鹅掌楸杂种优势分子机理研究
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摘要
鹅掌楸种间杂交具有明显的杂种优势，从DNA分子水平研究鹅掌楸杂种优势的遗传基础具有重要意义。本文以江西景德镇、南京江浦2个试验点3-7a生的杂交鹅掌楸子代测定林为材料，在分析子代幼林期生长性状遗传变异及杂种优势表现的基础上，利用SSR分子标记检测鹅掌楸交配亲本的遗传距离以及杂交组合子代的一般杂合度和特殊杂合度，进而分析鹅掌楸生长性状的杂种优势与亲本遗传距离及子代杂合度之间的相关性。同时，利用Tassel3.0软件对景德镇试验点鹅掌楸杂种子代7a生长性状与101个SSR位点进行关联分析，寻找与生长性状相关联的DNA分子标记，并对相关联分子标记所对应的EST序列进行生物信息学分析。主要研究结果如下：
鹅掌楸杂种子代生长表现与杂种优势分析。对景德镇试验点16个杂交组合3-7a的生长量进行了分析，发现树高和胸径在不同组合间差异较大。树高与胸径生长最快的两个组合分别是S×M、H×M；生长最慢的两个组合分别是L×C2、N1×H。方差分析结果表明，树高及胸径年生长量在不同林龄及不同杂交组合间均达到了极显著差异。对鹅掌楸杂交子代生长性状杂种优势分析发现，并非所有的杂种子代都表现出杂种优势。其中，12个杂交组合表现出正向杂种优势，其余4个杂交组合表现出负向杂种优势。方差分析结果表明，树高、胸径杂种优势在不同组合间差异均达到了极显著水平（P=0.0001），但树高超高亲优势、树高超中亲优势及胸径超高亲优势在在不同林龄间差异不明显，未达到显著性水平（P=0.4432，P=0.0819，P=0.1615），仅胸径超中亲优势在不同林龄间差异达到显著性水平（P=0.002）；。
鹅掌楸交配亲本遗传距离与杂种优势相关性。利用30个SSR分子标记分析了景德镇试验点16个杂交组合亲本间的遗传距离。各交配组合亲本间遗传距离存在较大的遗传差异，变幅为0.117～1.4。将子代杂种优势4个度量指标分别与亲本遗传距离进行相关性分析，发现二者存在相关性，但均未达到显著性水平（P=0.9504，P=0.9823，P=0.6422，P=0.4591）。进一步分析发现，当亲本间遗传距离<1.0时，随着遗传距离增加，子代杂种优势逐渐增加。在亲本遗传距离为1.0左右时，子代杂种优势达最大值。当遗传距离>1.0时，随着遗传距离增大，杂种优势反而下降。为了验证上述结论，用同一批SSR分子标记对定植于南京江浦的另一批鹅掌楸杂交组合材料进行分析，所得结论与此相似。因此，可以认为，在进行杂交育种亲本选配时，亲本对之间最适的遗传距离应为1.0左右。
鹅掌楸杂交子代群体杂合度与杂种优势相关性。利用同一批SSR分子标记分析了景德镇试验点16个杂交组合子代的一般杂合度。杂交组合子代具有较高的观测杂合度（0.6931～0.8278）。对杂种子代杂种优势表现与一般杂合度进行相关性分析，发现二者相关性均未达到显著性水平（P=0.8252，P=0.8846，P=0.7513，P=0.7286）。利用关联分析结果对杂种子代进行特殊杂合度分析，发现杂交组合子代特殊杂合度变化幅度较大（0.3741～0.8714）。对杂种子代杂种优势表现与特殊杂合度进行相关性分析，发现二者相关性也均未达到显著性水平（P=0.5694，P=0.716，P=0.447， P=0.5389），说明杂交子代杂合度可能并非鹅掌楸杂种优势形成的主要原因。
杂交鹅掌楸生长性状与SSR分子标记关联分析。利用Tassel3.0软件对101对SSR标记间的连锁不平衡（LD）进行分析，发现有40对位点间存在较强的连锁不平衡。利用筛选后的61个标记通过Structure2.3软件对景德镇试验点16个杂交组合进行群体结构分析，得出最佳模拟分群数目为8（K值）。通过计算不同杂交组合子代归入各亚群的概率(Q值)，对关联分析的结果进行矫正，再利用Tassel3.0软件对景德镇杂交组合7a生长量性状与101对SSR标记进行关联分析。发现在P＜0.01显著性水平下，共有6个标记与树高性状相关联，7个标记与胸径性状关联。而且，关联的标记中，除691号位点未与树高性状关联外，其余关联标记与两个性状均存在极显著相关，说明两个性状之间可能存在较高的相关性。进一步对树高与胸径相关性研究发现，二者相关系数r=0.9438，P=0.0001。
与鹅掌楸生长性状关联分子标记的生物信息学分析。基于上述关联分析结果，对每一个与生长性状相关联的SSR标记，从鹅掌楸EST数据库找出与其相对应的EST序列，登入NCBI网站进行序列比对。7个关联位点中，有4个可以搜索到与其EST相匹配的序列，并在其它物种中做过相关研究。另外3个EST序列在数据库中未找到与其相关序列。其中2号位点对应的EST序列在蓖麻研究中相关序列是有关叶绿体的分子伴侣（铜伴侣）；15号位点对应的EST序列在拟南芥研究中相关序列是有关核内小核糖核蛋白输入蛋白的；542号位点对应的EST序列在毛果杨研究中相关序列是有关组蛋白H1的；642号位点对应的EST序列在拟南芥研究中相关序列是有关顶端生长缺陷蛋白的。分析发现TIP1是促使顶端生长和维持细胞生长所必需的。
遗传多样性度量与SSR标记数量的关系。从实验室规模化开发的1000多对鹅掌楸属EST-SSR引物中筛选出100对多态性较好引物，以景德镇试验点16个鹅掌楸种间杂交组合为材料，以Na、Ne、I、Ho、Nei5个遗传多样性度量指标。以多态性为指标对SSR标记位点依次排序，5个位点为一组，分析SSR分子标记数量与遗传多样性度量的关系。研究发现，随着检测的标记位点数增加，估算的群体遗传多样性随之降低；但当标记位点数达到20个左右时，群体遗传多样性趋于稳定，表明在利用多态性较高的分子标记进行群体遗传多样性检测时，标记数量以20个左右为宜。
