风味甜瓜果实酸味形成机理及糖酸遗传和积累模拟研究
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摘要
本研究以风味甜瓜(新疆农科院哈密瓜研究中心选育出的酸味甜瓜)为材料,基于对风味甜瓜成熟时酸味的成分、酸味形成的生理机制、糖酸的遗传特点及与温光的关系等开展了以下四方面的研究工作:
1.利用首张欧美甜瓜果实基因芯片Melon cDNA array ver1.0检测新疆厚皮甜瓜成熟果实基因表达分析,并检测了60Co-γ辐照酸味抗病变异株成熟果实基因表达水平,从酸代谢的分子水平探讨风味甜瓜酸味形成机理;
2.利用盖钧镒具有主基因效应的数量性状理论,以60Co-γ射线诱变酸味突变自交系和‘黄皮脆’形成的P1、P2、F1、F2为试验材料,通过4个世代联合分析法,对果实糖酸含量性状的遗传进行了全面系统的研究;
3.利用灰色关联度法对‘风味3号’果实坐果期、果实快速生长期及成熟期糖酸增量与温光积累增量和发育成熟期糖酸积累及温光因子的关系进行了研究,探讨在甜瓜果实发育过程中影响果实糖和酸积累的主要影响因子;
4.利用大棚春、秋茬和温室春、秋茬栽培,对有效积温累积、日温差累积、光合有效辐射累积与甜瓜果实蔗糖、柠檬酸积累及糖酸比关系进行了模拟研究。
获得的主要结论如下:
1.风味甜瓜中的酸味主要是由于柠檬酸的积累所致;通过基因芯片转录谱检测,发现3类基因的上调表达可能和柠檬酸积累有关,分别是:柠檬酸合成酶、乙烯、Ca2+-CaM信号系统。
2.酸味突变自交系和黄皮脆杂交组合的果实总糖含量、葡萄糖含量、果糖含量、蔗糖含量、总酸含量、柠檬酸含量、酸度和糖酸比均受主基因+多基因控制,但主基因和多基因在性状遗传控制上所起的作用有所不同:总糖、葡萄糖、果糖、柠檬酸、糖酸比受两对主基因+多基因混合遗传模型控制;蔗糖、总酸、酸度受一对主基因+多基因混合遗传模型控制。糖含量、酸度和糖酸比性状受主基因的影响较酸含量大,酸含量性状受多基因影响较糖含量、酸度和糖酸比大。主基因+多基因效应决定了糖含量、酸含量和糖酸比性状变异的95.74% - 99.45%,还有很小的变异是由环境因素决定的。
葡萄糖与果糖之间、蔗糖与总糖之间、柠檬酸与总酸之间有高度的表型相关性;总糖与葡萄糖、总糖与果糖、总糖与可溶性固形物含量、糖酸比与果糖、蔗糖与柠檬酸、蔗糖与总酸、总酸与可溶性固形物含量之间、总酸与苹果酸之间、总糖与柠檬酸、总糖与总酸之间有较高的相关性。
葡萄糖含量性状和酸度(pH)性状之间、蔗糖含量性状和酸度(pH)性状之间、总酸含量性状和酸度(pH)性状之间、柠檬酸含量性状和酸度(pH)性状之间均受同一对主基因控制,表现为一因多效现象。
3.无论在坐果期、果实快速生长期还是成熟期光合有效辐射累积对‘风味3号’甜瓜果实的糖酸增量影响最大。在果实整个发育成熟期,对果实总糖影响最大的是日温差累积;对果实葡萄糖影响最大的是有效积温累积;对果糖影响最大的是光合有效辐射累积;对蔗糖影响最大的是光合有效辐射累积;对总酸含量影响最大的是日温差累积;对苹果酸和柠檬酸影响最大的是日温差累积。
4.通过回归分析,建立了甜瓜果实蔗糖、柠檬酸及糖酸比受有效积温与日温差累积的驱动模型。不同茬口果实蔗糖、柠檬酸及糖酸比,随有效积温与日温差累积增加都呈logistic曲线函数变化,但不同茬口函数常数项不同,常数项均由日温差累积累积驱动,它们之间为线性关系。
The feasibility of using Melon cDNA array ver1.0 to detect the gene expression of mature fruit of Cucumis melo var.ameri were studied. The results showed that the melon cDNA array ver1.0 could detect the expression of 2008 genes in the variety which was sourness taste in fruit and diseases resistance of Cucumis melo var.ameri. The detected genes accounted for 65.4% of the total 3068 probe groups. 251 up-regulated and 224 down-regulated were detected in the variety, accounting for 12.5% and 11.16% of total detected genes respectively. This finding included additional 62 up- and 33 down-regulated unknown (“no hit”) genes. Among the identified ones, 47 genes expressed significant (FDR<0.01 and fold change≧2) differential expressed genes. The identified genes include, citric synthase mitochondrial precursor and those associated with defense response, cell wall modification, plant hormones and signal transduction, structural constituent of ribosome,transcriptional factor/regulatory factor. The changes in mRNA abundance detected by microarray analysis was validated by using RT-PCR, results showed that melon cDNA array ver1.0 could efficiently be used for gene expression analysis of Cucumis melo var.ameri. Component and content of organic acid and citrate synthase (CS) activity during pre- and post-irradiation fruit development were measured by using high performance liquid chromatography and uv-spectrophotometer. The comparative transcription profiling found that several genes that could be related to traits of sourness, disease resistance in the fruit of post 60Co-γirradiation. Our results also suggest that 60Co-γirradiation mutation changes the mRNA levels for CS mitochondria, followed by changes in the protein and/or activity levels that relates with the changes in the citrate levels. It may be a primary trigger for citrate accumulation in the fruit of sour-flavor matant.
