RNA Interference of Myocyte Enhancer Factor 2A Accelerates Atherosclerosis in Apolipoprotein E-Deficient Mice

Objective

Myocyte enhancer factor-2A (MEF 2A) has been shown to be involved in atherosclerotic lesion development, but its role in preexisting lesions is still unclear. In the present study we aim to assess the role of MEF 2A in the progression of pre-existing atherosclerosis.

Methods

Eighty apolipoprotein E-deficient mice (APOE KO) were randomly allocated to control, scramble and MEF 2A RNA interference (RNAi) groups, and constrictive collars were used to induce plaque formation. Six weeks after surgery, lentiviral shRNA construct was used to silence the expression of MEF 2A. Carotid plaques were harvested for analysis 4 weeks after viral vector transduction. Inflammatory gene expression in the plasma and carotid plaques was determined by using ELISAs and real-time RT-PCR.

Results

The expression level of MEF 2A was significantly reduced in plasma and plaque in the RNAi group, compared to the control and NC groups, whereas the expression level of pro-inflammatory cytokines was markedly increased. Silencing MEF 2A using lentiviral shRNA significantly reduced the plaque collagen content and fibrous cap thickness, as well as increased plaque area. However, silencing MEF 2A had no obvious effect on plaque lipid content.

Conclusions

Lentivirus-mediated MEF 2A shRNA accelerates inflammation and atherosclerosis in APOE KO mice, but has no effect on lipoprotein levels in plasma.     

Identifying Quantitative Trait Loci Affecting Resistance to Congenital Hypothyroidism in 129+Ter/SvJcl Strain Mice

Tyrosylprotein sulfotransferase 2 (TPST2) is one of the enzymes responsible for tyrosine O-sulfation and catalyzes the sulfation of the specific tyrosine residue of thyroid stimulating hormone receptor (TSHR). Since this modification is indispensable for the activation of TSH signaling, a non-functional TPST2 mutation (Tpst2^grt) in DW/J-grt mice leads to congenital hypothyroidism (CH) characterized by severe thyroid hypoplasia and dwarfism related to TSH hyporesponsiveness. Previous studies indicated that the genetic background of the 129^+Ter/SvJcl (129) mouse strain ameliorates Tpst2^grt-induced CH. To identify loci responsible for CH resistance in 129 mice, we performed quantitative trait locus (QTL) analysis using backcross progenies from susceptible DW/J and resistant 129 mice. We used the first principal component calculated from body weights at 5, 8 and 10 weeks as an indicator of CH, and QTL analysis mapped a major QTL showing a highly significant linkage to the distal portion of chromosome (Chr) 2; between D2Mit62 and D2Mit304, particularly close to D2Mit255. In addition, two male-specific QTLs showing statistically suggestive linkage were also detected on Chrs 4 and 18, respectively. All QTL alleles derived from the 129 strain increased resistance to growth retardation. There was also a positive correlation between recovery from thyroid hypoplasia and the presence of the 129 allele at D2Mit255 in male progenies. These results suggested that the major QTL on Chr 2 is involved in thyroid development. Moreover, since DW/J congenic strain mice carrying both a Tpst2^grt mutation and 129 alleles in the major QTL show resistance to dwarfism and thyroid hypoplasia, we confirmed the presence of the resistant gene in this region, and that it is involved in thyroid development. Further genetical analysis should lead to identification of genes for CH tolerance and, from a better understanding of thyroid organogenesis and function, the subsequent development of new treatments for thyroid disorders.     

Exercise Capacity and Response to Training Quantitative Trait Loci in a NZW X 129S1 Intercross and Combined Cross Analysis of Inbred Mouse Strains

