Resistance in sorghum to shoot fly Atherigona soccata: Evidence for the source of leaf surface wetness

Leaf surface wetness (LSW) of the central whorl leaf of sorghum seedlings has been associated with susceptibility to shoot fly. Previous physical and physiological evidence suggested that LSW originates from the plant. This was confirmed by radioactive labelling methods using tritium and carbon-14. Tritiated water applied to the soil of potted seedlings was translocated to the surface of the whorl leaf. There were significant differences in the amount of tritiated water collected from susceptible (CSH 5) and resistant (IS 18551) genotypes. Studies with carbon-14 labelling of sorghum seedlings indicated the presence of (small amounts of) solutes in the surface water that may affect larval movement and survival.     

The dynamics of leaf surface wetness of sorghum seedlings in relation to resistance to the shoot fly, Atherigona soccata

In quantitative measurements of leaf surface wetness (LSW) of the central whorl leaf of sorghum seedlings in August (rainy season) and November (post-rainy season), the highest amount (6.29 mg of water) was recorded in August in the shoot fly Atherigona soccata (Diptera: Muscidae), susceptible sorghum genotype CSH 1, while the lowest (0.07 mg) was recorded in November in the resistant genotype IS 18551. Studies on diurnal fluctuation revealed that LSW was lowest at sunset, was highest between 02.00 and 04.00 h (closely corresponding with hatching of shoot fly eggs) and dropped before sunrise. This fluctuation was associated with the evaporation of water from the plant during the night. More LSW accumulation occurred during the main crop season (June-October) than in the post-rainy season (November-April). Annual fluctuation of LSW followed trends similar to the population dynamics of shoot fly and crop infestation and were correlated with rainfall, temperature and relative humidity. Measurements of leaf temperature and the vapour pressure gradient between the leaf and the air indicated that leaf surface water originates from the plant. This was further supported by the different amounts of LSW on susceptible and resistant cultivars with similar microclimatic conditions.     

Leaf surface wetness in sorghum and resistance to shoot fly, Atherigona soccata: role of soil and plant water potentials

In experiments with potted plants, the relationships between soil matric potential, plant water potential and production of water droplets (leaf surface wetness) on the folded central whorl leaf of seedlings of sorghum genotypes that are either resistant or susceptible to shoot fly (Atherigona soccata) damage were investigated. Differences in soil matric potentials in the pots affected the plant water status, which in turn had profound effects on the production of water droplets on the central whorl leaf of the sorghum genotype susceptible to shoot fly. There was no consistent variation in the relationship between plant water potential and soil matric potential of resistant and susceptible sorghum genotypes. However, there was very little or practically no water droplets on the central whorl leaf of the resistant genotypes, indicating that the production of water droplets is not solely the result of internal water status of the plant. It is suggested that leaf surface wetness is genetically controlled and that an understanding of the mechanism by which water is transferred to the leaf surface will enhance breeding for resistance to shoot fly.     

Resistance in sorghum to the shoot fly, Atherigona soccata: epicuticular wax and wetness of the central whorl leaf of young seedlings

Sorghum genotypes known to be resistant or susceptible to shoot fly, Atherigona soccata Rondani were examined by scanning electron microscopy for differences in epicuticular wax structure and wetness of the central leaf whorl. Two major types of wax structures were observed: shoot fly resistant and moderately resistant genotypes were characterised by a smooth amorphous wax layer and sparse wax crystals while susceptible genotypes possessed a dense meshwork of crystalline epicuticular wax. The density of wax crystals decreased from the third leaf to the seventh leaf stage and was related to both seedling age and leaf position. Water droplets on susceptible genotypes with dense wax crystals showed spreading at the edges indicating a tendency to wet easily. In resistant genotypes with less dense wax crystals the droplets remained intact and did not spread.     

