树干液流径向分布格局研究进展

树干液流径向分布的不均匀性是引起热技术估算单株乃至林分蒸腾误差的主要来源。因此, 了解树干液流径向分布格局并将其定量化, 成为利用热扩散和热脉冲技术准确估算森林蒸腾的必要条件。该文详细介绍了树干液流径向分布格局的研究方法, 总结了目前4种常见的树干液流的径向分布格局, 分析了影响树干液流径向分布格局的内部结构因素和外部环境因素, 阐明了树干液流径向分布格局的时间动态特征及其影响因素, 并提出目前研究中存在的问题和可能的解决方法。     

休眠前期玉兰树干液流的变化及其对环境因子的响应

2011年9月26日-11月5日,利用热平衡技术连续测算了引种北京4年的红花玉兰[抗寒型(HK)和非抗寒型(HF)]、黄玉兰(HY)和二乔玉兰(EQ)的树干液流,结合同步观测的环境因子数据,分析休眠前玉兰的树干液流和综合环境因子的变化,以及在0.5h尺度和日尺度下液流对各环境因子的响应.结果表明:休眠前,玉兰的树干液流呈明显的逐日下降趋势.影响玉兰树干液流的7个环境因子可分为气象因子(MI)和土壤因子(SI)两类.树干液流与MI具有同步的单峰节律性变化规律,与SI呈同步降低趋势.MI和SI对树干液流的共同影响程度为69.8％ ～73.2％.0.5 h尺度和日尺度下,玉兰树干液流与总辐射(Rs)、空气水汽压亏缺(D)、相对湿度(RH)、气温(Ta)和风速(w)极显著相关;在0.5h尺度下与土壤温度(Ts)和土壤含水量(SWC)的相关性不显著,在日尺度下与Ts、SWC和昼长(Z)极显著相关,相关系数在0.8左右.日尺度下进入各回归模型的环境因子有Rs、Z和D,而0.5h尺度下除SWC和Ts外其余环境因子均进入模型,且日尺度逐步回归决定系数(0.92 ～0.96)大于0.5h尺度(0.77 ～0.87)(除HF外).除HF外,其余玉兰的液流变化规律与各单一环境因子间的相关程度一致,这与HF无法正常越冬而其余玉兰在北京顺利越冬的实际情况相一致.     

黄土丘陵区刺槐和侧柏人工林树干液流特征及其对降水的响应

水分供应不足及水热不同步常导致黄土丘陵地区在春末和夏初出现季节性干旱。为阐明该地区主要造林树种的蒸腾耗水特征及其对降水的响应, 使用热扩散式树干茎流计(TDP)于2009年4-10月对黄土丘陵区安塞国家生态试验站刺槐(Robinia pseudoacacia)和侧柏(Platycladus orientalis)的树干液流密度(F_d)进行连续观测, 并同步测定了气象、土壤水分等环境因子。结果表明: 刺槐和侧柏在生长季内不同生长时期的F_d均表现为单峰型日变化特征, 刺槐最高液流峰值为0.12068 m³·m^-2·h^-1, 是侧柏最高液流值(0.03737 m³·m^-2·h^-1)的3.23倍。除生长旺盛季(7-8月)外, 刺槐和侧柏降水后的F_d明显高于降水前。同时反映水汽压差(VPD)和太阳辐射(R_s)的蒸腾变量(VT)能够很好地模拟F_d, 且两者呈显著的指数正相关关系, 随VT的增加F_d逐渐增大, VT增加到50 kPa (W·m^-2)^1/2左右时, F_d的变化趋于稳定; 通过对降水前后两个树种水力导度(拟合参数b值)分析, 相对于侧柏, 刺槐更易受降水的影响(p < 0.001)。因此, 可认为刺槐是降水敏感型植物, 而侧柏是降水不敏感型植物。该研究通过分析黄土丘陵区人工林树种对降水的差异性响应, 从树木水分利用方面能够为当地生态恢复过程中人工林的管理提供科学依据。     

