甘蓝型油菜分枝角度主基因+多基因混合遗传模型及遗传效应

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汪文祥, 胡琼, 梅德圣, 李云昌, 周日金, 王会, 成洪涛, 付丽, 刘佳. . 甘蓝型油菜分枝角度主基因+多基因混合遗传模型及遗传效应. 作物学报, 2016, 42(8): 1103-1111
WANG Wen-Xiang, HU Qiong, MEI De-Sheng, LI Yun-Chang, ZHOU Ri-Jin, WANG Hui, CHENG Hong- Tao, FU Li, LIU Jia. . Genetic Effects of Branch Angle Using Mixture Model of Major Gene Plus Polygene in Brassica napus L. . ACTA AGRONOMICA SINICA, 2016, 42(8): 1103-1111 复制到剪切板

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甘蓝型油菜分枝角度主基因+多基因混合遗传模型及遗传效应

汪文祥, 胡琼, 梅德圣, 李云昌, 周日金, 王会, 成洪涛, 付丽, 刘佳^*

中国农业科学院油料作物研究所 / 农业部油料作物生物学与遗传育种重点实验室, 湖北武汉430062

^*通讯作者(Corresponding author): 刘佳, E-mail: liujia02@caas.cn, Tel: 027-86711556

第一作者联系方式: E-mail:wangwx8@gmail.com

收稿日期:2016-01-14 接受日期:2016-05-09 网络出版日期:2016-05-30

基金:本研究由中国农业科学院科技创新工程项目(Group No.118), 国家农业现代产业技术体系建设专项(CARS-13), 国家自然科学基金项目(31471535), 湖北省自然科学基金项目(2014CFB156)和湖北省科技创新工程项目资助

摘要

分枝角度是油菜株型重要性状, 是油菜品种高产及适合机械化收获理想株型的基本组成之一。为明确油菜分枝角度的遗传, 本研究选用油菜分枝角度大的松散型材料6098B和分枝角度小的紧凑型材料Purler配制杂交组合, 采用主基因+多基因混合遗传模型方法对该组合6世代(P₁、P₂、F₁、F₂、BCP₁和BCP₂)的分枝角度进行了遗传分析。结果表明, 上部第一分枝(顶枝)和基部第一分枝(基枝)角度的最适合遗传模型均为D-0 (1对加性-显性主基因+加性-显性-上位性多基因)。顶枝角的主基因加性效应值为4.939º, 显性效应值为-4.156º, 主基因遗传率在BCP₁、BCP₂和F₂中分别是34.08%、1.40%和14.99%, 多基因遗传率分别为24.43%、61.72%和63.98%; 而基枝角的主基因加性效应值为2.217º, 显性效应值为-1.941º, 主基因遗传率在BCP₁、BCP₂和F₂中分别是7.86%、1.24%和4.84%, 多基因遗传率分别为66.46%、58.49%和73.96%。结果发现油菜分枝角度明显存在主效基因, 为油菜分枝角度的遗传改良奠定了基础。

关键词: 油菜; 分枝角度; 理想株型; 主基因+多基因; 遗传效应

Genetic Effects of Branch Angle Using Mixture Model of Major Gene Plus Polygene in Brassica napus L.

WANG Wen-Xiang, HU Qiong, MEI De-Sheng, LI Yun-Chang, ZHOU Ri-Jin, WANG Hui, CHENG Hong- Tao, FU Li, LIU Jia^*

Oil Crops Research Institute of Chinese Academy of Agricultural Sciences / Key Laboratory of Biology and Genetic Improvement of Oil Crops of the Ministry of Agriculture, Wuhan 430062, China

Fund:This study was support by the Science and Technology Innovation Project of Chinese Academy of Agricultural Sciences (Group No.118), the Earmarked Fund for China Agriculture Research System (CARS-13), the Natural Science Foundation of China (31471535), the Natural Science Foundation of Hubei Province (2014CFB156) and the Hubei Agricultural Science and Technology Innovation Center

Abstract

Branch angle is one of important plant architectural traits in rapeseed ( Brassica napus), which is the basic requirement for high-density cultivation and mechanical harvesting. To reveal the genetic mechanism of branch angle in rapeseed, we produced six generations (P₁, P₂, F₁, F₂, BCP₁, and BCP₂) from a cross between loose-type 6098B and compact-type Purler to analyze genetic effects of branch angle in rapeseed by the mixed major gene plus polygene inheritance model. The branch angle showed a continuous normal distribution, where both the basal branch angle and the top branch angle were dominated by a pair of major gene with additive-dominant effects plus polygenes with additive-dominance-epistasis effects (D-0 model). The additive effect and dominant effect of major gene of the top branch angle were 4.939 and -4.156, the heritability in BCP₁, BCP₂, and F₂ was 34.08%, 1.40%, and 14.99%, respectively for the major gene, and 24.43%, 61.72%, and 63.98%, respectively for the polygenes. The additive effect and dominant effect of major gene of the basal branch angle were 2.217 and -1.941, the heritability in BCP₁, BCP₂, and F₂ was 7.86%, 1.24%, and 4.84%, respectively for the major gene, and 66.46%, 58.49%, and 73.96%, respectively for the polygenes. Results of this study found that the branch angle traits of 6098B×Purler is controlled by polygenes, but there is a major gene among them. The research may be useful for further understanding and genetic improvement of plant architecture in rapeseed.

Keyword: Brassica napus L.; Branch angle; Ideal plant architecture; Major gene plus polygene; Genetic effects

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油菜是我国种植面积最大的油料作物, 年种植面积约730万公顷(2015中国统计年鉴)。油菜籽不仅仅用来榨油, 菜籽饼粕也是重要的工业原料, 并与畜牧和水产养殖业发展高度关联。近年来, 油料供给短缺的状态一直未得到有效解决, 甚至日趋紧张^[1]。单产水平低和机械化生产普及率低是限制我国油菜产业发展的主要瓶颈。培育适合高效机械化生产油菜品种, 构建油菜理想株型, 改良油菜在密植环境下的生长发育情况, 是提高油菜产量的基础, 也是促进机械化收获提高油菜生产效益的关键。

株高、分枝部位高、分枝数和主花序有效长是油菜株型重要的组成部分, 相关的研究较多^{[2, 3, 4, 5, 6, 7]}, 但关于油菜分枝角度的研究较少。分枝角度主要指有效分枝与主茎之间所成的夹角。分枝角度大, 植株松散, 增加了单个植株的占地面积, 降低了叶面积指数及单位面积上光合作用面积, 相应降低了产量; 植株间相互交织, 影响光能透射和通风, 不仅因降低植株中下部绿色组织的光能利用率而降低群体产量, 同时增加了病害的风险及机械化收割时刮碰造成的产量损失。减小分枝角度对增加油菜种植密度, 实现高产高效生产意义重大。

油菜分枝角度的遗传研究, 对培育紧凑株型油菜品种具有重要理论指导作用。前人的研究基本局限于分枝角度与产量性状的相关性^{[8, 9]}, 而对油菜分枝角度基因的效应大小及其作用形式很少报道。张文学等^[10]首次研究了甘蓝型油菜紧凑型和松散型杂种一代分枝性状的遗传效应, 结果表明紧凑型与松散型正反交F₁代分枝角度无差异, 表现双亲平均值, 说明细胞质效应不影响分枝角度。王俊生等^[11]利用6个油菜品种构建30个完全双列杂交组合, 对分枝角度、一次分枝数、分枝长等10多个株型性状进行了遗传效应及其与产量性状的相关性分析, 结果表明分枝角度狭义遗传率较高, 显性效应对分枝角度有减效作用, 并与单株产量特别是单株角果数存在显著的遗传负相关。近几年, 为构建机械化密植条件下的油菜理想株型, 分枝角度的研究也逐渐得到重视^{[12, 13]}, Liu等^[14]利用油菜自然群体, 定位了6个与分枝角度显著关联的基因组区域, 并获得其中4个区域内候选基因。但迄今为止仍缺少利用遗传群体对该性状进行传统遗传研究的报道。

主基因+多基因混合遗传模型是由盖钧镒^[15]提出的数量性状遗传分析方法, 在多种植物的遗传研究中得到应用^[16], 在油菜性状的遗传研究中也广泛应用^{[9, 17, 18, 19, 20, 21]}。本文以6个世代的研究群体(P₁、P₂、F₁、F₂、BCP₁和BCP₂), 利用数字图像采集法鉴定油菜上部第一分枝角(顶枝角)和基部第一分枝角(基枝角), 运用主基因+多基因混合遗传模型分析分枝角度的遗传模式和遗传效应, 评估单个主基因不同的遗传效应和各分离世代的遗传率, 揭示分枝角度的遗传规律。研究结果不仅为改良油菜株型性状提供理论指导, 也为进一步QTL定位和相关基因的克隆奠定基础。

1 材料与方法

1.1 试验材料

亲本材料分别为6098B (P₁)和Purler (P₂), 6098B是中国农业科学院油料作物研究所杂种优势利用课题组选育, 属于松散型; Purler是引自澳大利亚的油菜品种, 属于紧凑型。2013年秋, 在中国农业科学院油料作物研究所阳逻试验基地种植亲本, 2014年春手工去雄授粉杂交获得F₁(P₁× P₂)种子; 同年, 将亲本P₁、P₂和F₁种植于青海大学试验基地, 将F₁套袋自交收获F₂种子, 再以2个亲本分别与F₁回交获得BCP₁(6098B/Purler//6098B)和BCP₂ (6098B/Purler// Purler)。从而获得6098B/Purler杂交组合的6个基本世代群体, 即P₁、P₂、F₁、F₂、BCP₁和BCP₂群体。亲本材料由中国农业科学院油料作物研究所油菜杂种优势利用课题组保存提供。

