Allocation characteristics and prediction models of water storage capacity among different tree species in Jiaohe, Jilin Province of northeastern China
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摘要:目的
分析吉林蛟河12个乔木树种整株及各组分含水率和储水量分配特征，构建并筛选各树种最优储水量预测模型，探讨不同树种储水量随森林发育阶段的变化，为该地区森林树种储水量估算提供模型参考。
方法采用单因素方差分析，对比12个树种不同器官含水率和储水量占比的差异，并通过多重比较法进行显著性检验。应用肯德尔秩相关分析法，以胸径（D）、树高（H）、D2H为模型自变量，整株及各器官储水量为因变量，构建多种形式的储水量预测模型，并通过模型决定系数、参数显著性以及赤池信息准则筛选最优模型。结合林地信息，计算不同发育阶段树种的储水量。
结果（1）12个树种器官平均含水率顺序为树叶 > 树根 > 树枝 > 树干。除千金榆外，其余树种在各器官储水量分配上普遍呈现树干 > 树根 > 树枝 > 树叶的趋势。随胸径增大，树枝储水量占比增大，而树干与树叶的储水量占比减小，树根储水量变化不显著。（2）12个树种的储水量预测模型均为对数函数形式，不同树种器官的最优模型自变量各异。（3）随着森林演替，单位面积乔木的储水量和生物量均增加。
结论本研究揭示了储水量占比和含水率在器官间与物种间存在显著差异，其中储水量与树高、胸径之间存在种间特异性，且不同器官储水量占比随胸径增长呈现不同变化趋势。所筛选的储水量最优模型均为对数函数形式，其中单树种储水量预测模型具有较高的拟合精度，而全树种模型更适用于估算区域性储水量。本文阐明了吉林蛟河树木水分状况在不同时空尺度上的变化规律，有助于加深对生态系统动态过程的理解，并为该地区森林树种储水量的精确估算提供了可靠的模型参考。
Abstract:ObjectiveThis paper analyzed the distribution characteristics of moisture contents of 12 tree species in northeastern China and species-specific allometric equations of 12 tree species were established to explore the differences in water storage capacity characteristics among different tree species with forest developing, as well as providing model reference for the estimation of water storage capacity in this area.
MethodOne-way ANOVA and multiple comparison methods were used to contrast differences in moisture content and water storage capacity proportion among various organs across the 12 tree species. Utilizing Kendall’s rank correlation analysis to identify DBH (D), tree height (H), and D2H as predictor variables in water storage capacity prediction models with whole-tree and organ-specific water storage capacity serving as response variables. Different forms of water storage capacity prediction models were constructed based on these relationships. Optimal models were selected through evaluation using the coefficient of determination, parameter significance level, and Akaike’s information criterion. Integrating stand information, this approach was employed to calculate the water storage capacity of trees across varied developing stages.
Result(1) Overall, average moisture content was highest in leaves, followed by roots, branches, and stems. Except for Carpinus cordata, all other species showed a consistent pattern in water allocation across organs: stem > root > branch > leaf. As D increased, the proportion of branch water storage capacity increased, while the proportion of stem and leaf water storage capacity decreased, with no significant changes in root water storage capacity. (2) The water storage capacity prediction models for all 12 tree species were best represented by logarithmic functions. The optimal independent variables for organ moisture content models of different tree species were different. (3) With forest succession, both water storage capacity and biomass per unit area increased.
ConclusionThe study highlights significant differences in water storage capacity and distribution among organs and tree species, with species-specific relationship between water storage capacity and D, as well as H. The percentage of water storage capacity of different organs shows different trends with the increase of breast diameter. The water content prediction models for all 12 tree species were best represented by logarithmic functions. The single-species models have higher fitting accuracy, while the multi-species model has broader application. This research elucidates the spatiotemporal dynamics of water status in temperate-boreal tree species, contributing to a deeper understanding of ecosystem dynamics. It provides a scientific basis for accurate estimation of tree water storage capacity in the forest region of Jiaohe, Jilin Province of northeastern China.
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图 1 不同树种器官含水率分布格局
不同小写字母表示不同树种的同一器官间含水率差异显著（P < 0.05）。下同。Different lowercase letters indicate significant differences in moisture content between the same organ of varied tree species (P < 0.05). The same below.
