植物生态学报 ›› 2025, Vol. 49 ›› Issue (2): 356-366.DOI: 10.17521/cjpe.2024.0042 cstr: 32100.14.cjpe.2024.0042
• 研究论文 • 上一篇
陈文义1,2, 王智勇1,2, 周梦岩1, 麻文俊1, 王军辉1, 罗志斌1,3,4, 周婧1,*()
收稿日期:
2024-02-07
接受日期:
2024-08-23
出版日期:
2025-02-20
发布日期:
2025-02-20
通讯作者:
*周婧: (gaha2008@126.com)基金资助:
CHEN Wen-Yi1,2, WANG Zhi-Yong1,2, ZHOU Meng-Yan1, MA Wen-Jun1, WANG Jun-Hui1, LUO Zhi-Bin1,3,4, ZHOU Jing1,*()
Received:
2024-02-07
Accepted:
2024-08-23
Online:
2025-02-20
Published:
2025-02-20
Supported by:
摘要:
为了探究幼龄楸树(Catalpa bungei)主干、枝条、叶、粗根、细根、整株、地上部分和地下部分各组分生物量分配规律并建立相应的异速生长模型, 在3个相邻省份4个取样点的3-8年幼龄期楸树人工林中, 选取41株胸径(D)范围为3.2-24.8 cm的样木, 采用全称质量法测量楸树各组分生物量并分析其分配规律。分别以D、树高(H)及其复合形式D2H为预测变量, 利用简单幂函数的形式, 构建楸树主干、枝条、叶、粗根、细根、整株、地上部分和地下部分的生物量模型并验证其准确性。幼龄期楸树各组分生物量存在明显异速生长关系。地上部分生物量平均占比为80.54%, 其中主干生物量平均占比为49.29%, 远高于地下部分, 而细根生物量仅占整株的0.29%。在D ≤ 10 cm时, 随D增大, 枝条生物量占比逐渐增大, 而粗根生物量占比减小, 导致地上和地下生物量差距增大; 10 cm < D < 25 cm时, 各组分生物量占比变化放缓。在构建的各组分生物量模型中, 3个预测变量预测精度排序为D > D2H > H, 以D为单一预测变量拟合的主干、枝条、叶、粗根、整株、地上部分和地下部分异速生长模型精度较高, 以D2H为预测变量拟合的细根异速生长模型精度较高; 以不同径级楸树进行抽样, 验证模型准确性的结果显示, 各组分最优预测变量所构建的异速生长模型估测准确度高。幼龄期楸树各组分生物量平均占比顺序为: 主干>枝条>粗根>叶>细根; 随着楸树D增大, 地上部分生物量分配比例呈上升趋势。综合评估异速生长模型的拟合效果可知, D是预测楸树除细根外其他各组分生物量的最可靠变量, D2H是估算细根生物量的可靠变量。利用构建的异速生长模型可预测幼龄楸树的生长规律, 为选育优良楸树无性系提供重要参考。
陈文义, 王智勇, 周梦岩, 麻文俊, 王军辉, 罗志斌, 周婧. 幼龄楸树生物量分配规律与异速生长模型. 植物生态学报, 2025, 49(2): 356-366. DOI: 10.17521/cjpe.2024.0042
CHEN Wen-Yi, WANG Zhi-Yong, ZHOU Meng-Yan, MA Wen-Jun, WANG Jun-Hui, LUO Zhi-Bin, ZHOU Jing. Biomass allocation and allometric growth model of young Catalpa bungei. Chinese Journal of Plant Ecology, 2025, 49(2): 356-366. DOI: 10.17521/cjpe.2024.0042
样地 Sample site | 株数 Number of plants | 胸径 D (cm) | 年龄 Age (a) | 造林密度 Afforestation density (ind.·hm-2) | 年降水量 Mean annual precipitation (mm) | 年平均气温 Mean annual air temperature (℃) | 土壤类型 Soil type | ||||
---|---|---|---|---|---|---|---|---|---|---|---|
0-5 | 5-10 | 10-15 | 15-20 | 20-25 | |||||||
山东菏泽 Heze, Shandong | 9 | 1 | 2 | 2 | 3 | 1 | 5 | 833 | 782.6 ± 69.5 | 15.58 ± 0.05 | 冲积土 Fluvisol |
河南永城 Yongcheng, Henan | 8 | 1 | 2 | 3 | 2 | 0 | 4 | 833 | 888.1 ± 72.6 | 16.67 ± 0.11 | 冲积土 Fluvisol |
湖北襄阳 Xiangyang, Hubei | 12 | 1 | 3 | 3 | 2 | 3 | 5 | 833 | 908.6 ± 100.4 | 16.40 ± 0.09 | 始成土 Inceptisol |
湖北石首 Shishou, Hubei | 12 | 2 | 2 | 3 | 2 | 3 | 3, 8 | 833 | 1 222.8 ± 77.8 | 17.77 ± 0.13 | 淋溶土 Alfisol |
总计 Total | 41 | 5 | 9 | 11 | 9 | 7 | - | - | - | - | - |
建模组 Modeling group | 28 | 3 | 7 | 7 | 6 | 5 | - | - | - | - | - |
