青稞抗倒伏性状的基因型差异

doi:10.3864/j.issn.0578-1752.2019.02.004

摘要/Abstract

摘要：

【目的】倒伏是影响青稞生产和产量的主要原因之一。筛选与青稞抗倒伏性密切相关的性状并构建抗倒伏评价体系,是开展青稞抗倒伏品种选育的重要理论依据。【方法】通过对35份青稞种质资源根系、茎秆和穗部23个农艺性状进行方差分析,明确不同基因型间各性状的差异性。采用相关分析筛选出与青稞抗倒伏性密切相关的指标,并通过主成分分析和线性回归分析构建完成青稞抗倒伏评价体系。【结果】不同基因型青稞材料在同一性状间表现出较大差异,且农艺性状在基因型间的差异均极显著,农艺性状中倒伏率的遗传变异最丰富;同一参试材料的农艺性状在两个生态区间差异较大,海北高寒生态区试点各基因型的遗传变异较丰富;各性状的基因型与环境因素间存在显著互作效应（P<0.05）。相关分析结果表明茎秆强度同青稞抗倒伏性关系最为密切,并通过抗倒伏指标构建青稞抗倒伏评价体系。青稞分蘖数过多、第三和第四茎节过长易引起植株倒伏。根干重大、茎秆重、茎秆壁厚、茎秆强度大是植株固持能力强、抗倒伏伏性优异的原因。【结论】分蘖数、穗重、茎长、茎重、茎秆强度适合作为青稞抗倒伏性评价指标,验证结果表明抗倒伏评价体系较可靠,可用于青稞种质的抗倒伏性评价。

关键词: 青稞, 倒伏, 主成分分析, 茎秆强度, 评价体系

Abstract:

【Objective】 Lodging has become one of the main factors affecting the production and yield of hulless barley. Screening the traits closely related to the lodging resistance and constructing the lodging resistance evaluation system is an important theoretical basis for the breeding of hulless barley varieties. 【Method】 23 agronomic traits in roots, stems, and ears related to the lodging of 35 hulless barley germplasms were analyzed by statistical method. The variance analysis was carried out to identify the differences among different traits, and correlation analysis was used to screen out the traits which closely related to the lodging resistance. The indexes were finally constructed by principal component analysis and linear stepwise regression analysis to construct a hulless barley resistance evaluation system. 【Result】 The results showed that different genotypes of barley cultivars had larger differences among the same traits, and the differences of genotypes among the phenotypic traits were extremely significant, among this, the genetic variation of lodging rate was the most abundant. The agronomic traits of the same genotypes were greatly affected between different ecoregion, and the genetic variation of the genotypes in Haibei alpine farming-pastoral ecotone was abundant. There were significant interaction effects between genotypes of various traits and environmental factors (P<0.05). Correlation analysis showed that the stem strength was most closely related to the lodging resistance of the hulless barley, which was used lodging resistance index to construct the lodging resistance evaluation system. When the plant had more tiller numbers, longer third and fourth stem segments, which caused the lodging, and made the number of grains reduced, and the ear weight became lighter. When the root dry weight was heavier, the stem was heavier, the wall thickness was thicker, and the stem strength was larger, which made the plants have the stronger ability to retain and lodging resistance. 【Conclusion】 Based on tiller numbers, ear weight, stem length, stem weight and stem strength, a comprehensive evaluation system for the resistance indexes of barley was constructed. The results showed that the system was reliable and could be used for the evaluation of lodging resistance of hulless barley germplasms.

Key words: hulless barley, lodging, principal component analysis, stem strength, evaluation system

白羿雄,姚晓华,姚有华,吴昆仑. 青稞抗倒伏性状的基因型差异[J]. 中国农业科学, 2019, 52(2): 228-238.

BAI YiXiong,YAO XiaoHua,YAO YouHua,WU KunLun. Difference of Traits Relating to Lodging Resistance in Hulless Barley Genotypes[J]. Scientia Agricultura Sinica, 2019, 52(2): 228-238.