1.2 田间试验

2014年9月, 将6个世代群体种子同期播种在中国农业科学院油料作物研究所阳逻试验基地, 行距33 cm, 株距11 cm, 每行15株, 田间管理与普通大田生产基本相同。种植P₁、P₂和F₁群体各6行, F₂群体18行, BCP₁和BCP₂群体各12行。试验地肥力均匀一致, 参照当地大田栽培的施肥水平。2015年5月8日避开行两端的边株分单株收获群体待表型考察。

1.3 分枝角度鉴定方法

2015年6月在室内采用数字图像处理法^[22]鉴定6世代群体分枝角度: 油菜成熟后, 剪取连有油菜上部第一分枝(顶枝)和基部第一分枝(基枝)的茎段, 采集图像。将该图像文件导入AutoCAD软件, 利用角度工具标注夹角角度, 将分枝角度结果记录到Excel文档中。

1.4 数据统计分析

利用SAS 9.3统计分析软件进行基本统计量、相关系数及多组数据差异显著性比较。利用植物数量性状主基因+多基因混合遗传模型分析方法^[15], 并结合软件^[23]数据处理进行遗传模型分析(分析软件由南京农业大学章元明教授提供)。通过极大似然分析和IECM (iterated expectation and conditional maximization)算法对混合分布中的相关分布参数作出估计, 再采用AIC (Akaike’ s information criterion)值选择最佳模型, 并进行适合性检验, 包括均匀性U₁²、U₂²和U₃²检验, Smirnov检验(nW²)和Kolmogorov检验(D_n), 根据检验结果选择最优遗传模型。最后采用最小二乘法依据最优模型的各成分分布参数估计各基因效应值、方差等遗传参数。群体的多基因方差σ _pg²= σ _p²-σ _mg²-σ ², h_mg²= σ _mg²/σ _p²× 100%, 多基因遗传率h_pg²= σ _pg²/σ _p²× 100%, 其中σ _mg²、σ _pg²、σ _p²和σ ²分别表示主基因方差、多基因方差、表型方差和环境方差。

2 结果与分析

2.1 表型数据分析

由表1可知, 亲本P₁(6098B)的顶枝角和基枝角分别为71.7° ± 5.8° 和50.6° ± 6.8° ; 亲本P₂(Purler)的顶枝角度和基枝角分别为26.2° ± 4.5° 和15.4° ± 2.7° ; 2个亲本2个部位的分枝角间差异均达到显著水平(P< 0.05)。F₁的顶枝角和基枝角均值分别为41.6° ± 6.2° 和38.0° ± 5.4° , 介于双亲之间, 略偏向高值亲本。F₂群体的顶枝角和基枝角均值分别为39.2º ± 10.9º 和34.5º ± 8.9º , 其分布为偏态分布, 表明分枝角度受到多基因的控制。F₁与亲本(P₁)回交的BCP₁群体的顶枝角和基枝角均值分别为47.0° ± 7.8° 和41.0° ± 8.1° , 偏向于P₁亲本(图1)。F₁与亲本(P₂)回交的BCP₂群体的顶枝角度和基枝角均值分别为32.2° ± 8.2° 和26.4° ± 7.2° , 偏向于亲本P₂(图2)。利用所有单株进行统计分析, 结果表明顶枝角与基枝角的相关系数为0.611 (P < 0.01, n = 419)。

表1 6098B× Purler 各世代分枝角度统计及变异系数 Table 1 Performance of branch angle in six generations from 6098B× Purler in B. napus

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	图1 6098B× Purler杂交组合顶枝角的频次分布 A: BCP₁分离群体的频次分布; B: BCP₂分离群体的频次分布; C: F₂分离群体的频次分布。Fig. 1 Frequency distribution of top branch angle from 6098B× Purler A: Frequency distribution in the BCP₁populations; B: Frequency distribution in the BCP₂ populations; C: Frequency distribution in the F₂ populations.

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图2 6098B× Purler杂交组合基枝角的频次分布
A: BCP₁分离群体的频次分布; B: BCP₂分离群体的频次分布; C: F₂分离群体的频次分布。Fig. 2 Frequency distribution of basal branch angle from 6098B× Purler
A: Frequency distribution in the BCP₁populations; B: Frequency distribution in the BCP₂ populations; C: Frequency distribution in the F₂ populations.

2.2 分枝角度主基因+多基因遗传分析

2.2.1 遗传模型的选择利用植物数量性状主基因+多基因混合遗传模型的6世代(P₁、P₂、F₁、BCP₁、BCP₂和F₂)联合分析, 估算获得1对主基因(A类模型)、2对主基因(B类模型)、多基因(C类模型)、1对主基因+多基因(D类模型)、2对主基因+多基因(E类模型)共5类24种遗传模型的极大似然函数值和AIC值(表2)。根据遗传模型的选取原则, 即AIC值最小准则, 选取AIC值最小及与最小AIC值比较接近的一组遗传模型作为备选模型。结果显示24种混合模型中, 油菜顶枝角和基枝角的D-0 (1对加性-显性主基因+加性-显性-上位性多基因)和E-1 (2对加性-显性-上位性主基因+加性-显性多基因)模型的AIC值相对较小, 可作为备选模型。

2.2.2 候选模型的适合性测验对备选模型进行一组(U₁²、U₂²、U₃²、_nW²和D_n)适合性检验(表3), 结果表明, 顶枝角的D-0模型和E-1模型没有统计量的差异达到显著水平(P < 0.05), 根据AIC值大小确定顶枝角的最适模型为D-0 (1对加性-显性主基因+加性-显性-上位性多基因); 而基枝角的2个备选模型中D-0模型的1个统计量达到显著水平, E-1模型没有统计量达到显著水平, 根据AIC值和达到显著水平统计量最少的原则, 基枝角的最适模型为D-0 (1对加性-显性主基因+加性-显性-上位性多基因)。

表2 6098B× Purler组合后代各遗传模型的AIC值 Table 2 AIC values of the genetic models from 6098B× Purler

模型代码 Model code	模型含义 Implication of model	AIC值 AIC value		极大似然函数值log_Max_likelihood_value
模型代码 Model code	模型含义 Implication of model	顶枝角 Top branch angle	基枝角 Basal branch angle	顶枝角 Top branch angle	基枝角 Basal branch angle
A-1	1MG-AD	2891.41	2801.24	-1441.70	-1396.62
A-2	1MG-A	2896.16	2806.81	-1445.08	-1400.40
A-3	1MG-EAD	2919.70	2796.71	-1456.85	-1395.35
A-4	1MG-AEND	2912.20	2858.57	-1453.10	-1426.29
B-1	2MG-ADI	2836.36	2717.54	-1408.18	-1348.77
B-2	2MG-AD	2879.57	2771.67	-1433.78	-1379.83
B-3	2MG-A	2885.88	2738.07	-1438.94	-1365.04
B-4	2MG-EA	2866.24	2737.06	-1430.12	-1365.53
B-5	2MG-AED	2973.21	2797.38	-1482.61	-1394.69
B-6	2MG-EEAD	2971.21	2795.38	-1482.61	-1394.69
C-0	PG-ADI	2846.25	2735.20	-1413.13	-1357.60
C-1	PG-AD	2860.88	2711.22	-1423.44	-1348.61
D-0	MX1-AD-ADI	2832.34	2704.68	-1404.17	-1340.34
D-1	MX1-AD-AD	2863.60	2710.13	-1422.80	-1346.06
D-2	MX1-A-AD	2860.09	2708.13	-1422.05	-1346.07
D-3	MX1-EAD-AD	2862.28	2708.21	-1423.14	-1346.11
D-4	MX1-AEND-AD	2851.54	2707.49	-1417.77	-1345.74
E-0	MX2-ADI-ADI	2840.74	2713.49	-1402.37	-1338.75
E-1	MX2-ADI-AD	2835.83	2706.85	-1402.92	-1338.43
E-2	MX2-AD-AD	2868.25	2712.78	-1423.12	-1345.39
E-3	MX2-A-AD	2847.70	2706.95	-1414.85	-1344.48
E-4	MX2-EA-AD	2862.21	2708.11	-1423.10	-1346.05
E-5	MX2-AED-AD	2864.28	2710.21	-1423.14	-1346.11
E-6	MX2-EEAD-AD	2862.28	2708.22	-1423.14	-1346.11

MG: major gene model; MX: mixed major gene and polygene model; PG: polygene model; A: additive effect; D: dominance effect; I: interaction; N: negative; E: equal; e.g. Model E-1 = MX2-ADI-AD, means mixed model with two major genes of additive-dominance- epistasis effects plus additive-dominance polygene. Minimum AIC value in each cross is underlined, showing the good-fitting model.

