Photosynthetic limitation in soybean in response to soil water-nitrogen interactions and its relationship with leaf water use efficiency
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摘要:
本文以我国北方旱地大豆（Glycine max （L.） Merr.）为材料，以光合限制作用为研究对象，设计土壤水-氮交互实验，探究其光合限制作用的水-氮交互响应及其与叶片瞬时水分利用效率（WUEins）和内在水分利用效率（WUEint）的相关性。结果显示：（1）大豆叶内CO2扩散性限制作用（叶肉限制，lm；气孔限制，lsc）随水分胁迫逐渐增大，生化限制（lb）逐渐减小；（2）土壤水-氮交互后，植株lm和lsc均减小，而lb显著增大；（3）lm和lb分别与WUEins及WUEint呈显著的负、正相关（P<0.05），而lsc与二者的相关性不显著；（4）相较lsc和lb，lm对植株光合碳同化与叶片水分利用效率的贡献率最大，为大豆光合与水分利用能力变化的主导限制因子。研究结果旨在揭示土壤干旱与大气氮沉降双因子交互调控作物光合碳同化的生态效应，并分析该效应与作物水分利用能力的内在关系。
Abstract:Glycine max (L.) Merr. in northern China were selected as the experimental materials, with photosynthetic limitation as the primary focus of this study. A soil water-nitrogen interaction experiment was designed to explore the relationship between photosynthetic limitation and leaf instantaneous (WUEins) and intrinsic water use efficiency (WUEint). Results showed that: (1) CO2 diffusion limitations, including mesophyll limitation (lm) and stomatal limitation (lsc), gradually increased with water stress, while biochemical limitation (lb) gradually decreased. (2) The interaction between water and nitrogen reduced lm and lsc, while lb significantly increased. (3) lm and lb were negatively and positively correlated with WUEins and WUEint, respectively (P<0.05), while lsc showed no significant correlation with either. (4) Among the limiting factors, lm contributed most to soybean photosynthetic carbon assimilation and leaf water use efficiency, making it the dominant constraint on photosynthetic and water use capacity. These findings reveal the ecological effects of soil drought and atmospheric nitrogen deposition on crop photosynthetic carbon assimilation and their intrinsic relationship with water use capacity.
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图 1 不同控水处理土壤水势（Ψsoil）变化
CK为对照，MW为轻度水分胁迫，SW为重度水分胁迫。下同。不同小写字母表示不同处理间差异显著（n=3，P<0.05）。
Figure 1. Changes in soil water potential (Ψsoil) under different water control treatments
CK: Control; MW: Moderate water stress; SW: Severe water stress. Same below. Lowercase letters indicate significant differences (n=3, P<0.05).
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图 2 大豆光合相对限制作用随土壤水、氮含量的变化
MWLN为轻度水分胁迫，低氮添加；MWMN为轻度水分胁迫，中氮添加；MWHN为轻度水分胁迫，高氮添加；SWLN为重度水分胁迫，低氮添加；SWMN为重度水分胁迫，中氮添加；SWHN为重度水分胁迫，高氮添加。下同。*表示不同处理间在P<0.05水平上（n=3）差异显著，ns表示在该水平下差异不显著。
Figure 2. Changes in relative limitations of gsc, gm, and biochemical capacity with soil water and nitrogen in Glycine max seedlings
MWLN: Moderate water stress×low N addition; MWMN: Moderate water stress×medium N addition; MWHN: Moderate water stress×high N addition; SWLN: Severe water stress×low N addition; SWMN: Severe water stress×medium N addition; SWHN: Severe water stress×high N addition. Same below. * indicates significant difference between treatments at P<0.05 (n=3), ns indicates no significant difference at this level.
