小麦茎秆实心度对茎秆强度的影响及相关性状QTL分析

引用本文

潘婷, 胡文静, 李东升, 程晓明, 吴荣林, 程顺和. 小麦茎秆实心度对茎秆强度的影响及相关性状QTL分析.作物学报, 2016, 43(1): 9-18
PAN Ting, HU Wen-Jing, LI Dong-Sheng, CHENG Xiao-Ming, WU Rong-Lin, CHENG Shun-He. Influence of Stem Solidness on Stem Strength and Stem Solidness Associated QTLs in Bread Wheat. Acta Agronomica Sinica, 2016, 43(1): 9-18 复制到剪切板

Doi:10.3724/SP.J.1006.2017.00009
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作物学报编辑部

小麦茎秆实心度对茎秆强度的影响及相关性状QTL分析

潘婷^1,^*, 胡文静^2,^*, 李东升², 程晓明², 吴荣林², 程顺和^1,^2,^*

¹南京农业大学作物遗传与种质创新国家重点实验室, 江苏南京210095

²江苏里下河地区农业科学研究所 / 国家小麦改良中心扬州分中心, 江苏扬州225007

^*通讯作者(Corresponding author): 程顺和, E-mail:13905273926@126.com ^**同等贡献(Contributed equally to this work)

第一作者联系方式: E-mail:862430412@qq.com, Tel: 18550707316

收稿日期:2016-09-18

基金:本研究由国家现代农业产业技术体系建设专项(CARS-03-03B, CARS-3-2-11)和江苏省农业科技自主创新资金项目[CX(13)2022]资助

摘要

在QTL水平上研究茎秆实心度与强度的遗传关系及实心度对茎秆强度的影响, 为小麦抗倒伏育种提供依据。利用普通小麦宁麦18 与实秆小麦种质“武云实秆”的F₂群体和F_2:3家系, 对小麦茎秆强度、实心度及影响实心度的相关性状包括厚径比、壁厚、茎粗和髓腔直径进行了相关分析, 并对茎秆强度相关性状QTL进行分子标记定位及遗传效应分析。结果表明小麦茎秆强度与厚径比、壁厚均呈极显著正相关, 与髓腔直径呈极显著负相关。基于复合区间作图法进行QTL定位, 检测到与茎秆强度、厚径比、壁厚、茎粗和髓腔直径相关QTL共23个, 分布在1B、3B、4A、4B、5A上, 表型贡献率3.5%~44.0%。在染色体3B、4A和5A上的标记区间 gwm547-gwm247、 wmc718-wmc468和 gwm156-gwm443均检测到贡献率很高的茎秆实心度相关QTL, 说明在这3条染色体上可能存在控制茎秆强度的主效QTL。用普通小麦宁麦13 (N13)×武云实秆的24个F₇家系检验分子标记 gwm247的可靠性表明利用标记 gwm247选育茎秆实心度优于宁麦13的概率较大。研究结果为进一步精细定位相关主效QTL以及分子标记辅助改良小麦茎秆强度奠定了基础。

关键词: 小麦; 茎秆强度; 实心度; QTL定位

Influence of Stem Solidness on Stem Strength and Stem Solidness Associated QTLs in Bread Wheat

PAN Ting^1,^**, HU Wen-Jing^2,^**, LI Dong-Sheng², CHENG Xiao-Ming², WU Rong-Lin², CHENG Shun-He^1,^2,^*

¹ State Key Laboratory of Crop Genetics and Germplasm Enhancement, Nanjing Agricultural University, Nanjing 210095, China

² Institute of Agriculture Sciences of the Lixiahe District / Branch of National Wheat Improvement Center, Yangzhou 225007, China

Fund:This study was supported by the China Agriculture Research System (CARS-03-03B, CARS-3-2-11) and the Independent Innovation Fund for Agricultural Science and Technology in Jiangsu Province [CX(13)2022]

Abstract

Improving stem strength is an important way to increase lodging resistance of wheat. Solid stem traits, including thickness-diameter ratio of culm, stem diameter, culm wall thickness and pith diameter, are the main targets for improvement of stem strength in wheat. In the present study, an F₂ population and F_2:3 families derived from the cross N18 × WYSG were used to study the influence of stem solidness on stem strength and to identify QTLs for solidness related traits. The thickness-diameter ratio of culm, stem diameter, culm wall thickness and pith diameter were significantly correlated with stem strength. By using composite interval mapping (CIM), a total of 23 QTLs for the five traits were detected on chromosomes 1B, 3B, 4A, 4B, and 5A. Phenotypic variations explained by individual QTL ranged from 3.5% to 44.0%. Pleiotropic QTLs were found on chromosomes 3B ( gwm547-gwm247), 4A ( wmc718-wmc468), and 5A ( gwm156-gwm443), all contributing very high phenotypic variations. The reliability of the marker gwm247 was confirmed using 24 F₇ lines derived from the N13 (hollow stem) × WYSG (solid stem) cross. The markers linked to major QTLs could be valuable in wheat breeding for lodging resistance with marker assisted selection technology.

Keyword: Wheat; Stem strength; Stem solidness; Quantitative trait loci

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随着小麦产量不断提高同时伴随施肥量的增加, 倒伏成为威胁我国小麦高产稳产的重要因素之一^{[1, 2]}。植株矮化可以有效缓解小麦倒伏危害的发生^{[3, 4]}, 但一般会导致生物量减少, 群体郁闭, 通风透光差, 病害加重, 影响产量^[5], 并且植株矮化后茎秆强度可能减弱, 仍然会发生倒伏。因此, 改良茎秆强度提高抗倒性是解决小麦倒伏的理想途径^[6]。

目前, 抗倒性相关研究, 包括茎秆形态结构^{[7, 8, 9, 10, 11]}、解剖结构^{[12, 13]}、化学成分^[7]等对抗倒性的影响。汪灿等^[12]关于荞麦茎秆解剖结构及木质素代谢与抗倒性关系的研究指出茎秆中木质素含量、机械组织厚度、茎壁厚度、大维管束数目和维管束面积可以作为禾本科茎秆抗倒伏育种的重要指标。有学者认为小麦茎秆强度与中下部茎粗及鲜重呈显著正相关, 而与各节间长度呈显著负相关^{[14, 15, 16]}。但是也有一些学者认为茎秆粗度与秆强之间没有显著相关性^[17]。实秆小麦在抗倒伏育种中起很重要的作用。陈华华等^[18]和Kong等^[19]研究发现, 实秆小麦较普通小麦具有极小的髓腔, 腔中有髓, 其机械组织含量显著高于普通小麦, 因而实秆小麦茎秆强度较高。

