The effects of low levels of nitrogen addition (7·7 and 15·4 kg N ha−1 yr−1) on plant sensitivity to biotic and abiotic stress were studied at a lowland heath in the south of England that has received N treatments since 1989. Larval growth rates and adult weights of heather beetles were found to be significantly higher when insects were reared on plants that had received additional N, with implications for insect survival and reproductive success. Electrolyte leakage measurements failed to reveal any significant impact of N addition on plant sensitivity to frost episodes in early winter. In April, however, there was some evidence of slightly decreased frost hardiness in plants receiving additional N. Accelerated spring bud burst also suggested earlier physiological activity in N-treated plots. The rate of water loss from excised shoots of Calluna vulgaris (L.) Hull was significantly faster in plants receiving additional N, although no difference in plant water potential was measured in the field after a prolonged dry spell. Whilst experimental N addition had only a small effect on plant sensitivity to abiotic stress, the relationship between enhanced deposition and increased insect performance was clear, with the potential for substantially increased insect damage at deposition rates around the critical load contributing to the formation of gaps in the Calluna canopy.

Fungal growth within reaction zones of beech (Fagus sylvatica) challenged by three basidiomycetes, Inonotus hispidus, Ganoderma adspersum, Fomitopsis pinicola, and one ascomycete, Ustulina deusta, was studied in naturally colonized and artificially inoculated wood. All the fungi, except F. pinicola, breached reaction zones, but the mechanisms involved were all somewhat different. Both I. hispidus and U. deusta bypassed blocked cell lumina by tunnelling through cell walls (soft-rot mode), but the latter caused far more decomposition of cell walls. Degradation of polyphenols was slight with I. hispidus and absent with U. deusta. By contrast, G. adspersum preferentially degraded the polyphenolic occlusions in the cell lumina. The failure of F. pinicola to invade reaction zones was typical of a brown rot fungus having limited enzymatic potential and a uniform growth pattern. Mechanisms of lesion expansion, illustrated and summarized in schematic diagrams, are consistent with earlier observations that reaction zones in beech sapwood are static boundaries, which may be successfully breached by white and soft-rot fungi.

The relative ozone sensitivities of 25 German native herbaceous plant species, representative of arable field margins or disturbed habitats, were examined over three consecutive growing seasons. Plants were grown from seed, potted into natural soils and exposed in open-top chambers for the entire season to different ozone-exposure regimes covering a range of concentrations from <5 to 48 ppb (seasonal 8 h daily mean). The assessment of ozone effects was carried out by recording the first day of visible symptom appearance and the percentage of injured leaves at the end of vegetative growth. Species exhibited contrasting patterns of symptom expression under ozone stress, with either ozone-specific symptoms or ozone-enhanced foliar pigmentation and senescence. Classifications of species according to their ozone susceptibility varied depending on whether measurement was of the total extent of visible injury, ozone threshold doses for the incidence of symptoms, or modelled exposure–response relationships. The most sensitive species exhibiting ozone-specific symptoms were Cirsium arvense and Sonchus asper, which responded to accumulated ozone exposures <1500 ppb.h (AOT40). For these and three other species, an AOT40 peak of a single day was found to be responsible for the incidence of ozone-specific symptoms, i.e. injury occurred rapidly within a few days of the day with the highest AOT40, while other species responded only to longer-term ozone exposures. The relative ozone sensitivity of the species was calculated by combining the different sensitivity criteria, and possible systematic trends (taxonomic or evolutionary features) are pointed out. The results suggest it may be possible to use a particular group of native herbaceous plant species with contrasting patterns of ozone sensitivity as a biomonitoring system in the field. This allows plant responses to be related either to peak values or to prolonged ozone exposure, making it possible to distinguish between short- and long-term effects of ozone.

