水磷耦合对冬小麦水、磷利用与产量的影响及其生理基础
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摘要
供试品种为济麦20（JM20）和山农16（SN16），设置四个灌水水平：W0（不灌水）、W1（拔节水60mm）、W2（拔节水＋开花水、每次灌水60mm）、W3（拔节水+开花水+灌浆水、每次灌水60mm）；每个灌溉处理下设置不施磷（P0）、施90kg/hm2（P1）P2O5、施180kg/hm2（P2）P2O53个施磷量处理，研究水磷耦合对冬小麦水、磷利用和产量的影响及其生理基础。结果如下：1水磷耦合对小麦耗水特性和水分利用效率的影响
随着施磷量和灌水量的增加，旗叶水势和相对含水量均升高，提高灌浆后期旗叶的保水能力，延缓了旗叶衰老。两品种间比较，W0、W1处理，SN16旗叶抗失水能力显著高于JM20，利于籽粒产量的形成。
随着灌水量的增加，麦田总耗水量显著增加，土壤水的消耗量显著降低，土壤供水占总耗水量的比例显著下降，不利于小麦对降水和土壤水的利用。施磷显著提高了土壤贮水的利用率， W1、W2、W3处理的总耗水量和土壤供水量P2处理显著高于P1处理。品种间的比较，SN16总耗水量和土壤供水量及其占总耗水量的比例显著大于JM20，但灌水量和降水量占总耗水量的比例低于JM20。
增加灌水量会显著降低小麦的灌水利用效率，而增施磷肥有效提高灌水利用效率。过多灌水和施磷肥均降低了水分利用效率。两品种间比较，SN16能在干旱条件更好的利用土壤水和灌水，水分利用效率较高。在本试验条件下，灌溉量和施磷量在降水利用效率、灌水利用效率和水分利用效率的影响中存在极显著的互作效应。
小麦全生育期耗水量随生育期的进程呈现先低后高的变化趋势，P1水平下，拔节至成熟期的耗水量占到整个生育期的66.18%~79.39%。在P0条件下灌水显著增加了拔节后的阶段耗水量和日耗水量。在P1、P2条件下，在W2基础上继续增加灌水，拔节后的耗水模系数和日耗水量都略有增加但不显著，甚至略有下降，说明W2处理保持了较低的耗水量和耗水模系数，有利于水分的高效利用。
JM20和SN16两品种品种在W2处理籽粒产量和水分利用效率较高，其中SN16品种总耗水量为355~387mm，耗水量的水分来源分别为：降水占42.64%~45.80%，灌水占31.45%~33.78%，土壤供水占20.42%~25.93%； JM20品种总耗水量为337~394mm，耗水量的水分来源分别为：降水占41.33%~48.26%，灌水占30.48%~35.59%，土壤供水占16.10%~28.19%。2水磷耦合对小麦磷素利用的影响
灌水可以显著影响土壤速效磷的含量，拔节期和开花期灌水可以显著提高土壤有效磷的含量，在同时灌拔节水和开花水的条件下（W2）0～40cm土层土壤有效磷的含量随着施磷量的增加而显著增加，其他灌水处理则略有增加但不显著。
增施磷肥和灌水均增加冬小麦植株磷素积累量和植株磷积累百分率，同时提高冬小麦开花期前植株吸收磷素比例，同时提高了冬小麦干物质磷素生产效率、籽粒磷素生产效率。虽然随着施磷量和灌水量的增加，冬小麦磷素积累量增加，但在过多灌水和施肥的情况下（W3P2）小麦籽粒磷素生产效率有所下降，即在过多灌水和施磷时灌水与施磷对冬小麦籽粒产量的降低效应大于灌水与施磷对磷素积累量的增加效应。
籽粒中活性有机磷和磷脂含量较高，而冬小麦茎秆中活性有机磷含量最高，磷脂磷含量最低。施用磷肥和增加灌水都能提高冬小麦籽粒中各形态磷和全磷的积累，随着施磷量和灌水量的增加籽粒中活性无机磷、活性有机磷、全磷的含量增大。
成熟期磷素积累量、籽粒全磷和磷脂磷含量以及籽粒和干物质磷素生产效率的灌水量和施磷量互作效应均达到极显著水平。
在本试验条件下，灌溉量和施磷量在对小麦吸收磷素和磷素利用效率的影响中存在极显著互作效应。3水磷耦合对小麦产量形成的影响
增施磷肥能够提高小麦对氮素的吸收及在籽粒中的分配，但在灌水过多时磷肥的增加效应不明显。不同磷素水平对不同品种蛋白质含量的影响不一，且与对旗叶氮代谢关键酶活性的影响存在着对应关系。
在P1处理条件下，W2处理的籽粒产量高于W0、W1处理，在W2处理基础上再增加灌水量，籽粒产量无显著提高，且显著降低了水分利用效率；施磷显著提高了籽粒产量，但P2与P1处理相比，籽粒产量无显著提高，且显著降低了W2处理的水分利用效率。说明灌水和施磷显著提高籽粒的产量，但过量灌水会导致籽粒产量下降。适量灌水，增施磷肥可以显著提高籽粒产量。
在W0和W1处理条件下，增施磷肥有利于籽粒产量和水分利用效率的同步提高；而在W3处理条件下条件下，增施磷肥籽粒产量无显著提高，灌溉效益和磷素利用效率均降低。
在本试验条件下，施磷90kg/hm~2，拔节水和开花水分别灌60mm的W2处理，获得高的籽粒产量、水分和磷素利用效率，农田耗水量较低。
Effects of irrigation and phosphorus fertilizer application on water use efficiency andphosphorus use efficiency in wheat were studied in the field experiments which were carriedout on low phosphorus using the cultivar Jimai20(JM20) and Shannong16(SN16). Fourirrigation amounts were set with no water (W0), jointing water at60mm (W1),jointing+anthesis at60mm (W2), jointing+anthesis+filling water at60mm (W3) every time,and three phosphorus fertilizer (P2O5) rates were set with0(P0),90(P1) and180(P2)P2O5kg/ha, respectively.1. Effects coupling of irrigation and phosphorus on water consumption characteristicsand water use efficiency in winter wheat
