The influence of soil temperature and water content on belowground hydraulic conductance and leaf gas exchange in mature trees of three boreal species

Understanding stomatal regulation is fundamental to predicting the impact of changing environmental conditions on vegetation. However, the influence of soil temperature (ST) and soil water content (SWC) on canopy conductance (g_s) through changes in belowground hydraulic conductance (k_bg) remains poorly understood, because k_bg has seldom been measured in field conditions. Our aim was to (a) examine the dependence of k_bg on ST and SWC, (b) examine the dependence of g_s on k_bg and (c) test a recent stomatal optimization model according to which g_s and soil-to-leaf hydraulic conductance are strongly coupled. We estimated k_bg from continuous sap flow and xylem diameter measurements in three boreal species. k_bg increased strongly with increasing ST when ST was below +8°C, and typically increased with increasing SWC when ST was not limiting. g_s was correlated with k_bg in all three species, and modelled and measured g_s were well correlated in Pinus sylvestris (a model comparison was only possible for this species). These results imply an important role for k_bg in mediating linkages between the soil environment and leaf gas exchange. In particular, our finding that ST strongly influences k_bg in mature trees may help us to better understand tree behaviour in cold environments.     

华北落叶松冠层平均气孔导度模拟及其对环境因子的响应

冠层气孔导度是生态水文学研究中的一个重要参数,研究其对环境因子的响应,能为建立机理性的森林蒸腾模型提供理论依据.本文利用热扩散探针,于2005年5-9月,测定了六盘山叠叠沟小流域华北落叶松人工林树干液流及其同步的环境因子,计算了林分冠层平均气孔导度(gc)并构建了Jarvis形式的冠层平均气孔导度模型,分析了gc对光合有效辐射(PAR)、空气水汽压亏缺(DVP)和土壤相对有效含水率(REW)的响应.结果表明:该模型能有效地计算gc的日变化特征,计算与观测的gc决定系数R2为0.76 (n=952).gc对各环境因子有不同响应关系,并表现为非线性特征.其中PAR是gc的驱动因子,当PAR＜0.35 mmol·m-2·s-1时驱动作用明显,大于此值则驱动作用变小;DVP是gc的限制因子,随着DVP的增加gc降低;REW=41%是gc对土壤水分响应的一个关键阈值,当REW＞41%时,土壤水分对gc的影响较小,当REW＜41%时,土壤水分则成为gc的限制因子.     

Survey and synthesis of intra- and interspecific variation in stomatal sensitivity to vapour pressure deficit

Responses of stomatal conductance (g_s) to increasing vapour pressure deficit (D) generally follow an exponential decrease described equally well by several empirical functions. However, the magnitude of the decrease – the stomatal sensitivity – varies considerably both within and between species. Here we analysed data from a variety of sources employing both porometric and sap flux estimates of g_s to evaluate the hypothesis that stomatal sensitivity is proportional to the magnitude of g_s at low D ( ≤ 1 kPa). To test this relationship we used the function g_s = g_sref–m· lnD where m is the stomatal sensitivity and g_sref = g_s at D = 1 kPa. Regardless of species or methodology, m was highly correlated with g_sref (average r² = 0·75) with a slope of approximately 0·6. We demonstrate that this empirical slope is consistent with the theoretical slope derived from a simple hydraulic model that assumes stomatal regulation of leaf water potential. The theoretical slope is robust to deviations from underlying assumptions and variation in model parameters. The relationships within and among species are close to theoretical predictions, regardless of whether the analysis is based on porometric measurements of g_s in relation to leaf-surface D (D_s), or on sap flux-based stomatal conductance of whole trees (G_Si), or stand-level stomatal conductance (G_S) in relation to D. Thus, individuals, species, and stands with high stomatal conductance at low D show a greater sensitivity to D, as required by the role of stomata in regulating leaf water potential.     

