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The arbuscular mycorrhiza (AM) is one of the most important symbioses on earth, formed between about 80% of vascular plants and AM fungi (Smith & Read, 1997), which are placed in the order Glomales (Morton & Benny, 1990). A discussion has recently arisen about the rDNA-gene and general genome organization of this fungal group, which, as far as is known, does not possess sexuality (Sanders, 1999). This discussion mainly arose as a result of the assumption that one spore of an AM fungus (Scutellospora castanea) contains small subunit (SSU) rRNA genes that belong, phylogenetically, to different families within the Glomales, namely the Gigasporaceae and Glomaceae (Hosny et al., 1999). Questions about genetic drift, infrequent recombination, concerted evolution and the validity of phylogenetic techniques have arisen (Hosny et al., 1999; Sanders, 1999). However, this assumption might well be based on the sequence of an Ascomycete contaminant.

The male sterile mutant, ms35, of Arabidopsis thaliana was produced by X-irradiation of seeds. The mutant produces fertile pollen, but is male sterile because the anthers do not dehisce. Anther development in ms35 plants occurs as in wild-type Arabidopsis until shortly after microspores are released from meiotic tetrads. Thereafter, in the wild type, bands of lignified, cellulosic secondary wall thickenings are laid down around the cells of the anther endothecium. In contrast, wall thickenings are not formed in the endothecium of the ms35 mutant. Development of other lignified tissues, for example the vascular tissue of the stamen, occurs normally in ms35 plants. In mutant anthers, as pollen maturation is completed, the stomium is cleaved but the anther wall does not retract to release pollen. The block in anther dehiscence in ms35 plants is specifically correlated with the absence of endothecial wall thickenings. The ms35 mutation represents the first genetic evidence in support of the proposed role of the endothecium in anther dehiscence. The ms35 gene was mapped to the top arm of chromosome 3 (hy2-(4.17±2.31 cM)-ms35-(32.14±5.45 cM)-gl1).

The results presented in this paper describe the short- and long-term toxicity of arsenate in Silene vulgaris. Short-term toxicity, measured as inhibition of root elongation, depended on phosphate nutrition, arsenate being much less toxic at high phosphate supply. At low phosphate levels more arsenic was taken up by the plants. Under chronic exposure, toxicity (measured as inhibition of biomass production) did not increase with time. In addition, the accumulation of phytochelatins (PCs) as a function of toxicity and duration of exposure was studied. Short-term PC accumulation (over a 3 d period) was positively correlated with exposure. Isolation of peptide complexes from prolongedly exposed plants showed that PC2, PC3 and PC4 were present, although the latter not until at least 3 d exposure. Arsenic co-eluted mainly with PC2 and PC3. Fractions containing PC4 were devoid of As, probably due to dissociation of the complexes during extraction or elution. The breakdown of PCs after arresting As exposure was very slow. This could explain the continuous accumulation of PCs throughout longer periods of As exposure.

Branch bags were used to expose branches on mature Sitka spruce trees to either ambient [CO2] (A) or elevated [CO2] (E) for 4 yr. This paper reports the effects of this treatment on the growth, development and phenology of the branches, including shoot expansion, shoot numbers, needle dimensions, needle numbers and stomatal density. The effect of elevated [CO2] on the relationship between leaf area and sapwood area was investigated. Exposure to elevated [CO2] doubled photosynthetic rates in current-year shoots and, despite some down-regulation, 1-yr-old E shoots also had higher rates of photosynthesis than their A counterparts. Thus, the amount of assimilate fixed by E branches was substantially more than that fixed by A branches; however, this increase in the local production of assimilate did not lead to an increase in non-structural carbohydrate or stimulate growth or meristematic activity within the E branches. There was a very consistent relationship between leaf area and stem cross-sectional area that was not influenced by [CO2]. However, unbagged branches had thicker stems than bagged branches, resulting in a slightly lower ratio of leaf area to cross-sectional area. The implications of the results for the modelling of growth and allocation and the potential utility of the branch bag technique are discussed.

