胚和胚乳基因效应对水稻幼苗生物量的影响研究

本文利用和胚乳遗传模型[研究了水稻早期胚后生长生物量性状的遗传控制,结果表明：在两个时期水稻幼苗生物量性状中,除了第16天时根鲜重（RFW）主要受到胚乳显性效应控制外,叶鲜重（LFW）,叶干重（LDW）,根干重（RDW）主要受到胚基因显性效应和胚乳基因加性效应的控制,胚加性和乳乳加性效应占总遗传方差的40－54％,说明对生物量性状进行早期选择有效,各个性状都检测到显著的胚狭义遗传率和胚乳狭义遗传率,说明在早期世代即可估计选择进程,对亲本的遗传效应值的预测表明,对根部性状的选择在第8天进行比较合适,并以亲本P1,P3和P6较好,它们既可提高RFW又可提高RDW,而对地上部分性状的选择在第16天时进行比较合适,并以P4,P9和P10为最好。     

基于F3种子的胚乳性状QTL区间定位

文章提出了包括胚乳效应和母体效应的胚乳性状QTL定位的统计方法,该方法的实验设计是分子标记基因型信息来自F2母体植株和F3种子胚(或植株),胚乳性状表型值来自F3单粒种子胚乳,称之为两步等级设计。同时,用计算机全面模拟以验证该模型的可行性,模拟结果表明,只要群体足够大,该模型能较有效地进行胚乳性状QTL定位并精确地估计出胚乳QTL的各种遗传效应和母体效应。     

麦芽品质性状的遗传模型及其分析方法

本文提出分析大麦芽品质性状世代平均数的遗传模型,根据麦芽性状的特点,该模型将控制麦芽性状的总遗传效应分解为萌动胚基因效应和胚乳基因效应。     

胚乳性状的世代遗传方差

本文论述由两个纯系杂交而衍生的各种世代群体的胚乳性状的遗传方差分量。根据有关世代胚乳的遗传组成和加性-显性模型下的基因效应,导出了它们的总的、株间的和自交世代的遗传方差,并分别列于表1—3。     

胚乳性状数量基因的定位方法

将三倍体胚乳性状的数量遗传模型和二倍体性状数量基因（QTL）图构建方法相结合,导出双侧标记基因型下有关胚乳性状QTL的遗传组成、平均数和遗传方差分量,据之提出以某一区间双侧标记基因型胚乳性状的平均值为依变数,以该区间内任一点假定存在的QTL的加性效应d、显性效应h1和／或h2的系数为自变数,进行有重复观察值的多元线性回归分析,根据多元线性回归的显著性测验该点是否存在QTL,并估计出QTL的遗传效应。给定区间内任一点,皆可以此进行分析,从而可在整条染色体上作图,并以之确定QTL的数目和最可能位置,同时,在检测某一区间时,利用多元线性回归方法将该区间外可能存在的QTL的干扰进行统计控制,以提高QTL检测的精度。此外,还讨论了如何将之推广应用于其他类型的DNA不对应资料以及具复杂遗传模型的胚乳性状资料。     

基于BC_1F_（1：2）种子的胚乳QTL区间定位

提出了基于分子标记基因型信息来自BC_1F_1母体植株,胚乳性状表型值来自BC_1F_（1：2）单粒种子胚乳的试验设计的胚乳QTL定位的区间作图方法.同时,用计算机全面模拟以验证该模型的可行性,模拟结果表明,只要群体足够大,该模型能有效地进行胚乳性状QTL定位并能估计出胚乳QTL的各种遗传效应和母体效应.     

玉米籽粒性状的遗传模型研究

用１０个遗传上和籽粒形态性状上具有差异的玉米自交系,依多种可能的交配方法获得亲本Ｐ１、Ｐ２、Ｆ１（Ｐ１× Ｐ２）、Ｆ２、Ｂ１（Ｆ１×Ｐ１）、Ｂ２（Ｆ１× Ｐ２）及其相应反交ＲＦ１、ＲＦ２、ＲＢ１、ＲＢ２共１０个种子世代。种植２年。依广义遗传模型建立包括种子胚乳加性、胚乳显性、母体加性、母体显性和细胞质效应的遗传模型,运用种子数量性状的精细鉴别法［１］和混合模型分析法［２,３］,对粒长、粒宽、粒长宽比、粒厚及百粒重作了性状表达遗传机制的鉴别与探讨。单个组合的遗传模型精细测验表明,５个籽粒性状的遗传主要受母体显性和胚乳基因型（包括加性和灵性）的控制,一个组合的粒宽、粒厚和百粒重上还检测到细胞质效应。对２５对 Ｆ１正反交组合世代均值依ＭＩＮＱＵＥ法分析的结果表明,５个籽粒性状的遗传方差中,母体遗传方差占６０％以上,胚乳基因型方差低于４０％,粒长和百粒重还有细胞质效应,约占１０％～３０％。可见,籽粒性状的遗传特点是受多套遗传系统控制,其中以母体基因型的作用最大。     

