小麦苗期耐低磷相关基因位点的挖掘与候选基因分析

doi:10.3724/SP.J.1006.2025.51033

Abstract

Abstract:

Phosphorus (P) is an essential macronutrient for crop growth and development. However, only a small fraction of the phosphorus present in soil is effectively utilized by plants, and long-term application of P fertilizers can lead to environmental pollution. Therefore, it is of great significance to screen for varieties with strong tolerance to low-P conditions and to identify associated QTLs and candidate genes. In this study, a natural population consisting of 389 wheat varieties was used as experimental material. A three-year hydroponic experiment (2022, 2023, and 2024) was conducted under both normal-P (control) and low-P conditions at the wheat seedling stage. Nine traits were measured, including seedling height, main root length, root number, shoot dry weight, root dry weight, total root length, root surface area, root diameter, and number of root tips. For each trait, the low-P tolerance coefficient and BLUP values were calculated, and comprehensive evaluation D-values were derived based on these coefficients. Correlation analysis of BLUP values across the three environments revealed significantly positive correlations among most traits, except for root diameter, under both low-P and control conditions. Cluster analysis of the comprehensive D-values identified ‘Fengdecunmai 1' as a strongly low-P-tolerant variety across all four environments (2022, 2023, 2024, and BLUP). A genome-wide association study (GWAS) was carried out using low-P tolerance coefficients and D-values for the nine traits across the four environments, based on the 660K SNP chip. A total of 1,197 significant SNP markers were detected, forming 464 QTLs. Among these, 20 QTLs were repeatedly detected in two environments, and 7 QTLs were detected in three or four environments, with the phenotypic variance explained (R²) ranging from 4.09% to 10.58%. Based on previously published transcriptomic data and gene functional annotation, three candidate genes associated with low-P tolerance were identified within the regions of the seven QTLs. TraesCS4D02G022900 and TraesCS4D02G023300 encode F-box family proteins. Their Arabidopsis ortholog, At5g21040, encodes a protein containing both WD40 and F-box domains and functions as a negative regulator of the P starvation response. TraesCS6D02G154700 encodes a receptor-like protein kinase involved in plant growth, development, and responses to stress and disease. Further analysis of the expression patterns of these three candidate genes in both leaves and roots under low-P stress revealed differential expression consistent with previous transcriptomic results. These findings provide a solid foundation for the development of low-P-tolerant wheat cultivars and for elucidating the functions and regulatory mechanisms of genes associated with low-P stress.

Key words: wheat, low phosphorus stress, GWAS, SNP, candidate genes

CAO Zhi-Yang, GAO Li-Feng, JIANG Dong-Yan, WANG Shu-Guang, YANG Jin-Wen, JIA Ji-Zeng, LI Ning, SUN Dai-Zhen. Identification of genetic loci related to low phosphorus tolerance at the seedling stage in wheat and analysis of candidate genes[J].Acta Agronomica Sinica, 2025, 51(10): 2632-2651.

Figures/Tables 16

Table 1

Table 2

Table 3

Statistical analysis of phenotypic traits under different phosphorus treatments"

