Elevated CO2 does not affect stem CO2 efflux nor stem respiration in a dry Eucalyptus woodland,but it shifts the vertical gradient in xylem [CO2]

To quantify stem respiration (R_S) under elevated CO₂ (eCO₂), stem CO₂ efflux (E_A) and CO₂ flux through the xylem (F_T) should be accounted for, because part of respired CO₂ is transported upwards with the sap solution. However, previous studies have used E_A as a proxy of R_S, which could lead to equivocal conclusions. Here, to test the effect of eCO₂ on R_S, both E_A and F_T were measured in a free‐air CO₂ enrichment experiment located in a mature Eucalyptus native forest. Drought stress substantially reduced E_A and R_S, which were unaffected by eCO₂, likely as a consequence of its neutral effect on stem growth in this phosphorus‐limited site. However, xylem CO₂ concentration measured near the stem base was higher under eCO₂, and decreased along the stem resulting in a negative contribution of F_T to R_S, whereas the contribution of F_T to R_S under ambient CO₂ was positive. Negative F_T indicates net efflux of CO₂ respired below the monitored stem segment, likely coming from the roots. Our results highlight the role of nutrient availability on the dependency of R_S on eCO₂ and suggest stimulated root respiration under eCO₂ that may shift vertical gradients in xylem [CO₂] confounding the interpretation of E_A measurements.     

Effects of xylem sap flow on carbon dioxide efflux from stems of birch (Betula pendula Roth)

The effect of xylem sap flow in stems of mature Betula pendula Roth on radial CO₂ efflux was studied from April to October 2001. Temperature-controlled respiration cuvettes allowed measurements of CO₂ efflux without interference from temperature gradients between stem surface and sapwood. Variations of sap flow in different stem sectors, and in a given sector at different heights were analysed. Daytime reduction of CO₂ efflux caused by sap flow was expressed as the difference between gross and apparent CO₂ release. Gross CO₂ release was calculated from Arrhenius-equations derived from night-time data records of the same day, which were free from interference by sap flow. In mid-July, daytime reductions of CO₂ efflux reached 1.8–3.9 μmol CO₂ m⁻² g⁻¹ xylem sap transpired. Assuming tree-specific maximum transpiration rates of 30 kg H₂O d⁻¹ this is up to 40% of gross CO₂ release. In relation to photosynthetic CO₂ fixation the endogenous supply of dissolved CO₂ to the leaves acccounted for 0.5–3.7%. This study indicates a negative correlation between sap flow velocity and radial CO₂ efflux from B. pendula stems. Periods of unbalanced CO₂ partial pressures between aqueous and gaseous pathways during increase and decrease of sap flow seem to affect gaseous CO₂ release through lenticels. It is concluded that CO₂ efflux rates are not simply equivalent to respiration rates because of the interference of aqueous CO₂ transport by xylem sap flow in the wood-body of trees.     

Stem CO2 efflux of ten species in temperate forests in Northeastern China

Stem CO₂ efflux (E _S) is an important component of forest ecosystem carbon budgets and net ecosystem CO₂ exchange, but little is known about E _S in temperate forests in Northeastern China, an area with a large extent of forest. We measured E _S along with stem temperature at 1?cm depth (Ts) over a 9?month period in 2007 on ten dominant tree species of secondary forests of the region. Other measurements included the autotrophic component of soil CO₂ efflux (E _A) and stem diameter at breast height (DBH). Our objectives were to (1) examine the seasonal patterns and species differences in E _S, and (2) determine the correlations between E _S and Ts, DBH and E _A. Mean E _S for the measurement period ranged from 1.09 to 1.74?μmol?CO₂?m^?2?s^?1 among the ten species. The sensitivity of E _S to Ts (Q ₁₀ ) ranged from 1.87 to 2.61. Across the ten species 57–89% of variation in E _S was explained by T _S and DBH. There was also a linear relationship between mean E _S and E _A. E _S was better predicted by Ts in the dormant season than the growing season, indicating that additional factors such as growth respiration and internal transport of CO₂ in the xylem became more important contributors to E _S during the growing season. Stem CO₂ efflux increased, and Q ₁₀ decreased, with increasing DBH in all species. Although temperature exerts strong control on the rate of cellular respiration, we conclude that in tree stems in situ, T _S, DBH and many other factors affect the relationship between CO₂ evolution by respiring cells and the diffusion of CO₂ to the stem surface.     

