植物生态学报 ›› 2014, Vol. 38 ›› Issue (9): 929-940.DOI: 10.3724/SP.J.1258.2014.00087
所属专题: 生态化学计量
收稿日期:
2014-03-11
接受日期:
2014-06-03
出版日期:
2014-03-11
发布日期:
2014-09-22
通讯作者:
张元明
基金资助:
XIAO Yao1,2,TAO Ye1,ZHANG Yuan-Ming1,*()
Received:
2014-03-11
Accepted:
2014-06-03
Online:
2014-03-11
Published:
2014-09-22
Contact:
ZHANG Yuan-Ming
摘要:
荒漠草本植物是荒漠生态系统物种多样性的主体, 对其生物量分配及叶片化学计量特征随植物生长的变化规律的研究有助于深入了解荒漠草本植物生存策略和功能特征。该文选择古尔班通古特沙漠4种优势草本(2种短命植物, 2种一年生长营养期植物)为研究对象, 通过野外原位多时段取样, 对比研究了四者生物量分配、叶片N-P化学计量学随植物生长的变化特征, 以及二者之间的关系。结果表明, 在生物量累积过程中, 4种植物根冠比逐渐减小, 地上与地下生物量间的相关生长关系也发生变化, 其中琉苞菊(Hyalea pulchella)和角果藜(Ceratocarpus arenarius)的相关生长指数先增加后减小, 并趋于稳定, 而尖喙牻牛儿苗(Erodium oxyrrhynchum)和沙蓬(Agriophyllum squarrosum)的相关生长指数由小到大并趋于等速生长。琉苞菊叶片N、P含量呈逐渐增长趋势, 而另外3种植物呈下降趋势, 表明所研究的荒漠植物在生长过程中, 叶片N-P化学计量发生改变, 叶片化学计量特征与生物量指标的相关性较弱。
肖遥,陶冶,张元明. 古尔班通古特沙漠4种荒漠草本植物不同生长期的生物量分配与叶片化学计量特征. 植物生态学报, 2014, 38(9): 929-940. DOI: 10.3724/SP.J.1258.2014.00087
XIAO Yao,TAO Ye,ZHANG Yuan-Ming. Biomass allocation and leaf stoichiometric characteristics in four desert herbaceous plants during different growth periods in the Gurbantünggüt Desert, China. Chinese Journal of Plant Ecology, 2014, 38(9): 929-940. DOI: 10.3724/SP.J.1258.2014.00087
物种 Species | 生长期 Growth period | 样本量 No. of samples | 地上生物量 Aboveground biomass (g) | 地下生物量 Belowground biomass (g) | 总生物量 Total biomass (g) |
---|---|---|---|---|---|
琉苞菊 Hyalea pulchella | April 18 | 33 | 0.026 4 ± 0.003 7a | 0.007 7 ± 0.000 7a | 0.034 1 ± 0.004 3a |
April 28 | 35 | 0.094 4 ± 0.006 8ab | 0.010 2 ± 0.000 7a | 0.104 6 ± 0.007 2ab | |
May 8 | 33 | 0.212 4 ± 0.016 4ab | 0.031 0 ± 0.003 3b | 0.243 4 ± 0.019 4ab | |
May 18 | 40 | 0.328 0 ± 0.023 2b | 0.055 4 ± 0.003 9c | 0.383 3 ± 0.026 8b | |
May 31 | 40 | 1.431 0 ± 0.179 8c | 0.145 9 ± 0.014 2d | 1.577 0 ± 0.192 8c | |
尖喙牻牛儿苗 Erodium oxyrrhynchum | April 28 | 35 | 0.018 4 ± 0.001 4a | 0.004 9 ± 0.000 3a | 0.023 3 ± 0.001 6a |
May 8 | 40 | 0.051 3 ± 0.004 7a | 0.010 8 ± 0.000 9b | 0.062 1 ± 0.005 4a | |
May 18 | 40 | 0.113 6 ± 0.008 6b | 0.025 1 ± 0.002 1c | 0.138 7 ± 0.010 6b | |
May 31 | 36 | 0.265 4 ± 0.011 3c | 0.024 3 ± 0.003 0c | 0.289 7 ± 0.012 3c | |
角果藜 Ceratocarpus arenarius | April 28 | 29 | 0.009 1 ± 0.001 0a | 0.001 9 ± 0.000 1a | 0.010 9 ± 0.001 1a |
May 8 | 40 | 0.017 8 ± 0.001 4a | 0.003 9 ± 0.000 3a | 0.021 7 ± 0.001 7a | |