Genetic factors determining exercise capacity and the magnitude of the response to exercise training are poorly understood. The aim of this study was to identify quantitative trait loci (QTL) associated with exercise training in mice. Based on marked differences in training responses in inbred NZW (-0.65 ± 1.73 min) and 129S1 (6.18 ± 3.81 min) mice, a reciprocal intercross breeding scheme was used to generate 285 F₂ mice. All F₂ mice completed an exercise performance test before and after a 4-week treadmill running program, resulting in an increase in exercise capacity of 1.54 ± 3.69 min (range = -10 to +12 min). Genome-wide linkage scans were performed for pre-training, post-training, and change in run time. For pre-training exercise time, suggestive QTL were identified on Chromosomes 5 (57.4 cM, 2.5 LOD) and 6 (47.8 cM, 2.9 LOD). A significant QTL for post-training exercise capacity was identified on Chromosome 5 (43.4 cM, 4.1 LOD) and a suggestive QTL on Chromosomes 1 (55.7 cM, 2.3 LOD) and 8 (66.1 cM, 2.2 LOD). A suggestive QTL for the change in run time was identified on Chromosome 6 (37.8 cM, 2.7 LOD). To identify shared QTL, this data set was combined with data from a previous F₂ cross between B6 and FVB strains. In the combined cross analysis, significant novel QTL for pre-training exercise time and change in exercise time were identified on Chromosome 12 (54.0 cM, 3.6 LOD) and Chromosome 6 (28.0 cM, 3.7 LOD), respectively. Collectively, these data suggest that combined cross analysis can be used to identify novel QTL and narrow the confidence interval of QTL for exercise capacity and responses to training. Furthermore, these data support the use of larger and more diverse mapping populations to identify the genetic basis for exercise capacity and responses to training.     

Overexpression of CCN3 Inhibits Inflammation and Progression of Atherosclerosis in Apolipoprotein E-Deficient Mice

Background

Cysteine-rich 61/connective tissue growth factor/nephroblastoma overexpressed (CCN) 3 has been recently reported to play a role in regulating inflammation of vascular endothelial cells. However, the role of CCN3 in atherosclerosis, which is characterized by vascular inflammation, remains unclear.

Hypothesis and Objectives

Overexpression of CCN3 may relieve the inflammation response in and inhibit the progress of atherosclerosis. We aimed to explore the potential roles of CCN3 in inflammation in atherosclerosis.

Strategy and Main Results

In in vitro studies using cultured human aortic endothelial cells and human umbilical vein endothelial cells, CCN3 mRNA and protein expression significantly decreased in response to tumor necrosis factor-α and interleukin-1β treatments (p<0.05), when analyzed by quantitative real-time polymerase chain reaction and Western blot. Using a mouse model of atherosclerosis, the mRNA and protein levels of CCN3 decreased by 72.2% (p = 0.041) and 86.4% (p = 0.036), respectively, compared with levels in wild-type control mice, respectively. Overexpression of CCN3 by adenovirus-mediated gene overexpression decreased low-density lipoprotein cholesterol by 48.9% (p = 0.017), total cholesterol by 58.9% (p = 0.031), and triglycerides by 56.8% (p = 0.022), and it increased high-density lipoprotein cholesterol level by 2.16-fold (p = 0.039), compared with control groups. Additionally, a reduced plaque area and increased fibrous cap were observed (p<0.05). Furthermore, CCN3 overexpression decreased cell adhesion molecule-1 mRNA expression by 84.7% (p = 0.007) and intercellular adhesion molecule-1 mRNA expression by 61.2% (p = 0.044). Inflammatory factors, including matrix metalloproteinases, cyclooxygenase 2, and tissue factor also significantly (p<0.05) decreased with CCN3 overexpression in the atherosclerotic mouse model. Additionally, CCN1 and CCN2, which have been reported to be highly expressed in aortic atherosclerotic plaques, were significantly downregulated (p<0.05) by CCN3 overexpression.

Conclusion

CCN3 overexpression is associated with control of inflammatory processes and reversion of dyslipidemia in the process of atherosclerosis, which implies that CCN3 may be a promising target in the treatment of atherosclerosis.     

Identification of Quantitative Trait Loci Controlling Levels of Radiation-Induced Thymocyte Apoptosis in Mice

Thymocyte apoptosis levels are higher in C57BL/6J mice than in C3Hf/Kam mice. Low-dose irradiation increases the numbers of thymocytes undergoing apoptosis, but the strain difference persists. We mapped three loci controlling radiation-induced thymocyte apoptosis levels in F2 intercross progeny of these strains. The strongest association of a genomic region with an apoptosis level occurred in a region of chromosome 11 known to harbor a locus (or loci) important in the pathogenesis of several rodent models of autoimmune disease. Additional loci influencing radiation-induced thymocyte apoptosis were identified on chromosomes 9 and 16. The genetic polymorphisms underlying these loci may have an evolutionary role in fine-tuning the apoptotic response in T cells and may be important in the etiology of lymphoproliferative disorders and autoimmunity.     

