小麦籽粒钙和钾含量的QTL定位及遗传效应分析

doi:10.7668/hbnxb.20194687

摘要/Abstract

摘要：

钙和钾是小麦重要的矿质营养元素,发掘和探索与其含量相关的遗传机制及效应对于人体营养健康具有重要意义。旨在为小麦籽粒矿质元素生物强化育种提供理论基础,以Avocet/Chilero(AC)构建的164份F₆重组自交系(RILs)和以Avocet/Huites(AH)构建的175份F₆RILs为材料,分析了2个群体在5个环境下籽粒钙(GCa)和钾(GK)含量的表型变异,结合DArT芯片分型结果,共定位到19个与籽粒钙含量相关的QTL,分别位于1A、1D、2A、2B、3A、3D、4A、4B、4D、5A、5B、7A、7B和7D染色体上,解释了3.23%~16.29%的表型变异,同时共定位到23个与籽粒钾含量相关的QTL,分别位于1B、2A、2B、3A、3B、4A、4D、5A、6A、6B和7D染色体上,解释了3.31%~24.66%的表型变异。其中,QGCa.haust-1A、QGCa.haust-AC-5A和QGK.haust-AC-2A.2可在多环境下被定位到,QGCa.haust-1A和QGCa.haust-AC-5A分别解释了表型变异的7.82%~12.72%和9.68%~15.57%,其物理区间为498.67~532.21 Mb和461.52~486.26 Mb,QGK.haust-AC-2A.2解释了表型变异的8.15%~15.20%,其物理区间为354.61~462.37 Mb。通过对3个位点的遗传效应分析可知,分别携带QGCa.haust-1A、QGCa.haust-AC-5A和QGK.haust-AC-2A.2效应位点能有效增加小麦籽粒中的钙和钾含量,聚合效应分析表明,同时携带QGCa.haust-1A效应位点和QGCa.haust-AC-5A效应位点的株系钙含量极显著高于仅携带单一位点的株系。综上,在1A、2A和5A染色体上定位到3个与籽粒钙和钾含量相关的稳定主效位点,其显著提高了小麦籽粒钙和钾含量。

关键词: 小麦, 钙元素, 钾元素, QTL, 遗传效应

Abstract:

Calcium and potassium are important mineral nutrient elements in wheat.It is significant to explore the related genetic mechanisms and effects on human nutritional health.To provide a theoretical basis for biofortification breeding of trace elements in wheat grains,we used 164 F₆ recombinant inbred lines(RILs)derived from Avocet/Chilero(AC)and 175 F₆ RILs derived from Avocet/Huites(AH).Our investigation focused on phenotypic variations in grain calcium(GCa)and grain potassium(GK)content in five environments.QTL mapping was conducted with diversity arrays technology(DArT)chip.Nineteen QTLs associated with grain calcium content were identified,distributed on chromosomes 1A,1D,2A,2B,3A,3D,4A,4B,4D,5A,5B,7A,7B,and 7D,explaining 3.23%—16.29% of phenotypic variation.Simultaneously,23 QTLs linked to grain potassium content were identified on chromosomes 1B,2A,2B,3A,3B,4A,4D,5A,6A,6B,and 7D,explaining 3.31%—24.66% of phenotypic variation.QGCa.haust-1A,QGCa.haust-AC-5A and QGK.haust-AC-2A.2 were located in multiple environments.QGCa.haust-1A and QGCa.haust-AC-5A explained 7.82%—12.72% and 9.68%—15.57% of phenotypic variation,and the physical intervals were 498.67—532.21 Mb and 461.52—486.26 Mb,respectively.QGK.haust-AC-2A.2 explained 8.15%—15.20% of phenotypic variation,with a physical range of 354.61—462.37 Mb.The genetic effect analysis of QGCa.haust-1A,QGCa.haust-AC-5A,and QGK.haust-AC-2A.2 showed that each locus effectively increased the calcium and potassium content in wheat grain.Aggregation effect analysis indicated that the lines with QGCa.haust-1A and QGCa.haust-AC-5A effect loci had highly significantly higher calcium content than those with only a single locus.In summary,three stable loci of grain calcium and potassium content are mapped on chromosomes 1A,2A,and 5A,which could significantly increase calcium and potassium content in wheat grain.

