早实核桃花器形态发育和果实性状遗传特性研究
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摘要
本文对核桃早实类群和晚实类群以及早实类群二次花的花芽形态发育规律进行了解剖观察。还以早实核桃品种元林和晚实核桃品种青林及其杂交获得的60个F1后代单株为试材,对果实性状的变异系数、广义遗传力、相关遗传力、遗传相关系数、相对遗传进度等遗传参数进行了估算。结果表明:
1.核桃花芽发育研究:
(1)核桃雌花芽的发育
核桃雌花芽的分化过程可以划分为:生理分化期、形态分化临界期、雌花序原基和花柄原基分化期、雌花原基分化期、苞片原基分化期、花萼原基和雌蕊原基分化期、幼胚分化期和雌花成熟期8个时期。在山东泰安地区,雌花于4月中旬至5月中上旬开始分化至竖年4月中旬至4月下旬成熟开放,历时约10个月。
(2)核桃雄花芽的发育
雄花芽分化主要经历以下7个阶段:鳞片分化期、雄花序分化期、雄花原基出现期、雄花花被分化期、雄蕊原基分化期、花药分化期、雄花成熟期。
在山东泰安地区雄花序鳞片分化期3月下旬在芽内开始;雄花序分化期在4月上、中旬随新稍伸长而出现:雄花原基出现期随着雄花序的分化,在基部鳞片状组织的基部出现雄花原基突起;雄花花被分化期5月上旬出现;雄蕊原基分化期在花被分化之后5天左右出现;花药分化期在6月下旬;雄花成熟期在翌年4月初萌发后形成。研究认为,雄花序分化期形成的鳞片状组织,其基部分化花原基,是“拟花托”组织,其前端具有花被的保护功能。
2.核桃果实性状的遗传特性
(1)核桃果实性状的变异
核桃杂交群体中果实性状存在着广泛的变异,尤其是单株结果数、单株坚果产量和单株果仁产量的变异系数高达90%以上。
(2)遗传参数
核桃果实性状的遗传力普遍较高,壳厚、仁重、干果横径、出仁率、单株坚果产量、内种皮色泽、单株果仁产量、青果横径、缝横径、单株结果数、青果鲜重、青果纵径的遗传力在0.716—0.998之间,湿坚果重、干果重、干果横径和壳重的遗传力在0.339—0.478之间。单株结果数、单株坚果产量和单株果仁产量的遗传变异系数(GCV)最大,果仁重、壳厚和鲜果重也具有较大的遗传变异系数。
(3)相关遗传力
核桃果实性状间的遗传相关系数多数达到显著或极显著,但性状间的相关遗传力普遍小于目标性状遗传力。单株坚果产量与单株果仁产量相关遗传力比果仁产量遗传力略大,单株坚果产量和单株果仁产量与单株结果数的相关遗传力比单株结果数的遗传力略大。
(4)通径分析
通径分析结果表明:单株干果产量、单株结果数、单果仁重、缝横径、青果纵径、干果纵径、壳厚对果仁产量的直接作用达到极显著或显著水平,干果产量和结果数的直接作用明显大于其它性状,壳厚对于果仁产量是负作用;多数性状主要通过干果产量对果仁产量产生作用,其次是通过结果数。选择果仁丰产单株时,干果产量和结果数是最重要的原因指标,通径分析的决策系数也表明干果产量是主要决策性状,而青果纵径是主要限制性状。
(5)选择结果
结合指数选择和直接选择得到1—6、5—16、4—2、5—11共4个果仁产量、出仁率、壳厚、内种皮色泽等性状优良的单株。
This paper studied the process of floral bud development of precocious walnut and later mature walnut as well as the secdonary bud development on morphology.We also studied the development of walnut complete flower in our country.Coefficient of variability,broad-sense heritability,correlated heritability,genetic correlation coefficients,relative genetic schedule were calculated for analyzing the genetic character of walnut fruits which were gotten from precocious walnut variety Lüyuan,later mature variety Lüfeng and their 60 F1 hybrid plants.The result showed that:
1.the reasearch about walnut floral bud development
(1) process of female bud development
The process of female bud development lasted about 10 months in Taian city,which started physiological differentiation from mid-April to early May or mid-May and blossomed in mid-April or late -April in second year can be divided into 8 stages:physiological differentiation stage,morphological differentiation critical stage,appearing stage of female inflorescence primordium and pedicel primordium,appearing stage of female flower primordium,appearing stage of bract primordium,appearing stage of calyx primordium and pistil primordia,appearing stage of immature embryo, mature period of female flower.
(2) process of male bud development
The process of male bud development can be divided into 7 stages: squama differentiation stage,male inflorescence differentiation stage, appearance of male flower primordium,appearance of male flower perianth, appearance of male flower primordium,anther differentiation stage,mature period of male flower.In taian city,the squama started differentiated in mixed bud in late March;Male inflorescence differentiated with new shoot elongation in early or mid-April;With the male inflorescence primordium developmented male flower primordium appeared on proximal of squama; Male flower perianth differentiated in early May;Stamen primordium differentiated after male flower perianth primordium appeared 5 days.Anther differentiated in late june;Male flower matured in mid-April or late April.We beieve:the squama organization formed in male inflorescence differentiation stage was similar to receptacle,on proximal of the squama the male flower primordium appeared,the front-end of the squama played the function of flower perianth.