Under the same phosphorus rate, with increasing irrigation amount, water consumptionamount significantly increased, but the ratio of precipitation amount and soil water amount towater consumption amount reduced significantly, and also improved the proportion of soilwater consumption for total water consumption. At the base of treatment W2, more irrigationrate decreased the ratio of precipitation and soil water consumption to water consumptionamount, but significantly increased the water consumption amount.
Compared with the P0treatment, additation of phosphorus fertilizer can improve the useof soil water, increase the proportion of soil water consumption for total water consumption.The results in this experiment also suggested that there was a significant interaction betweenirrigation rate and phosphorus rate for water use efficiency(WUEY) and irrigation water useefficiency(WUEI).
Under the treatment W0and W1, the grain yield and water use efficiency of SN16washigher than JM20. Under the treatment W2, water consumption amount of SN16was355~387mm, the percentage of irrigation rate to water consumption amount was31.45%~33.78%, the percentage of precipitation rate to water consumption amount was42.64%~45.80%, the percentage of soil water consumption rate to water consumption amountwas20.42%~25.93%. Under the treatment W2, water consumption amount of JM20was337~394mm, the percentage of irrigation rate to water consumption amount was30.48%~35.59%, the percentage of precipitation rate to water consumption amount was41.33%~48.26%, the percentage of soil water consumption rate to water consumption amountwas16.10%~28.19%.
Based on these results, it is concluded that treatment with phosphorus application at P2O590kg/ha or180kg/ha, irrigation at120mm could get high grain yield and water useefficiency. The treatment P1W2had the higher grain yield, water use efficiency(WUEY),while over-irrigation increased the water consumption amount, and decreased the WUEYandirrigation water use efficiency(WUEI).2. Effects coupling of irrigation and phosphorus on phosphorus uptake and phosphorususe efficiency in winter wheat
The amount of soil available P was significantly increased with increasing P fertilizerapplications when less irrigation levels (W0, W1and W2) were applied at different wheatgrowth stages except maturity, but under W3treatment there was only a slight increase and nosignificant difference between P1and P2treatment.
