Wound-induced green leaf volatiles cause the release of acetylated derivatives and a terpenoid in maize

Green leaf volatiles (GLVs), generally occurring C6 alcohols, aldehydes and acetates from plants, play an important role in plant-plant communication. These compounds induce intact plants to produce jasmonic acid, and induce defense-related gene expression and the release of volatile compounds. Here, we address wound-induced GLVs cause the release of acetylated derivatives and a terpenoid, (E)-4,8-dimethylnona-1,3,7-triene (DMNT) in intact maize, which may be a type of plant-plant interaction mediated by airborne GLVs. Upon exposure of intact maize seedlings to wound-induced GLVs, (Z)-3-hexenyl acetate was consistently the most abundant compound released. Exogenous application of individual alcohols and aldehydes mostly resulted in the release of corresponding acetate esters. C6-alcohols with a double bond between the second and third, or the third and fourth carbon atoms, C5- or C6-aldehydes, and (Z)-3-hexenyl acetate triggered the release of DMNT. When (Z)-3-hexenyl acetate and hexyl acetate were used to treat maize seedlings, they were recovered from the plants. These data demonstrated that: (1) apart from direct adsorption and re-release of acetate esters, absorption and conversion of exogenous alcohols and aldehydes into acetate esters occurred, and (2) DMNT was induced by a range of aldehydes and unsaturated alcohols.     

Green leaf volatiles: hydroperoxide lyase pathway of oxylipin metabolism

Green leaf volatiles (GLVs) are C(6) aldehydes, alcohols, and their esters formed through the hydroperoxide lyase pathway of oxylipin metabolism. Plants start to form GLVs after disruption of their tissues and after suffering biotic or abiotic stresses. GLV formation is thought to be regulated at the step of lipid-hydrolysis, which provides free fatty acids to the pathway. Recently, studies dissecting the physiological significance of GLVs in plants have emerged, and it has been postulated that GLVs are important molecules both for signaling within and between plants and for allowing plants and other organisms surrounding them to recognize or compete with each other.     

Pine shoot beetle, Tomicus piniperda (Col., Scolytidae), responses to common green leaf volatiles

We tested the hypothesis that green leaf volatiles (GLVs) disrupt the response of overwintered pine shoot beetles, Tomicus piniperda (L.) to multiple-funnel traps baited with the attractive host volatile α-pinene. A combination of four GLV alcohols, 1-hexanol ( E )-2-hexen-1-ol ( Z )-2-hexen-1-ol, and ( Z )-3-hexen-1-ol, caused 54 and 36% reduction in the number of pine shoot beetles captured in two separate trapping experiments. Similarly, a combination of the four alcohols plus two GLV aldehydes, hexanal and ( E )-2-hexenal, caused 38% reduction in the number of pine shoot beetles captured compared with α-pinene alone. A blend of the two GLV aldehydes was not disruptive. None of the four GLV alcohols nor the two GLV aldehydes were disruptive when tested individually. The finding that the blend of four GLV alcohols reduced attraction of T. piniperda supports the general hypothesis that GLVs common to nonhost angiosperms are disruptive to conifer-attacking bark beetles (Scolytidae).     

绿叶挥发物产生特征及其生态生理作用研究进展

该文系统综述了植物绿叶挥发物(green leaf volatiles, GLVs)的形成特征、对植物的生态生理作用及调控机制等方面的最新研究进展。GLVs是指植物经十八碳烷酸途径过氧化物酶分支途径产生的含6个碳原子的醛、醇及其酯。除叶片外, 植物的根、茎、果实、种子等部位均可以合成该类化合物, 合成调控存在转录和转录后水平的调控。在特定植物中, GLVs合成及其组分还受到植物生长发育阶段和生长季节等外源环境的影响。昆虫啃食、微生物感染以及有益真菌的定植等生物逆境与缺氮等物理逆境均具有诱导GLVs合成的作用。除了参与植物特有气味形成外, GLVs在植物直接和间接防御应答中发挥着重要的作用。GLVs不仅具有抑制微生物和多种昆虫繁殖的作用, 还具有诱导多种防御化合物合成、预置(prime)有关信号途径的作用。基于GLVs在植物界的广泛存在性和在防御应答中的多层次作用, 该文作者提出了GLVs在道地药材品质形成中的潜在作用及开展相关研究的必要性和紧迫性。     