Respiratory carbon use and carbon storage in mid‐rotation loblolly pine (Pinus taeda L.) plantations: the effect of site resources on the stand carbon balance

We used estimates of autotrophic respiration (R_A), net primary productivity (NPP) and soil CO₂ evolution (S_ff), to develop component carbon budgets for 12‐year‐old loblolly pine plantations during the fifth year of a fertilization and irrigation experiment. Annual carbon use in R_A was 7.5, 9.0, 15.0, and 15.1 Mg C ha^?1 in control (C), irrigated (I), fertilized (F) and irrigated and fertilized (IF) treatments, respectively. Foliage, fine root and perennial woody tissue (stem, branch, coarse and taproot) respiration accounted for, respectively, 37%, 24%, and 39% of R_A in C and I treatments and 38%, 12% and 50% of R_A in F and IF treatments. Annual gross primary production (GPP=NPP+R_A) ranged from 13.1 to 26.6 Mg C ha^?1. The I, F, and IF treatments resulted in a 21, 94, and 103% increase in GPP, respectively, compared to the C treatment. Despite large treatment differences in NPP, R_A, and carbon allocation, carbon use efficiency (CUE=NPP/GPP) averaged 0.42 and was unaffected by manipulating site resources. Ecosystem respiration (R_E), the sum of S_ff, and above ground R_A, ranged from 12.8 to 20.2 Mg C ha^?1 yr^?1. S_ff contributed the largest proportion of R_E, but the relative importance of S_ff decreased from 0.63 in C treatments to 0.47 in IF treatments because of increased aboveground R_A. Aboveground woody tissue R_A was 15% of R_E in C and I treatments compared to 25% of R_E in F and IF treatments. Net ecosystem productivity (NEP=GPP‐R_E) was roughly 0 in the C and I treatments and 6.4 Mg C ha^?1 yr^?1 in F and IF treatments, indicating that non‐fertilized treatments were neither a source nor a sink for atmospheric carbon while fertilized treatments were carbon sinks. In these young stands, NEP is tightly linked to NPP; increased ecosystem carbon storage results mainly from an increase in foliage and perennial woody biomass.     

Effects of sap velocity on the daytime increase of stem CO2 efflux from stems of Schima superba trees

Stem CO₂ efflux (E _s) has been estimated from a temperature-related equation, but sap flux often affects measurements of E _s, which leads to misunderstanding real stem respiration. In order to observe the relationship between E _s and stem temperature and to analyze the effect of sap velocity on E _s, stem temperature, E _s and sap flux were measured from a subtropical Schima superba plantation in South China on three trees for consecutive 3 days in July and October 2009. Stem temperature, E _s and sap velocity were significantly higher in July than in October. Stem temperature could explain 17–41 and 54–75% variations of E _s in July and October, respectively. A negative relationship between E _s and stem temperature was found during 1800–2300 hours in July. The daytime E _s was 9.2, 4.3 and 2.4% higher than the predicted for three trees in July, and this occurred only on Tree 1 in October. Sap velocity was positively correlated with E _s for three trees in July, and the increase of E _s with the increase of sap velocity was only observed on Tree 1 in October. These results demonstrated that the occurrence of sap flux could account for the increase of daytime E _s, and the effect of sap velocity on E _s varied with the seasons from the S. superba stem.     

