苜蓿根蘖性状发生及其调节机制研究
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摘要
苜蓿的根蘖性状是指在水平侧根的不定位置有根蘖芽发生从而形成远离母株的独立枝条,使植株水平发展的特性。本文系统研究了苜蓿根蘖性状发生、表现的规律及其生理和遗传机制,在对根蘖型苜蓿在不同生态条件下生长发育的展开多年定点调查基础上,系统地分析了苜蓿根蘖性状与土壤类型、生长年限等因子的关系,进一步明确了根蘖型苜蓿的干旱适应性。在对根蘖性状的形态发生和表现过程做出形态划分的基础上,从内源激素和等位酶角度对根蘖性状发生和表达的生理、遗传机制进行了系统研究,主要研究结论如下:
1.对内蒙古镶黄旗三年生苜蓿材料根系类型进行调查,筛选出Travois、Ladak、Teton、新疆大叶苜蓿分别作为根蘖型、侧根型、根茎型及轴根型苜蓿的典型试验材料;比较根蘖类型与其他根系类型苜蓿在遗传特性、产量及营养特性方面的差异,表明根蘖型苜蓿是进行抗逆、优质、高产聚合育种的优异遗传资源;通过层次分析法(AHP)确立不同根系类型苜蓿品种苗期的抗旱性排序依次为:Travois(根蘖型)>新疆大叶(轴根型)>Ladak(侧根型)>Teton(根茎型),表明根蘖型苜蓿有较强的干旱适应性。
2.苜蓿整个生长周期内对根蘖苜蓿根系的调查表明,根蘖性状发生及表现可划分为五个时期:根蘖节时期,根蘖芽时期,根蘖枝条时期,根蘖株丛时期和二级根蘖时期。前两个时期为根蘖发生时期,后三个时期为根蘖表现时期。对根蘖性状发生的解剖学研究表明根蘖芽起源于根部木栓形成层分生组织;秋季刈割后根蘖植株根部可溶性糖含量减少,根蘖芽萌发前后,可溶性蛋白含量呈现显著变化,表明营养物质的动态变化影响根蘖性状的发生及表现。
3.对内蒙和北京两地不同生境下同一份根蘖品种根蘖性状发生和表现做多年定点调查,结果表明两地根蘖发生率无明显差异,但内蒙地区平均根蘖幅度较北京地区更高;边际效应影响根蘖表现,但不影响根蘖发生;播种后第1年便有根蘖性状发生,第3年是根蘖发生的高峰期,第5年后趋于稳定。
4.根蘖植株中根颈直径、根颈入土深度及根颈分枝数变异较大,与根蘖地上部分表现性状无显著相关性,且在根蘖性状发生程度较高的植株中,根颈直径有减小的趋势;根蘖性状的发生与10～20cm土层中三级根的数量有关,其性状的表现则与0～10cm土层中根系总量相关,表明根蘖苜蓿不同土层中不同根系的功能有所差异。土壤水分含量与一级根数量呈正相关,与根蘖性状的表现呈负相关,土壤容重与10-20cm土层中三级根数量呈显著负相关。
5.内源激素的测定和与根蘖性状表现的相关分析表明:根蘖苜蓿根部膨大无芽处的GA和ABA含量显著高于膨大有芽处(P≤0.01),表明GA和ABA抑制了根蘖芽的发生;GA、IAA、ABA含量与根蘖性状表现有关,具体体现在GA含量高的苜蓿根蘖丛数多,根蘖幅度小,IAA含量高的苜蓿根蘖丛数少,根蘖幅度大,而ABA含量则与根蘖幅度和根蘖株丛数均呈负相关。ABA与0-20cm土层中的一级根数量呈显著正相关,且ABA在镶黄旗地区苜蓿根系组织中的含量明显高于北京地区相应材料,干旱诱导了根蘖苜蓿根部ABA的产生,而根蘖苜蓿由于其根蘖特性,根部ABA的积累量小于其它两种材料,表明其受到的干旱胁迫较小。
6.对根蘖、非根蘖及黄花苜蓿种群等位基因频率的分析表明,根蘖型与非根蘖型苜蓿群体等位基因频率的差异主要是在LAP-298.0和LAP-296.0上。其中LAP-298.0在根蘖苜蓿群体中的频率为6%,而在非根蘖型苜蓿中则为0,在黄花苜蓿种群中LAP-298.0的频率为10%,该基因可看着与根蘖性状相关的标记基因。基因频率的研究首次验证了黄花苜蓿的基因渗入对根蘖性状的发生起到重要作用。
Creeping-rooted character in alfalfa (Medicago sativa L.)was the character which can originate a shoot from the root system, and provide the plant with a rhizome-like ability to regenerate in stands. This habit of spreading by means of root proliferation provides alfalfa with a form of vegetative spread and multiplication.Through 6 years(2003-2007) experiments of the alfalfa germplasm evaluation and new variety breeding project,the mechanism of creeping rooted character’s ontogene and performance was systematically studied.The results are as following:
1. Root system type of three years old alfalfa materials was investigated in Xianghuang banner of inner Mongolia,Travois ,Ladak, Teton and Xinjiang big leaf were selected for representing the root system type of Creeping-rooted (CR), Branch rooted( BR ), Rhizomatous rooted( RR), and Tap rooted (TR) type respectively. Based on it, the genetic character, yield and nutrition character were analyzed between the 4 rooted types.,Which showed that CR is the important genetic resource in breeding for the drought-resistant, high quality and high quantity new varieties . AHP analysis method were used to analyzed the drought resistant of different root system types and the drought resistance ability at the Seedling Stage. Results showed that CR alfalfa has the gene from yellow followered alfalfa,and performed excellent in the steppe district. The synthetic evaluation values of the different experimental root type material showed that the drought dolerent ability in seeding stage is :CR>TR>BR>RR.