The addition of a P fertilizer and irrigation increased P accumulation amounts and thepercentage of P accumulation amounts at the different stage, as well as the proportion of Paccumulation amounts before the anthesis stage. The addition of a P fertilizer and irrigationalso increased the P production efficiency for grain and dry matter; moreover, there were verysignificant interactions between irrigation and P rate on P production efficiency for grain anddry matter. The content of P accumulation was increased accordingly by the addition of a Pfertilizer and irrigation, but overabundance of the P fertilizer and irrigation with180kg/haand180mm irrigation caused P efficiency of wheat grain to decrease.
The contents of lecithoid-P and labile organic-P are higher than other forms of P in grain.The application of a P fertilizer and the addition of irrigation can improve the levels ofvarious forms of P and total P accumulation in winter wheat grain. The levels of labileinorganic-P, labile organic-P, lecithoid-P and total-P increased according to the increase in theamount of P applied and the addition of irrigation. There was also a very significant positivecorrelation between soil available P content and P accumulation in wheat at various growthperiods. The treatment with an irrigation application at jointing and anthesis with90~180kg/ha P fertilizer can achieve the highest lecithoid-P level in grain and a high P useefficiency in grain. Correlation analysis showed that there was a significant positivecorrelation between the different forms of P and total P content in the grain (P<0.05). Therewas also a significant positive correlation between lecithoid-P and other forms of P (P<0.05),indicating that there is a dynamic balance between lecithoid-P and other P forms; interactionsand restrictions between these forms maintain the physiological requirements of crops.
The interaction effect analysis also showed that for the P accumulation, grain total P,grain phospholipid P and P production efficiency for dry matter, there were significantlyinteractions between irrigation and P rate (P<0.01).
Based on these results, the treatment with phosphorus application at P2O590kg/ha, andirrigation at120mm could get high total phosphorus and different forms phosphorus in winterwheat grain. And the treatment with phosphorus application at P2O5180kg/ha, and irrigation at60mm also could get high total phosphorus and different forms phosphorus in winter wheatgrain.3. Effects of irrigation and phosphorus fertilizer on grain yield
Under P1treatment, the total starch content and the components content of treatment W2was higher than treatment W0and W1. At the base of treatment W2, more irrigation ratedecreased the ratio of amylpectin to amylase significantly. Compared with treatment P0,phosphorus application increased the starch content, but had no significant effect on the ratioof amylpectin to amylose.There were no significantly difference between treatment P1and P2.
The synthesis and accumulation of protein in grains were increased by P applicationduring grain filling stage, especially in the early stage. The protein content in treatment P1and P2was significantly higher than that in P0during the early grain filling. When maturing,P1treatment had the highest total protein content in both cultivars. In P1treatment, there werelittle effect on the contents of albumin and globulin, but significantly increase the contents ofgliadin and gluten. These results suggested that P1treatment had little effect on wheatnutritional quality, but could improve its processing quality. Therefore, different cultivationmanagement should be adopted according to the different quality demands of different wheattypes.
At the base of treatment W2, more irrigation rate decreased the transformation and theamino acid from vegetative parts to grain during middle and late filling stage, and decreasedthe ratio of amylopectin to amylose significantly.
Under the treatment P1, the total starch content and its components content of treatmentW2was higher than that of treatment W0and W1.
The treatment W2had higher grain yield than W0and W1under the treatment P1. At thebase of treatment W2, more irrigation rate did not increase grain yield, but it decreased thewater use efficiency significantly. Phosphorus application improved the grain yieldsignificantly, but there was no significant difference between treatment P1and P2.