Fine‐mapping of Obesity‐related Quantitative Trait Loci in an F9/10 Advanced Intercross Line

Obesity develops in response to a combination of environmental effects and multiple genes of small effect. Although there has been significant progress in characterizing genes in many pathways contributing to metabolic disease, knowledge about the relationships of these genes to each other and their joint effects upon obesity lags behind. The LG,SM advanced intercross line (AIL) model of obesity has been used to characterize over 70 loci involved in fatpad weight, body weight, and organ weights. Each of these quantitative trait loci (QTLs) encompasses large regions of the genome and require fine‐mapping to isolate causative sequence changes and possible mechanisms of action as indicated by the genetic architecture. In this study we fine‐map QTLs first identified in the F₂ and F_2/3 populations in the combined F_9/10 advanced intercross generations. We observed significantly narrowed QTL confidence regions, identified many single QTL that resolve into multiple QTL peaks, and identified new QTLs that may have been previously masked due to opposite gene effects at closely linked loci. We also present further characterization of the pleiotropic and epistatic interactions underlying these obesity‐related traits.     

水稻F2不育和抽穗期QTL分析

对台中65（粳稻）/Bhadua（籼稻）杂交F2代群体构建了RFLP连锁图谱,含94个分布较为均匀的标记。对F2小穗不育性状进行单点分析和区间分析的结果基本一致：有两个F2小穗不育QTL座位分别位于染色体1的XNpb113～XNpb346之间和染色体8的G187～XNpb397之间,而且该两个QTL均为新检测出的座位;检测出5个抽穗期TQL,其中3个座位在单点分析和区间分析中的结果一致,分别位于染色体1的XNpb113～XNpb346,染色体4的C891～C335,染色体的8的C166～C1121,另外,染色体6的XNpb27为单点分析结果,染色体10的R716～C405为区间分析结果。由于染色体1上的F2不育QTL和抽穗期QTL重叠,该QTL座位是由于遗传效应所至还是由于环境因素（迟抽穗）所至有待构建近等基因系进一步研究。;位于染色体1和10上的抽穗期QTL座位为新检测的座位。对新检测的F2不育和抽穗期QTL座位正在建立相应的近等基因系以精确定位和克隆上述基因。     

Application of an Effective Statistical Technique for an Accurate and Powerful Mining of Quantitative Trait Loci for Rice Aroma Trait

When a phenotype of interest is associated with an external/internal covariate, covariate inclusion in quantitative trait loci (QTL) analyses can diminish residual variation and subsequently enhance the ability of QTL detection. In the in vitro synthesis of 2-acetyl-1-pyrroline (2AP), the main fragrance compound in rice, the thermal processing during the Maillard-type reaction between proline and carbohydrate reduction produces a roasted, popcorn-like aroma. Hence, for the first time, we included the proline amino acid, an important precursor of 2AP, as a covariate in our QTL mapping analyses to precisely explore the genetic factors affecting natural variation for rice scent. Consequently, two QTLs were traced on chromosomes 4 and 8. They explained from 20% to 49% of the total aroma phenotypic variance. Additionally, by saturating the interval harboring the major QTL using gene-based primers, a putative allele of fgr (major genetic determinant of fragrance) was mapped in the QTL on the 8th chromosome in the interval RM223-SCU015RM (1.63 cM). These loci supported previous studies of different accessions. Such QTLs can be widely used by breeders in crop improvement programs and for further fine mapping. Moreover, no previous studies and findings were found on simultaneous assessment of the relationship among 2AP, proline and fragrance QTLs. Therefore, our findings can help further our understanding of the metabolomic and genetic basis of 2AP biosynthesis in aromatic rice.     

Genetic Identification of Quantitative Trait Loci for Contents of Mineral Nutrients in Rice Grain

In present study, Fe, Zn, Mn, Cu, Ca, Mg, P and K contents of 85 introgression lines （ILs） derived from a cross between an elite indica cultivar Teqing and the wild rice （Oryza rufipogon） were measured by inductively coupled argon plasma （ICAP） spectrometry. Substantial variation was observed for all traits and most of the mineral elements were significantly positive correlated or independent except for Fe with Cu. A total of 31 putative quantitative trait loci （QTLs） were detected for these eight mineral elements by single point analysis. Wild rice （O. rufipogon） contributed favorable alleles for most of the QTLs （26 QTLs）, and chromosomes 1,9 and 12 exhibited 14 QTLs （45%） for these traits. One major effect of QTL for zinc content accounted for the largest proportion of phenotypic variation （11%-19%） was detected near the simple sequence repeats marker RM152 on chromosome 8. The co-locations of QTLs for some mineral elements observed in this mapping population suggested the relationship was at a molecular level among these traits and could be helpful for simultaneous improvement of these traits in rice grain by marker assisted selection.     

