我国北方玉米品种及自交系更替过程中产量和生理特性的演进
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摘要
本试验于2009～2010年在辽宁省铁岭市昌图县金家镇、沈阳农业大学科研基地及沈阳市特种玉米遗传育种与生理重点实验室进行。选用1950年代以来生产上大面积推广的4个OPV品种和31玉米杂交种，以及亲本自交系为试材，探讨我国玉米品种及其亲本自交系更替过程中农艺性状、产量杂种优势、干物质积累特性、生理特性及品质的演变规律，明确农艺性状及品质性状对产量形成的重要性。主要研究结果如下：
1.不同年代品种随着年代的推进产量逐渐增加，低、中、高密度下年代间平均增长比率分别为16.7%、15.7%和21.6%，且各年代间差异均达显著或极显著水平，表明近年代品种产量改良效果显著。随产量的提高，不同年代品种穗长、穗粗、穗行数、百粒重明显增加，近年代品种秃尖度较大、出籽率较低，说明这两项指标正向不利方向发展。1970S和2000S品种的株高、穗位高显著高于其它年代，且1970S品种穗位/株高显著高于其它年代，表明1970S品种重心较高，而1970S后品种重心则逐渐降低。年代更替过程中，各品种茎叶夹角逐渐减小、叶向值显著提高、持绿性明显增加；近年代品种雄穗分支数显著减少，倒伏率、倒折率明显降低。相关和通径分析表明，低、中、高密度下百粒重、穗行数、穗粒数、穗长、穗粗、粒长、叶向值与产量均呈极显著正相关关系；对产量的贡献率居前三位的为粒长、穗粗和穗长。
2.自交系随着年代的推进产量逐渐增加，三个密度下年代间增加幅度在3.1%-37.3%之间；穗长、穗粒数变化不明显；穗粗、粒长、百粒重及叶向值呈不同程度的增加趋势，倒伏率、倒折率、雄穗长、雄穗分支数呈降低趋势，近年代品种与早年代间差异达显著或极显著水平，说明这些性状的正向改良是产量提高的主要原因。相关和通径分析结果显示，三个密度下百粒重、穗粗、粒长、叶向值均呈显著正相关关系，对产量贡献率较大的为穗粗、穗长和叶向值。
3.我国不同年代单交种产量及中亲值产量均随年代的推进整体呈增加趋势。绝对杂种优势随年代的推进逐渐增加，低密度、中密度、高密度下绝对杂种优势平均增加比率分别为3.9%、7.9%和18.8%，而相对杂种优势、中亲优势及杂种优势指数在低密度、中密度下略有降低趋势，在高密度下略有增加趋势；超高亲优势年代间变化较为平缓，1980S-1990S品种超低亲优势显著低于1970S。通过以上分析表明，低密度下杂种优势对产量的贡献率较大，随着密度的增加杂种优势越来越小，说明非杂种优势因素对产量的贡献率越来越大。
4.不同年代品种干物质积累量随年代的推进呈增加–降低–增加趋势。1950S品种各个生育时期干物质积累量均小于其它年代品种；2000S年代品种灌浆中后期干物质积累速率高于其它年代品种，成熟时茎、叶、鞘干物质积累量高于其它年代品种，但其籽粒干物重所占比例较低，经济系数较小。各年代品种比较，茎的转运量、转运率及对籽粒贡献率均以1990S品种最高；叶的转运量以1980S品种最高，转运率和对籽粒贡献率以1950S品种最高；鞘的转运量和转运率以1980S品种最高，对籽粒贡献率以1950S品种最高。1950S和1990S品种灌浆强度较大，但灌浆持续期短，1970S和1980S品种灌浆持续期较长，但灌浆强度较小，2000S品种灌浆强度较强，灌浆持续期较长，利于高产。1950S-1980S品种脱水速率逐渐加快，而1990S和2000S品种脱水速率明显低于其它年代品种。
5.对不同年代品种灌浆期穗位叶净光合速率、叶绿素含量、保护酶活性及膜脂过氧化产物的含量的测定结果表明，光合速率随年代的推进呈增加趋势；叶绿素a、叶绿素b及叶绿素总量呈增加趋势，这种趋势在灌浆后期越发明显；类胡萝卜素/叶绿素呈降低趋势，说明近期品种持绿性优于早期品种，利于后期光合作用；超氧化物歧化酶（SOD）活性呈增加趋势，膜脂过氧化产物（MDA）含量逐渐降低，这种趋势在灌浆后期尤为突出，表明近期品种具有较强的清除活性氧的能力。
6.1970S-2000S品种淀粉含量逐渐增加，低、中、高密度下年代间平均增长比率分别为0.88%、0.81%和0.77%，且1970S品种与其它年代间差异达极显著水平；脂肪含量、蛋白质含量与淀粉含量均呈负相关关系，说明1970S-2000S品种更替过程中籽粒淀粉含量逐渐增加，而脂肪和蛋白质含量则呈降低趋势。容重的变化整体较为平缓，年代间差异不大。相关分析表明淀粉含量、籽粒容重与产量呈正相关关系，蛋白质、脂肪、赖氨酸含量与产量呈负相关关系。以上分析说明我国北方地区玉米品种更替过程中品质逐渐下降。
7.自交系更替过程中，1980S自交系淀粉和脂肪含量高于其它年代，蛋白质含量则显著低于其它年代；1970S-1990S自交系籽粒赖氨酸含量逐渐降低。
The experiment was carried out in Jin Jia town of Changtu County of Liaoning ProvinceTie Ling City, scientific research base of Shen Yang Agricultural University and keyLaboratory of Specialty Maize Genetics Breeding and Physiology of Shen Yang City from2009to2010. Using four large scale OPV varieties on production since1950s and thirty-onemaize hybrids and their parental inbred lines as experimental materials, evolutions ofagronomic traits, yield heterosis, dry material accumulation, physiological characteristics andqualities of Chinese maize varieties and their parental inbred lines during the yearreplacement were explored to define the importance of agronomic and quality traits to yieldformation. The main results were as follows:
1. Yields of different year varieties increased with the advancing of era, and the averagegrowth rate among years were respectively16.7%,15.7%and21.6%under the low, mediumand high densities and the differences among years were significant or highly significant,which showed that effects of yield improvement were significant in recent years.Ear length,ear diameter, rows per ear and100-kernel weight of different year varieties increasedsignificantly with the increase of yield. Greater bald degree and lower seeding rate of recentyear varieties indicated that these two indexes developed in negative direction. Plant heightand ear height of1970S and2000S varieties were significantly higher than other years, andthe ratio of ear height/plant height of1970S varieties of were significantly higher than otheryears which indicated gravity centers of1970S varieties were higher, hovever, the gravitycenters of other years varieties were gradually reduced.During the year replacement, theangles of stem and leaf of all varieties gradually reduced while the LOV and value of greenduration obviously increased; branch number of tassel of recent year varieties significantlyreduced and rates of lodging and discount fell greatly. Correlation and path analysis showedthat100-kernel weight, rows per ear, grain number, ear length, ear diameter, grain length andLOV were highly significant positive correlation with yields under the low, medium and highdensity; the first three contribution rate on Yield were grain length, ear diameter and earlength.
2. Yields of inbred lines increased with the increase of era, increasing margins of differentyear were from3.1%to37.3%under the three densities; ear length and grain number had notobvious change; ear diameter, grain length,100-grain weight and LOV showed increasing trends of different extent while lodging rate, discount rate, tassel length and branch number oftassel showed decreasing trends and the differences between recent year varieties and formeryear varieties were significant or highly significant, which indicated that the positiveimprovement of these traits was the main reason for yield increasing. Correlation and pathanalysis showed that100-kernel weight, ear diameter, grain length and LOV under the threedensity were significant positive correlation and ear width, ear length and LOV did morecontribution to yield.
3.Yelds of different year single cross hybrids and their mid-parent values all took onincreasing trends with the advancing of era. Absolute heterosis increased with the advancingof era and average ratios of the absolute heterosis increased by3.9%,7.9%and18.8%respectively under the low, medium and high density, while the relative heterosis, mid-parentheterosis and heterosis indexes decreased slightly under the low and medium density but tooka slight upward trend under the high density; ultra-high-parent heterosis varied gentlybetween years and ultra-low-parent heterosis of1980S-1990S varieties was significantlylower than1970S. Through the analysis above showed that heterosis made more contributionsto yields under the low density, while heterosis got smaller and smaller with the increase ofdensities, which indicated that explaining factors made more contributions to yields.