Traumatin- and Dinortraumatin-containing Galactolipids in Arabidopsis: THEIR FORMATION IN TISSUE-DISRUPTED LEAVES AS COUNTERPARTS OF GREEN LEAF VOLATILES*

Green leaf volatiles (GLVs) consisting of six-carbon aldehydes, alcohols, and their esters, are biosynthesized through the action of fatty acid hydroperoxide lyase (HPL), which uses fatty acid hydroperoxides as substrates. GLVs form immediately after disruption of plant leaf tissues by herbivore attacks and mechanical wounding and play a role in defense against attackers that attempt to invade through the wounds. The fates and the physiological significance of the counterparts of the HPL reaction, the 12/10-carbon oxoacids that are formed from 18/16-carbon fatty acid 13-/11-hydroperoxides, respectively, are largely unknown. In this study, we detected monogalactosyl diacylglycerols (MGDGs) containing the 12/10-carbon HPL products in disrupted leaf tissues of Arabidopsis, cabbage, tobacco, tomato, and common bean. They were identified as an MGDG containing 12-oxo-9-hydroxy-(E)-10-dodecenoic acid and 10-oxo-7-hydroxy-(E)-8-decenoic acid and an MGDG containing two 12-oxo-9-hydroxy-(E)-10-dodecenoic acids as their acyl groups. Analyses of Arabidopsis mutants lacking HPL indicated that these MGDGs were formed enzymatically through an active HPL reaction. Thus, our results suggested that in disrupted leaf tissues, MGDG-hydroperoxides were cleaved by HPL to form volatile six-carbon aldehydes and non-volatile 12/10-carbon aldehyde-containing galactolipids. Based on these results, we propose a novel oxylipin pathway that does not require the lipase reaction to form GLVs.     

Auxin-induced leaf blade expansion in Arabidopsis requires both wounding and detachment

Elevation of leaf auxin (indole-3-acetic acid; IAA) levels in intact plants has been consistently found to inhibit leaf expansion whereas excised leaf strips grow faster when treated with IAA. Here we test two hypothetical explanations for this difference in growth sensitivity to IAA by expanding leaf tissues in vivo versus in vitro. We asked if, in Arabidopsis, IAA-induced growth of excised leaf strips results from the wounding required to excise tissue and/or results from detachment from the plant and thus loss of some shoot or root derived growth controlling factors. We tested the effect of a range of exogenous IAA concentrations on the growth of intact attached, wounded attached, detached intact, detached wounded as well as excised leaf strips. After 24 h, the growth of intact attached, wounded attached, and detached intact leaves was inhibited by IAA concentrations as little as 1 µM in some experiments. Growth of detached wounded leaves and leaf strips was induced by IAA concentrations as low as 10 µM. Stress, in the form of high light, increased the growth response to IAA by leaf strips and reduced growth inhibition response by intact detached leaves. Endogenous free IAA content of intact attached leaves and excised leaf strips was found not to change over the course of 24 h. Together these results indicate growth induction of Arabidopsis leaf blade tissue by IAA requires both substantial wounding as well as detachment from the plant and suggests in vivo that IAA induces parallel pathways leading to growth inhibition.     

Emission of herbivore-induced volatiles in absence of a herbivore--response of Zea mays to green leaf volatiles and terpenoids

Green leaf volatiles (GLV), a series of saturated and monounsaturated six-carbon aldehydes, alcohols, and esters are emitted by plants upon mechanical damage. Evidence is increasing that intact plants respond to GLV by activating their own defense mechanisms, thus suggesting that they function in plant-plant communication. The present paper demonstrates that exposure of maize plants to naturally occurring GLV, including (Z)-3-, (E)-2- and saturated derivatives, induce the emission of volatile blends typically associated with herbivory. Position or configuration of a double bond, but not the functional group of the GLV influenced the strength of the emissions. (Z)-3-Configured compounds elicited stronger responses than (E)-2- and saturated derivatives. The response to (Z)-3-hexen-1-ol increased linearly with the dose between 200 and 1000 nmol per plant. Not only the naturally occurring (E)-2-hexenal, but also (E)-2-pentenal and (E)-2-heptenal induced maize plants, although to a lesser extent. Externally applied terpenoids [(3E)-4,8-dimethyl-1,3,7-nonatriene, beta-caryophyllene, and (E)-beta-farnesene] did not significantly increase the total amount of inducible volatiles in maize. Of three tested maize cultivars Delprim and Pactol responded much stronger than Attribut. Recovery experiments in the presence and absence of maize plants demonstrated that large proportions of externally applied GLV were assimilated by the plants, whereas (3E)-4,8-dimethyl-1,3,7-nonatriene was recovered in much higher amounts. The results furthermore suggested that plants converted a part of the assimilated leaf aldehydes and alcohols to the respective acetates. We propose that GLV not only can alert neighboring plants, but may facilitate intra-plant information transfer and can help mediate the systemic defense response in a plant.     