水稻幼苗活力性状的低温反应数量性状基因座检测

以籼粳交“密阳23/吉冷1号”的F2：3代200个家系作为作图群体,在12℃冷水胁迫下,进行苗高、苗鲜重和苗干重等水稻幼苗活力性状的低温反应鉴定,并利用由SSR标记构建的分子连锁图谱为基础,对冷水胁迫下苗高、苗鲜重和苗干重以及它们的低温反应指数进行了数量性状基因座（QTLs）检测。研究结果表明,低温胁迫下上述幼苗活力性状在F3家系群中均表现为接近正态的连续分布,表现为由多基因控制的数量性状;在第1、2、7、8和12染色体上,检测到与幼苗活力性状的低温反应相关的QTL共12个,对表型变异的贡献率范围为5．2％-17．9％,其中位于第2染色体RM262-RM263区间和第12染色体RM270-RM17区间的与低温下苗高相关的qCSH2和qCSH12,以及位于第12染色体RM19-RM270区间和第1染色体RM129-RM9区间的分别控制低温下苗干重及其低温反应指数的qSDW12和qCSDW1对表型变异的贡献率较大,分别为16．6％、17．9％、15．9％和16．2％。其增效等位基因均来自吉冷1号,前两者均表现为加性效应,后两者分别表现为显性和超显性。     

Genetic Identification of Quantitative Trait Loci for Contents of Mineral Nutrients in Rice Grain

In present study, Fe, Zn, Mn, Cu, Ca, Mg, P and K contents of 85 Introgression linee (ILs) derived from a cross between an elite indica cultivar Teqing and the wild rice (Oryza rufipogon) were measured by inductively coupled argon plasma (ICAP) spectrometry. Substantial variation was observed for all traits and most of the mineral elements were significantly positive correlated or independent except for Fe with Cu. A total of 31 putative quantitative trait loci (QTLs) were detected for these eight mineral elements by single point analysis. Wild rice (O. rufipogon) contributed favorable alleles for most of the QTLs (26 QTLs), and chromosomes 1, 9 and 12 exhibited 14 QTLs (45%) for these traits. One major effect of QTL for zinc content accounted for the largest proportion of phenotypic variation (11%-19%) was detected near the simple sequence repeats marker RM152 on chromosome 8. The co-locations of QTLs for some mineral elements observed in this mapping population suggested the relationship was at a molecular level among these traits and could be helpful for simultaneous improvement of these traits in rice grain by marker assisted selection.     

Identification of Quantitative Trait Loci Influencing Wood Specific Gravity in an Outbred Pedigree of Loblolly Pine

We report the identification of quantitative trait loci (QTL) influencing wood specific gravity (WSG) in an outbred pedigree of loblolly pine (Pinus taeda L.). QTL mapping in an outcrossing species is complicated by the presence of multiple alleles (>2) at QTL and marker loci. Multiple alleles at QTL allow the examination of interaction among alleles at QTL (deviation from additive gene action). Restriction fragment length polymorphism (RFLP) marker genotypes and wood specific gravity phenotypes were determined for 177 progeny. Two RFLP linkage maps were constructed, representing maternal and paternal parent gamete segregations as inferred from diploid progeny RFLP genotypes. RFLP loci segregating for multiple alleles were vital for aligning the two maps. Each RFLP locus was assayed for cosegregation with WSG QTL using analysis of variance (ANOVA). Five regions of the genome contained one or more RFLP loci showing differences in mean WSG at or below the P = 0.05 level for progeny as grouped by RFLP genotype. One region contained a marker locus (S6a) whose QTL-associated effects were highly significant (P > 0.0002). Marker S6a segregated for multiple alleles, a prerequisite for determining the number of alleles segregating at the linked QTL and analyzing the interactions among QTL alleles. The QTL associated with marker S6a appeared to be segregating for multiple alleles which interacted with each other and with environments. No evidence for digenic epistasis was found among the five QTL.     

