Patterns of wood carbon dioxide efflux across a 2,000-m elevation transect in an Andean moist forest

During a 1-year measurement period, we recorded the CO₂ efflux from stems (R _S) and coarse woody roots (R _R) of 13–20 common tree species at three study sites at 1,050, 1,890 and 3,050 m a.s.l. in an Andean moist forest. The objective of this work was to study elevation changes of woody tissue CO₂ efflux and the relationship to climate variation, site characteristics and growth. Furthermore, we aim to provide insights into important respiration–productivity relationships of a little studied tropical vegetation type. We expected R _S and R _R to vary with dry and humid season conditions. We further expected R _S to vary more than R _R due to a more stable soil than air temperature regime. Seasonal variation in woody tissue CO₂ efflux was indeed mainly attributable to stems. At the same time, temperature played only a small role in triggering variations in R _S. At stand level, the ratio of C release (g C m^?2 ground area year^?1) between stems and roots varied from 4:1 at 1,050 m to 1:1 at 3,050 m, indicating the increasing prevalence of root activity at high elevations. The fraction of growth respiration from total respiration varied between 10 (3,050 m) and 14% (1,050 m) for stems and between 5 (1,050 m) and 30% (3,050 m) for roots. Our results show that respiratory activity and hence productivity is not driven by low temperatures towards higher elevations in this tropical montane forest. We suggest that future studies should examine the limitation of carbohydrate supply from leaves as a driver for the changes in respiratory activity with elevation.     

Stem hydraulic capacitance decreases with drought stress: implications for modelling tree hydraulics in the Mediterranean oak Quercus ilex

Hydraulic modelling is a primary tool to predict plant performance in future drier scenarios. However, as most tree models are validated under non‐stress conditions, they may fail when water becomes limiting. To simulate tree hydraulic functioning under moist and dry conditions, the current version of a water flow and storage mechanistic model was further developed by implementing equations that describe variation in xylem hydraulic resistance (R_X) and stem hydraulic capacitance (C_S) with predawn water potential (Ψ_PD). The model was applied in a Mediterranean forest experiencing intense summer drought, where six Quercus ilex trees were instrumented to monitor stem diameter variations and sap flow, concurrently with measurements of predawn and midday leaf water potential. Best model performance was observed when C_S was allowed to decrease with decreasing Ψ_PD. Hydraulic capacitance decreased from 62 to 25 kg m^?3 MPa^?1 across the growing season. In parallel, tree transpiration decreased to a greater extent than the capacitive water release and the contribution of stored water to transpiration increased from 2.0 to 5.1%. Our results demonstrate the importance of stored water and seasonality in C_S for tree hydraulic functioning, and they suggest that C_S should be considered to predict the drought response of trees with models.     

中国南亚热带森林不同演替阶段土壤呼吸的分离量化

量化森林土壤呼吸(R_S)及其组分对准确地评估森林土壤碳吸存极其重要。该文以鼎湖山南亚热带季风常绿阔叶林及其演替系列针阔叶混交林和马尾松(Pinus massoniana)林为研究对象, 采用挖壕沟法结合静态气室CO₂测定法对这3种林分类型的R_S进行分离量化。结果表明: 鼎湖山3种森林演替系列上的森林R_S及其组分(自养呼吸R_A、异养呼吸R_H)均呈现出明显的季节动态, 表现为夏季最高、冬季最低的格局。在呼吸总量上, 季风常绿阔叶林显著高于针阔叶混交林和马尾松林, 但混交林与马尾松林之间差异不显著; R_A除季风常绿阔叶林显著大于针阔叶混交林外, 其余林分之间差异不显著; 对于R_H来说, 3个林分之间均无显著差异。随着森林正向演替的进行, 由马尾松林至针阔叶混交林至季风常绿阔叶林, R_A对土壤总呼吸的年平均贡献率分别为(39.48 ± 15.49)%、(33.29 ± 17.19)%和(44.52 ± 10.67)%, 3个林分之间差异不显著。方差分析结果表明, 土壤温度是影响R_S及其组分的主要环境因子, 温度与R_S及其组分呈显著的指数关系; 土壤含水量对R_S的影响不显著, 甚至表现为轻微的抑制现象, 但未达到显著性水平。对温度敏感性指标Q₁₀值的分析表明, 3个林分均为R_A的温度敏感性最大, R_H的温度敏感性最小。     

