Exposure of Chlorella vulgaris to elevated concentrations of copper, chromium, nickel and zinc led to intracellular accumulation of high concentrations of these metals. Concomitantly, accumulation of free proline occurred, depending on the concentration of metals in the external medium or in the cell. The greater the toxicity or accumulation of a metal, the greater the amount of intracellular proline in algal cells. However, higher concentrations of copper and chromium were inhibitory to proline accumulation by the test organism. The accumulation of proline was triggered within a few hours of metal treatment. Test metals also induced lipid peroxidation; copper was the most efficient inducer whereas zinc was the least. Pretreatment of C. vulgaris with proline counteracted metal-induced lipid peroxidation and potassium ion efflux. Thus the present work shows a protective effect of proline on metal toxicity through inhibition of lipid peroxidation.

Oxygen evolution was measured from mesophyll tissues in spinach leaves using a photoacoustic technique. The photosynthetic capacity of individual cell layers was measured by directing microscopic beams of light, 40 μm wide, to cells exposed within a leaf cross section. The resulting profile for oxygen-evolution potential was relatively flat, indicating a uniform capacity for photosynthesis in leaf mesophyll tissues. Two experimental approaches were used to estimate the photosynthetic performance of individual mesophyll cell layers when white light was applied to the adaxial leaf surface. These experiments indicated that oxygen was produced relatively uniformly across the mesophyll and that oxygen evolution increased with irradiance of the white light applied to the leaf surface. The measured profiles for oxygen evolution and capacity are flatter than previous measurements of profiles of fixed carbon and estimates of profiles for absorbed light within spinach leaves.

Feeding dynamics in the spittlebug Philaenus spumarius were analysed to determine the food source. Evidence from previous reports indicates that this species and similar insects feed from the xylem. However, it is not clear whether they extract sap from the main transpiration stream, or from some subsidiary compartment of xylem, such as immature or cavitated vessels. In feeding from such subsidiary compartments, the insects could avoid the large hydraulic tensions thought to prevail in xylem on the main transpiration stream. Philaenus, for example, is shown here to initiate feeding, and to feed rapidly, from plants in which the full xylem tension is estimated at −10 bar. From measurements of feeding rates and feeding patterns in stressed and unstressed plants, it is shown that Philaenus cannot feed from subsidiary xylem compartments. It is concluded that this insect feeds directly from the main transpiration stream, and at the full hydraulic tension which prevails there. The anatomy of the feeding pump in Philaenus appears consistent with this conclusion. Spittlebugs therefore offer an experimental tool for quasi- non-invasive sampling of the transpiration stream.

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.

The function of a leaf is photosynthesis, which requires the interception of light and access to atmospheric CO2 while controlling water loss. This paper examines the influence of leaf anatomy on both light capture and CO2 diffusion. As photosynthetic metabolism is spread between many chloroplasts, a leaf faces the challenge of matching light capture by a given chloroplast with the metabolic capacity of that chloroplast. Chloroplasts nearest the leaf surface receive the greatest irradiance and therefore absorb more light per unit chlorophyll than chloroplasts in the centre of a leaf. Electron transport and carbon fixation capacities per unit of chlorophyll decline with increasing depth in the leaf, to compensate for the decline in light absorbed per unit chlorophyll. Many key photosynthetic protein complexes in chloroplasts have nuclear encoded genetic information. Consequently, all chloroplasts within a given cell have a similar metabolic complement, which limits the potential gradient of photosynthetic capacity per unit chlorophyll across the leaf. A simple model couples light absorption through the leaf (based on the Beer–Lambert law) with the profile of chlorophyll through a leaf and the gradient in photosynthetic capacity. It is validated by comparison with 14CO2 fixation profiles through spinach leaves obtained in various studies. The model can account for published 14C fixation profiles obtained with blue, red and green light of different irradiances and white light applied in different combinations to the adaxial and abaxial surfaces of spinach leaves. The model confirms that spongy mesophyll increases the apparent extinction coefficient of chlorophyll compared to palisade tissue. The palisade tissue nearest the surface which receives light facilitates the penetration of light to a greater depth, while spongy mesophyll promotes scattering to enhance light absorption, thus reducing the gradient in light absorbed per unit chlorophyll through a leaf. CO2 fixation faces a diffusional limitation, which necessitates Rubisco to be spread evenly across the cell walls exposed to intercellular airspace. Mesophyll cell structure reflects the need to have a large cell surface per unit volume exposed to airspaces. The regular array of columnar cells in palisade tissue, or cell lobing in monocot leaves, results in greater exposed surface per unit tissue volume than spongy mesophyll. The exposed surface area per unit leaf area scales with photosynthetic capacity such that the difference in CO2 partial pressure between substomatal cavities and the sites of carboxylation within chloroplasts is, on average, independent of photosynthetic capacity of the leaf. However, Rubisco specific activity declines as the Rubisco content per unit leaf area increases due to greater internal diffusional limitations.

