Environmentally induced maternal effects on offspring phenotype are well known in plants. When genotypes or maternal lineages are replicated and raised in different environmental conditions, the phenotype of their offspring often depends on the environment in which the parents developed. However, the degree to which such maternal effects are maintained over subsequent generations has not been documented in many taxa. Here we report the results of a study designed to assess the effects of parental environment on vegetative and reproductive traits, using glasshouse-raised maternal lines sampled from natural populations of Arabidopsis thaliana. Replicates of five highly selfed lines from each of four wild populations were cultivated in two abiotic environments in the glasshouse, and the quality and performance of seeds derived from these two environments were examined over two generations. We found that offspring phenotype was strongly influenced by parental environment, but because the parental environments differed with respect to the time of seed harvest, it was not possible to distinguish clearly between parental environmental effects and the possible (but unlikely) effects of seed age on offspring phenotype. We observed a rapid decline in the expression of ancestral environmental effects, and no main environmental effects on progeny phenotype persisted in the second generation. The mechanism of transmission of environmental effects did not appear to be associated with the quantity or quality of reserves in the seeds, suggesting that environmental effects may be transmitted across subsequent generations via some mechanism that generates environment-specific gene expression.

Rice (Oryza sativa) plants were grown with their roots sandwiched between thin layers of phosphorus-deficient soil from which they were separated by fine mesh, and root-induced changes in the soil affecting phosphate solubility were measured. The concentrations of low molecular weight organic anions in the thin layers, particularly citrate, increased in the presence of the plants. Apparent rates of citrate excretion from the roots, calculated from the quantities in the soil and rates of decomposition calculated with a first order rate constant measured independently, varied from 337–155 nmol g−1 root f. wt h−1 over the course of plant growth, equivalent to 2–3% of plant d. wt. Rates of excretion were similar for NH4+ and NO3−-fed plants. The soil pH decreased from its initial value by up to 0.6 units for the NH4+-fed plants and increased by up to 0.4 units for the NO3−-fed ones. The contribution of organic anion excretion to the pH changes was small compared with that of the inorganic cation-anion balance in the plants. The extent to which the observed excretion of citrate and root-induced pH changes could account for the observed phosphate solubilization and uptake was assessed using a mathematical model. Previous work had shown that phosphate solubilization by rice in this soil could not be explained by enhanced phosphatase activity in the rhizosphere, and the roots were not infected with mycorrhizas. The model allows for the diffusion of the solubilizing agent (citrate or H+) away from the roots, its decomposition by soil microbes (citrate only); its reaction with the soil in solubilizing phosphate and diffusion of the solubilized phosphate to the roots. The model contains no arbitrary assumptions and uses only independently measured parameter values. The agreement between the measured time course of phosphorus uptake and that predicted for solubilization by citrate was good. Root-induced acidification by NH4+-fed plants resulted in additional solubilization, the acidification enhancing the solubilizing effect of citrate. However, the final phosphorus uptake by NH4+-fed plants was no greater than that of NO3−-fed plants, presumably because the acidification inhibited plant growth. The mechanism of solubilization by citrate involved formation of soluble metal-citrate chelates rather than displacement of phosphate from adsorption sites.

Symptoms which are associated with die-back in Phragmites: growth inhibition, root and bud death, premature shoot senescence, blocked aeration and vascular systems, especially in rhizomes and roots, and abnormal surface and internal cell-wall lignification and suberization of roots were induced by each of three of the lower volatile organic acids, propionic, butyric and caproic. These acids were applied in nutrient media in concentrations similar to those previously associated with die-back sites and/or in sediments containing rotting rhizomes and roots of the plant. At concentrations of 1.4 and 0.56 mM, respectively, butyric and caproic acids were each found to be highly toxic at pH 4.5, but relatively innocuous at pH 6. Propionic acid, applied at a much higher concentration of 10.4 mM, was highly toxic at both pH 4.5 and 6. The results support previous findings that the undissociated forms of the organic acids are the more toxic. Rhizomes and roots, rotting in water or waterlogged sand, released cocktails of acids and produced pH in the range 4.8–5.4. Phragmites seedlings planted in these media died within 12 h. Overall, the results support the theory that die-back in Phragmites can be induced and/or perpetuated by organic acids released from the decaying underground parts of the plant or other sources of organic matter.

