北京林业大学森林资源与生态系统过程北京市重点实验室, 北京林业大学林学院, 北京 100083
Changes in leaf nitrogen and phosphorus stoichiometry of woody plants along an altitudinal gradient in Changbai Mountain, China
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氮(N)和磷(P)是植物生长所必需的营养元素, 对植物个体的生长发育起着关键性的作用。由于N和P的养分获得和利用效率往往受到自然环境的限制, N、P成为陆地生态系统中的限制营养元素(Koerselman & Meuleman, 1996; Güsewell, 2004)。叶片是植物进行光合作用的主要器官, 也是植物N、P周转利用的活跃部位, 研究叶片N、P含量和N:P在地理和环境梯度上的变异规律有着重要意义。
海拔梯度上气候、土壤以及植被物种组成急剧变化, 能够综合反映气温、降水、土壤、植被等环境因子的作用, 使海拔梯度成为研究叶片N、P化学计量特征的理想场所。目前关于叶片N、P含量在海拔梯度上的空间格局尚未得出一致的结论。有的研究表明乔木、灌木、草本三种生活型的植物叶片N含量均随海拔的上升而增加, 并认为高海拔地区的低N供应不影响成熟叶片的代谢活性(K?rner, 1989)。而在热带山地雨林生态系统的研究中发现, 由于高海拔地区的低温和低N矿化率, 单位质量的叶片N含量随海拔的上升显著下降; P含量、N:P则没有表现出明显的垂直变异(van de Weg et al., 2009)。有研究发现在同样的生态系统中叶片N:P随海拔的上升而下降, 认为高海拔地区植物生长受到N限制(Fisher et al., 2013); 有的研究则发现叶片N、P含量随海拔的升高先增加后降低(Shi et al., 2012; Fisher et al., 2013)。这些研究之间的差异表明, 关于植物叶片N、P含量和N:P在海拔梯度上的变化格局及其驱动因子的研究还有待深入。
以往研究对叶片性状空间格局及其机制进行了深入的探讨。一般认为, 在大尺度上气候是影响叶N、P含量的一个因素。如Reich和Oleksyn (2004)在全球尺度上的研究表明, 植物叶片的N、P含量随年平均气温的升高而降低, 而N:P增加。其他研究也证实了气候对叶N、P化学计量特征有显著影响, 但发现气候的解释力往往不高(He et al., 2010), 说明还有其他因素起重要作用。不同植物功能类群的叶片N、P含量具有显著差异。例如草本植物的N、P含量显著高于木本植物, 落叶树种高于常绿树种, 阔叶树种高于针叶树种(Han et al., 2005; 吴统贵等, 2010)。具有长寿命叶片的植物具有较低的N、P含量(Reich & Oleksyn, 2004), 因此常绿植物能适应低营养状况的生存环境。同低海拔地区植物相比, 高海拔地区的植物常绿树种所占的比例较大, 因此N、P含量较低。一些研究认为, 气候改变群落的物种组成是导致叶片N、P含量地理变异的一个重要因素(He et al., 2008, 2010)。在系统发育关系上, 亲缘关系越远的植物叶片N和P含量差异往往越大(如蕨类植物和种子植物), 亲缘关系越近的植物叶片N和P含量差异较小(如双子叶和单子叶植物)(任书杰等, 2007)。关于气候和物种(谱系关系)的相对作用大小, 近年来多数研究表明, 对叶性状变异起主导作用的是系统发育关系, 其解释力明显高于气候等环境因子(He et al., 2010; Liu et al., 2013)。
叶片N、P含量受气候、植物功能型、系统发育关系等多种因素的影响(Reich & Oleksyn, 2004; Han et al., 2005, 2011; He et al., 2010; Liu et al., 2013; 樊江文等, 2014)。而且以上因子之间可能存在相互影响, 共同作用于植物叶片N、P含量。例如气候可能导致植物功能型、物种组成改变, 从而影响叶N、P含量(He et al., 2008, 2010)。本文研究长白山海拔梯度上叶片N、P化学计量特征, 主要探究以下两个问题: 1)海拔梯度上木本植物叶片N、P含量和N:P的空间格局; 2)气候、植物功能型、谱系对叶片N、P含量以及N:P的作用大小, 以进一步揭示叶片N、P元素在海拔梯度上的变化机制。
1 研究方法
1.1 研究地点
长白山(41.38°-42.60° N, 126.92°-129.00° E), 位于我国吉林省东南部, 与朝鲜相邻, 是欧亚大陆东岸的最高山, 也是东北植物区系唯一拥有明显植被垂直带谱的山脉(赵淑清等, 2004)。长白山地区属温带大陆性山地气候, 随着海拔的升高(海拔500-2 744 m), 年降水量逐渐增加(720-1 400 mm), 而年平均气温从3.5 ℃下降到-7.4 ℃。长白山海拔梯度上气候条件的巨大差异, 使长白山地区的植被具有典型的垂直结构, 由下至上依次形成了阔叶红松混交林(Pinus koraiensis and broadleaf mixed forest, 海拔500-1 100 m)、云冷杉林(Picea and Abies forest, 海拔1 100-1 800 m)、亚高山岳桦林带(Betula ermanii forest, 海拔1 800-2 000 m)和高山苔原带(alpine tundra, 海拔2 000 m以上) 4个植被垂直带(Zhu et al., 2010)。土壤类型也有着明显的垂直分异, 其中, 海拔1 000 m以下为暗棕色森林土, 海拔1 000-1 800 m为棕色针叶林土, 海拔1 800-2 000 m为石灰性钙化土, 海拔2 000 m以上为苔原土(王战等, 1980)。
1.2 叶片采集及性状测定
2013年的7、8月, 我们在长白山沿海拔梯度从阔叶红松林带至岳桦林带(海拔530-1 940 m), 选择典型林分设置了14块20 m × 50 m的样地进行群落调查, 具体操作方法见方精云等(2009)制定的植物群落调查方法和技术规范。同时利用GPS、海拔表记录样地的经纬度、海拔等地理信息。样地基本情况见附录I。对每块样地中乔木、灌木层的优势种, 选择健康成熟、长势均匀的5株个体, 在树冠外层向阳面的枝条上, 每个植株摘取5-10片充分展开、成熟向阳、无病虫害的叶片。叶片在80 ℃烘箱中烘干至恒质量, 将烘干的叶片粉碎、过筛、装袋, 以备化学分析。采用C/N元素分析仪(2400 II CHNS/O Element Analyzer, Perkin-Elmer, Boston, USA)测定样品全N含量(g·kg-1), 采用钼蓝比色法测定样品全P含量(g·kg-1), 叶片N:P =叶片全N含量/叶片全P含量。
1.3 气候数据的计算
我们采用以下公式来推算各样地的气温和降水量:T (P) = a + b × Lat + c × Long + d × Alt其中, T和P分别为1-12月的月平均气温和月降水量, a、b、c、d为回归常数, Lat、Long和Alt分别为纬度、经度和海拔(方精云, 1992)。采用Wang等(2008)利用国家气象台站数据建立的模型估算各样地的月气候指标, 在此基础上计算年平均气温(MAT)和年降水量(MAP)。对这些模型的验证结果表明, 年平均气温的预测精度为96%, 年降水量的预测精度为79%, 足以满足本研究的需要。在数据的预分析中, 我们还考虑了生长季平均气温、生长季降水量、最冷月平均气温和温暖指数等多种气候指标。这些气候指标和MAT、MAP的相关性很强(R2 > 0.9), 对叶N、P含量的解释力也与MAT、MAP接近。由于以往多数大尺度上的研究使用的是MAT和MAP (Reich & Oleksyn, 2004; Han et al., 2005, 2011), 为了便于和这些研究结果进行比较, 以及减少变量间的共线性, 在最终分析中我们采用了MAT和MAP进行统计分析。
1.4 植物功能型和谱系组划分
本研究共采集了431份植物样品, 共获得19科30属48个物种的叶功能性状数据。通过查阅《东北植物志》, 将长白山木本植物划分为以下3种植物功能型: 生活型(分为乔木、灌木植物)、叶型(分为针叶、阔叶植物)、叶片习性(分为常绿、落叶植物)。
为了检验谱系关系对叶片N、P化学计量特征的影响, 我们基于Zanne等(2014)的进化树, 利用R软件中的phylomatic函数, 建立了所采样物种的系统发育树(附录II)。为了比较气候和谱系关系的解释力的大小, 我们采用He等(2009)建立的方法, 在系统发育树的90 Mya (million years ago)和60 Mya两个进化时间节点处将物种划分为入不同的谱系组。选择这两个时间节点是因为它们所划分成的谱系组数目(90 Mya 谱系组包括19类, 60 Mya谱系组包括30类), 与传统分类学上的分类数目(19科30属)一致, 便于两种分类方式结果的比较(He et al., 2010; Liu et al., 2013; Sun et al., 2017)。
