小麦矮秆突变体的鉴定及其突变性状的关联分析

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贺军与^,, 尹顺琼, 陈云琼, 熊静蕾, 王卫斌, 周鸿斌, 陈梅, 王梦玥, 陈升位^,^*云南农业大学农学与生物技术学院, 云南昆明 650201

Identification of wheat dwarf mutants and analysis on association between the mutant traits of the dwarf plants

HE Jun-Yu^,, YIN Shun-Qiong, CHEN Yun-Qiong, XIONG Jing-Lei, WANG Wei-Bin, ZHOU Hong-Bin, CHEN Mei, WANG Meng-Yue, CHEN Sheng-Wei^,^*Collage of Agronomy and Biotechnology, Yunnan Agricultural University, Kunming 650201, Yunnan, China

通讯作者: *陈升位, E-mail: ynkmcsw@126.com

收稿日期:2020-08-16接受日期:2020-11-13网络出版日期:2021-05-12

基金资助:

国家自然科学基金项目.31660434
国家自然科学基金项目.32060457

Received:2020-08-16Accepted:2020-11-13Online:2021-05-12

Fund supported:

National Natural Science Foundation of China.31660434
National Natural Science Foundation of China.32060457

作者简介 About authors
E-mail: etfiee_coisini@126.com

摘要
矮秆突变体是小麦育种和株高遗传研究的重要基因资源。通过‘云麦53’成熟种子的EMS (Ethyl methyl sulfonate)诱变及诱变植株连续自交, 获得了33个M₃代候选突变体。通过诱变亲本与M₂和M₃代候选植株的株高差异分析, 筛选到26个矮秆突变体, 其株高变幅为(13.61±0.11)~(44.08±1.73) cm。基于8个矮秆基因的12个特异性标记检测发现, 26个矮秆突变体至少携带2个矮秆基因标记位点。除株高外, 26个矮秆突变体还携带穗长、小穗密度、节间数和平均节间长4个不同突变性状。26个矮秆突变体可聚为5个亚类, 第1亚类的小穗数和小花数最少; 第2亚类的株高最矮, 穗长和平均节间长最短, 小穗密度最高; 第3亚类突变体的节间数最少。株高与平均节间长和节间数呈极显著相关, 偏相关系数分别为0.94、0.58, 但与穗长、小穗数、小花数和小穗密度4个性状无相关性。26个矮秆突变体的株高与平均节间长和节间数关联遗传, 携带不同的突变基因组合, 可用于小麦矮化育种, 以及株高、穗长和小穗密度等性状的遗传机制研究。
关键词： 小麦;突变体;矮秆基因;分子标记;遗传关联

Abstract

Dwarf mutant is an important gene resource in wheat breeding and plant height genetic research. In this paper, EMS (ethyl methyl sulfonate) was used to mutate the mature seeds of ‘Yunmai 53’, and 33 candidate dwarf mutants of M₃generation were finally obtained by self-bred. Twenty-six dwarf mutants were selected by analyzing differences between the mutant parent in two years and that of M₂ and M₃ generation candidate plants, and the variation range of their plant height was from (13.61 ± 0.11) cm to (44.08 ± 1.73) cm. Twenty-six dwarf mutants with at least 2 mutant sites were verified based on 12 specific markers of 8 dwarf genes. In addition to plant height, 26 dwarf mutants also carried four mutational traits, spike length, spikelet density, internode number, and average internode length. The 26 dwarf mutants could be clustered into 5 subgroups. Among them, the first subgroup was the least in spikelets and florets, the second subgroup was the shortest in plant height, spike length and average internode length, and the highest spikelet density, while the third subgroup was the least in internode number. Plant height was significantly correlated with average internode length and internode number with partial correlation coefficients of 0.94 and 0.58, respectively, but not correlated with spike length, spikelet number and spikelet density. The plant height was genetically associated with internode length and internode number in 26 dwarf mutants. The mutants carried different combinations of the mutant gene, and can be useful in wheat dwarf breeding and the studies on genetic mechanism of the traits, such as plant height, spike length and spikelet density.

