西葫芦杂种一代产量优势形成机理的研究
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摘要
本试验选取遗传差异显著的7个自交系以及配制的7个杂交组合为试材，研究了西葫芦杂种一代产量优势形成的农艺学、生理学和遗传学基础，以期为杂种优势利用、亲本选择选配和优势预测提供理论依据。主要研究结果如下：
1.农艺学性状相关分析及杂种一代优势表现分析表明，单株结果数、单果重、果实发育速度、雌花数、座果率、果长、果粗、早期单株产量、早期单株结果数、早期单果重等性状与单株产量呈显著或极显著正相关，且杂种一代均表现为正向优势，以单株产量、果实发育速度两个性状优势尤为明显，平均超亲优势指数均为1.61。始花期、始收期与单株产量呈显著负相关，杂种一代表现为负向优势，平均超中优势指数分别为0.96和0.97。杂种一代与单株产量密切相关的农艺学性状较其亲本均有不同程度的优势表现，构成产量优势形成的农艺学基础。
2.苗期、结果中期、结果末期光合速率与单株产量呈显著或极显著正相关，且杂种一代均表现为正向优势，以苗期和结果末期优势尤为明显。不同生长发育期叶面积与单株产量相关性均达显著或极显著水平，且杂种一代叶面积较其亲本均表现出明显的正向优势。结果中期叶绿素b含量、结果末期叶绿素a/b与单株产量的相关性达显著水平，而且杂种一代较其亲本表现为正向优势。杂种一代较其亲本拥有更大的光合面积，对光照尤其是弱光利用率高，整个生长发育期植株均能维持较高的光合速率，可能是杂种优势形成的重要生理基础之一。
整个生长发育期，杂种一代植株中干物质含量较其亲本均表现出不同程度的正向优势，以结果期优势更为明显,优势指数均在1.40以上。生长前期杂种一代干物质在营养器官中积累分配和结果中期、结果末期在果实中积累分配较亲本均表现出明显优势，可能是产量优势形成的又一重要原因。
从氮、磷、钾养分积累分配分析可以看出，苗期杂种一代与其亲本无明显差异，结果期，杂种一代均表现出较其亲本更快的营养元素积累能力，
生长前期营养器官和后期果实中养分积累分配强优势表现表明，杂种一代较其亲本能够更为合理的分配氮、磷、钾营养成分，养分利用率高，有助于产量优势形成。
3.遗传距离与单株产量的优势指数呈不显著程度的负相关，表明遗传距离并非越大越好。
亲子相关分析表明，杂种一代农艺学性状均不同程度的受到亲本影响，可以通过亲本性状表现对杂种一代与亲本密切相关的性状进行初步预测，育种时应注意以适当的品种作亲本。
In this experiment, seven inbreds which had different genetic backgroud together with seven hybrids breded with them were studied to find the agronomical, physiological and genetic mechanism of heterosis performance. The aim was to provide theoretical basis for heterosis utility, parent selection and heterosis prodiction. The result were as follows:
1. Correlation analysis between agronomical characters and heterosis performance in the hybrids indicated that the fruit number per plant, fruit weight, fruit growth rate, female flower number, percentage of fruit setting, early yield per plant, early fruit number per plant, early fruit weight length of fruit and diameter of fruit all had positive correlation with the yield per plant at significant or very significant level. The hybrids showed positive mid-parent heterosis and even super-parent heterosis in these characters. Among them, the heterosis of the yield per plant and the fruit growth rate were much higher, whose mid-parent heterosis index reached 1.61, and super-parent heterosis index reached 1.53 and 1.47 respectively. However, the period from planting to flowering and from planting to first harvesting. had negative relationship with the yield per plant and the hybrids showed negative heterosis. Their mid-parent heterosis index were 0.96 and 0.97 respectively. In a word, all of the characters which were closely correlated with the yield per plant had heterosis performance compared with their parents, this may be the agronomical basis of heterosis forming.
