群体密度对玉米茎秆及根系抗倒伏特性的影响
摘要
高密度栽培已经成为目前玉米高产潜力挖掘的关键措施之一,但是群体密度的增加增加了玉米倒伏的风险。吉林省年平均降水量654.3毫米,且降雨多集中在7-8月,正值玉米生长抽雄-灌浆的关键时期,在高温、多雨多风的气候影响下,倒伏已成为影响耐密性玉米实现高产的重要原因之一。关于玉米倒伏的研究已有一些研究,但关于群体密度对玉米地上部茎秆和地下部根系倒伏特性的系统研究并不多见。本研究主要探讨不同密度对玉米茎秆及根系形态、干物质及相关生理特性的变化,探明其和倒伏率的相关关.系,以揭示玉米的倒伏机理,为吉林省玉米抗倒伏栽培及育种提供量化参考依据。
本研究选择7个种植密度,2009-2011年选择4个抗倒伏性能不同的品种,采用随机区组设计,通过茎秆和根系形态、干物质及相关生理指标的测定,分析其和茎秆强度及倒伏率的相关性。2012年,以吉林省11个主导品种为试验材料,从形态学和生理学角度,研究“布拉万”台风对不同玉米品种的影响,探明不同玉米品种的倒伏特征。研究结果表明：
1、茎秆农艺性状的结果为：穗位高和穗位高系数随着密度的增加逐渐增加,株高在7万株／公顷的密度下达到最大；郑单958的穗位高和穗位高系数对密度的变化不敏感。先玉420和先玉335倒伏率与穗位高和穗位高系数呈现极显著正相关,益丰29和郑单958则与株高、穗位高和穗位高系数均无显著相关性。同一品种相同节间粗随密度增加逐渐降低、节间长度无规律性变化。节间干物质百分比、单位茎长干重、节间强度均随密度增加先升高后降低,在7和8万株／公顷出现最大值,各品种表现不同。两个玉米品种其倒伏率与第3、4节茎长、茎粗、单位茎长干重、茎秆强度均显著相关。品种间比较,益丰29和郑单958第2-6节间干物质百分比、单位茎长干重、节间强度、穗位高及穗位高系数均高于先玉335和先玉420,株高则相反。
2、茎秆木质素及关键合成酶活性结果为：不同密度处理下,平均节间纤维素含量、木质素含量、PAL活性、TAL活性和CAD活性均为益丰29高于先玉420。益丰29节间PAL、TAL活性节间表现为第3节间>第4节间>第5节间。CAD活性各节间变化因品种而不同,先玉420第4节>第3节>第5节,益丰29第3节>第4节>第5节。先玉420节间木质素含量与CAD活性显著正相关,益丰29则与PAL活性和TAL活性呈显著正相关。在抽雄期和灌浆初期,益丰29节间强度与木质素含量、PAL活性、TAL活性显著正相关,先玉420与TAL呈显著正相关。
3、茎秆化学成分及矿质元素结果为：随着密度的增加,节间N、K、Si、Ca、Mg、Zn含量均先升高后降低,最高点均出现在7万株／公顷和8万株／公顷,因品种和生育时期而不同。品种间益丰29高于先玉420。玉米茎秆强度与K、Si和淀粉含量呈现显著正相关,与N、Ca、Mg含量呈现正相关,但不显著,与Zn和可溶性糖含量相关性不大。益丰29和先玉420第3、4、5节间茎秆强度和Si含量均呈现显著正相关,与其他指标相关性因节间和生育时期而不同。
4、根系特征及生理特性结果为：气生根和总根数量随着密度的增加先微升高后降低,在7万株/公顷数量最多。先玉420根系干重随着密度的变化先增加后降低,益丰29、先玉335和先玉420均呈降低-升高-降低的变化,低谷和高峰分别出现在6万株/公顷和7万株/公顷。从密度8万株/公顷,各品种根系活力开始下降,郑单958降幅最大。在玉米抽雄期,根系SOD、POD和CAT活性在7万株/公顷达最大值后,密度继续增加,酶活性降低；在灌浆初期,先玉335的SOD和POD活性随密度变化呈现“M,,型曲线,其他品种均为单峰曲线。各项指标均为郑单958和先玉335高于益丰29和先玉420,丙二醛相反。玉米倒伏率与气生根和总根数量、干物质百分比和根系活力呈现极显著负相关,与其他指标相关性不大。
5、“布拉万”台风对玉米倒伏及产量的影响结果为：供试11个玉米品种以良玉99倒伏能力最强,其穗位高系数、第1-6节间平均直径、节间干重、单位茎长干重、茎秆可溶性糖含量、木质素含量、Si含量及茎秆强度均处于较高水平；其气生根数目、根系干重及干物质比较高。银河32和银河33抗倒伏能力最差,各项指标均处于最低水平。尽管良玉99抗倒伏能力较强,但其产量显著低于郑单958和先玉335。2012年,郑单958产量仅比2009-2011年三年平均产量降低5.13%,表现出较强的产量稳定性。
综上所述,群体密度对玉米茎秆和根系形态、干物质及相关生理特性均产生一定的影响。在有限的资源下,密度过高或过低均不利于生长,产生一定的密度效应。玉米倒伏是品种特性、栽培技术及土壤、气候等内外因素综合作用的结果。因此,在选择玉米抗倒伏品种时,应综合考虑形态和生理指标,合理密植,合理施肥,适量追施微量元素肥料,以降低倒伏的风险,保证玉米生产潜力得到充分发挥。此外,抗倒伏能力和产量并非呈现极显著的正相关,如何选育出抗倒伏能力强且产量高的玉米品种,或是如何通过栽培技术措施使得抗倒伏玉米品种能够最大程度的发挥出高产潜力,实现良种良法的配套是未来育种及栽培工作者共同面临的挑战和研究方向。本研究仅为玉米抗倒伏形态及生理机制提供科学的理论依据。
The planting of high density had been one of the key measures for improving yield of maize, while the increasing of population density increased the risk of lodging. The average precipitation each year was654.3mm and centralized in July and August, which was the key stage during tasseling to filling stage. Under the effects of high temperature and rainy, lodging had become one of the most important reason of reducing yield of maize. In the past, there were some studies about lodging of maize, while the systematic study about the effects of population density on stalk and root was infrequent. In this study, the effects of different population density on the morphology, dry matter and physiological characteristics of stalk and root were studied, and the relationship between lodging percentage and all indexes was determined to reveal the mechanism of lodging of maize, and provide theoretical foundation for breeding and cultivate of lodging-resistance.
