Search

Ecohydrology and invasive ecology have become increasingly important in the context of global climate change. This study presents the first in-depth analysis of the water use of invasive and native plants of the same growth form at multiple scales: leaf, plant, and ecosystem. We reanalyzed data for several hundred native and invasive species from over 40 published studies worldwide to glean global trends and to highlight how patterns vary depending on both scale and climate. We analyzed all pairwise combinations of co-occurring native and invasive species for higher comparative resolution of the likelihood of an invasive species using more water than a native species and tested for significance using bootstrap methods. At each scale, we found several-fold differences in water use between specific paired invasive and native species. At the leaf scale, we found a strong tendency for invasive species to have greater stomatal conductance than native species. At the plant scale, however, natives and invasives were equally likely to have the higher sap flow rates. Available data were much fewer for the ecosystem scale; nevertheless, we found that invasive-dominated ecosystems were more likely to have higher sap flow rates per unit ground area than native-dominated ecosystems. Ecosystem-scale evapotranspiration, on the other hand, was equally likely to be greater for systems dominated by invasive and native species of the same growth form. The inherent disconnects in the determination of water use when changing scales from leaf to plant to ecosystem reveal hypotheses for future studies and a critical need for more ecosystem-scale water use measurements in invasive- vs. native-dominated systems. The differences in water use of native and invasive species also depended strongly on climate, with the greater water use of invasives enhanced in hotter, wetter climates at the coarser scales.

Leaf venation architecture is tremendously diverse across plant species. Understanding the hydraulic functions of given venation traits can clarify the organization of the vascular system and its adaptation to environment. Using a spatially explicit model (the program K_leaf), we subjected realistic simulated leaves to modifications and calculated the impacts on xylem and leaf hydraulic conductance (K(x) and K(leaf), respectively), important traits in determining photosynthesis and growth. We tested the sensitivity of leaves to altered vein order conductivities (1) in the absence or (2) presence of hierarchical vein architecture, (3) to major vein tapering, and (4) to modification of vein densities (length/leaf area). The K(x) and K(leaf) increased with individual vein order conductivities and densities; for hierarchical venation systems, the greatest impact was from increases in vein conductivity for lower vein orders and increases in density for higher vein orders. Individual vein order conductivities were colimiting of K(x) and K(leaf), as were their densities, but the effects of vein conductivities and densities were orthogonal. Both vein hierarchy and vein tapering increased K(x) relative to xylem construction cost. These results highlight the important consequences of venation traits for the economics, ecology, and evolution of plant transport capacity.

Mosses are an understudied group of plants that can potentially confirm or expand principles of plant function described for tracheophytes, from which they diverge strongly in structure. We quantified 35 physiological and morphological traits from cell-, leaf- and canopy-level, for 10 ground-, trunk- and branch-dwelling Hawaiian species. We hypothesized that trait values would reflect the distinctive growth form and slow growth of mosses, but also that trait correlations would be analogous to those of tracheophytes. The moss species had low leaf mass per area and low gas exchange rate. Unlike for tracheophytes, light-saturated photosynthetic rate per mass (A(mass)) did not correlate with habitat irradiance. Other photosynthetic parameters and structural traits were aligned with microhabitat irradiance, driving an inter-correlation of traits including leaf area, cell size, cell wall thickness, and canopy density. In addition, we found a coordination of traits linked with structural allocation, including costa size, canopy height and A(mass). Across species, A(mass) and nitrogen concentration correlated negatively with canopy mass per area, analogous to linkages found for the 'leaf economic spectrum', with canopy mass per area replacing leaf mass per area. Despite divergence of mosses and tracheophytes in leaf size and function, analogous trait coordination has arisen during ecological differentiation.