Reduction of shoot fly damage in irrigated post-rainy season sorghum by manipulating irrigation

Soil moisture was manipulated in an attempt to control shoot fly (Atherigona soccata Rondani) incidence in irrigated post-rainy season sorghum grown under a rainout shelter (ROS) and in field conditions. After uniform irrigation at sowing, the plants were subjected to water stress at young seedling stage (7–28 days after emergence, DAE) for different lengths of time. Soil water had profound effects on the production of water droplets on the surface of the central whorl leaf of seedlings (leaf surface wetness, LSW) of sorghum genotypes. LSW, which facilitates movement of the larvae, was more drastically affected in susceptible (CSH 5) than in moderately resistant (IS 1054) sorghum genotypes. Shoot fly oviposition (infestation) and deadhearts (crop damage) were much higher in treatments with full irrigation (control) than in treatments to which less water was applied during the first 3 wk after seedling emergence. This resulted in higher plant biomass and overall grain yield in the latter treatments than in the control. Using insecticides to control shoot fly infestation, it was shown that a simple cultural practice of inducing plant stress by reduced soil moisture content during early plant growth gave the same or better control of shoot fly damage and the same or higher grain yield than insecticide-protected plots with full irrigation. Thus the costs associated with irrigation requirement and insecticide can be greatly reduced in the former management option compared with the latter. It is suggested that manipulation of soil water content during the vulnerable early stages of crop growth can reduce shoot fly damage in irrigated post-rainy season sorghum.     

空气湿度与土壤水分胁迫对紫花苜蓿叶表皮蜡质特性的影响

选用2个抗旱性不同的紫花苜蓿品种,敖汉(强抗旱)和三得利(弱抗旱),设置空气湿度(45%-55%和75%-85%)和土壤水分胁迫(75%和35%田间持水量)处理,分析紫花苜蓿叶表皮蜡质含量、组分及晶体结构、气体交换参数、水势及脯氨酸含量的变化规律。结果表明,单独土壤水分胁迫时,紫花苜蓿叶表皮蜡质晶体结构及蜡质总量无显著变化;敖汉蜡质组分中烷类、酯类含量增加,醇类含量下降;三得利醇类含量下降,烷类、酯类含量变化不显著。低空气湿度胁迫时,两品种蜡质总量无显著变化,烷类和酯类含量显著增加,醇类含量显著下降,叶表皮片状蜡质晶体结构熔融呈弥漫性,扩大了对叶表面积的覆盖,其蒸腾速率显著低于正常湿度。复合胁迫处理时,叶表皮片状蜡质晶体结构继续呈弥漫性,烷类、酯类、未知蜡质组分含量均高于单独胁迫处理,醇类含量最低,而蜡质总量除三得利显著高于对照外,其余均无显著差异。紫花苜蓿叶表皮蜡质各组分含量(除醇类)及蜡质总量与光合速率呈显著负相关,与蒸腾速率无显著相关关系。蜡质总量与叶水势呈显著正相关。总体上,敖汉蜡质总量显著高于三得利,蜡质组分中烷类物质的增加有助于提高植株的抗旱性。在复合胁迫下,强抗旱品种主要通过气孔因素控制水分散失,而弱抗旱品种通过气孔和非气孔因素共同控制植物水分散失。     

Leaf surface chemistry of sorghum seedlings influencing expression of resistance to sorghum shoot fly, Atherigona soccata

Sorghum shoot fly, Atherigona soccata is one of the serious constraints to sorghum production, and host plant resistance is an important component for controlling this pest. We studied the expression of resistance to A. soccata in a diverse array of sorghum genotypes in relation to composition of leaf surface chemicals during the seedling stage. The sorghum genotypes IS 1054, IS 1057, IS 2146, IS 4664, IS 2312, IS 2205, SFCR 125, SFCR 151, ICSV 700, and IS 18551 exhibited antixenosis for oviposition, and suffered less deadhearts due to sorghum shoot fly, A. soccata. Compounds undecane 5- methyl, decane 4- methyl, hexane 2, 4- methyl, pentadecane 8- hexyl, and dodecane 2, 6, 11- trimethyl, present on the leaf surface of sorghum seedlings, were associated with susceptibility to shoot fly; while 4, 4- dimethyl cyclooctene was associated with resistance to shoot fly. The compounds associated with resistance/susceptibility to shoot fly, can be used as marker traits to select for resistance as well as for diversifying and increasing the levels of resistance to this pest. The role of biochemical compounds for developing sorghum varieties with resistance to shoot fly, A. soccata has been discussed.     