For three decades, hypotheses relating to the occurrence and function of cyclic nucleotides in higher plants have been highly controversial. Although cyclic nucleotides had been shown to have key regulatory roles in animals and bacteria, investigations with higher plants in the 1970s and early 1980s were criticized on the basis of (i) a lack of specificity of effects apparently elicited by cyclic nucleotides, (ii) the equivocal identification of putative endogenous cyclic nucleotides and (iii) ambiguity in the identification of enzymes connected with cyclic nucleotide. More recent evidence based on more rigorous identification procedures has demonstrated conclusively the presence of cyclic nucleotides, nucleotidyl cyclases and cyclic nucleotide phosphodiesterases in higher plants, and has identified plant processes subject to regulation by cyclic nucleotides. Here we review the history of the debate, the recent evidence establishing the presence of these compounds and their role; future research objectives are discussed.

Mycorrhizal mycelium was allowed to grow into a compartment from which roots were excluded. Ammonium or nitrate was supplied exclusively to the fungus and the respiration rate before and after nitrogen addition was measured. Mycelial respiration in the fungal compartment represented 11–29% of total fungal and root respiration. The mycelium in the fungal compartment received 20–29% of shoot net assimilation, and 43–64% of the carbon allocated to the mycelium was respired. The nitrogen-induced respiration increase ranged, in the fungal compartment, from 54 to 180%.

Swards of Dactylis glomerata cultivars (cvs) KM2 and Lutetia and of Lolium perenne cvs Aurora and Vigor were grown under full irrigation or prolonged summer drought (80 d) in a field experiment in the South of France.

After irrigation was withheld, leaf extension rates of all cvs fell by 90% within 9–12 d, and rapid scorching of laminae followed. Tiller mortality at the end of the drought was very different in the cocksfoot cvs (4% for KM2 and 76% for Lutetia) and intermediate (41%) for both ryegrass cvs. Following re-watering, rates of herbage regrowth were closely correlated with tiller survival. Measured minerals contributed c. 0·52 MPa to osmotic potential in all treatments, whereas water-soluble carbohydrates (WSC) contributed 0·25 MPa under irrigation and 0·46 MPa during drought.

There was no systematic difference between the two species for summer survival under severe drought, but large differences between the cocksfoot cvs. The traits most strongly associated with superior survival were: (a) a deep root system and greater water uptake at depth; (b) low water and osmotic potentials in surviving laminae, i.e. better tolerance to dehydration; (c) large pool-size of WSC reserves (fructans having degree of polymerization >4) in entire tiller bases (stubble); (d) low accumulation of proline in stubble; (e) rapid nitrogen uptake after rewatering.

An early successional plant community was exposed to various ozone concentrations for two growing seasons (1994–1995) in open-top chambers in Auburn, Alabama, USA. The ozone treatments were: AA, ambient air (open plots); CF, carbon-filtered air (c. 0·5×ambient air), 1×, non-filtered air, and 2×, twice ambient air. Vegetative canopy cover exhibited a pattern of accumulation in the spring, with maximum canopy cover attained in summer, then senescence of foliage in the autumn 1994. This pattern was not observed in 1995 as a result of a drought during the spring and summer. Varying ozone exposures caused shifts in the competitive interactions between plants, thereby altering community structure. Higher canopy cover, vertical canopy density (layers of foliage), species richness, diversity, and evenness existed in the CF treatments than in the other treatments. In addition, winged sumac (Rhus copallina L.) became a major component of the CF treatments only during 1995. Surprisingly, blackberry (Rubus cuneifolius Pursh.), a species considered ozone-sensitive, based on visible injury, dominated canopy cover within the 2× treatments, 41 and 33% of total canopy cover in 1994 and 1995, respectively. From these results it is concluded that plant communities existing in areas where lower ozone concentrations are prevalent might be more complex and diverse than those existing in areas with higher ozone concentrations.