A model of seed germination is proposed which uses a variable with the units of an osmotic potential (virtual osmotic potential) to integrate the effect of a constant or a varying water potential. This differs from existing models that describe the effects of fixed water potentials on germination, or the effects of fixed priming water potentials on the subsequent germination at a fixed water potential. When a seed is sown, the virtual osmotic potential is assumed to fall at a rate that depends on the ambient water potential, and on the difference between its current and a minimum value. Radicle growth is assumed to initiate when the difference between the ambient water potential and the virtual osmotic potential exceeds a threshold. The germination of carrot and onion seeds at various fixed potentials below 0 MPa was well described by the virtual osmotic potential model. The model was also used to simulate the results of experiments in which seeds were given a single step change in water potential.

Chlorophyll fluorescence measurements were used to test whether externally applied abscisic acid (ABA) has an effect on the desiccation tolerance of lichens and their sensitivity to constant water saturation. A surplus of ABA did not decrease the time required to regain full photosynthetic capacity after prolonged dry periods. However, an effect of ABA could be observed when lichens were permanently hydrated at moderately high temperatures. Lichens suffered less from constant saturation if ABA was added to the incubation medium. These results suggest a positive effect of ABA on membrane function.

The relative rates of ammonium and nitrate-N uptake and assimilation by creeping bent (Agrostis stolonifera), were investigated for plants grown in soil and supplied with three different ratios of ammonium and nitrate-N. Following two preliminary defoliations, plants were supplied with the equivalent of 150 kg N ha−1, given as 15N-(differentially) labelled NH4+ and NO3−-N in three different ratios (20:80, 50:50 and 80:20), followed by sequential destructive harvests of shoots and roots at four points during a 35-d regrowth period. Maximum use of labelled nitrogen and ‘exhaustion’ of soil mineral nitrogen reserves occurred much earlier when plants were supplied with half or more of their nitrogen as ammonium, than occurred when they were supplied predominately with nitrate-N. The lack of consistency in the patterns of ammonium and nitrate-N absorption, however, implied that the plants had no specific preference for either nitrogen form. Supplying plants with different combinations of ammonium and nitrate produced distinctive differences in plant morphology. In the high nitrate treatment, plants preferentially partitioned resources into shoot and stolon formation, whereas in the high ammonium treatment, resources were preferentially partitioned into root production. These changes in plant morphology might be adaptations to aid species survival in environments associated with a predominance of either nitrogen form.

This study explored reflectance indices sampled with a ‘leaf reflectometer’ as measures of pigment content for leaves of contrasting light history, developmental stage and functional type (herbaceous annual versus sclerophyllous evergreen). We employed three reflectance indices: a modified normalized difference vegetation index (NDVI), an index of chlorophyll content; the red/green reflectance ratio (RRED[ratio ]RGREEN), an index of anthocyanin content; and the change in photochemical reflectance index upon dark–light conversions (ΔPRI), an index of xanthophyll cycle pigment activity. In Helianthus annuus (sunflower), xanthophyll cycle pigment amounts were linearly related to growth light environment; leaves in full sun contained approximately twice the amount of xanthophyll cycle pigments as leaves in deep shade, and at midday a larger proportion of these pigments were in the photoprotective, de-epoxidized forms relative to shade leaves. Reflectance indices also revealed contrasting patterns of pigment development in leaves of contrasting structural types (annual versus evergreen). In H. annuus sun leaves, there was a remarkably rapid increase in amounts of both chlorophyll and xanthophyll cycle pigments along a leaf developmental sequence. This pattern contrasted with that of Quercus agrifolia (coast live oak, a sclerophyllous evergreen), which exhibited a gradual development of both chlorophyll and xanthophyll cycle pigments along with a pronounced peak of anthocyanin pigment content in newly expanding leaves. These temporal patterns of pigment development in Q. agrifolia leaves suggest that anthocyanins and xanthophyll cycle pigments serve complementary photoprotective roles during early leaf development. The results illustrate the use of reflectance indices for distinguishing divergent patterns of pigment activity in leaves of contrasting light history and functional type.