Figure 1. Distribution patterns of moisture content in organs of different tree species
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图 2 不同树种各器官储水量的分配格局
Figure 2. Allocation patterns of water storage capacity in organs of different tree species
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图 3 树木各器官储水量占比随胸径变化的分配格局
Figure 3. Allocation patterns of water storage capacity in organs with DBH
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图 4 12个树种整株（a）、树叶（b）、树枝（c）、树干（d）、树根（e）及地上部分（f）储水量随胸径变化的分配格局
Figure 4. Allocation patterns of water storage capacity of whole plant (a), leaf (b), branch (c), stem (d), root (e) and aboveground (f) of 12 tree species with DBH
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图 5 12个树种整株（a）、树叶（b）、树枝（c）、树干（d）、树根（e）及地上部分（f）储水量随树高变化的分配格局
Figure 5. Allocation patterns of water storage capacity of whole plant (a), leaf (b), branch (c), stem (d), root (e) and aboveground (f) of 12 tree species with H
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图 6 12个树种整株（a）、树叶（b）、树枝（c）、树干（d）、树根（e）及地上部分（f）储水量随胸径树高组合变化的分配格局
Figure 6. Allocation patterns of water storage capacity of whole plant (a), leaf (b), branch (c), stem (d), root (e) and aboveground (f) of 12 tree species with D2H
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图 7 不同发育阶段林地树种储水量的分布格局
Figure 7. Distribution patterns of water storage capacity of tree species at different developing stages
表 1 不同发育阶段林分树种组成及相关指标
Table 1 Composition of tree species and related indicators in different developing stages of stand
林分
Stand胸高断面积和物种重要值
Basal area and species important value平均胸径
Average DBH/cm平均树高
Average
tree height/m密度/（株·hm−2）
Density/
(tree·ha−1)树种
Tree species胸高断面积/（m2·hm−2）
Basal area/(m2·ha−1)重要值
Important
value中龄林
Middle-aged forest胡桃楸 Juglans mandshurica 11.35 25.12 9.20 10.52 1 405 东北槭 Acer mandshuricum 1.95 15.58 五角槭Acer pictum subsp. mono 2.06 12.99 裂叶榆 Ulmus laciniata 1.05 8.58 春榆 Ulmus japonica 1.34 8.40 近熟林
Near-mature forest水曲柳 Fraxinus mandshurica 6.52 11.58 9.52 10.85 1 539 五角槭 Acer pictum subsp. mono 4.61 11.07 红松 Pinus koraiensis 3.42 7.05 紫椴 Tilia amurensis 4.29 6.52 春榆 Ulmus japonica 1.16 6.07 成熟林
Mature forest五角槭 Acer pictum subsp. mono 5.91 8.86 10.87 14.38 1 004 红松 Pinus koraiensis 4.91 7.41 紫椴 Tilia amurensis 4.15 6.52 水曲柳 Fraxinus mandshurica 4.24 6.26 东北槭 Acer mandshuricum 2.29 5.19 老龄林
Old growth forest千金榆 Carpinus cordata 1.70 10.83 11.20 15.43 980 五角槭 Acer pictum subsp. mono 4.03 10.63 裂叶榆 Ulmus laciniata 4.48 9.56 东北槭 Acer mandshuricum 1.91 8.98 紫椴 Tilia amurensis 4.92 7.90 注：表中仅列出各发育阶段林分中物种重要值排名前5位的树种。引自文献[21−22]。Notes: the table only lists the top 5 tree species with the highest species importance values in each developing stage of stand. Cited from reference [21−22]. 
