粉单竹遗传多样性和麻竹种内杂交子代遗传变异研究
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摘要
粉单竹(Bambusa chungii)和麻竹(Dendrocalamus latiflorus)分别是我国南方地区重要的材用竹种和笋竹两用竹种。中国林科院亚林所有关研究人员于2001年收集了粉单竹自然分布区内(广东、广西和海南)的8个产地在广东茂名建立了粉单竹产地试验林。同时在福建南靖和广东南雄设置了通过人工杂交得到的不同产地麻竹种内杂交子代30个家系的对比试验。本文以此为材料，对粉单竹的遗传多样性和麻竹种内杂交子代遗传变异情况进行了研究。
1．粉单竹群体表型遗传多样性：研究发现所调查的表型性状在产地群体间存在显著或极显著差异；同一产地内各性状变异幅度也较大，具有较为丰富的表型遗传多样性，表明长期的地理隔离和缺乏基因交流使粉单竹群体产生了遗传分化。粉单竹的主要性状受中度以上遗传控制，胸径、竹高、枝下高、枝下平均节数和胸径处节长等重要的生长性状之间存在紧密的正向遗传相关，非常有利于多性状的同时改良。表型性状与环境因子的相关分析表明，粉单竹的地理变异主要反应冷—暖的气候生态模式，其生长与分布范围主要受热量因子的限制。利用综合评定法以用材为目的对粉单竹进行产地选择，认为广东增城、怀集、信宜和广西桂林为广东西南部地区发展粉单竹林可选用的较好产地。
2．粉单竹纸浆材材性变异及优良产地选择：粉单竹的纤维长度、纤维素含量和1％NaOH抽出物含量在产地间存在极显著或显著差异。木素含量和纤维长度的绝对变化量很小，无选择意义。苯醇抽出物在个体间变化很大，可进行优良单丛的选择。纤维长度及纤维长宽比呈现纬度上从南到北(海拔上从低到高)逐渐减小的地理变异模式。
纤维素含量和纤维长度的遗传力高，受遗传控制强；1％NaOH抽出物受中等强度遗传控制；木素含量和纤维长宽比受遗传控制最弱。1％NaOH抽出物和苯醇抽出物与产量指标呈显著负相关，纤维素含量与产量指标呈显著正相关；纤维形态与生长、产量指标关系密切。本文所考察的材性与生长、产量性状间的相关关系多对生长与材性的联合选择有利。利用主成分分析对材性和生长、产量性状联合选择，初步选出广东怀集、广东信宜、广西南宁和广西桂林4个优良纸浆材产地。
3．麻竹种内杂交子代生长性状遗传变异分析：对发笋量、地径、苗高性状的调查研究发现，3个性状在家系间存在显著或极显著差异，变异系数为13.69％-44.33％，遗传变异丰富。利用隶属函数法进行单点选择，以20％的入选率在两试验点分别选出6个优良家系。南靖试验点选择后平均发笋量、平均地径、平均竹高分别比未选择前提高了25.3％、11.69％和18.38％。南雄试验点选择后平均发笋量、平均地径、平均竹高分别比选择前提高了23.8％，14.1％和12.9％。联合分析发现上述3性状的基因型与环境互作效应(GEI)极显著，南靖点各性状的遗传力高于南雄点。单点选择后，发笋量、地径和竹高等性状在南靖点获得的遗传增益为15.94％，10.29％，18.01％，高于南雄点的9.05％，7.03％，5.58％。
4．麻竹种内杂交子代群体遗传变异的ISSR分析：利用9个ISSR引物对22个杂种家系样本的扩增结果表明，家系间的基因多样度(He)和Shannon多样性指数分别是0.3655和0.5442。22个家系间遗传相似系数值变化范围为0.3953-0.7907，平均为0.6130；遗传距离变化范围为0.2348-0.928，平均为0.4996，杂种群体的遗传多样性较高，说明通过杂交可以拓宽和丰富竹子的种内基因资源。
5．麻竹种内杂交子代竹笋营养成分分析：对30个家系竹笋营养成分的分析表明，可溶性总糖含量、淀粉含量、游离氨基酸总量、蛋白质含量在家系间的变异幅度都很大，变异系数依次为23.69％；26.70％；38.26％；15.76％。蛋白质水解氨基酸中必需氨基酸的比例都在30％以上，变异幅度为33.8％-42.97％。根据竹笋可溶性糖、淀粉、游离氨基酸总量、蛋白质及必需氨基酸比例等指标对参试30个家系进行聚类分析，选出了氨基酸和蛋白质含量较高的两个类群。这两个类群共包括11个家系，其中10个家系是来自JD或HK两个产地麻竹为亲本的杂交子代。说明麻竹竹笋的营养品质主要受遗传的影响，通过合适的亲本选配可以得到营养品质优良的杂种后代。
6．麻竹杂交子代几个重要酶活性的初步研究：蔗糖磷酸合成酶(SPS)和蔗糖合成酶(SS)，谷氨酰胺合成酶(GS)和谷氨酸合成酶(GOGAT)活性在家系间差异很大，SS、SPS、GS和GOGAT活性变异幅度依次为24.72μmolsucrose·g~(-1)·h~(-1)-68.87μmolsucrose·g~(-1)·h~(-1)；44.91μmolsucrose·g~(-1)·h~(-1)-267.55μmolsucrose·g~(-1)·h~(-1)；3.48μmol·g~(-1)·h~(-1)-14.78μmol·g~(-1)·h~(-1)；0.0111-0.2806个酶单位。竹笋游离氨基酸总量与GOGAT活性之间存在显著正相关，说明GOGAT活性高有利于氨的同化和氨基酸的合成积累，可考虑将其作为评价麻竹氨同化能力的生理指标之一。