Tropical forests are becoming increasingly alien-dominated through the establishment of timber plantations and secondary forests. Despite widespread recognition that afforestation results in increased evapotranspiration and lower catchment yields, little is known of the impacts of timber plantations on water balance relative to native forest. Native forest trees have been claimed to use water conservatively and enhance groundwater recharge relative to faster-growing alien species, and this argument should motivate native forest preservation and restoration. However, data have been available primarily for leaf-level gas exchange rather than for whole-plant and stand levels. We measured sap flow of dominant tree and tree fern species over eight weeks in native Metrosideros polymorpha forest and adjacent alien timber plantations on the island of Hawai'i and estimated total stand transpiration. Metrosideros polymorpha had the lowest values of sap flux density and whole-tree water use (200 kg m(-2) sapwood d(-1), or 8 kg/d for trees of 35 cm mean diameter at breast height, D), substantially less than timber species Eucalyptus saligna or Fraxinus uhdei (33 and 34 kg/d for trees of 73 and 30 cm mean D, respectively). At the stand level, E. saligna and F. uhdei trees had three- and ninefold higher water use, respectively, than native M. polymorpha trees. Understory Cibotium tree ferns were most abundant in M. polymorpha-dominated forest where they accounted for 70% of water use. Overall, F. uhdei plantation had the highest water use at 1.8 mm/d, more than twice that of either E. saligna plantation or M. polymorpha forest. Forest water use was influenced by species composition, stem density, tree size, sapwood allocation, and understory contributions. Transpiration varied strongly among forest types even within the same wet tropical climate, and in this case, native forest had strikingly conservative water use. Comparisons of vegetation cover in water use should provide additional resolution to ecosystem valuation and land management decisions.

Previous studies have shown a rapid enhancement in leaf hydraulic conductance (K(leaf)) from low to high irradiance (from <10 to >1000 micromol photons m(-2) s(-1)), using the high-pressure flow meter (HPFM), for 7 of 14 tested woody species. However, theoretical suggestions have been made that this response might arise as an artifact of the HPFM. We tested the K(leaf) light response for six evergreen species using refined versions of the rehydration kinetics method (RKM) and the evaporative flux method (EFM). We found new evidence for a rapid, 60% to 100% increase in K(leaf) from low to high irradiance for three species. In the RKM, the leaf rehydration time constant declined by up to 70% under high irradiance relative to darkness. In the EFM, under higher irradiance, the flow rate increased disproportionately to the water potential gradient. Combining our data with those of previous studies, we found that heterobaric species, i.e. those with bundle sheath extensions (BSEs) showed a twofold greater K(leaf) light response on average than homobaric species, i.e. those without BSEs. We suggest further research to characterize this substantial dynamic at the nexus of plant light- and water-relations.

Understanding the impacts of combined resource supplies on seedlings is critical to enable prediction of establishment growth, and forest dynamics. We investigated the effects of irradiance and water treatments on absolute growth, and relative growth rate (RGR) and its components, for seedlings of four Quercus species differing in leaf habit and with a wide variation in seed mass. Plants were grown for 6.5 months at three levels of irradiance (100, 27, and 3% daylight), and treated during the last 2.5 months with two watering treatments (frequent watering v. suspended watering). Both shade and drought reduced seedling growth rates, with a significant interaction: under full irradiance the drought treatment had a stronger impact on RGR and final biomass than under deep shade. For three species, seed mass was positively related to absolute growth, with stronger correlations at lower irradiance. The evergreen species grew faster than the deciduous species, though leaf habit accounted for a minor part of the interspecific variation in absolute growth. Seedling biomass was determined positively either by RGR or seed mass; RGR was positively linked with net assimilation rate (NAR) and leaf mass fraction (LMF), and seed mass was negatively linked with RGR and LMF, but positively linked with NAR. Seedling RGR was not correlated with light-saturated net photosynthetic rate, but was strongly correlated with the net carbon balance estimated, from photosynthetic light-response curves, considering daily variation in irradiance. These findings suggest an approach to applying short-term physiological measurements to predict the RGR and absolute growth rate of seedlings in a wide range of combinations of irradiance and water supplies.

General models of plant vascular architecture, based on scaling of pipe diameters to remove the length dependence of hydraulic resistance within the xylem, have attracted strong interest. However, these models have neglected to consider the leaf, an important hydraulic component; they assume all leaves to have similar hydraulic properties, including similar pipe diameters in the petiole. We examine the scaling of the leaf xylem in 10 temperate oak species, an important hydraulic component. The mean hydraulic diameter of petiole xylem vessels varied by 30% among the 10 oak species. Conduit diameters narrowed from the petiole to the midrib to the secondary veins, consistent with resistance minimization, but the power function scaling exponent differed from that predicted for stems. Leaf size was an organizing trait within and across species. These findings indicate that leaf vasculature needs to be included in whole-plant scaling models, for these to accurately reflect and predict whole-plant transport and its implications for performance and ecology.

