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A plant cuticle is a protecting film covering the outermost skin layer (
epidermis The epidermis is the outermost of the three layers that comprise the skin, the inner layers being the dermis and hypodermis. The epidermis layer provides a barrier to infection from environmental pathogens and regulates the amount of water relea ...
) of
leaves A leaf ( : leaves) is any of the principal appendages of a vascular plant stem, usually borne laterally aboveground and specialized for photosynthesis. Leaves are collectively called foliage, as in "autumn foliage", while the leaves, ste ...
, young shoots and other aerial plant organs (aerial here meaning all plant parts not embedded in soil or other substrate) that have no ''
periderm Bark is the outermost layers of stems and roots of woody plants. Plants with bark include trees, woody vines, and shrubs. Bark refers to all the tissues outside the vascular cambium and is a nontechnical term. It overlays the wood and consist ...
''. The film consists of lipid and hydrocarbon polymers impregnated with wax, and is synthesized exclusively by the epidermal cells. Kolattukudy, PE (1996) Biosynthetic pathways of cutin and waxes, and their sensitivity to
environmental stresses Stress, either physiological, biological or psychological, is an organism's response to a stressor such as an environmental condition. Stress is the body's method of reacting to a condition such as a threat, challenge or physical and psycholog ...
. In: Plant Cuticles. Ed. by G. Kerstiens, BIOS Scientific publishers Ltd., Oxford, pp 83-108


Description

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 A sporophyte () is the diploid multicellular stage in the life cycle of a plant or alga which produces asexual spores. This stage alternates with a multicellular haploid gametophyte phase. Life cycle The sporophyte develops from the zygote p ...
generation of
hornwort Hornworts are a group of non-vascular Embryophytes (land plants) constituting the division Anthocerotophyta (). The common name refers to the elongated horn-like structure, which is the sporophyte. As in mosses and liverworts, hornworts have a ...
s, and in both sporophyte and
gametophyte A gametophyte () is one of the two alternating multicellular phases in the life cycles of plants and algae. It is a haploid multicellular organism that develops from a haploid spore that has one set of chromosomes. The gametophyte is the se ...
generations of
moss Mosses are small, non-vascular flowerless plants in the taxonomic division Bryophyta (, ) ''sensu stricto''. Bryophyta (''sensu lato'', Schimp. 1879) may also refer to the parent group bryophytes, which comprise liverworts, mosses, and hornw ...
es 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 Pectinases are a group of enzymes that breaks down pectin, a polysaccharide found in plant cell walls, through hydrolysis, transelimination and deesterification reactions. Commonly referred to as pectic enzymes, they include pectolyase, pectozyme ...
and
cellulase Cellulase (EC 3.2.1.4; systematic name 4-β-D-glucan 4-glucanohydrolase) is any of several enzymes produced chiefly by fungi, bacteria, and protozoans that catalyze cellulolysis, the decomposition of cellulose and of some related polysaccharid ...
.