2.In the stage of seedling, middle fruiting and end fruiting, the correlation coefficient between the photosynthetic rate and the yield per plant reached significant even very significant level. The hybrids all performed positive heterosis and the heterosis were especially prominent in the stage of seedling and end fruiting. In different stages, the leaf area all correlated with the yield character at significant or even very significant level. The hybrids showed positive heterosis in the whole life. The character of chlorophyll b in the middle fruiting stage and the chlorophyll a/b in the end fruiting stage were significantly and positively correlated with the yield character and the hybrids had
heterosis performance. In conclusion, the hybrids had larger leaf area, they could utilize light especially weak light more usefully, and in the growth stage long the plant could always maintain higher photosynthesis rate, which may become one of the important physiological basis of heterosis performing.
As for the dry matter content in the plant, he hybrids all showed heterosis performance compared with their parents at different level in the whole growth stage. The heterosis performance were especially obvious in the three fruiting stage, whose heterosis index were all above 1.40. In the former growth stage, the dry matter in the vegetation organ of the hybrids showed strong heterosis compared with their parent, so did that in the fruit organ during the middle and end fruiting stage, which may be another important course of yield hetersosis forming.
From the analysis of the nitrogen, phosphorus and potassium nutrition accumulation and division, we could see that during the former growth stage, there were not much difference between hybrids and their parents, while in the fruit stage, the hybrids absorbed the nutrition faster and more stably than their parents. In the early growth stage, the nutrition accumulated in the vegetation organ and in the latter stage that in the fruit of hybrids all showed great heterosis. So, the hybrids could utilize the nutrition more rationally and efficently. These may help to form yield heterosis.
3. The negative but not significant correlation between the genetic distance and parents showed that it wasn't the case that the greater the genetic distance were, the better.
The relationship between hybrids and their parents indicated that the performance of agronomical characters in the hybrids were all affected by their parents at different level, so the heterosis
1.农艺学性状相关分析及杂种一代优势表现分析表明，单株结果数、单果重、果实发育速度、雌花数、座果率、果长、果粗、早期单株产量、早期单株结果数、早期单果重等性状与单株产量呈显著或极显著正相关，且杂种一代均表现为正向优势，以单株产量、果实发育速度两个性状优势尤为明显，平均超亲优势指数均为1.61。始花期、始收期与单株产量呈显著负相关，杂种一代表现为负向优势，平均超中优势指数分别为0.96和0.97。杂种一代与单株产量密切相关的农艺学性状较其亲本均有不同程度的优势表现，构成产量优势形成的农艺学基础。
2.苗期、结果中期、结果末期光合速率与单株产量呈显著或极显著正相关，且杂种一代均表现为正向优势，以苗期和结果末期优势尤为明显。不同生长发育期叶面积与单株产量相关性均达显著或极显著水平，且杂种一代叶面积较其亲本均表现出明显的正向优势。结果中期叶绿素b含量、结果末期叶绿素a/b与单株产量的相关性达显著水平，而且杂种一代较其亲本表现为正向优势。杂种一代较其亲本拥有更大的光合面积，对光照尤其是弱光利用率高，整个生长发育期植株均能维持较高的光合速率，可能是杂种优势形成的重要生理基础之一。
整个生长发育期，杂种一代植株中干物质含量较其亲本均表现出不同程度的正向优势，以结果期优势更为明显,优势指数均在1.40以上。生长前期杂种一代干物质在营养器官中积累分配和结果中期、结果末期在果实中积累分配较亲本均表现出明显优势，可能是产量优势形成的又一重要原因。
从氮、磷、钾养分积累分配分析可以看出，苗期杂种一代与其亲本无明显差异，结果期，杂种一代均表现出较其亲本更快的营养元素积累能力，
生长前期营养器官和后期果实中养分积累分配强优势表现表明，杂种一代较其亲本能够更为合理的分配氮、磷、钾营养成分，养分利用率高，有助于产量优势形成。
3.遗传距离与单株产量的优势指数呈不显著程度的负相关，表明遗传距离并非越大越好。
亲子相关分析表明，杂种一代农艺学性状均不同程度的受到亲本影响，可以通过亲本性状表现对杂种一代与亲本密切相关的性状进行初步预测，育种时应注意以适当的品种作亲本。
In this experiment, seven inbreds which had different genetic backgroud together with seven hybrids breded with them were studied to find the agronomical, physiological and genetic mechanism of heterosis performance. The aim was to provide theoretical basis for heterosis utility, parent selection and heterosis prodiction. The result were as follows:
1. Correlation analysis between agronomical characters and heterosis performance in the hybrids indicated that the fruit number per plant, fruit weight, fruit growth rate, female flower number, percentage of fruit setting, early yield per plant, early fruit number per plant, early fruit weight length of fruit and diameter of fruit all had positive correlation with the yield per plant at significant or very significant level. The hybrids showed positive mid-parent heterosis and even super-parent heterosis in these characters. Among them, the heterosis of the yield per plant and the fruit growth rate were much higher, whose mid-parent heterosis index reached 1.61, and super-parent heterosis index reached 1.53 and 1.47 respectively. However, the period from planting to flowering and from planting to first harvesting. had negative relationship with the yield per plant and the hybrids showed negative heterosis. Their mid-parent heterosis index were 0.96 and 0.97 respectively. In a word, all of the characters which were closely correlated with the yield per plant had heterosis performance compared with their parents, this may be the agronomical basis of heterosis forming.
