烟草微核心种质构建及相关性状数量遗传分析
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摘要
微核心种质是核心种质的核心种质，是核心种质的代表性子集，具有最小的遗传重复，能够最大程度地代表原资源群体的遗传多样性和遗传结构。烟草微核心种质研究的最终目的是为了促进烟草种质资源的高效管理、深入评价和充分利用，最大程度地满足生产和科研的需要。本研究以烟草原核心种质为材料，比较研究数量性状数据和分子标记数据不同整合比例及不同抽样策略构建微核心种质的遗传多样性保有量，构建了烟草微核心种质。在烟草微核心种质的基础上，筛选2份有代表性的烤烟微核心种质，运用植物数量性状“主基因+多基因”混合遗传模型，进行烤烟重要植物学及生理性状的遗传研究，以期为烤烟数量遗传育种提供一定的理论指导。主要研究结论如下：
（1）通过12份不同烟草种质类型，筛选均匀分布于烟草24条连锁群上的700对SSR标记，综合考虑PIC值、扩增带型统计的难易及引物的重复性，获得54对SSR核心引物，建立了核心引物数据库。本研究有利于规范引物筛选程序、提高引物筛选效率，对烟草种质资源的遗传多样性分析、核心种质构建、指纹图谱构建以及种子真伪性和纯度的鉴定等都具有现实意义。
（2）以我国2001年构建的446份烟草核心种质为材料，根据23个数量性状资料和54对SSR引物的分子标记数据，开展微核心种质构建研究。不同取样比例（40%、30%、25%、20%、15%、10%和5%）分析表明，15%的取样比例可获得普通烟草90%以上的多态位点百分率，30%的取样比例可获得黄花烟草80%以上的多态位点百分率，是较好的微核心种质取样规模；表型数据和分子数据整合后的数据要比不整合好，且表型数据与分子数据的比例为0.1:0.9时，整合效果最好。对于连续型数据遗传距离估算的2种距离（欧氏距离和马氏距离）中，欧氏距离效果最优；对于离散型数据遗传距离估算的4种距离（Simple matching、Jaccard、Nei-Li和主成分距离）中，Jaccard距离效果最优。对4种分组取样策略（简单比例法、平方根比例法、对数比例法和多样性比例法）和3种取样方法（随机取样法、优先取样法和变异度取样法）比较表明，总体取样方法中变异度取样法最优，在组内样本含量较大时，简单比例法和多样性比例法获得的微核心种质代表性最好，为最优取样策略；而当组内样本含量较小时，对数比例法获得的微核心种质代表性最好，为最优取样策略。最后，在简单比例法和对数比例法取样筛选出的117份核心样品中，又通过定向选择补充了10份具有优异表型性状的材料，构建了含127份烟草的微核心种质。分子和表型检验都表明本研究所构建的微核心种质具有较好的代表性。
（3）利用“主基因+多基因”混合遗传模型的6个世代联合分离分析方法，分析烤烟组合丸叶×Coker319几个重要植物学性状的遗传效应。结果表明，烤烟的株高、叶数、叶面积和鲜叶重受2对加性-显性-上位性主基因+加性-显性-上位性多基因控制，株高与叶数遗传以加性效应及显性×显性上位性效应为主，叶面积和鲜叶重各遗传效应相差不多，其上位性效应>加性效应>显性效应，F2世代的主基因遗传率分别为57.53%、42.63%、30.32%和44.26%。移栽至中心花开放天数受2对加性-显性-上位性主基因+加性-显性多基因控制，以加性×加性上位性效应、加性效应及显性×显性上位性效应为主，主基因遗传率为64.79%。茎围和比叶重均受1对完全显性主基因+加性-显性多基因控制，茎围遗传以多基因为主，其多基因加性效应和显性效应大小相当，比叶重遗传主基因、多基因的加性效应和显性效应大致相当，主基因遗传率分别为2.48%和38.71%。叶形指数受1对加性-显性主基因+加性-显性-上位性多基因控制，主基因加性效应与显性效应基本相当，主基因遗传率为49.64%。叶长、叶宽、节距和蒴果重受加性-显性-上位性多基因控制，多基因遗传率分别为60.75%、62.14%、75.08%和82.34%。
（4）应用主基因+多基因6个世代联合分析方法对烤烟丸叶×Coker319组合的几个生理性状进行了分析。结果表明，丸叶×Coker319组合的总叶绿素含量受1对加性-显性主基因+加性-显性-上位性多基因控制，主基因加性效应为-5.89，主基因显性效应为-3.47；B1、B2和F2世代总叶绿素含量的主基因遗传率分别为3.61%、46.11%和48.94%；多基因遗传率分别为52.04%、8.25%和0.00%，说明F2世代总叶绿素含量表现出较高的主基因遗传率，并受环境影响。对烤烟总叶绿素含量的改良要以主基因为主，同时注意环境的影响。光合速率和蒸腾速率受加性-显性-上位性多基因控制，多基因遗传率分别为20.69%和13.56%。气孔导度受2对加性-显性-上位性主基因+加性-显性-上位性多基因控制，以显性×显性上位性效应为主，B1、B2和F2世代气孔导度的主基因遗传率分别为58.03%、35.11%和40.59%；多基因遗传率分别为1.43%、21.34%和2.03%。
Mini core collection is a core of the core collection and a representative subset of core collection,consisted of1imited accessions with minimum genetic redundantcy and retained most genetic diversityand genetic structure in initial collection. The ultimate goal of constructing mini core collection of to-bacco is to study its genetic diversity, promote its effective management, deeply evaluation and makingfull use of, thus it will meet the needs of production and scientific research to the greatest degree. Accu-rate measure in genetic similarity among accessions and specifying an appropriate samlpling stratey is akey step in developing a mini core collection. This study will focus on evaluating the properties of dif-ferent integrating ratio between molecular marker and quantitative trait and different sampling strategiesby comparing the genetic