The study reported was conducted to establish the impacts of photochemical oxidants (primarily ambient ozone, O3) on the yield of watermelon (Citrullus lanatus) at a site on the east coast of Spain. Fruit yield and quality were monitored in plots established in a commercial watermelon field exposed, in open-top chambers (OTCs), to non-filtered air (NFA; near-ambient levels of ozone) or charcoal-filtered air (decreased levels of photochemical oxidants including O3; CFA), or to ambient air (AA), during the 1988 and 1989 growing seasons. Ambient levels of O3 were found to exceed present UN-ECE (United Nations Economic Commission for Europe; Convention on Long-Range Transboundary Air Pollution) critical level guidelines for the protection of crop yield by approx. twofold in 1988 and by approx. fivefold in 1989. Plants exposed to NFA and AA developed visible O3 injury on the upper surface of sun-exposed older leaves, and fruit yield (annual marketable fruit weight and number) was found to be depressed in OTCs ventilated with NFA in comparison with those receiving CFA. Consistent with inter-annual variations in O3 exposure, greater yield losses were experienced in 1989 (39%) than in 1988 (19%), an effect mediated predominantly by a decline in fruit number rather than average fruit weight. Exposure to ambient levels of ozone also slightly decreased fruit quality (4–8% decline in soluble solids content). Leaf gas exchange measurements made in the field in 1988 revealed effects of O3 on fruit yield and quality to be associated with a decline in the net CO2 assimilation rate per unit leaf area under light saturation (Asat) and stomatal conductance to water vapour (gs), and enhanced rates of dark respiration. A/ci curves (where A is the net CO2 assimilation rate per unit leaf area and ci is the mole fraction of CO2 in the leaf intercellular air space) constructed for plants grown in laboratory-based closed chambers, and exposed to an accumulated O3 exposure similar to that experienced by plants in the field, suggested that the likely cause of the decline in photosynthetic capacity was (1) a decrease in the amount and/or activity of Rubisco and (2) an impaired capacity for regeneration of ribulose 1,5-bisphosphate, which was not mediated through changes in the photochemical efficiency of photosystem II (Fv/Fm, where Fv is variable chlorophyll a fluorescence and Fm is maximum chlorophyll a fluorescence). No shift in the relative stomatal limitation to photosynthesis was observed under the influence of O3, suggesting that the decline in gs induced by the pollutant in both field and laboratory was the result, and not the cause, of the decrease in Asat. Ozone exposure also caused a decrease in C isotope discrimination (approx. 0.5‰), a shift that revealed a departure from predicted theory based on supporting leaf gas exchange measurements. The study demonstrates that ambient levels of photochemical oxidants on the Spanish Mediterranean coast are high enough to adversely influence the yield and physiology of an economically important crop grown in the region, and the magnitude of the effects was greater than would be predicted from exposure–yield-loss relationships for grain and fodder crops in central and northern Europe. Implications for the derivation of UN-ECE ozone standards are discussed, along with the mechanistic basis of the observed yield decreases.

Seedlings of 12 provenances of European beech (Fagus sylvatica) were exposed to ambient, non-filtered air (NF) or NF+50 nl l−1 ozone (NF50) for 8 h d−1 in open-top chambers (OTCs), from 1 June to 4 October 1995. In 1996 exposure was continued from 31 May to 1 October at four levels: charcoal-filtered air (CF), NF, NF50 and NF+100 nl l−1 ozone (NF100). Provenances were grown for both seasons in outside reference plots. All treatments were replicated. Ozone did not affect gas exchange in the provenances until late in the second season. NF100 reduced photosynthesis by 18% in August 1996 compared to CF. In September, photosynthesis was reduced by 22% in NF50 and by 29% in NF100. After two seasons, ozone reduced the root:shoot ratio by 24% when comparing CF and NF100; this was caused by reductions in the root biomass. Ozone did not affect height growth or stem diameter, and there were no ozone×provenance interactions for any growth parameter. There was, however, a significant ozone×provenance interaction for photosynthesis, showing northwest European provenances to be more sensitive to ozone than southeast European provenances when comparing dose–response estimates. This is interpreted in terms of genetic adaptation of the photosynthetic apparatus to regional growing conditions. Seedlings in the chambers grew 45% taller, and had 28% more shoot biomass and 29% smaller root biomass, resulting in a 44% reduction of root:shoot ratios compared to seedlings outside. Increased temperature and decreased PAR inside the chambers relative to the outside were probably the main causes for the differences. The magnitude of the chamber effects in OTCs raises doubts about conclusions drawn from ozone exposures in such chambers. This and previous ozone experiments with OTCs may have reached inaccurate conclusions concerning the size of ozone responses due to chamber effects.