Bambusa chungii and Dendrocalamus latiflorus are important timber bamboo and ediblebamboo species in south of China. 8 provenances of B. chungii were collected from it naturallydistributing geographical region and a provenance testing with 8 provenances was establishedby a study team from the Research Institute of Subtropical Forestry, CAF. 30 hybrid familiesobtained via artificial crossing between different provenances of D. latiflorus were planted inNanjin, Fujian provenance and Nanxiong, Guangdong provenance respectively for comparing.Studies on gene-diversity of B. chungii and genetic variations in hybrid offspring of D.latiflorus populations were developed based on material above.
1. Morphological gene-diversity analysis for B. chungii: The analysis showed that there wereabundant variances in main morphological traits among the different provenances and all thevariances reached significant or great significant level. Genetic variations emerged because thepopulations growing in segregated regions for long time and low gene flow among and withinthe populations. There also a higher variable ranges within the same population. The geneticdiversity of B. chungii population was abundant. Heritabilities of the main characteristics of B.chungii were strong and stronger. The correlative coefficients among DBH, height, clear height,nodes number under the first branch, and node length at DBH were significant and positive,that would benefit for multi-traits improvement synchronously. The correlation analysisrevealed that thermal factors were limiting factors for growth and distribution of B. chungii,which reflected climate changes from cold to warm in the distributing area of this species.Selection for all the populations was given on the basis of synthesis evaluation. Theprovenances i. e. Zengcheng, Huaiji, Xinyi and Guilin were considered the optimal provenancesfor extension in the region around southwest of Guandong Province.
2. Variations analysis of pulpbamboo properties of B. chungii and the optimum sources screen:fibre length, fibrin and 1% NaOH extraction contents had significant difference among 8provenances. The absolute variations of lignin content and fibre length in different provenanceswere low, which suggested that those two traits were unnecessary to be included in theselection program. Superior clump with lower benzo-alcohol extractive content could be selected because large differences among all clumps tested in this study. A decrease tendencyof fibre length and fibre length-width ratio from south to north (high altitude to low altitude)was found.
