北方杂交粳稻异交结实机制及高产制种技术研究
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摘要
利用奥林帕斯IX81荧光倒置显微镜和DP71单芯片彩色CCD相机软件,研究了杂交粳稻不育系柱头长度、宽度、面积、体积、开颖外露率、闭颖外露率和花柱长度、夹角、不育系开颖角度、午前花率及父本恢复系的产粉量;结合人工气候箱对恢复系花粉活力进行检测;并在杂交粳稻制种田调查了不育系和恢复系开花习性,分析了花期预测与调节技术,对不育系的柱头活力进行测定。在此基础上系统研究了北方杂交粳稻制种中对异交结实的影响因素和形成机制以及异交结实率的变化规律、大面积杂交粳稻制种的花期预测和花期调节方法,旨在为北方杂交粳稻高产制种提供一定的理论指导。主要研究结果如下:
1.不育系间和恢复系间的花时和花期存在着品种间差异,在选择确定一对母本不育系和恢复系制种时,要实现不育系和恢复系的花时、花期相遇的优化组合,以保证花期相遇。不同不育系和不同恢复系花期性状存在着明显的品种间差异,因此,应改变过去传统的花期相遇标准即以始穗期来衡量花期是否相遇,而是采用开花高峰期相遇为标准。
2.参试不育系的柱头生活力系数的大小依次为:辽846A、辽02A、辽60A、辽99A、辽30A、辽39A、辽105A、辽5216A、屉锦A、秀岭A,相应的不育系异交结实率的趋势与此相近。说明不育系的柱头生活力系数越强,不育系异交结实率越高。
3.不育系花柱夹角与柱头闭颖外露率呈显著线性正相关,不育系柱头开颖外露率与柱头闭颖外露率呈极显著线性正相关,柱头面积、柱头长度、柱头宽度、花柱长度、颖花开颖角度与柱头闭颖外露率均呈正相关。不育系花柱夹角越大,其柱头闭颖外露率越高;不育系柱头开颖外露率越高,其柱头闭颖外露率越高。不育系柱头闭颖外露率和柱头开颖外露率的高低并非决定于某一个性状,而是决定于各性状综合作用的结果。
不育系柱头闭颖外露率、柱头开颖外露率、午前花率、柱头体积与异交结实率之间存在显著或极显著的正相关。异交结实率与柱头闭颖外露率、柱头开颖外露率、午前花率、柱头体积之间的复相关系数达到显著水平。不育系的异交结实率与柱头闭颖外露率、柱头开颖外露率、午前花率、柱头体积的多元回归方程为:y=-9.44+0.36x1+0.17x2+0.20x3+1.41x4(P＜0.01,回归方程极显著)。要达到较高不育系异交结实率,应加强对柱头闭颖外露率、柱头开颖外露率、午前花率、柱头体积的正向选择,特别加强对柱头闭颖外露率的选择。不育系柱头闭颖外露率对异交结实率的直接通径系数最大,通过柱头开颖外露率、午前花率、柱头体积的间接通径系数较小,柱头闭颖外露率可以在较大程度上直接影响不育系异交结实率;不育系柱头开颖外露率、午前花率、柱头体积对异交结实率的直接效应较小,而通过柱头闭颖外露率对异交结实率的间接效应较大。在杂交粳稻不育系选育和高产制种方面,要把重点放在提高不育系的柱头闭颖外露率上,同时兼顾提高柱头开颖外露率、午前花率和选择较大体积柱头。
4.恢复系花粉活力随离体时间的延长而降低,且在33℃以上高温胁迫下,处理时间越长,下降幅度也越大。不同杂交粳稻恢复系高温伤害的程度存在明显差异,其原因可能与恢复系的籼性程度有关。水稻花粉不耐高温,花粉活力随离体时间的延长而降低,且在33℃以上高温胁迫下,处理时间越长,下降幅度也越大。
恢复系产粉量与花药长度呈极显著线性正相关,与花药宽度正相关系数虽然也很高,但未达到显著水平;恢复系产粉量与花药面积呈极显著线性正相关,恢复系产粉量与花药体积呈极显著线性正相关。这说明花药的长度、宽度、面积以及体积越大,恢复系的产粉量越大,同时说明杂交粳稻花药的长度、宽度、面积以及体积与品种遗传性关系很大。
5.采用幼穗剥查法、叶龄回归预测法、叶龄余数预测法、叶片日均长速预测法对花期进行预测,取得了很好效果。应用叶龄余数预测,要根据第二双零叶和伸长叶枕距来判断倒三叶,就能避免叶龄余数预测的不准确性。
6.在杂交粳稻父本穗分化Ⅲ期,通过控水、增N、施用多效唑、割叶等多种处理延缓穗分化进程,最大可调节花期7d左右。单项措施的效应为割叶＞施多效唑＞增N＞排水。多项措施的累加效应大于单项措施效应,但其总效应小于各单项措施效应之和。割叶、施多效唑皆对父本植株高度和单株花粉量有显著负向影响。过量增施多效唑,导致颍花退化,造成穗顶部秃尖。割叶、施多效唑与增施N肥结合,有互补效应。
Using OlympusⅨ81 fluorescence covert microscope with its DP71 single-chip color CCD camera software,the physical characteristics of female parent sterile lines of pistils and glumes were measured,pollen amount of male parent restorer lines were analyzed and calculated,and the pollen viability of male parent restorer lines were investigated in artificial climate box.Stigma vigor of sterile lines were determined,and the flowering habits,the prediction and adjustment of flowering period of sterile lines in hybrid seed field and male restorer lines were investigated also.Influencing factors and formation mechanismes of ourcrossing and variation of outcrossing rate were systematicly studied on hybrid japonica rice seed production in Northern China.Meanwhile the outcrossing characteristics,and the methods of prediction and adjustments of flowering with large-area hybrid japonica rice seed production and were studied,with a view to provide a theoretical guidance for high-yielding hybrid rice seed production.
1.There were variety differences in blooming time and blooming period among different sterile lines and male restorer lines.If a couple of CMS(sytoplasm male sterile) and male restorer line were chosen,that the optimum combination of the parents' blooming time and blooming period should been realized to ensure synchronization of flowering.
