水分胁迫及复水对作物生长及养分吸收补偿效应的研究
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摘要
干旱地区降雨量少且时空分布不均，作物很难有效利用土壤中有限的水分和养分。干旱胁迫已成为农业减产的主要原因。作物受旱后补充供水可在一定程度上产生补偿生长，使产量得到部分恢复。同时，施肥对作物抵抗干旱及促进复水后的恢复效应有一定的作用。本研究以冬小麦及夏玉米作试材，在盆栽条件下控制水分和养分的供应，以研究其对作物的生长发育及水分、养分吸收规律的影响，旨在查明作物的水分敏感期、复水最大效应期以及施肥对作物抵御干旱、恢复生长的作用，以求为干旱地区节水农业的发展提供有益的理论依据。所得主要结论如下：
（1）不同时期、不同程度水分胁迫对冬小麦生长发育及吸收能力造成一定的伤害。总的来看，分蘖期适应水分胁迫能力强于拔节期，因此受胁迫伤害较轻；重度胁迫受害较中度胁迫严重，但复水后小麦生长及生理反应的补偿效应也相对明显。分蘖期根冠比远大于拔节期，这是分蘖期适应干旱能力强的重要原因之一。复水后地上部生长速率显著大于根系，使根冠比降低。由于恢复供水后耗水量的增幅大于干物质生产的增幅，所以复水后水分利用效率（WUE）是显著降低的。
（2）施肥可增强小麦抗旱性，使分蘖期小麦干物质积累几乎不受水分胁迫的影响，高肥处理对拔节期受旱小麦复水后籽粒及总干物质生产的恢复有促进作用。另外，施肥还可降低拔节期冬小麦的气孔导度和蒸腾速率，减少水分无效散失。在中度胁迫下，施肥可以提高小麦的WUE。
（3）水分胁迫在不同程度上抑制了玉米的生长发育，重度胁迫影响普遍大于中度胁迫，苗前期受水分影响大于苗后期。土壤水分条件对养分吸收的影响顺序为磷>氮>钾。WUE并未简单地随水分胁迫的加重而提高，在苗前期以充分供水的对照为最高，苗后期中度胁迫WUE较高。从复水后总的恢复效果来看，苗前期中度胁迫及苗后期重度胁迫复水后恢复效应较明显，可以作为经济补水的重要时期。
（4）从苗期生长及养分吸收上看，施氮的效应尚不明显，但从生理反应上表现出了一定的作用。高氮处理对改善玉米苗前期植株水分状况有一定作用，但对苗后期效果不明显。施氮同样能降低玉米气孔导度及蒸腾速率，提高水分利用的经济性。
（5）从水分胁迫及复水后夏玉米生长发育及水分、养分吸收动态来看，胁迫5天后复水的处理短期内恢复较快，后期逐渐缓慢；胁迫10天后复水的处理恢复相对缓慢，但始终保持着持续增长的趋势，最终各种生长及吸收指标接近于胁迫5天的处理。无论何种水分条件，根冠比均随生育期的延长而降低，复水后降低趋势较明显，而充分供水条件下根冠比随生育期的延长降低较少。WUE随生育期延长而增高，前期增长快，后期减缓，呈“S”形曲线。相比之下，未复水的处理WUE一直高于充分供水的处理，复水后WUE的变化趋势与充分供水的处理较为相似。
（6）施氮对玉米苗期生长发育动态影响不太明显，低氮处理在复水初期恢复速度有略高于高氮处理的倾向，直到后期才逐渐被高氮处理所超过。
总之，不同时期、不同程度的水分胁迫对不同作物所造成的伤害不同，复水后所产生的补偿效应也不相同，同一作物复水后不同生长发育指标的恢复也有较大差别，复水可以使受旱作物在一定程度上解除胁迫，恢复生理功能，增加产量，但水分利用效率有所降低。施肥在一定程度上提高了作物的抗旱性。
Lack of rainfall and its uneven distribution are main reasons to affect the efficient use of water and nutrient in soil by crops in dry land areas. Water has become the main cause of reduction of yield in this area. It is found that if a crop suffered from the drought rewatered, there would be some compensatory growth, and the yield also would have compensatory. In the same time, fertilization can play an important role in crop drought resistance and the formation of compensatory effect after rewatering. In this research, winter wheat and summer corn were selected as experimental crops. Our purpose is to investigate the sensitive period of drought resistance of the crop and the most effective period of water recovery, and their relationship with the fertilization to provide useful information for the water saving technology in agriculture in dry land areas. The main results were as follows:
(1) The time and duration of water stress will affect the growth and nutrient uptake of winter wheat. In general, the adapting capacity of wheat to water stress in tillering stage was stronger than in jointing stage, so it suffered less injury. Serious water stress did more harm to wheat than middle water stress，and also showed a greater recovery in growth than middle water stress when water supply was recovered. At tillering stage, the ratio of root/shoot was higher than that at jointing stage, which was one of the most important reasons that wheat had a strong drought resistance capacity at tillering stage. After rewatering, the growth rate of overground part was faster the root, therefore，the ratio of root / shoot decreased；the increase of water consumption was greater than dry matter when rewatering, therefore, its water use efficiency (WUE) declines.
