Water vapour isotopic exchange by epiphytic bromeliads in tropical dry forests reflects niche differentiation and climatic signals

The 18O signals in leaf water (delta18O(lw)) and organic material were dominated by atmospheric water vapour 18O signals (delta18O(vap)) in tank and atmospheric life forms of epiphytic bromeliads with crassulacean acid metabolism (CAM), from a seasonally dry forest in Mexico. Under field conditions, the mean delta18O(lw) for all species was constant during the course of the day and systematically increased from wet to dry seasons (from 0 to +6 per thousand), when relative water content (RWC) diminished from 70 to 30%. In the greenhouse, progressive enrichment from base to leaf tip was observed at low night-time humidity; under high humidity, the leaf tip equilibrated faster with delta18O(vap) than the other leaf sections. Laboratory manipulations using an isotopically depleted water source showed that delta18O(vap) was more rapidly incorporated than liquid water. Our data were consistent with a Craig-Gordon (C-G) model as modified by Helliker and Griffiths predicting that the influx and exchange of delta18O(vap) control delta18O(lw) in certain epiphytic life forms, despite progressive tissue water loss. We use delta18O(lw) signals to define water-use strategies for the coexisting species which are consistent with habitat preference under natural conditions and life form. Bulk organic matter (delta18O(org)) is used to predict the deltaO18(vap) signal at the time of leaf expansion.     

Components of ecosystem evaporation in a temperate coniferous rainforest, with canopy transpiration scaled using sapwood density

Here we develop and test a method to scale sap velocity measurements from individual trees to canopy transpiration (E(c)) in a low-productivity, old-growth rainforest dominated by the conifer Dacrydium cupressinum. Further, E(c) as a component of the ecosystem water balance is quantified in relation to forest floor evaporation rates and measurements of ecosystem evaporation using eddy covariance (E(eco)) in conditions when the canopy was dry and partly wet. Thermal dissipation probes were used to measure sap velocity of individual trees, and scaled to transpiration at the canopy level by dividing trees into classes based on sapwood density and canopy position (sheltered or exposed). When compared with ecosystem eddy covariance measurements, E(c) accounted for 51% of E(eco) on dry days, and 22% of E(eco) on wet days. Low transpiration rates, and significant contributions to E(eco) from wet canopy evaporation and understorey transpiration (35%) and forest floor evaporation (25%), were attributable to the unique characteristics of the forest: in particular, high rainfall, low leaf area index, low stomatal conductance and low productivity associated with severe nutrient limitation.     

deltaO of water vapour, evapotranspiration and the sites of leaf water evaporation in a soybean canopy

Stable isotopes in water have the potential to diagnose changes in the earth's hydrological budget in response to climate change and land use change. However, there have been few measurements in the vapour phase. Here, we present high-frequency measurements of oxygen isotopic compositions of water vapour ( δ _v) and evapotranspiration ( δ _ET) above a soybean canopy using the tunable diode laser (TDL) technique for the entire 2006 growing season in Minnesota, USA. We observed a large variability in surface δ _v from the daily to the seasonal timescales, largely explained by Rayleigh processes, but also influenced by vertical atmospheric mixing, local evapotranspiration (ET) and dew formation. We used δ _ET measurements to calculate the isotopic composition at the sites of evaporative enrichment in leaves ( δ _L,e) and compared that with the commonly used steady-state prediction ( δ _L,s). There was generally a good agreement averaged over the season, but larger differences on individual days. We also found that vertical variability in relative humidity and temperature associated with canopy structure must be addressed in canopy-scale leaf water models. Finally, we explored this data set for direct evidence of the Péclet effect.     

西双版纳热带季节雨林水热通量

利用西双版纳热带季节雨林2003和2004年常规气象、生物量以及水热通量观测资料,对该林地两年内各能量分量的数值大小和变化规律、能量分配以及水量平衡特征等进行了分析研究。结果表明,2003和2004年净辐射总量分别为3516.4MJ/(m.2a)和3516.6MJ/(m.2a)。在能量分配过程中潜热通量占优势,2003年和2004年的总量分别是相应年份净辐射总量的46%和44%,显热通量则分别只有12%和11%。2003年和2004年林冠传导率均值分别为10.3mm/s和10.0mm/s,其中干热季期间的林冠传导率明显低于雾凉季和雨季。林冠传导率与叶面积指数和空气饱和水汽压差值之间分别呈极显著的正、负线性相关关系;它基本上不受土壤含水量的影响,只是当长期无雨或雨量很小导致土壤含水量低于0.15m3/m3时,林冠传导率才与土壤含水量间存在极显著的相关关系。西双版纳热带雨林2003和2004年的蒸散量分别是663mm和634mm,受浓雾和林冠传导率的综合影响,该森林生态系统干季蒸散量低于雨季,这是西双版纳热带季节雨林能够在水热极限条件下生存并良好发育的重要原因。     

Reconstructing the δ18O of atmospheric water vapour via the CAM epiphyte Tillandsia usneoides: seasonal controls on δ18O in the field and large‐scale reconstruction of δ18Oa

