Mielichhoferia elongata (Hornsch.) Hornsch. and Saelania glaucescens (Hedw.) Broth. in Scotland

Abstract

Lamella margins of Polytriehadelphus from throughout the range of the genus have been examined by SEM. All specimens clearly referable on other characters to the genus have characteristically crenate lamella margins covered with angular flakes of epicuticular wax. In this respect they resemble species of Dawsonia but differ from all other Polytrichaceae examined. The lamella margin characters are useful for critical identification of non-fruiting plants of Polytriehadelphus from other vegetatively similar Polytrichaceae. They confirm that P. erieoides and the North American P. lyallii Mitt. do not belong to this genus. Polytriehadelphus has a trans-Pacific and essentially Southern Hemisphere distribution extending from Sulawesi and New Guinea through New Zealand and the Fuegian region to the northern Andes, with a north-western outpost in Costa Rica. It is likely that this distribution is derived from a continuous Antarctic range in the late Cretaceous, and that much of the radiation of the genus at the two northern ends of its present range took place in the mid to late Tertiary. Outlying stations on Tristan da Cunha and the Marquesas probably result from long-distance dispersal.     

Isozyme evidence regarding the origins of three allopolyploid species ofPolytrichastrum (Polytrichaceae, Bryophyta)

Electrophoretic data show thatPolytrichastrum pallidisetum, P. ohioense, andP. sexangulare are allopolyploids. They display fixed, heterozygous banding patterns at five to six of the 11 enzyme loci that we screened. In total, we sampled 304 populations representing three genera (Polytrichastrum, Polytrichum, andPogonatum) and 18 species in our examination of the allopolyploids and their putative haploid progenitors. There were no extant species that fit perfectly as a progenitor of any allopolyploid. BothP. pallidisetum andP. ohioense appear to have originated as intergeneric hybrids between one progenitor with aPolytrichastrum-type genome and another with aPolytrichum-type genome. The extant haploid species with the most similar genomes to the putative progenitors ofP. pallidisetum werePolytrichastrum appalachianum andPolytrichum commune, whereas forPolytrichastrum ohioense the species werePolytrichastrum formosum (orP. longisetum) andPolytrichum commune. Polytrichastrum sexangulare was genetically more similar to species ofPogonatum, as was one taxon that appears to be one of its progenitors,P. sexangulare var.vulcanicum ( =Polytrichum/Pogonatum sphaerothecium). The other progenitor also must have possessed alleles that are common in species ofPogonatum. The Polytrichaceae are a relatively ancient group of mosses, and the hybridizations that gave rise to these allopolyploids may have occurred long ago. It is likely that the genomes of the original progenitors have changed over time or that those progenitors are now extinct.     

Wandlabyrinthe im Sporophyten von Polytrichum

Zusammenfassung Die Wandlabyrinthe im Sporophyten dreier Polytrichum-Arten (P. commune, P. formosum und P. juniperinum) wurden im Elektronenmikroskop untersucht.In den äußeren 4–6 Zellreihen des Sporogonfußes nimmt der Differenzierungsgrad des Labyrinthes radial von innen nach außen zu.Bei Polytrichum formosum wurde eine Beziehung der Protuberanzenentstehung zur Lyse der Zellwand gefunden.Die Protuberanzen besitzen Schichtbau, ebenso wie die Wand, aus der sie hervorgehen. Sie sind auch in der späteren, mächtigen Wandauflagerung an ihrem Kontrast kenntlich (Polytrichum commune). Bei Polytrichum formosum entstehen, ausgehend von diesen ursprünglich isolierten Protuberanzen, in den Wandvorsprüngen Zonen starken Kontrastes, deren Umrisse jeweils den Appositionsgrenzen zu folgen scheinen. Im Gegensatz dazu treten bei Polytrichum juniperinum in der Wandsubstanz kontrastreiche Einschlüsse auf, die keinerlei Beziehung zum ursprünglichen Bau des Wandlabyrinths zeigen. Die Menge dieser Einschlüsse scheint von der Wachstumsintensität des Sporophyten abzuhängen, und es wurde vermutet, daß es sich dabei um Exkrete handelt.In der Wandsubstanz waren die verschiedensten Strukturen zu beobachten, deren Herkunft und Entstehungsmöglichkeiten diskutiert wurden.