It has been proved that an obvious heterosis occurred in interspecific hybrid ofLiriodendron, while its genetic basis has not been clear yet. In this paper, the relationshipbetween the heterosis of growth and the genetic distance among mating parents and theheterozygosity of hybrids was explored based on the evaluation on both growth traits and SSRmarkers. Two batch of crossing combinations were taken as experimental populations, onecontains16interspecific hybrid combinations and their parental half-sibs and was establishedin2006at Jingdezhen, the other covers17interspecific hybrid combinations,7intraspecifichybrid combinations and their parental half-sibs in2010at Nanjing. Furthermore, theassociation between the growth traits of hybrid Liriodendron and SSR markers were analyzed.Meanwhile, the relationship between the number of SSR markers and the measurement ongenetic diversity was also explored. The main results are as follows:
There were significant variation existing among combinations when analysing3-7aheight and DBH of16hybrid combinations at Jingdezhen trial site. Two combinations withfastest growth are S×M and H×M; while two combinations with slowest growth are L×C2andN1×H. Analysis of variance revealed significant variations of annual growth existing amongboth tree ages and combinations. Relatively large difference of heterosis among hybridcombinations was found by comparing the growth of hybrids with parental half-sibs. Not allthe hybrids showed heterosis,12hybrid combinations showed positive heterosis and4hybridcombinations showed negative heterosis. According to the analysis of variance, heterosischange on DBH and heigh was clear among mating combinations that the results reach ahighly significant leve（lP=0.0001，P=0.0001),while heterosis change on height and over BPHfor DBH of mating combinations was not clear among forest ages that the results did not reacha significant level（P=0.4432,P=0.0819,P=0.1615), however, heterosis change on over MPHfor DBH of mating combinations was clear among forest ages that the results reach a highlysignificant level（P=0.002).
Based on the analysis of the parental genetic distance, relatively large genetic variationsamong parents were found with variation range of0.117～1.4at Jingdezhen. According toresults of correlation analysis, significant correlations were not found between the geneticdistance and the heterosis of growth（P=0.9504,P=0.9823,P=0.6422，P=0.4591).Furtherstudies showed that heterosis reached the maximum when the genetic distance was about1,heterosis showed downward trend with further increase of genetic distance. In order to verifythe above conclusions, the correlation of genetic distance and heterosis was analyzed with thesame molecular marker at Nanjing Qiaolin trial site,and similar conclusion was gotten,Parental genetic distance1could be used as indicator of parent selection.
