Effects of wax bloom variation in Brassica oleracea on foraging by a vespid wasp

Foraging by individual Polistes dominulus (Christ) (Hymenoptera: Vespidae) was observed and compared on Brassica oleracea L. plants that differed in surface wax bloom. Twenty-six wasps, previously trained to forage on plants for 4th instar Trichoplusia ni (Walker), were presented in the greenhouse with mixtures of plants with normal wax bloom or genetically reduced wax bloom, and on which T. ni caterpillars had been placed. During foraging, the wasps were observed to slip significantly more frequently from the leaf surfaces of normal-wax bloom plants than from reduced-wax bloom plants (129 vs 63 occurrences) and to retrieve significantly fewer pieces of caterpillar after attack on normal-wax bloom plants (151 vs 223 pieces). Altogether the wasps retrieved 333 caterpillar pieces from reduced-wax bloom plants and 248 pieces from normal-wax bloom plants. Despite these differences, the number of caterpillars attacked and killed did not differ between the two wax bloom types (116 vs 121), nor did handling time for individual attacks (time from contact with prey until prey piece was carried to the nest) (170±12.5 s vs 180±10.5 s). Thus, in contrast with previous reports for smaller predators, wax bloom variation in B. oleracea did not influence the effectiveness of P. dominulus as a predator of T. ni.     

Leaf surface wax and plant morphology of peas influence insect density

Insect predators and parasitoids adhere better, forage more effectively, and take more aphid prey on pea plants (Pisum sativum L.) (Leguminosae) with mutations that reduce the crystalline wax bloom on the plant surface. To assess the agronomic potential of this trait for pest management, abundance of pea aphids (Acyrthosiphon pisum L.) (Homoptera: Aphididae) and coccinellid predators, and percent parasitism of the aphids were evaluated on pea lines differing in wax bloom and plant architecture over two field seasons. Three pairs of pea lines were evaluated, each pair with a different architecture and differing within the pair in the amount of surface wax bloom (reduced or normal). The trials included plots treated with a narrow spectrum insecticide (pymetrozine) to reduce aphid populations and untreated controls. Reduced wax peas had significantly fewer aphids per plant in 2002 but not in 2003. Total natural enemy abundance was greater on reduced wax than on normal wax pea lines in both years of the study. Pymetrozine reduced aphid densities significantly in both years. Among the four pea lines evaluated for yield, seed yield per plant was affected by plant morphology and insecticide treatment. Yield was greatest on semileafless plants and on pymetrozine sprayed plots in both years. Yield of the reduced wax line in the semileafless background was similar to or exceeded yield in its normal wax sister line, suggesting that this morphological type was best for an agronomically viable reduced wax phenotype. Pea weevil (Bruchus pisorum L.) (Coleoptera: Bruchidae) damage to seed was overall more frequent on seeds from reduced wax varieties than from normal wax varieties. The results illustrate the trade‐offs associated with a reduced wax trait in peas but also show that certain combinations of reduced wax and gross morphology lead to reduced pea aphid populations and yields similar to those of normal wax peas.     

Dynamics of the wax bloom of a seasonal Namib Desert tenebrionid,Cauricara phalangium (Coleoptera: Adesmiini)

Summary The dynamics of the extracuticular was bloom in a winter species of tenebrionid beetle, Cauricara phalangium, from the Namib Desert were monitored in the field and under laboratory conditions. The beetles possessed a full complement of the white wax material after adult emergence. The amount of this material on the integument declined towards the end of the season. The wax bloom was regenerated in both the field and laboratory, with high temperatures and low humidities bringing about greatest renewal. End of the season decline appears tobe related to the senescence of these seasonal beetles. Water loss rates differed significantly for individuals collected in May, when fully bloomed, and in August when little or no wax bloom was present. The wax bloom material contributes to the protection of these diurnal beetles against the high temperatures and radiant heat loads in the Namib Desert.     

