Phosphate Transport and Homeostasis in Arabidopsis

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Phosphorus (P) is an essential macronutrient for all living organisms. It serves various basic biological functions as a structural element in nucleic acids and phospholipids, in energy metabolism, in the activation of metabolic intermediates, as a component in signal transduction cascades, and the regulation of enzymes. Of the major nutrients, P is the most dilute and the least mobile in soil. High sorbing capacity for P in the soil (e.g. sorbtion to metal oxides), P mineralization (e.g. calcium phosphates such as apatite), and/or fixation of P in organic soil matter (by converting soluble P into organic molecules) result in low availability of this macronutrient for uptake into plants (Marschner, 1995). P is absorbed by plants as orthophosphate (Pi, inorganic phosphate). Pi concentration in the soil solution hardly reaches 10 µM and may even drop to submicromolar levels at the root/soil interface, where Pi uptake by plants and root surface-colonizing microorganisms leads to the generation of a zone of Pi depletion around the root cylinder that is maintained due to slow diffusion of Pi from regions distant to the root surface (Figure 1). Fig. 1. A transverse section through the tip of a primary root. The dotted line indicates the outer border of the P depletion zone. The arrow indicates the direction of growth. In industrialized countries, low P availability in agricultural soils is compensated by a high input of P fertilizer to guarantee high crop productivity and yield. Water run-off, soil erosion and leakage in highly fertilized agricultural soils may cause environmental problems such as eutrophication of lakes and rivers. As forecasted by Tilman et al. (2001), during the next 50 years, which is likely to be the final period of rapid agricultural expansion, demand for food by global population will be a major driver of global environmental change. Conversion of natural ecosystems to agriculture by 2050 will be accompanied by an approximate 2.5-fold increase in nitrogen- and P- driven eutrophication of terrestrial, freshwater, and near-shore marine ecosystems. Modern agricultural soils are almost universally maintained at high fertilization. Selection of new cultivars is usually made under such conditions and will not normally distinguish between plants varying in nutrient efficiency (Stevens and Rick, 1986). To alleviate the forecasted adverse negative effects of agricultural expansion, scientists have started to use classical breeding strategies and biotechnology to improve crop plants, based on the current knowledge and aiming at an improved crop yield with a lower input of fertilizer, thus protecting the environment. In contrast, in many developing tropical countries, subsistence farmers can not buy enough fertilizer due to limited financial capacities or poor infrastructure (Sanchez et al., 1997). As a consequence, P deprivation dramatically limits crop yield and is one of the reasons for poverty and malnutrition. In the future, agriculture from both developed as well as developing countries could thus benefit from modern crop varieties with enhanced P efficiency, thus leading to improved fertilizer management and increased crop yield on low-P soils. Thorough knowledge of the plant's response to P deprivation stress will contribute to the rational and targeted breeding of P efficient crop plants. Therefore, the authors of this chapter focus on summarizing the current state of research covering physiology, biochemistry, and molecular genetics of P acquisition and allocation, and P homeostasis within the plant. Although this review will mainly focus on knowledge acquired on Arabidopsis thaliana, some specific results obtained with other plant species will also be included in this work. For example, formation of mycorrhizae, which is observed in most vascular plants and strongly contributes to plant P nutrition, does not occur in Brassicaceae and therefore Arabidopsis is not a suitable model for mycorrhizae studies.