红榉种源变异
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摘要
红榉（Zelkova schneideriana Hand-Mazz．）材质优良，花纹美丽，抗压能力、耐水湿性和耐腐朽性强，是室内装饰和高档家具的首选材料，也是船舰、桥梁、建筑、车辆等的高档用材，深受国际国内欢迎，市场价格昂贵。本文研究了红榉不同种源的种子、叶片形态变异、苗期生长节律、木材材性差异及其光合特性和抗旱性，并用ITS序列比较分析了红榉种源的起源与分化。主要研究结果如下：
（1）红榉种源间(单株间)种子长、宽、长宽比和千粒重等性状的差异均达极显著水平（P<0.01），且种源水平上各性状的遗传力分别为0.9018、0.9210、0.9221和0.9008，单株水平上的遗传力分别为0.9260、0.9792、0.9326和0.9925，表明红榉种子形态性状存在着丰富的遗传变异，且受到中等程度以上的遗传控制；相关分析表明，红榉种子长、宽及千粒重之间存在着极显著相关关系，且红榉种子性状无明显的经向和纬向变异，种子长主要受年均温和无霜期影响，种子宽主要受1月均温影响，而千粒重主要受年均温的影响；聚类分析表明，红榉种子形态特征地理变异呈现区域板块变异模式和随机变异模式等。
（2）红榉种源间各叶片形态性状的环境方差、表型方差均大于或等于其遗传方差，遗传方差分量所占百分比少，说明红榉叶片形态性状受环境的影响很大；红榉叶片叶面积与叶长、叶宽呈极显著正相关，与周长呈显著相关，且各种源的叶面积拟合方程相关系数均在0.98以上，拟合效果良好；红榉种源间叶片长宽比、形状因子和叶厚的广义遗传力分别为0.7101、0.7064和0.7595，且遗传变异系数小，说明这些性状受到中等程度以上的遗传控制，能够比较稳定地遗传给后代。
（3）红榉不同种源苗期苗高和地径生长均为全期生长类型，且符合“S”型曲线，并能用Logistic生长方程对其生长节律动态很好的拟合；红榉种源幼苗苗高、地径的物候期参数和生长参数均存在比较明显的差异，各种源间线性生长量占总生长量百分率为54.60%～66.55%。
（4）红榉不同种源间木材基本密度的差异达到极显著水平，而种源内木材基本密度比较稳定，变异较小；不同种源间木材径向全干缩率不存在显著差异，但种源内不同个体间的差异显著；红榉木材基本密度与径向全干缩率相关关系不显著，表明基本上相互独立遗传，且两者与各地理气候因子相关关系也不显著。
（5）红榉的净光合速率(Pn)日变化呈双峰型，呈轻微的光合“午休”现象，其峰值分别出现在上午8：00和下午14:00。根据日变化参数Pn、胞间CO_2浓度（Ci）和气孔限制值（Ls）的变化方向，推断红榉光合“午休”主要受非气孔因素限制。相关分析和多元逐步回归分析表明，影响红榉光合速率日变化的主要生理生态因子为Tr、Vpdl、Ca和Ci等。
（6）当大气CO_2浓度为400μmol·mol~(-1)时，红榉叶片净光合速率(Pn)与光合有效辐射（PAR）之间回归方程为Pn=－9×10~(-6)PAR~2+0.0212PAR+0.592，光饱和点、补偿点及表观量子效率分别为1177.8μmol·m~(-2)·s~(-1)、20.0μmol·m~(-2)·s~(-1)、0.0345；叶片蒸腾速率(Tr)与PAR间回归方程：Tr=0.025PAR＋4.535(R~2=0.8169,n=48)；叶片水分利用效率（WUE）和PAR之间的回归方程：WUE=~(-2)×10~(-6)PAR~2+0.0033PAR+0.1939，相关系数为0.8222(n=48)；在1200μmol·m~(-2)·s~(-1)PAR条件下，Pn与CCO_2之间回归方程为Pn=~(-2)×10~(-5)CCO_22＋0.0621CCO_2－5.5018，R~2=0.936（n=45），CO_2饱和点、补偿点、羧化速率分别为1552.5μmol·mol~(-1)、84.8μmol·mol~(-1)、0.0403；Tr随CCO_2增强基本保持稳定不变。WUE与CCO_2之间回归方程为：WUE=~(-3)×10~(-6)CCO_22+0.0105CCO_2–0.9442，R~2=0.8945（n=45）。
（7）红榉不同种源叶片净光合速率日变化趋势并不一致，其中怀化种源表现为单峰曲线，而湖州等其他3个种源则表现为双峰曲线，且湖州种源的光合日均值显著高于其他种源；相关分析和逐步回归分析表明，不同红榉种源净光合速率日变化的影响因子大体相似，其中蒸腾速率和光合有效辐射为最稳定的影响因子；红榉不同种源的净光合速率的光响应曲线和CO_2响应曲线的总体趋势相同，但不同种源间光补偿点、光饱和点、CO_2补偿点及CO_2饱和点差异较大。
（8）红榉叶片中脯氨酸、丙二醛、可溶性蛋白质、可溶性糖和叶绿素含量等生理指标在不同种源间存在着极显著差异；隶属函数法综合评价表明，5个红榉种源的抗旱性顺序为赣州种源>怀化种源>滁州种源>湖州种源>南京种源。这说明红榉不同种源间存在着较大的抗旱性能力差异，具有较大的种源选择潜力。
(9)红榉5个种源的ITS序列、ITS1和ITS2长度变异范围分别为561～623bp、157～210bp和215～222bp；软件DNAMAN、遗传距离和UPGMA法系统发育树表明，南京种源与赣州种源的亲缘关系较近，湖州种源与怀化种源的亲缘关系较近，而滁州种源则单独聚为一类。
As an endemic species, Zelkova schneideriana Hand.-Mazz. is valuable for economy andfor landscape designing. Because of the over exploitation, the resources become less and less,so that it was listed in Red Dated Book with second grade. In this paper, The seed and leafmorphologic characteristics, seedling growth rhythm, wood characteristics, photosynthesis anddrought-tolerance, and ITS sequence analysis of different provenances were conducted, and theresults were as following:
(1) Highly significant differences were found among different provenances (individuals)in seed length, width, ratio of length to width, and1000-grain weight, and theirs heritabilitywere0.9018,0.9210,0.9221and0.9008at the provenance level, and0.9260,0.9792,0.9326and0.9925at the individual level, respectively. This indicated that these seed morphologiccharacters lied abundant genetic variation, and were in moderate to higher genetic controlled.There were highly significant correlation among seed length, width and1000-grain weight, andthese seed morphologic characters demonstrated no obvious longitude or latitude variationtendency, seed length was mainly affected by annual average temperature and frostless days,seed width was average temperature in January, While1000-grain weight was annual averagetemperature; Cluster analysis suggested that the geographic variation of seed morphologiccharacters of some distribution zones represented regional block variation pattern, while theothers showed random variation pattern.
(2)The environmental variance (VE), phenotypic variance (VP) of leaf traits amongdifferent provenances were equal to, or greater than the genetic variance(VG), this indicatedthat these traits were mainly affected by the environment; And there were highly significantdifference between leaf length, width and area, and theirs coefficients of the regressionequation were greater than0.98; And the broad-sense heritability of length/width, form factorand thickness were0.7101,0.7064and0.7595, respectively, and theirs genetic variation coefficients were rather smaller, these revealed that these traits were in moderate to highergenetic controlled.
(3) The growth rhythm of the seedling height and basal diameter belonged to full-timegrowth type, and matched perfectly with “S” curve, and was fitted by Logistic GrowthEquation at significant level; Compared among different provenances, there existed significantdifference among theirs phenophase and growth parameters of height and basal diameter, andthe ratio of total linear growth reached from54.60%to66.55%.
(4)There were highly significant differences of the basic density among differentprovenances, While theirs CV were rather stable; There were no significant differences ofradial dry shrinkage among different provenances, While the individuals within oneprovenance revealed rather higher difference; Basic density was not closely related to radialdry shrinkage. Based on this, independent selection of basic density and radial shrinkage wereexpected to generate improved effects. And both of them had no significant correlation to thegeographic and meteorological factors too.
(5)The diurnal change of net photosynthetic rate (Pn) showed double-peak curve with aslightly midday depression, with maximum values occurred at about8:00am and14:00pm.And the midday depression was mainly controlled by non-stomatal factors judging from thechanging tendency of net photosynthetic rate, intercellular CO_2concentration (Ci) and stomatallimitation (Ls). Correlation and regression analyses showed that Tr、Vpdl、Ca and Ci wereimportant affecting factors on the photosynthesis.
