植物生态学报 ›› 2012, Vol. 36 ›› Issue (12): 1277-1285.DOI: 10.3724/SP.J.1258.2012.01277
陈卫英1,2,*(
), 陈真勇1, 罗辅燕1, 彭正松1, 余懋群2
收稿日期:2012-09-11
接受日期:2012-11-01
出版日期:2012-09-11
发布日期:2012-11-28
通讯作者:
陈卫英
作者简介:* E-mail: chen_weiy@163.com
CHEN Wei-Ying1,2,*(
), CHEN Zhen-Yong1, LUO Fu-Yan1, PENG Zheng-Song1, YU Mao-Qun2
Received:2012-09-11
Accepted:2012-11-01
Online:2012-09-11
Published:2012-11-28
Contact:
CHEN Wei-Ying
摘要:
光响应曲线的参数是研究植物生理状态的重要指标, 常用的光响应曲线模型无法准确地计算出光饱和点和最大净光合速率。该文利用光响应曲线新模型——指数改进模型、直角双曲线模型、直角双曲线修正模型、非直角双曲线模型和指数模型, 拟合高粱(Sorghum bicolor)、苋(Amaranthus tricolor)、大麦(Hordeum vulgare)和半夏(Pinellia ternata)的光响应曲线, 并随机选取部分数据进行检验, 得到了各模型计算出的主要生理参数, 并对这些数据进行了比较分析, 讨论了各模型之间的优缺点和准确性, 描述了C3、C4植物光响应的适宜性。结果表明, 基于C3植物得到的指数改进模型和直角双曲线修正模型能较准确地计算出C3、C4植物饱和光强和最大净光合速率, 并在描述光响应曲线时比另外3个模型具有更高的精确性和适宜性。实验结果可为光响应曲线模型在C3和C4光合途径植物中的应用提供参考。
陈卫英, 陈真勇, 罗辅燕, 彭正松, 余懋群. 光响应曲线的指数改进模型与常用模型比较. 植物生态学报, 2012, 36(12): 1277-1285. DOI: 10.3724/SP.J.1258.2012.01277
CHEN Wei-Ying, CHEN Zhen-Yong, LUO Fu-Yan, PENG Zheng-Song, YU Mao-Qun. Comparison between modified exponential model and common models of light-response curve. Chinese Journal of Plant Ecology, 2012, 36(12): 1277-1285. DOI: 10.3724/SP.J.1258.2012.01277
| 材料 Material | 模型 Model | 拟合值的MSE Fitted MSE | 拟合值的MAE Fitted MAE | 预测值的MSE Test MSE | 预测值的MAE Test MAE |
|---|---|---|---|---|---|
| 高粱 Sorghum bicolor | 指数改进模型 Modified exponential model | 0.092 | 0.239 | 0.070 | 0.239 |
| 直角双曲线修正模型 Modified rectangular hyperbola model | 0.159 | 0.320 | 0.112 | 0.295 | |
| 直角双曲线模型 Rectangular hyperbola model | 0.329 | 0.476 | 0.363 | 0.527 | |
| 非直角双曲线模型 Nonrectangular hyperbola model | 0.052 | 0.192 | 0.217 | 0.383 | |
| 指数模型 Exponential model | 0.098 | 0.243 | 0.065 | 0.204 | |
| 苋 Amaranthus tricolor | 指数改进模型 Modified exponential model | 0.043 | 0.186 | 0.119 | 0.272 |
| 直角双曲线修正模型 Modified rectangular hyperbola model | 0.049 | 0.186 | 0.208 | 0.357 | |
| 直角双曲线模型 Rectangular hyperbola model | 0.325 | 0.458 | 1.113 | 0.757 | |
| 非直角双曲线模型 Nonrectangular hyperbola model | 0.194 | 0.363 | 0.217 | 0.409 | |
| 指数模型 Exponential model | 0.053 | 0.215 | 0.235 | 0.320 | |
| 半夏 Pinellia ternata | 指数改进模型 Modified exponential model | 0.116 | 0.288 | 0.404 | 0.308 |
| 直角双曲线修正模型 Modified rectangular hyperbola model | 0.042 | 0.160 | 0.190 | 0.235 | |
| 直角双曲线模型 Rectangular hyperbola model | 0.494 | 0.569 | 1.569 | 0.912 | |
| 非直角双曲线模型 Nonrectangular hyperbola model | 0.261 | 0.402 | 1.026 | 0.628 | |
