西藏半野生小麦Q1028种子强休眠特性研究
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摘要
西藏半野生小麦(Triticum aestivum ssp.tibetanum Shao)是中国西藏特有的一种小麦资源。它与我国栽培小麦地方品种相比较的特异性——脆穗和包壳特性已有许多研究,而它的另一重要特性——种子强休眠特性则报道很少。经过室内测试证明,所有西藏半野生小麦成熟期的种子发芽率为0%或接近于0%。本研究以Q1028为对象,对西藏半野生小麦种子强休眠特性进行分析。通过对Q1028蜡熟期的整穗发芽、颖壳提取液的种子发芽、常规种子发芽、破皮的种子发芽、种子切胚发芽及不同后熟时期的发芽测试,结果表明:Q1028的种子休眠性极强。蜡熟期的种子发芽率为0%,其休眠特性主要由于胚自身的休眠引起。
西藏半野生小麦Q1028与弱休眠小麦郑麦9023杂交组合的P_1、P_2、F_1、F_2、BC_1、BC_2六个世代进行了种子休眠遗传分析,结果表明:Q1028的种子休眠性极强,蜡熟期平均种子发芽率为0.75%,后熟8周的种子平均发芽率也仅有2.25%。遗传分析显示,西藏半野生小麦的强休眠特性主要受两对部分显性的互补主效基因控制,其中,至少有一对基因与红皮有关。根据后熟8周新增发芽率的测试与分析推测,除蜡熟期的两对基因控制种子的休眠强度外,可能还有另外的基因控制种子休眠维持的时间。
通过对易发芽和不易发芽极端群体的分子标记分析,将这两个休眠基因定位到3AL和1B上。位于3AL上的基因,可能已有研究者报道,暂命名为SD-1;位于1B上休眠基因则未见前人报道,暂时命名为SD-2。利用SSR标记将SD-1定位于Xgwm5、Xwmc651、Xwmc664、Xgwm155和SD-1的同一侧,他们的排列顺序是Xgwm5、Xwmc651、Xwmc664、Xgwm155和SD-1,该基因与这些引物的连锁距离分别是4.2CM、6.9CM、20.9CM、10.9CM。同时,分离群体上验证结果表明Xgwm498与休眠基因连锁,并将该休眠位点初步定位于1B上。用该标记对F_2代167个单株进行PCR扩增,根据扩增结果,采用Mapmaker3.0软件计算遗传距离,结果表明,该休眠位点与Xgwm498连锁距离为30.6CM。
根据F_2代蜡熟和后熟发芽发芽情况,选择出了30株种子强休眠单株,这些单株性状已经较Q1028有明显的改良,尤其是单株序号为123、135、257、278和279的单株,它们同时具有了种子强休眠、颖壳较软、和不易断穗等特性。这为以后选育具有种子强休眠特性的品种提供了宝贵的遗传材料。
T.aestivum ssp.tibetanum is a kind of special semi-wild hexaploid wheat resources,only distributing in Tibet of China.It has special characters,such as hulled glume and brittle spikelet.However,seed dormancy,another important character,was little reported.We have had observed and tested more than 10 accessions of them for dormancy,and found that every of them was 0 or near 0 percentages both with threshed seeds' and intact spikes' germinations.T.aestivum ssp.tibetanum cv.Q1028 was identified for seed dormant property by testing its' seed germination percentages of intact spikes,seeds with bract powder,normal seeds,seeds of pierced coat, sectioned embryos,and germination percentages of intact spikes and seeds at various stage after harvest.The result showed that the seed dormancy of Q1028 was very high. Its germination percentage of seed was 0%after harvest.The dormancy of Q1028 came from embryo dormancy itself mainly.
The cross of Q1028 with ZhengMai9023,which was non-dormant,was analyzed on P_1,P_2,F_1,F_2,BC_1 and BC_2 plants.The result showed that the seed dormancy of Q1028 was very high.Its germination percentage of seed was 0.75%at wax maturity.At eight weeks after maturity,its germination percentage of seed was only 2.25%.Its seed dormancy was controlled as partial dominant trait by two complementary genes.At least one gene between them was related with red capsule of seed.Besides,it was possible that other genes controlled maintaining time of seed dormancy.
Molecular markers analysis in the extremeness population,Using SSR markers, the two loci were located on 3AL and 1B,respectively.The one located on 3AL was named as SD-1,which possibly has been located by other researchers;the other one located on 1B was named as SD-2,which hasn't been reported before.Further molecular mapping showed that SD-1 was linked with SSR marker in the order of Xgwm5-Xwmc651-Xwmc664-Xgwm155-SD-1 with genetic distances of 4.2 cM、6.9 cM、20.9 cM、10.9 cM,respectively.At the same time,another one pair of primer located on chromosome arm 1BL,Xgwm498,were found being linked to strong seed dormancy.The 167 F_2 individuals were then amplified with marker Xgwm498.The statistic genetic distance between Xgwm498 and dormancy locus was calculated to be 30.6 cM using the software Mapmaker3.0.
