Isotopic composition of transpiration and rates of change in leaf water isotopologue storage in response to environmental variables.

During daylight hours, the isotope composition of leaf water generally approximates steady-state leaf water isotope enrichment model predictions. However, until very recently there was little direct confirmation that isotopic steady-state (ISS) transpiration in fact exists. Using isotope ratio infrared spectroscopy (IRIS) and leaf gas exchange systems we evaluated the isotope composition of transpiration and the rate of change in leaf water isotopologue storage (isostorage) when leaves were exposed to variable environments. In doing so, we developed a method for controlling the absolute humidity entering the gas exchange cuvette for a wide range of concentrations without changing the isotope composition of water vapour. The measurement system allowed estimation of (18)O enrichment both at the evaporation site and for bulk leaf water, in the steady state and the non-steady state. We show that non-steady-state effects dominate the transpiration isoflux even when leaves are at physiological steady state. Our results suggest that a variable environment likely prevents ISS transpiration from being achieved and that this effect may be exacerbated by lengthy leaf water turnover times due to high leaf water contents.

Spatial variation in photosynthetic CO(2) carbon and oxygen isotope discrimination along leaves of the monocot triticale (Triticum × Secale) relates to mesophyll conductance and the Péclet effect.

Carbon and oxygen isotope discrimination of CO(2) during photosynthesis (Δ(13)C(obs) and Δ(18)O(obs)) were measured along a monocot leaf, triticale (Triticum × Secale). Both Δ(13)C(obs) and Δ(18)O(obs) increased towards the leaf tip. While this was expected for Δ(18)O(obs) , because of progressive enrichment of leaf water associated with the Péclet effect, the result was surprising for Δ(13) C(obs). To explore parameters determining this pattern, we measured activities of key photosynthetic enzymes [ribulose bis-phosphate carboxylase-oxygenase (Rubisco), phosphoenolpyruvate carboxylase (PEPC) and carbonic anhydrase) as well as maximum carboxylation and electron transport rates (V(cmax) and J(max)) along the leaf. Patterns in leaf internal anatomy along the leaf were also quantified. Mesophyll conductance (g(m)) is known to have a strong influence on Δ(13)C(obs) , so we used three commonly used estimation methods to quantify variation in g(m) along the leaf. Variation in Δ(13)C(obs) was correlated with g(m) and chloroplast surface area facing the intercellular air space, but unrelated to photosynthetic enzyme activity. The observed variation could cause errors at higher scales if the appropriate portion of a leaf is not chosen for leaf-level measurements and model parameterization. Our study shows that one-third of the way from the base of the leaf represents the most appropriate portion to enclose in the leaf chamber.

Cellulose (delta)18O is an index of leaf-to-air vapor pressure difference (VPD) in tropical plants.

Cellulose in plants contains oxygen that derives in most cases from precipitation. Because the stable oxygen isotope composition, δ(18)O, of precipitation is associated with environmental conditions, cellulose δ(18)O should be as well. However, plant physiological models using δ(18)O suggest that cellulose δ(18)O is influenced by a complex mix of both climatic and physiological drivers. This influence complicates the interpretation of cellulose δ(18)O values in a paleo-context. Here, we combined empirical data analyses with mechanistic model simulations to i) quantify the impacts that the primary climatic drivers humidity (e(a)) and air temperature (T(air)) have on cellulose δ(18)O values in different tropical ecosystems and ii) determine which environmental signal is dominating cellulose δ(18)O values. Our results revealed that e(a) and T(air) equally influence cellulose δ(18)O values and that distinguishing which of these factors dominates the δ(18)O values of cellulose cannot be accomplished in the absence of additional environmental information. However, the individual impacts of e(a) and T(air) on the δ(18)O values of cellulose can be integrated into a single index of plant-experienced atmospheric vapor demand: the leaf-to-air vapor pressure difference (VPD). We found a robust relationship between VPD and cellulose δ(18)O values in both empirical and modeled data in all ecosystems that we investigated. Our analysis revealed therefore that δ(18)O values in plant cellulose can be used as a proxy for VPD in tropical ecosystems. As VPD is an essential variable that determines the biogeochemical dynamics of ecosystems, our study has applications in ecological-, climate-, or forensic-sciences.