西安市常见绿化植物叶片润湿性能及其影响因素

利用接触角测定仪测定了西安市21种常见绿化植物叶片表面的接触角,探讨了叶片表面特性如蜡质、绒毛、气孔对接触角的影响。结果表明,植物叶片正背面、物种间的接触角差异均显著,叶片正面和背面接触角大小在40°～140°。接触角大小与变异系数呈负相关,可能由于接触角小的润湿叶片在不同的生境和位置下,受到环境条件的影响较大而出现大的变异;接触角较大的非润湿性叶片,环境物质持留时间较短,对叶片形态和组成影响较小,因而出现小的变异。植物叶片表面的接触角随蜡质含量的升高而增大。表皮蜡质去除后大部分叶片接触角明显降低,尤其是疏水性较强的银杏(Ginkgo biloba)、月季(Ro-sa chinensis)和紫叶小檗(Berberis thunbergii)。女贞(Ligustrum lucidum)正背面、加杨(Popu-lus canadensis)背面等亲水型的叶片蜡质去除后接触角反而增大。叶片绒毛的多少及其形态、分布方式对接触角具有重要的影响,不同的作用方式表现出润湿和不润湿的特征,人为将其去除可以增加叶片的润湿性。背面气孔密度与气孔长度、保卫细胞长度呈负相关;接触角则与气孔密度呈负相关,与气孔长度呈正相关。     

Physico‐chemical mechanisms of resistance to shoot fly,Atherigona soccata in sorghum,Sorghum bicolor

Sorghum shoot fly, Atherigona soccata is an important pest of sorghum, and host plant resistance is one of the important components for minimizing the losses due to this pest. Therefore, we evaluated a diverse array of sorghum genotypes to identify physico‐chemical characteristics conferring resistance to A. soccata. Susceptibility to shoot fly was associated with high amounts of soluble sugars, fats, leaf surface wetness and seedling vigour; while leaf glossiness, plumule and leaf sheath pigmentation, trichome density and high tannin, Mg and Zn showed resistance to shoot fly. Stepwise regression indicated that Mg, Zn, soluble sugars, tannins, fats, leaf glossiness, leaf sheath and plumule pigmentation and trichome density explained 99.8% of the variation in shoot fly damage. Path coefficient analysis suggested that leaf glossiness, trichome density, Mg and fat content and plant plumule pigmentation can be used as markers traits to select for shoot fly resistance in sorghum.     

Functional interaction between leaf trichomes, leaf wettability and the optical properties of water droplets

Abstract. Because CO₂ diffuses 10000 times more slowly through water than air, there may be strong selective pressure for increased water repellency in terrestrial plant leaves. In the present study, leaf trichomes appeared to have a strong influence on leaf water repellency (i.e. degree of water droplet formation on the leaf surface) as well as the retention of droplets on the leaf. Based upon evaluation of 38 plant species from 21 families, we found that leaves with trichomes were more water repellent, especially where trichome density was greater than 25mm². However, droplet repellency and retention were both high in some species where trichomes entrapped droplets. Finally, the lensing effects of water droplets on leaf surfaces increased incident sunlight by over 20-fold directly beneath individual droplets. These results may have important implications for such processes as stomatal function, whole leaf photosynthesis, and transpiration for a large variety of plant species.     