（6）Under the condition of400μmol·mol~(-1)CO_2, the regression equations between Pn andPAR is Pn=－9×10~(-6)PAR~2+0.0212PAR+0.592; the light saturation point (LSP)， lightcompensation point(LCP) and apparent quantum yield (AQY)are1177.8μmol·m~(-2)·s~(-1),20.0μmol·m~(-2)·s~(-1),0.0345, respectively; the regressive equation between transpirationrate(Tr)and PAR is Tr=0.025PAR＋4.535, and the regressive equation between waterutilization efficiency(WUE) and PAR is WUE=~(-2)×10~(-6)PAR~2+0.0033PAR+0.1939withR~2=0.8222. Under the condition of1200μmol·m~(-2)·s~(-1)of PAR, the regressive equation betweenPn and CO_2contents is Pn=~(-2)×10~(-5)CCO_22＋0.0621CCO_22－5.5018with R~2=0.936(n=45), CO_2 saturation point (CSP), CO_2compensation point (CCP) and Carboxylation efficiency (CE) are1552.5μmol·mol~(-1)、84.8μmol·mol~(-1)、0.0403, respectively; The regressive equation betweenWUE and CO_2contents is WUE=~(-3)×10~(-6)CCO_22+0.0105CCO_2–0.9442。However, Tr keepssteadily with the increase of CO_2contents.
(7) The diurnal variation of Pn of different provenances revealed different changetendency, and the Huaihua provenance showed single-peak curve, while the other threedemonstrated double-peak one, and the later all were limited by stomata factors, and thediurnal average of Pn of Huzhou provenance were higher significantly than the others; Thecorrelation analysis and stepwise regression analysis showed that the main affecting factorswere transpiration rate (Tr) and photosynthetic active radiation (PAR), and so on; The testedprovenances showed similar responses of Pn to PAR and Ci with evident provenance differenceof light compensation point (LCP), light saturation point (LSP), CO_2compensation point (CCP)and CO_2saturation point.
(8) There were highly significant difference of Pro, MDA, Soluble protein, Soluble sugarand Chl contents among different provenances, and the comprehensive evaluation of thedetermination of membership function revealed that the order of drought-tolerance of differentprovenances were Ganzhou>Huaihua>Chuzhou>Huzhou>Nanjing.
(9) The length of ITS sequence, ITS1and ITS2ranged61～623bp,157～210bp and215～222bp, respectively. And the DNAMAN, genetic distance and UPGMA analysis showedthat the relationships between Nanjing provenance and Ganzhou provenance, and betweenHuzhou provenance and Huaihua provenance were closer than the others, and the Chuzhouprovenances was clustered one single groups.
（1）红榉种源间(单株间)种子长、宽、长宽比和千粒重等性状的差异均达极显著水平（P<0.01），且种源水平上各性状的遗传力分别为0.9018、0.9210、0.9221和0.9008，单株水平上的遗传力分别为0.9260、0.9792、0.9326和0.9925，表明红榉种子形态性状存在着丰富的遗传变异，且受到中等程度以上的遗传控制；相关分析表明，红榉种子长、宽及千粒重之间存在着极显著相关关系，且红榉种子性状无明显的经向和纬向变异，种子长主要受年均温和无霜期影响，种子宽主要受1月均温影响，而千粒重主要受年均温的影响；聚类分析表明，红榉种子形态特征地理变异呈现区域板块变异模式和随机变异模式等。
（2）红榉种源间各叶片形态性状的环境方差、表型方差均大于或等于其遗传方差，遗传方差分量所占百分比少，说明红榉叶片形态性状受环境的影响很大；红榉叶片叶面积与叶长、叶宽呈极显著正相关，与周长呈显著相关，且各种源的叶面积拟合方程相关系数均在0.98以上，拟合效果良好；红榉种源间叶片长宽比、形状因子和叶厚的广义遗传力分别为0.7101、0.7064和0.7595，且遗传变异系数小，说明这些性状受到中等程度以上的遗传控制，能够比较稳定地遗传给后代。