Leaf Cuticular Waxes of Bermudagrass Response to Environment-Driven Adaptations of Climate Effect Inferred from Latitude and Longitude Gradient in China

Bermudagrass (Cynodon dactylon) is a widely used warm season lawn grass. Cuticular wax covering the surface of plant leaves plays an important role in helping plants resist biotic and abiotic stresses. We analyzed the changes of cuticle wax in 25 bermudagrass populations from different longitude and latitude gradients, in order to verify how environmental conditions affect the structure and chemical composition of cuticle wax. Five wax components were identified, including alkanes, esters, alkenes, aldehydes and primary alcohols. The wax characteristics were divided into two principal components, explaining 58.2 % and 66.7 % of the total variability in latitude and longitude, even some populations had a certain correlation with each other. Pearson correlation analysis further showed that the total wax coverage, wax component content and antioxidant enzyme activity of bermudagrass populations on the latitudinal gradient had different responses to environmental factors. Finally, nineteen key genes involved in wax biosynthesis, redox and photosynthesis were identified and verified by RT-qPCR. The results showed that the responses of bermudagrass in different populations to climate change were quite different, which was of great significance for the evolution of bermudagrass populations.     

Waxy bloom in peas influences the performance and behavior of Aphidius ervi, a parasitoid of the pea aphid

Leaf surface waxy bloom can influence the predator–prey interactions that take place on peas. We tested whether the interaction between the pea aphid Acyrthosiphon pisum Harris (Homoptera: Aphidae) and a parasitoid, Aphidius ervi Haliday (Hymenoptera: Aphidiidae), is affected by reduced wax. We performed greenhouse experiments comparing aphid parasitism by individual A. ervi on two varieties of reduced wax peas to two normal wax sister varieties. We also observed the behavior of individual A. ervi in the greenhouse and measured field parasitism in small plots of reduced wax and normal wax peas. In the greenhouse, individual A. ervi parasitized more aphids on the reduced wax varieties than on their normal wax counterparts. Wasps spent more time actively foraging on reduced wax pea plants, which may contribute to the higher parasitism observed on those varieties. The greenhouse results suggested that the improved performance of individual A. ervi on reduced wax peas might contribute to a higher parasitism on reduced wax peas in the field. Field parasitism was significantly higher in reduced wax pea plots during 2000. Overall parasitism was higher in 2000 than in 2001 and 2002. In the latter years, parasitism was higher on reduced wax plants, but not significantly different from normal wax plants. Improved foraging by individual A. ervi resembles improved foraging by other carnivorous insects on reduced wax peas. The advantages of reduced wax for biological control of the pea aphid may hold when any of several different natural enemies is abundant.     

Effects of leaf epicuticular wax on the movement, foraging behavior, and attack efficacy of Diaeretiella rapae

Plant morphological characters can affect the foraging abilities of natural enemies. Heavy wax blooms have been shown to impede predators searching for herbivores on various species in the genus Brassica (Brassicaceae). This study determined whether epicuticular wax affected the foraging behavior of the braconid wasp Diaeratiella rapae (M’Intosh) (Hymenoptera: Braconidae) as it searched for its aphid host Brevicoryne brassicae L. (Homoptera: Aphididae) on the leaves of cauliflower varieties with heavy or light wax blooms. Wasps on the variety with a heavier wax bloom foraged more slowly, groomed more often and for longer periods of time, fell from the leaves more often, took longer to find colonies of aphids, and attacked them at a lower rate than wasps foraging on the variety with a lighter wax bloom. When epicuticular wax was removed from the leaves, the wasp's foraging efficiency and efficacy improved significantly on the cauliflower variety with a heavy wax bloom. The amount of epicuticular wax present on a leaf was shown to impede the ability of a parasitoid to forage, locate, and attack its host.     

Plant waxy bloom on peas affects infection of pea aphids by Pandora neoaphidis

This study examined the effects of the surface wax bloom of pea plants, Pisum sativum, on infection of pea aphids, Acyrthosiphon pisum, by the fungal pathogen Pandora neoaphidis. In prior field surveys, a higher proportion of P. neoaphidis-killed pea aphids (cadavers) had been observed on a pea line with reduced wax bloom, as compared with a sister line with normal surface wax bloom. Laboratory bioassays were conducted in order to examine the mechanisms. After plants of each line infested with aphids were exposed to similar densities of conidia, the rate of accumulation of cadavers on the reduced wax line was significantly greater than on the normal wax bloom line; at the end of the experiment (13d), the proportion of aphid cadavers on the reduced wax line was approximately four times that on the normal wax bloom line. When plants were exposed to conidia first and then infested with aphids, the rate of accumulation of cadavers was slightly but significantly greater on the reduced wax line, and infection at the end of the experiment (16d) did not differ between the lines. When aphids were exposed first and then released onto the plants, no differences in the proportion of aphid cadavers were observed between the pea lines. Greater infection of pea aphid on reduced wax peas appears to depend upon plants being exposed to inoculum while aphids are settled in typical feeding positions on the plant. Additional experiments demonstrated increased adhesion and germination by P. neoaphidis conidia to leaf surfaces of the reduced wax line as compared with normal wax line, and this could help explain the higher infection rate by P. neoaphidis on the reduced wax line. In bioassays using surface waxes extracted from the two lines, there was no effect of wax source on germination of P. neoaphidis conidia.     