Effect of alterations in cuticular wax biosynthesis on the physicochemical properties and topography of maize leaf surfaces

The leaf surface properties of 11 cuticular wax mutants of maize were characterized, and this information was used to identify the quantitative relations among distinct leaf surface traits. Compared with the wild‐type maize, these mutants were reduced 3–24% in their leaf surface hydrophobicity, 20–88% in the mass of cuticular waxes on their leaves, and 52–94% in the percentage of planar leaf surface area covered with epicuticular crystalline waxes. They also differed in the presence and abundance of the epicuticular crystalline waxes in each of seven structural classes. With the exception of one mutant, the mass of cuticular waxes produced by these mutants was positively correlated with the number of epicuticular crystalline waxes per unit area on their leaves. Furthermore, an increase of 0·4 mg of cuticular wax per gram of leaf (dry weight) was associated with a 1% increase in leaf surface area covered by epicuticular crystalline waxes, and this 1% increase was associated with a 2° increase in the contact angle of a water droplet on the leaf surface. Linear differences in the leaf surface hydrophobicity were associated with exponential differences in the mass of the cuticular waxes produced. Quantitative knowledge of these leaf surface properties is highly relevant to the interactions of leaves with environmental factors such as microbes, insects, agricultural chemicals, and pollutants.     

The citrus leaf cuticle in relation to measurement of leaf water potential using thermocouple psychrometers*

Abstract. Cuticular resistance to water vapour diffusion is an important aspect of thermocouple psychrometry and may introduce significant error in the measurement of leaf water potential (Ψ). The effect of the citrus (Citrus mitis Blanco) leaf cuticle on water vapour movement was studied using the times required for vapour pressure equilibration during thermocouple psychrometric measurement of Ψ. Cuticular abrasion with various carborundum powders was used to reduce the diffusive resistance of both the adaxial and abaxial leaf surfaces, and the extent of the disruption to the leaf was investigated with light and electron microscopy. Cuticular abrasion resulted in reduced equilibration times due to decreased cuticular resistance and greater water vapour movement between the leaf and the psychrometer chamber. Equilibration times were reduced from over 5 h in the unabraded control leaves to 1 h with cuticle abrasion. This was associated with the decrease in diffusive resistance with cuticular abrasion from over 55 s cm^?1 to less than 8 s cm^?1 for both the adaxial and abaxial leaf surfaces. Scanning electron micrographs of the abraded leaf tissue revealed considerable disruption of the stomatal ledge and of the guard cells, surface smoothing and displacement of waxes into the stomatal aperture, and damage to veins. Observations with the transmission electron microscope revealed frequent disruption of epidermal cell walls, and damage to both the cytoplasmic and vacuolar membranes.     

Shift in birch leaf metabolome and carbon allocation during long-term open-field ozone exposure

Current and future ozone concentrations have the potential to reduce plant growth and increase carbon demand for defence and repair processes, which may result in reduced carbon sink strength of forest trees in long‐term. Still, there is limited understanding regarding the alterations in plant metabolism and variation in ozone tolerance among tree species and genotypes. Therefore, this paper aims to study changes in birch leaf metabolome due to long‐term realistic ozone stress and to relate these shifts in the metabolism with growth responses. Two European white birch (Betula pendula Roth) genotypes showing different ozone sensitivity were growing under 1.4–1.7 × ambient ozone in open‐field conditions in Central Finland. After seven growing seasons, the trees were analysed for changes in leaf metabolite profiling, based on 339 low molecular weight compounds (including phenolics, polar and lipophilic compounds, and pigments) and related whole‐tree growth responses. Genotype caused most of the variance of metabolite concentrations, while ozone concentration was the second principal component explaining the metabolome profiling. The main ozone caused changes included increases in quercetin‐phenolic compounds and compounds related to leaf cuticular wax layer, whereas several compounds related to carbohydrate metabolism and function of chloroplast membranes and pigments (such as chlorophyll‐related phytol derivatives) were decreasing. Some candidate compounds such as surface wax‐related squalene, 1‐dotriacontanol, and dotriacontane, providing growth‐related tolerance against ozone were demonstrated. This study indicated that current growth‐based ozone risk assessment methods are inadequate, because they ignore ecophysiological impacts due to alterations in leaf chemistry.     