4. Dry matter accumulation of different year varieties increased wavely with theadvancing of era. Dry matter accumulation of1950S varieties were less than the other yearsvarieties in all stages; Dry matter accumulation of2000S varieties were higher than the otheryears varieties in late filling stage and dry matter accumulation of stem, leaf and sheath werehigher than the other years varieties in mature while the proportion of grain dry weight waslower and the economy coefficient was smaller. Comparisons of different year varieties, thestem translocation, transfer rate and grain contribution rate of1990S varieties were all thehighest; leaf translocation of1980S varieties were the highest while leaf transfer rate andgrain of1950S varieties were the highest; sheath translocation and transfer rate of1980Svarieties were the highest while grain contribution rate of1950S varieties were the highest.Filling intensities of1950S and1990S varieties were stronger but their duration were shorter;filling durations of1970S and1980S varieties were longer but their filling intensities weresmaller; varieties of2000S had stronger filling intensities and longer filling durations goodfor high yield. Dehydration rates from1950S to1980S varieties gradually accelerated whiledehydration rates of1990S and2000S varieties were significantly lower than other yearsvarieties.
5Determination results of net photosynthetic rate, chlorophyll content, protectiveenzymes and membrane lipid peroxidation content of ear leaves of different year varietiesshowed that with the advancing of era, the net photosynthetic rate increased; content ofchlorophyll a, chlorophyll b and chlorophyll increased and the trend were more obvious inlate filling stage; carotenoid/chlorophyll decreased, which indicated that green-stay of latervarieties were better that of early varieties, which would be helpful to later photosynthesis;activities of superoxide dismutase (SOD) increased and content of membrane lipidperoxidation (MDA) decreased, and the trend was particularly prominent in late filling stage,which indicated that the later varieties had stronger abilities of scavenging reactive oxygenspecies.
6. Content of Starch from1970S to2000S varieties increased and average growth rateamong years were0.88%,0.81%and0.77%under the low, medium and high density, anddifferences between1970S and other year varieties were significant; content of fat, proteinand starch showed a negative correlation,which indicated that content of starch increasedwhile content of fat and protein decreasd during the variety replacement from1970S to2000S.Bulk density changed more gently and the differences among years were not obvious.Correlation analysis showed that content of starch and grain bulk density were positivelycorrelated with yield, while content of protein, fat and lysine were negatively correlated withyield. The analysis above showed that qualities gradually declined during the replacements ofmaize varieties in northern China.
7. During the replacements of Inbred lines, content of starch and fat of1980S inbred lineswere higher than that of other years inbred lines while content of protein was significantlylower than other years inbred lines; content of lysine from1970S to1990S inbred linesgradually decreased.
1.不同年代品种随着年代的推进产量逐渐增加，低、中、高密度下年代间平均增长比率分别为16.7%、15.7%和21.6%，且各年代间差异均达显著或极显著水平，表明近年代品种产量改良效果显著。随产量的提高，不同年代品种穗长、穗粗、穗行数、百粒重明显增加，近年代品种秃尖度较大、出籽率较低，说明这两项指标正向不利方向发展。1970S和2000S品种的株高、穗位高显著高于其它年代，且1970S品种穗位/株高显著高于其它年代，表明1970S品种重心较高，而1970S后品种重心则逐渐降低。年代更替过程中，各品种茎叶夹角逐渐减小、叶向值显著提高、持绿性明显增加；近年代品种雄穗分支数显著减少，倒伏率、倒折率明显降低。相关和通径分析表明，低、中、高密度下百粒重、穗行数、穗粒数、穗长、穗粗、粒长、叶向值与产量均呈极显著正相关关系；对产量的贡献率居前三位的为粒长、穗粗和穗长。
