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Plants prevent uncontrolled water loss by synthesizing, depositing and maintaining a hydrophobic layer over their primary aerial organs—the plant cuticle. Quercus coccifera L. can plastically respond to environmental conditions at the cuticular level. When exposed to hot summer conditions with high vapour-pressure deficit (VPD) and intense solar radiation (Mediterranean atmospheric conditions; MED), this plant species accumulates leaf cuticular waxes even over the stomata, thereby decreasing transpirational water loss. However, under mild summer conditions with moderate VPD and regular solar radiation (temperate atmospheric conditions; TEM), this effect is sharply reduced. Despite the ecophysiological importance of the cuticular waxes of Q. coccifera , the wax composition and its contribution to avoiding uncontrolled dehydration remain unknown. Thus, we determined several leaf traits for plants exposed to both MED and TEM conditions. Further, we qualitatively and quantitatively investigated the cuticular lipid composition by gas chromatography. Finally, we measured the minimum leaf conductance (gmin) as an indicator of the efficacy of the cuticular transpiration barrier. The MED leaves were smaller, stiffer and contained a higher load of cuticular lipids than TEM leaves. The amounts of leaf cutin and cuticular waxes of MED plants were 1.4 times and 2.6 times higher than that found for TEM plants, respectively. In detail, MED plants produced higher amounts of all compound classes of cuticular waxes, except for the equivalence of alkanoic acids. Although MED leaves contained higher cutin and cuticular wax loads, the gmin was not different between the two habitats. Our findings suggest that the qualitative accumulation of equivalent cuticular waxes might compensate for the higher wax amount of MED plants, thereby contributing equally to the efficacy of the cuticular transpirational barrier of Q. coccifera. In conclusion, we showed that atmospheric conditions profoundly affect the cuticular lipid composition of Q. coccifera leaves, but do not alter its transpiration barrier properties. [ABSTRACT FROM AUTHOR]

Background and Aims The cuticle of a limited number of plant species contains cutan, a chemically highly resistant biopolymer. As yet, the biosynthesis of cutan is not fully understood. Attempting to further unravel the origin of cutan, we analysed the chemical composition of enzymatically isolated cuticular membranes of Agave americana leaves. Methods Cuticular waxes were extracted with organic solvents. Subsequently, the dewaxed cuticular membrane was depolymerized by acid-catalysed transesterification yielding cutin monomers and cutan, a non-hydrolysable, cuticular membrane residue. The cutan matrix was analysed by thermal extraction, flash pyrolysis and thermally assisted hydrolysis and methylation to elucidate the monomeric composition and deduce a putative biosynthetic origin. Key Results According to gas chromatography–mass spectrometry analyses, the cuticular waxes of A. americana contained primarily very-long-chain alkanoic acids and primary alkanols dominated by C32, whereas the cutin biopolyester of A. americana mainly consisted of 9,10-epoxy ω-hydroxy and 9,10,ω-trihydroxy C18 alkanoic acids. The main aliphatic cutan monomers were alkanoic acids, primary alkanols, ω-hydroxy alkanoic acids and alkane-α,ω-diols ranging predominantly from C28 to C34 and maximizing at C32. Minor contributions of benzene-1,3,5-triol and derivatives suggested that these aromatic moieties form the polymeric core of cutan, to which the aliphatic moieties are linked via ester and possibly ether bonds. Conclusions High similarity of aliphatic moieties in the cutan and the cuticular wax component indicated a common biosynthetic origin. In order to exclude species-specific peculiarities of A. americana and to place our results in a broader context, cuticular waxes, cutin and cutan of Clivia miniata , Ficus elastica and Prunus laurocerasus leaves were also investigated. A detailed comparison showed compositional and structural differences, indicated that cutan was only found in leaves of perennial evergreen A. americana and C. miniata , and made clear that the phenomenon of cutan is possibly less present in plant species than suggested in the literature. [ABSTRACT FROM AUTHOR]