在20世纪40年代, 为了预防麦茎蜂(Cephus cinctus Norton)的危害, 加拿大推广了实秆小麦Rescue和Chinook^{[20, 21]}。McNeal等^[22]和Wellace等^[23]提出实秆性状可能对小麦产量造成影响。Adrianna等^[24]认为实秆麦是到目前为止预防麦茎蜂危害最有效的方法, 其实验表明实秆基因不会影响籽粒品质, 但对产量有影响, 当麦茎蜂数量很少时, 实秆麦的产量显著低于空秆麦, 而当麦茎蜂数量大到足以影响小麦产量时, 实秆麦的产量与空秆麦之间没有显著差别。但是也有实验表明, 实秆基因与产量性状之间没有显著关系^[17]。McNeal等^[25]以6个实秆小麦与3个普通小麦杂交得到的17个系为材料, 证明实秆基因并不是影响小麦高产的因素。

国内、外学者利用分子标记技术对小麦茎秆强度相关性状进行了基因定位。Hai等^[14]利用CA9613 × H1488创建的DH群体对控制茎秆强度、茎壁厚度、髓腔直径、茎粗进行基因定位, 其中控制茎秆强度的基因位于3A和3B上, 并分别解释10.6%和16.6%的表型变异。郭会君等^[26]研究发现控制小麦茎秆强度的基因位于2D和5B上, 共同解释24%的表型变异。Verm等^[27]利用Milan × Catbird产生的DH群体将控制茎秆抗折力的基因定位在4B和2D上, 表型贡献率分别为11%和13%。

国内研究茎秆强度与茎壁厚、茎粗、节间长度等关系较多, 较少研究实秆小麦与茎秆强度的关系。本实验旨在研究小麦茎秆实心度对茎秆强度的影响, 并定位相关基因进行, 为培育小麦抗倒伏新品种奠定基础。

1 材料与方法

1.1 试验材料

武云实秆是四川省农业科学院采用实秆小麦86-741^[28]与川麦42 ^[29]杂交、回交育成, 其茎秆基部有极小髓腔, 髓腔中有髓。以空心高秆种质“ 宁麦18” 为母本, 矮秆实心种质“ 武云实秆” 为父本。于江苏里下河地区农业科学研究所万福基地试验田, 2013— 2014年生长季获得F₂群体, 2014— 2015年生长季获得的杂交F₁代和F_2:3家系。验证群体是宁麦13和武云实秆的杂交F₇代高代系。

1.2 实心度的测定

2014年和2015年, 分别选取成熟期小麦武云实秆、宁麦18和F₁植株各20株, 取F₂群体184个单株及对应F_2:3家系每行20株。用手术刀切开基部第1节间和第2节间, 用游标卡尺测量外茎及内径的长度, 每个单株成“ 米” 字形测3次取均值。外径记为茎粗, 内径记为髓腔直径, 秆壁厚度为外径与内径差值的一半。实心度用厚径比来表示, 厚径比为秆壁厚度与茎粗的比值^{[24, 30]}。

1.3 茎秆强度的测定

2015年度, 分别选取成熟期小麦武云实秆、宁麦18及F₁植株各20株, 取对应184个F_{2: 3}家系每行20株采用茎秆强度检测仪YY1-a测力计(浙江托普仪器有限公司), 垂直作用于茎秆基部第1和第2节间的中间位置, 缓缓用力, 茎秆折断时的最大读数即为该茎的茎秆强度。

1.4 QTL作图及遗传分析

从1872对SSR引物中筛选出221对具有多态性的引物。利用JoinMap4.0构建遗传图谱, 其中170个SSR标记基本均匀分布在18条染色体上(1D、4D、7D未覆盖到), 覆盖小麦染色体组1482.15 cM, 标记间平均遗传距离为8.72 cM。利用WinQTLCart 2.5软件在全基因组范围对控制小麦茎秆强度及实心度相关性状的QTL进行检测, 采用复合区间作图法进行QTL定位^[31], 当阈值(LOD)大于2.5时, 就认为可能存在一个QTL, 同时计算每个QTL的贡献率和加性效应。

2 结果与分析

2.1 亲本间差异显著性分析

武云实秆和F₁植株基部第1和第2节间的秆强、厚径比、壁厚、髓腔直径均与宁18之间存在极显著差异, 武云实秆的茎秆强度和厚径比分别是宁麦18的2.20倍和2.97倍(表1)。而武云实秆基部第1节间的茎粗与宁麦18之间差异并不显著。在这些性状中, 茎秆强度的变异系数都高于10%, 基部第2节间变异系数较大, 说明茎秆强度在品种间有较高的遗传变异。

表1 亲本宁麦18、武云实秆及其F₁代植株基部第1和第2节间茎秆强度相关性状 Table 1 Stem strength related traits for the first and second stem internodes of Ningmai 18, Wuyunshigan, and their F₁

2.2 各性状在F_2:3家系中的变异

F_2:3家系茎秆第2节间的强度、厚径比、茎粗、秆壁厚度和髓腔直径均呈现连续分布(图1), 表明这些性状均属多基因控制的数量性状。其中茎秆强度变幅为11.0~49.4 N, 厚径比的变幅为0.13~0.44, 茎粗的变幅为3.43~4.65 mm, 壁厚的变幅为0.58~1.69 mm, 髓腔直径的变幅为0.45~3.09 mm。这几个性状均表现超亲分离, 其中茎秆强度和茎粗表现更明显, 表明实秆的遗传基础更复杂。

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	图1 宁麦18 (P₁)与武云实秆(P₂)杂交F_2:3家系的表型分布Fig. 1 Phenotypic variations in the F_2:3 lines derived from the cross between Ning 18 (P₁) and Wuyunshigan (P₂)

2.3 茎秆强度与茎秆性状的相关分析

F_2:3家系基部第2节间茎秆强度与厚径比、壁厚呈极显著正相关(表2), 而与髓腔直径呈极显著负相关, 即小麦秆壁越厚、髓腔越小, 其实心度越高, 茎秆强度也就越大。而茎秆粗度与茎秆强度并没有显著相关性。

茎秆基部第2节间的实心度与茎壁厚度呈极显著正相关, 与茎粗和髓腔直径呈显著负相关, 即茎秆越细、茎壁越厚髓腔直径越小, 茎秆的实心度就越高。

表2 小麦茎秆强度与性状的相关系数 Table 2 Correlation coefficients between stem strength and culm traits in wheat

