高梁子粒淀粉遗传特性及积累规律的研究
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摘要
我国80%的粒用高粱用于酿酒业,出酒率的高低主要取决于淀粉含量。淀粉是高粱子粒的主要成分,其含量高低是决定高粱酿造产量和品质的重要因素。目前,在粒用高粱的研究和应用上主要以高淀粉酿造型高粱为主。对高粱子粒淀粉含量遗传特性、积累规律及调控技术进行深入研究,无疑对高粱育种、栽培的理论和实践上都有重要的现实意义。但有关高粱子粒淀粉遗传特性、淀粉积累规律及其与叶片生理关系,以及钾肥调控对高粱子粒淀粉积累特性的影响研究很少或尚未见系统报道。
本研究以高淀粉和低淀粉高粱不育系和恢复系,利用NCⅡ设计的不完全双列杂交设计组配杂交组合,通过对杂交种的淀粉含量进行测定,分析淀粉在亲本和杂交种间的遗传表现。以高淀粉和低淀粉高粱为试验材料,研究了高淀粉和低淀粉高粱淀粉积累特性及其与叶片生理关系,以及钾肥调控对淀粉积累特性的影响。主要结果与结论如下。
1、高粱总淀粉含量、直链淀粉含量和支链淀粉含量的一般配合力效应和特殊配合力效应对3个性状的表型值均有重要作用。在高淀粉高粱杂交种的选育中,首先要重视对双亲一般配合力的选择。在一般配合力高的基础上更多的选择有较高的特殊配合力的组合,才能使杂交种获得较高的淀粉含量。淀粉含量占高粱籽粒产量的70～80%,对高粱的品质性状有着决定性的影响。亲本会对杂交后代总淀粉中的直(支)链淀粉比例产生显著影响。高淀粉含量主要与高支链淀粉含量有关。由于支链淀粉的合成需要消耗更多的能量,因此对这个性状的改良难度更大。本研究结果表明,恢复系群体的一般配合力效应值多为正值,通过加强对这个群体的性状改进,选育出支链淀粉含量更高的恢复系是可行的。
2、高淀粉和低淀粉高粱子粒淀粉积累特性研究结果表明,高粱子粒总淀粉、支链淀粉和直链淀粉含量随子粒灌浆过程推进均呈不断上升趋势,至成熟期达到最大值。在子粒灌浆过程中,高淀粉高粱和低淀粉高粱子粒淀粉含量变化趋势均呈“S”型曲线变化；高淀粉高粱和低淀粉高粱在灌浆初期子粒直链淀粉含量并无显著差异,而在灌浆中后期才逐渐表现差异；与直链淀粉积累不同,高淀粉高粱子粒支链淀粉含量在灌浆初期就显著高于低淀粉高粱子粒支链淀粉含量；子粒总淀粉积累的差异主要是由于整个灌浆期支链淀粉积累和灌浆中后期直链淀粉积累的不同造成的。
3、高淀粉和低淀粉高粱开花后叶片光合指标及其与子粒淀粉含量关系研究结果表明,高淀粉高粱叶片比叶重、叶绿素含量和净光合速率均高于低淀粉高粱叶片。在灌浆中后期,子粒淀粉含量与叶片比叶重、叶绿素含量、净光合速率之间存在显著正相关关系。其中子粒淀粉含量和净光合速率之间的相关性要比淀粉含量与比叶重、叶绿素含量之间的相关性密切。由于现代技术手段的改进,可方便地进行叶片净光合速率的瞬时、准确测定,因此可以把净光合速率作为选择高淀粉高粱的一个有效指标。
4、高淀粉和低淀粉高粱开花后叶片保护酶活性、丙二醛含量及其与子粒淀粉含量关系研究结果表明,高粱开花后,叶片SOD、POD和CAT活性均呈单峰曲线变化,SOD和CAT活性峰值出现在花后28天,POD活性峰值出现在花后21天。高淀粉与低淀粉高粱叶片SOD和CAT活性在花后14天内差异不显著；在开花21天后,高淀粉高粱叶片SOD和CAT活性显著高于低淀粉高粱叶片SOD和CAT活性。在开花后35天内,高淀粉高粱叶片POD活性显著高于低淀粉高粱叶片POD活性,42天后,高淀粉与低淀粉高粱叶片POD活性差异不大。在开花21天后,子粒淀粉含量与叶片SOD、CAT活性之间存在显著正相关关系；在开花35天内,子粒淀粉含量与叶片POD活性之间存在显著正相关关系。高淀粉与低淀粉高粱叶片MDA含量在花后14天内差异不显著；在开花21天后,高淀粉高粱叶片MDA含量显著低于低淀粉高粱叶片MDA含量,子粒淀粉含量与叶片MDA含量之间存在显著负相关关系。高淀粉高粱开花后,叶片保持较高的保护酶活性和较低的丙二醛含量,使叶片衰老减缓,为保持较高光合作用强度奠定基础。
5、钾肥调控对高粱植株干物质积累与分配的影响研究结果表明,施用钾肥增加高粱植株有效绿叶叶面积和干物质积累量,使干物质向经济器官积累增多,提高经济系数,增加子粒产量。其中以每公顷基施钾肥90kg,追施钾肥30kg产量最高。
6、钾肥调控对高粱子粒淀粉积累特性的影响研究结果表明,施用钾肥明显提高子粒总淀粉、支链淀粉和直链淀粉含量；施钾肥提高灌浆期高粱总淀粉积累速率和支链淀粉积累速率,以及灌浆中后期直链淀粉积累速率。其中以每公顷基施钾肥90kg,追施钾肥30kg子粒淀粉含量最高。
7、钾肥调控对高粱叶片光合生理指标的影响研究结果表明,施用钾肥可以增加高粱花后叶片比叶重、叶绿素含量和净光合速率,这是施钾增产量的原因之一。其中以每公顷基施钾肥90kg,再追施钾肥30kg效果最好。
8、钾肥调控对高粱叶片保护酶活性和丙二醛含量的影响研究结果表明,施钾肥调控能显著提高高粱叶片SOD、POD和CAT活性,降低叶片MDA含量,有利于活性氧的清除,延缓叶片衰老进程,这是施用钾肥提高子粒产量的原因之一。