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 higher-order 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.

Coordinated variation has been reported for leaf structure, composition and function, across and within species, and theoretically should occur across populations of a species that span an extensive environmental range. We focused on Hawaiian keystone tree species Metrosideros polymorpha, specifically, 13-year old trees grown (2-4 m tall) in a common garden (approximately 1 ha field with 2-3 m between trees) from seeds collected from 14 populations along an altitude-soil age gradient. We determined the genetic component of relationships among specific leaf area (SLA), the concentrations of nitrogen (N) and pigments (chlorophylls, carotenoids, and anthocyanins), and photosynthetic light-use efficiency. These traits showed strong ecotypic variation; SLA declined 35% with increasing source elevation, and area-based concentrations of N, Chl a + b and Car increased by 50, 109 and 96%, respectively. Concentrations expressed on a mass basis were not well related to source elevation. Pigment ratios expressed covariation that suggested an increased capacity for light harvesting at higher source elevation; Chl/N and Car/Chl increased with source elevation, whereas Chl a/b declined; Chl a/b was higher for populations on younger soil, suggesting optimization for low N supply. Parallel trends were found for the photosynthetic reactions; light-saturated quantum yield of photosystem II (Phi (PSII)) and electron transport rate (ETR) increased with source elevation. Correlations of the concentrations of photosynthetic pigments, pigment ratios, and photosynthetic function across the ecotypes indicated a stoichiometric coordination of the components of the light-harvesting antennae and reaction centers. The constellation of coordinated morphological, biochemical and physiological properties was expressed in the leaf reflectance and transmittance properties in the visible and near-infrared wavelength region (400-950 nm), providing an integrated metric of leaf status among and between plant phenotypes.

Inter- and intraspecific variation in hydraulic traits was investigated in nine Cordia (Boraginaceae) species growing in three tropical rainforests differing in mean annual precipitation (MAP). Interspecific variation was examined for the different Cordia species found at each site, and intraspecific variation was studied in populations of the widespread species Cordia alliodora across the three sites. Strong intra- and interspecific variation were observed in vulnerability to drought-induced embolism. Species growing at drier sites were more resistant to embolism than those growing at moister sites; the same pattern was observed for populations of C. alliodora. By contrast, traits related to hydraulic capacity, including stem xylem vessel diameter, sapwood specific conductivity (K(s)) and leaf specific conductivity (K(L)), varied strongly but independently of MAP. For C. alliodora, xylem anatomy, K(s), K(L) and Huber value varied little across sites, with K(s) and K(L) being consistently high relative to other Cordia species. A constitutively high hydraulic capacity coupled with plastic or genotypic adjustment in vulnerability to embolism and leaf water relations would contribute to the ability of C. alliodora to establish and compete across a wide precipitation gradient.

The hydraulic resistance of the leaf (R1) is a major bottleneck in the whole plant water transport pathway and may thus be linked with the enormous variation in leaf structure and function among tropical rain forest trees. A previous study found that R1 varied by an order of magnitude across 10 tree species of Panamanian tropical lowland rain forest. Here, correlations were tested between R1 and 24 traits relating to leaf venation and mesophyll structure, and to gross leaf form. Across species, R1 was related to both venation architecture and mesophyll structure. R1 was positively related to the theoretical axial resistivity of the midrib, determined from xylem conduit numbers and dimensions, and R1 was negatively related to venation density in nine of 10 species. R1 was also negatively related to both palisade mesophyll thickness and to the ratio of palisade to spongy mesophyll. By contrast, numerous leaf traits were independent of R1, including area, shape, thickness, and density, demonstrating that leaves can be diverse in gross structure without intrinsic trade-offs in hydraulic capacity. Variation in both R1-linked and R1-independent traits related strongly to regeneration irradiance, indicating the potential importance of both types of traits in establishment ecology.

