Characterization and Distribution of Water-repellent, Self-cleaning Plant Surfaces

During the last 20 years, a wealth of data dealing with scanningelectron microscopy of plant surfaces has been published. Theultrastructure of epidermal surfaces has been investigated withrespect to taxonomic, as well as functional aspects. Withinthe latter, water-repellency has received much attention andhas been well documented. Water-repellency is based on surfaceroughness caused by different microstructures (trichomes, cuticularfolds and wax crystals), together with the hydrophobic propertiesof the epicuticular wax. In addition, contaminating particlesare carried away by water droplets, resulting in a cleaned surface(Lotus-effect). Therefore, rough, waxy leaves are not only water-repellentbut anti-adhesive with respect to particulate contamination.Based on 200 water-repellent plant species, the present papersurveys micromorphological characteristics of anti-adhesiveplant surfaces. Leaves that are permanently water-repellentcan be differentiated by distictively convex to papillose epidermalcells and a very dense layer of epicuticular waxes. Leaves thatare water-repellent for only a limited period of time have onlyslightly convex epidermal cells and often have a less densewax layer. Finally, an overview is given on the occurrence ofwater-repellency among different life forms and within differenthabitats. Water-repellency is concentrated in herbaceous species,while it is rare in trees. Among different habitats, subtropicalregions, wetlands and disturbed areas appear to have more specieswith water-repellent leaves. The importance of roughness andwater-repellency, respectively, as the basis of an anti-adhesive,self-cleaning surface, in comparison to other functions of microstructures,is discussed.     

Seasonal changes of leaf surface contamination in beech, oak, and ginkgo in relation to leaf micromorphology and wettability

The leaf surfaces of beech, oak and ginkgo have been investigated with respect to contamination with particles during one growing season. Based on the observation that particles are removed from water-repellent leaves by rain (Lotus effect) the three species were selected because they differ in leaf surface micromorphology and wettability. Leaves of beech are smooth, lacked wax crystals and were ±wettable. Those of ginkgo were rough because their cells were convex and were densely covered by wax crystals, resulting in permanent water repellency. Leaves of oak were covered by waxes and were water repellent when young, but, a few weeks after leaf expansion had ceased the waxes were rapidly eroded. These differences in wettability resulted in different amounts of contamination. Ginkgo collected a very small number of particles during the whole vegetation period. In beech the contamination was significantly higher, but fairly constant, whereas oak leaves accumulated particles with age.     

On the epidermal structure ofboleophthalmus andscartelaos mudskippers with reference to their adaptation to terrestrial life

The skin structures of 4 species of oxudercine gobies (3 species ofBoleophthalmus and 1 species ofScartelaos) were investigated in relation to the terrestrial exposure of these fishes. These species have similarities in both lifestyle and skin structure. The specializations for terrestrial life mainly include the presence of dermal bulges, a thick middle cell layer, and a vascularized epidermis. Moreover, mucous cells are distributed only on the epidermis where the capillaries are undeveloped. In all species, the dermal bulges are large on the head and dorsal body, which are most often exposed to the air, and push up a thin epidermis, forming so-called papillae. Capillaries are densely distributed on the apical area of each papilla. InBoleophthalmus, the middle cell layer is thicker, the bulges are larger and distributed over a greater part of the body, and the area of the skin surface having the papillae is larger than it is inScartelaos. These differences suggest that the contribution of the skin to respiration is comparatively large inBoleophthalmus species, reflecting their more frequent activities on mudflats relative to the activities of theScartelaos species, which prefer to stay in the water. Mucous cells are abundantly distributed on the epidermis surface between the papillae in all species. The separation of the capillaries and the mucous cells may be due to an impeding of gas exchange by the mucus.     

