普通丝瓜种质资源评价体系及主要农艺性状遗传规律研究
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摘要
丝瓜种质资源是丝瓜新品种选育、遗传理论研究、生物技术研究和农业生产的重要基础。目前我国已收集到丝瓜种质资源500余份,其中普通丝瓜[Luffa cylindrica (Linn.) Roem.]462份,有棱丝瓜[Luffa acutangula (Linn.) Roxb.]40多份。收集的种质资源数量较多、类型也较丰富,但对收集的种质资源研究较少,缺乏一套科学规范的种质资源鉴定和评价体系,以致影响到种质资源的整理、品种选育和相互引种等工作的开展。丝瓜种质主要农艺性状遗传规律的研究起步晚,研究成果少而不系统,不但影响了丝瓜优异种质资源的有效利用,而且制约了新品种的育种进程。
本研究基于课题组前期收集的丝瓜种质资源,通过对普通丝瓜种质主要农艺性状及其变异情况、种质间亲缘关系等的研究,以期建立普通丝瓜种质鉴定、评价体系；同时利用在主要农艺性状方面有梯度差异的普通丝瓜的高代自交系配制相应的杂交组合,以各自杂交组合的P1、P2、F1、B1、B2和F26个世代为材料,利用主基因+多基因混合遗传模型联合分离分析其遗传规律。期望通过以下几个方面的研究,为普通丝瓜优异种质资源的利用及遗传育种奠定基础。具体如下：
1.普通丝瓜主要农艺性状的调查和统计分析
对102份普通丝瓜种质的长势、叶色、叶形、裂叶度、雌(雄)花的花色、叶横径、叶纵径、叶柄长、茎粗、分枝数、第1雌花节位、雄花花冠直径、雌花花冠直径及商品瓜的瓜皮色、瓜面手感、瓜皮皱缩、瓜面的茸毛疏密、凸(凹)纹条数、凸瘤多寡、凹纹色、凸纹色、瓜长、瓜粗、瓜形指数、平均单瓜重和瓜形等性状进行调查和统计分析。结果表明：28个性状能初步反映出普通丝瓜种质主要农艺性状的特性,各性状可细分成3级(或5级、或6级)。各性状不同级别分布频次的差异可反映出种质资源基因型的丰富程度,提出的分级标准可为普通丝瓜种质的鉴定提供了较为客观的评价标准,与之配套的参照品种能客观地反映环境造成的差异。
2.普通丝瓜主要农艺性状变异情况分析及相关性研究
对102份普通丝瓜种质的11个性状(瓜长、雄花花冠直径、雌花花冠直径、瓜长、瓜径、单瓜重、叶横径、叶纵径、叶柄长、茎粗、瓜形指数和分枝数)的数据进行统计分析,探讨了这些性状间的相关性及变异情况。结果表明：瓜形指数和分枝数变异系数大,叶横径和叶纵径变异系数小。瓜长与叶柄长显著正相关,与瓜径显著负相关；瓜径与与分枝数显著正相关；单瓜重与瓜长、瓜径极显著正相关,且与叶柄长显著正相关。
3.普通丝瓜种质熟性的评价标准、评价指标及相关性分析
研究了102份普通丝瓜种质的♂1～♂3(第1-3雄花蕾节位)、♀1～♂3(第1-3雌花蕾节位)、♂1天数(从出苗到开放第1朵雄花的天数)、♀1天数(从出苗到开放第1朵雌花的天数)之间的相关性。结果表明,♂1～♂3之间及♂1～♂3之间均达极显著正相关；♂1～♂3和♂1天数间、♀1～♂3和♀1天数间均达极显著正相关。研究认为,可以以第1雌花蕾节位的高、低作为普通丝瓜熟性迟、早的评价标准,本研究提出普通丝瓜熟性迟早(南京地区春季)的评价指标：早熟材料的♀1≤7.6节,中熟材料的♀1是7.7节～12.9节,晚熟材料的♀1≥13.0节。
4.普通丝瓜种质性型分化的特点和表现形式
对78份普通丝瓜种质30节内的雌花蕾、雄花蕾着生状况进行调查,研究了无性节、纯雄节、纯雌节和双性节的分布规律。结果表明：普通丝瓜的性型分化比较复杂。无性节一般分布在起始5节内,少量品种在高节位还有零星分布；双性节中雌花蕾和雄花蕾有11种组合方式,以多雄+单雌节最常见；雄花蕾分布在纯雄节和双性节之中,雄花蕾在纯雄节中存在6种表现形式；早熟材料的有雄节数(包括纯雄节和双性节)为24.1节,中熟材料的有雄节数为23.6节,晚熟材料的有雄节数为21.3节,且熟性越早有雄节数越多；雌花蕾在纯雌节中有4种表现形式,以单雌形式存在最普遍；早熟材料的有雌节数(包括纯雌节和双性节)为23节,中熟材料的有雌节数为18.7节,晚熟材料的有雌节数为13.5节；无性节、纯雄节、纯雌节和双性节的节数与品种来源地、果形、植株长势和叶色均未表现出相关性。
5.丝瓜种质亲缘关系的ISSR分析
利用ISSR标记对来源于不同地区的丝瓜种质资源的亲缘关系进行分析。从80条ISSR引物中筛选出多态性强、重复性好的9条引物,对43份丝瓜种质基因组DNA进行扩增,共扩增出60条谱带,平均每个引物扩增出6.67条带,其中多态性带47个,多态性位点百分率为78.3%。丝瓜种质间遗传相似系数变化范围在0.37-0.98之间,暗示了丝瓜栽培种内的遗传基础相对狭窄。利用UPGMA聚类分析,发现43个丝瓜种质划分为6个类群,类群的划分与形态学性状较吻合,而且与地理来源也有较高的相关性。
6.普通丝瓜种质第1雌花节位的遗传规律研究