In this paper, four different varieties were used as the test material and seven density treatment levels were applied, and the random block design was used. The morphology, dry matter and physiological characteristics of stalk and root were measured, and the relationship between lodging percentage and all indexes was determined. In2012,11leading varieties of Jilin province were used. And the effects of Blavin typhoon on lodging and yield of maize were studied from the point of morphology and physiology.
1、The results of stalk agronomic traits showed that:The ear height and ear height coefficient were increased with the increasing of density, and that of zhengdan958were not sensitive to the changes of density. Lodging percentage of xianyu335and xianyu420showed positive significantly correlation with ear height and ear height coefficient, while that of yifeng29and zhengdan958had not correlation significantly. The circumference of internode was decreased with the increasing of density, while the length of internode had not regular changes. Dry matter percentage and strength of stalk were increased first and then decreased with the increase of population density. The lodging percentage showed significantly correlated with the length, circumference, dry weight and strength of internode. Except plant height, other indexes of yifeng29and zhengdan958were higher than those of xianyu335and xianyu420. The stalk strength was positively correlated with K, Si and starch content, and was not positively correlated with Zn and soluble sugar content.
2、The results of ligin and its key synthetase showed that:under different density, the content of ligin and cellulose, the activities of phenylalanine ammonia-lyase(PAL), tyrosine ammonia-lyase(TAL) and cinnamyl alcohol dehydrogenase(CAD) of yifent29were higher than those of xianyu420. The order of PAL and TAL activities was3th>4th>5th internode. The order of CAD activities in xianyu420was4th>3th>5th internode, while that of yifeng29was3th>4th>5th internode. The ligin content of xianyu420had a positive significantly correlation with CAD activity, while the ligin content of yifeng29had a positive significantly correlation with PAL and TAL activities. At tasseling and filling stage, the stalk strength of Yifeng29had a positive significantly correlation with ligin content, activity of PAL and TAL yet that of Xianyu420had only significant and positive correlation with TAL activity.
