宁夏六盘山主要树种及典型森林植被的水分利用效率研究
摘要
针对我国温带干旱缺水区林水矛盾的问题，本文以六盘山区几种典型森林植被为研究对象，在半湿润气候的香水河小流域和半干旱气候的叠叠沟小流域，采用稳定性碳同位素技术及树木年轮测定仪、树干径向变化记录仪和SF-L树干液流仪等仪器，通过估算典型植被叶片的13C、生物量与生产力以及耗水，研究了六盘山主要森林植被水分利用效率（WUE）的部位、种间、生活型和水分生境等方面的差异及不同时间（年内、年际）的变化规律，同时在个体和林分尺度上分析了主要环境因子对WUE的影响。本研究结果将有助于系统认识森林植被WUE的特征和变化规律，在半干旱地区植被建设中的节水树种选择、树种合理配置和森林经营管理措施制定等方面具有一定的指导意义。本论文的主要结论如下：
1.主要树种叶片水平上的水分利用效率
华北落叶松、油松、华山松、辽东栎、少脉椴、白桦、沙棘和山桃8个六盘山主要树种的稳定性碳同位素技术研究表明：同一树种的叶片WUE存在冠层部位差异，但各树种表现各异。华北落叶松、白桦和油松表现为：上部＞中部＞下部；而华山松和辽东栎为：下部>上部>中部。香水河小流域各树种叶片的WUE分布范围为3.62～7.09mmol/mol，平均值为5.35（±0.66）mmol/mol；叠叠沟小流域各树种叶片WUE分布范围是10.42～13.14mmol/mol，平均为11.47（±0.88）mmol/mol；香水河小流域各树种叶片WUE的大小顺序为：油松＞山桃＞沙棘＞辽东栎＞华北落叶松＞华山松＞少脉椴＞白桦；不同生活型树木叶片WUE基本上呈现出灌木>小乔木>常绿针叶树>落叶针叶树>落叶阔叶树的变化规律。8个树种叶片WUE在生长季初期较高，而在生长季中、后期较低，但季节变化幅度随树种而异。不同水分生境下的华北落叶松、山桃和沙棘等三种树种叶片WUE都存在着明显的种内个体差异，但总体上呈现出半干旱区主要树种叶片WUE极显著地（P＜0.01）大于半湿润区的趋势。
2.主要树种个体水平上的水分利用效率——以华北落叶松为例
在生长季内，华北落叶松个体WUE与蒸腾量的变化表现一致，均为先略有升高后下降的变化规律，6月份最高。个体WUE在0.88~9.73g/kg之间，平均值为3.97g/kg，6月份的WUE最大。除空气湿度外，华北落叶松个体水平的WUE与太阳辐射、降水、温度、风速等气象因子都呈现出正相关关系，但与太阳辐射和潜在蒸发散达到显著性相关。
3.典型植被群落水平的水分利用效率
1）华北落叶松林的WUETr（生物量增量/蒸腾量）和WUEET（生物量增量/蒸散量）在初期最高，然后持续降低；降水利用效率（RUE）的变化规律是先略有升高后下降。华北落叶松林三种不同表达形式的WUE变化幅度存在差异。其中，WUETr在0.016~14.373g·m~(-2)·mm~(-1)变化，均值为5.72g·m~(-2)·mm~(-1)，整个生长季WUETr为6.933g·m~(-2)·mm~(-1)；WUEET在0.008~10.718g·m~(-2)·mm~(-1)之间，平均为3.84g·m~(-2)·mm~(-1)，整个生长季WUEET为4.035g·m~(-2)·mm~(-1)；RUE变化幅度是0.002~5.126g·m~(-2)·mm~(-1)，均值为1.81g·m~(-2)·mm~(-1)，整个生长季RUE为1.787g·m~(-2)·mm~(-1)。
2）华山松林RUE的年际变化表现出“快速上升-缓慢上升-缓慢下降”趋势，但年际之间的波动幅度存在差异；而华北落叶松林和油松林的RUE呈现出波动快速上升的趋势。华北落叶松林和油松林与华山松林的RUE存在极显著差异（P＜0.01），但华北落叶松林和油松林的差异不显著。多年平均RUE（g·m~(-2)·mm~(-1)）的大小依次为：华北落叶松林（1.12）＞油松林（0.97）＞华山松林（0.45）；华北落叶松人工林的RUE年际变异最大（0.54），华山松林次之（0.45），油松林最小（0.41）。
三种针叶林的RUE受降水量年内分配格局的影响。华山松天然林的RUE与上年8月降水量呈显著正相关，与当年9~11月降水量呈显著或极显著负相关；华北落叶松和油松林的RUE与上年9月降水量显著正相关，与当年4月降水量显著负相关，且华北落叶松林的RUE还与当年9月降水量显著正相关。华北落叶松和油松林的RUE与大多月份温度成正比，而华山松林的RUE与大多月份温度成反比。3、6月份温度显著或极显著的影响三种针叶林的RUE，6月温度还影响上年的RUE；华山松与2月月均温和月最高温度呈显著性负相关；华北落叶松林的RUE与4、5月份的温度呈显著正相关，而油松林的RUE与4月温度、5-8月份最低温极显著相关。华山松林的RUE与3月份的湿度极显著正相关，而华北落叶松和油松林的RUE与4月空气湿度显著负相关。
4.几点认识
1）随年降水量的增加，华山松林的RUE逐渐减小，而华北落叶松林和油松林的RUE均先略有升高后降低；在湿润年份，三种林分的RUE趋向于相同的最小值；而在干旱年份，三种林分也趋向于相同的RUE，但不一定是最大值。
2）树木WUE随着研究尺度的扩大呈递减的趋势，这可能与光合产物和水分的无效消耗增多有关。同时，不同尺度影响WUE的关键环境因子不同。因此，在实践生产中要综合分析不同尺度上WUE及其主导环境因子，可能才能做到适地适树的原则。
