John S Sperry

University of Utah, Salt Lake City, UT, USA

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Publications (54)282.44 Total impact

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    Article: Embolism resistance as a key mechanism to understand adaptive plant strategies.
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    ABSTRACT: One adaptation of plants to cope with drought or frost stress is to develop wood that is able to withstand the formation and distribution of air bubbles (emboli) in its water conducting xylem cells under negative pressure. The ultrastructure of interconduit pits strongly affects drought-induced embolism resistance, but also mechanical properties of the xylem are involved. The first experimental evidence for a lower embolism resistance in stems of herbaceous plants compared to stems of their secondarily woody descendants further supports this mechanical-functional trade-off. An integrative approach combining (ultra)structural observations of the xylem, safety-efficiency aspects of the hydraulic pipeline, and xylem-phloem interactions will shed more light on the multiple adaptive strategies of embolism resistance in plants.
    Current opinion in plant biology 02/2013; · 10.33 Impact Factor
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    Dataset: nature11688-s2
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    Article: Global convergence in the vulnerability of forests to drought.
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    ABSTRACT: Shifts in rainfall patterns and increasing temperatures associated with climate change are likely to cause widespread forest decline in regions where droughts are predicted to increase in duration and severity. One primary cause of productivity loss and plant mortality during drought is hydraulic failure. Drought stress creates trapped gas emboli in the water transport system, which reduces the ability of plants to supply water to leaves for photosynthetic gas exchange and can ultimately result in desiccation and mortality. At present we lack a clear picture of how thresholds to hydraulic failure vary across a broad range of species and environments, despite many individual experiments. Here we draw together published and unpublished data on the vulnerability of the transport system to drought-induced embolism for a large number of woody species, with a view to examining the likely consequences of climate change for forest biomes. We show that 70% of 226 forest species from 81 sites worldwide operate with narrow (<1 megapascal) hydraulic safety margins against injurious levels of drought stress and therefore potentially face long-term reductions in productivity and survival if temperature and aridity increase as predicted for many regions across the globe. Safety margins are largely independent of mean annual precipitation, showing that there is global convergence in the vulnerability of forests to drought, with all forest biomes equally vulnerable to hydraulic failure regardless of their current rainfall environment. These findings provide insight into why drought-induced forest decline is occurring not only in arid regions but also in wet forests not normally considered at drought risk.
    Nature 11/2012; · 36.28 Impact Factor
  • Article: Vulnerability curves by centrifugation: is there an open vessel artefact, and are 'r' shaped curves necessarily invalid?
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    ABSTRACT: Vulnerability curves using the 'Cavitron' centrifuge rotor yield anomalous results when vessels extend from the end of the stem segment to the centre ('open-to-centre' vessels). Curves showing a decline in conductivity at modest xylem pressures ('r' shaped) have been attributed to this artefact. We determined whether the original centrifugal method with its different rotor is influenced by open-to-centre vessels. Increasing the proportion of open-to-centre vessels by shortening stems had no substantial effect in four species. Nor was there more embolism at the segment end versus centre as seen in the Cavitron. The dehydration method yielded an 'r' shaped curve in Quercus gambelii that was similar to centrifuged stems with 86% open-to-centre vessels. Both 'r' and 's' (sigmoidal) curves from Cercocarpus intricatus were consistent with each other, differing only in whether native embolism had been removed. An 'r' shaped centrifuge curve in Olea europaea was indistinguishable from the loss of conductivity caused by forcing air directly across vessel end-walls. We conclude that centrifuge curves on long-vesselled material are not always prone to the open vessel artefact when the original rotor design is used, and 'r' shaped curves are not necessarily artefacts. Nevertheless, confirming curves with native embolism and dehydration data is recommended.