The general heterozygosity of hybrids were analyzed with the same molecular marker atJing dezhen, the range of the general heterozygosity of hybrids was0.6931～0.8278.The correlation of the general heterozygosity and heterosis did not reach a significant level（P=0.8252,P=0.8846,P=0.7513,P=0.7286);The special heterozygosity of hybrids wereanalyzed according to the results of association analysis, the range of the specialheterozygosity of hybrids was0.3741～0.8714. Furthermore, the correlation was analyzedbetween special heterozygosity of hybrids and heterosis,the results did not reach a significantlevel too（P=0.5694,P=0.716,P=0.447,P=0.5389),It can be deduced that heterozygosity ofhybrids is probably not the main reason for the formation of Liriodendron heterosis.
LD among101pairs of markers was analyzed using Tassel3.0Software,we found thatstrong linkage disequilibrium between the40pairs of loci. Population structure of16hybridcombinations was also explored based on the results of LD screening, we found that8was thethe best analog clustering number，and calculated the probability of hybrid combinationoffspring which attributed to the subsets. Furthermore, association analysis was investigatedbetween7-year-old growth traits of16hybrid combinations at Jing dezhen and101pairs ofmolecular markers,the result showed that6markers associated with height trait and7markersassociated with DBH trait when the correlation between markers and traits was highlysignificant (P＜0.01＝, and the associated markers were consistent in addition to the691sitedid not associate with height trait, Further speculated that there may be a high correlationbetween the two traits, Further studies showed that the correlation of height and DBH reacheda highly significant level (r=0.9438, P=0.0001).
EST sequences which corresponded to the associated markers were identified accordingto the correlation results，the result of sequence alignment showed that EST sequences of the4sites matched with sequences which came from NCBI，EST sequences of the3sites were notfound in the database，EST sequence of the2ndsite corresponded to sequence of chloroplastchaperone (copper companion) in Ricinus communis；EST sequence of the fifteenth No. sitecorresponded to sequence of snurportin-1in Arabidopsis thaliana；EST sequence of No.542site corresponded to sequence of histone H1in Populus trichocarpa；EST sequence of No.642site corresponded to sequence of tip growth defective in Arabidopsis thaliana; Further studiesshowed that TIP1was necessary to promote tip growth and cell growth,however, the aboverelated genes function needed to be verified in Liriodendron.
The relationship between the number of SSR markers and the measurement on geneticdiversity was explored，SSR sites were rearranged in accordance with the number of allelesand effective alleles，considered five sites as a gradient based on the order, the metrics ofgenetic diversity (Na, Ne, I, Ho, Nei) were selected to analyze genetic diversity of hybridcombinations. Further studies showed that the genetic diversity of hybrid combinations wasstable when the number of loci was about20.
鹅掌楸杂种子代生长表现与杂种优势分析。对景德镇试验点16个杂交组合3-7a的生长量进行了分析，发现树高和胸径在不同组合间差异较大。树高与胸径生长最快的两个组合分别是S×M、H×M；生长最慢的两个组合分别是L×C2、N1×H。方差分析结果表明，树高及胸径年生长量在不同林龄及不同杂交组合间均达到了极显著差异。对鹅掌楸杂交子代生长性状杂种优势分析发现，并非所有的杂种子代都表现出杂种优势。其中，12个杂交组合表现出正向杂种优势，其余4个杂交组合表现出负向杂种优势。方差分析结果表明，树高、胸径杂种优势在不同组合间差异均达到了极显著水平（P=0.0001），但树高超高亲优势、树高超中亲优势及胸径超高亲优势在在不同林龄间差异不明显，未达到显著性水平（P=0.4432，P=0.0819，P=0.1615），仅胸径超中亲优势在不同林龄间差异达到显著性水平（P=0.002）；。