Seeds of the wetland legume, Lotus uliginosus, were germinated and grown in vermiculite which was either continuously flooded or well-drained. Plants from both treatments were infected by Mesorhizobium loti strain DUS341 via a ‘classical’ root hair pathway, although some flooded plants appeared to be infected via enlarged epidermal cells. Subsequent to infection by M. loti, nodule meristems, which had developed within the root outer cortex, were penetrated by infection threads that released bacteria into the meristematic cells. The infection threads and infection droplets were immunogold labelled with monoclonal antibodies (MAC265 and MAC236) that recognize epitopes (at approx. 155/170 and 170/210 kDa, respectively) on a glycoprotein component of the matrix that surrounded the bacteria within the threads or droplets. Although labelling of infection threads or infection droplets with MAC236 was stronger than that with MAC265, both antibodies strongly labelled material occluding intercellular spaces in the cortices of developing nodules that had not yet expressed nitrogenase (as determined by a lack of signal after immunogold labelling with an antibody raised against nitrogenase component II). After 60 d, nitrogenase activity, shoot and root dry weights, and nodule fresh weight per plant did not differ between the treatments. After a further 30 d submergence, the flooded stems developed extensive aerenchyma and there was profuse development of (nodulated) adventitious roots. Nodules also formed at the junction of adventitious roots and the subtending stem and these were connected vascularly to a small stalk of tissue which gave rise to both a nodule and an adventitious root. The flooded nodules had prominent lenticels, and possible air pathways from the atmosphere to the nitrogen-fixing bacteroids are discussed.

We previously found that the ethylene inhibitor Ag+ could overcome the inhibitory effect of nitrate on nodulation of soybean (Glycine max) cv. Bragg. The same treatment increased nodulation quantitatively under non-inhibitory conditions, strongly suggesting involvement of ethylene in the control of nodulation in this species. Supernodulation mutants that lack internal autoregulation of nodulation, however, had biosynthesis capacity similar to the wild type. In the present work, the effects of ethylene on nodulation of ‘Bragg’ and two separate, but allelic, supernodulating mutants (nts382 and nts1007) were compared. The nodulation process appeared much more sensitive than plant growth and development to ethylene, which reduced the number of nodules per plant, but nearly twofold more in the wild type than in the supernodulation mutants. The cause–effect relationship is established by the counteracting effect of Ag+ and the fact that the stronger the inhibition by ethylene, the higher the recovery of nodulation ability with the ethylene antagonist. This higher tolerance of or lower sensitivity to ethylene in nts382 persists even under low inoculum dose, where nodule number and mass could be decreased to wild-type levels. Differences between the mutant and the wild type in the triple response test do not appear to support differences in ethylene perception on a whole-plant basis. The results suggest that sensitivity of nodulation to ethylene might have been affected in supernodulation mutants.

Mature heather (Calluna vulgaris) and bracken (Pteridium aquilinum) turfs, transplanted from the field, were subjected to factorially combined experimental treatments for three consecutive years. Summer drought had the greatest effect, decreasing photosynthesis, growth and reproductive output in both species, and opening the bracken canopy. The timing of the drought relative to plant development was critical to which species was worst affected; bracken was worst affected by an early drought, heather by a later drought. Both species showed physiological damage during drought but, as predicted on the basis of their functional types, heather showed greater acclimation of water-use efficiency to drought stress. Contrary to expectations based on functional types, heather responded more rapidly than bracken to increased nitrogen supply (50 kg N ha−1 yr−1). Added nitrogen caused both species to start above-ground growth earlier in the spring. For bracken this stimulation was short- lived; added nitrogen might be preferentially allocated to the rhizome and the longer-term consequences of this are unknown. For heather, nitrogen promoted growth and flowering throughout the season. There was no positive effect on the photosynthetic physiology of either species; changes in resource partitioning, and thus photosynthate production at the canopy level, are the most likely mechanism for the increase in heather shoot growth. Warmer temperatures increased heather shoot growth from early spring onwards but did not advance bracken crozier emergence, although frond height and the proportion of fertile fronds were subsequently increased. No significant effects of warming on the photosynthetic physiology of either species were found. Predictions of responses of heather and bracken to environmental change are complicated by the strong interactive effects of unpredictable climatic events such as drought and extreme winter temperatures. When drought was imposed, damage to heather was much greater in plants receiving increased nitrogen supply. Stimulation of growth by nitrogen resulted in a water demand that was unsustainable in drought conditions, leading to wilting, reduced shoot growth and some acclimation of water-use efficiency. Additionally, a very cold winter spell proved most damaging to heather that had been droughted in the previous summer. For bracken, winter damage occurred in plants that had been warmed, with significantly fewer fronds emerging in the next spring and thus canopy photosynthetic potential being reduced. We predict that positively managed heather has the potential to limit the bracken problem in conditions of environmental change, provided that high levels of nitrogen deposition do not coincide with increased drought frequency.