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图/表 8

表1

表2

参试青稞基因型各茎节农艺性状"

指标（西宁点） Index (Xining point)	第二节 S					第三节 T					第四节 F
指标（西宁点） Index (Xining point)	茎长 PL (cm)	壁厚 WT (mm)	茎粗 ST (mm)	茎重 SW (g)	秆强 SS (N)	茎长 PL (cm)	壁厚 WT (mm)	茎粗 ST (mm)	茎重 SW (g)	秆强 SS (N)	茎长 PL (cm)	壁厚 WT (mm)	茎粗 ST (mm)	茎重 SW (g)	秆强 SS (N)
最小值 Minimum	8.19	0.63	3.51	0.16	4.86	10.78	0.62	3.87	0.18	3.71	11.08	3.49	0.17	0.56	3.99
最大值 Maximum	14.47	1.41	6.01	0.64	22.07	20.15	1.07	5.81	0.66	16.95	26.52	6.38	0.62	1.11	15.46
均值 Mean	11.09	0.95	4.65	0.29	11.86	15.04	0.82	4.83	0.32	9.86	17.74	4.58	0.31	0.74	8.18
标准差 Standard deviation	1.65	0.17	0.65	0.10	4.54	2.22	0.11	0.54	0.09	3.17	3.73	0.60	0.10	0.10	2.62
方差 Variance	2.71	0.03	0.43	0.01	20.57	4.94	0.01	0.29	0.01	10.06	13.94	0.36	0.01	0.01	6.88
偏度 Skewness	0.05	0.66	0.21	1.26	0.47	0.44	0.38	0.22	1.37	0.40	0.65	0.57	1.26	1.17	0.65
峰度 Kurtosis	-0.98	0.21	-0.81	2.05	-0.44	-0.41	-0.48	-0.78	3.34	-0.62	-0.13	0.74	1.23	2.11	0.05
变异系数 CV (%)	14.85	18.38	14.08	34.61	38.24	14.79	13.36	11.11	28.92	32.15	21.05	13.11	32.63	13.98	32.07
指标（海北点） Index (Haibei point)	第二节 S					第三节 T					第四节 F
指标（海北点） Index (Haibei point)	茎长 PL (cm)	壁厚 WT (mm)	茎粗 ST (mm)	茎重 SW (g)	秆强 SS (N)	茎长 PL (cm)	壁厚 WT (mm)	茎粗 ST (mm)	茎重 SW (g)	秆强 SS (N)	茎长 PL (cm)	壁厚 WT (mm)	茎粗 ST (mm)	茎重 SW (g)	秆强 SS (N)
最小值 Minimum	5.80	0.15	1.05	0.06	2.09	7.10	0.15	1.08	0.11	2.09	6.10	1.07	0.15	0.02	1.23
最大值 Maximum	23.50	1.94	4.99	0.85	11.73	31.50	1.94	4.98	1.00	11.73	32.50	5.81	1.93	0.89	13.11
均值 Mean	12.03	0.73	2.75	0.32	5.35	16.18	0.73	2.82	0.37	5.35	19.32	2.75	0.74	0.36	4.11
标准差 Standard deviation	3.13	0.45	1.20	0.15	1.75	3.57	0.45	1.23	0.18	1.75	5.62	1.26	0.48	0.16	1.86
方差 Variance	9.80	0.20	1.44	0.02	3.07	12.75	0.20	1.51	0.03	3.07	31.54	1.60	0.23	0.03	3.46
偏度 Skewness	0.40	0.71	0.24	1.13	0.62	0.84	0.71	0.16	0.88	0.62	0.15	0.34	0.61	0.71	1.69
峰度 Kurtosis	0.36	-0.34	-1.45	1.51	0.47	2.49	-0.34	-1.49	0.46	0.47	-0.19	-1.23	-0.58	0.15	5.40
变异系数 CV (%)	26.02	62.12	43.53	47.70	32.77	22.08	62.12	43.59	47.28	32.77	29.07	45.97	64.88	45.51	45.27

表2

表3

表4

表5

性状间相关分析"

