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.     

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.     

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.     

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.

     

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.     

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.     

Non-glaucousness, a probable aphid-resistance character of wheat

Cuticular waxes from wheat cultivars that have a bluish-green waxy bloom under field conditions were compared with those of non-glaucous (NG) variants that appeared dark green and glossy in the field. Cuticular wax from NG variants was deficient in diketones and hydroxydiketones, and did not form the micro-spicules of epicuticular wax which were characteristic of the normal bloom. Typically, NG wheats were resistant to Sitobion avenae in the field, but in environments where wax secretion was diminished, resistance was reduced or absent. Possible causes of this resistance, which appears likely to be easy to exploit, are discussed.     

种皮蜡质与乌拉尔甘草种子抗菌及抗老化能力的关系

本文研究了乌拉尔甘草种子种皮蜡质的厚度与蜡质抑菌效果、蜡质与种子抗老化能力和种子吸水能力的关系。结果表明：用浓硫酸浸泡2-3min可完全除去蜡质,蜡质的厚度约为30gm;接种菌液后,脱蜡种子在PDA培养基上形成的青霉和曲霉菌斑的直径显著大于不脱蜡的种子形成的菌斑直径,但蜡质的体外抗菌能力不明显：随着老化时间的延长,脱蜡种子的含水量增加速度和发芽率下降速度明显快于未脱蜡种子,且在湿度较高的环境下,脱蜡种子的平衡水分比未脱蜡种子显著升高,说明蜡质在一定程度上可以保护种子不易受外界湿度的影响。     

Functional biodiversity of lipids in Antarctic zooplankton: Calanoides acutus, Calanus propinquus, Thysanoessa macrura and Euphausia crystallorophias

Zooplankton samples were collected in January 1993 off Dronning Maud Land along a transect from open waters to the marginal ice zone close to the Antarctic ice shelf. Thysanoessa macrura was caught in open waters while Calanoides acutus and Calanus propinquus were mainly sampled between ice floes in the marginal ice zone. The “ice-krill”Euphausia crystallorophias was found over the shelf directly associated with ice floes. T. macrura had a lipid content up to 36% of its dry weight with the dominant lipid class, wax ester, accounting for 45–50% of the total lipid. The predominance of 18:1 fatty alcohols is the striking characteristic of the wax esters. Small specimens of E. crystallorophias had lipid levels up to 26% of their dry weight with, unexpectedly, triacylglycerols being the dominant lipid (up to 41% of total lipid). The small levels of wax esters in these animals (3–6% of total lipid) had phytol as a major constituent. Large specimens of E. crystallorophias had up to 34% of their dry weight as lipid, with wax esters (47% of total lipid) dominated by 16:0 and 14:0 fatty alcohols as the major lipid. Calanus propinquus had lipid levels of up to 34% of their dry weight, with triacylglycerols (up to 63% of total lipid) being the dominant lipid. High levels of 22:1 (n-9) fatty acid were present in the triacylglycerols. Calanoides acutus had lipid levels up to 35% of the dry weight with wax esters accounting for up to 83% of total lipid. High levels of (n-3) polyunsaturated fatty acids were recorded with 20:5(n-3), 22:6(n-3) and 18:4(n-3) being the dominant moieties. On the basis of their lipid compositions we deduce that: (1) Calanoides acutus is the strictest herbivore among the four species studied, heavily utilizing the typical spring bloom; (2) T. macrura is essentially omnivorous, probably utilizing the less defined bloom situations found in oceanic waters; (3) E. crystallorophias is an omnivore well adapted to utilize both a bloom situation and to feed on ice algae and micro-zooplankton associated with the ice; (4) Calanus propinquus seems to be the most opportunistic feeder of the four species studied, probably grazing heavily on phytoplankton during a bloom and, during the rest of the year, feeding on whatever material is available, including particulates, flagellates and other ice-associated algae. We conclude that the different biochemical pathways generating large oil reserves of different compositions, enabling species to utilize different ecological niches, are major determinants of biodiversity in polar zooplankton. Accepted: 22 June 1998     

六类常见泌蜡昆虫的蜡腺和蜡泌物研究进展

泌蜡昆虫体表除了具有和其他昆虫一样的薄蜡层之外,还具有特殊的蜡腺,在生长发育过程中能够分泌蜡质覆盖于体表,从而有利于昆虫防止体内水分过度蒸发、抵御外界多种不利条件、阻碍病原物入侵和躲避天敌寄生或捕食,对昆虫的生存和繁衍具有重要的保护作用。本文概述了蚧虫、蚜虫、蜡蝉、木虱、粉虱和粉蛉等6类常见泌蜡昆虫的蜡腺和蜡泌物的超微形态结构、蜡泌物的主要化学组成及泌蜡的分子机制等方面的研究进展,以期为今后其在昆虫分类学和系统发育学的研究以及在害虫防治中的应用提供参考。     

