Vascular tissues and guard cells play a central role in conducting water and controlling the transpiration stream, but to get in and out of vascular tissues water has to flow through living cells. When water flows across living tissues, it can follow different routes: across cell walls (apoplastic path), from cell to cell across either the plasmodesmata (symplastic path), or traversing the cell membranes (transcellular path). Biophysical criteria have been developed to distinguish between the apoplastic and cell-to-cell paths (16). These studies will now benefit from recent biochemical and genetic descriptions of critical cell barriers, such as the root endodermis with its Casparian bands, and from new insights into the molecular mechanisms of plasmodesmatal gating. The most significant breakthrough to date in understanding conductance of living cells has come from the discovery of a class of water channel proteins named aquaporins (1). Aquaporins have now been found in nearly all living organisms. They belong to a larger family of membrane proteins homologous to bovine major intrinsic protein (MIP) and exhibit a typical structure with six membrane-spanning domains and an internal symmetry showing two highly conserved Asn-Pro-Ala motifs (Fig. 1 ). Because of their abundance, plant MIP homologs were
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