Branching patterns of clonal root of Populus euphratica and its associations with soil factors
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摘要:目的胡杨根系克隆繁殖对胡杨林更新及其群落维持具有重要作用，而胡杨根克隆器官——繁殖根系的扩展分枝是实现克隆繁殖的重要生态过程。本文以明确胡杨繁殖根系分枝特征(节间长、分支强度、分支垂向和水平向夹度)在异质生境下的可塑性和主要分枝类型，探究胡杨繁殖根系特征与土壤因子的关系，找到影响胡杨繁殖根系变化的关键环境因子。方法采用典型抽样对16个样点的一段跨度10 m以上的繁殖根分枝以及对应的土壤因子进行挖掘和调查。运用相关性分析(CA)和主成分分析(PCA)方法分析繁殖根特征之间的相关性和分布规律。运用冗余分析(RDA)和方差分解探究根分枝特征与土壤因子的关联性。结果(1) CA和PCA结果显示，4个分枝特征之间的相关性较高(P＜0.05)，PCA第1轴解释了繁殖根特征变异的66.88%，基本代表了从占据型(分枝强、短、垂向角小、水平角大)到游击型(分枝弱、长、垂向角大、水平角小)两种极端分枝类型的变化。占据型分枝更多出现在林缘，而游击型分枝更多出现在靠近河道处。(2)RDA结果显示，垂向分枝夹角主要受土壤紧实度的正影响(R2=0.64，P＜0.05)，节间长度主要受紧实度的负向影响(R2=0.87，P＜0.05)。土壤全碳和全氮主要对分枝强度和分枝水平夹角有正向作用，但土壤含砂量对它们具有负向作用。主轴分布深度主要受毛管水埋深影响(R2=0.62，P＜0.05)。(3)方差分解结果显示，三类土壤特征共解释了胡杨分枝特征变异的68.3%，土壤物理特征(土壤质地、紧实度)的独立作用最大(R2=0.12，P＜0.05)，土壤水分(土壤含盐量、含水量)和养分特征(全N、C)的独立作用较小(R2=0.07，P＜0.05；R2=0.03)，但与土壤物理特征的共同作用较大(R2=0.23, P＜0.05)。结论(1) 在异质性的河漫滩环境中，胡杨繁殖根系分枝在两种极端分枝类型——游击型和占据型分枝形成的连续谱之间变化。(2)土壤物理特征是影响胡杨繁殖根分枝的最关键因素，胡杨繁殖根系主要选择占据土壤质地良好，紧实度小的斑块。单纯水分或养分丰富的斑块不会引起胡杨繁殖根分枝明显变异，但是在土壤质地和紧实度良好的土壤斑块中，水分和养分的变化会对繁殖根分枝产生显著影响。(3)胡杨繁殖根的垂向分枝夹角具有明显的生态塑性，是胡杨克隆器官的重要适应特征。Abstract:ObjectiveThe clonal growth of Populus euphratica plays an important role to the recruitment and persistence of the population in arid-climate region, and the change in branching pattern of clonal roots is a key process for P. euphratica to achieve the clonal growth in heterogeneous environment. This paper attempts to clarify the branching pattern of clonal root system of P. euphratica in heterogeneous riparian habitat, to estimate the correlations between branching traits of clonal roots including internal length (IL), branching intensity (BI), vertical-and horizontal-branching angle (VBA and HBA), and soil factors, and then to find the key factors driving changes of the branching pattern.MethodWe investigated 16 clonal root branches over 10 m by manual digging and collected below-ground environmental factors correspondingly in a natural floodplain of Ejin Oasis, Inner Mongolia of northern China. Correlation and ordination of the branching traits of clonal root were analyzed using correlation analysis (CA) and principal component analysis (PCA). Redundancy analysis (RDA) and variation partitioning (VP) were applied to explore the relationship between root branching traits and soil factors.Result(1) The results of CA and PCA showed that there was a high correlation between the four branching traits (P < 0.05), and the first axis, reflecting 66.88% variation of the branching traits, can represent the changes from guerrilla-branching type (with sparser and longer branches, smaller branching angle in vertical and horizontal) to occupation-branching type (with denser and shorter branches, steeper branching angle in vertical and wider branching angle in horizontal). The former more occur on the edge of the river, and the later more on the edge of the forest. (2) The results of RDA revealed that VBA was mainly affected by soil compactness positively (R2=0.64, P < 0.05), and IL was mainly affected by it negatively (R2=0.87, P < 0.05). Soil total carbon and total nitrogen had a major