2.4 QTL定位及遗传效应分析

共检测到4个控制茎秆强度的QTL (表3和图2), 位于3B、4A、4B和5A染色体上, 分别解释表型变异的20.7%、4.9%、6.9%和5.0%。其中QSs-3B、QSs-4B和QSs-5A-2增效基因来自武云实秆, 而QSs-4A增效基因来自宁麦18。

在3B、4A和5A染色体上共检测到4个控制小麦厚径比的QTL (表3和图2)。其中, QTdr-3B-2在两年中均被检测到, 且分别解释表型贡献率为28.9%和43.9%, 遗传效应很大, 增效基因均来自为武云实秆。

与茎秆粗度相关的2个QTL分别位于1B和3B染色体上, 其中QSd-1B在两年中均被检测到, 分别可解释表型贡献率的20.7%和9.1%。

在3B、4A和5A染色体上检测到4个控制小麦髓腔直径的QTL, 单个QTL可解释表型变异的4.0%~41.9%, 其中QPd-3B-2和QPd-4A加性效应为正, 增效基因均来自宁麦18。

在3B、4A、4B和5A染色体上检测到5个控制茎壁厚度的QTL, 加性效应均来自武云实杆。位于3B上的QCwt-3B-2在两年度均被检测到, 表型贡献率分别为22.3%和44.0%; 而位于4A和5A上的QTL对表型性状的贡献率分别为24.5%、10.9%和10.3%。

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	图2 小麦茎秆强度相关性状QTL在染色体上的分布Fig. 2 Chromosome location of putative QTL associated with traits related to stem strength in wheat

表3 小麦茎秆强度相关性状的QTL Table 3 Putative QTLs for traits related to stem strength

2.5 分子标记gwm247的可靠性检验

用24个宁麦13× 武云实秆F₇代高代系验证分子标记gwm247的可靠性, SSR标记gwm247在宁麦13× 武云实秆F₇家系中扩增结果显示, 除了23号家系, 其余家系在此位点的基因型均与武云实秆相同(图3), 其茎秆厚径比除21号和23号家系外, 其余家系均比宁麦13显著较大(图4和表4), 表明利用标记gwm247选育出茎秆实心度优于宁麦13的茎秆类型概率较大。

	Figure Option View Download New Window
	图3 SSR标记gwm247在宁麦13 (P₃) × 武云实秆(P₂) 24个F₇家系中的扩增结果Fig. 3 Amplification patterns of the SSR marker gwm247 in 24 F₇ lines derived from the cross of Ningmai 13 (P₃)× Wuyunshigan (P₂)

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	图4 宁麦13 (P₃) × 武云实秆(P₂) F₇家系茎秆强度与厚径比分布图Fig. 4 Histogram of stem strength and thickness-diameter ratio of culm for 24 F₇ lines derived from Ningmai 13 (P₃) and Wuyunshigan (P₂)

表4 各家系与宁麦13茎秆强度与厚径比的差异显著性比较 Table 4 Difference in stem strength and thickness-diameter ratio of culm between each lines and Ningmai 13

3 讨论

植株高度^{[32, 33]}、重心高度^[34]、茎秆节间长度^[17]、髓腔直径^[18]、茎壁厚度^[35]、节间干重^[11]、维管束量^[36]、木质素、纤维素含量^[37]等都是影响植株抗倒性的重要因素。本研究发现茎秆强度与基部节间第1和第2节间实心度、秆壁厚度和髓腔直径均呈极显著相关。对于茎秆粗度与抗倒性的关系, 不同学者观点分歧。部分学者认为茎秆越粗, 植株抗倒伏能力越强^{[5, 11]}, 也有不少学者认为茎秆粗度与抗倒伏能力关系不密切^[17]。而本研究表明, 小麦茎秆强度与基部第1节间茎秆粗度关系不显著, 与基部第2节间关系显著, 这说明基部第2节间粗度可作为衡量茎秆强度的一个指标。因此在小麦育种过程中选择茎秆粗、秆壁厚、髓腔小、实心度高的小麦品种有助于提高小麦茎秆强度从而提高抗倒性以达到稳产的目的。

与Hai等^[14]的报道一致, 本研究发现小麦茎秆基部第2节间强度、实心度、壁厚、茎粗和髓腔直径在F_2:3家系中均呈现连续分布, 表明这些性状均属多基因控制的数量性状。共检测到23个与这5个性状关联的QTL, 分布于5条染色体上, 其中在3B染色体标记区间gwm547-gwm247同时检测到控制5个性状的QTL, 解释的表型变异率为8.14%~44.00%, 说明该区域在控制小麦抗倒伏方面起着重要作用。茎秆强度基因被定位在3B、4A、4B和5A染色体上, 而位于3B上的QSs-3B-2解释表型变异率较大, 该位点与Hai等^[14]将控制秆强的基因QSs-3B定位在标记gwm108-wmc291之间结果相近, 可能是控制茎秆强度的主效QTL。

茎秆实心性状受到复杂的数量性状控制, 在回交育种过程中常常丢失实心性状, 所以仅通过观察表型来改良小麦品种的茎秆实心度很困难, 分子标记辅助育种可以将分子标记应用于作物改良。早在1959年就有关于实秆基因的报道, Larson和MacDonald等^{[38, 39]}利用S-615的单体系推断出3B和5A染色体上有控制小麦茎秆实心性状的位点。到目前为止, 关于小麦茎秆实心性状的QTL大多定位在3BL上, 并与标记gwm247、gwm547、gwm340和barc77紧密连锁^{[18, 19, 40]}。本研究验证了前人的这些报道, 而且还发现了一些茎秆实心度相关的新位点(表3和图2), 例如在4A染色体标记wmc718- wmc468之间同时定位了控制茎秆实心度、壁厚和髓腔直径的QTL, 在一定程度上解释了茎秆壁厚、髓腔直径与实心度存在显著相关性。其中位于3B染色体的QTdr-3B-2表型贡献率很大, 该位点很可能是控制茎秆实心度的主效QTL。

本研究在分子标记gwm247附近检测到多个与抗倒性相关的QTL, 且对表型贡献率都很大, gwm247可用作分子标记辅助抗倒伏育种的重要标记。本研究用普通小麦宁麦13× 武云实秆的24个F₇家系检验分子标记gwm247的可靠性表明利用标记gwm247选育出茎秆实心度优于宁麦13的概率较大(图3、图4和表4)。但要开发可靠分子标记就需要进一步定位或克隆该基因。本研究将为进一步精细定位实秆基因, 选育抗倒小麦新品种奠定基础。

4 结论

检测到与茎秆强度、厚径比、壁厚、茎粗和髓腔直径相关QTL共23个, 分布在1B、3B、4A、4B、5A上, 其中在3B染色体标记区间gwm547-gwm247同时检测到控制5个性状的QTL, 解释的表型变异率为8.14%~44.00%。小麦茎秆强度与厚径比、壁厚均呈极显著正相关, 与髓腔直径呈极显著负相关。利用gwm247选育茎秆实心度优于宁麦13的可靠性较高。

The authors have declared that no competing interests exist.