Elevated CO<Subscript>2</Subscript> reduces sap flux in mature deciduous forest trees

We enriched in CO₂ the canopy of 14 broad-leaved trees in a species-rich, ca. 30-m-tall forest in NW Switzerland to test whether elevated CO₂ reduces water use in mature forest trees. Measurements of sap flux density (J_S) were made prior to CO₂ enrichment (summer 2000) and throughout the first whole growing season of CO₂ exposure (2001) using the constant heat-flow technique. The short-term responses of sap flux to brief (1.5–3 h) interruptions of CO₂ enrichment were also examined. There were no significant a priori differences in morphological and physiological traits between trees which were later exposed to elevated CO₂ (n=14) and trees later used as controls (n=19). Over the entire growing season, CO₂ enrichment resulted in an average 10.7% reduction in mean daily J_S across all species compared to control trees. Responses were most pronounced in Carpinus, Acer, Prunus and Tilia, smaller in Quercus and close to zero in Fagus trees. The J_S of treated trees significantly increased by 7% upon transient exposure to ambient CO₂ concentrations at noon. Hence, responses of the different species were, in the short term, similar in magnitude to those observed over the whole season (though opposite because of the reversed treatment). The reductions in mean J_S of CO₂-enriched trees were high (22%) under conditions of low evaporative demand (vapour pressure deficit, VPD <5 hPa) and small (2%) when mean daily VPD was greater than 10 hPa. During a relatively dry period, the effect of elevated CO₂ on J_S even appeared to be reversed. These results suggest that daily water savings by CO₂-enriched trees may have accumulated to a significantly improved water status by the time when control trees were short of soil moisture. Our data indicate that the magnitude of CO₂ effects on stand transpiration will depend on rainfall regimes and the relative abundance of the different species, being more pronounced under humid conditions and in stands dominated by species such as Carpinus and negligible in mono-specific Fagus forests.     

Hydraulic properties and photosynthetic rates in co-occurring lianas and trees in a seasonal tropical rainforest in southwestern China

In this study, we examined wood anatomy, hydraulic properties, photosynthetic rate, and water status and osmotic regulation in three liana species and three tree species co-occurring in a seasonal tropical rain forest. Our results showed that the three liana species had larger vessel diameter, lower sapwood density, and consequently higher branch sapwood specific hydraulic conductivity (K _S) than the three tree species. Across species, K _S was positively correlated with leaf nitrogen concentration and maximum net CO₂ assimilation rate. However, it was also positively correlated with xylem water potential at 50% loss of hydraulic conductivity, indicating a trade-off between hydraulic efficiency and safety. Compared to the tree species, the liana species had higher predawn leaf water potential and lower osmotic adjustment in the dry season. The combination of more efficient water transport, higher photosynthetic rates, and their ability to access to more reliable water source at deeper soil layers in the dry season in the lianas should contribute to their fast growth.     