性状 Trait	环境 Environment	对照处理 CK			低磷处理 Low-phosphorus			耐低磷系数 Low-phosphorus tolerance coefficient
性状 Trait	环境 Environment	平均值 Mean	标准差 SD	变异系数CV (%)	平均值 Mean	标准差 SD	变异系数CV (%)	平均值 Mean	标准差 SD	变异系数CV (%)
苗高 SH (cm)	E1	29.18	6.25	21.42	19.02	4.10	21.56	0.66	0.10	15.54
	E2	27.64	4.91	17.75	18.43	2.84	15.41	0.68	0.11	16.70
	E3	29.79	5.23	17.56	19.35	3.23	16.68	0.66	0.11	16.99
	BLUP	28.94	3.94	13.61	19.01	2.27	11.96	0.66	0.07	10.70
主根长 MRL (cm)	E1	22.39	5.31	23.71	25.90	5.53	21.35	1.18	0.20	17.04
	E2	22.79	5.40	23.68	25.71	5.31	20.64	1.15	0.21	18.47
	E3	23.60	5.30	22.46	27.86	5.75	20.64	1.22	0.30	24.86
	BLUP	22.84	3.18	13.94	26.42	3.36	12.71	1.17	0.15	12.64
根条数 RN	E1	5.96	1.30	21.83	4.93	0.93	18.79	0.84	0.14	16.80
	E2	5.83	1.09	18.72	4.89	0.87	17.83	0.86	0.18	20.46
	E3	6.52	1.31	20.15	5.25	1.04	19.86	0.83	0.21	25.70
	BLUP	6.12	0.59	9.60	5.01	0.49	9.84	0.82	0.08	9.18
茎叶干重 SDW (g)	E1	0.043	0.017	41.08	0.023	0.008	33.33	0.58	0.17	29.30
	E2	0.047	0.019	41.18	0.027	0.009	31.55	0.62	0.17	27.32
	E3	0.058	0.025	43.13	0.026	0.010	40.92	0.51	0.28	55.81
	BLUP	0.048	0.011	22.27	0.025	0.004	17.71	0.53	0.10	18.76
根干重 RDW(g)	E1	0.010	0.005	47.66	0.014	0.005	35.43	1.52	0.57	37.46
	E2	0.010	0.005	49.06	0.016	0.005	33.84	1.73	0.75	43.16
	E3	0.012	0.006	47.45	0.015	0.007	46.63	1.48	0.78	52.59
	BLUP	0.011	0.002	21.08	0.015	0.003	20.93	1.37	0.28	20.55
总根长 TRL (cm)	E1	262.0	89.4	34.12	308.4	107.4	34.82	1.26	0.49	38.63
	E2	254.4	124.9	49.10	312.9	118.8	37.96	1.43	0.71	49.92
	E3	259.2	89.7	34.60	292.1	120.7	41.32	1.22	0.57	46.91
	BLUP	255.9	34.6	13.52	309.0	36.3	11.76	1.22	0.18	14.71
根面积 RSA (cm²)	E1	17.37	8.07	46.46	20.15	7.03	34.87	1.33	0.59	44.62
	E2	16.83	7.37	43.80	21.45	8.21	38.28	1.44	0.70	48.78
	E3	18.35	5.74	31.30	20.93	7.86	37.54	1.22	0.53	43.02
	BLUP	17.31	1.91	11.01	20.65	1.98	9.57	1.20	0.14	11.73
根直径 RD (mm)	E1	0.234	0.029	12.21	0.205	0.023	11.13	0.88	0.08	9.11
	E2	0.224	0.032	14.28	0.200	0.029	14.62	0.90	0.11	12.26
	E3	0.229	0.033	14.21	0.218	0.029	13.35	0.96	0.11	11.86
	BLUP	0.229	0.016	7.07	0.208	0.012	5.89	0.91	0.05	5.20
根尖数 RTN	E1	465.2	255.8	54.99	531.0	257.8	48.54	1.34	0.74	54.89
	E2	486.8	216.4	44.45	584.5	297.2	50.84	1.32	0.65	49.27
	E3	469.9	227.1	48.33	616.4	300.7	48.77	1.54	0.94	61.33
	BLUP	458.6	96.8	21.10	562.6	136.3	24.23	1.25	0.30	23.73

Table 3

Fig. 1

Table 4

Fig. 2

Fig. 3

Table S1

Test wheat materials"