Changes in leaf trichomes and epicuticular flavonoids during leaf development in three birch taxa

BACKGROUND AND AIMS: Changes in number of trichomes and in composition and concentrations of their exudates throughout leaf development may have important consequences for plant adaptation to abiotic and biotic factors. In the present study, seasonal changes in leaf trichomes and epicuticular flavonoid aglycones in three Finnish birch taxa (Betula pendula, B. pubescens ssp. pubescens, and B. pubescens ssp. czerepanovii) were followed. METHODS: Trichome number and ultrastructure were studied by means of light, scanning and transmission electron microscopy, while flavonoid aglycones in ethanolic leaf surface extracts were analysed by high-pressure liquid chromatography. KEY RESULTS: Density of both glandular and non-glandular trichomes decreased drastically with leaf expansion while the total number of trichomes per leaf remained constant, indicating that the final number of trichomes is established early in leaf development. Cells of glandular trichomes differentiate before those of the epidermis and produce secreted material only during the relatively short period (around 1-2 weeks) of leaf unfolding and expansion. In fully expanded leaves, glandular trichomes appeared to be at the post-secretory phase and function mainly as storage organs; they contained lipid droplets and osmiophilic material (probably phenolics). Concentrations (mg g(-1) d. wt) of surface flavonoids decreased with leaf age in all taxa. However, the changes in total amount ( microg per leaf) of flavonoids during leaf development were taxon-specific: no changes in B. pubescens ssp. czerepanovii, increase in B. pendula and in B. pubescens ssp. pubescens followed by the decline in the latter taxon. Concentrations of most of the individual leaf surface flavonoids correlated positively with the density of glandular trichomes within species, suggesting the participation of glandular trichomes in production of surface flavonoids. CONCLUSIONS: Rapid decline in the density of leaf trichomes and in the concentrations of flavonoid aglycones with leaf age suggests that the functional role of trichomes is likely to be most important at the early stages of birch leaf development.     

A new technique for measurement of water permeability of stomatous cuticular membranes isolated from Hedera helix leaves

Transpiration of cuticular membranes isolated from the lower stomatous surface of Hedera helix (ivy) leaves was measured using a novel approach which allowed a distinction to be made between gas phase diffusion (through stomatal pores) and solid phase diffusion (transport through the polymer matrix membrane and cuticular waxes) of water molecules. This approach is based on the principle that the diffusivity of water vapour in the gas phase can be manipulated by using different gases (helium, nitrogen, or carbon dioxide) while diffusivity of water in the solid phase is not affected. This approach allowed the flow of water across stomatal pores ('stomatal transpiration') to be calculated separately from the flow across the cuticle (cuticular transpiration) on the stomatous leaf surface. As expected, water flux across the cuticle isolated from the astomatous leaf surface was not affected by the gas composition since there are no gas-filled pores. Resistance to flux of water through the solid cuticle on the stomatous leaf surface was about 11 times lower than cuticular resistance on the astomatous leaf surface, indicating pronounced differences in barrier properties between cuticles isolated from both leaf surfaces. In order to check whether this difference in resistance was due to different barrier properties of cuticular waxes on both leaf sides, mobility of 14C-labelled 2,4-dichlorophenoxy-butyric acid 14C-2,4-DB) in reconstituted cuticular wax isolated from both leaf surfaces was measured separately. However, mobility of 14C-2,4-DB in reconstituted wax isolated from the lower leaf surface was 2.6 times lower compared with the upper leaf side. The significantly higher permeability of the ivy cuticle on the lower stomatous leaf surface compared with the astomatous surface might result from lateral heterogeneity in permeability of the cuticle covering normal epidermal cells compared with the cuticle covering the stomatal cell surface.     