作者已声明无竞争性利益关系。

参考文献

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2014

0.0

... 随着小麦产量不断提高同时伴随施肥量的增加, 倒伏成为威胁我国小麦高产稳产的重要因素之一^[1,2] ...

2012

0.0

... 随着小麦产量不断提高同时伴随施肥量的增加, 倒伏成为威胁我国小麦高产稳产的重要因素之一^[1,2] ...

2014

0.0

许琦, 杨娜, 柴永峰, 杨淑巧, 赵智勇, 裴蕾, 郭文治, 刘跃鹏. 中国小麦主要矮秆基因的分布及其对株高的影响. 西北农业学报 2014, 23(5): 59-64

, Yang

, Chai Y

, Yang S

, Zhao Z

, Pei

, Guo W

, Liu Y

Distribution and impact on plant height of major wheat dwarfing genes in China.

Acta Agric Boreali-Occids Sin, 2014, 23(5): 59-64 (in Chinese with English abstract)

In order to find out the distribution and utilization of the dwarf genes in Chinese winter wheat region and clarify their impact on wheat plant height and to improve wheat’s water and fertilizer use efficiency, the dwarf gene molecular markers developed by predecessors and 210 winter wheat materials were utilized to studied the distribution and utilization of the Rht-B1b , Rht-D1b and Rht8 genes in Chinese winter wheat region. The research results showed that 51 wheat varieties/lines contained the dwarf gene Rht-B1b , accounting for 24.3% of the total 210 wheat varieties/lines; 40 wheat varieties/lines contained the dwarf gene Rht-D1b , accounting for 19.0%; 94 wheat varieties/lines contained the dwarf gene Rht8 , accounting for 44.8%. The average plant height of wheat varieties/lines containing the dwarf gene Rht-B1b was 84.0 cm, but that without the Rht-B1b dwarf gene was 93.7cm; the average plant height of wheat varieties/lines containing the dwarf gene Rht-D1b was 77.9 cm, but that without the Rht-D1b dwarf gene was 94.6cm; similarly, the average plant height of wheat varieties/lines containing the dwarf gene Rht8 was 90.0 cm, but that without the Rht-B1b dwarf gene was 92.5 cm. The average plant height of wheat containing single dwarf gene of Rht-B1b, Rht-D1b and Rht8 declined 9.7 cm, 16.7 cm and 2.5 cm, respectively. Based on the above study, further analysis was conducted the combined distribution of three dwarf genes and the effects of different combination of dwarf genes on lowering plant height, the result showed that the combined effect of three dwarf genes was greater than the combined effect of two dwarf genes, and the combined effect of two dwarf genes was greater than the effect of single dwarf genes. Moreover, the utilization of wheat dwarf gene molecular markers in wheat breeding was discussed.

为了了解矮秆基因在中国冬麦区的分布及利用情况和明确矮秆基因对小麦株高的影响，进一步提高小麦的水肥利用效率，利用前人开发的矮秆基因分子标记和收集的210份冬小麦材料研究矮秆基因 Rht-B1b 、 Rht-D1b 和 Rht8 在中国冬麦区的分布和利用情况。结果表明：210份冬小麦材料含有矮秆基因 Rht-B1b 的材料有51份，占总材料的24.3%；含有矮秆基因 Rht-D1b 的材料有40份，占19.0%；含有 Rht8 矮秆基因的材料有94份，占44.8%。含有 Rht-B1b 的小麦材料平均株高为84.0 cm，不含有 Rht-B1b 的小麦材料的平均株高为93.7 cm；含有 Rht-D1b 的小麦材料平均株高为77.9 cm，不含有 Rht-D1b 的小麦材料的平均株高为94.6 cm；含有 Rht8 的小麦材料平均株高为90.0 cm，不含有 Rht8 的小麦材料的平均株高为92.5 cm； Rht-B1b 、 Rht-D1b 和 Rht8 对小麦平均株高的降秆作用分别为9.7 cm、16.7 cm和2.5 cm。进一步分析不同矮秆基因组合的联合分布情况，同时分析不同矮秆基因的联合降秆作用，结果表明3个矮秆基因的联合降秆作用大于2个矮秆基因的降秆作用，2个矮秆基因的联合降秆作用大于单个矮秆基因的降秆作用；另外还对分子标记在小麦矮秆基因的选择利用方面进行了探讨。

... 植株矮化可以有效缓解小麦倒伏危害的发生^[3,4], 但一般会导致生物量减少, 群体郁闭, 通风透光差, 病害加重, 影响产量^[5], 并且植株矮化后茎秆强度可能减弱, 仍然会发生倒伏 ...

2014

0.0

崔淑佳, 潘晓萍, 高居荣, 王洪刚, 李兴锋. 不同小麦品种(系)株高及节间长度研究. 山东农业科学, 2014, 46(10): 19-22

Cui S

, Pan X

, Gao J

, Wang H

, Li X

Analysis of plant height and internode length of different wheat varieties (lines).

Shand ong Agric Sci, 2014, 46(10): 19-22 (in Chinese with English abstract)

对矮秆、中秆、高秆共20个小麦品种（系）的株高及各节间长进行研究，结果表明：矮秆小麦主要是通过缩短不同节间长度来降低株高，特别是基部节间；株高相近的品种（系）同一节间占株高的比例基本相同，株高类型不同的品种（系）其节间比例的变化相对较大；两个小偃麦矮秆种质31504-1和31505-1株高构成指数IL 明显大于其它小麦，说明其抗倒伏性更稳定，这对小麦株高的遗传改良具有重要意义。

2002

0.0

肖世和, 张秀英, 闫长生, 张文祥, 海林, 郭会君. 小麦茎秆强度的鉴定方法研究. 中国农业科学, 2002, 35: 7-11

Xiao S

, Zhang X

, Yan C

, Zhang W

, Hai

, Guo H

Determination of resistance to lodging by stem strength in wheat.

