中国东北主要造林树种细根寿命及影响因子研究
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摘要
树木细根(直径<2 mm)的生产和死亡过程(即周转),对森林生态系统碳(C)分配格局和养分循环过程具有重要影响。细根的寿命(从出生至死亡的时间),是准确估计细根周转的重要参数。然而,由于概念与研究方法的限制,导致人们对细根寿命与周转过程及影响因素,还缺乏全面的了解。中国东北部的温带森林,分别占全国森林总面积和蓄积量的35.3%和34.5%,在区域和全球尺度的C平衡中发挥着重要的作用。因此,在该地区开展树木细根周转及影响因素的深入研究,对于准确估计区域水平上的森林C储存与释放,具有重要的意义。为此,本研究以中国东北5个典型造林树种的人工同龄纯林(1986年造林),即胡桃楸(Juglans mandshurica)、黄波罗(Phellodendron amurense)、红松(Pinus koraiensis)、樟子松(Pinus sylvestris var. mongolica)、和红皮云杉(Picea koraiensis)为研究对象,在2年的时间里(2008-2009年),采用微根管(Minirhizotron)技术定量研究了细根的生产和死亡过程,估计了细根的寿命,同期测定了相关环境因子与细根的结构与生理特征。主要目的是：(1)揭示5树种人工林细根的生产和死亡的季节格局,确定哪些环境因子对中国东北温带森林细根周转过程具有重要的影响；(2)估计和比较不同树种细根寿命的差异,探索细根直径、根序(个体根在根系分支中的位置)、细根出生的季节和土壤深度对细根寿命的影响和潜在的机制；(3)结合同期水曲柳(Fraxinus mandshurica)和兴安落叶松(Larix gmelinii)的研究结果,选取生理功能一致、具有种间可比性的1级根为研究单元,探索细根寿命与其他根系结构、生理特征在树种之间的关联性。研究结果表明：
1、细根总根长的生产量在种间存在明显差异。5树种细根生产均表现为春、秋季低,夏季高的季节格局。云杉年根长生产量最大,2年平均生产量为1.41 mm cm-2a-1,而黄波罗生产量最低(0.23 mm cm-2 a-1),其余树种根长生产的大小顺序为红松<胡桃楸<樟子松。细根的死亡与生产过程是同时发生的。但与生产不同,总根长的死亡量高峰存在树种及年际间的差异。不同直径大小(0-0.5 mm,0.5-1.0 mm,>1 mm)细根类群的生产过程与总根长的季节的动态类似,但是细根的死亡动态随直径大小出现差异。回归分析显示,细根总根长的生产量与大气温度、土壤温度(10 cm深度)和降雨量存在不同程度的相关性,大气温度对根长生产量的影响最强,而降雨量的影响最弱。在不同直径细根类群中,以直径0-0.5 mm根群同环境变量的关系最密切,表现出与环境变量之间的普遍相关(出现相关性的频次高),以及较高的回归方程决定系数。但是,总根长及不同直径根长的死亡量与环境变量的联系均很弱。
2、不同树种细根寿命存在着显著的差异。其中红松寿命最长(平均寿命=434天),而黄波罗寿命最短(平均寿命=279天),但常绿针叶树种(红松、樟子松和云杉)与落叶阔叶树种(胡桃楸和黄波罗)之间并没有恒定的区别。多变量回归(Cox比例风险回归)分析显示,不同生活型树木细根的寿命也没有明显的区别。存活分析显示,随着直径增加、根序增高、出生土壤深度增大,细根寿命普遍有延长的趋势,而且夏、秋季出生的细根倾向于比春季细根寿命长(Log-rank test方法)。但在控制其他因素影响条件下,Cox比例风险回归分析表明,仅针叶树细根寿命随着直径增加而显著地延长；根序对树种细根寿命增加的作用仅在云杉中是显著的；而出生季节和土壤深度的影响相对普遍,5个树种中,有3个树种的夏、秋季节出生细根比春季具有更长的寿命,有4个树种细根寿命随着土壤深度增加而延长。
3、种间1级根寿命、形态、组织化学和生理特征分析显示,常绿针叶树种(红松、樟子松和云杉)并没有比落叶阔叶(水曲柳、胡桃楸和黄波罗)或落叶针叶树种(落叶松)的根系寿命长。所有针叶树种均具有较高的组织密度,低的N含量、N：C比和维持呼吸速率。各根系特征在树种之间的变异程度不同,单根长度的变异系数最大(CV=0.701),而N：C比则具有最小的变异(CV=0.156)。种间1级根寿命与其他形态、组织化学和生理特征之间均不存在显著的相关性。但是,比根长(1 g根干生物量的长度)与分支比之间,N含量与N：C比和维持呼吸速率三者之间,均显示出显著的正相关根系(p<0.05)；而组织密度与N含量、N：C比或维持呼吸速率之间,直径与比根长、分支比之间,均表现出显著的负相关关系。
总之,本研究证实,温带地区树木细根生产与死亡过程具有明显的季节动态。相比较,细根的生产受到环境因素,特别是大气温度的强烈影响,而死亡受环境因素影响较弱。直径细小、以吸收功能为主的细根组分对环境变化最为敏感。树种水平上,细根的寿命受内在与外在因素的共同影响。各树种低级根(前2或3级)之间寿命差异较小暗示了这些细根个体倾向于作为一个整体同时死亡,因此具有模块(module)特征。季节和土壤深度对细根寿命具有普遍影响,但是细根结构及个体的发育与寿命的联系仍然值得深入探索。种间细根寿命的变异具有一定的独立性。根系不同结构特征之间的紧密相关,则显示了树木根系在养分获取与器官维持之间的权衡,以及树种间资源利用策略的差异。这些研究结果,将为了解树种水平上的细根动态机制,准确估计我国东北地区森林地下C分配和养分循环过程提供必要的参考。
Fine root (diameter<2 mm) production and mortality are defined as turnover. Fine root turnover plays a key role in regulating carbon (C) allocation and nutrients cycles in forest ecosystem. Fine root longevity is the important parameter for the precise estimate of fine root turnover. However, conceptual and methodological constraints may have limited our understanding of fine root longevity. The temperate forest in northeastern China accounts