About 80% of sorghum grains is used to brew. The alcohol yield mainly depends on starch content. And starch is the main component of grain sorghum. Its high-low contents are the important factors which decide sorghum brewing quality. At present, in grain sorghum research and application, we mainly use the brewery sorghum with high starch. The relevant in-depth researches, such as in sorghum starch content, genetic characteristics and starch accumulation regulation, have no doubt important practical significance on sorghum breeding, cultivation theory and practice. But the researches are rare or have not been reported in the genetic characteristics of sorghum grain starch, starch accumulation regulation, its physiological relationship with leaves and the influence of potassium control on starch accumulating characteristics. In sorghum sterility and restorers of high starch and low starch, through the use of B. Griffing incomplete double row cross to design hybrids and measuring the starch content of cross seeds, the study analyzed the genetic performances of starches between parents and hybrids.
With high starch and low starch sorghum as test materials, it studied the sorghum starch accumulation properties of high starch and low starch and their relations with leaf physiological properties, and the influence of potassium regulation on starch accumulation. The main results and conclusions were as followed.
1. The content of parental sorghum amylopectin and amylose would have a significant effect on the proportion of amylopectin and amylose of the total starch in the offspring. The general combining ability and specific combining ability for the contents of total sorghum starch, amylopectin and amylose were highly significant. It indicated that both additive and non-additive effects had important roles on the three phenotypic traits. We should attach importance to the choice of GCA in sorghum hybrid breeding of the high starch. In order to make hybrids get a higher starch content, we should choose higher combination of specific combining ability on the basis of general combining ability. It will have a decisive influence on the quality and traits of sorghum if starch content occupies 70 to 80% of sorghum grain yield, especially amylopectin. Since the synthesis of amylopectin need to consume more energy, the improvement of this trait is more difficult. The results showed that the value of general combining ability of the restorer line group is more positive and we should strengthen to improve the traits of this group. It is feasible to breed a restorer with a higher amylopectin content.