May 18 | 40 | 0.068 3 ± 0.004 9ab | 0.009 8 ± 0.000 7a | 0.078 1 ± 0.005 4ab | |
May 31 | 40 | 0.152 7 ± 0.016 4b | 0.018 5 ± 0.001 6b | 0.171 1 ± 0.017 8b | |
June 10 | 40 | 0.407 3 ± 0.044 3c | 0.037 1 ± 0.003 2c | 0.444 4 ± 0.047 4c | |
June 27 | 32 | 1.215 0 ± 0.100 9d | 0.091 5 ± 0.006 6d | 1.307 0 ± 0.106 4d | |
沙蓬 Agriophyllum squarrosum | May 8 | 38 | 0.027 7 ± 0.002 5a | 0.005 8 ± 0.000 3a | 0.033 4 ± 0.002 8a |
May 18 | 40 | 0.125 2 ± 0.010 0a | 0.019 9 ± 0.001 5a | 0.145 1 ± 0.011 4a | |
May 31 | 40 | 0.587 3 ± 0.054 0a | 0.091 2 ± 0.008 0ab | 0.678 5 ± 0.061 4a | |
June 10 | 17 | 2.364 0 ± 0.320 3b | 0.254 6 ± 0.032 3b | 2.618 0 ± 0.351 2b | |
June 27 | 24 | 14.720 0 ± 1.486 0c | 1.598 0 ± 0.207 1c | 16.320 0 ± 1.680 0c |
表1 4种植物不同生长期的生物量
Table 1 Biomass of the four plants in different growth periods
物种 Species | 生长期 Growth period | 样本量 No. of samples | 地上生物量 Aboveground biomass (g) | 地下生物量 Belowground biomass (g) | 总生物量 Total biomass (g) |
---|---|---|---|---|---|
琉苞菊 Hyalea pulchella | April 18 | 33 | 0.026 4 ± 0.003 7a | 0.007 7 ± 0.000 7a | 0.034 1 ± 0.004 3a |
April 28 | 35 | 0.094 4 ± 0.006 8ab | 0.010 2 ± 0.000 7a | 0.104 6 ± 0.007 2ab | |
May 8 | 33 | 0.212 4 ± 0.016 4ab | 0.031 0 ± 0.003 3b | 0.243 4 ± 0.019 4ab | |
May 18 | 40 | 0.328 0 ± 0.023 2b | 0.055 4 ± 0.003 9c | 0.383 3 ± 0.026 8b | |
May 31 | 40 | 1.431 0 ± 0.179 8c | 0.145 9 ± 0.014 2d | 1.577 0 ± 0.192 8c | |
尖喙牻牛儿苗 Erodium oxyrrhynchum | April 28 | 35 | 0.018 4 ± 0.001 4a | 0.004 9 ± 0.000 3a | 0.023 3 ± 0.001 6a |
May 8 | 40 | 0.051 3 ± 0.004 7a | 0.010 8 ± 0.000 9b | 0.062 1 ± 0.005 4a | |
May 18 | 40 | 0.113 6 ± 0.008 6b | 0.025 1 ± 0.002 1c | 0.138 7 ± 0.010 6b | |
May 31 | 36 | 0.265 4 ± 0.011 3c | 0.024 3 ± 0.003 0c | 0.289 7 ± 0.012 3c | |
角果藜 Ceratocarpus arenarius | April 28 | 29 | 0.009 1 ± 0.001 0a | 0.001 9 ± 0.000 1a | 0.010 9 ± 0.001 1a |
May 8 | 40 | 0.017 8 ± 0.001 4a | 0.003 9 ± 0.000 3a | 0.021 7 ± 0.001 7a | |
May 18 | 40 | 0.068 3 ± 0.004 9ab | 0.009 8 ± 0.000 7a | 0.078 1 ± 0.005 4ab | |
May 31 | 40 | 0.152 7 ± 0.016 4b | 0.018 5 ± 0.001 6b | 0.171 1 ± 0.017 8b | |
June 10 | 40 | 0.407 3 ± 0.044 3c | 0.037 1 ± 0.003 2c | 0.444 4 ± 0.047 4c | |
June 27 | 32 | 1.215 0 ± 0.100 9d | 0.091 5 ± 0.006 6d | 1.307 0 ± 0.106 4d | |
沙蓬 Agriophyllum squarrosum | May 8 | 38 | 0.027 7 ± 0.002 5a | 0.005 8 ± 0.000 3a | 0.033 4 ± 0.002 8a |
