气孔对土壤和空气湿度的反应及其模拟
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摘要
陆生植物在进行光合作用时通过叶片上的气孔吸入二氧化碳，但同时体内水分也从气孔向外散逸即气孔蒸腾。对叶的水分代谢来说，蒸腾是需求，根茎输送水分是供应。供水动力源于蒸腾对水的需求，供应受需求控制；另一方面，需求依赖供应并且受到有限供应的制约。植物通过调控气孔导度来调控需求。土壤湿度和空气湿度是影响气孔导度的重要环境条件。
气孔对空气湿度的反应本质上是气孔对蒸腾速率的反应，现象上可以分成三类。Buckley等人的模型和Dewar的模型都能模拟气孔对叶表水汽压亏缺（D_s）的普遍的A类反应：在D_s逐渐增大的过程中，气孔导度随着蒸腾速率的增大线性减小。但是在Buckley等人的模型中土壤到叶片表皮的有效水力学阻力（R）增大会使斜率绝对值增大，而在Dewar的模型中斜率与表皮细胞到保卫细胞的水力学阻力有关，与R无关。在这个问题上，有实验支持Buckley等人的模型。植物对土壤干旱最灵敏的适应反应之一是气孔导度减小。根源化学信号理论和水力学信号（hydraulic signal）理论各自都有实验依据，但都存在不足，虽然对立但是可以也应该统一起来。Buckley等人的模型提供了一个建立全面系统的气孔导度模型的基础。
Buckley等人的模型，其核心假设是水力-主动局部反馈假说：保卫细胞对表皮上保卫细胞紧邻区域水分状况的变化有主动调节胞内渗透压的反应。文献中以蚕豆为材料的实验支持这一假说。但是Grantz和Schwartz以鸭跖草为材料的实验不支持这一假说。他们把离体表皮浸泡在较高浓度的KCl溶液中，气氛为空气。本文的实验中，离体表皮浸泡在40mM KCl溶液中，气氛为除去CO_2的空气。施加渗透胁迫时，甘露醇通过溶液替换的方式加入溶液从而始终保持除去CO_2的空气气氛。结果表明保卫细胞对渗透胁迫有减少胞内渗透物质积累的主动反应。ABA大幅增强这一主动反应。并且，在ABA的作用下已张开气孔的保卫细胞对渗透胁迫有主动降低胞内渗透压的反应。这些结果支持水力-主动局部反馈假说，并且提示在模型中引入ABA反应时，ABA应该增强保卫细胞对表皮水势的敏感性。
Buckley等人的模型有很强的包容能力，在其基础上可以引入的因素包括：根系和地上部ABA的合成、ABA与水力学信号的交互作用、ABA之外的其它化学信号、pH对ABA运动的影响、包括木质部栓塞在内的植株水导变化、植株体内储水等。本文尝试在模型中引入根源ABA信号以及ABA与水力学信号的交互作用，新模型能够模拟等水行为（isohydric behavior）和气孔对D_s的C类反应：气孔导度不随着D_s和蒸腾速率的增大而减小。接着再引入木质部栓塞，新模型可以模拟气孔对D_s的B类反应：气孔导度和蒸腾速率均随着D_s的增大而减小。适度的木质部栓塞还可以促进气孔关闭，增强叶水势的稳定性。
Terrestrial plants inhale CO_2through stomata on leaves in photosynthesis, andmeanwhile water dissipate out of stomata as stomatal transpiration. Water consumed bytranspiration is supplied by the root and stem. The driving force for the water supplyderives from transpiration, i.e. water supply is controlled by water demand. On the otherhand, transpiration relies on water supply that is usually restricted. Plants regulate stomatalconductance to control transpiration. Soil humidity and air humidity are two importantenvironmental factors relative to stomatal regulation.
