植物整体水分平衡的生理生态调控机制研究
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摘要
水资源不足是干旱半干旱地区生态环境和农业生产的主要限制因素。植物水分平衡的生理生态基础研究是发展节水农业和生态建设的重要依据。植物可以通过调节根茎枝叶等部位的导水特性以维持整体水分平衡，而水通道蛋白的深入研究将有助于揭示植物水分关系的生理生态机制。本论文选用玉米（Zea mays L.）、小麦（Triticum spp.）和甜高粱（Sorghum bicolor (L.) Moench）为实验材料，通过人工控制环境条件下以聚乙二醇诱导形成短期或长期水分胁迫处理，研究了水通道蛋白转录调节、从细胞到整株各尺度导水特性响应以及叶片水氮利用能力变化。取得主要结果如下：
(1).恒定环境条件下玉米478的叶蒸腾速率、根导水率、叶导水率和整株导水率均表现明显日变化，且各参数间均存在显著正相关关系。这对于维持植株整体水分平衡是必要的。水分胁迫处理2小时后，玉米478根导水率显著降低而叶导水率却显著升高；叶片中ZmPIPs基因的转录量也显著增加，尤其是ZmPIP1;2；而根中仅ZmPIP2;5的转录水平显著下调，表明ZmPIPs可能参与了根叶导水特性的短期调节过程。
(2).干旱敏感玉米478的叶蒸腾速率呈明显的日变化规律，而抗旱性较强的天四较低且维持恒定；478在叶片水力结构、根叶形态和根导水率等方面均表现出适应高蒸腾耗水的特征；478和天四通过不同的叶内部水势差调节策略维持叶片水分平衡。短期水分胁迫下，478的根导水率下降而叶导水率增加，且叶蒸腾降低幅度却较小导致其体内水分亏缺；而天四的叶蒸腾速率大幅下降且根导水率在胁迫8小时后部分恢复。低叶蒸腾、水分胁迫后蒸腾迅速降低以及根导水率能部分恢复等特性可能是天四保持水分平衡的重要原因。
(3).正常供水天四和478的根叶中各ZmPIPs基因的转录水平存在明显差异，其中ZmPIP1;5在478中转录量较大而在天四中检测不到。PEG诱导水分胁迫处理2小时内，天四根中各ZmPIPs转录水平总体上调而叶片中总体下调，可能有助于其增加根系吸水同时减少叶片失水；478根中和叶片中各ZmPIPs的mRNA相对含量表现短暂的升高过程，这可能与其叶片蒸腾失水降低幅度较小有关。
(4).随着染色体倍性增加，小麦叶蒸腾失水和根系吸水能力均不断增大，而皮层细胞体积却逐渐减小。在正常供水或水分胁迫条件下，叶蒸腾速率、单根导水率、根细胞导水率以及TaPIPs转录水平之间均存在显著正相关关系。皮层细胞体积可能参与调节根径向水流各途径的相对贡献。在小麦进化过程中，根中TaPIP基因转录水平、细胞-细胞途径和质外体途径均显著增加，进而增强根系吸水能力以满足冠层耗水增加从而维持整株水分平衡。
(5).既然植物的基因表达、不同尺度导水特性和叶蒸腾之间存在一致性关系，那么叶片水分利用特性可能对整株耗水起控制作用。高抗旱性甜高粱对长期水分胁迫的适应过程中，叶片瞬时水分利用效率（WUE）显著增加，叶面积和单株耗水均显著降低，而干物质积累却变化不显著；叶氮积累显著降低；而光合氮利用效率（PNUE）和氮利用效率（NUE）显著增加。在干旱适应过程中，甜高粱叶片PNUE提高可能是WUE提高并减少植株耗水的原因之一。
本论文通过分析控制环境条件下所取得的实验数据，较深入探讨了植物水分平衡生理生态机制中的一些科学问题。研究结果有助于深化植物整体抗旱性的生物学基础的认识，为深入研究水-氮-碳三者间协调关系的生理机制提供基础，并为通过基因改良提高植物水分利用效率提供实验依据。
Water shortage often limits the ecological environment and agricultural productionin the arid and semi-arid area. Understanding the ecophysiological basis of highefficiency of plant water use is very important for developing water-saving agricultureand constructing ecological civilization. Plant can maintain water balance with variablehydraulic properties and aquaporins could be involved, which may help to understandthe ecophysiological mechanisms of plant water relations. This dissertation used thehydroponically grown maize (Zea mays L.), wheat (Triticum spp.) and sweet sorghum(Sorghum bicolor (L.) Moench) seedlings as experimental materials, and used cell-androot-pressure probes, high pressure flow meter and quantitive real-time PCR tomeasure the responses of cell, single root, root and shoot and whole-plant hydraulicsand aquaporin genes transcription to short-term and long-term water stress induced byPEG6000and root excision. The main results are as follows:
(1). The leaf hydraulic conductivity (Kleaf) of Line478varied diurnally andcorrelated with whole plant hydraulic conductivity. Similar diurnal rhythms of Kleafandthe root hydraulic conductivity (K_(root)) could be important to maintaining whole plantwater balance. Krootsignificantly correlated with leaf transpiration rate (E). After2h ofosmotic stress, the Krootof stressed plants significantly declined but K_(leaf) increased; thetranscription of four ZmPIPs was significantly up regulated in leaves, especially forZmPIP1;2, and ZmPIP2;5was down regulated in roots. The up-regulated Kleafanddown-regulated Krootmay break the water balance; and ZmPIPs genes may get involvedin the hydraulics changes during short-term water stress.