Four generations of inbred line with sour-flavor matant and Huangpicui P1, P2, F1 and F2 were used to study the inheritance of melon sugar content、sour content and sugar acid radio traits by joint analysis method of multiple generations. The results showed that the total sugar content was controlled by two pairs of equal additive major genes plus additive-dominant polygene (E-4 model). The major gene heritability was 88.8%; the polygene heritability was 6.94%. The glucose content fitted two pairs of additive-dominance-epitasis major genes plus additive-dominant-epitasis polygene (E-0 model). The flucrose content fitted two pairs of additive-dominance-epitasis major genes plus additive-dominant polygene (E-1 model). The inheritance of sucrose content fitted one pair of additive–dominance major gene plus additive–dominance-epitasis polygene (D-0 model). The major gene heritability was 74.76%; the polygene heritability was 25.04%. The inheritance of total sour content fitted one pair of additive–dominance major gene plus additive–dominance-epitasis polygene (D-0 model). The major gene heritability was 26.68%; the polygene heritability was 72.77%. The citrate content fitted two pairs of additive-dominance major genes plus additive-dominant polygene (E-2 model). The major gene heritability was 90.13%; the polygene heritability was 8.78%. Sugar acid radio trait fitted two pairs of additive-dominance-epitasis major genes plus additive-dominant polygene (E-1 model). The major gene heritability was 82.86%; the polygene heritability was 16.02%. The inheritance of acidity fitted one pair of additive–dominance major gene plus additive–dominance-epitasis polygene (D-0 model). The major gene heritability was 84.21%; the polygene heritability was 14.13%. There were highly correlated between glucose content and fructose, sucrose content and total sugar, citrate content and total sour. It had high relativity between total sugar and glucose content, total sugar and fructose, sucrose content and citrate, citrate content and total sugar, total sugar and total sour, total sour and malate, sucrose content and total sour, sugar acid radio and fructose. The glucose content and acidity traits refer to only one pleiotropic major gene. The additive effect and dominance effect of the major gene were estimated as -2.53037 mg g-1 FW and 3.1681 mg g-1 FW on glucose, 0.6659 and -0.6325 on acidity, respectively. The major gene showed overdominance for glucose content and close to complete dominance for acidity. The sucrose content and acidity traits refered to only one pleiotropic major gene. The additive effect and dominance effect of the major gene were estimated as- 13.5563 mg g-1 FW and–1.5948 mg g-1 FW on sucrose, and–0.6122 and -0.7451 on acidity, respectively. The major gene showed incomplete dominance for sucrose content and complete dominance for acidity. The citrate content and acidity traits refered to only one pleiotropic major gene. The additive effect and dominance effect of the major gene were estimated as -1.0855 mg g-1 FW and-2.8537 mg g-1 FW on citrate, and 0.6476 and–0.6708 on acidity, respectively. The major gene shows overdominance for citrate content and close to complete dominance for acidity. The total sour content and acidity traits refered to only one pleiotropic major gene. The additive effect and dominance effect of the major gene are estimated as1.2735 mg g-1 FW and -2.9746 mg g-1 FW on total sour content, and -0.647and–0.6697 on acidity, respectively, The major gene showed overdominance for glucose content and close to complete dominance for acidity.
