1 Plant ecology at high elevations.- The concept of limitation.- A regional and historical account.- The challenge of alpine plant research.- 2 The alpine life zone.- Altitudinal boundaries.- Global alpine land area.- Alpine plant diversity.- Origin of alpine floras.- Alpine growth forms.- 3 Alpine climate.- Which alpine climate.- Common features of alpine climates.- Regional features of alpine climates.- 4 The climate plants experience.- Interactions of relief, wind and sun.- How alpine plants influence their climate.- The geographic variation of alpine climate.- 5 Life under snow: protection and limitation.- Temperatures under snow.- Solar radiation under snow.- Gas concentrations under snow.- Plant responses to snowpack.- 6 Alpine soils.- Physics of alpine soil formation.- The organic compound.- The interaction of organic and inorganic compounds.- 7 Alpine treelines.- About trees and lines.- Current altitudinal positions of climatic treelines.- Treeline-climate relationships.- Intrazonal variations and pantropical plateauing of alpine treelines.- Treelines in the past.- Attempts at a functional explanation of treelines.- A hypothesis for treeline formation.- Growth trends near treelines.- Evidence for sink limitation.- 8 Climatic stress.- Survival of low temperature extremes.- Avoidance and tolerance of low temperature extremes.- Heat stress in alpine plants.- Ultraviolet radiation - a stress factor.- 9 Water relations.- Ecosystem water balance.- Soil moisture at high altitudes.- Plant water relations - a brief review of principles.- Water relations of alpine plants.- Desiccation stress.- Water relations of special plant types.- 10 Mineral nutrition.- Soil nutrients.- The nutrient status of alpine plants.- Nutrient cycling and nutrient budgets.- Nitrogen fixation.- Mycorrhiza.- Responses of vegetation to variable nutrient supply.- 11 Uptake and loss of carbon.- Photosynthetic capacity of alpine plants.- Photosynthetic responses to the environment.- Daily carbon gain of leaves.- The seasonal carbon gain of leaves.- C4 and CAM photosynthesis at high altitudes.- Tissue respiration of alpine plants.- Ecosystem carbon balance.- 12 Carbon investments.- Non-structural carbohydrates.- Lipids and energy content.- Carbon costs of leaves and roots.- Whole plant carbon allocation.- 13 Growth dynamics and phenology.- Seasonal growth.- Diurnal leaf extension.- Rates of plant dry matter accumulation.- Functional duration of leaves and roots.- 14 Cell division and tissue formation.- Cell size and plant size.- Mitosis and the cell cycle.- From meristem activity to growth control.- 15 Plant biomass production.- The structure of alpine plant canopies.- Primary productivity of alpine vegetation.- Plant dry matter pools.- Biomass losses through herbivores.- 16 Plant reproduction.- Flowering and pollination.- Seed development and seed size.- Germination.- Alpine seed banks and natural recruitment.- Clonal propagation.- Alpine plant age.- Community processes.- 17 Global change at high elevation.- Alpine land use.- The impact of altered atmospheric chemistry.- Climatic change and alpine ecosystems.- References (with chapter annotation).- Taxonomic index (genera).- Geographical index.- Color plates.- Plant life forms.- The alpine life zone.- Environmental stress.- The human dimension.

Alpine plant life

The rhizosphere is the interface between plant roots and soil where interactions among a myriad of microorganisms and invertebrates affect biogeochemical cycling, plant growth and tolerance to biotic and abiotic stress. The rhizosphere is intriguingly complex and dynamic, and understanding its ecology and evolution is key to enhancing plant productivity and ecosystem functioning. Novel insights into key factors and evolutionary processes shaping the rhizosphere microbiome will greatly benefit from integrating reductionist and systems-based approaches in both agricultural and natural ecosystems. Here, we discuss recent developments in rhizosphere research in relation to assessing the contribution of the micro- and macroflora to sustainable agriculture, nature conservation, the development of bio-energy crops and the mitigation of climate change.

Going back to the roots: the microbial ecology of the rhizosphere.

https://cbs.umn.edu/sites/cbs.umn.edu/files/public/downloads/Nguyenetal2015b.pdf

FUNGuild: An open annotation tool for parsing fungal community datasets by ecological guild

Evidence is mounting that the immense diversity of microorganisms and animals that live belowground contributes significantly to shaping aboveground biodiversity and the functioning of terrestrial ecosystems. Our understanding of how this belowground biodiversity is distributed, and how it regulates the structure and functioning of terrestrial ecosystems, is rapidly growing. Evidence also points to soil biodiversity as having a key role in determining the ecological and evolutionary responses of terrestrial ecosystems to current and future environmental change. Here we review recent progress and propose avenues for further research in this field.