The facultative hemiparasite Rhinanthus minor was grown alone, or with either Hordeum vulgare or Trifolium alpestre as a host. All plants were fed 5 mM nitrate. In the parasites, successful attachment led to dramatic increases in growth (particularly with barley as host) and in tissue concentrations of total N, NO3−, H2PO4− and K+, as well as moderate increases in Cl− and SO42−. Mg2+ and Ca2+ concentrations either remained unchanged, increased or decreased, depending on the host. Xylem sap was collected from leaf veins (barley) or the stem (Rhinanthus and clover), by applying pneumatic pressure to the rooted soil to raise the soil water potential so that in vivo xylem sap exuded from small incisions once the applied pressure balanced the tension exerted by transpiration. High balancing pressures were needed for unattached Rhinanthus, whereas simultaneous collection of xylem sap from attached Rhinanthus and its host, at a much lower balancing pressure, was possible only when the parasite was enclosed in polyethylene film and high transpiration abolished. From plots of xylem sap volume flow versus applied pressure, the hydraulic conductivity Lp and the overall hydraulic resistance r were obtained. The value of r was high in unattached Rhinanthus and dramatically decreased by attachment to a host, highlighting the improved access to water gained by Rhinanthus when parasitizing a host. For the parasites, attachment resulted in greatly increased concentrations and solute flow rates in xylem of K+, NO3−, H2PO4− and amino acids, and decreased concentrations of Ca2+ and Mg2+. Estimates of NO3− reduction in the parasite showed that 99% was reduced in unattached plants, 85% in those attached to clover and 52% in those attached to barley. In the parasitized hosts, ion concentrations in xylem sap were somewhat increased relative to unparasitized controls. Conversely, xylem sap amino-acid concentrations in infected hosts were decreased. Glutamine (Gln) was the principal amino acid in xylem sap of unattached Rhinanthus, but after attachment to both hosts asparagine (Asn) predominated. In clover, Asn was the major transport amino acid, suggesting massive transfer of Asn from host to parasite. In unparasitized barley, however, Gln was the major xylem sap amino acid, but unexpectedly, in this species Asn was induced as the principal amino acid when parasitized. The results are discussed in terms of mutual host–parasite interactions.

Leaching of nitrate through soil profiles is a major cause of soil acidification. Legume species differ in their ability to take up nitrate (NO3−) and in the degree to which soil NO3− impairs legume nodulation and N2 fixation. This pot experiment examined species variation in N2 fixation and acid production in grain legumes grown for 6 wk on a sandy soil with Ca(NO3)2 added at 0–56.7 mg N kg−1 soil. The eight legumes studied were Lupinus angustifolius (‘Gungurru’), Lupinus albus (‘Kiev mutant’), Lupinus luteus (‘Teo’), Cicer arietinum (Selection T1587), Pisum sativum (‘Dundale’ and ‘Wirrega’), Vicia faba (‘Fiord’) and Lathyrus sativus (Selection 453). While number of nodules was not significantly affected by treatment with NO3−, increasing supply of NO3− decreased nodule mass, with P. sativum ‘Wirrega’ being most sensitive and C. arientinum and V. faba least sensitive. The concentrations of nitrogen in plants generally decreased as the supply of NO3− increased. Increasing supply of NO3− decreased the percentage of N2 fixation, most significantly in L. sativus and least in L. albus. In the absence of added NO3−, the amounts of H+ produced per unit biomass (specific acid production) differed by as much as 2.2-fold among the species, with C. arientinum generating most H+, followed by L. angustifolius, L. luteus, L. albus, V. faba, L. sativus, P. sativum ‘Dundale’ and P. sativum ‘Wirrega’. This species variation was even greater at higher NO3− supply. In the presence of NO3− at up to 14.2 mg N kg−1, acid production by these legumes correlated well with uptake of excess cations but not with the concentration of nitrogen in plants. For all species, release of H+ declined with increasing supply of NO3−, decreasing most in L. sativus and least in C. arientinum. The results suggest that there might be scope for selecting grain legume species in order to minimize soil acidification by decreasing leaching of NO3−.