Leaves are extraordinarily variable in form, longevity, venation architecture, and capacity for photosynthetic gas exchange. Much of this diversity is linked with water transport capacity. The pathways through the leaf constitute a substantial (>or=30%) part of the resistance to water flow through plants, and thus influence rates of transpiration and photosynthesis. Leaf hydraulic conductance (K(leaf)) varies more than 65-fold across species, reflecting differences in the anatomy of the petiole and the venation architecture, as well as pathways beyond the xylem through living tissues to sites of evaporation. K(leaf) is highly dynamic over a range of time scales, showing circadian and developmental trajectories, and responds rapidly, often reversibly, to changes in temperature, irradiance, and water supply. This review addresses how leaf structure and physiology influence K(leaf), and the mechanisms by which K(leaf) contributes to dynamic functional responses at the level of both individual leaves and the whole plant.

The transport of water, sugar, and nutrients in trees is restricted to specific vascular pathways, and thus organs may be relatively isolated from one another (i.e. sectored). Strongly sectored leaf-to-leaf pathways have been shown for the transport of sugar and signal molecules within a shoot, but not previously for water transport. The hydraulic sectoriality of leaf-to-leaf pathways was determined for current year shoots of six temperate deciduous tree species (three ring-porous: Castanea dentata, Fraxinus americana, and Quercus rubra, and three diffuse-porous: Acer saccharum, Betula papyrifera, and Liriodendron tulipifera). Hydraulic sectoriality was determined using dye staining and a hydraulic method. In the dye method, leaf blades were removed and dye was forced into the most proximal petiole. For each petiole the vascular traces that were shared with the proximal petiole were counted. For other shoots, measurements were made of the leaf-area-specific hydraulic conductivity for the leaf-to-leaf pathways (k(LL)). In five out of the six species, patterns of sectoriality reflected phyllotaxy; both the sharing of vascular bundles between leaves and k(LL) were higher for orthostichous than non-orthostichous leaf pairs. For each species, leaf-to-leaf sectoriality was determined as the proportional differences between non-orthostichous versus orthostichous leaf pairs in their staining of shared vascular bundles and in their k(LL); for the six species these two indices of sectoriality were strongly correlated (R2=0.94; P <0.002). Species varied 8-fold in their k(LL)-based sectoriality, and ring-porous species were more sectored than diffuse-porous species. Differential leaf-to-leaf sectoriality has implications for species-specific co-ordination of leaf gas exchange and water relations within a branch, especially during fluctuations in irradiance and water and nutrient availability.

The leaf hydraulic conductance (K(leaf)) is a major determinant of plant water transport capacity. Here, we measured K(leaf), and its basis in the resistances of leaf components, for fully illuminated leaves of five tree species that regenerate in deep shade, and five that regenerate in gaps or clearings, in Panamanian lowland tropical rainforest. We also determined coordination with stomatal characters and leaf mass per area. K(leaf) varied 10-fold across species, and was 3-fold higher in sun- than in shade-establishing species. On average, 12% of leaf hydraulic resistance (= 1/K(leaf)) was located in the petiole, 25% in the major veins, 25% in the minor veins, and 39% outside the xylem. Sun-establishing species had a higher proportion of leaf resistance in the xylem. Across species, component resistances correlated linearly with total leaf resistance. K(leaf) correlated tightly with indices of stomatal pore area, indicating a coordination of liquid- and vapor-phase conductances shifted relative to that of temperate woody species. Leaf hydraulic properties are integrally linked in the complex of traits that define differences in water use and carbon economy across habitats and vegetation zones., (Copyright New Phytologist (2005).)

This paper examines the dependence of whole leaf hydraulic conductance to liquid water (K(L)) on irradiance when measured with a high pressure flowmeter (HPFM). During HPFM measurements, water is perfused into leaves faster than it evaporates hence water infiltrates leaf air spaces and must pass through stomates in the liquid state. Since stomates open and close under high versus low irradiance, respectively, the possibility exists that K(L) might change with irradiance if stomates close tightly enough to restrict water movement. However, the dependence of K(L) on irradiance could be due to a direct effect of irradiance on the hydraulic properties of other tissues in the leaf. In the present study, K(L) increased with irradiance for 6 of the 11 species tested. Whole leaf conductance to water vapour, g(L), was used as a proxy for stomatal aperture and the time-course of changes in K(L) and g(L) was studied during the transition from low to high irradiance and from high to low irradiance. Experiments showed that in some species K(L) changes were not paralleled by g(L) changes. Measurements were also done after perfusion of leaves with ABA which inhibited the g(L) response to irradiance. These leaves showed the same K(L) response to irradiance as control leaves. These experimental results and theoretical calculations suggest that the irradiance dependence of K(L) is more consistent with an effect on extravascular (and/or vascular) tissues rather than stomatal aperture. Irradiance-mediated stimulation of aquaporins or hydrogel effects in leaf tracheids may be involved.