油菜油分、蛋白质和硫苷含量相关性分析及QTL 定位

为定位与油分、蛋白质和硫苷含量等品质性状相关的数量性状位点(QTL), 以2个含油量较高的甘蓝型油菜(Brassica napus)品系8908B和R1为研究材料, 配置正反交组合。在正反交F₂代群体中, 含油量和蛋白质含量都存在极显著的负相关, 相关系数分别为-0.68和-0.81, 含油量和硫苷含量相关性不显著; 蛋白质含量和硫苷含量在正交群体中相关性不显著, 但在反交群体中存在显著负相关(相关系数r =-0.45)。利用正交F₂代群体中的118个单株, 构建了包含121个标记的遗传连锁图谱, 图谱长1 298.7 cM, 有21个连锁群(LGs)。采用复合区间作图法, 在连锁图上定位了2个与含油量有关的QTL, 分别位于LG8和LG10, 其贡献率分别为4.8%和13.7%, 增效基因都来源于R1; 定位了2个与蛋白质含量有关的QTL: pro1 和 pro2, 分别位于LG1和LG3, 其贡献率分别为15.2%和14.1%, 位点pro1由8908B提供增效基因, pro2则由R1提供增效基因; 定位了4个与硫苷含量有关的QTL, 其中LG20上有2个, LG4和LG8上各1个, 它们的贡献率在1.9%-25.4%之间, 除LG20上glu1的增效基因来自R1外, 其余3个QTL位点均由8908B提供增效基因。     

Identification of Quantitative Trait Loci (QTLs) for Fruit Quality Traits in Apple

A segregating mapping population of “Co-op 17” × “Co-op 16” was used to identify quantitative trait loci (QTLs) associated with various fruit quality traits in apple. Phenotypic data were collected over 2 years for fruit circumference (in centimeter), diameter at midpoint (in centimeter), length (in centimeter), weight (in gram), total soluble solids (in degree Brix), and total titratable acids (in percent) for the segregating population. The phenotypic data along with a previously constructed genetic map, based on simple sequence repeat markers derived from expressed sequence tag and bacterial artificial chromosome end sequence databases, were used in marker–trait association analysis. Interval mapping identified two QTLs linked to fruit size components on linkage groups 03 and 05 with limit of detection scores of 3.27–4.06 and 3.29–4.02 along with phenotypic variation accounting for 15.4–46.4 and 18.3–21.9 %, respectively.     

油菜油分、蛋白质和硫苷含量相关性分析及QTL定位

为定位与油分、蛋白质和硫苷含量等品质性状相关的数量性状位点（QTL）,以2个含油量较高的甘蓝型油菜（Brassica napus）品系8908B和R1为研究材料,配置正反交组合。在正反交F2代群体中,含油量和蛋白质含量都存在极显著的负相关,相关系数分别为-0．68和-0．81,含油量和硫苷含量相关性不显著：蛋白质含量和硫苷含量在正交群体中相关性不显著,但在反交群体中存在显著负相关（相关系数r=-0．45）。利用正交F2代群体中的118个单株,构建了包含121个标记的遗传连锁图谱,图谱长1298．7cM,有21个连锁群（LGs）。采用复合区间作图法,在连锁图上定位了2个与含油量有关的QTL,分别位于LG8和LG10,其贡献率分别为4．8％和13．7％,增效基因都来源于R1;定位了2个与蛋白质含量有关的QTL：pr01和pr02,分别位于LG1和LG3,其贡献率分别为15．2％和14．1％,位点pr07由8908B提供增效基因,pro2则由R1提供增效基因：定位了4个与硫苷含量有关的QTL,其中LG20上有2个,LG4和LG8上各1个,它们的贡献率在1．9％-25．4％之间,除LG20上glu7的增效基因来自R1外,其余3个QTL位点均由8908B提供增效基因。     

Statistical Parameters in Behavioral Tasks and Implications for Sample Size of C57BL/6J:129S6/SvEvTac Mixed Strain Mice

Most mixed strain progeny from gene-knockout experiments typically originate from C57BL/6J and one of the 129 substrains, frequently 129S6/SvEvTac. The results of this behavioral survey suggest that C57BL/6J:129S6/SvEvTac mixed strain mice are amenable to behavioral testing. The variability in behavioral tasks for subjects arising from this mixed strain genetic background does not preclude screening with a battery of behavioral tests. With clues provided by a screen of mixed strain subjects, follow-up analyses with isogenic, congenic, or F1 hybrid animals may be targeted to specific behavioral themes.     