Birch (Betula pendula Roth) seedlings were grown under enhanced u.v.-B radiation and simulated forest-soil conditions, after which individual secondary metabolites were determined in the leaves. It was found that not all of the u.v.-absorbing secondary metabolites of the seedlings responded to supplementary u.v.-B radiation. Under increased u.v.-B radiation, significant increases in concentration were observed only for the major flavonoid, quercitrin, the minor flavonoid, myricetin-3-galactoside, and for chlorogenic acid. On the other hand, 3,4′dihydroxypropiophenone-3-β-D-glucopyranoside decreased under u.v.-B irradiation. The concentration of phenolic compounds in the leaves changed during the growing season (between two harvests) but this change was not related to u.v.-B enhancement. A low availability of mineral nutrients did not impair the capacity of the seedlings to accumulate u.v.-protecting phenolic compounds under increased u.v.-B radiation. The growth conditions used might have affected the intracellular concentrations of secondary metabolites, and thus furnished the birch seedlings with an increased tolerance of u.v.-B radiation. These findings point to the significance of certain phenolic components in the protection of deciduous trees against u.v.-B radiation.

Hakea psilorrhyncha R. M. Barker seedlings were subjected to a two-way interaction experiment with two levels of water availability and two levels of light. Physiological drought was imposed by adding 6000 MW polyethylene glycol (PEG), at an osmotic potential of −0·56 MPa, in a continuous-flow drip irrigation system. Unstressed plants (-PEG) were watered with distilled water (osmotic potential of −0·01 MPa). Seedlings were grown under natural light (120–500 μmol m−2 s−1 sunlight at midday +L) or shaded (<150 μmol m−2 s−1 sunlight at midday; −L. Plant morphology tissue water relations and carbon isotope composition (δ13C) were measured after 12 wk growth. The smallest leaves had the highest mass per unit area (LMA, an index of sclerophylly) and were present in the +PEG+L treatment, whereas the largest, thickest leaves had the lowest density and were produced in the −PEG+L treatment. Plants in shade were smaller and less sclerophyllous. There was a linear decrease in osmotic potential at full turgor and turgor loss point, and an increase in elastic modulus and δ13C, as level of sclerophylly increased, with −PEG−L leaves at one extreme and +PEG+L at the other. We conclude that high levels of sclerophylly induced by low water availability and high light intensity are associated with substantial drought tolerance in H. psilorrhyncha.

The effect of simultaneous N2 fixation and light limitation on the growth of two strains of Anabaena sp. Bory de St. Vincent and Aphanizomenon flos-aquae (L.) Ralfs was investigated using continuous cultures. Under severely light-limited conditions, Aphanizomenon showed a broader absorption spectrum (due to the presence of phycoerythrin), a higher maximum efficiency of photosynthesis, a higher steady-state N2 fixation activity and a higher growth affinity for light than did Anabaena. On the other hand, under light saturation, Anabaena showed a higher maximum rate of O2 production and a higher maximum specific growth rate than Aphanizomenon. These monoculture results characterize Anabaena and Aphanizomenon, in relative terms, as a ‘sun’ and a ‘shade’ species respectively, and are in accordance with field observations. The difference between the two species in their acclimatory response is discussed in terms of a species-specific alteration of the PSI[ratio ]PSII stoichiometry. Besides the species-specific modulation of the accessory pigments, such an acclimation would provide a biochemical basis for the observed physiological differences. The monoculture results were used to differentiate the niches of the two species and suggested that Aphanizomenon would competitively displace Anabaena under N2-fixing, light-limited conditions. However, when both species were grown together, Anabaena became dominant and seemed to be the superior competitor for light. In order to explain this finding, the possible effects of release of allelopathic compounds, or dynamic aspects of light supply, are discussed.

