The present study of structural and physiological changes during the development of the cushion moss, Grimmia pulvinata, quantifies the size-dependence of various parameters of water relations such as changes in surface: volume ratio (S/V) or water loss rates, and also measures net CO2 gas exchange in the light and the dark. Larger cushions had lower S/V values than smaller ones and featured lower rates of area-based evapotranspiration, owing to higher boundary-layer resistance, but did not differ in relative water storage capacity (expressed as a percentage of d. wt). In combination, this leads to considerably longer hydration periods in larger cushions. By contrast, CO2 gas-exchange parameters were negatively correlated with size : larger cushions showed significantly lower (mass-based) rates of net photosynthesis and dark respiration. Using these data, we estimated carbon budgets during a drying cycle as a function of cushion size. When including alternations of dark and light periods, the relationship proved to be rather complicated. Depending on the time of hydration, net carbon budgets not only varied quantitatively with size but sometimes took on both positive and negative values depending on cushion size. We conclude that neglecting plant size can lead to unrepeatable or even misleading results in comparative ecophysiological studies, and therefore urge for adequate attention to be paid to size in these studies.

The presence of copper-binding proteins produced in response to added copper was examined in isolates of Laccaria laccata (Scop. ex Fr.) Cooke and Paxillus involutus (Batsch ex Fr.) Fr. taken from copper-contaminated and uncontaminated sites, and in a single isolate of Scleroderma citrinum Pers. from a contaminated site. Two isolates of Laccaria (GLac4 and ELac1) grew better in 1·5 mM and 2·5 mM copper than a third (Lac3G) and were considered to be more tolerant. Amongst five isolates of P. involutus, three (WJPax2R, GPaxRSp2 and Pax4) were capable of growth in media containing 4·0 mM copper and were regarded as tolerant. All isolates of both Laccaria and Paxillus were capable of some growth in 2·5 mM copper, but S. citrinum was much more copper-sensitive and the concentration had to be reduced at least 10-fold before any growth occurred. Tolerance of isolates was not related to whether they were taken from copper-contaminated or uncontaminated sites. Copper-binding proteins were detected in response to copper in the culture media in the two tolerant isolates of Laccaria (GLac4 and ELac1) but not in the least tolerant isolate. In Paxillus, similar proteins were found in two tolerant isolates (GPaxRSp2 and Pax4) but not in WJPax2R, which was also regarded as tolerant, nor in any of the less tolerant isolates. Copper-binding proteins were not detected in S. citrinum. The copper-binding protein purified from the Laccaria isolate ELac1 appeared as a single band in modified SDS–PAGE electrophoresis. Its molecular mass and spectral characteristics were consistent with it being a metallothionein.

Young trees of European beech (Fagus sylvatica) acclimated for one growing season to ambient (c. 367 μl l−1) or elevated CO2 levels (c. 660 μl l−1) were exposed during the subsequent year to combinations of the same CO2 regimes and ambient or twice-ambient ozone (O3) levels (generated from the database of a rural site). By the end of June, before the development of macroscopic leaf injury, the raised O3 levels had not affected the light and dark reactions of photosynthesis. However, acclimation to elevated CO2 had resulted in lowered chlorophyll and nitrogen concentrations, whereas photosynthetic performance, examined over a wide range of parameters from light and dark reactions, remained unchanged or showed only slight reductions (e.g. apparent electron transport rate, ETR; apparent quantum yield of CO2 gas exchange, ΦCO2; apparent carboxylation efficiency, CE; and photosynthetic capacity at light and CO2 saturation, PC). In August, after the appearance of leaf necroses, plants grown under ambient CO2 and twice-ambient O3 conditions declined in both the photosynthetic light reactions (optimum electron quantum yield, Fv/Fm, non-photochemical energy quenching, NPQ, reduction state of QA, apparent electron quantum yield, ΦPSII, maximum electron transport rates) and the dark reactions as reflected by CE, ΦCO2, as well as the maximum CO2 uptake rate (i.e. PC). CE, ΦCO2 and PC were reduced by c. 75, 40 and 75%, respectively, relative to plants exposed to ambient CO2 and O3 levels. By contrast, plants exposed to twice-ambient O3 and elevated CO2 levels maintained a photosynthetic performance similar to individuals grown either under ambient CO2 and ambient O3, or elevated CO2 and ambient O3 conditions. The long-term exposure to elevated CO2 therefore tended to counteract adverse chronic effects of enhanced O3 levels on photosynthesis. Possible reasons for this compensatory effect in F. sylvatica are discussed.