MG: 主基因模型; MX: 主基因+多基因混合模型; PG: 多基因遗传模型; A: 加性效应; D: 显性效应; I: 互作; N: 负向; E: 相等; 例如E-1模型MX2-ADI-AD, 表示2对加性-显性-上位性主基因+加性-显性多基因混合遗传模型; 下画线表示AIC值最小, 用于最适遗传模型检验。

表2 6098B× Purler组合后代各遗传模型的AIC值 Table 2 AIC values of the genetic models from 6098B× Purler

表3 6098B× Purler群体适合性检验 Table 3 Fitness tests of selected models in 6098B× Purler

2.2.3 遗传参数估计根据选择的最优遗传模型估计顶枝角和基枝角的最优遗传模型一阶、二阶遗传参数(表4)。结果表明, 顶枝角和基枝角的主基因加性效应(d_a)均是正值; 而显性效应(h_a)分别为-4.156和-1.941, 均为负值, 主基因显性效应与加性效应的比值均为︱h_a/d_a︱< 1, 即显性效应都小于相应的加性效应, 表明控制分枝角度的1对主基因以加性效应为主。在显性效应的作用方向上, 1对主基因表现为分枝角度大对分枝角度小为部分显性。显性效应增大分枝角度, 表现为部分平生型分枝。

对于油菜顶枝角, BCP₁、BCP₂和F₂分离世代群体遗传率分别为58.51%、63.12%和78.97%, 其中主基因遗传率分别为34.08%、1.40%和14.99%, 多基因遗传率分别为24.43%、61.72%和63.98%, 各世代的多基因遗传率都大于相应世代的主基因遗传率, 可见油菜分枝角度的遗传体系中, 微效多基因的遗传贡献占主要部分; 3个分离世代群体环境因素引起的变异分别占11.23%、35.97%和7.10%; 可见油菜分枝角性状受基因型和环境双重影响。同样, 对于油菜基枝角度, BCP₁、BCP₂和F₂分离世代群体遗传率分别为74.32%、59.73%和78.80%, 其中主基因遗传率分别为7.86%、1.24%和4.84%, 多基因遗传率分别为66.46%、58.49%和73.96%, 各世代的多基因遗传率都大于相应世代的主基因遗传率。可见基枝角的遗传体系中, 微效多基因的遗传贡献占主要部分; 3个分离世代群体环境因素引起的变异分别占19.35%、39.54%和17.21%。

表4 6098B× Purler杂交组合分枝角度的一阶遗传参数估计值 Table 4 First order genetic parameters estimated for the fitting models of branch angle from 6098B× Purler

表5 6098B× Purler杂交组合分枝角度的二阶遗传参数估计值 Table 5 Second order genetic parameters estimated for the fitting models of branch angle from 6098B× Purler

3 讨论

分枝角度是油菜株型重要的性状, 与产量、耐密性及机械化收获紧密相关。提高种植密度对提高油菜单产和机收适应性都具有非常重要的作用。密植除了增加植株的叶面积指数和光能利用效率外^[24], 还能提高油菜氮肥利用效率^[25], 促进氮素向籽粒转移^[26], 起到增产效果。密植还可以提高油菜群体的整齐度, 使茎杆变细、分枝部位提高、分枝长度变短及成熟期相对集中, 有利于机械联合收获, 显著降低机收损失率^[27]。密植栽培要求选择株型紧凑、分枝角度较小的品种, 避免植株间相互交织, 有利于光能透射和通风, 提高植株中下部绿色组织的光能利用率, 从而提高群体产量, 也降低了病害的风险。王俊生等^[11]的研究发现分枝角度与单株产量特别是单株角果数存在显著的遗传负相关, 说明分枝角度小的植株在生长中后期的物质积累中占有优势, 由于中下部光合作用受到的影响较小, 促进了有效分枝数和有效角果数的提高。紧凑型油菜在密植条件下植株间不相互交叉, 可以降低机械收割时刮碰造成的产量损失, 更有利于机械联合收割。因此, 改良油菜株型, 不仅能促进油菜产量的提高也可以推动机械化生产的实施, 符合我国油菜发展的长远目标。

分枝角度形成的直接原因是分枝感受向重力性, 受到植株内部生长素和独角金内酯等激素的调节^{[28, 29]}。油菜具多分枝习性, 常规种植密度下(23~30万株 hm^-2), 油菜的平均分枝数为5~10个。根据田间观察, 油菜植株从下到上的分枝角度一般逐渐加大, 本研究获得的数据证实了这一趋势。从试验结果看, 顶枝和基枝分枝角度相关性极显著, 相关系数为0.611, 说明顶枝和基枝都能反映油菜品种的分枝角度习性。但顶枝角的遗传效应值(加性效应和显性效应)大于基枝角, 基枝角度因环境引起的变异大于顶枝角度, 说明基枝角度受到植株间互相交织和遮蔽的影响更大。因此, 在当前的种植密度下, 顶枝角度更适合作为分枝角度性状指标用于定量遗传分析。

由于顶、基枝角在BCP₁、BCP₂和F₂分离群体中都表现连续正态分布, 选用主基因+多基因模型进行遗传模型和效应分析是恰当的。本研究结果表明, 油菜顶枝角和基枝角均受1对加性-显性主基因+加性-显性-上位性多基因控制(D-0模型), 主基因加性效应使分枝角度增加, 而主基因显性效应使分枝角度减少。其中显性效应为负值与王俊生等^{[10, 11]}的显性效应对分枝角度有减效作用的结论相同。主基因遗传率较小, 多基因遗传率较大, 表明油菜分枝角度性状被主基因调控, 但主要受多基因控制, 并受环境因素的影响, 与前人研究^{[10, 11]}相比, 本研究更进一步获得主基因和微效多基因的遗传效应值, 为进一步利用高世代群体克隆相关基因奠定了基础。在选育株型紧凑(分枝角度< 30° )油菜品种的过程中需要采用联合选育, 即采用轮回选择或聚合的方法来累积增效基因, 以提高选择效率^[30]。同时在油菜杂种优势利用中, 需充分考虑亲本产量性状杂种优势, 同时选取分枝角度均较小的亲本配组, 才能获得杂交种的高产兼顾紧凑株型。

本研究利用6098B和Purler分枝角度差异显著的亲本构建6个世代遗传群体进行遗传分析, 并且分析样本足够大, 对于油菜分枝角度的鉴定采用数字图像处理分析方法, 从而大大提高结果的准确性^[22]; 利用主基因+多基因混合遗传模型分析油菜分枝角度的遗传, 从研究的油菜组合中检测出1对主基因的存在与效应, 同时还分析了主基因之外的微效基因的综合表现。下一步需要在不同种植密度环境下, 将分枝角度、分枝数、角果数以及单株产量等性状结合分子标记进行遗传定位分析, 从分子水平上阐述分枝角度的基因及其遗传调控网络, 并与产量等相关基因位点作元分析, 发掘密植环境下的与分枝角度及产量相关性状紧密连锁的分子标记, 实现油菜紧凑株型高产稳产的分子标记辅助育种。

4 结论

甘蓝型油菜上分枝角和下分枝角性状均受1对加性-显性主基因+加性-显性-上位性多基因控制(D-0模型), 显性效应值和加性效应值不同向, 表现为部分显性; 遗传率在3个分离世代中都表现为中等或者中等偏下、说明在其后代选育相对株型紧凑(分枝角度小)的高产育种品系中, 需要考虑兼顾主基因、多基因和环境因素的影响。

The authors have declared that no competing interests exist.

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2014

0.0

王汉中, 殷艳. 我国油料产业形势分析与发展对策建议. 中国油料作物学报, 2014, 36: 414-421

Wang H

, Yin

Analysis and strategy for oil crop industry in China.

Chin J Oil Crop Sci, 2014, 36: 414-421 (in Chinese with English abstract)

Currently, Chinese edible oils were highly dependent on import. After analyzing the demand, supply and trade of oil crops in the world as well as in China, we discussed the major problems in current development of Chinese oil crops industry. The strategies and suggestions were made to solve the problems related to policy making, technology, etc.

Oil Crops Reasearch Institute, CAAS, Key Laboratory of Biology and Genetic Improvement of Oil Crops, Ministry of Agriculture, Wuhan 430062, China

在当前我国食用植物油对外依存度居高不下的严峻形势下，本文通过对世界及我国油料产需及贸易形势进行分析，明确了当前我国油料产业发展存在的主要问题，针对存在的问题从政策、技术等方面提出了相应的对策建议。

... 近年来, 油料供给短缺的状态一直未得到有效解决, 甚至日趋紧张^[1] ...

2015

0.0

唐敏强, 程晓晖, 童超波, 刘越英, 赵传纪, 董彩华, 于景印, 马小艮, 黄军艳, 刘胜毅. 甘蓝型油菜株高性状的全基因组关联分析. 作物学报, 2015, 41: 1121-1126

Tang M

, Cheng X

, Tong C

, Liu Y

, Zhao C

, Dong C

, Yu J

, Ma X

, Huang J

, Liu S

Genome-wide association analysis of plant height in rapeseed (Brassica napus).

Acta Agron Sin, 2015, 41: 1121-1126 (in Chinese with English abstract)

Appropriate plant height is a trait related to lodging resistance and mechanical harvest, and ideal plant height is a basis of ideotype for high yield. The plant height of 371 rapeseed accessions collected from different growing regions in the world was measured in field experiments, all accessions were genotyped using a 60K SNP chip and 21856 informative SNPs were acquired, then the data of SNPs and plant height were used for genome-wide association analysis. Major results were as follows: 1) 4 significant SNPs related to plant height were detected, including Bn-A07-p3583161, Bn-scaff_22790_1-p1271170, Bn-scaff_20735_1-p42779, and Bn-scaff_18702_1-p589589, which located in A07, C01 and C02 chromosomes, and explained 11.33%, 11.75%, 12.31%, and 10.97% of phenotypic variation, respectively; 2) According to the reference genome annotation information, three candidate genes corresponding to the first three significant SNP were predicted, each gene was locates in 492.0, 9.5, and 69.5 kb away from the three SNP, respectively ; 3) The significant (two-tailed t-test) difference of plant height between means of two allelic groups of accessions corresponding to each significant SNP was 11.09, 15.12, 10.48, and 8.93 cm, respectively.