选用第1雌花节位有差异的普通丝瓜品种的高代自交系配制L1xL2(‘五叶香丝瓜’ב短圆筒丝瓜’)、L2xL3(‘短圆筒丝瓜’ב蛇形丝瓜’)和L3×L4(‘蛇形丝瓜’ב长沙肉丝瓜’)3套组合,通过调查3套组合各自的P1、P2、F1、B1、B2和F2植株的第1雌花节位,利用主基因+多基因混合遗传模型联合分离分析了第1雌花节位遗传规律。结果表明：L1×L2第1雌花节位遗传符合2对加性-显性-上位性主基因+加性-显性多基因遗传模型,L2×L3的遗传符合1对加性主基因+加性-显性多基因遗传模型,L3×L4的遗传符合2对加性主基因+加-显性多基因遗传模型；L1×L2组合的B1、B2和F2群体遗传率(主基因+多基因)分别为66.13%、51.29%和68.27%,L2×L3组合的B1、B2和F2群体遗传率(主基因+多基因)分别为82.02%、64.87%和65.62%,L3×L4组合的B1、B2和F2群体遗传率(主基因+多基因)分别为54.45%、61.88%和58.91%；L1×L2组合B1、B2和F2群体的环境方差占总表型方差的比例分别是23.43%、48.69%和31.73%,L2×L3组合B1、B2和F2群体的环境方差占总表型方差的比例分别是34.27%、55.40%和34.38%,L3×L4组合B1、B2和F2群体的环境方差占总表型方差的比例分别是45.1%、38.02%和40.71%。以上结果表明,普通丝瓜第1雌花节位是由主基因和多基因控制的数量性状,早熟性(较低的第1雌花节位)难以通过杂种优势来实现；第1雌花节位遗传不稳定,易受环境因素的影响,但定向选择会有较好的效果。
7.普通丝瓜种质果实性状的遗传规律研究
应用植物数量性状主基因+多基因混合遗传模型对普通丝瓜品种‘50-5'(‘黑籽短圆筒’)×'20-4'(、‘桂林水瓜’)杂交组合6个世代群体5个果实性状(瓜柄长、瓜长、瓜径、瓜形指数和单瓜重)进行了联合分析,结果表明：‘50-5’×'20-4'组合果实的瓜柄长遗传符合2对加性-显性-上位性主基因+加性-显性多基因遗传模型；其B1、B2和F2群体遗传率(主基因+多基因)分别为68.49%、70.53%和82.07%,环境方差占总表型方差的比例分别是31.5%、29.47%和17.92%。瓜长遗传符合2对加性-显性-上位性主基因+加性-显性-上位性多基因遗传模型；B1、B2和F2群体遗传率(主基因+多基因)分别为68.85%、84.55%和81.68%,环境方差占总表型方差的比例分别是31.15%、15.44%和18.32%。瓜径遗传符合2对加性-显性-上位性主基因+加性-显性-上位性多基因遗传模型；B1、B2和F2群体遗传率(主基因+多基因)分别为65.23%、73.06%和73.82%,环境方差占总表型方差的比例分别是34.62%、26.94%和26.13%。瓜形指数遗传符合2对加性-显性-上位性主基因+加性-显性-上位性多基因遗传模型；B1、B2和F2群体遗传率(主基因+多基因)分别为65.23%、62.8%和78.89%,环境方差占总表型方差的比例分别是34.76%、37.19%和21.11%。单瓜重遗传符合2对加性-显性-上位性主基因+加性-显性-上位性多基因遗传模型；B1、B2和F2群体遗传率(主基因+多基因)分别为70.71%、85.35%和89.64%,环境方差占总表型方差的比例分别是29.29%、14.64%和10.36%。以上结果分析表明,瓜柄长性状的主基因遗传率较小,宜采用个体选择法(基因型选择法),在分离晚代进行选择；瓜长、瓜径、瓜形指数和单瓜重性状的主基因遗传率较大,宜采取混合选择法(表型选择法),在分离早代进行选择。5个果实性状的环境方差占总表型方差的比例均较高,故在育种过程中要尽量采取措施以减少环境影响。
Germplasm of luffa [Luffa cylindrical L.] is the important basic of new luffa varieties breeding, genetic theory studying, and biotechnology studying and agricultural production. Although,500luffa germplasm were collected in our country, among which462were luffa [Luffa cylindrical (L.) Roem.] and about40were luffa [Luffa acutangula (Linn.) Roxb.], the studies on these germplasm were relatively less, and no normative evaluation system was constructed, therefore, affecting the tidying of germplasm, breeding of variety and introduction of new variety. The study on inheritance of main agronomic characters was relatively late and poor and unsystematic, not only restricting the utilization of excellent germplasm, but also the efficiency of breeding.