鹅掌楸交配亲本遗传距离与杂种优势相关性。利用30个SSR分子标记分析了景德镇试验点16个杂交组合亲本间的遗传距离。各交配组合亲本间遗传距离存在较大的遗传差异，变幅为0.117～1.4。将子代杂种优势4个度量指标分别与亲本遗传距离进行相关性分析，发现二者存在相关性，但均未达到显著性水平（P=0.9504，P=0.9823，P=0.6422，P=0.4591）。进一步分析发现，当亲本间遗传距离<1.0时，随着遗传距离增加，子代杂种优势逐渐增加。在亲本遗传距离为1.0左右时，子代杂种优势达最大值。当遗传距离>1.0时，随着遗传距离增大，杂种优势反而下降。为了验证上述结论，用同一批SSR分子标记对定植于南京江浦的另一批鹅掌楸杂交组合材料进行分析，所得结论与此相似。因此，可以认为，在进行杂交育种亲本选配时，亲本对之间最适的遗传距离应为1.0左右。
鹅掌楸杂交子代群体杂合度与杂种优势相关性。利用同一批SSR分子标记分析了景德镇试验点16个杂交组合子代的一般杂合度。杂交组合子代具有较高的观测杂合度（0.6931～0.8278）。对杂种子代杂种优势表现与一般杂合度进行相关性分析，发现二者相关性均未达到显著性水平（P=0.8252，P=0.8846，P=0.7513，P=0.7286）。利用关联分析结果对杂种子代进行特殊杂合度分析，发现杂交组合子代特殊杂合度变化幅度较大（0.3741～0.8714）。对杂种子代杂种优势表现与特殊杂合度进行相关性分析，发现二者相关性也均未达到显著性水平（P=0.5694，P=0.716，P=0.447， P=0.5389），说明杂交子代杂合度可能并非鹅掌楸杂种优势形成的主要原因。
杂交鹅掌楸生长性状与SSR分子标记关联分析。利用Tassel3.0软件对101对SSR标记间的连锁不平衡（LD）进行分析，发现有40对位点间存在较强的连锁不平衡。利用筛选后的61个标记通过Structure2.3软件对景德镇试验点16个杂交组合进行群体结构分析，得出最佳模拟分群数目为8（K值）。通过计算不同杂交组合子代归入各亚群的概率(Q值)，对关联分析的结果进行矫正，再利用Tassel3.0软件对景德镇杂交组合7a生长量性状与101对SSR标记进行关联分析。发现在P＜0.01显著性水平下，共有6个标记与树高性状相关联，7个标记与胸径性状关联。而且，关联的标记中，除691号位点未与树高性状关联外，其余关联标记与两个性状均存在极显著相关，说明两个性状之间可能存在较高的相关性。进一步对树高与胸径相关性研究发现，二者相关系数r=0.9438，P=0.0001。
与鹅掌楸生长性状关联分子标记的生物信息学分析。基于上述关联分析结果，对每一个与生长性状相关联的SSR标记，从鹅掌楸EST数据库找出与其相对应的EST序列，登入NCBI网站进行序列比对。7个关联位点中，有4个可以搜索到与其EST相匹配的序列，并在其它物种中做过相关研究。另外3个EST序列在数据库中未找到与其相关序列。其中2号位点对应的EST序列在蓖麻研究中相关序列是有关叶绿体的分子伴侣（铜伴侣）；15号位点对应的EST序列在拟南芥研究中相关序列是有关核内小核糖核蛋白输入蛋白的；542号位点对应的EST序列在毛果杨研究中相关序列是有关组蛋白H1的；642号位点对应的EST序列在拟南芥研究中相关序列是有关顶端生长缺陷蛋白的。分析发现TIP1是促使顶端生长和维持细胞生长所必需的。
遗传多样性度量与SSR标记数量的关系。从实验室规模化开发的1000多对鹅掌楸属EST-SSR引物中筛选出100对多态性较好引物，以景德镇试验点16个鹅掌楸种间杂交组合为材料，以Na、Ne、I、Ho、Nei5个遗传多样性度量指标。以多态性为指标对SSR标记位点依次排序，5个位点为一组，分析SSR分子标记数量与遗传多样性度量的关系。研究发现，随着检测的标记位点数增加，估算的群体遗传多样性随之降低；但当标记位点数达到20个左右时，群体遗传多样性趋于稳定，表明在利用多态性较高的分子标记进行群体遗传多样性检测时，标记数量以20个左右为宜。
It has been proved that an obvious heterosis occurred in interspecific hybrid ofLiriodendron, while its genetic basis has not been clear yet. In this paper, the relationshipbetween the heterosis of growth and the genetic distance among mating parents and theheterozygosity of hybrids was explored based on the evaluation on both growth traits and SSRmarkers. Two batch of crossing combinations were taken as experimental populations, onecontains16interspecific hybrid combinations and their parental half-sibs and was establishedin2006at Jingdezhen, the other covers17interspecific hybrid combinations,7intraspecifichybrid combinations and their parental half-sibs in2010at Nanjing. Furthermore, theassociation between the growth traits of hybrid Liriodendron and SSR markers were analyzed.Meanwhile, the relationship between the number of SSR markers and the measurement ongenetic diversity was also explored. The main results are as follows:
There were significant variation existing among combinations when analysing3-7aheight and DBH of16hybrid combinations at Jingdezhen trial site. Two combinations withfastest growth are S×M and H×M; while two combinations with slowest growth are L×C2andN1×H. Analysis of variance revealed significant variations of annual growth existing amongboth tree ages and combinations. Relatively large difference of heterosis among hybridcombinations was found by comparing the growth of hybrids with parental half-sibs. Not allthe hybrids showed heterosis,12hybrid combinations showed positive heterosis and4hybridcombinations showed negative heterosis. According to the analysis of variance, heterosischange on DBH and heigh was clear among mating combinations that the results reach ahighly significant leve（lP=0.0001，P=0.0001),while heterosis change on height and over BPHfor DBH of mating combinations was not clear among forest ages that the results did not reacha significant level（P=0.4432,P=0.0819,P=0.1615), however, heterosis change on over MPHfor DBH of mating combinations was clear among forest ages that the results reach a highlysignificant level（P=0.002).
