广适性玉米杂交种抗逆稳产生理机制研究
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摘要
广适性玉米杂交种的主要特点是产量水平高且稳产性好、年际间变化小,适宜种植区域广、年种植面积大、甚至能够跨生态区种植。它们是推动玉米品种更新换代的主力,是提高玉米单产水平的最有效途径。品种的适应性比杂种优势更重要,玉米品种的产量高低取决于对当地生态条件的适应能力。干旱、肥力不足、花期阴雨是是黄淮海地区影响玉米产量的重要非生物逆境因子。本研究在人工控制条件下,选用适应性不同的玉米品种,研究了干旱、低肥、花期遮阴等3个胁迫因子单一及两、三重复合胁迫(共7种逆境)对玉米生长发育的影响,以探索广适性品种抗逆稳产的生理机制。主要研究结果如下：
1.多胁迫因子间存在明显的交互作用,不是单因子效应的累加。低肥、干旱和花期遮阴等因子对产量影响存在交互作用。在两因素的交互作用中,肥力、干旱、遮阴等三者间的交互作用均达到显著或极显著水平。
2.花期遮阴对玉米生长发育及产量的影响大于低肥和干旱。花期遮阴、低肥和干旱单胁迫造成玉米产量的损失分别为84.48%、51.23%和18.26%。本研究中,7种逆境胁迫降低了13.62%～91.55%的玉米产量。玉米产量在7种逆境胁迫下依次表现为干旱>低肥>低肥干旱>遮阴>干旱遮阴>低肥遮阴>低肥干旱遮阴。复合胁迫对产量的影响大于单一逆境胁迫。
3.广适性玉米杂交品种对多种逆境具有较高的抗性,没有明显的缺陷。对照处理ZD958(郑单958)的单株产量最高,且在7种胁迫条件下的平均单株产量最高,变异系数最小,稳产性最好；ND108(农大108)的变异系数居第二位,YY22(豫玉22)和ZY9(张玉9号)的变异系数最大。品种的稳产性与它们的抗逆性密切相关。ZD958和ND108对低肥、干旱和遮阴胁迫均具有较好的抗性,因此,产量在不同胁迫间波动小。YY22对遮阴胁迫敏感,ZY9对遮阴和低肥胁迫都较敏感,故产量在不同胁迫间波动大。因此,广适性玉米杂交种对逆境胁迫没有明显的敏感性,表现出广泛的适应性。
4.广适性玉米杂交种器官发育受胁迫影响小。雌雄穗协调发育是完成受精、籽粒建成的前提。逆境胁迫对不同适应性品种散粉时间的影响很小,对花丝抽出时间的影响大。广适性玉米杂交种品种ZD958受逆境胁迫花丝抽出时间推迟0.75-7.95d,雌雄穗开花间隔(ASI)增加了1.15～6.60d,在7种逆境胁迫间变异系数最小,分别为28.13%和22.89%。穗发育也表现相同的特点,吐丝后2周时7种胁迫条件下ZD958果穗占穗部总鲜重(果穗+苞叶)的平均比例为34.66%,居第三位,胁迫间的变异系数仅高于ND108。第五次取样时,ZD958果穗占穗部总鲜重的平均比例最高,为42.87%,变异系数仍然较小,居第二位。
5.广适性玉米杂交种的叶绿素含量高且胁迫处理间波动小。逆境胁迫下,5个玉米品种的叶绿素含量降低了11.82%-48.99%,叶绿素a含量变化小于叶绿素b。逆境胁迫解除后,叶绿素含量略有回升。叶绿素含量的降低主要在于叶绿素b含量下降。这也导致逆境胁迫处理后叶绿素a/b值显著升高。低肥、干旱、遮阴等3种逆境因子及其交互作用对玉米叶片中叶绿素含量的影响基本一致。ZD958的叶绿素含量高且胁迫处理间波动小。
6.广适性玉米杂交种物质积累快、籽粒形成关键期的植株生长率高。干物质积累和植株生长速率是植株个体生理功能的最终体现。对照处理ZD958的单株干物质积累量最大,收获时较其它品种高10.4%-17.3%,籽粒形成关键期的植株生长率最高,为3.32g/(株·d),高于其它品种4.75%-15.15%,且穗物质分配指数高。逆境胁迫期间(吐丝期)和逆境胁迫处理后(成熟期)广适性玉米杂交种ZD958的单株干物质在7种逆境条件下的变异系数均为最小,分别为3.94%和8.34%。胁迫处理后各品种穗物质分配指数减小了7.53%～79.0%,ZD958的穗物质分配指数在7种胁迫处理间波动最小。
7.广适性玉米杂交种物质转运调控能力强。逆境胁迫处理后,茎、叶贮存物质都向穗部转运。茎中物质转运晚于叶片,且叶片在8种环境条件下均存在物质转运,而茎中的物质只有在对照、干旱、低肥和低肥干旱4种环境条件下存在物质转运。广适性玉米杂交种ZD958叶片对照处理仅有6.52%的物质转运,而其它4个品种叶片物质的转运率达18.73%～30.58%。逆境条件下,ZD958最多可调控叶片32.27%的物质的转运,调控能力大高于其它4个品种。
8.广适性玉米杂交种的碳氮代谢协调。茎、叶、穗中C/N的变化可以反映胁迫对碳氮代谢的影响。茎C/N在低肥遮阴、低肥干旱遮阴、干旱胁迫吐丝期以及遮阴和干旱遮阴胁迫吐丝期和吐丝后3周时降低,说明胁迫对这些时期茎碳代谢的影响大于氮代谢,其它时期胁迫对氮代谢的影响大于碳代谢。叶C/N在干旱遮阴吐丝期和吐丝后3周时降低,胁迫氮代谢的影响大于氮代谢；其它时期胁迫处理的C/N均升高,说明胁迫主要影响了叶片中的氮代谢。穗C/N在吐丝后3周和收获期胁迫处理下增加,氮代谢受胁迫处理的影响大于碳代谢。广适性玉米杂交种ZD958在各环境条件下叶、茎中可溶性糖和全氮含量较高,C/N居中,碳氮代谢协调,逆境胁迫后恢复速度快于其它品种。
ABSTRACT The main characteristics of maize hybrids with wide adaptability is of high and stable yield, small changes between years, extensive areas suitable for planting, planting large area yearly, even to be able to grow across ecological zones. They were often the main force to promote the replacement of maize varieties, were the most effective way to improve the level of maize yield. The adaptability of varieties is more important than heterosis. The level of maize production depends on its ability to adapt to local ecological conditions. Drought, inadequate fertility, low light were the main abiotic stress factors for yield and stability in maize producing areas of the Huanghuaihai maize belt and the maize producing area. In this study, under controlled conditions, the different maize varieties with adaptability were used to study the lack of fertility, the effect of flowering drought, low light stress factors and three double, triple Stress (7stresses, totally) on maize growth to explore physiological mechanism of the wide varieties with wide adaptability response to these stress factors. The main results are as follows:
Low light on corn growth and yield was large than that of fertility and flowering drought. In this study,7Stresses decreased yield by13.62%～91.55%. The effect of7kinds of stress on the five yield of maize hybrids in order were shade> low fertility and shade>low fertility, drought and shade> drought and shade>low fertility>low fertility and drought> drought. Shading at flower period significantly reduced the yield of5varieties by84.48%than non-shading treatments. Low fertility and drought treatment caused51.23%and18.26%yield loss, respectively. Shading had a great influence on flowering time. The silking time of5varieties was delayed by more than7d in treatments of shade, low fertility and shade, drought and shade, low fertility, drought and shade.
There were more significant interactions between the factors, not just the cumulative effect of single stress factors. The interactions of low-fat, flowering shade and drought on the yield of maize yield existed. Among the interactions of two factors, the interactions of fertility, drought, and shade were significant or very significant. The interaction had very obvious effect on characteristics of species. The resistance of Zhangyu-9to low fertility was significantly higher than other varieties. Shading significantly increased the degree of sensitivity of maize to fertility, Zhangyu-9was the most sensitive and its barren-resistant index changed from the highest to the lowest.
The corn hybrids variety with wide adaptability had high resistance to stresses, and no obvious defects. The yield of Nongda108was the highest in the treatment of drought and shade, was the lowest in the treatment of drought, drought and low fertility. Zhang Yu-9yielded most in the treatment of low fertility, and yielded lowest in the treatment of shade and low fertility. The yield per plant Yuyu22in low-fertility and drought conditions was higher than Zhengdan958, and it was the second highest in high fertilizer and drought conditions. However, the yield per plant of Yuyu22was the lowest in the conditions of shade, drought and shade, low fertility conditions. Luoyu-4yielded lowest in the condition of low fertility, drought, shade. Zhengdan958yielded highestly in the conditions of shade, drought, and low fertility and drought-shade. And it yielded the second highestly in the other four stresses. Therefore, the wide adaptability of maize hybrids did not significantly sensitive to stress, showing a wide range of adaptability to stresses.