降雨减少对刺槐树干液流特征的影响

在全球气候变化背景下,区域降水格局正在发生改变,对森林生态系统生产力和水文过程将产生重要影响。为揭示降雨量减少对树木蒸腾耗水特征及其响应环境因子的影响,本研究以黄土高原半湿润区主要造林树种刺槐(Robinia pseudoacacia)为对象,在减雨样地林分行间布设人工减雨板减少约47%的降雨输入,采用Granier型热扩散探针测定树干液流动态,并同步监测太阳辐射、空气温度、相对湿度、降雨事件和土壤含水量等环境因子,分析了树干液流对减雨处理及季节性土壤水分变化的响应特征。结果表明：减雨处理显著降低了刺槐液流通量密度标准化值,处理第3年标准化液流通量密度显著低于对照样地;减雨样地刺槐标准化液流日变化峰值时间早于对照样地,表明减雨样地刺槐较早实施了气孔调节,改变了其与气象因子的时滞时间;标准化液流通量密度响应蒸腾驱动因子的拟合方程参数值在样地间呈极显著差异,显示减雨样地刺槐响应气象因子的敏感性有所降低。降雨量的改变不仅影响林地土壤水分状况,还会对刺槐蒸腾耗水及其对环境因子的响应产生影响。     

树木胸径大小对树干液流变化格局的偏度和时滞效应

通过分析具不同水力结构的马占相思、荷木和粉单竹液流变化格局的偏度和时滞,探讨液流的空间分布特征及对冠层蒸腾的影响。结果表明:荷木的液流格局偏度和时滞随树木胸径的增加呈减小的趋势,但马占相思由于冠层开阔和林分分化程度高而规律不明显,粉单竹液流偏度随胸径减少,由于冠幅较小,接受的光照较均匀,个体间的时滞差异不明显,但时滞值比胸径近似的荷木小。树干水分传输过程中存在液流再分配的现象,边材的导水效率可能是影响时滞的重要原因。冠层蒸腾的空间异质性与树木储存水有关,大树储存水较多,冠层蒸腾的异质性小;小树储存水较少,液流被优先分配到光照充足的东南方位,导致冠层蒸腾较高的异质性。旱季受土壤水分的限制,大树储存水对蒸腾的贡献大于湿季,而小树蒸腾由于受到储水容量的制约,储存水对蒸腾的贡献小于湿季。冠层接受光照的迟或早以及辐射量的大小是引起蒸腾时间变化格局和树干不同方位液流格局差异的重要原因,但液流的横向交换弱化了这种现象,往往是个体间的差异掩盖了方位的差异。湿季较小胸径的树木比偏值(枝下高与胸高处液流偏度的比值)大于旱季,而较大胸径的树木比偏值恰好相反,总体而言,比偏值随着胸径的增加而逐渐下降。     

三种温带树种树干储存水对蒸腾的贡献

树干储存水在协调叶片水分和碳平衡、维持树木水分收支平衡中起着重要作用。以无孔材红松(Pinus koraiensis)、散孔材山杨(Populus davidiana)和环孔材蒙古栎(Quercus mongolica)为对象, 于2010年8月中旬至9月末(生长季后期)测定其冠基和干基树干液流通量以及树干储存水的日变化过程, 量化分析树干储存水对日蒸腾量的贡献及其生物影响因子。结果表明: 冠基的液流比干基启动得早, 而且两者在一天中存在显著的时滞。树干储存水的日进程总体上可分为: 完全释放、以释放为主补充为辅、以补充为主释放为辅、饱和稳定等4个阶段, 但每个阶段的持续时间和变化格局随树种而变。红松的树干储存水在一天内表现出两个释放-补充周期, 而两种阔叶树种均只经历了一次释放-补充过程。在测定时段内红松、山杨、蒙古栎标准化到平均木(边材体积为0.29 m³)的树干储存水释放量分别为: (3.4 ± 1.5)、(2.4 ± 0.6)和(1.5 ± 0.4) kg·d ^-1, 分别占日蒸腾量的18.9%、17.1%和8.8%。树干储存水释放量与日蒸腾量呈显著的指数函数关系, 而与干基的边材面积和树高呈正相关关系。该研究突显了树木大小(树高和边材面积)和材性特征对树干储存水释放量及其对蒸腾量贡献的重要影响。     