1 Oil Crops Research Institute, Chinese Academy of Agricultural Sciences / Key Laboratory of Biology and Genetic Improvement of Oil Crops, Ministry of Agriculture, Wuhan 430062, China?; 2 Hubei university / Hubei Collaborative Innovation Center for Green Transformation of Bio-Resources ,Wuhan 430062, China

适当的株高不仅是抗倒性和机械化收获的需要，而且是建立高产理想株型的需要。本研究测定来自世界油菜主产国的 371 份种质资源材料构成的自然群体的株高，利用 60K SNP 芯片对该群体进行基因型检测，获得了 21 856 个 SNP 标记，利用这些标记和株高数据进行全基因组关联分析结果表明： ( 1 ) 检测到 4 个与株高显著关联的 SNP ，即 Bn-A07-p3583161 、 Bn-scaff_22790_1-p1271170 、 Bn-scaff_20735_1-p42779 和 Bn-scaff_18702_1-p589589 ，分别分布于 A07 、 C01 和 C02 染色体；它们对表型变异的解释率分别为 11.33% 、 11.75% 、 12.31% 和 10.97% ； ( 2 ) 参考基因组信息，在上述前 3 个 SNP 附近的 492 、 9.5 和 69.5 kb 处分别找到一个与株高性状相关的候选基因； ( 3 ) 4 个显著标记 SNP 各自的等位基因间表型均值差异分别为 11.09 、 15.12 、 10.48 和 8.93 cm ，双尾 t 测验结果表明各等位基因组间差异显著。

... 株高、分枝部位高、分枝数和主花序有效长是油菜株型重要的组成部分, 相关的研究较多^{[2,3,4,5,6,7]}, 但关于油菜分枝角度的研究较少 ...

2015

0.0

王嘉, 荆凌云, 荐红举, 曲存民, 谌利, 李加纳, 刘列钊. 甘蓝型油菜株高、第一分枝高和分枝数的QTL检测及候选基因筛选. 作物学报, 2015, 41: 1027-1038

Wang

, Jing L

, Jian H

, Qu C

, Chen

, Li J

, Liu L

Quantitative trait loci mapping for plant height, the first branch height, and branch number and possible cand idate genes screening in Brassica napus L.

Acta Agron Sin, 2015, 41: 1027-1038 (in Chinese with English abstract)

Plant height, the first branch height and branch number are important agronomic traits in rapeseed. In our study, QTL mapping of plant height, the first branch height and branch number in Brassica napus was tested by using the high density SNP genetic map constructed from the high generation RIL population with the rapeseed 60K chip array. The reference SNP genetic map contains 2795 SNP markers, covering 1832.9 cM with an average distance of 0.66 cM in B. napus genome. Totally, 24 putative QTLs were identified for plant height, the first branch height and branch number by using the complex interval mapping. The phenotypic variation was explained by individual QTL ranged from 5.00% to 15.26% for 11 QTLs of plant height, from 5.04% to 12.99% for seven QTLs of the first branch height, and from 5.95% to 8.14% for six QTLs of branch number. We collected 156 genes associated with plant height, 10 genes associated with the first branch height and 148 genes associated with branch number in Arabidopsis thaliana and searched the homology region of the QTL confidence interval E-value ATGID1B/GID1B and WRI1 were found in QTL confidence interval on A07 and SLR/IAA14 and AXR2/IAA7 were found in QTL confidence interval on A08 in 2013 and 2014. The first branch height candidate gene PHT1;8 was found in QTLs q2013FBH-C05-1 and q2014FBH-C05-2 that had an overlapping confidence interval . Moreover, we found four genes associated with branch number that E-value from 0 to 3E–56 on A03 and C06, respectively.

College of Agronomy and Biotechnology, Southwest University, Chongqing 400715, China

株高、分枝数及第 1 分枝高是油菜重要的农艺性状。本研究利用甘蓝型油菜 GH06 和 P174 杂交 , F 2 通过单粒法连续自交至 F 11 构建重组自交系群体 , 利用油菜 60K 芯片对该群体进行基因分型 , 构建高密度遗传连锁图谱。结果表明 , 该图谱包含 2795 个 SNP 多态性标记位点 , 总长 1832.9 cM , 相邻标记间平均距离为 0.66 cM 。在此图谱基础上采用复合区间作图法 (CIM) , 检测到 3 个农艺性状的 24 个 QTL 。其中 11 个株高 QTL 分别位于 A01 、 A06 、 A07 、 A08 、 A10 和 C06 染色体 , 单个 QTL 解释 5.00%~15.26% 的表型变异 ; 7 个第 1 分枝高 QTL 分别位于 A06 、 C05 和 C06 染色体 , 单个 QTL 解释 5.04%~12.99% 的表型变异 ; 6 个分枝数 QTL 分别位于 A03 、 A07 、 C01 、 C04 和 C06 染色体 , 单个 QTL 解释 5.95%~8.14% 的表型变异。将 156 个拟南芥株高相关基因、 10 个拟南芥第 1 分枝高相关基因和 148 个拟南芥分枝数相关基因与 QTL 对应置信区间序列进行同源比较分析 (E , 分别找出了 2 0 个株高候选基因、 3 个第 1 分枝高候选基因以及 12 个分枝数候选基因。 2 个环境中在 A07 染色体上重复检测到的 QTL 置信区间检测到与株高相关的候选基因 ATGID1B/GID1B 和 WRI1 , A08 染色体上重复检测到的 QTL 置信区间检测到 SLR/IAA14 和 AXR2/IAA7 2 个与株高相关的候选基因。在具有部分置信区间重叠的 q2013FBH-C05-1 和 q2014FBH-C05-2 区间均检测到第 1 分枝高候选基因 PHT1;8 , 在 A03 和 C06 染色体上的 QTL 置信区间内 , 分别检测到 4 个分枝数候选基因 , 匹配 E 值介于 0~3E–56 之间。

... 株高、分枝部位高、分枝数和主花序有效长是油菜株型重要的组成部分, 相关的研究较多^{[2,3,4,5,6,7]}, 但关于油菜分枝角度的研究较少 ...

2014

0.0

张凤启, 刘越英, 程晓辉, 童超波, 董彩华, 唐敏强, 黄军艳, 刘胜毅. 利用高密度SNP标记定位甘蓝型油菜株高QTL. 中国油料作物学报, 2014, 36: 695-700

Zhang F

, Liu Y

, Cheng X

, Tong C

, Dong C

, Tang M

, Huang J

, Liu S

QTL mapping of plant heigh using high desity SNP markers in Brassica napus.

Chin J Oil Crop Sci, 2014, 36: 695-700 (in Chinese with English abstract)

Plant height is an important factor affecting the yield of oilseed rape. In order to find the markers linked with plant height and to increase the efficiency of plant height improvement by molecular marker-assisted selection, identification of quantitative trait loci (QTL) with single environment and multi-environment detection methods for plant height was conducted in a recombinant inbred lines (RILs) population derived from a cross between 888-5 (a dwarf line) and M083 (a tall line) by employing the newly developed Brassica napus 60K SNP array. The results indicated that a total of 27 putative QTLs for plant height in the 4 environments were detected on the chromosomes A2, A4, A5, A6, A9, A10, C3 and C7, each accounting for 0.70% - 26.10% of the phenotypic variation. Among them, 6 QTLs showed interaction with the environment, and their effect values in the 4 environments were different, but their directions were consistent. QTL qPHE2-4 which had the largest effect in the 27 QTLs was detected in 2 environments, explaining 17.96%-26.10% of the phenotypic variation, and 2 linkage SNP markers in this QTL region had less than 0.2 cM distance from the peak of the QTL．These QTLs provided useful information for improving the plant height in oilseed rape breeding.

Oil Crops Research Institute, Chinese Academy of Agricultural Sciences, Key Laboratory of Biology and Genetic Improvement of Oil Crops, Ministry of Agriculture, Wuhan 430062, China

为了提高甘蓝型油菜株高的分子标记辅助育种效率，以888-5（矮秆）×M083（高秆）构建的重组自交系为材料，通过利用新开发的油菜60K SNP芯片对群体进行SNP标记分型，并结合单环境和多环境2种QTL检测方法对4个环境下株高进行了QTL定位及其与环境互作分析。结果表明：2种QTL定位方法，共检测出27个油菜株高QTL，分布于8个连锁群（A2、A4、A5、A6、A9、A10、C3和C7），单个QTL可解释的表型变异为0.70%~26.10%。其中6个QTL与环境存在互作效应，在4个环境下效应大小不一，但效应方向表现一致。主效QTL qPH2-4在2个环境下重复检测到，对表型变异解释为17.96%~26.10%，而且有2个SNP标记距离该QTL的最大峰值小于0.2cM。这些QTL或标记为油菜株高的遗传改良提供了有用的遗传信息。

... 株高、分枝部位高、分枝数和主花序有效长是油菜株型重要的组成部分, 相关的研究较多^{[2,3,4,5,6,7]}, 但关于油菜分枝角度的研究较少 ...

2012

0.0

谢田田, 陈玉波, 黄吉祥, 张尧锋, 徐爱遐, 陈飞, 倪西源, 赵坚义. 甘蓝型油菜不同发育时期株高QTL的动态分析. 作物学报, 2012, 38: 1802-1809

Xie T

, Chen Y

, Huang J

, Zhang Y

, Xu A

, Chen

, Ni X

, Zhao J

Dynamic analysis of QTL for plant height of rapeseed at different developmental stages.