Using both oxygen isotope ratios of leaf water (δ¹⁸O_L) and cellulose (δ¹⁸O_C) of Tillandsia usneoides in situ, this paper examined how short‐ and long‐term responses to environmental variation and model parameterization affected the reconstruction of the atmospheric water vapour (δ¹⁸O_a). During sample‐intensive field campaigns, predictions of δ¹⁸O_L matched observations well using a non‐steady‐state model, but the model required data‐rich parameterization. Predictions from the more easily parameterized maximum enrichment model (δ¹⁸O_L–M) matched observed δ¹⁸O_L and observed δ¹⁸O_a when leaf water turnover was less than 3.5 d. Using the δ¹⁸O_L–M model and weekly samples of δ¹⁸O_L across two growing seasons in Florida, USA, reconstructed δ¹⁸O_a was ?12.6 ± 0.3‰. This is compared with δ¹⁸O_a of ?12.4 ± 0.2‰ resolved from the growing‐season‐weighted δ¹⁸O_C. Both of these values were similar to δ¹⁸O_a in equilibrium with precipitation, ?12.9‰. δ¹⁸O_a was also reconstructed through a large‐scale transect with δ¹⁸O_L and the growing‐season‐integrated δ¹⁸O_C across the southeastern United States. There was considerable large‐scale variation, but there was regional, weather‐induced coherence in δ¹⁸O_a when using δ¹⁸O_L. The reconstruction of δ¹⁸O_a with δ¹⁸O_C generally supported the assumption of δ¹⁸O_a being in equilibrium with precipitation δ¹⁸O (δ¹⁸O_ppt), but the pool of δ¹⁸O_ppt with which δ¹⁸O_a was in equilibrium – growing season versus annual δ¹⁸O_ppt – changed with latitude.     

On the difference in the net ecosystem exchange of CO2 between deciduous and evergreen forests in the southeastern United States

The southeastern United States is experiencing a rapid regional increase in the ratio of pine to deciduous forest ecosystems at the same time it is experiencing changes in climate. This study is focused on exploring how these shifts will affect the carbon sink capacity of southeastern US forests, which we show here are among the strongest carbon sinks in the continental United States. Using eight‐year‐long eddy covariance records collected above a hardwood deciduous forest (HW) and a pine plantation (PP) co‐located in North Carolina, USA, we show that the net ecosystem exchange of CO₂ (NEE) was more variable in PP, contributing to variability in the difference in NEE between the two sites (ΔNEE) at a range of timescales, including the interannual timescale. Because the variability in evapotranspiration (ET) was nearly identical across the two sites over a range of timescales, the factors that determined the variability in ΔNEE were dominated by those that tend to decouple NEE from ET. One such factor was water use efficiency, which changed dramatically in response to drought and also tended to increase monotonically in nondrought years (P < 0.001 in PP). Factors that vary over seasonal timescales were strong determinants of the NEE in the HW site; however, seasonality was less important in the PP site, where significant amounts of carbon were assimilated outside of the active season, representing an important advantage of evergreen trees in warm, temperate climates. Additional variability in the fluxes at long‐time scales may be attributable to slowly evolving factors, including canopy structure and increases in dormant season air temperature. Taken together, study results suggest that the carbon sink in the southeastern United States may become more variable in the future, owing to a predicted increase in drought frequency and an increase in the fractional cover of southern pines.     

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.     

Seasonal controls on interannual variability in carbon dioxide exchange of a near-end-of rotation Douglas-fir stand in the Pacific Northwest, 1997–2006

This study analyzes 9 years of eddy‐covariance (EC) data carried out in a Pacific Northwest Douglas‐fir (Pseudotsuga menzesii) forest (58‐year old in 2007) on the east coast of Vancouver Island, Canada, and characterizes the seasonal and interannual variability in net ecosystem productivity (NEP), gross primary productivity (GPP), and ecosystem respiration (R_e) and primary climatic controls on these fluxes. The annual values (± SD) of NEP, GPP and R_e were 357 ± 51, 2124 ± 125, and 1767 ± 146 g C m^?2 yr^?1, respectively, with ranges of 267–410, 1592–2338, and 1642–2071 g C m^?2 yr^?1, respectively. Spring to early summer (March–June) accounted for more than 80% of annual NEP while late spring to early autumn (May–August) was mainly responsible for its interannual variability (～80%). The major drivers of interannual variability in annual carbon (C) fluxes were annual and spring mean air temperatures (T_a) and water deficiency during late summer and autumn (July–October) when this Douglas‐fir forest growth was often water‐limited. Photosynthetically active radiation (Q), and the combination of Q and soil water content (θ) explained 85% and 91% of the variance of monthly GPP, respectively; and 91% and 96% of the variance of monthly R_e was explained by T_a and the combination of T_a and θ, respectively. Annual net C sequestration was high during optimally warm and normal precipitation years, but low in unusually warm or severely dry years. Excluding 1998 and 1999, the 2 years strongly affected by an El Niño/La Niña cycle, annual NEP significantly decreased with increasing annual mean T_a. Annual NEP will likely decrease whereas both annual GPP and R_e will likely increase if the future climate at the site follows a trend similar to that of the past 40 years.     

