In a recent Forum article, Tibbett (2000) argued that mycorrhizal symbionts are more important than root proliferation for the exploitation of nutrient-rich zones or ‘patches’ in soil. There is little doubt that mycorrhizal fungi can enhance nutrient capture for their host. Both ecto- and ericoid mycorrhizal fungi can access nutrients (e.g. organic N) that would otherwise be unavailable to the plant, and through enzymatic production may enhance organic matter degradation. But this paper addresses the role of arbuscular mycorrhizas (AMs) in patch exploitation, as AMs are the most common of mycorrhizal associations, occurring in two-thirds of all land plant species. Tibbett’s case for mycorrhizas competing better than roots for nutrients needs critical examination. He stated that ‘we may come to perceive the role of roots, in terms of acquiring nutrients from organic patches, as relatively inert structures from which symbiotic organs, that can better compete with the wider microbiota are dispatched.’ Thus fungal, not root, proliferation would be expected. In support of this view, Tibbett (2000) cited experiments where root proliferation took a long time to occur in complex organic patches of a high C : N ratio. Other studies however, reveal that: • Plant N capture from N-rich patches of lower C : N ratio (<4) can be substantial (45–54%) even if root proliferation occurs only after c. 25 days (Hodge, Robinson & Fitter 2000). • Mycorrhizal colonization had little effect on the root foraging responses of seven co-occurring herbaceous species, and only one (Oxalis acetosella) showed a nutritional benefit from colonization (Farley & Fitter 1999). • Root responses are demonstrably more important for plant N capture than those of AM fungi. Lolium perenne seedlings supplied with l-lysine patches had the same slight (<20%) AM colonization that did not vary among treatments nor differ from controls (Hodge et al. 1999a). By contrast, roots responded strongly to the treatments. Duke et al. (1994) concluded that ‘increased plant root proliferation and uptake capacity are likely to be more important for the exploitation of temporary nutrient pulses or patches than is increased mycorrhizal activity.’ In contrast, Tibbett (2000) stated that ‘current emphasis on plant roots may have over estimated their significance in patch exploitation.’ Which view is correct? It is not yet possible to give a definitive answer. More experiments must be carried out on the ecology of AM fungi, in particular on the role of the external mycelium, before we can determine their importance in patch exploitation and their influence on root proliferation. Furthermore, AM fungi are multifunctional, and it cannot be assumed that all will necessarily enhance nutrient capture equally. Meanwhile, it should be remembered that the context in which roots proliferate may be as important as the response itself, for example, proliferation is relatively unimportant for nitrate capture by isolated individuals, but is important when individuals compete for nitrate (Hodge et al. 1999b). This context dependency may also apply to AM fungi. For example, inoculation of Plantago lanceolata seedlings with an AM fungus did not increase N capture from an organic patch, but it did increase root production, most pronouncedly within the patch zone (Hodge et al. 2000). Had the plants been grown in competition, however, the AM-related increase in root production may have enhanced their N capture. Root proliferation in N-rich patches would still give a plant a competitive advantage over its neighbours, whatever their mycorrhizal status.
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