验证组 Testing group | 13 | 2 | 2 | 4 | 3 | 2 | - | - | - | - | - |
表1 不同胸径等级楸树样本量以及样地环境条件
Table 1 Sample size of Catalpa bungei with different D grades and environmental conditions of the sample sites
样地 Sample site | 株数 Number of plants | 胸径 D (cm) | 年龄 Age (a) | 造林密度 Afforestation density (ind.·hm-2) | 年降水量 Mean annual precipitation (mm) | 年平均气温 Mean annual air temperature (℃) | 土壤类型 Soil type | ||||
---|---|---|---|---|---|---|---|---|---|---|---|
0-5 | 5-10 | 10-15 | 15-20 | 20-25 | |||||||
山东菏泽 Heze, Shandong | 9 | 1 | 2 | 2 | 3 | 1 | 5 | 833 | 782.6 ± 69.5 | 15.58 ± 0.05 | 冲积土 Fluvisol |
河南永城 Yongcheng, Henan | 8 | 1 | 2 | 3 | 2 | 0 | 4 | 833 | 888.1 ± 72.6 | 16.67 ± 0.11 | 冲积土 Fluvisol |
湖北襄阳 Xiangyang, Hubei | 12 | 1 | 3 | 3 | 2 | 3 | 5 | 833 | 908.6 ± 100.4 | 16.40 ± 0.09 | 始成土 Inceptisol |
湖北石首 Shishou, Hubei | 12 | 2 | 2 | 3 | 2 | 3 | 3, 8 | 833 | 1 222.8 ± 77.8 | 17.77 ± 0.13 | 淋溶土 Alfisol |
总计 Total | 41 | 5 | 9 | 11 | 9 | 7 | - | - | - | - | - |
建模组 Modeling group | 28 | 3 | 7 | 7 | 6 | 5 | - | - | - | - | - |
验证组 Testing group | 13 | 2 | 2 | 4 | 3 | 2 | - | - | - | - | - |
胸径 D (cm) | 主干 Trunk (kg·plant-1) | 枝条 Branch (kg·plant-1) | 叶 Leaf (kg·plant-1) | 粗根 Coarse root (kg·plant-1) | 细根 Fine root (kg·plant-1) | |||||
---|---|---|---|---|---|---|---|---|---|---|
范围 Range | 平均值 Mean | 范围 Range | 平均值 Mean | 范围 Range | 平均值 Mean | 范围 Range | 平均值 Mean | 范围 Range | 平均值 Mean | |
0-5 | 0.83-6.56 | 2.52Ad | 0.15-5.46 | 1.55ABc | 0.11-0.55 | 0.27Bd | 0.51-3.21 | 1.39ABd | 0.01-0.04 | 0.02Bc |
5-10 | 3.47-17.30 | 8.35Ad | 1.12-6.15 | 3.23Bc | 0.28-3.41 | 1.30BCcd | 1.63-10.95 | 3.71Bcd | 0.01-0.07 | 0.05Cc |
10-15 | 14.45-37.18 | 26.49Ac | 7.33-20.09 | 13.04Bc | 1.29-6.92 | 4.06Dbc | 4.75-14.74 | 8.60Cc | 0.04-0.15 | 0.11Ec |
15-20 | 41.87-73.16 | 55.63Ab | 16.95-76.11 | 32.92Bb | 1.79-16.09 | 7.21CDab | 8.40-25.44 | 16.52Cb | 0.10-0.27 | 0.19Db |
20-25 | 53.91-116.49 | 89.22Aa | 35.50-73.64 | 56.43Ba | 2.87-19.19 | 9.61Da | 19.78-42.27 | 31.02Ca | 0.13-0.56 | 0.30Da |
表2 不同径级楸树主干、枝条、叶、粗根、细根生物量
Table 2 Biomass of trunk, branch, leaf, coarse root and fine root of different diameter classes of Catalpa bungei
胸径 D (cm) | 主干 Trunk (kg·plant-1) | 枝条 Branch (kg·plant-1) | 叶 Leaf (kg·plant-1) | 粗根 Coarse root (kg·plant-1) | 细根 Fine root (kg·plant-1) | |||||
---|---|---|---|---|---|---|---|---|---|---|
