Soil Chemical Analysis

Almost all land plants form symbiotic associations with mycorrhizal fungi. These below-ground fungi play a key role in terrestrial ecosystems as they regulate nutrient and carbon cycles, and influence soil structure and ecosystem multifunctionality. Up to 80% of plant N and P is provided by mycorrhizal fungi and many plant species depend on these symbionts for growth and survival. Estimates suggest that there are c. 50 000 fungal species that form mycorrhizal associations with c. 250 000 plant species. The development of high-throughput molecular tools has helped us to better understand the biology, evolution, and biodiversity of mycorrhizal associations. Nuclear genome assemblies and gene annotations of 33 mycorrhizal fungal species are now available providing fascinating opportunities to deepen our understanding of the mycorrhizal lifestyle, the metabolic capabilities of these plant symbionts, the molecular dialogue between symbionts, and evolutionary adaptations across a range of mycorrhizal associations. Large-scale molecular surveys have provided novel insights into the diversity, spatial and temporal dynamics of mycorrhizal fungal communities. At the ecological level, network theory makes it possible to analyze interactions between plant-fungal partners as complex underground multi-species networks. Our analysis suggests that nestedness, modularity and specificity of mycorrhizal networks vary and depend on mycorrhizal type. Mechanistic models explaining partner choice, resource exchange, and coevolution in mycorrhizal associations have been developed and are being tested. This review ends with major frontiers for further research.

https://nph.onlinelibrary.wiley.com/doi/epdf/10.1111/nph.13288

Mycorrhizal ecology and evolution: the past, the present, and the future

A comprehensive appraisal of the mycorrhizal literature provides data for 336 plant families representing 99% of flowering plants, with regard to mycorrhizas and other nutritional adaptations. In total, arbuscular (AM), orchid, ectomycorrhizas (EM) and ericoid mycorrhizas and nonmycorrhizal (NM) roots occur in 74%, 9%, 2%, 1% and 6% of Angiosperm species respectively. Many families of NM plants have alternative nutritional strategies such as parasitism, carnivory, or cluster roots. The remaining angiosperms (8%) belong to families reported to have both AM and NM species. These are designated as NM-AM families here and tend to occur in habitats considered non-conducive to mycorrhizal fungi, such as epiphytic, aquatic, extremely cold, dry, disturbed, or saline habitats. Estimated numbers of species in each category of mycorrhizas is presented with lists of NM and EM families. Evolutionary trends are also summarised by providing data on all clades and orders of flowering and non-flowering vascular plants on a global scale. A case study of Western Australian plants revealed that plants with specialised nutritional modes such as carnivory, cluster roots, or EM were much more diverse in this ancient landscape with infertile soils than elsewhere. Detailed information on the mycorrhizal diversity of plants presented here is linked to a website (mycorrhizas.info) to allow data to remain current. Over a century of research effort has resulted in data on mycorrhizal associations of >10,000 plant species that are of great value, but also somewhat of a liability due to conflicting information about some families and genera. It is likely that these conflicts result in part from misdiagnosis of mycorrhizal associations resulting from a lack of standardisation in criteria used to define them. Families that contain both NM and AM species provide a second major source of inconsistency, but even when these are excluded there is a ∼10% apparent error rate in published lists of mycorrhizal plants. Arbuscules are linked to AM misdiagnosis since they are used less often than vesicles to recognise AM associations in roots and apparently occur sporadically in NM plants. Key issues with the diagnosis of mycorrhizal plants are discussed using the Cyperaceae as a case study. Detailed protocols designed to consistently distinguish AM from endophytic Glomeromycotan Fungus Colonisation (GFC) are provided. This review aims to stimulate debate and provide advice to researchers delving into root biology.

Mycorrhizal associations and other means of nutrition of vascular plants: understanding the global diversity of host plants by resolving conflicting information and developing reliable means of diagnosis

Heavy metals in soils

The role of mycorrhizae and plant growth promoting rhizobacteria (PGPR) in improving crop productivity under stressful environments.

Chenopods are generally regarded as non-host plants for mycorrhizal fungi and are believed not to benefit from colonization by mycorrhizal fungi. Perennial Atriplex nummularia Lindl., growing under field conditions, showed a relatively high level of colonization by mycorrhizal fungi (10–30% of root length colonized) in spring and summer. Accordingly, two glasshouse experiments were designed to assess the effects of inoculation with mycorrhizal fungi (with a single species or a mixture of different species) on growth, nutrient uptake, and rhizosphere bacterial community composition of A. nummularia at high and low salinity levels (2.2 and 12 dSm−1). Only low and patchy colonization by mycorrhizal fungi (1–2 of root length colonized) was detected in inoculated plants under glasshouse conditions which was unaffected by salinity. Despite the low colonization, inoculation increased plant growth and affected nutrient uptake at both salinity levels. The effects were higher at an early stage of plant development (6 weeks) than at a later stage (9–10 weeks). Salinity affected the bacterial community composition in the rhizosphere as examined by ribosomal intergenic spacer amplification (RISA) of 16S rDNA, digitization of the band patterns and multivariate analysis. The effects of inoculation with mycorrhizal fungi on growth of A. nummularia may be attributed to (i) direct effects of mycorrhizal fungi on plant nutrient uptake and/or (ii) indirect effects via mycorrhizal-induced changes in the bacterial community composition.