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表 2 12个优势树种样本数量和测定指标
Table 2 Sample size and measurement indicators of 12 dominant tree species
树种
Tree species样本数
Sample size胸径
DBH/cm树高
Tree height/m整株储水量
Water storage capacity of whole plant/kg白桦 Betula platyphylla 10 5.7 ~ 40.0 9.3 ~ 22.8 8.34 ~ 1 027.83 东北槭 Acer mandshuricum 10 7.8 ~ 35.9 9.1 ~ 18.5 18.07 ~ 725.20 春榆 Ulmus japonica 9 5.6 ~ 39.9 6.8 ~ 20.1 10.17 ~ 1 343.24 红松 Pinus koraiensis 11 8.4 ~ 44.0 6.7 ~ 22.3 14.77 ~ 1 132.20 胡桃楸 Juglans mandshurica 10 6.5 ~ 42.5 8.2 ~ 23.0 15.03 ~ 1 169.22 朝鲜槐 Maackia amurensis 9 4.9 ~ 25.4 7.0 ~ 18.2 7.73 ~ 391.08 蒙古栎 Quercus mongolica 9 8.0 ~ 41.2 8.4 ~ 22.8 16.83 ~ 1 036.42 千金榆 Carpinus cordata 9 5.1 ~ 13.4 7.9 ~ 11.9 6.04 ~ 67.91 青杨 Populus ussuriensis 10 9.1 ~ 47.1 10.5 ~ 26.4 20.78 ~ 1 329.98 五角槭 Acer pictum subsp. mono 12 6.4 ~ 45.3 8.5 ~ 20.6 10.72 ~ 1 105.85 水曲柳 Fraxinus mandshurica 10 10.7 ~ 41.4 10.9 ~ 23.7 16.21 ~ 1 371.45 紫椴 Tilia amurensis 10 7.0 ~ 42.2 9.6 ~ 22.5 10.57 ~ 950.29
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表 3 储水量方程式
Table 3 Water storage capacity equation
方程代码
Equation code方程类型
Equation type方程形式
Equation form方程代码
Equation code方程类型
Equation type方程形式
Equation formEq.1 一元线性 Univariate linear y = aD + b Eq.8 多元二次 Multivariate quadratic y = aD2 + bH + c Eq.2 一元线性 Univariate linear y = aH + b Eq.9 多元二次 Multivariate quadratic y = aD + bH2 + c Eq.3 多元线性 Multivariate linear y = aD + bH + c Eq.10 多元二次 Multivariate quadratic y = aD2 + bH2 + c Eq.4 一元二次 Quadratic y = aD2 + b Eq.11 对数 Logarithmic ln y = ln a + bln D Eq.5 一元二次 Quadratic y = aH2 + b Eq.12 对数 Logarithmic ln y = ln a + bln H Eq.6 一元二次 Quadratic y = aD2 + bD + c Eq.13 对数 Logarithmic ln y = ln a + bln（D2H） Eq.7 一元二次 Quadratic y = aH2 + bH + c Eq.14 对数 Logarithmic ln y = ln a + bln D + cln H 注：y代表组分储水量（kg），包括整株、叶、茎干、根和地上储水量。H表示树高（m），D表示胸径（cm）。a、b、c为回归方程参数。下同。Notes: y, water storage capacity of component (kg), including leaf water storage capacity, stem water storage capacity, root water storage capacity, aboveground water storage capacity and water storage capacity of whole plant. H, tree height (m); D, diameter at breast height (cm). a, b, c are regression equation parameters, respectively. The same below.
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表 4 肯德尔秩相关性分析
Table 4 Kendall_tau correlation analysis
储水量
Water storage capacityD H D2H 冠幅
Crown width整株 Whole plant 0.922*** 0.709*** 0.910*** 0.213 叶 Leaf 0.753*** 0.583*** 0.728*** 0.109 枝 Branch 0.802*** 0.609*** 0.777*** 0.138 干 Stem 0.881*** 0.733*** 0.889*** 0.232 根 Root 0.881*** 0.674*** 0.859*** 0.156 注：***表示极显著相关（P < 0.001）。 Note: *** indicates extremely significant correlation (P < 0.001).