We tested a hypothesis that elevated ozone was an eliciting or contributing factor in outbreaks of the ‘top dying’ (or ‘subtop dying’) syndrome in Norway spruce (Picea abies). Progeny were used from open-pollinated trees within a stand with the ‘top dying’ syndrome. The mother trees were classified in relation to the expression of the ‘top dying’ syndrome, and progeny from the healthiest and least healthy thirds of the population were exposed to high and low concentrations of ozone for three seasons. Elevated ozone did not affect height growth of the trees. It did not measurably affect net photosynthesis, stomatal conductance or instantaneous water use efficiency. Chlorophyll and carotenoid contents were also not significantly affected by ozone concentration. In the first year, instantaneous water use efficiency was lower in the progeny of the unhealthy mother trees than in the healthy mother trees. Furthermore, the unhealthy mother trees tended to produce longer annual shoots and showed more winter damage at the end of the experiment. None of these parameters were related to ozone concentration in the atmosphere. These results do not support a hypothesis that elevated ozone is a significant contributory factor or an eliciting factor in the development of the syndrome.

A range of plant species typical of semi-natural grasslands were tested for their sensitivity to short-term ozone injury under normal and reduced irrigation, and in relationship to air vapour pressure deficit. Potted specimens of 24 herbs, legumes and grasses were exposed during two seasons to four O3 treatments in open-top chambers. The ozone treatments were: (a) charcoal-filtered air; (b) charcoal-filtered air plus ozone to match ambient levels; (c) charcoal-filtered air plus O3 to ambient levels 1.5 and (d) charcoal-filtered air with ozone added to twice ambient levels during selected episodes of 7–13 d. During these ozone episodes, half of the plants in each ozone treatment received reduced irrigation (dry treatment) while the rest was kept under full irrigation (wet treatment). Type and date of first occurrence of leaf injury were noted during individual growth periods. Plants were harvested three times per year, and the percentage of injured leaves was recorded. Depending on species, injury symptoms were expressed as flecking (O3-specific injury), leaf yellowing or anthocyanin formation. Carum carvi and most species of the Fabaceae family (Onobrychis sativa, Trifolium repens, Trifolium pratense) were found to be most responsive to O3, injury occurring after only a few days of exposure in treatment (b). An episodic reduction in irrigation tended to reduce the expression of O3-specific symptoms, but only in species for which a reduction in soil moisture potential and an associated reduction in stomatal conductance during the dry episodes were observed. In other species, the protection from O3 injury seemed to be of little importance. Using artificial neural networks the injury response of nine species was analysed in relation to Species, stomatal conductance, ozone as AOT40 (accumulated exposure above a threshold of 0.04 ppm for periods with global radiation [ges ]50 W m−2 (Fuhrer et al., 1997)), mean relative growth rate, air vapour pressure deficit and global radiation. In the model with all factors, Species was most important, and when Species was omitted, stomatal conductance was the most important determinant for leaf injury to occur, whereas mean relative growth rate was less important. With no plant-related factors included, air vapour pressure deficit and AOT40 were of highest importance. Only in eight species was a positive relationship found between these two factors during the five days before the onset of injury, indicating increasing protection from ozone with increasing air vapour pressure deficit in some but not all species. These data show that across a range of grassland species, leaf injury caused by elevated levels of ozone is most likely to occur in species with high stomatal conductance and that protection from ozone during dry periods is species-specific and depends on a reduction in stomatal conductance due to a decrease in soil moisture potential. Protection under increased vapour pressure deficit can occur in some but not all species, depending on the relationship between stomatal conductance and air vapour pressure deficit.