The effects of enhanced UV-B radiation were investigated in the carnivorous plant Pinguicula vulgaris in a field experiment performed in Abisko, North Sweden (68° 21′ N, 18° 49′ E, 380 m above sea level). Potted plants were exposed to either ambient or ambient plus supplemental UV-B radiation, simulating a 15% ozone depletion. No effect was observed on either the epicuticular (external) or cellular (internal) UV absorbing capacity of the leaves. However, the anthocyanin content was more than doubled by supplemental UV-B radiation. In laboratory experiments, the anthocyanin rich, UV-B treated leaves were less susceptible to a low temperature/high light photoinhibitory treatment, as judged by in vivo chlorophyll fluorescence measurements. Yet, this potential benefit did not considerably affect the growth of the plant in the field (leaf area and dry mass, reproductive dry mass, flowering frequency, senescence rates, dry mass of winter buds). However, there was a marginally significant increase in root dry mass and in the root to shoot ratio, which may underlie the significant increase in the nitrogen content of the leaves. We suggest that P. vulgaris is resistant against UV-B radiation damage and that the possible negative effects of additional UV-B radiation on the growth of these plants may have been effectively counterbalanced by the lower risk of photoinhibition, due to the concomitant increase in anthocyanins.

The nodulation status and nodule morphology of 62 taxa of Leguminosae in a rain forest in French Guiana are reported according to the taxonomy of the family. The N2-fixing species are then fitted into ‘functional groups’ according to their behaviour towards illumination, in order to evaluate their importance in the global dynamics of the stand. The results showed that 67% of the observed species were nodulated (50, 71 and 77% of the Caesalpiniaceae, Mimosaceae and Papilionaceae, respectively). In the Caesalpiniaceae, nodule-like structures were reported in the genus Crudia and in the species Senna quinquangulata, although this needs to be confirmed. All the nodules studied in this subfamily were astragaloid and mucunoid. In the Mimosaceae, the ability of a new genus (Balizia) to form nodules was reported, as well as nodulation on aerial roots in Inga stipularis. The nodules studied were mainly mucunoid. In the Papilionaceae, nodulation on aerial roots in Poecilanthe hostmannii and on conventional roots of the genus Paramachaerium were reported for the first time. All types of nodular structures were found in this subfamily but the structures were quite uniform at the tribal level. These are consistent with suggestions that nodule morphology has a taxonomic value. Eight functional groups of N2-fixing species are proposed, ranging from light dependance to shade tolerance. These results indicate the important role played by N2-fixing species in the global dynamics of the stand and that N inputs by N2 fixation were continuous along the gradient of energetic resources that characterizes the silvigenetic process. The interactions between the photosynthetic capacities of the species and the ability to fix N2 in low light conditions are discussed.