Acta Agron Sin, 2012, 38: 1802-1809 (in Chinese with English abstract)

The main objective of this study was to explore the genetic control mechanism of plant height in rapeseed and to reveal its dynamic gene expression in specific growing period. Taking advantage of the updated SG map, we determined phenotypic data of plant height from 282 SG-DH lines in three environments and eight development stages, and analyzed both static and dynamic quantitative trait locus (QTL) with unconditional and conditional mapping approach. The results showed that when alleles from Gaoyou existed in two major QTLs PHA3 and PHC6 synchronously, the plant height could decrease about 20 cm; when alleles of plant height came from Sollux in PHA9 and PHC1, and Gaoyou in PHA1, PHA3 and PHC6 , the plantsfell 40 cm in height. The mechanism of QTL for plant height was less affected by environment, but various QTLs showed difference in gene expression patterns: plant height was dominated by alleles from one of parents or from two parents by turns at different stages. In general, the QTL for plant height could be detected from the middle to late stages, while the gene expression appeared shortly and only during the most active growing period, in congruent with the rule of “trait appearance goes after genetic expression”.

1 College of Chemistry and Life Sciences, Zhejiang Normal University, Jinhua 321000, China; 2 Institute of Crop and Nuclear Technology Utilization, Zhejiang Academy of Agricultural Sciences, Hangzhou 310021, China; 3 College of Life and Environmental Sciences, Hangzhou Normal University, Hangzhou 310036, China; 4 College of Agronomy, Northwest Agricultural and forestry University, Yangling 712100, China

利用新版SG遗传图谱和282个SG-DH株系在中国西安、杭州和德国哥廷根3个生长环境下8个发育时期测定的株高数据，运用WinQTLCart2.5复合区间作图法以及结合条件遗传分析方法对其进行静态和动态QTL分析。结果显示, 来自品种Gaoyou等位基因在 PHA3 和 PHC6 两个QTL上同时存在时，可降低株高约20 cm；而当植株整合来自冬性品种Sollux的 PHA9 、 PHC1 和来自半冬性品种Gaoyou的 PHA1 、 PHA3 、 PHC6 时，株高可相应下降40 cm；环境对株高QTL的作用机理影响不大，但不同QTL的基因表达模式不同，存在来自双亲之一的等位基因控制株高和双亲等位基因在不同生长时期交替控制株高两种情况；通常株高QTL在中后期才能被检测到，但基因多在生长最为旺盛的短时期内表达，符合基因表达在先，性状表现在后的规律。解析株高性状在不同发育时期基因的累加效应和特定时段内的净表达效应，对克隆油菜株高基因和指导生产实践都将提供富有价值的科学信息和理论依据。

... 株高、分枝部位高、分枝数和主花序有效长是油菜株型重要的组成部分, 相关的研究较多^{[2,3,4,5,6,7]}, 但关于油菜分枝角度的研究较少 ...

2014

0.0

... 株高、分枝部位高、分枝数和主花序有效长是油菜株型重要的组成部分, 相关的研究较多^{[2,3,4,5,6,7]}, 但关于油菜分枝角度的研究较少 ...

2015

0.0

... 株高、分枝部位高、分枝数和主花序有效长是油菜株型重要的组成部分, 相关的研究较多^{[2,3,4,5,6,7]}, 但关于油菜分枝角度的研究较少 ...

2006

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王俊生, 李少钦, 张耀文. 不同栽培密度对紧凑型油菜产量和主要性状的影响. 耕作与栽培, 2006, (3): 25-26

Wang J

, Li S

, Zhang Y

Effects of different planting density on yield and main characters of compact in rapeseed.

Gengzuo yu Zaipei, 2006, (3): 25-26 (in Chinese)

研究以3个紧凑株型和1个普通株型油菜品种"秦优7号"为材料,研究了它们在不同密度下的个体与群体的产量结构,结果表明,在密度为210000株/hm2时,紧凑型油菜品种的个体与群体的协调性最佳,可以获得该类品种的最高产量;"秦优7号"的产量在150000株/hm2的密度下最高,在210000株/hm2密度下,产量有不显著减产;紧凑型油菜品种达到最高产量时的密度高于"秦优7号".随密度从150000/hm2增加到270000株/hm2时,紧凑型油菜品种和"秦优7号"品种的单株角果数都减少,但紧凑型品种的角果数缓慢较少,而"秦优7号"较少幅度较大;两类品种千粒重的增加和角果粒数的减少均较平缓.在较高密度下,个体与群体协调性好,群体角果数多,这是紧凑型油菜在较高密度下能获得较高产量的主要原因.

... 前人的研究基本局限于分枝角度与产量性状的相关性^[8,9], 而对油菜分枝角度基因的效应大小及其作用形式很少报道 ...

2013

0.0

... 前人的研究基本局限于分枝角度与产量性状的相关性^[8,9], 而对油菜分枝角度基因的效应大小及其作用形式很少报道 ...

... 主基因+多基因混合遗传模型是由盖钧镒^[15]提出的数量性状遗传分析方法, 在多种植物的遗传研究中得到应用^[16], 在油菜性状的遗传研究中也广泛应用^{[9,17,18,19,20,21]} ...

2000

0.0

... 张文学等^[10]首次研究了甘蓝型油菜紧凑型和松散型杂种一代分枝性状的遗传效应, 结果表明紧凑型与松散型正反交F₁代分枝角度无差异, 表现双亲平均值, 说明细胞质效应不影响分枝角度 ...

... 其中显性效应为负值与王俊生等^[10,11]的显性效应对分枝角度有减效作用的结论相同 ...

... 主基因遗传率较小, 多基因遗传率较大, 表明油菜分枝角度性状被主基因调控, 但主要受多基因控制, 并受环境因素的影响, 与前人研究^[10,11]相比, 本研究更进一步获得主基因和微效多基因的遗传效应值, 为进一步利用高世代群体克隆相关基因奠定了基础 ...

2005

0.0

... 王俊生等^[11]利用6个油菜品种构建30个完全双列杂交组合, 对分枝角度、一次分枝数、分枝长等10多个株型性状进行了遗传效应及其与产量性状的相关性分析, 结果表明分枝角度狭义遗传率较高, 显性效应对分枝角度有减效作用, 并与单株产量特别是单株角果数存在显著的遗传负相关 ...

... 王俊生等^[11]的研究发现分枝角度与单株产量特别是单株角果数存在显著的遗传负相关, 说明分枝角度小的植株在生长中后期的物质积累中占有优势, 由于中下部光合作用受到的影响较小, 促进了有效分枝数和有效角果数的提高 ...

... 其中显性效应为负值与王俊生等^[10,11]的显性效应对分枝角度有减效作用的结论相同 ...

2012

0.0

... 近几年, 为构建机械化密植条件下的油菜理想株型, 分枝角度的研究也逐渐得到重视^[12,13], Liu等^[14]利用油菜自然群体, 定位了6个与分枝角度显著关联的基因组区域, 并获得其中4个区域内候选基因 ...

2014

0.0

2016

0.0

2003

0.0

... 利用植物数量性状主基因+多基因混合遗传模型分析方法^[15], 并结合软件^[23]数据处理进行遗传模型分析(分析软件由南京农业大学章元明教授提供) ...

2008

0.0

李余生, 朱镇, 张亚东, 赵凌, 王才林. 水稻稻曲病抗性的主基因+多基因混合遗传模型分析. 作物学报, 2008, 34: 1728-1733

Li Y

, Zhu

, Zhang Y

, Zhao

, Wang C

Genetic analysis of rice false smut resistance using major gene plus polygene mixed genetic model.

Acta Agron Sin, 2008, 34: 1728-1733 (in Chinese with English abstract)

Rice false smut, a fungus disease caused by Ustilaginoidea virens (Cooke) Takahashi, has become one of the most important rice diseases in China. It is of practical guiding significance to probe resistant mechanism for programming rice breeding for resistance to false smut. The present study was aimed to estimate the genetic model and genetic parameters of rice false smut resistance using the artificial inoculation technique to induce the rice false smut efficiently, and the disease rate index scored based on percentage of the infected grains. The experiment was conducted using 157 recombinant inbred lines (RILs) population of F10 derived from a cross of IR28 ( indica ) ´ Daguandao ( japonica ), two parents exhibited resistant and susceptible to rice false smut respectively. The results showed that the rice false smut resistance was controlled by two major genes plus polygene mixed ge-netic model (model E-1-3), and two major genes had equivalent additive effect of 11.41, their heritability was about 76.67%, while the heritability of polygene was about 22.86%. The resistance to rice false smut had significant major gene effect, and the results implied that not only major gene’s effect but also polygene’s effect should be considered in breeding for resistance to rice false smut.

Institute of Food Crops, Jiangsu Academy of Agricultural Science / Jiangsu High Quality Rice Research and Development Center, Nanjing 210014, Jiangsu, China

稻曲病是由稻曲病菌引起的水稻真菌病害, 目前已上升为我国水稻主要病害之一, 探明水稻稻曲病的抗性遗传机制对制定抗病育种策略具有现实指导意义。本研究以病情指数为指标, 利用高效引发稻曲病人工接种方法对来自抗病亲本IR28(籼稻)与感病亲本大关稻(粳稻)组合的亲本及其衍生的F 10 代重组自交系群体(RIL)进行抗病性评价, 采用数量性状的主基因+多基因混合遗传模型对抗性的遗传模型进行判别与遗传参数的估计。结果表明, 水稻稻曲病抗性的遗传除受主基因控制外还受多基因的影响, 符合E-1-3遗传模型, 即2对主基因+多基因混合遗传模型, 2对主基因间表现为等加性作用, 加性效应为11.41; 主基因遗传率为76.67%, 多基因遗传率为22.86%, 抗性遗传存在明显的主基因效应。提示抗稻曲病育种时不仅要考虑主基因对抗病性的作用, 也不能忽视基因对抗性的影响。

2014

0.0

周清元, 崔翠, 阴涛, 陈东亮, 张正圣, 李加纳. 甘蓝型油菜角果长度的主基因+多基因混合遗传模型. 作物学报, 2014, 40: 1493-1500

Zhou Q

, Cui

, Yin

, Chen D

, Zhang Z

, Li J

Genetic analysis of silique length using mixture model of major gene plus polygene in Brassica napus L.

