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.

Twenty-one isolates of the ectomycorrhizal fungus Suillus luteus were screened for their tolerance to the heavy metals Zn, Cd, Cu and Ni, measured as inhibition of radial growth and biomass production. Two populations from even-aged pine stands were investigated: 10 isolates were obtained from an area polluted with high levels of Zn, Cd and Cu, and 11 isolates were obtained from a control population located in a nearby unpolluted area. RFLP patterns of the internal transcribed spacer region of the isolates confirmed the morphological identification of the carpophores. All isolates were maintained on basic medium without elevated metals to avoid phenotypically acquired metal tolerance. The in vitro Zn and Cd tolerance of the S. luteus isolates from the polluted habitat were significantly higher than the tolerances measured in the isolates from the nonpolluted site. This observation suggests that the elevated soil metal concentrations might be responsible for the evolution of adaptive Zn and Cd tolerance. Tolerance was maintained in an isolate not exposed to elevated metals for 3 yr. The two S. luteus populations did not differ in tolerance to Cu and Ni. The mechanisms for the adaptive Zn and Cd tolerance are not identical as there was no correlation between response to the two metals; the most Zn-tolerant isolate was the most sensitive for Cd in the metal-tolerant population. Zinc did not accumulate in basidiocarp tissue, whereas Cd levels in basidiocarps were significantly higher in the population on the polluted site. Inter-simple sequence-repeat fingerprints showed that 90% of the isolates were from different individuals. The genetic variation in the population from the unpolluted site was considerably larger than that observed at the polluted site.

The hydraulic conductance (L0) of detached, exuding root systems from melon (Cucumis melo cv. Amarillo oro) was measured. All plants received a half-strength Hoagland nutrient solution, and plants stressed either solely with NaCl (50 mM) or with NaCl (50 mM) following treatment (2 d) with CaCl2 (10 mM) were compared with controls and CaCl2-treated (10 mM) plants. The L0 of NaCl-treated plants was markedly decreased when compared to control and CaCl2-treated plants, but the decrease was smaller when NaCl was added to plants previously treated with CaCl2. A similar effect was observed when the flux of Ca2+ into the xylem and the Ca2+ concentration in the plasma membrane of the root cells were determined. In control, CaCl2- and NaCl + CaCl2-treated plants, HgCl2 treatment (50 μM) caused a sharp decline in L0 to values similar to those of NaCl-stressed roots, but L0 was restored by treatment with 5 mM DTT. However, in NaCl roots only a slight effect of Hg2+ and DTT were observed. The effect of all treatments on L0 was similar to that on osmotic water permeability (Pf) of individual protoplasts isolated from roots. The results suggest that NaCl decreased the passage of water through the membrane and roots by reducing the activity of Hg-sensitive water channels. The ameliorative effect of Ca2+ on NaCl stress could be related to water-channel function.

The daily integral of photosynthesis for a population of planktonic cyanobacteria in the Baltic Sea is calculated by numerical analysis based on the photosynthesis–irradiance curve of the organism, the vertical distribution of the population and details of the underwater light field. Variations in the light field are calculated from continuous recordings of surface irradiance and measurements of vertical light attenuation, with corrections for losses by reflection at the water surface that depend on the sun's elevation and roughening by wind. Integrals calculated using models in which simplifying assumptions are made about the surface irradiance overestimate the calculated value by 21% on a cloudless day and by 75%, or more, on an overcast day. Even larger discrepancies (>100%) result when the population, which is concentrated in near-surface layers, is assumed to be evenly distributed with depth. Errors of up to 5% result from neglect of surface reflection, and up to 12% from neglect of changes in phytoplankton distribution over 24 h. Details are given of the method of numerical analysis, using standard computer spreadsheets. Versions of the spreadsheet are described in which effects of changes in the vertical distribution of the phytoplankton and light attenuation can be calculated and the effects of water temperature and photoinhibition can be modelled. The method will improve estimates of primary productivity in natural waters. It can also be used for calculating values of the euphotic depth integrated over 24 h. The spreadsheets are available on http://www.bio.bris.ac.uk/research/walsby/integral.htm.