3）沙棘、山桃、油松和华北落叶松等4个人工主要造林树种在干旱条件和湿润条件下都具有较高的叶片WUE，是具有较高的抗旱能力的生态型节水树种，在干旱缺水地区可选为主要的造林树种。同时，不同的水分生境下，灌木和小乔木的WUE均大于乔木的，故干旱缺水区可适当减少乔木比例；树木的WUE具有保守性和变异性，可采取适度的抗旱锻炼等措施提高造林树种的WUE。
Based on the measurements of vegetation structure, stable carbon isotope, dendrometer,sap flow and tree ring width, we measured the leaf13C, biomass, productivity, transpirationand evapotranspiration of dominant tree species to calculate water use efficiency (WUE) atleaf-, tree-and stand scales in2010in the south (Xiangshuihe Watershed, semi-humid area)and north (Diediegou Watershed, semi-arid area) of Liupanshan Mountain, Ningxia. Thevariations in temporal and spatial heterogeneity of WUE were analyzed, we also analyzed howWUE responds to climate change at different scales. This study is helpful to systematicunderstand the variation characteristics of WUE, and to select tree species for the afforestaionand recovery of forest vegetation in the semi-arid and semi-humid area, and to offer scientificguidance for the harmonious and integrated management of forest and water. The mainconclusions were as follows:
1. Leaf level of water use efficiency
Leaf WUE of Larix principris-rupprechtii, Betula platyphylla, Pinus tablaeformisincreased with increasing canopy height, but the leaf WUE in the low canopy of Pinusarmandii and Quercus liaotungensis were higher compared with those of the mid-and upperparts in the canopy. The range of leaf WUE values was3.62～7.09mmol/mol, with the average5.35（±0.66）mmol/mol in Xiangshuihe Watershed, and10.42～13.14mmol/mol, with average11.47（±0.88）mmol/mol in Diediegou Watershed. The order of mean WUE of each treespecies is as P. tablaeformis＞Prunus davidiana＞Hippophae rhamnoides＞Q. liaotungersis＞L. principis-rupprechtii＞P. armandii＞Tilia panciostata＞B. Platyphylla. The leaf WUEfor life forms were significantly different, with a sequence of shrubs＞evergreen trees＞leavedtrees.The leaf WUE of species flucutated with the time during the growing season, beingrelatively higher WUE in the early stage than that in the middle and late phases. The leaf WUEamong individual trees of L. principris-rupprechtii, P. davidiana and H. rhamnoides in twodifferent water-environmental regions were significantly different during the growing season,IV and semi-arid area with the higher values. It showed that the ability of water use efficiency ofthe same tree species could be improved through long-term adaptation to drought.