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表 5 各组分储水量最优方程形式
Table 5 Optimal equation forms for the water storage capacity of each component
树种 Tree species 组分 Component 方程形式 Equation form R2 P AIC 白桦 Betula platyphylla 整株 Whole plant ln y = 1.042ln（D2H） − 3.965 0.98 < 0.01 −4.140 叶 Leaf ln y = 1.031ln（D2H） − 7.353 0.90 < 0.01 18.269 枝 Branch ln y = 1.441ln（D2H） − 9.520 0.92 < 0.01 22.886 干 Stem ln y = 0.939ln（D2H） − 3.631 0.99 < 0.01 −18.812 根 Root ln y = 2.780ln D − 4.467 0.98 < 0.01 5.694 地上 Aboveground ln y = 1.014ln（D2H） − 3.993 0.99 < 0.01 −9.207 东北槭 Acer mandshuricum 整株 Whole plant ln y = 2.425ln D − 2.101 0.99 < 0.01 −12.626 叶 Leaf ln y = 2.686ln D − 2.832ln H + 1.28 0.76 < 0.01 15.751 枝 Branch ln y = 3.914ln D − 2.367ln H − 1.611 0.94 < 0.01 11.654 干 Stem ln y = 0.955ln（D2H） − 3.981 0.99 < 0.01 −11.171 根 Root ln y = 2.303ln D − 3.010 0.98 < 0.01 −3.235 地上 Aboveground ln y = 2.471ln D − 2.567 0.99 < 0.01 −13.765 春榆 Ulmus japonica 整株 Whole plant ln y = 2.446ln D − 2.200 0.98 < 0.01 1.746 叶 Leaf ln y = 2.308ln D − 4.717 0.98 < 0.01 3.353 枝 Branch ln y = 4.033ln D − 2.317ln H − 1.967 0.97 < 0.01 12.564 干 Stem ln y = 0.948ln（D2H） − 3.571 0.96 < 0.01 9.203 根 Root ln y = 2.633ln D − 3.786 0.98 < 0.01 2.582 地上 Aboveground ln y = 2.487ln D − 2.318 0.98 < 0.01 4.405 红松 Pinus koraiensis 整株 Whole plant ln y = 0.984ln（D2H） − 3.377 0.97 < 0.01 6.370 叶 Leaf ln y = 0.8497ln（D2H） − 4.8269 0.91 < 0.01 15.640 枝 Branch ln y = 2.652ln D − 4.920 0.86 < 0.01 24.353 干 Stem ln y = 0.937ln D + 2.698ln H − 5.283 0.99 < 0.01 −0.603 根 Root ln y = 0.952ln（D2H） − 4.663 0.96 < 0.01 9.206 地上 Aboveground ln y = 0.992ln（D2H） − 3.690 0.97 < 0.01 6.555 胡桃楸 Juglans mandshurica 整株 Whole plant ln y = 2.6679ln D − 0.5497ln H − 1.201 3 0.99 < 0.01 −11.226 叶 Leaf ln y = 3.2676ln D − 2.73ln H + 0.395 6 0.97 < 0.01 1.119 枝 Branch ln y = 4.861 8ln D − 3.8021ln H − 0.1967 0.93 < 0.01 19.893 干 Stem ln y = 0.893 9ln（D2H） − 3.407 6 0.99 < 0.01 −13.070 根 Root ln y = 2.543ln D − 3.949 0.97 < 0.01 5.653 地上 Aboveground ln y = 2.635ln D − 0.558ln H − 1.326 0.99 < 0.01 −12.311 朝鲜槐 Maackia amurensis 整株 Whole plant ln y = 2.328ln D − 1.812 0.98 < 0.01 −1.580 叶 Leaf ln y = 1.502ln D − 3.100 0.78 < 0.01 14.688 枝 Branch ln y = 2.952ln D − 5.301 0.86 < 0.01 22.856 干 Stem ln y = 0.855 5ln（D2H） − 3.021 7 0.99 < 0.01 −12.734 根 Root ln y = 2.391ln D − 3.456 0.97 < 0.01 2.387 地上 Aboveground ln y = 0.904 5ln（D2H） − 3.072 8 0.97 < 0.01 0.724 千金榆 Carpinus cordata 整株 Whole plant ln y = 2.389ln D − 2.072 0.97 < 0.01 −7.642 叶 Leaf ln y = 2.917 1ln D-2.309 7ln H − 0.344 3 0.86 < 0.01 7.819 枝 Branch ln y = 3.476ln D − 5.862 0.93 < 0.01 7.884 干 Stem ln y = 1.936ln D − 1.842 0.95 < 0.01 −6.045 根 Root ln y = 1.006ln（D2H） − 5.366 0.78 < 0.01 12.272 地上 Aboveground ln y = 2.393ln D − 2.267 0.98 < 0.01 −10.507 青杨 Populus ussuriensis 整株 Whole plant ln y = 1.007ln（D2H） − 3.776 0.99 < 0.01 −26.517 叶 Leaf ln y = 2.85ln D − 1.89ln H − 