The role of leaf surface wetness in larval behaviour of the sorghum shoot fly, Atherigona soccata

The susceptibility of sorghum to the shoot fly Atherigona soccata Rondani, (Diptera: Muscidae) is affected by seedling age and is highest when seedlings are 8–12 days old. This corresponds with high moisture accumulation on the central leaf which is the path of newly hatched larva as it moves downwards from the oviposition site, towards the growing apex. Studies showed that leaf surface wetness (LSW) of the central shoot leaf was higher in 10-day old seedlings than in seedlings of other ages. Similarly, LSW was much higher in the susceptible sorghum genotype CSH 1 than in the resistant genotype IS 2146. Larvae moved faster towards the growing point and produced deadhearts much earlier in CSH 1 than in IS 2146. They also moved faster in 10-day old seedlings than in seedlings of other ages. It was also shown that the leaf surface wetness of the central shoot leaf is a more reliable parameter of resistance than the glossy leaf trait or trichome density.

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L'influence de la humidité de la surface foliaire sur le comportement de la mouche des pousses du sorgho

Résumé La sensibilité du sorgho à la mouche des pousses du sorgho, Atherigona soccata Rondani, est liée à l'âge de la plantule. Elle est plus forte lorsque la plantule est âgée de 8 à 12 jours et la sensibilité est maximale à 10 jours. A ce stade de croissance on observe une forte accumulation d'humidité sur la feuille centrale de la tige. Les jeunes larves traversent cette zone humide lorsqu'elles descendent vers la zone de croissance à partir des pontes déposées sur la face ventrale des feuilles déroulées.Des études ont été menées à l'ICRISAT (Inde) sur la relation entre l'humidité de la feuille centrale de la tige des plantules du sorgho et les dégâts provoqués par la mouche des pousses. L'humidité de la surface des feuilles (HSF) a été estimée grâce à une échelle visuelle graduée 1 à 5 où, 1 = pas d'humidité apparente et 5 = surface de la feuille recouverte de gouttes d'eau. La HSF est plus élevée sur des pousses de sorgho âgées de 10 j que sur les pousses appartenant à d'autres classes d'âge. Les valeurs observées sont également plus fortes pour les variétés non résistantes à ce ravageur (CSH 1,4.8) que pour les variétés résistantes (IS 2146, (2)). La vitesse du déplacement larvaire entre le cornet et la zone de la croissance varie en fonction de l'âge de la plante et des cultivars. Les larves migrent plus rapidement vers la zone de croissance et provoquent la mort du coeur du sorgho plus tôt dans la variété CSH 1 que dans IS 2146. Les larves se déplacent plus rapidement dans les pousses âgées de 10 j que dans les pousses appartenant à d'autres classes d'âge.Des études ont également démontré que la HSF n'est pas directement liée au caractère feuille lisse où à la densité des trichomes. La HSF est faible pour les génotypes résistants présentent où non le caractère feuille lisse. Par contre la HSF est élevée pour les génotypes non résistants présentant le caractère feuille lisse ou non. Aucune relation directe entre la densité des trichomes et les dégâts provoqués par la mouche des pousses n'a pu être mise en évidence. L'analyse des correlations établie pour les caractères de surface des feuilles avec la mort du cur des sorghos indique que les correlations sont faibles et non-significatives pour le caractère feuille lisse (0.49) et la densité des trichomes (0.39 et 0.2). Par contre les correlations sont fortes et significatives pour la HSF (0.82).On conclue que la HSF de la feuille centrale de la tige est un facteur important dans le déterminisme de la résistance du sorgho vis à vis de la mouche des pousses. Les relations entre les processus physiologiques de la plante et les facteurs impliquées dans l'accumulation d'eau sur la surface des feuilles font actuellement l'objet d'études détaillées.