2. Individual level of water use efficiency
The seasonal pattern of WUE value of individual tree of L. principis-rupprechtii exhibitedan initial increase and a subsequent decrease, the value ranged from0.88to9.73g/kg duringgrowing season in2010, having the maximum in June. The growing season WUE is4.85g/kg.Individual level of water use efficiency of L. principis-rupprechtii was mainly affected by solarradiation and the potential evaporation. WUE exhibited positive associations with precipitation,temperature and wind speed, except air humidity.
3. Estimation of water use efficiency at stand level
1) The WUETr(Productivity/Transpiration) and WUEET(Productivity/Evapotranspiration)of L.principis-rupprechtii plantation had higher values in the early stage of growing seasonthan that in the middle and late phases. The RUE(Productivity/Precipitation) values ofL.principis-rupprechtii plantation exhibited an initial increase and a subsequent decrease. TheWUETrvalues ranged from0.016to14.373g·m~(-2)·mm~(-1), and the WUEETvalues ranged from0.008to10.718g·m~(-2)·mm~(-1), and the RUE values ranged from0.002to5.126g·m~(-2)·mm~(-1).
2) The interannual changes of RUE of P.armandii natural forest exhibited “fast increase-slow increase-slow decrease”, but variation amplitude was different. The interannual changesof RUE of L.principis-rupprechtii and P.tablaeformis plantations showed rapid rising trendwith fluctuations, and the greater fluctuations in L.principis-rupprechtii plantation. The RUEof two artificial forests and natural forest had extremely remarkable difference, with the orderof L.principis-rupprechtii plantation(1.12t· hm~(-2)·a~(-1))＞P.tablaeformis plantation(0.97t·hm~(-2)·a~(-1))＞P.armandii Natural Forest(0.45t·hm~(-2)·a~(-1)). The coefficient of variation in RUEwas as L.principis-rupprechtii plantation(0.54)＞P.armandii natural forest(0.45)＞P.tablaeformis plantation(0.41).
The RUE of P.armandii natural forest was significantly positively correlated withprecipitation in August of the previous year and that from September to November of thecurrent year, but the RUE of two artificial forests was significantly positively correlated with precipitation in September of the previous year and negatively correlated with precipitation inApril of the current year, and the productivity of L.principis-rupprechtii plantation was alsosignificantly positively correlated with precipitation in September of the current year. So theRUE of the three forests was affected by the distribution pattern of precipitation during theyear.Most of monthly temperature negatively affected the RUE of P.armandii natural forest,but positively affected the RUE of two artificial forests, and the RUE values of three forestswas significantly correlated with temperature in June of the previous year and that in Marchand June of the current year. There were significantly negative correlations between RUE ofP.armandii Natural Forest and mean and minimum temperature in February, and the RUE ofL.principis-rupprechtii plantation was also significantly correlated with mean-and minimum-temperature in April and May of the current year, but P.tablaeformis plantation wassignificantly correlated with mean temperature in April and minimum temperature from May toAugust of the current year.There were significantly correlations between RUE of P.armandiinatural forest and mean humidity in March, and that between productivity of artificial forestand mean humidity in April.
4. Some innovative viewpoint in the WUE research
1) we showed that RUE of P.armandii forest decreases as mean annual precipitationincreases, but RUE of L.principis-rupprechtii and P.tablaeformis plantation increased at firstand then decreased. During the driest years, there was convergence to common RUE that wastypical of three coniferous tree plantions, but in most moisture years, all forests exhibited thesame minimum rate of biomass production per unit rainfall.
2) With the expansion of the research scale, the WUE of trees showed a decreasing trend,and was affected by different environmental factors. In forest management, we shouldcomprehensive analysis the WUE and its impact factors at different scales to select suitabletree species in the afforestaion and recovery of forest vegetation.