A centrifugal method for extracting apoplastic sap from roots of lupin (Lupinus angustifolius) and pea (Pisum sativum) plants, and a method to analyse malic dehydrogenase in the sap using capillary electrophoresis, are described. Osmolality of apoplastic sap was relatively constant at relative centrifugal forces (RCFs) between 600 and 3000 g for lupin, and between 600 and 4000 g for pea. Electropherograms of a marker enzyme (malic dehydrogenase) and other components in apoplastic and symplastic saps revealed that contamination occurred at 7000 g. It is suggested that apoplastic sap expelled from plant roots at RCF between 600 and 3000 g is free from symplastic contamination, and is regarded as being of apoplastic origin. The proposed method was used to measure apoplastic pH changes in the plant roots in response to external pH, ammonium, nitrate and vanadate.

The relationships between leaf structure, nitrogen concentration and CO2 assimilation rate (A) were studied for 14 grass species grown in the laboratory under non-limiting nutrient conditions. Structural features included leaf thickness and density, and the proportion of leaf volume occupied by different types of tissue (mesophyll, epidermis, vessels and sclerenchyma). Relationships were assessed for data expressed per unit leaf area and fresh mass. The latter was found to be closely related to leaf volume, which allowed us to use A per unit leaf fresh mass (Afm) as a surrogate of A per unit leaf volume. Assimilation rate per unit leaf area (Aa) was positively correlated with leaf thickness and with the amount of mesophyll per unit leaf area; the relationship with leaf nitrogen content per unit area was only marginally significant. Afm was negatively correlated with leaf thickness and positively with fresh mass-based leaf organic nitrogen concentration. A multiple regression involving these two variables explained 81% of the variance in Afm. The value of Afm was also significantly related to the proportion of mesophyll in the leaf volume, but surprisingly the correlation was negative. This was because thin leaves with high Afm and nitrogen concentration had proportionally more mechanically supportive tissues than thick ones; as a consequence, they also had a lower proportion of mesophyll. These data suggest that, in addition to leaf nitrogen, leaf thickness has a strong impact on CO2 assimilation rate for the grass species studied.

Both sensitive and tolerant clones of aspen (Populus tremuloides) were exposed to ozone using four different exposure regimes under controlled environmental conditions. Based on data on ambient ozone from 10 cities in the USA, three treatments of 4-wk exposure to the same SUM06 (an accumulation of hourly O3 concentrations greater than 0.06 ml l−1) were constructed. The regimes allowed us to investigate: (a) the importance of long (3 wk, treatment 1) versus short (1 wk, treatment 2) duration of regimes with high peaks; (b) the effect of treatments with variable peak occurrence (treatments 1 and 2) versus uniform peak occurrence (treatment 3) during the exposure period. Nonfumigated control plants were maintained at ozone concentrations <10 nl l−1. Bifacial black necrosis, a typical symptom of ozone injury on aspen leaves, occurred on both clones after 2 wk exposure. Up to 60% of the leaves on the sensitive clone were injured, with an average of 6% of total leaf area injured. In the tolerant clone only 10% of the leaves were injured, with less than 1% of the total leaf area symptomatic. The severity of injury was consistently greatest in treatment 2, followed by treatments 1 and 3, respectively. The interval between peak exposures was less important than the occurrence of peaks versus a stable maximum concentration. Premature leaf abscission occurred in the sensitive clone. Measures of gas exchange demonstrated reduced photosynthesis under ozone fumigation, but exposure regime was not a significant factor. Concentrations of two antioxidants, ascorbic acid and glutathione, were almost always greater in the resistant than in the sensitive clone, but the differences were not statistically significant. The levels of these antioxidants in aspen leaves did not change with ozone fumigation or leaf age.