1.5 数据分析
本研究中将影响叶片N、P元素变异的因子分为3类: 一是气候因子, 包括MAT、MAP; 二是植物功能型, 包括生活型、叶型、叶片习性; 三是谱系组, 包括90 Mya、60 Mya谱系组和物种。叶片N、P含量和N:P数据在分析前均经log10对数转换, 以提高解释模型残差的正态性(He et al., 2009)。对解释模型残差的Q-Q图检验表明, 残差基本呈正态分布, 因此解释模型是可接受的(Crawley, 2007)。我们使用独立样本t检验比较不同植物功能型(乔木与灌木、针叶与阔叶、常绿与落叶)之间各性状的差异; 采用一般线性模型(GLMs)和混合模型的方差分析检验不同因子对叶片N、P含量和N:P的解释力大小(Balvanera et al., 2006; He et al., 2009)。为了探究气候和谱系是如何共同作用于叶片N、P元素变异的, 我们采用变异分离的方法, 将N、P元素变异划分为3部分: (1) a, b分别表示气候、谱系的单独解释力; (2) c表示气候和谱系的协同作用; (3)未被解释的变异: 除了气候和谱系外, 其他未在本研究中涉及的因素(He et al., 2009; Liu et al., 2013)。采用F检验对a、b解释力的显著性进行检验。
所有的数据分析均在R 3.3.2中完成。
2 结果
2.1 长白山叶片N、P含量和N:P沿海拔梯度的变化
由表1可以看出, 长白山地区叶片N含量的范围是5.10-37.90 g·kg-1, 叶片P含量0.66-6.75 g·kg-1, 叶片N:P值为2.47-33.70。不同植物功能型的叶片N、P含量具有显著差异。灌木的叶片N含量(22.37 g·kg-1)显著高于乔木(20.74 g·kg-1), 两者的P含量、N:P差异不显著; 阔叶植物的叶片N、P含量和N:P (分别为23.02 g·kg-1、2.32 g·kg-1、11.19)均显著高于针叶植物(分别为15.50 g·kg-1、1.91 g·kg-1、9.35); 落叶植物叶片N、P含量和N:P显著高于常绿植物。由图1、图2可以看出, 叶片N含量随海拔升高而下降(R2 = 0.012, p < 0.05), 随气温的升高而增加(R2 = 0.012, p < 0.05), 随降水量的增加而下降(R2 = 0.011, p < 0.05); 叶片P含量与海拔关系不显著, 并且与气温、降水量均无显著关系; 叶片N:P随海拔的升高而下降(R2 = 0.016, p < 0.01), 与气温正相关(R2 = 0.016, p < 0.01), 与降水量负相关(R2 = 0.016, p < 0.01)。
2.2 气候、植物功能型、谱系对叶片N、P含量和N:P的影响
由表2可以看出, 对于叶片N含量, 气候(MAT、MAP)、3种植物功能型(生活型、叶型、叶片习性)和谱系(90 Mya谱系组、60 Mya谱系组、物种)均具有显著的解释力; 对于叶片P含量, 仅叶型、叶片习性和谱系对其有显著作用, 而气候、生活型不影响叶片P含量; 叶片N:P不受生活型的影响, 其他因子作用均显著。“种”水平的解释力均最大。Table 1
表1
表1长白山木本植物叶片氮、磷含量和氮磷比统计特征
Table 1Statistics of leaf nitrogen (N), phosphorus (P) concentrations and N:P of the examined plant species in Changbai Mountain
| 叶性状 Leaf trait | 功能群 Functional group | 样本量 n | 平均值 Mean | 最大值 Max | 最小值 Min | 标准偏差 SD | 标准误差 SE | 变异系数 CV | |
|---|---|---|---|---|---|---|---|---|---|
| 氮含量 N concentration (g·kg-1) | 全部样品 All samples | 431 | 21.46 | 37.90 | 5.10 | 6.75 | 0.33 | 0.31 | |
| 生活型 Life form | 乔木 Tree | 240 | 20.74b | 37.90 | 5.10 | 6.80 | 0.44 | 0.33 | |
| 灌木 Shrub | 191 | 22.37a | 37.40 | 6.80 | 6.59 | 0.48 | 0.29 | ||
| 叶型 Leaf shape | 针叶 Coniferous | 89 | 15.50b | 36.80 | 8.50 | 4.91 | 0.52 | 0.32 | |
| 阔叶 Broad-leaved | 342 | 23.02a | 37.90 | 5.10 | 6.29 | 0.34 | 0.27 | ||
| 叶片习性 Leaf habit | 常绿 Evergreen | 87 | 14.49b | 35.00 | 8.44 | 4.42 | 0.47 | 0.31 | |
| 落叶 Deciduous | 344 | 23.23a | 37.90 | 5.10 | 6.06 | 0.33 | 0.26 | ||
| 磷含量 P concentration (g·kg-1) | 全部样品 All samples | 431 | 2.23 | 6.75 | 0.66 | 0.87 | 0.04 | 0.39 | |
| 生活型 Life form | 乔木 Tree | 240 | 2.22 | 6.75 | 0.66 | 0.93 | 0.06 | 0.42 | |
| 灌木 Shrub | 191 | 2.25 | 5.29 | 0.77 | 0.80 | 0.06 | 0.35 | ||
| 叶型 Leaf shape | 针叶 Coniferous | 89 | 1.91b | 4.13 | 0.66 | 0.77 | 0.08 | 0.40 | |
| 阔叶 Broad-leaved | 342 | 2.32a | 6.75 | 0.77 | 0.88 | 0.05 | 0.38 | ||
| 叶片习性 Leaf habit | 常绿 Evergreen | 87 | 1.74b | 3.30 | 0.66 | 0.66 | 0.07 | 0.38 | |
| 落叶 Deciduous | 344 | 2.36a | 6.75 | 0.99 | 0.87 | 0.05 | 0.37 | ||
| N:P | 全部样品 All samples | 431 | 10.81 | 33.70 | 2.47 | 4.77 | 0.23 | 0.44 | |
| 生活型 Life form | 乔木 Tree | 240 | 10.60 | 33.70 | 2.47 | 4.89 | 0.32 | 0.46 | |
| 灌木 Shrub | 191 | 11.07 | 26.34 | 3.12 | 4.62 | 0.33 | 0.42 | ||
| 叶型 Leaf shape | 针叶 Coniferous | 89 | 9.35b | 25.92 | 3.90 | 4.68 | 0.50 | 0.50 | |
| 阔叶 Broad-leaved | 342 | 11.19a | 33.70 | 2.47 | 4.72 | 0.26 | 0.42 | ||
| 叶片习性 Leaf habit | 常绿 Evergreen | 87 | 9.75b | 25.92 | 3.28 | 4.93 | 0.53 | 0.51 | |
| 落叶 Deciduous | 344 | 11.08a | 33.70 | 2.47 | 4.70 | 0.25 | 0.42 | ||
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显示原图|下载原图ZIP|生成PPT图1叶片氮(N) (A)、磷(P) (B)含量和氮磷比(N:P) (C)与海拔的关系。
-->Fig. 1Leaf nitrogen (N) (A), phosphorus (P) (B) concentrations and N:P (C) in relation to altitude.
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显示原图|下载原图ZIP|生成PPT图2叶片氮(N)、磷(P)含量和氮磷比(N:P)与年平均气温和年降水量的关系。
-->Fig. 2Leaf nitrogen (N), phosphorus (P) concentrations and N:P in relation to mean annual temperature and mean annual precipitation.