Keywords：wheat;mutant;dwarf genes;molecular marker;genetic association

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本文引用格式
贺军与, 尹顺琼, 陈云琼, 熊静蕾, 王卫斌, 周鸿斌, 陈梅, 王梦玥, 陈升位. 小麦矮秆突变体的鉴定及其突变性状的关联分析[J]. 作物学报, 2021, 47(5): 974-982. doi:10.3724/SP.J.1006.2021.01066
HE Jun-Yu, YIN Shun-Qiong, CHEN Yun-Qiong, XIONG Jing-Lei, WANG Wei-Bin, ZHOU Hong-Bin, CHEN Mei, WANG Meng-Yue, CHEN Sheng-Wei. Identification of wheat dwarf mutants and analysis on association between the mutant traits of the dwarf plants[J]. Acta Agronomica Sinica, 2021, 47(5): 974-982. doi:10.3724/SP.J.1006.2021.01066

自“绿色革命”以来, 全球种植的小麦品种中有90%含有矮秆和半矮秆基因^[1,2,3,4,5]。迄今为止, 已报道的矮秆和半矮秆基因超过25个, 但可用于小麦育种的矮秆基因局限于Rht-B1b、Rht-D1b、Rht8和Rht9^[6,7]。矮秆基因Rht-B1b、Rht-D1b和Rht8在降低小麦株高时对其他性状无显著不良影响, 且Rht-D1b、Rht8和Rht9可显著增加穗粒数^{[8,9,10,11,12]}。李杏普等^[13]发现Rht10和Rht12的降秆作用大于Rht8, 但其株系的地上部分生物量低于Rht8株系。王清海等^[14]研究发现Rht14、Rht16和Rht18的降秆效应分别为47.3%、39.9%和36.0%。携带Rht14、Rht16和Rht18的矮秆株系的小穗小花数变化不大, 但穗长变长。Rht5在降低小麦株高的同时会减少每穗小穗数和穗粒数^[15]。前人研究表明, 大多数小麦推广品种携带的矮秆基因为Rht-B1b、Rht-D1b和Rht8^[3-6,16-17], 可有效利用的矮秆亲本不足, 育成品种遗传背景狭窄。创制新突变体是解决该问题的有效策略。本文采用甲基磺酸乙酯(Ethyl methyl sulfonate, EMS)诱变‘云麦53’ (国审品种)成熟种子, 通过诱变植株的连续3年套袋自交成功获得了一批株高、小穗密度和小花数等性状不同的候选突变体。基于连续2年的田间观察和常见矮秆基因的12个特异性分子标记检测鉴定了26株矮秆突变株, 分析了26个突变体的株高差异及其与穗长、小穗数和小穗密度等农艺性状的遗传关联性, 评估了矮秆突变体的育种价值。

1 材料与方法

1.1 试验材料

诱变亲本‘云麦53’由云南省农业科学研究院粮食作物研究所提供, 突变体由课题组创制。所有材料由课题组繁育并保存。

1.2 试验材料的种植及取样

1.2.1 材料诱变和繁育将15,000粒‘云麦53’种子置于三角瓶中, 用pH 8.0的0.05 mol L^-1磷酸缓冲液(PBS)震荡浸泡4 h, 再用0.6% EMS室温震荡浸泡16 h, 然后用自来水冲洗4 h。诱变种子及其后代均播种于云南农业大学昆明市盘龙区试验教学农场。分别套袋自交M₁、M₂和M₃候选突变材料, 按单株收获种子。

1.2.2 农艺性状调查按随机排列种植试验材料, 每个材料种植1行, 5行后种植1行诱变亲本。行距20cm, 株距5 cm, 3次重复。其他田间管理措施与常规大田管理一致。于2018—2019年度和2019—2020年度分别在开花期调查穗长、小穗数和小花数, 待种子成熟后调查株高、平均节间长和节间数。M₂代调查单株性状, M₃株系随机调查5株, 以5株均值代表株系性状的表型值。按下列公式计算小穗密度、平均节间长。