How aphids lose their marbles

Insects provide examples of many cunning stratagems to cope with the challenges of living in a world dominated by surface forces. Despite being the current masters of the land environment, they are at constant risk of being entrapped in liquids, which they prevent by having waxy and hairy surfaces. The problem is particularly acute in an enclosed space, such as a plant gall. Using secreted wax to efficiently parcel and transport their own excrement, aphids were able to solve this problem 200 Myr ago. Here, we report on the physical and physiological significance of this ingenious solution. The secreted powdery wax has three distinct roles: (i) it is hydrophobic, (ii) it creates a microscopically rough inner gall surface made of weakly compacted wax needles making the gall ultra-hydrophobic, and (iii) it coats the honeydew droplets converting them into liquid marbles, that can be rapidly and efficiently moved.     

Influence of atmospheric CO2 enrichment, soil N, and water stress on needle surface wax formation in Pinus palustris (Pinaceae)

Interactive effects of increasing atmospheric CO2 with resource limitations on production of surface wax in plants have not been studied. Pinus palustris seedlings were grown for 1 yr at two levels of soil N (40 or 400 kg N_ha-1_yr-1) and water stress (-0.5 or -1.5 MPa xylem pressure potential) in open-top field chambers under two levels of CO2 (365 or 720 mumol/mol). Needle surface wax content was determined at 8 mo (fall) and 12 mo (spring) and epicuticular wax morphology was examined using scanning electron microscopy (SEM) at 12 mo. Wax content expressed on both a leaf area and dry mass basis was increased due to main effects of low N and water stress. No main effects of CO2 were observed; however, a CO2 x N interaction at 12 mo indicated that under low soil N the elevated CO2 treatment had less wax (surface area or dry mass basis) compared to its ambient counterpart. Morphologically, low N needle surfaces appeared rougher compared to those of high N needles due to more extensive wax ridges. Although the main effect of water treatment on wax density was not reflected by changes in wax morphology, the CO2 x N interaction was paralleled by alterations in wax appearance. Decreases in density and less prominent epicuticular wax ridges resulting from growth under elevated CO2 and limiting N suggest that dynamics of plant/atmosphere and plant/pathogen interactions may be altered.     

垂叶榕叶片表面蜡质化学组分及蓟马锉吸胁迫对其含量的影响

为明确垂叶榕(Ficus benjamina)叶片表面蜡质的化学组分和蓟马(Thrips spp.)锉吸胁迫对蜡质含量的影响,采用气相色谱-质谱联用仪(GC-MS)对不同危害等级蓟马锉吸叶片的表面蜡质化学组分及含量进行了检测和比较分析。结果表明,垂叶榕叶片的表面蜡质中含有32种化合物,包含有9种酸、8种醇、5种酯、5种酮和5种烃。不同危害等级叶片的表面蜡质总量、已分离化合物的总离子峰面积、各类化合物的相对含量、各类化合物及其主要组分的离子峰面积均不同。垂叶榕叶片的表面蜡质的化学组分和含量可直接或间接反映叶片受蓟马锉吸危害的程度。这说明叶片表面蜡质的化学组分及含量在垂叶榕与蓟马的互作和协同进化中扮演着重要角色。     

南洋臀纹粉蚧雌成虫主要器官及其蜡泌物的扫描电镜观察

为明确南洋臀纹粉蚧Planococcus lilacinus(Cockerell)雌成虫及其蜡泌物的结构特征,利用扫描电镜观察该虫体表主要器官、蜡质及泌蜡腺体的超微结构.结果表明:南洋臀纹粉蚧雌成虫外覆白色粉状厚蜡被,体缘具18对蜡棒,触角8节、口器和足发达且分布有不同长度的毛形和刺形感受器,眼为单眼,腹脐和背孔唇形、...     

Attachment force of the beetle Cryptolaemus montrouzieri (Coleoptera, Coccinellidae) on leaflet surfaces of mutants of the pea Pisum sativum (Fabaceae) with regular and reduced wax coverage