Sci Agric Sin, 2002, 35: 7-11 (in Chinese with English abstract)

茎秆强度决定小麦品种的抗倒伏性能,但现有鉴定茎秆强度的方法还不完善。用秆强测定器测定了661个小麦品种和1183个F2代单株,结果表明,乳熟至蜡熟期是鉴定茎秆强度的适宜时期,利用秆强测定器可从F2代分离群体中鉴定出强秆单株。成熟时茎秆强度降低,不易区分茎秆强度的品种间差异。通过分析30个品种的茎秆强度值与茎秆解剖性状的相关性,发现上部节间的髓腔大小与茎秆强度呈极显著负相关,而下部节间的茎秆粗度与茎秆强度呈极显著正相关。由于穗部重量和生物产量与茎秆强度呈极显著正相关,育种家应选择秆强与穗重相适应的品种以提高产量潜力和稳产性。

... 部分学者认为茎秆越粗, 植株抗倒伏能力越强^{[5, 11]}, 也有不少学者认为茎秆粗度与抗倒伏能力关系不密切^[17] ...

2007

0.0

... 因此, 改良茎秆强度提高抗倒性是解决小麦倒伏的理想途径^[6] ...

2004

0.0

... 目前, 抗倒性相关研究, 包括茎秆形态结构^{[7,8,9,10,11]}、解剖结构^[12,13]、化学成分^[7]等对抗倒性的影响 ...

2015

0.0

韦爽, 万燕, 晏林, 向达兵, 赵钢, 黄凯丰, 谭茂玲, 宋月, 张莹. 不同苦荞品种茎秆强度和植株性状的差异及其相关性. 作物杂志, 2015, (2): 59-63

Wei

, Wan

, Yan

, Xiang D

, Zhao

, Huang K

, Tan M

, Song

, Zhang

Difference and correlation of different tartary buckwheat cultivars in stem strength and plant characters.

Crops, 2015, (2): 59-63 (in Chinese with English abstract)

... 目前, 抗倒性相关研究, 包括茎秆形态结构^{[7,8,9,10,11]}、解剖结构^[12,13]、化学成分^[7]等对抗倒性的影响 ...

2014

0.0

... 目前, 抗倒性相关研究, 包括茎秆形态结构^{[7,8,9,10,11]}、解剖结构^[12,13]、化学成分^[7]等对抗倒性的影响 ...

2015

0.0

... 目前, 抗倒性相关研究, 包括茎秆形态结构^{[7,8,9,10,11]}、解剖结构^[12,13]、化学成分^[7]等对抗倒性的影响 ...

2001

0.0

... 目前, 抗倒性相关研究, 包括茎秆形态结构^{[7,8,9,10,11]}、解剖结构^[12,13]、化学成分^[7]等对抗倒性的影响 ...

... 3 讨论植株高度^[32,33]、重心高度^[34]、茎秆节间长度^[17]、髓腔直径^[18]、茎壁厚度^[35]、节间干重^[11]、维管束量^[36]、木质素、纤维素含量^[37]等都是影响植株抗倒性的重要因素 ...

... 部分学者认为茎秆越粗, 植株抗倒伏能力越强^{[5, 11]}, 也有不少学者认为茎秆粗度与抗倒伏能力关系不密切^[17] ...

2014

0.0

汪灿, 阮仁武, 袁晓辉, 胡丹, 杨浩, 林婷婷, 何沛龙, 李燕, 易泽林. 荞麦茎秆解剖结构和木质素代谢及其与抗倒性的关系. 作物学报, 2014, 40: 1846-1856

Wang

, Ruan R

, Yuan X

, Hu

, Yang

, Lin T

, He P

, Li

, Yi Z

Relationship of anatomical structure and lignin metabolism with lodging resistance of culm in buckwheat.

Acta Agron Sin, 2014, 40: 1846-1856 (in Chinese with English abstract)

Lodging is a major problem in buckwheat ( Fagopyrum esculentum Moench ) production. In this study, we investigated the relationship between anatomical structure together with lignin metabolism and lodging resistance. The results indicated that the lodging resistance of culm in buckwheat was closely related to the anatomical structure and lignin metabolism of culm. The lodging percentage was negatively correlated with snapping resistance parameter of culm ( r = –0.907, P , lignin content ( r = –0.844, P , mechanical tissue thickness ( r = –0.881, P , culm wall thickness ( r = –0.947, P , vascular bundle area ( r = –0.846, P , mechanical tissue layer number ( r = –0.806, P r = –0.709, P , but positively correlated with lodging index ( r = 0.842 , P . The lignin content was positively correlated with activities of phenylalanine ammonialyase ( r = 0.984, P 4-coumarate: CoA ligase ( r = 0.927, P cinnam yl alcohol dehydrogenase ( r = 0.862, P Therefore, lignin content in culm, mechanical tissue layer number, mechanical tissue thickness, culm wall thickness, large vascular bundle number, and vascular bundle area can be used as main indicators to evaluate lodging resistance in buckwheat. Buckwheat cultivars with high resistance to culm snapping and lodging usually have high lignin content, large number of mechanical tissue layer, and large number of vascular bundle, and thick mechanical tissue and culm wall, and large vascular bundle area.

以抗倒伏能力不同的 4 个荞麦品种研究表明，荞麦茎秆抗倒伏能力与茎秆解剖结构和木质素代谢密切相关。倒伏率与茎秆抗折力参数 ( r = –0.907, P 、木质素含量 ( r = –0.844, P 、机械组织厚度 ( r = –0.881, P 、茎壁厚度 ( r = –0.947, P 、维管束面积 ( r = –0.846, P 、机械组织层数 ( r = –0.806, P 和大维管束数目 ( r = –0.709, P 呈显著负相关，而与倒伏指数 ( r = 0.842, P ) 呈显著正相关。木质素含量与苯丙氨酸解氨酶 (PAL) 活性 ( r = 0.984, P 、 4- 香豆酸 :CoA 连接酶 (4CL) 活性 ( r = 0.927, P 和肉桂醇脱氢酶 (CAD) 活性 ( r = 0.862, P 呈显著正相关。茎秆木质素含量、机械组织层数、机械组织厚度、茎壁厚度、大维管束数目和维管束面积可以作为荞麦茎秆抗倒伏能力的重要评价指标。茎秆木质素含量高、机械组织层数多、机械组织和茎壁厚、大维管束数目多且维管束面积大的荞麦品种，其茎秆抗折力参数大、倒伏指数小、抗倒伏能力强。

... 目前, 抗倒性相关研究, 包括茎秆形态结构^{[7,8,9,10,11]}、解剖结构^[12,13]、化学成分^[7]等对抗倒性的影响 ...