小穗密度=单株小穗数/单株穗长

平均节间长=单株节间长/单株节间数

1.2.3 矮秆基因的分子标记检测于拔节期剪取M₂单株及其诱变亲本的幼嫩叶片, 置于-80℃保存备用。采用CTAB法提取基因组DNA。采用Ellis等^[18,19]报道的BF-WR1、BF-MR1、BF-WR2、BF-MR2、WMC317、BARC102、BARC151、WMC410和WMS577, 以及XBARC3^[20]、Xgwm261^[20]和WMC503^[21]特异性标记检测突变体携带的矮秆基因(表1), 其中BF-WR1和BF2-WR2为野生型(株高正常)基因标记, 其余标记为突变型(矮秆)基因标记。由上海生工生物技术有限公司合成所有引物。参照前人报道的PCR扩增条件扩增各标记, 采用8%聚丙烯酰胺凝胶电泳检测PCR扩增产物。上述标记均重复检测3次, 3次检测结果一致。

Table 1
表1
表1矮秆基因及其特异性标记的引物序列和片段长度
Table 1Specific molecular markers of dwarf genes and the fragment length and primers sequence of the markers

基因 Gene	标记 Marker	正向引物 Forward sequence (5′-3′)	反向引物 Reverse sequence (5′-3′)	片段长度 Product size (bp)
Rht-B1a	BF-WR1	GGTAGGGAGGCGAGAGGCGAG	CATCCCCATGGCCATCTCGAGCTG	237
Rht-B1b	BF-MR1	GGTAGGGAGGCGAGAGGCGAG	CATCCCCATGGCCATCTCGAGCTA	237
Rht-D1a	BF2-WR2	GGCAAGCAAAAGCTTCGCG	GGCCATCTCGAGCTGCAC	264
Rht-D1b	BF-MR2	CGCGCAATTATTGGCCAGAGATAG	CCCCATGGCCATCTCGAGCTGCTA	254
Rht4	WMC317	TGCTAGCAATGCTCCGGGTAAC	TCACGAAACCTTTTCCTCCTCC	170
Rht5	BARC102	GGAGAGGACCTGCTAAAATCGAAGACA	GCGTTTACGGATCAGTGTTGGAGA	200
Rht8	Xgwm261	CTCCCTGTACGCCTAAGGC	CTCGCGCTACTAGCCATTG	192
Rht8	WMC503	GCAATAGTTCCCGCAAGAAAAG	ATCAACTACCTCCAGATCCCGT	225
Rht9	BARC151	TGAGGAAAATGTCTCTATAGCATCC	CGCATAAACACCTTCGCTCTTCCACTC	220
Rht12	WMC410	GGACTTGAAAGGAAGCTTGTGA	CATGGATGGCATGCAGTGT	114
Rht13	WMS577	ATGGCATAATTTGGTGAAATTG	TGTTTCAAGCCCAACTTCTATT	130
Rht14/16/18	XBARC3	TTCCCTGTGTCTTTCTAATTTTTTTT	GCGAACTCCCGAACATTTTTAT	210

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1.2.4 农艺性状的关联分析基于SPSS 25.0软件的t测验、聚类、方差、多重比较和相关分析模块默认参数, 分析突变体株高差异及其与穗长、小穗数和小穗密度等性状的遗传关联性。显著和极显著水平分别为5%和1%。

2 结果与分析

2.1 候选矮秆突变体的农艺性状差异分析

本研究采用t测验检测了‘云麦53’与33个候选材料的株高(M₂单株表型值, M₃株系均值)差异(表2)。结果表明, mutant 2、mutant 4、mutant 5等26个材料与‘云麦53’的株高存在显著或极显著差异, 为矮秆突变体。在26个矮秆突变体中mutant 16、mutant 23、mutant 25和mutant 26的株高值低于20 cm, mutant 18、mutant 19和mutant 20的株高值大于40 cm, 其余19个突变体的株高值介于20~40 cm之间(图1)。与‘云麦53’相比, mutant 4、mutant 5和mutant 7等18个突变体的穗长显著或极显著缩短; mutant 4、mutant 5和mutant 7等11个突变体的小穗密度显著或极显著增加; mutant 18的小花数显著减少; mutant 5、mutant 11、mutant 14等13个突变体的平均节间长显著或极显著缩短; mutant 5、mutant 10和mutant 14等6个突变体的节间数显著或极显著减少(表2和图2)。mutant 7、mutant 12、mutant 24、mutant 28和mutant 29的节间数在M₂和M₃代中未发生分离, 均低于‘云麦53’的节间数, 但因方差为0、未做t测验。26个矮秆突变体携带株高、穗长和小穗密度等多个突变性状。

图1

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图1‘云麦53’与部分矮秆突变体的株高差异

A: 突变体31; B: 突变体17; C: 突变体10; D: 突变体16; E: 突变体23; F: 突变体25。
Fig. 1Differences of plant height between ‘Yunmai 53’ and other dwarf mutants

A: mutant 31; B: mutant 17; C: mutant 10; D: mutant 16; E: mutant 23; F: mutant 25.