2.自交系随着年代的推进产量逐渐增加，三个密度下年代间增加幅度在3.1%-37.3%之间；穗长、穗粒数变化不明显；穗粗、粒长、百粒重及叶向值呈不同程度的增加趋势，倒伏率、倒折率、雄穗长、雄穗分支数呈降低趋势，近年代品种与早年代间差异达显著或极显著水平，说明这些性状的正向改良是产量提高的主要原因。相关和通径分析结果显示，三个密度下百粒重、穗粗、粒长、叶向值均呈显著正相关关系，对产量贡献率较大的为穗粗、穗长和叶向值。
3.我国不同年代单交种产量及中亲值产量均随年代的推进整体呈增加趋势。绝对杂种优势随年代的推进逐渐增加，低密度、中密度、高密度下绝对杂种优势平均增加比率分别为3.9%、7.9%和18.8%，而相对杂种优势、中亲优势及杂种优势指数在低密度、中密度下略有降低趋势，在高密度下略有增加趋势；超高亲优势年代间变化较为平缓，1980S-1990S品种超低亲优势显著低于1970S。通过以上分析表明，低密度下杂种优势对产量的贡献率较大，随着密度的增加杂种优势越来越小，说明非杂种优势因素对产量的贡献率越来越大。
4.不同年代品种干物质积累量随年代的推进呈增加–降低–增加趋势。1950S品种各个生育时期干物质积累量均小于其它年代品种；2000S年代品种灌浆中后期干物质积累速率高于其它年代品种，成熟时茎、叶、鞘干物质积累量高于其它年代品种，但其籽粒干物重所占比例较低，经济系数较小。各年代品种比较，茎的转运量、转运率及对籽粒贡献率均以1990S品种最高；叶的转运量以1980S品种最高，转运率和对籽粒贡献率以1950S品种最高；鞘的转运量和转运率以1980S品种最高，对籽粒贡献率以1950S品种最高。1950S和1990S品种灌浆强度较大，但灌浆持续期短，1970S和1980S品种灌浆持续期较长，但灌浆强度较小，2000S品种灌浆强度较强，灌浆持续期较长，利于高产。1950S-1980S品种脱水速率逐渐加快，而1990S和2000S品种脱水速率明显低于其它年代品种。
5.对不同年代品种灌浆期穗位叶净光合速率、叶绿素含量、保护酶活性及膜脂过氧化产物的含量的测定结果表明，光合速率随年代的推进呈增加趋势；叶绿素a、叶绿素b及叶绿素总量呈增加趋势，这种趋势在灌浆后期越发明显；类胡萝卜素/叶绿素呈降低趋势，说明近期品种持绿性优于早期品种，利于后期光合作用；超氧化物歧化酶（SOD）活性呈增加趋势，膜脂过氧化产物（MDA）含量逐渐降低，这种趋势在灌浆后期尤为突出，表明近期品种具有较强的清除活性氧的能力。
6.1970S-2000S品种淀粉含量逐渐增加，低、中、高密度下年代间平均增长比率分别为0.88%、0.81%和0.77%，且1970S品种与其它年代间差异达极显著水平；脂肪含量、蛋白质含量与淀粉含量均呈负相关关系，说明1970S-2000S品种更替过程中籽粒淀粉含量逐渐增加，而脂肪和蛋白质含量则呈降低趋势。容重的变化整体较为平缓，年代间差异不大。相关分析表明淀粉含量、籽粒容重与产量呈正相关关系，蛋白质、脂肪、赖氨酸含量与产量呈负相关关系。以上分析说明我国北方地区玉米品种更替过程中品质逐渐下降。
7.自交系更替过程中，1980S自交系淀粉和脂肪含量高于其它年代，蛋白质含量则显著低于其它年代；1970S-1990S自交系籽粒赖氨酸含量逐渐降低。
The experiment was carried out in Jin Jia town of Changtu County of Liaoning ProvinceTie Ling City, scientific research base of Shen Yang Agricultural University and keyLaboratory of Specialty Maize Genetics Breeding and Physiology of Shen Yang City from2009to2010. Using four large scale OPV varieties on production since1950s and thirty-onemaize hybrids and their parental inbred lines as experimental materials, evolutions ofagronomic traits, yield heterosis, dry material accumulation, physiological characteristics andqualities of Chinese maize varieties and their parental inbred lines during the yearreplacement were explored to define the importance of agronomic and quality traits to yieldformation. The main results were as follows:
1. Yields of different year varieties increased with the advancing of era, and the averagegrowth rate among years were respectively16.7%,15.7%and21.6%under the low, mediumand high densities and the differences among years were significant or highly significant,which showed that effects of yield improvement were significant in recent years.Ear length,ear diameter, rows per ear and100-kernel weight of different year varieties increasedsignificantly with the increase of yield. Greater bald degree and lower seeding rate of recentyear varieties indicated that these two indexes developed in negative direction. Plant heightand ear height of1970S and2000S varieties were significantly higher than other years, andthe ratio of ear height/plant height of1970S varieties of were significantly higher than otheryears which indicated gravity centers of1970S varieties were higher, hovever, the gravitycenters of other years varieties were gradually reduced.During the year replacement, theangles of stem and leaf of all varieties gradually reduced while the LOV and value of greenduration obviously increased; branch number of tassel of recent year varieties significantlyreduced and rates of lodging and discount fell greatly. Correlation and path analysis showedthat100-kernel weight, rows per ear, grain number, ear length, ear diameter, grain length andLOV were highly significant positive correlation with yields under the low, medium and highdensity; the first three contribution rate on Yield were grain length, ear diameter and earlength.