Effects of Methanol on Wettability of the Non-Smooth Surface on Butterfly Wing

The contact angles of distilled water and methanol solution on the wings of butterflies were determined by a visual contact angle measuring system. The scale structures of the wings were observed using scanning electron microscopy, The influence of the scale micro- and ultra-structure on the wettability was investigated. Results show that the contact angle of distilled water on the wing surfaces varies from 134.0° to 159.2°. High hydrophobicity is found in six species with contact angles greater than 150°. The wing surfaces of some species are not only hydrophobic but also resist the wetting by methanol solution with 55% concentration. Only two species in Parnassius can not resist the wetting because the micro-structure （spindle-like shape） and ultra-structure （pinnule-like shape） of the wing scales are remarkably different from that of other species. The concentration of methanol solution for the occurrence of spreading/wetting on the wing surfaces of different species varies from 70% to 95%. After wetting by methanol solution for 10 min, the distilled water contact angle on the wing surface increases by 0.8°-2.1°, showing the promotion of capacity against wetting by distilled water.     

The development of the cuticle in Phormium tenax

Summary As seen in the scanning electron microscope the surface wax of leaves of Phormium tenax L. consists of vertical, plate-like crystals. These increase in size and number and undergo a change in form during development. The abaxial surface has a dense covering of wax crystals, but none are present on the ridges over vascular tissues. Numerous papillae are found between these ridges in later stages of development. On the adaxial surface both wax crystals and papillae are present only around infrequent stomata.When viewed in section normal to the leaf surface the cuticle is first apparent as a thin, lamellate layer. Another layer containing a reticulum of electrondense material increases in thickness beneath the lamellae during development. This layer eventually becomes the most extensive component of the cuticle. Both the adaxial and abaxial cuticles show a similar pattern of development.     

Interfacial Effects of Superhydrophobic Plant Surfaces： A Review

Nature is a huge gallery of art involving nearly perfect structures and properties over the millions of years of development. Many plants and animals show water-repellent properties with fine micro-structures, such as lotus leaf, water skipper and wings of butterfly. Inspired by these special surfaces, the artificial superhydrophobic surfaces have attracted wide attention in both basic research and industrial applications. The wetting properties of superhydrophobic surfaces in nature are affected by the chemical compositions and the surface topographies. So it is possible to realize the biomimetic superhydrophobic surfaces by tuning their surface roughness and surface free energy correspondingly. This review briefly introduces the physical-chemical basis of superhydrophobic plant surfaces in nature to explain how the superhydrophobicity of plant surfaces can be applied to different biomimetic functional materials with relevance to technological applications. Then, three classical effects of natural surfaces are classified： lotus effect, salvinia effect, and petal effect, and the promising strategies to fabricate biomimetic su- perhydrophobic materials are highlighted. Finally, the prospects and challenges of this area in the future are proposed.     

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。     

The effects of superficial wax on leaf wettability

Experiments are described which provide more information on the role played by superficial waxes in the natural water-repellency of leaf surfaces. Contact angles of water were measured on a variety of leaf surfaces, before and after removal of wax, and on smooth films of the isolated superficial waxes. The differences in wettability of leaf surfaces are not wholly accounted for by differences which occur in the chemical and hydrophobic properties of their superficial waxes. Waxes isolated from leaves exhibiting contact angles less than 90° are usually more hydrophobic than the leaf surface itself. On most leaves exhibiting angles greater than 90° wax is the dominant factor governing water-repellency, the isolated wax normally making at least a 60 % contribution to the contact angle measured on the leaf surface. Additional factors, such as roughness, responsible for the occurrence of contact angles greater than 110° on certain leaf surfaces, reside in the wax layer. The hydrophobic properties of some leaves are unaffected by chloroform washing, revealing that superficial waxes play little part in their wettability.     

A fast, precise and low-cost replication technique for nano- and high-aspect-ratio structures of biological and artificial surfaces

Biological surfaces are multifunctional interfaces between the organisms and their environment. Properties such as the wettability and adhesion of particles are linked to the micro- and nanostructures of their surfaces. In this study, we used plant and artificial surfaces covered with wax crystals to develop a low-cost replication technique with high resolution. The technique is applicable for fragile surface structures, as demonstrated for three-dimensional wax crystals, and is fast to prevent shrinking of the biological material by water loss during the molding process. Thermal evaporation of octacosan-1-ol has been used to create microstructured surfaces with small platelets as templates for molding. Epoxy resin as filling material provided the smallest deviations from the original surface structures and can be used for replication of nanostructures as small as 4.5 nm. Contact angle measurements of leaves and their replicas show that this technique can be used to develop biomimetic surfaces with similar wettability as in the plant surfaces.     