三种植被恢复树种的冠层气孔导度特征及其对环境因子的敏感性

应用Granier热消散探针,长期监测华南地区荷木、大叶相思和柠檬桉林不同径级样树的树干液流,结合同步观测的气象数据,求算冠层气孔导度(g_c),并分析其对环境因子的响应方式及敏感性.结果表明: 不同季节荷木林日间平均g_c显著高于大叶相思和柠檬桉(P<0.05)（除3月外）.在干季和湿季,g_c与光合有效辐射(PAR)呈现对数正相关关系(P<0.001),湿季g_c对PAR响应比干季更敏感.g_c与水汽压亏缺(VPD)在干湿季均呈现对数负相关关系(P<0.001),同样在湿季表现出更高的敏感性.湿季g_c与VPD的偏相关系数高于干季,VPD对气孔行为的调控作用在湿季更为明显.随着土壤含水量的降低,g_c对VPD的敏感性下降,荷木和柠檬桉林下降的幅度大于大叶相思林,荷木和柠檬桉林下降的幅度相当.通过综合分析g_c对环境因子(PAR和VPD)的敏感性及其对土壤含水量变化的响应规律,发现乡土树种荷木作为植被恢复树种比外来引种的大叶相思和柠檬桉更为适宜.     

Responses of stomatal conductance to light, humidity and temperature in winter wheat and barley grown at three concentrations of carbon dioxide in the field

Responses of leaf stomatal conductance to light, humidity and temperature were characterized for winter wheat and barely grown at ambient (about 350 μmol mol^?1 in the daytime), ambient + 175 and ambient + 350 μmol mol^?1 concentrations of carbon dioxide in open‐topped chambers in field plots over a three year period. Stomatal responses to environment were determined by direct manipulation of single environmental factors, and those results were compared with responses derived from natural day to day variation in mid‐day stomatal conductance. The purpose of these experiments was to determine the magnitude of reduction in stomatal conductance at elevated [CO₂], and to assess whether the relative response of conductance to elevated [CO₂] was constant across light, humidity and temperature conditions. The results indicated that light, humidity and temperature all significantly affected the relative decrease in stomatal conductance at elevated [CO₂]. The relative decrease in conductance with elevated [CO₂] was greater at low light, low water vapour pressure difference, and high temperature in both species. For measurements made at saturating light near mid‐day, the ratio of mid‐day stomatal conductances at doubled [CO₂] to that at ambient [CO₂] ranged from 0.42 to 0.86, with a mean of 0.66 in barley, and from 0.33 to 0.80, with a mean of 0.56 in wheat. Day‐to‐day variation in the relative effect of elevated [CO₂] on conductance was correlated with the relative stimulation of [CO₂] assimilation rate and with temperature. Some limitations of multiple linear regression, multiplicative, and ‘Ball–Berry' models as summaries of the data are discussed. In barley, a better fit to the models occurred in individual years than for the combined data, and in wheat a better fit to the models occurred when data from near the end of the season were removed.     

Stomatal response to air humidity and its relation to stomatal density in a wide range of warm climate species

The gas exchange of 19 widely different warm climate species was observed at different leaf to air vapour pressure deficits (VPD). In all species stomata tended to close as VPD increased resulting in a decrease in net photosynthesis. The absolute reduction in leaf conductance per unit increase in VPD was greatest in those species which had a large leaf conductance at low VPDs. This would be expected even if stomata of all species were equally sensitive. However the percentage reduction in net photosynthesis (used as a measure of the relative sensitivity of stomata of the different species) was also closely related to the maximal conductance at low VPD. Similarily the relative sensitivity of stomata to changes in VPD was closely related to the weighted stomatal density or crowding index.The hypothesis is presented that stomatal closure at different VPDs is related to peristomatal evaporation coupled with a high resistance between the epidermis and the mesophyll and low resistance between the stomatal apparatus and the epidermal cells. This hypothesis is consistent with the greater relative sensitivity of stomata on leaves with a high crowding index.The results and the hypothesis are discussed in the light of selection, for optimal productivity under differing conditions of relative humidity and soil water availablility, by observation of stomatal density and distribution on the two sides of the leaf.Visiting scientist, plant physiologist and research assitant of the Cassava Program     

Application of a Coupled Photosynthesis–Stomatal Conductance Model to Analysis of Carbon Assimilation by Spruce and Larch Trees in the Forests of Russia