Plant species typical of semi-natural grasslands were tested for their sensitivity to long-term ozone effects on growth under two irrigation regimes, and the relationship between ozone sensitivity, relative growth rate, and plant strategy was analysed. Individuals of 24 grasses, herbs and legumes were grown in pots and exposed for two seasons to four levels of ozone in open-top chambers: (a) CF, charcoal-filtered air, (b) CFAA, charcoal-filtered air plus ozone to match ambient levels, (c) CF+, charcoal-filtered air plus ozone added to match 1.5 times ambient levels and (d) CF++, charcoal-filtered air with ozone added to match twice ambient levels during selected episodes of 7–13 d. During the ozone episodes, one half of the plants received reduced irrigation (dry treatment) while the rest was kept well watered (wet treatment). The effect of ozone on the relative growth rate was determined during the initial growth period during the first season. Above-ground plant dry weight was determined three times during each season, and stubble dry weight at the end of the experiment. In most species, the effect of ozone on relative growth rate was not significant, except in Centaurea jacea and Lychnis flos-cuculi for which a significant stimulation of relative growth rate was found. There was a significant negative relationship between percentage relative growth rate and relative growth rate measured in the CF treatment, indicating that in species growing faster in clean air relative growth rate tended to be more sensitive to ozone than in slower-growing species. After the first re-growth period during the first season, a significant effect of ozone on above-ground biomass was found in five species, i.e. a decrease in Chrysanthemum leucanthemum, Trifolium pratense and T. repens, and an increase in Silene dioïca. During the second season no significant ozone effect was detected, except for an increase in stubble mass in C. leucanthemum. In 10 species, the dry treatment caused a significant decrease in biomass, but in Plantago lanceolata during the first year, and in Knautia arvensis and T. repens during the second year, an ozone×irrigation interaction was observed. Exposure–response patterns in the wet treatment varied between species. In some species growth tended to be increased in charcoal-filtered air plus ozone to ambient levels, in others either a continuous increase or decrease in biomass was observed with increasing ozone. Trifolium repens showed the lowest AOT40 (ozone accumulated above a threshold of 40 ppb) corresponding to a 10% change in biomass. The comparison between exposure–response pattern between the wet and dry treatments revealed reduced, increased, or unchanged ozone sensitivity. The shift was related to plant strategy and not to the specific sensitivity to the dry treatment. The specific response to both ozone and dry treatment tended to be related to Grime's triangular classification of plant strategies, i.e. the C-S-R model. In species with a large component of C (competitor strategy), and/or R (ruderal strategy), biomass was more sensitive to both stresses, compared with species with a high component of S (stress tolerators), and ozone×drought interactions were more pronounced. The results indicate that specific relative growth rate is an important determinant for the potential sensitivity of grassland species to long-term effects of ozone, and that reduced irrigation causes no general protection from the effects of ozone on growth.

Simulated swards of perennial ryegrass (Lolium perenne) growing in 1-m3 soil blocks in the glasshouse were either well watered or deprived of water for 57 d and then rewatered. The first aim was to measure effects of drought on sugar (water-soluble carbohydrate) composition of laminae and sheaths of mature laminae, and bases and laminae of young (growing) leaves. The second aim was to use pulse labelling with 14CO2 to follow the partitioning of recently-fixed assimilates, and the assembly and consumption of reserve sugars (fructans). Over the last 7 d of drought growth almost stopped, old leaves died faster than they were replaced, and total sugar (which had doubled in concentration during drought) was rapidly consumed. Leaf laminae had lower content of total sugars and of large fructan (DP>5) than did growing bases and sheaths. Drought greatly reduced the rate at which sugar was exported from the laminae to the sheaths and growing leaf bases, and the rate at which it was converted to fructan. Nevertheless, fructan accumulated over the first 50 d of drought. Rewatering did not result in depolymerization and remobilization of sugars that had been formed during the last 7 d of drought, but stimulated their further assembly into high-DP fructans. Our hypothesis, that accumulation of neo-kestose (a DP-3 fructan) in droughted laminae was a symptom of sugar remobilization just before death, was disproved. It is concluded that sugar reserves contribute to drought resistance only under extreme conditions. The specific role of fructan in dry environments might be to improve regrowth when drought is relieved, rather than to enhance growth during drought.