... 汪灿等^[12]关于荞麦茎秆解剖结构及木质素代谢与抗倒性关系的研究指出茎秆中木质素含量、机械组织厚度、茎壁厚度、大维管束数目和维管束面积可以作为禾本科茎秆抗倒伏育种的重要指标 ...

2010

0.0

桑云, 赵亮, 张坤普, 田纪春, 叶宝兴. 小麦DH群体穗下节间直径、茎壁厚及茎壁面积的QTL定位. 作物学报, 2010, 36: 61-67

Sang

, Zhao

, Zhang K

, Tian J

, Ye B

Mapping QTLs for uppermost internode diameter and thickness and area of culm wall with doubled-haploid population in wheat.

Acta Agron Sin, 2010, 36: 61-67 (in Chinese with English abstract)

A population of 168 doubled haploid derived from two elite Chinese wheat ( Triticum aestivum L.) cultivars Huapei 3 and Yumai 57, grown in two continuous cropping seasons in Tai’an, Shandong province, was used to investigate the genetic basis of uppermost internode diameter (UID) and thickness (CWT) and area (CWA) of culm wall in wheat. The quantitative trait loci (QTLs) for each trait were analyzed based on the constructed molecular linkage map of this population, including 324 SSR markers covering the whole wheat genome. The additive effects, epistatic effects, and their interactions with environment were identified by using the mixed linear model approach. A total of ten additive QTLs and six pairs of epistatic QTLs were detected. Among the ten QTLs, three had interactions with environment. For UID, four QTLs were detected on chromosomes 2D, 3D, and 5D (two regions), and explained 14.98% of phenotypic variation. Three QTLs for CWT were identified and accounted for 14.41% of the variation in the same or close region chromosome of UID, which showed pleiotropisms or tight linkages. Three QTLs on chromosome 3D, 4B and 5D controlling CWA explained 19.03% of the variation. Two intervals of XCFD53–XWMC18 and XWMC215–XBARC34 on chromosomes 2D and 5D for thickness and area of culm wall explained 11.37% and 10.98% phenotypic variation, respectively. They could be used in marker-assisted selection. For the six epistatic QTLs, the epistatic effects were not significant and without epistatic × environment interactions. These QTLs ac counted for 19.01% of the observed phenotypic variation, without any additive effects (except qCWA-3D ) of their own but involved in epistatic interactions. Such loci might play the role of modifying agents that tend to activate other loci or modify the action of other loci.

由小麦品种花培 3 号和豫麦 57 杂交获得 168 个 DH 株系，连续两年在山东泰安种植，利用 324 个 SSR 标记构建遗传连锁图谱，并基于混合线性模型对控制穗下节间直径、茎壁厚及茎壁面积的 QTL 遗传效应和环境互作效应进行分析。共检测到 10 个加性效应位点和 6 对上位效应位点，其中 3 个加性位点参与环境互作效应。检测到位于染色体 2D 、 3D 和 5D(2 个 ) 控制穗下节间直径的 4 个加性 QTLs ，与控制茎壁厚的 3 个加性位点相同或相邻，表现出一因多效或紧密连锁效应。两个位于染色体 2D 和 5D 控制茎壁厚和茎壁面积 QTL 有较大遗传贡献率，分别为 11.37% 和 10.98% ，适用于分子标记辅助育种和聚合育种。 6 对上位性效应遗传贡献率较小、无环境互作效应。

... 目前, 抗倒性相关研究, 包括茎秆形态结构^{[7,8,9,10,11]}、解剖结构^[12,13]、化学成分^[7]等对抗倒性的影响 ...

2005

0.0

... 有学者认为小麦茎秆强度与中下部茎粗及鲜重呈显著正相关, 而与各节间长度呈显著负相关^[14,15,16] ...

... Hai等^[14]利用CA9613 #cod#x000D7 ...

... 与Hai等^[14]的报道一致, 本研究发现小麦茎秆基部第2节间强度、实心度、壁厚、茎粗和髓腔直径在F_2:3家系中均呈现连续分布, 表明这些性状均属多基因控制的数量性状 ...

... 茎秆强度基因被定位在3B、4A、4B和5A染色体上, 而位于3B上的QSs-3B-2解释表型变异率较大, 该位点与Hai等^[14]将控制秆强的基因QSs-3B定位在标记gwm108-wmc291之间结果相近, 可能是控制茎秆强度的主效QTL ...

2004

0.0

... 有学者认为小麦茎秆强度与中下部茎粗及鲜重呈显著正相关, 而与各节间长度呈显著负相关^[14,15,16] ...

2002

0.0

... 有学者认为小麦茎秆强度与中下部茎粗及鲜重呈显著正相关, 而与各节间长度呈显著负相关^[14,15,16] ...

2003

0.0

董琦, 王爱萍, 梁素明. 小麦基部茎节形态结构特征与抗倒性的研究. 山西农业大学学报, 2003, (3): 188-191

Dong

, Wang A

, Liang S

Study on the architectural characteristics of wheat stalks.

J Shanxi Agric Univ, 2003, (3): 188-191 (in Chinese with English abstract)

通过不同栽培密度条件下小麦茎秆形态结构特征的观察,明确了小麦茎秆的生长状况与植株的抗倒性密切相关.基部节间的健壮程度与小麦抗倒伏相关性大小为:第二节间＞第一节间＞第三节间.茎秆健壮程度因素对茎秆倒伏指数的影响大小为:基部节间壁厚＞基部节间长度＞节间内径大小,而节间外径大小与植株抗倒关系不密切.

... 但是也有一些学者认为茎秆粗度与秆强之间没有显著相关性^[17] ...

... 但是也有实验表明, 实秆基因与产量性状之间没有显著关系^[17] ...

... 部分学者认为茎秆越粗, 植株抗倒伏能力越强^{[5, 11]}, 也有不少学者认为茎秆粗度与抗倒伏能力关系不密切^[17] ...

2008

0.0

陈华华, 李俊, 万洪深, 王玲玲, 彭正松, 杨武云. 实心小麦86-741茎秆的解剖分析及壁厚特性的SSR标记. 作物学报, 2008, 34: 1381-1385

Chen H

, Li

, Wan H

, Wang L

, Peng Z

, Yang W

Microsatellite markers for culm wall thickness and anatomical features of solid stem Wheat 86-741.