According to F_2 generation germination data,the agronomical characters of 30 dormancy plants has been significantly improved than of Q1028,especially for No.135,No.123,No.135,No.257,No.278 and No.279.They have these characters such as flexible hull,dense spike and not brittle spikelet,beside strong seed dormancy. They are the precious genetic material for breeding.
西藏半野生小麦Q1028与弱休眠小麦郑麦9023杂交组合的P_1、P_2、F_1、F_2、BC_1、BC_2六个世代进行了种子休眠遗传分析,结果表明:Q1028的种子休眠性极强,蜡熟期平均种子发芽率为0.75%,后熟8周的种子平均发芽率也仅有2.25%。遗传分析显示,西藏半野生小麦的强休眠特性主要受两对部分显性的互补主效基因控制,其中,至少有一对基因与红皮有关。根据后熟8周新增发芽率的测试与分析推测,除蜡熟期的两对基因控制种子的休眠强度外,可能还有另外的基因控制种子休眠维持的时间。
通过对易发芽和不易发芽极端群体的分子标记分析,将这两个休眠基因定位到3AL和1B上。位于3AL上的基因,可能已有研究者报道,暂命名为SD-1;位于1B上休眠基因则未见前人报道,暂时命名为SD-2。利用SSR标记将SD-1定位于Xgwm5、Xwmc651、Xwmc664、Xgwm155和SD-1的同一侧,他们的排列顺序是Xgwm5、Xwmc651、Xwmc664、Xgwm155和SD-1,该基因与这些引物的连锁距离分别是4.2CM、6.9CM、20.9CM、10.9CM。同时,分离群体上验证结果表明Xgwm498与休眠基因连锁,并将该休眠位点初步定位于1B上。用该标记对F_2代167个单株进行PCR扩增,根据扩增结果,采用Mapmaker3.0软件计算遗传距离,结果表明,该休眠位点与Xgwm498连锁距离为30.6CM。
根据F_2代蜡熟和后熟发芽发芽情况,选择出了30株种子强休眠单株,这些单株性状已经较Q1028有明显的改良,尤其是单株序号为123、135、257、278和279的单株,它们同时具有了种子强休眠、颖壳较软、和不易断穗等特性。这为以后选育具有种子强休眠特性的品种提供了宝贵的遗传材料。
T.aestivum ssp.tibetanum is a kind of special semi-wild hexaploid wheat resources,only distributing in Tibet of China.It has special characters,such as hulled glume and brittle spikelet.However,seed dormancy,another important character,was little reported.We have had observed and tested more than 10 accessions of them for dormancy,and found that every of them was 0 or near 0 percentages both with threshed seeds' and intact spikes' germinations.T.aestivum ssp.tibetanum cv.Q1028 was identified for seed dormant property by testing its' seed germination percentages of intact spikes,seeds with bract powder,normal seeds,seeds of pierced coat, sectioned embryos,and germination percentages of intact spikes and seeds at various stage after harvest.The result showed that the seed dormancy of Q1028 was very high. Its germination percentage of seed was 0%after harvest.The dormancy of Q1028 came from embryo dormancy itself mainly.
The cross of Q1028 with ZhengMai9023,which was non-dormant,was analyzed on P_1,P_2,F_1,F_2,BC_1 and BC_2 plants.The result showed that the seed dormancy of Q1028 was very high.Its germination percentage of seed was 0.75%at wax maturity.At eight weeks after maturity,its germination percentage of seed was only 2.25%.Its seed dormancy was controlled as partial dominant trait by two complementary genes.At least one gene between them was related with red capsule of seed.Besides,it was possible that other genes controlled maintaining time of seed dormancy.
Molecular markers analysis in the extremeness population,Using SSR markers, the two loci were located on 3AL and 1B,respectively.The one located on 3AL was named as SD-1,which possibly has been located by other researchers;the other one located on 1B was named as SD-2,which hasn't been reported before.Further molecular mapping showed that SD-1 was linked with SSR marker in the order of Xgwm5-Xwmc651-Xwmc664-Xgwm155-SD-1 with genetic distances of 4.2 cM、6.9 cM、20.9 cM、10.9 cM,respectively.At the same time,another one pair of primer located on chromosome arm 1BL,Xgwm498,were found being linked to strong seed dormancy.The 167 F_2 individuals were then amplified with marker Xgwm498.The statistic genetic distance between Xgwm498 and dormancy locus was calculated to be 30.6 cM using the software Mapmaker3.0.
According to F_2 generation germination data,the agronomical characters of 30 dormancy plants has been significantly improved than of Q1028,especially for No.135,No.123,No.135,No.257,No.278 and No.279.They have these characters such as flexible hull,dense spike and not brittle spikelet,beside strong seed dormancy. They are the precious genetic material for breeding.
引文
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