Water relations, xylem embolism, root and shoot hydraulic conductance of both young plants in the field and potted seedlings of Quercus pubescens have been studied with the aim of investigating whether these variables may account for the well known adaptation of this oak species to arid habitats. Our data revealed that Q. pubescens is able to maintain high leaf relative water contents under water stress conditions. In fact, relative water contents measured in summer (July) did not differ from those recorded in April. This was apparently achieved by compensating water loss by an equal amount of water uptake. Such a drought avoidance strategy was made possible by the recorded high hydraulic efficiency of stems and roots under water stress. In fact, root hydraulic conductance of field-grown plants was maintained high in summer when the percentage loss of hydraulic conductance of stems was lowest. The hydraulic architecture of young plants of Q. pubescens measured in terms of partitioning of hydraulic resistances along the water pathway revealed that the highest hydraulic resistance was located in stems of the current year's growth. This hydraulic architecture is interpreted as consistent with the adaptation of Q. pubescens to arid habitats as a consequence of the recorded seasonal changes in water relation parameters as well as in root and stem hydraulics.

Most work on root proliferation to a localized nutrient supply has ignored the possible role of mycorrhizal fungi, despite their key role in nutrient acquisition. Interactions between roots of Plantago lanceolata, an added arbuscular mycorrhiza (AM) inoculum and nitrogen capture from an organic patch (Lolium perenne shoot material) dual-labelled with 15N and 13C were investigated, to determine whether root proliferation and nitrogen (N) capture was affected by the presence of AM fungi. Decomposition of the organic patch in the presence and absence of roots peaked in all treatments at day 3, as shown by the amounts of 13CO2 detected in the soil atmosphere. Plant N concentrations were higher in the treatments with added inoculum 10 d after patch addition, but thereafter did not differ among treatments. Plant phosphorus concentrations at the end of the experiment were depressed by the addition of the organic residue in the absence of mycorrhizal inoculum. Although uninoculated plants were also colonized by mycorrhizal fungi, colonization was enhanced at all times by the added inoculum. Addition of the AM inoculum increased root production, observed in situ by the use of minirhizotron tubes, most pronouncedly within the organic patch zone. Patch N capture by the end of the experiment was c. 7.5% and was not significantly different as a result of adding an AM inoculum. Furthermore, no 13C enrichments were detected in the plant material in any of the treatments showing that intact organic compounds were not taken up. Thus, although the added AM fungal inoculum benefited P. lanceolata seedlings in terms of P concentrations of tissues it did not increase total N capture or affect the form in which N was captured by P. lanceolata roots.

Cherry seedlings (Prunus avium) were grown from seed for two growing seasons in three ambient [CO2] (∼350 μmol mol−1) and three elevated [CO2] (ambient+∼350 μmol mol−1) open-top chambers, and in three outside blocks. A drying cycle was imposed in both the growing seasons to half the seedlings: days 69–115 in the first growing season, and in the second growing season days 212–251 on the same seedlings which had already experienced drought. Stomatal conductance was significantly reduced in elevated [CO2]-grown, unstressed seedlings in both the first and second growing seasons, but was not caused by a decrease in stomatal density. Droughted seedlings showed little or no reduction in stomatal conductance in response to elevated [CO2]. However, stomatal conductance was highly correlated with soil water status. Photosynthetic rate increased significantly in response to elevated [CO2] in both water regimes, leading to improvement in instantaneous transpiration efficiency over the whole duration of the experiment, but there was no relationship between instantaneous transpiration efficiency and long-term water use efficiency. The Amax was strongly reduced in the second growing season, but unaffected by [CO2] treatment. Although photosynthetic rate was not down-regulated, Rubisco activity was decreased by elevated [CO2], possibly because of the increased leaf carbon: nitrogen ratio which had occurred by the ends of the two growing seasons. Elevated [CO2] did not improve plant water relations (for example, bulk leaf – water potential, osmotic potentials at full and zero turgor, relative water content at zero turgor, bulk modulus of elasticity of the cell) and thus did not increase water-stress tolerance of cherry seedlings.