2. The research on the accumulation characteristics of sorghum grain starch of high starch and low starch. The results showed that total starch, amylopectin and amylose contents increased continuously along with the advance of grain filling process and reached the maximum at maturity stage. The changes of the starch accumulation in both high and low starch sorghum during grain filling stage showed as "S" curve. The amylose accumulation rate was not significantly different between the high starch sorghum and low starch sorghum at the early stage of grain filling. The amylose accumulation rate was significantly different between the high starch sorghum and low starch sorghum at the middle and later stages of grain filling. The difference in amylopectin content appeared at the early stage of grain filling, and amylopectin of high starch sorghum were constantly higher than that of low starch sorghum during grain filling period. The amylopectin accumulation depended on the average filling rate and the high filling rate at the early stage of grain filling. The difference in the total starch content was mainly ascribed to amylopectin content at the all stages of grain filling, and amylose content at the middle and later stages of grain filling.
3. The researches on the leaf photosynthesis index after high starch and low starch sorghum flowering and its relationship with grain starch content. The results showed that the weights, chlorophyll contents and net photosynthetic rates of high-starch sorghum leaves were higher than the low starch sorghum leaves. In the late filling stage, there was a significant positive correlation among grain starch content, leaf weight, chlorophyll content and net photosynthetic rate. Among them, the correlation between grain starch content and net photosynthetic rate was closer than the correlation among the starch content, specific leaf weight and chlorophyll content. Because of the improvement of modern technological means, we can easily make Instantaneous and accurate determination to leaf net photosynthetic rate. So we take the net photosynthetic rate as an effective indicator of high sorghum starch.
4. The research on the relationship among the leaf protective enzyme activities, MDA content and grain starch content after flowering of high and low starch sorghum. The results showed that after sorghum flowering, the activity of leaf SOD. POD and CAT showed a single peak curve. The activity peak of SOD and CAT appeared at 28 days after flowering and the activity peak of SOD appeared at 21 days after flowering. There were no significant differences about the activity of high starch and low starch sorghum leaves SOD and CAT within 14 days after flowering.21 days after flowering later, the activities of high-starch sorghum leaves SOD and CAT were significantly higher than those of the low-starch SOD and CAT. In 35 days after flowering, the activity of leaf POD of the high-starch sorghum was significantly higher than the activity of leaf POD of the low-starch of sorghum.42 days later, sorghum leaf POD activities between high starch and low starch were similar.21 days after flowering, there was a significant positive correlation between grain starch content and the activity of leaf SOD, CAT; In 35 days after flowering, there was a significant positive correlation between grain starch content and the activity of leaf POD.14 days after flowering, there were not significant differences in MDA content between high starch and low starch sorghum; 21 days after flowering, the content of high starch sorghum leaves MDA was significantly lower than the content of low starch sorghum leaves MDA and there was a significant negative correlation between grain starch content and the leaf MDA content. After high-starch sorghum flowering, the leaves keep a higher enzymes activity and a lower MDA content to slow leaf senescence and lay a foundation in maintaining a high photosynthesis.
5. The research on the influence of regulation of potassium on dry matter accumulation and distribution of sorghum. The results showed that using fertilizer effectively increased the green leaf area of sorghum plants and the accumulation of dry matter and made dry matter increase the accumulation of economic organ, improved economic factors and increased grain yield. The yield will be the highest if we use 90kg of Potassium in per hectare, and then add another 30kg potassium fertilizer to it.
6. The research on the influence of potassium regulation on the accumulation characteristics of sorghum grain starch. The results showed that using K fertilizer significantly increased the total sorghum starch accumulation rate during the filling stage, accumulation rate of amylopectin and accumulation rate of amylose in mid or late filling stage. using K fertilizer increased filling rate of and and amylose in the late filling accumulation rate. The starch content will be the highest if we use 90kg of Potassium in per hectare, and then add another 30kg potassium fertilizer to it.
7. The research on the influence of potassium regulation on physiological index of sorghum leaves. The results showed that using K fertilizer could increase the sorghum leaf weight after flowering, chlorophyll content and net photosynthetic rate. It was one of the reasons which using K fertilizer could increase production. The effect will be the best if we use 90kg of Potassium, and then add another 30kg potassium fertilizer to it.