The influence of temperature and solar on sugar and acid content of fruit FN3 melon cultured in spring film house, autumn film house and autumu green house, were analyzed by using grey relational grade analysis method. The results showed that the influence rankings on sugar content were: accumulative solar radiation > accumulative daily temperature difference > effective accumulative temperature. The influence of temperature and solar on acid content of FN3 fruit were in order of accumulative daily temperature difference > accumulative solar radiation > effective accumulative temperature.
With different crops for rotation, the effects of air temperature and accumulated daily difference in temperature on the sucrose, citrate accumulation and sugar acid radio of greenhouse melon fruit were studied, with related simulation model established. The results showed that the sucrose, citrate accumulation and sugar acid radio model were logistic function, all of which were driven by effective accumulative temperature. The constant term in the functions was driven by accumulative daily difference in temperature, and the correlation was a linear function. Model test showed that the models were able to objectively simulate and predict the changes of fruit sucrose, citrate accumulation and sugar acid radio with fruit development, and possessed practical value for the accurate estimates to melon fruit inherent quality and optimization to determine harvest of melon fruit.
1.利用首张欧美甜瓜果实基因芯片Melon cDNA array ver1.0检测新疆厚皮甜瓜成熟果实基因表达分析,并检测了60Co-γ辐照酸味抗病变异株成熟果实基因表达水平,从酸代谢的分子水平探讨风味甜瓜酸味形成机理;
2.利用盖钧镒具有主基因效应的数量性状理论,以60Co-γ射线诱变酸味突变自交系和‘黄皮脆’形成的P1、P2、F1、F2为试验材料,通过4个世代联合分析法,对果实糖酸含量性状的遗传进行了全面系统的研究;
3.利用灰色关联度法对‘风味3号’果实坐果期、果实快速生长期及成熟期糖酸增量与温光积累增量和发育成熟期糖酸积累及温光因子的关系进行了研究,探讨在甜瓜果实发育过程中影响果实糖和酸积累的主要影响因子;
4.利用大棚春、秋茬和温室春、秋茬栽培,对有效积温累积、日温差累积、光合有效辐射累积与甜瓜果实蔗糖、柠檬酸积累及糖酸比关系进行了模拟研究。
获得的主要结论如下:
1.风味甜瓜中的酸味主要是由于柠檬酸的积累所致;通过基因芯片转录谱检测,发现3类基因的上调表达可能和柠檬酸积累有关,分别是:柠檬酸合成酶、乙烯、Ca2+-CaM信号系统。
2.酸味突变自交系和黄皮脆杂交组合的果实总糖含量、葡萄糖含量、果糖含量、蔗糖含量、总酸含量、柠檬酸含量、酸度和糖酸比均受主基因+多基因控制,但主基因和多基因在性状遗传控制上所起的作用有所不同:总糖、葡萄糖、果糖、柠檬酸、糖酸比受两对主基因+多基因混合遗传模型控制;蔗糖、总酸、酸度受一对主基因+多基因混合遗传模型控制。糖含量、酸度和糖酸比性状受主基因的影响较酸含量大,酸含量性状受多基因影响较糖含量、酸度和糖酸比大。主基因+多基因效应决定了糖含量、酸含量和糖酸比性状变异的95.74% - 99.45%,还有很小的变异是由环境因素决定的。
葡萄糖与果糖之间、蔗糖与总糖之间、柠檬酸与总酸之间有高度的表型相关性;总糖与葡萄糖、总糖与果糖、总糖与可溶性固形物含量、糖酸比与果糖、蔗糖与柠檬酸、蔗糖与总酸、总酸与可溶性固形物含量之间、总酸与苹果酸之间、总糖与柠檬酸、总糖与总酸之间有较高的相关性。