Heritabilities of fibrin and fibre length were strong, while 1% NaOH extraction contents'Heritabilities was medium, and lignin and fibre width got trivial genetic control. The results ofcorrelation analysis between culm properties and growth or yield traits presented there werehighlighted negative correlation between 1% NaOH extraction, benzo-alcohol extractivecontents and yield but significant positive correlation between fibrin content and yield. Tenselyrelationship was detected between fibre form indices and growth or yield traits. Most of theresults discovered were favorable for optimum pulpbamboo selections associating with growthand yield characters. 4 provenances i. e. Huaiji, Xinyi, Nanning and Guilin have been selectedas superior sources based on principle component analysis combined pulpbamboo properties,growth and yield traits.
3. Genetic variation analysis of growth traits for hybrid offspring of D. latiflorus: For shootnumber, collar diameter and height, there were significant differences among 30 familiesbecause of rich genetic variance, while the coefficient of variation (CV) of the 3 traits was13.69%-44.33% in two testing sites. Each 6 families have been selected as superior familiesbased on subordinate function value from two sites. The raise of shoot number, collar diameterand height were25.3%, 11.69%, 18.38% at Nanjin and 23.8%, 14.1%, 12.9% at Nanxiong.Combined variation analysis on 3 traits showed that the interaction between genotype andenvironment was highly significant. Heritabilities of 3 traits and the expectant genetic gain atsingle site at Nanjin were higher than at Nanxiong.
4. Evaluation of the genetic variation of the hybrid descendants of D. latiflorus based on ISSRmarkers: 9 ISSR primers were utilized to analyze the genetic variance of 22 hybrid familieswith ISSR techniques. The results demonstrated that Nei's gene diversity(He) and Shannon'sinformation index were 0.3655 and 0.5442 at the species level respectively. Genetic similaritycoefficient ranged from 0.3953 to 0.7907 and genetic distance ranged from 0.2348 to 0.928 in22 hybrid family populations. The mean of genetic similarity coefficient and genetic distancewere 0.6130 and 0.4996 respectively, which proved that the hybrid population had a highergenetic diversity and gene resources could be enriched by crossbreeding.
5. Bamboo shoot nutrient content variation of the hybrid descendants of D. latiflorus: Therewere great variance ranges in content of Water-soluble carbohydrates, starch, dissociate aminoacid and protein among different family, the CV of these indexes were 23.69%, 26.70%, 38.26%and 15.76% in turn. Proportion of essential amino acid exceeded 30% in all families andfluctuated between 33.8% and 42.97%. All of the families could be classed into 3 clusters onbasis of similarities of contents of Water-soluble carbohydrates, starch, dissociate amino acid,protein and Proportion of essential amino acid. 11 families included in clusterⅠand clusterⅡwere excellent families at nutrient quality with higher contents of dissociate amino acid,protein and Proportion of essential amino acid. 10 of them had one common parent JD or HKprovenance at least. It indicated that bamboo shoot nutrition of D. latiflorus was controlled bygenetic, excellent hybrids at nutrient quality could be obtained by crossbreeding with suitableparents.
6. A first studies on activities of several enzymes of the hybrid descendants of D latiflorus:Activities of sucrose phosphate synthase(SPS), sucrose synthase(SS), glutamine synthetase(GS)and glutamate synthase(GOGAT) were investigated in this research. The results indicated greatdifference of the activities of 4 enzymes in all families. The number of activities of SS, SPS,GS and GOGAT varied from 24.72μmolsucrose.g~(-1)·h~(-1) to 68.87μmolsucrose·g~(-1)·h~(-1),44.91μmolsucrose·g~(-1)·h~(-1) to 267.55μmolsucrose·g~(-1)·h~(-1), 3.48μmol·g~(-1)·h~(-1) to 14.78μmol·g~(-1)·h~(-1),0.0111 to 0.2806 unit in turn. Correlation coefficient between dissociate amino acid andGOGAT activity was significant, which expressed higher activity of GOGAT would promoteassimilation of ammonium and accumulation of amino acid. GOGAT could be considered asone of the physiological index to evaluate the capacity of ammonium assimilation of Dlatiflorus.
