小麦籽粒多酚氧化酶生化特性及其控制基因的研究
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摘要
多酚氧化酶(Polyphenol Oxidase,PPO)是引起亚洲面条和其它面制食品酶促褐变的主要因素,发生褐变的食品是不受消费者欢迎的,所以如何选育低PPO活性品种是育种家们非常感兴趣的问题,而小麦籽粒PPO活性如何被快速、准确地检测,PPO的生化特性以及控制基因与活性的关系是育种家们急需解决的问题。本文选用24个小麦品种初步探讨了以邻苯二酚代替L-DOPA,按照AACC 22-85的程序检测小麦籽粒PPO活性的可能性,并选用196份小麦品种作了进一步验证;选用3个代表性品种,初步分析了小麦PPO的酶学特性;选用19、54个PPO活性差异较大的品种,分别分析了籽粒发育、萌发过程中同工酶的变化;分析了11种化学试剂在5种浓度下对100个小麦品种籽粒PPO活性的抑制和激活效应;对43条EST和7条mRNA进行了序列组装;采用2A和2D染色体上STS标记分别检测了300个F_4单株和362份小麦品种的PPO等位基因变异,并分析了等位基因2AL(2A染色体上低PPO活性基因)、2AH(2A染色体上高PPO活性基因)、2DL(2D染色体上低PPO活性基因)、2DH(2D染色体上高PPO活性基因)及等位基因组成类型与籽粒PPO活性的关系,初步探讨了内含子变异对PPO活性的影响,并提出了内含子参与PPO基因表达调控的可能性。通过以上试验,得出以下结论:
1.以邻苯二酚代替L-DOPA,按照AACC 22-85的程序检测小麦籽粒PPO活性,是一个成本较低、准确可靠的小麦籽粒PPO活性检测方法。
2.小麦PPO的最适反应温度为65℃,最适pH值为4.0～4.6,Km=0.19mol/L,Vmax=4.04×10~2mol/min。
3.干种子中没有检测到同工酶。同工酶的数目在种子萌发过程中和灌浆期逐渐增加,而在从乳熟期到完熟期间逐渐减少。籽粒在吸水12h后PPO活性迅速升高,24h时达到峰值,然后逐渐趋向平缓。
4.巯基乙醇、EDTA、饱和NaCl和SDS、亚硫酸氢钠、铜试剂、谷胱甘肽和维生素C对小麦PPO产生抑制效应,CuCl_2产生激活效应。抑制剂和激活剂浓度与小麦PPO活性之间呈对数关系,小麦PPO的抑制和激活效应也受添加剂种类和基因型的影响。
5.小麦中至少存在15种PPO基因,这些基因可分为3大类。第Ⅰ类为组织特异表达基因,与组织阶段发育生理功能有关;第Ⅱ类为种子储藏基因,参与完成种子萌发过程中某项生理功能;第Ⅲ类为抗耐性基因,亦可称为防御基因,在外界因素诱导下表达。
6.约50%的籽粒PPO活性变异是由2A、2D染色体上等位基因变异提供的,2A染色体上等位基因变异对籽粒PPO活性的影响是2D染色体等位变异的2-3倍。
7.大多数品种籽粒PPO活性处于中低水平。四种基因型2AL/2DL、2AL/2DH、2AH/2DL、2AH/2DH的籽粒PPO活性逐渐升高一个档次(40-60AU/min·g),且其活性分别处于低偏中、中等、中偏高、高偏中水平,可分别使活性降低30.1%、降低7.8%、升高8.3%、升高30.0%。
8.2AL和2DL基因分别使籽粒PPO活性降低25%和10%。2AH和2DH基因均可使籽粒PPO活性增加17%左右。2AL、2DL、2AH、2DH基因分别使籽粒PPO活性维持在低、中等、中高、中高水平。因此,在低PPO活性小麦育种中,应注重对2AL基因的选择。
9.DQ889708可能是位于2B染色体上控制籽粒高PPO活性的主效基因片段。
Polyphenol Oxidase(PPO) is the major cause of enzymatic discoloration in Asian noodles and other wheat-based end products,Darkening and discoloration affect consumer acceptance of wheat products,especially yellow alkaline and white salted noodles,so it becomes very important to select low PPO activity variety for breeders,however,there are needs for a rapid,accurate assay for determining PPO activity and relationships between controlling genes and activities.24 cultivars are used to explore the possibility of with catechol instead of L-DOPA to detect wheat kernel PPO activity according to AACC 22-85,and it is further validated by 196 cultivars;3 representative cultivars are used to study the characteristics of PPO for enzymology;changes of isoenzymes are analyzed during kernel development and grain germination use 19 and 54 cultivars which PPO activities have more differences;the inhibition and activation of 11 chemical agents under 5 kinds of concentrations on PPO activities are researched by 100 cultivars;43 ESTs and 7 mRNAs are assemblied with DNAMAN software;the allelic variations of PPO are detected use STS markers located on chromosome 2A and 2D in 300 F_4 single lines and 362 cultivars,and the effections of allele including 2AL(low PPO activity on chromosome 2A),2AH(high PPO activity on chromosome 2A),2DL(low PPO activity on chromosome 2D),2DH(high PPO activity on chromosome 2D) and allelic genotypes on PPO activities are studied,so it is for variation of introns either,the introns may influence gene transcription and expression.The results from above experiments showed as follows:
1.According to AACC 22-85,with catechol instead of L-DOPA to detect wheat kernel PPO activity is one low-cost and credible mothed.
