Tropical montane forests (TMF) are associated with a widely observed suite of characteristics encompassing forest structure, plant traits and biogeochemistry. With respect to nutrient relations, montane forests are characterized by slow decomposition of organic matter, high investment in below-ground biomass and poor litter quality, relative to tropical lowland forests. However, within TMF there is considerable variation in substrate age, parent material, disturbance and species composition. Here we emphasize that many TMFs are likely to be co-limited by multiple nutrients, and that feedback among soil properties, species traits, microbial communities and environmental conditions drive forest productivity and soil carbon storage. To date, studies of the biogeochemistry of montane forests have been restricted to a few, mostly neotropical, sites and focused mainly on trees while ignoring mycorrhizas, epiphytes and microbial community structure. Incorporating the geographic, environmental and biotic variability in TMF will lead to a greater recognition of plant–soil feedbacks that are critical to understanding constraints on productivity, both under present conditions and under future climate, nitrogen-deposition and land-use scenarios.

Relations between soil nutrient availability, the occurrence of root symbioses and tree species composition in tropical Africa are discussed in the light of new evidence. Transects from humid areas at the equator towards the deserts can be summarized as follows:

High availability of nitrogen, low availability of phosphorus, low pH and aluminium toxicity probably restrict the competitiveness of nitrogen-fixing species in the forests on highly weathered and leached soils in humid west and central Africa. Ectomycorrhizal species attain dominance in species-poor forests on the soils poorest in phosphorus, while species with vesicular-arbuscular (VA) mycorrhiza form diverse forests on more fertile soils. Nitrogen-fixing species, which mostly are VA mycorrhizal, are more common in woodlands, where nitrogen is lost in grass fires, than in forests, which have a more closed nitrogen cycle. Low availability of phosphorus, however, limits the growth of nitrogen-fixing species in moist savannas. Ectomycorrhizal species are dominant, possibly because of a competitive advantage conferred by the large storage capacity of the fungal mantles under conditions of discontinuous nutrient supply. Nitrogen-fixing species become more abundant towards drier areas with an increase in soil phosphorus and a decline in soil nitrogen. These conditions culminate at the desert fringe, where nitrogen-fixing species reach their most conspicuous development.

It is concluded that soil nutrient availability and root symbioses interact and strongly affect the composition of vegetation communities.

Root biomass, root lengths, and mycorrhizal associations were compared in a series of primary and Acacia-dominated secondary rainforest stands on nutrient-poor, red podzolic soils developed from low grade Palaeozoic metasediments. Five soil cores to 200 mm depth were collected at random locations from each of 20 sites. Ten of these sites were in 20–25 m high closed secondary forest (30–40 y old) dominated by Acacia aulacocarpa and ten sites were located in primary, selectively-logged, rainforest (28–32 m tall). Arbuscular mycorrhizas were the only form of association found in the primary forest sites. Ectomycorrhizas dominated the secondary forest sites although arbuscular mycorrhizas were also present. The primary forest sites had significantly higher root biomass (34.4 ± 17.8 t ha-1) and root length (33,400 ± 3,200 km ha-1) than the secondary forests (11.6 ± 4.6 t ha-1 and 25,200 ± 4,800 km ha-1 respectively), and this was interpreted as a reflection of the greater allocation of biomass to roots necessary to support the greater above ground biomass. The specific root length in the secondary forest (340 ± 119 cm g-1) was twice that of the primary forest (154 ± 65 cm g-1) indicating that the trees in the secondary forests achieved a degree of soil exploration which was comparable to that in the primary forest with less than half the biomass allocation to roots. The dominance of ectomycorrhizas in the secondary forest was associated with the prevalence of Acacia aulacocarpa, and the results cannot be extended to other secondary forests in the region. The implications that the dominant ectomycorrhizal associations have for the patterns of successional development and the patterns of species colonization in these Acaria-dominated secondary forests are discussed.

Changes in the production and turnover of roots in forests and grasslands in response to rising atmospheric CO2 concentrations, elevated temperatures, altered precipitation, or nitrogen deposition could be a key link between plant responses and longer-term changes in soil organic matter and ecosystem carbon balance. Here we summarize the experimental observations, ideas, and new hypotheses developed in this area in the rest of this volume. Three central questions are posed. Do elevated atmospheric CO2, nitrogen deposition, and climatic change alter the dynamics of root production and mortality? What are the consequences of root responses to plant physiological processes? What are the implications of root dynamics to soil microbial communities and the fate of carbon in soil? Ecosystem-level observations of root production and mortality in response to global change parameters are just starting to emerge. The challenge to root biologists is to overcome the profound methodological and analytical problems and assemble a more comprehensive data set with sufficient ancillary data that differences between ecosystems can be explained. The assemblage of information reported herein on global patterns of root turnover, basic root biology that controls responses to environmental variables, and new observations of root and associated microbial responses to atmospheric and climatic change helps to sharpen our questions and stimulate new research approaches. New hypotheses have been developed to explain why responses of root turnover might differ in contrasting systems, how carbon allocation to roots is controlled, and how species differences in root chemistry might explain the ultimate fate of carbon in soil. These hypotheses and the enthusiasm for pursuing them are based on the firm belief that a deeper understanding of root dynamics is critical to describing the integrated response of ecosystems to global change.