范围 Range | 平均值 Mean | 范围 Range | 平均值 Mean | 范围 Range | 平均值 Mean | 范围 Range | 平均值 Mean | 范围 Range | 平均值 Mean | |
0-5 | 0.83-6.56 | 2.52Ad | 0.15-5.46 | 1.55ABc | 0.11-0.55 | 0.27Bd | 0.51-3.21 | 1.39ABd | 0.01-0.04 | 0.02Bc |
5-10 | 3.47-17.30 | 8.35Ad | 1.12-6.15 | 3.23Bc | 0.28-3.41 | 1.30BCcd | 1.63-10.95 | 3.71Bcd | 0.01-0.07 | 0.05Cc |
10-15 | 14.45-37.18 | 26.49Ac | 7.33-20.09 | 13.04Bc | 1.29-6.92 | 4.06Dbc | 4.75-14.74 | 8.60Cc | 0.04-0.15 | 0.11Ec |
15-20 | 41.87-73.16 | 55.63Ab | 16.95-76.11 | 32.92Bb | 1.79-16.09 | 7.21CDab | 8.40-25.44 | 16.52Cb | 0.10-0.27 | 0.19Db |
20-25 | 53.91-116.49 | 89.22Aa | 35.50-73.64 | 56.43Ba | 2.87-19.19 | 9.61Da | 19.78-42.27 | 31.02Ca | 0.13-0.56 | 0.30Da |
胸径 D (cm) | 整株 Total tree (kg·plant -1) | 地上 Aboveground (kg·plant -1) | 地下 Belowground (kg·plant-1) | 根冠比 Root to shoot ratio | ||||
---|---|---|---|---|---|---|---|---|
范围 Range | 平均值 Mean | 范围 Range | 平均值 Mean | 范围 Range | 平均值 Mean | 范围 Range | 平均值 Mean | |
0-5 | 1.69-15.79 | 5.76 | 1.17-12.57 | 4.34 | 0.52-3.22 | 1.42 | 0.26-0.64 | 0.42a |
5-10 | 7.61-34.82 | 16.63 | 4.87-23.82 | 12.87 | 1.63-11.00 | 3.76 | 0.15-0.56 | 0.32a |
10-15 | 35.13-68.06 | 52.29 | 28.74-55.48 | 43.58 | 4.79-14.90 | 8.71 | 0.14-0.28 | 0.20b |
15-20 | 77.64-180.77 | 112.47 | 62.55-111.58 | 95.76 | 8.63-25.68 | 16.71 | 0.08-0.24 | 0.18b |
20-25 | 129.19-248.95 | 186.59 | 101.02-209.32 | 155.27 | 20.08-42.40 | 31.32 | 0.15-0.28 | 0.20b |
表3 不同径级楸树整株、地上部分和地下部分生物量
Table 3 Biomass of total tree, above- and below-ground parts of different diameter classes of Catalpa bungei
胸径 D (cm) | 整株 Total tree (kg·plant -1) | 地上 Aboveground (kg·plant -1) | 地下 Belowground (kg·plant-1) | 根冠比 Root to shoot ratio | ||||
---|---|---|---|---|---|---|---|---|
范围 Range | 平均值 Mean | 范围 Range | 平均值 Mean | 范围 Range | 平均值 Mean | 范围 Range | 平均值 Mean | |
0-5 | 1.69-15.79 | 5.76 | 1.17-12.57 | 4.34 | 0.52-3.22 | 1.42 | 0.26-0.64 | 0.42a |
5-10 | 7.61-34.82 | 16.63 | 4.87-23.82 | 12.87 | 1.63-11.00 | 3.76 | 0.15-0.56 | 0.32a |
10-15 | 35.13-68.06 | 52.29 | 28.74-55.48 | 43.58 | 4.79-14.90 | 8.71 | 0.14-0.28 | 0.20b |
15-20 | 77.64-180.77 | 112.47 | 62.55-111.58 | 95.76 | 8.63-25.68 | 16.71 | 0.08-0.24 | 0.18b |
20-25 | 129.19-248.95 | 186.59 | 101.02-209.32 | 155.27 | 20.08-42.40 | 31.32 | 0.15-0.28 | 0.20b |
图1 楸树各组分生物量占比平均值。图例中标出了具体计算结果, 格式为平均值±标准误。
Fig. 1 Average biomass proportion of various components of Catalpa bungei. The results (mean ± SE) are highlighted in the legend.