positive influence on BI and HBA, but soil sandy content had a negative effects on them. The depth of branching points of clonal root was positively controlled by capillary water depth (R2=0.62, P < 0.05). (3) VP results showed that the three set of soil variables including soil nutrient-related, water-related and physical factors explained a total of 68.3% of the variance of P. euphratica branching traits. The independent effects of soil physical factors (soil texture, compactness) were the highest (R2=0.12, P < 0.05). Nutrient- and water-related factors had relative smaller independent effects (R2=0.07, P < 0.05; R2=0.03, respectively), but they had a greatest together influences with soil physical factors to variance of the branching traits (R2=0.23, P < 0.05).Conclusion(1) Guerrilla-branching type and occupation-branching type are two main branching pattern of clonal roots for P. euphratica to vegetative propagate in heterogeneous floodplain of arid region. (2) Soil physical factor is the most important factor to drive changes in branching pattern of the clonal roots. P. euphratica tends to develop an occupation-branching clonal roots in soil patches with less sandy and smaller compactness to establish its ramets. (3) The vertical branching angle of clonal roots of P. euphratica expresses a significant ecological plasticity, it may be an important adaptive trait of clonal roots for woody plant that should be paid attention in the future researches of clonal plant.
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图 1 距河道不同距离样点繁殖根特征的主成分分析
字母A~E分别代表 5个不同取样带的样点，数字1~4表示距河道远近，1最近、4最远，如A1表示A样带距河道最近的样点。
Figure 1. Principal component analysis (PCA) for clonal root traits in 16 samples of different distance to the river
Letters A to E represent different transects, and numbers 1 to 4 represent samples in different distances to the river, in which 1 represents the farthest and 4 represents the nearest.
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图 2 繁殖根特征和土壤因子的冗余分析
实线向量为土壤因子，虚线向量为根特征。il.节间长; mrd.主根分布深度; vba.垂向分枝夹角; bi.分枝强度; hba.水平向分枝夹角; SC.土壤紧实度; SdR.土壤砂粒百分含量; CWD.毛管水埋深; TOC.土壤全碳含量; TON.土壤全氮含量; WC.土壤质量含水量; SS.土壤含盐量。*表示该因子为某根特征的主要影响因子(P＜0.05)。
Figure 2. Redundancy analyses (RDAs) calculated from the six clonal root traits of P. euphratica and the below-ground environmental factors
Bold line vector represents soil factor, dashed line vector represents root trait. il, internal length; bi, branch intensity; hba, horizontal branch angle; vba, vertical branch angle; mrd; axis depth; CWD, soil capillary water depth; WC, soil mass water content; SS, soil salinity; SdR, the ratio of sand; SC, soil compactness; TOC, total C; TON, total N; * represents a major influence factor (P < 0.05).
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图 3 方差分解分析土壤物理性质、土壤养分和土壤水分可利用性对根特征影响的纯效应和共同效应
*表示因子对根特征的独立作用具有显著性(*, P＜0.05);f.部分解释率小于0，不显示值。
Figure 3. Individual and shared effects of soil physical properties, nutrient and water availability on the variance of branching traits for clonal root
* denotes that the individual effect is statistical significance (*, P < 0.05); the value of f. less than 0 is null.