A low auxin concentration (0.1% naphthalene acetic acid) induced tracheids with longitudinal polarity parallel to the hypocotyl axis in young Pinus pinea seedlings. Application of 0.1% ethrel laterally and 0.1% naphthalene acetic acid apically disturbed axial tracheid polarity and promoted the differentiation of tracheids with a lateral orientation. Ethrel by itself, with no auxin background, did not affect tracheid differentiation. Apical application of 1% gibberellic acid with the low auxin, reversed the polarity disorder induced by ethrel. Disturbance of axial tracheid polarity was observed under a high auxin concentration (0.5% naphthalene acetic acid) which was similar to the combined effect of ethrel and auxin. The high auxin concentration increased tracheid number significantly. This effect was curtailed following treatment with inhibitors of ethylene formation (Co2+; 1-aminoethoxy- vinylglycine) and action (Ag2+). The role of ethylene in controlling the differentiation of radial tracheids, which characterize the vascular rays of pines, is discussed.

Although mat-forming cyanobacteria dominate many freshwater ecosystems in the Arctic and Antarctic, their optimal temperature for growth (Topt) is usually much higher than the temperature range of their native habitat. The present study compared the temperature dependence of growth, pigment composition and absorbance, photosynthesis and photosynthate partitioning for two strains of cyanobacteria with contrasting Topt values; Phormidium subfuscum, isolated from McMurdo Ice Shelf, Antarctica, and Phormidium tenue, collected from the Kuparuk River in the tundra region of northern Alaska. Phormidium subfuscum grew between 5 and 20°C with a Topt of 15°C whereas P. tenue showed detectable growth from 10 to 40°C and a Topt of 30°C. Light utilization efficiency, photosynthetic capacity and the irradiance at the onset of light saturation increased with increasing temperature up to Topt in both strains. The cellular concentrations of chlorophyll a (Chl a) and carotenoid (CAR) and the in vivo absorbance maxima for Chl a, CAR, C-phycocyanin and allophycocyanin changed little for P. subfuscum but all these variables increased across the temperature range up to Topt for P. tenue. Neither P. subfuscum nor P. tenue showed changes in relative carbon allocation with varying temperature, suggesting that gross biochemical alterations are not a characteristic of temperature acclimation in these cyanobacteria. We conclude that the eurythermal cyanobacterium P. tenue optimizes growth over a wide range of temperatures by adjusting its light-capturing as well as carbon fixation characteristics, whereas stenothermal P. subfuscum relies on changes in carbon fixation without concomitant shifts in pigment content.

Mycelial cord systems, up to 50 cm in diameter, of the basidiomycete Phanerochaete velutina, a common woodland saprotroph, were grown on unsterile soil in model laboratory microcosms from either 4 or 16 cm3 wood inocula. After 48 d, systems were supplied at 10-d intervals with pairs of new 4 cm3 beechwood resources placed behind the foraging colony margin, where possible on opposite sides of the system. Image analysis was used to quantify radial extension, hyphal cover, the mass (DBM) and surface fractal (DBS) dimensions of the mycelial systems and wood-resource bleaching activity. Mycelial systems developing from small inocula had significantly (P[les ]0.05) lower radial extension rates, hyphal cover and DBM[ratio ]DBS ratio than those from large inocula. Initially, systems developing from small inocula also displayed significantly (P<0.05) slower wood-resource bleaching activity than those from large inocula, suggesting that carbon limitation affected both foraging behaviour and resource utilization. A separate central compartment containing the inoculum was supplied with 32P orthophosphate and its partitioning amongst wood resources was monitored nondestructively for 44 d. Total 32P acquisition by wood resources was not significantly (P>0.05) affected by inoculum resource size. However, the proportion of total acquired P allocated to resources, which varied according to the length of time that resources had been in contact with the mycelium, was dependent upon inoculum size. The results support the hypothesis that phosphorus translocation is not a demand-driven process. We suggest that the time taken before the greatest rate of phosphorus withdrawal from a uniform translocation stream is dependent upon prior availability of carbon within colonized resources.