1．粉单竹群体表型遗传多样性：研究发现所调查的表型性状在产地群体间存在显著或极显著差异；同一产地内各性状变异幅度也较大，具有较为丰富的表型遗传多样性，表明长期的地理隔离和缺乏基因交流使粉单竹群体产生了遗传分化。粉单竹的主要性状受中度以上遗传控制，胸径、竹高、枝下高、枝下平均节数和胸径处节长等重要的生长性状之间存在紧密的正向遗传相关，非常有利于多性状的同时改良。表型性状与环境因子的相关分析表明，粉单竹的地理变异主要反应冷—暖的气候生态模式，其生长与分布范围主要受热量因子的限制。利用综合评定法以用材为目的对粉单竹进行产地选择，认为广东增城、怀集、信宜和广西桂林为广东西南部地区发展粉单竹林可选用的较好产地。
2．粉单竹纸浆材材性变异及优良产地选择：粉单竹的纤维长度、纤维素含量和1％NaOH抽出物含量在产地间存在极显著或显著差异。木素含量和纤维长度的绝对变化量很小，无选择意义。苯醇抽出物在个体间变化很大，可进行优良单丛的选择。纤维长度及纤维长宽比呈现纬度上从南到北(海拔上从低到高)逐渐减小的地理变异模式。
纤维素含量和纤维长度的遗传力高，受遗传控制强；1％NaOH抽出物受中等强度遗传控制；木素含量和纤维长宽比受遗传控制最弱。1％NaOH抽出物和苯醇抽出物与产量指标呈显著负相关，纤维素含量与产量指标呈显著正相关；纤维形态与生长、产量指标关系密切。本文所考察的材性与生长、产量性状间的相关关系多对生长与材性的联合选择有利。利用主成分分析对材性和生长、产量性状联合选择，初步选出广东怀集、广东信宜、广西南宁和广西桂林4个优良纸浆材产地。
3．麻竹种内杂交子代生长性状遗传变异分析：对发笋量、地径、苗高性状的调查研究发现，3个性状在家系间存在显著或极显著差异，变异系数为13.69％-44.33％，遗传变异丰富。利用隶属函数法进行单点选择，以20％的入选率在两试验点分别选出6个优良家系。南靖试验点选择后平均发笋量、平均地径、平均竹高分别比未选择前提高了25.3％、11.69％和18.38％。南雄试验点选择后平均发笋量、平均地径、平均竹高分别比选择前提高了23.8％，14.1％和12.9％。联合分析发现上述3性状的基因型与环境互作效应(GEI)极显著，南靖点各性状的遗传力高于南雄点。单点选择后，发笋量、地径和竹高等性状在南靖点获得的遗传增益为15.94％，10.29％，18.01％，高于南雄点的9.05％，7.03％，5.58％。
4．麻竹种内杂交子代群体遗传变异的ISSR分析：利用9个ISSR引物对22个杂种家系样本的扩增结果表明，家系间的基因多样度(He)和Shannon多样性指数分别是0.3655和0.5442。22个家系间遗传相似系数值变化范围为0.3953-0.7907，平均为0.6130；遗传距离变化范围为0.2348-0.928，平均为0.4996，杂种群体的遗传多样性较高，说明通过杂交可以拓宽和丰富竹子的种内基因资源。
5．麻竹种内杂交子代竹笋营养成分分析：对30个家系竹笋营养成分的分析表明，可溶性总糖含量、淀粉含量、游离氨基酸总量、蛋白质含量在家系间的变异幅度都很大，变异系数依次为23.69％；26.70％；38.26％；15.76％。蛋白质水解氨基酸中必需氨基酸的比例都在30％以上，变异幅度为33.8％-42.97％。根据竹笋可溶性糖、淀粉、游离氨基酸总量、蛋白质及必需氨基酸比例等指标对参试30个家系进行聚类分析，选出了氨基酸和蛋白质含量较高的两个类群。这两个类群共包括11个家系，其中10个家系是来自JD或HK两个产地麻竹为亲本的杂交子代。说明麻竹竹笋的营养品质主要受遗传的影响，通过合适的亲本选配可以得到营养品质优良的杂种后代。
6．麻竹杂交子代几个重要酶活性的初步研究：蔗糖磷酸合成酶(SPS)和蔗糖合成酶(SS)，谷氨酰胺合成酶(GS)和谷氨酸合成酶(GOGAT)活性在家系间差异很大，SS、SPS、GS和GOGAT活性变异幅度依次为24.72μmolsucrose·g~(-1)·h~(-1)-68.87μmolsucrose·g~(-1)·h~(-1)；44.91μmolsucrose·g~(-1)·h~(-1)-267.55μmolsucrose·g~(-1)·h~(-1)；3.48μmol·g~(-1)·h~(-1)-14.78μmol·g~(-1)·h~(-1)；0.0111-0.2806个酶单位。竹笋游离氨基酸总量与GOGAT活性之间存在显著正相关，说明GOGAT活性高有利于氨的同化和氨基酸的合成积累，可考虑将其作为评价麻竹氨同化能力的生理指标之一。
Bambusa chungii and Dendrocalamus latiflorus are important timber bamboo and ediblebamboo species in south of China. 