Well watered (WW) or drought-stressed (DS) saplings of ozone-sensitive and ozone-tolerant (less sensitive) birch (Betula pendula Roth) clones were exposed for 43 d to 0 nl l−1 or 100 nl l−1 ozone. Relative growth rates of leaves, stem, and roots, leaf discolouration, stomatal conductance and induction of genes encoding stress-related proteins PR-10, PAL and a LEA-group protein BP8 were determined. In general, both ozone and drought stress, singly and in combination, increased transcript levels of PR-10 in both clones. This was related to lower induction of PAL (except in older leaves of the tolerant clone), and increased proportions of visibly injured and yellowed leaves in ozone-exposed plants. The clones differed in their stomatal conductance and growth responses. In the less sensitive clone 2, ozone did not affect growth rates, but high stomatal conductance was observed in WW ozone-exposed plants. The more sensitive clone 5 showed, on the contrary, reduced growth rates and low stomatal conductance in WW ozone plants. Interestingly, clone 2 was sensitive to drought stress alone, whereas clone 5 was highly sensitive to ozone and drought stress experienced together. The results show that appearance of visible injuries (necrotic flecks) and enhanced yellowing of leaves coincided with the induction of genes for stress proteins PR-10 and PAL. The short-term growth responses, however, seemed to be separate processes. Additionally, stomatal conductance was related to leaf injuries and growth rates in a complicated manner, emphasizing the complex nature of ozone sensitivity/tolerance mechanisms in birch.

Cadmium accumulation and tolerance are discussed in a New Phytologist article by Krämer (2000), which comments on a paper by Lombi et al. (2000). In this context, a number of additional points should be made, putting the role of humans in the evolution of metal resistance into context and emphasizing what is the ‘hype’ of phytoremediation.

It is important that sites created by humans should not be overemphasized in considering the evolution of metal resistance. Plants resistant to heavy metals have their primary sites not on these, but on soils where ores are outcropping, the so-called metalliferous or orogenic soils (Ernst, 1974). Over thousands of years, natural exposure to a surplus of various metals, depending on the mineralization process, has driven the evolution of metal resistance in many plant species under the local environmental conditions. Many publications have shown that Thlaspi caerulescens can hyperaccumulate Zn (e.g. Vázquez et al., 1992), and accumulate other heavy metals such as Cu and Pb depending on soil chemistry (e.g. Baker et al., 1994). It has been known for more than 30 years that T. caerulescens gives a good response to experimentally supplied high Zn levels (Ernst, 1968). One of the ecological effects of hyperaccumulation of heavy metals is a defence against herbivorous insects (Boyd & Martens, 1994). This effect is enhanced by a preferential accumulation of heavy metals in the epidermal leaf layer (Heath et al., 1997).

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

Predictions of root architecture have been tested using only a narrow set of environmental conditions and species. There is little information on root architecture of plants growing on semi-arid grasslands where soil nutrient heterogeneity and defoliation might impose particular restrictions on root growth. We conducted a split-root study to evaluate the effect of nonuniform nutrient supply and clipping on root architecture and biomass allocation of three range grasses found in the Intermountain West of the USA: ‘Whitmar’, a cultivar of Pseudoroegneria spicata, Bromus tectorum (cheatgrass), and ‘Hycrest’, a hybrid between Agropyron desertorum and A. cristatum. Root systems of individual seedlings were equally divided between the two divisions of a partitioned 10-l pot, and each of the divisions was independently supplied with ‘high or ‘low’ nutrient solution. Plants were either clipped or not. Characteristics of root architecture (topology and link length), root size (root length and number of root tips), and biomass distribution (specific root length and root mass[ratio ]root volume ratio) were evaluated. Root topology (branching distribution) changed with nutrient treatments and defoliation in all three species. Length of links (a segment of root between two branches, or between a node and a root tip) responded to nutrient treatment only in defoliated plants of ‘Whitmar’, in which relative growth rate was associated with changes in architectural traits (external–external and internal–internal link length), while biomass accumulation and relative growth rate in cheatgrass and ‘Hycrest’ maintained allometric relationships with root length. Our results suggest that large plasticity in root architecture, together with inherently low relative growth rate are adaptations that allow ‘Whitmar’ to grow well in heterogeneous and poor soil-nutrient conditions. Cheatgrass and ‘Hycrest’ exhibited characteristics suggesting capability for rapid exploitation of nutrient patches.