葡萄糖含量性状和酸度(pH)性状之间、蔗糖含量性状和酸度(pH)性状之间、总酸含量性状和酸度(pH)性状之间、柠檬酸含量性状和酸度(pH)性状之间均受同一对主基因控制,表现为一因多效现象。
3.无论在坐果期、果实快速生长期还是成熟期光合有效辐射累积对‘风味3号’甜瓜果实的糖酸增量影响最大。在果实整个发育成熟期,对果实总糖影响最大的是日温差累积;对果实葡萄糖影响最大的是有效积温累积;对果糖影响最大的是光合有效辐射累积;对蔗糖影响最大的是光合有效辐射累积;对总酸含量影响最大的是日温差累积;对苹果酸和柠檬酸影响最大的是日温差累积。
4.通过回归分析,建立了甜瓜果实蔗糖、柠檬酸及糖酸比受有效积温与日温差累积的驱动模型。不同茬口果实蔗糖、柠檬酸及糖酸比,随有效积温与日温差累积增加都呈logistic曲线函数变化,但不同茬口函数常数项不同,常数项均由日温差累积累积驱动,它们之间为线性关系。
The feasibility of using Melon cDNA array ver1.0 to detect the gene expression of mature fruit of Cucumis melo var.ameri were studied. The results showed that the melon cDNA array ver1.0 could detect the expression of 2008 genes in the variety which was sourness taste in fruit and diseases resistance of Cucumis melo var.ameri. The detected genes accounted for 65.4% of the total 3068 probe groups. 251 up-regulated and 224 down-regulated were detected in the variety, accounting for 12.5% and 11.16% of total detected genes respectively. This finding included additional 62 up- and 33 down-regulated unknown (“no hit”) genes. Among the identified ones, 47 genes expressed significant (FDR<0.01 and fold change≧2) differential expressed genes. The identified genes include, citric synthase mitochondrial precursor and those associated with defense response, cell wall modification, plant hormones and signal transduction, structural constituent of ribosome,transcriptional factor/regulatory factor. The changes in mRNA abundance detected by microarray analysis was validated by using RT-PCR, results showed that melon cDNA array ver1.0 could efficiently be used for gene expression analysis of Cucumis melo var.ameri. Component and content of organic acid and citrate synthase (CS) activity during pre- and post-irradiation fruit development were measured by using high performance liquid chromatography and uv-spectrophotometer. The comparative transcription profiling found that several genes that could be related to traits of sourness, disease resistance in the fruit of post 60Co-γirradiation. Our results also suggest that 60Co-γirradiation mutation changes the mRNA levels for CS mitochondria, followed by changes in the protein and/or activity levels that relates with the changes in the citrate levels. It may be a primary trigger for citrate accumulation in the fruit of sour-flavor matant.
