Aplant cuticle is a protecting film covering the outermost skin layer (epidermis) ofleaves, young shoots and other aerial plant organs (aerial here meaning all plant parts not embedded in soil or other substrate) that have noperiderm. The film consists of lipid and hydrocarbon polymers infused with wax, and is synthesized exclusively by the epidermal cells.[1]
The plant cuticle is a layer composed of lipid polymers and waxes that is present on the outer surfaces of the primary organs of all vascular land plants. It is also present in thesporophyte generation ofhornworts, and in both sporophyte andgametophyte generations ofmosses.[2] The plant cuticle forms a coherent outer covering of the plant that can be isolated intact by treating plant tissue with enzymes such aspectinase andcellulase.[citation needed]
Plant cuticles are composed primarily of the insoluble lipid polymerscutin andcutan, as well as solublewaxes.Cutin, apolyesterpolymer composed of inter-esterifiedomega hydroxy acids which are cross-linked byester andepoxide bonds, is the best-known structural component of the cuticular membrane.[3][4] The cuticle can also contain a non-saponifiablehydrocarbon polymer known ascutan.[5]
The cuticular membrane also contains waxes, which are the main mechanism for conferring water resistance in the plant cuticle.[6][7] Theseepicuticular waxes are mixtures ofhydrophobicaliphatic compounds, hydrocarbons with chain lengths typically in the range C16 to C36.[8]
Cuticular wax is known to be largely composed of compounds which derive fromvery-long-chain fatty acids (VLCFAs), such asaldehydes,alcohols,alkanes,ketones, andesters.[9][10] Also present are other compounds in cuticular wax which are not VLCFA derivatives, such asterpenoids,flavonoids, andsterols,[10] 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 theendoplasmic reticulum ofepidermal cells.[10] An important catalyzer thought to be in this process is the fatty acid elongase (FAE) complex.[9][10][11]
To form cuticular wax components, VLCFAs are modified through either two identified pathways, an acyl reduction pathway or adecarbonylation pathway.[10] In the acyl reduction pathway, areductase converts VLCFAs into primary alcohols, which can then be converted to wax esters through awax synthase.[10][11] In the decarbonylation pathway, aldehydes are produced and decarbonylated to form alkanes, and can be subsequently oxidized to form secondary alcohols and ketones.[9][10][11] The wax biosynthesis pathway ends with the transportation of the wax components from the endoplasmic reticulum to the epidermal surface.[10]
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.[12] 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).[13] Thelotus effect has applications inbiomimetic technical materials.
Dehydration protection provided by a maternal cuticle improves offspring fitness in the mossFunaria hygrometrica[2] and in the sporophytes of allvascular plants. Inangiosperms the cuticle tends to be thicker on the top of the leaf (adaxial surface), but is not always thicker. The leaves ofxerophytic plants adapted to drier climates have more equal cuticle thicknesses compared to those ofmesophytic plants from wetter climates that do not have a high risk of dehydration.[citation needed]
"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".[14]
The plant cuticle is one of a series ofinnovations, together withstomata,xylem andphloem and intercellular spaces instem and laterleaf mesophyll tissue, thatplants evolved more than 450 million years ago during the transition between life in water and life on land.[12] 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, thestomatal guard cells, which regulate the rates oftranspiration and CO2 exchange.
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