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我国东北地区水稻产量和品质及遗传多样性研究
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摘要
东北地区是我国重要的商品粮生产基地,水稻播种面积和总产量已占到北方粳稻区的68%左右,商品率则占到65%以上。东北地区地理纬度跨度大、气候生态环境差异明显,而对这一区域水稻的产量、品质、形态生理特性及遗传多样性等仍缺乏系统研究。本研究以东北3省水稻区域试验品种(品系)和有代表性推广品种为试材,研究东北地区水稻产量和品质的变化规律及遗传多样性,主要试验结果如下:
     1.3省各自产量及其构成因素的品种间差异均达到极显著水平,辽宁产量极显著高于吉林和黑龙江,并且吉林显著高于黑龙江。辽宁穗数和每穗粒数分别极显著低于和高于另两省,千粒重以吉林最高,结实率则以黑龙江最高。辽宁每穗粒数显著高于吉林和黑龙江,而穗长却与吉林和黑龙江差异不显著,因此,辽宁着粒密度显著高于吉林和黑龙江。
     2.东北地区生产上应用的和育种单位选育的品种中,超高产和低产品种较少,多数品种处于中低产到高产之间。单位面积穗数在不同产量水平间无明显规律性。穗粒数随产量的提高呈现增加趋势,二次枝梗数和二次枝梗粒数的增加是穗粒数提高的关键。超高产类型结实率和千粒重低于其他类型。所以,在稳定穗数和千粒重的同时,提高穗粒数和结实率是东北地区超高产水稻品种获得高产的重要条件。
     3.辽宁出糙率最高,蛋白质和游离脂肪酸含量也最高;黑龙江精米率、整精米率最高,粒长、长宽比也最高,直链淀粉含量和食味值与吉林相近,并且显著高于辽宁;吉林垩白粒率、垩白度、白度最高,粒宽也最高。产量及其相关性状基本都与出糙率正相关。穗数、穗型指数、结实率及与其密切相关的一二次枝梗结实率对精米率、整精米率有正面影响,穗粒数及与其密切相关的着粒密度、一二次枝梗数、一二次枝梗粒数对精米率、整精米率有负面影响。垩白米率和垩白度与结实率,尤其是二次枝梗结实率关系密切,因此,提高品种的结实率有利于垩白性状的改善。
     4.产量与生物产量和后期物质生产量极显著正相关,与前期物质生产量相关不显著,所以尽管超高产类型前期物质量较高,但是茎鞘物质输出量、贡献率和转运率与产量无明显关系,超高产主要是依靠后期光合生产能力的改善实现的。
     5.东北地区1971年以来水稻产量的年增长率为0.94%,穗数和结实率的提高是黑龙江产量提高的关键原因,吉林和辽宁产量提高则是依靠穗粒数的增加。品种随年代演替过程中,通过提高生物产量和降低收获指数来增加产量,要得益于结实阶段较高的群体生长率和茎鞘物质输出量。随着时间的演进,稻米加工品质、外观品质和食味品质均得到明显改善。
     6.3省籽粒充实率以黑龙江最高,辽宁和吉林差异不显著。饱谷千粒重是吉林>黑龙江>辽宁,相互差异显著。受精粒千粒重是吉林和黑龙江极显著高于辽宁,吉林与黑龙江差异不显著。饱粒率和结实率是辽宁、黑龙江>吉林。充实率与产量正相关不显著,与着粒密度、二次枝梗数、二次枝梗粒数和二次粒率显著或极显著负相关,与穗型指数、一次枝梗结实率和二次枝梗结实率显著或极显著正相关。充实率与碾磨品质和外观品质关系密切,对其有正面影响。RVA谱特征是黑龙江峰值粘度、热浆粘度、冷胶粘度、回复值、消减值和起浆温度最大,辽宁崩解值最大,吉林峰值粘度时间最大,并且充实率与RVA谱特征值关系密切。
     7.东北地区多数品种(系)是穗型直到半直立(82.6%)、着粒密度中到稀(80.2%)、穗重轻到中(72.8%)、二次枝梗分布下部优势型(98.8%)。黑龙江主要是弯、半直立穗型品种,吉林以半直立穗型品种为主,辽宁则以直立穗型品种为主。吉林和黑龙江品种多数是中、稀穗型,辽宁主要是中、密穗型。吉林和黑龙江多数品种是中、轻穗型,辽宁则以中、重穗型为主。穗颈大、小维管束数和第2节间大、小维管束数平均值表现为辽宁>吉林>黑龙江。辽宁上三叶长、宽大于吉林和黑龙江,但叶基角较小。穗颈大、小维管束数和第2节间大、小维管束数与一、二次枝梗数和一、二次枝梗粒数及二次粒率呈极显著正相关,与一、二次枝梗结实率和千粒重呈负相关。总体上有穗型直立、着粒密度大、重穗型、穗颈维管束数多、上三叶长且宽而角度小、株高和生物产量高有利于高产的趋势。
     8.东北地区水稻推广品种表型遗传多样性高于DNA水平,时期间表型与微卫星标记的遗传多样性具有较高的相似性,而地区间无明显相似性。东北水稻推广品种遗传多样性比较狭窄,随着时间推移增加的新等位基因多于消失的旧等位基因数。三省水稻推广品种遗传多样性呈现黑龙江>吉林>辽宁的趋势,但由于地区间频繁引种,使东北水稻品种在区域上的遗传分化已不明显,其差异主要来源于品种间的基因型差异。不同省份存在较多的互补等位变异数,最多的在黑龙江和辽宁之间。不同省份和不同时期间分别拥有各自特有和特缺的等位变异。基于遗传距离的Neighbor-Joining聚类显示参试品种可分为5个类群,每一类群均有2个以上时期(省份)品种分布。
     9.利用40个SSR标记分析18份我国东北水稻品种和13份日本品种的遗传多样性,共检测到108个等位变异,其中中国东北等位变异数为103个(平均2.54个),日本品种等位变异数为94个(2.32个)。遗传变异主要存在于品种间(95%),群体之间(5%)差异较小,聚类分析将供试材料分为5类,日本引进品种没有形成独立的类群。中国东北选育品种的遗传多样性高于日本引进品种,并且保留了日本品种94.7%的等位变异。日本品种中可利用的特异等位基因十分有限,已经不能满足东北地区水稻育种发展的需求。
Northeast China is an important commercial grain production base,and its rice sowing area and yield had accounted for about 68%of Northern Japonica rice,while the rate of merchandise accounted for more than 65%.Geographic latitude span larger,and climatic ecological environment is different obviously in Northeast area,however,there was no systematic research for yield and quality,morphological and physiological characteristics and genetic diversity of rice.The changing regulation of yield and quality and genetic diversity were studied using cultivars and lines attended regional trials and commercial varieties area as materials in Northeast.The major results were as follows:
     1.There was significant difference in grain yield and yield components among different cultivars and lines,and grain yield of Liaoning province was significantly higher than that of Jilin and Heilongjiang province,and grain yield of Jilin province was significantly higher than that of Heilongjiang.The number of panicle per unit area and grains per panicle in Liaoning province were significantly lower and higher than those in Jilin and Heilongjiang province, respectively,1000-grain weight was the highest in Jilin province,and seed-setting rate was the highest in Heilongjiang province.The grains per panicle of Liaoning was significantly higher than that of Jilin and Heilongjiang province,but the panicle length was not significant difference among different provinces,therefore,grain density in Liaoning province was very significantly higher than that of Jilin and Heilongjiang province.
     2.The super-high-yield and low-yield varieties were less,most varieties in mid-low-yielding to high-yielding in China Northeast area.Panicles per unit area showed no obvious regulary in different yield types.Grains per panicle showed an increasing trend with yield improving,and grains per panicle was increased by increasing the number of secondary branches and secondary branches grains.The seed-setting rate and 1000-grain weight of super-high-yield rice were lower than other yield types.Therefore,improving grains per panicle and seed-setting rate was an important condition for achieving super-high-yield in rice.
     3.Liaoning province had the highest brown rice rate,protein content and dissociated fatty acid content;Heilongjiang province had the highest milled rice rate,head rice rate,grain length and length-width ratio of grain.The amylose content and palatability value of rice in Heilongjiang were similar to Jilin province,and were significantly higher than those of Liaoning.Yield,panicle number,panicle type index,seed-setting rate,grain density,the number of primary branches and secondary branches,the number of primary branch grains and secondary branch grains,the seed-setting rate of primary branches and secondary branches were negatively correlated to milled rice rate and head rice rate.The chalky grain percentage and chalkiness degree were closely related to seed-setting rate,especially seed-setting rate of secondary branches,therefore,improving seed-setting rate was conducive to the improvement of traits chalkiness.
     4.Economic yield was very significant positive correlation with biological yield,and was not significant correlation with material production in early growth stage.Therefore,even though material production in early growth stage of super-high-yield rice was higher than other yield type,dry matter exportation from vegetative organs,apparent conversion percentage and apparent translocation percentage had no significant correlation with economic yield.Improving yield for super-high-yield rice depended mainly on the photosynthetic capacity on early stage.