Carbon dioxide transport inxylem causes errors in estimation of rates of respiration in stemsand branches of trees

Complementary laboratory and field experiments showed that theinternal transport of carbon dioxide (CO₂) in the xylemof trees is an important pathway for carbon movement. Carbon dioxidereleased by respiration dissolves in sap and moves upward in thetranspirational stream. The concentration of CO₂ in xylemsap can be up to three orders of magnitude greater than that foundin the atmosphere. In the present experiments, diffusion outwardof a portion of xylem‐transported CO₂ caused a substantialoverestimation of the apparent rate of stem and branch respiration.Rates of CO₂ efflux were linearly related to sap CO₂ concentration.Direct manipulations of xylem sap CO₂ concentration producedrapid and reversible changes in CO₂ efflux from stemsand branches, in some cases quadrupling the rate of efflux. Theseresults demonstrated that apparent rates of stem and branch respirationof trees are in large part a by‐product of the rate of CO₂ diffusionfrom xylem sap.     

Diverging temperature response of tree stem CO2 release under dry and wet season conditions in a tropical montane moist forest

It is commonly presumed that plant respiratory CO₂ release increases with increasing temperature. However, we report on very contrasting stem CO₂ release (R _S)–temperature relationships of trees in a species-rich tropical montane forest of southern Ecuador under dry and wet season conditions. Rates of R _S were low and completely uncoupled from the dial temperature regime during the humid season. In contrast, during the dry season, R _S was generally higher and temperature sensitivity of R _S differed greatly in degree and even in the direction of response, indicating that temperature might not be the only determinant of R _S. In order to explain the heterogeneity of R _S, we related R _S to vapour pressure deficit, wind speed and solar radiation as important abiotic drivers influencing transpiration and photosynthesis. Stepwise multiple regression analyses with these meteorological predictors either were biased by high collinearity of the independent variables or could not enhance the ability to explain the variability of R _S. We assume maintenance respiration to dominate under humid conditions unfavourable for energy acquisition of the tree, thus explaining the pronounced uncoupling of R _S from atmospheric parameters. In contrast, the drier and hotter climate of the dry season seems to favour R _S via enhanced assimilatory substrate delivery and stem respiratory activity as well as elevated xylem sap CO₂ imports with increased transpiration. In addition, tree individual differences in the temperature responses of R _S may mirror diverging climatic adaptations of co-existing moist forest tree species which have their distribution centre either at higher or lower elevations.     

The concentration and efflux of tree stem CO2 and the role of xylem sap flow

The accurate assessment of actual tree stem respiration and its relation with temperature plays a considerable role in investigating the forest carbon cycle. An increasing number of research reports have indicated that tree stem respiration determined with the commonlyapplied chamber gas exchange measuring system does not follow expectations regarding temperature relationships. This method is based on the nowadays widely-accepted theory that the respired CO₂ in a tree stem would all diffuse outward into the atmosphere. However, it neglects partial CO₂ that is dissolved in the xylem sap and is carried away by the transpirational stream. Scientists have started to realize that the respired CO₂ measured with the chamber gas exchange method is only a portion of the total stem respiration (CO₂ efflux), while the other portion, which is sometimes very substantial in quantity (thought to occupy maybe 15%–75% of the total stem respiration), is transported to the upper part of the stem and to the canopy by sap flow. This suggests that the CO₂ produced by respiration is re-allocated within the stem. Accordingly, the change in CO₂ efflux could be reflected in the rates of sap flow in addition to its dependence on temperature. Proper methods and instruments are required to quantify the internal and external CO₂ fluxes in the trunk and their interaction with related environmental factors.     

