穗重型小麦发育特性及穗粒重性状的遗传模型研究
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摘要
小麦是我国的第二大作物，对国家粮食安全具有举足轻重的地位。选育高产小麦新品种是增加小麦产量的有效途径。穗重型小麦具有突出的穗部性状和较高的单穗粒重，为我国超级小麦育种带来希望与曙光。然而，作为一个品种类型，穗重型小麦在性状发育特性、穗部性状的遗传传递规律及其分蘖和粒重等性状在发育过程中的动态遗传表现等方面尚缺乏系统的研究。针对这些问题，本研究以近年来黄淮麦区选育和引进的穗重型小麦基因型为试材，对穗重型小麦性状发育特性、穗重型小麦穗部性状遗传分析及分蘖、粒重性状的动态发育遗传规律进行了探讨，以期对穗重型小麦种质资源的合理利用与超级小麦新品种选育提供依据与参考。所取得的主要结果如下：
1、在群体发育方面，穗重型小麦冬季分蘖、春季分蘖及单位面积穗数都低于对照（多穗型品种），通过密度和播期调节的范围较窄；分蘖偏少的原因是形成一个分蘖所需要的积温更多，对低温的适应性更差。穗重型小麦形成较多小穗的原因在于：幼穗分化时期较短，表现为单棱期短，进入二棱期早，二棱期较长；幼穗分化速率快。穗重型小麦单茎干物质始终高于对照，单株干物质前期（开花前）高后期低，单位面积干物质积累偏少，且茎叶鞘输出的干物质多，向籽粒输入的少，输出输入效率较低。穗重型小麦净光合速率前（灌浆以前）高后低，呼吸速率与蒸腾速率均高于对照，根系吸收能力较强。穗重型小麦籽粒平均灌浆速率、最大灌浆速率均低于对照，灌浆持续期持平，粒鲜重和籽粒体积最大日增量不小于对照，决定穗重型小麦粒重的主要因素是灌浆速率，灌浆持续期起次要作用，平均灌浆速率、叶面积持续期与籽粒饱满指数显著正相关。穗重型小麦群体源相对不足，库容量较大，源、库之间的关系不够协调。穗重型小麦源不足表现为：群体偏小，群体生长率降低；光合产物大部分供应籽粒，叶片营养亏缺，导致叶片功能下降，光合产率降低。穗重型小麦冠层温度偏高，旗叶叶绿素水平偏低且下降迅速，叶片输出功能较低，无法适应潜在库容对光合产物的需要。
2、利用主基因-多基因混合遗传模型，对穗重型小麦3组合6世代的主要农艺性状分析表明，各性状的遗传模型因组合不同而表现有同有异，相同的情况是所有组合的单株穗数、株高和单穗粒重均一致地符合多基因的加性-显性-上位性遗传模型；符合这一遗传模型的还有：组合Ⅱ、组合Ⅲ的穗长，组合Ⅰ的每穗小穗数，组合Ⅱ的每穗粒数，这些性状均受多基因位点控制，且基因间存在上位性互作。表现各异的情况是：组合Ⅰ的穗长由2对具有加性-显性-上位性互作的主基因所控制，组合Ⅱ、Ⅲ的每穗小穗数由1对负向完全显性主基因+加性-显性多基因控制，组合Ⅰ、Ⅲ的每穗粒数由2对加性-显性-上位性主基因+加性-显性-上位性多基因共同控制，组合间出现的差异可能与亲本类型和遗传背景有关。
3、穗重型小麦分蘖性状同时受到遗传主效应和基因型×环境互作效应的共同控制，且以遗传主效应为主，其中加性效应和显性效应在分蘖形成的早期（0-7天）、早中期（8-14天）、中期（15-21天）表现得比较活跃，可以检测到显著的条件遗传效应方差分量。净光合速率的遗传以基因型方差为主，发育的中期（5月16日）以前，显性方差是遗传变异的主要分量，中后期加性方差为主要分量，灌浆（5月2日起）开始以后，大量的微效多基因被激活表达，从5月2日到5月16日，是控制净光合速率性状表现的基因表达最为活跃的时期；粒重形成以基因的加性效应、显性效应，显性与环境互作为主，环境对粒重形成的影响也达到显著水平，说明粒重极易受环境条件的影响；灌浆开始，加性效应初现，上位性效应依然存在，而环境效应最大；灌浆初期（5.10︱5.7），加性显性效应短暂消失，上位性效应持续增大；灌浆中期（5.13︱5.105.16︱5.13），加性显性效应再次出现，上位性效应短暂消失，但上位与环境互作却达到最大，说明此时粒重对环境影响非常敏感。
Wheat is the second largest crop planted in China, and plays a very important role fornational food security. The breeding of high-yielding varieties is an effective way to increasewheat production. The high spike weight wheat shows prominent ear characteristics and highgrain weight per ear, which bring hope and dawn for super wheat breeding in China. However,as a cultivar type, some traits of high spike weight wheat have not been systematic studied,such as the features of character development, genetic laws of typical panicle traits passing tothe offspring, dynamic genetic rule on grain weight and tiller traits in the developmentalprocess and so on. In response to these problems, we used high spike weight wheat genotype,breed or introduced to Huanghuai wheat area, as tested materials in this study to investigatetraits development features, spike genetic analysis, tillers and grain weight traits dynamicdevelopmental genetics rule of spike wheat, aiming to provide basis and references for therational use of high spike weight wheat germplasm resources and breeding of new superwheat varieties. The results obtained are as follows:
1. In Population development,the winter tillers, spring tillers and panicles in high spikeweight wheat were lower than those of in the control (panicle varieties), the scope ofregulation was more narrow by the density and sowing date, formation of a tiller needed moreaccumulated temperature, and adaptability to low temperature was worse, controlled bymulti-panicle varieties. High spike weight wheat spike differentiation period was generallyless than that of the control, a short single ridge stage period, the two-ridge stage period wasearlier and last longer. The spike differentiation rate also was faster than the control, whichresulted in more spikes of the high spike weight wheat. The single stem dry matteraccumulation was higher than that of the control, but single plant dry matter accumulationearly stage (before flowering) was higher than late stage, dry matter accumulation of per unitarea had no advantage. Most varieties of high spike weight wheat outputted more dry matterfrom the stem sheath and leaf sheath, but inputted less to the grain, input-output rate also waslow, indicating that some problems exited in its material flow. The net photosynthetic rate ofhigh spike weight wheat was higher than that of the control before grain filling and then reduced, but both of the respiratory rate and transpiration rate were higher, meaning that highspike weight wheat belonged to