表 1 距河道不同距离样点的土壤理化性质和根特征描述统计和方差分析
Table 1 Summary and comparison of soil properties and clonal root traits for samples of different distance to the rive
项目
Item距河道不同距离样点
Samples of different distance to riverway远(n=5)
Far(n=5)中(n=5)
Middle(n=5)近(n=6)
Close(n=6)土壤因子
Soil factor土壤水分可利
用性 Soil water
availability毛管水埋深 CWD/cm 106.4±6.2 97.8±3.4 93.5±1.3 含水量 WC/% 22.3±1.7 21.1±1.9 22.8±1.8 含盐量 SS/ % 3.6±0.9 3.5±0.7 2.7±0.9 土壤物理性质
Soil physical
property土壤中值粒径 D0.5/μm 74.0±13.6 b 131.6±17.0 ab 200.1±44.1 a 砂粒含量 SdR/% 61.9±6.4 c 85.3±4.5 b 89.3±4.9 a 粉粒含量 SlR/% 34.7±5.8 a 13.6±4.2 b 9.9±4.4 c 粘粒含量 ClR/% 3.3±0.58 a 1.1±0.35 b 0.8±0.49 c 土壤密度 BD/(g·cm-3) 1.51±0.02 b 1.50±0.01 b 1.54±0.03 a 紧实度 SC/(kg·cm-2) 1.4±0.31 1.1±0.19 0.6±0.10 土壤养分
Soil nutrient全碳量 TOC/(g·kg-1) 6.66±1.20 5.56±1.30 3.79±0.42 全氮量 TON/(g·kg-1) 0.58±0.081 a 0.42±0.044 b 0.34±0.032 b 全磷量 TOP/(g·kg-1) 0.202±0.025 0.181±0.037 0.195±0.032 繁殖根特征
Clonal root traits节间长 il/m 1.66±0.18 b 2.24±0.21 b 3.14±0.25 a 分支强度 bi 1.16±0.15 a 1.09±0.04 a 0.82±0.09 b 水平分枝角度 hba (正弦值Sine value) 0.531±0.024 0.506±0.022 0.486±0.018 垂直分枝角度 vba (正弦值Sine value) 0.118±0.018 a 0.094±0.009 a 0.063±0.001 b 主轴分布深度 mrd/cm 88.8±10.6 82.0±4.9 69.0±5.6 注：CWD.毛管水埋深; WC.土壤质量含水量; SS.土壤含盐量; D0.5.土壤中值粒径; SdR.土壤砂粒百分含量; SlR.土壤粉粒百分含量; ClR.土壤粘粒百分含量; BD.土壤密度; SC.紧实度; TOC.土壤全碳含量; TON.土壤全氮含量; TOP.土壤全磷含量; il.平均节间长; bi.分枝强度; hba.水平向分枝角度; vba.垂向分枝角度; mrd.主根分布深度。同行中的不同小写字母表示不同距离样点差异显著(P＜0.05)。Notes: CWD, soil capillary water depth; WC, soil mass water content; SS, soil salinity; D0.5, the mid-value of grain diameter; SdR, the ratio of sand; SlR, the ratio of silt; ClR, the ratio of clay; BD, soil bulk density; SC, soil compactness; TOC, soil total C content; TON, soil total N content; TOP, soil total P content; il, mean internal length; bi, branch intensity; hba, horizontal branch angle; vba, vertical branch angle; mrd, axis depth. Different letters in a same row denote significant difference between samples in different distances (P＜0.05). 
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表 2 成对根特征之间的Pearson相关性系数和显著性检验
Table 2 Coefficients of Pearson's correlation for pairwise clonal root traits with original dat
变量 Variable 节间长 il 水平分支角度 hba 分支强度 bi 垂向分支角度 vba 水平分支角度 hba -0.662** 分支强度 bi -0.844** 0.629** 垂向分支角度 vba -0.786** 0.531** 0.787** 主轴分布深度 mrd -0.401 0.486* 0.474 0.297 注：*表示根特征之间的相关性显著(*, P＜0.05;**, P < 0.01)。Notes: correlations were significant: **, P < 0.01; *, P＜0.05.
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表 3 根特征前两轴的载荷得分
Table 3 Loadings scores of root variables in the first two axe
变量
Variable第1轴
PCA1第2轴
PCA2节间长 il -0.913 0.215 分枝强度 bi 0.931 -0.048 垂向分枝角度 vba 0.917 -0.340 水平分枝角度 hba 0.803 0.266 主轴分布深度 mrd 0.593 0.827
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