There were obvious variety differences in blooming period characteristics among different sterile lines and restorer lines.The traditional standard of synchronization of flowering which using initial heading as judgement should be changed.And full-bloom stage should be used as the new standard for synchronization flowering.
2.The coefficients of stigma vigour of sterile lines were as followes from strong to weak: Liao 846A,Liao 02A,Liao 60A,Liao 99A,Liao 30A,Liao 39A,Liao 105A,Liao 5216A, Tijin A,Xiuling A.The trend of outcrossing rate of corresponding sterile lines was similar with the coefficients of stigma vigour.It means that the stronger the coefficients of stigma,the higher the outcrossing rate of sterile lines.
3.Correlative analysis showed that there was a significantly positive correlation between stigma angle and stigma exsertion rate under closed-glume of sterile lines.There was a linear positive correlation between stigma exsertion rate under opened-glume and stigma exsertion rate under closed-glume of sterile lines.There were positive correlations between sigma area, stigma length,stigma width,opened-glume angle and stigma exsertion rate under closed-glume.Truthes were like that the bigger the stigma angle of sterile,the higher the stigma exsertion rate under closed-glume,and the higher the stigma exsertion rate under opened-glume,the higher the stigma exsertion rate under closed-glume.The stigma exsertion rate under closed-glume and the stigma exsertion rate under opened-glume didn't depend on someone characteristic but on comprehensive effects of all characteristics.
There were significantly and extremely significant positive correlations between stigma exsertion rate under closed-glume,stigma exsertion rate under opened-glume,flowering rate before noon,stigma volume of sterile lines and outcrossing rate.There were negative correlations between outcrossing rate and stigma exsertion rate under closed-glume,stigma exsertion rate under opened-glume,flowering rate before noon,stigma volume.The outcrossing rate,stigma exsertion rate under closed-glume,stigma exsertion rate under opened-glume,flowering rate before noon and stigma volume of sterile lines built up a multiple regression equation as:y=-9.44+0.36x1+0.17x2+0.20x3+1.41x4(p＜0.01,the regression equation was extremely significant).To achieve a higher outcrossing rate of sterile lines,the stigma exsertion rate under opened-glume,flowering rate before noon and stigma volume,especially stigma exsertion rate under closed-glume should be positively chosen.The direct path coefficient of stigma exsertion rate under closed-glume to outcrossing rate was the biggest.Because the indirect path coefficients of stigma exsertion rate under opened-glume, flowering rate before noon and stigma volume were smaller,so the outcrossing rate of sterile lines in a greater degree.The direct effects of stigma exsertion rate under opened-glume, flowering rate before noon and stigma volume of sterile lines on outcrossing rate were smaller, but the indirect effect of stigma exsertion rate under closed-glume of sterile lines on outcrossing rate was bigger.In the aspects of sterile lines choosing and high-yielding seed production of hybrid japonica rice,the key point was improving the stigma exsertion rate under closed-glume of sterile lines,meanwhile improving stigma exsertion rate under opened-glume,flowering rate and choosing stigma with bigger volume.