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表3的结果显示, 气候(MAT、MAP)对叶片N含量和N:P均有显著影响。所有的植物功能型对N含量有显著作用, 叶型对N含量的解释力最大(22.60%); 仅叶型和叶片习性对P含量有显著作用, 解释力分别为5.70%和6.46%; 仅有叶型对N:P有显著作用, 解释力为2.24%。谱系对叶片N、P含量和N:P均有较强的解释力, 其中90 Mya谱系组的解释力最大, 为12.06%-13.18%, 60 Mya谱系组的解释力为3.82%-8.10%, “种”的解释力为5.01%-7.32%。这和采用科属种分类的分析结果(附录III)相似: “科”的解释力为9.81%-11.50%, “属”的解释力为3.11%- 8.79%, “种”的解释力为6.29%-11.73%。90 Mya谱系组和60 Mya谱系组对叶片N、P含量和N:P变异的解释力和科属的解释力基本上是相同的(He et al., 2010)。
Table 2
表2
表2各变量对氮(N)、磷(P)含量和氮磷比(N:P)的单因子解释力
Table 2Contributions of each factor to the variations of leaf nitrogen (N), phosphorus (P) concentrations and N:P
| 叶性状 Leaf traits | 年平均气温 Mean annual temperature | 年降水量 Mean annual precipitation | 生活型 Life form | 叶型 Leaf shape | 叶片习性 Leaf habit | 90 Mya谱系组 90 Mya division | 60 Mya谱系组 60 Mya division | 物种 Species | |
|---|---|---|---|---|---|---|---|---|---|
| 氮含量 N concentration | R2 | 0.012 | 0.012 | 0.016 | 0.229 | 0.324 | 0.415 | 0.459 | 0.536 |
| p | 0.023* | 0.025* | 0.008** | 0.000*** | 0.000*** | 0.000*** | 0.000*** | 0.000*** | |
| 磷含量 P concentration | R2 | 0.003 | 0.003 | 0.002 | 0.049 | 0.115 | 0.216 | 0.306 | 0.391 |
| p | 0.233 | 0.247 | 0.342 | 0.000*** | 0.000*** | 0.000*** | 0.000*** | 0.000*** | |
| N:P | R2 | 0.016 | 0.016 | 0.003 | 0.028 | 0.018 | 0.152 | 0.215 | 0.303 |
| p | 0.008** | 0.009** | 0.257 | 0.000*** | 0.005** | 0.000*** | 0.000*** | 0.000*** |
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Table 3
表3
表3气候、植物功能型、谱系对叶片氮(N)、磷(P)含量和氮磷比(N:P)影响的一般线性模型
Table 3Summary of results of general linear model analysis for the effects of environmental factors, plant functional groups and phylogeny (90 and 60 Mya phylogenetic group, species) on leaf nitrogen (N), phosphorus (P) concentrations and N:P
| N | P | N:P | |||||||
|---|---|---|---|---|---|---|---|---|---|
| df | %SS | p | df | %SS | p | df | %SS | p | |
| 气候 Climate | |||||||||
| 年平均气温 Mean annual temperature | 1 | 1.21 | 0.001** | 1 | 0.33 | 0.146 | 1 | 1.65 | 0.002** |
| 年降水量 Mean annual precipitation | 1 | 0.61 | 0.022* | 1 | 0.45 | 0.089 | 1 | 1.28 | 0.007** |
| 植物功能型 Plant functional groups | |||||||||
| 生活型 Life form | 1 | 1.54 | 0.000*** | 1 | 0.22 | 0.237 | 1 | 0.27 | 0.214 |
| 叶型 Leaf shape | 1 | 22.60 | 0.000*** | 1 | 5.70 | 0.000*** | 1 | 2.24 | 0.000*** |
| 叶片习性 Leaf habit | 1 | 9.34 | 0.000*** | 1 | 6.46 | 0.000*** | 1 | 0.02 | 0.767 |
| 谱系 Phylogeny | |||||||||
| 90 Mya谱系组 90 Mya division | 18 | 12.06 | 0.000*** | 18 | 13.03 | 0.000*** | 18 | 13.18 | 0.000*** |
| 60 Mya谱系组 60 Mya division | 10 | 3.82 | 0.000*** | 10 | 8.10 | 0.000*** | 10 | 7.34 | 0.000*** |
| 物种 Species | 17 | 5.01 | 0.000*** | 17 | 6.33 | 0.002** | 17 | 7.32 | 0.001** |
| 残差 Residuals | 380 | 43.81 | 380 | 59.36 | 380 | 66.72 | |||
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从方差分解的结果(图3)可以看出, 气候和谱系均对叶片N、P含量和N:P具有显著作用。气候的独立作用显著但解释力不高, 仅占1.50%-2.98%; 谱系的独立解释力高达30.36%-54.38%, 而气候和谱系的协同作用不显著。另外, 未被解释的变异占43.81%- 66.72%。
3 讨论
3.1 叶片N、P含量与海拔、气候因子的关系
长白山地区木本植物叶片N含量的平均值为21.46 g·kg-1、叶片P含量为2.23 g·kg-1, 叶片N:P为10.81 (表1)。对全球1 280种陆生植物的研究发现, 它们叶片N含量的平均值为20.1 g·kg-1, 叶片P含量的平均值为1.80 g·kg-1, N:P平均为13.8 (贺金生和韩兴国, 2010)。Han等(2005)研究中国753种陆生植物得出, 叶片N、P含量和N:P的平均值分别为18.6 g·kg-1、1.21 g·kg-1和14.4。Zhao等(2014)对长白山地区175种植物的研究发现, 该地区植物的叶片N、P含量(分别为24.13 g·kg-1和2.22 g·kg-1)高于中国植物的叶片N、P含量, N:P (11.54)则低于全国尺度。本研究结果与之一致。Zhao等(2014)通过对长白山地区土壤养分含量的分析认为, 长白山地区叶片N、P含量高于全国水平的原因是长白山地区的土壤养分含量高于全国水平。
显示原图|下载原图ZIP|生成PPT图3气候和物种(即谱系的总解释力, 见
A, 叶片N含量。B, 叶片P含量。C, 叶片N:P。a和b分别为气候和谱系的单独解释力, c为两者的协同作用。采用F检验对a、b的显著性进行检验。***, p < 0.001; **, p < 0.01。
-->Fig. 3Variation partitioning analysis for leaf nitrogen (N), phosphorus (P) concentrations and N:P by three types of factors: Climate (mean annual temperature and mean annual precipitation), species (phylogenetic differences among species,
A, leaf N concentration. B, leaf P concentration. C, N:P. a and b are the pure effects of climate and species, respectively; c is their synergistic effects. The significances of a and b were evaluated with F test. ***, p < 0.001; **, p < 0.01.
-->
本文结果表明, 长白山地区的木本植物叶片N含量、N:P均在海拔梯度上呈现出显著变化, 并且受气温、降水的显著影响, N含量、N:P高的植物出现在气温高、降水量少、环境较为适宜的低海拔地区; 而叶片P含量与海拔和气候因子的关系均不显著(图1, 图2)。已有的大尺度上的研究发现, 纬度梯度上的从北向南, 经度梯度上的从西到东, 热量和水分均呈现出逐渐增加的趋势, 叶片N、P含量的变化趋势在这两个梯度上呈现出一致性, 均从北到南、从西向东下降(Han et al., 2011)。但是在长白山海拔梯度上, 叶片P含量并未呈现出显著的变化。
本研究中叶片N含量随海拔的增加而降低, 与气温正相关, 与降水负相关, 这与部分海拔梯度上的研究结果(van de Weg et al., 2009; Zhao et al., 2014)一致。但是在水平(纬度、经度)梯度上, 全球以及全国尺度上的大部分研究表明, 叶片N含量与气温呈负相关关系(Reich & Oleksyn, 2004; Han et al., 2005, 2011; Chen et al., 2013)。本文的结果表明叶N含量在海拔梯度上的变化机制与水平梯度上可能存在差异。高海拔地区的低温和多降水带来的淋溶作用导致N的矿化率较低(Reich & Oleksyn, 2004; Zhao et al., 2014), 抑制了植物根系对N的吸收利用效率, 因此叶N含量随海拔的升高而降低。而且高海拔地区气候寒冷、条件恶劣、植物生长慢, 相比低海拔地区的植物不需要进行大量的代谢活动(Elser et al., 2003), 也会导致叶N含量随海拔的升高而降低。
本研究中叶片P含量与海拔没有显著的相关关系。陆地生态系统中的P元素不会通过微生物活动进入生态系统, 几乎所有的活性P都来源于岩石风化(Smeck, 1985), 然后通过与其他元素结合形成难以分解的化合物进入土壤中。相比N元素, P在土壤溶液中的扩散更不易(Aerts & Champion, 1999)。本研究中, 叶片P含量与气温、降水的关系均不显著。N、P均为植物生长的限制性元素, 但是同叶片N含量相比, 叶P含量在海拔梯度上更稳定。这个结果与水平梯度上的结论相反, 有研究认为, 纬度梯度上叶片P含量更为活跃, 与气候、土壤等环境因子的关系更为密切(Lovelock et al., 2007; Chen et al., 2013)。
长白山地区叶片N:P随海拔的升高而降低。叶片N:P的驱动因子更为复杂。有研究认为物种由于其特有的营养平衡机制会选择其最适N:P (Kay et al., 2005), 环境的筛选作用在一定的地理区域内形成物种库, 从而导致群落尺度上的N:P差异(He et al., 2008)。在不考虑物种组成的情况下, 土壤可能直接控制植物N:P。在极端环境中, 植物倾向于积累过量的营养元素(Chapin et al., 1990), 有研究证实施肥后植物会积累过量的P而不是过量的N (Cordell et al., 2001)。这种存储过程可能导致了低的N:P (He et al., 2008)。
3.2 气候、功能型、谱系对叶片N、P含量和N:P的作用
除了气候等环境因子, 叶片N、P含量和N:P还受植物功能型、物种组成的强烈影响(Reich & Oleksyn, 2004; Han et al., 2011)。在本文中, 气候、植物功能型和谱系的一般线性模型表明这三者能够解释大部分的叶片N、P含量在海拔梯度上的变异(表3)。我们的结果表明: 不同功能型植物叶片的N、P含量和N:P差异显著, 灌木物种的叶N含量显著高于乔木, 两者的叶片P含量差异不显著; 阔叶、落叶物种的N、P含量分别显著高于针叶、常绿树种(表1), 与以往的研究结果(Han et al., 2005; Chen et al., 2013)一致。可见植物功能型是叶片N、P含量变异的重要影响因子。在不考虑谱系的研究中, 植物功能型的解释力所占比例最大(Han et al., 2011; Chen et al., 2013)。本研究也显示植物功能型对叶片N、P含量和N:P的解释力远大于气候(表3)。植物功能型对叶片N含量的解释力为33.48%, 而对于叶P含量、N:P仅为12.38%、2.53% (表3)。Chen等(2013)研究我国东部的386种木本植物发现, 叶片N含量主要取决于植物功能型(常绿阔叶、落叶阔叶、针叶植物)沿纬度梯度的变化, 而叶片P含量、N:P与气候、植物功能型和它们之间的相互作用有关。叶片N含量在不同的功能型之间差异较大, 而P含量、N:P更多地受谱系关系影响。我们的结果还表明叶型对叶片N、P含量和N:P均有显著的解释力, 对叶片N含量的解释力高达22.60%, 叶型(针叶、阔叶)可能是比植物生活型(乔、灌木)对叶片N、P含量影响更大的因子。
关于谱系和气候的相对作用大小, 本文结果表明谱系(90 Mya 谱系组、60 Mya 谱系组、物种)对叶片N、P含量和N:P的作用是最显著的(表2, 表3), “物种”的单独解释力为30.36%-54.38% (图3), 远大于气候的解释力, 与已有的研究结果一致, 系统发育关系是叶片N、P元素格局形成的最主要的因素(He et al., 2010; Liu et al., 2013)。
气候对叶N、P含量和N:P的作用均显著(p < 0.05), 但解释力不高(1.50%-2.98%)(图3)。海拔梯度上的研究结果与以往水平梯度上的研究结论一致: 气候对叶片N、P含量的影响是“适中” (modest), 而不是如预想的那样强烈(Wright et al., 2005; He et al., 2008)。在本文中, 气候和“物种”对叶片N、P含量和N:P的共同作用均不显著。以往的研究认为气候对陆地生态系统的结构和功能的地理格局(例如植物功能型、物种分布、生物多样性、植被初级生产力、植物生态学特性等)具有重要作用(Hedin, 2004; Wright et al., 2004), 并认为气候导致的物种组成和生活型变化是叶性状地理变化的重要驱动力(He et al., 2008)。比如, He等(2006)研究中国草原群落发现, 气温对叶片N含量的作用主要是通过改变物种组成, 而不是直接作用于叶性状。本研究结果和上述结果有明显差异: 海拔梯度上的气候和物种组成变化对叶片N、P含量和N:P没有显著的协同作用(图3)。为什么会有这种差异还有待于进一步研究。但本文在海拔梯度上的研究结果, 与以往水平梯度上的结果表现出诸多差异。这可能说明在纬度梯度上得到的结果并不能直接推广到海拔梯度上。今后需要系统开展海拔和纬度梯度上的比较研究, 以深入理解叶片化学计量特征地理格局的影响机制。