3) The L. principis-rupprechtii, P. tablaeformis, P. davidiana and H. rhamnoides that hadhigher WUE values whether they were in dry condition or wet conditions, were ecologicalwater-saving species, having high drought tolerance, so they can be chosen as the main afforestation tree for vegetation restoration and construction in arid and semi-arid regions.Secondly, the WUE values of shrubs and small trees were more than trees in different waterconditions, so we could rationally configure the proportion of different tree species. The WUEof trees had the characteristics of conservative and variability, so moderate drought exercisemeasures could be taken to improve the WUE of tree species in afforestations.
1.主要树种叶片水平上的水分利用效率
华北落叶松、油松、华山松、辽东栎、少脉椴、白桦、沙棘和山桃8个六盘山主要树种的稳定性碳同位素技术研究表明：同一树种的叶片WUE存在冠层部位差异，但各树种表现各异。华北落叶松、白桦和油松表现为：上部＞中部＞下部；而华山松和辽东栎为：下部>上部>中部。香水河小流域各树种叶片的WUE分布范围为3.62～7.09mmol/mol，平均值为5.35（±0.66）mmol/mol；叠叠沟小流域各树种叶片WUE分布范围是10.42～13.14mmol/mol，平均为11.47（±0.88）mmol/mol；香水河小流域各树种叶片WUE的大小顺序为：油松＞山桃＞沙棘＞辽东栎＞华北落叶松＞华山松＞少脉椴＞白桦；不同生活型树木叶片WUE基本上呈现出灌木>小乔木>常绿针叶树>落叶针叶树>落叶阔叶树的变化规律。8个树种叶片WUE在生长季初期较高，而在生长季中、后期较低，但季节变化幅度随树种而异。不同水分生境下的华北落叶松、山桃和沙棘等三种树种叶片WUE都存在着明显的种内个体差异，但总体上呈现出半干旱区主要树种叶片WUE极显著地（P＜0.01）大于半湿润区的趋势。
2.主要树种个体水平上的水分利用效率——以华北落叶松为例
在生长季内，华北落叶松个体WUE与蒸腾量的变化表现一致，均为先略有升高后下降的变化规律，6月份最高。个体WUE在0.88~9.73g/kg之间，平均值为3.97g/kg，6月份的WUE最大。除空气湿度外，华北落叶松个体水平的WUE与太阳辐射、降水、温度、风速等气象因子都呈现出正相关关系，但与太阳辐射和潜在蒸发散达到显著性相关。
3.典型植被群落水平的水分利用效率
1）华北落叶松林的WUETr（生物量增量/蒸腾量）和WUEET（生物量增量/蒸散量）在初期最高，然后持续降低；降水利用效率（RUE）的变化规律是先略有升高后下降。华北落叶松林三种不同表达形式的WUE变化幅度存在差异。其中，WUETr在0.016~14.373g·m~(-2)·mm~(-1)变化，均值为5.72g·m~(-2)·mm~(-1)，整个生长季WUETr为6.933g·m~(-2)·mm~(-1)；WUEET在0.008~10.718g·m~(-2)·mm~(-1)之间，平均为3.84g·m~(-2)·mm~(-1)，整个生长季WUEET为4.035g·m~(-2)·mm~(-1)；RUE变化幅度是0.002~5.126g·m~(-2)·mm~(-1)，均值为1.81g·m~(-2)·mm~(-1)，整个生长季RUE为1.787g·m~(-2)·mm~(-1)。
2）华山松林RUE的年际变化表现出“快速上升-缓慢上升-缓慢下降”趋势，但年际之间的波动幅度存在差异；而华北落叶松林和油松林的RUE呈现出波动快速上升的趋势。华北落叶松林和油松林与华山松林的RUE存在极显著差异（P＜0.01），但华北落叶松林和油松林的差异不显著。多年平均RUE（g·m~(-2)·mm~(-1)）的大小依次为：华北落叶松林（1.12）＞油松林（0.97）＞华山松林（0.45）；华北落叶松人工林的RUE年际变异最大（0.54），华山松林次之（0.45），油松林最小（0.41）。
三种针叶林的RUE受降水量年内分配格局的影响。华山松天然林的RUE与上年8月降水量呈显著正相关，与当年9~11月降水量呈显著或极显著负相关；华北落叶松和油松林的RUE与上年9月降水量显著正相关，与当年4月降水量显著负相关，且华北落叶松林的RUE还与当年9月降水量显著正相关。华北落叶松和油松林的RUE与大多月份温度成正比，而华山松林的RUE与大多月份温度成反比。3、6月份温度显著或极显著的影响三种针叶林的RUE，6月温度还影响上年的RUE；华山松与2月月均温和月最高温度呈显著性负相关；华北落叶松林的RUE与4、5月份的温度呈显著正相关，而油松林的RUE与4月温度、5-8月份最低温极显著相关。华山松林的RUE与3月份的湿度极显著正相关，而华北落叶松和油松林的RUE与4月空气湿度显著负相关。
4.几点认识
1）随年降水量的增加，华山松林的RUE逐渐减小，而华北落叶松林和油松林的RUE均先略有升高后降低；在湿润年份，三种林分的RUE趋向于相同的最小值；而在干旱年份，三种林分也趋向于相同的RUE，但不一定是最大值。