(2) At tillering stage, water stress had limited effects on wheat dry matter production when the soil fertility was high. During the jointing stage, the grain and total dry matter production was increased when water was supplied again. In addition, fertilization reduced the stomatal conductance and the rate of transpiration of winter wheat, so water dissipation was reduced. Under the condition of middle water stress, fertilization improved the water use efficiency of winter wheat.
(3) Water stress restrained the growth of summer corn，and serious water stress had a more significant influence to corn than the middle water stress. In prophase of seeding stage, water played a more important role than anaphase of seeding stage. Soil water
condition influenced the uptake of nutrient in soil, the order of influence degree was phosphorus>nitrogen>potassium. Water use efficiency was the highest under of the condition of full water supply. In anaphase of seeding stage, the WUE was in a relative high level. When water supply was recovered, the injury of middle stress in prophase of seeding stage and serious in anaphase of seeding stage was recovered significantly.
(4) High level nitrogen treatment could play some action in improving the water status of injured corn in prophase of seeding stage, but it had little effect in anaphase of seeding stage. Fertilization reduced the stomatal conductance and the rate of transpiration of summer corn，therefore，water was saved.
(5) In the angle of trends of corn growth under water stress and rewatering condition, treatments of 5 days water stress recovered growth in a short time, treatments of 10 days stress recovered slowly, but they kept the increasing for a long time；finally, they catched up with the treatments of 5 days stress. No matter what water condition, the ratio of root/shoot declined along with corn growth. After rewatered, it showed an obvious trend of decreasing. On the contrary, the treatments of full water supplied showed a little decrease. With corn growth, WUE was increased; it increases fast in prophase and slower in anaphase of seeding stage. After water recovery, WUE of corn showed the same trends with the treatments of full water supplied.