Norway spruce (Picea abies (L.) Karst.) seedlings were inoculated with the ectomycorrhizal fungus Laccaria bicolor ((Marie) Orton), strain S238 N, in axenic conditions. The presence of the fungus slowed tap–root elongation by 26% during the first 15 d after inoculation and then stimulated it by 136%. In addition, it multiplied in vitro lateral root formation by 4.3, the epicotyl growth of the seedlings by 8.4 and the number of needles by 2. These effects were maintained when the fungus was separated from the roots by a cellophane membrane preventing symbiosis establishment, thus suggesting that the fungus acted by non-nutritional effects. We tested the hypothesis that IAA produced by L. bicolor S238 N would be responsible for the stimulation of fungal induced rhizogenesis. We showed in previous work that L. bicolor S238 N can synthesize IAA in pure culture. Exogenous IAA supplies (100 and 500 μm) reproduced the stimulating effect of the fungus on root branching but inhibited root elongation. The presence of 2,3,5-triiodobenzoic acid (TIBA) in the culture medium significantly depressed lateral root formation of inoculated seedlings. As TIBA had no significant effect on IAA released in the medium by L. bicolor S238 N, but counteracted the stimulation of lateral rhizogenesis induced by an exogenous supply of IAA, we suggest that TIBA inhibited the transport of fungal IAA in the root. Furthermore TIBA blocked the colonization of the main root cortex by L. bicolor S238 N and the formation of the Hartig net. These results specified the role of fungal IAA in the stimulation of lateral rhizogenesis and in ectomycorrhizal symbiosis establishment.

The identity of black alder (Alnus glutinosa (L.) Gaertn.) ectomycorrhizas was investigated using PCR/RFLP analysis of the ITS region from 16 morphotypes sampled at a 60-yr-old black alder stand. A comparison was made with restriction patterns from sporocarps of 28 mycobionts, of which 16 originated from the same stand, the remaining 12 came from two geographically distant alder stands. Eight of the mycorrhizal types could thus be identified, whereas eight mycorrhizal types remained unidentified. The identified mycorrhizas belonged to the genera Russula, Lactarius, Naucoria and Cortinarius. Four of the identified ectomycorrhizal types had identical PCR/RFLP profiles to corresponding fruit bodies from all investigated stands with no detectable intraspecific variation, despite the geographical distance of c. 300 km between the sampling locations. By contrast, intraspecific variation between sporocarps from the different locations was detected in Paxillus rubicundulus, mycorrhizas of which were not found. The diversity of fruiting alder mycobionts at the main experimental plot only partly matched the diversity observed from mycorrhizas when comparing their PCR/RFLP profiles. The results are discussed regarding sampling techniques, PCR/RFLP analyses and ecological aspects.

A total of 107 putative ericoid mycorrhizal endophytes were isolated from hair roots of Calluna vulgaris from two abandoned arsenic/copper mine sites and a natural heathland site in southwest England. The endophytes were initially grouped as 14 RFLP types, based on the results of ITS-RFLP analysis using the restriction endonucleases Hinf I, Rsa I and Hae III. ITS sequences were obtained for representative isolates from each RFLP type and compared phylogenetically with sequences for known ericoid mycorrhizal endophytes and selected ascomycetes. The majority of endophyte isolates (62–92%) from each site were identified as Hymenoscyphus ericae, but a number of other less common mycorrhizal RFLP types were also identified, all of which appear to have strong affinities with the order Leotiales. None of the less common RFLP types was isolated from C. vulgaris at more than one field site. Neighbour-joining analysis indicated similarities between the endophytes from C. vulgaris and mycorrhizal endophytes isolated from other Ericaceae and Epacridaceae hosts in North America and Australia.