Based on these results, it is concluded that the treatment with phosphorus application atP2O590kg/ha, irrigation at180mm could get high grain yield, water use efficiency and P use efficency. And the treatment with phosphorus application at P2O5180kg/ha, and irrigation at60mm also could get high grain yield and P use efficency,its water use efficiency was higherthan treatment W2P2, but its water consumption amount was lower than treatment W2P2,which has practical implications for regions with water resource shortage.
随着施磷量和灌水量的增加，旗叶水势和相对含水量均升高，提高灌浆后期旗叶的保水能力，延缓了旗叶衰老。两品种间比较，W0、W1处理，SN16旗叶抗失水能力显著高于JM20，利于籽粒产量的形成。
随着灌水量的增加，麦田总耗水量显著增加，土壤水的消耗量显著降低，土壤供水占总耗水量的比例显著下降，不利于小麦对降水和土壤水的利用。施磷显著提高了土壤贮水的利用率， W1、W2、W3处理的总耗水量和土壤供水量P2处理显著高于P1处理。品种间的比较，SN16总耗水量和土壤供水量及其占总耗水量的比例显著大于JM20，但灌水量和降水量占总耗水量的比例低于JM20。
增加灌水量会显著降低小麦的灌水利用效率，而增施磷肥有效提高灌水利用效率。过多灌水和施磷肥均降低了水分利用效率。两品种间比较，SN16能在干旱条件更好的利用土壤水和灌水，水分利用效率较高。在本试验条件下，灌溉量和施磷量在降水利用效率、灌水利用效率和水分利用效率的影响中存在极显著的互作效应。
小麦全生育期耗水量随生育期的进程呈现先低后高的变化趋势，P1水平下，拔节至成熟期的耗水量占到整个生育期的66.18%~79.39%。在P0条件下灌水显著增加了拔节后的阶段耗水量和日耗水量。在P1、P2条件下，在W2基础上继续增加灌水，拔节后的耗水模系数和日耗水量都略有增加但不显著，甚至略有下降，说明W2处理保持了较低的耗水量和耗水模系数，有利于水分的高效利用。
JM20和SN16两品种品种在W2处理籽粒产量和水分利用效率较高，其中SN16品种总耗水量为355~387mm，耗水量的水分来源分别为：降水占42.64%~45.80%，灌水占31.45%~33.78%，土壤供水占20.42%~25.93%； JM20品种总耗水量为337~394mm，耗水量的水分来源分别为：降水占41.33%~48.26%，灌水占30.48%~35.59%，土壤供水占16.10%~28.19%。2水磷耦合对小麦磷素利用的影响
灌水可以显著影响土壤速效磷的含量，拔节期和开花期灌水可以显著提高土壤有效磷的含量，在同时灌拔节水和开花水的条件下（W2）0～40cm土层土壤有效磷的含量随着施磷量的增加而显著增加，其他灌水处理则略有增加但不显著。
增施磷肥和灌水均增加冬小麦植株磷素积累量和植株磷积累百分率，同时提高冬小麦开花期前植株吸收磷素比例，同时提高了冬小麦干物质磷素生产效率、籽粒磷素生产效率。虽然随着施磷量和灌水量的增加，冬小麦磷素积累量增加，但在过多灌水和施肥的情况下（W3P2）小麦籽粒磷素生产效率有所下降，即在过多灌水和施磷时灌水与施磷对冬小麦籽粒产量的降低效应大于灌水与施磷对磷素积累量的增加效应。
籽粒中活性有机磷和磷脂含量较高，而冬小麦茎秆中活性有机磷含量最高，磷脂磷含量最低。施用磷肥和增加灌水都能提高冬小麦籽粒中各形态磷和全磷的积累，随着施磷量和灌水量的增加籽粒中活性无机磷、活性有机磷、全磷的含量增大。
成熟期磷素积累量、籽粒全磷和磷脂磷含量以及籽粒和干物质磷素生产效率的灌水量和施磷量互作效应均达到极显著水平。
在本试验条件下，灌溉量和施磷量在对小麦吸收磷素和磷素利用效率的影响中存在极显著互作效应。3水磷耦合对小麦产量形成的影响
增施磷肥能够提高小麦对氮素的吸收及在籽粒中的分配，但在灌水过多时磷肥的增加效应不明显。不同磷素水平对不同品种蛋白质含量的影响不一，且与对旗叶氮代谢关键酶活性的影响存在着对应关系。
在P1处理条件下，W2处理的籽粒产量高于W0、W1处理，在W2处理基础上再增加灌水量，籽粒产量无显著提高，且显著降低了水分利用效率；施磷显著提高了籽粒产量，但P2与P1处理相比，籽粒产量无显著提高，且显著降低了W2处理的水分利用效率。说明灌水和施磷显著提高籽粒的产量，但过量灌水会导致籽粒产量下降。适量灌水，增施磷肥可以显著提高籽粒产量。
在W0和W1处理条件下，增施磷肥有利于籽粒产量和水分利用效率的同步提高；而在W3处理条件下条件下，增施磷肥籽粒产量无显著提高，灌溉效益和磷素利用效率均降低。
在本试验条件下，施磷90kg/hm~2，拔节水和开花水分别灌60mm的W2处理，获得高的籽粒产量、水分和磷素利用效率，农田耗水量较低。
Effects of irrigation and phosphorus fertilizer application on water use efficiency andphosphorus use efficiency in wheat were studied in the field experiments which were carriedout on low phosphorus using the cultivar Jimai20(JM20) and Shannong16(SN16). Fourirrigation amounts were set with no water (W0), jointing water at60mm (W1),jointing+anthesis at60mm (W2), jointing+anthesis+filling water at60mm (W3) every time,and three phosphorus fertilizer (P2O5) rates were set with0(P0),90(P1) and180(P2)P2O5kg/ha, respectively.1. Effects coupling of irrigation and phosphorus on water consumption characteristicsand water use efficiency in winter wheat