Leaf and stem epicuticular wax structures in Lonicera species (Caprifoliaceae)

Epicuticular waxes are found on the surface of the plant epidermis as crystal structures which show morphological diversity and may be useful in plant systematics. The aim of this study was to recognise the diversity of the epicuticular wax cover on leaves and stems of Lonicera species (Caprifoliaceae) and to estimate the taxonomic value of wax traits. Adaxial and abaxial sides of the leaves and the stem surface in 35 taxa were investigated through a scanning electron microscope (SEM). The assessment was prepared on the basis of the presence or absence of basic types of wax structures and their dimensions, using categorical response analysis, multiple correspondence analysis and cluster analysis. In the analysed species, we found differences in the wax layer structure on the surfaces of the leaves and stems. Apart from the smooth layer, tubules and platelets can be found. Tubules are considerably more frequent than platelets. In many cases, the same plant part produces more than just one wax type. Our analysis indicates that wax structures are not very informative for the taxonomy of Lonicera, because we observed only a weak correspondence between the presence or absence of some types of wax tubules and the infrageneric classification of the genus. However, in particular cases these new data can be used for species identification.     

A new species of Polymastia (Porifera, Hadromerida, Polymastiidae) from the Aleutian Islands, Alaska, USA

Polymastia fluegeli n. sp. is described from deep water off the Aleutian Islands (Alaska, USA). P. fluegeli is disc-shaped and lives partly buried in the sediment, with only the papillae protruding above the surface. This new species has a basal layer of agglutinated sediment particles occurring between the choanosome and the ectosomal lower layer. This conspicuous sediment layer is not described for other species of Polymastia. Occurring spicule types and sizes are different from known species of the area.     

Surface wax,structure and function in leaves of Polytrichaceae

Abstract

The form and distribution of epicuticular wax has been examined by SEM in 35 species of Polytrichaceae. Australian and New Zealand species of Dawsonia have a dense covering of prominent angular platelets (or more irregular wax) on the lamella-margins; Polytrichadelphus magellanicus is very similar. In Polytrichum the epicuticular wax forms an angular to rounded granularity which varies in quantity from species to species; abundant and prominent on the lamella-margins of, e.g., P. formosum and P. commune, hardly detectable in, e.g., P. sexangulare. Pogonatum spp. show sparse to dense angular to rounded granularity on both surfaces, generally more abundant on the upper surface but (with the exceptions of P. grandifolium and (especially) P. urnigerum) less differentiated on the lamella-margins than in Dawsonia or Polytrichum. The parallel between Polytrichum alpinum, Pogonatum grandifolium, P. urnigerum, Psilopilum australe and P. crispulum is noteworthy. Oligotrichum, Batramiopsis, Atrichum and the other species of Psilopilum show either no apparent surface wax or only a fine surface granularity, although their leaves are generally water-repellent, sometimes strongly so. Dendroligotrichum dendroides has almost smooth lamella-margins; D. squamosum has abundant and prominent surface wax.

A major function of epicuticular wax in Polytrichaceae is probably to exclude water from the ‘pseudo-mesophyll’ formed by the lamellae, a function supplemented by the inflexed leaf margins of Polytrichum section Juniperifolia. It is best developed on the lamella-margins of the species of Dawsonia, Polytrichadelphus, Polytrichum and Pogonaturn which probably most nearly approach mesophytic vascular plants in their pattern of adaptation.     