黄土高原半湿润区刺槐树干液流对人工截留降雨输入及环境因子的响应

随着全球气候变化加剧, 局部地区温度上升和降水量改变将对区域植被的分布与生长产生重要影响。在黄土高原半湿润及半干旱地区植被恢复中, 刺槐(Robinia pseudoacacia)是大面积种植的人工林树种。为探究该树种蒸腾耗水特征对降水量改变及水分条件差异的响应, 于2015年4月起, 在地处黄土高原半湿润区的陕西省永寿县槐平林场, 于35年生刺槐人工林样地中布设了人工截留降雨试验, 减少了47.5%的降雨输入。处理当年生长季内, 截留降雨处理区0-100 cm土层的平均土壤含水量相对于对照区(23.76%)有明显降低(22.59%)。采用Granier热扩散探针对截留降雨处理区和对照区的样树树干液流动态进行连续监测, 并同步监测主要气象环境因子(太阳辐射、空气温度和湿度)和林地土壤含水量, 分析了截留降雨处理区与对照区树干液流通量密度动态特征及其对环境因子的响应。结果表明: 截留降雨输入处理降低了刺槐树干液流通量密度, 截留降雨处理期间典型天气的平均液流通量密度(1.64 mL·m ^-2·s ^-1)不仅低于同组样树在处理前一年同期的水平(2.42 mL·m ^-2·s ^-1), 而且远低于试验期间对照区样树的平均水平(3.38 mL·m ^-2·s ^-1); 同时, 截留降雨处理还降低了刺槐液流通量密度对气象因子变化的敏感性, 截留降雨处理区样树液流通量密度响应空气水汽压亏缺的拟合方程参数值与对照区样树差异显著。分析可知, 降水量水平不仅影响土壤水分状况, 而且影响刺槐对气象环境因子响应的敏感性, 降水量减少导致的土壤含水量整体降低会使得该区域刺槐蒸腾耗水量下降, 显示其对环境因子的适应性, 但最终会导致生产力的大幅度降低。     

环境和冠层结构对华北落叶松林树干液流的影响

准确量化环境和冠层结构变化对树干液流的影响,对于深入理解变化环境下的树木水分利用机制具有重要意义.在六盘山香水河小流域于2019年6-9月利用热扩散式探针监测了华北落叶松林的树干液流,同步观测林外气象、林内根系层土壤含水量和冠层结构动态,分析树干液流速率与潜在蒸散(PET)、土壤相对可利用水分(REW)和林冠层叶面积指...     

毛红椿人工林树干液流动态变化对坡位的响应

2012年7-10月,采用液流测定系统监测了位于浙江省开化县的毛红椿人工林上下坡位的树干液流,并同步监测生态环境因子,分析了毛红椿人工林树干液流与土壤含水量、温度和水势等环境因子的关系.结果表明: 研究区上、下坡位树干液流日变化均呈典型的“昼高夜低”单峰曲线;下坡位毛红椿树干液流速率平均值显著高于上坡位;上坡位土壤温度显著高于下坡位,而下坡位土壤含水量和土壤水势显著高于上坡位;下坡位土壤含水量和土壤水势是影响毛红椿树干液流速率的主要因子,而上坡位土壤温度和土壤水势对毛红椿树干液流速率有较大影响.     