In four field and glasshouse experiments designed to alter the supply of resources through manipulation of nutrients, root tissue, leaf area and fruit number in Alliaria petiolata (Brassicaceae), more than 99% of ovules per plant showed signs of fertilization, suggesting that seed production in this plant was not pollen limited. However, in all treatments a significant proportion of fruits and seeds did not develop to maturity. Total fruit and seed production did not differ significantly from controls when plants were given nutrient supplements at flowering. Removal of 50–75% of the root tissue in 1-yr-old plants significantly reduced fruit set, but had no effect on individual seed development. Removal of cauline leaves significantly reduced most measures of fruit and seed production, suggesting that current photosynthate is critical for fruit and seed filling. Seed maturation was significantly affected by both fruit position within an infructescence and ovule position within a fruit. Basally located fruits and ovules (within fruits) developed more mature seeds than distally positioned fruits and ovules. Plants responded to removal of basal fruits by re-allocating resources to distal fruits that would normally have aborted. Our results suggest that fruits and seeds act as reproductive sinks competing for parental photosynthate. Patterns of resource allocation within infructescences and fruits were also modified by our experiments.

Under ecologically realistic environmental conditions, the water-use efficiency (WUE) of Peperomia scandens, a CAM plant, was higher than that of the C3 congener P. obtusifolia. This difference has been attributed to differences in stomatal activity between C3 and CAM plants, coupled with differences in the evaporative demand of the atmosphere during which the stomata are open. This explanation has apparently not, however, been experimentally tested. Thus, WUEs were compared in these species in two experiments in which the atmospheric evaporative demand was identical (or nearly so) during the period of stomatal opening (i.e. during the night for the CAM plant and during the day for the C3 species). In both experiments, the WUE of the CAM species was higher than that of the C3 species. These results suggest that factors other than differences in atmospheric environmental conditions must also be responsible for the observed differences in WUE. Because CO2 uptake rates of the CAM species were substantially lower than those of the C3 species, the lower WUE in the CAM species resulted primarily from lower transpiration rates. Lower rates of water loss in P. scandens, relative to rates in P. obtusifolia, were ascribed, in part, to lower stomatal densities. Thus, leaf morphological differences, in addition to differences in atmospheric evaporative demand, help to explain the high WUE typically measured in CAM plants.

Resistance to complete submergence was tested in three Rumex species that occur in the Dutch river forelands. The species differ in both habitat and life history characteristics. The annual or biennial R. maritimus and the biennial or short lived perennial R. palustris grow on frequently flooded mud flats of low elevation, while the perennial R. thyrsiflorus can be found on dykes and river dunes that are seldom flooded. The flooding characteristics of the habitats of the three species were determined. These data were used to design experiments to determine the survival and biomass development of the three species during submergence and the influence of plant size and light level on these parameters. It was shown in all three species that plants submerged during daytime were much more resistant to flooding than those submerged at night. This is most probably due to the generation of oxygen or carbohydrates by underwater photosynthesis. Mature plants of the three species showed higher survival after submergence than juvenile plants, which might be caused by higher carbohydrate levels in the taproots of mature plants. In addition, the three species clearly differed in survival and biomass development during submergence. Rumex thyrsiflorus, the species least subjected to flooding, is least tolerant to complete submergence. Rumex maritimus, which can avoid the floods by having a short life cycle, is less tolerant to submergence than R. palustris, which has to survive the floods as a vegetative plant. It was noted that some plants that survived the flooding period itself, still died in the following period of drained conditions, possibly due to post-anoxic injury.

The natural abundance of 13C and 15N was measured in basidiocarps of at least 115 species in 88 genera of ectomycorrhizal, wood-decomposing and litter-decomposing fungi from Japan and Malaysia. The natural abundance of 13C and 15N was also measured in leaves, litter, soil and wood from three different sites. 15N and 13C were enriched in ectomycorrhizal and wood-decomposing fungi, respectively, relative to their substrates. Ectomycorrhizal and wood-decomposing fungi could be distinguished on the basis of their δ13C and δ15N signatures. Although there was high variability in the isotopic composition of fungi, the following isotope- enrichment factors (ε, mean±SD) of the fungi relative to substrates were observed:

εectomycorrhizal fungi/litter = 6.1±0.4‰ 15N

εectomycorrhizal fungi/wood = 1.4±0.8‰ 13C

εwood-decomposing fungi/wood = −0.6±0.7‰ 15N

εwood-decomposing fungi/wood = 3.5±0.9‰ 13C

The basis of isotope fractionation in C metabolism from wood to wood-decomposing fungus is discussed.