The efficacy of the cuticular transpiration barrier and its resistance to elevated temperatures are significantly higher in a typical water-saver than in a water-spender plant growing in hot desert. Water-saver and water-spender strategies are successful adaptations allowing plants to cope with the limitations of hot desert habitats. We investigated whether the efficacy of the cuticular transpiration barrier and its susceptibility to high temperatures are ecophysiological traits differentially developed in the water-spender Citrullus colocynthis and the water-saver Phoenix dactylifera. Minimum leaf conductance (g min) at 25 °C was six times lower in P. dactylifera (1.1×10–5 m s–1) than in C. colocynthis (6.9×10–5 m s–1). Additionally, g min in the range 25–50 °C did not change in P. dactylifera but increased by a factor of 3.2 in C. colocynthis. Arrhenius formalism applied to the C. colocynthis g min led to a biphasic graph with a steep increase at temperatures ≥35 °C, whereas for P. dactylifera the graph was linear over all temperatures. Leaf cuticular wax coverage amounted to 4.2±0.4 µg cm–2 for C. colocynthis and 29.4±4.2 µg cm–2 for P. dactylifera. In both species, waxes were mainly composed of very-long-chain aliphatics. Midpoints of the wax melting ranges of P. dactylifera and C. colocynthis were 80 °C and 73 °C, respectively. We conclude that in P. dactylifera a particular wax and cutin chemistry prevents the rise of g min at elevated temperatures. [ABSTRACT FROM AUTHOR]

When the stomata are closed under drought, the only route for water loss from the leaf interior to the atmosphere is across the cuticle. Thus, the extent of cuticular transpiration in relation to the reservoirs of water in the plant and the water acquisition from the soil determines the fitness and survival of the plant. It is, therefore, widely assumed that the cuticular water permeability of plants regularly experiencing drought is comparatively low and, thus, adapted to the environment. To test this hypothesis, 382 measurements of cuticular permeability from 160 species were extracted from the literature published between 1996 and 2017. The data had been produced either by using isolated cuticles and astomatous leaf sides or by measuring the minimum leaf conductance under conditions assumed to induce maximum stomatal closure. The species were assigned to 11 life form groups. Except for two particular cases (epiphytes, and climbers and lianas), the cuticular permeabilities of all groups either did not differ significantly or the available data did not allow a statistical test. In conclusion, present knowledge either does not support the hypothesis that ecological adaptions of cuticular water permeability exist or the available data are insufficient to test it. [ABSTRACT FROM AUTHOR]

Plant cuticular waxes play a crucial role in limiting nonstomatal water loss. The goal of this study was to localize the transpiration barrier within the layered structure of cuticles of eight selected plant species and to put its physiological function into context with the chemical composition of the intracuticular and epicuticular wax layers. Four plant species (Tetrastigma voinierianum, Oreopanax guatemalensis, Monstera deliciosa, and Schefflera elegantissima) contained only very-long-chain fatty acid (VLCFA) derivatives such as alcohols, alkyl esters, aldehydes, and alkanes in their waxes. Even though the epicuticular and intracuticular waxes of these species had very similar compositions, only the intracuticular wax was important for the transpiration barrier. In contrast, four other species (Citrus aurantium, Euonymus japonica, Clusia flava, and Garcinia spicata) had waxes containing VLCFA derivatives, together with high percentages of alicyclic compounds (triterpenoids, steroids, or tocopherols) largely restricted to the intracuticular wax layer. In these species, both the epicuticular and intracuticular waxes contributed equally to the cuticular transpiration barrier. We conclude that the cuticular transpiration barrier is primarily formed by the intracuticular wax but that the epicuticular wax layer may also contribute to it, depending on species-specific cuticle composition. The barrier is associated mainly with VLCFA derivatives and less (if at all) with alicyclic wax constituents. The sealing properties of the epicuticular and intracuticular layers were not correlated with other characteristics, such as the absolute wax amounts and thicknesses of these layers. [ABSTRACT FROM AUTHOR]

Previously, it was shown that β-ketoacyl-coenzyme A synthase ECERIFERUM6 (CER6) is necessary for the biosynthesis of very-long-chain fatty acids with chain lengths beyond C28 in tomato (Solatium lycopersicum) fruits and C26 in Arabidopsis (Ambidopsis thaliana) leaves and the pollen coat. CER6 loss of function in Arabidopsis resulted in conditional male sterility, since pollen coat lipids are responsible for contact-mediated pollen hydration. In tomato, on the contrary, pollen hydration does not rely on pollen coat lipids. Nevertheless, mutation in S1CER6 impairs fertility and floral morphology. Here, the contribution of SICER6 to the sexual reproduction and flower development of tomato was addressed. Cytological analysis and cross-pollination experiments revealed that the slcer6 mutant has male sterility caused by (1) hampered pollen dispersal and (2) abnormal tapetum development. SlCER6 loss of function provokes a decrease of n- and iso-alkanes with chain lengths of C27 or greater and of anteiso-alkanes with chain lengths of C28 or greater in flower cuticular waxes, but it has no impact on flower cuticle ultrastructure and cutin content. Expression analysis confirmed high transcription levels of SlCER6 in the anther and the petal, preferentially in sites subject to epidermal fusion. Hence, wax deficiency was proposed to be the primary reason for the flower fusion phenomenon in tomato. The S1CER6 substrate specificity was revisited. It might be involved in elongation of not only linear but also branched very-long-chain fatty acids, leading to production of the corresponding alkanes. SlCER6 implements a function in the sexual reproduction of tomato that is different from the one in Arabidopsis: S1CER6 is essential for the regulation of timely tapetum degradation and, consequently, microgametogenesis. [ABSTRACT FROM AUTHOR]