4.The pollen viability of six male restorer lines was decreased with off-body time prolonged,and under the treatment of over 33℃high temperature stress,the longer the treatment time,the more descent range the viability.There were significant differences of injury degrees among different hybrid japonica rice restorer lines,cause of their different indica-compatibility.Rice pollen couldn't tolerate high temperature.Pollen viability was decreased with off-body time prolonged.Under the treatment of over 33℃high temperature stress,the longer was the treatment time,the more was the descent range.There were significant differences in heat tolerance of pollen viability among different hybrid japonica rice restorer lines.It could be used for improving the heat tolerance of hybrid japonica rice restorer lines and breeding new varieties.
There was a extremely significant linear positive correlation between pollen amount and stigma length of hybrid japonica rice restorer lines.Though the correlation coefficient of pollen amount and stigma width was big,it did not reach significant level.There was a very significantly linear positive correlation between pollen amount and anther area of hybrid japonica rice restorer lines.There was a very significantly linear positive correlation between pollen amount and anther volume of hybrid japonica rice restorer lines.It meant that the larger the anther length,anther width,anther area and anther volume,the more the pollen amount of hybrid japonica rice restorer lines.Meanwhile it meant that anther length,anther width,anther area and anther volume were closely related to heredity of varieties.
5.Hybrid japonica rice seed production was forecasted by stripping and counting young panicle,leaf-age regression,leaf-age remainder and daily growing velocity,which all obtained good results.By leaf-age remainder forecast,the top 3rd leaves should be judged by second bi-zero leaf and GDOP,to avoid the inaccuracy of leaf-age remainder forecast.
6.In the panicle initiation stageⅢof male parent hybrid japonica rice C4115,the spike differentiation process was delayed by controlling water,increasing N,using Paclobutrazol,picking off leaves treatment and so on.The effect of any single treatment was as followes:by picking off leaves＞by using Paclobutrazol＞by increasing N＞by controlling water.Additive effects of many treatments were better than that of any single one,but the additive effects were less than the sum of all the single treatment.There were significant negative effects on plant height and pollen amount per plant by picking off leaves and using Paclobutrzol.Using excessive Paclobutrazol resulted in spike degradation and bear top. Picking off leaves and using Paclobutrazol combined with increasing N showed complementary effects.
1.不育系间和恢复系间的花时和花期存在着品种间差异,在选择确定一对母本不育系和恢复系制种时,要实现不育系和恢复系的花时、花期相遇的优化组合,以保证花期相遇。不同不育系和不同恢复系花期性状存在着明显的品种间差异,因此,应改变过去传统的花期相遇标准即以始穗期来衡量花期是否相遇,而是采用开花高峰期相遇为标准。
2.参试不育系的柱头生活力系数的大小依次为:辽846A、辽02A、辽60A、辽99A、辽30A、辽39A、辽105A、辽5216A、屉锦A、秀岭A,相应的不育系异交结实率的趋势与此相近。说明不育系的柱头生活力系数越强,不育系异交结实率越高。
3.不育系花柱夹角与柱头闭颖外露率呈显著线性正相关,不育系柱头开颖外露率与柱头闭颖外露率呈极显著线性正相关,柱头面积、柱头长度、柱头宽度、花柱长度、颖花开颖角度与柱头闭颖外露率均呈正相关。不育系花柱夹角越大,其柱头闭颖外露率越高;不育系柱头开颖外露率越高,其柱头闭颖外露率越高。不育系柱头闭颖外露率和柱头开颖外露率的高低并非决定于某一个性状,而是决定于各性状综合作用的结果。
