alkane hydroxylase (MAH1; Greer et al., 2007). Changes in leaf epicuticular wax, gas exchange and biochemistry metabolism between Jatropha mollissima and Jatropha curcas under semi-arid conditions. Here, we review recent progress in the biochemistry and molecular biology of cuticle synthesis and function and highlight some of the major questions that will drive future research in this field. The ω-hydroxylase is encoded by members of the CYP86 subfamily of cytochrome P450s (CYP86A4 in Arabidopsis flowers; Li-Beisson et al., 2009), while the midchain hydroxylase is encoded by the CYP77 subfamily (CYP77A6 in Arabidopsis flowers; Li-Beisson et al., 2009). While mechanical rupture may be sufficient for cuticle penetration, particularly of thinner cuticles (Tenberge, 2007), most fungal pathogens also secrete cutinases, a class of small, nonspecific esterases that hydrolyze the cutin polyester and release free cutin monomers (Longhi and Cambillau, 1999). The fine structure of the plant cuticle Christopher E. Jeffree, Science Faculty Electron Microscope Facility, Edinburgh, UK 3. Ed. 1. in the alpine region of central Japan were not related to mechanical damage of cuticle and cuticle thickness 23) Volume 23 Edition by Markus Riederer (Editor), Caroline Muller (Editor) ISBN-13: 978-1405132688 The author responsible for distribution of materials integral to the findings presented in this article in accordance with the policy described in the Instructions for Authors (www.plantphysiol.org) is: Jocelyn K.C. On the other hand, a general role of WIN1/SHN1-related transcription factors in the regulation of cutin synthesis is indicated by studies of orthologous genes in barley (Hordeum vulgare; Taketa et al., 2008) and tomato (Shi et al., 2013). Through successive addition of two carbons per cycle derived from malonyl-CoA, the ultimate products of this complex are VLCFAs. by Markus Riederer and Caroline Muller. 4). Biology of the Plant Cuticle by Markus Riederer, 9780470988718, available at Book Depository with free delivery worldwide. While our understanding of cuticle biosynthesis at the molecular level remains incomplete, recent progress in deciphering these pathways is bringing us closer than ever to an ability to selectively modify cuticle properties in order to improve agricultural productivity. - Language: English. As described above, several key areas of cuticle biogenesis remain poorly understood. Joseph T. Wells, CPA, CFE, is the founder and Chairman of … The roles of the cuticle in plant development: organ adhesions and beyond. 1. The transition from an exclusively aquatic to a terrestrial life style, therefore, would have necessitated the evolution of a toolbox of morphological and physiological features, some of which are apparent through studies of the fossil record or by examining extant plant lineages. Benny Chefetz, Decomposition and sorption characterization of plant cuticles in soil, Plant and Soil, 10.1007/s11104-007-9318-1, 298, 1-2, (21-30), (2007). For a more detailed review of cuticle chemical ecology, see Müller and Riederer (2005). DCR encodes a protein of the BAHD acyltransferase family that localizes to the cytoplasm, and it has been proposed that it may be involved in acyl transfer of cutin monomers to form precursor intermediates or oligomeric structures (Panikashvili et al., 2009). Another example of the influence of waxes on pathogenicity is provided by the inhibitor of rust tube germination1 (irg1) mutant of M. truncatula, which exhibits decreased amounts of epicuticular wax crystals on the abaxial leaf surface, corresponding to a substantial decrease in wax primary alcohol groups. Monomeric composition can provide a “parts list,” but the relative abundance of possible linkages in the polymer is difficult to determine, largely due to the difficulty of solubilizing intact cutin (Serra et al., 2012). However, the most critical adaptive trait for survival during terrestrialization would have been the ability to retain water in increasingly dehydrating habitats. cyanobacteria Blue-green bacteria; unicellular or filamentous chains of cells that carry out photosynthesis. Differences in protodermal cell wall structure in zygotic and somatic embryos of Daucus carota (L.) cultured on solid and in liquid media. 