Functional morphological patterns in root apices of tomato (Lycopersicon esculentum) dependent on growth, ageing and infection by the arbuscular mycorrhizal (AM) fungus Glomus mosseae and/or by the soilborne pathogenic fungus Phytophthora nicotianae var parasitica (P. parasitica) were studied. Uninfected root apices were characterized by closed, tri-layered meristems with nonreticulate nuclei; however, some apices of each treatment lost their meristematic nature, stopped growing and differentiated, becoming ‘parenchymatized’. The pathogenic fungus reduced the apex diameter and the number of mitotically active and viable apices inducing plasmolysis, cell and nucleus degeneration, and necrosis. The AM fungus, on the other hand, produced an increase in apex size and reduced the percentage of necrosis both in uninfected roots and in roots infected by P. parasitica. Thus, the AM fungus protected the apices from the pathogenic infection, allowing normal root growth. Furthermore, larger apices, which produce thicker roots, might indirectly contribute to plant protection. Increased volumes of colonizable tissues favour the spreading of the symbiont, and P. parasitica hyphae are always excluded from arbuscule-containing cells.

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

Dark septate root endophytes (DSE) are conidial or sterile fungi (Deuteromycotina, Fungi Imperfecti) likely to be ascomycetous and colonizing plant roots. They have been reported for nearly 600 plant species representing about 320 genera and 100 families. DSE fungi occur from the tropics to arctic and alpine habitats and comprise a heterogeneous group that functionally and ecologically overlaps with soil fungi, saprotrophic rhizoplane-inhabiting fungi, obligately and facultatively pathogenic fungi and mycorrhizal fungi. Numerous species of undescribed sterile and anamorphic taxa may also await discovery. Although DSE are abundant in washed root and soil samples from various habitats, and are easily isolated from surface-sterilized roots of ecto-, ectendo-, endo- and non-mycorrhizal host species, their ecological functions are little understood. Studies of DSE thus far have yielded inconsistent results and only poorly illustrate the role of DSE in their natural habitats. These inconsistencies are largely due to the uncertain taxonomic affinities of the strains of DSE used. In addition, because different strains of a single anamorph taxon seem to vary greatly in function, no clear generalizations on their ecological role have been drawn. This paper reviews the current literature on DSE and the ecology and discusses the need for and direction of future research.

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.

One hundred and sixty-eight sterile endophytic mycelia were isolated from roots of four Woollsia pungens (Cav.) F. Muell. (Epacridaceae) plants collected from a field site in New South Wales, Australia. All isolates formed typical ericoid mycorrhizal structures when inoculated onto roots of Vaccinium macrocarpon Ait. (Ericaceae). Microsatellite-primed PCR fingerprints generated using the primers (GTG)5 and (GACA)4 indicated that considerable genetic diversity exists within the endophyte population. It was estimated that a minimum of 43 genetically distinct mycelial genets were present in the root systems of the sampled W. pungens population, with most genets confined to individual plants. Two genets, however, were present within the root systems of two adjacent plants. While most genets were represented by less than eight isolates, three genets contained up to 41 isolates, suggesting that root system colonization by some endophytic mycelia might be extensive.

Several temporal and spatial scales have been considered during the Symposium, and the presentations by Wallenda & Kottke (1998) and by Norby (1998) represent two extreme points in a wide range. They both deal with short and long-term impacts on the local as well as the global scale. I present here a few additional thoughts, and a subject that both papers have touched only slightly will be expanded: the differential impact of different forms of nitrogen deposition.

Seedlings of Betula pendula Roth. were grown in the presence of Paxillus involutus (Batsch) Fr., and metabolic changes during mycorrhiza formation were examined by measuring organic acid and amino acid pools and related enzyme activities, following sequential harvests. Glutamine, aspartate and asparagine pools were always lower in infected roots than in non-infected roots, especially during Hartig net initiation and formation. Glutamate concentration was similar in both tissues. Citrate and malate were the two major organic acids detected and their concentrations were equal in infected and non-infected roots. Aspartate aminotransferase, glutamine synthetase, NAD-dependent malate dehydrogenase and glucose-6-phosphate dehydrogenase activities were higher in infected roots than non-infected roots. For all enzymes revealed on polyacrylamide gels, both root and fungal isoforms were present in infected roots. Quantitative changes in enzyme capacities and metabolite pools indicated that mycorrhiza formation caused a re-arrangement of the main metabolic pathways during the very early stages following contact, which might be related to the structural changes.