8. The research on the influence of potassium regulation on protective enzyme activities of sorghum leaves and MDA content. The results showed that using K fertilizer could significantly improve the activity of sorghum leaf SOD, POD and CAT, decrease MDA content in leaves and be conducive to the removal of active oxygen, delay leaf senescence, which is one of the reasons to improve grain yield by using K fertilizer.
本研究以高淀粉和低淀粉高粱不育系和恢复系,利用NCⅡ设计的不完全双列杂交设计组配杂交组合,通过对杂交种的淀粉含量进行测定,分析淀粉在亲本和杂交种间的遗传表现。以高淀粉和低淀粉高粱为试验材料,研究了高淀粉和低淀粉高粱淀粉积累特性及其与叶片生理关系,以及钾肥调控对淀粉积累特性的影响。主要结果与结论如下。
1、高粱总淀粉含量、直链淀粉含量和支链淀粉含量的一般配合力效应和特殊配合力效应对3个性状的表型值均有重要作用。在高淀粉高粱杂交种的选育中,首先要重视对双亲一般配合力的选择。在一般配合力高的基础上更多的选择有较高的特殊配合力的组合,才能使杂交种获得较高的淀粉含量。淀粉含量占高粱籽粒产量的70～80%,对高粱的品质性状有着决定性的影响。亲本会对杂交后代总淀粉中的直(支)链淀粉比例产生显著影响。高淀粉含量主要与高支链淀粉含量有关。由于支链淀粉的合成需要消耗更多的能量,因此对这个性状的改良难度更大。本研究结果表明,恢复系群体的一般配合力效应值多为正值,通过加强对这个群体的性状改进,选育出支链淀粉含量更高的恢复系是可行的。
2、高淀粉和低淀粉高粱子粒淀粉积累特性研究结果表明,高粱子粒总淀粉、支链淀粉和直链淀粉含量随子粒灌浆过程推进均呈不断上升趋势,至成熟期达到最大值。在子粒灌浆过程中,高淀粉高粱和低淀粉高粱子粒淀粉含量变化趋势均呈“S”型曲线变化；高淀粉高粱和低淀粉高粱在灌浆初期子粒直链淀粉含量并无显著差异,而在灌浆中后期才逐渐表现差异；与直链淀粉积累不同,高淀粉高粱子粒支链淀粉含量在灌浆初期就显著高于低淀粉高粱子粒支链淀粉含量；子粒总淀粉积累的差异主要是由于整个灌浆期支链淀粉积累和灌浆中后期直链淀粉积累的不同造成的。
3、高淀粉和低淀粉高粱开花后叶片光合指标及其与子粒淀粉含量关系研究结果表明,高淀粉高粱叶片比叶重、叶绿素含量和净光合速率均高于低淀粉高粱叶片。在灌浆中后期,子粒淀粉含量与叶片比叶重、叶绿素含量、净光合速率之间存在显著正相关关系。其中子粒淀粉含量和净光合速率之间的相关性要比淀粉含量与比叶重、叶绿素含量之间的相关性密切。由于现代技术手段的改进,可方便地进行叶片净光合速率的瞬时、准确测定,因此可以把净光合速率作为选择高淀粉高粱的一个有效指标。
4、高淀粉和低淀粉高粱开花后叶片保护酶活性、丙二醛含量及其与子粒淀粉含量关系研究结果表明,高粱开花后,叶片SOD、POD和CAT活性均呈单峰曲线变化,SOD和CAT活性峰值出现在花后28天,POD活性峰值出现在花后21天。高淀粉与低淀粉高粱叶片SOD和CAT活性在花后14天内差异不显著；在开花21天后,高淀粉高粱叶片SOD和CAT活性显著高于低淀粉高粱叶片SOD和CAT活性。在开花后35天内,高淀粉高粱叶片POD活性显著高于低淀粉高粱叶片POD活性,42天后,高淀粉与低淀粉高粱叶片POD活性差异不大。在开花21天后,子粒淀粉含量与叶片SOD、CAT活性之间存在显著正相关关系；在开花35天内,子粒淀粉含量与叶片POD活性之间存在显著正相关关系。高淀粉与低淀粉高粱叶片MDA含量在花后14天内差异不显著；在开花21天后,高淀粉高粱叶片MDA含量显著低于低淀粉高粱叶片MDA含量,子粒淀粉含量与叶片MDA含量之间存在显著负相关关系。高淀粉高粱开花后,叶片保持较高的保护酶活性和较低的丙二醛含量,使叶片衰老减缓,为保持较高光合作用强度奠定基础。
5、钾肥调控对高粱植株干物质积累与分配的影响研究结果表明,施用钾肥增加高粱植株有效绿叶叶面积和干物质积累量,使干物质向经济器官积累增多,提高经济系数,增加子粒产量。其中以每公顷基施钾肥90kg,追施钾肥30kg产量最高。
6、钾肥调控对高粱子粒淀粉积累特性的影响研究结果表明,施用钾肥明显提高子粒总淀粉、支链淀粉和直链淀粉含量；施钾肥提高灌浆期高粱总淀粉积累速率和支链淀粉积累速率,以及灌浆中后期直链淀粉积累速率。其中以每公顷基施钾肥90kg,追施钾肥30kg子粒淀粉含量最高。
7、钾肥调控对高粱叶片光合生理指标的影响研究结果表明,施用钾肥可以增加高粱花后叶片比叶重、叶绿素含量和净光合速率,这是施钾增产量的原因之一。其中以每公顷基施钾肥90kg,再追施钾肥30kg效果最好。
8、钾肥调控对高粱叶片保护酶活性和丙二醛含量的影响研究结果表明,施钾肥调控能显著提高高粱叶片SOD、POD和CAT活性,降低叶片MDA含量,有利于活性氧的清除,延缓叶片衰老进程,这是施用钾肥提高子粒产量的原因之一。
