Hydraulic constraints modify optimal photosynthetic profiles in giant sequoia trees.

Optimality theory states that whole-tree carbon gain is maximized when leaf N and photosynthetic capacity profiles are distributed along vertical light gradients such that the marginal gain of nitrogen investment is identical among leaves. However, observed photosynthetic N gradients in trees do not follow this prediction, and the causes for this apparent discrepancy remain uncertain. Our objective was to evaluate how hydraulic limitations potentially modify crown-level optimization in Sequoiadendron giganteum (giant sequoia) trees up to 90 m tall. Leaf water potential (Ψ l ) and branch sap flow closely followed diurnal patterns of solar radiation throughout each tree crown. Minimum leaf water potential correlated negatively with height above ground, while leaf mass per area (LMA), shoot mass per area (SMA), leaf nitrogen content (%N), and bulk leaf stable carbon isotope ratios (δ(13)C) correlated positively with height. We found no significant vertical trends in maximum leaf photosynthesis (A), stomatal conductance (g s), and intrinsic water-use efficiency (A/g s), nor in branch-averaged transpiration (E L), stomatal conductance (G S), and hydraulic conductance (K L). Adjustments in hydraulic architecture appear to partially compensate for increasing hydraulic limitations with height in giant sequoia, allowing them to sustain global maximum summer water use rates exceeding 2000 kg day(-1). However, we found that leaf N and photosynthetic capacity do not follow the vertical light gradient, supporting the hypothesis that increasing limitations on water transport capacity with height modify photosynthetic optimization in tall trees.

Environmental controls in the water use patterns of a tropical cloud forest tree species, Drimys brasiliensis (Winteraceae).

Trees from tropical montane cloud forest (TMCF) display very dynamic patterns of water use. They are capable of downwards water transport towards the soil during leaf-wetting events, likely a consequence of foliar water uptake (FWU), as well as high rates of night-time transpiration (Enight) during drier nights. These two processes might represent important sources of water losses and gains to the plant, but little is known about the environmental factors controlling these water fluxes. We evaluated how contrasting atmospheric and soil water conditions control diurnal, nocturnal and seasonal dynamics of sap flow in Drimys brasiliensis (Miers), a common Neotropical cloud forest species. We monitored the seasonal variation of soil water content, micrometeorological conditions and sap flow of D. brasiliensis trees in the field during wet and dry seasons. We also conducted a greenhouse experiment exposing D. brasiliensis saplings under contrasting soil water conditions to deuterium-labelled fog water. We found that during the night D. brasiliensis possesses heightened stomatal sensitivity to soil drought and vapour pressure deficit, which reduces night-time water loss. Leaf-wetting events had a strong suppressive effect on tree transpiration (E). Foliar water uptake increased in magnitude with drier soil and during longer leaf-wetting events. The difference between diurnal and nocturnal stomatal behaviour in D. brasiliensis could be attributed to an optimization of carbon gain when leaves are dry, as well as minimization of nocturnal water loss. The leaf-wetting events on the other hand seem important to D. brasiliensis water balance, especially during soil droughts, both by suppressing tree transpiration (E) and as a small additional water supply through FWU. Our results suggest that decreases in leaf-wetting events in TMCF might increase D. brasiliensis water loss and decrease its water gains, which could compromise its ecophysiological performance and survival during dry periods.

Influence of stem temperature changes on heat pulse sap flux density measurements.

While natural spatial temperature gradients between measurement needles have been thoroughly investigated for continuous heat-based sap flow methods, little attention has been given to how natural changes in stem temperature impact heat pulse-based methods through temporal rather than spatial effects. By modelling the theoretical equation for both an ideal instantaneous pulse and a step pulse and applying a finite element model which included actual needle dimensions and wound effects, the influence of a varying stem temperature on heat pulse-based methods was investigated. It was shown that the heat ratio (HR) method was influenced, while for the compensation heat pulse and Tmax methods changes in stem temperatures of up to 0.002 °C s(-1) did not lead to significantly different results. For the HR method, rising stem temperatures during measurements led to lower heat pulse velocity values, while decreasing stem temperatures led to both higher and lower heat pulse velocities, and to imaginary results for high flows. These errors of up to 40% can easily be prevented by including a temperature correction in the data analysis procedure, calculating the slope of the natural temperature change based on the measured temperatures before application of the heat pulse. Results of a greenhouse and outdoor experiment on Pinus pinea L. show the influence of this correction on low and average sap flux densities.