2）树木WUE随着研究尺度的扩大呈递减的趋势，这可能与光合产物和水分的无效消耗增多有关。同时，不同尺度影响WUE的关键环境因子不同。因此，在实践生产中要综合分析不同尺度上WUE及其主导环境因子，可能才能做到适地适树的原则。
3）沙棘、山桃、油松和华北落叶松等4个人工主要造林树种在干旱条件和湿润条件下都具有较高的叶片WUE，是具有较高的抗旱能力的生态型节水树种，在干旱缺水地区可选为主要的造林树种。同时，不同的水分生境下，灌木和小乔木的WUE均大于乔木的，故干旱缺水区可适当减少乔木比例；树木的WUE具有保守性和变异性，可采取适度的抗旱锻炼等措施提高造林树种的WUE。
Based on the measurements of vegetation structure, stable carbon isotope, dendrometer,sap flow and tree ring width, we measured the leaf13C, biomass, productivity, transpirationand evapotranspiration of dominant tree species to calculate water use efficiency (WUE) atleaf-, tree-and stand scales in2010in the south (Xiangshuihe Watershed, semi-humid area)and north (Diediegou Watershed, semi-arid area) of Liupanshan Mountain, Ningxia. Thevariations in temporal and spatial heterogeneity of WUE were analyzed, we also analyzed howWUE responds to climate change at different scales. This study is helpful to systematicunderstand the variation characteristics of WUE, and to select tree species for the afforestaionand recovery of forest vegetation in the semi-arid and semi-humid area, and to offer scientificguidance for the harmonious and integrated management of forest and water. The mainconclusions were as follows:
1. Leaf level of water use efficiency
Leaf WUE of Larix principris-rupprechtii, Betula platyphylla, Pinus tablaeformisincreased with increasing canopy height, but the leaf WUE in the low canopy of Pinusarmandii and Quercus liaotungensis were higher compared with those of the mid-and upperparts in the canopy. The range of leaf WUE values was3.62～7.09mmol/mol, with the average5.35（±0.66）mmol/mol in Xiangshuihe Watershed, and10.42～13.14mmol/mol, with average11.47（±0.88）mmol/mol in Diediegou Watershed. The order of mean WUE of each treespecies is as P. tablaeformis＞Prunus davidiana＞Hippophae rhamnoides＞Q. liaotungersis＞L. principis-rupprechtii＞P. armandii＞Tilia panciostata＞B. Platyphylla. The leaf WUEfor life forms were significantly different, with a sequence of shrubs＞evergreen trees＞leavedtrees.The leaf WUE of species flucutated with the time during the growing season, beingrelatively higher WUE in the early stage than that in the middle and late phases. The leaf WUEamong individual trees of L. principris-rupprechtii, P. davidiana and H. rhamnoides in twodifferent water-environmental regions were significantly different during the growing season,IV and semi-arid area with the higher values. It showed that the ability of water use efficiency ofthe same tree species could be improved through long-term adaptation to drought.