3、The results of chemical components and mineral elements showed that:mineral elements including nitrogen(N),potassium(K),silicon(Si),calcium(Ca),magnesium(Mg) and zinc(Zn) were increased first and then decreased with the increase of population density. The peak occurred under the treatment of seven and eight thousand plants per hectare, which was different with variety and growth stage. The contents of N, K, Si, Ca, Mg and Zn of yifeng29were higher that those of xianyu420. The stalk strength was positively correlated with K, Si and starch content, and was not positively correlated with Zn and soluble sugar content.
4、The results of root characterstics showed:The numbers of brace root and total root were increased first and then decreased with the increase of population density, which were the highest under the treatment of70,000/ha. The root dry weight of xianyu420was increase first then reduced,while those of yifeng29, xianyu335,xianyu420were decreased first. then increased, and then decreased, and the trough and peak occurred under the treatment of60,000/ha and70,000/ha. The activity of root was decreased under the treatment of80,000/ha, and the decreased degree reached the highest. At the tasseling stage of maize, SOD, POD and CAT activity of root had maximum value under the treatment of70,000/ha. At the filling stage, SOD and POD activities of Xianyu335presented the "M" shaped curve with the increase of density, while presented single-peak curve. The indicators of zhengdan958and xianyu335were higher than those of yifeng29and xianyu420. The malondialdehyde content is the opposite. The lodging percentage showed negative significantly correlation with the number of brace root and total root, dry matter percentage and root activity.
5、Effects of the Blavin typhoon on lodging and yield of maize showed that:in the11test varieties, the ability of lodging-resistance, ear high coefficient, average diameter, dry weight of internode, soluble sugar content, ligin content, Si content and stalk strength were highest in liangyu99varieties. And the number of brace root, root dry weight and dry matter were all higher in liangyu99. The lodging-resistance of yihe32and yihe33were the poorest. Although liangyu99had strong ability of lodging-resistance, its yield was lower than zhengdan958and xianyu335. In2012, the yield of zhengdan958was decreased5.13percent compared to the average yield in2009-2011, which had strong yield stability.
In summary, there were some effects of population density on the morphology, dry matter and physiological characteristics of stalk and root. In limited resources, too high or too low density was all bad to the growth of corn, which may product the density effect. Lodging of maize was the results of comprehensive effects including variety, cultivation technique, soil and climate. Thus, we have to consider all effects, and have rational close planting and fertilizer, and dress the trace elements to reduce the risk of lodging. There was not all positive correlation between the ability of lodging-resistance and yield. It was a challenge to breed the maize variety which had strong lodging-resistance and high yield, and play high-yield potential through the measure of cultivation technique. The study provided scientific foundation for lodging mechanism of maize.
