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Plant cuticle

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Water beads on the waxy cuticle of kale leaves

A plant cuticle is a protecting film covering the outermost skin layer (epidermis) of leaves, young shoots and other aerial plant organs (aerial here meaning all plant parts not embedded in soil or other substrate) that have no periderm. The film consists of lipid and hydrocarbon polymers infused with wax, and is synthesized exclusively by the epidermal cells. [1]

Description

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Anatomy of a eudicot leaf

The plant cuticle is a layer of lipid polymers impregnated with waxes that is present on the outer surfaces of the primary organs of all vascular land plants. It is also present in the sporophyte generation of hornworts, and in both sporophyte and gametophyte generations of mosses.[2] The plant cuticle forms a coherent outer covering of the plant that can be isolated intact by treating plant tissue with enzymes such as pectinase and cellulase.[citation needed]

Composition

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The cuticle is composed of an insoluble cuticular membrane impregnated by and covered with soluble waxes. Cutin, a polyester polymer composed of inter-esterified omega hydroxy acids which are cross-linked by ester and epoxide bonds, is the best-known structural component of the cuticular membrane.[3][4] The cuticle can also contain a non-saponifiable hydrocarbon polymer known as cutan.[5] The cuticular membrane is impregnated with cuticular waxes[6] and covered with epicuticular waxes, which are mixtures of hydrophobic aliphatic compounds, hydrocarbons with chain lengths typically in the range C16 to C36.[7]

Cuticular wax biosynthesis

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Cuticular wax is known to be largely composed of compounds which derive from very-long-chain fatty acids (VLCFAs), such as aldehydes, alcohols, alkanes, ketones, and esters.[8][9] Also present are other compounds in cuticular wax which are not VLCFA derivatives, such as terpenoids, flavonoids, and sterols,[9] and thus have different synthetic pathways than those VLCFAs.

The first step of the biosynthesis pathway for the formation of cuticular VLCFAs, occurs with the de novo biosynthesis of C16 acyl chains (palmitate) by chloroplasts in the mesophyll,[1] and concludes with the extension of these chains in the endoplasmic reticulum of epidermal cells.[9] An important catalyzer thought to be in this process is the fatty acid elongase (FAE) complex.[8][9][10]

To form cuticular wax components, VLCFAs are modified through either two identified pathways, an acyl reduction pathway or a decarbonylation pathway.[9] In the acyl reduction pathway, a reductase converts VLCFAs into primary alcohols, which can then be converted to wax esters through a wax synthase.[9][10] In the decarbonylation pathway, aldehydes are produced and decarbonylated to form alkanes, and can be subsequently oxidized to form secondary alcohols and ketones.[8][9][10] The wax biosynthesis pathway ends with the transportation of the wax components from the endoplasmic reticulum to the epidermal surface.[9]

Functions

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The primary function of the plant cuticle is as a water permeability barrier that prevents evaporation of water from the epidermal surface, and also prevents external water and solutes from entering the tissues.[11] In addition to its function as a permeability barrier for water and other molecules (prevent water loss), the micro and nano-structure of the cuticle have specialised surface properties that prevent contamination of plant tissues with external water, dirt and microorganisms. Aerial organs of many plants, such as the leaves of the sacred lotus (Nelumbo nucifera) have ultra-hydrophobic and self-cleaning properties that have been described by Barthlott and Neinhuis (1997).[12] The lotus effect has applications in biomimetic technical materials.

Dehydration protection provided by a maternal cuticle improves offspring fitness in the moss Funaria hygrometrica[2] and in the sporophytes of all vascular plants. In angiosperms the cuticle tends to be thicker on the top of the leaf (adaxial surface), but is not always thicker. The leaves of xerophytic plants adapted to drier climates have more equal cuticle thicknesses compared to those of mesophytic plants from wetter climates that do not have a high risk of dehydration from the under sides of their leaves.