The permeabilities of amino acids for isolated cuticular membranes of ivy (Hedera helix L.) were measured at different pH. Cuticular permeances were lowest for the zwitterionic form at pH 6, followed by the cationic form at pH 1. Highest permeances were obtained for the anionic form at pH 11. Permeances were not correlated with octanol/water partition coefficients and decreased at a given pH with increasing molar volume of the solute. This finding suggests that permeation takes place in the polar cuticular pathways. The effect of pH on the cuticular transport properties was analysed according to the porous membrane model considering the polyelectrolytic character of the cuticle in terms of porosity, tortuosity, and size selectivity of the aqueous cuticular pathway which is altered by pH. An increase of water content and permeability of the cuticular membrane was caused by the dissociation of weak acidic groups with increasing pH leading to a swelling of the cuticle induced by fixed negative charges. In addition, the pH-dependent size of the hydration shell of the amino acids was identified as a secondary factor explaining the variability of cuticular permeances. [ABSTRACT FROM PUBLISHER]

Crown gall tumors induced by Agrobacterium tumefaciens represent a sink that has to be provided with nutrients and water by the host plant. The lack of an intact epidermis or cuticle results in uncontrolled loss of water. However, neither the tumor nor the host plant displays wilting. This phenomenon points to drought adaptation in both tumors and the crown gall host plant. To understand the underlying molecular mechanisms of protection against desiccation the gene expression pattern of Arabidopsis (Arabidopsis thaliana) tumors was integrated with the profile of stress metabolites: Arabidopsis tumors accumulated high amounts of abscisic acid (ABA), the ethylene precursor aminocyclopropyl carboxylic acid, osmoprotectants, and form a suberized periderm-like protective layer. Suberization of the outer tumor cell layers most likely is mediated by ABA since external application of ABA induced suberization of Arabidopsis roots. However, the expression level of the classical marker genes, known to respond to drought stress and/or ABA, was lower in tumors. Instead another set of drought and/or ABA- inducible genes was more highly transcribed. Elevated transcription of several ABA-dependent aquaporin genes might indicate that ABA controls the water balance of the tumor. The retarded tumor growth on abi and aba mutant plants underlined the importance of a tumor-specific ABA signaling pathway. Taken together, we propose that ABA is an important signal for protection of tumors against desiccation and thus supports tumor development. [ABSTRACT FROM AUTHOR]

Cuticular waxes play a pivotal role in limiting transpirational water loss across the primary plant surface. The astomatous fruits of the tomato (Lycopersicon esculentum) 'MicroTom' and its lecer6 mutant, defective in a β-ketoacyl-coenzyrne A synthase, which is involved in very-long-chain fatty acid elongation, were analyzed with respect to cuticular wax load and composition. The developmental course of fruit ripening was followed. Both the 'MicroTom' wild type and lecer6 mutant showed similar patterns of quantitative wax accumulation, although exhibiting considerably different water permeances. With the exception of immature green fruits, the Iecer6 mutant exhibited about 3- to 8-fold increased water loss per unit time and fruit surface area when compared to the wild type. This was not the case with immature green fruits. The differences in final cuticular barrier properties of tomato fruits in both lines were fully developed already in the mature green to early breaker stage of fruit development. When the qualitative chemical composition of fruit cuticular waxes during fruit ripening was investigated, the deficiency in a β-ketoacyl-coenzyme A synthase in the lecer6 mutant became discernible in the stage of mature green fruits mainly by a distinct decrease in the proportion of n-alkanes of chain lengths > C28 and a concomitant increase in cyclic triterpenoids. This shift in cuticular wax biosynthesis of the lecer6 mutant appears to be responsible for the simultaneously occurring increase of water permeance. Changes in cutin composition were also investigated as a function of developmental stage. This integrative functional approach demonstrates a direct relationship between cuticular transpiration barrier proper- ties and distinct chemical modifications in cuticular wax composition during the course of tomato fruit development. [ABSTRACT FROM AUTHOR]