The authors have declared that no competing interests exist.
作者声明没有竞争性利益冲突.
参考文献 原文顺序
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被引期刊影响因子
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| [5] | . Abstract Leaf-level studies of Metrosideros polymorpha Gaud. (Myrtaceae) canopy trees at both ends of a substrate age gradient in the Hawaiian Islands pointed to differential patterns of adjustment to both nutrient limitation and removal of this limitation by long-term (8-14 years) nitrogen (N), phosphorus (P) and N + P fertilizations. The two study sites were located at the same elevation, had similar annual precipitation, and supported forests dominated by M. polymorpha, but differed in the age of the underlying volcanic substrate, and in soil nutrient availability, with relatively low N at the young site (300 years, Thurston, Hawaii) and relatively low P at the oldest site (4,100,000 years, Kokee, Kauai). Within each site, responses to N and P fertilization were similar, regardless of the difference in soil N and P availability between sites. At the young substrate site, nutrient addition led to a larger mean leaf size (about 7.4 versus 4.8 cm2), resulting in a larger canopy leaf surface area. Differences in foliar N and P content, chlorophyll concentrations and carboxylation capacity between the fertilized and control plots were small. At the old substrate site, nutrient addition led to an increase in photosynthetic rate per unit leaf surface area from 4.5 to 7.6 micromol m(-2) s(-1), without a concomitant change in leaf size. At this site, leaves had substantially greater nutrient concentrations, chlorophyll content and carboxylation capacity in the fertilized plots than in the control plots. These contrasting acclimation responses to fertilization at the young and old sites led to significant increases in total carbon gain of M. polymorpha canopy trees at both sites. At the young substrate site, acclimation to fertilization was morphological, resulting in larger leaves, whereas at the old substrate site, physiological acclimation resulted in higher leaf carboxylation capacity and chlorophyll content. |
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| [10] | . . 植物群落是不同植物在长期环境变化中相互作用、相互适应而形成的组合.它提供着人类赖以生存的主要物质资源,维系着地球生态系统的健康和功能,也为各种动物和其他生物提供食物来源和栖息地,是人类生存和发展不可或缺的物质基础,具有不可替代的作用.我国植物群落类型多样,在世界上首屈一指,但我国至今尚没有一次全面和系统的植物群落清查,不仅影响了人们对我国植物资源的了解、利用和保护,也不利于我国生态学、环境科学和地理学等相关学科的发展.采用统一的方法体系和技术规范开展我国植物群落的清查工作势在必行,并具有紧迫性.本文基于作者长期的野外工作实践和国内外的群落调查方法,首先简要定义了与植物群落清查有关的重要概念,在此基础上,论述了调查样地的设置原则和体系、群落清查的技术指标和方法、主要优势种生态属性的测定方法和规范,并介绍了大样地调查的主要步骤.通过本文的介绍、归纳和总结,试图为制定我国植物群落清查的技术规范提供基础材料和技术储备. |
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| [15] | . Abstract Nitrogen (N) and carbon-nitrogen (C:N) ratio are key foliar traits with great ecological importance, but their patterns across biomes have only recently been explored. We conducted a systematic census of foliar C, N and C:N ratio for 213 species, from 41 families over 199 research sites across the grassland biomes of China following the same protocol, to explore how different environmental conditions and species composition affect leaf N and C:N stoichiometry. Leaf C:N stoichiometry is stable in three distinct climatic regions in Inner Mongolia, the Tibetan Plateau, and Xinjiang Autonomous Region, despite considerable variations among co-existing species and among different vegetation types. Our results also show that life form and genus identity explain more than 70% of total variations of foliar N and C:N ratio, while mean growing season temperature and growing season precipitation explained only less than 3%. This suggests that, at the biome scale, temperature affects leaf N mainly through a change in plant species composition rather than via temperature itself. When our data were pooled with a global dataset, the previously observed positive correlation between leaf N and mean annual temperature (MAT) at very low MATs, disappeared. Thus, our data do not support the previously proposed biogeochemical hypothesis that low temperature limitations on mineralization of organic matter and N availability in soils lead to low leaf N in cold environments. |
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| [17] | . Leaf N and P stoichiometry covaries with many aspects of plant biology, yet the drivers of this trait at biogeographic scales remain uncertain. Recently we reported the patterns of leaf C and N based on systematic census of 213 species over 199 research sites in the grassland biomes of China. With the expanded analysis of leaf P, here we report patterns of leaf P and N:P ratios, and analyze the relative contribution of climatic variables and phylogeny in structuring patterns of leaf N:P stoichiometry. Average values of leaf P and N:P ratio were 1.9/mg/g 1 and 15.3 (mass ratio), respectively, consistent with the previous observation of a higher N:P ratio in China flora than the global averages (ca. 13.8), resulting from a lower leaf P. Climatic variables had very little direct correlation with leaf P and N:P ratios, with growing season precipitation and temperature together explaining less than 2% of the variation, while inter-site differences and within-site phylogenetic variation explained 55 and 26% of the total variation in leaf P and N:P ratios. Across all sites and species, leaf N and P were highly positively correlated at all levels. However, the within-site, within-species covariations of leaf N and P were weaker than those across sites and across species. Leaf N and P relationships are driven by both variation between sites at the landscape scale (explaining 58% of the variance) and within sites at the local scale (explaining 24%), while the climatic factors exerted limited influence (explaining less than 3%). In addition, leaf N:P ratios in two dominant genera Kobresia and Stipa had different responses to precipitation. This study suggests that geographic variation and between-species variation, rather than climatic variation, are the major determinants of grassland foliar stoichiometry at the biome level. |