This paper reports the effects of nutrient magnesium (Mg) concentrations on the growth and photosynthetic physiology of clonal Pinus radiata from four female parents (families) known to differ in their tolerance to Mg deficiency and in their needle Mg concentrations. Plants were grown in flowing nutrient solutions with 2 mg l−1 (control) and 0.8 mg l−1 (low) Mg. Plant growth, needle Mg concentration, photosynthesis, chlorophyll fluorescence and carotenoid pigment content were measured. At low Mg, needle Mg concentration was about half that of control plants, height growth was reduced 15–25%, and the needles showed strong visual characteristics of Mg deficiency. Photosynthesis was also halved, and was associated with closure of the stomata under low Mg and with reductions in the residual conductance. In needles from plants grown at low Mg, photochemical yield was reduced both in the light and in the dark, and was strongly dependent on needle Mg concentrations below a threshold concentration of 0.02–0.025% (d. wt basis). The electron transport rate (ETR) at saturating photon flux density in low-Mg-grown needles was reduced to about half that of their Mg controls, but the photon efficiency of ETR was unaffected by the Mg concentration the plants were grown in. Photosynthetic quenching was markedly reduced and non-photosynthetic quenching was increased following growth in low Mg. Growth under low Mg also increased levels of zeaxanthin. Although family differences in growth and photosynthetic physiology were present, few family × Mg interactions were significant. We conclude that Mg deficiency probably affects growth through severe reductions in photosynthesis.

Embolisms in the vessels of maize axile roots of different types were observed directly after rapid freezing of intact, functioning roots in the field, by cryo-scanning electron microscopy. Quantification of the degree of embolization in each root was made by counting empty and full vessels of both the late and early metaxylem (LMX & EMX), and expressed as percent embolized vessels of the LMX, and %EMX poles containing embolized vessels. Contents of the connecting xylem (CX) at branch root junctions, and of xylem in branch roots were observed also, but not systematically quantified. Records of % embolized vessels were made from dawn to dusk on summer days in Ottawa under moderate irradiance, and in Canberra under high irradiance. Measurements in Canberra were supported by estimates of irradiance, of stomatal conductance, and of chamber balance pressure of bagged and unbagged leaves. Soon after sunrise embolisms appeared in all types of vessel, at balance pressures c. 300–400 kPa, and increased rapidly with increasing irradiance. During the middle of the day % embolized vessels reached a maximum (LMX ≈70% in Ottawa, and ≈80% in Canberra). At all times the EMX vessels were less embolized. The midday maximum was brief in Ottawa, and % embolized vessels fell to a low value during the afternoon. In Canberra the maximum was prolonged into late afternoon. By dusk nearly all vessels were once again filled with sap. The balance pressures measured during vessel refilling in Canberra ranged from 500 kPa to 1200 kPa. At all times of the day sap was seen entering some embolized vessels. Almost all were refilling by mid- to late-afternoon. Such refilling was especially frequent at junctions of branch roots with the axile roots. X-ray microanalysis of the sap entering the vessels, and of the liquid filling or partly filling vessels, showed the concentration of mineral solutes present in the sap was below the threshold of detection (≈12 mM). These results are discussed in relation to current opinions about embolisms and vessel refilling.