图2 幼龄楸树不同组分生物量比例随胸径的变化。
Fig. 2 Changes of biomass proportion of various components of young Catalpa bungei with increasing diameter at breast height (D).
组分生物量 Component biomass | 降水量 Precipitation | 气温 Air temperature | 年龄 Age |
---|---|---|---|
整株 Total tree | -0.029 | -0.110 | 0.496** |
地上部分 Aboveground | -0.054 | -0.129 | 0.487** |
地下部分 Belowground | 0.043 | -0.071 | 0.566** |
主干 Trunk | -0.015 | -0.115 | 0.507** |
枝条 Branch | -0.085 | -0.126 | 0.427** |
叶 Leaf | -0.215 | -0.157 | 0.141 |
粗根 Coarse root | 0.048 | -0.066 | 0.562** |
细根 Fine root | 0.054 | -0.110 | 0.481** |
表4 生物量与降水量、气温及楸树年龄间相关系数
Table 4 Correlation coefficient between component biomass of Catalpa bungei., precipitation, air temperature and the tree age
组分生物量 Component biomass | 降水量 Precipitation | 气温 Air temperature | 年龄 Age |
---|---|---|---|
整株 Total tree | -0.029 | -0.110 | 0.496** |
地上部分 Aboveground | -0.054 | -0.129 | 0.487** |
地下部分 Belowground | 0.043 | -0.071 | 0.566** |
主干 Trunk | -0.015 | -0.115 | 0.507** |
枝条 Branch | -0.085 | -0.126 | 0.427** |
叶 Leaf | -0.215 | -0.157 | 0.141 |
粗根 Coarse root | 0.048 | -0.066 | 0.562** |
细根 Fine root | 0.054 | -0.110 | 0.481** |
组分生物量 Component biomass | 回归模型 Regression model | a | b | R2 | RMSE | AICc |
---|---|---|---|---|---|---|
主干 Trunk | ln W = a + bln D | -2.521 | 2.257 | 0.949 3 | 0.296 8 | -62.04 |
ln W = a + b ln H | -3.978 | 3.400 | 0.748 9 | 0.660 2 | -17.27 | |
ln W = a + b ln (D2H) | -3.123 | 0.881 | 0.935 2 | 0.335 3 | -55.21 | |
枝条 Branch | ln W = a + bln D | -4.439 | 2.716 | 0.902 7 | 0.507 4 | -32.01 |
ln W = a + b ln H | -5.898 | 3.948 | 0.663 0 | 0.944 1 | 2.76 | |
ln W = a + bln (D2H) | -5.113 | 1.053 | 0.876 9 | 0.570 6 | -25.44 | |
叶 Leaf | ln W = a + bln D | -4.502 | 2.203 | 0.839 2 | 0.548 6 | -27.64 |
ln W = a + bln H | -5.761 | 3.239 | 0.630 6 | 0.831 5 | -4.35 | |
ln W = a + bln (D2H) | -5.061 | 0.856 | 0.818 8 | 0.582 4 | -24.29 | |
粗根 Coarse root | ln W = a + bln D | -2.629 | 1.894 | 0.903 0 | 0.353 2 | -52.30 |
ln W = a + bln H | -3.895 | 2.875 | 0.722 8 | 0.597 2 | -22.89 | |
ln W = a + bln (D2H) | -3.146 | 0.741 | 0.893 6 | 0.369 9 | -49.72 | |
细根 Fine root | ln W = a + bln D | -6.298 | 1.565 | 0.777 4 | 0.476 6 | -35.52 |
ln W = a + bln H | -7.786 | 2.591 | 0.740 3 | 0.514 8 | -31.21 | |
ln W = a + bln (D2H) | -6.805 | 0.624 | 0.798 4 | 0.453 5 | -38.30 | |
整株 Total tree | ln W = a + bln D | -1.836 | 2.264 | 0.946 0 | 0.307 8 | -60.00 |
ln W = a + bln H | -3.203 | 3.364 | 0.726 3 | 0.692 7 | -14.58 | |
ln W = a + bln (D2H) | -2.424 | 0.882 | 0.927 1 | 0.357 5 | -51.62 | |
地上 Aboveground | ln W = a + bln D | -2.313 | 2.368 | 0.943 6 | 0.329 2 | -56.23 |
ln W = a + bln H | -3.715 | 3.505 | 0.718 8 | 0.735 2 | -11.24 | |
ln W = a + bln (D2H) | -2.923 | 0.921 | 0.923 1 | 0.384 4 | -47.56 | |