本研究选择7个种植密度,2009-2011年选择4个抗倒伏性能不同的品种,采用随机区组设计,通过茎秆和根系形态、干物质及相关生理指标的测定,分析其和茎秆强度及倒伏率的相关性。2012年,以吉林省11个主导品种为试验材料,从形态学和生理学角度,研究“布拉万”台风对不同玉米品种的影响,探明不同玉米品种的倒伏特征。研究结果表明：
1、茎秆农艺性状的结果为：穗位高和穗位高系数随着密度的增加逐渐增加,株高在7万株／公顷的密度下达到最大；郑单958的穗位高和穗位高系数对密度的变化不敏感。先玉420和先玉335倒伏率与穗位高和穗位高系数呈现极显著正相关,益丰29和郑单958则与株高、穗位高和穗位高系数均无显著相关性。同一品种相同节间粗随密度增加逐渐降低、节间长度无规律性变化。节间干物质百分比、单位茎长干重、节间强度均随密度增加先升高后降低,在7和8万株／公顷出现最大值,各品种表现不同。两个玉米品种其倒伏率与第3、4节茎长、茎粗、单位茎长干重、茎秆强度均显著相关。品种间比较,益丰29和郑单958第2-6节间干物质百分比、单位茎长干重、节间强度、穗位高及穗位高系数均高于先玉335和先玉420,株高则相反。
2、茎秆木质素及关键合成酶活性结果为：不同密度处理下,平均节间纤维素含量、木质素含量、PAL活性、TAL活性和CAD活性均为益丰29高于先玉420。益丰29节间PAL、TAL活性节间表现为第3节间>第4节间>第5节间。CAD活性各节间变化因品种而不同,先玉420第4节>第3节>第5节,益丰29第3节>第4节>第5节。先玉420节间木质素含量与CAD活性显著正相关,益丰29则与PAL活性和TAL活性呈显著正相关。在抽雄期和灌浆初期,益丰29节间强度与木质素含量、PAL活性、TAL活性显著正相关,先玉420与TAL呈显著正相关。
3、茎秆化学成分及矿质元素结果为：随着密度的增加,节间N、K、Si、Ca、Mg、Zn含量均先升高后降低,最高点均出现在7万株／公顷和8万株／公顷,因品种和生育时期而不同。品种间益丰29高于先玉420。玉米茎秆强度与K、Si和淀粉含量呈现显著正相关,与N、Ca、Mg含量呈现正相关,但不显著,与Zn和可溶性糖含量相关性不大。益丰29和先玉420第3、4、5节间茎秆强度和Si含量均呈现显著正相关,与其他指标相关性因节间和生育时期而不同。
4、根系特征及生理特性结果为：气生根和总根数量随着密度的增加先微升高后降低,在7万株/公顷数量最多。先玉420根系干重随着密度的变化先增加后降低,益丰29、先玉335和先玉420均呈降低-升高-降低的变化,低谷和高峰分别出现在6万株/公顷和7万株/公顷。从密度8万株/公顷,各品种根系活力开始下降,郑单958降幅最大。在玉米抽雄期,根系SOD、POD和CAT活性在7万株/公顷达最大值后,密度继续增加,酶活性降低；在灌浆初期,先玉335的SOD和POD活性随密度变化呈现“M,,型曲线,其他品种均为单峰曲线。各项指标均为郑单958和先玉335高于益丰29和先玉420,丙二醛相反。玉米倒伏率与气生根和总根数量、干物质百分比和根系活力呈现极显著负相关,与其他指标相关性不大。
5、“布拉万”台风对玉米倒伏及产量的影响结果为：供试11个玉米品种以良玉99倒伏能力最强,其穗位高系数、第1-6节间平均直径、节间干重、单位茎长干重、茎秆可溶性糖含量、木质素含量、Si含量及茎秆强度均处于较高水平；其气生根数目、根系干重及干物质比较高。银河32和银河33抗倒伏能力最差,各项指标均处于最低水平。尽管良玉99抗倒伏能力较强,但其产量显著低于郑单958和先玉335。2012年,郑单958产量仅比2009-2011年三年平均产量降低5.13%,表现出较强的产量稳定性。
综上所述,群体密度对玉米茎秆和根系形态、干物质及相关生理特性均产生一定的影响。在有限的资源下,密度过高或过低均不利于生长,产生一定的密度效应。玉米倒伏是品种特性、栽培技术及土壤、气候等内外因素综合作用的结果。因此,在选择玉米抗倒伏品种时,应综合考虑形态和生理指标,合理密植,合理施肥,适量追施微量元素肥料,以降低倒伏的风险,保证玉米生产潜力得到充分发挥。此外,抗倒伏能力和产量并非呈现极显著的正相关,如何选育出抗倒伏能力强且产量高的玉米品种,或是如何通过栽培技术措施使得抗倒伏玉米品种能够最大程度的发挥出高产潜力,实现良种良法的配套是未来育种及栽培工作者共同面临的挑战和研究方向。本研究仅为玉米抗倒伏形态及生理机制提供科学的理论依据。
The planting of high density had been one of the key measures for improving yield of maize, while the increasing of population density increased the risk of lodging. The average precipitation each year was654.3mm and centralized in July and August, which was the key stage during tasseling to filling stage. Under the effects of high temperature and rainy, lodging had become one of the most important reason of reducing yield of maize. In the past, there were some studies about lodging of maize, while the systematic study about the effects of population density on stalk and root was infrequent. In this study, the effects of different population density on the morphology, dry matter and physiological characteristics of stalk and root were studied, and the relationship between lodging percentage and all indexes was determined to reveal the mechanism of lodging of maize, and provide theoretical foundation for breeding and cultivate of lodging-resistance.
In this paper, four different varieties were used as the test material and seven density treatment levels were applied, and the random block design was used. The morphology, dry matter and physiological characteristics of stalk and root were measured, and the relationship between lodging percentage and all indexes was determined. In2012,11leading varieties of Jilin province were used. And the effects of Blavin typhoon on lodging and yield of maize were studied from the point of morphology and physiology.