1.575 0.89 < 0.01 14.203 枝 Branch ln y = 3.895ln D − 2.074ln H − 2.540 0.96 < 0.01 10.814 干 Stem ln y = 1.557ln D + 1.900ln H − 5.451 0.99 < 0.01 −21.472 根 Root ln y = 2.469ln D − 3.831 0.98 < 0.01 −2.403 地上 Aboveground ln y = 1.004ln（D2H） − 3.974 0.99 < 0.01 −27.189 五角槭 Acer pictum subsp. mono 整株 Whole plant ln y = 0.979 7ln（D2H） − 3.492 4 0.99 < 0.01 −10.945 叶 Leaf ln y = 0.726 6ln（D2H） − 4.598 7 0.90 < 0.01 11.838 枝 Branch ln y = 1.132ln（D2H） − 6.653 0.91 < 0.01 21.919 干 Stem ln y = 0.937 8ln（D2H） − 3.824 7 0.99 < 0.01 −11.027 根 Root ln y = 2.535ln D − 3.801 0.98 < 0.01 −1.232 地上 Aboveground ln y = 0.955 6ln（D2H） − 3.616 5 0.99 < 0.01 −11.835 蒙古栎 Quercus mongolica 整株 Whole plant ln y = 2.155 0ln D + 0.572 3ln H − 2.847 4 0.99 < 0.01 −16.751 叶 Leaf ln y = 2.201 92ln D − 0.063 17ln H − 4.740 38 0.98 < 0.01 −2.433 枝 Branch ln y = 4.576ln D − 1.727ln H − 5.735 0.93 < 0.01 18.667 干 Stem ln y = 1.640ln D + 1.248ln H − 3.715 0.99 < 0.01 −12.174 根 Root ln y = 2.315ln D − 3.337 0.98 < 0.01 −2.664 地上 Aboveground ln y = 0.9912ln（D2H） − 3.737 9 0.99 < 0.01 −16.113 水曲柳 Fraxinus mandshurica 整株 Whole plant ln y = 2.865ln D − 3.470 0.97 < 0.01 3.883 叶 Leaf ln y = 0.785 2ln（D2H） − 4.784 3 0.91 < 0.01 8.269 枝 Branch ln y = 1.438ln（D2H） − 9.785 0.97 < 0.01 8.782 干 Stem ln y = 3.173ln D − 5.147 0.86 < 0.01 22.814 根 Root ln y = 3.09ln D − 0.991 1ln H − 2.552 4 0.99 < 0.01 −8.244 地上 Aboveground ln y = 2.985ln D − 4.186 0.96 < 0.01 8.710 紫椴 Tilia amurensis 整株 Whole plant ln y = 0.999 5ln（D2H） − 3.737 4 0.99 < 0.01 −14.284 叶 Leaf ln y = 0.913 9ln（D2H） − 6.460 3 0.93 < 0.01 12.766 枝 Branch ln y = 3.698ln D − 1.282ln H − 4.12 0.99 < 0.01 1.795 干 Stem ln y = 1.548ln D + 1.739ln H − 5.148 0.99 < 0.01 −5.491 根 Root ln y = 2.256ln D − 3.059 0.97 < 0.01 2.284 地上 Aboveground ln y = 1.024ln（D2H） − 4.246 0.99 < 0.01 −14.025 全树种 All tree species 整株 Whole plant ln y = 2.296 7ln D + 0.350 8ln H − 2.674 1 0.98 < 0.01 −44.547 叶 Leaf ln y = 2.449 9lnD − 0.920 2ln H − 2.853 2 0.85 < 0.01 179.610 枝 Branch ln y = 3.333 0lnD − 0.943 2ln H − 3.993 6 0.89 < 0.01 217.206 干 Stem ln y = 0.962 3ln（D2H） − 3.919 0 0.96 < 0.01 48.822 根 Root ln y = 2.456 8ln D + 0.165 4ln H − 4.098 4 0.96 < 0.01 57.071 地上 Aboveground ln y = 2.254 5ln D + 0.400 9ln H − 2.962 9 0.98 < 0.01 −16.184
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表 6 不同发育阶段林分单位面积乔木储水量
Table 6 Water storage capacity of trees per unit area in different developing stages of stand
指标
Index中龄林
Middle-aged forest近熟林
Near-mature forest成熟林
Mature forest老龄林
Old growth forest储水量/（t·hm−2） Water storage capacity/(t·ha−1) （194.37 ± 10.39）b （201.37 ± 6.58）b （271.88 ± 8.96）a （276.36 ± 10.70）a 生物量/（t·hm−2） Biomass/(t·ha−1) （248.02 ± 13.97）b （248.66 ± 7.31）b （299.78 ± 12.21）a （312.18 ± 12.65）a 注：同行不同小写字母表示差异显著（P < 0.05）。Notes: different lowercase letters in the same line indicate significant differences (P < 0.05).
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