Accelerated soil CO<Subscript>2</Subscript> efflux after conversion from secondary oak forest to pine plantation in southeastern China

Soil respiration (R _s) is an important component of the carbon cycle in terrestrial ecosystems, and changes in soil respiration with land cover alteration can have important implications for regional carbon balances. In southeastern China (Xiashu Experimental Forest, Jiangsu Province), we used an automated LI-8100 soil CO₂ flux system to quantify diurnal variation of soil respiration in a secondary oak forest and a pine plantation. We found that soil respiration in the pine plantation was significantly higher than that in the secondary oak forest. There were similar patterns of soil respiration throughout the day in both the secondary oak forest and the pine plantation during our 7-month study (March–September 2005). The maximum of R _s occurred between 4:00 pm and 7:00 pm. The diurnal variations of R _s were usually out of phase with soil surface (0.5 cm) temperature (T _g). However, annual variation in R _s correlated with surface soil temperature. Soil respiration reached to a maximum in June, and decreased thereafter. The Q₁₀ of R _s in the secondary oak forest was significantly higher than that in the pine plantation. The higher Q₁₀ value in the secondary oak forest implied that it might release more CO₂ than the pine plantation under a global-warming scenario. Our results indicated that land-use change from secondary forest to plantation may cause a significant increase in CO₂ emission, and reduce the temperature sensitivity of soil respiration in southeastern China.     

Woody tissue photosynthesis reduces stem CO2 efflux by half and remains unaffected by drought stress in young Populus tremula trees

A substantial portion of locally respired CO₂ in stems can be assimilated by chloroplast-containing tissues. Woody tissue photosynthesis (P_wt) therefore plays a major role in the stem carbon balance. To study the impact of P_wt on stem carbon cycling along a gradient of water availability, stem CO₂ efflux (E_A), xylem CO₂ concentration ([CO₂]), and xylem water potential (Ψ_xylem) were measured in 4-year-old Populus tremula L. trees exposed to drought stress and different regimes of light exclusion of woody tissues. Under well-watered conditions, local P_wt decreased E_A up to 30%. Axial CO₂ diffusion (D_ax) induced by distant P_wt caused an additional decrease in E_A of up to 25% and limited xylem [CO₂] build-up. Under drought stress, absolute decreases in E_A driven by P_wt remained stable, denoting that P_wt was not affected by drought. At the end of the dry period, when transpiration was low, local P_wt and D_ax offset 20% and 10% of stem respiration on a daily basis, respectively. These results highlight (a) the importance of P_wt for an adequate interpretation of E_A measurements and (b) homeostatic P_wt along a drought stress gradient, which might play a crucial role to fuel stem metabolism when leaf carbon uptake and phloem transport are limited.     

Responses of dryland soil respiration and soil carbon pool size to abrupt vs. gradual and individual vs. combined changes in soil temperature, precipitation, and atmospheric [CO2]: a simulation analysis

With the large extent and great amount of soil carbon (C) storage, drylands play an important role in terrestrial C balance and feedbacks to climate change. Yet, how dryland soils respond to gradual and concomitant changes in multiple global change drivers [e.g., temperature (T_s), precipitation (Ppt), and atmospheric [CO₂] (CO₂)] has rarely been studied. We used a process‐based ecosystem model patch arid land simulator to simulate dryland soil respiration (R_s) and C pool size (C_s) changes to abrupt vs. gradual and single vs. combined alterations in T_s, Ppt and CO₂ at multiple treatment levels. Results showed that abrupt perturbations generally resulted in larger R_s and had longer differentiated impacts than did gradual perturbations. R_s was stimulated by increases in T_s, Ppt, and CO₂ in a nonlinear fashion (e.g., parabolically or asymptotically) but suppressed by Ppt reduction. Warming mainly stimulated heterotrophic R_s (i.e., R_h) whereas Ppt and CO₂ influenced autotrophic R_s (i.e., R_a). The combined effects of warming, Ppt, and CO₂ were nonadditive of primary single‐factor effects as a result of substantial interactions among these factors. Warming amplified the effects of both Ppt addition and CO₂ elevation whereas Ppt addition and CO₂ elevation counteracted with each other. Precipitation reduction either magnified or suppressed warming and CO₂ effects, depending on the magnitude of factor's alteration and the components of R_s (R_a or R_h) being examined. Overall, Ppt had dominant influence on dryland R_s and C_s over T_s and CO₂. Increasing Ppt individually or in combination with T_s and CO₂ benefited soil C sequestration. We therefore suggested that global change experimental studies for dryland ecosystems should focus more on the effects of precipitation regime changes and the combined effects of Ppt with other global change factors (e.g., T_s, CO₂, and N deposition).     