(6) It seems that nitrogen treatment has no significant effect o
（1）不同时期、不同程度水分胁迫对冬小麦生长发育及吸收能力造成一定的伤害。总的来看，分蘖期适应水分胁迫能力强于拔节期，因此受胁迫伤害较轻；重度胁迫受害较中度胁迫严重，但复水后小麦生长及生理反应的补偿效应也相对明显。分蘖期根冠比远大于拔节期，这是分蘖期适应干旱能力强的重要原因之一。复水后地上部生长速率显著大于根系，使根冠比降低。由于恢复供水后耗水量的增幅大于干物质生产的增幅，所以复水后水分利用效率（WUE）是显著降低的。
（2）施肥可增强小麦抗旱性，使分蘖期小麦干物质积累几乎不受水分胁迫的影响，高肥处理对拔节期受旱小麦复水后籽粒及总干物质生产的恢复有促进作用。另外，施肥还可降低拔节期冬小麦的气孔导度和蒸腾速率，减少水分无效散失。在中度胁迫下，施肥可以提高小麦的WUE。
（3）水分胁迫在不同程度上抑制了玉米的生长发育，重度胁迫影响普遍大于中度胁迫，苗前期受水分影响大于苗后期。土壤水分条件对养分吸收的影响顺序为磷>氮>钾。WUE并未简单地随水分胁迫的加重而提高，在苗前期以充分供水的对照为最高，苗后期中度胁迫WUE较高。从复水后总的恢复效果来看，苗前期中度胁迫及苗后期重度胁迫复水后恢复效应较明显，可以作为经济补水的重要时期。
（4）从苗期生长及养分吸收上看，施氮的效应尚不明显，但从生理反应上表现出了一定的作用。高氮处理对改善玉米苗前期植株水分状况有一定作用，但对苗后期效果不明显。施氮同样能降低玉米气孔导度及蒸腾速率，提高水分利用的经济性。
（6）施氮对玉米苗期生长发育动态影响不太明显，低氮处理在复水初期恢复速度有略高于高氮处理的倾向，直到后期才逐渐被高氮处理所超过。
总之，不同时期、不同程度的水分胁迫对不同作物所造成的伤害不同，复水后所产生的补偿效应也不相同，同一作物复水后不同生长发育指标的恢复也有较大差别，复水可以使受旱作物在一定程度上解除胁迫，恢复生理功能，增加产量，但水分利用效率有所降低。施肥在一定程度上提高了作物的抗旱性。
Lack of rainfall and its uneven distribution are main reasons to affect the efficient use of water and nutrient in soil by crops in dry land areas. Water has become the main cause of reduction of yield in this area. It is found that if a crop suffered from the drought rewatered, there would be some compensatory growth, and the yield also would have compensatory. In the same time, fertilization can play an important role in crop drought resistance and the formation of compensatory effect after rewatering. In this research, winter wheat and summer corn were selected as experimental crops. Our purpose is to investigate the sensitive period of drought resistance of the crop and the most effective period of water recovery, and their relationship with the fertilization to provide useful information for the water saving technology in agriculture in dry land areas. The main results were as follows:
(1) The time and duration of water stress will affect the growth and nutrient uptake of winter wheat. In general, the adapting capacity of wheat to water stress in tillering stage was stronger than in jointing stage, so it suffered less injury. Serious water stress did more harm to wheat than middle water stress，and also showed a greater recovery in growth than middle water stress when water supply was recovered. At tillering stage, the ratio of root/shoot was higher than that at jointing stage, which was one of the most important reasons that wheat had a strong drought resistance capacity at tillering stage. After rewatering, the growth rate of overground part was faster the root, therefore，the ratio of root / shoot decreased；the increase of water consumption was greater than dry matter when rewatering, therefore, its water use efficiency (WUE) declines.
(2) At tillering stage, water stress had limited effects on wheat dry matter production when the soil fertility was high. During the jointing stage, the grain and total dry matter production was increased when water was supplied again. In addition, fertilization reduced the stomatal conductance and the rate of transpiration of winter wheat, so water dissipation was reduced. Under the condition of middle water stress, fertilization improved the water use efficiency of winter wheat.
(3) Water stress restrained the growth of summer corn，and serious water stress had a more significant influence to corn than the middle water stress. In prophase of seeding stage, water played a more important role than anaphase of seeding stage. Soil water
(4) High level nitrogen treatment could play some action in improving the water status of injured corn in prophase of seeding stage, but it had little effect in anaphase of seeding stage. Fertilization reduced the stomatal conductance and the rate of transpiration of summer corn，therefore，water was saved.
(5) In the angle of trends of corn growth under water stress and rewatering condition, treatments of 5 days water stress recovered growth in a short time, treatments of 10 days stress recovered slowly, but they kept the increasing for a long time；finally, they catched up with the treatments of 5 days stress. No matter what water condition, the ratio of root/shoot declined along with corn growth. After rewatered, it showed an obvious trend of decreasing. On the contrary, the treatments of full water supplied showed a little decrease. With corn growth, WUE was increased; it increases fast in prophase and slower in anaphase of seeding stage. After water recovery, WUE of corn showed the same trends with the treatments of full water supplied.
(6) It seems that nitrogen treatment has no significant effect o
引文
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