This study represents an investigation of surface-related plant–insect interactions. Surface micro-morphology of leaflets in pea (Pisum sativum) with wild-type crystalline surface waxes (waxy) and with reduced crystalline surface waxes (glossy) caused by a mutation (wel) were studied using various microscopy techniques. The free surface energy of these plant surfaces was estimated using contact angles of droplets of three different liquids. The morphological study of the attachment system in the ladybird beetle Cryptolaemus montrouzieri was combined with measurements of attachment (traction) forces, generated by beetles on these plant substrates. Differences were found in wax crystal shape, dimensions, and density between the adaxial and abaxial surfaces of waxy and glossy plants. The crystalline wax was not completely eliminated in the glossy plant: it was only slightly reduced on the adaxial side and underwent greater changes on the abaxial side. The free surface energy for both surfaces of both pea types was rather low with strongly predominating dispersion component. Insects generated low traction forces on all intact plant surfaces studied, except the abaxial surface of the glossy plant, on which the force was greater. After being treated with chloroform, all the surfaces allowed much higher traction forces. It is demonstrated that the difference in the crystal length and density of the epicuticular wax coverage within the observed range did not influence wettability of surfaces, but affected insect attachment. The reduction in insect attachment force on plant surfaces, covered with the crystalline wax, is explained by the decrease of the real contact area between setal tips of beetles and the substrate. Handling editor: Lars Chittka.     

Endogenous polyamines in apple and their relationship to fruit set and fruit growth

The distribution of free and bound polyamines was investigated from blooming until harvest on flower, fruitlets and fruits of Malus domestica Borkh cv. Golden Delicious, Relationships between polyamines and fruit set and growth were also investigated. The level of free polyamines was high only during the first weeks after full bloom and then decreased gradually. The amount of bound trichloroacetic acid-insoluble polyamines was much higher than free polyamines. Bound spermine in particular showed a high value for almost 40 days after full bloom, while spermidine and putrescine were no longer detectable even a few days after full bloom.
In relation to fruit set, it was possible to observe that abscission peaks took place when free polyamine levels were low or decreasing. Insofar as fruit growth is concerned, the most substantial variations in polyamine levels occurred very early in the season when fruit dry weight and protein amount were also changing rapidly and fruit diameter was almost impossible to measure.     

角倍蚜各虫态蜡腺的分布与结构

[目的]角倍蚜Schlechtendalia chinensis是五倍子的主要生产种,具有复杂的生活史,需要经历干母、干雌、秋迁蚜、越冬若蚜、春迁蚜和性蚜6种虫态,在盐肤木Rhus chinensis和藓类植物上转主寄生,经历有性和无性生殖、瘿内和瘿外世代的交替才能完成生活史.通过对角倍蚜蜡腺的研究,有助于深入了解角倍...     

Variations of cuticular wax in mulberry trees and their effects on gas exchange and post-harvest water loss

Though mulberry (Morus alba) tree shows great adaptations to various climate conditions, their leaf water status and photosynthesis are sensitive to climate changes. In the current study, seven widely planted mulberry cultivars in Chongqing, Southwest China, were selected to analyze leaf cuticular wax characteristics, gas exchange index, post-harvest leaf water status and their relationships, aiming to provide new theory in screening high resistant mulberry cultivars. Mulberry trees formed rounded cap-type idioblasts on the adaxial leaf surface. Film-like waxes and granule-type wax crystals covered leaf surfaces, varying in crystal density among cultivars. The stomatal aperture on the abaxial surface of cultivars with high wax amount was smaller than that of cultivars with low wax amount. The amount of total wax was negatively correlated with the net photosynthetic rate (P _N), transpiration rate (E) and stomatal conductance (g _s) and positively correlated with the moisture retention capacity. It suggested that both cuticular wax and stomatal factor might be involved in regulating water loss in mulberry leaves under field conditions. The variability in moisture retention capacity and cuticular wax characteristics might be important in evaluating and screening mulberry cultivars for increasing silk quality and silkworm productivity.     

Cuticular wax variation in the tomato (Solanum lycopersicum L.), related wild species and their interspecific hybrids

The tomato (Solanum lycopersicum L.) is one of the world's most important vegetable crop species. Among the many tomato accessions available, only a few are tolerant to abiotic stresses, which are responsible for the majority of the crop losses worldwide. Wild tomato species are then secondary gene pool in the breeding of more resistant tomato cultivars. In the current study, the composition of leaf cuticular waxes from fourteen tomato accessions, including S. lycopersicum, Solanum pennellii, Solanum pimpinellifolium, and their interspecific hybrids was studied in order to select the most adequate chemotaxonomic markers. Total cuticular wax load of S. pennellii plants was much higher than in the other plant species. Hydrocarbons were usually the most abundant wax components, followed by minor quantities of triterpenes and other compounds. Interspecific hybrids showed intermediate wax characteristics. The amount and composition of surface waxes were not correlated with the abiotic stress tolerance in S. lycopersicum. The composition of the hydrocarbon fraction was the least variable both within a single accession and between all the plants studied. Based on the results, cuticular hydrocarbons are proposed as potential chemotaxonomic markers in the classification of tomato and related species.