The water permeability of cuticles isolated from the leaves of 14 plant species was measured at temperatures from 10 °C to 55 °C at 5 K intervals. Permeances increased slightly with temperatures ≤35 °C and drastically in the higher temperature range. The data were analysed according to the Arrhenius formalism which led to distinct plots for the lower and higher temperature range, respectively. Activation energies of permeation for the lower temperature range were estimated to amount to 15.2–52.5 kJ mol−1, at higher temperature activation energies ranged from 52.2–117.3 kJ mol−1. This thermodynamics approach is used for further elucidating the pathway taken by water across the plant cuticle. Based on the results of this study it is hypothesized that the diffusion of water occurs along polysaccharide strands crossing the cuticle and that the transport properties of these polar pathways change with temperature. [ABSTRACT FROM PUBLISHER]

The permeability of astomatous leaf cuticular membranes of Hedera helix L. was measured for uncharged hydrophilic (octanol/water partition coefficient log KO/W ≤0) and lipophilic compounds (log KO/W >0). The set of compounds included lipophilic plant protection agents, hydrophilic carbohydrates, and the volatile compounds water and ethanol. Plotting the mobility of the model compounds versus the molar volume resulted in a clear differentiation between a lipophilic and a hydrophilic pathway. The size selectivity of the lipophilic pathway was described by the free volume theory. The pronounced tortuosity of the diffusional path was caused by cuticular waxes, leading to an increase in permeance for the lipophilic compounds after wax extraction. The size selectivity of the hydrophilic pathway was described by hindered diffusion in narrow pores of molecular dimensions. A distinct increase in size selectivity was observed for hydrophilic compounds with a molar volume higher than 110 cm3 mol−1. Correspondingly, the size distribution of passable hydrophilic pathways was interpreted as a normal distribution with a mean pore radius of 0.3 nm and a standard deviation of 0.02 nm. The increased permeance of the hydrophilic compounds by the removal of cuticular waxes was attributed to an increase in the porosity, a decrease in the tortuosity, and a widening of the pore size distribution. Cuticular transpiration resulted from the permeation of water across the hydrophilic pathway. The far-reaching implications of two parallel pathways for the establishment of correlations between cuticular structure, chemistry, and function are discussed. [ABSTRACT FROM PUBLISHER]

In the cuticular wax mixtures from leaves of pea (Pisum sativum) cv Avanta, cv Lincoln, and cv Maiperle, more than 70 individual compounds were identified. The adaxial wax was characterized by very high amounts of primary alcohols (71%), while the abaxial wax consisted mainly of alkanes (73%). An aqueous adhesive of gum arabic was employed to selectively sample the epicuticular wax layer on pea leaves and hence to analyze the composition of epicuticular crystals exposed at the outermost surface of leaves. The epicuticular layer was found to contain 74% and 83% of the total wax on adaxial and abaxial surfaces, respectively. The platelet-shaped crystals on the adaxial leaf surface consisted of a mixture dominated by hexacosanol, accompanied by substantial amounts of octacosanol and hentriacontane. In contrast, the ribbon-shaped wax crystals on the abaxial surface consisted mainly of hentriacontane (63%), with approximately 5% each of hexacosanol and octacosanol being present. Based on this detailed chemical analysis of the wax exposed at the leaf surface, their importance for early events in the interaction with host-specific pathogenic fungi can now be evaluated. On adaxial surfaces, approximately 80% of Erysiphe pisi spores germinated and 70% differentiated appressoria. In contrast, significantly lower germination efficiencies (57%) and appressoria formation rates (49%) were found for abaxial surfaces. In conclusion, the influence of the physical structure and the chemical composition of the host surface, and especially of epicuticular leaf waxes, on the prepenetration processes of biotrophic fungi is discussed. [ABSTRACT FROM AUTHOR]

Cuticular waxes play a pivotal role in limiting transpirational water loss across the plant surface. The correlation between the chemical composition of the cuticular waxes and their function as a transpiration barrier is still unclear. In the present study, intact tomato fruits (Lycopersicon esculentum) are used, due to their astomatous surface, as a novel integrative approach to investigate this composition– function relationship: wax amounts and compositions of tomato were manipulated before measuring unbiased cuticular transpiration. First, successive mechanical and extractive wax‐removal steps allowed the selective modification of epi‐ and intracuticular wax layers. The epicuticular film consisted exclusively of very‐long‐chain aliphatics, while the intracuticular compartment contained large quantities of pentacyclic triterpenoids as well. Second, applying reverse genetic techniques, a loss‐of‐function mutation with a transposon insertion in a very‐long‐chain fatty acid elongase β‐ketoacyl‐CoA synthase was isolated and characterized. Mutant leaf and fruit waxes were deficient in n‐alkanes and aldehydes with chain lengths beyond C30, while shorter chains and branched hydrocarbons were not affected. The mutant fruit wax also showed a significant increase in intracuticular triterpenoids. Removal of the epicuticular wax layer, accounting for one‐third of the total wax coverage on wild‐type fruits, had only moderate effects on transpiration. By contrast, reduction of the intracuticular aliphatics in the mutant to approximately 50% caused a 4‐fold increase in permeability. Hence, the main portion of the transpiration barrier is located in the intracuticular wax layer, largely determined by the aliphatic constituents, but modified by the presence of triterpenoids, whereas epicuticular aliphatics play a minor role. [ABSTRACT FROM PUBLISHER]