（3）红榉不同种源苗期苗高和地径生长均为全期生长类型，且符合“S”型曲线，并能用Logistic生长方程对其生长节律动态很好的拟合；红榉种源幼苗苗高、地径的物候期参数和生长参数均存在比较明显的差异，各种源间线性生长量占总生长量百分率为54.60%～66.55%。
（4）红榉不同种源间木材基本密度的差异达到极显著水平，而种源内木材基本密度比较稳定，变异较小；不同种源间木材径向全干缩率不存在显著差异，但种源内不同个体间的差异显著；红榉木材基本密度与径向全干缩率相关关系不显著，表明基本上相互独立遗传，且两者与各地理气候因子相关关系也不显著。
（5）红榉的净光合速率(Pn)日变化呈双峰型，呈轻微的光合“午休”现象，其峰值分别出现在上午8：00和下午14:00。根据日变化参数Pn、胞间CO_2浓度（Ci）和气孔限制值（Ls）的变化方向，推断红榉光合“午休”主要受非气孔因素限制。相关分析和多元逐步回归分析表明，影响红榉光合速率日变化的主要生理生态因子为Tr、Vpdl、Ca和Ci等。
（6）当大气CO_2浓度为400μmol·mol~(-1)时，红榉叶片净光合速率(Pn)与光合有效辐射（PAR）之间回归方程为Pn=－9×10~(-6)PAR~2+0.0212PAR+0.592，光饱和点、补偿点及表观量子效率分别为1177.8μmol·m~(-2)·s~(-1)、20.0μmol·m~(-2)·s~(-1)、0.0345；叶片蒸腾速率(Tr)与PAR间回归方程：Tr=0.025PAR＋4.535(R~2=0.8169,n=48)；叶片水分利用效率（WUE）和PAR之间的回归方程：WUE=~(-2)×10~(-6)PAR~2+0.0033PAR+0.1939，相关系数为0.8222(n=48)；在1200μmol·m~(-2)·s~(-1)PAR条件下，Pn与CCO_2之间回归方程为Pn=~(-2)×10~(-5)CCO_22＋0.0621CCO_2－5.5018，R~2=0.936（n=45），CO_2饱和点、补偿点、羧化速率分别为1552.5μmol·mol~(-1)、84.8μmol·mol~(-1)、0.0403；Tr随CCO_2增强基本保持稳定不变。WUE与CCO_2之间回归方程为：WUE=~(-3)×10~(-6)CCO_22+0.0105CCO_2–0.9442，R~2=0.8945（n=45）。
（7）红榉不同种源叶片净光合速率日变化趋势并不一致，其中怀化种源表现为单峰曲线，而湖州等其他3个种源则表现为双峰曲线，且湖州种源的光合日均值显著高于其他种源；相关分析和逐步回归分析表明，不同红榉种源净光合速率日变化的影响因子大体相似，其中蒸腾速率和光合有效辐射为最稳定的影响因子；红榉不同种源的净光合速率的光响应曲线和CO_2响应曲线的总体趋势相同，但不同种源间光补偿点、光饱和点、CO_2补偿点及CO_2饱和点差异较大。
（8）红榉叶片中脯氨酸、丙二醛、可溶性蛋白质、可溶性糖和叶绿素含量等生理指标在不同种源间存在着极显著差异；隶属函数法综合评价表明，5个红榉种源的抗旱性顺序为赣州种源>怀化种源>滁州种源>湖州种源>南京种源。这说明红榉不同种源间存在着较大的抗旱性能力差异，具有较大的种源选择潜力。
(9)红榉5个种源的ITS序列、ITS1和ITS2长度变异范围分别为561～623bp、157～210bp和215～222bp；软件DNAMAN、遗传距离和UPGMA法系统发育树表明，南京种源与赣州种源的亲缘关系较近，湖州种源与怀化种源的亲缘关系较近，而滁州种源则单独聚为一类。
As an endemic species, Zelkova schneideriana Hand.-Mazz. is valuable for economy andfor landscape designing. Because of the over exploitation, the resources become less and less,so that it was listed in Red Dated Book with second grade. In this paper, The seed and leafmorphologic characteristics, seedling growth rhythm, wood characteristics, photosynthesis anddrought-tolerance, and ITS sequence analysis of different provenances were conducted, and theresults were as following:
(1) Highly significant differences were found among different provenances (individuals)in seed length, width, ratio of length to width, and1000-grain weight, and theirs heritabilitywere0.9018,0.9210,0.9221and0.9008at the provenance level, and0.9260,0.9792,0.9326and0.9925at the individual level, respectively. This indicated that these seed morphologiccharacters lied abundant genetic variation, and were in moderate to higher genetic controlled.There were highly significant correlation among seed length, width and1000-grain weight, andthese seed morphologic characters demonstrated no obvious longitude or latitude variationtendency, seed length was mainly affected by annual average temperature and frostless days,seed width was average temperature in January, While1000-grain weight was annual averagetemperature; Cluster analysis suggested that the geographic variation of seed morphologiccharacters of some distribution zones represented regional block variation pattern, while theothers showed random variation pattern.