Acta Agron Sin, 2014, 40: 1493-1500 (in Chinese with English abstract)

Silique is one of major components for rapeseed yield. Inheritance of silique body length (SBL), valid silique length (VSL) and beak length (BL) in a cross of variety Zhongshuang 11 with long silique (P 1 ) and line 10D130 with short silique (P 2 ) was investigated by the mixed major gene plus polygene inheritance model. The results showed that SBL, VSL and BL in the populations of F 2 , BCP 1 , and BCP 2 were controlled by the major gene and polygenes. The SBL were dominated by two major genes with additive-dominance-epistasis effects plus polygenes with additive-dominance-epistasis effects (E-0 model). The heritability values of the major genes of SBL in BCP 1 , BCP 2 , and F 2 were estimated as 51.10%, 74.23%, and 66.93%, respectively, and the heritability values of the polygene were 29.16%, 17.11%, and 23.96% , respectively. The additive effects of two major genes of SBL were 1.75 and –0.06, and the dominant effects of two major genes were –0.59 and –0.86, respectively. The valid silique length was controlled by two major genes with additive-dominance-epistasis effects plus polygenes with additive-dominance effects (E-1 model). Heritability values of the major genes for SL in BCP 1 , BCP 2 , and F 2 generations were estimated as 47.63%, 68.51%, and 79.45%, respectively, and the heritability values of the polygene were 29.40%, 20.89%, and 12.47%, respectively. The additive effects of two major genes were equal (0.34) to there of the cross, but the dominant effects of the two major genes were –0.81 and –0.47, respectively. The beak length was dominated by two major gene with additive effects plus polygenes with additive- dominance effects (E-3 model). Heritability values of the major genes of BL in the cross were 33.71%, 72.75%, and 52.25%, respectively, and the heritability values of the polygene were 40.08%, 5.37%, and 27.60%, respectively. The additive effects of two major genes were 0.20 and –0.20, respectively.

College of Agronomy and Biotechnology, Southwest University / Engineering Research Center of South Upland Agriculture, Ministry of Education / Chongqing Engineering Research Center for Rapeseed, Southwest University, Chongqing 400716, China?

角果是油菜产量构成要素中重要的组成部分。本文以长角果品种中双11和短角果材料10D130为亲本配制杂交组合，采用主基因+多基因混合遗传模型分析方法对该组合6世代遗传群体(P 1 、P 2 、F 1 、BCP 1 、BCP 2 和F 2 )的果身长、角果长和果喙长进行遗传分析。结果表明，该组合的3个角果性状均呈连续分布，其中，果身长最适遗传模型为E-0 (2对加性-显性-上位性主基因+加性-显性-上位性多基因模型)，2对主基因加性效应值分别是1.75和–0.06，显性效应值分别是–0.59和–0.86，主基因遗传率在BCP 1 、BCP 2 和F 2 中分别是51.10%、74.23%和66.93%，多基因遗传率分别为29.16%、17.11%和23.96%。角果长的最适遗传模型为E-1 (2对加性-显性-上位性主基因+加性-显性多基因模型)，其中，第1对主基因加性效应为0.34，显性效应为–0.81，第2对主基因加性效应为0.34，显性效应为–0.47，主基因遗传率在BCP 1 、BCP 2 和F 2 中分别是47.63%、68.51%和79.45%，多基因遗传率分别为29.40%、20.89%和12.47%。果喙长的最适遗传模型为E-3模型(2对加性主基因+加-显多基因遗传模型)，2对主基因加性效应值分别是0.2和–0.2，主基因遗传率在BCP 1 、BCP 2 和F 2 中分别是33.71%、72.75%和52.25%，多基因遗传率分别为40.08%、5.37%和27.60%。

2014

0.0

金岩, 吕艳艳, 付三雄, 戚存扣. 甘蓝型油菜苗期耐淹性状主基因+多基因遗传分析. 作物学报, 2014, 40: 1964-1972

Jin

, Lü Y

, Fu S

, Qi C

Inheritance of major gene plus polygene of water-logging tolerance in Brassica napus L.

Acta Agron Sin, 2014, 40: 1964-1972 (in Chinese with English abstract)

The middle and lower reaches of Yangtze River is a main producing region of canola ( Brassica napus L.) in China. However, canola in this region is subjected to water-logging during planting period to reduce yield. It is of importance to study the inheritance of water-logging tolerance for canola. In this paper a family lines population of six generations of P 1 , F 1 , P 2 , B 1:2 , B 2:2 , F 2:3 derived from the cross of WR-4 (resistant)×WR-5 (non-resistant) was used to analyse genetic segregation by applying major gene plus polygene mixed inheritance model. The seedling mortality was recorded on the 7th day after logging-removing for plant recovery following six days full-submergence treatment of the seedlings in 2012 and 2013. The results showed that the seedling mortality was respectively fitted the genetic model of E-0 and B-3 i.e. two pairs of additive- dominant-epistatic major gene plus additive-dominant-epistatic polygene model and two pairs of additive major genes model. This result confirms that water-logging tolerance of seedling in this cross is controlled mainly by two major genes which expressed in the mode of additive-dominant-epistatic effects. While dominant effects expressed (2012) it gave a higher value of | h a |=0.3475, | h b |=0.0069 than the additive effect of the major genes which was | d a |=| d b |=0.0036. In the populations of B 1:2 , B 2:2 , and F 2:3 , h 2 mg was 36.25%, 61.40%, and 61.84% respectively with an average of 53.16% in 2012, and 8.30%, 30.48%, and 43.13% respectively with an average of 27.30% in 2013 . Variance from environment effects was 59.77% of the total phenotypic variance on an average in two years. A conclusion could be made that water-logging tolerance of seedling in B. napus is controlled by two major genes but heavily affected by environment. Since a higher value of h 2 mg was detected in F 2:3 populations, selection in early generations might be an effective way for waterlogging tolerance breeding in B. napus .

1 College of Agriculture, Nanjing Agricultural University, Nanjing 210095, China; 2 Institute of Industrial Crops, Jiangsu Academy of Agricultural Sciences, Nanjing 210014, China

长江中下游是中国油菜主产区，该地区油菜播栽期间雨水多，易产生湿害，造成产量下降。所以研究甘蓝型油菜苗期耐淹性的遗传规律，对选育耐淹性强油菜新品种，提高油菜产量意义重大。本文应用甘蓝型油菜品种 WR-4 ( 耐淹 ) 和 WR-5 ( 不耐淹 ) 杂交后代衍生的 6 个世代 ( P 1 、 F 1 、 P 2 、 B 1:2 、 B 2:2 、 F 2:3 ) 群体为材料，全淹 6 d 后去水恢复生长，去水后第 7 天调查死苗率，以此为耐淹性指标，于 2012 和 2013 年对 6 个世代群体家系进行耐淹性鉴定。应用植物数量性状主基因 + 多基因混合遗传模型多世代联合分析方法对耐淹性进行遗传分析。结果表明， 2 个年度该家系群体苗期耐淹性的最适遗传模型分别是 E-0 和 B-3 ，即 2 对加性 - 显性 - 上位性主基因 + 加性 - 显性 - 上位性多基因和 2 对加性主基因模型。由此可见，该家系群体甘蓝型油菜苗期耐淹性主要受 2 对主基因控制，主基因存在加性、显性和上位性效应。当有显性效应存在时 ( 2012 年 ) ，主基因显性效应值 | h a |=0.3475 ， | h b |=0.0069 ，大于主基因加性效应值 | d a |=| d b |=0.0036 。 B 1:2 、 B 2:2 和 F 2:3 群体的主基因遗传率 ( h 2 mg ) ， 2012 年分别为 36.25% 、 61.40% 和 61.84% ，平均为 53.16% ； 2013 年分别为 8.30% 、 30.48% 和 43.13% ；平均为 27.30% 。 2 年平均，环境变异占表型变异的 59.77% 。上述结果表明，甘蓝型油菜苗期耐淹性受 2 对主基因型控制，但环境对耐淹性状的表型影响较大。 F 2:3 家系群体苗期耐淹性遗传率较高，因此育种上可在早期世代对耐淹性状进行选择。

2009

0.0

田露申, 牛应泽, 余青青, 郭世星, 柳丽. 甘蓝型油菜白花性状的主基因+多基因遗传分析. 中国农业科学, 2009, 42: 3987-3995

Tian L

, Niu Y

, Yu Q

, Guo S

, Liu

Genetic analysis of white flower color with mixed model of major gene plus polygene in Brassica napus L.