2. Individual level of water use efficiency
The seasonal pattern of WUE value of individual tree of L. principis-rupprechtii exhibitedan initial increase and a subsequent decrease, the value ranged from0.88to9.73g/kg duringgrowing season in2010, having the maximum in June. The growing season WUE is4.85g/kg.Individual level of water use efficiency of L. principis-rupprechtii was mainly affected by solarradiation and the potential evaporation. WUE exhibited positive associations with precipitation,temperature and wind speed, except air humidity.
3. Estimation of water use efficiency at stand level
1) The WUETr(Productivity/Transpiration) and WUEET(Productivity/Evapotranspiration)of L.principis-rupprechtii plantation had higher values in the early stage of growing seasonthan that in the middle and late phases. The RUE(Productivity/Precipitation) values ofL.principis-rupprechtii plantation exhibited an initial increase and a subsequent decrease. TheWUETrvalues ranged from0.016to14.373g·m~(-2)·mm~(-1), and the WUEETvalues ranged from0.008to10.718g·m~(-2)·mm~(-1), and the RUE values ranged from0.002to5.126g·m~(-2)·mm~(-1).
2) The interannual changes of RUE of P.armandii natural forest exhibited “fast increase-slow increase-slow decrease”, but variation amplitude was different. The interannual changesof RUE of L.principis-rupprechtii and P.tablaeformis plantations showed rapid rising trendwith fluctuations, and the greater fluctuations in L.principis-rupprechtii plantation. The RUEof two artificial forests and natural forest had extremely remarkable difference, with the orderof L.principis-rupprechtii plantation(1.12t· hm~(-2)·a~(-1))＞P.tablaeformis plantation(0.97t·hm~(-2)·a~(-1))＞P.armandii Natural Forest(0.45t·hm~(-2)·a~(-1)). The coefficient of variation in RUEwas as L.principis-rupprechtii plantation(0.54)＞P.armandii natural forest(0.45)＞P.tablaeformis plantation(0.41).
The RUE of P.armandii natural forest was significantly positively correlated withprecipitation in August of the previous year and that from September to November of thecurrent year, but the RUE of two artificial forests was significantly positively correlated with precipitation in September of the previous year and negatively correlated with precipitation inApril of the current year, and the productivity of L.principis-rupprechtii plantation was alsosignificantly positively correlated with precipitation in September of the current year. So theRUE of the three forests was affected by the distribution pattern of precipitation during theyear.Most of monthly temperature negatively affected the RUE of P.armandii natural forest,but positively affected the RUE of two artificial forests, and the RUE values of three forestswas significantly correlated with temperature in June of the previous year and that in Marchand June of the current year. There were significantly negative correlations between RUE ofP.armandii Natural Forest and mean and minimum temperature in February, and the RUE ofL.principis-rupprechtii plantation was also significantly correlated with mean-and minimum-temperature in April and May of the current year, but P.tablaeformis plantation wassignificantly correlated with mean temperature in April and minimum temperature from May toAugust of the current year.There were significantly correlations between RUE of P.armandiinatural forest and mean humidity in March, and that between productivity of artificial forestand mean humidity in April.
4. Some innovative viewpoint in the WUE research
1) we showed that RUE of P.armandii forest decreases as mean annual precipitationincreases, but RUE of L.principis-rupprechtii and P.tablaeformis plantation increased at firstand then decreased. During the driest years, there was convergence to common RUE that wastypical of three coniferous tree plantions, but in most moisture years, all forests exhibited thesame minimum rate of biomass production per unit rainfall.
2) With the expansion of the research scale, the WUE of trees showed a decreasing trend,and was affected by different environmental factors. In forest management, we shouldcomprehensive analysis the WUE and its impact factors at different scales to select suitabletree species in the afforestaion and recovery of forest vegetation.
3) The L. principis-rupprechtii, P. tablaeformis, P. davidiana and H. rhamnoides that hadhigher WUE values whether they were in dry condition or wet conditions, were ecologicalwater-saving species, having high drought tolerance, so they can be chosen as the main afforestation tree for vegetation restoration and construction in arid and semi-arid regions.Secondly, the WUE values of shrubs and small trees were more than trees in different waterconditions, so we could rationally configure the proportion of different tree species. The WUEof trees had the characteristics of conservative and variability, so moderate drought exercisemeasures could be taken to improve the WUE of tree species in afforestations.
引文
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