Granier原始公式计算树干液流速率的适用性分析——以毛白杨为例

热扩散探针(TDP)在林木蒸腾研究中应用极广,其测定数据的计算精度直接影响对林木和林分耗水的精准定量。Granier原始公式(Fd=0.0119K1.231,Fd为液流速率(cm·s^-1),K为温差系数)是计算TDP测定数据的标准方法,但其准确性备受质疑。为系统了解Granier原始公式的适用性和明确对其校正的必要性,该研究以毛白杨(Populus tomentosa)为实验材料,采用室内茎段法和整树容器法,针对不同型号的TDP探针评估Granier原始公式的精度,并对比不同方法校正公式的应用效果。与茎段法实测值相比,Granier原始公式计算的液流速率平均低估52.3%–61.4%。通过茎段法和整树容器法得到的校正公式分别为Fd=0.0362K^1.870和F_d=0.0105K^0.976;且一种方法下得到的校正公式,在其他方法条件下应用时存在较大偏差。与Granier原始公式相比,整树容器法校正公式计算的大田生长状态下的7株林木的平均液流速率没有显著变化,但茎段法和其他研究中得到的毛白杨校正公式的计算结果均显著变大。以整树容器法做对比,Granier原始公式的精度明显高于其他校正公式,其相对平均绝对误差和均方根误差分别为10%和0.0005cm·s^-1。此外,校正公式的系数在不同林木间存在较大差异,但其数值与导水边材中探针长度所占比例呈显著负相关关系。综上,利用TDP测定液流时,可能有必要对Granier原始公式进行校正,但不同方法校正公式的应用效果差异巨大,表明以往研究中得到的校正公式具有很大局限性。同时,该研究未找到充足证据支撑“有必要采用校正公式以精确估算毛白杨液流速率”的观点,尤其是考虑到整树容器法校正公式对大田栽植的毛白杨的液流速率估算结果与Granier原始公式并无显著差异,因此建议继续对该树种延用Granier原始公式。     

Up-scaling to stand transpiration of an Asian temperate mixed-deciduous forest from single tree sapflow measurements

Species diversity in mixed forest stands is one of the factors that complicate up-scaling of transpiration from individual trees to stand level, since tree species are architecturally and functionally different. In this study, thermal dissipation probes were used to measure sap flow in five different tree species in a mixed-deciduous mountain forest in South Korea. Easily measurable tree characteristics that could serve to define individual tree water use among the different species were employed to scale up transpiration from single trees to stand level. Tree water use (TWU) was derived from sap flux density (SFD) and sapwood area (SA). Canopy transpiration E was scaled from TWU while canopy conductance (g _c) was computed from E and VPD. SFD, TWU and g _c were correlated with tree diameter at breast height (DBH) for all the five measured species (SFD: R ² = 0.21, P = 0.036; TWU: R ² = 0.83, P < 0.001; g _c: R ² = 0.63, P < 0.001). Maximum stand transpiration (E) during June, before the onset of the Asian monsoon rains, was estimated at 0.97 ± 0.12 mm per day. There was a good (R ² = 0.94, P < 0.0001) agreement between measured and estimated E using the relationship between TWU and DBH. Our study shows that using functional models that employ converging traits among species could help in estimating water use in mixed forest stands. Compared to SA, DBH is a better scalar for water use of mixed forest stands since it is non-destructive and easily obtainable.     

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.     

树木水分利用效率研究综述

在我国干旱半干旱地区,如何高效地利用极为有限的水资源来合理构建和恢复森林植被,并最大程度地发挥其多种功能,已经成为目前森林生态学研究中的一个热点,而对于树木水分利用效率(WUE)的研究则是其核心和关键内容之一。本文从介绍和剖析树木WUE的概念入手,分别在单叶、个体和群体等三个空间尺度上系统地阐述了WUE概念的内涵及其相应的计算方法,并且对各种方法的优缺点进行了评论。最后,在详细分析国内外树木WUE相关研究进展的基础上,针对我国干旱、半干旱地区水资源短缺和植被恢复与重建之间的矛盾,提出了今后树木WUE的研究发展趋势和应用前景。     

Stable carbon isotopes in tree rings indicate improved water use efficiency and drought responses of a tropical dry forest tree species