2. Rooted system of thecreeping rooted plant were investigated in the whole growth period.Thus the ontogenesis and performance of the creeping-rooted character including 5 stages: creeping rootedt swellon stage,creeping rooted bud stage,creeping rooted shoots stage,ceeping rooted clusters stage and further creeping rooted stage。The first 2 stages representated the ontogenesis stage,and the rest 3 stages as the performance stage.Anatomical research on CR character ontogenesis showed that the buds of root shoots grow originally from meristematic tissue of phellem cambium in root and after developing aboveground internodes, parts of its underground tissues put up root character, and then developing the offspring plant which will having the independent growth and developing function. Research on the dynamic change of root assimilation product of CR showed that the root soluble sugar content was probably related with the overgroud growth. .Starch content of CR alfalfa was higher than NCR and this could conclude that starch accumulation in root was the material basic to CR thrait happening. After bud germinating, soluble protein content changed evidently, this might be related with the synthesis of specific protein.
3. Based on the investigation and statistic of CR percentage rate and its expressional trait of different habitat in inner Mongolia and Beijing area, it was showed that percentage of creeping rooted plants of 3 years old materials was about 80%,and there was no evident difference in this two areas in the same plant age.The influence of the marginal effect on the performance of the creeping rooted character but has no effect on the ontogenesis of creeping rooted character.
4.Research on the root system structure of CR alfalfa indicated that the ontogenesis of CR was related to the third class root number in the 10~20cm soil layer, and the root number of each class in 0 ~10 cm was related to the expression of CR, which showed that the function of different root system in different soil layers was different.The function of different class of root in different soil layer. the studing also showed that with the increasing of CR trait, the root neck diameter had a decreased trend.Soil water content was positive correlated with the first class root number in the 0 ~ 20cm soil layer of CR alfalfa, and soil volume had evidently negative correlation with the third class root number in 10 ~ 20 cm soil layer.
5. Endogenous hormone including GA3,IAA,ABAand (Z+ZR) content in different part of the horizontal root in CR,NCR and the TR plant were measured by the method of HPLC. Result showed that endogenous hormone had different effect on the overground and underground character of the CR alfalfa: IAA and (Z+ZR) had the positive correlation with the plant height, CR cluster number and CR stem number, while GA had the negative correlation with this three characters; ABA content had the prominence positive correlation with the amount of the root, while had the prominence negative correlation with the creeping rooted extent and CR stem number. ABA of 3 types in Xianghuangqi spot were higher than that of the Beijing spot, and the content ranked as TR>NCR>CR,which was reverse to the Beijing spot. It showed that the CR alfalfa endured less drought stress in the aridity district than the TR and NCR alfalfa due to this character.