(2)The environmental variance (VE), phenotypic variance (VP) of leaf traits amongdifferent provenances were equal to, or greater than the genetic variance(VG), this indicatedthat these traits were mainly affected by the environment; And there were highly significantdifference between leaf length, width and area, and theirs coefficients of the regressionequation were greater than0.98; And the broad-sense heritability of length/width, form factorand thickness were0.7101,0.7064and0.7595, respectively, and theirs genetic variation coefficients were rather smaller, these revealed that these traits were in moderate to highergenetic controlled.
(3) The growth rhythm of the seedling height and basal diameter belonged to full-timegrowth type, and matched perfectly with “S” curve, and was fitted by Logistic GrowthEquation at significant level; Compared among different provenances, there existed significantdifference among theirs phenophase and growth parameters of height and basal diameter, andthe ratio of total linear growth reached from54.60%to66.55%.
(4)There were highly significant differences of the basic density among differentprovenances, While theirs CV were rather stable; There were no significant differences ofradial dry shrinkage among different provenances, While the individuals within oneprovenance revealed rather higher difference; Basic density was not closely related to radialdry shrinkage. Based on this, independent selection of basic density and radial shrinkage wereexpected to generate improved effects. And both of them had no significant correlation to thegeographic and meteorological factors too.
(5)The diurnal change of net photosynthetic rate (Pn) showed double-peak curve with aslightly midday depression, with maximum values occurred at about8:00am and14:00pm.And the midday depression was mainly controlled by non-stomatal factors judging from thechanging tendency of net photosynthetic rate, intercellular CO_2concentration (Ci) and stomatallimitation (Ls). Correlation and regression analyses showed that Tr、Vpdl、Ca and Ci wereimportant affecting factors on the photosynthesis.
（6）Under the condition of400μmol·mol~(-1)CO_2, the regression equations between Pn andPAR is Pn=－9×10~(-6)PAR~2+0.0212PAR+0.592; the light saturation point (LSP)， lightcompensation point(LCP) and apparent quantum yield (AQY)are1177.8μmol·m~(-2)·s~(-1),20.0μmol·m~(-2)·s~(-1),0.0345, respectively; the regressive equation between transpirationrate(Tr)and PAR is Tr=0.025PAR＋4.535, and the regressive equation between waterutilization efficiency(WUE) and PAR is WUE=~(-2)×10~(-6)PAR~2+0.0033PAR+0.1939withR~2=0.8222. Under the condition of1200μmol·m~(-2)·s~(-1)of PAR, the regressive equation betweenPn and CO_2contents is Pn=~(-2)×10~(-5)CCO_22＋0.0621CCO_22－5.5018with R~2=0.936(n=45), CO_2 saturation point (CSP), CO_2compensation point (CCP) and Carboxylation efficiency (CE) are1552.5μmol·mol~(-1)、84.8μmol·mol~(-1)、0.0403, respectively; The regressive equation betweenWUE and CO_2contents is WUE=~(-3)×10~(-6)CCO_22+0.0105CCO_2–0.9442。However, Tr keepssteadily with the increase of CO_2contents.
(7) The diurnal variation of Pn of different provenances revealed different changetendency, and the Huaihua provenance showed single-peak curve, while the other threedemonstrated double-peak one, and the later all were limited by stomata factors, and thediurnal average of Pn of Huzhou provenance were higher significantly than the others; Thecorrelation analysis and stepwise regression analysis showed that the main affecting factorswere transpiration rate (Tr) and photosynthetic active radiation (PAR), and so on; The testedprovenances showed similar responses of Pn to PAR and Ci with evident provenance differenceof light compensation point (LCP), light saturation point (LSP), CO_2compensation point (CCP)and CO_2saturation point.
(8) There were highly significant difference of Pro, MDA, Soluble protein, Soluble sugarand Chl contents among different provenances, and the comprehensive evaluation of thedetermination of membership function revealed that the order of drought-tolerance of differentprovenances were Ganzhou>Huaihua>Chuzhou>Huzhou>Nanjing.
(9) The length of ITS sequence, ITS1and ITS2ranged61～623bp,157～210bp and215～222bp, respectively. And the DNAMAN, genetic distance and UPGMA analysis showedthat the relationships between Nanjing provenance and Ganzhou provenance, and betweenHuzhou provenance and Huaihua provenance were closer than the others, and the Chuzhouprovenances was clustered one single groups.
引文
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