不育系柱头闭颖外露率、柱头开颖外露率、午前花率、柱头体积与异交结实率之间存在显著或极显著的正相关。异交结实率与柱头闭颖外露率、柱头开颖外露率、午前花率、柱头体积之间的复相关系数达到显著水平。不育系的异交结实率与柱头闭颖外露率、柱头开颖外露率、午前花率、柱头体积的多元回归方程为:y=-9.44+0.36x1+0.17x2+0.20x3+1.41x4(P＜0.01,回归方程极显著)。要达到较高不育系异交结实率,应加强对柱头闭颖外露率、柱头开颖外露率、午前花率、柱头体积的正向选择,特别加强对柱头闭颖外露率的选择。不育系柱头闭颖外露率对异交结实率的直接通径系数最大,通过柱头开颖外露率、午前花率、柱头体积的间接通径系数较小,柱头闭颖外露率可以在较大程度上直接影响不育系异交结实率;不育系柱头开颖外露率、午前花率、柱头体积对异交结实率的直接效应较小,而通过柱头闭颖外露率对异交结实率的间接效应较大。在杂交粳稻不育系选育和高产制种方面,要把重点放在提高不育系的柱头闭颖外露率上,同时兼顾提高柱头开颖外露率、午前花率和选择较大体积柱头。
4.恢复系花粉活力随离体时间的延长而降低,且在33℃以上高温胁迫下,处理时间越长,下降幅度也越大。不同杂交粳稻恢复系高温伤害的程度存在明显差异,其原因可能与恢复系的籼性程度有关。水稻花粉不耐高温,花粉活力随离体时间的延长而降低,且在33℃以上高温胁迫下,处理时间越长,下降幅度也越大。
恢复系产粉量与花药长度呈极显著线性正相关,与花药宽度正相关系数虽然也很高,但未达到显著水平;恢复系产粉量与花药面积呈极显著线性正相关,恢复系产粉量与花药体积呈极显著线性正相关。这说明花药的长度、宽度、面积以及体积越大,恢复系的产粉量越大,同时说明杂交粳稻花药的长度、宽度、面积以及体积与品种遗传性关系很大。
5.采用幼穗剥查法、叶龄回归预测法、叶龄余数预测法、叶片日均长速预测法对花期进行预测,取得了很好效果。应用叶龄余数预测,要根据第二双零叶和伸长叶枕距来判断倒三叶,就能避免叶龄余数预测的不准确性。
6.在杂交粳稻父本穗分化Ⅲ期,通过控水、增N、施用多效唑、割叶等多种处理延缓穗分化进程,最大可调节花期7d左右。单项措施的效应为割叶＞施多效唑＞增N＞排水。多项措施的累加效应大于单项措施效应,但其总效应小于各单项措施效应之和。割叶、施多效唑皆对父本植株高度和单株花粉量有显著负向影响。过量增施多效唑,导致颍花退化,造成穗顶部秃尖。割叶、施多效唑与增施N肥结合,有互补效应。
Using OlympusⅨ81 fluorescence covert microscope with its DP71 single-chip color CCD camera software,the physical characteristics of female parent sterile lines of pistils and glumes were measured,pollen amount of male parent restorer lines were analyzed and calculated,and the pollen viability of male parent restorer lines were investigated in artificial climate box.Stigma vigor of sterile lines were determined,and the flowering habits,the prediction and adjustment of flowering period of sterile lines in hybrid seed field and male restorer lines were investigated also.Influencing factors and formation mechanismes of ourcrossing and variation of outcrossing rate were systematicly studied on hybrid japonica rice seed production in Northern China.Meanwhile the outcrossing characteristics,and the methods of prediction and adjustments of flowering with large-area hybrid japonica rice seed production and were studied,with a view to provide a theoretical guidance for high-yielding hybrid rice seed production.
1.There were variety differences in blooming time and blooming period among different sterile lines and male restorer lines.If a couple of CMS(sytoplasm male sterile) and male restorer line were chosen,that the optimum combination of the parents' blooming time and blooming period should been realized to ensure synchronization of flowering.
There were obvious variety differences in blooming period characteristics among different sterile lines and restorer lines.The traditional standard of synchronization of flowering which using initial heading as judgement should be changed.And full-bloom stage should be used as the new standard for synchronization flowering.
2.The coefficients of stigma vigour of sterile lines were as followes from strong to weak: Liao 846A,Liao 02A,Liao 60A,Liao 99A,Liao 30A,Liao 39A,Liao 105A,Liao 5216A, Tijin A,Xiuling A.The trend of outcrossing rate of corresponding sterile lines was similar with the coefficients of stigma vigour.It means that the stronger the coefficients of stigma,the higher the outcrossing rate of sterile lines.
