核桃遗传多样性与优良类型选择研究
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摘要
本研究通过对我国核桃主产区陕西、新疆、云南等不同产地的核桃种质资源进行调查，并在陕西洛南、黄龙进行引种栽培，测定了核桃的13个形态指标、5个经济性状指标，以及不同类型DNA遗传标记，分析了不同类型核桃的生长、坚果品质、耐低温生理、生态适应性等差异；综合评价出优良的区域性种质资源，为核桃良种选育和区划提供了依据。主要结果如下：
1、对12个产地核桃的形态表型和核桃经济性状的分析表明，不同类型的形态和经济性状差异较大，多样性丰富。13个主要形态学性状的变异系数为2.8-19.1%，其中胸径、叶片厚度、坚果大小、侧花芽比例、平均枝长、产量等变异系数大于10%；5个经济指标的变异系数为9.7-37.14%。胸径、叶片厚度、侧花芽率与产量也具有显著相关性（p<0.05〉）。不同产地核桃生长性状、质量性状具有明显的地域性差异；核桃营养成分和土壤养分指标具有关联性。
2、对采自我国不同生态地理区的44个核桃样品、8个铁核桃样品和引自美国的7个核桃样品应用多态性较高的16个核桃SSR标记分析，共获得87个等位基因，扩增等位基因平均为5.4个。每个SSR位点的观察杂合度平均为0.3894，各位点PIC值平均为0.6632；核桃组59个样品的杂合度平均0.387，杂合度较低；对59个核桃组聚类分析表明：来自秦岭山地的核桃样品与华北核桃应属于同一生态类型；巩留的野核桃应属于与新疆生态型和华北生态型不同的一个独立的生态型；美国核桃和铁核桃应属于核桃种下不同的生态类型。
3、自然降温条件下核桃耐寒分析表明，新疆核桃类型最耐寒。将原产云南、四川、陕西、新疆的优良核桃类型，集中栽培在洛南山区，测定其在自然降温条件下的保护酶、可溶性蛋白等抗寒性指标，比较分析其耐寒性差异，不同产地核桃的抗寒性从强到弱依次为：新疆、陕西、四川、云南核桃。新疆核桃抗寒性较强，说明了核桃的抗寒能力和其原产地的气候条件有很大的关系。
4、核桃不同类型引种试验中，形态和经济性状指标与遗传参数的估算结果表明，12个核桃类型在陕西洛南均能正常生长和结果。2个试验点各地理类型的嫁接建园成活率差异不大，但核桃形态指标差异显著；营养期与结果期的树高和胸径生长类型间差异显著；各核桃类型稳产程度大小依次为A2、A3、A1、A5、A7、A9、A6、A4、A8；随着核桃园进入结果盛期后，各类型的核桃产量逐步趋向稳定；各类型年平均叶生长量、叶绿素含量、呼吸速率、光合速率等指标也存在一定的差异。运用主成分分析法，综合评价出适合在陕西省产区栽培的核桃优良类型，表现优异的地理类型有A1、A2、A3、A6、A7、A5，其经济性状的增益在10.68%~31.67%之间。
The test had conlected the most germplasm that cultivated in the walnut production areasof China, and had investigated13morphological growth index,5economical qualities of nuts,measured the DNA marker, analyzed the difference of growth，the nut qualities，the coldresistance physiological changes, and ecological adaptability, and comprehensively evaluedand choosed the relatively fine germplasm for production. The main results were as follows:
1. Morphological traits of walnut and qualities of kernel were measured and analyzedamong the12different original. The result show that walnut in China have very rich diversityof morphological and economic traits. The CV of13main Morphological index is range from2.8%to19.1%which indicated the difference of the traits from the different original. The CVof DBH, the thickness of leaf, the lateral fruit bud percentage, the average branch lenth, andthe yield are more than10%. The CV of5economical index is range from9.7-37.14%Thecorrelation between tree DBH, the thickness of leaf, the lateral fruit bud percentage and theyield are significant（p<0.05〉）. According to yeild analysis shows it is difference of growthtraits of walnut and qualities of kernel form different original, and also reflects the broadlygeographic distribution of walnut depend on soil and other climate facters.