the high photosynthetic and high breathing consumption type,while its root absorption ability was better. The average grain filling rate and the maximumgrain-filling rate of high spike weight wheat were lower, grain-filling period equal to thecontrol, the maximum daily increment of fresh weight and grain volume were close to orgreater. Analysis showed that, the main factor deciding high spike weight wheat grain weightwas the grain-filling rate, the grain-filling duration played a secondary role. Average fillingrate and leaf duration positively correlated to grain full index. High spike weight wheat had arelatively inadequate population source, a large storage capacity, but the relationship betweensource and storage was not coordination enough, which was the main reason for its grainunsaturated. There were two possible reasons for inadequate source of high spike weightwheat: A. the small population and low population growth rate, B. most of photosyntheticproducts was supplied to grain due to its large storage, while leaves got less nutrients causingthe blade decline and functional period shortened as well as the photosynthetic yield declined.High spike weight wheat had the higher canopy temperature, the shorter period of leaves’output function and the lower level of the flag leaf chlorophyll, which could not fulfill theneeds of photosynthetic products of the potential storage capacity.
2. According to the major gene and poly-genes mixed inheritance model, we analyzedhigh spike weight wheat’s main agronomic traits of its six generations of the threecombinations. The analysis showed that they are in line with the polygenic additive–dominance-epistasis model, including plant ears, plant height and grain weight per ear of allcombinations, the ear length of combination II and III, spikelets per ear of combination Iand grains per ear of combination II, being controlled by multiple loci and gene epistasis. Earlength of combinatio I was under the control of two pairs of additive–dominant–epistaticmajor genes. Spikelets per ear of combination II and III is controlled by one pair of negativecomplete dominant major gene and additive-dominant polygenes. Grains per ear ofcombination I and III was determined by two pairs additive-dominant-epistatic major geneand additive-dominant-epistatic polygenes. The appearing of differences betweencombinations may be related to the type of parent.
3. The tillering traits of high spike weight wheat were jointly controlled by genetic maineffects and genotype-environment interaction effects, mainly genetic main effects, andadditive and dominant effects were dominant at different developmental stages. Genescontrolling tillers were more active in the early formation of tillers (0-7days), the earlymidterm (8-14days) and medium (15-21days), according to the detected variancecomponents of the conditional genetic effect. Net photosynthetic rate was calculated mainly by genotype variance. Before the middle of the development (May16), dominance variance isthe main component of genetic variation, and additive variance was the main component atthe middle and late phase. From the start of grain-filling (May2), a large number of minorgenes are activated and express. From May2to May16, the genes controlling netphotosynthetic rate expressed most actively. Grain Weight Formation was controlled mainlyby additive, dominant effects, as well as dominant and environment interactions, and theimpact of the environment on Grain Weight Formation also reached a significant level,indicating that grain weight were highly vulnerable to the impact of environmental conditions.When grouting begined, the additive effect occurs, the epistatic effect still existed, butenvironment effect was the main. At the early grain filling stage (5.10|5.7), the additivedominance effected briefly disappeared, and the epistatic effects continuously increased. Atthe middle grouting phase (5.13|5.105.16|5.13), the additive dominant effects appearedagain, the epistatic effects briefly disappeared, but the interaction between the epistatic andthe environment reached the maximum, illustrating that the grain weight was very sensitive tothe environmental impact now.