Composition

The cuticle is composed of an insoluble cuticular membrane impregnated by and covered with soluble
wax Waxes are a diverse class of organic compounds that are lipophilic, malleable solids near ambient temperatures. They include higher alkanes and lipids, typically with melting points above about 40 °C (104 °F), melting to give low ...
es.
Cutin Cutin is one of two waxy polymers that are the main components of the plant cuticle, which covers all aerial surfaces of plants. It is an insoluble substance with waterproof quality. Cutin also harbors cuticular waxes, which assist in cuticle stru ...
, a
polyester Polyester is a category of polymers that contain the ester functional group in every repeat unit of their main chain. As a specific material, it most commonly refers to a type called polyethylene terephthalate (PET). Polyesters include natural ...
polymer A polymer (; Greek '' poly-'', "many" + ''-mer'', "part") is a substance or material consisting of very large molecules called macromolecules, composed of many repeating subunits. Due to their broad spectrum of properties, both synthetic and ...
composed of inter-esterified
omega hydroxy acid Omega hydroxy acids (also known as ω- hydroxycarboxylic acids) are a class of naturally occurring straight-chain aliphatic organic acids ''n'' carbon atoms long with a carboxyl group at position 1 (the starting point for the family of carboxylic ac ...
s which are cross-linked by
ester In chemistry, an ester is a compound derived from an oxoacid (organic or inorganic) in which at least one hydroxyl group () is replaced by an alkoxy group (), as in the substitution reaction of a carboxylic acid and an alcohol. Glycerides are f ...
and
epoxide In organic chemistry, an epoxide is a cyclic ether () with a three-atom ring. This ring approximates an equilateral triangle, which makes it strained, and hence highly reactive, more so than other ethers. They are produced on a large scale f ...
bonds, is the best-known structural component of the cuticular membrane.Holloway, PJ (1982) The chemical constitution of plant cutins. In: Cutler, DF, Alvin, KL and Price, CE The Plant Cuticle. Academic Press, pp. 45-85Stark, RE and Tian, S (2006) The cutin biopolymer matrix. In: Riederer, M & Müller, C (2006) Biology of the Plant Cuticle. Blackwell Publishing The cuticle can also contain a non-saponifiable
hydrocarbon In organic chemistry, a hydrocarbon is an organic compound consisting entirely of hydrogen and carbon. Hydrocarbons are examples of group 14 hydrides. Hydrocarbons are generally colourless and hydrophobic, and their odors are usually weak or ...
polymer known as Cutan.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 The cuticular membrane is impregnated with cuticular waxesJetter, 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 and covered with
epicuticular wax Epicuticular wax is a coating of wax covering the outer surface of the plant cuticle in land plants. It may form a whitish film or bloom on leaves, fruits and other plant organs. Chemically, it consists of hydrophobic organic compounds, mainly st ...
es, which are mixtures of
hydrophobic In chemistry, hydrophobicity is the physical property of a molecule that is seemingly repelled from a mass of water (known as a hydrophobe). In contrast, hydrophiles are attracted to water. Hydrophobic molecules tend to be nonpolar and, t ...
aliphatic compound In organic chemistry, hydrocarbons ( compounds composed solely of carbon and hydrogen) are divided into two classes: aromatic compounds and aliphatic compounds (; G. ''aleiphar'', fat, oil). Aliphatic compounds can be saturated, like hexane, ...
s, hydrocarbons with chain lengths typically in the range C16 to C36.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


Cuticular wax biosynthesis

Cuticular wax is known to be largely composed of compounds which derive from very-long-chain fatty acids (VLCFAs), such as
aldehydes In organic chemistry, an aldehyde () is an organic compound containing a functional group with the structure . The functional group itself (without the "R" side chain) can be referred to as an aldehyde but can also be classified as a formyl group ...
,
alcohols In chemistry, an alcohol is a type of organic compound that carries at least one hydroxyl () functional group bound to a saturated carbon atom. The term ''alcohol'' originally referred to the primary alcohol ethanol (ethyl alcohol), which is ...
,
alkanes In organic chemistry, an alkane, or paraffin (a historical trivial name that also has other meanings), is an acyclic saturated hydrocarbon. In other words, an alkane consists of hydrogen and carbon atoms arranged in a tree structure in whic ...
,
ketones In organic chemistry, a ketone is a functional group with the structure R–C(=O)–R', where R and R' can be a variety of carbon-containing substituents. Ketones contain a carbonyl group –C(=O)– (which contains a carbon-oxygen double bo ...
, and
esters In chemistry, an ester is a compound derived from an oxoacid (organic or inorganic) in which at least one hydroxyl group () is replaced by an alkoxy group (), as in the substitution reaction of a carboxylic acid and an alcohol. Glycerides are f ...
. Also present are other compounds in cuticular wax which are not VLCFA derivatives, such as terpenoids,
flavonoids Flavonoids (or bioflavonoids; from the Latin word ''flavus'', meaning yellow, their color in nature) are a class of polyphenolic secondary metabolites found in plants, and thus commonly consumed in the diets of humans. Chemically, flavonoids ...
, and
sterols Sterol is an organic compound with formula , whose molecule is derived from that of gonane by replacement of a hydrogen atom in position 3 by a hydroxyl group. It is therefore an alcohol of gonane. More generally, any compounds that contain the ...
, 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, and concludes with the extension of these chains in the
endoplasmic reticulum The endoplasmic reticulum (ER) is, in essence, the transportation system of the eukaryotic cell, and has many other important functions such as protein folding. It is a type of organelle made up of two subunits – rough endoplasmic reticulum ( ...
of epidermal cells. An important catalyzer thought to be in this process is the fatty acid elongase (FAE) complex. To form cuticular wax components, VLCFAs are modified through either two identified pathways, an acyl reduction pathway or a
decarbonylation Decarbonylation is a type of organic reaction that involves loss of CO. It is often an undesirable reaction since it represents a degradation. In the chemistry of metal carbonyls, decarbonylation describes a substitution process, whereby a CO lig ...
pathway. In the acyl reduction pathway, a
reductase A reductase is an enzyme that catalyzes a reduction reaction. Examples * 5α-Reductase * 5β-Reductase * Dihydrofolate reductase * HMG-CoA reductase * Methemoglobin reductase * Ribonucleotide reductase * Thioredoxin reductase * ''E. coli'' ni ...
converts VLCFAs into primary alcohols, which can then be converted to wax esters through a wax synthase. In the decarbonylation pathway, aldehydes are produced and decarbonylated to form alkanes, and can be subsequently oxidized to form secondary alcohols and ketones. The wax biosynthesis pathway ends with the transportation of the wax components from the endoplasmic reticulum to the epidermal surface.