	X1	X2	X3	X4	X5	X6	X7	X8	X9	X10	X11	X12	X13	X14	X15	X16	X17	X18	X19	X20	X21	X22	X23
X1	1
X2	-0.020	1
X3	-0.207^**	0.591^**	1
X4	0.043	0.062	0.023	1
X5	0.183^**	0.431^**	0.358^**	0.061	1
X6	-0.039	0.063	0.113	0.561^**	0.123	1
X7	0.046	-0.111	-0.157^*	0.089	-0.088	0.058	1
X8	-0.019	-0.212^**	-0.381^**	0.167^**	-0.531^**	-0.146^*	0.181^**	1
X9	-0.123	0.488^**	0.628^**	0.062	0.542^**	0.232^**	-0.151^*	-0.551^**	1
X10	-0.228^**	-0.095	-0.035	0.220^**	-0.258^**	0.162^*	0.381^**	0.293^**	0.004	1
X11	-0.537^**	0.467^**	0.591^**	-0.052	0.237^**	0.099	-0.238^**	-0.404^**	0.649^**	0.083	1
X12	0.212^**	-0.088	-0.259^**	0.173^**	-0.016	0.065	0.427^**	0.074	-0.195^**	0.155^*	-0.219^**	1
X13	-0.010	-0.248^**	-0.413^**	0.180^**	-0.551^**	-0.151^*	0.166^**	0.968^**	-0.592^**	0.253^**	-0.440^**	0.084	1
X14	-0.112	0.496^**	0.621^**	0.076	0.562^**	0.230^**	-0.150^*	-0.579^**	0.985^**	-0.023	0.634^**	-0.203^**	-0.612^**	1
X15	-0.261^**	-0.145^*	-0.112	0.346^**	-0.330^**	0.125	0.226^**	0.356^**	-0.087	0.771^**	0.029	0.159^*	0.366^**	-0.114	1
X16	-0.612^**	0.348^**	0.569^**	0.015	0.252^**	0.192^**	-0.132^*	-0.401^**	0.719^**	0.180^**	0.848^**	-0.235^**	-0.435^**	0.707^**	0.099	1
X17	0.247^**	-0.039	-0.236^**	0.217^**	-0.024	0.101	0.218^**	0.134^*	-0.160^*	0.236^**	-0.146^*	0.535^**	0.116	-0.166^**	0.281^**	-0.228^**	1
X18	-0.176^**	0.402^**	0.468^**	0.235^**	0.245^**	0.221^**	-0.077	0.006	0.776^**	0.221^**	0.470^**	-0.180^**	-0.033	0.766^**	0.182^**	0.583^**	-0.138^*	1
X19	-0.041	-0.090	-0.252^**	0.224^**	-0.401^**	-0.156^*	0.109	0.825^**	-0.344^**	0.249^**	-0.308^**	0.019	0.835^**	-0.361^**	0.357^**	-0.296^**	0.046	0.166^**	1
X20	-0.230^**	0.355^**	0.552^**	0.153^*	0.380^**	0.243^**	-0.081	-0.412^**	0.851^**	0.246^**	0.607^**	-0.131^*	-0.434^**	0.836^**	0.247^**	0.705^**	-0.044	0.738^**	-0.254^**	1
X21	-0.434^**	0.304^**	0.599^**	-0.022	0.288^**	0.163^*	-0.141^*	-0.434^**	0.713^**	0.166^**	0.749^**	-0.269^**	-0.444^**	0.710^**	0.075	0.830^**	-0.261^**	0.592^**	-0.288^**	0.733^**	1
X22	-0.316^**	0.026	0.221^**	0.013	0.005	0.174^**	0.016	-0.069	0.377^**	0.238^**	0.345^**	-0.053	-0.134^*	0.371^**	0.176^**	0.405^**	0.049	0.386^**	-0.078	0.402^**	0.359^**	1
X23	-0.262^**	0.056	0.288^**	-0.019	-0.076	0.168^**	0.001	-0.029	0.323^**	0.170^**	0.321^**	-0.071	-0.100	0.319^**	0.083	0.375^**	0.016	0.366^**	-0.093	0.341^**	0.323^**	0.835^**	1