To investigate the extent and size of root–soil air gaps that develop during soil drying, resin casts of roots of the desert succulent Agave deserti Engelm. were made in situ for container-grown plants and in the field. Plants that were droughted in containers for 7 and 14 d had 24 and 34% root shrinkage, respectively, leading to root–soil air gaps that would reduce the hydraulic conductivity at the root–soil interface by a factor of about 5. When containers were vibrated during drought, root–soil air gaps were greatly diminished, and the predicted conductivity at the interface was similar to that of the control (moist soil). For plants in the field (4 and 6 wk after the last rainfall), root shrinkage was greater than for container-grown plants, but root–soil contact on the root periphery was greater, which led to a higher predicted hydraulic conductivity at the root–soil interface. To test the hypothesis that root–soil air gaps would help to limit water efflux from roots in drying soil, the water potentials of the soil, root, and shoot of plants from vibrated containers (with gaps eliminated or reduced) and non-vibrated containers were compared. The soil water potential was lower for vibrated containers after 14 d of drought, suggesting more rapid depletion of soil water due to better root–soil contact, and the root water potential was lower as well, suggesting greater water loss by roots in the absence of root–soil air gaps. Thus, air gaps could benefit A. deserti by helping to maintain a higher root water potential in the early stages of drought and later by limiting root water loss at the root–soil interface when the water potential exceeds that of the soil.

We investigated the effect of elevated CO2 on the growth and mycorrhizal colonization of three tree species native to north-eastern American forests (Betula papyrifera Marsh., Pinus strobus L. and Tsuga canadensis L. Carr). Saplings of the tree species were collected from Harvard Forest, Massachusetts, and grown in forest soil under ambient (c. 375 ppm) and elevated (700 ppm) atmospheric CO2 concentrations for 27–35 wk.

In all three species there was a trend to increasing whole-plant, total-root and fine-root biomass in elevated CO2, and a significant increase in the degree of ectomycorrhizal colonization in B. papyrifera and P. strobus, but not in T. canadensis. However, in T. canadensis the degree of colonization with arbuscular mycorrhizas increased significantly. In both the ambient and elevated environments, on the roots of B. papyrifera and P. strobus 12 distinct ectomycorrhizal morphotypes were identified. Distinct changes in the ectomycorrhizal morphotype assemblage of B. papyrifera were observed under CO2 enrichment. This change resulted in an increase in the frequency of ectomycorrhizas with a higher incidence of emanating hyphae and rhizomorphs, and resulted in a higher density of fungal hyphae in a root exclusion chamber.

Sclerophylly and synthesis of phenolic compounds are active responses of plants subjected to environmental stress (drought, low nutrient supply, u.v.-B radiation, ozone). Here we describe the morphological and histochemical alterations occurring in field-grown leaves of Fagus sylvatica L. from three sites located along an ecological gradient: from a site in cool and protected conditions to one located on a mountain ridge, where the trees grow on a thin layer of soil and are exposed to the wind and to intense solar radiation in summer. The morphological data show that, as the ecological conditions of the stand worsen, individual leaf surface decreases, while the thickness of the leaves and their specific d. wt (i.e. d. wt per unit leaf area) increases. Histochemical and ultrastructural tests show a marked increase of phenolics during the course of the year. These substances, present primarily in the leaves of trees growing in stress conditions, have been identified mainly as tannins. They accumulate in the vacuoles, especially those of the upper epidermal layer and the palisade mesophyll; at a later stage they appear to be solubilized in the cytoplasm and retranslocated, eventually impregnating the outer wall of the epidermal cells amidst the cellulose fibrils, where they cluster together and form an electron-opaque layer between the wall and the cuticle. Observation of the epidermal cells also reveals that the outer cell wall is thicker. The paper discusses the roles of secondary metabolites in protection and detoxification processes; the possible ecological significance of these alterations in the ecophysiology of beech trees.