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| [19] | . Abstract Assessing the influence of climate, soil fertility, and species identity on leaf trait relationships is crucial for understanding the adaptations of plants to their environment and for interpreting leaf trait relationships across spatial scales. In a comparative field study of 171 plant species in 174 grassland sites across China, we examined the trade-offs, defined as negative covariance between two traits, between leaf persistence (leaf mass per area, LMA) and leaf productivity (mass-based photosynthetic rate, Amass, N and P content, and photosynthetic N use efficiency, PNUE). We asked to which extent these trade-offs were influenced by: (1) variation among sites within species, decomposed into variation due to climatic and soil variables; (2) variation among species within sites, decomposed into variation among taxonomic, functional, or phylogenetic groups; and (3) the joint contribution of variation among species and sites. We used mixed-model analysis of covariance to partition bivariate relationships between leaf traits into trade-off components. We found significant mass-based persistence-productivity trade-offs of LMA-Amass, LMA-N, LMA-P, and LMA-PNUE consistent with previous broadscale findings. Overall, (1) variation among sites within species explained 14-23%, (2) variation among species within sites explained 20-34%, and (3) the two together explained 42-63% of the total covariance between leaf traits. Interspecific trade-offs of LMA-Amass, LMA-N, and LMA-P were stronger than inter-site ones. A relatively low amount of covariance was explained by climatic and soil variables. However, we found the trade-offs were stronger for LMA-N and LMA-P at higher precipitation and for LMA-PNUE at greater soil fertility, if displayed by major axis regression, which combined both intra- and interspecific variation. Residual trade-offs within species and sites were weak, suggesting that intraspecific, intra-site variation in physiology was less important than variation imposed by species identity or environmental differences among sites. Our results from grassland biomes add evidence for the fundamental nature of productivity-persistence trade-offs in plants. No individual factor emerged as the single major cause for these trade-offs. Rather, the total covariance between leaf traits was explained by a combination of factors, each contributing a range of explanatory power. |
| [20] | . PubMed comprises more than 23 million citations for biomedical literature from MEDLINE, life science journals, and online books. Citations may include links to full-text content from PubMed Central and publisher web sites. |
| [21] | . Abstract Ecological stoichiometry, the study of the balance of energy and materials in living systems, may serve as a useful synthetic framework for evolutionary biology. Here, we review recent work that illustrates the power of a stoichiometric approach to evolution across multiple scales, and then point to important open questions that may chart the way forward in this new field. At the molecular level, stoichiometry links hereditary changes in the molecular composition of organisms to key phenotypic functions. At the level of evolutionary ecology, a simultaneous focus on the energetic and material underpinnings of evolutionary tradeoffs and transactions highlights the relationship between the cost of resource acquisition and the functional consequences of biochemical composition. At the macroevolutionary level, a stoichiometric perspective can better operationalize models of adaptive radiation and escalation, and elucidate links between evolutionary innovation and the development of global biogeochemical cycles. Because ecological stoichiometry focuses on the interaction of energetic and multiple material currencies, it should provide new opportunities for coupling evolutionary dynamics across scales from genomes to the biosphere. |
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| [24] | . Aims Understanding the drivers for leaf traits is critical to improving our predictions on ecosystems’ responses to global changes. Geographic patterns of leaf traits are shaped by phylogenetic, biological and environmental factors simultaneously. However, till now few studies have examined how these factors influenced leaf traits together, and how their effects differed at the within- and among-site levels. Methods We sampled leaf traits from a 1100 km shrub-biome transect across central Inner-Mongolia, including leaf mass per area (LMA), mass-based photosynthetic rate, nitrogen (N) and phosphorus (P) concentrations. We examined the effects of phylogenetic, biological (height and growth rate) and environmental (climate and soil) factors on leaf traits with mixed-model analyses of variance. Variation partitioning method was used to separate the joint and independent effects of these three types of factors. Important Findings (i) Climate and soil fertility (total or available nutrient concentrations) together explained 11.4–41.4% of among-site variations, with remarkable difference among traits. (ii) Height and height growth rate together explained 0.4–31.9% of trait variations (mostly among-site variations). Our results could only weakly support the ability of leaf traits as predictors for whole-plant growth. (iii) LMA was negatively related to height, which was consistent with the resource-use strategy hypothesis but inconsistent with the hypotheses proposed for coexisting trees, suggesting that the LMA–height relationship is shaped by rather different mechanisms between the within- and among-communities levels. (iv) The variation partitioning analysis showed that, the relationships between leaf traits and biological characters largely reflected the differences in both leaf traits and biological characters among species that occupying different sites. The relative importance of phylogenetic, biological and environmental factors differed remarkably among leaf traits, between the within- and among-communities levels, and between different biomes. (v) Our results strongly suggest the necessity of examining the three types of factors simultaneously, and at both the within- and among-communities levels, for a better understanding of the drivers for leaf traits patterns. |
| [25] | . |
| [26] | . A global data set including 5,087 observations of leaf nitrogen (N) and phosphorus (P) for 1,280 plant species at 452 sites and of associated mean climate indices demonstrates broad biogeographic patterns. In general, leaf N and P decline and the N/P ratio increases toward the equator as average temperature and growing season length increase. These patterns are similar for five dominant plant groups, coniferous trees and four angiosperm groups (grasses, herbs, shrubs, and trees). These results support the hypotheses that (i) leaf N and P increase from the tropics to the cooler and drier midlatitudes because of temperature-related plant physiological stoichiometry and biogeographical gradients in soil substrate age and then plateau or decrease at high latitudes because of cold temperature effects on biogeochemistry and (ii) the N/P ratio increases with mean temperature and toward the equator, because P is a major limiting nutrient in older tropical soils and N is the major limiting nutrient in younger temperate and high-latitude soils. |