Eucalyptus coccifera Hook., a plant capable of forming both arbuscular mycorrhizas and ectomycorrhizas, was used to compare the effects of the two mycorrhizal types on phosphorus uptake and C allocation. Seedlings were grown in a P-deficient soil/sand mixture inoculated with peat/vermiculite spawn of Laccaria bicolor (Maire) Orton or Thelephora terrestris (Ehrh.) Fr.; or with 250-μm sievings from leek colonized by Glomus caledonium (Nicol. & Gerd.) Trappe & Gerde., Glomus sp. type E3 or Glomus mosseae (Nicol. & Gerd.) Gerd. & Trappe or with autoclaved spawn (non-mycorrhizal control). Before the 89-d harvest, a subset of the harvested plants was labelled with 14C (45–60-min pulse, 202-h chase). Growth promotion and the increase in seedling P content was largest in the two ectomycorrhizal treatments. Production of fluorescein diacetate-stained external hyphae was three to seven times higher by ectomycorrhizal (ECM) fungi compared with arbuscular mycorrhizal (AM) fungi and was highly correlated with P uptake and shoot weight. Phosphorus inflow rates of ECM and AM seedlings were 3·8 times, and 2·0–2·7 times those of non-mycorrhizal seedlings. Phosphorus acquisition efficiencies were similar (11·2 and 10·0 μmol P mmol−1C for T. terrestris and Glomus E3 plants, respectively) for the two mycorrhizal types, and appeared to be greater than in uninoculated plants (7.2 μmol P mmol−1C) grown at the same P level.

High loads of nitrogen to spruce and beech forests can result in a complete inhibition of NO3− uptake by the roots of the trees. This conclusion is based on (a) a comparison of a field site continuously exposed to high loads of N and a N-limited site, (b) the results of N fertilization of a N-limited field site, and (c) laboratory experiments under controlled environmental conditions. From fertilization experiments in the field it appears that NH4+ uptake might become inhibited subsequent to an excessive uptake of NH4+. Apparently, the inhibition of NO3− uptake by high loads of N to forests is a consequence of an accumulation of organic amino compounds in the roots originating from phloem transport from the shoot to the roots. These amino compounds seem to signal the N demand of the shoot to the roots. At present this function cannot be attributed to an individual organic amino compound in beech or spruce, but Gln is a likely candidate in both species among other compounds, e.g. Glu in spruce or Asp in beech trees. Direct inhibition of NO3− uptake by NH4+ can be excluded from the present studies. The mechanism(s) by which elevated levels of particular organic amino compounds interact with NO3− uptake remains to be elucidated. This (these) mechanism(s) seem to affect NO3− influx rather than NO3− efflux. As a consequence of this (these) mechanism(s), spruce and beech trees can prevent, within a certain physiological window, N over-nutrition when the roots are exposed to excessive amounts of inorganic N. However, inhibition of NO3− and NH4+ uptake by the roots makes more N available for leaching into the ground water and, in addition, for soil microbial processes that result in the production and re-emission of volatile N compounds into the atmosphere.

At the ‘Höglwald’ site, continuously exposed to high loads of N, >20% of the N input from throughfall into the spruce and beech plots is re-emitted as NO and N2O. However, the NO to N2O ratio is highly dependent on the tree species, with a preference for NO in the spruce and a preference for N2O in the beech plot. Since at least part of the NO emitted from the soil will be converted inside the canopy in the presence of ozone to NO2 that might then be absorbed by the leaves, the portion of the N in the throughfall that will be released from the forest by gaseous N emission is higher in the beech than in the spruce plot. Leaching of NO3− into the ground water is high in the spruce, but minute in the beech plot. However, this positive effect of beech on ground water quality is achieved at the expense of an enhanced release of radiatively active N gases into the troposphere.

The flooding tolerance of Carex species was studied in relation to their field distribution and their capacity to form root aerenchyma under controlled conditions. In an alpine meadow, six Carex species were selected which were distributed in a clear zonation correlating with water content of the soil. Carex sempervirens and C. ferruginea were only found on nonflooded soil, the latter species preferring moister conditions. Carex davalliana and C. nigra were both associated with water-saturated soil, whereas C. limosa and C. rostrata preferred partially submerged conditions. Carex davalliana and C. limosa were bound to flooded soils with a relatively high redox potential and horizontally flowing groundwater. Carex rostrata and C. nigra grew in stagnant soil-flooded conditions with low soil redox potentials. The amount of aerenchyma in the roots of all species increased when grown in oxygen-deficient stagnant agar. This increase in root porosity, combined with increased root diameter, presumably improved internal aeration of the roots. Although all species survived experimental soil flooding, partial submergence was lethal to C. sempervirens and, surprisingly, also to the wetland species C. davalliana. Carex ferruginea showed a reduced growth rate during partial submergence. The three other species, all wetland plants, reached highest biomass production under soil-flooded and partially submerged conditions, with slower growth on free-draining soil. It is concluded that aerenchyma is not constitutive in the Carex species under study, and is best developed in Carex species from wetlands. Species with less aerenchyma perform poorly when soil-flooded, but conditions of partial submergence could even affect species with a considerable amount of root aerenchyma.