for 35.3% and 34.5% of national total area and stocking volume, respectively, thus play a key role in C budgets at regional level and global scale. However, to date, little information about fine root dynamics and longevity of Chinese temperate tree species has been reported. In this study, the minirhizotron approach was used in five temperate tree species in northeastern China to determine the dynamics of fine root production and mortality, and to estimate the root longevity during two growth seasons from 2008 and 2009. Climate factors including air temperature, soil temperature at 10 cm depth and precipitation, and some root structural and physiological traits were measured concurrently. The five species include two deciduous hardwoods Juglans mandshurica and Phellodendron amurense; and three evergreen conifers Pinus koraiensis, Pinus sylvestris var. mongolica, and Picea koraiensis. The aims of this study were:(1) to quantify the seasonal patterns of fine root production and mortality in five tree species, and determine what climate factors may have strong effects on root dynamics; (2) to estimate and compare root longevity across species, and determine how fine root diameter, branch order, season and soil depth of root birth would influence the longevity; (3) to investigate the potential correlation between longevity of the first order root and root structural or physiological traits, as well as the potential correlation between root structural or physiological traits. Our results show that:
(1) Total root length production differed markedly among species. The seasonal patterns of root production were similar across tree species, with the peaks consistently occurring in summer in two years. Picea koraiensis had the largest production on the annual basis (1.41 mm cm-2 a-1), while P. amurense was lowest (0.23 mm cm-2 a-1), leaving Pinus koraiensis, J. mandshurica and P. sylvestris var. mongolica in the mediate. Fine root mortality occurred simultaneously with production, but which peaks differed among species and years. Three diameter classes root population, i.e.0-0.5 mm,0.5-1.0 mm and>1 mm, showed similar seasonal pattern in production with total roots, but their mortalities are different between each other. Regression analysis showed that there were positive correlations between total root length production and air temperature, soil temperature at 10 cm depth and rainfall, in which air temperature had the strongest but precipitation weakest impacts. Roots with 0-0.5 mm diameter size showed the most sensitivity to environmental variables, exhibiting the consistant and close correlation with environmental variables. However, there were weak correlations between length mortality of total roots or differed diameter size roots and climate factors.