variation captured by subsets of tobacco, setting up a mini core collection oftobacco. Two accessions of flue-cured tobacco belong to mini core collection were screened to sudy theinheritance of important botany and physiological traits with the joint segregation analysis method ofmixed major gene plus polygene genetic mode, so as to provide some theoretical guidance for the quan-titative genetic breeding of flue-cured tobacco. The main conclusions of this study were as follows:
(1)The conception of core primer was defined.12different tobacco accessions was used to screen700SSR primer pairs which were equally distributed in24linkage groups of tobacco, and finally54core primer pairs fitting for tobacco DNA fingerprint pool were screened. Here we publish54core pri-mer pairs and their polymorphic information, which will be helpful to regulate the procedure and pro-mote the efficiency for screening SSR primers, and will play an important role in tobacco genetic diver-sity evaluation, core collection construction, fingerprint construction, seed genuineness and purity iden-tification of the tobacco varieties.
(2)446accessions of tobacco core collection which were constrcted in2001were characterized by23agronomic traits and molecular markers on54SSR core primers to establish a mini core collection.Seven different sampling ratios of accessions were tested, including5%to40%with an interval of5%.The results showed that15%and30%of samples could conserve more than90%and80%of variationsin N.tabacum and N.rustica respectively, which indicated that15%and30%was an adequate samplingratio for establishing the mini core collection in N.tabacum and N.rustica respectively. The intergratingdata between quantitative and molecular data is better than each other, and the proportion of quantitativeand molecular data for0.1:0.9was optimal. A comparision of the efficiencies of two genetic distance(Euclidean distance and Mahalanobis distance) for the continuous data and foue genetic distance (Sim-ple matching, Jaccard, Nei-Li and Principal component distance), showed that Euclidean distance andJaccard were optimal genetic distance. A comparison of the efficiencies of four sampling strategies bygeographical grouping (proportional, square root, logarithmic, and genetic diversity-depend) and threesampling mehtods (random sampling, preferred sampling and maximization), indicated proportional andlogarithmic strategy was optimal in term of establishing the most representative mini core collection,when the quantity of samples to be located is significant, the former is better, whereas the latter is better. Thus, a mini core collection of127accessions, including117selected based on proportional and loga-rithmic strategy and10accessions selected by their specific agro-morphological traits, was constructed.The tests on SSR data and agro-morphological characters demonstrated that the mini core collection hashigh genetic diversity and a good representative to the entire collection.