Relationship Between Cuticular Waxes and Storage Quality Parameters of Korla Pear Under Different Storage Methods

Cuticular wax is an important factor that affects storage quality of fruits and vegetables. Previous studies have shown that cuticular wax of pears changes significantly during storage, whereas there are few studies on the effects of different storage methods on the wax changes and the relationship with storage quality. Cuticular wax of Korla pear stored using different methods, was measured to analyze its total wax content, chemical compositions and their relationship with storage quality. At the end of storage, the highest cuticular wax content was observed in controlled atmosphere (CA) storage and the lowest in room temperature storage. The substances of the primary components with higher contents were nonacosane, (E, E)-ɑ-farnesene, dodecan-1-ol, 1,1-dimethoxynonane, nonanal, palmitic acid, and oleic acid. Total wax content, olefins and fatty acids were most significantly with the storage quality, followed by alkanes and esters. Moreover, total wax content, wax composition and weight loss were closely related to postharvest senescence. Overall, an understanding of variations in the cuticular wax under different storage methods could provide theoretical basis for further study on the storage and preservation technology of pears.

     

Effects of Water Stress on Cuticular Transpiration Rate and Amount and Composition of Epicuticular Wax in Seedlings of Six Oat Varieties

Six varieties of oat (Avena sativa L. cv. Stormogul II, Risto, Sol II, Selma, Sang and Pendek, arranged according to decreasing drought resistance) were cultivated under controlled conditions and exposed to water stress on 4 consecutive days. Seven-day-old seedlings were stressed by cooling the roots for 3 h to 1.0°C. During this treatment the leaf water potential decreased from -7 to -12 bars. Cuticular transpiration rate, total amount of epicuticular wax and amounts of some wax components (primary alcohols, alkanes, fatty acids) were determined. Unstressed seedlings of the most drought resistant variety (Stormogul II) showed the highest cuticular transpiration rate. After stress treatment the cuticular transpiration rate was most strongly reduced in this variety and at the same time it showed the largest increase in amount of epicuticular wax of the tested varieties. In Pendek and Sang, showing the least increase in epicuticular wax, the cuticular transpiration rate was only 5% lower after stress treatment. In all varieties the primary alcohol content of the epicuticular wax was slightly higher in stressed seedlings than in controls. Further, in Stormogul and Risto the content of the predominant alkanes was much lower in stressed seedlings than in controls. On the contrary, in Pendek the stressed seedlings showed a higher alkane content. In Stormogul II, Risto and Sol II the total amount of fatty acids was higher in stressed seedlings than in controls while the opposite was true in Sang. The relation between the epicuticular wax (amount and composition) and the cuticular transpiration rate is discussed as well as the possibility of using the tested parameters in a screening test for drought resistance.     

Will the climate of plant origins influence the chemical profiles of cuticular waxes on leaves of Leymus chinensis in a common garden experiment?

Cuticular wax covering the leaf surface plays important roles in protecting plants from biotic and abiotic stresses. Understanding the way in which plant leaf cuticles reflect their growing environment could give an insight into plant resilience to future climate change. Here, we analyzed the variations of cuticular waxes among 59 populations of Leymus chinensis in a common garden experiment, aiming to verify how environmental conditions influence the chemical profiles of cuticular waxes. In total, eight cuticular wax classes were identified, including fatty acids, aldehydes, primary alcohols, alkanes, secondary alcohols, ketones, β‐diketones, and alkylresorcinols, with β‐diketones the predominant compounds in all populations (averaged 67.36% across all populations). Great intraspecific trait variations (ITV) were observed for total wax coverage, wax compositions, and the relative abundance of homologues within each wax class. Cluster analysis based on wax characteristics could separate 59 populations into different clades. However, the populations could not be separated according to their original longitudes, latitudes, annual temperature, or annual precipitation. Redundancy analysis showed that latitude, arid index, and the precipitation from June to August were the most important parameters contributing to the variations of the amount of total wax coverage and wax composition and the relative abundance of wax classes. Pearson's correlation analysis further indicated that the relative abundance of wax classes, homologues in each wax class, and even isomers of certain compound differed in their responses to environmental factors. These results suggested that wax deposition patterns of L. chinensis populations formed during adaptations to their long‐term growing environments could inherit in their progenies and exhibit such inheritance even these progenies were exported to new environments.     