In this work, on the basis of luffa germplasm that collected, the studies on main agronomic characters and its variation and their phylogenetic relationship were carried in order to establish the evaluation system of luffa germplasm. In addition, the P1, P2, F1, B1, B2and F2groups of the first female flower node and/or five characters of fruit were constructed, and the inheritance of them were studied by using the mixed major gene plus polygene-gene inheritance model, expecting promote the utilization of excellent luffa germplasm and breeding. The detailed results as follows:
1. The studies on main agronomic characters and statistic analysis of luffa
The characters of102Luffa germplasm, such as growth vigor, leaf color, leaf shape, deep of leaf teeth, female/male flower color, fruit peel color, feeling of fruit surface by hand, shrinking of fruit peel, fuzz density on fruit peel, number of raised lines on fruit peel, number of concave lines on fruit peel, raised tumor density on fruit peel, raised lines color, concave lines color leaf width, leaf length, leaf stalk length, stem diameter, number of branch bine, the nodal position for first female flower bud, male flower cap diameter, female flower cap diameter, fruit length, fruit diameter, fruit index, mean fruit weight and fruit shape have been studied and analyzed. The results showed that these characters have a preliminary representative on main agronomic characteristics. In addition, These characters have been divided into different grades, frequency distribution and percentage of each grades have been investigated, reference varieties also have been given respectively.
2. Correlation analysis and variation of main agronomic characters of Luffa
The correlation analysis and variation range of eleven characters of102Luffa varieties, such as male flower cap diameter, female flower cap diameter, fruit length, fruit diameter, mean fruit weight, leaf width, leaf length, leaf stalk length, stem diameter, fruit index and number of branch bine have been studied. The results showed that the coefficients of variation of fruit index and number of branch bine were large, while, the coefficients of variation of leaf width and leaf length were small. The results of correlation analysis showed that significant positive correlation were found between the fruit length and leaf stalk length; the significant negative correlation were found between the fruit length and fruit diameter, while, positive correlation between the fruit diameter and the number of branch bine; the significant positive correlation were investigated among the mean fruit weight, fruit length and fruit diameter, while, the significant positive correlation between the mean fruit weigh and leaf stalk length.
3. Criterion and index of evaluation and correlation analysis of maturation traits of Luffa
The correlation between the node of the first to the third female flower bud and the first to the third male flower bud, the days of the first male and female flower of102luffa germplasm were analyzed. The results indicated that significant positive correlation was found between the node of the first to the third male flower bud, the node of the first to the third female flower bud, the node of the first to the third male flower bud and the days of the first male flower, the node of the first to the third female flower bud and the days of the first female flower, respectively. The node of the first female flower bud could be considered the standard to evaluate the maturation trait of luffa. The evaluation index as follows:less than7.6nodes of the first female flower were considered to be early maturity materials;7.7to12.9nodes were considered to be middle maturity materials and more than13.0nodes were late maturity materials.