Availability of assimilates and formation of aroma compounds in apples as affected by the fruit/leaf ratio

Availability of assimilates in apple trees ( Malus domestica cv. Jonagored) was affected by removing young fruits to obtain 3 ranges of fruit/leaf ratios with average values of 130, 268 and 381 fruits per kg leaf dry matter. Fruit analyses were carried out at fruit harvest and 4 times during a 3-week ripening period. The analyses included detection of volatile aroma components from the juice by headspace gas chromatography. At a low fruit/leaf ratio, higher concentrations of total dry matter, soluble solid and titrateable acids were found. The flesh was also firmer, and ethylene development proceeded at a lower rate and reached a lower maximum value. Aroma compounds consisted of ca 20% esters, 73% alcohols and 6% C-6 aldehydes. The production of butylacetate and hexylacetate, which were the dominating esters, peaked during the ripening period and was most pronounced at the lowest fruit/leaf ratios. At the last sampling date this was also the case for butanol, which was the dominating alcohol. Other esters and alcohols behaved similarly, while C-6 aldehydes showed no significant differences in the fruit/leaf ratio. We suggest that the greater availability of assimilates when internal competition is relieved at a low fruit/leaf ratio causes increased accumulation of fatty acid aroma precursors and aroma compounds as well as of sugars, acids and other compounds in the fruits.     

Green leaf volatiles protect maize (Zea mays) seedlings against damage from cold stress

Although considerable evidence has accumulated on the defensive activity of plant volatile organic compounds against pathogens and insect herbivores, less is known about the significance of volatile organic compounds emitted by plants under abiotic stress. Here, we report that green leaf volatiles (GLVs), which were previously shown to prime plant defences against insect herbivore attack, also protect plants against cold stress (4 °C). We show that the expression levels of several cold stress‐related genes are significantly up‐regulated in maize (Zea mays) seedlings treated with physiological concentrations of the GLV, (Z)‐3‐hexen‐1‐yl acetate (Z‐3‐HAC), and that seedlings primed with Z‐3‐HAC exhibit increased growth and reduced damage after cold stress relative to unprimed seedlings. Together, these data demonstrate the protective and priming effect of GLVs against cold stress and suggest an activity of GLVs beyond the activation of typical plant defence responses against herbivores and pathogens.     

Ontogenetic variation in chemical and physical characteristics of adaxial apple leaf surfaces

The reaction of plants to environmental factors often varies with developmental stage. It was hypothesized, that also the cuticle, the outer surface layer of plants is modified during ontogenesis. Apple plantlets, cv. Golden Delicious, were grown under controlled conditions avoiding biotic and abiotic stress factors. The cuticular wax surface of adaxial apple leaves was analyzed for its chemical composition as well as for its micromorphology and hydrophobicity just after unfolding of leaves ending in the seventh leaf insertion. The outer surface of apple leaves was formed by a thin amorphous layer of epicuticular waxes. Epidermal cells of young leaves exhibited a distinctive curvature of the periclinal cell walls resulting in an undulated surface of the cuticle including pronounced lamellae, with the highest density at the centre of cells. As epidermal cells expanded during ontogenesis, the upper surface showed only minor surface sculpturing and a decrease in lamellae. With increasing leaf age the hydrophobicity of adaxial leaf side decreased significantly indicated by a decrease in contact angle. Extracted from plants, the amount of apolar cuticular wax per area unit ranged from only 0.9 microgcm(-2) for the oldest studied leaf to 1.5 microgcm(-2) for the youngest studied leaf. Differences in the total amount of cuticular waxes per leaf were not significant for older leaves. For young leaves, triterpenes (ursolic acid and oleanolic acid), esters and alcohols were the main wax components. During ontogenesis, the proportion of triterpenes in total mass of apolar waxes decreased from 32% (leaf 1) to 13% (leaf 7); absolute amounts decreased by more than 50%. The proportion of wax alcohols and esters, and alkanes to a lesser degree, increased with leaf age, whereas the proportion of acids decreased. The epicuticular wax layer also contained alpha-tocopherol described for the first time to be present also in the epicuticular wax. The modifications in the chemical composition of cuticular waxes are discussed in relation to the varying physical characteristics of the cuticle during ontogenesis of apple leaves.     