    Plant Cell and Environment 03/2012; 35(3):601-10. · 5.22 Impact Factor
  • Article: A species-level model for metabolic scaling in trees I. Exploring boundaries to scaling space within and across species
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    ABSTRACT: Summary 1. Metabolic scaling theory predicts how tree water flow rate (Q) scales with tree mass (M) and assumes identical scaling for biomass growth rate (G) withM. Analytic models have derived general scaling expectations from proposed optima in the rate of axial xylem conduit taper (taper function) and the allocation of wood space to water conduction (packing function). Recent predictions suggest G and Q scale withMto the � 0·7 power with 0·75 as an upper bound. 2. We complement this a priori optimization approach with a numerical model that incorporates species-specific taper and packing functions, plus additional empirical inputs essential for predicting Q (effects of gravity, tree size, heartwood, bark, and hydraulic resistance of leaf, root and interconduit pits). Traits are analysed individually, and in ensemble across tree types, to define a 2D ‘scaling space’ of absolute Q vs. its scaling exponent with tree size. 3. All traits influenced Q and many affected its scaling with M. Constraints driving the optimization of taper or packing functions, or any other trait, can be relaxed via compensatory changes in other traits. 4. The scaling space of temperate trees overlapped despite diverse anatomy and winter-adaptive strategies. More conducting space in conifer wood compensated for narrow tracheids; extensive sapwood in diffuse-porous trees compensated for narrow vessels; and limited sapwood in ring-porous trees negated the effect of large vessels. Tropical trees, however, achieved the greatest Q and steepest size-scaling by pairing large vessels with extensive sapwood, a combination compatible with minimal water stress and no freezing-stress. 5. Intraspecific scaling across all types averaged Q / M 0·63 (maximum = Q / M 0·71) for size-invariant root–shoot ratio. Scaling reached Q / M 0·75 only if conductance increased faster in roots than in shoots with size. Interspecific scaling could reach Q / M 0·75, but this may require the evolution of size-biased allometries rather than arising directly from biophysical constraints. 6. Our species-level model is more realistic than its analytical predecessors and provides a tool for interpreting the adaptive significance of functional trait diversification in relation to wholetree water use and consequent metabolic scaling. Key-words: ecological wood anatomy, functional tree types, hydraulic architecture, Metabolic scaling theory, plant allometry, tree water transport, vascular network theory, West Brown and Enquist
    Functional Ecology 01/2012; 26(5):1054-1065.. · 4.57 Impact Factor
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    Article: A species‐level model for metabolic scaling of trees II. Testing in a ring‐and diffuse‐porous species.
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    ABSTRACT: Summary 1. A 17-parameter ‘species model’ that predicts metabolic scaling from vascular architecture was tested in a diffuse-porous maple (Acer grandidentatum) and a ring-porous oak (Quercus gambelii). Predictions of midday water transport (Q) and its scaling with above-ground mass (M) were compared with empirical measurements. We also tested the assumption that Q was proportional to the biomass growth rate of the shoot (G). 2. Water transport and biomass growth rate were measured on 18 trees per species that spanned a broad range in trunk diameter (4–26 cm). Where possible, the same trees were used for obtaining the 17 model parameters that concern external branching, internal xylem conduit anatomy, and soil-to-canopy sap pressure drop. 3. The model succeeded in predicting the Q by Mb scaling exponent, b, being within 8% (maple) and 6% (oak) of measured exponents from sap flow data. In terms of absolute Q, the model was better in maple (16% Q overestimate) than oak (128% overestimate). The overestimation of Q was consistent with the model not accounting for cavitation, which is reportedly more prevalent in oak than in maple at the study site. 4. The modelled and measured Q by M b exponents averaged within 3·6% of the measured G by M b exponents, supporting the assumption that G / Q1. The average b exponent was 0·62 ± 0·016 (mean ± SE) across species, rejecting b = 0·75 for intraspecific scaling. 5. The performance of this species model, both for scaling purposes as well as for predicting rates of water consumption within and between species, argues for its further refinement and wider application in ecology and ecosystem biology. Key-words: allometry, ecohydrology, hydraulic architecture, metabolic scaling theory, plant water transport, ring-porous and diffuse-porous trees, sap flow, vascular networks, WBE model
    Functional Ecology 01/2012; 26:1066-1076. · 4.57 Impact Factor
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    Article: The roles of hydraulic and carbon stress in a widespread climate-induced forest die-off.