Table 2
表2
表2‘云麦53’及候选突变体的7个农艺性状均值及其显著性(t测验)
Table 2Mean values of 7 agronomic traits of ‘Yunmai 53’ and candidate mutants, and the difference between the mean values (t-test)

材料 Material	株高 Plant height (cm)	穗长 Spike length (cm)	每穗小穗数 Spikelets number per spike	小穗密度 Spikelet density	小花数 Number of florets	平均节间长 Average internode length (cm)	节间数 Number of internodes
云麦53 Yunmai 53	74.89±4.03	10.30±0.31	19.25±1.21	1.88±0.18	59.35±5.23	11.10±1.04	6.00±0.00
mutant 1	56.95±6.95	10.49±1.52	20.50±3.50	2.05±0.63	60.34±9.34	7.50±3.33	5.00±0.00
mutant 2	39.52±2.02^*	8.40±1.10	17.30±3.30	2.15±0.68	58.20±16.20	7.98±0.78	4.50±0.50
mutant 3	63.99±2.51	6.11±0.59^*	17.10±1.10	2.84±0.46	51.70±3.70	11.23±0.96	5.40±0.60
mutant 4	37.24±0.44^*	4.44±0.34^**	19.30±1.30	4.35±0.04^**	55.00±1.00	7.21±0.61	4.50±0.50
mutant 5	21.94±3.66^*	3.72±0.02^**	20.00±1.00	5.37±0.24^**	58.30±1.30	4.88±0.62^*	3.80±0.20^**
mutant 6	39.85±4.39^*	6.01±0.45^*	19.10±2.10	3.73±1.27	55.50±4.50	6.87±0.44	4.50±0.50
mutant 7	33.89±2.89^*	4.71±0.41^**	21.00±2.00	4.46±0.04^**	63.40±6.40	7.22±0.39	4.00±0.00
mutant 8	34.92±2.58^*	4.40±0.10^**	22.50±0.50	5.12±0.01^**	67.50±1.50	7.16±0.44	4.40±0.60
mutant 9	31.81±6.79^*	4.18±0.58^*	21.30±2.30	5.12±0.16^**	61.20±4.20	6.20±0.95	4.40±0.60
mutant 10	25.55±7.85^*	4.24±0.35^**	19.40±1.40	4.62±0.71	54.70±0.70	5.44±1.77	3.90±0.10^**
mutant 11	29.78±1.92^*	8.20±0.70	16.10±0.90	1.97±0.06	48.00±3.00	5.49±0.24^*	4.50±0.50
mutant 12	38.79±2.61^*	4.63±0.13^**	19.30±2.30	4.16±0.38^*	54.40±3.40	7.30±0.08	5.00±0.00
mutant 13	38.32±8.48	3.98±0.38^**	15.90±1.90	3.98±0.10^**	44.90±2.90	6.65±0.85	4.50±0.50
mutant 14	33.18±6.02^*	4.36±0.24^**	18.80±0.80	4.34±0.43^*	53.00±1.00	5.95±0.03^*	3.90±0.01^**
mutant 15	30.94±2.26^*	4.00±0.50^**	17.90±0.10	4.55±0.59^*	49.50±4.50	5.68±0.24^*	4.80±0.20^*
mutant 16	13.61±0.11^**	3.39±0.11^**	18.40±1.40	5.45±0.59^*	45.70±5.30	2.26±0.63^*	4.80±1.20
mutant 17	32.62±4.98^*	3.96±0.26^**	17.50±0.50	4.43±0.16^**	49.10±1.90	6.16±0.62	4.50±0.50
mutant 18	44.08±1.73^*	7.08±0.48^*	13.50±2.50	1.90±0.23	33.25±0.25^*	7.87±0.97	5.50±0.50
mutant 19	42.41±12.19	10.58±1.52	21.40±2.40	2.10±0.53	69.10±12.10	6.10±1.01	5.30±0.70
mutant 20	42.18±6.58	7.97±0.63	15.30±0.30	1.94±0.20	43.10±1.90	7.31±1.89	4.80±0.20^*
mutant 21	31.26±4.44^*	4.02±0.48^**	15.60±1.60	3.99±0.88	40.40±1.60	7.04±0.79	3.80±0.20^**
mutant 22	52.28±5.48	8.45±0.95	19.50±0.50	2.34±0.33	56.30±0.70	8.44±0.94	5.50±0.50
材料 Material	株高 Plant height (cm)	穗长 Spike length (cm)	每穗小穗数 Spikelets number per spike	小穗密度 Spikelet density	小花数 Number of florets	平均节间长 Average internode length (cm)	节间数 Number of internodes
mutant 23	15.80±0.60^**	3.70±0.35^**	19.80±1.80	5.28±0.01^*	58.80±4.80	2.29±0.41^*	4.40±0.60
mutant 24	30.07±4.43^*	6.76±1.04	16.90±2.90	2.63±0.84	49.70±7.70	5.26±0.05^*	5.00±0.00
mutant 25	18.16±4.44^*	3.77±0.17^**	20.20±0.20	5.37±0.19^**	58.80±1.20	3.93±0.93^*	4.50±0.50
mutant 26	19.75±1.35^**	4.60±1.00^*	18.60±1.60	4.17±0.56	53.20±2.20	4.16±0.46^*	3.80±0.20^**
mutant 27	34.54±4.74^*	7.49±0.89	14.80±3.80	1.95±0.28	50.00±17.00	7.13±1.40	4.50±0.50
mutant 28	33.76±10.16	8.86±0.16	15.90±1.90	1.79±0.18	53.90±11.90	6.59±3.49	5.00±0.00
mutant 29	35.01±4.79^*	8.20±0.40	16.40±0.40	2.01±0.15	49.30±1.30	6.18±0.02^*	5.00±0.00
mutant 30	30.09±6.01^*	8.57±1.13	16.10±1.10	1.93±0.38	48.60±3.60	5.92±0.00^*	4.30±0.70
mutant 31	33.96±0.36^*	8.70±1.00	17.20±0.20	2.01±0.26	49.20±1.80	5.60±0.94	4.50±0.50
mutant 32	31.50±0.70^**	8.02±0.28^*	16.50±1.50	2.07±0.26	46.30±1.30	5.11±0.53^*	4.50±0.50
mutant 33	24.02±3.38^*	10.04±1.94	16.63±1.63	1.69±0.17	47.75±2.75	3.95±0.12^*	5.00±1.00

