3 SUMMARYA great diversity of plants and fungi engage in mycorrhizal associations. In natural habitats, and in an ecologically meaningful time span, these associations have evolved to improve the fitness of both plant and fungal symbionts. In systems managed by humans, mycorrhizal associations often improve plant productivity, but this is not always the case. Mycorrhizal fungi might be considered to be parasitic on plants when net cost of the symbiosis exceeds net benefits. Parasitism can be developmentally induced, environmentally induced, or possibly genotypically induced. Morphological, phenological, and physiological characteristics of the symbionts influence the functioning of mycorrhizas at an individual scale. Biotic and abiotic factors at the rhizosphere, community, and ecosystem scales further mediate mycorrhizal functioning. Despite the complexity of mycorrhizal associations, it might be possible to construct predictive models of mycorrhizal functioning. These models will need to incorporate variables and parameters that account for differences in plant responses to, and control of, mycorrhizal fungi, and differences in fungal effects on, and responses to, the plant. Developing and testing quantitative models of mycorrhizal functioning in the real world requires creative experimental manipulations and measurements. This work will be facilitated by recent advances in molecular and biochemical techniques. A greater understanding of how mycorrhizas function in complex natural systems is a prerequisite to managing them in agriculture, forestry, and restoration.

Root systems of most land plants form arbuscular mycorrhizal (AM) symbioses in the field, and these contribute to nutrient uptake. AM roots have two pathways for nutrient absorption, directly through the root epidermis and root hairs and via AM fungal hyphae into root cortical cells, where arbuscules or hyphal coils provide symbiotic interfaces. New physiological and molecular evidence shows that for phosphorus the mycorrhizal pathway (MP) is operational regardless of plant growth responses (positive or negative). Amounts delivered cannot be determined from plant nutrient contents because when responses are negative the contribution of the direct pathway (DP) is reduced. Nitrogen (N) is also delivered to roots via an MP, but the contribution to total N requirement and the costs to the plant are not clear. The functional interplay between activities of the DP and MP has important implications for consideration of AM symbioses in ecological, agronomic, and evolutionary contexts.

Arbuscular mycorrhizal (AM) fungi are vital components of nearly all terrestrial ecosystems, forming mutually beneficial (mutualistic) symbioses with the roots of around 80% of vascular plants and often increasing phosphate (P) uptake and growth. We present novel data showing that AM fungi can provide the dominant route for plant P supply, even when overall growth or P uptake remains unaffected. The results will change our understanding of the roles of AM fungi in agricultural and natural ecosystems; they also predict that mycorrhiza-specific plant P transporters must play a major role in plant P uptake regardless of whether the plants respond to AM colonization by taking up more P per plant or by increased dry weight, compared with nonmycorrhizal (NM) control plants.

In this Update, we review new findings about the roles of the arbuscular mycorrhizas (mycorrhiza = fungus plus root) in plant growth and phosphorus (P) nutrition. We focus particularly on the function of arbuscular mycorrhizal (AM) symbioses with different outcomes for plant growth (from positive to negative) and especially on the interplay between direct P uptake via root epidermis (including root hairs when present) and uptake via the AM fungal pathway. The results are highly relevant to many aspects of AM symbiosis, ranging from signaling involved in the development of colonization and the regulation of P acquisition to the roles of AM fungi in determining the composition of natural plant assemblages in ecological settings and their changes with time.