Soil respiration in six temperate forests in China

Scaling soil respiration (R_S), the major CO₂ source to the atmosphere from terrestrial ecosystems, from chamber‐based measurements to ecosystems requires studies on variations and correlations of R_S from various biomes and across geographic regions. However, few studies on R_S are available for Chinese temperate forest despite the importance of this forest in the national and global carbon budgets. In this study, we conducted 18‐month R_S measurements during 2004–2005 in six temperate forest types, representing the typical secondary forest ecosystems across various site conditions in northeastern China: Mongolian oak (Quercus mongolica Fisch.), aspen‐birch (Populous davidiana Dode and Betula platyphylla Suk.), mixed deciduous (no dominant tree species), hardwood (dominated by Fraxinus mandshurica Rupr., Juglans mandshurica Maxim., and Phellodendron amurense Rupr.) forests, Korean pine (Pinus koraiensis Sieb. et Zucc.) and Dahurian larch (Larix gmelinii Rupr.) plantations. Our specific objectives were to: (1) explore relationships of R_S against soil temperature and water content for the six forest ecosystems, (2) quantify annual soil surface CO₂ flux and its relations to belowground carbon storage, (3) examine seasonal variations in R_S and related environmental factors, and (4) quantify among‐ and within‐ecosystem variations in R_S. The R_S was positively correlated to soil temperature in all forest types, and was significantly influenced by the interactions of soil temperature and water content in the pine, larch, and mixed deciduous forests. The sensitivity of R_S to soil temperature at 10 cm depth (Q₁₀) ranged from 2.61 in the oak forest to 3.75 in the aspen‐birch forests. The Q₁₀ tended to increase with soil water content until reaching a threshold, and then decline. The annual R_S for the larch, pine, hardwood, oak, mixed deciduous, and aspen‐birch forests averaged 403, 514, 781, 785, 786, and 813 g C m^?2 yr^?1, respectively. The annual R_S of the broadleaved forests was 72% greater than that of the coniferous forests. The annual R_S was positively correlated to soil organic carbon (SOC) concentration at O horizon (R²=0.868) and total biomass of roots <0.5 cm in diameter (R²=0.748). The coefficient of variation (CV) of R_S among forest types averaged 25% across the 18‐month measurements. The CV of R_S within plots varied from 20% to 27%, significantly (P<0.001) greater than those among plots (9–15%), indicating the importance of the fine‐scaled heterogeneity in R_S. This study emphasized that variations in soil respiration and potential sampling bias should be appropriately tackled for accurate soil CO₂ flux estimates.     

Annual wood production in a tropical rain forest in NE Costa Rica linked to climatic variation but not to increasing CO2

Increased atmospheric [CO₂] could theoretically lead to increased forest productivity (‘CO₂ fertilization’). This mechanism was hypothesized as a possible explanation for biomass increases reported from tropical forests in the last 30+ years. We used unique long‐term records of annually measured stands (eighteen 0.5 ha plots, 10 years) and focal tree species (six species, 24 years) to assess the effects of rainfall, temperature, and atmospheric [CO₂] on annual wood production in a neotropical rain forest. Our study area was a meso‐scale section (600 ha) of old‐growth Tropical Wet Forest in NE Costa Rica. Using the repeated remeasurements we directly assessed the relative effects of interannual climatic variation and increasing atmospheric [CO₂] on wood production. A remarkably simple two‐factor model explained 91% of the interannual variance in stand‐level tree growth; the statistically independent factors were total dry season rainfall (positive effect, r²=0.85) and night‐time temperature (negative effect, r²=0.42). Stand‐level tree mortality increased significantly with night‐time temperature. After accounting for dry season rainfall and night‐time temperature, there was no effect of annual [CO₂] on tree growth in either the stand or focal species data. Tree growth in this Tropical Wet Forest was surprisingly sensitive to the current range of dry season conditions and to variations in mean annual night‐time temperature of 1–2°. Our results suggest that wood production in the lowland rainforests of NE Costa Rica (and by extension in other tropical regions) may be severely reduced in future climates that are only slightly drier and/or warmer.     

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.     

Stem respiration and carbon dioxide efflux of young <Emphasis Type="Italic">Populus deltoides</Emphasis> trees in relation to temperature and xylem carbon dioxide concentration