Photosynthetic activities of common spruce (Picea abies (L.) Karst.) and Dahurian larch (Larix gmelinii (Rupr) Rupr ex Kuzen) were analyzed on the basis of datasets obtained for 110- to 130-year-old forest stands in Middle Russia and East Siberia. Using a Li-Cor 6200 gas analyzer, photosynthesis was measured in parallel with transpiration, stomatal conductance, CO₂ concentrations in ambient air and intercellular spaces, the photosynthetically active radiation, air temperature, and air humidity. The data were examined within the framework of a biochemical model of photosynthesis of Farquhar et al. [1] in combination with the stomatal conductance model proposed by Jarvis [2]. The species-specific differences in carbon assimilation rates were discovered, and dependences of photosynthesis on the needle age, light regime, and growth conditions were revealed. The model parameters obtained were used to simulate the photosynthetic rates in spruce and larch trees at various weather conditions.     

Physiological acclimation dampens initial effects of elevated temperature and atmospheric CO2 concentration in mature boreal Norway spruce

Physiological processes of terrestrial plants regulate the land–atmosphere exchange of carbon, water, and energy, yet few studies have explored the acclimation responses of mature boreal conifer trees to climate change. Here we explored the acclimation responses of photosynthesis, respiration, and stomatal conductance to elevated temperature and/or CO₂ concentration ([CO₂]) in a 3‐year field experiment with mature boreal Norway spruce. We found that elevated [CO₂] decreased photosynthetic carboxylation capacity (?23% at 25 °C) and increased shoot respiration (+64% at 15 °C), while warming had no significant effects. Shoot respiration, but not photosynthetic capacity, exhibited seasonal acclimation. Stomatal conductance at light saturation and a vapour pressure deficit of 1 kPa was unaffected by elevated [CO₂] but significantly decreased (?27%) by warming, and the ratio of intercellular to ambient [CO₂] was enhanced (+17%) by elevated [CO₂] and decreased (?12%) by warming. Many of these responses differ from those typically observed in temperate tree species. Our results show that long‐term physiological acclimation dampens the initial stimulation of plant net carbon assimilation to elevated [CO₂], and of plant water use to warming. Models that do not account for these responses may thus overestimate the impacts of climate change on future boreal vegetation–atmosphere interactions.     

Seasonal and within‐canopy variation in shoot‐scale resource‐use efficiency trade‐offs in a Norway spruce stand

Previous leaf‐scale studies of carbon assimilation describe short‐term resource‐use efficiency (RUE) trade‐offs where high use efficiency of one resource requires low RUE of another. However, varying resource availabilities may cause long‐term RUE trade‐offs to differ from the short‐term patterns. This may have important implications for understanding canopy‐scale resource use and allocation. We used continuous gas exchange measurements collected at five levels within a Norway spruce, Picea abies (L.) karst., canopy over 3 years to assess seasonal differences in the interactions between shoot‐scale resource availability (light, water and nitrogen), net photosynthesis (A_n) and the use efficiencies of light (LUE), water (WUE) and nitrogen (NUE) for carbon assimilation. The continuous data set was used to develop and evaluate multiple regression models for predicting monthly shoot‐scale A_n. These models showed that shoot‐scale A_n was strongly dependent on light availability and was generally well described with simple one‐ or two‐parameter models. WUE peaked in spring, NUE in summer and LUE in autumn. However, the relative importance of LUE for carbon assimilation increased with canopy depth at all times. Our results suggest that accounting for seasonal and within‐canopy trade‐offs may be important for RUE‐based modelling of canopy carbon uptake.     