(3)The joint segregation analysis method of mixed major gene plus polygene genetic model wasused to study the inheritance of important botanical traits in flue-cured tobacco. Six generations（P1, P2,F1, B1, B2and F2） from the crosses （Wanye×Coker319） were investigated. It was found that plantheight, leaf number, leaf area and fresh leaf weight in flue-cured tobacco appeared to be a quantitativetrait and their inheritances fitted to a mixed genetic model of two major genes with addi-tive-dominant-epistatic effects plus polygenes with additive-dominant-epistatic effects （E0model）.Plant height and leaf number was chiefly controlled by the additive effect and the epistatic effect of do-minance×dominance. The additive, dominant and epistatic effects which the epistatic effect> the addi-tive effect> the dominance effect of leaf area and fresh leaf weight were all important. Heritabilities ofthe major genes were estimated to be57.53%,42.63%,30.32%and44.26%in F2. The inheritance ofdays of transplanting to flowering fitted to a mixed genetic model of two major genes with addi-tive-dominant-epistatic effects plus polygenes with additive-dominant effects （E1model） and it wasmainly dominated by the epistatic effect of additive×additive, the additive effect, and the epistatic ef-fect of dominance×dominance, the heritability of the major gene was64.79%in F2. The inheritance ofstem girth and leaf mass per area fitted to a mixed genetic model of one major gene with complete do-minant effects plus polygenes with additive-dominant effects （D3model）, stem girth was principallycontrolled by polygenes which additive effect was equal to its dominance effect, the heritability of themajor gene was2.48%and38.71%in F2. The inheritance of leaf index fitted to a mixed genetic modelof one major gene with additive-dominant effects plus polygenes with additive-dominant-epistatic ef-fects （D0model）, the additive effect and the dominance effect of the major gene was almost equal, theheritability of the major gene was49.64%in F2. Leaf length, leaf width, internodal distance and capsuleweight fitted to a mixed genetic model of polygenes with additive-dominant-epistatic effects （C0model）, the heritability of the polygenes were60.75%,62.14%,75.08%and82.34%in F2.
(4)The mixed major-gene plus polygene inheritance model in six generations （P1, P2, F1, B1, B2,and F2）was used to analyze the inheritance of several physiological traits in the flue-cured tobacco（N.tabacum） cross between WANYE and Coker319. The results showed that the SPAD reading wascontrolled by one gene with additive-dominance effect plus polygenes with addi-tive-dominance-epistatic effect （D0model）. The additive effect for major gene was-5.89, the do-minant effect for major gene was-3.47. The major gene heritabilities in B1, B2and F2were3.61%,46.11%and48.94%, respectively, and the corresponding polygenic heritabilities were52.04%,8.25%and0.00%, respectively. These results indicated that major gene in F2was a key factor and environment fac-tor was also relatively important. This implies that in the genetic improvement of SPAD reading ma-jor-gene is a main factor whereas environmental effect should be taken care of. Photosynthetic rate andtranspiration rate fitted to a mixed genetic model of polygenes with additive-dominant-epistatic effects （C0model）, the heritability of the polygenes were20.69%and13.56%in F2. It was also found thatstomatal conductance in flue-cured tobacco appeared to be a quantitative trait and their inheritances fit-ted to a mixed genetic model of two major genes with additive-dominant-epistatic effects plus poly-genes with additive-dominant-epistatic effects （E0model）. Stomatal conductance was chiefly con-trolled by the epistatic effect of dominance×dominance. Heritabilities of the major genes were esti-mated to be58.03%,35.11%and40.59%in B1, B2and F2, and the heritabilities of the polygenes wereestimated to be1.43%,21.34%and2.03%in B1, B2and F2.