We have studied the chitin and ergosterol contents of ectomycorrhizal roots in three sets of experiments to evaluate them as indicators of fungal biomass. The first set of experiments showed that ageing had a marked effect on ergosterol concentrations. The ergosterol content of 7-month-old, brown, shrunken Pinus sylvestris L.–Paxillus involutus (Fr.) Fr. mycorrhizas was found to be only 10% of that found in white, turgid, 1- or 4-month-old specimens. This supports the hypothesis that the compound is a good indicator of living fungal biomass. Ageing had a lesser effect on chitin concentrations since the chitin levels found in 7-month-old mycorrhizas were still 60% of the levels found in 1- and 4-month-old specimens.

Consequently, the chitin[ratio ]ergosterol ratio increased from about 14 to 19 in 1- and 4-month-old mycorrhizas respectively to about 110 in 7-month-old mycorrhizas. In the second set of experiments, we found that variation in plant growth had no effect on the chitin[ratio ]ergosterol ratio in whole root systems of either Alnus incana (L.) Moench or Pinus sylvestris mycorrhizal with Paxillus involutus. In the third set of experiments, we found a constant relationship between the two marker concentrations in 10-month-old root systems of Pinus sylvestris, regardless of fungal species involved, using Paxillus involutus, Piloderma croceum Erikss. & Hjorts and Suillus variegatus (Fr.) O. Kuntze as test organisms. Taken together, the results of this study suggest that both chitin and ergosterol give reliable, but different, relative measures of fungal biomass in mycorrhizal roots. Furthermore, we demonstrate that, in combination, the two chemical markers can be used to estimate both total and living fungal biomass (derived from the chitin[ratio ]ergosterol ratio).

Evidence that enhanced reactive nitrogen deposition is affecting semi-natural terrestrial ecosystems comes from historic increases in plant tissue N concentrations, correlations between tissue N concentrations and present-day total atmospheric N deposition, changes in plant amino-acid composition and effects on N assimilation. The ecological significance of such changes in biomarkers is uncertain. This paper explores the ecological significance of reactive atmospheric N deposition through a review of previous experimental findings and new experimental evidence from an acidic and a calcareous grassland, both showing phosphorus limitation, and a N-limited Calluna vulgaris (L.) Hull heathland in upland Britain. Nitrogen addition in the range 0–20 g N m−2 yr−1 initially (years 0–4) increased the growth of Calluna and a decline in some subordinate species. In subsequent years, shoot extension was not stimulated, but winter injury was observed from 1993 onwards, suggesting a strong interaction between N supply and climatic conditions. By contrast, the grasslands showed a small decrease in the cover of higher plants in later years (6–7) of the experimental treatments (0–14 g N m−2 yr−1) and no growth stimulation. All N treatments reduced the bryophyte cover in the acidic grassland. There were marked effects on below-ground processes, including a sustained stimulation of N mineralization in the grassland soils, and an increase in the bacterial utilization of organic substrates in the heathland, as measured in BIOLOG plates. The results strongly suggest the importance of atmospheric N deposition on microbially driven processes in soils, and are discussed in relation to the scale of potential ecosystem changes and their reversibility by pollution abatement.