13C-labelling patterns of green leaf volatiles indicating different dynamics of precursors in Brassica leaves

To investigate the dynamics of precursor compounds of green leaf volatiles (GLV)s and other biogenic compounds released by mechanically damaged Brassica oleracea leaves, plants were exposed for two consecutive 16h light phases to highly enriched (13)CO(2). Analysis by GC-MS indicated (1) biogenic compounds released upon wounding, (2) a different labelling pattern between and (3) within compounds, and (4) evidence for spatial heterogeneity of the precursor pool extrapolated from points (1)-(3). First, GLVs comprised C(5) and C(6) molecules, with the GLV pentenyl acetate being reported here for the first time from higher plants. Second, the labelling pattern found in most GLVs indicates a low turnover of the precursor alpha-linolenic acid. Moderate labelling of dimethyldisulphide indicates a connection to an active plastidic methyl pool closely connected to CO(2) fixation, and very weak labelling of terpenes indicates a constitutive monoterpene pool. Third, not all GLVs exhibit similarly strong labelling patterns (hexenyl acetate vs. hexyl acetate), indicating different precursors. As the labelling patterns of alcohol and acetate moieties in the esters differ, with only the former being strongly labelled, the precursor of the acetate moiety, acetyl-CoA, is likely to derive from a different cellular pool to that used in chloroplastic fatty acid synthesis, or was rapidly synthesised after the end of labelling. Fourth, the exceptionally high relative abundance of labelled GLV and the low concentration of unlabelled molecules are likely to occur because recently synthesized alpha-linolenic acid is bound in lipids that are organised in distinct areas, or are chemically different from the older lipids. They must be preferentially used as precursors.     

Activated oxygen‐mediated metabolic functions of leaf peroxisomes

Peroxisomes are subcellular organelles with an essentially oxidative type of metabolism. The presence in these organelles of superoxide dismutases and the generation of superoxide radicals (O₂^??) was first demonstrated in plant tissues and in recent years different experimental evidence has suggested the existence of cellular functions related to activated oxygen species. Some of these functions are analyzed in this work. In purified intact peroxisomes from pea (Pisum sativum L.) leaves, xanthine oxidase and urate oxidase were found to be present. The occurrence and the level of the metabolites xanthine, hypoxanthine, uric acid, and allantoin were studied in extracts of pea leaf peroxisomes by HPLC. Xanthine, uric acid, and allantoin were detected in peroxisomes. These results suggest a cellular role for leaf peroxisomes in the catabolism of purines. In peroxisomal membranes, 3 polypeptides (PMPs) with molecular masses of 18, 29 and 32 kDa, respectively, have been shown to generate superoxide radicals. These PMPs were purified from pea leaf peroxisomal membranes and characterized. While the 18- and 32-kDa PMPs use NADH as electron donor for O₂^?? production, the 29-kDa PMP was clearly dependent on NADPH. Very recently, the occurrence in pea leaf peroxisomes of all the enzymes of the ascorbate-glutathione cycle has been demonstrated. NADPH is required for the glutathione reductase activity of the cycle and this implies the reduction of NADP⁺ to NADPH. This recycling function could be carried out by the NADP-dependent glucose-6-phosphate dehydrogenase (G6PDH), 6-phosphogluconate dehydrogenase (6PGDH), and isocitrate dehydrogenase (ICDH). These 3 dehydrogenases have been demonstrated to be present in the matrix of pea leaf peroxisomes. The catabolism of purines, the superoxide-generating PMPs, the ascorbate-glutathione cycle, and the dehydrogenase-mediated recycling of NADPH, are activated oxygen roles of leaf peroxisomes that add to other functions previously known for peroxisomes from eukaryotic cells.     