2.The optimum temperature and pH of wheat polyphonel oxidase was 65℃and pH4.0～4.6,respectively.Km=0.19mol/L,Vmax=4.04×10~2mol/min
3.There were not isoenzymes detected in the wheat dry grains.The number of isozymes increased gradually in wheat grain germinating period and filling stage,but decreased in the developing kernel during from milk stage to mature stage.The PPO activity got increased rapidly after 12 hours in soaking,reached peak value at 24 hours, then it became smooth.
4.The activity of wheat grain PPO could be inhibiting by mercaptoethanol、EDTA、saturated NaCl、SDS、sodium bisulfite、sodium diethyldithiocarbamate、glutathione and vitamin c,and activating by CuCl_2.It presents logarithmic relation between wheat PPO activity and concentration of inhibitors and activating agent,the type of additives and genotype of cultivars affected inhibition and activation either.
5.There are 15 PPO genes in wheat at least,and these genes fall into 3 distinct sequence groups.GroupⅠare tissue-specific expression genes,which have relation to the function of tissue development;groupⅡare genes of storage in seed,which participate in germination;groupⅢare resistant genes,also are defensive genes,it can be induced by outside factors.
6.About 50%variation of kernel PPO activities are supplied by allelic variation located on chromosome 2A and 2D,the influence of allelic variation on PPO activity which located on chromosome 2A is 2-3 times than that of chromosome 2D.
7.Kernel PPO activities in most of cultivars is low and medium level.The PPO activity of genotype 2AL/2DL,2AL/2DH,2AH/2DL,2AH/2DH get higher one by one for one degree(40-60AU/min·g);and their activity level is low medium,medium,medium high,high medium,respectively;and they make activity decrease 30.1%,decrease 7.8%, increase 8.3%,increase 30.0%,respectively.
8.2AL and 2DL gene make kernel PPO activity decrease 25%and 10%, respectively.2AH and 2DH gene make kernel PPO activity increase around 17%.2AL, 2DL,2AH,2DH keep PPO activity at low,medium,medium high,medium high level, respectively.So,it should be attentioned to select 2AL gene for low PPO activity breeding program.
9.GenBank accession DQ889708 may be the major gene fragment located on chromosome 2B,which controlling high PPO activity in wheat kernels.
1.以邻苯二酚代替L-DOPA,按照AACC 22-85的程序检测小麦籽粒PPO活性,是一个成本较低、准确可靠的小麦籽粒PPO活性检测方法。
2.小麦PPO的最适反应温度为65℃,最适pH值为4.0～4.6,Km=0.19mol/L,Vmax=4.04×10~2mol/min。
3.干种子中没有检测到同工酶。同工酶的数目在种子萌发过程中和灌浆期逐渐增加,而在从乳熟期到完熟期间逐渐减少。籽粒在吸水12h后PPO活性迅速升高,24h时达到峰值,然后逐渐趋向平缓。
4.巯基乙醇、EDTA、饱和NaCl和SDS、亚硫酸氢钠、铜试剂、谷胱甘肽和维生素C对小麦PPO产生抑制效应,CuCl_2产生激活效应。抑制剂和激活剂浓度与小麦PPO活性之间呈对数关系,小麦PPO的抑制和激活效应也受添加剂种类和基因型的影响。
5.小麦中至少存在15种PPO基因,这些基因可分为3大类。第Ⅰ类为组织特异表达基因,与组织阶段发育生理功能有关;第Ⅱ类为种子储藏基因,参与完成种子萌发过程中某项生理功能;第Ⅲ类为抗耐性基因,亦可称为防御基因,在外界因素诱导下表达。
6.约50%的籽粒PPO活性变异是由2A、2D染色体上等位基因变异提供的,2A染色体上等位基因变异对籽粒PPO活性的影响是2D染色体等位变异的2-3倍。
7.大多数品种籽粒PPO活性处于中低水平。四种基因型2AL/2DL、2AL/2DH、2AH/2DL、2AH/2DH的籽粒PPO活性逐渐升高一个档次(40-60AU/min·g),且其活性分别处于低偏中、中等、中偏高、高偏中水平,可分别使活性降低30.1%、降低7.8%、升高8.3%、升高30.0%。
8.2AL和2DL基因分别使籽粒PPO活性降低25%和10%。2AH和2DH基因均可使籽粒PPO活性增加17%左右。2AL、2DL、2AH、2DH基因分别使籽粒PPO活性维持在低、中等、中高、中高水平。因此,在低PPO活性小麦育种中,应注重对2AL基因的选择。
9.DQ889708可能是位于2B染色体上控制籽粒高PPO活性的主效基因片段。