Key words: Wheat, Calcium, Potassium, QTL, Genetic effect

洪壮壮, 曾占奎, 宋俊乔, 李琼, 颜群翔, 赵越, 毕俊鸽, 张伟, 王春平. 小麦籽粒钙和钾含量的QTL定位及遗传效应分析[J]. 华北农学报, 2024, 39(4): 37-46. doi: 10.7668/hbnxb.20194687.

HONG Zhuangzhuang, ZENG Zhankui, SONG Junqiao, LI Qiong, YAN Qunxiang, ZHAO Yue, BI Junge, ZHANG Wei, WANG Chunping. QTL Mapping and Genetic Effect Analysis of Calcium and Potassium Content in Wheat Grain[J]. Acta Agriculturae Boreali-Sinica, 2024, 39(4): 37-46. doi: 10.7668/hbnxb.20194687.

http://www.hbnxb.net/CN/Y2024/V39/I4/37

图/表 7

表1 微波消解条件参数

Tab.1 The parameters for microwave digestion

步骤 Step	温度/℃ Temperature	时间/ min Time	风扇等级 Fan grade	搅拌 Stir
温度爬坡	120	8	1	关闭
Temperature climbing
温度保持		5	1	关闭
Temperature holding
温度爬坡	180	8	1	关闭
Temperature climbing
温度保持		30	1	关闭
Temperature holding
温度冷却	60	20	3	关闭
Temperature cooling

表2 AC群体和AH群体及亲本在多环境下GCa和GK含量的统计分析

Tab.2 Statistical analysis of GCa and GK content in AC and AH populations and the parents in multiple environments

性状 Trait	环境 Environment	亲本Parent			RILs群体RILs population
性状 Trait	环境 Environment	Avocet	Chilero	Huites	均值±标准误 $X ¯$ ±s	变异范围 Range of variation	变异系数/% CV
GCa/(mg/kg)	E1	569.38±11.49A	429.38±9.72B		458.25±57.89	310.08~662.48	12.60
	E2	525.68±8.56a	473.15±9.94b		502.17±59.60	363.28~692.04	11.87
	E3	666.88±29.17A	611.25±14.14B		569.89±68.40	415.63~776.88	12.00
	A_C BLUE	592.70	538.71		498.34±53.35	357.55~683.43	10.71
	E4	569.38±11.49A		430.00±15.91B	467.09±60.84	360.28~723.20	13.03
	E5	437.91±4.80A		370.17±0.04B	461.95±49.18	343.96~659.61	10.65
	A_H BLUE	449.98		376.51	463.59±48.54	326.30~620.73	10.47
GK/(mg/kg)	E1	4 201.25±36.24a	3 511.25±119.32b		3 842.88±327.94	3 018.13~4 944.38	8.53
	E2	3 633.13±65.41a	3 255.63±10.61b		3 608.83±288.21	2 963.13~5 003.13	7.99
	E3	3 907.50±27.40a	3 343.13±98.99b		4 099.18±582.87	3 141.25~5 415.00	14.22
	A_C BLUE	3 714.14	3 251.18		3 806.84±283.62	3 030.28~4 760.85	7.45
	E4	4 201.25±36.24A		3 803.13±44.19B	3 936.10±344.46	3 194.68~5 243.60	8.75
	E5	4 204.97±17.74A		3 714.19±77.75B	4 058.94±327.16	3 467.59~5 225.85	8.06
	A_H BLUE	4 330.27		3 826.65	4 018.00±307.67	3 398.93~5 225.85	7.66

表2 AC群体和AH群体及亲本在多环境下GCa和GK含量的统计分析

Tab.2 Statistical analysis of GCa and GK content in AC and AH populations and the parents in multiple environments