Based on the analysis of the parental genetic distance, relatively large genetic variationsamong parents were found with variation range of0.117～1.4at Jingdezhen. According toresults of correlation analysis, significant correlations were not found between the geneticdistance and the heterosis of growth（P=0.9504,P=0.9823,P=0.6422，P=0.4591).Furtherstudies showed that heterosis reached the maximum when the genetic distance was about1,heterosis showed downward trend with further increase of genetic distance. In order to verifythe above conclusions, the correlation of genetic distance and heterosis was analyzed with thesame molecular marker at Nanjing Qiaolin trial site,and similar conclusion was gotten,Parental genetic distance1could be used as indicator of parent selection.
The general heterozygosity of hybrids were analyzed with the same molecular marker atJing dezhen, the range of the general heterozygosity of hybrids was0.6931～0.8278.The correlation of the general heterozygosity and heterosis did not reach a significant level（P=0.8252,P=0.8846,P=0.7513,P=0.7286);The special heterozygosity of hybrids wereanalyzed according to the results of association analysis, the range of the specialheterozygosity of hybrids was0.3741～0.8714. Furthermore, the correlation was analyzedbetween special heterozygosity of hybrids and heterosis,the results did not reach a significantlevel too（P=0.5694,P=0.716,P=0.447,P=0.5389),It can be deduced that heterozygosity ofhybrids is probably not the main reason for the formation of Liriodendron heterosis.
LD among101pairs of markers was analyzed using Tassel3.0Software,we found thatstrong linkage disequilibrium between the40pairs of loci. Population structure of16hybridcombinations was also explored based on the results of LD screening, we found that8was thethe best analog clustering number，and calculated the probability of hybrid combinationoffspring which attributed to the subsets. Furthermore, association analysis was investigatedbetween7-year-old growth traits of16hybrid combinations at Jing dezhen and101pairs ofmolecular markers,the result showed that6markers associated with height trait and7markersassociated with DBH trait when the correlation between markers and traits was highlysignificant (P＜0.01＝, and the associated markers were consistent in addition to the691sitedid not associate with height trait, Further speculated that there may be a high correlationbetween the two traits, Further studies showed that the correlation of height and DBH reacheda highly significant level (r=0.9438, P=0.0001).
EST sequences which corresponded to the associated markers were identified accordingto the correlation results，the result of sequence alignment showed that EST sequences of the4sites matched with sequences which came from NCBI，EST sequences of the3sites were notfound in the database，EST sequence of the2ndsite corresponded to sequence of chloroplastchaperone (copper companion) in Ricinus communis；EST sequence of the fifteenth No. sitecorresponded to sequence of snurportin-1in Arabidopsis thaliana；EST sequence of No.542site corresponded to sequence of histone H1in Populus trichocarpa；EST sequence of No.642site corresponded to sequence of tip growth defective in Arabidopsis thaliana; Further studiesshowed that TIP1was necessary to promote tip growth and cell growth,however, the aboverelated genes function needed to be verified in Liriodendron.
The relationship between the number of SSR markers and the measurement on geneticdiversity was explored，SSR sites were rearranged in accordance with the number of allelesand effective alleles，considered five sites as a gradient based on the order, the metrics ofgenetic diversity (Na, Ne, I, Ho, Nei) were selected to analyze genetic diversity of hybridcombinations. Further studies showed that the genetic diversity of hybrid combinations wasstable when the number of loci was about20.
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