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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.

Background Agricultural production is often limited by low phosphorus (P) availability. In developing countries, which have limited access to P fertiliser, there is a need to develop plants that are more efficient at low soil P. In fertilised and intensive systems, P-efficient plants are required to minimise inefficient use of P-inputs and to reduce potential for loss of P to the environment. Scope Three strategies by which plants and microorganisms may improve P-use efficiency are outlined: (i) Root-foraging strategies that improve P acquisition by lowering the critical P requirement of plant growth and allowing agriculture to operate at lower levels of soil P; (ii) P-mining strategies to enhance the desorption, solubilisation or mineralisation of P from sparingly-available sources in soil using root exudates (organic anions, phosphatases), and (iii) improving internal P-utilisation efficiency through the use of plants that yield more per unit of P uptake.Conclusions We critically review evidence that more P-efficient plants can be developed by modifying root growth and architecture, through manipulation of root exudates or by managing plant-microbial associations such as arbuscular mycorrhizal fungi and microbial inoculants. Opportunities to develop P-efficient plants through breeding or genetic modification are described and issues that may limit success including potential trade-offs and trait interactions are discussed. Whilst demonstrable progress has been made by selecting plants for root morphological traits, the potential for manipulating root physiological traits or selecting plants for low internal P concentration has yet to be realised.

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.