2.the syudy on genetic parameter of walnut fruit traits
(1) the variation of walnut fruit traits
There was wide variation of fruit traits in walnut hybridization colony,especially the variation coefficient of fruit number of per plant and the nut yield of per plant and the kernel yield of per plant reached above 90%.
(2) the genetic parameters for fruit traits of walnut
Heritability of fruits were high widely,the heritability of shell thickness, kernel weight,nut transverse diameter,percent of kernel,nut yield of per plant, endopleura colour,kernel yield of per plant,fresh fruit transverse diameter, fresh fruit slot transverse diameter,fruit number of per plant,fresh fruit weight and fresh fruit vertical diameter were between 0.716 to 0.998.Only the heritability of moist nut weight,nut weight,nut transverse diameter,and shell weight were between 0.339 to 0.478.The variation coefficient of fruit number of per plant,nut yield of per plant and kernel yield of per plant were most great,the variation coefficient of kernel weight,shell thickness and fresh fruit weight were more bigger too.
(3) correlation heritability
The genetic correlation coefficients between walnut fruit traits are mostly significance at 1%level or at 5%level,but the correlated heritability between walnut fruit traits less than broad-sense heritability of target trait.The correlated heritability between nut yield of per plant and kernel yield of per plant was a little bigger than the heritability of kernel yield of per plant,the correlated heritability between nut yield of per plant and fruit number of per plant was a little bigger than the beritability of fruit number of per plant,the correlated heritability between kernel yield of per plant and fruit number of per plant was a little bigger than the heritability of fruit number of per plant.
(4) path analysis
Path analysis show that:the direct effect of fruit number of per plant,nut yield of per plant,fresh fruit vertical diameter,nut vertical diameter,nut slot transverse diameter,shell thickness,kernel weight to kernel yield of per plant are significance at 1%level or at 5%level.The direct effect of fruit number of per plant,nut yield of per plant to kernel yield of per plant were obviously greater than the other traits,the shell thickness played negative effect to kernel yield of per plant.Most traits affected the kernel yield of per plant through nut yield of per plant,then through fruit number of per plant.For kernel yield of per plant,the nut yield of per plant and fruit number of per plant is most important reasonable factor.Decision coefficient also show that the nut yield of per plant was master decision factor,however the fruit vertical diameter was master restriction factor.
(5)result of selecting
We identified 1-6、5-16、4-2、5-11 were superior tree for fruit trait using selection index and direct selection.
1.核桃花芽发育研究:
(1)核桃雌花芽的发育
核桃雌花芽的分化过程可以划分为:生理分化期、形态分化临界期、雌花序原基和花柄原基分化期、雌花原基分化期、苞片原基分化期、花萼原基和雌蕊原基分化期、幼胚分化期和雌花成熟期8个时期。在山东泰安地区,雌花于4月中旬至5月中上旬开始分化至竖年4月中旬至4月下旬成熟开放,历时约10个月。
(2)核桃雄花芽的发育
雄花芽分化主要经历以下7个阶段:鳞片分化期、雄花序分化期、雄花原基出现期、雄花花被分化期、雄蕊原基分化期、花药分化期、雄花成熟期。
在山东泰安地区雄花序鳞片分化期3月下旬在芽内开始;雄花序分化期在4月上、中旬随新稍伸长而出现:雄花原基出现期随着雄花序的分化,在基部鳞片状组织的基部出现雄花原基突起;雄花花被分化期5月上旬出现;雄蕊原基分化期在花被分化之后5天左右出现;花药分化期在6月下旬;雄花成熟期在翌年4月初萌发后形成。研究认为,雄花序分化期形成的鳞片状组织,其基部分化花原基,是“拟花托”组织,其前端具有花被的保护功能。
2.核桃果实性状的遗传特性
(1)核桃果实性状的变异
核桃杂交群体中果实性状存在着广泛的变异,尤其是单株结果数、单株坚果产量和单株果仁产量的变异系数高达90%以上。
(2)遗传参数
核桃果实性状的遗传力普遍较高,壳厚、仁重、干果横径、出仁率、单株坚果产量、内种皮色泽、单株果仁产量、青果横径、缝横径、单株结果数、青果鲜重、青果纵径的遗传力在0.716—0.998之间,湿坚果重、干果重、干果横径和壳重的遗传力在0.339—0.478之间。单株结果数、单株坚果产量和单株果仁产量的遗传变异系数(GCV)最大,果仁重、壳厚和鲜果重也具有较大的遗传变异系数。
(3)相关遗传力
核桃果实性状间的遗传相关系数多数达到显著或极显著,但性状间的相关遗传力普遍小于目标性状遗传力。单株坚果产量与单株果仁产量相关遗传力比果仁产量遗传力略大,单株坚果产量和单株果仁产量与单株结果数的相关遗传力比单株结果数的遗传力略大。
(4)通径分析
通径分析结果表明:单株干果产量、单株结果数、单果仁重、缝横径、青果纵径、干果纵径、壳厚对果仁产量的直接作用达到极显著或显著水平,干果产量和结果数的直接作用明显大于其它性状,壳厚对于果仁产量是负作用;多数性状主要通过干果产量对果仁产量产生作用,其次是通过结果数。选择果仁丰产单株时,干果产量和结果数是最重要的原因指标,通径分析的决策系数也表明干果产量是主要决策性状,而青果纵径是主要限制性状。
(5)选择结果
结合指数选择和直接选择得到1—6、5—16、4—2、5—11共4个果仁产量、出仁率、壳厚、内种皮色泽等性状优良的单株。
This paper studied the process of floral bud development of precocious walnut and later mature walnut as well as the secdonary bud development on morphology.We also studied the development of walnut complete flower in our country.Coefficient of variability,broad-sense heritability,correlated heritability,genetic correlation coefficients,relative genetic schedule were calculated for analyzing the genetic character of walnut fruits which were gotten from precocious walnut variety Lüyuan,later mature variety Lüfeng and their 60 F1 hybrid plants.The result showed that:
1.the reasearch about walnut floral bud development