Understanding the responses of tropical trees to increasing [CO₂] and climate change is important as tropical forests play an important role in carbon and hydrological cycles. We used stable carbon isotopes (δ¹³C) in tree rings to study the physiological responses of a tropical dry forest tree species in southern Mexico, Mimosa acantholoba to changes in atmospheric [CO₂] and variation in climate. Based on annual records of tree ring δ¹³C, we calculated intrinsic water use efficiency (W _i) and intercellular [CO₂] (c _i). Our results showed that trees responded strongly to the increase in atmospheric [CO₂] over the last four decades; W _i increased dramatically by 40%, while c _i remained largely constant. The maintenance of a constant c _i indicates that photosynthetic rates are unlikely to have increased in response to higher [CO₂], and that improvements in W _i are probably due to a reduction in stomatal conductance. This may have large consequences for the hydrological cycle. Inter-annual variation in c _i was strongly correlated with total annual rainfall (r = 0.70), and not influenced by temperature, solar radiation or cloud cover. Our results show that δ¹³C in tree rings of tropical dry forest trees may be a powerful tool to evaluate long-term responses of trees to increasing [CO₂] and to variation in climate.     

Whole tree xylem sap flow responses to multiple environmental variables in a wet tropical forest

In order to quantify and characterize the variance in rainforest tree physiology, whole tree sap flow responses to local environmental conditions were investigated in 10 species of trees with diverse traits at La Selva Biological Station, Costa Rica. A simple model was developed to predict tree sap flow responses to a synthetic environmental variable generated by a principle components analysis. The best fit was obtained with a sigmoid function which explained between 74 and 93% of the variation in sap flux of individual trees. Sap flow reached an asymptote where higher light and evaporative demand did not cause sap flux to increase further. Soil moisture had little influence on sap flux. The morphological characteristics of the trees significantly affected sap flow; taller trees responded to changes in environmental variables sooner than shorter trees and high liana cover buffered tree sap flow responses to weather. The effect of species‐specific differences on the model was small; the mean effectiveness of the model was reduced by 6% when parameters were estimated from a single pool of measurements taken from all individuals. The results indicate that sap flow response could be effectively estimated using a simple general model and composite environmental index for these 10 diverse tree species.     

亚热带森林演替树种叶片气孔导度对环境水分的水力响应

利用LI-1600稳态气孔计和PMS压力室,在田间测定了群落演替早期强阳生性树种桃金娘(Rhodomyrtus tomentosa)和三叉苦(Evodia lepta)、偏中性的阳生性树种荷木(Schima superba)、群落演替后期的耐荫树种鸭脚木(Schefllera octophylla)和九节(Psychotrie rubra)的叶片气孔导度(gs)和叶片水势(ΨL),研究不同演替阶段树种的气孔导度对环境水分的响应.结果表明,早上叶片有较高的ΨL,随着时间推移ΨL逐渐降低,与此同时比叶水力导度(KL)随ΨL降低而下降,桃金娘、三叉苦、荷木、鸭脚木和九节水力导度初始最低值时的ΨL分别为-1.6、-1.42、-1.30、-0.9MPa和-1.05MPa.随着ΨL降低,田间测定的gs开始从上午的较低值上升至约中午时的最大值,随后开始降低,此时的ΨL分别为-1.58、-1.52、-1.35、-1.02MPa和-1.0MPa.不同植物种类有不同的KL初始最低值的ΨL和gs达到最大值的ΨL.但不论何种供试树种,KL最低值时的ΨL与gs开始从最大值下降时的ΨL相近;显示KL与gs在动态变化中存在协调关系.树种间的gs和KL对ΨL的不同响应显示桃金娘和三叉苦的KL最低值时和gs开始下降时的ΨL均较鸭脚木和九节对应的ΨL低(p<0.05),意味着演替早期树种能在较强水分胁迫下保持较高的气孔导度.这一水力特性保证树种在水分胁迫下维持叶片的光合速率,有利于其在群落中的生长和优势地位的维护,而演替后期树种在较高ΨL下气孔关闭,降低了光合速率.全球变暖和环境进一步干旱可能成为限制亚热带森林植物群落的正向演替进程的潜在因素之一.     