Linking variability of tree water use and growth with species resilience to environmental changes

Tree growth is an indicator of tree vitality and its temporal variability is linked to species resilience to environmental changes. Second-order statistics that quantify the cross-scale temporal variability of ecophysiological time series (statistical memory) could provide novel insights into species resilience. Species with high statistical memory in their tree growth may be more affected by disturbances, resulting in lower overall resilience and higher vulnerability to environmental changes. Here, we assessed the statistical memory, as quantified with the decay in standard deviation with increasing time scale, in tree water use and growth of co-occurring European larch Larix decidua and Norway spruce Picea abies along an elevational gradient in the Swiss Alps using measurements of stem radius changes, sap flow and tree-ring widths. Local-scale interspecific differences between the two conifers were further explored at the European scale using data from the International Tree-Ring Data Bank. Across the analysed elevational gradient, tree water use showed steeper variability decay with increasing time scale than tree growth, with no significant interspecific differences, highlighting stronger statistical memory in tree growth processes. Moreover, Norway spruce displayed slower decay in growth variability with increasing time scale (higher statistical memory) than European larch; a pattern that was also consistent at the European scale. The higher statistical memory in tree growth of Norway spruce in comparison to European larch is indicative of lower resilience of the former in comparison to the latter, and could potentially explain the occurrence of European larch at higher elevations at the Alpine treeline. Single metrics of resilience cannot often summarize the multifaceted aspects of ecosystem functioning, thus, second-order statistics that quantify the strength of statistical memory in ecophysiological time series could complement existing resilience indicators, facilitating the assessment of how environmental changes impact forest growth trajectories and ecosystem services.     

Relative importance of drought,soil quality,and plant species in determining the strength of plant–herbivore interactions

1. Although studies on plant–herbivore interactions comparing different plant species are common, little is known about the importance of environmental conditions in determining variation in herbivory within single plant species. 2. This study assessed the effects of experimentally manipulated nutrient and water availability on plant palatability, and compared these differences with differences among species. The extent to which these patterns can be explained by leaf toughness and specific leaf area was also investigated. Six plant species from the subfamily Carduoideae and four free‐living leaf chewing invertebrates were used in the study. 3. Herbivore preferences were significantly affected by soil nutrients and water regime and varied among plant as well as herbivore species. Generally, herbivores preferred watered plants and plants from nutrient‐poor soil. The effects of soil nutrients and water regime differed between the plant and herbivore species. The differences between the plant species were greater than those between the environmental treatments. Differences at both levels could be partly explained by leaf toughness and specific leaf area. Leaf toughness, in particular, turned to be an important predictor indicating that herbivores preferred species with softer leaves, and species from wetter conditions with reduced leaf toughness. 4. The environmental conditions in which plants are growing have significant effects on plant palatability. Between‐species comparisons thus need to pay attention to this variation. Future studies may consider how the effects of current conditions interact with conditions of plant origin to predict possible effects of changes in environmental conditions on the intensity of plant–herbivore interactions.     

华北山区侧柏冠层气孔导度特征及其对环境因子的响应

冠层气孔导度(g_s)是衡量冠层-大气界面水汽通量的重要生物学常数,研究其特征及对环境因子的响应,能为开展森林冠层水汽交换过程的机理性研究提供理论依据.于2014年利用SF-L热扩散式探针测定了侧柏的树干液流密度(J_s),同步监测光合有效辐射(PAR)、饱和水汽压差(VPD)、气温(T)等环境因子,计算侧柏的冠层气孔导度特征并分析其对各环境因子的响应.结果表明: 侧柏液流密度的日变化总体呈双峰曲线,生长季高于非生长季,且胸径越大液流密度越大;冠层气孔导度日变化与单位叶面积冠层蒸腾(E_L)趋势相近,均呈双峰曲线,生长季的冠层气孔导度和蒸腾较非生长季略高.侧柏冠层气孔导度与空气温度呈抛物线关系,在10 ℃左右冠层气孔导度达到峰谷;光合有效辐射以400 μmol·m^-2·s^-1为界,小于该阈值两者呈正相关关系,大于该阈值则冠层气孔导度受其影响较小;与饱和水汽压差呈负对数函数关系,随饱和水汽压差增大而逐渐降低.较高的空气温度和光合有效辐射、较低的饱和水汽压差有利于侧柏形成较大的冠层气孔导度,进而促进冠层蒸腾.     