表5

表6

表7

表8

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编号 Serial number	名称 Name	来源 Origin	编号 Serial number	名称 Name	来源 Origin
1	黄青1号 Huangqing 1	甘肃甘南 Gannan, Gansu	19	藏1257 Zang1257	西藏 Tibet
2	硕般多 Shuobanduo	西藏昌都 Changdu, Tibet	20	喜马拉14号 Ximala 14	西藏日喀则 Rikaze, Tibet
3	宕昌青稞（蓝） Dangchang hulless barley	甘肃陇南 Longnan, Gansu	21	藏902960 Zang 902960	西藏拉萨 Lasa, Tibet
4	东都红胶泥-1 Dongdu red clay-1	青海乐都 Ledu, Qinghai	22	IG107018	不详 Unknown
5	甘孜88140-5-6 Ganzi 88140-5-6	四川甘孜 Ganzi, Sichuan	23	昆仑14号 Kunlun 14	青海西宁 Xining, Qinghai
6	阿7019 A 7019	四川阿坝 Aba, Sichuan	24	昆仑15号 Kunlun 15	青海西宁 Xining, Qinghai
7	阿127 A127	四川阿坝 Aba, Sichuan	25	紫勾芒青稞 Purple hook hullessbarley	不详 Unknown
8	苏农0006 Sunong 006	江苏南山 Nanshan, Jiangsu	26	藏1373 Zang 1373	西藏 Tibet
9	北青9号 Beiqing 9	青海海北 Haibei, Qinghai	27	白金紫 Baijinzi	不详 Unknown
10	甘青5号 Ganqing 5	甘肃甘南 Gannan, Gansu	28	藏1673 Zang 1673	西藏 Tibet
11	洛隆宗 Luolongzong	西藏昌都 Changdu, Tibet	29	藏902861 Zang 902861	西藏 Tibet
12	乌表 Wubiao	江苏 Jiangsu	30	藏2503 Zang 2503	西藏 Tibet
13	光头大麦 Guangtou barley	不详 Unknown	31	汤堆小春青稞 Tangduixiaochun hulless barley	云南汤堆 Tangdui, Yunnan
14	紫青稞 Purple hulless barley	西藏贡嘎 Gongga, Tibet	32	汤满青稞 Tangman hulless barley	云南香格里拉 Xianggelila, Yunnan
15	北青1号 Beiqing 1	青海海北 Haibei, Qinghai	33	ZYM01909	西藏工布江达 Gongbujiangda, Tibet
16	青永1421 Qingyong 1421	西藏扎达 Zhada, Tibet	34	ZYM01915	西藏工布江达 Gongbujiangda, Tibet
17	青永4080 Qing yong 4080	西藏 Tibet	35	ZYM01922	西藏工布江达 Gongbujiangda, Tibet
18	紫壳扎尼玛 Purple shell zhanima	西藏拉萨 Lasa, Tibet

指标（西宁点） Index（Xining point）	株高 PH (cm)	有效分蘖数 TN	重心 GG (cm)	根干重 RDW (g)	根数 RN	穗重 PW (g)	穗粒数 KPS	倒伏率 LR (%)
最小值 Minimum	72.90	3.04	29.30	0.60	23.10	0.82	12.70	0.00
最大值 Maximum	115.90	12.79	48.30	2.11	64.50	4.48	76.90	1.00
均值 Mean	96.14	6.83	40.36	0.98	39.71	3.18	53.46	0.42
标准差 Standard deviation	8.98	2.17	4.19	0.32	9.27	0.71	13.21	0.33
方差 Variance	80.62	4.69	17.53	0.10	85.92	0.50	174.44	0.11
偏度 Skewness	-0.24	0.74	-0.16	1.87	0.58	-0.90	-1.18	0.12
峰度 Kurtosis	-0.29	0.52	-0.36	3.89	0.06	1.87	1.73	-1.40
变异系数 CV (%)	9.34	31.70	10.37	32.44	23.34	22.31	24.70	79.69
指标（海北点） Index（Haibei point）	株高 PH (cm)	有效分蘖数 TN	重心 GG (cm)	根干重 RDW (g)	根数 RN	穗重 PW (g)	穗粒数 KPS	倒伏率 LR (%)
最小值 Minimum	60.00	1.00	23.00	0.15	8.00	0.88	13.00	0.00
最大值 Maximum	192.00	12.00	57.00	1.40	50.00	6.59	89.00	0.98
均值 Mean	94.90	3.45	37.88	0.56	29.55	2.55	44.83	0.47
标准差 Standard deviation	16.03	1.96	5.51	0.24	9.40	1.11	17.39	0.38
方差 Variance	256.95	3.86	30.34	0.06	88.31	1.24	302.45	0.14
偏度 Skewness	1.13	1.56	0.06	1.03	0.26	1.20	0.40	-0.01
峰度 Kurtosis	8.72	3.51	0.35	1.20	-0.74	1.61	-0.41	-1.70
变异系数 CV (%)	16.89	56.95	14.54	43.65	31.80	43.57	38.79	81.29