Summary• We investigated structural and functional diversity in arbuscular mycorrhizal (AM) symbioses involving three plant species and three AM fungi and measured contributions of the fungi to P uptake using compartmented pots and 33 P. The plant / fungus combinations varied in growth and P responses. Flax ( Linum usitatissimum ) responded positively to all fungi, and medic ( Medicago truncatula ) to Glomus caledonium and G. intraradices , but not Gigaspora rosea . Tomato ( Lycopersicon esculentum ) showed no positive responses.• Hyphal growth in soil was very low for Gi. rosea and high for both Glomus spp. Hyphal lengths in root + hyphal compartment (RHC) and hyphal compartment (HC) were similar for G. intraradices , but much higher in HC for G. caledonium .• Specific activities of 33 P in plants and soil indicated that fungal P uptake made substantial contributions to five plant/fungus combinations and significant contributions to a further two. G. intraradices delivered close to 100% of the P in all three plants. G. caledonium and Gi. rosea delivered less P. The amount was not related to colonisation or to growth or P responses.• We conclude that: AM colonisation can result in complete inactivation of the direct P uptake pathway via root hairs and epidermis; calculations of AM contributions to P uptake from total plant P will often be highly inaccurate; and lack of plant responsiveness does not mean that an AM fungus makes no contribution to P uptake.

Summary• Arbuscular mycorrhizal fungal (AMF) communities were established in pots using fungal isolates from a single field in Switzerland. It was tested whether multispecies mixtures provided more phosphorus and supported greater plant growth than single AMF species.• Two host plants, medic (Medicago truncatula) and leek (Allium porrum), were inoculated with three AMF species (Glomus mosseae, G. claroideum and G. intraradices), either separately or in mixtures. The composition of the AMF communities in the roots was assessed using real-time PCR to determine the copy number of large ribosomal subunit genes.• Fungal communities in the roots were usually dominated by one AMF species (G. mosseae). The composition of the communities depended on both plant identity and the time of harvest. Leek colonized by a mixture of G. claroideum and G. intraradices acquired more P than with either of the two AMF separately.• Direct evidence is provided for functional complementarity among species within the AMF community colonizing a single root system. Competition among the species poses a major challenge in interpreting experiments with mixed inoculations, but this is greatly facilitated by use of real-time PCR.

Arbuscular mycorrhizal (AM) symbioses are formed by approximately 80% of vascular plant species in all major terrestrial biomes. In consequence an understanding of their functions is critical in any study of sustainable agricultural or natural ecosystems. Here we discuss the implications of recent results and ideas on AM symbioses that are likely to be of particular significance for plants dealing with abiotic stresses such as nutrient deficiency and especially water stress. In order to ensure balanced coverage, we also include brief consideration of the ways in which AM fungi may influence soil structure, carbon deposition in soil and interactions with the soil microbial and animal populations, as well as plantplant competition. These interlinked outcomes of AM symbioses go well beyond effects in increasing nutrient uptake that are commonly discussed and all require to be taken into consideration in future work designed to understand the complex and multifaceted responses of plants to abiotic and biotic stresses in agricultural and natural environments.

Even though ecologists and agronomists have considered the spatial root distribution of plants to be important for interspecific interactions in natural and agricultural ecosystems, few experimental studies have quantified patterns of root distribution dynamics and their impacts on interspecific interactions. A field experiment was conducted to investigate the relationship between root distribution and interspecific interactions between intercropped plants. Roots were sampled twice by auger and twice by the monolith method in wheat (Triticum aestivum L.)/maize (Zea mays L.) and faba bean (Vicia faba L.)/maize intercropping and in sole wheat, maize, and faba bean up to 100 cm depth in the soil profile. The results showed that the roots of intercropped wheat spread under maize plants, and had much greater root length density (RLD) at all soil depths than sole wheat. The roots of maize intercropped with wheat were limited laterally, but had a greater RLD than sole-cropped maize. The RLD of maize intercropped with faba bean at different soil depths was influenced by intercropping to a smaller extent compared to maize intercropped with wheat. Faba bean had a relatively shallow root distribution, and the roots of intercropped maize spread underneath them. The results support the hypotheses that the overyielding of species showing benefit in the asymmetric interspecific facilitation results from greater lateral deployment of roots and increased RLD, and that compatibility of the spatial root distribution of intercropped species contributes to symmetric interspecific facilitation in the faba bean/maize intercropping.