Ring widths of five Mediterranean forest tree species (Arbutus unedo, Fraxinus ornus, Quercus cerris, Quercus ilex and Quercus pubescens) growing close to a natural source of CO2 in Tuscany, Italy and at a nearby control site were compared. At the CO2-enriched site, trees have been growing for decades under elevated CO2 concentrations. They originated from parent trees that also grew under elevated CO2 in natural conditions, and they have been continuously exposed to elevated CO2 throughout their growth. Tree-ring series from each of the species were prepared. Assigning calendar dates to rings was difficult but possible, and ring-width series were built for all species. The ring-width data were analysed using a two-sided t-test to assess if there was a difference between the radial growth at the CO2-enriched site and the control site. The cumulative basal area at the same cambial age at both sites was also compared using a Wilcoxon test. Radial growth of trees at the CO2-enriched site was not significantly different from growth at the control site. For each species, year by year, radial growth at the CO2-enriched site was tested against the control site and significant differences were found in only a few years; these differences were not synchronous with extreme climatic events. The expected increase in above-ground productivity, as one of the ecosystem responses to increasing CO2 during drought stress, was not observed in this Mediterranean woody plant community, despite being water-limited. Other resource limitations, such as low nutrient availability (common in the Mediterranean region), may have counteracted the positive effect of elevated CO2 under drought stress, or trees may have acclimated to the high CO2.

Regular (monthly) additions of NH4NO3 (4–12 g N m−2 yr−1) were made over a period of 8 yr (1989–98) to areas of moorland in North Wales dominated by the ericaceous shrub Calluna vulgaris. Results from the early stages of the experiment (1990–94) have shown marked and dose-related increases in shoot extension and canopy height in response to the nitrogen treatments, with significantly higher shoot nitrogen contents. The nitrogen-related stimulation in the growth of the C. vulgaris canopy over this period has resulted in large accumulations of litter on the high-nitrogen-treated plots (6.6 kg m−2 in plots treated with 12 g N m−2 yr−1, compared with 3.8 kg m−2 for the water controls). Litter nitrogen concentrations were also significantly increased at the higher rates of nitrogen addition, leading to a doubling of the total return of nitrogen to the litter layer over the experimental period. These changes in vegetation structure were associated with large reductions in the abundance of the bryophyte and lichen species normally present under the untreated canopy. Results since 1994, however, show little increase in shoot extension in response to the nitrogen treatments, with no clear dose response to increasing levels of addition. These findings are associated with a dose-related increase in the susceptibility of the nitrogen-treated areas of the C. vulgaris canopy to late winter injury, characterized as browning of the shoot tips in early to late spring. These results indicate that deleterious effects are now accumulating as a result of the long-term addition of nitrogen to these moorland plots.

Sloughing of root cap cells and exudation of mucilage plays an important role in the penetration of compacted soils by roots. For the first time we have quantified the rate of sloughing of root cap cells in an abrasive growth medium that was compacted to create mechanical impedance to root growth. The number of maize (Zea mays) root cap cells sloughed into sand increased as a result of compaction, from 1930 to 3220 d−1 per primary root. This represented a 12-fold increase in the number of cells sloughed per mm root extension (from 60 to >700). We estimated that the whole of the cap surface area was covered with detached cells in compacted sand, compared with c. 7% of the surface area in loose sand. This lubricating layer of sloughed cells and mucilage probably decreases frictional resistance to soil penetration. The total carbon deposited by the root was estimated at c. 110 μg g−1 sand d−1. Sloughed cells accounted for <10% of the total carbon, the vast majority of carbon being contained in mucilage exudates.

Very little is known about the physiological interactions between plant hosts and symptomless endophytic fungi despite their widespread occurrence. We investigated the impact of two such fungi, Colletotrichum musae and Fusarium moniliforme, upon the photosynthetic capacity of two crop plants, banana and maize, respectively. Endophyte-free plants were obtained first and then infected with the fungi. Measurements of total chlorophyll content revealed very little difference between endophyte-free and infected plants of banana, whereas in maize they showed 50% reductions in the endophyte-infected plants. The maximum photochemical capacity (Fv/Fm ) was measured in order to determine if the plants had any photoinhibitory effect caused by biotic or abiotic factors. After 45 d of growth, endophyte-free banana plants had similar values of Fv/Fm to plants typical of nonstressed conditions, whereas the endophyte-infected plants showed a reduction of approx. 15%. Unlike banana, infected maize plants displayed values of Fv/Fm similar to those of control and endophyte-free plants, indicating that the maximum photochemical capacity was not affected by infection. The light response curves of both species showed that the photosynthethic capacity was severely reduced in endophyte-infected plants, reaching saturation at c. 400 μmol m−2s−1 whereas the control and endophyte-free plants were saturated at much higher photon flux densities. In banana the effect seemed to be due to an impairment of electron transport in the thylakoid membranes. By contrast, reduction of the photosynthetic capacity in maize was due to a reduction in chlorophyll content, leading to a decrease in the electron transport components and a consequent reduction in carbohydrate synthesis. It is possible that the reduction in the maximum yield of photosynthesis in both crops was caused by toxins produced by the fungi. Nevertheless there were no major macroscopic effects on the plants to indicate disease symptoms.