Similar tree seedling responses to shrubs and to simulated environmental changes at Pyrenean and subarctic treelines

Background: Climate, land-use and disturbance regimes are key drivers of treeline dynamics worldwide, but local and regional spatio-temporal patterns indicate that additional factors play an important role. Some studies suggest that shrub-tree interactions control tree seedling recruitment patterns across the treeline ecotone, but little is known about the generality of this interaction.

Aims: We established an experiment in a treeline ecotone in the central Pyrenees to investigate the role of such interactions and other environmental factors on tree seedling growth and survival. It was based on a similar experiment we completed recently in the subarctic Scandes. By comparing local phenomena between both experiments we assessed the generality of the findings across regions with different biogeographic histories and species characteristics.

Methods: We followed the survival and growth of transplanted Pinus uncinata seedlings during three growing seasons in a multi-factorial design (forest vs. treeline, +/– shrub removal, +/– temperature increase and +/– nutrient addition).

Results: There was better seedling growth at the treeline compared to the forest, and the presence of shrubs prevented seedling winter damage and herbivory. Both temperature and nutrient increase enhanced seedling performance.

Conclusions: Although some particular details were exclusive to the Pyrenees or to the Scandes, the similarities indicated general patterns and help to interpret the underlying mechanisms of treeline dynamics in both regions. This convergence of responses points to the potential of the development of a robust mechanistically founded predictive framework for scaling up shrub impacts on treeline dynamics to other regions.     

亚热带山地森林附生地衣的移植生长及其对环境变化的响应

本研究选取云南哀牢山亚热带森林系统6种常见的附生大型地衣进行为期2年的移植实验,分析其在原生林林外、林缘以及林内3种生境下的生物量增长速率和健康率等的差异,并探讨其生长对环境因子变化的响应。结果显示,网肺衣Lobaria retigera和平滑牛皮叶Sticta nylanderiana在林缘处生物量增长最快;而槽枝Sulcaria sulcata、多花松萝Usnea florida、皮革肾岛衣Nephromopsis pallescens、针芽肺衣Lobaria isidiophora 4种地衣则都是在林外光照更强、湿度更低的条件下生长得最好,林内最差。除网肺衣外,所有地衣的健康率均在林外较好,林缘次之,林内最差。非参数相关分析表明,6种地衣的生长速率和光照以及温度呈正相关,与大气湿度呈负相关;温度和光照是影响地衣生长最主要的生境因子。     

Origins of root-mediated pH changes in the rhizosphere and their responses to environmental constraints: A review

The aim of the present review is to define the various origins of root-mediated changes of pH in the rhizosphere, i.e., the volume of soil around roots that is influenced by root activities. Root-mediated pH changes are of major relevance in an ecological perspective as soil pH is a critical parameter that influences the bioavailability of many nutrients and toxic elements and the physiology of the roots and rhizosphere microorganisms. A major process that contributes root-induced pH changes in the rhizosphere is the release of charges carried by H⁺ or OH^– to compensate for an unbalanced cation–anion uptake at the soil–root interface. In addition to the ions taken up by the plant, all the ions crossing the plasma membrane of root cells (e.g., organic anions exuded by plant roots) should be taken into account, since they all need to be balanced by an exchange of charges, i.e., by a release of either H⁺ or OH^–. Although poorly documented, root exudation and respiration can contribute some proportion of rhizosphere pH decrease as a result of a build-up of the CO₂ concentration. This will form carbonic acid in the rhizosphere that may dissociate in neutral to alkaline soils, and result in some pH decrease. Ultimately, plant roots and associated microorganisms can also alter rhizosphere pH via redox-coupled reactions. These various processes involved in root-mediated pH changes in the rhizosphere also depend on environmental constraints, especially nutritional constraints to which plants can respond. This is briefly addressed, with a special emphasis on the response of plant roots to deficiencies of P and Fe and to Al toxicity. Finally, soil pH itself and pH buffering capacity also have a dramatic influence on root-mediated pH changes.