0 Ratings 0 Want to read; 0 Currently reading; 0 Have read; This edition published in 2006 by Blackwell Pub. This suggests that lipid-binding proteins or other factors are not necessary in order to facilitate the transport of this major precursor of cutin biosynthesis. This observation is further corroborated by studies of the maize mutant glossy1, which does not accumulate aldehydes in its wax complement. The waxes are either deposited within the cutin matrix (intracuticular wax) or accumulate on its surface as epicuticular wax crystals, or films. Annual Plant Reviews, Biology of the Plant Cuticle by Markus Riederer, Caroline Muller, 2008, Wiley & Sons, Incorporated, John edition, in English Classical biochemistry, using crude extracts from pea (Pisum sativum), indicated that the reaction likely occurs via the reduction of VLCFA-CoA to an aldehyde intermediate followed by decarbonylation, yielding an alkane that is 1C shorter (Cheesbrough and Kolattukudy, 1984; Schneider-Belhaddad and Kolattukudy, 2000). leaves and young shoots. Cloning of the CER9 gene revealed it to encode a protein with sequence similarity to yeast Doa10, an E3 ubiquitin ligase involved in ER-associated degradation of misfolded proteins (Lü et al., 2012). Scanning electron microscopy can reveal the elaborate and diverse morphologies of epicuticular wax crystals (Fig. Specifically, it has decreased levels of dicarboxylic acids and increased amounts of ω-hydroxy acids, leading the authors to suggest that HTH may have a role in the oxidation of ω-hydroxy fatty acids to the dicarboxylic acid cutin monomers that are characteristic of Arabidopsis stem and leaf cuticles (Kurdyukov et al., 2006b). 0 Ratings 0 Want to read; 0 Currently reading; 0 Have read; This edition published in 2006 by Blackwell Pub. Waxes reflect both UV and visible light, but not necessarily to the same extent, and the reflectance of UV has been reported to be greater in some cases (Holmes and Keiller, 2002). Note: Contents data are machine generated based on pre-publication provided by the publisher. Regulation of cuticle biosynthesis. Endocytotic uptake of nutrients in carnivorous plants. C, Transmission electron micrograph image of an Arabidopsis stem epidermal cell wall and cuticle. 55 terms. Genes encoding the remaining subunits of the FAE complex, represented by KCR1, PAS2, and CER10, respectively, are less redundant, and their pleiotropic mutant phenotypes underscore the shared importance of the FAE in generating VLCFA precursors for sphingolipid biosynthesis (Zheng et al., 2005; Bach et al., 2008; Beaudoin et al., 2009). It has also become clear that the physiological role of the cuticle extends well beyond its primary function as a transpiration barrier, playing important roles in processes ranging from development to interaction with microbes. This book treats the major functions in detail and, in most cases, devotes separate chapters to each of them. A model was proposed wherein CER7 is involved in the degradation of a small RNA species that negatively regulates the CER3 transcript. Aside from the SHN family, other AP2 domain transcription factors from different clades may also play a role in cuticle regulation. 3. Overexpression of this gene led to glossy leaves with a greater wax load than the wild type and lower transpiration, although this was likely due to a reduced density of stomata rather than the wax phenotype (Aharoni et al., 2004). In Arabidopsis, 21 genes are predicted to encode β-ketoacyl-CoA synthase, and for wax biosynthesis, the most important gene, based on the mutant phenotype, is CER6 (Fiebig et al., 2000). Plus, free two-day shipping for six months when you sign up for Amazon Prime for Students. Cutin is typically composed of interesterified hydroxy fatty acids, with lesser amounts of glycerol, phenylpropanoids, and dicarboxylic acids (Kolattukudy, 2001). Annual Plant Reviews, Volume 23 A much clearer picture is now emerging of the fine structure of the plant cuticle and its surface, the composition of cuticular waxes and the biosynthetic pathways leading to them. Consequently, the capacity to synthesize, deposit, and maintain a hydrophobic surface layer, or cuticle, over the surfaces of aerial organs was arguably one of the most important innovations in the history of plant evolution. Chemical processes that cleave ester bonds, such as saponification, readily release these monomeric constituents, although in some species an additional lipidic polymer, referred to as cutan, remains recalcitrant to such treatments. Get this from a library! 