2. In order to analyse the relatedness and the genetic diversity among germplasm in Sect.Juglans by SSR markers, the experiment use16SSR primer pairs to amplify SSR fragmentsof44samples of Juglans regia from different ecological types,8samples of J. sigillata, and7cultivars of Amerian walnut. The16pairs of SSR primers has generated87alleles，whichaveraged5.4. The observed heterozygosity(Ho) of16SSR loci ranged from0.000to0.931,which averaged0.3894. The value of PIC ranged from0.3953to0.7472, which averaged0.6632. The heterozygosity of59Sect. Juglans samples ranged from0.133to0.813, andaveraged0.387. It showed that the heterozygosity of Sect. Juglans samples was low. In thedendrogram, The walnut samples from Qinling Mountain and those from Huabei shouldbelong to the same ecological type. Wild walnut samples from Gongliu county, Xinjiangshould belong to the one absolute ecological type, which was different from Xinjiang ecological type and Huabei ecological type. The American walnuts and J. sigillata, should bedifferent ecological type in J. regia.
3. The activity of protective enzymes, soluble sugar, and other cold resistance index ofwalnut types from Yunnan, Sichuan,Shaanxi and Xinjiang were studied under the naturetemperature change condition. The range of cold resistance of geographical type of walnut areas follows: Xinjiang＞Shaanxi＞Sichuan＞Yunnan. The membership degree value fromWensu, Xinjiang showed more higher than that from other region, which can explan that thecold resistance from Xinjiang is much higher than from other region.
4.The traits and genetic parameter evaluation index shows that all of the selective walnuttypes accomplished normal growth in the test plot in Luonan, Shaanxi, and easily adapted tothe Shaanxi conditions similar to the original. The growth the height were great different fromthe diameter of the walnut in the vegetative growth and fruiting. The production stabilityvalue from high to low is:A2, A3, A1, A5, A7, A9, A6, A4, A8. The yield will be stable fromthe canopy closed. The leaf area and other photosynthetic index also be different form walnuttypes. Using the methode of principal component analysis, better walnut types which wereadapt to cultivating in Luonan,Shaanxi were selected, and the selections increased benefitsrange from10.68%~31.67%in average.
1、对12个产地核桃的形态表型和核桃经济性状的分析表明，不同类型的形态和经济性状差异较大，多样性丰富。13个主要形态学性状的变异系数为2.8-19.1%，其中胸径、叶片厚度、坚果大小、侧花芽比例、平均枝长、产量等变异系数大于10%；5个经济指标的变异系数为9.7-37.14%。胸径、叶片厚度、侧花芽率与产量也具有显著相关性（p<0.05〉）。不同产地核桃生长性状、质量性状具有明显的地域性差异；核桃营养成分和土壤养分指标具有关联性。
2、对采自我国不同生态地理区的44个核桃样品、8个铁核桃样品和引自美国的7个核桃样品应用多态性较高的16个核桃SSR标记分析，共获得87个等位基因，扩增等位基因平均为5.4个。每个SSR位点的观察杂合度平均为0.3894，各位点PIC值平均为0.6632；核桃组59个样品的杂合度平均0.387，杂合度较低；对59个核桃组聚类分析表明：来自秦岭山地的核桃样品与华北核桃应属于同一生态类型；巩留的野核桃应属于与新疆生态型和华北生态型不同的一个独立的生态型；美国核桃和铁核桃应属于核桃种下不同的生态类型。
3、自然降温条件下核桃耐寒分析表明，新疆核桃类型最耐寒。将原产云南、四川、陕西、新疆的优良核桃类型，集中栽培在洛南山区，测定其在自然降温条件下的保护酶、可溶性蛋白等抗寒性指标，比较分析其耐寒性差异，不同产地核桃的抗寒性从强到弱依次为：新疆、陕西、四川、云南核桃。新疆核桃抗寒性较强，说明了核桃的抗寒能力和其原产地的气候条件有很大的关系。
4、核桃不同类型引种试验中，形态和经济性状指标与遗传参数的估算结果表明，12个核桃类型在陕西洛南均能正常生长和结果。2个试验点各地理类型的嫁接建园成活率差异不大，但核桃形态指标差异显著；营养期与结果期的树高和胸径生长类型间差异显著；各核桃类型稳产程度大小依次为A2、A3、A1、A5、A7、A9、A6、A4、A8；随着核桃园进入结果盛期后，各类型的核桃产量逐步趋向稳定；各类型年平均叶生长量、叶绿素含量、呼吸速率、光合速率等指标也存在一定的差异。运用主成分分析法，综合评价出适合在陕西省产区栽培的核桃优良类型，表现优异的地理类型有A1、A2、A3、A6、A7、A5，其经济性状的增益在10.68%~31.67%之间。