Four generations of inbred line with sour-flavor matant and Huangpicui P1, P2, F1 and F2 were used to study the inheritance of melon sugar content、sour content and sugar acid radio traits by joint analysis method of multiple generations. The results showed that the total sugar content was controlled by two pairs of equal additive major genes plus additive-dominant polygene (E-4 model). The major gene heritability was 88.8%; the polygene heritability was 6.94%. The glucose content fitted two pairs of additive-dominance-epitasis major genes plus additive-dominant-epitasis polygene (E-0 model). The flucrose content fitted two pairs of additive-dominance-epitasis major genes plus additive-dominant polygene (E-1 model). The inheritance of sucrose content fitted one pair of additive–dominance major gene plus additive–dominance-epitasis polygene (D-0 model). The major gene heritability was 74.76%; the polygene heritability was 25.04%. The inheritance of total sour content fitted one pair of additive–dominance major gene plus additive–dominance-epitasis polygene (D-0 model). The major gene heritability was 26.68%; the polygene heritability was 72.77%. The citrate content fitted two pairs of additive-dominance major genes plus additive-dominant polygene (E-2 model). The major gene heritability was 90.13%; the polygene heritability was 8.78%. Sugar acid radio trait fitted two pairs of additive-dominance-epitasis major genes plus additive-dominant polygene (E-1 model). The major gene heritability was 82.86%; the polygene heritability was 16.02%. The inheritance of acidity fitted one pair of additive–dominance major gene plus additive–dominance-epitasis polygene (D-0 model). The major gene heritability was 84.21%; the polygene heritability was 14.13%. There were highly correlated between glucose content and fructose, sucrose content and total sugar, citrate content and total sour. It had high relativity between total sugar and glucose content, total sugar and fructose, sucrose content and citrate, citrate content and total sugar, total sugar and total sour, total sour and malate, sucrose content and total sour, sugar acid radio and fructose. The glucose content and acidity traits refer to only one pleiotropic major gene. The additive effect and dominance effect of the major gene were estimated as -2.53037 mg g-1 FW and 3.1681 mg g-1 FW on glucose, 0.6659 and -0.6325 on acidity, respectively. The major gene showed overdominance for glucose content and close to complete dominance for acidity. The sucrose content and acidity traits refered to only one pleiotropic major gene. The additive effect and dominance effect of the major gene were estimated as- 13.5563 mg g-1 FW and–1.5948 mg g-1 FW on sucrose, and–0.6122 and -0.7451 on acidity, respectively. The major gene showed incomplete dominance for sucrose content and complete dominance for acidity. The citrate content and acidity traits refered to only one pleiotropic major gene. The additive effect and dominance effect of the major gene were estimated as -1.0855 mg g-1 FW and-2.8537 mg g-1 FW on citrate, and 0.6476 and–0.6708 on acidity, respectively. The major gene shows overdominance for citrate content and close to complete dominance for acidity. The total sour content and acidity traits refered to only one pleiotropic major gene. The additive effect and dominance effect of the major gene are estimated as1.2735 mg g-1 FW and -2.9746 mg g-1 FW on total sour content, and -0.647and–0.6697 on acidity, respectively, The major gene showed overdominance for glucose content and close to complete dominance for acidity.
The influence of temperature and solar on sugar and acid content of fruit FN3 melon cultured in spring film house, autumn film house and autumu green house, were analyzed by using grey relational grade analysis method. The results showed that the influence rankings on sugar content were: accumulative solar radiation > accumulative daily temperature difference > effective accumulative temperature. The influence of temperature and solar on acid content of FN3 fruit were in order of accumulative daily temperature difference > accumulative solar radiation > effective accumulative temperature.
With different crops for rotation, the effects of air temperature and accumulated daily difference in temperature on the sucrose, citrate accumulation and sugar acid radio of greenhouse melon fruit were studied, with related simulation model established. The results showed that the sucrose, citrate accumulation and sugar acid radio model were logistic function, all of which were driven by effective accumulative temperature. The constant term in the functions was driven by accumulative daily difference in temperature, and the correlation was a linear function. Model test showed that the models were able to objectively simulate and predict the changes of fruit sucrose, citrate accumulation and sugar acid radio with fruit development, and possessed practical value for the accurate estimates to melon fruit inherent quality and optimization to determine harvest of melon fruit.
引文
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