4. Characters and manifestation of the sex differentiation in Luffa
The node of male and female flower bud distributed inside the30th nodes of78luffa germplasm and the distribution of asexual node, pure-male node, pure-female node and male/female node was investigated. The results indicated that the sex differentiation of luffa was complex. The asexual node always distributed under the fifth node, a little in higher node.11combinations existed in the male/female node, and the main combination was poly-male-single-female node. The male flower distributed in the pure-female node and male/female node, and there were6presentations of male flower in pure-male node. Early-maturing luffa have24.1male nodes (including pure-male node and male/female node), middle-maturing luffa have23.6male nodes, and late-maturing luffa have21.3male nodes. The results also indicated that the earlier-maturing luffa would have more number of male nodes.4manifestations of the female flower were existed in pure-female node, mainly in single female flower.23of female nodes (including pure-female node and male/female node) were in early-maturing luffa,18.7of female nodes in middle-maturing luffa, and13.5of female nodes in late-maturing luffa. The number of asexual node, pure-male node, pure-female node and male/female node were not correlated to origin of varieties, fruit shape, growth and color of leaf.
5. Analysis on the phylogenetic relationship of Luffa based on ISSR
Phylogenesis of43common luffa germplasm from different regions was analyzed with Inter-simple Sequence Repeat (ISSR).9primers were selected from80ISSR primers.60DNA fragments were amplified from43samples and each primer resulted in6.67DNA fragments.47fragments were polymorphic (percentage of polymorphic bands was78.3%). The value of Nei's genetic similarity (GS) indexes of43common luffa germplasm varied from0.37to0.98, suggesting that the genetic diversity among them is short. Cluster analysis with UPGMA method showed that the43germplasm could be divided into six groups. The results according to the pedigree relations had connections with morphological traits and regions of germplasm origin.
6. The inheritance of the first female flower node in Luffa
Inheritance of the node for first female flower in luffa(Luffa cylindrical Roem.) in P1、 P2、F1、B1、B2and F2from two crosses L1×L2(Wuyexiangsigua×Duanyuantongsigua) and L2×L3(Duanyuantongsigua×Shexingsigua) were investigated by the mixed major gene plus polygene-gene inheritance model of quantitative traits. The joint analysis results showed that the node for first female flower bud of the cross L1×L2was controlled by two adding-dominance-epistatic major gene plus adding-dominance polygene model, another cross L2×L3was controlled by one adding-dominance major gene plus adding-dominance polygene model, the third cross L3XL4was controlled by two adding major gene plus adding-dominance polygene model. Heritability values(of major gene+polygene) of the cross L1×L2of B1、B2and F2population was estimated as66.13%、51.29%and68.27%respectively; the cross L2×L3of B1、B2and F2population was estimated as82.02%、64.87%and65.62%respectively, the cross L3×L4of B1、B2and F2population was estimated as54.45%、61.88%and58.91%respectively. Environment VAR of the cross L1×L2of B1、B2and F2population was34.27%、55.40%and34.38%in total VAR respectively; while the cross L2×L3of B1、B2and F2population was34.27%、55.40%and34.38%respectively, the cross L3×L4of B1、B2and F2population was45.1%、38.02%and40.71%respectively. Conclusion:The node for first female flower bud in luffa(Luffa cylindrical Roem.) was quantitative traits and was controlled by major gene and polygene, early matured(lower node) was made by hybrid vigor impossibly, directional selection was effective on the node for first female flower bud, heritability values of the crosses was instability and environment play an important role in the inheritance.
7. The inheritance of fruit characters of Luffa
Inheritance of the5fruit characters(fruit stalk length、fruit length、fruit diameter fruit index and mean fruit weight) for luffa(Luffa cylindrical Roem.) in P1、P2、F1、B1、B2and F2from the cross '50-5'×'20-4'(Heiziduanyuanton) x(Guilinshuigua) were investigated by the mixed major gene plus polygene-gene inheritance model of quantitative traits. The joint analysis results showed that the fruit stalk length of the cross of'50-5'x'20-4'was controlled by two adding-dominance-epistatic major gene plus adding-dominance polygene model, the fruit length、fruit diameter、fruit index and mean fruit weight f the cross of '50-5'×'20-4' were controlled by two adding-dominance-epistatic major gene plus adding-dominance-epistatic polygene model. Heritability values(of major gene+polygene) of fruit stalk length in this cross of B1、B2and F2population was estimated as68.49%、70.53%and82.07%respectively; fruit length was estimated as68.85%、84.55%and81.68%respectively; fruit diameter was estimated as65.23%、73.06%and73.82%respectively; fruit index was estimated as65.23%、62.8%and78.89%respectively; and mean fruit weight was estimated as70.71%、85.35%and89.64%respectively. Environment VAR of fruit stalk length in this cross of B1. B2and F2population was31.5%.29.47%和17.92%in total VAR respectively; fruit length was31.15%、15.44%and18.32%respectively; fruit diameter was34.62%.26.94%and26.13%respectively; fruit index was34.76%.37.19%and21.11%respectively; and mean fruit weight was29.29%.14.64%and10.36%respectively. The individual selection (Genotype selection)of fruit character Of fruit stalk length on high separated generations was effective because of its small heritability values; while the mixed selection (Phenotype selection)of fruit character Of fruit length. fruit diameter, fruit index and mean fruit weight on young(low) separated generations was effective because of its big heritability values. Systematic consideration on5fruit characters was better. Environment VAR of5fruit characters has much proportion in total phenotype VAR, environment effects must be reduced to minimum in the luffa breeding research.