The test had conlected the most germplasm that cultivated in the walnut production areasof China, and had investigated13morphological growth index,5economical qualities of nuts,measured the DNA marker, analyzed the difference of growth，the nut qualities，the coldresistance physiological changes, and ecological adaptability, and comprehensively evaluedand choosed the relatively fine germplasm for production. The main results were as follows:
1. Morphological traits of walnut and qualities of kernel were measured and analyzedamong the12different original. The result show that walnut in China have very rich diversityof morphological and economic traits. The CV of13main Morphological index is range from2.8%to19.1%which indicated the difference of the traits from the different original. The CVof DBH, the thickness of leaf, the lateral fruit bud percentage, the average branch lenth, andthe yield are more than10%. The CV of5economical index is range from9.7-37.14%Thecorrelation between tree DBH, the thickness of leaf, the lateral fruit bud percentage and theyield are significant（p<0.05〉）. According to yeild analysis shows it is difference of growthtraits of walnut and qualities of kernel form different original, and also reflects the broadlygeographic distribution of walnut depend on soil and other climate facters.
2. In order to analyse the relatedness and the genetic diversity among germplasm in Sect.Juglans by SSR markers, the experiment use16SSR primer pairs to amplify SSR fragmentsof44samples of Juglans regia from different ecological types,8samples of J. sigillata, and7cultivars of Amerian walnut. The16pairs of SSR primers has generated87alleles，whichaveraged5.4. The observed heterozygosity(Ho) of16SSR loci ranged from0.000to0.931,which averaged0.3894. The value of PIC ranged from0.3953to0.7472, which averaged0.6632. The heterozygosity of59Sect. Juglans samples ranged from0.133to0.813, andaveraged0.387. It showed that the heterozygosity of Sect. Juglans samples was low. In thedendrogram, The walnut samples from Qinling Mountain and those from Huabei shouldbelong to the same ecological type. Wild walnut samples from Gongliu county, Xinjiangshould belong to the one absolute ecological type, which was different from Xinjiang ecological type and Huabei ecological type. The American walnuts and J. sigillata, should bedifferent ecological type in J. regia.
3. The activity of protective enzymes, soluble sugar, and other cold resistance index ofwalnut types from Yunnan, Sichuan,Shaanxi and Xinjiang were studied under the naturetemperature change condition. The range of cold resistance of geographical type of walnut areas follows: Xinjiang＞Shaanxi＞Sichuan＞Yunnan. The membership degree value fromWensu, Xinjiang showed more higher than that from other region, which can explan that thecold resistance from Xinjiang is much higher than from other region.
4.The traits and genetic parameter evaluation index shows that all of the selective walnuttypes accomplished normal growth in the test plot in Luonan, Shaanxi, and easily adapted tothe Shaanxi conditions similar to the original. The growth the height were great different fromthe diameter of the walnut in the vegetative growth and fruiting. The production stabilityvalue from high to low is:A2, A3, A1, A5, A7, A9, A6, A4, A8. The yield will be stable fromthe canopy closed. The leaf area and other photosynthetic index also be different form walnuttypes. Using the methode of principal component analysis, better walnut types which wereadapt to cultivating in Luonan,Shaanxi were selected, and the selections increased benefitsrange from10.68%~31.67%in average.
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