（1）通过12份不同烟草种质类型，筛选均匀分布于烟草24条连锁群上的700对SSR标记，综合考虑PIC值、扩增带型统计的难易及引物的重复性，获得54对SSR核心引物，建立了核心引物数据库。本研究有利于规范引物筛选程序、提高引物筛选效率，对烟草种质资源的遗传多样性分析、核心种质构建、指纹图谱构建以及种子真伪性和纯度的鉴定等都具有现实意义。
（2）以我国2001年构建的446份烟草核心种质为材料，根据23个数量性状资料和54对SSR引物的分子标记数据，开展微核心种质构建研究。不同取样比例（40%、30%、25%、20%、15%、10%和5%）分析表明，15%的取样比例可获得普通烟草90%以上的多态位点百分率，30%的取样比例可获得黄花烟草80%以上的多态位点百分率，是较好的微核心种质取样规模；表型数据和分子数据整合后的数据要比不整合好，且表型数据与分子数据的比例为0.1:0.9时，整合效果最好。对于连续型数据遗传距离估算的2种距离（欧氏距离和马氏距离）中，欧氏距离效果最优；对于离散型数据遗传距离估算的4种距离（Simple matching、Jaccard、Nei-Li和主成分距离）中，Jaccard距离效果最优。对4种分组取样策略（简单比例法、平方根比例法、对数比例法和多样性比例法）和3种取样方法（随机取样法、优先取样法和变异度取样法）比较表明，总体取样方法中变异度取样法最优，在组内样本含量较大时，简单比例法和多样性比例法获得的微核心种质代表性最好，为最优取样策略；而当组内样本含量较小时，对数比例法获得的微核心种质代表性最好，为最优取样策略。最后，在简单比例法和对数比例法取样筛选出的117份核心样品中，又通过定向选择补充了10份具有优异表型性状的材料，构建了含127份烟草的微核心种质。分子和表型检验都表明本研究所构建的微核心种质具有较好的代表性。
（3）利用“主基因+多基因”混合遗传模型的6个世代联合分离分析方法，分析烤烟组合丸叶×Coker319几个重要植物学性状的遗传效应。结果表明，烤烟的株高、叶数、叶面积和鲜叶重受2对加性-显性-上位性主基因+加性-显性-上位性多基因控制，株高与叶数遗传以加性效应及显性×显性上位性效应为主，叶面积和鲜叶重各遗传效应相差不多，其上位性效应>加性效应>显性效应，F2世代的主基因遗传率分别为57.53%、42.63%、30.32%和44.26%。移栽至中心花开放天数受2对加性-显性-上位性主基因+加性-显性多基因控制，以加性×加性上位性效应、加性效应及显性×显性上位性效应为主，主基因遗传率为64.79%。茎围和比叶重均受1对完全显性主基因+加性-显性多基因控制，茎围遗传以多基因为主，其多基因加性效应和显性效应大小相当，比叶重遗传主基因、多基因的加性效应和显性效应大致相当，主基因遗传率分别为2.48%和38.71%。叶形指数受1对加性-显性主基因+加性-显性-上位性多基因控制，主基因加性效应与显性效应基本相当，主基因遗传率为49.64%。叶长、叶宽、节距和蒴果重受加性-显性-上位性多基因控制，多基因遗传率分别为60.75%、62.14%、75.08%和82.34%。
（4）应用主基因+多基因6个世代联合分析方法对烤烟丸叶×Coker319组合的几个生理性状进行了分析。结果表明，丸叶×Coker319组合的总叶绿素含量受1对加性-显性主基因+加性-显性-上位性多基因控制，主基因加性效应为-5.89，主基因显性效应为-3.47；B1、B2和F2世代总叶绿素含量的主基因遗传率分别为3.61%、46.11%和48.94%；多基因遗传率分别为52.04%、8.25%和0.00%，说明F2世代总叶绿素含量表现出较高的主基因遗传率，并受环境影响。对烤烟总叶绿素含量的改良要以主基因为主，同时注意环境的影响。光合速率和蒸腾速率受加性-显性-上位性多基因控制，多基因遗传率分别为20.69%和13.56%。气孔导度受2对加性-显性-上位性主基因+加性-显性-上位性多基因控制，以显性×显性上位性效应为主，B1、B2和F2世代气孔导度的主基因遗传率分别为58.03%、35.11%和40.59%；多基因遗传率分别为1.43%、21.34%和2.03%。
Mini core collection is a core of the core collection and a representative subset of core collection,consisted of1imited accessions with minimum genetic redundantcy and retained most genetic diversityand genetic structure in initial collection. The ultimate goal of constructing mini core collection of to-bacco is to study its genetic diversity, promote its effective management, deeply evaluation and makingfull use of, thus it will meet the needs of production and scientific research to the greatest degree. Accu-rate measure in genetic similarity among accessions and specifying an appropriate samlpling stratey is akey step in developing a mini core collection. This study will focus on evaluating the properties of dif-ferent integrating ratio between molecular marker and quantitative trait and different sampling strategiesby comparing the genetic variation captured by subsets of tobacco, setting up a mini core collection oftobacco. Two accessions of flue-cured tobacco belong to mini core collection were screened to sudy theinheritance of important botany and physiological traits with the joint segregation analysis method ofmixed major gene plus polygene genetic mode, so as to provide some theoretical guidance for the quan-titative genetic breeding of flue-cured tobacco. The main conclusions of this study were as follows:
(1)The conception of core primer was defined.12different tobacco accessions was used to screen700SSR primer pairs which were equally distributed in24linkage groups of tobacco, and finally54core primer pairs fitting for tobacco DNA fingerprint pool were screened. Here we publish54core pri-mer pairs and their polymorphic information, which will be helpful to regulate the procedure and pro-mote the efficiency for screening SSR primers, and will play an important role in tobacco genetic diver-sity evaluation, core collection construction, fingerprint construction, seed genuineness and purity iden-tification of the tobacco varieties.
(2)446accessions of tobacco core collection which were constrcted in2001were characterized by23agronomic traits and molecular markers on54SSR core primers to establish a mini core collection.Seven different sampling ratios of accessions were tested, including5%to40%with an interval of5%.The results showed that15%and30%of samples could conserve more than90%and80%of variationsin N.tabacum and N.rustica respectively, which indicated that15%and30%was an adequate samplingratio for establishing the mini core collection in N.tabacum and N.rustica respectively. The intergratingdata between quantitative and molecular data is better than each other, and the proportion of quantitativeand molecular data for0.1:0.9was optimal. A comparision of the efficiencies of two genetic distance(Euclidean distance and Mahalanobis distance) for the continuous data and foue genetic distance (Sim-ple matching, Jaccard, Nei-Li and Principal component distance), showed that Euclidean distance andJaccard were optimal genetic distance. A comparison of the efficiencies of four sampling strategies bygeographical grouping (proportional, square root, logarithmic, and genetic diversity-depend) and threesampling mehtods (random sampling, preferred sampling and maximization), indicated proportional andlogarithmic strategy was optimal in term of establishing the most representative mini core collection,when the quantity of samples to be located is significant, the former is better, whereas the latter is better. Thus, a mini core collection of127accessions, including117selected based on proportional and loga-rithmic strategy and10accessions selected by their specific agro-morphological traits, was constructed.The tests on SSR data and agro-morphological characters demonstrated that the mini core collection hashigh genetic diversity and a good representative to the entire collection.