This study examines interrelationships between eight leaf attributes (specific leaf mass, area, dry mass, lamina thickness, mesophyll cell number per cm2, mesophyll cell volume, chloroplast volume, and number of chloroplasts per mesophyll cell) in field-grown plants of 94 species from the Eastern Pamir Mountains, at elevations between 3800 and 4750 m. Unlike most other mountain areas, the Eastern Pamirs, Karakorum system, Tadjikistan provide localities where low temperatures and radiation combine with moisture stress at high altitudes. For all the attributes measured, significant differences were found between plants with different mesophyll types. Leaves with dorsiventral palisade structure (dorsal palisade, ventral spongy mesophyll cells) had thicker leaves with larger but fewer mesophyll cells, containing more and larger chloroplasts. These differences in mesophyll type are reflected in differences in the total surface of mesophyll cells per unit leaf area (Ames/A) or volume (Ames/V). Plants with isopalisade leaf structure (palisade cells under both dorsal and ventral surfaces) are more commonly xerophytes and their increased values of Ames/A and Ames/V decrease CO2 mesophyll resistance, which is an important adaptation to drought. Path analysis shows the critical importance of mesophyll cell volume in leading to the covariance between the different leaf attributes and hence to specific leaf mass (SLM), even though mesophyll cell volume is not itself strongly correlated with SLM. This is because mesophyll cell volume increases SLM through its effects on leaf thickness and chloroplast number per cell, but decreases SLM through its negative effect on mesophyll cell density.

Microbial immobilization may decrease the inorganic nutrient concentrations of the soil to the extent of affecting plant nutrient uptake and growth. We have hypothesized that graminoids with opportunistic nutrient-acquisition strategies are strongly influenced by nutrient limitation imposed by microbes, whereas growth forms such as dwarf shrubs are less affected by the mobilization–immobilization cycles in microbes. By adding NPK fertilizer, labile C (sugar) and fungicide (benomyl) over a 5 yr period in a fully factorial design, we aimed to manipulate the sink–source potential for nutrients in a non-acidic heath tundra soil. After 2 yr, N and P accumulated in the microbial biomass after fertilization with no change in microbial C, which suggests that nutrients did not limit microbial biomass growth. After 5 yr, microbial C was enhanced by 60% in plots with addition of labile C, which points to C-limitation of the microbial biomass. Microbial biomass N and P tended to increase following addition of labile C, by 10 and 25%, respectively. This caused decreased availability of NH4+ and P, showing close microbial control of nutrient availability. The most common graminoid, Festuca ovina, responded to fertilizer addition with a strong increase, and to labile C addition with a strong decrease in cover, providing the first direct field evidence that nutrient limitation imposed by immobilizing microbes can affect the growth of tundra plants. Also in support of our hypothesis, following addition of labile C the concentrations of N and K in leaves and that of N in roots of F. ovina decreased, whilst the demand of roots for P increased. In contrast, the most common dwarf shrub, Vaccinium uliginosum, was only slightly sensitive to changes in resource availability, showing no cover change after 4 yr addition of labile C and fertilizer, and little change in leaf nutrient concentrations. We suggest that the differential responses of the two growth forms are due to differences in storage and nutrient uptake pathways, with the dwarf shrub having large nutrient storage capacity and access to organic forms of N through its mycorrhizal association. While the fungicide had no effect on ericoid mycorrhizal colonization of roots or symbiotic function inferred from plant 15N natural abundance, it decreased microbial biomass C and N after 2 yr. Throughout the fifth season, the availability of soil NO3− and inorganic P was decreased with no change in microbial biomass C, N or P, suggesting a negative impact of benomyl on N and P mineralization.

Potted subterranean clover (Trifolium subterraneum) plants of different ages were exposed to 70 nl l−1 ozone for 6 h, either during the light or during the dark period in a laboratory-based climate chamber. There was limited visible leaf injury on plants which were 14–20 and 28–34 d old and no significant decrease in biomass after daytime ozone exposure. The oldest leaves of 22–26 d old plants exhibited severe visible injury, which was associated with a significant reduction in biomass in 24–26 d old plants. Thus, ozone-induced visible injury of different magnitude developed in all plants, but was associated with biomass reduction only during a limited period of the plant's life-span. Apart from modifying ozone uptake by plants, climatic conditions are important as growth modifiers. It is suggested that subterranean clover plants of defined developmental stages should be used in bioindication of ozone. Night ozone exposure injured significantly fewer leaves than day exposure. However, some leaves developed visible injury even after night ozone exposure. Night uptake of ozone may be of more importance in northern than in central and southern Europe, because summer nights are short and, for a certain period, never completely dark.

An appendix is available for this article which was not included in the print edition of the Journal. The appendix may be found at the rear of the full text, PDF version of the article.