Sci Agric Sin, 2009, 42: 3987-3995 (in Chinese with English abstract)

【Objective】 White flower color in Brassica napus L. was observed quantitatively and the genetic characteristics were analyzed in order to provide a theoretical basis for its applications in rapeseed breeding. 【Method】 Six generations (P1, P2, F1, B1, B2 and F2) of two crosses (HW243×HZ21-1, HW243×Zhongyou 821) were used as genetic populations and fresh petals were taken and scanned with a computer scanner. The characteristic values（CIE RGB system）of petal color were obtained with a self-designed computer software. The genetic characteristics of the white flower color were analyzed with a mixed model of major genes plus polygenes. 【Result】 It was found that the white flower color in Brassica napus L. appeared to be a quantitative trait and its inheritance fitted to a mixed genetic model of two major genes with additive-dominant-epistatic effects plus polygenes with additive-dominant-epistatic effects (the E-0 model). The additive, dominant and epistatic effects of major genes were all important. Heritabilities of the major genes were estimated to be 96.94% and 95.83% in F2, 54.58% and 49.57% in B1, 98.14% and 97.67% in B2. Heritabilities of the polygenes were estimated to be 3.93% and 2.47% in F2, 35.64% and 46.9% in B1, 0.98% and 2.06% in B2. 【Conclusion】 The white flower color in B. napus L. appeared to be a quantitative trait. Its inheritance fitted to a genetic model of two major genes with additive, dominant and epistatic effects plus polygenes with additive, dominant and epistatic effects. The effects of major genes were dominant and the effects of polygenes were relatively small. Heritabilities of the major genes were high, and environmental influence was small.

（四川农业大学油菜研究中心/教育部作物基因资源与遗传改良重点实验室）

【目的】对甘蓝型油菜白花性状进行量化观察,研究其数量遗传特性,为育种利用提供理论依据。【方法】利用扫描仪和颜色提取软件对油菜新鲜花瓣进行处理,获得花瓣颜色特征值（CIE RGB值）,选择能反映花瓣颜色差异的B值,应用植物数量性状主基因+多基因混合遗传模型多世代联合分析方法,对甘蓝型油菜杂交组合（HW243×HZ21-1和HW243×中油821）的P1、P2、F1、B1、B2和F2世代群体进行分析。【结果】甘蓝型油菜白花性状表现为一数量性状,其遗传符合两对加性-显性-上位性主基因+加性-显性-上位性多基因遗传模型,以主基因作用为主,多基因的作用相对较小。两对主基因的加性、显性和上位性效应均具有较大的作用。在F2群体中主基因的遗传率为96.94%和95.83%,多基因遗传率为3.93%和2.47%;在B1群体中主基因的遗传率为54.58%和49.57%,多基因遗传率分别为35.64%和46.9%;在B2群体中主基因的遗传率为98.14%和97.67%,多基因遗传率分别为0.98%和2.06%。【结论】甘蓝型油菜白花性状具有数量性状的遗传特性,其遗传符合两对加性-显性-上位性主基因+加性-显性-上位性多基因遗传模型,以主基因效应为主,多基因效应相对较小。主基因的遗传力较高,受环境影响较小。

2008

0.0

顾慧, 戚存扣. 甘蓝型油菜(Brassica napus L. )抗倒伏性状的主基因+多基因遗传分析. 作物学报, 2008, 34: 376-381

, Qi C

Genetic analysis of lodging resistance with mixed model of major gene plus polygene in Brassica napus L.

Acta Agron Sin, 2008, 34: 376-381 (in Chinese with English abstract)

Rapeseed is one of the most important oil crops in the world. Lodging damage results in not only lower seed yield and oil quality, but also easier infection to the infection of Sclerotinia and more difficulties for mechanized harvest. So it is of a great significance to study lodging resistance in B. napus L. However, lodging is a complex trait associated with several factors. It is difficult to evaluate in field. In this paper, inheritance of lodging resistance was analyzed applying the joint segregation analysis of a mixed genetic model of major gene plus polygene in six generations (P1, P2, F1, B1, B2, and F2 ) derived from two crosses of Zheping 1 (lodging resistance) × 04Pb11 (lodging) (Cross I) and Ning 1243 (lodging resistance) × 04Pb11 (Cross II) in B. napus L. Resistant pressure per plant (RPPP) was measured at flowering time by Digital Force Gauge, which was used to assess the ability of lodging resistance, and the correlation between RPPP and lodging scale was scored at mature time (r = -0.38, P≤0.01). The results showed that lodging resistance was dominated by two major genes with additive-dominance-epistatic effects plus polygenes with additive-dominance-epistatic effects (E-0 model) in Cross I, and two major genes with additive-dominance-epistatic effects plus polygenes with additive-dominance effects (E-1 model) in Cross II. The additive effects of two major genes in two crosses were more powerful than the dominant one (︱h/d︱＜1). On average, Hereditability of major genes (hmg2) was 54.71% in F2 population, and that of polygenes (hpg2) only detected in B1 population was 10.56%. It indicated that the lodging resistance in B. napus L. was dominated by major genes. In addition, the environmental variance and the genetic variance accounted for 46.57% and 53.43% of the phenotypic variance in each population respectively, indicating that the environmental factors have a great effect on lodging in B. napus L.. However, higher hereditability of major genes can be favorable to an efficient selection in early generation of lodging resistance breeding.

1 Agronomy College of Nanjing Agricultural University, Nanjing 210095, Jiangsu; 2 Institute of Industrial Crops, Jiangsu Academy of Agricultural Sciences, Nanjing 210014, Jiangsu, China

应用植物数量性状主基因+多基因混合遗传模型多世代联合分析方法，研究了甘蓝型油菜浙平1号×04Pb11(I)和宁1243×04Pb11(II)的P1、P2、F1、B1、B2和F2 6个世代初花期单株抗压力的遗传。结果表明：抗倒伏性状的遗传在组合I受2对加性-显性-上位性主基因+加性-显性-上位性多基因控制，在组合II受2对加性-显性-上位性主基因+加性-显性多基因控制；2个组合中的2对主基因都以加性效应为主，都表现抗倒对易倒部分显性或完全显性，2对主基因间存在明显的基因互作效应；2个组合中，F2群体主基因遗传率平均为54.71%，而多基因遗传率只在B1群体中检测到，平均为10.56%，表明2个组合的抗倒伏性状是以主基因遗传为主，应在早期世代进行选择；2个组合各群体中，遗传变异平均占表型变异的53.43%，而环境变异平均占表型变异的46.57%，表明环境对油菜抗倒伏性状的影响比较大。

2014

0.0

王会, 桑世飞, 梅德圣, 李云昌, 刘佳, 付丽, 王军, 陈玉峰, 胡琼. 甘蓝型油菜DH群体抗裂角性遗传分析. 中国油料作物学报, 2014, 36: 437-449

Wang

, Sang S

, Mei D

, Li Y

, Liu

, Fu

, Wang

, Chen Y

, Hu

Inheritance analysis of pod shatter resistance using a DH population in Brassica napus L.

Chin J Oil Crop Sci, 2014, 36: 437-449 (in Chinese with English abstract)

Improvement of pod shatter resistance is the basic target for rapeseed breeding suitable for mechanized harvesting. To better understand the inheritance of pod shatter resistance in Brassica napus, a doubled haploid (DH) population was used in the study. Two parents of the population had contrasting pod shatter resistance levels. Pod shatter resistance index was determined using random impact test method in two years, and the inheritance was analyzed by major gene plus polygene genetic model for plant quantitative traits as well as the analysis of skewness and kurtosis. Results showed that the pod shatter resistance was controlled by 3 genes with major effects and 5-7 polygenes with minor effects. There were major genes additive effects and complementary gene interactions. The heritability of major genes was greater than 85.00% in both years, indicating pod shatter resistance was mainly controlled by major genes and less influenced by environments. Therefore, in breeding process for pod shatter resistance of rapeseed, pyramiding major genes from lines containing different major genes by hybridization and selection in early generation should be considered, and the influence of environmental factors can be ignored.

1. Oil Crops Research Institute of Chinese Academy of Agricultural Sciences / Key Laboratory for Biological Sciences and Genetic Improvement of Oil Crops, Ministry of Agriculture, Wuhan 430062, China 2. Graduate School of Chinese Academy of Agricultural Sciences, Beijing 100081, China 3. Oil Crops Research Institute of Guizhou Province, Guiyang 550006, China

提高抗裂角性是培育适合机械化收获油菜品种的基本要求。为了明确油菜抗裂角性的遗传规律，本研究利用抗裂角性差异显著的两个甘蓝型油菜品系配制杂交组合，通过小孢子培养构建了一个双单倍体（doubled haploid, DH）群体。采用随机碰撞法对该群体进行连续两年的抗裂角性鉴定，并利用植物数量性状的主基因+多基因混合遗传模型及偏度和峰度分析对抗裂角性进行遗传分析。结果表明，甘蓝型油菜的抗裂角性受3对主基因+多基因控制，涉及的基因数目为8～10对，存在基因的加性效应及基因间的互补作用;两年试验中抗裂角性的主基因遗传率均大于85.00%，主基因遗传率较高，表明该性状主要受主基因控制，受其它微效多基因及环境的影响有限。因此，在油菜抗裂角性遗传改良中，应通过杂交聚合不同抗性等级材料中的主效抗裂角基因，在早期世代加强对抗性的选择，并可以忽略环境因素的影响。

2015

0.0

汪文祥, 胡琼, 梅德圣, 李云昌, 王会, 王军, 付丽, 刘佳. 基于图像处理的油菜分枝及角果着生角度测量方法. 中国油料作物学报, 2015, 37: 566-570

Wang W

, Hu

, Mei D

, Li Y

, Wang

, Fu

, Liu

Evaluation of branch and pod angle measurement based on digital images from Brassica napus L.