变异来源 Variance sources	df	第二节茎长 SSL		第二节壁厚 SWT		第二节茎粗 SST		第二节茎重 SSW		第二节秆强 SSS		第三节茎长 TSL		第三节壁厚 TWT		第三节茎粗 TST
变异来源 Variance sources	df	F	P	F	P	F	P	F	P	F	P	F	P	F	P	F	P
基因型 Genotype (G)	34	1.44	0.005	1.08	0.00**	2.09	0	1.93	0.00^**	1.96	0.00^**	2.52	0.00^**	1.00	0.00^**	1.43	0.00^**
环境 Environment (E)	1	6.60	0.002	83.79	0.00^**	1296.69	0	0.74	0.043	58.44	0.00^**	5.84	0.00^**	92.42	0.00^**	1457.58	0.00^**
基因型×环境 G×E	34	1.71	0.016	24.57	0.00^**	16.44	0	5.93	0.00^**	16.66	0.00^**	3.14	0.00^**	48.22	0.00^**	36.90	0.00^**
变异来源 Variance sources	df	第三节茎重 TSW		第三节秆强 TSS		第四节茎长 FSL		第四节壁厚 FWT		第四节茎粗 FWT		第四节茎重 FSW		第四节秆强 FSS		倒伏率 LR
变异来源 Variance sources	df	F	P	F	P	F	P	F	P	F	P	F	P	F	P	F	P
基因型 Genotype (G)	34	1.41	0.00^**	2.20	0.00^**	1.85	0.00^**	1.42	0.00^**	0.86	0.00^**	2.08	0.00^**	1.82	0.00^**	2.63	0.00^**
环境 Environment (E)	1	3.44	0.00^**	87.03	0.00^**	3.78	0.00^**	76.58	0.00^**	25.82	0.00^**	295.53	0.00^**	96.19	0.00^**	0.50	0.00^**
基因型×环境 G×E	34	5.01	0.00^**	51.21	0.00^**	6.19	0.00^**	27.23	0.00^**	11.95	0.00^**	4.27	0.00^**	9.80	0.00^**	1411.03	0.00^**
变异来源 Variance sources	df	根数 RN		根干重 RDW		株高 PH		分蘖数 TN		重心 CG		穗重 PW		穗粒数 KPS
变异来源 Variance sources	df	F	P	F	P	F	P	F	P	F	P	F	P	F	P
基因型 Genotype (G)	34	0.72	0.00^**	0.69	0.00^**	1.09	0.00^**	2.07	0.00^**	1.46	0.00^**	2.68	0.00^**	2.25	0.00^**
环境 Environment (E)	1	22.41	0.00^**	36.46	0.00^**	0.24	0.254	85.01	0.00^**	6.67	0.00^**	15.19	0.00^**	9.06	0.00^**
基因型×环境 G×E	34	10.57	0.00^**	14.99	0.00^**	5.65	0.00^**	5.00	0.00^**	7.89	0.00^**	23.92	0.00^**	47.81	0.00^**

初始特征值 Initial eigenvalue			提取平方和载入 Extract square sum loading
特征值 EVl	方差贡献率 VCP (%)	累积贡献率 CP (%)	特征值 EVl	方差贡献率 VCP (%)	累积贡献率 CP (%)
5.00	38.46	38.46	5.00	38.46	38.46
2.40	18.43	56.89	2.40	18.43	56.89
1.36	10.44	67.33	1.36	10.44	67.33
1.27	9.80	77.13	1.27	9.80	77.13
0.71	5.43	82.56
0.49	3.74	86.30
0.47	3.64	89.94
0.43	3.33	93.26
0.25	1.89	95.16
0.23	1.74	96.90
0.15	1.16	98.06
0.14	1.06	99.12
0.11	0.88	100.00

	成份 Component
	1	2	3	4
根干重 RDW	0.714	-0.289	0.109	0.024
分蘖数 TN	0.327	-0.448	0.564	0.310
第二节茎重 SST	0.219	0.799	0.060	-0.363
第二节秆强 SSS	0.821	-0.103	0.091	-0.045
第三节茎长 TSL	-0.300	0.449	0.497	0.415
第三节茎重 TSW	0.136	0.821	0.082	-0.413
第三节秆强 TSS	0.893	-0.045	0.058	-0.102
第四节茎长 FSL	-0.226	0.569	0.449	0.417
第四节壁厚 FWT	0.762	0.089	0.089	-0.052
第四节茎重 FSW	0.851	0.103	0.269	-0.029
第四节秆强 FSS	0.880	-0.093	0.089	-0.144
穗重 PW	0.573	0.372	-0.440	0.483
穗粒数 KPS	0.544	0.308	-0.524	0.510