The rhizosphere is a complex environment where roots interact with physical, chemical and biological properties of soil. Structural and functional characteristics of roots contribute to rhizosphere processes and both have significant influence on the capacity of roots to acquire nutrients. Roots also interact extensively with soil microorganisms which further impact on plant nutrition either directly, by influencing nutrient availability and uptake, or indirectly through plant (root) growth promotion. In this paper, features of the rhizosphere that are important for nutrient acquisition from soil are reviewed, with specific emphasis on the characteristics of roots that influence the availability and uptake of phosphorus and nitrogen. The interaction of roots with soil microorganisms, in particular with mycorrhizal fungi and non-symbiotic plant growth promoting rhizobacteria, is also considered in relation to nutrient availability and through the mechanisms that are associated with plant growth promotion.

Acquisition of phosphorus and nitrogen in the rhizosphere and plant growth promotion by microorganisms

The Green Revolution boosted crop yields in developing nations by introducing dwarf genotypes of wheat and rice capable of responding to fertilisation without lodging. We now need a second Green Revolution, to improve the yield of crops grown in infertile soils by farmers with little access to fertiliser, who represent the majority of third-world farmers. Just as the Green Revolution was based on crops responsive to high soil fertility, the second Green Revolution will be based on crops tolerant of low soil fertility. Substantial genetic variation in the productivity of crops in infertile soil has been known for over a century. In recent years we have developed a better understanding of the traits responsible for this variation. Root architecture is critically important by determining soil exploration and therefore nutrient acquisition. Architectural traits under genetic control include basal-root gravitropism, adventitious-root formation and lateral branching. Architectural traits that enhance topsoil foraging are important for acquisition of phosphorus from infertile soils. Genetic variation in the length and density of root hairs is important for the acquisition of immobile nutrients such as phosphorus and potassium. Genetic variation in root cortical aerenchyma formation and secondary development (‘root etiolation’) are important in reducing the metabolic costs of root growth and soil exploration. Genetic variation in rhizosphere modification through the efflux of protons, organic acids and enzymes is important for the mobilisation of nutrients such as phosphorus and transition metals, and the avoidance of aluminum toxicity. Manipulation of ion transporters may be useful for improving the acquisition of nitrate and for enhancing salt tolerance. With the noteworthy exceptions of rhizosphere modification and ion transporters, most of these traits are under complex genetic control. Genetic variation in these traits is associated with substantial yield gains in low-fertility soils, as illustrated by the case of phosphorus efficiency in bean and soybean. In breeding crops for low-fertility soils, selection for specific root traits through direct phenotypic evaluation or molecular markers is likely to be more productive than conventional field screening. Crop genotypes with greater yield in infertile soils will substantially improve the productivity and sustainability of low-input agroecosystems, and in high-input agroecosystems will reduce the environmental impacts of intensive fertilisation. Although the development of crops with reduced fertiliser requirements has been successful in the few cases it has been attempted, the global scientific effort devoted to this enterprise is small, especially considering the magnitude of the humanitarian, environmental and economic benefits being forgone. Population growth, ongoing soil degradation and increasing costs of chemical fertiliser will make the second Green Revolution a priority for plant biology in the 21st century.

Roots of the Second Green Revolution

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.

Plant and microbial strategies to improve the phosphorus efficiency of agriculture

Publisher Summary This chapter discusses the competing theories of root lifespan and reviews the evidence available to support them. New methods of root observation and analysis produce data appropriate to testing these theories, but the results till date are few and often conflicting. Tentative generalizations include a suggestion that small diameter roots with low tissue density tend to have short lifespan. Root lifespan appears to be longest in cold environments, but data are lacking for tropical species. There is a strong seasonal variation in lifespan, with roots produced in the fall surviving longest, at least in temperate climates. Species differences are difficult to quantify because of seasonal and interannual variation, but root lifespan of deciduous fruit crops seems to be shorter than those of temperate deciduous forest trees or citrus, a broadleaf evergreen. The current model of root efficiency omits some important factors that may exert control over root lifespan. Fine roots have other functions in addition to absorption, including transport of water and nutrients. Furthermore, seasonality of climate and the need for carbon and nutrient storage could constrain the root lifespan that optimizes plant fitness to differ from that which maximizes root efficiency.

The Ecology of Root Lifespan

Publisher Summary This chapter discusses the ecological implications of mycorrhizal associations in natural ecosystems and the role of soil or environmental factors, mycorrhizal fungus characteristics or host plant properties. Mycorrhizal associations are regulated by features of the host plant and mycorrhizal fungus, as well as by soil conditions and environmental factors. Factors which can influence the occurrence and effectiveness of mycorrhizal associations include (1) root properties (2) edaphic or climatic factors (3) soil organisms, (4) soil disturbance, and (5) host-fungus compatibility. Several other complex ecological topics also discussed in the chapter include (1) mycorrhizal phenology , (2) factors responsible for varying degrees of mycorrhizal dependency in host plants, (3) the role of mycorrhizal hyphae in soil (4) nutrient competition involving mycorrhizal and non-mycorrhizal plants, and (5) mycorrhizal interactions involving pollution and other stresses, the rhizosphere, soil properties and allelopathy. Finally, the chapter also discusses the population ecology of mycorrhizal fungi and the influence of their associations on plant population ecology.

Mycorrhizas in Natural Ecosystems

Phosphorus Uptake by Six Plant Species as Related to Root Hairs1

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Phosphorus Uptake by Six Plant Species as Related to Root Hairs1