8 provenances of B. chungii were collected from it naturallydistributing geographical region and a provenance testing with 8 provenances was establishedby a study team from the Research Institute of Subtropical Forestry, CAF. 30 hybrid familiesobtained via artificial crossing between different provenances of D. latiflorus were planted inNanjin, Fujian provenance and Nanxiong, Guangdong provenance respectively for comparing.Studies on gene-diversity of B. chungii and genetic variations in hybrid offspring of D.latiflorus populations were developed based on material above.
1. Morphological gene-diversity analysis for B. chungii: The analysis showed that there wereabundant variances in main morphological traits among the different provenances and all thevariances reached significant or great significant level. Genetic variations emerged because thepopulations growing in segregated regions for long time and low gene flow among and withinthe populations. There also a higher variable ranges within the same population. The geneticdiversity of B. chungii population was abundant. Heritabilities of the main characteristics of B.chungii were strong and stronger. The correlative coefficients among DBH, height, clear height,nodes number under the first branch, and node length at DBH were significant and positive,that would benefit for multi-traits improvement synchronously. The correlation analysisrevealed that thermal factors were limiting factors for growth and distribution of B. chungii,which reflected climate changes from cold to warm in the distributing area of this species.Selection for all the populations was given on the basis of synthesis evaluation. Theprovenances i. e. Zengcheng, Huaiji, Xinyi and Guilin were considered the optimal provenancesfor extension in the region around southwest of Guandong Province.
2. Variations analysis of pulpbamboo properties of B. chungii and the optimum sources screen:fibre length, fibrin and 1% NaOH extraction contents had significant difference among 8provenances. The absolute variations of lignin content and fibre length in different provenanceswere low, which suggested that those two traits were unnecessary to be included in theselection program. Superior clump with lower benzo-alcohol extractive content could be selected because large differences among all clumps tested in this study. A decrease tendencyof fibre length and fibre length-width ratio from south to north (high altitude to low altitude)was found.