Identification of Quantitative Trait Loci for Grain Appearance and Milling Quality Using a Doubled-Haploid Rice Population

Improving grain quality, which is composed primarily of the appearance of the grain and its cooking and milling attributes, is a major objective of many rice-producing areas in China. In the present study, we conducted a marker-based genetic analysis of the appearance and milling quality of rice （Oryza sativa L.） grains using a doubled-haploid population derived from a cross between the indica inbred Zhenshan 97 strain and the japonica inbred Wuyujing 2 strain. Quantitative trait locus （QTL） analysis using a mixed linear model approach revealed that the traits investigated were affected by one to seven QTLs that individually explained 4.0%-30.7% of the phenotypic variation. Cumulatively, the QTL for each trait explained from 12.9% to 61.4% of the phenotypic variation. Some QTLs tended to have a pleiotropic or location-linked association as a cause of the observed phenotypic correlations between different traits. Improvement of the characteristics of grain appearance and grain weight, as well as an improvement in the milling quality of rice grains, would be expected by a recombination of different QTLs using marker-assisted selection.     

水稻种子萌发和苗期ABA敏感性的QTL定位分析

植物激素ABA参与不同的生理过程,尤其是在种子发育和非生物逆境的适应都需要ABA的调控.以水稻珍汕97和旱稻IRAT109为亲本的重组自交系群体为材料,分别调查种子发芽和苗期对ABA的敏感程度.种子发芽阶段以ABA处理下的相对发芽势（Relative germination vigor,RGV）和相对发芽率（Relative germination rate,RGR）为指标,苗期以ABA喷施处理下的卷叶程度（Leaf rolling scores by ABA spaying,LRS）和含ABA水培条件下的卷叶程度（Leaf rolling scores by ABA culturing,LRC）为指标.性状相关分析表明发芽阶段的相对发芽势与苗期卷叶程度呈显著正相关.用复合区间作图法和混合线性模型对ABA敏感性QTL定位和上位性效应分析.两种软件检测到的主效QTL位点大致相同.共检测到5个单位点QTL和6对上位性QTL与发芽阶段的ABA敏感性有关;8个单位点QTL和5对上位性QTL与水稻苗期对ABA的敏感性有关;在苗期,两种ABA处理条件下共检测到两个共同的QTL;仅一个共同的QTL同时控制发芽阶段和苗期对ABA的敏感性.这些研究结果说明,水稻对ABA的敏感性同时受单位点的多基因和上位性基因控制;而且控制种子萌发阶段发芽势和苗期对ABA敏感性的遗传基础有很大的不同.     

玉米雌雄开花间隔天数、结穗率与产量的数量性状位点（QTL）分析

采用SSR标记连锁图谱和复合区间作图法在山西灌溉和干旱胁迫条件下,对玉米(Zea mays L.)自交系黄早四×掖107组合的F_3群体雌雄开花间隔天数(ASI)、结穗率和籽粒产量进行了数量性状位点(QTL)定位及基因效应分析。结果表明,在两种水分处理下,ASI、结穗率与籽粒产量的相关性均达到显著水平(P<0.05)。在灌溉和干旱胁迫卜,分别检测到3个和2个控制ASI的QTL,位于第1、2、3和第2、5染色体上。在灌溉条件下,在第3和第6染色体上各检测到1个控制结穗率的QTL,基因作用方式呈加性或部分显性,可解释19.9％的表型变异;在干旱条件下,在第3、7、10染色体上共检测到4个控制结穗率的QTL,基因作用方式为显性或部分显性,可解释60.4％的表型变异。在灌溉和干旱胁迫下,控制产量的QTL分别定位在第3、6、7和第1、2、4、8染色体上,基因作用方式均以加性或部分显性为主,可解释的表型变异为7.3％～22.0％。在干旱条件下,借助连锁分子标记和基因效应分析,可构建包含ASI、结穗率和产量QTL的选择指数,用于分子标记辅助育种。