1、在群体发育方面，穗重型小麦冬季分蘖、春季分蘖及单位面积穗数都低于对照（多穗型品种），通过密度和播期调节的范围较窄；分蘖偏少的原因是形成一个分蘖所需要的积温更多，对低温的适应性更差。穗重型小麦形成较多小穗的原因在于：幼穗分化时期较短，表现为单棱期短，进入二棱期早，二棱期较长；幼穗分化速率快。穗重型小麦单茎干物质始终高于对照，单株干物质前期（开花前）高后期低，单位面积干物质积累偏少，且茎叶鞘输出的干物质多，向籽粒输入的少，输出输入效率较低。穗重型小麦净光合速率前（灌浆以前）高后低，呼吸速率与蒸腾速率均高于对照，根系吸收能力较强。穗重型小麦籽粒平均灌浆速率、最大灌浆速率均低于对照，灌浆持续期持平，粒鲜重和籽粒体积最大日增量不小于对照，决定穗重型小麦粒重的主要因素是灌浆速率，灌浆持续期起次要作用，平均灌浆速率、叶面积持续期与籽粒饱满指数显著正相关。穗重型小麦群体源相对不足，库容量较大，源、库之间的关系不够协调。穗重型小麦源不足表现为：群体偏小，群体生长率降低；光合产物大部分供应籽粒，叶片营养亏缺，导致叶片功能下降，光合产率降低。穗重型小麦冠层温度偏高，旗叶叶绿素水平偏低且下降迅速，叶片输出功能较低，无法适应潜在库容对光合产物的需要。
2、利用主基因-多基因混合遗传模型，对穗重型小麦3组合6世代的主要农艺性状分析表明，各性状的遗传模型因组合不同而表现有同有异，相同的情况是所有组合的单株穗数、株高和单穗粒重均一致地符合多基因的加性-显性-上位性遗传模型；符合这一遗传模型的还有：组合Ⅱ、组合Ⅲ的穗长，组合Ⅰ的每穗小穗数，组合Ⅱ的每穗粒数，这些性状均受多基因位点控制，且基因间存在上位性互作。表现各异的情况是：组合Ⅰ的穗长由2对具有加性-显性-上位性互作的主基因所控制，组合Ⅱ、Ⅲ的每穗小穗数由1对负向完全显性主基因+加性-显性多基因控制，组合Ⅰ、Ⅲ的每穗粒数由2对加性-显性-上位性主基因+加性-显性-上位性多基因共同控制，组合间出现的差异可能与亲本类型和遗传背景有关。
3、穗重型小麦分蘖性状同时受到遗传主效应和基因型×环境互作效应的共同控制，且以遗传主效应为主，其中加性效应和显性效应在分蘖形成的早期（0-7天）、早中期（8-14天）、中期（15-21天）表现得比较活跃，可以检测到显著的条件遗传效应方差分量。净光合速率的遗传以基因型方差为主，发育的中期（5月16日）以前，显性方差是遗传变异的主要分量，中后期加性方差为主要分量，灌浆（5月2日起）开始以后，大量的微效多基因被激活表达，从5月2日到5月16日，是控制净光合速率性状表现的基因表达最为活跃的时期；粒重形成以基因的加性效应、显性效应，显性与环境互作为主，环境对粒重形成的影响也达到显著水平，说明粒重极易受环境条件的影响；灌浆开始，加性效应初现，上位性效应依然存在，而环境效应最大；灌浆初期（5.10︱5.7），加性显性效应短暂消失，上位性效应持续增大；灌浆中期（5.13︱5.105.16︱5.13），加性显性效应再次出现，上位性效应短暂消失，但上位与环境互作却达到最大，说明此时粒重对环境影响非常敏感。
Wheat is the second largest crop planted in China, and plays a very important role fornational food security. The breeding of high-yielding varieties is an effective way to increasewheat production. The high spike weight wheat shows prominent ear characteristics and highgrain weight per ear, which bring hope and dawn for super wheat breeding in China. However,as a cultivar type, some traits of high spike weight wheat have not been systematic studied,such as the features of character development, genetic laws of typical panicle traits passing tothe offspring, dynamic genetic rule on grain weight and tiller traits in the developmentalprocess and so on. In response to these problems, we used high spike weight wheat genotype,breed or introduced to Huanghuai wheat area, as tested materials in this study to investigatetraits development features, spike genetic analysis, tillers and grain weight traits dynamicdevelopmental genetics rule of spike wheat, aiming to provide basis and references for therational use of high spike weight wheat germplasm resources and breeding of new superwheat varieties. The results obtained are as follows:
1. In Population development,the winter tillers, spring tillers and panicles in high spikeweight wheat were lower than those of in the control (panicle varieties), the scope ofregulation was more narrow by the density and sowing date, formation of a tiller needed moreaccumulated temperature, and adaptability to low temperature was worse, controlled bymulti-panicle varieties. High spike weight wheat spike differentiation period was generallyless than that of the control, a short single ridge stage period, the two-ridge stage period wasearlier and last longer. The spike differentiation rate also was faster than the control, whichresulted in more spikes of the high spike weight wheat. The single stem dry matteraccumulation was higher than that of the control, but single plant dry matter accumulationearly stage (before flowering) was higher than late stage, dry matter accumulation of per unitarea had no advantage. Most varieties of high spike weight wheat outputted more dry matterfrom the stem sheath and leaf sheath, but inputted less to the grain, input-output rate also waslow, indicating that some problems exited in its material flow. The net photosynthetic rate ofhigh spike weight wheat was higher than that of the control before grain filling and then reduced, but both of the respiratory rate and transpiration rate were higher, meaning that highspike weight wheat belonged to the high photosynthetic and high breathing consumption type,while its root absorption ability was better. The average grain filling rate and the maximumgrain-filling rate of high spike weight wheat were lower, grain-filling