Regiospecific assembly of gold nanoparticles around the pores of diatoms: toward three-dimensional nanoarrays.

The templated growth of gold nanoparticles in 3D arrays within the nanopores of unicellular diatoms involves pretreament of the skeletons with poly(vinylpyridine) which has a unique dewetting property. This self-assembly provides a nanochemical analogue of lithography for engineering complex nanostructures. The process can be universally applied to the many types of diatom skeletons which vary in size and structure.

Using multiple trait associations to define hydraulic functional types in plant communities of south-western Australia.

Assessing the hydrological imbalance and associated land degradation issues facing much of southern Australia and other parts of the world requires a better understanding of the defining features of ecosystem water use and the design of sustainable agroecosystems. Thus, by grouping species with similar water-use strategies into 'hydraulic functional types' (HFTs), we investigated the characteristics of water use in species-rich plant communities of south-western Australia. HFTs were determined using multiple-trait associations between morphological and physiological traits relating to water transport, water-use efficiency and response to water deficit. Sixteen traits were assessed from a subset of 21 species from three plant communities located along a topographically determined soil- and water-availability gradient. Multivariate analyses showed that trait variation was least at sites with shallower soils and putatively lower water availability, suggesting a convergence of water-use strategies at sites where plants are exposed to large seasonal water deficits. Stem hydraulic parameters, including stem-specific hydraulic conductivity, conduit diameter and maximum percentage embolism, were positively correlated, indicating the generality that larger conduit diameter permits greater hydraulic efficiency and is associated with greater seasonal reductions in hydraulic conductivity in this ecosystem. Wood density was not correlated with these traits, but closely associated with species' ability to withstand more negative water potentials during summer. Long-term integrated water-use efficiency was lower in shallow-rooted species that exhibited more negative summer water potentials. Specific leaf area and minimum leaf water potential were correlated with a number of separate traits, and appear to represent key axes of trait variation that describe the water-use strategies of different HFTs along the topographic gradient. Five HFTs were classified using a resemblance analysis according to combinations of traits that pertain to different water-use strategies among species; year-round active tree, year-round active shrub, hemiparasite, drought-suppressed broad-leaved shrub and drought-suppressed narrow-leaved shrub.

Leaf water relations during summer water deficit: differential responses in turgor maintenance and variation in leaf structure among different plant communities in south-western Australia.

We measured leaf water relations and leaf structural traits of 20 species from three communities growing along a topographical gradient. Our aim was to assess variation in seasonal responses in leaf water status and leaf tissue physiology between sites and among species in response to summer water deficit. Species from a ridge-top heath community showed the greatest reductions in pre-dawn leaf water potentials (Psi(leaf)) and stomatal conductance during summer; species from a valley-floor woodland and a midslope mallee community showed less reductions in these parameters. Heath species also displayed greater seasonal reduction in turgor-loss point (Psi(TLP)) than species from woodland or mallee communities. In general, species that had larger reductions in Psi(leaf) during summer showed significant shifts in either their osmotic potential at full turgor (Psi(pi 100); osmotic adjustment) or in tissue elasticity (epsilon(max)). Psi(pi 100) and epsilon(max) were negatively correlated, during both spring and summer, suggesting a trade-off between these different mechanisms to cope with water stress. Specific leaf area varied greatly among species, and was significantly correlated with seasonal changes in Psi(TLP) and pre-dawn Psi(leaf). These correlations suggest that leaf structure is a prerequisite for cellular mechanisms to be effective in adjusting to water deficit.

Predicting the limits to tree height using statistical regressions of leaf traits.

Leaf morphology and physiological functioning demonstrate considerable plasticity within tree crowns, with various leaf traits often exhibiting pronounced vertical gradients in very tall trees. It has been proposed that the trajectory of these gradients, as determined by regression methods, could be used in conjunction with theoretical biophysical limits to estimate the maximum height to which trees can grow. Here, we examined this approach using published and new experimental data from tall conifer and angiosperm species. We showed that height predictions were sensitive to tree-to-tree variation in the shape of the regression and to the biophysical endpoints selected. We examined the suitability of proposed end-points and their theoretical validity. We also noted that site and environment influenced height predictions considerably. Use of leaf mass per unit area or leaf water potential coupled with vulnerability of twigs to cavitation poses a number of difficulties for predicting tree height. Photosynthetic rate and carbon isotope discrimination show more promise, but in the second case, the complex relationship between light, water availability, photosynthetic capacity and internal conductance to CO(2) must first be characterized.

