植物生态学报 ›› 2009, Vol. 33 ›› Issue (1): 81-88.DOI: 10.3773/j.issn.1005-264x.2009.01.009
收稿日期:
2008-04-15
接受日期:
2008-08-22
出版日期:
2009-04-15
发布日期:
2009-01-30
通讯作者:
周志春
作者简介:
*E-mail: zczhou@fy.hz.zj.cn; zczhou_risf@163.com基金资助:
MA Xue-Hong, ZHOU Zhi-Chun*(), JIN Guo-Qing, ZHANG Yi
Received:
2008-04-15
Accepted:
2008-08-22
Online:
2009-04-15
Published:
2009-01-30
Contact:
ZHOU Zhi-Chun
摘要:
为了研究邻株竞争对马尾松(Pinus massoniana)和木荷(Schima superba)觅取异质分布养分机理和行为的影响, 我们设计单植、双株纯栽和两株混植3种栽植方式, 构建了同质和异质养分环境开展盆栽实验。结果表明: 单植时, 马尾松和木荷苗木生长对斑块养分反应敏感, 与同质养分环境相比, 两树种在异质养分环境中具有苗高生长量大、干物质积累量高、根系在富养斑块中大量增生, 根系N、P含量和吸收效率高等特点。在异质养分环境中, 木荷与马尾松邻株竞争时的生长表现优于双株纯栽模式而与单植处理相近, 根系形态可塑性和生理可塑性在其觅取斑块养分中的作用显著增强; 与木荷邻株竞争时, 马尾松苗高生长也表现出较单植和双株纯栽模式一定的优势, 这与其根系的广布性、觅养精确性和反应敏感度变化较小及富养斑块中根系P素含量和吸收效率较高等有关。相反, 同种邻株竞争则使得异质养分环境中马尾松和木荷的根系广布性减小, 反应敏感度减弱, 富养斑块中根系N、P含量降低, 苗高生长量和干物质积累量减小。与马尾松相比, 同种邻株竞争对异质养分环境中木荷生长的负向影响更为强烈。建议在生产中采用混交造林的方式促进马尾松和木荷生长。若要营造人工纯林, 可通过适当降低初植密度或及时调控林分密度促进林木生长。
马雪红, 周志春, 金国庆, 张一. 竞争对马尾松和木荷觅取异质分布养分行为的影响. 植物生态学报, 2009, 33(1): 81-88. DOI: 10.3773/j.issn.1005-264x.2009.01.009
MA Xue-Hong, ZHOU Zhi-Chun, JIN Guo-Qing, ZHANG Yi. EFFECTS OF COMPETITION ON FORAGING BEHAVIOR OF PINUS MASSONIANA AND SCHIMA SUPERBA IN A HETEROGENEOUS NUTRIENT ENVIRONMENT. Chinese Journal of Plant Ecology, 2009, 33(1): 81-88. DOI: 10.3773/j.issn.1005-264x.2009.01.009
养分环境 Nutrient environment | 栽植方式 Planting pattern | 马尾松 Pinus massoniana | 木荷 Schima superba | ||||||
---|---|---|---|---|---|---|---|---|---|
苗高 Seedling height (cm) | 干物质积累量 Dry matter accumulation (g) | 根冠比 Root-shoot ratio | 苗高 Seedling height (cm) | 干物质积累量 Dry matter accumulation (g) | 根冠比 Root-shoot ratio | ||||
异质 Heterogeneity | A | 19.3±3.6a | 1.820 9±0.708a | 0.389 9±0.234a | 18.2±4.9a | 5.038 8±2.951a | 0.534 1±0.134a | ||
B | 20.9±2.0a | 2.014 9±0.612a | 0.338 8±0.077a | 24.7±5.6b | 9.591 6±2.703Ab | 0.689 4±0.205a | |||
C | 17.3±4.9a | 4.002 6±2.553b | 0.300 4±0.132a | 23.8±8.8Ab | 10.682 3±6.241b | 0.682 5±0.102a | |||
同质 Homogeneity | A | 12.1±3.8a | 0.777 5±0.380a | 0.553 6±0.251a | — | — | — | ||
B | 10.0±2.3a | 0.614 4±0.165a | 0.675 5±0.159a | 23.6±6.9a | 7.069 8±1.458a | 0.507 2±0.119a | |||
C | 13.0±6.7a | 1.185 0±1.278a | 0.599 6±0.175a | 16.7±5.2b | 4.243 4±1.564b | 0.777 3±0.540a | |||
p | <0.000 1 | <0.000 1 | <0.000 1 | 0.260 6 | 0.026 5 | 0.727 0 |
表1 不同栽植方式下马尾松和木荷在异质和同质养分环境中的生长差异(平均值±标准偏差)
Table 1 Growth differences of Pinus massoniana and Schima superba in heterogeneous and homogeneous nutrient environments under different planting patterns (mean ± SD)