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Wall labyrinths in the sporophyte of Polytrichum

Summary In the foot of the Polytrichum sporophyte the outer walls of the peripheral cells are coated with a labyrinth. In the outer 4–6 cell layers protuberances arising from the lamellated cell wall were found which are interpreted as initial stages of the labyrinth. In Polytrichum formosum there is a correlation between the lysis of the cell wall and the initiation of new protuberances. In the outer cell layers the protuberances increase more and more and fuse. In the external cells large aggregations of wall substances are deposited, in which the protuberances are seen as electron dense cores. In the wall substances of the external cells there are large inclusions of an electron dense material, the accumulation of which seems to be in relation to the growth intensity of the sporophyt. Some structures enclosed in the wall material seem to be membraneous elements.

     

Scapania mucronata Buch in Scotland

Abstract

Regeneration from detached leaves at 20°C in 12 h days was studied in six species. Of the four species of Polytrichum and two of Pogonatum studied, only Pogonatum aloides did not regenerate from leaves. This result could be related to the fact that in nature only this species possesses persistent primary protonemata. Polytrichum commune, P. formosum and P. juniperinum developed long, muchbranched secondary protonemata which produced buds at intervals. Pogonatum urnigerum developed buds without an initial protonema, and Polytrichum piliferum short, unbranched protonemal threads, each bearing a single terminal bud. In all cases regenerants arose from either the large cells at the bases of lamellae, or less often, basal cells of the lamellae themselves. In Pogonatum urnigerum the amount of regeneration tended to decrease from apex to base of leaf, while in the Polytrichum species the reverse occurred. No definite conclusions could be drawn with regard to the effect, if any, of leaf age on regeneration. Possibly the temperature experienced beforehand or some other seasonal factor, affects regeneration, but the question needs further investigation.     

Effect of epicuticular wax crystals on the localization of artificially deposited sub-micron carbon-based aerosols on needles of <Emphasis Type="Italic">Cryptomeria japonica</Emphasis>

Elucidation of the mechanism of adsorption of particles suspended in the gas-phase (aerosol) to the outer surfaces of leaves provides useful information for understanding the mechanisms of the effect of aerosol particles on the growth and physiological functions of trees. In the present study, we examined the localization of artificially deposited sub-micron-sized carbon-based particles on the surfaces of needles of Cryptomeria japonica, a typical Japanese coniferous tree species, by field-emission scanning electron microscopy. The clusters (aggregates) of carbon-based particles were deposited on the needle surface regions where epicuticular wax crystals were sparsely distributed. By contrast, no clusters of the particles were found on the needle surface regions with dense distribution of epicuticular wax crystals. Number of clusters of carbon-based particles per unit area showed statistically significant differences between regions with sparse epicuticular wax crystals and those with dense epicuticular wax crystals. These results suggest that epicuticular wax crystals affect distribution of carbon-based particles on needles. Therefore, densely distributed epicuticular wax crystals might prevent the deposition of sub-micron-sized carbon-based particles on the surfaces of needles of Cryptomeria japonica to retain the function of stomata.     

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.     

Slippery ant-plants and skilful climbers: selection and protection of specific ant partners by epicuticular wax blooms in Macaranga (Euphorbiaceae)

 In many ant-plant species of the genus Macaranga in South-East Asia, conspicuous blooms of epicuticular wax crystals cover the stem surface. We found that many ant species were unable to walk on these surfaces. Only the specific ant partners of glaucous Macaranga host plants were capable of moving on the slippery stems without difficulty. Therefore, the epicuticular coatings of Macaranga myrmecophytes appear to have a selective function and protect the associated ants against competitors. The epicuticular aggregates function as a physical barrier; no evidence of chemical repellence was found. The extent to which ”foreign” ant species are excluded from a tree strongly depends on inclination, diameter and length of the glaucous stem sections. The particular growth form of some glaucous Macaranga ant-plants enhances the influence of the wax barriers. The ant associates of glaucous and glossy Macaranga ant-plants (genera Crematogaster and Camponotus) differ strongly in their capacity to adhere to the glaucous stems. For this reason, the wax blooms in Macaranga can act as an ecological isolation mechanism for the sympiotic ants. Within the genus Macaranga, we find a high correspondence between the occurrence of glaucousness and obligatory ant association (50% in ant-plants; 6.7% in non-myrmecophytes). The genus Macaranga thus represents one of the few cases known so far where epicuticular wax crystals are likely to have evolved in relation to insects. Received: 2 January 1997 / Accepted: 9 June 1997     