Heritabilities of fibrin and fibre length were strong, while 1% NaOH extraction contents'Heritabilities was medium, and lignin and fibre width got trivial genetic control. The results ofcorrelation analysis between culm properties and growth or yield traits presented there werehighlighted negative correlation between 1% NaOH extraction, benzo-alcohol extractivecontents and yield but significant positive correlation between fibrin content and yield. Tenselyrelationship was detected between fibre form indices and growth or yield traits. Most of theresults discovered were favorable for optimum pulpbamboo selections associating with growthand yield characters. 4 provenances i. e. Huaiji, Xinyi, Nanning and Guilin have been selectedas superior sources based on principle component analysis combined pulpbamboo properties,growth and yield traits.
3. Genetic variation analysis of growth traits for hybrid offspring of D. latiflorus: For shootnumber, collar diameter and height, there were significant differences among 30 familiesbecause of rich genetic variance, while the coefficient of variation (CV) of the 3 traits was13.69%-44.33% in two testing sites. Each 6 families have been selected as superior familiesbased on subordinate function value from two sites. The raise of shoot number, collar diameterand height were25.3%, 11.69%, 18.38% at Nanjin and 23.8%, 14.1%, 12.9% at Nanxiong.Combined variation analysis on 3 traits showed that the interaction between genotype andenvironment was highly significant. Heritabilities of 3 traits and the expectant genetic gain atsingle site at Nanjin were higher than at Nanxiong.
4. Evaluation of the genetic variation of the hybrid descendants of D. latiflorus based on ISSRmarkers: 9 ISSR primers were utilized to analyze the genetic variance of 22 hybrid familieswith ISSR techniques. The results demonstrated that Nei's gene diversity(He) and Shannon'sinformation index were 0.3655 and 0.5442 at the species level respectively. Genetic similaritycoefficient ranged from 0.3953 to 0.7907 and genetic distance ranged from 0.2348 to 0.928 in22 hybrid family populations. The mean of genetic similarity coefficient and genetic distancewere 0.6130 and 0.4996 respectively, which proved that the hybrid population had a highergenetic diversity and gene resources could be enriched by crossbreeding.
5. Bamboo shoot nutrient content variation of the hybrid descendants of D. latiflorus: Therewere great variance ranges in content of Water-soluble carbohydrates, starch, dissociate aminoacid and protein among different family, the CV of these indexes were 23.69%, 26.70%, 38.26%and 15.76% in turn. Proportion of essential amino acid exceeded 30% in all families andfluctuated between 33.8% and 42.97%. All of the families could be classed into 3 clusters onbasis of similarities of contents of Water-soluble carbohydrates, starch, dissociate amino acid,protein and Proportion of essential amino acid. 11 families included in clusterⅠand clusterⅡwere excellent families at nutrient quality with higher contents of dissociate amino acid,protein and Proportion of essential amino acid. 10 of them had one common parent JD or HKprovenance at least. It indicated that bamboo shoot nutrition of D. latiflorus was controlled bygenetic, excellent hybrids at nutrient quality could be obtained by crossbreeding with suitableparents.
6. A first studies on activities of several enzymes of the hybrid descendants of D latiflorus:Activities of sucrose phosphate synthase(SPS), sucrose synthase(SS), glutamine synthetase(GS)and glutamate synthase(GOGAT) were investigated in this research. The results indicated greatdifference of the activities of 4 enzymes in all families. The number of activities of SS, SPS,GS and GOGAT varied from 24.72μmolsucrose.g~(-1)·h~(-1) to 68.87μmolsucrose·g~(-1)·h~(-1),44.91μmolsucrose·g~(-1)·h~(-1) to 267.55μmolsucrose·g~(-1)·h~(-1), 3.48μmol·g~(-1)·h~(-1) to 14.78μmol·g~(-1)·h~(-1),0.0111 to 0.2806 unit in turn. Correlation coefficient between dissociate amino acid andGOGAT activity was significant, which expressed higher activity of GOGAT would promoteassimilation of ammonium and accumulation of amino acid. GOGAT could be considered asone of the physiological index to evaluate the capacity of ammonium assimilation of Dlatiflorus.
引文
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