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    ABSTRACT: Forest ecosystems store approximately 45% of the carbon found in terrestrial ecosystems, but they are sensitive to climate-induced dieback. Forest die-off constitutes a large uncertainty in projections of climate impacts on terrestrial ecosystems, climate-ecosystem interactions, and carbon-cycle feedbacks. Current understanding of the physiological mechanisms mediating climate-induced forest mortality limits the ability to model or project these threshold events. We report here a direct and in situ study of the mechanisms underlying recent widespread and climate-induced trembling aspen (Populus tremuloides) forest mortality in western North America. We find substantial evidence of hydraulic failure of roots and branches linked to landscape patterns of canopy and root mortality in this species. On the contrary, we find no evidence that drought stress led to depletion of carbohydrate reserves. Our results illuminate proximate mechanisms underpinning recent aspen forest mortality and provide guidance for understanding and projecting forest die-offs under climate change.
    Proceedings of the National Academy of Sciences 12/2011; 109(1):233-7. · 9.68 Impact Factor
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    Article: Rare pits, large vessels and extreme vulnerability to cavitation in a ring-porous tree species.
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    ABSTRACT: • The rare pit hypothesis predicts that the extensive inter-vessel pitting in large early-wood vessels of ring-porous trees should render many of these vessels extremely vulnerable to cavitation by air-seeding. This prediction was tested in Quercus gambelii. • Cavitation was assessed from native hydraulic conductivity at field sap tension and in dehydrated branches. Single-vessel air injections gave air-seeding pressures through vessel files; these data were used to estimate air-seeding pressures for inter-vessel walls and pits. • Extensive cavitation occurred at xylem sap tensions below 1 MPa. Refilling occurred below 0.5 MPa and was inhibited by phloem girdling. Remaining vessels cavitated over a wide range to above 4 MPa. Similarly, 40% of injected vessel files air-seeded below 1.0 MPa, whereas the remainder seeded over a wide range exceeding 5 MPa. Inter-vessel walls averaged 1.02 MPa air-seeding pressure, similar and opposite to the mean cavitation tension of 1.22 MPa. Consistent with the rare pit hypothesis, only 7% of inter-vessel pits were estimated to air-seed by 1.22 MPa. • The results confirm the rare pit prediction that a significant fraction of large vessels in Q. gambelii experience high probability of failure by air-seeding.
    New Phytologist 12/2011; 193(3):713-20. · 6.64 Impact Factor
  • Article: Comparative hydraulic architecture of tropical tree species representing a range of successional stages and wood density.
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    ABSTRACT: Plant hydraulic architecture (PHA) has been linked to water transport sufficiency, photosynthetic rates, growth form and attendant carbon allocation. Despite its influence on traits central to conferring an overall competitive advantage in a given environment, few studies have examined whether key aspects of PHA are indicative of successional stage, especially within mature individuals. While it is well established that wood density (WD) tends to be lower in early versus late successional tree species, and that WD can influence other aspects of PHA, the interaction of WD, successional stage and the consequent implications for PHA have not been sufficiently explored. Here, we studied differences in PHA at the scales of wood anatomy to whole-tree hydraulic conductance in species in early versus late successional Panamanian tropical forests. Although the trunk WD was indistinguishable between the successional groups, the branch WD was lower in the early successional species. Across all species, WD correlated negatively with vessel diameter and positively with vessel packing density. The ratio of branch:trunk vessel diameter, branch sap flux and whole-tree leaf-specific conductance scaled negatively with branch WD across species. Pioneer species showed greater sap flux in branches than in trunks and a greater leaf-specific hydraulic conductance, suggesting that pioneer species can move greater quantities of water at a given tension gradient. In combination with the greater water storage capacitance associated with lower WD, these results suggest these pioneer species can save on the carbon expenditure needed to build safer xylem and instead allow more carbon to be allocated to rapid growth.
    Oecologia 03/2011; 167(1):27-37. · 3.41 Impact Factor
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    Article: Linking irradiance-induced changes in pit membrane ultrastructure with xylem vulnerability to cavitation.
