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Concept: Cutin

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The plant cuticle is the outermost layer covering aerial tissues and is composed of cutin and waxes. The cuticle plays an important role in protection from environmental stresses and glaucousness, the bluish-white colouration of plant surfaces associated with cuticular waxes, has been suggested as a contributing factor in crop drought tolerance. However, the cuticle structure and composition is complex and it is not clear which aspects are important in determining a role in drought tolerance. Therefore, we analysed residual transpiration rates, cuticle structure and epicuticular wax composition under well-watered conditions and drought in five Australian bread wheat genotypes, Kukri, Excalibur, Drysdale, RAC875 and Gladius, with contrasting glaucousness and drought tolerance.

Concepts: Cutin, Epicuticular wax, Epidermis, Wax, Plant physiology, Leaf, Cuticle, Plant cuticle

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The climatic stressors that modify the chemical composition of plant tissues have been recently reviewed by Suseela and Tharayil (2017). In particular, the authors stated that the response of plants to global change effects (viz. increasing temperatures and frequent drought periods) can induce important modifications in plant cuticles such as an increase of the main cuticle components (cutin and polysaccharides), a preponderance of cutan, heavier wax loads, and changes in the chemical composition of waxes, mainly the accumulation of longer aliphatic compounds. In this letter, we would like to emphasize the biophysical consequences that this new scenario involves and how it would affect plant performance. This article is protected by copyright. All rights reserved.

Concepts: Omega hydroxy acid, Epidermis, All rights reserved, Copyright, Cuticle, Cutan, Plant cuticle, Cutin

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Cuticles envelope primary surfaces of the above-ground portion of plants. They function as barriers to water movement and to gas exchange, and in pathogen defense. To serve as a barrier on growing organs, cuticles must remain intact but at the same time must accommodate ongoing growth. Minimizing cuticle failure has stimulated significant research on the cuticle’s mechanical properties. The objective here is to review the literature on the mechanical properties of isolated fruit and leaf cuticles. Cuticles are viscoelastic polymers. Viscoelasticity results mainly from the cutin matrix. Impregnation by waxes, flavonoids, and cutan increases stiffness and strength but decreases extensibility. On the inner side, the cutin matrix is impregnated by cell wall polysaccharides, which are responsible for its elastic behavior. Across species, the maximum forces sustainable by hydrated cuticles in uniaxial tensile tests averaged 0.82 N (range 0.15-1.63 N), the maximum stresses averaged 13.2 MPa (range 2.0-29.0 MPa), the maximum strains averaged 8.8% (range 1.6-28.0%), and the moduli of elasticity averaged 224 MPa (range 60-730 MPa). Among the environmental factors, high temperature and hydration both decreased stiffness. Therefore, the mechanical properties of cuticles in vivo depend largely on the relative proportions of their constituents. These proportions change during development and are also affected by environmental factors such as temperature.

Concepts: Bacteria, Cell wall, Cutin, Plant cuticle, Cutan, Continuum mechanics, Viscoelasticity, Elasticity

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The assembly of the lipophilic cuticle layer and suberin lamellae, approximately 450 million years ago, was a major evolutionary development that enabled plants to colonize terrestrial habitats. The cuticle layer is composed of cutin polyester and embedded cuticular waxes, whereas the suberin lamellae consist of very long chain fatty acid derivatives, glycerol, and phenolics cross-linked with alkyl ferulate-embedded waxes. Due to their substantial biological roles in plant life, the mechanisms underlying the assembly of these structures have been extensively investigated. In the last decade, the introduction of ‘omics’ approaches, including genomics, transcriptomics, proteomics, and metabolomics, have been key in the identification of novel genetic and chemical elements involved in the formation and function of the cuticle layer and suberin lamellae. This review summarizes contemporary studies that utilized various large-scale, ‘omics’ strategies in combination with novel technologies to unravel how building blocks and polymers of these lipophilic barriers are made, and moreover linking structure to function along developmental programs and stress responses. We anticipate that the studies discussed here will inspire scientists studying lipophilic barriers to integrate complementary ‘omics’ approaches in their efforts to tackle as yet unresolved questions and engage the main challenges of the field to date.

Concepts: Chemical element, Biology, Evolution, Genetics, Proteomics, Cutin, Functional genomics, Fatty acid

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Aliphatic and aromatic lipids are both essential structural components of the plant cuticle, an important interface between the plant and environment. Although crosslinks between aromatic and aliphatic or other moieties are known to be associated with the formation of leaf cutin, root and seed suberin, the contribution of aromatic lipids to the biosynthesis of anther cuticles and pollen walls remains elusive. In this study, we characterized the rice male sterile mutant, defective pollen wall 2 (dpw2), which showed an abnormal anther cuticle, a defective pollen wall, and complete male sterility. Compared with the wild type (WT), dpw2 anthers have increased amounts of cutin and waxes and decreased levels of lipidic and phenolic compounds. DPW2 encodes a cytoplasmically localized BAHD acyltransferase. In vitro assays demonstrated that recombinant DPW2 specifically transfers hydroxycinnamic acid moieties, using ω-hydroxy fatty acids as acyl acceptors and hydroxycinnamoyl-CoAs as acyl donors. Thus, DPW2 plays a fundamental role in pollen development via the biosynthesis of key components of the anther cuticle and pollen wall.

Concepts: Cuticle, Plant cuticle, Pollination, Flower, Cutin, Plant anatomy, Fatty acid, Plant morphology

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The Arabidopsis cuticle, as observed by electron microscopy, consists primarily of the cutin/cutan matrix. The cuticle possesses a complex substructure, which is correlated with the presence of intracuticular waxes. The plant cuticle is composed of an insoluble polyester, cutin, and organic solvent soluble cuticular waxes, which are embedded within and coat the surface of the cutin matrix. How these components are arranged in the cuticle is not well understood. The Arabidopsis cuticle is commonly understood as ‘amorphous,’ lacking in ultrastructural features, and is often observed as a thin (~80-100 nm) electron-dense layer on the surface of the cell wall. To examine this cuticle in more detail, we examined cuticles from both rapidly elongating and mature sections of the stem and compared the preservation of the cuticles using conventional chemical fixation methods and high-pressure freezing/freeze-substitution (HPF/FS). We found that HPF/FS preparation revealed a complex cuticle substructure, which was more evident in older stems. We also found that the cuticle increases in thickness with development, indicating an accretion of polymeric material, likely in the form of the non-hydrolyzable polymer, cutan. When wax was extracted by chloroform immersion prior to sample preparation, the contribution of waxes to cuticle morphology was revealed. Overall, the electron-dense cuticle layer was still visible but there was loss of the cuticle substructure. Furthermore, the cuticle of cer6, a wax-deficient mutant, also lacked this substructure, suggesting that these fine striations were dependent on the presence of cuticular waxes. Our findings show that HPF/FS preparation can better preserve plant cuticles, but also provide new insights into the fine structure of the Arabidopsis cuticle.

Concepts: Polyester, Solubility, Polymer, Plant cuticle, Cutan, Cutin, Cuticle, Solvent