     

Interaction of bean leaf pubescence with rust urediniospore deposition and subsequent infection density

The role of dense straight trichomes on bean leaves associated with race-nonspecific resistance to rust was studied using a settling tower method for uniform inoculation. Abaxial leaf trichomes, which occur only on adult leaves, trapped significant numbers of urediniospores, especially those deposited in clusters, and resulted in reduced effective inoculum reaching the leaf surface. In the field where infection conditions are often less than optimum, leaf pubescence would partially explain the observed resistance. However, the observed increase in total number of spores as well as similar infection densities on fully expanded leaves of glabrous and pubescent genotypes cannot be explained by the trichomes. Other physical or chemical features of the leaf surface such as waxes or cations may play a role. The ecological fit of leaf pubescence is discussed.     

Exploiting physical defence traits for crop protection: leaf trichomes of Rubus idaeus have deterrent effects on spider mites but not aphids

Plants possess anti‐herbivore defences that could be exploited for crop protection. The potential for deploying physical defence traits for more sustainable pest management (i.e. reduced pesticide application) has not been fully realised. Using a perennial crop (red raspberry, Rubus idaeus), we take the novel approach of quantifying within‐ and between‐genotype variation in a resistance trait, leaf trichome density, to determine precisely the effect of trichomes on host plant preference and suitability for two shoot‐feeding arthropods, the European large raspberry aphid (Amphorophora idaei) and two‐spotted spider mite (Tetranychus urticae). Additionally, we tested whether this trait influenced searching behaviour of a generalist herbivore predator (lacewing larvae, Chrysoperla carnea). Although there was no consistent genotypic variation in R. idaeus suitability for T. urticae, our hypothesis that T. urticae would avoid high leaf trichome density was supported on certain genotypes. The deterrent effect was mainly on egg deposition rather than leaf selection by adults, with up to sixfold differences in leaf preference depending on the genotypes offered. By contrast, there was significant genotypic variation in R. idaeus suitability for A. idaei (10‐fold variation in aphid abundance), but, contrary to our prediction, aphid preference and infestation levels were unrelated to leaf trichome density. Instead, A. idaei performed best on vigorous genotypes, indicating that plant tolerance traits contributed to R. idaeus suitability for aphids. Leaf trichomes had little effect on the behaviour of the beneficial control agent C. carnea larvae. We conclude that physical anti‐herbivore defences, specifically leaf trichomes, could be deployed to deter particular arthropod pests. However, the mechanistic approach adopted here is necessary to avoid antagonistic effects on other pests or on natural enemies.     

Polar paths of diffusion across plant cuticles: new evidence for an old hypothesis

BACKGROUND: The plant cuticle is an extracellular lipophilic biopolymer covering leaf and fruit surfaces. Its main function is the protection of land-living plants from uncontrolled water loss. In the past, the permeability of the cuticle to water and to non-ionic lipophilic molecules (pesticides, herbicides and other xenobiotics) was studied intensively, whereas cuticular penetration of polar ionic compounds was rarely investigated. RECENT PROGRESS: Recent work measuring cuticular penetration of inorganic and organic ions is presented; the effects of molecular size of ions, temperature, wax extraction, humidity and plasticizers strongly support the conclusion that ions penetrate cuticles via water-filled pores. The cuticle covering stomata and trichomes forms the preferential site of ion penetration. This indicates that cuticles possess a pronounced lateral heterogeneity: the largest fraction of the cuticle surface is covered by the lipophilic domains of cutin and wax, but to a certain extent polar domains are also present in the cuticle, which form preferential sites of penetration for polar compounds. THE FUTURE: The chemical nature of these polar domains awaits detailed characterization, which will be of major importance in agriculture and green biotechnology, since polar paths of diffusion represent the most important transport routes for foliar-applied nutrients. Furthermore, many compounds acting as inducers of gene expression in transgenic plants are ionic and need to penetrate the cuticle via polar paths in order to be active.     

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.