(2) During two growth seasons from 2008 to 2009, median root longevity differed significantly among species, ranging from 276 d (P. amurense) to>430 d(Pinus koraiensis). However, conifers did not consistently have longer root longevity than hardwoods, and Cox proportional hazards regression analysis showed the similar results when controlling for other covariates (factors). Survival analysis (Log-rank test) showed that the longevity tend to be larger in roots with greater diameter, and higher branch order or born in deeper soil depth across species. In comparison, roots born in summer or autumn, more often had longer longevity than those born in spring. However, when controlling for other covariates, Cox analysis showed that only coniferous root longevity significantly increased with diameter increasing, and just root longevity in Picea koraiensis shifted with branch order. In contrast, influences of season and soil depth of root birth were more general. Three out of 5 species displayed significant longer root longevity in summer or autumn, and root longevities in 4 out of 5 species significantly increased with soil depth increasing. (3) Incorporating previous results from Fraxinus mandshurica and Larix gmelinii, the present study showed that even the first order roots, the evergreen confers did not have longer longevity than deciduous hardwoods or conifers. In comparison with three hardwood species, four coniferous species all had higher tissue density, lower nitrogen (N) content, N:C ratio, and maintaining root respiration rate. There were differed extents of the variation of root traits, with the largest CV (coefficient of variation,0.701) in root length and the lowest in N:C ratio (CV=0.156). We did not find the significant correlation between root longevity and any other root traits. In comparison, there were significant positive correlations between specific root length (SRL) and branching ratio (BR), so did to N, and N:C ratio or maintaining root respiration rate. However, there were significant negative correlation between tissue density and N content, N:C ratio or root respiration rate, and between diameter and SRL or BR.
In sum, this study reveals the strong seasonality of fine root production and mortality in temperate tree species. In comparison with root length mortality, production of root length is strongly affected by climate factors, especially by air temperature. The thinnest root population (diameter in 0-0.5 mm) primarily with uptakes function, were more sensitive to environmental change. At species level, tree root longevity is controlled by both endogenous and exogenous factors. The lower distal order roots may incline to die as an intact braches with similar and short longevity, indicating these roots may constitute the mainbody of ephemeral portion within tree root branching systems. Season and soil depth of root birth had general impacts on root survivorship across species, but the relationship between the ontogeny of individual roots and longevity still deserved further study. The longevity of the first order root tended to vary independently among tree species. However, the close correlations between root structural traits demonstrated the general trade-off between resource acquisition and conservation in tree root, as well as the difference in resource acquisition in roots among tree species. These findings may improve our understanding on the mechanism of fine root dynamics at species level, and estimation of the belowground C and nutrients cycles in forest ecosystems in northeastern China.