(2). The E and leaf water potential of478varied diurnally, but those of Tian4,which is more drought resistant, were more constant. Line478had advantages on leafhydraulic architecture, leaf and root morphology, K_(leaf)and K_(root), which may contribute to the higher E. The K_(leaf)and K_(root) of both Tian4and478varied diurnally. The K_(root)ofboth Tian4and478was reduced under osmotic stress, but the K_(root)of Tian4subsequently recovered. A lower and rapidly reduced leaf water loss and the recovery ofroot hydraulics during short-term osmotic stress may account for the ability ofdrought-resistant maize to maintain plant water balance.
(3). The transcription levels of ZmPIPs in mature leaves and roots of well wateredTian and478were significantly different. ZmPIP1;5was highly expressed in478but itsmRNA was not detected in Tian4. Within2h of water stress induced by PEG, thetranscription levels of ZmPIPs were up-regulated in roots but were down-regulated inleaves of Tian4, which may be help to increase water uptake and decrease water loss; inleaves and roots of478plants, the transcription levels of ZmPIPs both showed antemporary increase, which may contribute to the higher leaf water transpiration.
(4). The E, single root (Lp_(root)) and cell (Lp_(cell)) hydraulic conductivity of wheatincreased with increasing ploidy, but the V_(cell) was reduced. Osmotic stress significantlyreduced the E, Lp_(cell), Lp_(root), and the relative mRNA content of TaPIP1;2and TaPIP2;5in wheat. Under well-watered or osmotic stress conditions, Lp_(root)positively correlatedwith the E and Lp_(cell); the relative mRNA content of TaPIP1;2and TaPIP2;5significantly correlated with Lp_(cell)and Lp_(root), respectively. Lp_(cell)was reduced, but theLp_(cell)/Lp_(root)increased with increasing Vcell, suggesting that Vcellmay affect root radicalwater transport. Thus, the increased Lp_(cell)and transcription levels of TaPIP1;2andTaPIP2;5in wheat roots provides insight into the mechanisms underlying enhanced rootwater uptake during wheat evolution.
(5). The Pnof stressed plants totally recovered three days later while gswereconsistently lower than the controls, getting an improved instantaneous water-useefficiency (WUE). During prolonged water stress, the total water loss per plant ofstressed plants reduced significantly, while the dry mass of the whole plant did notchange, and leaf dry mass per unit area increased. The total leaf nitrogen, leaf nitrogenper unit area and leaf nitrogen concentration of stressed plants reduced significantly. Butthe photosynthetic nitrogen-use efficiency (PNUE) and dry mass based nitrogen-useefficiency (NUE) increased significantly. WUE positively correlated with PNUE. Bothimproved water-and nitrogen-use efficiencies of sweet sorghum under water stress maypartly explain its physiological acclimation to drought.