本研究基于课题组前期收集的丝瓜种质资源,通过对普通丝瓜种质主要农艺性状及其变异情况、种质间亲缘关系等的研究,以期建立普通丝瓜种质鉴定、评价体系；同时利用在主要农艺性状方面有梯度差异的普通丝瓜的高代自交系配制相应的杂交组合,以各自杂交组合的P1、P2、F1、B1、B2和F26个世代为材料,利用主基因+多基因混合遗传模型联合分离分析其遗传规律。期望通过以下几个方面的研究,为普通丝瓜优异种质资源的利用及遗传育种奠定基础。具体如下：
1.普通丝瓜主要农艺性状的调查和统计分析
对102份普通丝瓜种质的长势、叶色、叶形、裂叶度、雌(雄)花的花色、叶横径、叶纵径、叶柄长、茎粗、分枝数、第1雌花节位、雄花花冠直径、雌花花冠直径及商品瓜的瓜皮色、瓜面手感、瓜皮皱缩、瓜面的茸毛疏密、凸(凹)纹条数、凸瘤多寡、凹纹色、凸纹色、瓜长、瓜粗、瓜形指数、平均单瓜重和瓜形等性状进行调查和统计分析。结果表明：28个性状能初步反映出普通丝瓜种质主要农艺性状的特性,各性状可细分成3级(或5级、或6级)。各性状不同级别分布频次的差异可反映出种质资源基因型的丰富程度,提出的分级标准可为普通丝瓜种质的鉴定提供了较为客观的评价标准,与之配套的参照品种能客观地反映环境造成的差异。
2.普通丝瓜主要农艺性状变异情况分析及相关性研究
对102份普通丝瓜种质的11个性状(瓜长、雄花花冠直径、雌花花冠直径、瓜长、瓜径、单瓜重、叶横径、叶纵径、叶柄长、茎粗、瓜形指数和分枝数)的数据进行统计分析,探讨了这些性状间的相关性及变异情况。结果表明：瓜形指数和分枝数变异系数大,叶横径和叶纵径变异系数小。瓜长与叶柄长显著正相关,与瓜径显著负相关；瓜径与与分枝数显著正相关；单瓜重与瓜长、瓜径极显著正相关,且与叶柄长显著正相关。
3.普通丝瓜种质熟性的评价标准、评价指标及相关性分析
研究了102份普通丝瓜种质的♂1～♂3(第1-3雄花蕾节位)、♀1～♂3(第1-3雌花蕾节位)、♂1天数(从出苗到开放第1朵雄花的天数)、♀1天数(从出苗到开放第1朵雌花的天数)之间的相关性。结果表明,♂1～♂3之间及♂1～♂3之间均达极显著正相关；♂1～♂3和♂1天数间、♀1～♂3和♀1天数间均达极显著正相关。研究认为,可以以第1雌花蕾节位的高、低作为普通丝瓜熟性迟、早的评价标准,本研究提出普通丝瓜熟性迟早(南京地区春季)的评价指标：早熟材料的♀1≤7.6节,中熟材料的♀1是7.7节～12.9节,晚熟材料的♀1≥13.0节。
4.普通丝瓜种质性型分化的特点和表现形式
对78份普通丝瓜种质30节内的雌花蕾、雄花蕾着生状况进行调查,研究了无性节、纯雄节、纯雌节和双性节的分布规律。结果表明：普通丝瓜的性型分化比较复杂。无性节一般分布在起始5节内,少量品种在高节位还有零星分布；双性节中雌花蕾和雄花蕾有11种组合方式,以多雄+单雌节最常见；雄花蕾分布在纯雄节和双性节之中,雄花蕾在纯雄节中存在6种表现形式；早熟材料的有雄节数(包括纯雄节和双性节)为24.1节,中熟材料的有雄节数为23.6节,晚熟材料的有雄节数为21.3节,且熟性越早有雄节数越多；雌花蕾在纯雌节中有4种表现形式,以单雌形式存在最普遍；早熟材料的有雌节数(包括纯雌节和双性节)为23节,中熟材料的有雌节数为18.7节,晚熟材料的有雌节数为13.5节；无性节、纯雄节、纯雌节和双性节的节数与品种来源地、果形、植株长势和叶色均未表现出相关性。
5.丝瓜种质亲缘关系的ISSR分析
利用ISSR标记对来源于不同地区的丝瓜种质资源的亲缘关系进行分析。从80条ISSR引物中筛选出多态性强、重复性好的9条引物,对43份丝瓜种质基因组DNA进行扩增,共扩增出60条谱带,平均每个引物扩增出6.67条带,其中多态性带47个,多态性位点百分率为78.3%。丝瓜种质间遗传相似系数变化范围在0.37-0.98之间,暗示了丝瓜栽培种内的遗传基础相对狭窄。利用UPGMA聚类分析,发现43个丝瓜种质划分为6个类群,类群的划分与形态学性状较吻合,而且与地理来源也有较高的相关性。
6.普通丝瓜种质第1雌花节位的遗传规律研究
选用第1雌花节位有差异的普通丝瓜品种的高代自交系配制L1xL2(‘五叶香丝瓜’ב短圆筒丝瓜’)、L2xL3(‘短圆筒丝瓜’ב蛇形丝瓜’)和L3×L4(‘蛇形丝瓜’ב长沙肉丝瓜’)3套组合,通过调查3套组合各自的P1、P2、F1、B1、B2和F2植株的第1雌花节位,利用主基因+多基因混合遗传模型联合分离分析了第1雌花节位遗传规律。