| [27] | . . 大尺度的叶片的氮(N)和磷(P)的化学计量学特征是植物对环境条件的长期适应的结果,并能为大尺度模型的发展提供数据基础.通过对文献数据的搜集整理,对中国东部南北样带168个采样点的654种植物的N和P的化学计量学特征、空间格局及其与气候因子(年均温度)的关系进行了研究.结果表明,中国东部南北样带654种植物叶片的N和P的化学计量学特征存在很大的变异性.叶片N的变化范围为2.17～52.61 mg·g-1,几何平均数为17.55 mg·g-1,叶片P的变化范围为0.10-10.27 mg·g-1,几何平均数为1.28 mg·g-1,叶片N/P的变化范围为1.7～74.6,几何平均数为13.5.中国东部南北样带乃至中国区域叶片P的含量显著低于全球尺度的其它研究结果,这说明与全球尺度相比,中国区域植被生长更易受到P的限制.在所研究的不同功能群间,以叶片N含量的差异最大,P次之,N/P的差异最小,这与在多数的功能群内,其叶片N和P含量间存在显著相关性有关.另外在系统发育上亲缘越远的功能群N和P含量差异越大(蕨类植物与种子植物),亲缘越近的差异越小(双子叶植物与单子叶植物).叶片的N和P含量与纬度和年均温度间存在极显著的相关关系,随着纬度升高和年均温度的降低,叶片N和P含量极显著地增加(P《0.001).可是N/P与纬度和年均温度的相关性较弱(P=0.386和P=0.342),这可能是由于N和P含量随纬度和年均温度的变化趋势相同且变异性较大,并且相对于全球尺度而言本研究的区域范围相对较小等多种因素共同导致的. |
| [28] | . Mount Gongga spans 6500 m in elevation and has intact and continuous vertical vegetation belts, ranging from subtropical evergreen broad-leaved vegetation to an alpine frigid sparse grass and desert zone. Investigating the altitudinal trends in leaf nitrogen (N) on Mount Gongga can increase our understanding of the global biogeography of foliar N. In this study, 460 leaf samples from mosses, ferns, and seed plants were collected along an altitudinal gradient on the eastern slope of Mount Gongga, and the variation in leaf N concentration (mass basis) with elevation was analyzed. There are considerable differences in leaf N between mosses and ferns, mosses and seed plants, C4 and C3 plants, and evergreen and deciduous woody plants. The general altitudial pattern of leaf N in Mount Gongga plants was that leaf N kept increasing until an elevation of about 2200 m above sea level, with a corresponding mean annual temperature (MAT) of 8.5 C, and then decreased with increasing elevation. However, the evergreen woody plants displayed a decline trend in leaf N across the altitude gradient. Our findings provide an insight into the altitudinal variation in leaf N. |
| [29] | . Transformations of P along both biological and pedologic pathways are reviewed and integrated in order to develop a comprehensive appreciation of P dynamics in soil ecosystems. The biological P cycle is driven by the essential role P plays in energy transport in biological systems. Mineralization of organic P is influenced by a process referred to as biochemical mineralization wherein hydrolysis of P is repressed by an ample supply of labile P and induced by a limited supply. Pedological P transformations are driven by leaching and increasing acidity which result in the weathering of primary minerals and the formation of secondary minerals. Although the biological cycle is more dynamic than the sluggish pedologic pathway, long-term evolution of P forms in soil ecosystems is controlled by the latter which continually moves P into a sink consisting primarily of occluded P. Occurrence of soluble P as an intermediate during P transformations accommodates translocation of P within soils and landscapes. Because both P content and forms and classes of Soil Taxonomy are related to stage of soil development, a relationship is proposed between taxonomic classes and P forms in soils. |
| [30] | . Whether there is a general allometry law across plant species with different sizes and under different environment has long been controversial and shrubs are particularly useful to examine these questions. |
| [31] | . |
| [32] | . Temperate and boreal forests act as major sinks for atmospheric CO 2 . To assess the magnitude and distribution of the sinks more precisely, an accurate estimation of forest biomass is required. However, the determinants of large-scale biomass pattern (especially root biomass) are still poorly understood for these forests in China. In this study, we used 515 field measurements of biomass across the northeast part of China, to examine factors affecting large-scale biomass pattern and root–shoot biomass allocation. Our results showed that, Picea & Abies forest and coniferous & broadleaf mixed forest had the highest mean biomass (178–20202Mg/ha), while Pinus sylvestris forest the lowest (7802Mg/ha). The root:shoot (R/S) biomass ratio ranged between 0.09 and 0.67 in northeast China, with an average of 0.27. Forest origin (primary/secondary/planted forest) explained 31–37% of variation in biomass (total, shoot and root), while climate explained only 8–15%, reflecting the strong effect of disturbance on forest biomass. Compared with shoot biomass, root biomass was less limited by precipitation as a result of biomass allocation change. R/S ratio was negatively related to water availability, shoot biomass, stand age, height and volume, suggesting significant effects of climate and ontogeny on biomass allocation. Root–shoot biomass relationships also differed significantly between natural and planted forests, and between broadleaf and coniferous forests. Shoot biomass, climate and forest origin were the most important predictors for root biomass, and together explained 83% of the variation. This model provided a better way for estimating root biomass than the R/S ratio method, which predicted root biomass with a R 2 of 0.71. |
| [33] | . . |
| [34] | . ABSTRACT Aim68 Our aim was to quantify climatic influences on key leaf traits and relationships at the global scale. This knowledge provides insight into how plants have adapted to different environmental pressures, and will lead to better calibration of future vegetation–climate models. Location68 The data set represents vegetation from 175 sites around the world. Methods68 For more than 2500 vascular plant species, we compiled data on leaf mass per area (LMA), leaf life span (LL), nitrogen concentration (N mass ) and photosynthetic capacity (A mass ). Site climate was described with several standard indices. Correlation and regression analyses were used for quantifying relationships between single leaf traits and climate. Standardized major axis (SMA) analyses were used for assessing the effect of climate on bivariate relationships between leaf traits. Principal components analysis (PCA) was used to summarize multidimensional trait variation. Results68 At hotter, drier and higher irradiance sites, (1) mean LMA and leaf N per area were higher; (2) average LL was shorter at a given LMA, or the increase in LL was less for a given increase in LMA (LL–LMA relationships became less positive); and (3) A mass was lower at a given N mass , or the increase in A mass was less for a given increase in N mass . Considering all traits simultaneously, 18% of variation along the principal multivariate trait axis was explained by climate. Main conclusions68 Trait-shifts with climate were of sufficient magnitude to have major implications for plant dry mass and nutrient economics, and represent substantial selective pressures associated with adaptation to different climatic regimes. |