编号 Code	品种名称 Variety name	编号 Code	品种名称 Variety name
1	科农199 Kenong 199	196	晋麦31 Jinmai 31
2	晋麦2148 Jinmai 2148	197	临抗5027 Linkang 5027
3	山农205 Shannong 205	198	晋麦54 Jinmai 54
4	阿夫 Afu	199	晋麦47 Jinmai 47
5	博爱7023 Boai 7023	200	平阳181 Pingyang 181
6	郑麦9023 Zhengmai 9023	201	临麦4号 Linmai 4
7	郑州5号 Zhengzhou 5	202	临麦2号 Linmai 2
8	豫麦35 Yumai 35	203	矮抗58 Aikang 58
9	豫麦24 Yumai 24	204	平原50 Pingyuan 50
10	豫麦15 Yumai 15	205	百农3217 Bainong 3217
11	偃师4号 Yanshi 4	206	安阳一号 Anyang 1
12	中焦2号 Zhongjiao 2	207	矮粒多 Ailiduo
13	中洛08－2 Zhongluo 08－2	208	丰德存麦1号 Fengdecunmai 1
14	陕253 Shaan 253	209	百农791 Bainong 791
15	商洛81(2)4-19-23 Shangluo 81(2)4-19-23	210	藁优2018 Gaoyou 2018
16	咸农151 Xiannong 151	211	旱选3号 Hanxuan 3
17	小偃6号 Xiaoyan 6	212	旱选10号 Hanxuan 10
18	淮麦33 Huaimai 33	213	国麦301 Guomai 301
19	苏麦3号 Sumai 3	214	开麦21 Kaimai 21
20	生选3号 Shengxuan 3	215	焦麦266 Jiaomai 266
21	苏麦6号 Sumai 6	216	怀川916 Huaichuan 916
22	徐州21 Xuzhou 21	217	洛麦26 Luomai 26
23	扬麦13 Yangmai 13	218	洛麦23 Luomai 23
24	扬麦12 Yangmai 12	219	兰考90(6)52-24 Lankao 90(6)52-24
25	扬麦158 Yangmai 158	220	新麦16 Xinmai 16
26	扬麦17 Yangmai 17	221	温麦4号 Wenmai 4
27	扬麦4号 Yangmai 4	222	太学12 Taixue 12
28	扬麦2号 Yangmai 2	223	新乡9178 Xinxiang 9178
29	扬麦1号 Yangmai 1	224	新麦9号 Xinmai 9
30	镇麦168 Zhenmai 168	225	新麦26 Xinmai 26
31	扬麦5号 Yangmai 5	226	信阳12高 Xinyang 12 gao
32	安徽11 Anhui 11	227	信阳12矮 Xinyang 12 ai
33	龙辐91B-569 (龙辐10号) Longfu 91B-569 (Longfu 10)	228	兴义4号 Xingyi 4
34	西昌76-9 Xichang 76-9	229	许科718 Xuke 718
35	川麦107 Chuanmai 107	230	许农7号 Xunong 7
36	川麦22 Chuanmai 22	231	偃893选 Yan 893xuan
37	繁6 Fan 6	232	偃展4110 Yanzhan 4110
38	绵麦1403 Mianmai 1403	233	豫麦10号 Yumai 10
39	绵麦37 Mianmai 37	234	豫麦13 Yumai 13
40	新春8号 Xinchun 8	235	豫麦16 Yumai 16
41	新春2号 Xinchun 2	236	豫麦18(矮早781) Yumai 18 (Aizao 781)
42	辽春10号 Liaochun 10	237	豫麦19 Yumai 19
43	辽春9号 Liaochun 9	238	豫麦25 Yumai 25
44	新春11号 Xinchun 11	239	豫麦2号 Yumai 2
45	宁春13 Ningchun 13	240	豫麦47 Yumai 47
46	宁春4号 Ningchun 4	241	豫麦70-36 Yumai 70-36
47	吉春1016 Jichun 1016	242	豫农949 Yunong 949
48	品春16 Pinchun 16	243	早穗30 Zaosui 30
49	青春5号 Qingchun 5	244	郑麦1308 (17河南) Zhengmai 1308 (17Henan)
50	绵麦48 Mianmai 48	245	郑麦379 Zhengmai 379
51	绵阳15 Mianyang 15	246	郑麦9201 Zhengmai 9201
52	绵阳26 Mianyang 26	247	郑州17 Zhengzhou 17
53	绵阳79-2 Mianyang 79-2	248	郑州24 Zhengzhou 24
54	内乡182 Neixiang 182	249	邢麦1号 Xingmai 1
55	内麦19 Neimai 19	250	新乡展示田取样(大穗方穗大粒) Xinxiangzhanshitianquyang (dasuifangsuidali)
56	内江31 Neijiang 31	251	中焦1号 Zhongjiao 1
57	雅安早 Yaanzao	252	郑州8960 Zhengzhou 8960
58	镇麦6号 Zhenmai 6	253	郑州761 Zhengzhou 761
59	镇麦9号 Zhenmai 9	254	中洛08－1 Zhongluo 08－1
60	镇麦3号 Zhenmai 3	255	中焦3号 Zhongjiao 3
61	镇麦4号 Zhenmai 4	256	中优206 Zhongyou 206
62	镇麦5号 Zhenmai 5	257	中新78 Zhongxin 78
63	双吉4号 Shuangji 4	258	中育9398 Zhongyu 9398
64	皖麦31 Wanmai 31	259	中育12 Zhongyu 12