About 80% of sorghum grains is used to brew. The alcohol yield mainly depends on starch content. And starch is the main component of grain sorghum. Its high-low contents are the important factors which decide sorghum brewing quality. At present, in grain sorghum research and application, we mainly use the brewery sorghum with high starch. The relevant in-depth researches, such as in sorghum starch content, genetic characteristics and starch accumulation regulation, have no doubt important practical significance on sorghum breeding, cultivation theory and practice. But the researches are rare or have not been reported in the genetic characteristics of sorghum grain starch, starch accumulation regulation, its physiological relationship with leaves and the influence of potassium control on starch accumulating characteristics. In sorghum sterility and restorers of high starch and low starch, through the use of B. Griffing incomplete double row cross to design hybrids and measuring the starch content of cross seeds, the study analyzed the genetic performances of starches between parents and hybrids.
With high starch and low starch sorghum as test materials, it studied the sorghum starch accumulation properties of high starch and low starch and their relations with leaf physiological properties, and the influence of potassium regulation on starch accumulation. The main results and conclusions were as followed.
1. The content of parental sorghum amylopectin and amylose would have a significant effect on the proportion of amylopectin and amylose of the total starch in the offspring. The general combining ability and specific combining ability for the contents of total sorghum starch, amylopectin and amylose were highly significant. It indicated that both additive and non-additive effects had important roles on the three phenotypic traits. We should attach importance to the choice of GCA in sorghum hybrid breeding of the high starch. In order to make hybrids get a higher starch content, we should choose higher combination of specific combining ability on the basis of general combining ability. It will have a decisive influence on the quality and traits of sorghum if starch content occupies 70 to 80% of sorghum grain yield, especially amylopectin. Since the synthesis of amylopectin need to consume more energy, the improvement of this trait is more difficult. The results showed that the value of general combining ability of the restorer line group is more positive and we should strengthen to improve the traits of this group. It is feasible to breed a restorer with a higher amylopectin content.