Many hypotheses have been developed to explain the adaptive nature of insect galls. One of these, the nutrition hypothesis, states that gall formers have advantages over other insects because gall tissue provides a better (higher quality) food source than unmodified tissue. However, this has rarely been experimentally tested. In a test of this hypothesis, we grew plants of Cirsium arvense (L.) Scop. in a factorial design with two main treatments: the addition of nitrogen (to enhance foliar N levels) and of fungicide (to reduce colonization of roots by arbuscular mycorrhizal fungi). Mycorrhizal fungi have been shown previously to reduce the N concentration of host plants. Plants were exposed to adult gall flies, Urophora cardui L., and maintained through one season to allow maturation of galls.

Reduction of the percentage mycorrhizal colonization by fungicide resulted in an elevation of total stem N comparable to that achieved by N addition, but gall N concentration remained unchanged in all treatments. Nitrogen application elevated stem N levels when mycorrhizal fungi were present, but application of both compounds together did not result in any increase over either single treatment. Fungicide application resulted in larger galls, which contained more larval chambers, with more live, and heavier, larvae. However, the main effects of N were not significant, as N addition only increased fly performance on plants where mycorrhizas were not reduced.

It is suggested that U. cardui gall inhabitants can manipulate N at an optimal level and thus might conform to a modified version of the nutrition hypothesis. Mycorrhizal colonization might reduce gall fly performance by delaying the appearance, or impairing the quality, of secondary nutritive tissue in the gall. Future tests of the nutrition hypothesis should include a consideration of the plant's mycorrhizal status.

Xylem maturation in elongating leaf blades of tall fescue (Festuca arundinacea) was studied using staining and microcasting. Three distinctive regions were identified in the blade: (1) a basal region, in which elongation was occurring and protoxylem (PX) vessels were functioning throughout; (2) a maturation region, in which elongation had stopped and narrow (NMX) and large (LMX) metaxylem vessels were beginning to function; (3) a distal, mature region in which most of the longitudinal water movements occurred in the LMX. The axial hydraulic conductivity (Kh) was measured in leaf sections from all these regions and compared with the theoretical axial hydraulic conductivity (Kt) computed from the diameter of individual inner vessels. Kt was proportional to Kh throughout the leaf, but Kt was about three times Kh. The changes in Kh and Kt along the leaf reflected the different stages of xylem maturation. In the basal 60 mm region, Kh was about 0.30±0.07 mmol s−1 mm MPa−1. Beyond that region, Kh rapidly increased with metaxylem element maturation to a maximum value of 5.0±0.3 mmol s−1 mm MPa−1, 105 mm from the leaf base. It then decreased to 3.5±0.2 mmol s−1 mm MPa−1 near the leaf tip. The basal expanding region was observed to restrict longitudinal water movement. There was a close relationship between the water deposition rate in the elongation zone and the sum of the perimeters of PX vessels. The implications of this longitudinal vasculature on the partitioning of water between growth and transpiration is discussed.

Traits associated with leaf energy balance are especially important for desert plants because decoupling leaf and ambient temperatures can be critical for maximizing productivity and survival in these hot, dry regions. Deserts are also one of the most climatically unpredictable and heterogeneous biomes; thus, variation of energy balance characters due either to plasticity or genetic differentiation might also be crucial to the success of desert plant species. In a common garden environment we examined variation in the traits associated with energy balance and productivity in Encelia farinosa A. Gray (brittlebush) plants. Comparing two populations from contrasting rainfall and drought climates, we found that plants from the wetter site (Superior, Arizona) always maintained higher leaf absorptances than plants from the more xeric region (Oatman, Arizona) when at similar water potentials. Superior plants also increased stomatal conductance and photosynthesis in response to mid-spring rainfall, whereas Oatman plants did not. Oatman plants, however, tended to have greater leaf areas and continuous spring growth, which made them larger than Superior plants, yet both populations produced the same number of flower heads. The differences for these traits, and the associations among them, agree with predictions based on the contrasting drought and rainfall environments of these two populations. Our results suggest that the differences might represent alternative suites of characters of selective importance for maximizing carbon gain over the climatically variable range of this widespread species. Furthermore, these differences, detected in common environment conditions, might have a genetic basis, which might contribute to a greater potential for local adaptation in this species.