| 指数模型 Exponential model | 0.210 | 0.295 | 0.919 | 0.514 | |
| 大麦‘藏青320’ Hordeum vulgare ‘zangqing320’ | 指数改进模型 Modified exponential model | 0.070 | 0.237 | 0.250 | 0.307 |
| 直角双曲线修正模型 Modified rectangular hyperbola model | 0.023 | 0.122 | 0.107 | 0.214 | |
| 直角双曲线模型 Rectangular hyperbola model | 0.548 | 0.643 | 1.490 | 0.778 | |
| 非直角双曲线模型 Nonrectangular hyperbola model | 0.192 | 0.324 | 0.855 | 0.626 | |
| 指数模型 Exponential model | 0.114 | 0.224 | 0.586 | 0.453 |
表1 由均方误差(MSE)和平均绝对误差(MAE)表示的5个模型的精确度
Table 1 The accuracy of five models through the values of mean square error (MSE) and mean absolute error (MAE)
| 材料 Material | 模型 Model | 拟合值的MSE Fitted MSE | 拟合值的MAE Fitted MAE | 预测值的MSE Test MSE | 预测值的MAE Test MAE |
|---|---|---|---|---|---|
| 高粱 Sorghum bicolor | 指数改进模型 Modified exponential model | 0.092 | 0.239 | 0.070 | 0.239 |
| 直角双曲线修正模型 Modified rectangular hyperbola model | 0.159 | 0.320 | 0.112 | 0.295 | |
| 直角双曲线模型 Rectangular hyperbola model | 0.329 | 0.476 | 0.363 | 0.527 | |
| 非直角双曲线模型 Nonrectangular hyperbola model | 0.052 | 0.192 | 0.217 | 0.383 | |
| 指数模型 Exponential model | 0.098 | 0.243 | 0.065 | 0.204 | |
| 苋 Amaranthus tricolor | 指数改进模型 Modified exponential model | 0.043 | 0.186 | 0.119 | 0.272 |
| 直角双曲线修正模型 Modified rectangular hyperbola model | 0.049 | 0.186 | 0.208 | 0.357 | |
| 直角双曲线模型 Rectangular hyperbola model | 0.325 | 0.458 | 1.113 | 0.757 | |
| 非直角双曲线模型 Nonrectangular hyperbola model | 0.194 | 0.363 | 0.217 | 0.409 | |
| 指数模型 Exponential model | 0.053 | 0.215 | 0.235 | 0.320 | |
| 半夏 Pinellia ternata | 指数改进模型 Modified exponential model | 0.116 | 0.288 | 0.404 | 0.308 |
| 直角双曲线修正模型 Modified rectangular hyperbola model | 0.042 | 0.160 | 0.190 | 0.235 | |
| 直角双曲线模型 Rectangular hyperbola model | 0.494 | 0.569 | 1.569 | 0.912 | |
| 非直角双曲线模型 Nonrectangular hyperbola model | 0.261 | 0.402 | 1.026 | 0.628 | |
| 指数模型 Exponential model | 0.210 | 0.295 | 0.919 | 0.514 | |
| 大麦‘藏青320’ Hordeum vulgare ‘zangqing320’ | 指数改进模型 Modified exponential model | 0.070 | 0.237 | 0.250 | 0.307 |
| 直角双曲线修正模型 Modified rectangular hyperbola model | 0.023 | 0.122 | 0.107 | 0.214 | |
| 直角双曲线模型 Rectangular hyperbola model | 0.548 | 0.643 | 1.490 | 0.778 | |
| 非直角双曲线模型 Nonrectangular hyperbola model | 0.192 | 0.324 | 0.855 | 0.626 | |
| 指数模型 Exponential model | 0.114 | 0.224 | 0.586 | 0.453 |
| 材料 Material | 模型 Model | LCP | LSP | Pmax | Rdark | φ0 | θ |