Enhanced attraction of Plutella xylostella (Lepidoptera: Plutellidae) to pheromone-baited traps with the addition of green leaf volatiles

Diamondback moth, Plutella xylostella (L.) (Lepidoptera: Plutellidae), is one of the most serious pests of Brassicaceae crops worldwide. Electrophysiological and behavioral responses of P. xylostella to green leaf volatiles (GLVs) alone or together with its female sex pheromone were investigated in laboratory and field. GLVs 1-hexanol and (Z)-3-hexen-1-ol elicited strong electroantennographic responses from unmated male and female P. xylostella, whereas (Z)-3-hexenyl acetate only produced a relatively weak response. The behavioral responses of unmated moths to GLVs were further tested in Y-tube olfactometer experiments. (E)-2-Hexenal, (Z)-3-hexen-1-ol, and (Z)-3-hexenyl acetate induced attraction of males, reaching up to 50%, significantly higher than the response to the unbaited control arm. In field experiments conducted in 2008 and 2009, significantly more moths were captured in traps baited with synthetic sex pheromone with either (Z)-3-hexenyl acetate alone or a blend of (Z)-3-hexenyl acetate, (Z)-3-hexen-1-ol, and (E)-2-hexenal compared with sex pheromone alone and other blend mixtures. These results demonstrated that GLVs could be used to enhance the attraction of P. xylostella males to sex pheromone-baited traps.     

Effects of volatile compounds on arthrospore germination and mycelial growth ofGeotrichum candidum citrus race

Three groups of volatile compounds, i.e., alcohols, aldehydes and esters, were tested for their effects on arthrospore germination and mycelial growth ofGeotrichum candidum citrus race, the causal agent of citrus sour rot. Alcohols (heptanol, octanol, nonanol, decanol, geraniol, citronellol) at a concentration of 1.0 μl/ml showed 60% or more inhibitory effects on both germination and mycelial growth of this pathogen. Among aldehydes tested, only citral had an inhibitory effect of more than 50%, while esters had no effect. The chain length of straight-chain (C6–C12) alcohols correlated with inhibitory effect, and nonanol (C9) was most active. Treatment with alcohols or citral prior to inoculation reduced colonization and maceration of lemon peel by this pathogen by 70% or more. Results suggested that alcohols or citral can probably be used to prevent the development of citrus sour rot.     

Detection of plant volatiles after leaf wounding and darkening by proton transfer reaction "time-of-flight" mass spectrometry (PTR-TOF)

Proton transfer reaction-time of flight (PTR-TOF) mass spectrometry was used to improve detection of biogenic volatiles organic compounds (BVOCs) induced by leaf wounding and darkening. PTR-TOF measurements unambiguously captured the kinetic of the large emissions of green leaf volatiles (GLVs) and acetaldehyde after wounding and darkening. GLVs emission correlated with the extent of wounding, thus confirming to be an excellent indicator of mechanical damage. Transient emissions of methanol, C5 compounds and isoprene from plant species that do not emit isoprene constitutively were also detected after wounding. In the strong isoprene-emitter Populus alba, light-dependent isoprene emission was sustained and even enhanced for hours after photosynthesis inhibition due to leaf cutting. Thus isoprene emission can uncouple from photosynthesis and may occur even after cutting leaves or branches, e.g., by agricultural practices or because of abiotic and biotic stresses. This observation may have important implications for assessments of isoprene sources and budget in the atmosphere, and consequences for tropospheric chemistry.     

Very long chain alkylresorcinols accumulate in the intracuticular wax of rye (Secale cereale L.) leaves near the tissue surface

Alkylresorcinols (ARs) are bioactive compounds occurring in many members of the Poaceae, likely at or near the surface of various organs. Here, we investigated AR localization within the cuticular wax layers of rye (Secale cereale) leaves. The total wax mixture from both sides of the leaves was found to contain primary alcohols (71%), alkyl esters (11%), aldehydes (5%), and small amounts (<3%) of alkanes, steroids, secondary alcohols, fatty acids and unknowns. A homologous series of ARs (3%) was identified by GC-MS and comparison with a synthetic standard of nonadecylresorcinol. The alkyl side chains of the wax ARs contained odd numbers of carbons ranging from C19 to C27, with a prevalence of C21, C23 and C25. Waxes from both sides of the leaf, analyzed separately in a second experiment, comprised the same compound classes in similar relative amounts and with similar homolog patterns. Finally, the epicuticular and intracuticular wax layers were sampled separately from the abaxial side of the leaf. While ARs accounted for 2% of the intracuticular wax, they were not detectable in the epicuticular wax. The intracuticular wax was also slightly enriched in steroids, whereas the epicuticular layer contained more primary alcohols. All other wax constituents were distributed evenly between both wax layers.     