Growth response of Atriplex nummularia to inoculation with arbuscular mycorrhizal fungi at different salinity levels

Arbuscular mycorrhizal fungi (AMF) can increase plant growth and nutrition. However, their capacity to reduce the leaching of nutrients through the soil profile is less well understood. Here we present results of an experiment in which the effects of forming arbuscular mycorrhizas (AM) on plant growth and nutrition, nutrient depletion from soil, and nutrient leaching, were investigated in microcosms containing the grass Phalaris aquatica L. Mycorrhizal and non-mycorrhizal plants were grown in a mixture of riparian soil and sand under glasshouse conditions. The formation of AM by P. aquatica significantly increased plant growth and nutrient uptake. Lower levels of NO3–, NH4+ and plant available P in both soil and leachate were observed in columns containing mycorrhizal root systems. These differences in nutrient interception were proportionally greater than the increase in root biomass of the mycorrhizal plants, compared with their non-mycorrhizal counterparts. Taken together, these data indicate that mycorrhizal root systems have an important, but previously little considered, role to play reducing the net loss of nutrients via leaching.

Arbuscular mycorrhizas enhance plant interception of leached nutrients.

The capacity of mycorrhizal and non-mycorrhizal root systems to reduce nitrate (NO3−) and ammonium (NH4+) loss from soils via leaching was investigated in a microcosm-based study. A mycorrhiza defective tomato mutant and its mycorrhizal wildtype progenitor were used in this experiment in order to avoid the indirect effects of establishing non-mycorrhizal control treatments on soil nitrogen cycling and the wider soil biota. Mycorrhizal root systems dramatically reduced nitrate loss (almost 40 times less) via leaching, compared to their non-mycorrhizal counterparts, following a pulse application of ammonium nitrate to experimental microcosms. The capacity of AM to reduce nutrient loss via leaching has received relatively little attention, but as demonstrated here, can be significant. Taken together, these data highlight the need to consider the potential benefits of AM beyond improvements in plant nutrition alone.

/pdf/arbuscular-mycorrhizas-reduce-nitrogen-loss-via-leaching-4p55dsq0rh.pdf

Arbuscular mycorrhizas reduce nitrogen loss via leaching.

Two experiments with soil cores were carried out to investigate the effects of arbuscular mycorrhizal (AM) fungal colonization on mobility of phosphorus (P) during leaching of repacked columns of a soil with a loamy sand texture. Trifolium subterraneum plants inoculated with an AM fungus or not inoculated were grown in cores with low or high P concentrations for 8 or 10 weeks in the glasshouse. Cores were then irrigated with 2500 mL water and the leachate collected. Plant growth and the amounts of P removed by plants, remaining in soil as available P and removed dissolved in leachate were measured. Mycorrhizal fungal colonization and development of external hyphae were also determined. Inoculation and/or P application significantly increased plant growth and plant P removal and decreased P leaching. In low P soils AM fungal colonization significantly increased plant P uptake and decreased soil available P and total dissolved P in leachates. Lower P leaching from cores with AM plants under low P conditions was related to enhancement of plant growth and to scavenging and removal of P from the soil by roots and/or external hyphae. When P was applied AM effects were not observed and available P remaining in the soil after leaching was much higher, regardless of AM fungal colonization.

Influence of Arbuscular Mycorrhizal (AM) Symbiosis on Phosphorus Leaching through Soil Cores

Effects of the mycorrhizal fungus Glomus intraradices on establishment of subterranean clover (Trifolium subterraneum L.) seedlings in saline conditions were studied in a glasshouse experiment. Growth and nutrient uptake were determined 10, 20 and 30 days after transplanting of mycorrhizal and nonmycorrhizal matched seedlings into soils with five different levels of salinity. Mycorrhizal plants had greater shoot and root dry weight than nonmycorrhizal plants. The enhancement in seedling dry weight due to mycorrhizal fungi was greater under high salinity levels. The detrimental effects of salinity stress on plant growth were appeared immediately after application low salinity stress to nonmycorrhizal plants (3.5 dS/m), but it was only observed in mycorrhizal plants at 7.5 dS/m and above. Mycorrhizal fungi increased P concentrations in shoots and roots compared with nonmycorrhizal plants particularly at 12 dS/m. Root K/Na ratio was also increased in mycorrhizal plants, possibly contributing to salinity tolerance. Calculation of mycorrhizal responses in terms of plant dry weight, P and K contents showed that the beneficial effects of mycorrhizal fungi on seedling salinity tolerance are due to different mechanisms at different stage of growth: increased P uptake during early growth and increased K uptake at the later stages. Results are discussed in the context of application of mycorrhizal inoculation to revegetation of salt affected lands.

https://ijpp.gau.ac.ir/article_616_534ab87c88357c720a58b225bd86c772.pdf

Hamid Reza Asghari

Papers

Growth response of Atriplex nummularia to inoculation with arbuscular mycorrhizal fungi at different salinity levels

Arbuscular mycorrhizas enhance plant interception of leached nutrients.

Arbuscular mycorrhizas reduce nitrogen loss via leaching.

Influence of Arbuscular Mycorrhizal (AM) Symbiosis on Phosphorus Leaching through Soil Cores

Vesicular-arbuscular (VA) mycorrhizae improve salinity tolerance in pre-inoculation subterranean clover (Trifolium subterraneum) seedlings