Belowground biodiversity and ecosystem functioning

Progress towards understanding the extent to which mycorrhizal fungi are involved in the mobilization of nitrogen (N) and phosphorus (P) from natural substrates is reviewed here. While mycorrhiza research has emphasized the role of the symbiosis in facilitation of capture of these nutrients in ionic form, attention has shifted since the mid-1980s to analysing the mycorrhizal fungal abilities to release N and P from the detrital materials of microbial faunal and plant origins, which are the primary sources of these elements in terrestrial ecosystems. Ericoid, and some ectomycorrhizal fungi have the potential to be directly involved in attack both on structural polymers, which may render nutrients inaccessible, and in mobilization of N and P from the organic polymers in which they are sequestered. The advantages to the plant of achieving intervention in the microbial mobilization-immobilization cycles are stressed. While the new approaches may initially lack the precision achieved in studies of readily characterized ionic forms of N and P, they do provide insights of greater ecological relevance. The results support the hypothesis that selection has favoured ericoid and ectomycorrhizal systems with well developed saprotrophic capabilities in those ecosystems characterized by retention of N and P as organic complexes in the soil. The need for further investigation of the abilities of arbuscular mycorrhizal fungi to intervene in nutrient mobilization processes is stressed.

https://nph.onlinelibrary.wiley.com/doi/pdf/10.1046/j.1469-8137.2003.00704.x

Mycorrhizas and nutrient cycling in ecosystems – a journey towards relevance?

This study was conducted to evaluate the effects of wildfires on ectomycorrhizal (EM) fungal communities in Scots pine (Pinus sylvestris) stands. Below- and above-ground communities were analysed in terms of species richness and evenness by examining mycorrhizas and sporocarps in a chronosequence of burned stands in comparison with adjacent unburned late-successional stands. The internal transcribed spacer (ITS)-region (rDNA) of mycobionts from single mycorrhizas was digested with three restriction enzymes and compared with an ITS–restriction fragment length polymorphism (RFLP) reference database of EM sporocarps. Spatial variation seemed to be more prominent than the effects of fire on the EM fungal species composition. Most of the common species tended to be found in all sites, suggesting that EM fungal communities show a high degree of continuity following low-intensity wildfires. Species richness was not affected by fire, whereas the evenness of species distributions of mycorrhizas was lower in the burned stands. The diversity of EM fungi was relatively high considering that there were only three EM tree species present in the stands. In total, 135 EM taxa were identified on the basis of their RFLP patterns; 66 species were recorded as sporocarps, but only 11 of these were also recorded as mycorrhizas. The species composition of the below-ground community of EM fungi did not reflect that of the sporocarps produced. EM fungal species present in our ITS–RFLP reference database accounted for 54–99% of the total sporocarp production in the stands, but only 0–32% of the mycorrhizal abundance.

Ectomycorrhizal fungal communities in late‐successional Swedish boreal forests, and their composition following wildfire

SUMMARY The spatial distribution of genets in the ectomycorrhizal fungus Suillus bovinus (L.: Fr.) Roussel were studied in somatic incompatibility pairings of isolates from five Scots pine (Pinus sylvestris L.) stands differing in forest history and age. With increasing forest age, the size of genets increased while the number of genets and production of sporocarps per unit area decreased. There was an estimated 700-5700 genets ha-1 in younger forests and 30-120 ha-1 in older ones. The maximum size of genets was 1-7-5-3 m in the younger forest and to 17-5 m in the older ones. The production of sporocarps per unit area decreased with increasing forest age. Furthermore, production of sporocarps increased nonlinearly with the size of the genet, suggesting that genets become fragmented. Ergosterol measurements indicated that the fungal biomass of one genet consisted of 20-45 % sporocarps and 55-80 % mycorrhiza, not including extramatrical mycelia. Mycorrhizal aggregations in soil were mapped in two 5 m2 areas and, based on somatic incompatibility tests, all were found to belong to the same genet as sporocarps present above ground. Production of spores per sporocarp was estimated to be 1 1-12 8 x 108. The observations are discussed in terms of population ecology.

https://nph.onlinelibrary.wiley.com/doi/pdf/10.1111/j.1469-8137.1994.tb04006.x

Size, distribution and biomass of genets in populations of Suillus bovinus (L.: Fr) Roussel revealed by somatic incompatibility.