Binucleate Rhizoctonia endophytes from the roots of nursery-grown Pinus sylvestris (Scots pine) seedlings and the orchid Goodyera repens from Scots pine forests were characterized on the basis of morphological characters, anastomosis group membership and PCR-assisted ribosomal DNA fingerprinting. Common hyphal and colony morphological traits displayed by the Finnish binucleate Rhizoctonia isolates and a range of Canadian orchid root endophytes enabled them to be placed in the anamorphic genus Ceratorhiza. Five main anastomosis groups were identified and included groups that contained different combinations of Scots pine, G. repens and Canadian Ceratorhiza spp. isolates. Two Scots pine root endophytes anastomosed with a phytopathogenic Japanese tester isolate, confirming their membership of anastomosis group I, which is known to include the teleomorphic species Ceratobasidium cornigerum. Hierarchical cluster analysis of RFLPs in the internal transcribed spacer of ribosomal DNA enabled the division of isolates into one of five RFLP groups. The RFLP and anastomosis groupings were closely correlated; isolates within each of four RFLP groups, which shared 100% RFLP identity, anastomosed in the cross-pairing anastomosis group tests. However, all represented different vegetatively compatible populations (clones) because the diagnostic killing reaction, a cellular vegetative incompatibility response, was identified at hyphal fusion junctions. These findings indicate a high degree of intraspecific variation within Ceratobasidium cornigerum, which includes isolates able to enter into either mutualistic or pathogenic root association with susceptible host plants. The common anastomosis group/RFLP groupings identified also strongly support the hypothesis that conifer tree roots can act as large inoculum reservoirs for these orchid endophytes, allowing the development of inter-plant connections, via commonly shared hyphal linkages, in boreal forest ecosystems.

A cDNA coding for a fungal amino acid transporter (AmAAP1) was identified from Amanita muscaria ectomycorrhizas. The transporter gene was expressed at a basal level under all conditions investigated, but its expression was enhanced 10-fold in the absence of a N source utilized by the fungus. Nitrate was not a suitable N source for A. muscaria and resulted in maximal AmAAP1 expression. The expression of AmAAP1 in a yeast mutant revealed its function as a high-affinity amino acid transporter with a broad substrate spectrum. AmAAP1 takes up all investigated amino acids with Km values between 22 μM for histidine and up to 100 μM for proline. Gene expression and amino acid uptake data together indicate two main functions for AmAAP1: uptake of amino acids from soil for fungal nutrition, and prevention of an amino acid loss by hyphal leakage in the absence of a suitable N source.

This study investigated the relationships between ericoid mycorrhizal endophytes of the Ericaceae (Northern Hemisphere) and the Epacridaceae (Australia). Over 200 fungi were isolated from the roots of two species of Epacridaceae from Victoria, Australia. The isolates were divided into 12 groups by morphology on quarter-strength potato dextrose agar. All were slow-growing and most were dematiaceous, but groups varied from white through pink to dark olive. The ITS1–5.8S–ITS2 ribosomal DNA was amplified and sequenced from eight isolates, forming typical ericoid mycorrhizal morphology in Epacris impressa and one nonmycorrhizal isolate. Sequences were compared, by using similarities and maximum-parsimony analysis, with those of Hymenoscyphus ericae (Leotiales) and Oidiodendron species (Hyphomycetes), the most common endophytes of the Ericaceae. Maximum-parsimony analysis produced four clusters: (1) all Oidiodendron species (at least 90% similarity); (2) all five Victorian dark grey-olive isolates (at least 96% similarity); (3) one Victorian isolate and Cistella grevillei (88% similarity); (4) two light-coloured Victorian isolates and H. ericae (81% similarity). This suggests that these isolates from the Epacridaceae do not belong to the same species as those forming ericoid mycorrhiza in the Ericaceae.