Coupling between carbon cycling and climate in a high-elevation,subalpine forest: a model-data fusion analysis

Fundamental questions exist about the effects of climate on terrestrial net ecosystem CO₂ exchange (NEE), despite a rapidly growing body of flux observations. One strategy to clarify ecosystem climate–carbon interactions is to partition NEE into its component fluxes, gross ecosystem CO₂ exchange (GEE) and ecosystem respiration (R _E), and evaluate the responses to climate of each component flux. We separated observed NEE into optimized estimates of GEE and R _E using an ecosystem process model combined with 6 years of continuous flux data from the Niwot Ridge AmeriFlux site. In order to gain further insight into the processes underlying NEE, we partitioned R _E into its components: heterotrophic (R _H) and autotrophic (R _A) respiration. We were successful in separating GEE and R _E, but less successful in accurately partitioning R _E into R _A and R _H. Our failure in the latter was due to a lack of adequate contrasts in the assimilated data set to distinguish between R _A and R _H. We performed most model runs at a twice-daily time step. Optimizing on daily-aggregated data severely degraded the model’s ability to separate GEE and R _E. However, we gained little benefit from using a half-hourly time step. The model-data fusion showed that most of the interannual variability in NEE was due to variability in GEE, and not R _E. In contrast to several previous studies in other ecosystems, we found that longer growing seasons at Niwot Ridge were correlated with less net CO₂ uptake, due to a decrease of available snow-melt water during the late springtime photosynthetic period. Warmer springtime temperatures resulted in increased net CO₂ uptake only if adequate moisture was available; when warmer springtime conditions led into mid-summer drought, the annual net uptake declined.     

Global patterns and predictors of stem CO2 efflux in forest ecosystems

Stem CO₂ efflux (E_S) plays an important role in the carbon balance of forest ecosystems. However, its primary controls at the global scale are poorly understood and observation‐based global estimates are lacking. We synthesized data from 121 published studies across global forest ecosystems and examined the relationships between annual E_S and biotic and abiotic factors at individual, biome, and global scales, and developed a global gridded estimate of annual E_S. We tested the following hypotheses: (1) Leaf area index (LAI) will be highly correlated with annual E_S at biome and global scales; (2) there will be parallel patterns in stem and root CO₂ effluxes (R_A) in all forests; (3) annual E_S will decline with forest age; and (4) LAI coupled with mean annual temperature (MAT) and mean annual precipitation (MAP) will be sufficient to predict annual E_S across forests in different regions. Positive linear relationships were found between E_S and LAI, as well as gross primary production (GPP), net primary production (NPP), wood NPP, soil CO₂ efflux (R_S), and R_A. Annual E_S was correlated with R_A in temperate forests after controlling for GPP and MAT, suggesting other additional factors contributed to the relationship. Annual E_S tended to decrease with stand age. Leaf area index, MAT and MAP, predicted 74% of variation in E_S at global scales. Our statistical model estimated a global annual E_S of 6.7 ± 1.1 Pg C yr⁻¹ over the period of 2000–2012 with little interannual variability. Modeled mean annual E_S was 71 ± 43, 270 ± 103, and 420 ± 134 g C m² yr⁻¹ for boreal, temperate, and tropical forests, respectively. We recommend that future studies report E_S at a standardized constant temperature, incorporate more manipulative treatments, such as fertilization and drought, and whenever possible, simultaneously measure both aboveground and belowground CO₂ fluxes.     