Under the same phosphorus rate, with increasing irrigation amount, water consumptionamount significantly increased, but the ratio of precipitation amount and soil water amount towater consumption amount reduced significantly, and also improved the proportion of soilwater consumption for total water consumption. At the base of treatment W2, more irrigationrate decreased the ratio of precipitation and soil water consumption to water consumptionamount, but significantly increased the water consumption amount.
Compared with the P0treatment, additation of phosphorus fertilizer can improve the useof soil water, increase the proportion of soil water consumption for total water consumption.The results in this experiment also suggested that there was a significant interaction betweenirrigation rate and phosphorus rate for water use efficiency(WUEY) and irrigation water useefficiency(WUEI).
Under the treatment W0and W1, the grain yield and water use efficiency of SN16washigher than JM20. Under the treatment W2, water consumption amount of SN16was355~387mm, the percentage of irrigation rate to water consumption amount was31.45%~33.78%, the percentage of precipitation rate to water consumption amount was42.64%~45.80%, the percentage of soil water consumption rate to water consumption amountwas20.42%~25.93%. Under the treatment W2, water consumption amount of JM20was337~394mm, the percentage of irrigation rate to water consumption amount was30.48%~35.59%, the percentage of precipitation rate to water consumption amount was41.33%~48.26%, the percentage of soil water consumption rate to water consumption amountwas16.10%~28.19%.
Based on these results, it is concluded that treatment with phosphorus application at P2O590kg/ha or180kg/ha, irrigation at120mm could get high grain yield and water useefficiency. The treatment P1W2had the higher grain yield, water use efficiency(WUEY),while over-irrigation increased the water consumption amount, and decreased the WUEYandirrigation water use efficiency(WUEI).2. Effects coupling of irrigation and phosphorus on phosphorus uptake and phosphorususe efficiency in winter wheat
The amount of soil available P was significantly increased with increasing P fertilizerapplications when less irrigation levels (W0, W1and W2) were applied at different wheatgrowth stages except maturity, but under W3treatment there was only a slight increase and nosignificant difference between P1and P2treatment.