Acta Agron Sin, 2008, 34: 1381-1385 (in Chinese with English abstract)

Lodging is a big problem in further increasing yield of wheat ( Triticum aestivum L.). Development of variety with thickwall of basal internodes is an effective way to improve lodging resistance in wheat. A solid stem line “86-741” has been developed from the cross between hexaploid Triticale and common wheat, and acted as parent in lodging resistance breeding. To detect the gene(s) of culm wall thickness in 86-741, we made a cross between 86-741 and a stem hollow wheat line “Xinjiang Daomai” [ Triticum petropavlovskyi Udacz. et Migusch] in 2004–2005 growth season, and developed its F 2 population in 2005–2006 growth season. The F 2 population was used for bulked segregation analysis (BSA) of the gene related to culm wall thickness by SSR markers. Anatomical analysis revealed that 86-741 had very smaller stem pith cavity with pith, thicker mechanical tissue and more vascular bundles than Xinjiang Daomai. The F 1 progeny of the cross showed some tendencies to its parent 86-741: culm with smaller diameters of pith cavity than the parent Xinjiang Daomai. The percentage of mechanical tissue in F 1 was smaller than that in 86-741, and the number of vascular bundles in F 1 was less than that in 86-741, but more than that in Xinjiang Daomai. Two recessive genes relating to pith and a single dominant gene on 3BL relating to culm wall thickness were detected in 86-741, respectively. The gene controlling culm wall thickness was temporarily designated as Cwt-1. We found 3 microsatellite (SSR) markers Xgwm-247, Xgwm-340, and Xgwm-547 linking to Cwt-1, in which Xgwm-247 was the closest to the gene with genetic distance of 5.3 cM. Our results indicated that Xgwm-247 can be used in marker-assisted selection (MAS) for lodging resistance breeding by selecting culm wall thickness trait in wheat.

以实心小麦86-741、新疆稻麦及其杂种F 1 为材料, 利用石蜡切片和SSR标记方法对86-741的实心特性进行了分析。结果表明, 实心小麦86-741茎内仅有极小髓腔, 髓腔中有髓, 机械组织中有维管束的分布; 杂种F1偏向实心亲本86-741, 有髓腔, 但髓腔的直径小于新疆稻麦, 机械组织所占的比例小于亲本86-741; 86-741维管束数最多, F 1 代其次, 新疆稻麦最少。遗传分析与SSR标记结果表明, 86-741髓的有无是由双隐性基因控制的; 而其壁厚性状由位于3BL染色体上的单显性基因控制, 与SSR标记Xgwm-547的遗传距离为5.3 cM。暂将该壁厚基因定名为Cwt-1。Xgwm-547可用于辅助转育壁厚特性, 提高小麦品种的抗倒伏能力。

... 陈华华等^[18]和Kong等^[19]研究发现, 实秆小麦较普通小麦具有极小的髓腔, 腔中有髓, 其机械组织含量显著高于普通小麦, 因而实秆小麦茎秆强度较高 ...

... 到目前为止, 关于小麦茎秆实心性状的QTL大多定位在3BL上, 并与标记gwm247、gwm547、gwm340和barc77紧密连锁^{[18,19, 40]} ...

2013

0.0

... 到目前为止, 关于小麦茎秆实心性状的QTL大多定位在3BL上, 并与标记gwm247、gwm547、gwm340和barc77紧密连锁^{[18,19, 40]} ...

1946

0.0

... 在20世纪40年代, 为了预防麦茎蜂(Cephus cinctus Norton)的危害, 加拿大推广了实秆小麦Rescue和Chinook^[20,21] ...

1948

0.0

... 在20世纪40年代, 为了预防麦茎蜂(Cephus cinctus Norton)的危害, 加拿大推广了实秆小麦Rescue和Chinook^[20,21] ...

1965

0.0

... McNeal等^[22]和Wellace等^[23]提出实秆性状可能对小麦产量造成影响 ...

1966

0.0

... McNeal等^[22]和Wellace等^[23]提出实秆性状可能对小麦产量造成影响 ...

2015

0.0

... Adrianna等^[24]认为实秆麦是到目前为止预防麦茎蜂危害最有效的方法, 其实验表明实秆基因不会影响籽粒品质, 但对产量有影响, 当麦茎蜂数量很少时, 实秆麦的产量显著低于空秆麦, 而当麦茎蜂数量大到足以影响小麦产量时, 实秆麦的产量与空秆麦之间没有显著差别 ...

... 实心度用厚径比来表示, 厚径比为秆壁厚度与茎粗的比值^[24,30] ...

1979

0.0

... McNeal等^[25]以6个实秆小麦与3个普通小麦杂交得到的17个系为材料, 证明实秆基因并不是影响小麦高产的因素 ...

2002

0.0

郭会君. 小麦茎秆强度及其相关性状的QTL分析. 中国农业科学院硕士学位论文, 北京, 2002

QTL Analysis for Stem Strength and Its Correlated Traits in Wheat. MS Thesis of The Chinese Academy of Agricultural Sciences. Beijing, China, 2002

通过重塑茎秆质量改良小麦抗倒性和稳产性,进而提高产量潜力和籽粒品质是实现下一个育种突破的重要途径.该研究采用以H1488和CA9613为亲本构建的DH群体对茎秆强度及其相关性状进行QTLs定位和遗传效应分析.

... 郭会君等^[26]研究发现控制小麦茎秆强度的基因位于2D和5B上, 共同解释24%的表型变异 ...

2005

0.0

... Verm等^[27]利用Milan #cod#x000D7 ...

1999

0.0

廖晓虹, 蒲宗君, 杨武云. 小麦遗传材料86-741与黑麦的可杂交性研究. 云南农业大学报, 1999, 14(1): 37-39

Liao X

, Pu Z

, Yang W

Crossability of common wheat line 86-741 with rye.

J Yunnan Agric Univ, 1999, 14(1): 37-39 (in Chinese with English abstract)

用小麦遗传材料８６－７４１与黑麦杂交，分析其可杂交性，结果表明：８６－７４１与秦岭黑麦，土耳其黑麦２年的杂交结实率超过５０％，推测其可杂交性基因型为ｋｒ１ｋｒ１ｋｒ２ｋｒ２．８６－７４１可作为远缘杂交的桥梁材料，转育小麦近缘属种优良基因，改良和创新小麦种质资源。

... 1 试验材料武云实秆是四川省农业科学院采用实秆小麦86-741^[28]与川麦42 ^[29]杂交、回交育成, 其茎秆基部有极小髓腔, 髓腔中有髓 ...

