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Construction of the Model for Soil Moisture Effects on Leaf Photosynthesis Rate of Winter Wheat
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摘要: 植物叶片光合速率是表征植物光合能力的重要参数，对土壤水分反应敏感，建立不同土壤水分对冬小麦叶片光合速率影响模型，有助于准确理解冬小麦的光合作用和产量形成。该文收集整理了1996—2017年我国冬小麦主产区11个试验地点、17个冬小麦品种的干旱和渍水试验数据共64组310个样本，分别构建干旱和渍水对冬小麦叶片光合速率影响的分段式和指数型模型，形成土壤水分对冬小麦叶片光合速率影响模型（the model for Soil Moisture Effects on leaf Photosynthesis rate of winter wheat，SMEP）。结果表明：随着土壤相对湿度增加，冬小麦叶片光合速率系数呈稳定低值-线性增加-稳定高值-缓慢下降的特点；随着渍水时间延长，冬小麦叶片光合速率系数呈缓慢下降-快速下降的特点。对SMEP模型进行回代检验、外推检验、单点验证、单发育期验证发现，模型模拟结果与文献数据有较好的一致性，回归系数在1.0附近，且均达到0.01显著性水平。SMEP模型将嵌入中国农业气象模式（CAMM1.0），为CAMM不断完善提供科技支撑。Abstract: The rate of leaf photosynthesis, which is sensitive to soil moisture, is one of the most important parameters to characterize the photosynthetic capacity of plants. Constructing a model which can reveal effects of soil moisture on leaf photosynthesis rate of winter wheat will be helpful to accurately understand the photosynthesis and yield formation. A total of 310 photosynthesis rate samples under different soil moistures, including 227 drought stress samples in 50 tests and 83 waterlogging stress samples in 14 tests, are jointly collected from 17 winter wheat cultivars at 11 experimental sites via the published references. However, photosynthesis rates of winter wheat are quite different between different cultivars, different developmental stages, and different experimental sites. Normalized photosynthesis rate coefficients for winter wheat are derived by calculating the ratio of leaf photosynthesis rate under different water stresses and CK. And then, segmental and exponential models are established for effects of drought and waterlogging stresses on leaf photosynthesis rate of winter wheat, respectively. The model for soil moisture effects on leaf photosynthesis rate of winter wheat (SMEP) is correspondingly constructed. Photosynthesis rate coefficients of winter wheat leaves show the trend of "stable low value-linear increase-stable high value-slow decrease" with the increase of soil relative moisture. Meanwhile, photosynthesis rate coefficients exhibit characteristics of "slow decline-rapid decline" with the prolongation of waterlogging stress. Four tests, including back-training test, extrapolation test, single-site test and certain developmental stage test, are also done to validate the SMEP model. Generally, the results simulated by the SMEP model are in good agreement with the records in the literatures. The linear regression coefficients are all around 1.0, and the regression equations all pass the significant test of 0.01. SMEP model will be coupled to Chinese Agro-Meteorological Model (CAMM1.0), providing scientific and technological supports for the continuous improvement of CAMM1.0.
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图 1 文献收集水分胁迫试验数据的站点分布
Fig. 1 Site distribution for water stress experiments reported in references
图 2 土壤相对湿度对冬小麦叶片光合速率影响模型示意图
Fig. 2 Schematic diagram of soil relative moisture effects on leaf photosynthesis rate of winter wheat
图 3 干旱对冬小麦叶片光合速率的影响
(a)冬小麦叶片光合速率系数随土壤相对湿度的变化动态,
(b)文献数据和SMEP模型模拟的冬小麦叶片光合速率系数对比验证Fig. 3 Effects of drought stresses on leaf photosynthesis rate of winter wheat