Histochemistry of the tongue epithelium in four mammals with respect to keratinization

The tongue epithelium was examined in the laboratory rat, guinea pig, rabbit and Domestic cat, using light microscopical, histological fluorescent and histochemical methods. The distributions of the enzymes, acid and alkaline phosphatase were examined. Protein-bound phospholipid and calcium were investigated, together with thiol sulphydryl groups and cysteine disulphide bonds of proteins. A variety of different types of keratinization were shown in the various species, as well as in the same species in different regions of the tongue. The most strongly keratinized structures were the filiform and conical papillae which varied widely from species to species. Those of the rat dorsum were similar to papillae described previously in the House mouse and have strongly keratinized spines. The guinea pig showed some differences but also had keratinized spines. In contrast the rabbit papillae did not have spines but the horny layer over the posterior sides was hardened instead to form pointed edges. Human filiform papillae are similar to the rabbit without spines but the horny layer is less strongly keratinized. In the Domestic cat the conical papillae were also without spines but the horny layer on the anterior and posterior surface was hardened to form claw-like structures.     

Attachment forces of pea aphids (Acyrthosiphon pisum) on different legume species

1. Sympatric populations of insects adapted to different host plants are good model systems not only to study how they adapt to the chemistry of their food plant, but also to investigate whether morphological modifications evolved enabling them to live successfully on a certain plant species. 2. The pea aphid, Acyrthosiphon pisum (Harris) encompasses at least 11 genetically distinct sympatric host races, each showing a preference for a certain legume species. The leaflet surfaces of these legumes differ considerably in their wax coverage. 3. It was investigated whether the attachment structures of three pea aphid genotypes from different host races are adapted to the different surface properties of their host plants and whether they show differences in their attachment ability on the respective host and non‐host plants. 4. The surface morphology of plants and aphid tarsi was examined using SEM (scanning electron microscopy). The ability of the aphids to walk on specific surfaces was tested using traction force measurements. 5. The presence of wax blooms on the leaflets lowers the aphids' attachment ability considerably and diminishes their subsequent attachment on ‘neutral’ surfaces like glass. The pea aphid host races differ in their ability to walk on certain surfaces. However, the genotype from the adapted aphid host race was not necessarily the one with the best walking performance on their host plant. All aphids, regardless of the original host plant, were most efficient on the neutral control surface glass. The general host plant Vicia faba was the plant with the most favourable surface for all aphid host races.     

Epicuticular wax structures on stems and comparison between stems and leaves – A survey

The cuticle, forming the outermost layer of plant tissues and being in direct contact with the environment, consists of waxes and cutin. Waxes are hydrophobic substances that are divided in two groups: intra- and epicuticular, depending on their localisation. Epicuticular waxes appear as smooth coverings, however, many plants also produce superimposed wax structures of a crystalline nature. While studies of waxes have almost exclusively focused on leaves, here a survey of epicuticular wax structures on stems is presented. The stem surface of 343 higher plant taxa, representing 80 families, was examined using scanning electron microscopy. The adaxial and abaxial surfaces of leaves of 319 taxa were also examined to determine the relationship between wax structures on stems and leaves. Wax structures are classified, described and discussed. The results of the study indicate that stems exhibit the same main wax crystal types that have been described for leaves. Seventy percent of the examined taxa produced wax crystals on their stems. In ∼24% of the taxa, wax crystals were absent on leaves and found only on stems. In plant taxa that produce wax crystals, 40% exhibit the same type on either side of their leaves and on their stem. However, a much stronger morphological similarity exists between crystal shapes present on the adaxial and abaxial surfaces of leaves than between those present on the stem and those on leaves. In general, these observations suggest that stems are quite different than leaves in terms of their epicuticular wax structures.     

Fine structure and distribution of epidermal projections associated with taste buds on the oral papillae in some loricariid catfishes (Siluroidei: Loricariidae)

Two morphologically distinct structures occur on the surfaces of the oral papillae in several loricariid catfish species; namely, (1) typical vertebrate taste buds composed of receptor and sustentacular cells and (2) brushlike projections, termed epidermal brushes, that represent specialized epidermal cells containing keratin. Both of these structures were studied with the combined use of light microscopy and scanning and transmission electron microscopy. The general body surface, fins, and rostral cutaneous processes of some loricariid catfishes are covered with taste or terminal buds but lack the epidermal brushes. It is suggested that the epidermal brushes found on the oral papillae serve as protective devices for the taste buds and as abrasive surfaces for substrate scraping during feeding. The taste buds on the oral papillae may detect any gustatory stimuli from the resulting substrate disturbance. Comparative studies reveal many differences in the number and spatial arrangement of these two structures on the oral papillae among the several species of the Loricariidae examined. These differences may represent functional adaptations to the various modes of life in the Loricariidae.     