Effect of changes in water content on photosynthesis,transpiration and discrimination against 13CO2 and C18O16O in Pleurozium and Sphagnum

Photosynthetic gas exchange characteristics of two common boreal forest mosses, Sphagnum (section acutifolia) and Pleurozium schreberi, were measured continuously during the time required for the moss to dry out from full hydration. Similar patterns of change in CO₂ assimilation with variation in water content occurred for both species. The maximum rates of CO₂ assimilation for Sphagnum (approx. 7 mol m^–2 s^–1) occurred at a water content of approximately 7 (fresh weight/dry weight) while for Pleurozium the maximum rate (approx. 2 mol m^–2 s^–1) occurred at a water content of approximately 6 (fresh weight/dry weight). Above and below these water contents CO₂ assimilation declined. In both species total conductance to water vapour (expressed as a percentage of the maximum rates) remained nearly constant at a water content above 9 (fresh weight/dry weight), but below this level declined in a strong linear manner. Short-term, on-line ¹³CO₂ and C¹⁸O¹⁶O discrimination varied substantially with changes in moss water content and associated changes in the ratio of chloroplast CO₂ to ambient CO₂ partial pressure. At full hydration (maximum water content) both Sphagnum and Pleurozium had similar values of ¹³CO₂ discrimination (approx. 15). Discrimination against ¹³CO₂ increased continuously with reductions in water content to a maximum of 27 in Sphagnum and 22 in Pleurozium. In a similar manner C¹⁸C¹⁶O discrimination increased from approximately 30 at full hydration in both species to a maximum of 150 in Sphagnum and 90 in Pleurozium, at low water content. The observed changes in C¹⁸O¹⁶O were strongly correlated to predictions of a mechanistic model of discrimination processes. Field measurements of moss water content suggested that photosynthetic gas exchange by moss in the understory of a black spruce forest was regularly limited by low water content.     

The role of effective leaf mixing length in the relationship between the δ18O of stem cellulose and source water across a salinity gradient

Previous mangrove tree ring studies attempted, unsuccessfully, to relate the δ¹⁸O of trunk cellulose (δ¹⁸O_CELL) to the δ¹⁸O of source water (δ¹⁸O_SW). Here, we tested whether biochemical fractionation associated with one of the oxygen in the cellulose glucose moiety or variation in leaf water oxygen isotope fractionation (Δ_LW) can interfere with the δ¹⁸O_SW signal as it is recorded in the δ¹⁸O_CELL of mangrove (saltwater) and hammock (freshwater) plants. We selected two transects experiencing a salinity gradient, located in the Florida Keys, USA. The δ¹⁸O_CELL throughout both transects did not show the pattern expected based on that of the δ¹⁸O_SW. We found that in one of the transects, biochemical fractionation interfered with the δ¹⁸O_SW signal, while in the other transect Δ_LW differed between mangrove and hammock plants. Observed differences in Δ_LW between mangroves and hammocks were caused by a longer effective leaf mixing length (L) of the water pathway in mangrove leaves compared to those of hammock leaves. Changes in L could have caused the δ¹⁸O_CELL to record not only variations in the δ¹⁸O_SW but also in Δ_LW making it impossible to isolate the δ¹⁸O_SW signal.     

Isotopic heterogeneity of water in transpiring leaves: identification of the component that controls the δ18O of atmospheric O2 and CO2

Two direct but independent approaches were developed to identify the average δ¹⁸O value of the water fraction in the chloroplasts of transpiring leaves. In the first approach, we used the δ¹⁸O value of CO₂ in isotopic equilibrium with leaf water to reconstruct the δ¹⁸O value of water in the chloroplasts. This method was based on the idea that the enzyme carbonic anhydrase facilitates isotopic equilibrium between CO₂ and H₂O predominantly in the chloroplasts, at a rate that is several orders of magnitude faster than the non-catalysed exchange in other leaf water fractions. In the second approach, we measured the δ¹⁸O value of O₂ from photosynthetic water oxidation in the chloroplasts of intact leaves. Since O₂ is produced from chloroplast water irreversibly and without discrimination, the δ¹⁸O value of the O₂ should be identical to that of chloroplast water. In intact, transpiring leaves of sunflower (Helianthus annuus cv. giant mammoth) under the experimental conditions used, the average δ¹⁸O value of chloroplasts water was displaced by 3—10 % (depending on relative humidity and atmospheric composition) below the value predicted by the conventional Craig & Gordon model. Furthermore, this δ¹⁸O value was always lower than the δ¹⁸O value that was measured for bulk leaf water. Our results have implications for a variety of environmental studies since it is the δ¹⁸O value of water in the chloroplasts that is the relevant quantity in considering terrestrial plants influence on the δ¹⁸O values of atmospheric CO₂ and O₂, as well as in influencing the δ¹⁸O of plant organic matter.