    Lenka Plavcová, Uwe G Hacke, John S Sperry
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    ABSTRACT: The effect of shading on xylem hydraulic traits and xylem anatomy was studied in hybrid poplar (Populus trichocarpa x deltoides, clone H11-11). Hydraulic measurements conducted on stem segments of 3-month-old saplings grown in shaded (SH) or control light (C) conditions indicated that shading resulted in more vulnerable and less efficient xylem. Air is thought to enter vessels through pores in inter-vessel pit membranes, thereby nucleating cavitation. Therefore, we tested if the ultrastructure and/or chemistry of pit membranes differed in SH and C plants. Transmission electron micrographs revealed that pit membranes were thinner in SH, which was paralleled by lower compound middle lamella thickness. Immunolabelling with JIM5 and JIM7 monoclonal antibodies surprisingly indicated that pectic homogalacturonans were not present in the mature pit membrane regardless of the light treatment. Porosity measurements conducted with scanning electron microscopy were significantly affected by the method used for sample dehydration. Drying through a gradual ethanol series seems to be a better alternative to drying directly from a hydrated state for pit membrane observations in poplar. Scanning electron microscopy based estimates of pit membrane porosity probably overestimated real porosity as suggested by the results from the 'rare pit' model.
    Plant Cell and Environment 03/2011; 34(3):501-13. · 5.22 Impact Factor
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    Article: Calibration of thermal dissipation sap flow probes for ring- and diffuse-porous trees.
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    ABSTRACT: Thermal dissipation probes (the Granier method) are routinely used in forest ecology and water balance studies to estimate whole-tree transpiration. This method utilizes an empirically derived equation to measure sap flux density, which has been reported as independent of wood characteristics. However, errors in calculated sap flux density may occur when large gradients in sap velocity occur along the sensor length or when sensors are inserted into non-conducting wood. These may be conditions routinely associated with ring-porous species, yet there are few cases in which the original calibration has been validated for ring-porous species. We report results from laboratory calibration measurements conducted on excised stems of four ring-porous species and two diffuse-porous species. Our calibration results for ring-porous species were considerably different compared with the original calibration equation. Calibration equation coefficients obtained in this study differed by as much as two to almost three orders of magnitude when compared with the original equation of Granier. Coefficients also differed between ring-porous species across all pressure gradient conditions considered; however, no differences between calibration slopes were observed for data collected within the range of expected in situ pressure gradients. In addition, dye perfusions showed that in three of the four ring-porous species considered, active sapwood was limited to the outermost growth ring. In contrast, our calibration results for diffuse-porous species showed generally good agreement with the empirically derived Granier calibration, and dye perfusions showed that active sapwood was associated with many annual growth rings. Our results suggest that the original calibration of Granier is not universally applicable to all species and xylem types and that previous estimates of absolute rates of water use for ring-porous species obtained using the original calibration coefficients may be associated with substantial error.
    Tree Physiology 12/2010; 30(12):1545-54. · 2.88 Impact Factor
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    Article: Testing hypotheses that link wood anatomy to cavitation resistance and hydraulic conductivity in the genus Acer.
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    ABSTRACT: • Vulnerability to cavitation and conductive efficiency depend on xylem anatomy. We tested a large range of structure-function hypotheses, some for the first time, within a single genus to minimize phylogenetic 'noise' and maximize detection of functionally relevant variation. • This integrative study combined in-depth anatomical observations using light, scanning and transmission electron microscopy of seven Acer taxa, and compared these observations with empirical measures of xylem hydraulics. • Our results reveal a 2 MPa range in species' mean cavitation pressure (MCP). MCP was strongly correlated with intervessel pit structure (membrane thickness and porosity, chamber depth), weakly correlated with pit number per vessel, and not related to pit area per vessel. At the tissue level, there was a strong correlation between MCP and mechanical strength parameters, and some of the first evidence is provided for the functional significance of vessel grouping and thickenings on inner vessel walls. In addition, a strong trade-off was observed between xylem-specific conductivity and MCP. Vessel length and intervessel wall characteristics were implicated in this safety-efficiency trade-off. • Cavitation resistance and hydraulic conductivity in Acer appear to be controlled by a very complex interaction between tissue, vessel network and pit characteristics.