(a)dynamics of photosynthesis rate coefficients with soil relative moisture,
(b)comparisons of photosynthesis rate coefficients between records from references and SMEP simulations
图 4 渍水持续时间对冬小麦叶片光合速率的影响
(a)冬小麦叶片光合速率系数随渍水持续日数的变化动态,
(b)文献数据和SMEP模型模拟的冬小麦叶片光合速率系数对比验证Fig. 4 Effect of waterlogging duration on leaf photosynthesis rate of winter wheat
(a)dynamics of photosynthesis rate coefficients with waterlogging durations,
(b)comparisons of photosynthesis rate coefficients between records from references and SMEP simulations
图 6 干旱对冬小麦叶片光合速率系数的外推检验结果
(a)光合速率系数随土壤相对湿度变化, (b)文献数据与SMEP模型模拟的光合速率系数对比
Fig. 6 Independent validations of photosynthesis rate coefficients of winter wheat leaves under drought stresses
(a)dynamics of photosynthesis rate coefficients with soil relative moisture,
(b)comparisons of photosynthesis rate coefficients between records from references and SMEP simulations
图 7 渍水对冬小麦叶片光合速率系数的集合验证结果
(a)光合速率系数随渍水持续日数变化, (b)文献数据与SMEP模型模拟的光合速率系数对比
Fig. 7 Independent validations of photosynthesis rate coefficients for winter wheat leaves under waterlogging stresses
(a)dynamics of photosynthesis rate coefficients with waterlogging duration,
(b)comparisons of photosynthesis rate coefficients between records from references and SMEP simulations
图 8 栾城(a)、新乡(b)干旱和合肥(c)渍水下冬小麦叶片光合速率系数单点验证结果
Fig. 8 Single-site validations of photosynthesis rate coefficients for winter wheat leaves under drought and waterlogging stresses (a)drought stress at Luancheng, (b)drought stress at Xinxiang, (c)waterlogging stress at Hefei
图 9 冬小麦拔节期(a)和灌浆期(b)光合速率系数随土壤相对湿度变化动态
Fig. 9 Dynamics of photosynthesis rate coefficients for winter wheat leaves with soil relative moisture at jointing(a) and booting(b) stages
表 1 文献收集的水分胁迫条件下冬小麦旗叶叶片光合速率数据汇总
Table 1 Summary of photosynthesis rate for winter wheat flag leaves under different water stresses reported in references
胁迫试验 试验品种 试验地点 省份 组数 样本量 发育阶段 光强/(μmol·m-2·s-1) 文献 新冬2 阜康 新疆 1 5 灌浆期 1400 [28] 高优503 栾城 山东 5 25 拔节-蜡熟期 [29] 高优503 栾城 山东 8 39 拔节-灌浆期 [30] 小偃22 杨凌 陕西 1 4 三叶期 [31] 京麦9428 大兴 北京 1 2 灌浆期 饱和光强 [32] 扬麦9 南京 江苏 4 12 灌浆期 [33] 周麦27 鹤壁 河南 3 9 灌浆期 [34] 干旱 山农20 泰安 山东 1 4 灌浆期 [35] 山农21 泰安 山东 1 4 灌浆期 [35] 扬麦10 南京 江苏 5 43 起身-灌浆期 1000±20 [18] 鲁麦7 莱阳 山东 7 27 孕穗-灌浆期 [36] 郑麦98 新乡 河南 3 15 拔节期 [37] 长武 陕西 9 27 [38] 鲁麦23 泰安 山东 1 11* 拔节期 饱和光强 [20] 烟农19 合肥 安徽 4 28 开花-灌浆期 [25] 汶农17 南京 江苏 1 3 开花-灌浆期 [39] 扬麦16 南京 江苏 1 3 开花-灌浆期 [39] 渍水 豫麦34 南京 江苏 1 2 开花-灌浆期 1100 [24] 扬麦9 南京 江苏 1 2 开花-灌浆期 1100 [24] 扬麦13 南京 江苏 1 4 开花-灌浆期 1100 [40] 扬麦10 南京 江苏 5 41* 起身-灌浆期 1000±20 [18] 注：*表示建模数据，其余为验证数据；光强数值空缺表示测量条件是晴朗无云的09:00—11:00(北京时，下同)。 
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表 2 水分胁迫下文献数据与SMEP模型模拟的站点尺度冬小麦叶片光合速率系数统计
Table 2 Photosynthesis rate coefficients for winter wheat leaves between records from references and SMEP simulations
站点 线性回归方程 决定系数 均方根误差 样本量 栾城 y=1.3219x-0.3130 0.9791 0.005 5 新乡 y=0.9922x+0.0182 0.9342 0.003 5 合肥 y=0.9008x-0.0299 0.9078 0.018 7 注：x为文献数据，y为SMEP模型模拟结果，且所有方程均达到0.01显著性水平。
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表 3 水分胁迫下文献数据与SMEP模型模拟的冬小麦拔节和灌浆期叶片光合速率系数
Table 3 Photosynthesis rate coefficients for winter wheat leaves between records from references and SMEP simulations at jointing and booting stages
发育期 线性回归方程 决定系数 均方根误差 样本量 拔节期 y=1.0753x-0.0860 0.8372 0.0067 9 灌浆期 y=0.9545x+0.0255 0.6106 0.0113 12
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