The water permeability of the leaves of three deciduous plants (Acer campestre, Fagus sylvatica, Quercus petraea) and two evergreen plants (Hedera helix, Ilex aquifolium) was analysed in order to assess its role as a mechanism of drought resistance. Cuticular permeances were determined by measurement of the water loss through adaxial, astomatous leaf surfaces. Minimum conductances after complete stomatal closure were obtained by leaf drying curves. The comparison of the water permeabilities determined with these two experimental systems revealed good agreement in the case of Acer, Fagus, Quercus, and Ilex. For Hedera the minimum conductance was 3‐fold higher than the cuticular permeance indicating a significant contribution of residual stomatal transpiration. The leaf water potential was measured as a function of water content and analysed by pressure–volume curves. The influence of water potential as a component of the driving force for transpirational water loss was assessed in order to identify modifications of the cuticular barrier by the leaf water content. The ecophysiological meaning of the water relations parameters describing transpiration under drought conditions (cuticular transpiration, minimum transpiration, residual stomatal transpiration, effect of leaf water content on transpiration) and the water relations parameters derived from pressure–volume curves (osmotic potential at full saturation, turgor loss point, bulk modulus of elasticity) are discussed with regard to adaptations for drought resistance. [ABSTRACT FROM PUBLISHER]

The behaviour of (semi‐)volatile organic compounds at the interface between the leaf surface and the atmosphere was investigated by finite‐element numerical simulation. Three model systems with increasing complexity and closeness to the real situation were studied. The three‐dimensional model systems were translated into appropriate grid structures and diffusive and convective transport in the leaf/atmosphere interface was simulated. Fenpropimorph (cis‐4‐[3‐(4‐tert‐butylphenyl)‐2‐methylpropyl]‐2,6‐dimethylmorpholine) and Kresoxim‐methyl ((E)‐methyl‐2‐methoxyimino‐2‐[2‐(o‐tolyloxy‐methyl)phenyl] acetate) were used as model compounds. The simulation showed that under still and convective conditions the vapours emitted by a point source rapidly form stationary envelopes around the leaves. Vapour concentrations within these unstirred layers depend on the vapour pressure of the compound in question and on its affinity to the lipoid surface layers of the leaf (cuticular waxes, cutin). The rules deduced from the numerical simulation of organic vapour behaviour in the leaf/atmosphere interface are expected to help in assessing how (semi‐)volatile plant products (e.g. hormones, pheromones, secondary metabolites) and xenobiotics (e.g. pesticides, pollutants) perform on plant surfaces. [ABSTRACT FROM PUBLISHER]

The cuticle is the major barrier against uncontrolled water loss from leaves, fruits and other primary parts of higher plants. More than 100 mean values for water permeabilities determined with isolated leaf and fruit cuticles from 61 plant species are compiled and discussed in relation to plant organ, natural habitat and morphology. The maximum barrier properties of plant cuticles exceed that of synthetic polymeric films of equal thickness. Cuticular water permeability is not correlated to the thickness of the cuticle or to wax coverage. Relationships between cuticular permeability, wax composition and physical properties of the cuticle are evaluated. Cuticular permeability to water increases on the average by a factor of 2 when leaf surface temperature is raised from 15 °C to 35 °C. Organic compounds of anthropogenic and biogenic origin may enhance cuticular permeability. The pathway taken by water across the cuticular transport barrier is reviewed. The conclusion from this discussion is that the bulk of water diffuses as single molecules across a lipophilic barrier while a minor fraction travels along polar pores. Open questions concerning the mechanistic understanding of the plant cuticular transport barrier and the role the plant cuticle plays in ensuring the survival and reproductive success of an individual plant are indicated. [ABSTRACT FROM AUTHOR]

Provides an overview of the depth and breadth of plant stress biology with updates on knowledge in selected fields of the physiology and ecology of stress in plants. Sources of plant stress; Stress tolerance of plants.