| [35] | . |
| [36] | . . |
| [37] | . |
| [38] | . |
| [39] | . . 通过沿海拔梯度的系统调查，运用TWINSPAN分类方法，划分出长白山北坡的主要植物群落类型，对这些群落的组成和结构进行了分析。利用物种丰富度、α多样性和β多样性等指标，研究了群落多样性随海拔梯度的变化趋势。结果表明，乔木层植物可分为4个群落类型：从低海拔到高海拔依次为红松(Pinus koraiensis)针阔混交林、红松针阔叶树种与云冷杉组成的过渡群落、云冷杉暗针叶林以及岳桦林(Betula ermanii)。乔木层优势种重要值的分析清楚地反映出长白山北坡植被垂直带谱的优势成分。径级频度分布的分析表明该区域主要群落的自我更新状况良好。植物物种多样性随海拔梯度的变化趋势为：随海拔升高，乔木层和灌木层的物种丰富度呈下降趋势，但草本层的变化趋势不明显；乔木层和灌木层的α多样性(Shannon-Wiener指数)呈下降趋势，草本层则变化不明显；无论是木本层还是草本层的Pielou均匀度指数均没有明显的变化；乔木层、灌木层和草本层植物的β多样性(Cody指数)均随着海拔的升高而下降。 |
| [40] | . A number of studies have investigated regional and continental scale patterns of carbon (C) stocks in forest ecosystems; however, the altitudinal changes in C storage in different components (vegetation, detritus, and soil) of forest ecosystems remain poorly understood. In this study, we measured C stocks of vegetation, detritus, and soil of 22 forest plots along an altitudinal gradient of 700-2,000 m to quantify altitudinal changes in carbon storage of major forest ecosystems (Pinus koraiensis and broadleaf mixed forest, 700-1,100 m; Picea and Abies forest, 1,100-1,800 m; and Betula ermanii forest, 1,800-2,000 m) on Mt Changbai, Northeast China. Total ecosystem C density (carbon stock per hectare) averaged 237 t C ha(-1) (ranging from 112 to 338 t C ha(-1)) across all the forest stands, of which 153 t C ha(-1) (52-245 t C ha(-1)) was stored in vegetation biomass, 14 t C ha(-1) (2.2-48 t C ha(-1)) in forest detritus (including standing dead trees, fallen trees, and floor material), and 70 t C ha(-1) (35-113 t C ha(-1)) in soil organic matter (1-m depth). Among all the forest types, the lowest vegetation and total C density but the highest soil organic carbon (SOC) density occurred in Betula ermanii forest, whereas the highest detritus C density was observed in Picea and Abies forest. The C density of the three ecosystem components showed distinct altitudinal patterns: with increasing altitude, vegetation C density decreased significantly, detritus C density first increased and then decreased, and SOC density exhibited increasing but insignificant trends. The allocation of total ecosystem C to each component exhibited similar but more significant trends along the altitudinal gradient. Our results suggest that carbon storage and partitioning among different components in temperate forests on Mt Changbai vary greatly with forest type and altitude. |
1
1999
... 本研究中叶片P含量与海拔没有显著的相关关系.陆地生态系统中的P元素不会通过微生物活动进入生态系统, 几乎所有的活性P都来源于岩石风化(
Quantifying the evidence for biodiversity effects on ecosystem functioning and services
1
2006
... 我们使用独立样本t检验比较不同植物功能型(乔木与灌木、针叶与阔叶、常绿与落叶)之间各性状的差异; 采用一般线性模型(GLMs)和混合模型的方差分析检验不同因子对叶片N、P含量和N:P的解释力大小(
The ecology and economics of plants
1
1990
... 长白山地区叶片N:P随海拔的升高而降低.叶片N:P的驱动因子更为复杂.有研究认为物种由于其特有的营养平衡机制会选择其最适N:P (
Leaf nitrogen and phosphorus concentrations of woody plants differ in responses to climate, soil and plant growth form
4
2013
... 本研究中叶片N含量随海拔的增加而降低, 与气温正相关, 与降水负相关, 这与部分海拔梯度上的研究结果(
... 本研究中叶片P含量与海拔没有显著的相关关系.陆地生态系统中的P元素不会通过微生物活动进入生态系统, 几乎所有的活性P都来源于岩石风化(
... 我们的结果表明: 不同功能型植物叶片的N、P含量和N:P差异显著, 灌木物种的叶N含量显著高于乔木, 两者的叶片P含量差异不显著; 阔叶、落叶物种的N、P含量分别显著高于针叶、常绿树种(
... ;
Morphological and physiological adjustment to N and P fertilization in nutrient-limited Metrosideros polymorpha canopy trees in Hawaii
1
2001
... 长白山地区叶片N:P随海拔的升高而降低.叶片N:P的驱动因子更为复杂.有研究认为物种由于其特有的营养平衡机制会选择其最适N:P (
The R Book
2007
Growth rate-stoichiometry couplings in diverse biota
1
2003
... 本研究中叶片N含量随海拔的增加而降低, 与气温正相关, 与降水负相关, 这与部分海拔梯度上的研究结果(
中国草地样带植物氮磷元素空间格局及其与气候因子的关系
1
2014
... 叶片N、P含量受气候、植物功能型、系统发育关系等多种因素的影响(
地理要素对我国气温分布影响的数量评价
1
1992
... T (P) = a + b × Lat + c × Long + d × Alt其中, T和P分别为1-12月的月平均气温和月降水量, a、b、c、d为回归常数, Lat、Long和Alt分别为纬度、经度和海拔(
植物群落清查的主要内容、方法和技术规范
2009
Nutrient limitation in rainforests and cloud forests along a 3,000-m elevation gradient in the Peruvian Andes
2
2013
... 海拔梯度上气候、土壤以及植被物种组成急剧变化, 能够综合反映气温、降水、土壤、植被等环境因子的作用, 使海拔梯度成为研究叶片N、P化学计量特征的理想场所.目前关于叶片N、P含量在海拔梯度上的空间格局尚未得出一致的结论.有的研究表明乔木、灌木、草本三种生活型的植物叶片N含量均随海拔的上升而增加, 并认为高海拔地区的低N供应不影响成熟叶片的代谢活性(
... ;
N:P ratios in terrestrial plants: Variation and functional significance
1
2004
... 氮(N)和磷(P)是植物生长所必需的营养元素, 对植物个体的生长发育起着关键性的作用.由于N和P的养分获得和利用效率往往受到自然环境的限制, N、P成为陆地生态系统中的限制营养元素(
Leaf nitrogen and phosphorus stoichiometry across 753 terrestrial plant species in China
5
2005
... 以往研究对叶片性状空间格局及其机制进行了深入的探讨.一般认为, 在大尺度上气候是影响叶N、P含量的一个因素.如Reich和Oleksyn (2004)在全球尺度上的研究表明, 植物叶片的N、P含量随年平均气温的升高而降低, 而N:P增加.其他研究也证实了气候对叶N、P化学计量特征有显著影响, 但发现气候的解释力往往不高(
... 叶片N、P含量受气候、植物功能型、系统发育关系等多种因素的影响(
... T (P) = a + b × Lat + c × Long + d × Alt其中, T和P分别为1-12月的月平均气温和月降水量, a、b、c、d为回归常数, Lat、Long和Alt分别为纬度、经度和海拔(
... 本研究中叶片N含量随海拔的增加而降低, 与气温正相关, 与降水负相关, 这与部分海拔梯度上的研究结果(
... 我们的结果表明: 不同功能型植物叶片的N、P含量和N:P差异显著, 灌木物种的叶N含量显著高于乔木, 两者的叶片P含量差异不显著; 阔叶、落叶物种的N、P含量分别显著高于针叶、常绿树种(
Biogeography and variability of eleven mineral elements in plant leaves across gradients of climate, soil and plant functional type in China
6
2011
... 叶片N、P含量受气候、植物功能型、系统发育关系等多种因素的影响(
... T (P) = a + b × Lat + c × Long + d × Alt其中, T和P分别为1-12月的月平均气温和月降水量, a、b、c、d为回归常数, Lat、Long和Alt分别为纬度、经度和海拔(
... 本文结果表明, 长白山地区的木本植物叶片N含量、N:P均在海拔梯度上呈现出显著变化, 并且受气温、降水的显著影响, N含量、N:P高的植物出现在气温高、降水量少、环境较为适宜的低海拔地区; 而叶片P含量与海拔和气候因子的关系均不显著(
... 本研究中叶片N含量随海拔的增加而降低, 与气温正相关, 与降水负相关, 这与部分海拔梯度上的研究结果(
... 除了气候等环境因子, 叶片N、P含量和N:P还受植物功能型、物种组成的强烈影响(
... 我们的结果表明: 不同功能型植物叶片的N、P含量和N:P差异显著, 灌木物种的叶N含量显著高于乔木, 两者的叶片P含量差异不显著; 阔叶、落叶物种的N、P含量分别显著高于针叶、常绿树种(
Stoichiometry and large-scale patterns of leaf carbon and nitrogen in the grassland biomes of China
2006
生态化学计量学: 探索从个体到生态系统的统一化理论
8
2010
... 以往研究对叶片性状空间格局及其机制进行了深入的探讨.一般认为, 在大尺度上气候是影响叶N、P含量的一个因素.如Reich和Oleksyn (2004)在全球尺度上的研究表明, 植物叶片的N、P含量随年平均气温的升高而降低, 而N:P增加.其他研究也证实了气候对叶N、P化学计量特征有显著影响, 但发现气候的解释力往往不高(
... ,
... 叶片N、P含量受气候、植物功能型、系统发育关系等多种因素的影响(
... ,
... 为了检验谱系关系对叶片N、P化学计量特征的影响, 我们基于Zanne等(2014)的进化树, 利用R软件中的phylomatic函数, 建立了所采样物种的系统发育树(附录II).为了比较气候和谱系关系的解释力的大小, 我们采用He等(2009)建立的方法, 在系统发育树的90 Mya (million years ago)和60 Mya两个进化时间节点处将物种划分为入不同的谱系组.选择这两个时间节点是因为它们所划分成的谱系组数目(90 Mya 谱系组包括19类, 60 Mya谱系组包括30类), 与传统分类学上的分类数目(19科30属)一致, 便于两种分类方式结果的比较(