Stomata respond to transpiration rate rather than air humidity per se. The model ofBuckley et al. and Dewar’s model both can model the general stomatal response to watervapor deficit at leaf surface (D_s): stomatal conductance decreases linearly as transpirationrate increases because of rising D_s. The absolute value of the slope increases with theeffective hydraulic resistance from soil to leaf epidermis in the model of Buckley et al. InDewar’s model, however, the absolute value of the slope is determined by the hydraulicresistance from epidermal cells to guard cells instead. Several researches support the modelof Buckley et al. in this regard.
The most sensitive response of plants to drought is stomatal closing. As regards themechanism of this response, there are two contrasting theories: the theory of root-sourcedchemical signal and that of hydraulic signal. Both of them have convincing evidence whileneither of them is complete. They should be integrated. The model of Buckley et al. offersa basis for an integrated model of stomatal conductance.
The core hypothesis of the model of Buckley et al. is “hydro-active local feedback”:guard cell osmotic pressure is actively regulated in response to water status in theimmediate vicinity of guard cells in the epidermis. Evidence had indicated that guard cellsof Vicia faba respond actively to osmotic stress. However, Grantz and Schwartz found thatguard cells of Commelina communis L. did not respond actively to osmotic stress in detached epidermis. They had incubated detached epidermis in relatively highconcentration of KCl in ambient air. We incubated detached epidermis in40mM KCl inCO_2-free air. Mannitol was added by solution replacement without interrupting CO_2-freeair. Solute accumulation in guard cells was restricted in response to osmotic stress. ABAgreatly enhanced this active response. Furthermore, ABA stimulated active response ofguard cells of initially open stomata to osmotic stress: guard cell osmotic pressure wasreduced actively. Our results support “hydro-active local feedback” hypothesis, andsuggest that ABA should enhance guard cell sensitivity to epidermal water potential ifABA is to be introduced into the model.
The model of Buckley et al. can accommodate physiological properties concerningstomatal regulation, such as ABA synthesis in the shoot and the root, the interaction ofABA and hydraulic signal, some potential chemical signals, the effect of pH on themovement of ABA, changes in the hydraulic conductance of plants including xylemembolism, and water storage in plants. We introduced root-sourced ABA and theinteraction between ABA and hydraulic signal into the model, and successfully modeledisohydric behavior and stomatal response to D_sin Regime C. After xylem embolism wasintroduced in addition, stomatal response to D_sin Regime B was modeled. Simulation alsoshowed that moderate xylem embolism enhanced stomatal closing and the homeostasis ofleaf water potential.
气孔对空气湿度的反应本质上是气孔对蒸腾速率的反应，现象上可以分成三类。Buckley等人的模型和Dewar的模型都能模拟气孔对叶表水汽压亏缺（D_s）的普遍的A类反应：在D_s逐渐增大的过程中，气孔导度随着蒸腾速率的增大线性减小。但是在Buckley等人的模型中土壤到叶片表皮的有效水力学阻力（R）增大会使斜率绝对值增大，而在Dewar的模型中斜率与表皮细胞到保卫细胞的水力学阻力有关，与R无关。在这个问题上，有实验支持Buckley等人的模型。植物对土壤干旱最灵敏的适应反应之一是气孔导度减小。根源化学信号理论和水力学信号（hydraulic signal）理论各自都有实验依据，但都存在不足，虽然对立但是可以也应该统一起来。Buckley等人的模型提供了一个建立全面系统的气孔导度模型的基础。
Buckley等人的模型，其核心假设是水力-主动局部反馈假说：保卫细胞对表皮上保卫细胞紧邻区域水分状况的变化有主动调节胞内渗透压的反应。文献中以蚕豆为材料的实验支持这一假说。但是Grantz和Schwartz以鸭跖草为材料的实验不支持这一假说。他们把离体表皮浸泡在较高浓度的KCl溶液中，气氛为空气。本文的实验中，离体表皮浸泡在40mM KCl溶液中，气氛为除去CO_2的空气。施加渗透胁迫时，甘露醇通过溶液替换的方式加入溶液从而始终保持除去CO_2的空气气氛。结果表明保卫细胞对渗透胁迫有减少胞内渗透物质积累的主动反应。ABA大幅增强这一主动反应。并且，在ABA的作用下已张开气孔的保卫细胞对渗透胁迫有主动降低胞内渗透压的反应。这些结果支持水力-主动局部反馈假说，并且提示在模型中引入ABA反应时，ABA应该增强保卫细胞对表皮水势的敏感性。