2. Yields of inbred lines increased with the increase of era, increasing margins of differentyear were from3.1%to37.3%under the three densities; ear length and grain number had notobvious change; ear diameter, grain length,100-grain weight and LOV showed increasing trends of different extent while lodging rate, discount rate, tassel length and branch number oftassel showed decreasing trends and the differences between recent year varieties and formeryear varieties were significant or highly significant, which indicated that the positiveimprovement of these traits was the main reason for yield increasing. Correlation and pathanalysis showed that100-kernel weight, ear diameter, grain length and LOV under the threedensity were significant positive correlation and ear width, ear length and LOV did morecontribution to yield.
3.Yelds of different year single cross hybrids and their mid-parent values all took onincreasing trends with the advancing of era. Absolute heterosis increased with the advancingof era and average ratios of the absolute heterosis increased by3.9%,7.9%and18.8%respectively under the low, medium and high density, while the relative heterosis, mid-parentheterosis and heterosis indexes decreased slightly under the low and medium density but tooka slight upward trend under the high density; ultra-high-parent heterosis varied gentlybetween years and ultra-low-parent heterosis of1980S-1990S varieties was significantlylower than1970S. Through the analysis above showed that heterosis made more contributionsto yields under the low density, while heterosis got smaller and smaller with the increase ofdensities, which indicated that explaining factors made more contributions to yields.
4. Dry matter accumulation of different year varieties increased wavely with theadvancing of era. Dry matter accumulation of1950S varieties were less than the other yearsvarieties in all stages; Dry matter accumulation of2000S varieties were higher than the otheryears varieties in late filling stage and dry matter accumulation of stem, leaf and sheath werehigher than the other years varieties in mature while the proportion of grain dry weight waslower and the economy coefficient was smaller. Comparisons of different year varieties, thestem translocation, transfer rate and grain contribution rate of1990S varieties were all thehighest; leaf translocation of1980S varieties were the highest while leaf transfer rate andgrain of1950S varieties were the highest; sheath translocation and transfer rate of1980Svarieties were the highest while grain contribution rate of1950S varieties were the highest.Filling intensities of1950S and1990S varieties were stronger but their duration were shorter;filling durations of1970S and1980S varieties were longer but their filling intensities weresmaller; varieties of2000S had stronger filling intensities and longer filling durations goodfor high yield. Dehydration rates from1950S to1980S varieties gradually accelerated whiledehydration rates of1990S and2000S varieties were significantly lower than other yearsvarieties.
5Determination results of net photosynthetic rate, chlorophyll content, protectiveenzymes and membrane lipid peroxidation content of ear leaves of different year varietiesshowed that with the advancing of era, the net photosynthetic rate increased; content ofchlorophyll a, chlorophyll b and chlorophyll increased and the trend were more obvious inlate filling stage; carotenoid/chlorophyll decreased, which indicated that green-stay of latervarieties were better that of early varieties, which would be helpful to later photosynthesis;activities of superoxide dismutase (SOD) increased and content of membrane lipidperoxidation (MDA) decreased, and the trend was particularly prominent in late filling stage,which indicated that the later varieties had stronger abilities of scavenging reactive oxygenspecies.
6. Content of Starch from1970S to2000S varieties increased and average growth rateamong years were0.88%,0.81%and0.77%under the low, medium and high density, anddifferences between1970S and other year varieties were significant; content of fat, proteinand starch showed a negative correlation,which indicated that content of starch increasedwhile content of fat and protein decreasd during the variety replacement from1970S to2000S.Bulk density changed more gently and the differences among years were not obvious.Correlation analysis showed that content of starch and grain bulk density were positivelycorrelated with yield, while content of protein, fat and lysine were negatively correlated withyield. The analysis above showed that qualities gradually declined during the replacements ofmaize varieties in northern China.
7. During the replacements of Inbred lines, content of starch and fat of1980S inbred lineswere higher than that of other years inbred lines while content of protein was significantlylower than other years inbred lines; content of lysine from1970S to1990S inbred linesgradually decreased.
引文
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