Soil carbon stocks and fluxes in a warm-temperate oak chronosequence in China

Soil respiration (R_S) and soil carbon stocks, as well as stand properties were investigated in a warm-temperate oak chronosequence in order to understand the age effect on soil CO₂ efflux. The chronosequence consisted of three 40-year-old, 48-year-old, 80-year-old, and 143-year-old oak stands, respectively. R_S measurements were conducted using a Li-8100 soil CO₂ flux system from October 2008 to October 2009. Temporal variations of R_S of all the four forests largely depended on soil temperature of 5 cm depth (T₅) (R²?=?0.738?C0.825). The mean R_S for 40-year-old, 48-year-old, 80-year-old, and 143-year-old forests were 2.37, 2.59, 2.99, and 3.32 ??mol CO₂ m^-2 s^-1 respectively. Both top soil organic carbon (SOC) and light fraction organic carbon (LFOC) stocks were significantly correlated to R_S variation, while only significant different LFOC among stands was found. This indicated that cumulated labile organic carbon was a better indicator on R_S variation, which was further illustrated by a better relationship between R ₁₀ and LFOC than that of R₁₀ and SOC. We found that the variation of mean R_S among stands was well correlated with basal area (BA). Marginal correlation between R_S and fine root biomass (FR) demonstrated the relationship between R_S and belowground metabolism. We also found total porosity (TP) negatively influenced the mean R_S and this negative effect may mainly be attributed to the capillary porosity (CP). Forest growth and yield could be contributed to R_S variation among stands. Forest succession also changed soil labile carbon stock and soil physical properties that influenced the CO₂ efflux.     

Soil CO2 efflux and soil carbon balance of a tropical rubber plantation

Natural rubber is a valuable source of income in many tropical countries and rubber trees are increasingly planted in tropical areas, where they contribute to land-use changes that impact the global carbon cycle. However, little is known about the carbon balance of these plantations. We studied the soil carbon balance of a 15-year-old rubber plantation in Thailand and we specifically explored the seasonal dynamic of soil CO₂ efflux (F _S) in relation to seasonal changes in soil water content (W _S) and soil temperature (T _S), assessed the partitioning of F _S between autotrophic (R _A) and heterotrophic (R _H) sources in a root trenching experiment and estimated the contribution of aboveground and belowground carbon inputs to the soil carbon budget. A multiplicative model combining both T _S and W _S explained 58 % of the seasonal variation of F _S. Annual soil CO₂ efflux averaged 1.88 kg C m^?2 year^?1 between May 2009 and April 2011 and R _A and R _H accounted for respectively 63 and 37 % of F _S, after corrections of F _S measured on trenched plots for root decomposition and for difference in soil water content. The 4-year average annual aboveground litterfall was 0.53 kg C m^?2 year^?1 while a conservative estimate of belowground carbon input into the soil was much lower (0.17 kg C m^?2 year^?1). Our results highlighted that belowground processes (root and rhizomicrobial respiration and the heterotrophic respiration related to belowground carbon input into the soil) have a larger contribution to soil CO₂ efflux (72 %) than aboveground litter decomposition.     

Direct effects of atmospheric carbon dioxide concentration on whole canopy dark respiration of rice

The purpose of this study was to test for direct inhibition of rice canopy apparent respiration by elevated atmospheric carbon dioxide concentration ([CO₂]) across a range of short‐term air temperature treatments. Rice (cv. IR‐72) was grown in eight naturally sunlit, semiclosed, plant growth chambers at daytime [CO₂] treatments of 350 and 700 μmol mol^?1. Short‐term night‐time air temperature treatments ranged from 21 to 40 °C. Whole canopy respiration, expressed on a ground area basis (R_d), was measured at night by periodically venting the chambers with ambient air. This night‐time chamber venting and resealing procedure produced a range of increasing chamber [CO₂] which we used to test for potential inhibitory effects of rising [CO₂] on R_d. A nitrous oxide leak detection system was used to correct R_d measurements for chamber leakage rate (L) and also to determine if apparent reductions in night‐time R_d with rising [CO₂] could be completely accounted for by L. The L was affected by both CO₂ concentration gradient between the chamber and ambient air and the inherent leakiness of each individual chamber. Nevertheless, after correcting R_d for L, we detected a rapid and reversible, direct inhibition of R_d with rising chamber [CO₂] for air temperatures above 21 °C. This effect was larger for the 350 compared with the 700 μmol mol^?1 daytime [CO₂] treatment and was also increased with increasing short‐term air temperature treatments. However, little difference in R_d was found between the two daytime [CO₂] treatments when night‐time [CO₂] was at the respective daytime [CO₂]. These results suggest that naturally occurring diurnal changes in both ambient [CO₂] and air temperature can affect R_d. Because naturally occurring diurnal changes in both [CO₂] and air temperature can be expected in a future higher CO₂ world, short‐term direct effects of these environmental variables on rice R_d can also be expected.     