养分环境 Nutrient environment | 栽植方式 Planting pattern | 马尾松 Pinus massoniana | 木荷 Schima superba | ||||||
---|---|---|---|---|---|---|---|---|---|
苗高 Seedling height (cm) | 干物质积累量 Dry matter accumulation (g) | 根冠比 Root-shoot ratio | 苗高 Seedling height (cm) | 干物质积累量 Dry matter accumulation (g) | 根冠比 Root-shoot ratio | ||||
异质 Heterogeneity | A | 19.3±3.6a | 1.820 9±0.708a | 0.389 9±0.234a | 18.2±4.9a | 5.038 8±2.951a | 0.534 1±0.134a | ||
B | 20.9±2.0a | 2.014 9±0.612a | 0.338 8±0.077a | 24.7±5.6b | 9.591 6±2.703Ab | 0.689 4±0.205a | |||
C | 17.3±4.9a | 4.002 6±2.553b | 0.300 4±0.132a | 23.8±8.8Ab | 10.682 3±6.241b | 0.682 5±0.102a | |||
同质 Homogeneity | A | 12.1±3.8a | 0.777 5±0.380a | 0.553 6±0.251a | — | — | — | ||
B | 10.0±2.3a | 0.614 4±0.165a | 0.675 5±0.159a | 23.6±6.9a | 7.069 8±1.458a | 0.507 2±0.119a | |||
C | 13.0±6.7a | 1.185 0±1.278a | 0.599 6±0.175a | 16.7±5.2b | 4.243 4±1.564b | 0.777 3±0.540a | |||
p | <0.000 1 | <0.000 1 | <0.000 1 | 0.260 6 | 0.026 5 | 0.727 0 |
养分环境 Nutrient environment | 栽植方式 Planting pattern | 马尾松 Pinus massoniana | 木荷 Schima superba | ||||||||
---|---|---|---|---|---|---|---|---|---|---|---|
总根长 Total root length (mm) | 总根表面积 Total root surface area (cm2) | 根体积 Root volume (cm3) | 根系干物质量 Root dry matter accumulation (g) | 总根长 Total root length (mm) | 总根表面积 Total root surface area (cm2) | 根体积 Root volume (cm3) | 根系干物质量 Root dry matter accumulation (g) | ||||
异质 Heterogeneity | A | 677.38 ±311.4a | 109.956 1 ±56.041a | 0.942 6 ±0.214a | 1.440 9 ±0.842a | 3 864.79 ±2 209.7a | 473.388 6 ±285.712a | 0.998 6 ±0.410a | 4.676 0 ±3.021a | ||
B | 858.15 ±317.7ab | 142.710 7 ±54.809ab | 0.999 4 ±0.170ab | 1.905 4 ±0.778ab | 7 405.54 ±2 007.6b | 1 082.548 8 ±330.710b | 2.003 2 ±0.723b | 12.750 3 ±4.493b | |||
C | 987.32 ±602.5b | 167.104 5 ±102.854b | 1.187 4 ±0.389b | 2.271 1 ±1.440b | 7 780.97 ±5 648.1b | 1 024.766 9 ±715.828b | 1.750 8 ±0.930b | 10.825 9 ±7.542b | |||
同质 Homogeneity | A | 518.07 ±242.4a | 84.178 5 ±40.179a | 0.958 4 ±0.203a | 1.094 8 ±0.546a | - | - | - | - | ||
B | 686.45 ±260.9a | 103.647 0 ±37.331a | 0.962 8 ±0.094a | 1.256 7 ±0.444a | 5 886.63 ±1 644.8a | 775.254 2 ±166.937a | 1.336 8 ±0.305a | 8.274 9 ±1.923a | |||
C | 847.96 ±407.7a | 141.382 1 ±71.819a | 1.051 6 ±0.066a | 1.882 4 ±1.009a | 5 301.41 ±2 577.1a | 718.543 8 ±358.061a | 1.568 2 ±0.518a | 7.846 9 ±4.113a | |||
p | 0.032 4 | 0.021 0 | 0.079 2 | 0.015 7 | 0.409 0 | 0.364 8 | 0.584 9 | 0.339 8 |