(1) process of female bud development
The process of female bud development lasted about 10 months in Taian city,which started physiological differentiation from mid-April to early May or mid-May and blossomed in mid-April or late -April in second year can be divided into 8 stages:physiological differentiation stage,morphological differentiation critical stage,appearing stage of female inflorescence primordium and pedicel primordium,appearing stage of female flower primordium,appearing stage of bract primordium,appearing stage of calyx primordium and pistil primordia,appearing stage of immature embryo, mature period of female flower.
(2) process of male bud development
The process of male bud development can be divided into 7 stages: squama differentiation stage,male inflorescence differentiation stage, appearance of male flower primordium,appearance of male flower perianth, appearance of male flower primordium,anther differentiation stage,mature period of male flower.In taian city,the squama started differentiated in mixed bud in late March;Male inflorescence differentiated with new shoot elongation in early or mid-April;With the male inflorescence primordium developmented male flower primordium appeared on proximal of squama; Male flower perianth differentiated in early May;Stamen primordium differentiated after male flower perianth primordium appeared 5 days.Anther differentiated in late june;Male flower matured in mid-April or late April.We beieve:the squama organization formed in male inflorescence differentiation stage was similar to receptacle,on proximal of the squama the male flower primordium appeared,the front-end of the squama played the function of flower perianth.
2.the syudy on genetic parameter of walnut fruit traits
(1) the variation of walnut fruit traits
There was wide variation of fruit traits in walnut hybridization colony,especially the variation coefficient of fruit number of per plant and the nut yield of per plant and the kernel yield of per plant reached above 90%.
(2) the genetic parameters for fruit traits of walnut
Heritability of fruits were high widely,the heritability of shell thickness, kernel weight,nut transverse diameter,percent of kernel,nut yield of per plant, endopleura colour,kernel yield of per plant,fresh fruit transverse diameter, fresh fruit slot transverse diameter,fruit number of per plant,fresh fruit weight and fresh fruit vertical diameter were between 0.716 to 0.998.Only the heritability of moist nut weight,nut weight,nut transverse diameter,and shell weight were between 0.339 to 0.478.The variation coefficient of fruit number of per plant,nut yield of per plant and kernel yield of per plant were most great,the variation coefficient of kernel weight,shell thickness and fresh fruit weight were more bigger too.
(3) correlation heritability
The genetic correlation coefficients between walnut fruit traits are mostly significance at 1%level or at 5%level,but the correlated heritability between walnut fruit traits less than broad-sense heritability of target trait.The correlated heritability between nut yield of per plant and kernel yield of per plant was a little bigger than the heritability of kernel yield of per plant,the correlated heritability between nut yield of per plant and fruit number of per plant was a little bigger than the beritability of fruit number of per plant,the correlated heritability between kernel yield of per plant and fruit number of per plant was a little bigger than the heritability of fruit number of per plant.
(4) path analysis
Path analysis show that:the direct effect of fruit number of per plant,nut yield of per plant,fresh fruit vertical diameter,nut vertical diameter,nut slot transverse diameter,shell thickness,kernel weight to kernel yield of per plant are significance at 1%level or at 5%level.The direct effect of fruit number of per plant,nut yield of per plant to kernel yield of per plant were obviously greater than the other traits,the shell thickness played negative effect to kernel yield of per plant.Most traits affected the kernel yield of per plant through nut yield of per plant,then through fruit number of per plant.For kernel yield of per plant,the nut yield of per plant and fruit number of per plant is most important reasonable factor.Decision coefficient also show that the nut yield of per plant was master decision factor,however the fruit vertical diameter was master restriction factor.
(5)result of selecting
We identified 1-6、5-16、4-2、5-11 were superior tree for fruit trait using selection index and direct selection.
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