结果表明：L1×L2第1雌花节位遗传符合2对加性-显性-上位性主基因+加性-显性多基因遗传模型,L2×L3的遗传符合1对加性主基因+加性-显性多基因遗传模型,L3×L4的遗传符合2对加性主基因+加-显性多基因遗传模型；L1×L2组合的B1、B2和F2群体遗传率(主基因+多基因)分别为66.13%、51.29%和68.27%,L2×L3组合的B1、B2和F2群体遗传率(主基因+多基因)分别为82.02%、64.87%和65.62%,L3×L4组合的B1、B2和F2群体遗传率(主基因+多基因)分别为54.45%、61.88%和58.91%；L1×L2组合B1、B2和F2群体的环境方差占总表型方差的比例分别是23.43%、48.69%和31.73%,L2×L3组合B1、B2和F2群体的环境方差占总表型方差的比例分别是34.27%、55.40%和34.38%,L3×L4组合B1、B2和F2群体的环境方差占总表型方差的比例分别是45.1%、38.02%和40.71%。以上结果表明,普通丝瓜第1雌花节位是由主基因和多基因控制的数量性状,早熟性(较低的第1雌花节位)难以通过杂种优势来实现；第1雌花节位遗传不稳定,易受环境因素的影响,但定向选择会有较好的效果。
7.普通丝瓜种质果实性状的遗传规律研究
应用植物数量性状主基因+多基因混合遗传模型对普通丝瓜品种‘50-5'(‘黑籽短圆筒’)×'20-4'(、‘桂林水瓜’)杂交组合6个世代群体5个果实性状(瓜柄长、瓜长、瓜径、瓜形指数和单瓜重)进行了联合分析,结果表明：‘50-5’×'20-4'组合果实的瓜柄长遗传符合2对加性-显性-上位性主基因+加性-显性多基因遗传模型；其B1、B2和F2群体遗传率(主基因+多基因)分别为68.49%、70.53%和82.07%,环境方差占总表型方差的比例分别是31.5%、29.47%和17.92%。瓜长遗传符合2对加性-显性-上位性主基因+加性-显性-上位性多基因遗传模型；B1、B2和F2群体遗传率(主基因+多基因)分别为68.85%、84.55%和81.68%,环境方差占总表型方差的比例分别是31.15%、15.44%和18.32%。瓜径遗传符合2对加性-显性-上位性主基因+加性-显性-上位性多基因遗传模型；B1、B2和F2群体遗传率(主基因+多基因)分别为65.23%、73.06%和73.82%,环境方差占总表型方差的比例分别是34.62%、26.94%和26.13%。瓜形指数遗传符合2对加性-显性-上位性主基因+加性-显性-上位性多基因遗传模型；B1、B2和F2群体遗传率(主基因+多基因)分别为65.23%、62.8%和78.89%,环境方差占总表型方差的比例分别是34.76%、37.19%和21.11%。单瓜重遗传符合2对加性-显性-上位性主基因+加性-显性-上位性多基因遗传模型；B1、B2和F2群体遗传率(主基因+多基因)分别为70.71%、85.35%和89.64%,环境方差占总表型方差的比例分别是29.29%、14.64%和10.36%。以上结果分析表明,瓜柄长性状的主基因遗传率较小,宜采用个体选择法(基因型选择法),在分离晚代进行选择；瓜长、瓜径、瓜形指数和单瓜重性状的主基因遗传率较大,宜采取混合选择法(表型选择法),在分离早代进行选择。5个果实性状的环境方差占总表型方差的比例均较高,故在育种过程中要尽量采取措施以减少环境影响。
Germplasm of luffa [Luffa cylindrical L.] is the important basic of new luffa varieties breeding, genetic theory studying, and biotechnology studying and agricultural production. Although,500luffa germplasm were collected in our country, among which462were luffa [Luffa cylindrical (L.) Roem.] and about40were luffa [Luffa acutangula (Linn.) Roxb.], the studies on these germplasm were relatively less, and no normative evaluation system was constructed, therefore, affecting the tidying of germplasm, breeding of variety and introduction of new variety. The study on inheritance of main agronomic characters was relatively late and poor and unsystematic, not only restricting the utilization of excellent germplasm, but also the efficiency of breeding.