Based on the experimental data obtained from maize, wheat and sweet sorghum grown under controlled conditions, some important issues on the mechanism ofwhole-plant water balance were concerned in this dissertation. The results cancontribute to understand the biological bases of plant integrative drought-resistance, toclarify the physiological mechanisms of water-nitrogen-carbon relations, and to improvethe water use efficiency of plant by gene modification.
(1).恒定环境条件下玉米478的叶蒸腾速率、根导水率、叶导水率和整株导水率均表现明显日变化，且各参数间均存在显著正相关关系。这对于维持植株整体水分平衡是必要的。水分胁迫处理2小时后，玉米478根导水率显著降低而叶导水率却显著升高；叶片中ZmPIPs基因的转录量也显著增加，尤其是ZmPIP1;2；而根中仅ZmPIP2;5的转录水平显著下调，表明ZmPIPs可能参与了根叶导水特性的短期调节过程。
(2).干旱敏感玉米478的叶蒸腾速率呈明显的日变化规律，而抗旱性较强的天四较低且维持恒定；478在叶片水力结构、根叶形态和根导水率等方面均表现出适应高蒸腾耗水的特征；478和天四通过不同的叶内部水势差调节策略维持叶片水分平衡。短期水分胁迫下，478的根导水率下降而叶导水率增加，且叶蒸腾降低幅度却较小导致其体内水分亏缺；而天四的叶蒸腾速率大幅下降且根导水率在胁迫8小时后部分恢复。低叶蒸腾、水分胁迫后蒸腾迅速降低以及根导水率能部分恢复等特性可能是天四保持水分平衡的重要原因。
(3).正常供水天四和478的根叶中各ZmPIPs基因的转录水平存在明显差异，其中ZmPIP1;5在478中转录量较大而在天四中检测不到。PEG诱导水分胁迫处理2小时内，天四根中各ZmPIPs转录水平总体上调而叶片中总体下调，可能有助于其增加根系吸水同时减少叶片失水；478根中和叶片中各ZmPIPs的mRNA相对含量表现短暂的升高过程，这可能与其叶片蒸腾失水降低幅度较小有关。
(4).随着染色体倍性增加，小麦叶蒸腾失水和根系吸水能力均不断增大，而皮层细胞体积却逐渐减小。在正常供水或水分胁迫条件下，叶蒸腾速率、单根导水率、根细胞导水率以及TaPIPs转录水平之间均存在显著正相关关系。皮层细胞体积可能参与调节根径向水流各途径的相对贡献。在小麦进化过程中，根中TaPIP基因转录水平、细胞-细胞途径和质外体途径均显著增加，进而增强根系吸水能力以满足冠层耗水增加从而维持整株水分平衡。
(5).既然植物的基因表达、不同尺度导水特性和叶蒸腾之间存在一致性关系，那么叶片水分利用特性可能对整株耗水起控制作用。高抗旱性甜高粱对长期水分胁迫的适应过程中，叶片瞬时水分利用效率（WUE）显著增加，叶面积和单株耗水均显著降低，而干物质积累却变化不显著；叶氮积累显著降低；而光合氮利用效率（PNUE）和氮利用效率（NUE）显著增加。在干旱适应过程中，甜高粱叶片PNUE提高可能是WUE提高并减少植株耗水的原因之一。
本论文通过分析控制环境条件下所取得的实验数据，较深入探讨了植物水分平衡生理生态机制中的一些科学问题。研究结果有助于深化植物整体抗旱性的生物学基础的认识，为深入研究水-氮-碳三者间协调关系的生理机制提供基础，并为通过基因改良提高植物水分利用效率提供实验依据。
Water shortage often limits the ecological environment and agricultural productionin the arid and semi-arid area. Understanding the ecophysiological basis of highefficiency of plant water use is very important for developing water-saving agricultureand constructing ecological civilization. Plant can maintain water balance with variablehydraulic properties and aquaporins could be involved, which may help to understandthe ecophysiological mechanisms of plant water relations. This dissertation used thehydroponically grown maize (Zea mays L.), wheat (Triticum spp.) and sweet sorghum(Sorghum bicolor (L.) Moench) seedlings as experimental materials, and used cell-androot-pressure probes, high pressure flow meter and quantitive real-time PCR tomeasure the responses of cell, single root, root and shoot and whole-plant hydraulicsand aquaporin genes transcription to short-term and long-term water stress induced byPEG6000and root excision. The main results are as follows:
(1). The leaf hydraulic conductivity (Kleaf) of Line478varied diurnally andcorrelated with whole plant hydraulic conductivity. Similar diurnal rhythms of Kleafandthe root hydraulic conductivity (K_(root)) could be important to maintaining whole plantwater balance. Krootsignificantly correlated with leaf transpiration rate (E). After2h ofosmotic stress, the Krootof stressed plants significantly declined but K_(leaf) increased; thetranscription of four ZmPIPs was significantly up regulated in leaves, especially forZmPIP1;2, and ZmPIP2;5was down regulated in roots. The up-regulated Kleafanddown-regulated Krootmay break the water balance; and ZmPIPs genes may get involvedin the hydraulics changes during short-term water stress.