2.In the stage of seedling, middle fruiting and end fruiting, the correlation coefficient between the photosynthetic rate and the yield per plant reached significant even very significant level. The hybrids all performed positive heterosis and the heterosis were especially prominent in the stage of seedling and end fruiting. In different stages, the leaf area all correlated with the yield character at significant or even very significant level. The hybrids showed positive heterosis in the whole life. The character of chlorophyll b in the middle fruiting stage and the chlorophyll a/b in the end fruiting stage were significantly and positively correlated with the yield character and the hybrids had
heterosis performance. In conclusion, the hybrids had larger leaf area, they could utilize light especially weak light more usefully, and in the growth stage long the plant could always maintain higher photosynthesis rate, which may become one of the important physiological basis of heterosis performing.
As for the dry matter content in the plant, he hybrids all showed heterosis performance compared with their parents at different level in the whole growth stage. The heterosis performance were especially obvious in the three fruiting stage, whose heterosis index were all above 1.40. In the former growth stage, the dry matter in the vegetation organ of the hybrids showed strong heterosis compared with their parent, so did that in the fruit organ during the middle and end fruiting stage, which may be another important course of yield hetersosis forming.
From the analysis of the nitrogen, phosphorus and potassium nutrition accumulation and division, we could see that during the former growth stage, there were not much difference between hybrids and their parents, while in the fruit stage, the hybrids absorbed the nutrition faster and more stably than their parents. In the early growth stage, the nutrition accumulated in the vegetation organ and in the latter stage that in the fruit of hybrids all showed great heterosis. So, the hybrids could utilize the nutrition more rationally and efficently. These may help to form yield heterosis.
3. The negative but not significant correlation between the genetic distance and parents showed that it wasn't the case that the greater the genetic distance were, the better.
The relationship between hybrids and their parents indicated that the performance of agronomical characters in the hybrids were all affected by their parents at different level, so the heterosis
引文
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78. M.F.Zhao, Relationship between molecular marker heterozygosity and hybrid performance in intra- and inter-subspecific crosses of rice. Plant Breeding, 1999(118):139-144
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79. 1993,63(8):484-488
80. Mitchell, Interval mappong of vability loci causing beterosis in Arabidopsis. Genetics, 1995(140):1105-1109
81. Murayama, Agronomic performance of F1 hybrids of rice (Oryza sativa L) in japonica-indica crosses: Heterosis for and relationship between grain yield and related characters. Plant Production Science, 2002, 5(3):203-210.
82. Perenzin,F1 hybrid performance and parental diversity estimated using molecular markers. Euphytica, 1998, 100(1-3):;273-279
83. R.L.Lower, Gene action and heterosis for yield and vegetative characteristic in a cross between a gynoecious Picking cucumber inbred and a cucumis Sativus bar Fandwick inline, J.Amer Soc. Hort-Sci. 1982 ,107(1):75-78
84. Saghai, Correlation between molelcular marker distance and hybrid performance in US southern long grain rice. Crop Science, 1997(37):145-150
85. Saratha, Genetic inprovement is short season-soybean: Ⅰ Dry matter accumulation , partition and leaf area duration. Crop Science, 2001,41(2):1037-1048
86. Sarker , Heterosis for yield and yield attributes in 'WA' type rice hybrids (Oryza sativa L.) on multilocation performance. Crop Research (Hisar). 2001, 22(2): 262-266.