地下 Belowground | ln W = a + bln D | -2.602 | 1.889 | 0.903 8 | 0.350 6 | -52.70 |
ln W = a + bln H | -3.863 | 2.866 | 0.723 4 | 0.594 5 | -23.14 | |
ln W = a + bln (D2H) | -3.117 | 0.739 | 0.894 4 | 0.367 3 | -50.11 |
表5 楸树整株及各组分的异速生长模型
Table 5 Allometric growth models of whole Catalpa bungei and each component
组分生物量 Component biomass | 回归模型 Regression model | a | b | R2 | RMSE | AICc |
---|---|---|---|---|---|---|
主干 Trunk | ln W = a + bln D | -2.521 | 2.257 | 0.949 3 | 0.296 8 | -62.04 |
ln W = a + b ln H | -3.978 | 3.400 | 0.748 9 | 0.660 2 | -17.27 | |
ln W = a + b ln (D2H) | -3.123 | 0.881 | 0.935 2 | 0.335 3 | -55.21 | |
枝条 Branch | ln W = a + bln D | -4.439 | 2.716 | 0.902 7 | 0.507 4 | -32.01 |
ln W = a + b ln H | -5.898 | 3.948 | 0.663 0 | 0.944 1 | 2.76 | |
ln W = a + bln (D2H) | -5.113 | 1.053 | 0.876 9 | 0.570 6 | -25.44 | |
叶 Leaf | ln W = a + bln D | -4.502 | 2.203 | 0.839 2 | 0.548 6 | -27.64 |
ln W = a + bln H | -5.761 | 3.239 | 0.630 6 | 0.831 5 | -4.35 | |
ln W = a + bln (D2H) | -5.061 | 0.856 | 0.818 8 | 0.582 4 | -24.29 | |
粗根 Coarse root | ln W = a + bln D | -2.629 | 1.894 | 0.903 0 | 0.353 2 | -52.30 |
ln W = a + bln H | -3.895 | 2.875 | 0.722 8 | 0.597 2 | -22.89 | |
ln W = a + bln (D2H) | -3.146 | 0.741 | 0.893 6 | 0.369 9 | -49.72 | |
细根 Fine root | ln W = a + bln D | -6.298 | 1.565 | 0.777 4 | 0.476 6 | -35.52 |
ln W = a + bln H | -7.786 | 2.591 | 0.740 3 | 0.514 8 | -31.21 | |
ln W = a + bln (D2H) | -6.805 | 0.624 | 0.798 4 | 0.453 5 | -38.30 | |
整株 Total tree | ln W = a + bln D | -1.836 | 2.264 | 0.946 0 | 0.307 8 | -60.00 |
ln W = a + bln H | -3.203 | 3.364 | 0.726 3 | 0.692 7 | -14.58 | |
ln W = a + bln (D2H) | -2.424 | 0.882 | 0.927 1 | 0.357 5 | -51.62 | |
地上 Aboveground | ln W = a + bln D | -2.313 | 2.368 | 0.943 6 | 0.329 2 | -56.23 |
ln W = a + bln H | -3.715 | 3.505 | 0.718 8 | 0.735 2 | -11.24 | |
ln W = a + bln (D2H) | -2.923 | 0.921 | 0.923 1 | 0.384 4 | -47.56 | |
地下 Belowground | ln W = a + bln D | -2.602 | 1.889 | 0.903 8 | 0.350 6 | -52.70 |
ln W = a + bln H | -3.863 | 2.866 | 0.723 4 | 0.594 5 | -23.14 | |
ln W = a + bln (D2H) | -3.117 | 0.739 | 0.894 4 | 0.367 3 | -50.11 |
图3 幼龄楸树的最优异速生长模型。D, 胸径; H, 树高; W, 生物量。
Fig. 3 Optimal allometric growth model for young Catalpa bungei. D, diameter at breast height; H, tree height; W, biomass.
图4 最优异速生长模型估测值与实测值的关系。图中实线为估测值与实测值线性拟合的结果, slope为拟合直线的斜率; 黑色虚线表示1:1的关系。MAPE, 平均绝对百分比误差。
Fig. 4 Relationships between the biomass observed and predicted by the optimal growth models. The solid line in the figure is the result of linear fitting between the predicted value and the observed value; slope, the slope of the fitted line. The black dashed line indicates a 1:1 relationship. MAPE, mean absolute percentage error.
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