1A). In the case of the dihydroxyacyl cutin precursor 2-MHG, the glycerol moiety imparts sufficient polarity to allow aqueous solubility at low millimolar concentrations (Yeats et al., 2012b). The fine structure of the plant cuticle. 23) Volume 23 Edition by Markus Riederer (Editor), Caroline Muller (Editor) ISBN-13: 978-1405132688 Nevertheless, a complex regulatory network that responds to developmental and environmental cues, mediated by hormones, transcription factors, and posttranscriptional regulation, is beginning to emerge. The plant cuticle, a multi-layered membrane that covers plant aerial surfaces to prevent desiccation, consists of the structural polymer cutin and surface-sealing waxes. Huihui Bi, Sukanya Luang, Yuan Li, Natalia Bazanova, Nikolai Borisjuk, Maria Hrmova, Sergiy Lopato, Wheat drought-responsive WXPL transcription factors regulate cuticle biosynthesis genes, Plant Molecular Biology, 10.1007/s11103-017-0585-9, 94, 1-2, (15-32), (2017). SA, an important regulator of systemic acquired resistance (SAR), is preferentially transported from pathogen-infected to uninfected parts via the apoplast. Despite the clear genetic evidence supporting a role for ABC transporters in cuticular lipid export, the substrate specificity of these transporters has not yet been demonstrated in vitro. Biology of the plant cuticle by , 2006, Blackwell Pub. ABCG11 is also required for cutin accumulation, and since it is also able to dimerize with itself, it has been proposed that this homodimer is the functional complex responsible for cutin export (McFarlane et al., 2010). All Rights Reserved. Please check your email for instructions on resetting your password. L.) Skin: Structure and Antioxidant Content UV light in the UV-B spectrum is a considerable portion of the daylight that reaches the terrestrial surface, and it can threaten plant life by damaging DNA, the photosynthetic apparatus, and membrane lipids (Rozema et al., 1997). This layer may, as in the arthropods, contain pigments and chitin; in humans the cuticle is the epidermis. Bar = 5 μm. A common perception is that a thick cuticle is associated with a lower water permeability and thus increased tolerance to water stress. In this regard, an interesting area of future research might to determine whether relative proportions of UV and visible light reflection can be predictively changed by altering the composition of epicuticular waxes. Introduction: Biology of the plant cuticle. However, it has evolved a number of secondary functions that are consistent with its place as the outermost layer of primary aerial organs: it forms a physical barrier that is the first line of defense against pests and pathogens; in many species, elaborate epicuticular crystals help to form a self-cleaning surface, preventing dust and other debris from blocking sunlight; in some cases, it can act to screen excessive UV light; finally, as a defining feature of the epidermis, it plays a central role in development by physically establishing organ boundaries. The first plant colonizers of land, approximately 450 million years ago in the mid-Paleozoic era, faced a daunting set of challenges associated with their new terrestrial environment, including desiccation, temperature extremes, gravity, and increased exposure to UV radiation (Waters, 2003; Leliaert et al., 2011). Another example of an “orphan” cuticle-associated protein resulted from analysis of the Arabidopsis bodyguard (bdg) mutant, which exhibits a microscopically disorganized cuticle with increased permeability but significantly increased levels of wax and cutin (Kurdyukov et al., 2006a). and you may need to create a new Wiley Online Library account. Given the ER localization of wax and cutin biosynthetic processes, the authors proposed a role for CER9 in the homeostasis of key cuticle biosynthetic enzyme levels. Treats the major functions in detail and, ultimately, alkanes can undergo further modifications the initial contact between! 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