In this work, on the basis of luffa germplasm that collected, the studies on main agronomic characters and its variation and their phylogenetic relationship were carried in order to establish the evaluation system of luffa germplasm. In addition, the P1, P2, F1, B1, B2and F2groups of the first female flower node and/or five characters of fruit were constructed, and the inheritance of them were studied by using the mixed major gene plus polygene-gene inheritance model, expecting promote the utilization of excellent luffa germplasm and breeding. The detailed results as follows:
1. The studies on main agronomic characters and statistic analysis of luffa
The characters of102Luffa germplasm, such as growth vigor, leaf color, leaf shape, deep of leaf teeth, female/male flower color, fruit peel color, feeling of fruit surface by hand, shrinking of fruit peel, fuzz density on fruit peel, number of raised lines on fruit peel, number of concave lines on fruit peel, raised tumor density on fruit peel, raised lines color, concave lines color leaf width, leaf length, leaf stalk length, stem diameter, number of branch bine, the nodal position for first female flower bud, male flower cap diameter, female flower cap diameter, fruit length, fruit diameter, fruit index, mean fruit weight and fruit shape have been studied and analyzed. The results showed that these characters have a preliminary representative on main agronomic characteristics. In addition, These characters have been divided into different grades, frequency distribution and percentage of each grades have been investigated, reference varieties also have been given respectively.
2. Correlation analysis and variation of main agronomic characters of Luffa
The correlation analysis and variation range of eleven characters of102Luffa varieties, such as male flower cap diameter, female flower cap diameter, fruit length, fruit diameter, mean fruit weight, leaf width, leaf length, leaf stalk length, stem diameter, fruit index and number of branch bine have been studied. The results showed that the coefficients of variation of fruit index and number of branch bine were large, while, the coefficients of variation of leaf width and leaf length were small. The results of correlation analysis showed that significant positive correlation were found between the fruit length and leaf stalk length; the significant negative correlation were found between the fruit length and fruit diameter, while, positive correlation between the fruit diameter and the number of branch bine; the significant positive correlation were investigated among the mean fruit weight, fruit length and fruit diameter, while, the significant positive correlation between the mean fruit weigh and leaf stalk length.
3. Criterion and index of evaluation and correlation analysis of maturation traits of Luffa
The correlation between the node of the first to the third female flower bud and the first to the third male flower bud, the days of the first male and female flower of102luffa germplasm were analyzed. The results indicated that significant positive correlation was found between the node of the first to the third male flower bud, the node of the first to the third female flower bud, the node of the first to the third male flower bud and the days of the first male flower, the node of the first to the third female flower bud and the days of the first female flower, respectively. The node of the first female flower bud could be considered the standard to evaluate the maturation trait of luffa. The evaluation index as follows:less than7.6nodes of the first female flower were considered to be early maturity materials;7.7to12.9nodes were considered to be middle maturity materials and more than13.0nodes were late maturity materials.
4. Characters and manifestation of the sex differentiation in Luffa
The node of male and female flower bud distributed inside the30th nodes of78luffa germplasm and the distribution of asexual node, pure-male node, pure-female node and male/female node was investigated. The results indicated that the sex differentiation of luffa was complex. The asexual node always distributed under the fifth node, a little in higher node.11combinations existed in the male/female node, and the main combination was poly-male-single-female node. The male flower distributed in the pure-female node and male/female node, and there were6presentations of male flower in pure-male node. Early-maturing luffa have24.1male nodes (including pure-male node and male/female node), middle-maturing luffa have23.6male nodes, and late-maturing luffa have21.3male nodes. The results also indicated that the earlier-maturing luffa would have more number of male nodes.4manifestations of the female flower were existed in pure-female node, mainly in single female flower.23of female nodes (including pure-female node and male/female node) were in early-maturing luffa,18.7of female nodes in middle-maturing luffa, and13.5of female nodes in late-maturing luffa. The number of asexual node, pure-male node, pure-female node and male/female node were not correlated to origin of varieties, fruit shape, growth and color of leaf.
