1995; Van der Weerd et al. 2001, 2002). Diffusive exchange within compartments and exchange between compartments, passing membranes, affect the observed relaxation times (Van As 2007; Van As and Windt 2008). The observed T 2 (and T 1) of vacuolar water has been demonstrated to depend on the bulk T 2 in the vacuole (T 2, bulk), and the surface-to-volume ratio, S/V, of the vacuole (van der Weerd et al. 2001): $$ 1/T_2,\;\textobs = (H\; \times \; S/V)\; + \; 1/T_2,\;\textbulk $$ (6)The proportionality constant H is directly related to the actual tonoplast membrane permeability
for water (van der Weerd et al. 2002; Van As 2007). Equation 6 holds also for water in (xylem) vessels, where H now represents the
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MacFall and Van As 1996; Köckenberger 2001; Van As 2007; Van As and Windt 2008). In general, spatial resolution will not be high enough to resolve individual phloem or xylem vessels. As a consequence, pixels that contain flowing water will also contain a significant amount of stationary water. When vessels are very small, as is the case in phloem tissue, the relative amount of flowing water per pixel can be as small as a few percent. The greatest challenge in measuring phloem water transport, therefore, is to distinguish displacement of a small amount of very slowly moving water from a (very) large amount of stationary water showing displacements due to random movement as a result of Brownian motion.