TRANSLOCATION IN POLYTRICHUM COMMUNE (BRYOPHYTA). III. LOADING OF SUGARS IN SOURCE LEAVES

Polytrichum commune, the common hair-cap moss, possesses a system of long-distance food-conducting cells that can be traced from leaves into stems and down through underground rhizomes. Cytochemical analysis indicates that high adenosine triphosphatase activity is associated with the membranes of sugar-conducting deuter cells in Polytrichum leaves. Incipient plasmolysis determinations reveal high solute concentrations of leaf deuters. These two lines of evidence suggest that long-distance transport of photosynthate is initiated in this species by a process analogous to phloem loading of minor veins in leaves of flowering plants. Two sets of experimental observations suggest that sugar loading in Polytrichum is coupled with the transport of protons: the moderating effects of N-ethylmaleimide, p-chloromercuribenzenesulfonic acid, sodium orthovanadate, and fusicoccin on labeled sucrose uptake (as determined by liquid scintillation techniques), and correlated effects on acid flux from isolated leaves (as determined by computer-aided titrimetry).     

Chemical composition of epicuticular wax crystals on the slippery zone in pitchers of five <Emphasis Type="Italic">Nepenthes</Emphasis> species and hybrids

Plants of the carnivorous genus Nepenthes efficiently trap insects in leaf pitchers, mostly employing epicuticular wax crystals on the pitcher walls to make them slippery for the prey. In the present study, the compositions and micromorphologies of the wax crystals of five Nepenthes species and hybrids were analysed in order to test whether the chemical principles underlying this ecological function are widespread within the genus. Three wax layers could be distinguished within the Nepenthes pitcher cuticles: (1) the outermost part of the crystals forming the platelets visible in standard scanning electron microscopy, (2) the bottom portion of the epicuticular wax crystals, and (3) an intracuticular wax layer. The composition of the intracuticular wax differed significantly from that of the neighbouring epicuticular layer. The compositions of corresponding wax mixtures from all five Nepenthes species and hybrids were very similar, with almost equal amounts of very long chain aldehydes and primary alcohols. While triacontanal (C₃₀ aldehyde) was prevailing in the epicuticular crystals of Nepenthes albomarginata and Nepenthes x intermedia, Nepenthes x superba and Nepenthes x henriana were found to have especially high percentages of dotriacontanal (C₃₂ aldehyde). Nepenthes “khasiana” had an intermediate aldehyde composition with almost equal amounts of both chain lengths.     

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.     

EVIDENCE FOR THE PRESENCE OF LIGNIN IN MOSS GAMETOPHYTES

Lignin has been established as a constituent of the gametophyte axes in the giant New Zealand mosses Dawsonia sp. and Dendroligotrichum sp. Isolated products comprised ca. 6-10% of the dry weight of gametophyte axes and contained 61-62% C, 6.4-6.8% H, and 5.1-7.9% OCH₃. Characteristic color reaction and ultraviolet spectra were observed, and alkaline nitrobenzene oxidation yielded perhaps 14-18% of mixed aldehydes as their 2,4-dinitrophenylhydrazones. The presence of substantial lignin in these exceptionally tall upright moss gametophytes contrasts strikingly with north temperate species such as Polytrichum, Funaria, Bryum and others, and lends support to the hypothesis that lignification is a mechanically and/or gravitationally regulated process.     

Studies on stomatal function, epicuticular wax and stem-root transition region of polyethylene glycol-treated and nontreatedin vitro grape plantlets

Summary Scanning electron microscopy, light microscopy, and gravimetric analysis was used to evaluate stomatal function, epicuticular wax, and the stem-root transition region of grape (Vitis sp. ‘Valiant’) plantlets grownin vitro, polyethylene glycoltreatedin vitro, and greenhouse-grown plants. Scanning electron microscopic studies of leaf surfaces ofin vitro-grown plants showed widely open stomata as compared to leaf stomata of polyethylene glycol-treatedin vitro-cultured and greenhouse-grown plants. Ultrastructurally, leaf epicuticular wax ofin vitro plants was less dense than in their polyethylene-treated and greenhouse counterparts. Quantitatively,in vitro-grown plants had reduced epicuticular was as compared to polyethylene glycol-treated and greenhouse-grown plants. Light microscopic studies showed no obvious differences in the vascular connections in the stem-root transition region ofin vitro-cultured, polyethylene glycol-treatedin vitro-cultured, and greenhouse-grown plants. It is therefore likely that the rapid wilting and desiccation observed after transplantingin vitro grape plantlets is due to their defective stomatal function and reduced epicuticular wax and may not be due to poor water transport associated with vascular connection.     