表2 不同栽植方式下马尾松和木荷在异质和同质养分环境中的根系参数差异(平均值±标准偏差)
Table 2 Root parameter differences of Pinus massoniana and Schima superba in heterogeneous and homogeneous nutrient environments under different planting patterns (mean ± SD)
养分环境 Nutrient environment | 栽植方式 Planting pattern | 马尾松 Pinus massoniana | 木荷 Schima superba | ||||||||
---|---|---|---|---|---|---|---|---|---|---|---|
总根长 Total root length (mm) | 总根表面积 Total root surface area (cm2) | 根体积 Root volume (cm3) | 根系干物质量 Root dry matter accumulation (g) | 总根长 Total root length (mm) | 总根表面积 Total root surface area (cm2) | 根体积 Root volume (cm3) | 根系干物质量 Root dry matter accumulation (g) | ||||
异质 Heterogeneity | A | 677.38 ±311.4a | 109.956 1 ±56.041a | 0.942 6 ±0.214a | 1.440 9 ±0.842a | 3 864.79 ±2 209.7a | 473.388 6 ±285.712a | 0.998 6 ±0.410a | 4.676 0 ±3.021a | ||
B | 858.15 ±317.7ab | 142.710 7 ±54.809ab | 0.999 4 ±0.170ab | 1.905 4 ±0.778ab | 7 405.54 ±2 007.6b | 1 082.548 8 ±330.710b | 2.003 2 ±0.723b | 12.750 3 ±4.493b | |||
C | 987.32 ±602.5b | 167.104 5 ±102.854b | 1.187 4 ±0.389b | 2.271 1 ±1.440b | 7 780.97 ±5 648.1b | 1 024.766 9 ±715.828b | 1.750 8 ±0.930b | 10.825 9 ±7.542b | |||
同质 Homogeneity | A | 518.07 ±242.4a | 84.178 5 ±40.179a | 0.958 4 ±0.203a | 1.094 8 ±0.546a | - | - | - | - | ||
B | 686.45 ±260.9a | 103.647 0 ±37.331a | 0.962 8 ±0.094a | 1.256 7 ±0.444a | 5 886.63 ±1 644.8a | 775.254 2 ±166.937a | 1.336 8 ±0.305a | 8.274 9 ±1.923a | |||
C | 847.96 ±407.7a | 141.382 1 ±71.819a | 1.051 6 ±0.066a | 1.882 4 ±1.009a | 5 301.41 ±2 577.1a | 718.543 8 ±358.061a | 1.568 2 ±0.518a | 7.846 9 ±4.113a | |||
p | 0.032 4 | 0.021 0 | 0.079 2 | 0.015 7 | 0.409 0 | 0.364 8 | 0.584 9 | 0.339 8 |
养分斑块 Nutrient patch | 栽植方式 Planting pattern | 马尾松Pinus massoniana | 木荷Schima superba | |||||||
---|---|---|---|---|---|---|---|---|---|---|
总根长 Total root length (mm) | 总根表面积 Total root surface area (cm2) | 根体积 Root volume (cm3) | 根系干物质量 Root dry matter accumulation (g) | 总根长 Total root length (mm) | 总根表面积 Total root surface area (cm2) | 根体积 Root volume (cm3) | 根系干物质量 Root dry matter accumulation (g) | |||
富养 Rich | A | 460.87 ±223.2a | 74.962 2 ±39.917 a | 0.510 7 ±0.062a | 0.985 7 ±0.598a | 2 044.12 ±1 042.1a | 261.708 0 ±143.108a | 0.512 3 ±0.241a | 2.706 4 ±1.626a | |
B | 619.87 ±258.9a | 102.235 9 ±44.158ab | 0.522 1 ±0.036a | 1.345 7 ±0.603ab | 4 568.88 ±1 941.3b | 669.857 2 ±281.542b | 1.221 9 ±0.535b | 7.917 2 ±3.486b | ||