性状 Trait	环境 Environment	亲本Parent			RILs群体RILs population
性状 Trait	环境 Environment	Avocet	Chilero	Huites	均值±标准误 $X ¯$ ±s	变异范围 Range of variation	变异系数/% CV
GCa/(mg/kg)	E1	569.38±11.49A	429.38±9.72B		458.25±57.89	310.08~662.48	12.60
	E2	525.68±8.56a	473.15±9.94b		502.17±59.60	363.28~692.04	11.87
	E3	666.88±29.17A	611.25±14.14B		569.89±68.40	415.63~776.88	12.00
	A_C BLUE	592.70	538.71		498.34±53.35	357.55~683.43	10.71
	E4	569.38±11.49A		430.00±15.91B	467.09±60.84	360.28~723.20	13.03
	E5	437.91±4.80A		370.17±0.04B	461.95±49.18	343.96~659.61	10.65
	A_H BLUE	449.98		376.51	463.59±48.54	326.30~620.73	10.47
GK/(mg/kg)	E1	4 201.25±36.24a	3 511.25±119.32b		3 842.88±327.94	3 018.13~4 944.38	8.53
	E2	3 633.13±65.41a	3 255.63±10.61b		3 608.83±288.21	2 963.13~5 003.13	7.99
	E3	3 907.50±27.40a	3 343.13±98.99b		4 099.18±582.87	3 141.25~5 415.00	14.22
	A_C BLUE	3 714.14	3 251.18		3 806.84±283.62	3 030.28~4 760.85	7.45
	E4	4 201.25±36.24A		3 803.13±44.19B	3 936.10±344.46	3 194.68~5 243.60	8.75
	E5	4 204.97±17.74A		3 714.19±77.75B	4 058.94±327.16	3 467.59~5 225.85	8.06
	A_H BLUE	4 330.27		3 826.65	4 018.00±307.67	3 398.93~5 225.85	7.66

图1 小麦GCa和GK含量在RILs群体中的分布

Fig.1 Distribution of GCa and GK content in wheat RILs populations

表3 AC群体和AH群体GCa和GK含量的变异分析

Tab.3 Variance analysis of GCa and GK content in AC and AH populations

群体 Population	性状 Trait	变异来源 Source of variation	自由度 Degree of freedom	均方 Mean square	F值 F-value	P值 P-value	遗传力 Heritability
AC	GCa	基因型	163	15 848.31	46.29	0.00	0.82
		环境	2	963 254.75	2 813.33	0.00
		重复	2	379.00	1.11	0.35
		基因型×环境	305	3 370.06	9.84	0.00
	GK	基因型	163	450 795.69	28.27	0.00	0.83
		环境	2	13 324 130.00	835.52	0.00
		重复	2	60 006.11	3.76	0.88
		基因型×环境	303	89 304.38	5.60	0.00
AH	GCa	基因型	174	8 649.52	17.50	0.00	0.71
		环境	1	4 020.06	8.13	0.00
		重复	2	161.91	0.33	0.72
		基因型×环境	153	3 300.23	6.68	0.00
	GK	基因型	174	345 028.84	26.86	0.00	0.78
		环境	1	2 490 041.25	193.87	0.00
		重复	2	676.63	0.05	0.96
		基因型×环境	153	91 888.67	7.15	0.00