Fungal endophytes associated with the palm, Trachycarpus fortunei, within and outside its natural geographic range were investigated. Endophytes were relatively common with colonization rates of 23–57% at the four sites sampled. The endophyte assemblages at the different sites were diverse with 75 fertile species of ascomycetes and mitosporic fungi. The assemblage composition at each site was similar and between seven and 13 species comprised 81–89% of the taxa present in relative frequencies of >1%. Glomerella cingulata and Phomopsis spp., were consistently dominant, and a large number of rare species were recorded. The diversity at each site was similar in number, but the abundance of isolates varied. The results obtained were comparable to those of previous studies of palm endophyte assemblages, but the assemblages showed more affinity with unrelated temperate hosts than with tropical palm hosts. Quantitative and qualitative differences in endophyte assemblages from old and young tissues were observed, and more isolates were recovered from old tissues independent of the age of the palm. The composition of the assemblage varied with several taxa being exclusively or more commonly isolated from old tissues (e.g. xylariaceous taxa, Oxydothis sp. nov.) or young tissues (e.g. Stagonospora spp., Phoma multirostrata). Some differences in the composition of the assemblage and in relative frequencies of various species were observed in trees and saplings. Significantly more isolates were recovered from the vein than intervein tissues, independent of leaf age or tree age. Tissue specificity was not exhibited by any taxa isolated from either leaf or petiole tissues, except for xylariaceous taxa in leaf tissues. Some other taxa showed a preference for leaf tissues or petioles, whereas others were equally distributed amongst all tissues. Endophyte assemblages of palms from continuous distributions were similar, but those from disjunct distributions (i.e. outside the natural geographic range of the palm, such as Australia and Switzerland), differed significantly from each other and from assemblages within continuous distributions. The relative importance values of endophyte species at the two sites in China indicates the site-specific nature of the endophyte assemblages. Most previous studies on the endophytes of palm hosts have concentrated on tropical palms. However, this study examines the endophytes of the warm temperate palm Trachycarpus fortunei, and investigates the relative importance of host and climate related processes.

Genomic DNA from spores of the endomycorrhizal fungus Gigaspora margarita was amplified with two sets of degenerate primers designed on conserved regions of chitin synthase polypeptides. Three chitin synthase (chs) gene portions were identified: Gimchs1, Gimchs2 and Gimchs3, encoding one class II and two class IV chitin synthases, respectively. Gimchs1 and Gimchs3 transcripts were detected by reverse transcription polymerase chain reaction (RT-PCR) during root colonization, whereas none of three mRNAs was revealed during spore germination. The potential role of the genes expressed is discussed in the light of morphogenetic and signalling events.

The morphology and anatomy of ectomycorrhizas of Rhizopogon arctostaphyli, R. ellenae, R. flavofibrillosus, R. occidentalis, R. rubescens, R. smithii, R. subcaerulescens and R. truncatus synthesized on Ponderosa pine (Pinus ponderosa) in glasshouse conditions using spore slurries, are described and compared. All species produced a well-developed Hartig net, and a well-developed fungal mantle. The mantles of R. arctostaphyli, R. smithii and R. subcaerulescens ectomycorrhizas were two-layered with outer mantle hyphae of wider diameter than inner mantle hyphae. The mantle of R. subcaerulescens ectomycorrhizas also had distinctive peg-like structures (cystidia) along peripheral hyphae. Rhizopogon truncatus ectomycorrhizas were tuberculate in morphology and had a rind-like mantle enclosing adjacent roots. In addition, several species exhibited crystal inclusions in the outer mantle, presumably at the interface between mantle and soil.

In the December 1998 issue of New Phytologist, we published the research paper entitled ‘Natural occurrence of Ampelomyces intracellular mycoparasites in mycelia of powdery mildew fungi’ by Levente Kiss (New Phytol. (1998) 140, 709–714). Since its publication, an error in the published title has been brought to our attention: this refers to the ampelomyces, but the name should be Ampelomyces, meaning the genus.

We apologise to the author and to our readers for this mistake.