Cuticular wax composition in Cocos nucifera L.: physicochemical analysis of wax components and mapping of their QTLs onto the coconut molecular linkage map

Cuticular waxes were extracted from the leaves of a coconut mapping population generated by the controlled cross of an East African Tall and a Rennell Island Tall genotype for the construction of molecular linkage maps. The wax composition was analyzed by capillary gas chromatography/mass spectrometry, and for eight of the wax compounds, their absolute and relative amounts were determined. As reported previously for a different coconut ecotype (Malayan Yellow Dwarf), lupeol methyl ether, isoskimmiwallin, and skimmiwallin were identified as the major components of coconut cuticular wax. The additional compounds were characterized as 3-β-methoxy lupane (lupane methyl ether), lupeol and the acetic acid esters of lupeol, skimmiwallinol, and isoskimmiwallinol, respectively. Minor, nonidentified compounds amounted to some 5% of total wax content and included triterpenoids, sterols, primary alcohols, and fatty acids. The variation detected for parents and progeny with respect to the wax components allowed quantitative trait locus (QTL) analyses for their biosynthetic pathways. A total of 46 QTLs could be mapped onto the coconut linkage map which was extended by amplified fragment length polymorphism and single sequence repeat markers into a high density map with more than 1,000 mapped DNA markers. Several colocated QTLs for different traits were detected reflecting the observed correlations among characters.     

Comparative analysis of the chemical composition and water permeability of the cuticular wax barrier in Welsh onion ( Allium fistulosum L.)

Cuticular wax is a hydrophobic barrier between the plant surface and the environment that effectively reduces the loss of water. The surface of Welsh onion leaves is covered with wax. To explain the relationship between wax composition and water loss, we conducted this experiment. The water permeability and wax composition of leaves were determined by chemical and GC-MS methods. We performed a comparative analysis of the differences between the two cultivars and analyzed the relationship between water permeability and waxy components. Overall, the permeability to water was higher in ‘Zhangqiu’ than in ‘Tenko’. The wax amount of ‘Tenko’ was 1.28-fold higher than that of ‘Zhangqiu’ and was primarily explained by the much larger amounts of ketones and alcohols in the former. Among the waxy components, C29 ketones were most abundant. There were substantial discrepancies in wax composition, total wax content, and water permeability between the two cultivars. The main reason for the discrepancy in water permeability may be the significantly lower aliphatic fraction in ‘Zhangqiu’ than in ‘Tenko’. This study makes a vital contribution to drought resistance research on allium plants.     

The Arabidopsis cer26 mutant,like the cer2 mutant,is specifically affected in the very long chain fatty acid elongation process

Plant aerial organs are covered by cuticular waxes, which form a hydrophobic crystal layer that mainly serves as a waterproof barrier. Cuticular wax is a complex mixture of very long chain lipids deriving from fatty acids, predominantly of chain lengths from 26 to 34 carbons, which result from acyl‐CoA elongase activity. The biochemical mechanism of elongation is well characterized; however, little is known about the specific proteins involved in the elongation of compounds with more than 26 carbons available as precursors of wax synthesis. In this context, we characterized the three Arabidopsis genes of the CER2‐like family: CER2, CER26 and CER26‐like . Expression pattern analysis showed that the three genes are differentially expressed in an organ‐ and tissue‐specific manner. Using individual T–DNA insertion mutants, together with a cer2 cer26 double mutant, we characterized the specific impact of the inactivation of the different genes on cuticular waxes. In particular, whereas the cer2 mutation impaired the production of wax components longer than 28 carbons, the cer26 mutant was found to be affected in the production of wax components longer than 30 carbons. The analysis of the acyl‐CoA pool in the respective transgenic lines confirmed that inactivation of both genes specifically affects the fatty acid elongation process beyond 26 carbons. Furthermore, ectopic expression of CER26 in transgenic plants demonstrates that CER26 facilitates the elongation of the very long chain fatty acids of 30 carbons or more, with high tissular and substrate specificity.     