May 18 | 40 | 0.125 2 ± 0.010 0a | 0.019 9 ± 0.001 5a | 0.145 1 ± 0.011 4a | |
May 31 | 40 | 0.587 3 ± 0.054 0a | 0.091 2 ± 0.008 0ab | 0.678 5 ± 0.061 4a | |
June 10 | 17 | 2.364 0 ± 0.320 3b | 0.254 6 ± 0.032 3b | 2.618 0 ± 0.351 2b | |
June 27 | 24 | 14.720 0 ± 1.486 0c | 1.598 0 ± 0.207 1c | 16.320 0 ± 1.680 0c |
图2 4种荒漠草本植物不同生长期根冠比(R/S)变化(平均值±标准误差)。同一物种不同字母表示差异显著(p < 0.05)。
Fig. 2 Changes in root to shoot ratio (R/S) of the four desert herbs in different growth periods (mean ± SE). Different letters indicate significant differences within species (p < 0.05).
图3 4种荒漠草本植物不同生长期地上生物量(AGB)与地下生物量(BGB)间的相关生长关系。
Fig. 3 Allometric relationships between aboveground (AGB) and belowground biomass (BGB) of the four desert herbs in different growth periods.
物种 Species | 生长期 Growth period | R2 | p | 相关生长指数(平均值±标准误差) Allometric scaling exponent α (mean ± SD) | 等速生长检验 Test of isometry | |
---|---|---|---|---|---|---|
F | p | |||||
琉苞菊 Hyalea pulchella | April 18 | 0.461 | <0.000 1 | 0.542 ± 0.146c | 24.440 | 0.000 |
April 28 | 0.425 | <0.000 1 | 0.821 ± 0.220b | 2.273 | 0.141 | |
May 8 | 0.768 | <0.000 1 | 1.536 ± 0.271a | 26.161 | 0.000 | |
May 18 | 0.794 | <0.000 1 | 0.996 ± 0.149b | 0.003 | 0.960 | |
May 31 | 0.816 | <0.000 1 | 0.942 ± 0.133b | 0.739 | 0.395 | |
尖喙牻牛儿苗 Erodium oxyrrhynchum | April 28 | 0.444 | <0.000 1 | 0.700 ± 0.185b | 7.872 | 0.008 |
May 8 | 0.672 | <0.000 1 | 0.958 ± 0.180a | 0.215 | 0.646 | |
May 18 | 0.744 | <0.000 1 | 1.148 ± 0.191a | 2.838 | 0.100 | |
May 31 | 0.869 | <0.000 1 | 1.047 ± 0.132a | 0.558 | 0.460 | |
角果藜 Ceratocarpus arenarius | April 28 | 0.707 | <0.000 1 | 0.721 ± 0.154b | 10.221 | 0.004 |
May 8 | 0.692 | <0.000 1 | 1.058 ± 0.193a | 0.388 | 0.537 | |
May 18 | 0.618 | <0.000 1 | 0.852 ± 0.173ab | 2.582 | 0.116 | |
May 31 | 0.846 | <0.000 1 | 0.840 ± 0.108b | 7.520 | 0.009 | |
June 10 | 0.844 | <0.000 1 | 0.703 ± 0.091b | 31.576 | 0.000 | |
June 27 | 0.736 | <0.000 1 | 0.854 ± 0.163ab | 2.868 | 0.101 | |
沙蓬 Agriophyllum squarrosum | May 8 | 0.813 | <0.000 1 | 0.699 ± 0.102b | 25.809 | 0.000 |
May 18 | 0.739 | <0.000 1 | 0.854 ± 0.143b | 3.682 | 0.063 | |
May 31 | 0.839 | <0.000 1 | 0.954 ± 0.126ab | 0.529 | 0.472 | |
June 10 | 0.907 | <0.000 1 | 0.937 ± 0.157ab | 0.695 | 0.418 | |