The difference of dry matter of5corn hybrids at silking was small. From silking to harvest the dry matter accumulation of maize hybrids with wide adaptability Zhengdan958was faster10.4%to17.3%than other varieties. At silking and harvest, the dry matter per plant of maize hybrids with wide adaptability Zhengdan958changed smallest among7kind of stresses, and the variation coefficient were3.94%and8.34%respectively. Under normal conditions, plant growth rate of maize hybrids with wide adaptability Zhengdan958during seed formation critical period was the highest, being4.03g/(plant· d), higher27.57%～57.37%than other varieties.
The maize hybrids with wide adaptability Zhengdan958was stronger ability for controlling matter distribution. In this study, stem was sink organs to accumulate storage material at pre-or pro-silking, and2weeks after silking, storage material in stem began to transport and redistribute in the treatment of drought, low fertility, drought and low fertility. During stresses the redistribution of the storage material in leaves was of great important for maize response to stresses. The maize hybrids with wide adaptability Zhengdan958transported only6.52%leaf material under normal conditions, while the transfer rate of the other four varieties changed from18.73%to30.58%. But under stress conditions, Zhengdan958could transported32.27%mostly of the material in leaves, and its regulatory capacity was much higher than the other four varieties.
The organ development of maize hybrids with wide adaptability was influenced small under by stresses. The effect of stresses on dusting time of varieties with different adaptability was very small, less than2d; but its effect on silking time was very large. The silking time of wide adaptability Zhengdan958was put off0.75～7.95d by stresses, and its ASI increased1.15～6.60d. Their variation coefficients among7kinds of stresses were minimum in5varieties, being28.13%and22.89%respectively. Ear growth also showed the same characteristics. The ratio of ear to total ear (ear+bract) of Zhengdan958was34.66%under7kinds of stresses at2weeks after silking (third sample), ranking third among5varieties. The variation coefficient of Zhengdan958between stresses was only higher than Nongda108. The ratio of Zhengdan958was the highest at fifth sampling, being42.87%. its variation coefficient was also small and being in second place.
Characteristics of carbon and nitrogen metabolism of maize hybrids with wide adaptability. After silking, the soluble sugar content, C/N in leaves, stem and ear, and total nitrogen content in leaves of5maize varieties changed as single-peak curve, and was the highest at3weeks after silking (5th sample). The performance of organs in order was stem> ear> leaf. The total nitrogen content in stem and ear was the lowest at3weeks after silking, and it was higher at harvest than at silking. The content of soluble sugar and total nitrogen in leaves and stem of the maize hybrids Zhengdan958with wide adaptability under normal conditions was higher. After stresses relieved, it could restored quickly than other varieties.
1.多胁迫因子间存在明显的交互作用,不是单因子效应的累加。低肥、干旱和花期遮阴等因子对产量影响存在交互作用。在两因素的交互作用中,肥力、干旱、遮阴等三者间的交互作用均达到显著或极显著水平。
2.花期遮阴对玉米生长发育及产量的影响大于低肥和干旱。花期遮阴、低肥和干旱单胁迫造成玉米产量的损失分别为84.48%、51.23%和18.26%。本研究中,7种逆境胁迫降低了13.62%～91.55%的玉米产量。玉米产量在7种逆境胁迫下依次表现为干旱>低肥>低肥干旱>遮阴>干旱遮阴>低肥遮阴>低肥干旱遮阴。复合胁迫对产量的影响大于单一逆境胁迫。