    New Phytologist 11/2010; 190(3):709-23. · 6.64 Impact Factor
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    Article: Contrasting drought tolerance strategies in two desert annuals of hybrid origin.
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    ABSTRACT: Woody plants native to mesic habitats tend to be more vulnerable to drought-induced cavitation than those in xeric habitats. Cavitation resistance in herbaceous plants, however, is rarely studied and whether or not annual plants in arid habitats conform to the trends observed in woody plants is unknown. This question is addressed by comparing the hydraulic properties of annual plants endemic to relatively mesic and seasonally xeric habitats in the Great Basin Desert, in both native and experimental settings. Vulnerability to cavitation between species differed as predicted when vulnerability curves of similar-sized native individuals were compared. Contrary to expectations, Helianthus anomalus from the relatively mesic dune sites, on average, exhibited higher native embolism, lower soil-to-leaf hydraulic conductance (k(L)) and lower transpiration rates, than its xeric analogue, H. deserticola. In transplant gardens, H. anomalus' vulnerability to cavitation was unaffected by transplant location or watering treatment. In H. deserticola, however, vulnerability to cavitation varied significantly in response to watering in transplant gardens and varied as a function of stem water potential (Psi(stem)). H. deserticola largely avoided cavitation through its higher water status and generally more resistant xylem, traits consistent with a short life cycle and typical drought-escape strategy. By contrast, H. anomalus' higher native embolism is likely to be adaptive by lowering plant conductance and transpiration rate, thus preventing the loss of root-to-soil hydraulic contact in the coarse sand dune soils. For H. anomalus this dehydration avoidance strategy is consistent with its relatively long 3-4 month life cycle and low-competition habitat. We conclude that variance of hydraulic parameters in herbaceous plants is a function of soil moisture heterogeneity and is consistent with the notion that trait plasticity to fine-grained environmental variation can be adaptive.
    Journal of Experimental Botany 06/2010; 61(10):2769-78. · 5.36 Impact Factor
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    Article: Moving water well: comparing hydraulic efficiency in twigs and trunks of coniferous, ring-porous, and diffuse-porous saplings from temperate and tropical forests.
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    ABSTRACT: *Coniferous, diffuse-porous and ring-porous trees vary in their xylem anatomy, but the functional consequences of these differences are not well understood from the scale of the conduit to the individual. *Hydraulic and anatomical measurements were made on branches and trunks from 16 species from temperate and tropical areas, representing all three wood types. Scaling of stem conductivity (K(h)) with stem diameter was used to model the hydraulic conductance of the stem network. *Ring-porous trees showed the steepest increase in K(h) with stem size. Temperate diffuse-porous trees were at the opposite extreme, and conifers and tropical diffuse-porous species were intermediate. Scaling of K(h) was influenced by differences in the allometry of conduit diameter (taper) and packing (number per wood area) with stem size. *The K(h) trends were mirrored by the modeled stem-network conductances. Ring-porous species had the greatest network conductance and this value increased isometrically with trunk basal area, indicating that conductance per unit sapwood was independent of tree size. Conductances were lowest and most size-dependent in conifers. The results indicate that differences in conduit taper and packing between functional types propagate to the network level and have an important influence on metabolic scaling concepts.
    New Phytologist 02/2010; 186(2):439-50. · 6.64 Impact Factor
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    Article: Single-vessel flow measurements indicate scalariform perforation plates confer higher flow resistance than previously estimated.