Chin J Oil Crop Sci, 2015, 37: 566-570 (in Chinese with English abstract)

Branch and pod angles are important plant architecture traits, which are closely related to ideal plant type, yield, high-density cultivation and mechanical harvesting. For simple and accurate collection of these angles, digital image method was used to set up an angle measurement method for rapeseed. According to their branch angles, 22 materials were measured and grouped into 3 typical types: compact ( 40º, 11) and intermediate (30º-40º, 8). According to pod angles, the materials were grouped into 2 types: horizontal (close to 90º, 3) and inclined (

1.Oil Crops Research Institute of Chinese Academy of Agricultural Sciences，Key Laboratory of Biology and Genetic Improvement of Oil Crops of the Ministry of Agriculture，Wuhan 430062，China; 2. Oil Crops Research Institute of Guizhou Province，Guiyang 550006，China

以数字图像处理技术为基础，建立了一套油菜株型角度测量方法，并根据分枝角度将22份材料分为紧凑型（小于30º）3份，中间型（30～40º）8份和松散型（大于40º）11份；根据角果着生状态将22份材料分为平生型（趋近于90º）3份和斜生型（小于80º）19份。对其中5份材料进行数字图像处理测量值（x）和传统的手工方法测量值（y）比较，经显著性t测验和方差分析，测定结果符合曲线为y=1.019 1x-1.363 2，相关系数为0.991 9。可见，这一套数字图像技术具有较高的重复性、精确性，更具有较好的可操作性，可为油菜分枝角度和角果着生角度的遗传研究提供技术支撑。

... 3 分枝角度鉴定方法2015年6月在室内采用数字图像处理法^[22]鉴定6世代群体分枝角度: 油菜成熟后, 剪取连有油菜上部第一分枝(顶枝)和基部第一分枝(基枝)的茎段, 采集图像 ...

... 本研究利用6098B和Purler分枝角度差异显著的亲本构建6个世代遗传群体进行遗传分析, 并且分析样本足够大, 对于油菜分枝角度的鉴定采用数字图像处理分析方法, 从而大大提高结果的准确性^[22] ...

2013

0.0

2000

0.0

... 密植除了增加植株的叶面积指数和光能利用效率外^[24], 还能提高油菜氮肥利用效率^[25], 促进氮素向籽粒转移^[26], 起到增产效果 ...

2009

0.0

马霓, 张春雷, 李俊, 李光明. 种植密度对直播油菜结实期源库关系及产量的调节. 中国油料作物学报, 2009, 31: 180-184

, Zhang C

, Li

, Li G

Regulation of planting density on source-sink relationship and yield at seed-set stage of rapeseed (Brassica napus L. ).

Chin J Oil Crop Sci, 2009, 31: 180-184 (in Chinese with English abstract)

对6种种植密度下直播甘蓝型油菜结实期的源库关系及产量进行了研究，结果表明：角果长、宽分别在开花后19d和25d左右达到最大值，角果大小有随密度增加而减少的趋势；单位角果皮面积承担的籽粒数（SNPA）及单位角果皮面积的籽粒重（PPA）都随密度的增大呈先增后减的趋势；每角粒数和角果长度或角果面积都符合二次曲线的关系；随着种植密度的增加，单株株高、茎粗、一次有效分枝数、单株角果数、每果粒数和千粒重均下降，但群体单位面积角果数、角果面积增大，产量增高。密度继续增加，群体单位面积角果数和角果面积减少，产量下降。

... 密植除了增加植株的叶面积指数和光能利用效率外^[24], 还能提高油菜氮肥利用效率^[25], 促进氮素向籽粒转移^[26], 起到增产效果 ...

2013

0.0

朱珊, 李银水, 余常兵, 谢立华, 胡小加, 张树杰, 廖星, 廖祥生, 车志. 密度和氮肥用量对油菜产量及氮肥利用率的影响. 中国油料作物学报, 2013, 35: 179-184

Zhu

, Li Y

, Yu C

, Xie L

, Hu X

, Zhang S

, Liao

, Liao X

, Che

Effects of planting density and nitrogen application rate on rapeseed yield and nitrogen efficiency.

Chin J Oil Crop Sci, 2013, 35: 178-184 (in Chinese with English abstract)

For rapeseed mechanization production, effects of rape density and nitrogen (N) application rate on growth and N use efficiency were investigated under field condition. Two planting density (15?104 and 45?104 plant per hectare) and four N rates (0, 90, 180 and 270kg per hectare) were applied to study the interaction between N and density on agronomic characteristics, yield, N use efficiency and N fertilizer requirements for Zhongshuang 11. Results showed that plant height, branch number, effective siliques per plant and seeds per pod significantly decreased at higher density, but branch height and 1 000-seeds weight increased significantly. Under higher density, the plant-type was uniform and ideal for mechanized harvesting. Yield of high density rape was higher. Both yields on high and low density increased at 0 to 180kg/hm2 N rates. But when N rates were up to 270kg/hm2, both yields decreased. N use efficiency decreased at higher N rates and lower density. It indicated that higher density could promote seed N accumulation, decrease N fertilizer requirement and improve the N use efficiency. For the same targets yield (compared to the top economic yield at low density, the average yield of Hubei Province and that of China), the N fertilizer requirement significantly decreased by 22.9% to 30.6% after increasing planting density. Key words: Rapeseed (Brassica napus L.); Planting density; Nitrogen fertilizer; Yield; Nitrogen use efficiency

Oil Crops Research Institute of the Chinese Academy of Agricultural Sciences, Key Laboratory of Biology and Genetic Improvement of Oil Crops, Ministry of Agriculture, Wuhan 430062, China

为机械化种植油菜，研究不同施氮水平下密度对直播油菜生长发育和氮肥利用效率的影响。以甘蓝型油菜品种中双11号为材料，在低密（15万株/hm2）和高密（45万株/hm2）两种密度下，设施氮0、90、180和270kg/hm2处理，分析施氮量与密度对油菜农学性状、产量、氮肥利用效率的影响，并对比分析了密度与油菜需氮量之间的关系。结果表明，增密后油菜的株高、分枝数、单株角果数和每角粒数显著降低，分枝高度和千粒重显著增加。高密油菜群体株型结构整齐均一，更能满足机械化收获需要。高密处理油菜籽粒产量高于低密处理。施氮0～180kg/hm2时，两个密度下的籽粒产量均随施氮量增加而显著提高，但在270kg/hm2时两密度下油菜产量均有所下降。油菜氮肥表观利用率随着施氮量的增加而下降，但是又随种植密度的增加而增加。说明增加密度能促进氮素向油菜籽粒转移，降低每生产100kg籽粒油菜需氮量，提高氮肥表观利用率。在相同的目标产量（低密下的经济产量、湖北省油菜平均单产和全国油菜平均单产）下，油菜增密后可以节约氮肥用量22.9%~30.6%，增密减氮效果明显。

... 密植除了增加植株的叶面积指数和光能利用效率外^[24], 还能提高油菜氮肥利用效率^[25], 促进氮素向籽粒转移^[26], 起到增产效果 ...

2010

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马霓, 李云昌, 胡琼, 李俊, 李光明, 张春雷. 我国南方冬油菜机械化生产农艺研究进展. 中国油料作物学报, 2010, 32: 451-456

, Li Y

, Hu

, Li

, Li G

, Zhang C

Research of agronomic techniques form mechanical production of winter rape (Brassica napus L. ).

Chin J Oil Crop Sci, 2010, 32: 451-456 (in Chinese with English abstract)

油菜机械化生产是我国油菜生产发展的方向，近年来在南方冬油菜区进行了适合机械化生产的油菜品种筛选和鉴定，农机和农艺配套技术试验示范。本文综述了在油菜机械直播和机械移栽、抗倒伏措施及理想株型构建、施肥技术、病虫草害综合防治、化学催熟技术、机械收获技术等方面的研究概况，并探讨了存在的问题及今后的研究方向。

... 密植还可以提高油菜群体的整齐度, 使茎杆变细、分枝部位提高、分枝长度变短及成熟期相对集中, 有利于机械联合收获, 显著降低机收损失率^[27] ...

2014

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王玫, 陈洪伟, 王红利, 刘克锋. 独脚金内酯调控植物分枝的研究进展. 园艺学报, 2014, 41: 1924-1934

Wang

, Chen H

, Wang H

, Liu K

Research progress in regulatory role of strigolactones in shoot branching.

Acta Hort Sin, 2014, 41: 1924-1934 (in Chinese with English abstract)

The plant architecture of higher plants is regulated by environmental factors，genetic factors and phytohormones as well. Phytohormones play a critical role in regulating shoot branching. In addition to auxin and cytokinin，strigolactones are thought to be a new kind of hormone that regulate plant shoot branching. This review covers the regulatory roles for strigolactones，and its interaction with auxin and cytokinin in shoot branching regulation in higher plants. We also discuss the prospects in the gardening.

1College of Landscape，Beijing University of Agriculture，Beijing 102206，China；2College of Urban and Rural Development，Beijing University of Agriculture，Beijing 102206，China

高等植物株形的形成受外界环境、遗传以及植物激素等多因素的影响。植物激素对调控植物地上部分枝起着重要作用，除公认调控植物分枝的生长素和细胞分裂素以外，独脚金内酯是一种新被发现的调控植物分枝的激素。对独脚金内酯调控植物分枝及其与生长素、细胞分裂素协同作用的分子机理进行了综述，并对其在园艺方面所具有的潜在应用价值进行了展望。

... 分枝角度形成的直接原因是分枝感受向重力性, 受到植株内部生长素和独角金内酯等激素的调节^[28,29] ...

2015

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... 分枝角度形成的直接原因是分枝感受向重力性, 受到植株内部生长素和独角金内酯等激素的调节^[28,29] ...

2009

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... )油菜品种的过程中需要采用联合选育, 即采用轮回选择或聚合的方法来累积增效基因, 以提高选择效率^[30] ...