C | 710.27 ±509.1 a | 118.715 5 ±82.382b | 0.693 2 ±0.283b | 1.593 0 ±1.095b | 4 848.76 ±3 500.6b | 641.528 5 ±453.018b | 1.053 0 ±0.577b | 6.861 5 ±4.745b | ||
贫养 Poor | A | 258.85 ±110.3 a | 41.620 3 ±17.680a | 0.506 2 ±0.051a | 0.538 6 ±0.294a | 1 929.48 ±1 630.3a | 224.857 9 ±201.900a | 0.518 1 ±0.240a | 2.098 8 ±1.999a | |
B | 300.27 ±208.5 a | 50.698 4 ±34.235a | 0.529 5 ±0.085a | 0.694 3 ±0.483ab | 2 836.66 ±1 380.6a | 412.691 6 ±202.825a | 0.781 4 ±0.278a | 4.833 1 ±2.479b | ||
C | 307.83 ±96.8 a | 53.765 7 ±19.607a | 0.548 9 ±0.048a | 0.753 4 ±0.323b | 2 962.21 ±2 246.8a | 383.238 5 ±297.158a | 0.697 9 ±0.408a | 3.964 4 ±3.160ab | ||
p | <0.000 1 | 0.000 1 | 0.193 5 | 0.000 3 | 0.018 4 | 0.008 8 | 0.008 8 | 0.004 7 |
表3 不同栽植方式下马尾松和木荷在异质养分环境之富养和贫养斑块间的根系形态参数差异(平均值±标准偏差)
Table 3 Root parameter differences of Pinus massoniana and Schima superba in between rich- and poor-nutrient patch of heterogeneous nutrient environments under different planting patterns (mean±SD)
养分斑块 Nutrient patch | 栽植方式 Planting pattern | 马尾松Pinus massoniana | 木荷Schima superba | |||||||
---|---|---|---|---|---|---|---|---|---|---|
总根长 Total root length (mm) | 总根表面积 Total root surface area (cm2) | 根体积 Root volume (cm3) | 根系干物质量 Root dry matter accumulation (g) | 总根长 Total root length (mm) | 总根表面积 Total root surface area (cm2) | 根体积 Root volume (cm3) | 根系干物质量 Root dry matter accumulation (g) | |||
富养 Rich | A | 460.87 ±223.2a | 74.962 2 ±39.917 a | 0.510 7 ±0.062a | 0.985 7 ±0.598a | 2 044.12 ±1 042.1a | 261.708 0 ±143.108a | 0.512 3 ±0.241a | 2.706 4 ±1.626a | |
B | 619.87 ±258.9a | 102.235 9 ±44.158ab | 0.522 1 ±0.036a | 1.345 7 ±0.603ab | 4 568.88 ±1 941.3b | 669.857 2 ±281.542b | 1.221 9 ±0.535b | 7.917 2 ±3.486b | ||
C | 710.27 ±509.1 a | 118.715 5 ±82.382b | 0.693 2 ±0.283b | 1.593 0 ±1.095b | 4 848.76 ±3 500.6b | 641.528 5 ±453.018b | 1.053 0 ±0.577b | 6.861 5 ±4.745b | ||
贫养 Poor | A | 258.85 ±110.3 a | 41.620 3 ±17.680a | 0.506 2 ±0.051a | 0.538 6 ±0.294a | 1 929.48 ±1 630.3a | 224.857 9 ±201.900a | 0.518 1 ±0.240a | 2.098 8 ±1.999a | |
B | 300.27 ±208.5 a | 50.698 4 ±34.235a | 0.529 5 ±0.085a | 0.694 3 ±0.483ab | 2 836.66 ±1 380.6a | 412.691 6 ±202.825a | 0.781 4 ±0.278a | 4.833 1 ±2.479b | ||
C | 307.83 ±96.8 a | 53.765 7 ±19.607a | 0.548 9 ±0.048a | 0.753 4 ±0.323b | 2 962.21 ±2 246.8a | 383.238 5 ±297.158a | 0.697 9 ±0.408a | 3.964 4 ±3.160ab | ||
p | <0.000 1 | 0.000 1 | 0.193 5 | 0.000 3 | 0.018 4 | 0.008 8 | 0.008 8 | 0.004 7 |