|---|---|---|---|---|---|---|---|
| 高粱 Sorghum bicolor | 观测值 Measured value | 40-50 | 2 200 | 19.362 | 1.616 | - | - |
| 指数改进模型 Modified exponential model | 42.815 | 1 952 | 16.944 | 1.911 | 0.047 | - | |
| 直角双曲线修正模型 Modified rectangular hyperbola model | 41.033 | 2 071 | 18.873 | 1.981 | 0.051 | - | |
| 直角双曲线模型 Rectangular hyperbola model | 40.600 | - | 26.028 | 2.347 | 0.064 | - | |
| 非直角双曲线模型 Nonrectangular hyperbola model | 41.398 | - | 22.043 | 1.585 | 0.039 | 0.776 | |
| 指数模型 Exponential model | 41.438 | - | 20.892 | 1.835 | 0.046 | - | |
| 苋 Amaranthus tricolor | 观测值 Measured value | 80-100 | 2 200 | 39.618 | 5.626 | - | - |
| 指数改进模型 Modified exponential model | 95.385 | 2 295 | 40.720 | 5.390 | 0.060 | - | |
| 直角双曲线修正模型 Modified rectangular hyperbola model | 94.607 | 2 574 | 40.099 | 5.484 | 0.062 | - | |
| 直角双曲线模型 Rectangular hyperbola model | 90.161 | - | 65.378 | 6.135 | 0.075 | - | |
| 非直角双曲线模型 Nonrectangular hyperbola model | 97.738 | - | 50.000 | 4.719 | 0.049 | 0.809 | |
| 指数模型 Exponential model | 94.924 | - | 48.382 | 5.502 | 0.062 | - | |
| 半夏 Pinellia ternata | 观测值 Measured value | 0-20 | 1 200 | 11.762 | 0.630 | - | - |
| 指数改进模型 Modified exponential model | 8.012 | 967 | 11.619 | 0.422 | 0.054 | - | |
| 直角双曲线修正模型 Modified rectangular hyperbola model | 9.839 | 1 122 | 11.729 | 0.610 | 0.064 | - | |
| 直角双曲线模型 Rectangular hyperbola model | 11.167 | - | 13.424 | 0.987 | 0.095 | - | |
| 非直角双曲线模型 Nonrectangular hyperbola model | 8.680 | - | 11.822 | 0.392 | 0.045 | 0.906 | |
| 指数模型 Exponential model | 10.910 | - | 11.882 | 0.651 | 0.061 | - | |
| 大麦‘藏青320’ Hordeum vulgare ‘zangqing320’ | 观测值 Measured value | 20-40 | 1 600 | 19.953 | 1.399 | - | - |
| 指数改进模型 Modified exponential model | 21.611 | 1 577 | 19.601 | 1.150 | 0.055 | - | |
| 直角双曲线修正模型 Modified rectangular hyperbola model | 23.327 | 1 580 | 19.743 | 1.386 | 0.062 | - | |
| 直角双曲线模型 Rectangular hyperbola model | 25.536 | - | 25.439 | 2.061 | 0.088 | - | |
| 非直角双曲线模型 Nonrectangular hyperbola model | 23.828 | - | 22.182 | 1.237 | 0.053 | 0.768 | |
| 指数模型 Exponential model | 23.897 | - | 21.083 | 1.381 | 0.060 | - |
表2 不同模型计算出的4种材料的生理参数和观测值的比较
Table 2 Comparison of photosynthetic parameters calculated by the fitted formulations and measured values
| 材料 Material | 模型 Model | LCP | LSP | Pmax | Rdark | φ0 | θ |
|---|---|---|---|---|---|---|---|