^*和^**分别代表0.05和0.01显著水平。
^* and ^** represent significant differences at the 0.05 and 0.01 probability levels, respectively.

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图2

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图2部分矮秆突变体及其诱变亲本的穗部形态特征

(a): ‘云麦53’; (b), (c), (d)和(e): 候选矮秆突变体。
Fig. 2Spike characteristic of some dwarf mutants and their parents

(a): ‘Yunmai 53’; (b), (c), (d), and (e): candidate dwarf mutants.

2.2 候选矮秆突变体的分子标记检测

为了进一步验证26个矮秆材料, 本文采用前人报道的矮秆基因的12个特异性标记检测了33个候选材料及其诱变亲本。结果表明, 在‘云麦53’中未检测到BF-MR1 (Rht-B1b)、BF2-WR2 (Rht-D1a)、BARC102 (Rht5)和WMS577 (Rht13)标记, 但检测到其他9个标记。与‘云麦53’相比, mutant 1等32个候选突变体的BARC151 (Rht9)标记, mutant 2等5个候选突变体的Xgwm261 (Rht8)、WMC503 (Rht8)标记均具有差异, 表明其Rht9和Rht8基因发生了突变。在33个候选突变体中, mutant 3与‘云麦53’的共有扩增标记较多(7个), mutant 11与‘云麦53’仅有2个共同扩增标记(表3和图3), mutant 1、mutant 2、mutant 3、mutant 27和mutant 28的Xgwm261 (Rht8)、WMC503 (Rht8)和BARC151 (Rht9)均发生变异, 其中BARC151 (Rht9)标记位点的突变频率最高(表3)。