The addition of a P fertilizer and irrigation increased P accumulation amounts and thepercentage of P accumulation amounts at the different stage, as well as the proportion of Paccumulation amounts before the anthesis stage. The addition of a P fertilizer and irrigationalso increased the P production efficiency for grain and dry matter; moreover, there were verysignificant interactions between irrigation and P rate on P production efficiency for grain anddry matter. The content of P accumulation was increased accordingly by the addition of a Pfertilizer and irrigation, but overabundance of the P fertilizer and irrigation with180kg/haand180mm irrigation caused P efficiency of wheat grain to decrease.
The contents of lecithoid-P and labile organic-P are higher than other forms of P in grain.The application of a P fertilizer and the addition of irrigation can improve the levels ofvarious forms of P and total P accumulation in winter wheat grain. The levels of labileinorganic-P, labile organic-P, lecithoid-P and total-P increased according to the increase in theamount of P applied and the addition of irrigation. There was also a very significant positivecorrelation between soil available P content and P accumulation in wheat at various growthperiods. The treatment with an irrigation application at jointing and anthesis with90~180kg/ha P fertilizer can achieve the highest lecithoid-P level in grain and a high P useefficiency in grain. Correlation analysis showed that there was a significant positivecorrelation between the different forms of P and total P content in the grain (P<0.05). Therewas also a significant positive correlation between lecithoid-P and other forms of P (P<0.05),indicating that there is a dynamic balance between lecithoid-P and other P forms; interactionsand restrictions between these forms maintain the physiological requirements of crops.
The interaction effect analysis also showed that for the P accumulation, grain total P,grain phospholipid P and P production efficiency for dry matter, there were significantlyinteractions between irrigation and P rate (P<0.01).
Based on these results, the treatment with phosphorus application at P2O590kg/ha, andirrigation at120mm could get high total phosphorus and different forms phosphorus in winterwheat grain. And the treatment with phosphorus application at P2O5180kg/ha, and irrigation at60mm also could get high total phosphorus and different forms phosphorus in winter wheatgrain.3. Effects of irrigation and phosphorus fertilizer on grain yield
Under P1treatment, the total starch content and the components content of treatment W2was higher than treatment W0and W1. At the base of treatment W2, more irrigation ratedecreased the ratio of amylpectin to amylase significantly. Compared with treatment P0,phosphorus application increased the starch content, but had no significant effect on the ratioof amylpectin to amylose.There were no significantly difference between treatment P1and P2.
The synthesis and accumulation of protein in grains were increased by P applicationduring grain filling stage, especially in the early stage. The protein content in treatment P1and P2was significantly higher than that in P0during the early grain filling. When maturing,P1treatment had the highest total protein content in both cultivars. In P1treatment, there werelittle effect on the contents of albumin and globulin, but significantly increase the contents ofgliadin and gluten. These results suggested that P1treatment had little effect on wheatnutritional quality, but could improve its processing quality. Therefore, different cultivationmanagement should be adopted according to the different quality demands of different wheattypes.
At the base of treatment W2, more irrigation rate decreased the transformation and theamino acid from vegetative parts to grain during middle and late filling stage, and decreasedthe ratio of amylopectin to amylose significantly.
Under the treatment P1, the total starch content and its components content of treatmentW2was higher than that of treatment W0and W1.
The treatment W2had higher grain yield than W0and W1under the treatment P1. At thebase of treatment W2, more irrigation rate did not increase grain yield, but it decreased thewater use efficiency significantly. Phosphorus application improved the grain yieldsignificantly, but there was no significant difference between treatment P1and P2.
Based on these results, it is concluded that the treatment with phosphorus application atP2O590kg/ha, irrigation at180mm could get high grain yield, water use efficiency and P use efficency. And the treatment with phosphorus application at P2O5180kg/ha, and irrigation at60mm also could get high grain yield and P use efficency,its water use efficiency was higherthan treatment W2P2, but its water consumption amount was lower than treatment W2P2,which has practical implications for regions with water resource shortage.
引文
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