Interspecific C transfer was studied in laboratory microcosms containing pairs of 6-month-old Betula papyrifera Marsh. and Pseudotsuga menziesii (Mirb.) Franco seedlings growing in individual, root-restrictive (28 μm pore size) pouches filled with field soil. Interspecific transfer was examined by reciprocal labelling of seedlings with 13CO2(gas) and 14CO2(gas). At the time of labelling, the root zones of ectomycorrhizal (EM) B. papyrifera and P. menziesii were interconnected by an extensive network of EM mycelium. Carbon transferred through EM connections was distinguished from that through soil pathways by comparing microcosms where interconnecting hyphae were left intact vs. those where they were severed immediately before labelling.

Transfer was bidirectional, and represented 5% of total isotope uptake by both B. papyrifera and P. menziesii together. P. menziesii received on average 50% more 14C and 66% more 13C from paper birch than vice versa, however, differences between species were not statistically significant. Neither net nor bidirectional transfer differed between severing treatments, leaving in question the relative importance of EM hyphae versus soil transfer pathways. The tendency for P. menziesii to receive more isotope than B. papyrifera corresponded with a 10-fold greater net photosynthetic rate per seedling and two-fold greater foliar N concentration of B. papyrifera than P. menziesii.

The host range of the soilborne obligate biotroph, Plasmodiophora brassicae was investigated. Evidence is presented that infection by P. brassicae might occur in non-Brassica species, leading to the potential formation of resting spores. Structures resembling P. brassicae were found in the root cortex of Tropaeolum majus, Carica papaya, Reseda alba and Beta vulgaris as demonstrated by scanning electron microscopy. Inoculation of Brassica rapa roots with spores extracted from either T. majus or B. vulgaris roots which had been previously inoculated with P. brassicae led to development of clubroot in the roots of B. rapa. It was also shown that the development of the symptom might be correlated with glucosinolate content, although other host factors are implicated in the B. vulgaris interaction with P. brassicae. In the glucosinolate-containing non-Brassicas, T. majus and C. papaya, the concentrations of benzylglucosinolate increased markedly in roots inoculated with P. brassicae, compared with the controls. There were also increases in concentrations of benzylglucosinolate in leaves of T. majus after P. brassicae infection. However, in R. alba roots, the total glucosinolate content decreased after inoculation with P. brassicae compared with the controls. High root concentrations of 2-OH-2-phenylethylglucosinolate (glucobarbarin) compared with low root indole glucosinolates in this species might limit P. brassicae infection and development. The importance of our investigations in relation to cultivation of non-Brassica species on fields infested with P. brassicae is discussed.

A field and laboratory based bioassay has been developed to investigate the effects of the quantity and duration of simulated pollutant nitrogen (N) deposition on root-surface phosphomonoesterase (PME) activities in calcareous and acid grasslands. Seedlings of Plantago lanceolata were transplanted to a calcareous grassland and Agrostis capillaris seedlings were grown in microcosms containing soil from an acid grassland that had received either 7 yr (long-term) N additions or 18 months (short-term) N and phosphorus (P) additions. The bioassay revealed that short-term N treatments had little effect on the enzyme activity, whereas long-term N additions significantly increased PME activity within 7 d of transplanting into the field plots. Root-surface PME activity of A. capillaris was significantly reduced in soil that received additions of P. In the plots receiving long-term additions of N, a strong relationship was observed between extractable soil ammonium and root-surface PME activity. Soil ammonium concentrations accounted for 67% of the variation in PME activity of P. lanceolata in the calcareous grassland, and 86% of the variation in PME activity of A. capillaris in the acid grassland. These results provide evidence that N deposition may have considerable effects on the demand and turnover of P in ecosystems that are approaching or have reached N saturation.

