Summary Here, the coevolution of mycorrhizal fungi and roots is assessed in the light of evidence now available, from palaeobotanical and morphological studies and the analysis of DNA-based phylogenies. The first bryophyte-like land plants, in the early Devonian (400 million years ago), had endophytic associations resembling vesicular‐ arbuscular mycorrhizas (VAM) even before roots evolved. Mycorrhizal evolution would have progressed from endophytic hyphae towards balanced associations where partners were interdependent due to the exchange of limiting energy and nutrient resources. Most mycorrhizas are mutualistic, but in some cases the trend for increasing plant control of fungi culminates in the exploitative mycorrhizas of achlorophyllous, mycoheterotrophic plants. Ectomycorrhizal, ericoid and orchid mycorrhizas, as well as nonmycorrhizal roots, evolved during the period of rapid angiosperm radiation in the Cretaceous. It is hypothesised that roots gradually evolved from rhizomes to provide more suitable habitats for mycorrhizal fungi and provide plants with complex branching and leaves with water and nutrients. Selection pressures have caused the morphological divergence of roots with different types of mycorrizas. Root cortex thickness and exodermis suberization are greatest in obligately mycorrhizal plants, while nonmycorrhizal plants tend to have fine roots, with more roots hairs and relatively advanced chemical defences. Major coevolutionary trends and the relative success of plants with different root types are discussed.

Coevolution of roots and mycorrhizas of land plants

The loss of carbon from roots (rhizodeposition) and the consequent proliferation of microorganisms in the surrounding soil, coupled with the physical presence of a root and processes associated with nutrient uptake, gives rise to a unique zone of soil called the rhizosphere. In this review, we bring together evidence to show that roots can directly regulate most aspects of rhizosphere C flow either by regulating the exudation process itself or by directly regulating the recapture of exudates from soil. Root exudates have been hypothesized to be involved in the enhanced mobilization and acquisition of many nutrients from soil or the external detoxification of metals. With few exceptions, there is little mechanistic evidence from soil-based systems to support these propositions. We conclude that much more integrated work in realistic systems is required to quantify the functional significance of these processes in the field. We need to further unravel the complexities of the rhizosphere in order to fully engage with key scientific ideas such as the development of sustainable agricultural systems and the response of ecosystems to climate change. Contents I. Introduction 460 II. What is rhizodeposition? 460 III. Regulation of rhizodeposition 460 IV. How large is the root exudation C flux? 463 V. How responsive is the root exudation C flux? 463 VI. How responsive is the microbial community to root exudation? 464 VII. The role of root exudates in nutrient acquisition 464 VIII. Mycorrhizal fungi and rhizodeposition 471 IX. Future thoughts 474 Acknowledgements 474 References 474.

https://nph.onlinelibrary.wiley.com/doi/epdf/10.1111/j.1469-8137.2004.01130.x

Plant and mycorrhizal regulation of rhizodeposition

▪ Abstract Arbuscular mycorrhizae are symbiotic associations formed between a wide range of plant species including angiosperms, gymnosperms, pteridophytes, and some bryophytes, and a limited range of fungi belonging to a single order, the Glomales. The symbiosis develops in the plant roots where the fungus colonizes the apoplast and cells of the cortex to access carbon supplied by the plant. The fungal contribution to the symbiosis is complex, but a major aspect includes the transfer of mineral nutrients, particularly phosphate from the soil to the plant. Development of this highly compatible association requires the coordinate molecular and cellular differentiation of both symbionts to form specialized interfaces over which bi-directional nutrient transfer occurs. Recent insights into the molecular events underlying these aspects of the symbiosis are discussed.

Molecular and cellular aspects of the arbuscular mycorrhizal symbiosis

Secondary plant compounds are important signals in several symbiotic and pathogenic plant-microbe interactions. The present review is limited to two groups of secondary plant compounds, flavonoids and strigolactones, which have been reported in root exudates. Data on flavonoids as signaling compounds are available from several symbiotic and pathogenic plant-microbe interactions, whereas only recently initial data on the role of strigolactones as plant signals in the arbuscular mycorrhizal symbiosis have been reported. Data from other plant-microbe interactions and strigolactones are not available yet. In the present article we are focusing on flavonoids in plant-fungal interactions such as the arbuscular mycorrhizal (AM) association and the signaling between different Fusarium species and plants. Moreover the role of strigolactones in the AM association is discussed and new data on the effect of strigolactones on fungi, apart from arbuscular mycorrhizal fungi (AMF), are provided.