树种 Species | 栽植方式 Planting pattern | 觅养精确性 Foraging precision | 反应敏感度 Sensitivity |
---|---|---|---|
马尾松 Pinus massoniana | A | 0.348 1 | 2.341 8 |
B | 0.353 4 | 3.286 9 | |
C | 0.348 6 | 3.377 8 | |
木荷 Schima superba | A | 0.244 7 | — |
B | 0.221 1 | 1.356 7 | |
C | 0.261 0 | 2.517 4 |
表4 不同栽植方式下马尾松和木荷对异质养分环境的敏感度和觅养精确性
Table 4 Sensitivity to heterogeneous nutrient environment and foraging precision of Pinus massoniana and Schima superba under different planting patterns
树种 Species | 栽植方式 Planting pattern | 觅养精确性 Foraging precision | 反应敏感度 Sensitivity |
---|---|---|---|
马尾松 Pinus massoniana | A | 0.348 1 | 2.341 8 |
B | 0.353 4 | 3.286 9 | |
C | 0.348 6 | 3.377 8 | |
木荷 Schima superba | A | 0.244 7 | — |
B | 0.221 1 | 1.356 7 | |
C | 0.261 0 | 2.517 4 |
栽植方式 Planting pattern | 马尾松Pinus massoniana | 木荷Schima superba | ||||
---|---|---|---|---|---|---|
N | P | N | P | |||
A | 10.458 7±0.003ab | 0.744 8±0.348a | 12.303 1±1.787a | 1.068 8±0.246a | ||
B | 8.373 6±2.389a | 0.843 4±0.504a | 9.749 1±1.488b | 0.918 5±0.090a | ||
C | 12.456 6±2.642b | 1.208 7±0.618a | 15.254 2±1.943a | 0.978 7±0.291a |
表5 不同栽植方式下马尾松和木荷在异质养分环境中叶片N、P吸收效率(平均值±标准偏差)
Table 5 N and P absorption efficiency of Pinus massoniana and Schima superba leaves in the heterogeneous nutrient environments under different planting patterns (mean±SD, mg·plant-1)
栽植方式 Planting pattern | 马尾松Pinus massoniana | 木荷Schima superba | ||||
---|---|---|---|---|---|---|
N | P | N | P | |||
A | 10.458 7±0.003ab | 0.744 8±0.348a | 12.303 1±1.787a | 1.068 8±0.246a | ||
B | 8.373 6±2.389a | 0.843 4±0.504a | 9.749 1±1.488b | 0.918 5±0.090a | ||
C | 12.456 6±2.642b | 1.208 7±0.618a | 15.254 2±1.943a | 0.978 7±0.291a |
树种 Species | 栽植方式 Planting pattern | N | P | |||||||
---|---|---|---|---|---|---|---|---|---|---|
富养 Rich patch | 贫养 Poor patch | 比值 Ratio | 富养 Rich patch | 贫养 Poor patch | 比值 Ratio | |||||
养分含量 Nutrient content (mg·g-1) | ||||||||||
马尾松 Pinus massoniana | A | 2.177 8±2.633a | 0.845 0±0.598a | 2.58 | 0.175 4±0.164a | 0.070 2±0.033a | 2.50 | |||
B | 2.429 8±1.208a | 1.680 4±1.430b | 1.45 | 0.242 3±0.127b | 0.136 2±0.164b | 1.78 | ||||
C | 4.477 1±3.216b | 0.977 4±0.489a | 4.58 | 0.247 6±0.179b | 0.106 0±0.079b | 2.34 | ||||
木荷 Schima superba | A | 4.943 5±3.691a | 2.950 4±3.223a | 1.68 | 0.778 1±0.660a | 0.515 1±0.655a | 1.51 | |||
B | 14.599 6±5.462b | 3.364 9±1.940a | 4.34 | 2.071 2±1.459a | 0.681 4±0.675a | 3.04 | ||||
C | 9.710 1±6.516a | 2.426 3±2.589a | 4.00 | 1.203 1±1.199a | 0.510 0±0.488a | 2.41 | ||||