65	鄂麦11 Emai 11	260	中优9507 Zhongyou 9507
66	鄂麦12 Emai 12	261	周麦12 Zhoumai 12
67	鄂麦17 Emai 17	262	周8425B Zhou 8425B
68	鄂麦9号 Emai 9	263	中运1号 Zhongyun 1
69	鄂西84-1031 Exi 84-1031	264	周麦13 Zhoumai 13
70	荆州66 Jingzhou 66	265	周麦23 Zhoumai 23
71	察雅折达29 Chayazheda 29	266	周麦20 Zhoumai 20
72	滇662-525-2 Dian 662-525-2	267	周麦19 Zhoumai 19
73	黑宝 Heibao	268	周麦26 Zhoumai 26
74	甘肃96 Gansu 96	269	周麦25 Zhoumai 25
75	东3-6 Dong 3-6	270	周麦24 Zhoumai 24
76	北京10号 Beijing 10	271	周麦32 Zhoumai 32
77	北京0045 Beijing 0045	272	周麦28 Zhoumai 28
78	CA9722	273	周麦27 Zhoumai 27
79	昌乐5号 Changle 5	274	昌农339-5-1 Changnong 339-5-1
80	北京8号 Beijing 8	275	矮丰3号 Aifeng 3
81	北京837 Beijing 837	276	周麦9号 Zhoumai 9
82	丰抗8号 Fengkang 8	277	秦麦4号 Qinmai 4
83	丰抗7号 Fengkang 7	278	秦麦1号 Qinmai 1
84	东方红3号 Dongfanghong 3	279	泾阳60 Jingyang 60
85	京花2号 Jinghua 2	280	陕229 Shaan 229
86	京冬22 Jingdong 22	281	陕354 Shaan 354
87	京411 Jing 411	282	陕旱8675 Shaanhan 8675
88	轮选987 Lunxuan 987	283	陕合6号 Shaanhe 6
89	京双16 Jingshuang 16	284	陕麦150 Shaanmai 150
90	京农79-13 Jingnong 79-13	285	陕麦159 Shaanmai 159
91	农大1108 Nongda 1108	286	陕农1号 Shaannong 1
92	农大211 Nongda 211	287	陕农7859 Shaannong 7859
93	农大183 Nongda 183	288	陕优225 Shaanyou 225
94	品冬34 Pindong 34	289	渭麦4号 Weimai 4
95	农大139 Nongda 139	290	西安8号 Xi'an 8
96	农大36 Nongda 36	291	西北612 Xibei 612
97	农大311 Nongda 311	292	西农88 Xinong 88
98	中麦415 Zhongmai 415	293	咸农39 Xiannong 39
99	中麦175 Zhongmai 175	294	小山2134 Xiaoshan 2134
100	原冬3号 Yuandong 3	295	小偃22 Xiaoyan 22
101	高优503 Gaoyou 503	296	小偃4号 Xiaoyan 4
102	中农28 Zhongnong 28	297	小偃759 Xiaoyan 759
103	中麦9号 Zhongmai 9	298	淮麦20 Huaimai 20
104	华北187 Huabei 187	299	淮麦22 Huaimai 22
105	邯99-6143 Han 99-6143	300	连麦2号 Lianmai 2
106	邯6172 Han 6172	301	南大2419 Nanda 2419
107	冀5265 Ji 5265	302	宛原-66 Wanyuan-66
108	冀麦15 Jimai 15	303	宛7107 Wan 7107
109	冀麦19 Jimai 19	304	潍麦8号 Weimai 8
110	冀麦1号 Jimai 1	305	潍麦6号 Weimai 6
111	冀麦20 Jimai 20	306	万年2号 Wannian 2
112	冀麦22 Jimai 22	307	徐州24 Xuzhou 24
113	冀麦23 Jimai 23	308	徐州14 Xuzhou 14
114	冀麦24 Jimai 24	309	徐州8号 Xuzhou 8
115	冀麦26 Jimai 26	310	徐州438 Xuzhou 438
116	冀麦36 Jimai 36	311	徐州25 Xuzhou 25
117	冀麦37 Jimai 37	312	扬麦14 Yangmai 14
118	冀麦38新系 Jimai 38 xinxi	313	扬麦18 Yangmai 18
119	冀麦3号 Jimai 3	314	扬麦16 Yangmai 16
120	冀麦6号(衡水714) Jimai 6 (Hengshui 714)	315	浙麦1号 Zhemai 1
121	冀麦7号 Jimai 7	316	宁麦13 Ningmai 13
122	津丰1号 Jinfeng 1	317	宁麦14 Ningmai 14
123	科红1号 Kehong 1	318	宁麦15 Ningmai 15
124	科信9号 Kexin 9	319	宁麦9 Ningmai 9
125	科遗26 Keyi 26	320	宁糯麦1号 Ningnuomai 1
126	莱州953 Laizhou 953	321	川麦36 Chuanmai 36
127	师栾02－1 Shiluan 02－1	322	川麦42 Chuanmai 42
128	石11-045(史) Shi 11-045 (Shi)	323	川麦8号 Chuanmai 8
129	石11-045(王) Shi 11-045 (Wang)	324	川育20 Chuanyu 20
130	石11-5139(史) Shi 11-5139 (Shi)	325	川育6号 Chuanyu 6
131	石11-5139(王) Shi 11-5139 (Wang)	326	绵麦39 Mianmai 39