Table 3
表3
表3‘云麦53’和候选突变体中检测到的矮秆基因
Table 3Dwarf genes detected in the genome of ‘Yunmai 53’ and candidate mutants

材料 Material	检测到的基因 Genes detected by 13 markers
云麦53 Yunmai 53	Rht-B1a, Rht-D1b, Rht4, Rht8, Rht9, Rht12, Rht14, Rht16, Rht18
mutant 1	Rht-B1a, Rht-D1a, Rht4, Rht5, Rht8^#, Rht9^#, Rht13, Rht14, Rht16, Rht18
mutant 2	Rht-B1a, Rht-D1b, Rht4, Rht5, Rht8^#, Rht9^#, Rht13, Rht14, Rht16, Rht18
mutant 3	Rht-B1a, Rht-D1b, Rht4, Rht8^#, Rht9^#, Rht12, Rht14, Rht16, Rht18
mutant 10	Rht-B1a, Rht-D1a, Rht4, Rht8, Rht9^#
mutant 11	Rht-B1a, Rht-D1a, Rht5, Rht12
mutant 19	Rht-B1a, Rht-D1a, Rht4, Rht5, Rht8, Rht9^#, Rht14, Rht16, Rht18
mutant 26	Rht-B1a, Rht-D1b, Rht4, Rht8, Rht9, Rht13
mutant 27 & mutant 28	Rht-B1a, Rht-D1b, Rht4, Rht8^#, Rht9^#, Rht13
Other mutants	Rht-B1a, Rht-D1a, Rht4, Rht5, Rht8, Rht9^#

“^#”:多态性标记; Rht8两个标记(Xgwm261、WMC503)在试验材料间的多态性一致。
“^#”: polymorphism markers in the fragment. The polymorphism of Xgwm261 and WMC503 of Rht8 show consistent among the experimental materials.

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图3

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图3‘云麦53’和候选突变体的部分特异性扩增标记

CK: 对照; M: marker; 53: 云麦53; 1~33: 突变体1~突变体33; A和B分别是‘云麦53’和候选突变体中扩增的WMC503和BARC151标记; 红色箭: 差异片段。
Fig. 3Specific markers amplified from the genomes of ‘Yunmai 53’ and candidate mutants

CK: control; M: marker; 53: Yunmai 53; 1-33: mutant 1-mutant 33; A-B were WMC503 and BARC151 markers amplified in ‘Yunmai 53’ and candidate mutants, respectively. The red arrows indicate differential fragments.

2.3 基于7个性状的矮秆突变体聚类分析

基于株高、穗长和小穗密度等7个性状的聚类分析结果表明, 所有材料可聚为2个大类, 第1大类包括诱变亲本, 第2大类包括所有突变体(图4)。第2大类可分为5个亚类, 其中mutant 18和mutant 16分别聚为第1、2亚类, mutant 5、mutant 10和mutant 23等5个突变体为第3亚类, mutant 2、mutant 4和mutant 6等7个突变体为第4亚类, 其余12个突变体为第5亚类(图4)。与‘云麦53’相比, 5个亚类的突变体均表现植株矮化、穗长和平均节间长缩短, 但其余4个性状出现了分化。其中第1亚类突变体的小穗数和小花数极显著减少, 但小穗密度和节间数没有差异; 第2~4亚类突变体的小穗密度极显著增加, 小花数和节间数极显著减少, 但小穗数没有明显变化(图5)。在5个亚类中, 第1亚类突变体的小穗数和小花数降幅最大; 第2亚类的株高、穗长和平均节间长的降幅和小穗密度的增幅最大, 第3亚类突变体的节间数降幅最大(图5和附表1)。

图4

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图4基于7个农艺性状的26个突变体和诱变亲本聚类结果

Fig. 4Cluster result of 26 mutants and their parent based on seven agronomic traits

除株高外, 26个矮秆突变体还携带穗长缩短、小花密度增加和节间数减少等不同突变性状。通过分子检测本研

图5

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图5不同类群材料的农艺性状均值及其差异

I: ‘云麦53’类群; II~VI: 突变体的第1亚类~第5亚类。不同小写字母和大写字母分别代表0.05和0.01显著水平。
Fig. 5Mean values and differences of agronomic traits in different groups materials

I: group ‘Yunmai 53’; II-VI: the subclass 1-subclass 5 of mutants. Different lowercase letters and capital letters represent significant differences at the 0.05 and 0.01 probability levels, respectively.