"The waxy sheet of cuticle also functions in defense, forming a physical barrier that resists penetration by virus particles, bacterial cells, and the spores and growing filaments of fungi".[13]

Evolution

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The plant cuticle is one of a series of innovations, together with stomata, xylem and phloem and intercellular spaces in stem and later leaf mesophyll tissue, that plants evolved more than 450 million years ago during the transition between life in water and life on land.[11] Together, these features enabled upright plant shoots exploring aerial environments to conserve water by internalising the gas exchange surfaces, enclosing them in a waterproof membrane and providing a variable-aperture control mechanism, the stomatal guard cells, which regulate the rates of transpiration and CO2 exchange.

References

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  1. ^ a b Kolattukudy, PE (1996) Biosynthetic pathways of cutin and waxes, and their sensitivity to environmental stresses. In: Plant Cuticles. Ed. by G. Kerstiens, BIOS Scientific publishers Ltd., Oxford, pp 83-108
  2. ^ a b Budke, J.M.; Goffinet, B.; Jones, C.S. (2013). "Dehydration protection provided by a maternal cuticle improves offspring fitness in the moss Funaria hygrometrica". Annals of Botany. 111 (5): 781–789. doi:10.1093/aob/mct033. PMC 3631323. PMID 23471009.
  3. ^ Holloway, PJ (1982) The chemical constitution of plant cutins. In: Cutler, DF, Alvin, KL and Price, CE The Plant Cuticle. Academic Press, pp. 45-85
  4. ^ Stark, RE and Tian, S (2006) The cutin biopolymer matrix. In: Riederer, M & Müller, C (2006) Biology of the Plant Cuticle. Blackwell Publishing
  5. ^ Tegelaar, EW, et al. (1989) Scope and limitations of several pyrolysis methods in the structural elucidation of a macromolecular plant constituent in the leaf cuticle of Agave americana L., Journal of Analytical and Applied Pyrolysis, 15, 29-54
  6. ^ Jetter, R, Kunst, L & Samuels, AL (2006) Composition of plant cuticular waxes. In: Riederer, M & Müller, C (2006) Biology of the Plant Cuticle. Blackwell Publishing, 145-181
  7. ^ Baker, EA (1982) Chemistry and morphology of plant epicuticular waxes. In: Cutler, DF, Alvin, KL and Price, CE The Plant Cuticle. Academic Press, 139-165
  8. ^ a b c Yeats, Trevor H.; Rose, Jocelyn K.C. (September 2013). "The Formation and Function of Plant Cuticles". Plant Physiology. 163 (1): 5–20. doi:10.1104/pp.113.222737. ISSN 0032-0889. PMC 3762664. PMID 23893170.
  9. ^ a b c d e f g h Kunst, L; Samuels, A. L (2003-01-01). "Biosynthesis and secretion of plant cuticular wax". Progress in Lipid Research. 42 (1): 51–80. doi:10.1016/S0163-7827(02)00045-0. ISSN 0163-7827. PMID 12467640.
  10. ^ a b c Suh, Mi Chung; Kim, Hae Jin; Kim, Hyojin; Go, Young Sam (2014-04-01). "Arabidopsis Cuticular Wax Biosynthesis Is Negatively Regulated by the DEWAX Gene Encoding an AP2/ERF-Type Transcription Factor". The Plant Cell. 26 (4): 1666–1680. doi:10.1105/tpc.114.123307. ISSN 1040-4651. PMC 4036578. PMID 24692420.
  11. ^ a b Raven, J.A. (1977). "The evolution of vascular land plants in relation to supracellular transport processes". Advances in Botanical Research. 5: 153–219. doi:10.1016/S0065-2296(08)60361-4. ISBN 9780120059058.
  12. ^ Barthlott, W.; Neinhuis, C (1997). "Purity of the sacred lotus, or escape from contamination in biological surfaces". Planta. 202: 1–8. doi:10.1007/s004250050096. S2CID 37872229.
  13. ^ Freeman, S. (2002). Biological Science. New Jersey: Prentice-Hall, Inc. ISBN 978-0130819239.
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