1、The results of stalk agronomic traits showed that:The ear height and ear height coefficient were increased with the increasing of density, and that of zhengdan958were not sensitive to the changes of density. Lodging percentage of xianyu335and xianyu420showed positive significantly correlation with ear height and ear height coefficient, while that of yifeng29and zhengdan958had not correlation significantly. The circumference of internode was decreased with the increasing of density, while the length of internode had not regular changes. Dry matter percentage and strength of stalk were increased first and then decreased with the increase of population density. The lodging percentage showed significantly correlated with the length, circumference, dry weight and strength of internode. Except plant height, other indexes of yifeng29and zhengdan958were higher than those of xianyu335and xianyu420. The stalk strength was positively correlated with K, Si and starch content, and was not positively correlated with Zn and soluble sugar content.
2、The results of ligin and its key synthetase showed that:under different density, the content of ligin and cellulose, the activities of phenylalanine ammonia-lyase(PAL), tyrosine ammonia-lyase(TAL) and cinnamyl alcohol dehydrogenase(CAD) of yifent29were higher than those of xianyu420. The order of PAL and TAL activities was3th>4th>5th internode. The order of CAD activities in xianyu420was4th>3th>5th internode, while that of yifeng29was3th>4th>5th internode. The ligin content of xianyu420had a positive significantly correlation with CAD activity, while the ligin content of yifeng29had a positive significantly correlation with PAL and TAL activities. At tasseling and filling stage, the stalk strength of Yifeng29had a positive significantly correlation with ligin content, activity of PAL and TAL yet that of Xianyu420had only significant and positive correlation with TAL activity.
3、The results of chemical components and mineral elements showed that:mineral elements including nitrogen(N),potassium(K),silicon(Si),calcium(Ca),magnesium(Mg) and zinc(Zn) were increased first and then decreased with the increase of population density. The peak occurred under the treatment of seven and eight thousand plants per hectare, which was different with variety and growth stage. The contents of N, K, Si, Ca, Mg and Zn of yifeng29were higher that those of xianyu420. The stalk strength was positively correlated with K, Si and starch content, and was not positively correlated with Zn and soluble sugar content.
4、The results of root characterstics showed:The numbers of brace root and total root were increased first and then decreased with the increase of population density, which were the highest under the treatment of70,000/ha. The root dry weight of xianyu420was increase first then reduced,while those of yifeng29, xianyu335,xianyu420were decreased first. then increased, and then decreased, and the trough and peak occurred under the treatment of60,000/ha and70,000/ha. The activity of root was decreased under the treatment of80,000/ha, and the decreased degree reached the highest. At the tasseling stage of maize, SOD, POD and CAT activity of root had maximum value under the treatment of70,000/ha. At the filling stage, SOD and POD activities of Xianyu335presented the "M" shaped curve with the increase of density, while presented single-peak curve. The indicators of zhengdan958and xianyu335were higher than those of yifeng29and xianyu420. The malondialdehyde content is the opposite. The lodging percentage showed negative significantly correlation with the number of brace root and total root, dry matter percentage and root activity.
5、Effects of the Blavin typhoon on lodging and yield of maize showed that:in the11test varieties, the ability of lodging-resistance, ear high coefficient, average diameter, dry weight of internode, soluble sugar content, ligin content, Si content and stalk strength were highest in liangyu99varieties. And the number of brace root, root dry weight and dry matter were all higher in liangyu99. The lodging-resistance of yihe32and yihe33were the poorest. Although liangyu99had strong ability of lodging-resistance, its yield was lower than zhengdan958and xianyu335. In2012, the yield of zhengdan958was decreased5.13percent compared to the average yield in2009-2011, which had strong yield stability.
In summary, there were some effects of population density on the morphology, dry matter and physiological characteristics of stalk and root. In limited resources, too high or too low density was all bad to the growth of corn, which may product the density effect. Lodging of maize was the results of comprehensive effects including variety, cultivation technique, soil and climate. Thus, we have to consider all effects, and have rational close planting and fertilizer, and dress the trace elements to reduce the risk of lodging. There was not all positive correlation between the ability of lodging-resistance and yield. It was a challenge to breed the maize variety which had strong lodging-resistance and high yield, and play high-yield potential through the measure of cultivation technique. The study provided scientific foundation for lodging mechanism of maize.
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