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图 3 gsc、gm及生化能力对植株饱和光碳同化贡献率随土壤水、氮含量的变化
SCL、MCL和BL分别为gsc、gm和生化能力对叶片dAn/An的贡献率。
Figure 3. Changes in contributions of gsc, gm, and biochemical capacity to light-saturated assimilation with soil water and nitrogen in Glycine max seedlings
SCL, MCL, and BL are contributions of gsc, gm, and biochemical capacity to dAn/An, respectively.
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图 4 WUEins和WUEint分别与lm（A, B）、lsc（C, D）及lb（E, F）的相关性
Figure 4. Relationships of WUEins and WUEint with lm, lsc, and lb, respectively
表 1 叶片气孔形态特征值
Table 1 Anatomical morphological features of leaf stomata
处理
Treatment气孔密度
Ds / ind./mm2气孔大小
Ssize / μm2气孔开度
SS / μm2叶片厚度
Tleaf / μmCK 291.7±87.4a 141.2±34.4a 17.8±3.6a 192.7±24.1a MW 266.7±37.3a 141.4±42.2a 14.0±5.3ab 158.6±26.1b SW 250.0±96.2a 129.7±9.7a 8.6±1.5b 160.6±12.1b MWLN 208.3±45.0a 139.0±5.7a 11.3±2.2b 187.3±20.9ab MWMN 215.3±42.9a 147.6±37.5a 7.8±2.0bc 182.6±9.7ab MWHN 233.8±34.2a 103.7±10.8a 4.3±1.2d 192.2±18.8a SWLN 277.8±43.0a 130.4±15.8a 4.6±1.6cd 171.9±16.0ab SWMN 263.9±34.0a 145.3±16.4a 6.1±0.9cd 196.4±12.8a SWHN 222.2±96.2a 114.1±32.6a 2.2±0.1e 170.6±16.1ab 注：MWLN为轻度水分胁迫，低氮添加；MWMN为轻度水分胁迫，中氮添加；MWHN为轻度水分胁迫，高氮添加；SWLN为重度水分胁迫，低氮添加；SWMN为重度水分胁迫，中氮添加；SWHN为重度水分胁迫，高氮添加。同列不同小写字母表示处理间在P<0.05水平上差异显著。下同。 Notes: MWLN: Moderate water stress×low N addition; MWMN: Moderate water stress×medium N addition; MWHN: Moderate water stress×high N addition; SWLN: Severe water stress×low N addition; SWMN: Severe water stress×medium N addition; SWHN: Severe water stress×high N addition.Different lowercase letters in same column indicate significant differences at P<0.05 level. Same below. 
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表 2 不同水氮条件下土壤全氮、硝态氮、全磷、全钾、速效钾含量
Table 2 Key soil nutrient content of nitrate N, total P, total K, and available K under different water and nitrogen conditions
处理
Treatment全氮
Total nitrogen / g/kg硝态氮
Nitrate nitrogen / mg/kg全磷
Total P / g/kg全钾
Total K / g/kg速效钾
Available K / mg/kgCK 1.12±0.00a 12.4±0.3d 2.49±0.18b 30.3±1.9c 157.4±5.3d MW 1.06±0.15a 11.2±0.5d 2.37±0.15bc 29.5±1.2c 150.4±2.1d SW 1.02±0.14ab 11.0±0.4d 2.22±0.08c 29.0±1.8c 147.4±4.0d MWLN 1.26±0.14a 33.2±0.4c 2.92±0.06a 33.3±0.1b 200.7±30.9c MWMN 0.84±0.00bc 49.1±13.6b 2.91±0.02a 32.7±0.4b 163.3±3.8d MWHN 1.12±0.28a 58.6±4.6a 2.87±0.12a 35.3±0.7a 289.7±5.9a SWLN 1.12±0.00a 44.8±1.9b 2.79±0.29a 32.8±0.5b 239.9±13.2b SWMN 0.70±0.14c 55.3±2.4ab 2.91±0.16a 32.8±0.8b 222.9±6.3bc SWHN 1.12±0.00a 51.1±3.8ab 2.83±0.03a 33.6±0.2ab 240.3±20.0b
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