87. Sarker, Effect of nitrogen fertilization on photosynthetic characters and dry matter production in F1 hybrids of rice (Oryza sativa L.). Plant Production Science, 2002, 5(2) 131-138.
88. Shi Chunhai, Anslysis of genetic and genotype and environment interaction effect on heterosis of nutrientquality traits in indica hybrid rice. International-rice-research-notes, 1997,22(3):7-12
89. Smith, Similarities among a group of elite maize inbreds as measured by pedigree,F1 grain yield, heterosis and RFLPs. Theor. Appl. Genet., 1990(80):16-187
90. Srivastava, Heterosis and intergenomic complementation mitochondria, chlroplast and nucleus. Heterosis, reappraisal of theory and practice,1983:260-286
91. Stuer, Identification of genetic factors contributing to heterosis I a hybrid from two elite maize inbred using molecular markers.1992 (132):823-839
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99. Yu, Importance of epistasis as the genetic basis of F1 heterosis in an elite rice hybrid. Acad Sci., 1997,(94):9226-9231
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101. Zhang, Molecular divergence and hybrid performance in rice. Mol. Breed., 1995(1):133-142
102. Zhang, Molecular marker heterozygosity and hybrid performance in indic and japonica rice. Ther. Appl. Genet., 1996(93):1218-1224
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64. C.H.SHI, Analysis of imbryo,endosperm, cytoplamic and maternal effects for heterosis of protein and lysine content in indica hybrid rice. Plant Breeding, 1999(118):574-576
65. C.L.Morgan, Mid-parent advantage and heterosis in F1 hybrids of wheat from crosses among old and modern varieties. Journal of Agricultural Science, 1998,(130):287-295
66. G.J.Schwab, Growth and nutrient uptaken by cotton root under field conditions. Commu. Soil. Sci. Plant Anal., 2000,31(1/2):149-164
Grant, Heterosis and combining ability estimates in spring oilseed
67. rape(Brassica napus L.) Can J Genet Cyto, 1985 (27):472-478
68. H.Caballos, Additive ,dorminant and epistatic effects for maize grain yield in acid and non-acid soils. Theor.Appl.Genet., 1998(96):662-668
69. Harsh Mehta, 玉米光合作用、产量和产量组合的杂种优势. 国外农学-杂粮作物，1993（3）：8-11
70. Hirao, Characteristic of leaf photosynthesis in Chinese F1 hybrid rice Oryzasativa]cultivar,Shanyou63.Japanese-Journal-of-Crop-Science(Japan). 1995,64(2):209-215
71. Jinshui Yang, Molecular basis of heterosis in hybrid rice and hybrid maize relealed by mRNA amplification. IRRI, 1996,21(1):12-13
72. K.C.Falk, Performance of inter-cultivar summer tunnip rape hybrids in Saskatchewan. Canadian Journal of Plant Science, 1994(74):441-445
73. K.S.Randhawa, Assessment of combining ability, heterosis and genetic variance for fruit characters in muskmelon. Indian J. Genet.,1990,50(2):127-130
74. Kidwell, Forage yield of alfalfa population derived from parent selected on the basis of molecular marker diversity. Crop Science,1999,(39):223-227
75. Kirk,The plastids, their chemistry, structure growth and inheritance. Elsevier,1978
76. L.L.Benchimol, Genetic diversity in tropical maize inbred lines: heterotic group assignment and hybrid performance determined by RFLP markers. Plant Breeding, 2000 (119):491-496