(3)The joint segregation analysis method of mixed major gene plus polygene genetic model wasused to study the inheritance of important botanical traits in flue-cured tobacco. Six generations（P1, P2,F1, B1, B2and F2） from the crosses （Wanye×Coker319） were investigated. It was found that plantheight, leaf number, leaf area and fresh leaf weight in flue-cured tobacco appeared to be a quantitativetrait and their inheritances fitted to a mixed genetic model of two major genes with addi-tive-dominant-epistatic effects plus polygenes with additive-dominant-epistatic effects （E0model）.Plant height and leaf number was chiefly controlled by the additive effect and the epistatic effect of do-minance×dominance. The additive, dominant and epistatic effects which the epistatic effect> the addi-tive effect> the dominance effect of leaf area and fresh leaf weight were all important. Heritabilities ofthe major genes were estimated to be57.53%,42.63%,30.32%and44.26%in F2. The inheritance ofdays of transplanting to flowering fitted to a mixed genetic model of two major genes with addi-tive-dominant-epistatic effects plus polygenes with additive-dominant effects （E1model） and it wasmainly dominated by the epistatic effect of additive×additive, the additive effect, and the epistatic ef-fect of dominance×dominance, the heritability of the major gene was64.79%in F2. The inheritance ofstem girth and leaf mass per area fitted to a mixed genetic model of one major gene with complete do-minant effects plus polygenes with additive-dominant effects （D3model）, stem girth was principallycontrolled by polygenes which additive effect was equal to its dominance effect, the heritability of themajor gene was2.48%and38.71%in F2. The inheritance of leaf index fitted to a mixed genetic modelof one major gene with additive-dominant effects plus polygenes with additive-dominant-epistatic ef-fects （D0model）, the additive effect and the dominance effect of the major gene was almost equal, theheritability of the major gene was49.64%in F2. Leaf length, leaf width, internodal distance and capsuleweight fitted to a mixed genetic model of polygenes with additive-dominant-epistatic effects （C0model）, the heritability of the polygenes were60.75%,62.14%,75.08%and82.34%in F2.
(4)The mixed major-gene plus polygene inheritance model in six generations （P1, P2, F1, B1, B2,and F2）was used to analyze the inheritance of several physiological traits in the flue-cured tobacco（N.tabacum） cross between WANYE and Coker319. The results showed that the SPAD reading wascontrolled by one gene with additive-dominance effect plus polygenes with addi-tive-dominance-epistatic effect （D0model）. The additive effect for major gene was-5.89, the do-minant effect for major gene was-3.47. The major gene heritabilities in B1, B2and F2were3.61%,46.11%and48.94%, respectively, and the corresponding polygenic heritabilities were52.04%,8.25%and0.00%, respectively. These results indicated that major gene in F2was a key factor and environment fac-tor was also relatively important. This implies that in the genetic improvement of SPAD reading ma-jor-gene is a main factor whereas environmental effect should be taken care of. Photosynthetic rate andtranspiration rate fitted to a mixed genetic model of polygenes with additive-dominant-epistatic effects （C0model）, the heritability of the polygenes were20.69%and13.56%in F2. It was also found thatstomatal conductance in flue-cured tobacco appeared to be a quantitative trait and their inheritances fit-ted to a mixed genetic model of two major genes with additive-dominant-epistatic effects plus poly-genes with additive-dominant-epistatic effects （E0model）. Stomatal conductance was chiefly con-trolled by the epistatic effect of dominance×dominance. Heritabilities of the major genes were esti-mated to be58.03%,35.11%and40.59%in B1, B2and F2, and the heritabilities of the polygenes wereestimated to be1.43%,21.34%and2.03%in B1, B2and F2.
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