Physiological consequences of height-related morphological variation in Sequoia sempervirens foliage.

This study examined relationships between foliar morphology and gas exchange characteristics as they vary with height within and among crowns of Sequoia sempervirens D. Don trees ranging from 29 to 113 m in height. Shoot mass:area (SMA) ratio increased with height and was less responsive to changes in light availability as height increased, suggesting a transition from light to water relations as the primary determinant of morphology with increasing height. Mass-based rates of maximum photosynthesis (A(max,m)), standardized photosynthesis (A(std,m)) and internal CO(2) conductance (g(i,m)) decreased with height and SMA, while the light compensation point, light saturation point, and mass and area-based rates of dark respiration (R(m)) increased with height and SMA. Among foliage from different heights, much of the variation in standardized photosynthesis was explained by variation in g(i,) consistent with increasing limitation of photosynthesis by internal conductance in foliage with higher SMA. The syndrome of lower internal and stomatal conductance to CO(2) and higher respiration may contribute to reductions in upper crown growth efficiency with increasing height in S. sempervirens trees.

Effects of environmental parameters, leaf physiological properties and leaf water relations on leaf water delta18O enrichment in different Eucalyptus species.

Stable oxygen isotope ratios (delta18O) have become a valuable tool in the plant and ecosystem sciences. The interpretation of delta18O values in plant material is, however, still complicated owing to the complex interactions among factors that influence leaf water enrichment. This study investigated the interplay among environmental parameters, leaf physiological properties and leaf water relations as drivers of the isotopic enrichment of leaf water across 17 Eucalyptus species growing in a common garden. We observed large differences in maximum daily leaf water delta18O across the 17 species. By fitting different leaf water models to these empirical data, we determined that differences in leaf water delta18O across species are largely explained by variation in the Péclet effect across species. Our analyses also revealed that species-specific differences in transpiration do not explain the observed differences in delta18O while the unconstrained fitting parameter 'effective path length' (L) was highly correlated with delta18O. None of the leaf morphological or leaf water related parameters we quantified in this study correlated with the L values we determined even though L was typically interpreted as a leaf morphological/anatomical property. A sensitivity analysis supported the importance of L for explaining the variability in leaf water delta18O across different species. Our investigation highlighted the importance of future studies to quantify the leaf properties that influence L. Obtaining such information will significantly improve our understanding of what ultimately determines the delta18O values of leaf water across different plant species.

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.

Seasonality of photosynthetic parameters in a multi-specific and vertically complex forest ecosystem in the Sierra Nevada of California.

Understanding seasonal variations of photosynthetic parameters is critical for accurate modeling of carbon dioxide (CO2) uptake by ecosystems. Maximum carboxylation velocity (Vcmax), maximum rate of electron transport (Jmax), leaf respiration in the light (R(day)), light-saturated assimilation (Amax) and maximum quantum yield (Phi) were calculated from leaf gas exchange measurements made monthly throughout the year on leaves of three co-occuring evergreen species in a Pinus ponderosa Dougl. ex P. Laws. & C. Laws. forest with shrubs in the understory (Arctostaphylos manzanita Parry and Ceanothus cordulatus Kellogg.). The seasonality and relationships of the photosynthetic parameters with environmental and physiological variables differed among the species. The nitrogen-fixing species, C. cordulatus had the highest values of the parameters and the largest seasonal variation, whereas A. manzanita exhibited the lowest seasonality and weaker correlations with environmental variables. In general, variations in Vcmax were highly correlated with light, leaf mass per area and leaf nitrogen content on an area basis. Temporal scaling of the parameters with each other seemed possible for C. cordulatus and P. ponderosa. However, lags between these variables and Vcmax likely reflect the influences of other factors. The acclimation relationships found along vertical light gradients within canopies in other studies cannot be applied to seasonal variations. The Jmax to Vcmax ratio varied seasonally for P. ponderosa and A. manzanita, being lower at high light, high air temperature and low soil water content.