77. Le Gouis J, Diallel analysis of winter wheat at two nitrogen levels.Crop Science, 2002, 42(4). 1129-1134.
78. M.F.Zhao, Relationship between molecular marker heterozygosity and hybrid performance in intra- and inter-subspecific crosses of rice. Plant Breeding, 1999(118):139-144
Madrap.I.A., Heterosis in relation to combine-ability effect and phynotypic stability in sunflower. Indian J. Agric. Sci.,
79. 1993,63(8):484-488
80. Mitchell, Interval mappong of vability loci causing beterosis in Arabidopsis. Genetics, 1995(140):1105-1109
81. Murayama, Agronomic performance of F1 hybrids of rice (Oryza sativa L) in japonica-indica crosses: Heterosis for and relationship between grain yield and related characters. Plant Production Science, 2002, 5(3):203-210.
82. Perenzin,F1 hybrid performance and parental diversity estimated using molecular markers. Euphytica, 1998, 100(1-3):;273-279
83. R.L.Lower, Gene action and heterosis for yield and vegetative characteristic in a cross between a gynoecious Picking cucumber inbred and a cucumis Sativus bar Fandwick inline, J.Amer Soc. Hort-Sci. 1982 ,107(1):75-78
84. Saghai, Correlation between molelcular marker distance and hybrid performance in US southern long grain rice. Crop Science, 1997(37):145-150
85. Saratha, Genetic inprovement is short season-soybean: Ⅰ Dry matter accumulation , partition and leaf area duration. Crop Science, 2001,41(2):1037-1048
86. Sarker , Heterosis for yield and yield attributes in 'WA' type rice hybrids (Oryza sativa L.) on multilocation performance. Crop Research (Hisar). 2001, 22(2): 262-266.
87. Sarker, Effect of nitrogen fertilization on photosynthetic characters and dry matter production in F1 hybrids of rice (Oryza sativa L.). Plant Production Science, 2002, 5(2) 131-138.
88. Shi Chunhai, Anslysis of genetic and genotype and environment interaction effect on heterosis of nutrientquality traits in indica hybrid rice. International-rice-research-notes, 1997,22(3):7-12
89. Smith, Similarities among a group of elite maize inbreds as measured by pedigree,F1 grain yield, heterosis and RFLPs. Theor. Appl. Genet., 1990(80):16-187
90. Srivastava, Heterosis and intergenomic complementation mitochondria, chlroplast and nucleus. Heterosis, reappraisal of theory and practice,1983:260-286
91. Stuer, Identification of genetic factors contributing to heterosis I a hybrid from two elite maize inbred using molecular markers.1992 (132):823-839
92. T.A.Campbell, Melocular analysis of genetic relateness among alfalfa clones differing in levels of self-incomparibility. Canadian Journal of Plant Science, 2000,80(3):551-557
93. Tsaftaris, Studying the expression of genes in maize parental inbreds and their heterotic and nonheterotic hybrids. In proc XII Eucarpia maize and sorghum conference. Bergamo Italy, 1993:283-292
94. WELLSR, Heterosis in upland cotton.Ⅰ.Growth and leaf area partition. Crop Science, 1986(26):1119-1123
95. WELLSR, Heterosis in upland cotton.Ⅱ. Relationship of leaf area to plant photosynthesis. Crop Science, 1988(28):522-525
96. Wolf D P, Triple testcross analysis to detecte epistasis in maize. Crop Sci., 1997(37):763-770
97. Xiao J H, Dominace is the major genetic basis of heterosis in rice as revealed by QTL analysis using melocular markers. Genetics, 1995(140):745-754
98. Xiong, Relationships of differential gene expression in leaves with heterosis and heterozygosity in a rice diallel cross. Melocular Breeding, 1998(4):129-136
99. Yu, Importance of epistasis as the genetic basis of F1 heterosis in an elite rice hybrid. Acad Sci., 1997,(94):9226-9231
100. Zhang, A diallel analysis of heterosis in elite hybrid rice based on RFLPs and microsatellites. Ther. Appl. Genet., 1994(83):185-192
101. Zhang, Molecular divergence and hybrid performance in rice. Mol. Breed., 1995(1):133-142
102. Zhang, Molecular marker heterozygosity and hybrid performance in indic and japonica rice. Ther. Appl. Genet., 1996(93):1218-1224