籼稻稻米外观品质的细胞质,母体和胚乳遗传效应分析

利用浙协2号A等9个籼型不育系和T49等5个籼型恢复系进行不完全双列杂交,研究了籼稻稻米外观品质的遗传效应．结果表明,稻米外观品质性状的表现受制于胚乳、母体和细胞质三套遗传体系．糙米长、长宽比和长厚比等性状以母体遗传率为主,而糙米宽和糙米厚则以胚乳直接遗传率为主,糙米长和长宽比等性状的细胞质遗传率亦很重要．结果还发现外观品质性状间存在着较强的遗传相关,其中糙米长与糙米宽、糙米长与糙米厚、糙米宽与糙米厚、糙米宽与长宽比、糙米厚与长厚比以及糙米长宽比与长厚比性状间以胚乳直接加性和母体加性相关为主．而糙米长与长宽比、糙米长与长厚比、糙米宽与长厚比以及糙米厚与长宽比性状间则以胚乳直接显性和母体显性相关为主．就外观品质的总体情况而言,遗传效应预测值表明参试亲本以V20A、作5A和测早2-2较好,其各种遗传效应能够显著改善稻米品质性状。V20A／102和作5A／测早2-2等组合具有较好的稻米外观品质．     

水稻亚种间杂交粒重及加工品质性状的遗传效应分析

用4个广亲和粳型品种和5个籼型品种为材料,按NCⅡ设计配制杂交组合,获得同一环境下的亲本及F_1植株上的籽粒群体(F_2),对其稻谷千粒重、糙米千粒重、出糙率、总精米率以及整精米率等粒重和加工品质性状进行测定,并按胚乳性状遗传模型和混合线性模型的分析方法对籼粳亚种间杂交稻粒重及加工品质性状的遗传效应进行了研究,结果表明:籼粳交籽粒的粒重及各加工品质性状同时受到胚乳直接基因效应、母体基因效应以及微弱的细胞质效应的影响,但其主要受制于母体加性效应,并且存在一定的胚乳杂种优势和母体杂种优势;不同亲本品种对于粒重及加工品质性状的遗传改良具有不同的作用.     

谷类作物胚乳的遗传

胚乳是谷类作物种子的重要组成部分。一个完整的种子除了胚、种皮和果皮外,其余部分就是胚乳。由于它们最初发起的组织部位不同,其遗传行为也大不相同。有时仅从种子本身就可以观察到几种不同的遗传现象,如花粉直感、不同位点数基因的剂量效应和体细胞遗传等。正因为有这些复杂的遗传形式,在遗传学教学和实际应用中胚乳遗传常被忽视,并与2n体细胞的性状遗传相混淆。为了弄清这些复杂的遗传现象及种子各组成彼此间在遗传行为上的内在关系,本文特就这方面的内容作较系统介绍。 (一)胚乳的遗传及其与胚遗传的区别胚乳是双受精的产物。在正常情况下,谷类     

Multiple-interval mapping for quantitative trait loci controlling endosperm traits

