Leaves constitute a substantial fraction of the total resistance to water flow through plants. A key question is how hydraulic resistance within the leaf is distributed among petiole, major veins, minor veins, and the pathways downstream of the veins. We partitioned the leaf hydraulic resistance (R leaf ) for sugar maple (Acer saccharum) and red oak (Quercus rubra) by measuring the resistance to water flow through leaves before and after cutting specific vein orders. Simulations using an electronic circuit analog with resistors arranged in a hierarchical reticulate network justified the partitioning of total R leaf into component additive resistances. On average 64% and 74% of the R leaf was situated within the leaf xylem for sugar maple and red oak, respectively. Substantial resistance-32% and 49%-was in the minor venation, 18% and 21% in the major venation, and 14% and 4% in the petiole. The large number of parallel paths (i.e. a large transfer surface) for water leaving the minor veins through the bundle sheath and out of the leaf resulted in the pathways outside the venation comprising only 36% and 26% of R leaf . Changing leaf temperature during measurement of R leaf for intact leaves resulted in a temperature response beyond that expected from changes in viscosity. The extra response was not found for leaves with veins cut, indicating that water crosses cell membranes after it leaves the xylem. The large proportion of resistance in the venation can explain why stomata respond to leaf xylem damage and cavitation. The hydraulic importance of the leaf vein system suggests that the diversity of vein system architectures observed in angiosperms may reflect variation in whole-leaf hydraulic capacity.Water flow through the leaf is one of the most important but least understood components of the whole-plant hydraulic system. The leaf hydraulic resistance (R leaf ) constitutes a significant hydraulic bottleneck, correlates with leaf structure, and apparently constrains gas exchange (Tyree and Zimmermann, 2002;Sack et al., 2003b;Sack and Tyree, 2004). R leaf is an aggregate measure; once past the petiole, water flows through a reticulate network of veins, across the bundle sheath cells, and through or around mesophyll cells before evaporation and diffusion from the stomata (Esau, 1965). Recent work has focused primarily on measurement of R leaf and its functional correlates. Few studies have attempted to determine quantitatively how the various components of the water flow pathway through the leaf contribute to its total resistance.Partitioning of R leaf is a crucial step for understanding leaf hydraulic design. Cavitation in the leaf vein xylem can substantially increase R leaf , as can physical damage to major veins; both drive reductions of leaf water potential and gas exchange Nardini et al., 2001Cochard et al., 2002;Huve et al., 2002;Sack et al., 2003a). These observations suggest that a large proportion of R leaf is situated in the veins. However, a number of studies have reported that most of the leaf resista...

Leaf venation is a showcase of plant diversity, ranging from the grid-like network in grasses, to a wide variety of dendritic systems in other angiosperms. A principal function of the venation is to deliver water; however, a hydraulic significance has never been demonstrated for contrasting major venation architectures, including the most basic dichotomy, ''pinnate'' and ''palmate'' systems. We hypothesized that vascular redundancy confers tolerance of vein breakage such as would occur during mechanical or insect damage. We subjected leaves of woody angiosperms of contrasting venation architecture to severing treatments in vivo, and, after wounds healed, made detailed measurements of physiological performance relative to control leaves. When the midrib was severed near the leaf base, the pinnately veined leaves declined strongly in leaf hydraulic conductance, stomatal conductance, and photosynthetic rate, whereas palmately veined leaves were minimally affected. Across all of the species examined, a higher density of primary veins predicted tolerance of midrib damage. This benefit for palmate venation is consistent with its repeated evolution and its biogeographic and habitat distribution. All leaves tested showed complete tolerance of damage to second-and higherorder veins, demonstrating that the parallel flow paths provided by the redundant, reticulate minor vein network protect the leaf from the impact of hydraulic disruption. These findings point to a hydraulic explanation for the diversification of low-order vein architecture and the commonness of reticulate, hierarchical leaf venation. These structures suggest roles for both economic constraints and risk tolerance in shaping leaf morphology during 130 million years of flowering plant evolution.herbivory ͉ evolution ͉ physiology ͉ plant traits ͉ hydraulic architecture

In order to enhance biomedical research and development efficiency and innovation, nontraditional research collaborations have emerged that feature the sharing of information, resources, and capabilities. Although many of these so-called precompetitive collaborations are in the field of oncology, the lessons they offer are broadly applicable to other subfields of translational medicine.by guest on May 12, 2018