The effects of iron plaque on the growth of Typha latifolia L. and accumulation of copper and nickel in T. latifolia were investigated under laboratory conditions in nutrient solution cultures. Seedlings with and without iron plaque on their roots were exposed to 0·05 μg ml−1 Cu and 0·10 μg ml−1 Ni solutions for 72 d. The results showed no differences in root and shoot d. wt and leaf elongation when Cu or Ni were added to the solution and in the presence or absence of plaque. However, root length was reduced by Cu and Ni, and the reduction in root length was greater in the presence of plaque. Some Cu and Ni was adsorbed on root surfaces; roots with plaque took up more Cu, but less Ni than those without. The presence of plaque did not alter Cu uptake and translocation but increased Ni uptake and translocation. Most of the Cu and Ni taken up was retained in the roots, suggesting that the root tissue rather than the root surface or plaque is the main barrier for Cu and Ni transport. The results differ from those reported for other species.

Growth of crops in CO2-enriched atmospheres typically results in significant changes in root growth and development. Increased root carbohydrates stimulate root growth either directly (functioning as substrates) or indirectly (functioning as signal molecules) by enhancing cell division or cell expansion, or both. Although highly variable, the literature suggests that, generally, initiation and stimulation of lateral roots is favored over the elongation of primary roots, leading to more highly branched, shallower root systems. Such architectural shifts can render root systems less efficient, perhaps contributing to the lower specific root activities often reported. Allocation of carbon (C) to roots fluctuates through the life of the plant; root functional and growth responses should therefore not be viewed as static. In annual crops, C allocation to belowground processes changes as vegetative growth switches to reproduction and maturation. Reductions in C allocation to roots over time might cause temporal shifts in root deployment, perhaps affecting root demography. However, significant changes in root turnover (defined here as root flux or mortality relative to total root pool size) as a result of decreased root longevities in crop plants are unlikely. Consideration of changing C allocation to roots, a more thorough understanding of the mechanistic controls on root longevity, and a better characterization of the rooting habits (life histories) of different crop species will further our understanding of how increasing atmospheric [CO2] will affect root demography. This knowledge will lead the way toward a more thorough understanding of the linkage of atmosphere with belowground plant function and also that of plant function with soil biology and structure. Ultimately, successful modeling of global C and nitrogen (N) cycles will require empirical data concerning spatial and temporal deployment of roots for a range of crop species grown under different agricultural management systems.

Concentrations of starch in roots of seeder species of Erica from the Cape Floristic Region, South Africa were found to be considerably less than in resprouters. Shoot starch was highly variable but mean values were similar in both seeder and resprouter species of Erica. Three distinct patterns of starch storage in roots were recognized. All seeder species fell within definitions of Categories 1 (narrow major and minor parenchymatous rays, one to two cells wide with no inter-ray storage) or 2 (thick major rays up to seven cells wide and thin minor rays with small amounts of inter-ray storage) whereas resprouter species were consistently within Categories 2 or 3 (broad major and minor rays, up to eight cells wide and conspicuous inter-ray starch storage). Results are discussed in light of similar studies of the related Epacridaceae. ‘Mixed’ species (i.e. with seeder or resprouter individuals present, often in distinct populations) were always classified as belonging to Category 2. Studies of populations of three ‘mixed’ species confirmed that seeder forms had consistently lower amounts of root starch than resprouters. Rays of xylem parenchyma were the main sites for starch storage in roots of both seeders and resprouters and greater proportions of cross-sectional area of roots were consistently devoted to such storage tissues in resprouter forms of the three ‘mixed’ species. Analyses of a number of seeder and resprouter Erica coccinea populations showed that differences in amounts of realised and potential root starch storage are best explained by the effect of regeneration behaviour rather than by among-population variability.