Endosperm traits are trisomic inheritant and are of great economic importance because they are usually directly related to grain quality. Mapping for quantitative trait loci (QTL) underlying endosperm traits can provide an efficient way to genetically improve grain quality. As the traditional QTL mapping methods (diploid methods) are usually designed for traits under diploid control, they are not the ideal approaches to map endosperm traits because they ignore the triploid nature of endosperm. In this article, a statistical method considering the triploid nature of endosperm (triploid method) is developed on the basis of multiple-interval mapping (MIM) to map for the underlying QTL. The proposed triploid MIM method is derived to broadly use the marker information either from only the maternal plants or from both the maternal plants and their embryos in the backcross and F2 populations for mapping endosperm traits. Due to the use of multiple intervals simultaneously to take multiple QTL into account, the triploid MIM method can provide better detection power and estimation precision, and as shown in this article it is capable of analyzing and searching for epistatic QTL directly as compared to the traditional diploid methods and current triploid methods using only one (or two) interval(s). Several important issues in endosperm trait mapping, such as the relation and differences between the diploid and triploid methods, variance components of genetic variation, and the problems if effects are present and ignored, are also addressed. Simulations are performed to further explore these issues, to investigate the relative efficiency of different experimental designs, and to evaluate the performance of the proposed and current methods in mapping endosperm traits. The MIM-based triploid method can provide a powerful tool to estimate the genetic architecture of endosperm traits and to assist the marker-assisted selection for the improvement of grain quality in crop science. The triploid MIM FORTRAN program for mapping endosperm traits is available on the worldwide web (http://www.stat.sinica.edu.tw/chkao/).     

Mapping quantitative trait loci underlying triploid endosperm traits

Endosperm, which is derived from two polar nuclei fusing with one sperm, is a triploid tissue in cereals. Endosperm tissue determines the grain quality of cereals. Improving grain quality is one of the important breeding objectives in cereals. However, current statistical methods for mapping quantitative trait loci (QTL) under diploid genetic control have not been effective for dealing with endosperm traits because of the complexity of their triploid inheritance. In this paper, we derive for the first time the conditional probabilities of F(3) endosperm QTL genotypes given different flanking marker genotypes in F(2) plants. Using these probabilities, we develop a multiple linear regression method implemented via the iteratively reweighted least-squares (IRWLS) algorithm and a maximum likelihood method (ML) implemented via the expectation-maximization (EM) algorithm to map QTL underlying endosperm traits. We use the mean value of endosperm traits of F(3) seeds as the dependent variable and the expectations of genotypic indicators for additive and dominance effect of a putative QTL flanked by a pair of markers as independent variables for IRWLS mapping. However, if an endosperm trait is measured quantitatively using a single endosperm sample, the ML mapping method can be used to separate the two dominance effects. Efficiency of the methods is verified through extensive Monte Carlo simulation studies. Results of simulation show that the proposed methods provide accurate estimates of both the QTL effects and locations with very high statistical power. With these methods, we are now ready to map endosperm traits, as we can for regular quantitative trait under diploid control.     

Statistical methods for dissecting triploid endosperm traits using molecular markers. An autogamous model

The endosperm, a result of double fertilization in flowering plants, is a triploid tissue whose genetic composition is more complex than diploid tissue. We present a new maximum-likelihood-based statistical method for mapping quantitative trait loci (QTL) underlying endosperm traits in an autogamous plant. Genetic mapping of quantitative endosperm traits is qualitatively different from traits for other plant organs because the endosperm displays complicated trisomic inheritance and represents a younger generation than its mother plant. Our endosperm mapping method is based on two different experimental designs: (1) a one-stage design in which marker information is derived from the maternal genome and (2) a two-stage hierarchical design in which marker information is derived from both the maternal and offspring genomes (embryos). Under the one-stage design, the position and additive effect of a putative QTL can be well estimated, but the estimates of the dominant and epistatic effects are upward biased and imprecise. The two-stage hierarchical design, which extracts more genetic information from the material, typically improves the accuracy and precision of the dominant and epistatic effects for an endosperm trait. We discuss the effects on the estimation of QTL parameters of different sampling strategies under the two-stage hierarchical design. Our method will be broadly useful in mapping endosperm traits for many agriculturally important crop plants and also make it possible to study the genetic significance of double fertilization in the evolution of higher plants.     

Mapping epistatic quantitative trait loci underlying endosperm traits using all markers on the entire genome in a random hybridization design

Triploid endosperm is of great economic importance owing to its nutritious quality. Mapping endosperm trait loci (ETL) can provide an efficient way to genetically improve grain quality. However, most triploid ETL mapping methods do not produce unbiased estimates of the two dominant effects of ETL. A random hybridization design is an alternative method that may be used to overcome this problem. However, epistasis has an important role in the dissection of genetic architecture for complex traits. In this study, therefore, an attempt was made to map epistatic ETL (eETL) under a triploid genetic model of endosperm traits in a random hybridization design. The endosperm trait means of random hybrid lines, together with known marker genotype information from their corresponding parental F(2) plants, were used to estimate, efficiently and without bias, the positions and all of the effects of eETL using a penalized maximum likelihood method. The method proposed in this article was verified by a series of Monte Carlo simulation experiments. Results from the simulated studies show that the proposed method provides accurate estimates of eETL parameters with a low false-positive rate and a relatively short running time. This new method enables us to map triploid eETL in the same way as diploid quantitative traits.     