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.     

不同生境下十七种藓类植物叶的比较解剖学

使用石蜡切片法,对不同生境下的17种藓类植物的叶片进行了解剖观察和比较分析,结果表明不同种类的藓类植物在中肋导水主细胞的有无、厚壁细胞是否分化、中肋细胞层数及细胞密度、叶片细胞层数、叶表附属物、叶片细胞密度等方面存在显著差异。藓类植物叶的解剖结构具有生态适应意义,旱生环境下的藓类植物,叶片细胞胞壁具不同程度的增厚,有些藓类植物叶片具附属结构,藓类植物中肋的有无,反映了对水分吸收和运输方式的不同。例如,荫湿生环境下的羽枝青藓Brachythecium plumosum,其中肋细胞胞壁较薄,无导水主细胞和副细胞的分化,也没有厚壁细胞分化,能够在阴湿环境下吸收水分和养分;钝叶匍灯藓Plagiom niumrostratum具有与旱生藓类植物相似的中肋结构,叶片较厚,中肋具导水主细胞,中肋背面具厚壁细胞,这些特点使该种藓类植物能够分布于间隙性干旱胁迫的环境中;水灰藓Hygro-hypnum luridum叶片纤细柔弱,仅1层细胞,细胞胞壁薄,叶表无附属结构,中肋细胞层数少,无导水主细胞分化,也没有厚壁细胞,这些特点使得水灰藓能够生长在水生环境中;东亚小金发藓Pogonatum inflexum和波叶仙鹤藓Atrich umundulatum的叶腹面覆盖着栉片,东亚砂藓Racomitrium japonicum、大羽藓Thuidium cym-bifolium、福氏蓑藓Macromitrium ferriei、东亚短颈藓Diphyscium fulvifolium、扭口藓Barbula unguiculata和角齿藓Ceratodon purpureus的叶片表面有乳头状突起或疣状物,这些附属结构使它们能够适应于旱生的环境中。     

Purity of the sacred lotus, or escape from contamination in biological surfaces

The microrelief of plant surfaces, mainly caused by epicuticular wax crystalloids, serves different purposes and often causes effective water repellency. Furthermore, the adhesion of contaminating particles is reduced. Based on experimental data carried out on microscopically smooth (Fagus sylvatica L., Gnetum gnemon L., Heliconia densiflora Verlot, Magnolia grandiflora L.) and rough water-repellent plants (Brassica oleracea L., Colocasia esculenta (L.) Schott., Mutisia decurrens Cav., Nelumbo nucifera Gaertn.), it is shown here for the first time that the interdependence between surface roughness, reduced particle adhesion and water repellency is the keystone in the self-cleaning mechanism of many biological surfaces. The plants were artificially contaminated with various particles and subsequently subjected to artificial rinsing by sprinkler or fog generator. In the case of water-repellent leaves, the particles were removed completely by water droplets that rolled off the surfaces independent of their chemical nature or size. The leaves of N. nucifera afford an impressive demonstration of this effect, which is, therefore, called the “Lotus-Effect” and which may be of great biological and technological importance. Received: 19 August 1996 / Accepted: 12 November 1996     

An analysis of the epicuticular wax of Chenopodium album leaves in relation to environmental change, leaf wettability and the penetration of the herbicide bentazone

The fine-structure, epicuticular wax and contact angle characteristics of Chenopodium album leaf surfaces have been examined with respect to the control of this weed by the herbicide bentazone. Epicuticular wax analysis and contact angle measurements suggest a relative impermeability to bentazone (BAS 35107H), that may be enhanced by the inclusion of the oil adjuvant Actipron (BP Oil Ltd) to the formulation. Under different regimes of light, temperature and relative humidity the chemical composition of the epicuticular wax layer remained relatively constant, whereas herbicide efficacy was greatly altered. These findings are discussed in relation to the mode of action and penetration of bentazone in this species.