June 27 | 0.802 | <0.000 1 | 1.166 ± 0.229a | 2.632 | 0.119 |
表2 4种荒漠草本植物不同生长期地上与地下生物量间的相关生长指数及等速生长检验
Table 2 Allometric scaling exponents and the test of isometry between above- and belowground biomass of the four desert herbs in different growth periods
物种 Species | 生长期 Growth period | R2 | p | 相关生长指数(平均值±标准误差) Allometric scaling exponent α (mean ± SD) | 等速生长检验 Test of isometry | |
---|---|---|---|---|---|---|
F | p | |||||
琉苞菊 Hyalea pulchella | April 18 | 0.461 | <0.000 1 | 0.542 ± 0.146c | 24.440 | 0.000 |
April 28 | 0.425 | <0.000 1 | 0.821 ± 0.220b | 2.273 | 0.141 | |
May 8 | 0.768 | <0.000 1 | 1.536 ± 0.271a | 26.161 | 0.000 | |
May 18 | 0.794 | <0.000 1 | 0.996 ± 0.149b | 0.003 | 0.960 | |
May 31 | 0.816 | <0.000 1 | 0.942 ± 0.133b | 0.739 | 0.395 | |
尖喙牻牛儿苗 Erodium oxyrrhynchum | April 28 | 0.444 | <0.000 1 | 0.700 ± 0.185b | 7.872 | 0.008 |
May 8 | 0.672 | <0.000 1 | 0.958 ± 0.180a | 0.215 | 0.646 | |
May 18 | 0.744 | <0.000 1 | 1.148 ± 0.191a | 2.838 | 0.100 | |
May 31 | 0.869 | <0.000 1 | 1.047 ± 0.132a | 0.558 | 0.460 | |
角果藜 Ceratocarpus arenarius | April 28 | 0.707 | <0.000 1 | 0.721 ± 0.154b | 10.221 | 0.004 |
May 8 | 0.692 | <0.000 1 | 1.058 ± 0.193a | 0.388 | 0.537 | |
May 18 | 0.618 | <0.000 1 | 0.852 ± 0.173ab | 2.582 | 0.116 | |
May 31 | 0.846 | <0.000 1 | 0.840 ± 0.108b | 7.520 | 0.009 | |
June 10 | 0.844 | <0.000 1 | 0.703 ± 0.091b | 31.576 | 0.000 | |
June 27 | 0.736 | <0.000 1 | 0.854 ± 0.163ab | 2.868 | 0.101 | |
沙蓬 Agriophyllum squarrosum | May 8 | 0.813 | <0.000 1 | 0.699 ± 0.102b | 25.809 | 0.000 |
May 18 | 0.739 | <0.000 1 | 0.854 ± 0.143b | 3.682 | 0.063 | |
May 31 | 0.839 | <0.000 1 | 0.954 ± 0.126ab | 0.529 | 0.472 | |
June 10 | 0.907 | <0.000 1 | 0.937 ± 0.157ab | 0.695 | 0.418 | |
June 27 | 0.802 | <0.000 1 | 1.166 ± 0.229a | 2.632 | 0.119 |
图4 4种荒漠草本植物不同生长期叶片N、P含量及N:P (平均值±标准误差)。不同字母表示不同生长期同一指标差异显著(p < 0.05)。
Fig. 4 Leaf N, P contents and N:P of the four desert herbs in different growth periods (mean ± SE). Different letters indicate significant differences among different growth periods for a given index (p < 0.05).
图5 4种荒漠草本植物不同生长期叶片N:P (平均值±标准误差)。不同字母表示同一物种不同生长期N:P差异显著(p < 0.05)。
Fig. 5 Leaf N:P of the four desert herbs in different growth periods (mean ± SE). Different letters indicate significant differences among different growth periods within species (p < 0.05).