132	石12-4025(史大) Shi 12-4025 (Shida)	327	绵麦41 Mianmai 41
133	石12-4025(王) Shi 12-4025 (Wang)	328	绵麦45 Mianmai 45
134	石14-5628(13-4365)(史) Shi 14-5628 (13-4365) (Shi)	329	绵阳11 Mianyang 11
135	石14-5628(王) Shi 14-5628 (Wang)	330	绵阳19 Mianyang 19
136	石4056(17石) Shi 4056 (17Shi)	331	绵阳20 Mianyang 20
137	石-4185 Shi-4185	332	绵阳38 Mianyang 38
138	石4056-3(17石) Shi 4056-3 (17Shi)	333	绵阳86-11 Mianyang 86-11
139	石4056-1(17石) Shi 4056-1 (17Shi)	334	内乡5号 Neixiang 5
140	石麦12 Shimai 12	335	内乡188 Neixiang 188
141	石家庄54 Shijiazhuang 54	336	内乡185 Neixiang 185
142	石家庄407 Shijiazhuang 407	337	克丰3号 Kefeng 3
143	石麦19 Shimai 19	338	克全 Kequan
144	石麦18 Shimai 18	339	皖麦19 Wanmai 19
145	石麦15 Shimai 15	340	马场2号 Machang 2
146	矮孟牛Ⅳ Aimengniu Ⅳ	341	皖麦50 Wanmai 50
147	石优20 Shiyou 20	342	皖麦33 Wanmai 33
148	石优17号 Shiyou 17	343	鄂恩1号 E'en 1
149	济麦19号 Jimai 19	344	鄂麦15 Emai 15
150	蒿城9411(c02667) Haocheng 9411(c02667)	345	鄂麦16 Emai 16
151	矮孟牛Ⅴ Aimengniu Ⅴ	346	鄂麦19 Emai 19
152	济麦20号 Jimai 20	347	鄂麦6号 Emai 6
153	济南13 Jinan 13	348	抗辉县红 Kanghuixianhong
154	济麦22号 Jimai 22	349	碧蚂4号 Bima 4
155	济麦21号 Jimai 21	350	长丰6号 Changfeng 6
156	济南4号 Jinan 4	351	东1-23 Dong 1-23
157	济南2号 Jinan 2	352	东1-32 Dong 1-32
158	济南16 Jinan 16	353	东2-23 Dong 2-23
159	济宁12 Jining 12	354	东2-8 Dong 2-8
160	济南9号 Jinan 9	355	高38 Gao 38
161	济南8号 Jinan 8	356	高原602(青602) Gaoyuan 602(Qing 602)
162	良星66 Liangxing 66	357	天平3号 Tianping 3
163	济宁16号 Jining 16	358	黑辐84S1378 Heifu 84S1378
164	济宁13 Jining 13	359	有芒红7号 Youmanghong 7
165	鲁麦11号 Lumai 11	360	新冬2号 Xindong 2
166	鲁215953 Lu 215953	361	新冬20 Xindong 20
167	聊麦16号 Liaomai 16	362	中任2号 Zhongren 2
168	鲁麦12 Lumai 12	363	丰产3号 Fengchan 3
169	鲁麦14(烟中1604) Lumai 14 (Yanzhong 1604)	364	波他姆S70 BotamuS 70
170	鲁麦15 Lumai 15	365	合作2号 Hezuo 2
171	鲁麦21 Lumai 21	366	松花江1号 Songhuajiang 1
172	鲁麦3号 Lumai 3	367	克丰2号 Kefeng 2
173	鲁麦5号 Lumai 5	368	克丰6号 Kefeng 6
174	鲁麦7号 Lumai 7	369	克旱7号 Kehan 7
175	鲁原301 Luyuan 301	370	克旱10号 Kehan 10
176	山农12号 Shannong 12	371	克强 Keqiang
177	山农15 Shannong 15	372	克壮 Kezhuang
178	山农辐63 Shannongfu 63	373	垦九10号 Kenjiu 10
179	泰农18 Tainong 18	374	龙麦26 Longmai 26
180	泰山1号 Taishan 1	375	龙麦33 Longmai 33
181	泰山23 Taishan 23	376	宁麦12 Ningmai 12
182	泰山4号 Taishan 4	377	定西24 Dingxi 24
183	泰山5号 Taishan 5	378	咸83(104)-11中“s” Xian 83(104)-11zhong “s”
184	泰山9818 Taishan 9818	379	川84-7045 Chuan 84-7045
185	腾S15 TengS 15	380	绵麦40 Mianmai 40
186	汶农六号 Wennong 6	381	绵麦43 Mianmai 43
187	烟农15 Yannong 15	382	绵麦46 Mianmai 46
188	烟农19号 Yannong 19	383	大头黄 Datouhuang
189	烟农22号 Yannong 22	384	鄂麦18 Emai 18
190	烟农23号 Yannong 23	385	南大96co76 Nanda 96co76
191	烟农24号 Yannong 24	386	长丰4号 Changfeng 4
192	洲元9369 Zhouyuan 9369	387	克旱13 Kehan 13
193	淄麦12号 Zimai 12	388	垦红14 Kenhong 14
194	晋麦21 Jinmai 21	389	合作4号 Hezuo 4
195	晋麦30 Jinmai 30