In many plants, mucilage permeates the interface between root and soil, yet little is known about its physical properties or its influence on the physical properties of the rhizosphere. Mucilage was collected from 3–4-d-old, axenically-grown maize (Zea mays L. cv. Freya) and lupin (Lupinus angustifolius L. cv. Merrit) seedlings. Surface tension and viscosity were measured over a range of mucilage hydration, and neutral sugar analyses of the hydrolysed mucilages were obtained by gas chromatography. Surface tension of both maize and lupin mucilage was reduced to ∼48 mN m−1 at total solute concentrations >0·7 mg ml−1, indicating the presence of powerful surfactants. Mucilage viscosity increased with increasing solute concentration and decreasing temperature. At a total solute concentration of 0·7 mg ml−1, the viscosity of maize mucilage at 20°C was 2·1 mPa s (approx. double that of pure water), increasing to 3·3 mPa s at 5°C. Both maize and lupin mucilage showed viscoelastic behaviour. The major component of maize mucilage was found to be glucose, but in lupin it was fucose.

The surface tension and viscosity results support the idea that mucilage plays a major role in the maintenance of root–soil contact in drying soils. As surface tension decreases, the ability of the mucilage to wet the surrounding soil particles becomes greater. Also, as viscosity and elasticity increase, the resistance to movement of any soil particles in contact with the mucilage increases, a degree of stabilization of the rhizosphere structure is achieved and hydraulic continuity is maintained. It is unclear whether the surfactant is actively secreted by the root or is present simply as a result of leakage from root cells.

Due to their different physiological effects, elevated carbon dioxide and elevated ozone might have interactive impacts on plants, and differentially so on plants differing in photosynthetic pathway and growth rate. To test several hypotheses related to these issues, we examined the physiological, morphological and growth responses of six perennial species grown at various atmospheric concentrations of carbon dioxide and ozone. The species involved (two C3 trees: Populus tremuloides Michx., Quercus rubra L.; two C3 grasses: Agropyron smithii Rybd., Koeleria cristata L.; two C4 grasses: Bouteloua curtipendula Michx., Schizachyrium scoparium Michx.) differed in growth form, stomatal conductance and photosynthetic pathway. In situ photosynthesis, relative growth rate (RGR) and its determinants (leaf area ratio, specific leaf area, leaf weight ratio and root weight ratio) were determined via sequential harvests of seedlings that were grown in all combinations of 366 or 672 μmol mol−1 CO2 and 3 or 95 nmol mol−1 O3 over a 101-d period. Elevated CO2 had minimal effect on either photosynthesis or RGR. By contrast, RGR for all six species was lower in high O3 concentrations at ambient CO2, significantly so in A. smithii and P. tremuloides. Five of the six species also exhibited reductions in in situ photosynthesis at ambient CO2 in high-O3-grown compared with low-O3-grown plants. For all species, these O3-induced reductions in RGR and photosynthesis were absent in the elevated CO2 environment. Root weight ratio was significantly reduced by elevated O3 in A. smithii and P. tremuloides in ambient but not elevated CO2. Species with high stomatal conductance were the most susceptible to oxidant injury, while those with low stomatal conductance, such as the C4 species and Q. rubra, were not as detrimentally affected by O3. Elevated levels of CO2 will reduce stomatal conductance and O3 uptake, and might therefore reduce the potential for oxidant damage. However, there was a stronger relationship of the percent reduction in whole-plant mass due to O3, related to the ratio of photosynthesis to stomatal conductance. In general, results of this study of six functionally diverse plant species suggest that O3 pollution effects on carbon balance and growth are likely to be ameliorated by elevated concentrations of atmospheric CO2.