Oxidative respiration is strongly temperature driven. However, in woody stems, efflux of CO₂ to the atmosphere (E _A), commonly used to estimate the rate of respiration (R _S), and stem temperature (T _st) have often been poorly correlated, which we hypothesized was due to transport of respired CO₂ in xylem sap, especially under high rates of sap flow (f _s). To test this, we measured E _A, T _st, f _s and xylem sap CO₂ concentrations ([CO₂*]) in 3-year-old Populus deltoides trees under different weather conditions (sunny and rainy days) in autumn. We also calculated R _S by mass balance as the sum of both outward and internal CO₂ fluxes and hypothesized that R _S would correlate better with T _st than E _A. We found that E _A sometimes correlated well with T _st, but not on sunny mornings and afternoons or on rainy days. When the temperature effect on E _A was accounted for, a clear positive relationship between E _A and xylem [CO₂*] was found. [CO₂*] varied diurnally and increased substantially at night and during periods of rain. Changes in [CO₂*] were related to changes in f _s but not T _st. We conclude that changes in both respiration and internal CO₂ transport altered E _A. The dominant component flux of R _S was E _A. However, on a 24-h basis, the internal transport flux represented 9–18% and 3–7% of R _S on sunny and rainy days, respectively, indicating that the contribution of stem respiration to forest C balance may be larger than previously estimated based on E _A measurements. Unexpectedly, the relationship between R _S and T _st was sometimes weak in two of the three trees. We conclude that in addition to temperature, other factors such as water deficits or substrate availability exert control on the rate of stem respiration so that simple temperature functions are not sufficient to predict stem respiration.     

Genotypic variation in transpiration of coppiced poplar during the third rotation of a short‐rotation bio‐energy culture

The productivity of short‐rotation coppice (SRC) plantations with poplar (Populus spp.) strongly depends on soil water availability, which limits the future development of its cultivation, and makes the study of the transpirational water loss particularly timely under the ongoing climate change (more frequent drought and floods). This study assesses the transpiration at different scales (leaf, tree and stand) of four poplar genotypes belonging to different species and from a different genetic background grown under an SRC regime. Measurements were performed for an entire growing season during the third year of the third rotation in a commercial scale multigenotype SRC plantation in Flanders (Belgium). Measurements at leaf level were performed on specific days with a contrasted evaporative demand, temperature and incoming shortwave radiation and included stomatal conductance, stem and leaf water potential. Leaf transpiration and leaf hydraulic conductance were obtained from these measurements. To determine the transpiration at the tree level, single‐stem sap flow using the stem heat balance (SHB) method and daily stem diameter variations were measured during the entire growing season. Sap flow‐based canopy transpiration (E_c), seasonal dry biomass yield, and water use efficiency (WUE; g aboveground dry matter/kg water transpired) of the four poplar genotypes were also calculated. The genotypes had contrasting physiological responses to environmental drivers and to soil conditions. Sap flow was tightly linked to the phenological stage of the trees and to the environmental variables (photosynthetically active radiation and vapor pressure deficit). The total E_c for the 2016 growing season was of 334, 350, 483 and 618 mm for the four poplar genotypes, Bakan, Koster, Oudenberg and Grimminge, respectively. The differences in physiological traits and in transpiration of the four genotypes resulted in different responses of WUE.     

Seasonal variation in soil CO<Subscript>2</Subscript> efflux in evergreen coniferous and broad-leaved deciduous forests in a cool-temperate forest,central Korea

We measured the soil surface CO₂ efflux (R _S) from January 2005 to December 2006 in two neighboring stands in Gwangneung Forest, central Korea: evergreen coniferous forest (Abies holophylla, stand A) and broad-leaved deciduous forest (Quercus-dominated, stand Q). Regarding seasonal variation, R _S rate was low during the winter and early spring months in each stand and peaked in late July [1170 (stand A) and 1130 (stand Q) in 2005, and 1000 (stand A) and 740 (stand Q) mg CO₂ m⁻² h⁻¹ in 2006]. R _S rate was higher in stand A than in stand Q during most of the growing season. The pattern of summer rainfall differed between 2005 and 2006. R _S rate for both stands was suppressed significantly by the droughts in June 2005 and September 2006. After the heavy rainfall of July 2006, R _S rate was lower than in July 2005 in both stands, but this decrement was much greater in stand Q than in stand A. In midsummer (August) 2006, under higher soil temperature (ST) and lower soil water content (SWC) conditions than in August 2005, R _S rate of stand A was lower than that in August 2005, whereas stand Q showed no marked change. The exponential relationship between ST and R _S accounted for approximately 91–97% of the R _S variability in each stand and in each year. In stand A, the application of a second-order polynomial function indicated a significant correlation between SWC and R _S when the soil was warm (ST > 15°C). Our results suggest that the seasonality of R _S is strongly affected by the pattern of summer rainfall even in an Asia monsoon climate regime. In addition, the vegetation type (i.e., evergreen coniferous forest vs. broad-leaved deciduous forest) plays a significant role in response of R _S to various environmental fluctuations such as drought, heavy rainfall, and hot-dry condition.     