Short-term phosphate uptake rates were measured on intact ectomycorrhizal and non-mycorrhizal Pinus sylvestris seedlings using a new, non-destructive method. Uptake was quantified in semihydroponics from the depletion of Pi in a nutrient solution percolating through plant containers. Plants were grown for 1 or 2 months after inoculation at a low relative nutrient addition rate of 3% d−1 and under P limitation. Four ectomycorrhizal fungi were studied: Paxillus involutus, Suillus luteus, Suillus bovinus and Thelephora terrestris. The Pi-uptake capacity of mycorrhizal plants increased sharply in the month after inoculation. The increase was dependent on the development of the mycobionts. A positive correlation was found between the Pi-uptake rates of the seedlings and the active fungal biomass in the substrate as measured by the ergosterol assay. The highest Pi-uptake rates were found in seedlings associated with fungi producing abundant external mycelia. At an external Pi concentration of 10 μM, mycorrhizal seedlings reached uptake rates that were 2.5 (T. terrestris) to 8.7 (P. involutus) times higher than those of non-mycorrhizal plants. The increased uptake rates did not result in an increased transfer of nutrients to the plant tissues. Nutrient depletion was ultimately similar between mycorrhizal and non-mycorrhizal plants in the semihydroponic system. Net Pi absorption followed Michaelis–Menten kinetics: uptake rates declined with decreasing Pi concentrations in the nutrient solution. This reduction was most pronounced in non- mycorrhizal seedlings and plants colonized by T. terrestris. The results confirm that there is considerable heterogeneity in affinity for Pi uptake among the different mycobionts. It is concluded that the external mycelia of ectomycorrhizal fungi strongly influence the Pi-uptake capacity of the pine seedlings, and that some mycobionts are well equipped to compete with other soil microorganisms for Pi present at low concentrations in soil solution.

The possible involvement of polyamines in the tobacco mosaic virus (TMV)-induced hypersensitive reaction (HR) in Nicotiana tabacum (L.) cv. Samsun NN, which bears the N resistance gene, was investigated. Concentrations of free and conjugated polyamines, as well as the activities of their biosynthetic enzymes ornithine decarboxylase (ODC) and arginine decarboxylase (ADC), were analysed in the necrotic area (I, internal zone) and in the two concentric zones surrounding it (M, middle zone; E, external zone) during the progression of the HR (1, 3 and 5 d from TMV inoculation). Results were compared with those obtained in control plants (mock-inoculated) and with the pattern in susceptible TMV-infected tobacco plants.

In the HR, concentration increased towards zone I for free putrescine and spermidine. Highest concentrations in zones M and I occurred on day 3. Conjugated polyamines similarly showed an increasing concentration gradient towards zone I, with maximum accumulation on day 5. ODC and ADC activities also showed an increasing gradient towards zone I that could be related to the increase in free and conjugated polyamines. By contrast, in control plants concentrations of free and conjugated polyamines decreased from day 1 to day 5, ODC activity increased slightly, whereas ADC activity did not show significant changes. In the TMV-infected susceptible tobacco plants, polyamines tended to remain in the free state instead of being conjugated as infection progressed. These results suggest that polyamines could play a role in mechanisms of resistance to biotic stress.

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.