6. Through the starch gel electrophoresis of the monomeric allozymes LAP-2 alleles are found in the CR,NCR and medicago falcate populations .the results showed that the LAP-2 was a polymorphic loci。The allele frequencies of LAP-2100in the CR and NCR populationg were 91.0% and 99.9%respectively,nearly possessed by all the experimental materials。While the allele frequencies of LAP-298.0 in the CR and NCR population were 6% and 0% respectively.In the population of medicago falcate,the llele frequencies of LAP-295.5 is 10%,so this allele could be predicted as the medicago falcate gene,which originaled from the medicago falcate,and relatived with the CR character.
1.对内蒙古镶黄旗三年生苜蓿材料根系类型进行调查,筛选出Travois、Ladak、Teton、新疆大叶苜蓿分别作为根蘖型、侧根型、根茎型及轴根型苜蓿的典型试验材料;比较根蘖类型与其他根系类型苜蓿在遗传特性、产量及营养特性方面的差异,表明根蘖型苜蓿是进行抗逆、优质、高产聚合育种的优异遗传资源;通过层次分析法(AHP)确立不同根系类型苜蓿品种苗期的抗旱性排序依次为:Travois(根蘖型)>新疆大叶(轴根型)>Ladak(侧根型)>Teton(根茎型),表明根蘖型苜蓿有较强的干旱适应性。
2.苜蓿整个生长周期内对根蘖苜蓿根系的调查表明,根蘖性状发生及表现可划分为五个时期:根蘖节时期,根蘖芽时期,根蘖枝条时期,根蘖株丛时期和二级根蘖时期。前两个时期为根蘖发生时期,后三个时期为根蘖表现时期。对根蘖性状发生的解剖学研究表明根蘖芽起源于根部木栓形成层分生组织;秋季刈割后根蘖植株根部可溶性糖含量减少,根蘖芽萌发前后,可溶性蛋白含量呈现显著变化,表明营养物质的动态变化影响根蘖性状的发生及表现。
3.对内蒙和北京两地不同生境下同一份根蘖品种根蘖性状发生和表现做多年定点调查,结果表明两地根蘖发生率无明显差异,但内蒙地区平均根蘖幅度较北京地区更高;边际效应影响根蘖表现,但不影响根蘖发生;播种后第1年便有根蘖性状发生,第3年是根蘖发生的高峰期,第5年后趋于稳定。
4.根蘖植株中根颈直径、根颈入土深度及根颈分枝数变异较大,与根蘖地上部分表现性状无显著相关性,且在根蘖性状发生程度较高的植株中,根颈直径有减小的趋势;根蘖性状的发生与10～20cm土层中三级根的数量有关,其性状的表现则与0～10cm土层中根系总量相关,表明根蘖苜蓿不同土层中不同根系的功能有所差异。土壤水分含量与一级根数量呈正相关,与根蘖性状的表现呈负相关,土壤容重与10-20cm土层中三级根数量呈显著负相关。
5.内源激素的测定和与根蘖性状表现的相关分析表明:根蘖苜蓿根部膨大无芽处的GA和ABA含量显著高于膨大有芽处(P≤0.01),表明GA和ABA抑制了根蘖芽的发生;GA、IAA、ABA含量与根蘖性状表现有关,具体体现在GA含量高的苜蓿根蘖丛数多,根蘖幅度小,IAA含量高的苜蓿根蘖丛数少,根蘖幅度大,而ABA含量则与根蘖幅度和根蘖株丛数均呈负相关。ABA与0-20cm土层中的一级根数量呈显著正相关,且ABA在镶黄旗地区苜蓿根系组织中的含量明显高于北京地区相应材料,干旱诱导了根蘖苜蓿根部ABA的产生,而根蘖苜蓿由于其根蘖特性,根部ABA的积累量小于其它两种材料,表明其受到的干旱胁迫较小。
6.对根蘖、非根蘖及黄花苜蓿种群等位基因频率的分析表明,根蘖型与非根蘖型苜蓿群体等位基因频率的差异主要是在LAP-298.0和LAP-296.0上。其中LAP-298.0在根蘖苜蓿群体中的频率为6%,而在非根蘖型苜蓿中则为0,在黄花苜蓿种群中LAP-298.0的频率为10%,该基因可看着与根蘖性状相关的标记基因。基因频率的研究首次验证了黄花苜蓿的基因渗入对根蘖性状的发生起到重要作用。
Creeping-rooted character in alfalfa (Medicago sativa L.)was the character which can originate a shoot from the root system, and provide the plant with a rhizome-like ability to regenerate in stands. This habit of spreading by means of root proliferation provides alfalfa with a form of vegetative spread and multiplication.Through 6 years(2003-2007) experiments of the alfalfa germplasm evaluation and new variety breeding project,the mechanism of creeping rooted character’s ontogene and performance was systematically studied.The results are as following:
1. Root system type of three years old alfalfa materials was investigated in Xianghuang banner of inner Mongolia,Travois ,Ladak, Teton and Xinjiang big leaf were selected for representing the root system type of Creeping-rooted (CR), Branch rooted( BR ), Rhizomatous rooted( RR), and Tap rooted (TR) type respectively. Based on it, the genetic character, yield and nutrition character were analyzed between the 4 rooted types.,Which showed that CR is the important genetic resource in breeding for the drought-resistant, high quality and high quantity new varieties . AHP analysis method were used to analyzed the drought resistant of different root system types and the drought resistance ability at the Seedling Stage. Results showed that CR alfalfa has the gene from yellow followered alfalfa,and performed excellent in the steppe district. The synthetic evaluation values of the different experimental root type material showed that the drought dolerent ability in seeding stage is :CR>TR>BR>RR.