     5.Since 1971,the annual growth rate of rice yield was 0.94%in Northeast region of China.The increase of panicle per unit and seed-setting rate was the key reason for improving the yield,however in Jilin and Liaoning province was by improving grains per panicle.With the succession of cultivars,by increasing the biological yield and reducing harvest index to increase yield was necessary to obtain the high crop growth rate and dry matter exportation from stem and sheath in seed setting stage.As time evolution,the rice processing quality, appearance quality and eating value had been significantly improved.
     6.The grain plumpness of Heilongjiang province was significantly higher than that of Liaoning and Jilin province,but there was no significant difference in grain yield between Liaoning and Jilin province.1000-plump grain weight was fellowed by Jilin>Heilongjiang>Liaoning,and had significantly different among different provinces.1000-fertilized grain weight in Jilin and Heilongjiang province was very significantly higher than in Liaoning province,but there was no significant difference in grain yield between Jilin and Heilongjiang province.Plump grain rate and grain plumpness were in order of Liaoning>Heilongjiang>Jilin.Grain plumpness significantly correlated to yield,but significantly or very significantly negatively correlated with grain density,secondary branches,the number of secondary branch grains and ratio of secondary branch grains,and significantly or very significantly positively correlated with panicle trait index and seed-setting rate of primary branch and secondary branch.Grain plumpness had a positive effect on milling quality and appearance quality.Peak viscosity,hot paste viscosity,cool paste viscosity,consistence,setback and pasting temperature in Heilongjiang province were the highest among three provinces,and breakdown in Liaoning province and peak time in Jilin province were the highest,respectly. Except for hot paste viscosity and setback,grain plumpness had a significant relationship with other RVA profiles.
     7.The plant-type of most cultivars and lines at northeast region of China was EPT(erect panicle type) or SEPT(semi-erect panicle type) according to PNC(panicle-neck curvature), DPT(loose panicle type) or SDPC(semi-dense panicle type) according to grain density,MPT (middle panicle type) or HPT(heavy panicle type) according to panicle weight and superior-on-the-low-part type.Most rice cultivars(lines) were CPT(curve panicle type) or SEPT in Heilongjiang,SEPT in Jilin and EPT in Liaoning;semi-dense or loose panicle type in Jilin and Heilongjiang,and dense panicle type in Liaoning;MPT(middle panicle type) or TPT(thin panicle type) in Jilin and Heilongjiang,and HPT(heavy panicle type) in Liaoning. Large vascular bundles in neck(LVBN),Small vascular bundles in neck(SVBN),large vascular bundles in the second internode(LVBS) and small vascular bundles in the second internode(SVBS) were in order of Liaoning>Jilin>Heilongjiang;Length and width of the top three leaves was larger,but basic angle less than Jilin and Heilongjiang.LVBN,SVBN, LVBS and SVBS were closely positive correlated with primary branches,primary branch grains,secondary branches,secondary branch grains and secondary branch grains rate,but negative correlated with seed-setting rate and 1000-grain weight of primary branch and secondary branch.In general,there was a tendency that the more vascular bundles, length(width) of the top three leaves,plant height and biomass and smaller angle cultivation we cultivated,the higher yield we can obtained.
     8.The phenotypic genetic diversity in the Northeast China Commercial Rice population was higher than that of DNA level,among varieties of different years,there was high similarity in genetic diversity between morphological markers and SSR markers,but that was on the contrary among varieties of different provinces.There existed a scarcity of genetic diversity in Commercial population of northeast China.With advance of years,some old alleles immerged and some new ones appeared,and the news were more than the olds. Genetic diversity of commercial rice varieties in northeast China was in order of Heilongjiang>Jilin>liaoning.Because of frequent inter-regional introduction of varieties,the genetic differentiation was not obvious among regions,and mainly comes from genotype difference among varieties.There were some mutual complementary alleles among provincial subpopulations,especially more between Heilongjiang and Liaoning.There existed or deficient some alleles among varieties in different provinces and years.The commercial rice varieties in northeast China were clustered into 5 groups and every group included the varieties of 2 years or provinces.
     9.Rice varieties from China Northeast and 13 introduced varieties from Japan were studied on genetic diversity by 40 SSR markers.108 alleles were detected,among of that there were 103 alleles in China Northeast and 94 in Japan,and the average number of alleles per Primer was 2.54 and 2.32,respectively.More genetic variation existed in the species,but less between groups.Genetic diversity of China Northeast varieties was higher than Japan varieties and retained 94.68%theirs alleles.The available specific alleles were very limitation in Japan varieties and could no longer meet the rice breeding development of northeastern region of China.
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