Polyphenol Oxidase(PPO) is the major cause of enzymatic discoloration in Asian noodles and other wheat-based end products,Darkening and discoloration affect consumer acceptance of wheat products,especially yellow alkaline and white salted noodles,so it becomes very important to select low PPO activity variety for breeders,however,there are needs for a rapid,accurate assay for determining PPO activity and relationships between controlling genes and activities.24 cultivars are used to explore the possibility of with catechol instead of L-DOPA to detect wheat kernel PPO activity according to AACC 22-85,and it is further validated by 196 cultivars;3 representative cultivars are used to study the characteristics of PPO for enzymology;changes of isoenzymes are analyzed during kernel development and grain germination use 19 and 54 cultivars which PPO activities have more differences;the inhibition and activation of 11 chemical agents under 5 kinds of concentrations on PPO activities are researched by 100 cultivars;43 ESTs and 7 mRNAs are assemblied with DNAMAN software;the allelic variations of PPO are detected use STS markers located on chromosome 2A and 2D in 300 F_4 single lines and 362 cultivars,and the effections of allele including 2AL(low PPO activity on chromosome 2A),2AH(high PPO activity on chromosome 2A),2DL(low PPO activity on chromosome 2D),2DH(high PPO activity on chromosome 2D) and allelic genotypes on PPO activities are studied,so it is for variation of introns either,the introns may influence gene transcription and expression.The results from above experiments showed as follows:
1.According to AACC 22-85,with catechol instead of L-DOPA to detect wheat kernel PPO activity is one low-cost and credible mothed.
2.The optimum temperature and pH of wheat polyphonel oxidase was 65℃and pH4.0～4.6,respectively.Km=0.19mol/L,Vmax=4.04×10~2mol/min
3.There were not isoenzymes detected in the wheat dry grains.The number of isozymes increased gradually in wheat grain germinating period and filling stage,but decreased in the developing kernel during from milk stage to mature stage.The PPO activity got increased rapidly after 12 hours in soaking,reached peak value at 24 hours, then it became smooth.
4.The activity of wheat grain PPO could be inhibiting by mercaptoethanol、EDTA、saturated NaCl、SDS、sodium bisulfite、sodium diethyldithiocarbamate、glutathione and vitamin c,and activating by CuCl_2.It presents logarithmic relation between wheat PPO activity and concentration of inhibitors and activating agent,the type of additives and genotype of cultivars affected inhibition and activation either.
5.There are 15 PPO genes in wheat at least,and these genes fall into 3 distinct sequence groups.GroupⅠare tissue-specific expression genes,which have relation to the function of tissue development;groupⅡare genes of storage in seed,which participate in germination;groupⅢare resistant genes,also are defensive genes,it can be induced by outside factors.
6.About 50%variation of kernel PPO activities are supplied by allelic variation located on chromosome 2A and 2D,the influence of allelic variation on PPO activity which located on chromosome 2A is 2-3 times than that of chromosome 2D.
7.Kernel PPO activities in most of cultivars is low and medium level.The PPO activity of genotype 2AL/2DL,2AL/2DH,2AH/2DL,2AH/2DH get higher one by one for one degree(40-60AU/min·g);and their activity level is low medium,medium,medium high,high medium,respectively;and they make activity decrease 30.1%,decrease 7.8%, increase 8.3%,increase 30.0%,respectively.
8.2AL and 2DL gene make kernel PPO activity decrease 25%and 10%, respectively.2AH and 2DH gene make kernel PPO activity increase around 17%.2AL, 2DL,2AH,2DH keep PPO activity at low,medium,medium high,medium high level, respectively.So,it should be attentioned to select 2AL gene for low PPO activity breeding program.
9.GenBank accession DQ889708 may be the major gene fragment located on chromosome 2B,which controlling high PPO activity in wheat kernels.
引文
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