Table S1
附表1
附表1‘云麦53’类群和5个矮秆突变体亚类的方差分析结果
Table S1ANOVA result of ‘Yunmai 53’ and 5 subclasses dwarf mutants

性状 Traits	变异来源 SOV	平方和 SS	自由度 df	均方 MS	F值 F-value
株高Plant height (cm)	组间Between groups	11,981.952	5	2396.390	37.615^**
株高Plant height (cm)	组内Within group	4077.292	64	63.708
穗长Spike length (cm)	组间Between groups	119.531	5	23.906	4.603^**
穗长Spike length (cm)	组内Within group	332.417	64	5.194
小穗数Spikelets number per spike	组间Between groups	234.415	5	46.883	10.229^**
小穗数Spikelets number per spike	组内Within group	293.328	64	4.583
小穗密度Spikelet density	组间Between groups	48.716	5	9.743	6.920^**
小穗密度Spikelet density	组内Within group	90.110	64	1.408
小花数Number of florets	组间Between groups	3126.921	5	625.384	11.289^**
小花数Number of florets	组内Within group	3545.316	64	55.396
平均节间长Average internode length (cm)	组间Between groups	190.020	5	38.004	13.307^**
平均节间长Average internode length (cm)	组内Within group	182.776	64	2.865
节间数Number of Internode	组间Between groups	14.628	5	2.926	7.604^**
节间数Number of Internode	组内Within group	24.623	64	0.385

^**表示在0.01水平差异显著。
^**mean significant difference at the 0.01 probability level.

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2.4 突变农艺性状间的相关分析

基于26个突变体及其诱变亲本的株高等7个性状的二元相关和偏相关分析表明, 株高与穗长、小穗密度和平均节间长等4个性状的二元相关系数达到了显著或极显著水平, 但只有平均节间长和节间数与株高偏相关; 穗长与小穗数、小穗密度和平均节间长的二元相关系数达到了显著或极显著水平, 但只有小穗密度和平均节间长与穗长存在负偏相关; 小穗数与小穗密度和小花数的二元相关和偏相关系数均达到了极显著水平; 小穗密度与小花数和节间数的二元相关系数达到了显著或极显著水平, 但偏相关系数不显著; 虽然平均节间长与节间数的二元相关系数不显著, 但偏相关系数达到了显著水平(表4)。上述结果表明, 26个突变体的株高与平均节间长和节间数关联遗传, 其余4个性状与植株矮化没有遗传关联性。

Table 4
表4
表4不同突变性状的二元相关和偏相关系数及其显著性
Table 4Binary correlation, partial correlation coefficients and significances among the different mutant traits

性状 Trait	株高 Plant height (cm)	穗长 Spike length (cm)	小穗数 Spikelets number per spike	小穗密度 Spikelet density	小花数 Number of florets	平均节间长 Average internode length (cm)	节间数 Number of internode
株高 Plant height (cm)		0.51^**	-0.09	-0.47^*	0.06	0.92^**	0.59^**
穗长 Spike length (cm)	0.26		-0.54^*	-0.95^**	-0.25	0.36	0.53^**
小穗数 Spikelets number per spike	0.26	0.41		0.74^**	0.89^**	-0.07	-0.27
小穗密度 Spikelet density	-0.11	-0.95^**	0.64^**		0.47^*	-0.38	-0.48^**
小花数 Number of florets	-0.32	0.13	0.78^**	-0.09		0.10	-0.23
平均节间长 Average internode length (cm)	0.94^**	-0.43^*	-0.16	-0.29	0.35		0.35
节间数 Number of internode	0.58^**	0.16	-0.10	0.16	-0.08	-0.44^*

^*和^**分别代表0.05和0.01显著水平。
^* and ^** represent significant differences at the 0.05 and 0.01 probability levels, respectively.