2. The research on the accumulation characteristics of sorghum grain starch of high starch and low starch. The results showed that total starch, amylopectin and amylose contents increased continuously along with the advance of grain filling process and reached the maximum at maturity stage. The changes of the starch accumulation in both high and low starch sorghum during grain filling stage showed as "S" curve. The amylose accumulation rate was not significantly different between the high starch sorghum and low starch sorghum at the early stage of grain filling. The amylose accumulation rate was significantly different between the high starch sorghum and low starch sorghum at the middle and later stages of grain filling. The difference in amylopectin content appeared at the early stage of grain filling, and amylopectin of high starch sorghum were constantly higher than that of low starch sorghum during grain filling period. The amylopectin accumulation depended on the average filling rate and the high filling rate at the early stage of grain filling. The difference in the total starch content was mainly ascribed to amylopectin content at the all stages of grain filling, and amylose content at the middle and later stages of grain filling.
3. The researches on the leaf photosynthesis index after high starch and low starch sorghum flowering and its relationship with grain starch content. The results showed that the weights, chlorophyll contents and net photosynthetic rates of high-starch sorghum leaves were higher than the low starch sorghum leaves. In the late filling stage, there was a significant positive correlation among grain starch content, leaf weight, chlorophyll content and net photosynthetic rate. Among them, the correlation between grain starch content and net photosynthetic rate was closer than the correlation among the starch content, specific leaf weight and chlorophyll content. Because of the improvement of modern technological means, we can easily make Instantaneous and accurate determination to leaf net photosynthetic rate. So we take the net photosynthetic rate as an effective indicator of high sorghum starch.
4. The research on the relationship among the leaf protective enzyme activities, MDA content and grain starch content after flowering of high and low starch sorghum. The results showed that after sorghum flowering, the activity of leaf SOD. POD and CAT showed a single peak curve. The activity peak of SOD and CAT appeared at 28 days after flowering and the activity peak of SOD appeared at 21 days after flowering. There were no significant differences about the activity of high starch and low starch sorghum leaves SOD and CAT within 14 days after flowering.21 days after flowering later, the activities of high-starch sorghum leaves SOD and CAT were significantly higher than those of the low-starch SOD and CAT. In 35 days after flowering, the activity of leaf POD of the high-starch sorghum was significantly higher than the activity of leaf POD of the low-starch of sorghum.42 days later, sorghum leaf POD activities between high starch and low starch were similar.21 days after flowering, there was a significant positive correlation between grain starch content and the activity of leaf SOD, CAT; In 35 days after flowering, there was a significant positive correlation between grain starch content and the activity of leaf POD.14 days after flowering, there were not significant differences in MDA content between high starch and low starch sorghum; 21 days after flowering, the content of high starch sorghum leaves MDA was significantly lower than the content of low starch sorghum leaves MDA and there was a significant negative correlation between grain starch content and the leaf MDA content. After high-starch sorghum flowering, the leaves keep a higher enzymes activity and a lower MDA content to slow leaf senescence and lay a foundation in maintaining a high photosynthesis.
5. The research on the influence of regulation of potassium on dry matter accumulation and distribution of sorghum. The results showed that using fertilizer effectively increased the green leaf area of sorghum plants and the accumulation of dry matter and made dry matter increase the accumulation of economic organ, improved economic factors and increased grain yield. The yield will be the highest if we use 90kg of Potassium in per hectare, and then add another 30kg potassium fertilizer to it.
6. The research on the influence of potassium regulation on the accumulation characteristics of sorghum grain starch. The results showed that using K fertilizer significantly increased the total sorghum starch accumulation rate during the filling stage, accumulation rate of amylopectin and accumulation rate of amylose in mid or late filling stage. using K fertilizer increased filling rate of and and amylose in the late filling accumulation rate. The starch content will be the highest if we use 90kg of Potassium in per hectare, and then add another 30kg potassium fertilizer to it.
7. The research on the influence of potassium regulation on physiological index of sorghum leaves. The results showed that using K fertilizer could increase the sorghum leaf weight after flowering, chlorophyll content and net photosynthetic rate. It was one of the reasons which using K fertilizer could increase production. The effect will be the best if we use 90kg of Potassium, and then add another 30kg potassium fertilizer to it.
8. The research on the influence of potassium regulation on protective enzyme activities of sorghum leaves and MDA content. The results showed that using K fertilizer could significantly improve the activity of sorghum leaf SOD, POD and CAT, decrease MDA content in leaves and be conducive to the removal of active oxygen, delay leaf senescence, which is one of the reasons to improve grain yield by using K fertilizer.
引文
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