Direct and interaction-mediated effects of environmental changes on peatland bryophytes

Ecosystem processes of northern peatlands are largely governed by the vitality and species composition in the bryophyte layer, and may be affected by global warming and eutrophication. In a factorial experiment in northeast China, we tested the effects of raised levels of nitrogen (0, 1 and 2 g m⁻² year⁻¹), phosphorus (0, 0.1 and 0.2 g m⁻² year⁻¹) and temperature (ambient and +3°C) on Polytrichum strictum, Sphagnum magellanicum and S. palustre, to see if the effects could be altered by inter-specific interactions. In all species, growth declined with nitrogen addition and increased with phosphorus addition, but only P. strictum responded to raised temperature with increased production of side-shoots (branching). In Sphagnum, growth and branching changed in the same direction, but in Polytrichum, the two responses were uncoupled: with nitrogen addition there was a decrease in growth (smaller than in Sphagnum) but an increase in branching; with phosphorus addition growth increased but branching was unaffected. There were no two-way interactions among the P, N and T treatments. With increasing temperature, our results indicate that S. palustre should decrease relative to P. strictum (Polytrichum increased its branching and had a negative neighbor effect on S. palustre). With a slight increase in phosphorus availability, the increase in length growth and production of side-shoots in P. strictum and S. magellanicum may give them a competitive superiority over S. palustre. The negative response in Sphagnum to nitrogen could favor the expansion of vascular plants, but P. strictum may endure thanks to its increased branching.     

Spruce budworm feeding and oviposition are stimulated by monoterpenes in white spruce epicuticular waxes

Monoterpenes, source of the distinctive odor of conifers, are generally considered plant defensive compounds. However, they are also known to act as long‐range insect attractants, as they are volatile and permeate forest airspaces. Moreover, they are lipid soluble and can be absorbed into plant epicuticular waxes. We test their role in short‐range host plant choice by both adult females and larvae of a folivorous forest pest (Choristoneura fumiferana). We conducted laboratory assays testing the responses of Eastern spruce budworm to an artificial monoterpene mix (α‐pinene, β‐pinene, limonene, myrcene) and to white spruce (Picea glauca) epicuticular waxes in closed arenas. Ovipositing females preferred filter paper discs treated with P. glauca waxes to controls, and preferred the waxes + monoterpenes treatment to waxes alone. However, females showed no preference between the monoterpene‐treated disc and the control when presented without waxes. Feeding larvae prefered wax discs to control discs. They also consumed discs treated with realistic monoterpene concentrations and wax preferentially over wax‐only discs, but showed no preference between extremely high monoterpene concentrations and wax‐only controls. In an insect‐free assay, P. glauca epicuticular wax decreased monoterpene volatilization. These results suggest that P. glauca waxes and realistic concentrations of monoterpenes are stimulatory to both egg‐laying females and feeding larvae, and that their effects are synergistic.     

Two sides of a leaf blade: <Emphasis Type="Italic">Blumeria graminis</Emphasis> needs chemical cues in cuticular waxes of <Emphasis Type="Italic">Lolium perenne</Emphasis> for germination and differentiation

Plant surface characteristics were repeatedly shown to play a pivotal role in plant–pathogen interactions. The abaxial leaf surface of perennial ryegrass (Lolium perenne) is extremely glossy and wettable compared to the glaucous and more hydrophobic adaxial surface. Earlier investigations have demonstrated that the abaxial leaf surface was rarely infected by powdery mildew (Blumeria graminis), even when the adaxial surface was densely colonized. This led to the assumption that components of the abaxial epicuticular leaf wax might contribute to the observed impairment of growth and development of B. graminis conidia on abaxial surfaces of L. perenne. To re-assess this hypothesis, we analyzed abundance and chemical composition of L. perenne ab- and adaxial epicuticular wax fractions. While the adaxial epicuticular waxes were dominated by primary alcohols and esters, the abaxial fraction was mainly composed of n-alkanes and aldehydes. However, the major germination and differentiation inducing compound, the C₂₆-aldehyde n-hexacosanal, was not present in the abaxial epicuticular waxes. Spiking of isolated abaxial epicuticular Lolium waxes with synthetically produced n-hexacosanal allowed reconstituting germination and differentiation rates of B. graminis in an in vitro germination assay using wax-coated glass slides. Hence, the absence of the C₂₆-aldehyde from the abaxial surface in combination with a distinctly reduced surface hydrophobicity appears to be primarily responsible for the failure of normal germling development of B. graminis on the abaxial leaf surfaces of L. perenne.     