Table S1

Fig. 4

Fig. 5

Table 5

Table 6

QTL loci significantly associated with low-phosphorus tolerance related traits"

位点 Loci	置信区间 Confidence interval (bp)	性状 Trait	染色体 Chromosome	峰值SNP Peak SNP	物理位置 Position (bp)	P值 -log₁₀P	贡献率 R²(%)
LLPTT2A.1	172,021,064-173,698,784	E1-SH	2A	AX-111275849	172,859,924	1.00E-05	5.90
		E2-RD	2A	AX-111275849	172,859,924	4.69E-06	7.70
		E2-RTN	2A	AX-111275849	172,859,924	2.01E-06	7.91
LLPTT2A.2	651,505,509-653,078,373	E1-SH	2A	AX-109458041	652,291,941	3.57E-09	7.71
		E3-MRL	2A	AX-109458041	652,291,941	7.11E-05	4.20
		E3-RTN	2A	AX-109458041	652,291,941	7.03E-05	4.20
		BLUP-SH	2A	AX-109458041	652,291,941	3.61E-05	4.55
		BLUP-RN	2A	AX-109458041	652,291,941	2.42E-05	4.74
LLPTT2D.1	605,713,612-606,915,404	E3-SDW	2D	AX-110533652	606,693,178	6.71E-05	4.28
LLPTT2D.1	605,713,612-606,915,404	BLUP-SDW	2D	AX-110361781	605,852,678	2.99E-05	5.50
LLPTT3A.1	522,400,434-528,592,728	E1-RTN	3A	AX-109854803	523,151,974	5.00E-05	5.11
LLPTT3A.1	522,400,434-528,592,728	E3-RD	3A	AX-109039518	527,907,394	3.82E-05	5.49
LLPTT3A.2	528,596,882-532,633,996	E1-RTN	3A	AX-110185182	530,627,769	6.00E-05	4.34
LLPTT3A.2	528,596,882-532,633,996	E3-RD	3A	AX-109381718	530,590,416	2.40E-05	5.68
LLPTT3A.3	556,676,504-567,980,595	E1-RTN	3A	AX-110939649	567,143,453	1.00E-04	4.09
LLPTT3A.3	556,676,504-567,980,595	E3-RD	3A	AX-109505161	561,734,860	2.42E-05	5.97
LLPTT3B.1	708,368,449-719,084,208	E3-RN	3B	AX-94583652	717,304,474	4.11E-05	4.97
LLPTT3B.1	708,368,449-719,084,208	BLUP-RN	3B	AX-94583652	717,304,474	9.21E-05	4.38
LLPTT4A.1	564,593,256-566,270,976	E1-RSA	4A	AX-94925936	565,432,116	5.00E-05	4.79
		E1-RTN	4A	AX-94925936	565,432,116	1.58E-08	9.27
		E1-TRL	4A	AX-94925936	565,432,116	5.00E-05	4.77
		E2-SH	4A	AX-94925936	565,432,116	3.15E-08	9.51
LLPTT4B.1	142,811,405-172,520,416	E2-RD	4B	AX-108833248	145,374,185	4.20E-05	6.72
LLPTT4B.1	142,811,405-172,520,416	BLUP-MRL	4B	AX-108833248	145,374,185	7.63E-05	5.16
LLPTT4D.1	7,036,268-8,713,990	E1-RSA	4D	AX-109557110	7,875,129	1.20E-06	7.47
LLPTT4D.1	7,036,268-8,713,990	E2-SH	4D	AX-109557110	7,875,129	1.33E-06	8.20
LLPTT4D.2	9,410,565-11,088,287	E1-RSA	4D	AX-109005677	10,249,426	8.00E-05	5.22
		E1-RTN	4D	AX-109005677	10,249,426	4.00E-05	5.97
		E2-SH	4D	AX-109005677	10,249,426	1.01E-07	9.77
		BLUP-SH	4D	AX-109005677	10,249,426	1.78E-05	6.28
LLPTT5A.1	564,494,504-565,403,678	E1-RSA	5A	AX-109951485	564,534,428	6.00E-05	5.15
LLPTT5A.1	564,494,504-565,403,678	E2-SH	5A	AX-110511778	564,660,778	6.67E-05	4.77
LLPTT5A.2	667,361,083-669,563,683	E1-SH	5A	AX-108955444	668,223,726	3.00E-05	5.00
LLPTT5A.2	667,361,083-669,563,683	BLUP-SDW	5A	AX-108955444	668,223,726	9.38E-05	4.29
LLPTT5A.3	669,585,870-670,535,062	E1-SH	5A	AX-108803792	669,800,485	4.00E-05	4.68