    Mairgareth A Christman, John S Sperry
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    ABSTRACT: During vessel evolution in angiosperms, scalariform perforation plates with many slit-like openings transformed into simple plates with a single circular opening. The transition is hypothesized to have resulted from selection for decreased hydraulic resistance. Previously, additional resistivity of scalariform plates was estimated to be small - generally 10% or less above lumen resistivity - based on numerical and physical models. Here, using the single-vessel technique, we directly measured the hydraulic resistance of individual xylem vessels. The resistivity of simple-plated lumens was not significantly different from the Hagen-Poiseuille (HP) prediction (+6 + or - 3.3% mean deviation). In the 13 scalariform-plated species measured, plate resistivity averaged 99 + or - 13.7% higher than HP lumen resistivity. Scalariform species also showed higher resistivity than simple species at the whole vessel (+340%) and sapwood (+580%) levels. The strongest predictor of scalariform plate resistance was vessel diameter (r(2) = 0.84), followed by plate angle (r(2) = 0.60). An equation based on laminar flow through periodic slits predicted single-vessel measurements reasonably well (r(2) = 0.79) and indicated that Baileyan trends in scalariform plate evolution maintain an approximate balance between lumen and plate resistances. In summary, we found scalariform plates of diverse morphology essentially double lumen flow resistance, impeding xylem flow much more than previously estimated.
    Plant Cell and Environment 11/2009; 33(3):431-43. · 5.22 Impact Factor
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    Article: Murray's law, the 'Yarrum' optimum, and the hydraulic architecture of compound leaves.
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    ABSTRACT: There are two optima for maximizing hydraulic conductance per vasculature volume in plants. Murray's law (ML) predicts the optimal conduit taper for a fixed change in conduit number across branch ranks. The opposite, the Yarrum optimum (YO), predicts the optimal change in conduit number for a fixed taper. We derived the solution for YO and then evaluated compliance with both optima within the xylem of compound leaves, where conduits should have a minimal mechanical role. We sampled leaves from temperate ferns, and tropical and temperate angiosperms Leaf vasculature exhibited greater agreement with ML than YO. Of the 14 comparisons in 13 species, 12 conformed to ML. The clear tendency towards ML indicates that taper is optimized for a constrained conduit number. Conduit number may be constrained by leaflet number, safety requirements, and the fact that the number of conduits is established before their diameter during development. Within a leaf, ML compliance requires leaf-specific conductivity to decrease from petiole to petiolule with the decrease in leaf area supplied. A similar scaling applied across species, indicating lower leaf-specific petiole conductivity in smaller leaves. Small leaf size should offset lower conductivity, and petiole conductance (conductivity/length) may be independent of leaf size.
    New Phytologist 08/2009; 184(1):234-44. · 6.64 Impact Factor
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    Article: Testing the 'rare pit' hypothesis for xylem cavitation resistance in three species of Acer.
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    ABSTRACT: Eudicot angiosperms with greater vulnerability to xylem cavitation tend to have vessels with greater total area of inter-vessel pits, which inspired the 'rare pit' hypothesis: the more pits per vessel, by chance the leakier will be the vessel's single air-seeding pit and the lower the air-seeding threshold for cavitation to spread between vessels. Here, we demonstrate the feasibility of the hypothesis, using probability theory to model the axial propagation of air through air-injected stems. In the presence of rare, leaky pits, air-seeding pressures through short stems with few vessel ends in series should be low; pressures should increase in longer stems as more end-walls must be breached. Measurements on three Acer species conformed closely to model predictions, confirming the rare presence of leaky pits. The model indicated that pits air-seeding at or below the mean cavitation pressure (MCP) occurred at similarly low frequencies in all species. Average end-wall air-seeding pressures predicted by the model closely matched species' MCPs. Differences in species' vulnerability were primarily attributed to differences in frequency of the leakiest pits rather than pit number or area per vessel. Adjustments in membrane properties and extent of pitting per vessel apparently combine to influence cavitation resistance across species.
    New Phytologist 02/2009; 182(3):664-74. · 6.64 Impact Factor
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    Article: A case-study of water transport in co-occurring ring- versus diffuse-porous trees: contrasts in water-status, conducting capacity, cavitation and vessel refilling.