period equal to thecontrol, the maximum daily increment of fresh weight and grain volume were close to orgreater. Analysis showed that, the main factor deciding high spike weight wheat grain weightwas the grain-filling rate, the grain-filling duration played a secondary role. Average fillingrate and leaf duration positively correlated to grain full index. High spike weight wheat had arelatively inadequate population source, a large storage capacity, but the relationship betweensource and storage was not coordination enough, which was the main reason for its grainunsaturated. There were two possible reasons for inadequate source of high spike weightwheat: A. the small population and low population growth rate, B. most of photosyntheticproducts was supplied to grain due to its large storage, while leaves got less nutrients causingthe blade decline and functional period shortened as well as the photosynthetic yield declined.High spike weight wheat had the higher canopy temperature, the shorter period of leaves’output function and the lower level of the flag leaf chlorophyll, which could not fulfill theneeds of photosynthetic products of the potential storage capacity.
2. According to the major gene and poly-genes mixed inheritance model, we analyzedhigh spike weight wheat’s main agronomic traits of its six generations of the threecombinations. The analysis showed that they are in line with the polygenic additive–dominance-epistasis model, including plant ears, plant height and grain weight per ear of allcombinations, the ear length of combination II and III, spikelets per ear of combination Iand grains per ear of combination II, being controlled by multiple loci and gene epistasis. Earlength of combinatio I was under the control of two pairs of additive–dominant–epistaticmajor genes. Spikelets per ear of combination II and III is controlled by one pair of negativecomplete dominant major gene and additive-dominant polygenes. Grains per ear ofcombination I and III was determined by two pairs additive-dominant-epistatic major geneand additive-dominant-epistatic polygenes. The appearing of differences betweencombinations may be related to the type of parent.
3. The tillering traits of high spike weight wheat were jointly controlled by genetic maineffects and genotype-environment interaction effects, mainly genetic main effects, andadditive and dominant effects were dominant at different developmental stages. Genescontrolling tillers were more active in the early formation of tillers (0-7days), the earlymidterm (8-14days) and medium (15-21days), according to the detected variancecomponents of the conditional genetic effect. Net photosynthetic rate was calculated mainly by genotype variance. Before the middle of the development (May16), dominance variance isthe main component of genetic variation, and additive variance was the main component atthe middle and late phase. From the start of grain-filling (May2), a large number of minorgenes are activated and express. From May2to May16, the genes controlling netphotosynthetic rate expressed most actively. Grain Weight Formation was controlled mainlyby additive, dominant effects, as well as dominant and environment interactions, and theimpact of the environment on Grain Weight Formation also reached a significant level,indicating that grain weight were highly vulnerable to the impact of environmental conditions.When grouting begined, the additive effect occurs, the epistatic effect still existed, butenvironment effect was the main. At the early grain filling stage (5.10|5.7), the additivedominance effected briefly disappeared, and the epistatic effects continuously increased. Atthe middle grouting phase (5.13|5.105.16|5.13), the additive dominant effects appearedagain, the epistatic effects briefly disappeared, but the interaction between the epistatic andthe environment reached the maximum, illustrating that the grain weight was very sensitive tothe environmental impact now.
引文
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