帽儿山不同年龄森林土壤呼吸速率的影响因子

为探明东北温带森林恢复过程中土壤呼吸(R_S)的变化趋势及其影响因子,在帽儿山选取皆伐后天然更新恢复的4个年龄(1a、10a、25a和56a)林分进行了1年的野外原位测定。结果表明:(1)皆伐后天然更新恢复1年、10年、25年和56年林分的年R_S通量差异显著(P0.05),分别为686.5、639.7、733.3、762.3g C m~(-2)a~(-1);其中生长季(5月─10月)和非生长季的R_S通量也存在显著差异,均呈现出随林龄增加先减后增的趋势。全年、生长季和非生长季R_S随林龄变化的变异系数分别为7.6%、6.3%和21.1%,表明非生长季R_S通量的变异性加大了全年R_S通量的差异。(2)4个年龄林分的Rs季节变化趋势相似,且其主控因子均随季节而变:6月─8月Rs与土壤含水率呈二次函数关系(R~2波动在56%─79%之间),其余时段则与土壤温度呈指数函数关系(R~2波动在85%─93%之间)。(3)不同年龄林分生长季R_S与0─20cm土层有机碳(SOC)密度呈正相关关系(R~2=0.434,P0.05),而非生长季R_S与同期土壤5cm温度呈正相关关系(R~2=0.959,P0.01)。本研究区森林皆伐导致R_S降低,随皆伐后森林恢复R_S不断增加,其主导驱动因子是SOC密度的增加和非生长季土壤温度的变化。     

Seasonal differences in leaf-level physiology give lianas a competitive advantage over trees in a tropical seasonal forest

Lianas are an important component of most tropical forests, where they vary in abundance from high in seasonal forests to low in aseasonal forests. We tested the hypothesis that the physiological ability of lianas to fix carbon (and thus grow) during seasonal drought may confer a distinct advantage in seasonal tropical forests, which may explain pan-tropical liana distributions. We compared a range of leaf-level physiological attributes of 18 co-occurring liana and 16 tree species during the wet and dry seasons in a tropical seasonal forest in Xishuangbanna, China. We found that, during the wet season, lianas had significantly higher CO₂ assimilation per unit mass (A _mass), nitrogen concentration (N _mass), and δ¹³C values, and lower leaf mass per unit area (LMA) than trees, indicating that lianas have higher assimilation rates per unit leaf mass and higher integrated water-use efficiency (WUE), but lower leaf structural investments. Seasonal variation in CO₂ assimilation per unit area (A _area), phosphorus concentration per unit mass (P _mass), and photosynthetic N-use efficiency (PNUE), however, was significantly lower in lianas than in trees. For instance, mean tree A _area decreased by 30.1% from wet to dry season, compared with only 12.8% for lianas. In contrast, from the wet to dry season mean liana δ¹³C increased four times more than tree δ¹³C, with no reduction in PNUE, whereas trees had a significant reduction in PNUE. Lianas had higher A _mass than trees throughout the year, regardless of season. Collectively, our findings indicate that lianas fix more carbon and use water and nitrogen more efficiently than trees, particularly during seasonal drought, which may confer a competitive advantage to lianas during the dry season, and thus may explain their high relative abundance in seasonal tropical forests.     

Spatial and seasonal variations in stem CO2 efflux of Pinus canariensis at their upper distribution limit

We calculated stem CO₂ efflux (E _S) of Pinus canariensis at a timberline site in Tenerife, Canary Islands, from March 7, 2008 and February 9, 2009. E _S varied markedly throughout the year. Although E _S generally followed the seasonal trends in temperature, we observed seasonal adjustment of E _S in both E _S normalized to temperature (E _S10) and the temperature sensitivity (Q ₁₀) resulting in lower E _S10 values during the warm and dry season as compared during the cold and wet season; the latter corresponding with the period of stem growth. The Q ₁₀ by contrast, was higher during the warm and dry summer than during the cold and wet season, an observation suggesting that climate extremes such as summer drought in the Mediterranean may override the observation that Q ₁₀ declines with increasing temperature. As concurrent measurements estimating the potential impact of sap flow on E _S revealed no interaction in P. canariensis, the Q ₁₀ values were used along with E _S10 and stem temperature records to predict annual total E _S and for partitioning total E _S into its maintenance (E _m) and a growth (E _g) component.     