		E2-RDW	5A	AX-109879786	669,900,461	5.65E-05	5.36
		E3-RD	5A	AX-109350840	669,733,200	8.51E-05	5.24
		BLUP-SDW	5A	AX-110502745	669,624,184	7.53E-06	5.59
LLPTT5B.1	78,288,597-82,206,709	E1-SH	5B	AX-112289745	78,708,029	1.12E-09	10.57
		E3-MRL	5B	AX-112289745	78,708,029	9.37E-05	4.37
		E3-RD	5B	AX-111043315	81,787,277	7.04E-05	5.41
		BLUP-SH	5B	AX-112289745	78,708,029	3.47E-07	7.24
LLPTT5B.2	288,485,939-315,413,719	E2-RSA	5B	AX-111507396	308,618,491	5.82E-05	5.59
LLPTT5B.2	288,485,939-315,413,719	E3-RD	5B	AX-109867936	309,385,307	8.07E-05	5.01
LLPTT5B.3	403,717,589-407,635,520	E3-RD	5B	AX-109499442	404,998,534	6.77E-05	5.12
LLPTT5B.3	403,717,589-407,635,520	BLUP-RD	5B	AX-111557664	405,724,930	6.10E-05	4.26
LLPTT5B.4	489,841,220-490,759,761	E2-RD	5B	AX-108955618	490,738,277	4.08E-06	6.39
LLPTT5B.4	489,841,220-490,759,761	BLUP-RD	5B	AX-110507901	489,987,304	1.02E-05	5.95
LLPTT5D.1	224,743,268-226,420,990	E1-SH	5D	AX-111190797	225,582,129	3.57E-09	10.55
		E3-SH	5D	AX-111190797	225,582,129	2.50E-05	5.05
		BLUP-SH	5D	AX-111190797	225,582,129	2.50E-09	10.58
LLPTT5D.2	496,086,888-497,869,466	E3-MRL	5D	AX-89451602	496,978,177	4.28E-05	5.02
LLPTT5D.2	496,086,888-497,869,466	BLUP-MRL	5D	AX-89451602	496,978,177	7.69E-06	6.00
LLPTT6A.1	522,639,187-526,115,905	E1-MRL	6A	AX-111198037	524,832,883	1.00E-05	5.18
LLPTT6A.1	522,639,187-526,115,905	E3-RD	6A	AX-111056482	525,033,540	4.99E-05	5.37
LLPTT6B.1	526,476,834-534,466,335	E2-MRL	6B	AX-109512103	533,765,104	8.33E-05	4.49
LLPTT6B.1	526,476,834-534,466,335	E3-RD	6B	AX-110046271	529,767,951	9.87E-05	5.05
LLPTT6D.1	128,190,014-129,867,736	E1-SH	6D	AX-110373854	129,028,875	8.72E-07	6.72
		E2-RTN	6D	AX-110373854	129,028,875	8.41E-06	6.18
		E3-MRL	6D	AX-110373854	129,028,875	4.29E-05	4.66
		BLUP-SH	6D	AX-110373854	129,028,875	2.66E-05	4.80
		BLUP-RN	6D	AX-110373854	129,028,875	4.23E-05	4.59
LLPTT6D.2	266,362,121-268,039,843	E1-SH	6D	AX-94503640	267,200,982	3.69E-09	9.66
		E2-RD	6D	AX-94503640	267,200,982	2.68E-06	6.68
		E2-RTN	6D	AX-94503640	267,200,982	7.77E-08	8.92
		E3-SH	6D	AX-94503640	267,200,982	6.63E-05	4.36
		E3-MRL	6D	AX-94503640	267,200,982	4.12E-05	4.80
		BLUP-SH	6D	AX-94503640	267,200,982	4.18E-08	8.35
		BLUP-RN	6D	AX-94503640	267,200,982	1.87E-05	5.23
		BLUP-RDW	6D	AX-94503640	267,200,982	7.49E-05	4.33
LLPTT6D.3	428,891,281-428,923,196	E1-RTN	6D	AX-108961753	428,891,281	6.00E-05	4.99
LLPTT6D.3	428,891,281-428,923,196	E2-SH	6D	AX-108961753	428,891,281	7.70E-06	6.43
LLPTT7A.1	130,292,122-178,902,926	E3-RD	7A	AX-108941925	154,061,783	5.62E-05	5.32
LLPTT7A.1	130,292,122-178,902,926	BLUP-RTN	7A	AX-111767484	143,715,554	7.34E-05	4.24
LLPTT7A.2	681,702,374-685,505,945	E3-RN	7A	AX-111065932	681,754,446	5.86E-05	4.61
LLPTT7A.2	681,702,374-685,505,945	BLUP-RN	7A	AX-110533724	681,915,668	2.91E-05	4.83