Nighttime transpiration in woody plants from contrasting ecosystems.

It is commonly assumed that transpiration does not occur at night because leaf stomata are closed in the dark. We tested this assumption across a diversity of ecosystems and woody plant species by various methods to explore the circumstances when this assumption is false. Our primary goals were: (1) to evaluate the nature and magnitude of nighttime transpiration, E(n), or stomatal conductance, g(n); and (2) to seek potential generalizations about where and when it occurs. Sap-flow, porometry and stable isotope tracer measurements were made on 18 tree and eight shrub species from seven ecosystem types. Coupled with environmental data, our findings revealed that most of these species transpired at night. For some species and circumstances, nighttime leaf water loss constituted a significant fraction of total daily water use. Our evidence shows that E(n) or g(n) can occur in all but one shrub species across the systems we investigated. However, under conditions of high nighttime evaporative demand or low soil water availability, stomata were closed and E(n) or g(n) approached zero in eleven tree and seven shrub species. When soil water was available, E(n) or g(n) was measurable in these same species demonstrating plasticity for E(n) or g(n). We detected E(n) or g(n) in both trees and shrubs, and values were highest in plants from sites with higher soil water contents and in plants from ecosystems that were less prone to atmospheric or soil water deficits. Irrespective of plant or ecosystem type, many species showed E(n) or g(n) when soil water deficits were slight or non-existent, or immediately after rainfall events that followed a period of soil water deficit. The strongest relationship was between E(n) or g(n) and warm, low humidity and (or) windy (> 0.8 m s(-1)) nights when the vapor pressure deficit remained high (> 0.2 kPa in wet sites, > 0.7 kPa in dry sites). Why E(n) or g(n) occurs likely varies with species and ecosystem type; however, our data support four plausible explanations: (1) it may facilitate carbon fixation earlier in the day because stomata are already open; (2) it may enhance nutrient supply to distal parts of the crown when these nutrients are most available (in wet soils) and transport is rapid; (3) it may allow for the delivery of dissolved O(2) via the parenchyma to woody tissue sinks; or (4) it may occur simply because of leaky cuticles in older leaves or when stomata cannot close fully because of obstructions from stomatal (waxy) plugs, leaf endophytes or asymmetrical guard cells (all non-adaptive reasons). We discuss the methodological, ecophysiological, and theoretical implications of the occurrence of E(n) or g(n) for investigations at a variety of scales.

Hydraulic efficiency and safety of branch xylem increases with height in Sequoia sempervirens (D. Don) crowns.

The hydraulic limitation hypothesis of Ryan & Yoder (1997, Bioscience 47, 235-242) suggests that water supply to leaves becomes increasingly difficult with increasing tree height. Within the bounds of this hypothesis, we conjectured that the vertical hydrostatic gradient which gravity generates on the water column in tall trees would cause a progressive increase in xylem 'safety' (increased resistance to embolism and implosion) and a concomitant decrease in xylem 'efficiency' (decreased hydraulic conductivity). We based this idea on the historically recognized concept of a safety-efficiency trade-off in xylem function, and tested it by measuring xylem conductivity and vulnerability to embolism of Sequoia sempervirens branches collected at a range of heights. Measurements of resistance of branch xylem to embolism did indeed show an increase in 'safety' with height. However, the expected decrease in xylem 'efficiency' was not observed. Instead, sapwood-specific hydraulic conductivities (Ks) of branches increased slightly, while leaf-specific hydraulic conductivities increased dramatically, with height. The latter could be largely explained by strong vertical gradients in specific leaf area. The increase in Ks with height corresponded to a decrease in xylem wall fraction (a measure of wall thickness), an increase in percentage of earlywood and slight increases in conduit diameter. These changes are probably adaptive responses to the increased transport requirements of leaves growing in the upper canopy where evaporative demand is greater. The lack of a safety-efficiency tradeoff may be explained by opposing height trends in the pit aperture and conduit diameter of tracheids and the major and semi-independent roles these play in determining xylem safety and efficiency, respectively.

Hydraulic redistribution in three Amazonian trees.