2006

0.0

... 1 试验材料武云实秆是四川省农业科学院采用实秆小麦86-741^[28]与川麦42 ^[29]杂交、回交育成, 其茎秆基部有极小髓腔, 髓腔中有髓 ...

2006

0.0

... 实心度用厚径比来表示, 厚径比为秆壁厚度与茎粗的比值^[24,30] ...

1994

0.0

... 5软件在全基因组范围对控制小麦茎秆强度及实心度相关性状的QTL进行检测, 采用复合区间作图法进行QTL定位^[31], 当阈值(LOD)大于2 ...

2013

0.0

2012

0.0

冯素伟, 李淦, 胡铁柱, 姜小苓, 李小军, 董娜, 茹振钢. 不同小麦品种茎秆抗倒的研究. 麦类作物学报, 2012, 32: 1055-1059

Feng S

, Li

, Hu T

, Jiang X

, Li X

, Dong

, Ru Z

Study on the stem lodging resistance of different wheat varieties.

J Triticeae Crops, 2012, 32: 1055-1059 (in Chinese with English abstract)

Six high-yield wheat varieties widely applicated in production were selected, to investigate the lodging resistance of different varieties comprehensively,by combining field and laboratory methods.The results showed that the lodging index of Bainong AK 58 was smaller as 0.432, and which was significant difference with others.The research found that the stem mechanical strength of Bainong AK58 was larger, its second internode was shorter and thicker, and its culm wall was thicker, as the secondary root number was more, and its plant height and height of center of gravity were lower than others.Correlation analysis showed that lodging index had close relationship with the characteristics of second stem, the thickness of second section of the mechanical strength, the thickness of culm wall, the dry-matter weight of second stem and the root numbers, which were the main determinants of the lodging resistance.Therefore,reducing the plant height appropriately could reduce the gravity height of plant, and enhance the capability of plant lodging resistance.Stepwise regression analysis showed that the direct factors influencing lodging resistance were the mechanical strength of second stem, the dry-matter weight of second stem, the root numbers and the fresh plant weight of above ground, the most direct action on stem lodging index were the mechanical strength of the second stem with the direct path coefficient of -0.775.So, the stem lodging resistance strength of wheat varieties could be evaluated through the mechanical strength of second stem.

为了明确影响小麦抗倒伏性的主要因素，选用生产中应用较广泛的6个高产小麦品种，采用田间试验与实验室分析相结合的方法，对不同品种的抗倒伏性状进行研究。结果表明，百农矮抗58的倒伏指数较小，为0.432，与其他品种相比差异显著；百农矮抗58的茎秆机械强度大、茎秆基部第二节短粗、秆壁较厚、次生根数量较多、株高和重心高度较低。相关分析表明，倒伏指数与第二节关系密切，第二节机械强度越大、壁越厚、干重越大、地下部根量越多，倒伏指数就越小，茎秆抗倒性就越强；适当降低株高可以降低植株的重心高度，增强植株抗倒能力。逐步回归分析表明，影响茎秆抗倒性的直接因子为茎秆基部第二节机械强度、第二节干重、地下部根量及地上部鲜重，其中第二节机械强度对茎秆倒伏指数的直接作用最大，其直接通径系数为-0.775，可用第二节机械强度的大小评价品种茎秆抗倒性的强弱。

2006

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朱新开, 王祥菊, 郭凯泉, 郭文善, 封超年, 彭永欣. 小麦倒伏的茎秆特征及对产量与品质的影响. 麦类作物学报, 2006, 26: 87-92

Zhu X

, Wang X

, Guo K

, Guo W

, Feng C

, Peng Y

Stem characteristics of wheat with stem lodging and effects of lodging on grain yield and quality.

J Triticeae Crops, 2006, 26: 87-92 (in Chinese with English abstract)

Lodging is one of the key limitatton tactors in hign yield wheat of plants lodged on the 10th day after anthesis and the effects of lodging on grain yield and quality in six varieties were studied in this paper. The results showed that plants with short s

选用6个株高不同的小麦品种，在自然倒伏条件下，研究籽粒形成期倒伏植株的茎秆性状、倒伏对产量和品质的影响。结果表明，矮秆、基部节间较短、重心高度较低、穗下节间较长且株高构成指数大的植株有利于抗倒伏。小麦在籽粒形成期倒伏对产量、品质均有较大的影响，表现为产量下降，籽粒中蛋白质含量、湿面筋含量、沉降值有所上升，但蛋白质产量下降，支链淀粉、直链淀粉和总淀粉含量亦下降。

2003

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2000

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2009

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郭维俊, 王芬娥, 黄高宝, 张锋伟, 魏时来. 小麦茎秆力学性能与化学组分试验. 农业机械学报, 2009, 40: 110-114

Guo W

, Wang F

, Huang G

, Zhang F

, Wei S

Experiment on mechanical properties and chemical compositions of wheat stems.

Trans CSAM, 2009, 40: 110-114 (in Chinese with English abstract)

为分析茎秆的力学性能与主要化学成分之间的相互关系,试验研究了小麦成熟期茎秆的主要力学性能,测定了茎秆不同状态下纤维素、半纤维素、木质素,以及蛋白质、脂肪和糖分等主要化学成分含量.结果表明:小麦茎秆最大拉力为182.38～242.74N,拉伸强度为30.36～52.65MPa,弹性模量为1143.44～1985.86MPa;纤维素、半纤维素和木质素的质量分数分别为22.58%～48.18%、10.85%～22.45%、4.83%～10.59%.小麦茎秆是由纤维素、半纤维素和木质素组成的天然高分子复合材料,呈各向异性,其强度和刚度主要取决于纤维素、半纤维素和木质素的含量及其链接形式和排列方式.

1959

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... 早在1959年就有关于实秆基因的报道, Larson和MacDonald等^[38,39]利用S-615的单体系推断出3B和5A染色体上有控制小麦茎秆实心性状的位点 ...

1959

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... 早在1959年就有关于实秆基因的报道, Larson和MacDonald等^[38,39]利用S-615的单体系推断出3B和5A染色体上有控制小麦茎秆实心性状的位点 ...

2004

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... 到目前为止, 关于小麦茎秆实心性状的QTL大多定位在3BL上, 并与标记gwm247、gwm547、gwm340和barc77紧密连锁^{[18,19, 40]} ...