3.广适性玉米杂交品种对多种逆境具有较高的抗性,没有明显的缺陷。对照处理ZD958(郑单958)的单株产量最高,且在7种胁迫条件下的平均单株产量最高,变异系数最小,稳产性最好；ND108(农大108)的变异系数居第二位,YY22(豫玉22)和ZY9(张玉9号)的变异系数最大。品种的稳产性与它们的抗逆性密切相关。ZD958和ND108对低肥、干旱和遮阴胁迫均具有较好的抗性,因此,产量在不同胁迫间波动小。YY22对遮阴胁迫敏感,ZY9对遮阴和低肥胁迫都较敏感,故产量在不同胁迫间波动大。因此,广适性玉米杂交种对逆境胁迫没有明显的敏感性,表现出广泛的适应性。
4.广适性玉米杂交种器官发育受胁迫影响小。雌雄穗协调发育是完成受精、籽粒建成的前提。逆境胁迫对不同适应性品种散粉时间的影响很小,对花丝抽出时间的影响大。广适性玉米杂交种品种ZD958受逆境胁迫花丝抽出时间推迟0.75-7.95d,雌雄穗开花间隔(ASI)增加了1.15～6.60d,在7种逆境胁迫间变异系数最小,分别为28.13%和22.89%。穗发育也表现相同的特点,吐丝后2周时7种胁迫条件下ZD958果穗占穗部总鲜重(果穗+苞叶)的平均比例为34.66%,居第三位,胁迫间的变异系数仅高于ND108。第五次取样时,ZD958果穗占穗部总鲜重的平均比例最高,为42.87%,变异系数仍然较小,居第二位。
5.广适性玉米杂交种的叶绿素含量高且胁迫处理间波动小。逆境胁迫下,5个玉米品种的叶绿素含量降低了11.82%-48.99%,叶绿素a含量变化小于叶绿素b。逆境胁迫解除后,叶绿素含量略有回升。叶绿素含量的降低主要在于叶绿素b含量下降。这也导致逆境胁迫处理后叶绿素a/b值显著升高。低肥、干旱、遮阴等3种逆境因子及其交互作用对玉米叶片中叶绿素含量的影响基本一致。ZD958的叶绿素含量高且胁迫处理间波动小。
6.广适性玉米杂交种物质积累快、籽粒形成关键期的植株生长率高。干物质积累和植株生长速率是植株个体生理功能的最终体现。对照处理ZD958的单株干物质积累量最大,收获时较其它品种高10.4%-17.3%,籽粒形成关键期的植株生长率最高,为3.32g/(株·d),高于其它品种4.75%-15.15%,且穗物质分配指数高。逆境胁迫期间(吐丝期)和逆境胁迫处理后(成熟期)广适性玉米杂交种ZD958的单株干物质在7种逆境条件下的变异系数均为最小,分别为3.94%和8.34%。胁迫处理后各品种穗物质分配指数减小了7.53%～79.0%,ZD958的穗物质分配指数在7种胁迫处理间波动最小。
7.广适性玉米杂交种物质转运调控能力强。逆境胁迫处理后,茎、叶贮存物质都向穗部转运。茎中物质转运晚于叶片,且叶片在8种环境条件下均存在物质转运,而茎中的物质只有在对照、干旱、低肥和低肥干旱4种环境条件下存在物质转运。广适性玉米杂交种ZD958叶片对照处理仅有6.52%的物质转运,而其它4个品种叶片物质的转运率达18.73%～30.58%。逆境条件下,ZD958最多可调控叶片32.27%的物质的转运,调控能力大高于其它4个品种。
8.广适性玉米杂交种的碳氮代谢协调。茎、叶、穗中C/N的变化可以反映胁迫对碳氮代谢的影响。茎C/N在低肥遮阴、低肥干旱遮阴、干旱胁迫吐丝期以及遮阴和干旱遮阴胁迫吐丝期和吐丝后3周时降低,说明胁迫对这些时期茎碳代谢的影响大于氮代谢,其它时期胁迫对氮代谢的影响大于碳代谢。叶C/N在干旱遮阴吐丝期和吐丝后3周时降低,胁迫氮代谢的影响大于氮代谢；其它时期胁迫处理的C/N均升高,说明胁迫主要影响了叶片中的氮代谢。穗C/N在吐丝后3周和收获期胁迫处理下增加,氮代谢受胁迫处理的影响大于碳代谢。广适性玉米杂交种ZD958在各环境条件下叶、茎中可溶性糖和全氮含量较高,C/N居中,碳氮代谢协调,逆境胁迫后恢复速度快于其它品种。
ABSTRACT The main characteristics of maize hybrids with wide adaptability is of high and stable yield, small changes between years, extensive areas suitable for planting, planting large area yearly, even to be able to grow across ecological zones. They were often the main force to promote the replacement of maize varieties, were the most effective way to improve the level of maize yield. The adaptability of varieties is more important than heterosis. The level of maize production depends on its ability to adapt to local ecological conditions. Drought, inadequate fertility, low light were the main abiotic stress factors for yield and stability in maize producing areas of the Huanghuaihai maize belt and the maize producing area. In this study, under controlled conditions, the different maize varieties with adaptability were used to study the lack of fertility, the effect of flowering drought, low light stress factors and three double, triple Stress (7stresses, totally) on maize growth to explore physiological mechanism of the wide varieties with wide adaptability response to these stress factors. The main results are as follows:
Low light on corn growth and yield was large than that of fertility and flowering drought. In this study,7Stresses decreased yield by13.62%～91.55%. The effect of7kinds of stress on the five yield of maize hybrids in order were shade> low fertility and shade>low fertility, drought and shade> drought and shade>low fertility>low fertility and drought> drought. Shading at flower period significantly reduced the yield of5varieties by84.48%than non-shading treatments. Low fertility and drought treatment caused51.23%and18.26%yield loss, respectively. Shading had a great influence on flowering time. The silking time of5varieties was delayed by more than7d in treatments of shade, low fertility and shade, drought and shade, low fertility, drought and shade.