2. Rooted system of thecreeping rooted plant were investigated in the whole growth period.Thus the ontogenesis and performance of the creeping-rooted character including 5 stages: creeping rootedt swellon stage,creeping rooted bud stage,creeping rooted shoots stage,ceeping rooted clusters stage and further creeping rooted stage。The first 2 stages representated the ontogenesis stage,and the rest 3 stages as the performance stage.Anatomical research on CR character ontogenesis showed that the buds of root shoots grow originally from meristematic tissue of phellem cambium in root and after developing aboveground internodes, parts of its underground tissues put up root character, and then developing the offspring plant which will having the independent growth and developing function. Research on the dynamic change of root assimilation product of CR showed that the root soluble sugar content was probably related with the overgroud growth. .Starch content of CR alfalfa was higher than NCR and this could conclude that starch accumulation in root was the material basic to CR thrait happening. After bud germinating, soluble protein content changed evidently, this might be related with the synthesis of specific protein.
3. Based on the investigation and statistic of CR percentage rate and its expressional trait of different habitat in inner Mongolia and Beijing area, it was showed that percentage of creeping rooted plants of 3 years old materials was about 80%,and there was no evident difference in this two areas in the same plant age.The influence of the marginal effect on the performance of the creeping rooted character but has no effect on the ontogenesis of creeping rooted character.
4.Research on the root system structure of CR alfalfa indicated that the ontogenesis of CR was related to the third class root number in the 10~20cm soil layer, and the root number of each class in 0 ~10 cm was related to the expression of CR, which showed that the function of different root system in different soil layers was different.The function of different class of root in different soil layer. the studing also showed that with the increasing of CR trait, the root neck diameter had a decreased trend.Soil water content was positive correlated with the first class root number in the 0 ~ 20cm soil layer of CR alfalfa, and soil volume had evidently negative correlation with the third class root number in 10 ~ 20 cm soil layer.
5. Endogenous hormone including GA3,IAA,ABAand (Z+ZR) content in different part of the horizontal root in CR,NCR and the TR plant were measured by the method of HPLC. Result showed that endogenous hormone had different effect on the overground and underground character of the CR alfalfa: IAA and (Z+ZR) had the positive correlation with the plant height, CR cluster number and CR stem number, while GA had the negative correlation with this three characters; ABA content had the prominence positive correlation with the amount of the root, while had the prominence negative correlation with the creeping rooted extent and CR stem number. ABA of 3 types in Xianghuangqi spot were higher than that of the Beijing spot, and the content ranked as TR>NCR>CR,which was reverse to the Beijing spot. It showed that the CR alfalfa endured less drought stress in the aridity district than the TR and NCR alfalfa due to this character.
6. Through the starch gel electrophoresis of the monomeric allozymes LAP-2 alleles are found in the CR,NCR and medicago falcate populations .the results showed that the LAP-2 was a polymorphic loci。The allele frequencies of LAP-2100in the CR and NCR populationg were 91.0% and 99.9%respectively,nearly possessed by all the experimental materials。While the allele frequencies of LAP-298.0 in the CR and NCR population were 6% and 0% respectively.In the population of medicago falcate,the llele frequencies of LAP-295.5 is 10%,so this allele could be predicted as the medicago falcate gene,which originaled from the medicago falcate,and relatived with the CR character.
引文
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