(2). The E and leaf water potential of478varied diurnally, but those of Tian4,which is more drought resistant, were more constant. Line478had advantages on leafhydraulic architecture, leaf and root morphology, K_(leaf)and K_(root), which may contribute to the higher E. The K_(leaf)and K_(root) of both Tian4and478varied diurnally. The K_(root)ofboth Tian4and478was reduced under osmotic stress, but the K_(root)of Tian4subsequently recovered. A lower and rapidly reduced leaf water loss and the recovery ofroot hydraulics during short-term osmotic stress may account for the ability ofdrought-resistant maize to maintain plant water balance.
(3). The transcription levels of ZmPIPs in mature leaves and roots of well wateredTian and478were significantly different. ZmPIP1;5was highly expressed in478but itsmRNA was not detected in Tian4. Within2h of water stress induced by PEG, thetranscription levels of ZmPIPs were up-regulated in roots but were down-regulated inleaves of Tian4, which may be help to increase water uptake and decrease water loss; inleaves and roots of478plants, the transcription levels of ZmPIPs both showed antemporary increase, which may contribute to the higher leaf water transpiration.
(4). The E, single root (Lp_(root)) and cell (Lp_(cell)) hydraulic conductivity of wheatincreased with increasing ploidy, but the V_(cell) was reduced. Osmotic stress significantlyreduced the E, Lp_(cell), Lp_(root), and the relative mRNA content of TaPIP1;2and TaPIP2;5in wheat. Under well-watered or osmotic stress conditions, Lp_(root)positively correlatedwith the E and Lp_(cell); the relative mRNA content of TaPIP1;2and TaPIP2;5significantly correlated with Lp_(cell)and Lp_(root), respectively. Lp_(cell)was reduced, but theLp_(cell)/Lp_(root)increased with increasing Vcell, suggesting that Vcellmay affect root radicalwater transport. Thus, the increased Lp_(cell)and transcription levels of TaPIP1;2andTaPIP2;5in wheat roots provides insight into the mechanisms underlying enhanced rootwater uptake during wheat evolution.
(5). The Pnof stressed plants totally recovered three days later while gswereconsistently lower than the controls, getting an improved instantaneous water-useefficiency (WUE). During prolonged water stress, the total water loss per plant ofstressed plants reduced significantly, while the dry mass of the whole plant did notchange, and leaf dry mass per unit area increased. The total leaf nitrogen, leaf nitrogenper unit area and leaf nitrogen concentration of stressed plants reduced significantly. Butthe photosynthetic nitrogen-use efficiency (PNUE) and dry mass based nitrogen-useefficiency (NUE) increased significantly. WUE positively correlated with PNUE. Bothimproved water-and nitrogen-use efficiencies of sweet sorghum under water stress maypartly explain its physiological acclimation to drought.
Based on the experimental data obtained from maize, wheat and sweet sorghum grown under controlled conditions, some important issues on the mechanism ofwhole-plant water balance were concerned in this dissertation. The results cancontribute to understand the biological bases of plant integrative drought-resistance, toclarify the physiological mechanisms of water-nitrogen-carbon relations, and to improvethe water use efficiency of plant by gene modification.
引文
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