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.

Many leaves survive the severing of their major veins in apparently excellent health. According to the classical explanation, the leaf minor veins provide "conductive overload," an excess of parallel conductive paths, rendering the major veins hydraulically dispensable. Whether such an excess of conductive paths exists has important implications for vascular design and for leaf response to vascular damage. We subjected leaves of Acer saccharum and Quercus rubra to cutting treatments that disrupted the major vein system and determined leaf survival, stomatal conductance (g), quantum yield of photosystem II (Φ(PSII)), and leaf hydraulic conductance (K(leaf)). For A. saccharum, the cuts led to the death of distal lamina. For Q. rubra, however, the treated leaves typically remained apparently healthy. Despite their appearance, the treated Q. rubra leaves had a strongly reduced K(leaf), relative to control leaves, and g and Φ(PSII) were reduced distal to the cuts, respectively, by 75-97% and 48-76%. Gas exchange proximal to the cuts was unaffected, indicating the independence of lamina regions and their local stomata. Analogous results were obtained with excised Q. rubra leaves. These studies demonstrate an indispensable, vital role of the major veins in conducting water throughout the lamina.

To test whether the impact of drought on the growth and biomass allocation of first-season shade-tolerant woody seedlings in low irradiance differs from that in high irradiance, seedlings of Viburnum lantana, V. opulus, V. tinus and Hedera helix were grown in pots at two watering frequencies × three irradiances. Hypotheses in the recent literature variously predict that drought will have a stronger, weaker or equal impact on seedling relative growth rate (RGR) in deep shade relative to that in moderate shade. Experimental irradiance levels were selected in the typical range for temperate deciduous forest seedlings in either understorey or clearings: 3-4% daylight (low red: far-red shade), 3-4% daylight (neutral shade), and 30-40% daylight (neutral shade). Watering was 'frequent' (every 3-4 days) or 'infrequent' (five times during the 8-week experiment), producing soil matric potentials as low as -0.03 MPa, and -2 MPa. To prevent the interaction of irradiance and watering treatments, each seedling was grown in a 'shade tower' that was surrounded by an uncovered sward of grass (Festuca rubra), which depleted pot water at the same rate regardless of the species of seedling, or its irradiance treatment. Shading affected all species: seedlings in 3.5% daylight grew at 56-73% of their dry-mass RGR in 35% daylight. Low red: far-red shade reduced the RGR of Hedera to 68% of its value in neutral shade. Infrequent watering significantly reduced the RGR of only V. lantana and V. opulus, by approximately the same proportion across irradiance treatments. Infrequent watering did not significantly alter any species' biomass allocation across irradiance treatments. Such orthogonal impacts of deep shade and drought on seedling growth and biomass allocation indicate a large potential for niche differentiation at combinations of irradiance and water supply for species of forest seedlings, and suggest a multiplicative-effects approach for modelling seedling performance in microsites with different combinations of irradiance and water supply.

Recent cutbacks in private insurance benefits and publicly funded health programs are threatening the access of Americans, particularly the poor and the disabled, to private psychiatric care. The authors present the findings of three surveys that assessed the impact of threatened or actual cuts in third-party payments on the treatment provided by private practitioners. One survey indicated that, contrary to popular opinion, more than half of the psychiatrists in Northern California see severely disturbed Medicaid patients and that they provide outpatient psychiatric services to them for less cost than would public clinics. Two surveys conducted in the Washington, D.C., area indicated that both patients and psychiatrists have suffered from insurance cutbacks, with fewer patients being able to afford intensive private treatment and psychiatrists using reduced fees and less optimal treatment modalities. The fiscal and ethical dilemma posed by the cutbacks and further research needs are also explored.