表4 AC和AH群体中定位的GCa和GK含量相关的QTL

Tab.4 QTLs of GCa and GK content in AC and AH populations

性状 Trait	QTL	环境 Environment	物理区间/Mb Physical interval	标记区间 Marker interval		LOD值 LOD score	表型变异率/% Phenotypic variation rate	加性效应 Additive effect
GCa	QGCa.haust-AC-1A.1	E1	498.67~531.67	4404599	~2275809	3.69	9.74	-22.45
		A_C BLUE	498.67~531.67	4404599	~2275809	5.88	12.72	-25.27
	QGCa.haust-AC-1A.2	E2	498.82~499.84	3961251	~NP1204551	3.36	10.14	-21.29
	QGCa.haust-AH-1A	E4	527.79~532.21	100079848	~SNP100276833	2.88	7.82	-17.12
	QGCa.haust-AC-1D	E2	12.03~490.54	4991409	~SNP100509052	2.58	10.14	21.57
	QGCa.haust-AH-2A	A_H BLUE	258.35~433.32	100007140	~SNP1712854	3.92	6.57	15.49
	QGCa.haust-AH-2B	A_H BLUE	330.24~442.92	1712720	~SNP992800	3.08	3.97	-11.81
	QGCa.haust-AH-3A	E4	36.79~36.94	SNP1112192	~3028938	4.47	12.45	21.57
	QGCa.haust-AC-3D	E1	523.25~599.00	1235289	~3960257	3.29	11.84	23.00
		A_C BLUE	523.25~599.00	1235289	~3960257	3.39	10.49	21.24
	QGCa.haust-AC-4A	E3	617.91~642.95	984754	~SNP100643318	4.11	16.29	30.38
	QGCa.haust-AH-4B	E4	633.36~639.12	SNP1125612	~SNP1239576	3.34	10.51	22.90
	QGCa.haust-AC-5A	E1	461.52~486.26	1111766	~3384967	3.85	9.68	23.76
		E2	461.52~486.26	1111766	~3384967	5.13	15.57	27.77
		A_C BLUE	461.52~486.26	1111766	~3384967	6.57	13.69	27.81
	QGCa.haust-AH-5A	A_H BLUE	482.12~488.05	SNP2359596	~SNP2246034	2.63	3.23	10.75
	QGCa.haust-AH-5B	A_H BLUE	547.32~711.14	SNP1053823	~100092795	7.89	10.60	-19.42
	QGCa.haust-AC-7A	E1	29.12~53.57	1102212	~1121695	2.84	8.74	-19.32
	QGCa.haust-AH-7A.1	A_H BLUE	63.86~649.82	3959069	~1129617	5.97	7.95	-16.70
	QGCa.haust-AH-7A.2	E5	69.98~79.65	1005950	~SNP2279107	2.63	8.38	-14.97
	QGCa.haust-AH-7A.3	A_H BLUE	694.37~701.24	SNP1128104	~SNP1213152	5.16	6.53	-15.14
	QGCa.haust-AH-7B	A_H BLUE	718.08~726.98	1095778	~1066421	9.43	12.98	21.44
	QGCa.haust-AH-7D	E5	627.57~631.80	984331	~3945964	3.39	10.89	-17.09
GK	QGK.haust-AC-1B	A_C BLUE	504.51~544.13	2278693	~SNP2266225	5.55	7.37	-112.66
	QGK.haust-AC-2A.1	E3	62.36~299.77	1068736	~SNP1059865	3.28	14.28	-149.54
	QGK.haust-AC-2A.2	E1	354.61~462.37	1290197	~1217017	5.64	14.44	-147.75
		E2	354.61~462.37	1290197	~1217017	4.64	15.20	-141.97
		E3	354.61~462.37	1290197	~1217017	11.84	8.15	-340.26
	QGK.haust-AC-2B.1	E3	66.78~68.19	SNP1105506	~4991074	12.07	7.97	-283.43
	QGK.haust-AC-2B.2	E1	553.98~569.14	4984635	~SNP100261608	4.21	9.81	-101.71
	QGK.haust-AC-3A.1	E1	145.64~148.62	1103153	~3034124	9.57	24.66	172.29
		A_C BLUE	145.64~148.62	1103153	~3034124	14.86	23.56	208.90
	QGK.haust-AC-3A.2	E2	739.39~745.18	3942045	~3026156	3.46	10.16	-106.77
	QGK.haust-AH-3A	E5	51.38~54.94	2363021	~1131991	4.34	3.86	96.88
	QGK.haust-AC-3B	E3	707.93~733.62	1096061	~4004609	10.59	6.75	296.17
	QGK.haust-AH-3B.1	E5	244.23~819.90	SNP1125230	~3948912	7.02	8.90	147.02
	QGK.haust-AH-3B.2	E5	44.86~49.15	SNP1863107	~SNP4397490	3.89	3.48	100.29
	QGK.haust-AC-4A.1	A_C BLUE	239.41~584.04	3950319	~SNP100554548	7.80	10.60	-138.27
	QGK.haust-AC-4A.2	E3	617.05~617.36	SNP100419930	~SNP1131791	6.72	3.94	229.14
	QGK.haust-AC-4A.3	E3	577.51~581.47	100006450	~1057661	11.66	8.33	-307.89
	QGK.haust-AC-4D	E1	222.20~419.44	2248602	~2245206	2.56	5.62	-77.07
	QGK.haust-AH-5A.1	E5	528.52~572.04	1165946	~SNP4992747	8.81	8.93	-148.26
	QGK.haust-AH-5A.2	A_H BLUE	580.93~581.14	SNP100269435	~SNP100358982	6.70	8.12	-129.16
	QGK.haust-AH-5A.3	E5	595.39~596.03	SNP100116253	~1131350	16.50	19.31	216.57
		A_H BLUE	595.39~596.03	SNP100116253	~1131350	14.51	20.12	202.56
	QGK.haust-AH-6A	E5	530.14~607.41	SNP100414704	~SNP5001084	6.24	6.06	140.36
		A_H BLUE	530.14~607.41	SNP100414704	~SNP5001084	5.24	6.26	130.77
	QGK.haust-AH-6B	E5	112.94~707.72	4540911	~1233197	6.75	6.50	133.47
		A_H BLUE	112.94~707.72	4540911	~1233197	7.52	9.17	145.79
	QGK.haust-AC-7D.1	A_C BLUE	530.72~609.02	4543493	~SNP100360853	3.39	4.35	-84.49
	QGK.haust-AC-7D.2	E3	553.03~556.29	SNP2242097	~3961158	5.75	3.31	195.41
	QGK.haust-AC-7D.3	A_C BLUE	5.02~31.69	3384889	~SNP100273610	7.63	10.25	-139.95