Lack of photosynthetic or stomatal regulation after 9 years of elevated [CO2] and 4 years of soil warming in two conifer species at the alpine treeline

Alpine treelines are temperature‐limited vegetation boundaries. Understanding the effects of elevated [CO₂] and warming on CO₂ and H₂O gas exchange may help predict responses of treelines to global change. We measured needle gas exchange of Larix decidua Mill. and Pinus mugo ssp. uncinata DC trees after 9 years of free air CO₂ enrichment (575 µmol mol^?1) and 4 years of soil warming (+4 °C) and analysed δ¹³C and δ¹⁸O values of needles and tree rings. Tree needles under elevated [CO₂] showed neither nitrogen limitation nor end‐product inhibition, and no down‐regulation of maximal photosynthetic rate (A_max) was found. Both tree species showed increased net photosynthetic rates (A_n) under elevated [CO₂] (L. decidua: +39%; P. mugo: +35%). Stomatal conductance (g_H2O) was insensitive to changes in [CO₂], thus transpiration rates remained unchanged and intrinsic water‐use efficiency (iWUE) increased due to higher A_n. Soil warming affected neither A_n nor g_H2O. Unresponsiveness of g_H2O to [CO₂] and warming was confirmed by δ¹⁸O needle and tree ring values. Consequently, under sufficient water supply, elevated [CO₂] induced sustained enhancement in A_n and lead to increased C inputs into this ecosystem, while soil warming hardly affected gas exchange of L. decidua and P. mugo at the alpine treeline.     

The response of an Eastern Amazonian rain forest to drought stress: results and modelling analyses from a throughfall exclusion experiment

Warmer and drier climates over Eastern Amazonia have been predicted as a component of climate change during the next 50–100 years. It remains unclear what effect such changes will have on forest–atmosphere exchange of carbon dioxide (CO₂) and water, but the cumulative effect is anticipated to produce climatic feedback at both regional and global scales. To allow more detailed study of forest responses to soil drying, a simulated soil drought or 'throughfall exclusion' (TFE) experiment was established at a rain forest site in Eastern Amazonia, Brazil, for which time-series sap flow and soil moisture data were obtained. The experiment excluded 50% of the throughfall from the soil. Sap flow data from the forest plot experiencing normal rainfall showed no limitation of transpiration throughout the two monitored dry seasons. Conversely, data from the TFE showed large dry season declines in transpiration, with tree water use restricted to 20% of that in the control plot at the peak of both dry seasons. The results were examined to evaluate the paradigm that the restriction on transpiration in the dry season was caused by limitation of soil-to-root water transport, driven by low soil water potential and high soil-to-root hydraulic resistance. This paradigm, embedded in the soil–plant–atmosphere (SPA) model and driven using on-site measurements, provided a good explanation ( R ² > 0.69) of the magnitude and timing of changes in sap flow and soil moisture. This model-data correspondence represents a substantial improvement compared with other ecosystem models of drought stress tested in Amazonia. Inclusion of deeper rooting should lead to lower sensitivity to drought than the majority of existing models. Modelled annual GPP declined by 13–14% in response to the treatment, compared with estimated declines in transpiration of 30–40%.     

Late-stage canopy tree species with extremely low δ13C and high stomatal sensitivity to seasonal soil drought in the tropical rainforest of French Guiana

We assessed the daily time‐courses of CO₂ assimilation rate (A), leaf transpiration rate (E), stomatal conductance for water vapour (g_s), leaf water potential ( Ψ _w) and tree transpiration in a wet and a dry season for three late‐stage canopy rainforest tree species in French Guiana differing in leaf carbon isotope composition ( δ ¹³C). The lower sunlit leaf δ ¹³C values found in Virola surinamensis ( ? 29·9‰) and in Diplotropis purpurea ( ? 30·9‰), two light‐demanding species, as compared to Eperua falcata ( ? 28·6‰), a shade‐semi‐tolerant species, were clearly associated with higher maximum g_s values of sunlit leaves in the two former species. These two species were also characterized by a high sensitivity of g_s, sap flow density (Ju) and canopy conductance (g_c) to seasonal soil drought, allowing maintenance of high midday Ψ _w values in the dry season. The data for Diplotropis provided an original picture of increasing midday Ψ _w with increasing soil drought. In Virola, stomata were extremely sensitive to seasonal soil drought, leading to a dramatic decrease in leaf and tree transpiration in the dry season, whereas midday Ψ _w remained close to ? 0·3 MPa. The mechanisms underlying such an extremely high sensitivity of stomata to soil drought remain unknown. In Eperua, g_s of sunlit leaves was non‐responsive to seasonal drought, whereas Ju and g_c were lower in the dry season. This suggests a higher stomatal sensitivity to seasonal drought in shaded leaves than in sunlit ones in this species.     