A popular conceptual model asserts that shade tolerance is characterized by morphological and physiological traits that enhance the net rate of carbon capture in low light. We tested this model by quantitatively reviewing growth, leaf lifespan, CO2 exchange and morphological data from 76 studies on woody seedlings grown under conditions of low light. Data were placed into three tolerance categories (intolerant, intermediate, tolerant), two light categories (less than 4% and 4–12%) and two leaf phenology categories (broad-leaved evergreen and winter deciduous). For both evergreen and deciduous groups, intolerant species had traits conferring greater growth potential than tolerant species in both light categories. These traits included greater leaf mass ratio, leaf area ratio, specific leaf area and mass-based photosynthetic rates above light compensation. However, in 0–4% light, growth rates were similar for intolerant and tolerant species, because low light together with higher respiration rates for intolerant species limited the expression of their growth potential differences. Deciduous and evergreen intolerant species were similar in many respects. However, both intermediate and tolerant deciduous species had markedly lower leaf mass ratios and higher root mass ratios than intermediate and tolerant evergreen species. In addition, deciduous species and intolerant evergreens must cope with as much as sixfold higher leaf turnover rates than tolerant evergreen species. Thus, rather than maximizing growth rates in low light, tolerant evergreen species minimize biomass loss through long leaf lifespans and low respiration rates. Tolerant deciduous species also minimize biomass losses by minimizing whole-plant respiration rates but they accomplish low biomass turnover though low leaf mass ratio and not low leaf turnover rates. Furthermore, unlike most tropical evergreens, tolerant deciduous species can gain large fractions of their total growing season carbon during short periods when the overstory is leafless and then allocate this carbon to storage (as reflected by high root mass ratios) rather than new leaves. In conclusion, we found no support for the low-light-enhanced carbon capture model of shade tolerance as viewed strictly from the perspective of physiological growth capacity. This can be explained by the disadvantages to net growth and survival of maintaining a high growth potential at low light, because high growth potential results in greater rates of whole-plant respiration, tissue turnover, herbivory and mechanical damage and in decreased storage. Thus, shade tolerance can be characterized by traits that maximize survival and net growth, where net growth includes losses to all agents.

This study focuses on the following questions: (i) whether reductions in root:shoot ratio have a cost in terms of nutrient balance of the plant, and (ii) whether changes in resource-allocation patterns are proportional among different resources. Our approach was to analyse the variations in the allocation pattern induced by soil waterlogging. A pot experiment was conducted to analyse the effects of waterlogging on biomass, phosphorus (P) and nitrogen (N) accumulation of Paspalum dilatatum and Danthonia montevidensis, two waterlogging-tolerant grasses. When changing from oxic to anoxic conditions, a common response of these and other waterlogging-tolerant grasses is a reduction in allocation to below-ground resources. It was observed that (i) the reduction in root:shoot ratio caused by waterlogging did not have a cost in terms of capacity for nutrient uptake; (ii) resource partitioning within aerial parts was less sensitive to treatments than partitioning between roots and shoots; and (iii) biomass does not appear to be a useful currency for evaluating nutrient-allocation patterns, as the allocation of P and N was inadequately represented by biomass. The results presented here indicate that the existence of compensation mechanisms reduces the predictive value of the partition of resources for the capacity of plants to acquire resources. Data on the allocation of nutrients in relation to biomass suggest that the assumptions of independence in the allocation pattern between biomass and limiting nutrients under the effects of environmental factors can be extended.

The responses to CO2 of perennial grasses (Danthonia richardsonii and Phalaris aquatica) and legumes (Lotus pedunculatus and Trifolium repens) were compared under controlled conditions for isolated plants, monoculture stands and mixed-species stands along a N gradient to test whether: plant–plant interactions between species in mixed stands changed with concentration of CO2; responses to CO2 of species in mixtures could be related to their responses as single stands; responses of mixtures to CO2 could be related to the responses of individual species to CO2 and to competition. Plants were grown for 60 d in sand, using nutrient solutions (six nitrate concentrations from 0.25 to 16 mM NO3), at ambient (c. 357 μl l−1) or elevated CO2 (c. 712 μl l−1). Species dominance in the mixtures depended more on the range of N than of CO2 concentration provided: T. repens and L. pedunculatus dominated at low concentrations of N; L. pedunculatus and P. aquatica performed better at high concentrations. Responses of species in mixtures to CO2 were related to their responses in monocultures but not to those of isolated plants. Species biomass proportions in mixtures under ambient CO2 determined the outcome of mixture responses to CO2 more than of individual species responses to CO2. These results emphasize the influence of plant–plant interactions on community responses to CO2, since mixture behaviour under elevated CO2 could not be scaled-up from responses by isolated plants in this experiment.