表4 AC和AH群体中定位的GCa和GK含量相关的QTL

Tab.4 QTLs of GCa and GK content in AC and AH populations

性状 Trait	QTL	环境 Environment	物理区间/Mb Physical interval	标记区间 Marker interval		LOD值 LOD score	表型变异率/% Phenotypic variation rate	加性效应 Additive effect
GCa	QGCa.haust-AC-1A.1	E1	498.67~531.67	4404599	~2275809	3.69	9.74	-22.45
		A_C BLUE	498.67~531.67	4404599	~2275809	5.88	12.72	-25.27
	QGCa.haust-AC-1A.2	E2	498.82~499.84	3961251	~NP1204551	3.36	10.14	-21.29
	QGCa.haust-AH-1A	E4	527.79~532.21	100079848	~SNP100276833	2.88	7.82	-17.12
	QGCa.haust-AC-1D	E2	12.03~490.54	4991409	~SNP100509052	2.58	10.14	21.57
	QGCa.haust-AH-2A	A_H BLUE	258.35~433.32	100007140	~SNP1712854	3.92	6.57	15.49
	QGCa.haust-AH-2B	A_H BLUE	330.24~442.92	1712720	~SNP992800	3.08	3.97	-11.81
	QGCa.haust-AH-3A	E4	36.79~36.94	SNP1112192	~3028938	4.47	12.45	21.57
	QGCa.haust-AC-3D	E1	523.25~599.00	1235289	~3960257	3.29	11.84	23.00
		A_C BLUE	523.25~599.00	1235289	~3960257	3.39	10.49	21.24
	QGCa.haust-AC-4A	E3	617.91~642.95	984754	~SNP100643318	4.11	16.29	30.38
	QGCa.haust-AH-4B	E4	633.36~639.12	SNP1125612	~SNP1239576	3.34	10.51	22.90
	QGCa.haust-AC-5A	E1	461.52~486.26	1111766	~3384967	3.85	9.68	23.76
		E2	461.52~486.26	1111766	~3384967	5.13	15.57	27.77
		A_C BLUE	461.52~486.26	1111766	~3384967	6.57	13.69	27.81
	QGCa.haust-AH-5A	A_H BLUE	482.12~488.05	SNP2359596	~SNP2246034	2.63	3.23	10.75
	QGCa.haust-AH-5B	A_H BLUE	547.32~711.14	SNP1053823	~100092795	7.89	10.60	-19.42
	QGCa.haust-AC-7A	E1	29.12~53.57	1102212	~1121695	2.84	8.74	-19.32
	QGCa.haust-AH-7A.1	A_H BLUE	63.86~649.82	3959069	~1129617	5.97	7.95	-16.70
	QGCa.haust-AH-7A.2	E5	69.98~79.65	1005950	~SNP2279107	2.63	8.38	-14.97
	QGCa.haust-AH-7A.3	A_H BLUE	694.37~701.24	SNP1128104	~SNP1213152	5.16	6.53	-15.14
	QGCa.haust-AH-7B	A_H BLUE	718.08~726.98	1095778	~1066421	9.43	12.98	21.44
	QGCa.haust-AH-7D	E5	627.57~631.80	984331	~3945964	3.39	10.89	-17.09
GK	QGK.haust-AC-1B	A_C BLUE	504.51~544.13	2278693	~SNP2266225	5.55	7.37	-112.66
	QGK.haust-AC-2A.1	E3	62.36~299.77	1068736	~SNP1059865	3.28	14.28	-149.54
	QGK.haust-AC-2A.2	E1	354.61~462.37	1290197	~1217017	5.64	14.44	-147.75
		E2	354.61~462.37	1290197	~1217017	4.64	15.20	-141.97
		E3	354.61~462.37	1290197	~1217017	11.84	8.15	-340.26
	QGK.haust-AC-2B.1	E3	66.78~68.19	SNP1105506	~4991074	12.07	7.97	-283.43