高温干旱下两种冷季型草坪草叶片细胞超微结构的变化

多年生黑麦草和葡匐股颖在36℃下,控水处理6d,叶片细胞的超微结构与对照相比发生明显变化,叶绿体肿胀,被膜和片层破损,部分叶绿体变圆并向细胞中央移位,这可能有利于减少对阳光的吸收;线粒体和脂质球体数量增加;葡匐翦股颖的线粒体脊解体,但黑麦草的线粒体只受轻微影响。结果说明,两种草坪草的耐热性都较弱,但多年生黑麦草的耐热性稍强于匐匐剪胶颖。     

A comparison of sap flux and water relations of leaves of various isolated trees with special reference to foundation movement in clay soil

Diurnal variation in sap flux (S) through stems of six trees, two each of Ulmus procera SALISB., Melaleuca styphelioides SM. and Prunus cerasifera EHRH. ‘Nigra’ (referred to hereafter by their generic names), were estimated from measurements of heat pulse velocities. Leaf water potential (ψ), stomatal conductance (g _s ) and transpiration from leaves (T) of all replicate trees were measured at 1300–1500h, once during the summer. On two separate occasions measurements were made of S, ψ, (g _s ) and T for one each of Ulmus and Melaleuca trees to study diurnal variations in these parameters. A 12×12 m² area around each tree was kept covered to simulate the condition of trees growing on pavements adjacent to residential properties. Sap flux for these tree species was in the order Melaleuca>Ulmus>Prunus. It is suggested that the smaller canopy and sapwood area in Prunus compared to the other two species is responsible for lower water potential and lower transpiration rate than the other species. Detailed analysis of the diurnal variation in sap flux and water relation of leaves of Melaleuca and Ulmus indicated sap flux of Melaleuca to be greater than that of Ulmus at the same transpiration rate per unit leaf area although the sapwood area of the two species was marginally different. This may have been due either to the difference in canopy conductance or in leaf area between the two species. With the assumption that sap flux closely resembles the rate of soil water extraction for both species, results indicate that Melaleuca is likely to extract soil water at a higher rate than Ulmus and hence is capable of causing greater shrinkage and soil movement than Ulmus.     

Photosynthetic characteristics involved in adaptability to Karst soil and alien invasion of paper mulberry (Broussonetia papyrifera (L.) Vent.) in comparison with mulberry (Morus alba L.)

Unlike mulberry (Morus alba, M.a.), paper mulberry (Broussonetia papyrifera, B.p.) can acclimate to Karst soil and incline to alien invasion. The photosynthetic parameters, diurnal changes of carbonic anhydrase, and chlorophyll fluorescence induction, and water potential were measured on sunny days (SD) and cloudy days (CD). Photosynthetic midday depression occurred in B.p. but not in M.a. The irradiance-and CO₂-saturated photosynthetic rates of B.p. were significantly higher than those of M.a. There was no significant difference in water use efficiency between the two species on a SD. The maximum fluorescence, maximum quantum yield, photochemical quenching, and relative electron transport rate in the leaves of B.p. were much higher than those in M.a. The activity of carbonic anhydrase (CA) of B.p., on either an SD or a CD, was much greater than that of M.a. Higher transpiration rate (E) and net photosynthetic rate (P _N) of B.p. resulted in the lack of water in mesophyll cells. Although a higher CA activity of B.p. supplied both water and CO₂ for the photosynthesis of mesophyll cells, water in mesophyll cells was the factor limiting photosynthesis, and the intercellular CO₂ concentration of B.p. was high and stable.