5. Analysis on the phylogenetic relationship of Luffa based on ISSR
Phylogenesis of43common luffa germplasm from different regions was analyzed with Inter-simple Sequence Repeat (ISSR).9primers were selected from80ISSR primers.60DNA fragments were amplified from43samples and each primer resulted in6.67DNA fragments.47fragments were polymorphic (percentage of polymorphic bands was78.3%). The value of Nei's genetic similarity (GS) indexes of43common luffa germplasm varied from0.37to0.98, suggesting that the genetic diversity among them is short. Cluster analysis with UPGMA method showed that the43germplasm could be divided into six groups. The results according to the pedigree relations had connections with morphological traits and regions of germplasm origin.
6. The inheritance of the first female flower node in Luffa
Inheritance of the node for first female flower in luffa(Luffa cylindrical Roem.) in P1、 P2、F1、B1、B2and F2from two crosses L1×L2(Wuyexiangsigua×Duanyuantongsigua) and L2×L3(Duanyuantongsigua×Shexingsigua) were investigated by the mixed major gene plus polygene-gene inheritance model of quantitative traits. The joint analysis results showed that the node for first female flower bud of the cross L1×L2was controlled by two adding-dominance-epistatic major gene plus adding-dominance polygene model, another cross L2×L3was controlled by one adding-dominance major gene plus adding-dominance polygene model, the third cross L3XL4was controlled by two adding major gene plus adding-dominance polygene model. Heritability values(of major gene+polygene) of the cross L1×L2of B1、B2and F2population was estimated as66.13%、51.29%and68.27%respectively; the cross L2×L3of B1、B2and F2population was estimated as82.02%、64.87%and65.62%respectively, the cross L3×L4of B1、B2and F2population was estimated as54.45%、61.88%and58.91%respectively. Environment VAR of the cross L1×L2of B1、B2and F2population was34.27%、55.40%and34.38%in total VAR respectively; while the cross L2×L3of B1、B2and F2population was34.27%、55.40%and34.38%respectively, the cross L3×L4of B1、B2and F2population was45.1%、38.02%and40.71%respectively. Conclusion:The node for first female flower bud in luffa(Luffa cylindrical Roem.) was quantitative traits and was controlled by major gene and polygene, early matured(lower node) was made by hybrid vigor impossibly, directional selection was effective on the node for first female flower bud, heritability values of the crosses was instability and environment play an important role in the inheritance.
7. The inheritance of fruit characters of Luffa
Inheritance of the5fruit characters(fruit stalk length、fruit length、fruit diameter fruit index and mean fruit weight) for luffa(Luffa cylindrical Roem.) in P1、P2、F1、B1、B2and F2from the cross '50-5'×'20-4'(Heiziduanyuanton) x(Guilinshuigua) were investigated by the mixed major gene plus polygene-gene inheritance model of quantitative traits. The joint analysis results showed that the fruit stalk length of the cross of'50-5'x'20-4'was controlled by two adding-dominance-epistatic major gene plus adding-dominance polygene model, the fruit length、fruit diameter、fruit index and mean fruit weight f the cross of '50-5'×'20-4' were controlled by two adding-dominance-epistatic major gene plus adding-dominance-epistatic polygene model. Heritability values(of major gene+polygene) of fruit stalk length in this cross of B1、B2and F2population was estimated as68.49%、70.53%and82.07%respectively; fruit length was estimated as68.85%、84.55%and81.68%respectively; fruit diameter was estimated as65.23%、73.06%and73.82%respectively; fruit index was estimated as65.23%、62.8%and78.89%respectively; and mean fruit weight was estimated as70.71%、85.35%and89.64%respectively. Environment VAR of fruit stalk length in this cross of B1. B2and F2population was31.5%.29.47%和17.92%in total VAR respectively; fruit length was31.15%、15.44%and18.32%respectively; fruit diameter was34.62%.26.94%and26.13%respectively; fruit index was34.76%.37.19%and21.11%respectively; and mean fruit weight was29.29%.14.64%and10.36%respectively. The individual selection (Genotype selection)of fruit character Of fruit stalk length on high separated generations was effective because of its small heritability values; while the mixed selection (Phenotype selection)of fruit character Of fruit length. fruit diameter, fruit index and mean fruit weight on young(low) separated generations was effective because of its big heritability values. Systematic consideration on5fruit characters was better. Environment VAR of5fruit characters has much proportion in total phenotype VAR, environment effects must be reduced to minimum in the luffa breeding research.
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