Depressed hydraulic redistribution of roots more by stem refilling than by nocturnal transpiration for Populus euphratica Oliv. in situ measurement

During the night, plant water loss can occur either through the roots, as hydraulic redistribution (HR), or through the leaves via the stoma, as nocturnal transpiration (E_n), which was methodologically difficult to separate from stem refilling (R_e). While HR and E_n have been reported across a range of species, ecosystem, and climate zone, there is little understanding on the interactions between E_n and/or R_e and HR. As water movement at night occurs via gradients of water potential, it is expected that during periods of high atmospheric vapor pressure deficit (VPD), water loss via E_n will override water loss via HR. To test this hypothesis, sap flow in stems and roots of Populus euphratica Oliv. trees, growing in a riparian zone in a hyperarid climate, was measured once in a year. Nocturnal stem sap flow was separated into E_n and R_e using the “forecasted refilling” method. Substantial nocturnal sap flow (38% of 24‐hr flux on average) was observed and positively correlated with VPD; however, the strength of the correlation was lower (R² = .55) than diurnal sap flow (E_d) (R² = .72), suggesting that nocturnal stem sap flow was attributed to both water loss through the canopy and replenishment of water in stem tissues. Partitioning of nocturnal sap flow shows that R_e constituted approximately 80%, and E_n ~20%, of nocturnal sap flow. The amount of root sap flow attributed to redistribution was negatively related to E_d (R² = .69) and the amount of acropetally sap flow in stems, R_e (R² = .41) and E_n (R² = .14). It was suggested that the magnitude of HR is more strongly depressed by R_e that was recharge to the water loss via E_d than by E_n. It was consistent with whole‐tree water balance theory, that the nighttime upward sap flow to xylem, stem refilling and transpiration, may depress hydraulic redistribution of roots.     

Characteristics of leaf photosynthesis and simulated individual carbon budget in <Emphasis Type="Italic">Primula nutans</Emphasis> under contrasting light and temperature conditions

Primula nutans Georgi is widely distributed in hummock-and-hollow wetlands on the Qinghai-Tibetan Plateau. To assess the ecophysiology of this species in responding to microenvironments, we examined the photosynthetic characteristics and individual carbon gain of plants growing in different microsites from a hummock-and-hollow wetland on the Qinghai-Tibetan Plateau and under laboratory conditions. Plants from wetland hummock microsites showed significantly higher light-saturated photosynthetic CO₂ uptake (A _max) than those from microsites in hollows at a controlled temperature of 15°C in leaf chamber. Leaf dark respiration rate (R) was only significantly higher in plants from hummocks than hollows at the measuring temperature of 35°C. Optimum temperature for A _max was 15°C for all plants in the field despite different microsites. In plants growing under laboratory conditions differing in light and temperature, both A _max and R were significantly higher under higher growth light (photosynthetic photon flux density, PPFD: 800 or 400 μmol m⁻² s⁻¹) than low light of 90 μmol m⁻² s⁻¹. No statistically significant differences in A _max and R existed in plants differing in growing temperatures. Estimates derived from the photosynthetic parameters of field plants, and microsite environmental measures including PPFD, air temperature and soil temperature showed that the optimum mean daily temperature for net daily carbon gain was around 10°C and the net daily carbon gain was largely limited under lower daily total PPFD. These results suggest that the differences in A _max and R in P. nutans are strongly affected by growing light regimes but not by temperature regimes. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Stomatal responses of Douglas-fir seedlings to elevated carbon dioxide and temperature during the third and fourth years of exposure