Table 6

Table 7

Fig. 6

Fig. S1

Fig. 7

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[50]	Shi J C, Zhao B Y, Jin R, Hou L, Zhang X W, Dai H L, Yu N, Wang E T. A phosphate starvation response-regulated receptor-like kinase, OsADK1, is required for mycorrhizal symbiosis and phosphate starvation responses. New Phytol, 2022, 236: 2282-2293. doi: 10.1111/nph.18546 pmid: 36254112

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化学式 Chemical formula	浓度 Concentration (mmol L^-1)	化学式 Chemical formula	浓度 Concentration (mmol L^-1)
Ca(NO₃)₂·4H₂O	1.0	H₃BO₃	0.00100
KH₂PO₄	对照组0.2, 低磷组0.005 CK 0.2, LP 0.005	Na₂MOO₄·2H₂O	0.00015
MgSO₄·7H₂O	1.0	MnSO₄·H₂O	0.00100
KCl	1.5	ZnSO₄·7H₂O	0.00100
CaCl₂	1.5	CuSO₄·5H₂O	0.00050
Fe-EDTA	0.1

引物名称Primer name	引物序列Primer sequence (5'-3')
TraesCS4D02G022900-F	ACTGGAAAGGGGCGCTTT
TraesCS4D02G022900-R	GAACCATTCATCTTGCCCCC
TraesCS4D02G023300-F	GGGATGGAGGACTTCGACAA
TraesCS4D02G023300-R	ACTGTAGGGGACGCAAGGAA
TraesCS6D02G154700-F	ATATCGCGCACTGGACTCAC
TraesCS6D02G154700-R	ATATCGCGCACTGGACTCAC
TaActin-F	TGGTGTCATCAAGCCTGGTATGGT
TaActin-R	ACTCATGGTGCATCTCAACGGACT

性状Trait	处理Treatment (T)	基因型Genotype (G)	处理×基因型T×G
苗高SH	***	***	***
主根长MRL	***	***	***
根条数RN	***	***	***
茎叶干重SDW	***	***	***
根干重RDW	***	***	***
总根长TRL	***	***	***
根面积RSA	***	***	***
根直径RD	**	***	***
根尖数RTN	***	***	***

QTL	候选基因ID Candidate gene ID	基因位置 Gene position (bp)	基因功能注释 Gene function
LLPTT2A.2	TraesCS2A02G397700	651,504,966-651,510,544	组蛋白H1 Histone H1
	TraesCS2A02G398500	651,801,546-651,803,391	Gag多聚蛋白 Gag polyprotein
	TraesCS2A02G399900	652,999,871-653,002,090	解毒蛋白 Protein DETOXIFICATION
LLPTT4D.2	TraesCS4D02G022900	9,591,823-9,593,712	F-box家族蛋白 F-box family protein
	TraesCS4D02G023300	10,010,190-10,010,960	F-box家族蛋白 F-box family protein
	TraesCS4D02G025500	10,991,060-10,993,177	整合膜HPP家族蛋白 Integral membrane HPP family protein
LLPTT5B.1	TraesCS5B02G069300	79,081,294-79,082,277	HVA22-like蛋白 HVA22-like protein
LLPTT6D.1	TraesCS6D02G154700	129,785,419-129,791,207	类受体蛋白激酶 Receptor-like protein kinase
LLPTT6D.2	TraesCS6D02G193100	267,236,634-267,237,837	高亲和性硝酸盐转运蛋白 High affinity nitrate transporter
LLPTT6D.2	TraesCS6D02G193200	267,514,716-267,515,174	高亲和性硝酸盐转运蛋白 High affinity nitrate transporter

Identification of genetic loci related to low phosphorus tolerance at the seedling stage in wheat and analysis of candidate genes

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环境 Environment	SNP数量 Number of SNP	染色体 Chromosome	P值 -log₁₀P	贡献率 R²(%)	QTL数量 Number of QTL
E1	477	1A, 1B, 1D, 2A, 2B, 2D, 3A, 3B, 3D, 4A, 4B, 4D, 5A, 5B, 5D, 6A, 6B, 6D, 7A, 7B, 7D	9.99E-05-1.12E-09	4.01-10.57	90
E2	163	1A, 1B, 2A, 2B, 2D, 3A, 3B, 3D, 4A, 4B, 4D, 5A, 5B, 5D, 6A, 6B, 6D, 7A, 7B, 7D	9.94E-05-3.15E-08	4.32-9.77	86
E3	378	1A, 1B, 1D, 2A, 2B, 2D, 3A, 3B, 3D, 4A, 4B, 4D, 5A, 5B, 5D, 6A, 6B, 6D, 7A, 7B, 7D	9.99E-05-3.45E-07	4.02-8.04	215
BLUP	179	1A, 1B, 2A, 2B, 2D, 3B, 3D, 4A, 4B, 4D, 5A, 5B, 5D, 6A, 6B, 6D, 7A, 7B	9.99E-05-2.50E-09	4.02-10.58	73

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