1、细根总根长的生产量在种间存在明显差异。5树种细根生产均表现为春、秋季低,夏季高的季节格局。云杉年根长生产量最大,2年平均生产量为1.41 mm cm-2a-1,而黄波罗生产量最低(0.23 mm cm-2 a-1),其余树种根长生产的大小顺序为红松<胡桃楸<樟子松。细根的死亡与生产过程是同时发生的。但与生产不同,总根长的死亡量高峰存在树种及年际间的差异。不同直径大小(0-0.5 mm,0.5-1.0 mm,>1 mm)细根类群的生产过程与总根长的季节的动态类似,但是细根的死亡动态随直径大小出现差异。回归分析显示,细根总根长的生产量与大气温度、土壤温度(10 cm深度)和降雨量存在不同程度的相关性,大气温度对根长生产量的影响最强,而降雨量的影响最弱。在不同直径细根类群中,以直径0-0.5 mm根群同环境变量的关系最密切,表现出与环境变量之间的普遍相关(出现相关性的频次高),以及较高的回归方程决定系数。但是,总根长及不同直径根长的死亡量与环境变量的联系均很弱。
2、不同树种细根寿命存在着显著的差异。其中红松寿命最长(平均寿命=434天),而黄波罗寿命最短(平均寿命=279天),但常绿针叶树种(红松、樟子松和云杉)与落叶阔叶树种(胡桃楸和黄波罗)之间并没有恒定的区别。多变量回归(Cox比例风险回归)分析显示,不同生活型树木细根的寿命也没有明显的区别。存活分析显示,随着直径增加、根序增高、出生土壤深度增大,细根寿命普遍有延长的趋势,而且夏、秋季出生的细根倾向于比春季细根寿命长(Log-rank test方法)。但在控制其他因素影响条件下,Cox比例风险回归分析表明,仅针叶树细根寿命随着直径增加而显著地延长；根序对树种细根寿命增加的作用仅在云杉中是显著的；而出生季节和土壤深度的影响相对普遍,5个树种中,有3个树种的夏、秋季节出生细根比春季具有更长的寿命,有4个树种细根寿命随着土壤深度增加而延长。
3、种间1级根寿命、形态、组织化学和生理特征分析显示,常绿针叶树种(红松、樟子松和云杉)并没有比落叶阔叶(水曲柳、胡桃楸和黄波罗)或落叶针叶树种(落叶松)的根系寿命长。所有针叶树种均具有较高的组织密度,低的N含量、N：C比和维持呼吸速率。各根系特征在树种之间的变异程度不同,单根长度的变异系数最大(CV=0.701),而N：C比则具有最小的变异(CV=0.156)。种间1级根寿命与其他形态、组织化学和生理特征之间均不存在显著的相关性。但是,比根长(1 g根干生物量的长度)与分支比之间,N含量与N：C比和维持呼吸速率三者之间,均显示出显著的正相关根系(p<0.05)；而组织密度与N含量、N：C比或维持呼吸速率之间,直径与比根长、分支比之间,均表现出显著的负相关关系。
总之,本研究证实,温带地区树木细根生产与死亡过程具有明显的季节动态。相比较,细根的生产受到环境因素,特别是大气温度的强烈影响,而死亡受环境因素影响较弱。直径细小、以吸收功能为主的细根组分对环境变化最为敏感。树种水平上,细根的寿命受内在与外在因素的共同影响。各树种低级根(前2或3级)之间寿命差异较小暗示了这些细根个体倾向于作为一个整体同时死亡,因此具有模块(module)特征。季节和土壤深度对细根寿命具有普遍影响,但是细根结构及个体的发育与寿命的联系仍然值得深入探索。种间细根寿命的变异具有一定的独立性。根系不同结构特征之间的紧密相关,则显示了树木根系在养分获取与器官维持之间的权衡,以及树种间资源利用策略的差异。这些研究结果,将为了解树种水平上的细根动态机制,准确估计我国东北地区森林地下C分配和养分循环过程提供必要的参考。
Fine root (diameter<2 mm) production and mortality are defined as turnover. Fine root turnover plays a key role in regulating carbon (C) allocation and nutrients cycles in forest ecosystem. Fine root longevity is the important parameter for the precise estimate of fine root turnover. However, conceptual and methodological constraints may have limited our understanding of fine root longevity. The temperate forest in northeastern China accounts for 35.3% and 34.5% of national total area and stocking volume, respectively, thus play a key role in C budgets at regional level and global scale. However, to date, little information about fine root dynamics and longevity of Chinese temperate tree species has been reported. In this study, the minirhizotron approach was used in five temperate tree species in northeastern China to determine the dynamics of fine root production and mortality, and to estimate the root longevity during two growth seasons from 2008 and 2009. Climate factors including air temperature, soil temperature at 10 cm depth and precipitation, and some root structural and physiological traits were measured concurrently. The five species include two deciduous hardwoods Juglans mandshurica and Phellodendron amurense; and three evergreen conifers Pinus koraiensis, Pinus sylvestris var. mongolica, and Picea koraiensis. The aims of this study were:(1) to quantify the seasonal patterns of fine root production and mortality in five tree species, and determine what climate factors may have strong effects on root dynamics; (2) to estimate and compare root longevity across species, and determine how fine root diameter, branch order, season and soil depth of root birth would influence the longevity; (3) to investigate the potential correlation between longevity of the first order root and root structural or physiological traits, as well as the potential correlation between root structural or physiological traits. Our results show that:
(1) Total root length production differed markedly among species. The seasonal patterns of root production were similar across tree species, with the peaks consistently occurring in summer in two years. Picea koraiensis had the largest production on the annual basis (1.41 mm cm-2 a-1), while P. amurense was lowest (0.23 mm cm-2 a-1), leaving Pinus koraiensis, J. mandshurica and P. sylvestris var. mongolica in the mediate. Fine root mortality occurred simultaneously with production, but which peaks differed among species and years. Three diameter classes root population, i.e.0-0.5 mm,0.5-1.0 mm and>1 mm, showed similar seasonal pattern in production with total roots, but their mortalities are different between each other. Regression analysis showed that there were positive correlations between total root length production and air temperature, soil temperature at 10 cm depth and rainfall, in which air temperature had the strongest but precipitation weakest impacts. Roots with 0-0.5 mm diameter size showed the most sensitivity to environmental variables, exhibiting the consistant and close correlation with environmental variables. However, there were weak correlations between length mortality of total roots or differed diameter size roots and climate factors.