Two major components of climate change, increasing atmospheric [CO₂] and increasing temperature, may substantially alter the effects of water availability to plants through effects on the rate of water loss from leaves. We examined the interactive effects of elevated [CO₂] and temperature on seasonal patterns of stomatal conductance (g_s), transpiration (E) and instantaneous transpiration efficiency (ITE) in Douglas‐fir (Pseudotsuga menziesii (Mirb.) Franco) seedlings. Seedlings were grown in sunlit chambers at either ambient CO₂ (AC) or ambient + 180 µmol mol^?1 CO₂ (EC), and at ambient temperature (AT) or ambient + 3·5 °C (ET) in a full‐factorial design. Needle gas exchange at the target growth conditions was measured approximately monthly over 21 months. Across the study period and across temperature treatments, growth in elevated [CO₂] decreased E by an average of 12% and increased ITE by an average of 46%. The absolute reduction of E associated with elevated [CO₂] significantly increased with seasonal increases in the needle‐to‐air vapour pressure deficit (D). Across CO₂ treatments, growth in elevated temperature increased E an average of 37%, and did not affect ITE. Combined, growth in elevated [CO₂] and elevated temperature increased E an average of 19% compared with the ACAT treatment. The CO₂ supply and growth temperature did not significantly affect stomatal sensitivity to D or the relationship between g_s and net photosynthetic rates. This study suggests that elevated [CO₂] may not completely ameliorate the effect of elevated temperature on E, and that climate change may substantially alter needle‐level water loss and water use efficiency of Douglas‐fir seedlings.     

Annual cycles of water vapour and carbon dioxide fluxes in and above a boreal aspen forest

Water vapour and CO₂ fluxes were measured using the eddy correlation method above and below the overstorey of a 21-m tall aspen stand in the boreal forest of central Saskatchewan as part of the Boreal Ecosystem-Atmosphere Study (BOREAS). Measurements were made at the 39.5-m and 4-m heights using 3-dimensional sonic anemometers (Kaijo-Denki and Solent, respectively) and closed-path gas analysers (LI-COR 6262) with 6-m and 4.7-m long heated sampling tubing, respectively. Continuous measurements were made from early October to mid-November 1993 and from early February to late-September 1994. Soil CO₂ flux (respiration) was measured using a LI-COR 6000-09 soil chamber and soil evaporation was measured using Iysimetry. The leaf area index of the aspen and hazelnut understorey reached 1.8 and 3.3, respectively. The maximum daily evapotranspiration (E) rate was 5–6 mm d^?1. Following leaf-out the hazelnut and soil accounted for 22% of the forest E. The estimated total E was 403 mm for 1994. About 88% of the precipitation in 1994 was lost as evapotranspiration. During the growing season, the magnitude of half-hourly eddy fluxes of CO₂ from the atmosphere into the forest reached 1.2 mg CO₂ m^?2 s^?1 (33 μmol C m^?2 s^?1) during the daytime. Downward eddy fluxes at the 4-m height were observed when the hazelnut was growing rapidly in June and July. Under well-ventilated night-time conditions, the eddy fluxes of CO₂ above the aspen and hazelnut, corrected for canopy storage, increased exponentially with soil temperature at the 2-cm depth. Estimates of daytime respiration rates using these relationships agreed well with soil chamber measurements. During the 1994 growing season, the cumulative net ecosystem exchange (NEE) was -3.5 t C ha^?1 y^?1 (a net gain by the system). For 1994, cumulative NEE, ecosystem respiration (R) and gross ecosystem photosynthesis (GEP = R - NEE) were estimated to be -1.3, 8.9 and 10.2 t C ha^?1 y^?1 respectively. Gross photosynthesis of the hazelnut was 32% of GEP.