About half of the Amazon rainforest is subject to seasonal droughts of 3 months or more. Despite this drought, several studies have shown that these forests, under a strongly seasonal climate, do not exhibit significant water stress during the dry season. In addition to deep soil water uptake, another contributing explanation for the absence of plant water stress during drought is the process of hydraulic redistribution; the nocturnal transfer of water by roots from moist to dry regions of the soil profile. Here, we present data on patterns of soil moisture and sap flow in roots of three dimorphic-rooted species in the Tapajós Forest, Amazônia, which demonstrate both upward (hydraulic lift) and downward hydraulic redistribution. We measured sap flow in lateral and tap roots of our three study species over a 2-year period using the heat ratio method, a sap-flow technique that allows bi-directional measurement of water flow. On certain nights during the dry season, reverse or acropetal flow (i.e.,in the direction of the soil) in the lateral roots and positive or basipetal sap flow (toward the plant) in the tap roots of Coussarea racemosa (caferana), Manilkara huberi (maçaranduba) and Protium robustum (breu) were observed, a pattern consistent with upward hydraulic redistribution (hydraulic lift). With the onset of heavy rains, this pattern reversed, with continuous night-time acropetal sap flow in the tap root and basipetal sap flow in lateral roots, indicating water movement from wet top soil to dry deeper soils (downward hydraulic redistribution). Both patterns were present in trees within a rainfall exclusion plot (Seca Floresta) and to a more limited extent in the control plot. Although hydraulic redistribution has traditionally been associated with arid or strongly seasonal environments, our findings now suggest that it is important in ameliorating water stress and improving rain infiltration in Amazonian rainforests. This has broad implications for understanding and modeling ecosystem process and forest function in this important biome.

A validation, comparison and error analysis of two heat-pulse methods for measuring sap flow in Eucalyptus marginata saplings.

We validated and compared two heat-pulse methods for measuring sap flow in potted Eucalyptus marginata Donn ex. Smith (jarrah) saplings. During daylight hours and under well-watered conditions, rates of sap flow (0.1-0.5 kg h-1) measured by the established compensation heat-pulse method (CHPM) and the newly developed heat-ratio method (HRM) were similar to rates measured with a weighing lysimeter, and most of the time there was no significant difference (P<0.001) between methods. The HRM accurately described sap flow at night when rates of flow were low (< 0.1 kg h-1) or near zero, but the CHPM was unable to measure low rates of sap flow due to its inability to distinguish heat-pulse velocities below a threshold velocity of 0.1 kg h-1 (3-4 cm h-1). The greatest potential for error in the calculation of daily sap flow was associated with the misalignment of temperature sensors, the estimation of sapwood area and the method used to acquire total sap flow from point measurements of sap velocity. A direct comparison of the two heat-pulse methods (applied synchronously) revealed that the HRM had a more convincing mechanism for correcting spacing errors and was more resistant to random fluctuation in measurements than the CHPM. While we view the HRM more favourably than the CHPM in some key areas, both methods are valid and useful, within their constraints, for measuring transpiration in jarrah and other woody species.

Tree roots: conduits for deep recharge of soil water.

In previous work, we provided evidence from sap flow measurements that when root systems span soil layers of different moisture content, water is redistributed by roots in the direction of the difference in water potential. In addition to the phenomenon termed "hydraulic lift", where water is redistributed from depth to dry topsoil, the process of "hydraulic redistribution" includes downward transfer of water when the surface layers of soils with low permeability become wet after rainfall. In this paper, we support our previous findings with evidence from measurements of soil water and estimate the quantities of water transferred to depth following rain. Amounts of water stored at depth are not likely to be significant for drought avoidance by plants. However, downward transfer of water may be important to plant establishment and the reduction of waterlogging in certain soil types.

The redistribution of soil water by tree root systems.

Plant roots transfer water between soil layers of different water potential thereby significantly affecting the distribution and availability of water in the soil profile. We used a modification of the heat pulse method to measure sap flow in roots of Grevillea robusta and Eucalyptus camaldulensis and demonstrated a redistribution of soil water from deeper in the profile to dry surface horizons by the root system. This phenomenon, termed "hydraulic lift" has been reported previously. However, we also demonstrated that after the surface soils were rewetted at the break of season, water was transported by roots from the surface to deeper soil horizons - the reverse of the "hydraulic lift" behaviour described for other woody species. We suggest that "hydraulic redistribution" of water in tree roots is significant in maintaining root viability, facilitating root growth in dry soils and modifying resource availability.