Buckley等人的模型有很强的包容能力，在其基础上可以引入的因素包括：根系和地上部ABA的合成、ABA与水力学信号的交互作用、ABA之外的其它化学信号、pH对ABA运动的影响、包括木质部栓塞在内的植株水导变化、植株体内储水等。本文尝试在模型中引入根源ABA信号以及ABA与水力学信号的交互作用，新模型能够模拟等水行为（isohydric behavior）和气孔对D_s的C类反应：气孔导度不随着D_s和蒸腾速率的增大而减小。接着再引入木质部栓塞，新模型可以模拟气孔对D_s的B类反应：气孔导度和蒸腾速率均随着D_s的增大而减小。适度的木质部栓塞还可以促进气孔关闭，增强叶水势的稳定性。
Terrestrial plants inhale CO_2through stomata on leaves in photosynthesis, andmeanwhile water dissipate out of stomata as stomatal transpiration. Water consumed bytranspiration is supplied by the root and stem. The driving force for the water supplyderives from transpiration, i.e. water supply is controlled by water demand. On the otherhand, transpiration relies on water supply that is usually restricted. Plants regulate stomatalconductance to control transpiration. Soil humidity and air humidity are two importantenvironmental factors relative to stomatal regulation.
Stomata respond to transpiration rate rather than air humidity per se. The model ofBuckley et al. and Dewar’s model both can model the general stomatal response to watervapor deficit at leaf surface (D_s): stomatal conductance decreases linearly as transpirationrate increases because of rising D_s. The absolute value of the slope increases with theeffective hydraulic resistance from soil to leaf epidermis in the model of Buckley et al. InDewar’s model, however, the absolute value of the slope is determined by the hydraulicresistance from epidermal cells to guard cells instead. Several researches support the modelof Buckley et al. in this regard.
The most sensitive response of plants to drought is stomatal closing. As regards themechanism of this response, there are two contrasting theories: the theory of root-sourcedchemical signal and that of hydraulic signal. Both of them have convincing evidence whileneither of them is complete. They should be integrated. The model of Buckley et al. offersa basis for an integrated model of stomatal conductance.
The core hypothesis of the model of Buckley et al. is “hydro-active local feedback”:guard cell osmotic pressure is actively regulated in response to water status in theimmediate vicinity of guard cells in the epidermis. Evidence had indicated that guard cellsof Vicia faba respond actively to osmotic stress. However, Grantz and Schwartz found thatguard cells of Commelina communis L. did not respond actively to osmotic stress in detached epidermis. They had incubated detached epidermis in relatively highconcentration of KCl in ambient air. We incubated detached epidermis in40mM KCl inCO_2-free air. Mannitol was added by solution replacement without interrupting CO_2-freeair. Solute accumulation in guard cells was restricted in response to osmotic stress. ABAgreatly enhanced this active response. Furthermore, ABA stimulated active response ofguard cells of initially open stomata to osmotic stress: guard cell osmotic pressure wasreduced actively. Our results support “hydro-active local feedback” hypothesis, andsuggest that ABA should enhance guard cell sensitivity to epidermal water potential ifABA is to be introduced into the model.
The model of Buckley et al. can accommodate physiological properties concerningstomatal regulation, such as ABA synthesis in the shoot and the root, the interaction ofABA and hydraulic signal, some potential chemical signals, the effect of pH on themovement of ABA, changes in the hydraulic conductance of plants including xylemembolism, and water storage in plants. We introduced root-sourced ABA and theinteraction between ABA and hydraulic signal into the model, and successfully modeledisohydric behavior and stomatal response to D_sin Regime C. After xylem embolism wasintroduced in addition, stomatal response to D_sin Regime B was modeled. Simulation alsoshowed that moderate xylem embolism enhanced stomatal closing and the homeostasis ofleaf water potential.
引文
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