    Haruhiko Taneda, John S Sperry
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    ABSTRACT: Recent work has suggested that the large earlywood vessels of ring-porous trees can be extraordinarily vulnerable to cavitation making it necessary that these trees maintain a consistent and favorable water status. We compared cavitation resistance, vessel refilling, transport capacity and water status in a study of ring-porous Quercus gambelii Nutt. (oak) and diffuse-porous Acer grandidentatum Nutt. (maple). These species co-dominate summer-dry foothills in the western Rocky Mountains of the USA. Native embolism measurements, dye perfusions and balance pressure exudation patterns indicated that the large earlywood vessels of 2-3-year-old oak stems cavitated extensively on a daily basis as predicted from laboratory vulnerability curves, resulting in a more than 80% reduction in hydraulic conductivity. Maple branches showed virtually no cavitation. Oak vessels refilled on a daily basis, despite negative xylem pressure in the transpiration stream, indicating active pressurization of embo-lized vessels. Conductivity and whole-tree water use in oak were between about one-half and two-thirds that in maple on a stem-area basis; but were similar or greater on a leaf-area basis. Oak maintained steady and modest negative xylem pressure potentials during the growing season despite little rainfall, indicating isohydric water status and reliance on deep soil water. Maple was markedly anisohydric and developed more negative pressure potentials during drought, suggesting use of shallower soil water. Although ring porosity may have evolved as a mechanism for coping with winter freezing, this study suggests that it also has major consequences for xylem function during the growing season.
    Tree Physiology 12/2008; 28(11):1641-51. · 2.88 Impact Factor
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    Article: Safety and efficiency conflicts in hydraulic architecture: scaling from tissues to trees.
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    ABSTRACT: Tree hydraulic architecture exhibits patterns that propagate from tissue to tree scales. A challenge is to make sense of these patterns in terms of trade-offs and adaptations. The universal trend for conduits per area to decrease with increasing conduit diameter below the theoretical packing limit may reflect the compromise between maximizing the area for conduction versus mechanical support and storage. Variation in conduit diameter may have two complementary influences: one being compromises between efficiency and safety and the other being that conduit tapering within a tree maximizes conductance per growth investment. Area-preserving branching may be a mechanical constraint, preventing otherwise more efficient top-heavy trees. In combination, these trends beget another: trees have more, narrower conduits moving from trunks to terminal branches. This pattern: (1) increases the efficiency of tree water conduction; (2) minimizes (but does not eliminate) any hydraulic limitation on the productivity or tissue growth with tree height; and (3) is consistent with the scaling of tree conductance and sap flow with size. We find no hydraulic reason why tree height should scale with a basal diameter to the two-thirds power as recently claimed; it is probably another mechanical constraint as originally proposed. The buffering effect of capacitance on the magnitude of transpiration-induced xylem tension appears to be coupled to cavitation resistance, possibly alleviating safety versus efficiency trade-offs.
    Plant Cell and Environment 06/2008; 31(5):632-45. · 5.22 Impact Factor
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    Article: Wood anatomy constrains stomatal responses to atmospheric vapor pressure deficit in irrigated, urban trees.
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    ABSTRACT: Plant transpiration is strongly constrained by hydraulic architecture, which determines the critical threshold for cavitation. Because species vary greatly in vulnerability to cavitation, hydraulic limits to transpiration and stomatal conductance have not generally been incorporated into ecological and climate models. We measured sap flow, leaf transpiration, and vulnerability to cavitation of a variety of tree species in a well-irrigated but semi-arid urban environment in order to evaluate the generality of stomatal responses to high atmospheric vapor pressure deficit (D). We found evidence of broad patterns of stomatal responses to humidity based on systematic differences in vulnerability to cavitation. Ring-porous taxa consistently had vulnerable xylem and showed strong regulation of transpiration in response to D, while diffuse-porous taxa were less vulnerable and transpiration increased nearly linearly with D. These results correspond well to patterns in the distribution of the taxa, such as the prevalence of diffuse-porous species in riparian ecosystems, and also provide a means of representing maximum transpiration rates at varying D in broad categories of trees.
    Oecologia 06/2008; 156(1):13-20. · 3.41 Impact Factor