吸收效率 Nutrient absorption efficiency (mg·plant-1) | ||||||||||
马尾松 Pinus massoniana | A | 8.917 3±2.738a | 8.639 1±3.446a | 1.03 | 0.741 5±0.312a | 0.749 0±0.272a | 0.99 | |||
B | 8.311 6±1.494b | 11.861 2±2.770b | 0.70 | 0.848 4±0.309a | 0.972 4±0.576b | 0.87 | ||||
C | 13.631 9±1.173c | 6.999 8±2.096a | 1.95 | 0.754 4±0.442a | 0.688 1±0.275a | 1.10 | ||||
木荷 Schima superba | A | 9.621 9±3.747a | 7.879 0±4.383a | 1.22 | 1.414 3±0.587a | 1.875 7±2.754a | 0.75 | |||
B | 11.183 1±2.457a | 4.575 8±1.026b | 2.44 | 0.809 0±0.527b | 1.616 3±0.889a | 0.50 | ||||
C | 9.920 5±2.251a | 3.435 8±1.245b | 2.89 | 1.172 6±0.559a | 0.861 3±0.433b | 1.36 |
表6 不同栽植方式下马尾松和木荷在异质养分环境之富养和贫养斑块中根系N、P含量和吸收效率(平均值±标准偏差)
Table 6 N and P contents and absorption efficiency of Pinus massoniana and Schima superba roots in rich- and poor-nutrient patch of heterogeneous nutrient environment under different planting patterns (mean±SD)
树种 Species | 栽植方式 Planting pattern | N | P | |||||||
---|---|---|---|---|---|---|---|---|---|---|
富养 Rich patch | 贫养 Poor patch | 比值 Ratio | 富养 Rich patch | 贫养 Poor patch | 比值 Ratio | |||||
养分含量 Nutrient content (mg·g-1) | ||||||||||
马尾松 Pinus massoniana | A | 2.177 8±2.633a | 0.845 0±0.598a | 2.58 | 0.175 4±0.164a | 0.070 2±0.033a | 2.50 | |||
B | 2.429 8±1.208a | 1.680 4±1.430b | 1.45 | 0.242 3±0.127b | 0.136 2±0.164b | 1.78 | ||||
C | 4.477 1±3.216b | 0.977 4±0.489a | 4.58 | 0.247 6±0.179b | 0.106 0±0.079b | 2.34 | ||||
木荷 Schima superba | A | 4.943 5±3.691a | 2.950 4±3.223a | 1.68 | 0.778 1±0.660a | 0.515 1±0.655a | 1.51 | |||
B | 14.599 6±5.462b | 3.364 9±1.940a | 4.34 | 2.071 2±1.459a | 0.681 4±0.675a | 3.04 | ||||
C | 9.710 1±6.516a | 2.426 3±2.589a | 4.00 | 1.203 1±1.199a | 0.510 0±0.488a | 2.41 | ||||
吸收效率 Nutrient absorption efficiency (mg·plant-1) | ||||||||||
马尾松 Pinus massoniana | A | 8.917 3±2.738a | 8.639 1±3.446a | 1.03 | 0.741 5±0.312a | 0.749 0±0.272a | 0.99 | |||
B | 8.311 6±1.494b | 11.861 2±2.770b | 0.70 | 0.848 4±0.309a | 0.972 4±0.576b | 0.87 | ||||
C | 13.631 9±1.173c | 6.999 8±2.096a | 1.95 | 0.754 4±0.442a | 0.688 1±0.275a | 1.10 | ||||
木荷 Schima superba | A | 9.621 9±3.747a | 7.879 0±4.383a | 1.22 | 1.414 3±0.587a | 1.875 7±2.754a | 0.75 | |||
B | 11.183 1±2.457a | 4.575 8±1.026b | 2.44 | 0.809 0±0.527b | 1.616 3±0.889a | 0.50 | ||||
C | 9.920 5±2.251a | 3.435 8±1.245b | 2.89 | 1.172 6±0.559a | 0.861 3±0.433b | 1.36 |
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