	QGK.haust-AC-2B.2	E1	553.98~569.14	4984635	~SNP100261608	4.21	9.81	-101.71
	QGK.haust-AC-3A.1	E1	145.64~148.62	1103153	~3034124	9.57	24.66	172.29
		A_C BLUE	145.64~148.62	1103153	~3034124	14.86	23.56	208.90
	QGK.haust-AC-3A.2	E2	739.39~745.18	3942045	~3026156	3.46	10.16	-106.77
	QGK.haust-AH-3A	E5	51.38~54.94	2363021	~1131991	4.34	3.86	96.88
	QGK.haust-AC-3B	E3	707.93~733.62	1096061	~4004609	10.59	6.75	296.17
	QGK.haust-AH-3B.1	E5	244.23~819.90	SNP1125230	~3948912	7.02	8.90	147.02
	QGK.haust-AH-3B.2	E5	44.86~49.15	SNP1863107	~SNP4397490	3.89	3.48	100.29
	QGK.haust-AC-4A.1	A_C BLUE	239.41~584.04	3950319	~SNP100554548	7.80	10.60	-138.27
	QGK.haust-AC-4A.2	E3	617.05~617.36	SNP100419930	~SNP1131791	6.72	3.94	229.14
	QGK.haust-AC-4A.3	E3	577.51~581.47	100006450	~1057661	11.66	8.33	-307.89
	QGK.haust-AC-4D	E1	222.20~419.44	2248602	~2245206	2.56	5.62	-77.07
	QGK.haust-AH-5A.1	E5	528.52~572.04	1165946	~SNP4992747	8.81	8.93	-148.26
	QGK.haust-AH-5A.2	A_H BLUE	580.93~581.14	SNP100269435	~SNP100358982	6.70	8.12	-129.16
	QGK.haust-AH-5A.3	E5	595.39~596.03	SNP100116253	~1131350	16.50	19.31	216.57
		A_H BLUE	595.39~596.03	SNP100116253	~1131350	14.51	20.12	202.56
	QGK.haust-AH-6A	E5	530.14~607.41	SNP100414704	~SNP5001084	6.24	6.06	140.36
		A_H BLUE	530.14~607.41	SNP100414704	~SNP5001084	5.24	6.26	130.77
	QGK.haust-AH-6B	E5	112.94~707.72	4540911	~1233197	6.75	6.50	133.47
		A_H BLUE	112.94~707.72	4540911	~1233197	7.52	9.17	145.79
	QGK.haust-AC-7D.1	A_C BLUE	530.72~609.02	4543493	~SNP100360853	3.39	4.35	-84.49
	QGK.haust-AC-7D.2	E3	553.03~556.29	SNP2242097	~3961158	5.75	3.31	195.41
	QGK.haust-AC-7D.3	A_C BLUE	5.02~31.69	3384889	~SNP100273610	7.63	10.25	-139.95

图2 小麦GCa和GK含量的主要QTL定位

Fig.2 Main QTL mapping of GCa and GK content in wheat grain

图3 GCa和GK主效位点遗传效应分析 A.QGCa.haust-1A效应分析;B.QGCa.haust-AC-5A效应分析;C.QGK.haust-AC-2A.2效应分析;D.聚合效应分析。+.携带效应位点,-.未携带效应位点;大写字母表示P<0.01。

Fig.3 Genetic effect analysis of GCa and GK main effect loci A.QGCa.haust-1A effect analysis;B.QGCa.haust-AC-5A effect analysis;C.QGK.haust-AC-2A.2 effect analysis;D.Polymerization effect analysis.+.The effect locus is carried;-.The effect locus is not carried;capital letter represents P<0.01.

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