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3 讨论

通过诱变亲本和候选突变体连续2个年度的株高比对分析, 作者发现了26个矮秆突变体。基于前人报道的矮秆基因的12个特异性标记检测结果表明26个矮化材料至少携带有2个矮秆基因标记位点。虽然mutant 1、mutant 3和mutant 13等7个候选突变体中也检测到了矮秆基因突变, 但植株偏高, 为(33.76±10.16)~(63.99 ±2.51) cm, 且与诱变亲本‘云麦53’的株高没有显著差异。笔者认为这可能是2个年份间的株高波动降低了“t”值所致。在33个候选突变体中, 26个材料明显矮化、且携带不同的矮秆突变基因, 因此可判定为矮秆突变体。究发现26个突变体均携带与‘云麦53’不同的矮秆基因组合, 如mutant 11携带Rht5和Rht 12矮秆基因(可能还携带其他未被检测的矮秆基因), mutant 2携带Rht-D1b、Rht4、Rht5等9个矮秆基因。通过推广品种株高与穗长、小穗数和穗粒数的关联分析, 陈亮等^{[8,9,10,11,12]}发现矮秆突变基因Rht-D1b、Rht8和Rht9可增加穗粒数; 王清海等^[13,14]研究发现Rht8、Rht10、Rht12、Rht14、Rht16、Rht18等矮秆基因的效应不同, Rht10和Rht12的降秆效应大于Rht8的^[3,5]; Baloch等^[16]发现Rht14、Rht16和Rht18可增加穗长, Rht5可减少每穗小穗数。前人研究结果表明小麦矮秆基因对穗长、小穗数和穗粒数等性状可能具有多效性, 但在不同遗传背景中矮秆基因的遗传效应不同。因此作者认为是矮秆基因与穗长、小穗密度和小花数等性状的突变基因及其组合不同导致了26个突变体的农艺性状差异。本研究发现株高与平均节间长和节间数、以及穗长与小穗数、小穗数与小穗密度、小穗与小花数等性状关联遗传, 该结果与前人报道的基本一致^{[22,23,24,25,26,27,28,29,30,31,32]}。虽然株高降低后mutant 7等3个突变体的小穗数和小花数增多(均未达到显著水平), 其余突变体的穗长缩短、小穗数和小花数减少、小穗密度增加, 但株高与穗长等4个性状的偏相关系数均未达到显著水平。作者认为该结果可能与本文研究所用材料大多为相同亲本诱导的矮秆突变体有关, 可能是矮秆基因及其组合的多效性较低或者矮秆与其他性状的突变基因位点的连锁强度较低导致了株高与穗长、小穗数和小穗密度等性状不能关联遗传。通过进一步自交繁育, 可望从26份矮秆突变体中筛选到适合不同育种需求的矮秆亲本。通过聚类分析, 作者发现26个矮秆突变体可分为农艺特性不同的5个亚类, 不同亚类间株高、穗长和小穗数等农艺性状存在差异, 其中第2亚类和第3亚类的mutant 5、mutant 23和mutant 25等5个突变体株高不到20 cm, 均低于陆燕等的报道^{[12,17,33-35]}。虽然株高较低, 但mutant 11、mutant 27和mutant 28等突变体的穗长、小穗数和小花数没有变化, 且节间数只是导致部分突变体矮化的因素之一, 因此上述材料可用于小麦矮化育种。除此之外, 26个突变体携带的株高、穗长和小穗密度等突变性状基因及其组合类型不同, 如mutant 16携带株高较低(13.61 cm±0.11 cm), 穗长(3.72 cm±0.02 cm)和平均节间长(4.88 cm±0.62 cm)较短, 小穗密度(5.37±0.24)较高的基因组合; mutant 18具有植株偏高(44.08 cm±1.73 cm), 穗长偏长(7.08 cm±0.48 cm)小花数较少(33.25±0.25)的基因组合。基于上述突变体, 可望解析小麦株高及其构成因素、小穗密度和小花数等性状的遗传机制。

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