| 高粱 Sorghum bicolor | 观测值 Measured value | 40-50 | 2 200 | 19.362 | 1.616 | - | - |
| 指数改进模型 Modified exponential model | 42.815 | 1 952 | 16.944 | 1.911 | 0.047 | - | |
| 直角双曲线修正模型 Modified rectangular hyperbola model | 41.033 | 2 071 | 18.873 | 1.981 | 0.051 | - | |
| 直角双曲线模型 Rectangular hyperbola model | 40.600 | - | 26.028 | 2.347 | 0.064 | - | |
| 非直角双曲线模型 Nonrectangular hyperbola model | 41.398 | - | 22.043 | 1.585 | 0.039 | 0.776 | |
| 指数模型 Exponential model | 41.438 | - | 20.892 | 1.835 | 0.046 | - | |
| 苋 Amaranthus tricolor | 观测值 Measured value | 80-100 | 2 200 | 39.618 | 5.626 | - | - |
| 指数改进模型 Modified exponential model | 95.385 | 2 295 | 40.720 | 5.390 | 0.060 | - | |
| 直角双曲线修正模型 Modified rectangular hyperbola model | 94.607 | 2 574 | 40.099 | 5.484 | 0.062 | - | |
| 直角双曲线模型 Rectangular hyperbola model | 90.161 | - | 65.378 | 6.135 | 0.075 | - | |
| 非直角双曲线模型 Nonrectangular hyperbola model | 97.738 | - | 50.000 | 4.719 | 0.049 | 0.809 | |
| 指数模型 Exponential model | 94.924 | - | 48.382 | 5.502 | 0.062 | - | |
| 半夏 Pinellia ternata | 观测值 Measured value | 0-20 | 1 200 | 11.762 | 0.630 | - | - |
| 指数改进模型 Modified exponential model | 8.012 | 967 | 11.619 | 0.422 | 0.054 | - | |
| 直角双曲线修正模型 Modified rectangular hyperbola model | 9.839 | 1 122 | 11.729 | 0.610 | 0.064 | - | |
| 直角双曲线模型 Rectangular hyperbola model | 11.167 | - | 13.424 | 0.987 | 0.095 | - | |
| 非直角双曲线模型 Nonrectangular hyperbola model | 8.680 | - | 11.822 | 0.392 | 0.045 | 0.906 | |
| 指数模型 Exponential model | 10.910 | - | 11.882 | 0.651 | 0.061 | - | |
| 大麦‘藏青320’ Hordeum vulgare ‘zangqing320’ | 观测值 Measured value | 20-40 | 1 600 | 19.953 | 1.399 | - | - |
| 指数改进模型 Modified exponential model | 21.611 | 1 577 | 19.601 | 1.150 | 0.055 | - | |
| 直角双曲线修正模型 Modified rectangular hyperbola model | 23.327 | 1 580 | 19.743 | 1.386 | 0.062 | - | |
| 直角双曲线模型 Rectangular hyperbola model | 25.536 | - | 25.439 | 2.061 | 0.088 | - | |
| 非直角双曲线模型 Nonrectangular hyperbola model | 23.828 | - | 22.182 | 1.237 | 0.053 | 0.768 | |
| 指数模型 Exponential model | 23.897 | - | 21.083 | 1.381 | 0.060 | - |
图1 高粱、苋、大麦和半夏的光响应曲线。模型中PAR为2400、1000、400、80、40、20 μmol·m-2·s-1时, 对应点为预测值, 其余各点为拟合值。
Fig. 1 Light-response curves of Sorghum bicolor, Amaranthus tricolor, Hordeum vulgare and Pinellia ternata. PAR of 2400, 1000, 400, 80, 40, 20 μmol·m-2·s-1 are predicted values, and the rests are fitted values.
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