物种 Species | N (%) | P (%) | N:P (%) |
---|---|---|---|
琉苞菊 Hyalea pulchella | 34.74 | 29.43 | 13.39 |
尖喙牻牛儿苗 Erodium oxyrrhynchum | 33.49 | 23.32 | 14.45 |
角果藜 Ceratocarpus arenarius | 22.97 | 16.36 | 18.26 |
沙蓬 Agriophyllum squarrosum | 28.31 | 31.91 | 28.72 |
表3 4种荒漠草本植物不同生长期叶片化学计量特征的变异系数
Table 3 Coefficients of variation in the characteristics of leaf stoichiometry in the four desert herbs in different growth periods
物种 Species | N (%) | P (%) | N:P (%) |
---|---|---|---|
琉苞菊 Hyalea pulchella | 34.74 | 29.43 | 13.39 |
尖喙牻牛儿苗 Erodium oxyrrhynchum | 33.49 | 23.32 | 14.45 |
角果藜 Ceratocarpus arenarius | 22.97 | 16.36 | 18.26 |
沙蓬 Agriophyllum squarrosum | 28.31 | 31.91 | 28.72 |
指标 Index | 琉苞菊 Hyalea pulchella | 尖喙牻牛儿苗 Erodium oxyrrhynchum | 角果藜 Ceratocarpus arenarius | 沙蓬 Agriophyllum squarrosum | |||||||||||
---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|
N | P | N:P | N | P | N:P | N | P | N:P | N | P | N:P | ||||
P | 0.951* | - | - | 0.928 | - | - | 0.910* | - | - | 0.574 | - | - | |||
N:P | 0.602 | 0.330 | - | 0.836 | 0.572 | - | 0.790 | 0.464 | - | 0.455 | -0.466 | - | |||
BGB | 0.833 | 0.753 | 0.516 | -0.955* | -0.807 | -0.905 | -0.920** | -0.720 | -0.905* | -0.599 | -0.687 | 0.125 | |||
AGB | 0.743 | 0.652 | 0.486 | -0.936 | -0.995** | -0.601 | -0.894* | -0.671 | -0.922** | -0.586 | -0.680 | 0.131 | |||
TB | 0.752 | 0.661 | 0.489 | -0.952* | -0.995** | -0.636 | -0.896* | -0.675 | -0.921** | -0.587 | -0.681 | 0.131 | |||
R/S | -0.546 | -0.392 | -0.670 | 0.868 | 0.988* | 0.454 | 0.893* | 0.901* | 0.588 | 0.288 | 0.839 | -0.624 | |||
α | 0.379 | 0.124 | 0.906* | -0.791 | -0.644 | -0.766 | 0.221 | 0.435 | -0.147 | -0.688 | -0.904* | -0.260 |
表4 4种荒漠草本植物叶片化学计量特征与生物量间的相关系数
Table 4 Correlation coefficients between leaf stoichiometry and biomass of the four desert herbs
指标 Index | 琉苞菊 Hyalea pulchella | 尖喙牻牛儿苗 Erodium oxyrrhynchum | 角果藜 Ceratocarpus arenarius | 沙蓬 Agriophyllum squarrosum | |||||||||||
---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|
N | P | N:P | N | P | N:P | N | P | N:P | N | P | N:P | ||||
P | 0.951* | - | - | 0.928 | - | - | 0.910* | - | - | 0.574 | - | - | |||
N:P | 0.602 | 0.330 | - | 0.836 | 0.572 | - | 0.790 | 0.464 | - | 0.455 | -0.466 | - | |||
BGB | 0.833 | 0.753 | 0.516 | -0.955* | -0.807 | -0.905 | -0.920** | -0.720 | -0.905* | -0.599 | -0.687 | 0.125 | |||
AGB | 0.743 | 0.652 | 0.486 | -0.936 | -0.995** | -0.601 | -0.894* | -0.671 | -0.922** | -0.586 | -0.680 | 0.131 | |||
TB | 0.752 | 0.661 | 0.489 | -0.952* | -0.995** | -0.636 | -0.896* | -0.675 | -0.921** | -0.587 | -0.681 | 0.131 | |||
R/S | -0.546 | -0.392 | -0.670 | 0.868 | 0.988* | 0.454 | 0.893* | 0.901* | 0.588 | 0.288 | 0.839 | -0.624 | |||
α | 0.379 | 0.124 | 0.906* | -0.791 | -0.644 | -0.766 | 0.221 | 0.435 | -0.147 | -0.688 | -0.904* | -0.260 |
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