ACOMPARATIVE STUDY ON THE ELECTROANTENOGRAM RESPONSES OF THREE APHID SPECIES TO PLANT VOLATILES

Abstract  Electroantennogram responses (EAGs) to 39 plant volatiles from Aphis glycines, A. gossypii and Myzus persicae were recorded. The olfactory system of these three aphid species differentially responded to green leaf, aromatic, mono-and sequiterpene chemicals. The EAG depolarizatin to green leaf chemicals and their isomers were more pronounced than to terpenes, alcohols and aldehydes, 6 carbon atom compounds being the most effective. It was shown that there was certain difference in the olfactory spectra of these three aphid species to plant volatiles. Higher responses in A. glycines and A. gossypii were elicited by the leaf aldehydes and esters than by the corresponding alcohols, which are in contrast with those in M. persicue . The activity of aphid olfactory system was related to molecular structures of the compounds. The responses to saturated alcohols and aldehydes were higher than those to unsaturated. The responses of spring emigrants of A. glycines to terpene derivatives (alcohols, aldehydes and esters) were higher than those of A. gossypii , but were lower to terpene hydrocarbons. The dissimilarity in olfactory spectra of the aphid species reflected the evolution of their host plant selection specificity. In the evolution, their chemosensory system adapted to perceive host plants. The shapes of the EAGs evoked by the various components were consistently different, with the slowest recovery for geraniol, 1-decanol and (±)-β-cit-ronellol, and ware related to number of carbon atoms in straight chain saturated alcohols.     

Responses to ultraviolet-B radiation (280–315 nm) of pea (Pisum sativum) lines differing in leaf surface wax

To test the hypothesis that leaf surface wax influences plant responses to UV-B, 6 lines of cultivated pea (Pisum sativum L.), selected as having more or less wax, were grown at 0 or 6.5 kJ m^-2 day^-1 plant-weighted UV-B against a background of 850–950 μmol m^-2 s^-1 photosynthetically active radiation. In the 4 lines with least leaf surface wax the amount of wax on adaxial and abaxial leaf surfaces was increased following exposure to 6.5 kJ m^-2 day^-1 UV-B, but UV-B decreased surface wax in Scout, which had the greatest wax deposits. On the adaxial leaf surface, UV-B radiation caused a shift in wax composition from alcohols to esters and hydrocarbons and the ratio of short to long chain length alkyl ester homologues was increased. There was no evidence of a shortening in carbon chain length of hydrocarbons, primary alcohols or fatty acids due to UV-B and no significant correlation between wax amount and UV reflectance from leaves. UV-B induced significant increases in UV-absorbing compounds in the expanded leaves and buds of most lines. UV-B reduced the growth of all lines. Foliage area (leaves plus stipules) declined by 5–30%, plant dry weight by 12–30%, and plant height by 24–38%. Reductions in growth occurred in the absence of any changes in chlorophyll fluorescence or photosynthetic rate. UV-B also had no major effect on carbon allocation patterns. The effects of UV-B on growth appeared to be due to changes in tissue extension and expansion. Indeed, many of the responses to UV-B observed in this study of pea appear more consistent with indirect effects being expressed in developing tissues rather than through the direct action of UV-B on mature tissues. There was no evidence that wax amount or biochemistry was associated with the sensitivity of the lines to UV-B radiation. Furthermore, induction of pigments was not correlated with changes in growth. However, lines with the greatest constitutive amounts of pigments in unexpanded bud tissues were most tolerant of elevated UV-B.     

The composition of the lipid globules of Ilex aquifolium leaves and its variability with the age of the leaf and with season

The cytoplasmatic globules of Ilex aquifolium L. mainly consist of fatty acid esters of the triterpenoid alcohols α and β-amyrin. In very young leaves these are practically only palmitates. With increasing leaf age, increasing amounts of saturated and unsaturated C18 acids were found with comparatively high amounts of linoleic and linolenic acid in leaves of one year and older. Some accumulation of the triterpene precursor squalene was also observed with increasing age of the leaves. In addition to age effects, seasonal changes occur too. Independent of the age of the leaf, an increase in palmitic acid and a decrease in linoleic and especially linolenic acid was observed in the winter in mature leaves.     

Participation of Oryza sativa leaf wax in appressorium formation by Pyricularia oryzae

Appressorium formation of Pyricularia oryzae P2 on cover-glass coated with each of the components of rice leaf wax was examined. Wax esters, aldehydes and alcohols, having polar groups and low contact angles, promoted appressorium formation, but alkanes, non-polar molecules having high contact angles, had no effects. Germination of conidia, however, was not aftected with those constituents.