...
... 长白山地区木本植物叶片N含量的平均值为21.46 g·kg-1、叶片P含量为2.23 g·kg-1, 叶片N:P为10.81 (
... 关于谱系和气候的相对作用大小, 本文结果表明谱系(90 Mya 谱系组、60 Mya 谱系组、物种)对叶片N、P含量和N:P的作用是最显著的(
Leaf nitrogen:phosphorus stoichiometry across Chinese grassland biomes
6
2008
... 以往研究对叶片性状空间格局及其机制进行了深入的探讨.一般认为, 在大尺度上气候是影响叶N、P含量的一个因素.如Reich和Oleksyn (2004)在全球尺度上的研究表明, 植物叶片的N、P含量随年平均气温的升高而降低, 而N:P增加.其他研究也证实了气候对叶N、P化学计量特征有显著影响, 但发现气候的解释力往往不高(
... 叶片N、P含量受气候、植物功能型、系统发育关系等多种因素的影响(
... 长白山地区叶片N:P随海拔的升高而降低.叶片N:P的驱动因子更为复杂.有研究认为物种由于其特有的营养平衡机制会选择其最适N:P (
... ).这种存储过程可能导致了低的N:P (
... 气候对叶N、P含量和N:P的作用均显著(p < 0.05), 但解释力不高(1.50%-2.98%)(
... ), 并认为气候导致的物种组成和生活型变化是叶性状地理变化的重要驱动力(
Taxonomic identity, phylogeny, climate and soil fertility as drivers of leaf traits across Chinese grassland biomes
2010
Taxonomic, phylogenetic, and environmental trade-offs between leaf productivity and persistence
3
2009
... 本研究中将影响叶片N、P元素变异的因子分为3类: 一是气候因子, 包括MAT、MAP; 二是植物功能型, 包括生活型、叶型、叶片习性; 三是谱系组, 包括90 Mya、60 Mya谱系组和物种.叶片N、P含量和N:P数据在分析前均经log10对数转换, 以提高解释模型残差的正态性(
... 我们使用独立样本t检验比较不同植物功能型(乔木与灌木、针叶与阔叶、常绿与落叶)之间各性状的差异; 采用一般线性模型(GLMs)和混合模型的方差分析检验不同因子对叶片N、P含量和N:P的解释力大小(
... 表示气候和谱系的协同作用; (3)未被解释的变异: 除了气候和谱系外, 其他未在本研究中涉及的因素(
Global organization of terrestrial plant- nutrient interactions
1
2004
... 气候对叶N、P含量和N:P的作用均显著(p < 0.05), 但解释力不高(1.50%-2.98%)(
Toward a stoichiometric framework for evolutionary biology
1
2005
... 长白山地区叶片N:P随海拔的升高而降低.叶片N:P的驱动因子更为复杂.有研究认为物种由于其特有的营养平衡机制会选择其最适N:P (
The vegetation N:P: A new tool to detect the nature of nutrient limitation
1
1996
... 氮(N)和磷(P)是植物生长所必需的营养元素, 对植物个体的生长发育起着关键性的作用.由于N和P的养分获得和利用效率往往受到自然环境的限制, N、P成为陆地生态系统中的限制营养元素(
The nutritional status of plants from high altitudes
1
1989
... 海拔梯度上气候、土壤以及植被物种组成急剧变化, 能够综合反映气温、降水、土壤、植被等环境因子的作用, 使海拔梯度成为研究叶片N、P化学计量特征的理想场所.目前关于叶片N、P含量在海拔梯度上的空间格局尚未得出一致的结论.有的研究表明乔木、灌木、草本三种生活型的植物叶片N含量均随海拔的上升而增加, 并认为高海拔地区的低N供应不影响成熟叶片的代谢活性(
Relative effects of phylogeny, biological characters and environments on leaf traits in shrub biomes across central Inner Mongolia, China
5
2013
... 以往研究对叶片性状空间格局及其机制进行了深入的探讨.一般认为, 在大尺度上气候是影响叶N、P含量的一个因素.如Reich和Oleksyn (2004)在全球尺度上的研究表明, 植物叶片的N、P含量随年平均气温的升高而降低, 而N:P增加.其他研究也证实了气候对叶N、P化学计量特征有显著影响, 但发现气候的解释力往往不高(
... 叶片N、P含量受气候、植物功能型、系统发育关系等多种因素的影响(
... 为了检验谱系关系对叶片N、P化学计量特征的影响, 我们基于Zanne等(2014)的进化树, 利用R软件中的phylomatic函数, 建立了所采样物种的系统发育树(附录II).为了比较气候和谱系关系的解释力的大小, 我们采用He等(2009)建立的方法, 在系统发育树的90 Mya (million years ago)和60 Mya两个进化时间节点处将物种划分为入不同的谱系组.选择这两个时间节点是因为它们所划分成的谱系组数目(90 Mya 谱系组包括19类, 60 Mya谱系组包括30类), 与传统分类学上的分类数目(19科30属)一致, 便于两种分类方式结果的比较(
... 我们使用独立样本t检验比较不同植物功能型(乔木与灌木、针叶与阔叶、常绿与落叶)之间各性状的差异; 采用一般线性模型(GLMs)和混合模型的方差分析检验不同因子对叶片N、P含量和N:P的解释力大小(
... 关于谱系和气候的相对作用大小, 本文结果表明谱系(90 Mya 谱系组、60 Mya 谱系组、物种)对叶片N、P含量和N:P的作用是最显著的(
Testing the growth rate vs. geochemical hypothesis for latitudinal variation in plant nutrients
1
2007
... 本研究中叶片P含量与海拔没有显著的相关关系.陆地生态系统中的P元素不会通过微生物活动进入生态系统, 几乎所有的活性P都来源于岩石风化(
Global patterns of plant leaf N and P in relation to temperature and latitude
6
2004
... 以往研究对叶片性状空间格局及其机制进行了深入的探讨.一般认为, 在大尺度上气候是影响叶N、P含量的一个因素.如Reich和Oleksyn (2004)在全球尺度上的研究表明, 植物叶片的N、P含量随年平均气温的升高而降低, 而N:P增加.其他研究也证实了气候对叶N、P化学计量特征有显著影响, 但发现气候的解释力往往不高(
... 叶片N、P含量受气候、植物功能型、系统发育关系等多种因素的影响(
... T (P) = a + b × Lat + c × Long + d × Alt其中, T和P分别为1-12月的月平均气温和月降水量, a、b、c、d为回归常数, Lat、Long和Alt分别为纬度、经度和海拔(
... 本研究中叶片N含量随海拔的增加而降低, 与气温正相关, 与降水负相关, 这与部分海拔梯度上的研究结果(
... ).本文的结果表明叶N含量在海拔梯度上的变化机制与水平梯度上可能存在差异.高海拔地区的低温和多降水带来的淋溶作用导致N的矿化率较低(
... 除了气候等环境因子, 叶片N、P含量和N:P还受植物功能型、物种组成的强烈影响(
中国东部南北样带654种植物叶片氮和磷的化学计量学特征研究
1
2007
... 以往研究对叶片性状空间格局及其机制进行了深入的探讨.一般认为, 在大尺度上气候是影响叶N、P含量的一个因素.如Reich和Oleksyn (2004)在全球尺度上的研究表明, 植物叶片的N、P含量随年平均气温的升高而降低, 而N:P增加.其他研究也证实了气候对叶N、P化学计量特征有显著影响, 但发现气候的解释力往往不高(
Altitudinal variation in leaf nitrogen concentration on the eastern slope of Mount Gongga on the Tibetan Plateau, China
1
2012
... 海拔梯度上气候、土壤以及植被物种组成急剧变化, 能够综合反映气温、降水、土壤、植被等环境因子的作用, 使海拔梯度成为研究叶片N、P化学计量特征的理想场所.目前关于叶片N、P含量在海拔梯度上的空间格局尚未得出一致的结论.有的研究表明乔木、灌木、草本三种生活型的植物叶片N含量均随海拔的上升而增加, 并认为高海拔地区的低N供应不影响成熟叶片的代谢活性(
Phosphorus dynamics in soils and landscapes
1
1985
... 本研究中叶片P含量与海拔没有显著的相关关系.陆地生态系统中的P元素不会通过微生物活动进入生态系统, 几乎所有的活性P都来源于岩石风化(
Effects of biophysical constraints, climate and phylogeny on forest shrub allometries along an altitudinal gradient in Northeast China
1
2017
... 为了检验谱系关系对叶片N、P化学计量特征的影响, 我们基于Zanne等(2014)的进化树, 利用R软件中的phylomatic函数, 建立了所采样物种的系统发育树(附录II).为了比较气候和谱系关系的解释力的大小, 我们采用He等(2009)建立的方法, 在系统发育树的90 Mya (million years ago)和60 Mya两个进化时间节点处将物种划分为入不同的谱系组.选择这两个时间节点是因为它们所划分成的谱系组数目(90 Mya 谱系组包括19类, 60 Mya谱系组包括30类), 与传统分类学上的分类数目(19科30属)一致, 便于两种分类方式结果的比较(
Altitudinal variation in leaf mass per unit area, leaf tissue density and foliar nitrogen and phosphorus content along an Amazon-Andes gradient in Peru
2
2009
... 海拔梯度上气候、土壤以及植被物种组成急剧变化, 能够综合反映气温、降水、土壤、植被等环境因子的作用, 使海拔梯度成为研究叶片N、P化学计量特征的理想场所.目前关于叶片N、P含量在海拔梯度上的空间格局尚未得出一致的结论.有的研究表明乔木、灌木、草本三种生活型的植物叶片N含量均随海拔的上升而增加, 并认为高海拔地区的低N供应不影响成熟叶片的代谢活性(
... 本研究中叶片N含量随海拔的增加而降低, 与气温正相关, 与降水负相关, 这与部分海拔梯度上的研究结果(
Forest biomass and root-shoot allocation in northeast China
2008
长白山北坡主要森林类型及其群落结构特点
1
1980
... 长白山地区属温带大陆性山地气候, 随着海拔的升高(海拔500-2 744 m), 年降水量逐渐增加(720-1 400 mm), 而年平均气温从3.5 ℃下降到-7.4 ℃.长白山海拔梯度上气候条件的巨大差异, 使长白山地区的植被具有典型的垂直结构, 由下至上依次形成了阔叶红松混交林(Pinus koraiensis and broadleaf mixed forest, 海拔500-1 100 m)、云冷杉林(Picea and Abies forest, 海拔1 100-1 800 m)、亚高山岳桦林带(Betula ermanii forest, 海拔1 800-2 000 m)和高山苔原带(alpine tundra, 海拔2 000 m以上) 4个植被垂直带(
Modulation of leaf economic traits and trait relationships by climate
1
2005
... 气候对叶N、P含量和N:P的作用均显著(p < 0.05), 但解释力不高(1.50%-2.98%)(
The worldwide leaf economics spectrum
1
2004
... 气候对叶N、P含量和N:P的作用均显著(p < 0.05), 但解释力不高(1.50%-2.98%)(
珠江三角洲3种典型森林类型乔木叶片生态化学计量学
1
2010
... 以往研究对叶片性状空间格局及其机制进行了深入的探讨.一般认为, 在大尺度上气候是影响叶N、P含量的一个因素.如Reich和Oleksyn (2004)在全球尺度上的研究表明, 植物叶片的N、P含量随年平均气温的升高而降低, 而N:P增加.其他研究也证实了气候对叶N、P化学计量特征有显著影响, 但发现气候的解释力往往不高(
Three keys to the radiation of angiosperms into freezing environments
2014
The altitudinal patterns of leaf C:N:P stoichiometry are regulated by plant growth form, climate and soil on Changbai Mountain, China
2
2014
... 本研究中叶片N含量随海拔的增加而降低, 与气温正相关, 与降水负相关, 这与部分海拔梯度上的研究结果(
... ;
长白山北坡植物群落组成、结构及物种多样性的垂直分布
1
2004
... 长白山(41.38°-42.60° N, 126.92°-129.00° E), 位于我国吉林省东南部, 与朝鲜相邻, 是欧亚大陆东岸的最高山, 也是东北植物区系唯一拥有明显植被垂直带谱的山脉(
Altitudinal changes in carbon storage of temperate forests on Mt. Changbai, Northeast China
1
2010
... 长白山地区属温带大陆性山地气候, 随着海拔的升高(海拔500-2 744 m), 年降水量逐渐增加(720-1 400 mm), 而年平均气温从3.5 ℃下降到-7.4 ℃.长白山海拔梯度上气候条件的巨大差异, 使长白山地区的植被具有典型的垂直结构, 由下至上依次形成了阔叶红松混交林(Pinus koraiensis and broadleaf mixed forest, 海拔500-1 100 m)、云冷杉林(Picea and Abies forest, 海拔1 100-1 800 m)、亚高山岳桦林带(Betula ermanii forest, 海拔1 800-2 000 m)和高山苔原带(alpine tundra, 海拔2 000 m以上) 4个植被垂直带(