(2) During two growth seasons from 2008 to 2009, median root longevity differed significantly among species, ranging from 276 d (P. amurense) to>430 d(Pinus koraiensis). However, conifers did not consistently have longer root longevity than hardwoods, and Cox proportional hazards regression analysis showed the similar results when controlling for other covariates (factors). Survival analysis (Log-rank test) showed that the longevity tend to be larger in roots with greater diameter, and higher branch order or born in deeper soil depth across species. In comparison, roots born in summer or autumn, more often had longer longevity than those born in spring. However, when controlling for other covariates, Cox analysis showed that only coniferous root longevity significantly increased with diameter increasing, and just root longevity in Picea koraiensis shifted with branch order. In contrast, influences of season and soil depth of root birth were more general. Three out of 5 species displayed significant longer root longevity in summer or autumn, and root longevities in 4 out of 5 species significantly increased with soil depth increasing. (3) Incorporating previous results from Fraxinus mandshurica and Larix gmelinii, the present study showed that even the first order roots, the evergreen confers did not have longer longevity than deciduous hardwoods or conifers. In comparison with three hardwood species, four coniferous species all had higher tissue density, lower nitrogen (N) content, N:C ratio, and maintaining root respiration rate. There were differed extents of the variation of root traits, with the largest CV (coefficient of variation,0.701) in root length and the lowest in N:C ratio (CV=0.156). We did not find the significant correlation between root longevity and any other root traits. In comparison, there were significant positive correlations between specific root length (SRL) and branching ratio (BR), so did to N, and N:C ratio or maintaining root respiration rate. However, there were significant negative correlation between tissue density and N content, N:C ratio or root respiration rate, and between diameter and SRL or BR.
In sum, this study reveals the strong seasonality of fine root production and mortality in temperate tree species. In comparison with root length mortality, production of root length is strongly affected by climate factors, especially by air temperature. The thinnest root population (diameter in 0-0.5 mm) primarily with uptakes function, were more sensitive to environmental change. At species level, tree root longevity is controlled by both endogenous and exogenous factors. The lower distal order roots may incline to die as an intact braches with similar and short longevity, indicating these roots may constitute the mainbody of ephemeral portion within tree root branching systems. Season and soil depth of root birth had general impacts on root survivorship across species, but the relationship between the ontogeny of individual roots and longevity still deserved further study. The longevity of the first order root tended to vary independently among tree species. However, the close correlations between root structural traits demonstrated the general trade-off between resource acquisition and conservation in tree root, as well as the difference in resource acquisition in roots among tree species. These findings may improve our understanding on the mechanism of fine root dynamics at species level, and estimation of the belowground C and nutrients cycles in forest ecosystems in northeastern China.
引文
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