Determination of diffusion coefficients of octadecanoic acid in isolated cuticular waxes and their relationship to cuticular water permeabilities

Transport properties of cuticular waxes from 40 different plant species were investigated by measuring desorption rates of ¹⁴C-labelled octadecanoic acid from isolated and subsequently reconstituted wax. Diffusion coefficients (D) of octadecanoic acid in reconstituted waxes, calculated from the slopes of the regression lines fitted to the linearized portions of desorption kinetics, ranged from 1.2 × 10^?19 m² s^?1 (Senecio kleinia leaf) to 2.9 × 10^?17 m² s^?1 (Malus cf. domestica fruit). Cuticular water permeabilities (cuticular transpiration) measured with intact cuticular membranes isolated from 24 different species varied from 1.7 × 10^?11 m s^?1 (Vanilla planifolia leaf) up to 2.1 × 10^?9 m s^?1 (Malus cf. domestica fruit), thus covering a range of more than 2 orders of magnitude. Cuticular water permeabilities were highly correlated with diffusion coefficients of octadecanoic acid in isolated cuticular wax of the same species. It is therefore possible to estimate cuticular barrier properties of stomatous leaf surfaces or of leaves where isolation of the cuticle is impossible by measuring D of octadecanoic acid in isolated waxes of these leaves.     

Leaf cuticular waxes and physiological parameters in alfalfa leaves as influenced by drought

Drought significantly constrains higher yield of alfalfa (Medicago sativa L.) in arid and semiarid areas all over the world. This study evaluated the responses of leaf cuticular wax constituents to drought treatment and their relations to gas-exchange indexes across six alfalfa cultivars widely grown in China. Water deficit was imposed by withholding water for 12 d during branching stage. Cuticular waxes on alfalfa leaves were dominated by primary alcohols (41.7?C54.2%), alkanes (13.2?C26.9%) and terpenes (17.5?C28.9%), with small amount of aldehydes (1.4?C3.4%) and unknown constituents (4.5?C18.4%). Compared to total wax contents, the wax constituents were more sensitive to drought treatment. Drought decreased the contents of primary alcohol and increased alkanes in all cultivars. Alkane homologs, C25, C27, and C29, were all negatively correlated with photosynthetic rate, transpiration rate, stomatal conductance, and leaf water potential. Under drought conditions, both stomatal and nonstomatal factors were involved in controlling water loss from alfalfa leaves. No direct relationship was observed between wax contents and drought resistance among alfalfa cultivars. An increase in alkane content might be more important in improving drought tolerance of alfalfa under water deficit, which might be used as an index for selecting and breeding drought resistant cultivars of alfalfa.     

Molecular mapping and candidate gene analysis of a new epicuticular wax locus in sorghum (<Emphasis Type="Italic">Sorghum bicolor</Emphasis> L. Moench)

Key message

A new epicuticular wax (bloom) locus has been identified and fine mapped to the 207.89 kb genomic region on chromosome 1. A putative candidate gene, Sobic.001G269200, annotated as GDSL-like lipase/acylhydrolase, is proposed as the most probable candidate gene involved in bloom synthesis/deposition.

Abstract

Deposition of epicuticular wax on plant aerial surface is one strategy that plants adapt to reduce non-transpiration water loss. Epicuticular wax (bloom)-less mutants in sorghum with their glossy phenotypes exhibit changes in the accumulation of epicuticular wax on leaf and culm surfaces. We report molecular mapping of a new sorghum locus, bloomless mutant (bm39), involved in epicuticular wax biosynthesis in sorghum. Inheritance studies involving a profusely bloom parent (BTx623) and a spontaneous bloomless mutant (RS647) indicated that the parents differed in a single gene for bloom synthesis. Bloomless was recessive to bloom deposition. Genetic mapping involving F₂ and F₇ mapping populations in diverse genetic backgrounds (BTx623 × RS647; 296A × RS647 and 27A × RS647) identified and validated the map location of bm39 to a region of 207.89 kb on chromosome 1. SSR markers, Sblm13 and Sblm16, flanked the bm39 locus to a map interval of 0.3 cM on either side. Nine candidate genes were identified, of which Sobic.001G269200 annotated for GDSL-like lipase/acylhydrolase is the most likely gene associated with epicuticular wax deposition. Gene expression analysis in parents, isogenic lines and sets of near isogenic lines also confirmed the reduced expression of the putative candidate gene. The study opens possibilities for a detailed molecular analysis of the gene, its role in epicuticular wax synthesis and deposition, and may help to understand its function in moisture stress tolerance and insect and pathogen resistance in sorghum.