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.     

Temporal patterns of water flux in trees and lianas in a Panamanian moist forest

Using constant heat sap flow sensors, xylem water fluxes in ten tree species and two liana species were monitored for 5–10 days during the beginning of the wet season in May, 1993. For a subset of the trees, a branch was also monitored at the top of the crown for 5 days. Xylem flux (J _S) was related diurnally in all plants to vapor pressure deficit (D) measured within the upper-third of the canopy, and to incoming shortwave radiation R _S above the canopy. Cross-correlation analysis was used to estimate time lags between diurnal patterns of J _S and D or R _S, and between J _S in stems and branches. The maximum correlation coefficient from cross-correlation of J _S with R _S (range=0.57–0.92) was often higher than the maximum of J _S with D (range=0.43–0.89), indicating that diurnal J _S was more dependent on R _S than D. Time lags (lag corresponding to maximum correlation) of J _S at stem-base with D was shorter (0–45 min) than with radiation (5–115 min), highly variable within a species, and uncorrelated to the height or exposure of tree crowns or liana in the canopy. On a stand level, not accounting for the diel lag between stem sap flux and canopy flux resulted in errors in estimated canopy transpiration of up to 30%. Received: 19 October 1998 / Accepted: 8 June 1999     

Elevated CO2 ameliorates birch response to high temperature and frost stress: implications for modeling climate-induced geographic range shifts

Despite predictions that both atmospheric CO₂ concentrations and air temperature will rise together, very limited data are currently available to assess the possible interactive effects of these two global change factors on temperate forest tree species. Using yellow birch (Betula alleghaniensis) as a model species, we studied how elevated CO₂ (800 vs. 400 μl l⁻¹) influences seedling growth and physiological responses to a 5°C increase in summer air temperatures (31/26 vs. 26/21°C day/night), and how both elevated CO₂ and air temperature during the growing season influence seedling ability to survive freezing stress during the winter dormant season. Our results show that while increased temperature decreases seedling growth, temperature-induced growth reductions are significantly lower at elevated CO₂ concentrations (43% vs. 73%). The amelioration of high-temperature stress was related to CO₂-induced reductions in both whole-shoot dark respiration and transpiration. Our results also show that increased summer air temperature, and to a lesser degree CO₂ concentration, make dormant winter buds less susceptible to freezing stress. We show the relevance of these results to models used to predict how climate change will influence future forest species distribution and productivity, without considering the direct or interactive effects of CO_2. Received: 5 June 1997 / Accepted: 16 December 1997     

Stem CO<Subscript>2</Subscript> efflux of a <Emphasis Type="Italic">Populus nigra</Emphasis> stand: effects of elevated CO<Subscript>2</Subscript>, fertilization,and shoot size

To determine whether long-term growth in elevated atmospheric CO₂ concentration [CO₂] and nitrogen fertilization affects woody tissue CO₂ efflux, we measured stem CO₂ efflux as a function of temperature in three different size classes of shoots of Populus nigra L. (clone Jean Pourtet) on two occasions in 2004. Trees were growing in a short rotation coppice in ambient (370 μmol mol⁻¹) and elevated (550 μmol mol⁻¹, realised by a Free Air Carbon dioxide Enrichment system) [CO₂], and measurements were performed during the third growing season of the second rotation. Elevated CO₂ did not affect Q₁₀ or specific stem CO₂ efflux (E₁₀) of overall poplar shoots. The lack of any effect of N on stem CO₂ efflux indicated that nutrients were sufficient. Specific stem CO₂ efflux differed significantly between shoot sizes, emphasizing the importance of tree size when scaling-up respiration measurements to the stand level. Variation in stem CO₂ efflux could not be satisfactorily explained by temperature as the only driving variable. We hypothesize that transport of CO₂ with the sapflow might have confounded our results and could explain the high Q₁₀ values reported here. Predicting the respiratory carbon loss in a future elevated [CO₂] world must therefore move beyond the single-factor temperature dependent respiration model and involve multiple factors affecting stem CO₂ efflux rate.     