There were more significant interactions between the factors, not just the cumulative effect of single stress factors. The interactions of low-fat, flowering shade and drought on the yield of maize yield existed. Among the interactions of two factors, the interactions of fertility, drought, and shade were significant or very significant. The interaction had very obvious effect on characteristics of species. The resistance of Zhangyu-9to low fertility was significantly higher than other varieties. Shading significantly increased the degree of sensitivity of maize to fertility, Zhangyu-9was the most sensitive and its barren-resistant index changed from the highest to the lowest.
The corn hybrids variety with wide adaptability had high resistance to stresses, and no obvious defects. The yield of Nongda108was the highest in the treatment of drought and shade, was the lowest in the treatment of drought, drought and low fertility. Zhang Yu-9yielded most in the treatment of low fertility, and yielded lowest in the treatment of shade and low fertility. The yield per plant Yuyu22in low-fertility and drought conditions was higher than Zhengdan958, and it was the second highest in high fertilizer and drought conditions. However, the yield per plant of Yuyu22was the lowest in the conditions of shade, drought and shade, low fertility conditions. Luoyu-4yielded lowest in the condition of low fertility, drought, shade. Zhengdan958yielded highestly in the conditions of shade, drought, and low fertility and drought-shade. And it yielded the second highestly in the other four stresses. Therefore, the wide adaptability of maize hybrids did not significantly sensitive to stress, showing a wide range of adaptability to stresses.
The difference of dry matter of5corn hybrids at silking was small. From silking to harvest the dry matter accumulation of maize hybrids with wide adaptability Zhengdan958was faster10.4%to17.3%than other varieties. At silking and harvest, the dry matter per plant of maize hybrids with wide adaptability Zhengdan958changed smallest among7kind of stresses, and the variation coefficient were3.94%and8.34%respectively. Under normal conditions, plant growth rate of maize hybrids with wide adaptability Zhengdan958during seed formation critical period was the highest, being4.03g/(plant· d), higher27.57%～57.37%than other varieties.
The maize hybrids with wide adaptability Zhengdan958was stronger ability for controlling matter distribution. In this study, stem was sink organs to accumulate storage material at pre-or pro-silking, and2weeks after silking, storage material in stem began to transport and redistribute in the treatment of drought, low fertility, drought and low fertility. During stresses the redistribution of the storage material in leaves was of great important for maize response to stresses. The maize hybrids with wide adaptability Zhengdan958transported only6.52%leaf material under normal conditions, while the transfer rate of the other four varieties changed from18.73%to30.58%. But under stress conditions, Zhengdan958could transported32.27%mostly of the material in leaves, and its regulatory capacity was much higher than the other four varieties.
The organ development of maize hybrids with wide adaptability was influenced small under by stresses. The effect of stresses on dusting time of varieties with different adaptability was very small, less than2d; but its effect on silking time was very large. The silking time of wide adaptability Zhengdan958was put off0.75～7.95d by stresses, and its ASI increased1.15～6.60d. Their variation coefficients among7kinds of stresses were minimum in5varieties, being28.13%and22.89%respectively. Ear growth also showed the same characteristics. The ratio of ear to total ear (ear+bract) of Zhengdan958was34.66%under7kinds of stresses at2weeks after silking (third sample), ranking third among5varieties. The variation coefficient of Zhengdan958between stresses was only higher than Nongda108. The ratio of Zhengdan958was the highest at fifth sampling, being42.87%. its variation coefficient was also small and being in second place.
Characteristics of carbon and nitrogen metabolism of maize hybrids with wide adaptability. After silking, the soluble sugar content, C/N in leaves, stem and ear, and total nitrogen content in leaves of5maize varieties changed as single-peak curve, and was the highest at3weeks after silking (5th sample). The performance of organs in order was stem> ear> leaf. The total nitrogen content in stem and ear was the lowest at3weeks after silking, and it was higher at harvest than at silking. The content of soluble sugar and total nitrogen in leaves and stem of the maize hybrids Zhengdan958with wide adaptability under normal conditions was higher. After stresses relieved, it could restored quickly than other varieties.
引文
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