SUMMARY
Current knowledge of mycorrhizal diversity in arctic and alpine tundra is based mainly on static surveys of mycorrhizal associations of plant taxa and fruiting patterns of selected ectomycorrhizal fungal species in specific habitats. Within these limitations, it appears that: (1) non-mycorrhizal plants are widespread and predominate in certain plant communities; (2) typical arbuscular mycorrhizal associations are ubiquitous in low arctic and alpine areas but that the level of root colonization is highly variable; (3) root colonization by dark septate fungi is a common event but that their ecological significance is still unknown; (4) a large number of ectomycorrhizal fungal species are present as symbionts of a relatively few widely distributed shrubs and herbaceous plant taxa; (5) ericaceous plants with erieoid mycorrhizas dominate large arctic and alpine areas covered by heath communities. Physical environmental features strongly limit and shape species diversity in arctic and alpine tundra. Cold-dominated environments provide extreme conditions for the establishment and functioning of mycorrhizal associations. Therefore, such systems are simple models to address the ecology and evolution of mycorrhizal symbioses. Molecular methods to identify mycorrhizal fungi on plant roots will resolve questions related to the structure and dynamics of communities of mycorrhizal fungi in arctic and alpine tundra.

Mycorrhizal diversity in arctic and alpine tundra: an open question

Resupinate thelephoroid fungi (hereafter called tomentelloid fungi) have a world-wide distribution and comprise approximately 70 basidiomycete species with inconspicuous, resupinate sporocarps. It is only recently that their ability to form ectomycorrhizas (EM) has been realized, so their distribution, abundance and significance as mycobionts in forest ecosystems is still largely unexplored. In order to provide baseline data for future ecological studies of tomentelloid fungi, we explored their presence and abundance in nine Swedish boreal forests in which the EM communities had been analysed. Phylogenetic analyses were used to compare the internal transcribed spacer of nuclear ribosomal DNA (ITS rDNA) sequence data obtained from mycobionts on single ectomycorrhizal tips with that obtained from sporocarps of identified tomentelloid fungi. Five species of Tomentella and one species of Pseudotomentella were identified as ectomycorrhizal fungi. The symbiotic nature of Tomentella bryophila, T. stuposa, T. badia and T. atramentaria is demonstrated for the first time. T. stuposa and Pseudotomentella tristis were the most commonly encountered tomentelloid fungi, with the other species, including T. sublilacina, only being recorded from single stands. Overall, tomentelloid fungi were found in five of the studies, colonizing between 1 and 8% of the mycorrhizal root tips. Two of the five sites supported several tomentelloid species. Tomentelloid fungi appear to be relatively common ectomycorrhizal symbionts with a wide distribution in Swedish coniferous forests. The results are in accordance with accumulating data that fungal species which lack conspicuous sporocarps may be of considerable importance in EM communities.

Diversity and abundance of resupinate thelephoroid fungi as ectomycorrhizal symbionts in Swedish boreal forests.

Owing to previous methodological limitations, knowledge about the fine-scale distribution of fungal mycelia in decaying logs is limited. We investigated fungal communities in decaying Norway spruce logs at various spatial scales at two environmentally different locations in Sweden. On the basis of 454 pyrosequencing of the ITS2 region of rDNA, 1914 operational taxonomic units (OTUs) were detected in 353 samples. The communities differed significantly among logs, but the physical distance between logs was not found to have a significant effect on whether fungal communities had any resemblance to each other. Within a log, samples that were closer together generally had communities that showed more resemblance to each other than those that were further apart. OTUs characteristic for particular positions on the logs could be identified. In general, these OTUs did not overlap with the most abundant OTUs, and their ecological role was often unknown. Only a few OTUs were detected in the majority of logs, whereas numerous OTUs were rare and present in only one or a few logs. Wood-decaying Basidiomycetes were often represented by higher sequence reads in individual logs than Ascomycete OTUs, suggesting that Basidiomycete mycelia spread out more rapidly when established. OTU richness tended to increase with the decay stage of the sample; however, the known wood decayers were most abundant in less-decomposed samples. The fungi identified in the logs represented different ecological strategies. Our findings differ from previously published sporocarp studies, indicating that the highly abundant fruiting species may respond to environment in different ways than the rest of the fungal community.

Anders Dahlberg

Papers

Ectomycorrhizal fungal communities in late‐successional Swedish boreal forests, and their composition following wildfire

Size, distribution and biomass of genets in populations of Suillus bovinus (L.: Fr) Roussel revealed by somatic incompatibility.

Mycorrhizal diversity in arctic and alpine tundra: an open question

Diversity and abundance of resupinate thelephoroid fungi as ectomycorrhizal symbionts in Swedish boreal forests.

Patterns of fungal communities among and within decaying logs, revealed by 454 sequencing