3.Correlative analysis showed that there was a significantly positive correlation between stigma angle and stigma exsertion rate under closed-glume of sterile lines.There was a linear positive correlation between stigma exsertion rate under opened-glume and stigma exsertion rate under closed-glume of sterile lines.There were positive correlations between sigma area, stigma length,stigma width,opened-glume angle and stigma exsertion rate under closed-glume.Truthes were like that the bigger the stigma angle of sterile,the higher the stigma exsertion rate under closed-glume,and the higher the stigma exsertion rate under opened-glume,the higher the stigma exsertion rate under closed-glume.The stigma exsertion rate under closed-glume and the stigma exsertion rate under opened-glume didn't depend on someone characteristic but on comprehensive effects of all characteristics.
There were significantly and extremely significant positive correlations between stigma exsertion rate under closed-glume,stigma exsertion rate under opened-glume,flowering rate before noon,stigma volume of sterile lines and outcrossing rate.There were negative correlations between outcrossing rate and stigma exsertion rate under closed-glume,stigma exsertion rate under opened-glume,flowering rate before noon,stigma volume.The outcrossing rate,stigma exsertion rate under closed-glume,stigma exsertion rate under opened-glume,flowering rate before noon and stigma volume of sterile lines built up a multiple regression equation as:y=-9.44+0.36x1+0.17x2+0.20x3+1.41x4(p＜0.01,the regression equation was extremely significant).To achieve a higher outcrossing rate of sterile lines,the stigma exsertion rate under opened-glume,flowering rate before noon and stigma volume,especially stigma exsertion rate under closed-glume should be positively chosen.The direct path coefficient of stigma exsertion rate under closed-glume to outcrossing rate was the biggest.Because the indirect path coefficients of stigma exsertion rate under opened-glume, flowering rate before noon and stigma volume were smaller,so the outcrossing rate of sterile lines in a greater degree.The direct effects of stigma exsertion rate under opened-glume, flowering rate before noon and stigma volume of sterile lines on outcrossing rate were smaller, but the indirect effect of stigma exsertion rate under closed-glume of sterile lines on outcrossing rate was bigger.In the aspects of sterile lines choosing and high-yielding seed production of hybrid japonica rice,the key point was improving the stigma exsertion rate under closed-glume of sterile lines,meanwhile improving stigma exsertion rate under opened-glume,flowering rate and choosing stigma with bigger volume.
4.The pollen viability of six male restorer lines was decreased with off-body time prolonged,and under the treatment of over 33℃high temperature stress,the longer the treatment time,the more descent range the viability.There were significant differences of injury degrees among different hybrid japonica rice restorer lines,cause of their different indica-compatibility.Rice pollen couldn't tolerate high temperature.Pollen viability was decreased with off-body time prolonged.Under the treatment of over 33℃high temperature stress,the longer was the treatment time,the more was the descent range.There were significant differences in heat tolerance of pollen viability among different hybrid japonica rice restorer lines.It could be used for improving the heat tolerance of hybrid japonica rice restorer lines and breeding new varieties.
There was a extremely significant linear positive correlation between pollen amount and stigma length of hybrid japonica rice restorer lines.Though the correlation coefficient of pollen amount and stigma width was big,it did not reach significant level.There was a very significantly linear positive correlation between pollen amount and anther area of hybrid japonica rice restorer lines.There was a very significantly linear positive correlation between pollen amount and anther volume of hybrid japonica rice restorer lines.It meant that the larger the anther length,anther width,anther area and anther volume,the more the pollen amount of hybrid japonica rice restorer lines.Meanwhile it meant that anther length,anther width,anther area and anther volume were closely related to heredity of varieties.
5.Hybrid japonica rice seed production was forecasted by stripping and counting young panicle,leaf-age regression,leaf-age remainder and daily growing velocity,which all obtained good results.By leaf-age remainder forecast,the top 3rd leaves should be judged by second bi-zero leaf and GDOP,to avoid the inaccuracy of leaf-age remainder forecast.
6.In the panicle initiation stageⅢof male parent hybrid japonica rice C4115,the spike differentiation process was delayed by controlling water,increasing N,using Paclobutrazol,picking off leaves treatment and so on.The effect of any single treatment was as followes:by picking off leaves＞by using Paclobutrazol＞by increasing N＞by controlling water.Additive effects of many treatments were better than that of any single one,but the additive effects were less than the sum of all the single treatment.There were significant negative effects on plant height and pollen amount per plant by picking off leaves and using Paclobutrzol.Using excessive Paclobutrazol resulted in spike degradation and bear top. Picking off leaves and using Paclobutrazol combined with increasing N showed complementary effects.
引文
1.曹文亮,肖晓,王仁祥,袁志峰,肖层林.2007.气象因素对两系法杂交稻制种安全和产量的影响.作物研究.21(5):571-573
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