毛白杨树干呼吸及其温度敏感性季节变化特征

树干呼吸(E_s)是森林生态系统碳循环过程的重要组成部分,深入理解树干呼吸过程对未来气候变暖的响应及反馈机制有助于更加精确地估算森林生态系统碳储量。为揭示毛白杨树干呼吸及其温度敏感性的昼夜变化和季节动态规律,利用Li-Cor6400便携式光合作用测定系统及其配套使用的土壤呼吸测量气室(LI-6400-09)对冀南平原区毛白杨的树干呼吸和树干温度实施为期1年的连续监测。结果表明:(1)在生长季,毛白杨树干呼吸与树干温度之间在晚上呈现正相关的关系(R~2=0.88);相反,两者在白天为负相关的关系(R~2=0.96)。(2)整个观测期内,毛白杨树干呼吸和树干温度均呈现"钟形"的变化曲线,树干呼吸与树干温度之间存在着较好的指数函数关系(R~2=0.93),且树干呼吸的温度敏感性系数(Q_(10))为2.62;不同季节毛白杨树干呼吸的Q_(10)存在差异,生长季的Q_(10)(1.95)明显低于非生长季(3.00),表明生长呼吸和维持呼吸对温度的响应也并不相同。(3)温度矫正后的毛白杨树干呼吸(R_(15))在昼夜和季节尺度上均存在明显的变异,即夜晚的R_(15)显著高于白天(P0.01),生长季的R_(15)明显高于非生长季(P0.05);树干可溶性糖含量与生长季的R_(15)存在较好的相关性(R~2=0.52),而非生长季的R_(15)却主要受到树干淀粉含量的影响。研究结果表明,在生长季,毛白杨树干呼吸的在日变化主要受到温度的影响,而在季节尺度上Q_(10)的变异则与树干呼吸中维持呼吸所占比例及树干中非结构性碳水化合物(可溶性糖和淀粉)的含量及类型紧密相关。     

Tree growth variation in the tropical forest: understanding effects of temperature,rainfall and CO2

Tropical forest responses to climatic variability have important consequences for global carbon cycling, but are poorly understood. As empirical, correlative studies cannot disentangle the interactive effects of climatic variables on tree growth, we used a tree growth model (IBTREE) to unravel the climate effects on different physiological pathways and in turn on stem growth variation. We parameterized the model for canopy trees of Toona ciliata (Meliaceae) from a Thai monsoon forest and compared predicted and measured variation from a tree‐ring study over a 30‐year period. We used historical climatic variation of minimum and maximum day temperature, precipitation and carbon dioxide (CO₂) in different combinations to estimate the contribution of each climate factor in explaining the inter‐annual variation in stem growth. Running the model with only variation in maximum temperature and rainfall yielded stem growth patterns that explained almost 70% of the observed inter‐annual variation in stem growth. Our results show that maximum temperature had a strong negative effect on the stem growth by increasing respiration, reducing stomatal conductance and thus mitigating a higher transpiration demand, and – to a lesser extent – by directly reducing photosynthesis. Although stem growth was rather weakly sensitive to rain, stem growth variation responded strongly and positively to rainfall variation owing to the strong inter‐annual fluctuations in rainfall. Minimum temperature and atmospheric CO₂ concentration did not significantly contribute to explaining the inter‐annual variation in stem growth. Our innovative approach – combining a simulation model with historical data on tree‐ring growth and climate – allowed disentangling the effects of strongly correlated climate variables on growth through different physiological pathways. Similar studies on different species and in different forest types are needed to further improve our understanding of the sensitivity of tropical tree growth to climatic variability and change.