Genetic Evidence for the Allodiploid Origin of the Moss Species Polytrichum longisetum

Abstract: Fixed heterozygous banding patterns observed for 4 allozyme and 12 microsatellite loci, in combination with a chromosome number of 14, show that Polytrichum longisetum is an allodiploid species. Comparison of these banding patterns with those of related Polytrichaceae species suggest that Polytrichum formosum, or an ancestor taxon of this species, is one of the haploid progenitors of P. longisetum. The second progenitor species of P. longisetum could not be designated in this study as it was not among the examined possible progenitor species. DNA sequence data for two microsatellite loci, however, suggest that the second progenitor species should possibly be more closely related to P. formosum than any of the other haploid possible progenitor species examined in this study. As the current systematic literature does not mention the existence of such a species, this could indicate that the second progenitor species is already extinct.     

Cabbage waxes affect Trissolcus brochymenae response to short‐range synomones

We show that induced synomones, emitted as a consequence of Murgantia histrionica activity on Brassica oleracea, are adsorbed by the epicuticular waxes of leaves and perceived by the egg parasitoid Trissolcus brochymenae. Leaves were exposed to M. histrionica females placed on the abaxial leaf surface. After 24 h, the leaves were treated mechanically using gum arabic, or chemically using chloroform, on the adaxial surface, and finally the adaxial surface was assayed with T. brochymenae by two‐choice tests in a closed arena. Wasp females responded to mechanically dewaxed cabbage leaf portions with feeding punctures and footprints (Ff) and with feeding punctures, oviposition and footprints (FOf), showing no effect of wax removal. In contrast, the removal of the epicuticular waxes from leaf portions close to FOf, and from leaves with oviposition and footprints (Of), determined the lack of responses by T. brochymenae. Solvent extracts of different treatments were bioassayed, but only FOf triggered parasitoid response. Thus the detection of oviposition‐induced synomones by the parasitoid depends on their adsorption by the epicuticular waxes. Mechanical wax removal from leaf portions contaminated with host footprints (f) also determined a lack of wasp responses, suggesting that the footprints might trigger the induction of a “footprint‐induced synomone” adsorbed onto the epicuticular waxes and exploited by the parasitoid. Leaf portions with the abaxial lamina previously dewaxed and then contaminated by footprints (D+f) of M. histrionica did not affect the parasitoid response, indicating that the abaxial epicuticular waxes are not directly involved in the chemicals induced by M. histrionica footprints.     

Polytrichum Strictum as a Solution to Frost Heaving in Disturbed Ecosystems: A Case Study with Milled Peatlands

Substrate instability is a common problem in many disturbed ecosystems. In the case of milled harvested peatlands, the pioneer moss Polytrichum strictum is commonly found; it is well adapted to tolerate the harsh microclimatic conditions and peat instability of these sites. A field experiment was used to determine the effectiveness of P. strictum against frost heaving, a major type of disturbance on bare peat. Wooden dowels and fir trees (Abies balsamea) placed in a P. strictum carpet experienced almost no frost heaving, whereas heaving was severe on bare peat. Reintroduced P. strictum fragments thinly spread on bare peat reduced but did not eliminate frost heaving. Straw mulch (a protective cover often required in peatland restoration) effectively reduced heaving in the fall, but was less effective in the spring because it had partially decomposed. The P. strictum carpet, P. strictum fragments, and straw mulch reduced frost heaving by reducing the number of freeze–thaw cycles, by slowing the rate of ground thaw in the spring, and by reducing the unfrozen water content of the peat during the spring thaw. Different species of Polytrichum mosses should be considered for the restoration or regeneration of disturbed ecosystems where soil stability is problematic.     

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.     

Notes on water relations of Polytrichum commune Hedw.

Abstract

Water loss in the endohydric moss Polytrichum commune was found to be controlled by a complex series of leaf arrangement changes, and by changes in water potential deficit of the shoots. This contrasted with the water relations of the ectohydric moss Rhacomitrium lanuginosum in which there was apparently little control over loss. Water conduction in Polytrichum was predominantly internal under high evaporative flux, and external under moderate flux, but under many stress conditions both pathways would probably be necessary to maintain an optimum water balance.