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Flavonoids and strigolactones in root exudates as signals in symbiotic and pathogenic plant-fungus interactions.

Plants require adequate P from the very early stages of growth for optimum crop production. Phosphorus supply to the crop is affected by soil P, P fertilizer management and by soil and environmental conditions influencing P phytoavailability and root growth. Phosphorus uptake in many crops is improved by associations with arbuscular mycorrhizal fungi. Cropping system and long-term input of P through fertilizers and manures can influence the amount and phytoavailability of P in the system and the development of mycorrhizal associations. Optimum yield potential requires an adequate P supply to the crop from the soil or from P additions. Where early-season P supply is low, P fertilization may improve P nutrition and crop yield potential. Alternately, under low-P conditions, encouragement of arbuscular mycorrhizal associations may enhance P uptake by crops early in the growing season, improving crop yield potential and replacing starter fertilizer P applications. Soil P supply that exceeds P requirements of t...

Soil and fertilizer phosphorus: Effects on plant P supply and mycorrhizal development

The effect of solution phosphorus (P) concentration upon growth of pregerminated spores of the vesicular-arbuscular mycorrhizal fungus Gigaspora margarita was examined in vitro. P at 1 mM significantly inhibited branching of the primary germ tube. The number of branches and the total hyphal length were both significantly inhibited at 10 mM P. In addition, germinated spores exposed to exudates produced by Ri T-DNA-transformed roots of Daucus carota L. grown in the presence of P showed significantly less hyphal branching than those exposed to exudates produced by P-stressed roots. These phenomena could contribute to the observed inhibition of mycorrhiza formation by high P.

Phosphorus amendment inhibits hyphal branching of the VAM fungus Gigaspora margarita directly and indirectly through its effect on root exudation

Experiments were conducted to test the hypothesis that cellular compounds, especially wall-associated compounds, released during emergence of secondary roots, stimulate the growth of arbuscular mycorrhizal (AM) fungi. Purified cell walls, crude cell-wall extracts, crude cytoplasmic extracts, and phenolic compounds previously identified as cell wall-associated, from Ri T-DNA-transformed roots of host (Daucus carota L.) and non-host (Beta vulgaris L.) were incorporated into growth medium and tested for their effects upon growth of the AM fungi Gigaspora gigantea (Nicol. & Gerd.) Gerdemann and Trappe and Gigaspora margarita Becker and Hall. Purified cell walls of both plants had little effect on G. gigantea but non-host cell walls inhibited the growth of G. margarita. Ferulic acid, a major constituent of non-host root, depressed the growth of both fungi. Nothing tested which was unique to the non-host root affected hyphal growth to the point that contact would be prohibited. Caffeic acid, found in D. carota cytoplasm, also depressed growth of both fungi. Para-hydroxybenzoic acid, a constituent of D. carota roots, stimulated growth of G. margarita hyphae, but did not affect hyphal growth of G. gigantea. Vanillic acid, unique to D. carota root cell-wall extracts, stimulated hyphal growth and branching of both fungi, and should increase the probability of contact between fungus and host root.

https://nph.onlinelibrary.wiley.com/doi/pdf/10.1111/j.1469-8137.1996.tb01896.x

The differential effects of cell wall-associated phenolics, cell walls, and cytosolic phenolics of host and non-host roots on the growth of two species of AM fungi.

G. D. Abney

Papers

Phosphorus amendment inhibits hyphal branching of the VAM fungus Gigaspora margarita directly and indirectly through its effect on root exudation

The differential effects of cell wall-associated phenolics, cell walls, and cytosolic phenolics of host and non-host roots on the growth of two species of AM fungi.