Functions

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. 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 ''Nelumbo nucifera'', also known as sacred lotus, Laxmi lotus, Indian lotus, or simply lotus, is one of two extant species of aquatic plant in the family Nelumbonaceae. It is sometimes colloquially called a water lily, though this more often ref ...
'') have ultra-hydrophobic and self-cleaning properties that have been described by Barthlott and Neinhuis (1997). The
lotus effect The lotus effect refers to self-cleaning properties that are a result of ultrahydrophobicity as exhibited by the leaves of ''Nelumbo'', the lotus flower. Dirt particles are picked up by water droplets due to the micro- and nanoscopic architect ...
has applications in
biomimetic Biomimetics or biomimicry is the emulation of the models, systems, and elements of nature for the purpose of solving complex human problems. The terms "biomimetics" and "biomimicry" are derived from grc, βίος (''bios''), life, and μίμησ ...
technical materials. Dehydration protection provided by a maternal cuticle improves offspring fitness in the moss ''Funaria hygrometrica'' and in the sporophytes of all
vascular plant Vascular plants (), also called tracheophytes () or collectively Tracheophyta (), form a large group of land plants ( accepted known species) that have lignified tissues (the xylem) for conducting water and minerals throughout the plant. They al ...
s. In
angiosperm Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (), commonly called angiosperms. The term "angiosperm" is derived from the Greek words ('container, vessel') and ('seed'), and refers to those plants th ...
s the cuticle tends to be thicker on the top of the leaf ( adaxial surface), but is not always thicker. The leaves of
xerophytic A xerophyte (from Greek ξηρός ''xeros'' 'dry' + φυτόν ''phuton'' 'plant') is a species of plant that has adaptations to survive in an environment with little liquid water, such as a desert such as the Sahara or places in the Alps or the ...
plants adapted to drier climates have more equal cuticle thicknesses compared to those of
mesophytic Mesophytes are terrestrial plants which are neither adapted to particularly dry nor particularly wet environments. An example of a mesophytic habitat would be a rural temperate meadow, which might contain goldenrod, clover, oxeye daisy, and '' Rosa ...
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".


Evolution

The plant cuticle is one of a series of
innovations Innovation is the practical implementation of ideas that result in the introduction of new goods or services or improvement in offering goods or services. ISO TC 279 in the standard ISO 56000:2020 defines innovation as "a new or changed entit ...
, together with
stoma In botany, a stoma (from Greek ''στόμα'', "mouth", plural "stomata"), also called a stomate (plural "stomates"), is a pore found in the epidermis of leaves, stems, and other organs, that controls the rate of gas exchange. The pore is bor ...
ta,
xylem Xylem is one of the two types of transport tissue in vascular plants, the other being phloem. The basic function of xylem is to transport water from roots to stems and leaves, but it also transports nutrients. The word ''xylem'' is derived from ...
and
phloem Phloem (, ) is the living tissue in vascular plants that transports the soluble organic compounds made during photosynthesis and known as ''photosynthates'', in particular the sugar sucrose, to the rest of the plant. This transport process is ...
and intercellular spaces in
stem Stem or STEM may refer to: Plant structures * Plant stem, a plant's aboveground axis, made of vascular tissue, off which leaves and flowers hang * Stipe (botany), a stalk to support some other structure * Stipe (mycology), the stem of a mushro ...
and later
leaf A leaf ( : leaves) is any of the principal appendages of a vascular plant stem, usually borne laterally aboveground and specialized for photosynthesis. Leaves are collectively called foliage, as in "autumn foliage", while the leaves, ste ...
mesophyll tissue, that
plants Plants are predominantly photosynthetic eukaryotes of the kingdom Plantae. Historically, the plant kingdom encompassed all living things that were not animals, and included algae and fungi; however, all current definitions of Plantae exclude ...
evolved more than 450 million years ago during the transition between life in water and life on land. 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
stoma In botany, a stoma (from Greek ''στόμα'', "mouth", plural "stomata"), also called a stomate (plural "stomates"), is a pore found in the epidermis of leaves, stems, and other organs, that controls the rate of gas exchange. The pore is bor ...
tal guard cells, which regulate the rates of transpiration and CO2 exchange.


References

{{Botany Plant anatomy Plant physiology