The apoplast and its significance for plant mineral nutrition.

It has only recently become apparent that the apoplast plays a major role in a diverse range of processes, including intercellular signalling, plant-microbe interactions and both water and nutrient transport. Broadly defined, the apoplast constitutes all compartments beyond the plasmalemma - the interfibrillar and intermicellar space of the cell walls, and the xylem, including its gas- and water-filled intercellular space - extending to the rhizoplane and cuticle of the outer plant surface. The physico-chemical properties of cell walls influence plant mineral nutrition, as nutrients do not simply pass through the apoplast to the plasmalemma but can also be adsorbed or fixed to cell-wall components. Here, current progress in understanding the significance of the apoplast in plant mineral nutrition is reviewed. The contribution of the root apoplast to short-distance transport and nutrient uptakes is examined particularly in relation to Na toxicity and Al tolerance. The review extends to long-distance transport and the role of the apoplast as a habitat for microorganisms. In the leaf, the apoplast might have benefits over the vacuole as a site for short-term nutrient storage and solute exchange with the atmosphere. Contents Summary 167 I. Introduction 168 II. The properties of the apoplast and its implication for solute movement 168 1. The middle lamella 168 2. The primary wall 168 3. The secondary cell wall 169 III. The root apoplast - nutrient uptake and short-distance transport 170 IV. The apoplast as a compartment for long distance transport 174 V. The apoplast - habitat for microorganisms 175 VI. The apoplast of leaves - a compartment of storage and of reactions 177 1. Transport routes in the leaf apoplast 177 2. Methods of studying apoplastic solutes 177 3. Solute relations in the leaf apoplast 178 4. Concentration gradients in the leaf apoplast 179 5. Ion relations in the leaf apoplast and symptoms of deficiency and toxicity 179 6. Ion relations in the leaf apoplast - influence of nutrient supply 180 7. The leaf apoplast - compartment for transient ion storage 180 8. Ion fluxes between apoplast and symplast 181 9. Apoplastic ion balance 181 10. Leaf apoplast - interaction with the atmosphere 183 VII. Conclusions 183 Acknowledgements 183 References 183.