玉米籽粒性状的遗传效应分析

采用二倍体胚和三倍体胚乳种子遗传模型及其分析方法,以5个玉米自交系及其配制的F1,F2,BC1,BC2世代为材料,研究5个玉米种子性状的胚直接效应、胚乳直接效应、母体效应和细胞质效应。分析结果表明,除粒宽外,各性状的遗传同时由细胞质效应和胚、胚乳、母体基因效应所控制,百粒重主要受胚乳和母体效应的影响,粒长的遗传以母体效应为主,粒宽和粒厚以胚乳效应为主。各部位籽粒百粒重的胚乳直接加性效应与母体加性效应的协方差达到显著或极显著水平,其余性状的胚、胚乳直接效应与母体效应间的协方差均不显著,通过母体植株的遗传表现可以对这些性状进行有效的选择。S22 是改良百粒重的优良亲本。 Abstract：The embryo,endosperm and cytoplasm effects of seven seed traits were studied by genetic model for diploid embryo and triploid endosperm plant seeds using five inbreds and their F1, F2, BC1 and BC2 generations. The estimates of genetic variance components indicated that the inheritance of all other kernel traits was controlled by the four effects except kernel width. The 100?kernel weight was mainly controlled by endosperm and maternal effects , and kernel length was controlled by the maternal effects,while endosperm conrrolled kernel width and kernel thickness. Except the significant or highly significant covariances between the endosperm direct additive and maternal additive effects for 100－kernel weight,all other traits between the embryo or endosperm direct effect and the maternal were not significant. So,maize inbreds could be developed by direct selection based on maternal plants for these traits. S22 was the best inbred of the improvement for kernel weight in this study.     

The dynamic gene expression from different genetic systems for protein and lysine contents of indica rice

The dynamic expression of genes for protein and lysine contents of rice grain under different environments was carried out with time-dependent measures by using the developmental genetic models for quantitative traits of triploid endosperm in cereal crops. The results showed that the genetic effects, especially genotype × environment (GE) interaction effects from the genes expression of different genetic systems including triploid endosperm, cytoplasm and diploid maternal plant were important for the performance of both nutrient quality traits at all developmental times/stages of rice grain. The conditional genetic variance analysis found that the activation of quantitative genes especially from endosperm and maternal plant genetic systems for protein and lysine contents was gradually carried through the developmental process of rice grain. The net genetic effects showed that the new expression of quantitative genes for protein and lysine contents was more active at late filling stage (15–21 days after flowering) and maturity stage (22–28 days) of rice grain. Also the sequential expression of cytoplasmic genes cannot be ignorable for the development of nutrient quality traits. The phenomena that some genes could continuously express for several developmental stages or the genes expression could be interrupted among developmental stages of rice grain was detected especially for net endosperm additive main effects or maternal additive main effects. The differences of genetic relationships from different genetic systems were found for protein and lysine contents among developmental times of rice grain.     

Genetic analysis of non-essential amino acid contents in rice (Oryza sativa L.) across environments

Genetic main effects and genotype x environment (GE) interaction effects for 7 non-essential amino acids in milled rice were analyzed for two year data by using the genetic models based on mixed linear model approaches for quantitative traits of triploid endosperm. Nine cytoplasmic, male sterile lines as females and five restoring lines as males were introduced in a diallel cross in two environments. It was found that the content of non-essential amino acids including Asp, Ser, Glu, Gly and Tyr were mainly controlled by genetic main effects, whereas the content of Ala or Pro was mainly affected by GE effects. In genetic main effects, the cytoplasmic and maternal genetic effects were preponderant for all traits of non-essential amino acids, indicating that selection for improving these traits based on the maternal plant would be more effective than on seeds. The total narrow-sense heritabilities for non-essential amino acids were 70.9-85.9%. By predicating the genetic effects of parents, the total genetic effects from Xieqingzao, V20, Zuo 5 and Zhenshan 97 were mainly negative and these parents would decrease the content of most essential amino acids. Since parents of Zhenan 3, Yinchao 1, T49, 26715, 102 and 1391 had possessed a positive value of most total genetic effects, these parents could be chosen as optimal parents for increasing the content of most non-essential amino acids.