Three perennial grass species, Lolium perenne L., Agrostis capillaris L. and Festuca ovina L., were homogeneously labelled in phytotrons with 14CO2 at two CO2 concentrations (350 and 700 μl l−1). Plants were grown under two nitrogen regimes: one with a minor addition of 8 kg N ha−1, the other with an addition of 278 kg N ha−1. Carbon allocation over the different compartments of the plant/soil systems was measured: shoots, roots, rhizosphere soil (soil solution, microbial biomass and soil residue), and bulk soil. Elevated CO2 increased total net 14C recovery in all species by 14%, and significantly enhanced the below-ground 14C allocation by 26%, this enhancement was 24%, 39% and 21% for root, rhizosphere soil and bulk soil, respectively. Within the rhizosphere soil, the 14C amounts in the soil solution (+69%) and soil residue (+49%) increased significantly. Total microbial biomass-C in the rhizosphere soil was also increased (15%) by the elevated CO2 treatment, but only in proportion to the increased root mass. No interactions were observed between the elevated CO2 and N treatments. The N treatment increased total net 14C recovery by more than 300% and 14C was preferentially allocated to the shoots, leading to a significant increase in shoot-to-root ratio. However, N fertilization also increased (+111%) the absolute amount of 14C in soil. The three species behaved differently, but no interactions were observed between CO2 treatment and plant species. These results show that elevated CO2 induces an increased C input into soil for all three grass species at both N levels. However, the highest absolute amounts were found in the soils of the fastest growing species and at the highest N level.

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.

A glasshouse experiment was done to assess the development and phosphate metabolism of mycorrhizas formed by species of arbuscular mycorrhizal fungi (AMF) from two different genera, Gigaspora and Glomus on Desmodium ovalifolium plants at three concentrations of a phosphate source. The addition of phosphate (0–100 mg P kg−1) had no effect on the alkaline phosphatase activity, stained histochemically, in the intra-radical mycelium of Gigaspora rosea (BEG111), but decreased that of Glomus manihotis (BEG112) over a 10-wk period. The alkaline phosphatase activity of the extra-radical mycelium was unaffected by increasing phosphate addition (0–100 mg P kg−1) in both species of AMF over a 10-wk period. The extra-radical mycelium of Gi. rosea (BEG111) accumulated polyphosphate, determined by staining with 4′,6-diamidino-2-phenylindole, whereas polyphosphate was not detected in the extra-radical mycelium of G. manihotis (BEG112). This work indicates differences in the mechanisms of phosphate metabolism in the mycelium of AMF from different genera on a tropical host. This might be determined by the life-cycle strategies of these fungi, in particular the formation of auxiliary cells in Gigaspora. The possibility of a negative-feedback mechanism between alkaline phosphatase and polyphosphate in the extra-radical mycelium of Gi. rosea (BEG111) and the role of polyphosphate in the symbiosis are discussed.

The occurrence of crassulacean acid metabolism (CAM) among the epiphyte flora of the lowland forest on Barro Colorado Island (BCI), Panama, was investigated. A total of 116 species was included, i.e. about 2/3 of the known epiphyte taxa. As judged from the carbon isotope ratios and the absence of Kranz anatomy, indications of CAM were found in 29 species of three families, Orchidaceae (20), Bromeliaceae (7), and Cactaceae (2). We estimate that about 25% of the epiphyte flora of BCI are CAM plants. CAM was most prevalent in exposed sites, but even in the understorey two epiphyte species engage in CAM.

We performed two glasshouse experiments to determine whether the presence of arbuscular mycorrhizal symbiosis reduces the intensity of intraspecific competition at low concentrations of available phosphorus (P), and whether this effect is modified by a reduction in light intensity. In the first experiment, Trifolium subterraneum cv. Mt. Barker was grown at different densities in controlled conditions of light and temperature, and half of the pots were inoculated with spores of the arbuscular mycorrhizal fungus, Gigaspora margarita. In the second experiment, the plants were grown in similar controlled conditions but the light intensity received by half of the pots was reduced by >50%. The biomass and P content of individual mycorrhizal plants and the biomass response to mycorrhizal infection were drastically reduced as plant density increased. The effects of density on percentage infection, shoot and root P concentrations, and root: shoot ratios were inconsistent. Generally reduction in light intensity did not alter these effects. Mycorrhizal symbiosis increased intraspecific competition intensity through an increase in the availability of soil P. This increase in competition was reflected in the greater size inequality of low density mycorrhizal treatments. Our results emphasize that the main effects of mycorrhizas at the individual level cannot be expected to be apparent at the population level, because they are overridden by density-dependent processes.