水旱条件下小麦持绿相关性状QTL定位

doi:10.3724/SP.J.1006.2024.41022

Abstract

Abstract:

Drought is the primary abiotic stress that significantly affects wheat production. Extending the duration of leaf greenness, thereby increasing photosynthetic time and efficiency, is crucial for wheat in ensuring organic matter accumulation under drought stress and ultimately stabilizing wheat yield. Understanding the developmental and genetic characteristics of flag leaf greenness retention in wheat and identifying stable molecular markers closely associated with greenness-related genes, independent of environmental influences, can significantly accelerate the breeding process for drought resistance. In this study, we utilized a DH population consisting of 174 lines derived from Yangmai 16 and Zhongmai 895 as experimental materials. We conducted phenotypic evaluations of the percentage of green leaf area (GLA) and relative chlorophyll content (SPAD value) in flag leaves at 10 d, 14 d, 18 d, 22 d, 26 d, and 30 d post-flowering under two moisture environments: normal drip irrigation (NI) and drought stress (DS). We simulated the change in GLA using the Gompertz model and performed QTL mapping for eleven greenness-related traits, including time to maximum senescence rate (TMRS), green leaf area duration (GLAD), average senescence rate (ARS), time to the beginning of rapid senescence (T1), time to the end of rapid senescence (T2), and the dynamic SPAD value of the flag leaf. The results revealed a phenomenon of transgressive segregation in GLA, aging characteristic parameters, and dynamic SPAD values of the DH population and parents under both well-watered and drought conditions, demonstrating certain differences. A consistent trend of slow-rapid-slow decline in GLA was observed at different time points across all families and parents, with rapid senescence primarily occurring at 18 d, 22 d, and 26 d Except for the negative correlation between ARS and GLA-10D, positive correlations were observed among all other traits. The heritability of each trait ranged from 0.50 to 0.81. Linkage analysis identified a total of 27 stable loci associated with greenness retention in wheat across two or more environments. Among these, 11 loci were associated with wheat senescence characterization trait parameters, and 16 loci were associated with dynamic SPAD values of the flag leaf. These loci were distributed on chromosomes 1A, 1B, 4B, 4D, 5D, 7B, and 7D, explaining 3.86%-14.11% and 2.99%-17.45% of the phenotypic variance, respectively. One QTL regulating T1 and five SPAD values on chromosomes 4B, 4D, and 7B were consistently detected across the three environments, explaining 8.97%-14.11% and 6.85%-17.45% of the phenotypic variance, respectively. The results of QTL co-localization effect analysis revealed that loci containing Rht-D1b and AA genotypes exhibited significant enhancement and cumulative effects on SPAD-10D, while loci containing GG and TT genotypes showed significant enhancement and cumulative effects on SPAD-18D. Four QTL clusters with significant effects were identified on chromosomes 4B, 4D, 7B, and 7D. Notably, the segment of the QTL cluster on chromosome 4D (18.80-28.58 Mb) remained unaffected by water availability and contained the Rht-D1 gene. The loci regulating SPAD-0D and SPAD-10D might be influenced by the multiplicative effect of this gene. Candidate gene analysis identified eight candidate genes, such as TraesCS4D01G054000, TraesCS1B01G434300, and TraesCS7B01G010200, which are associated with greenness retention. These findings provide a theoretical foundation for molecular marker-assisted breeding aimed at improving greenness retention in wheat.

Key words: wheat, stay-green-related traits, QTL localization, convergent effect, candidate genes

CHEN Chen, CHENG Yu-Kun, WANG Wei, REN Yi, ZHANG Hai-Yan, CHEN Hui-Bo, GENG Hong-Wei. QTL mapping of stay-green-related traits in wheat under drought condition[J].Acta Agronomica Sinica, 2024, 50(11): 2684-2698.

Figures/Tables 10

Table S1

Statistical analysis of the proportion of green leaf area in the flag leaf of DH population and parents"

性状 Trait	环境 Environment	处理 Treatment	亲本 Parents		DH家系 DH Lines
性状 Trait	环境 Environment	处理 Treatment	扬麦16 Yangmai 16	中麦895 Zhongmai 895	最小值 Min.	最大值 Max.	平均值 Mean	标准差 SD	变异系数 CV	偏度 Skew.	峰度 Kurt.
GLA-10D	E1	NI	10.00	10.00	9.10	10.00	9.94^*	0.17	0.02	‒2.92	8.46
		DS	9.80	9.80	7.70	10.00	9.72	0.47	0.05	‒2.45	6.13
	E2	NI	10.00	10.00	9.10	10.00	9.98	0.09	0.01	‒6.80	54.47
		DS	9.80	10.00	8.80	10.00	9.97	0.14	0.01	‒6.21	43.19
	BLUE	NI	10.00	10.00	9.20	10.00	9.96^*	0.11	0.01	‒3.68	17.89
		DS	9.80	9.90	8.50	10.04	9.85	0.26	0.03	‒2.55	7.19
GLA-14D	E1	NI	9.60	10.00^*	7.20	10.00	9.51^*	0.58	0.06	‒1.82	3.23
		DS	7.60	8.80	4.00	10.00	8.63	1.30	0.15	‒1.50	2.17
	E2	NI	9.90	10.00	8.70	10.00	9.94	0.18	0.02	‒4.68	23.94
		DS	9.70	10.00^*	8.30	10.00	9.93	0.22	0.02	‒4.51	24.31
	BLUE	NI	9.75	10.00^*	8.60	10.00	9.73^*	0.31	0.03	‒1.74	2.76
		DS	8.65	9.40^*	6.95	10.09	9.28	0.67	0.07	‒1.47	2.01
GLA-18D	E1	NI	6.30	7.90^*	3.00	10.00	7.80^*	1.58	0.20	‒0.87	0.16
		DS	2.90	4.60^*	0.50	9.20	4.55	2.26	0.50	0.11	‒0.95
	E2	NI	9.40	10.00^*	6.60	10.00	9.72^*	0.49	0.05	‒3.35	14.61
		DS	8.80	10.00^*	7.50	10.00	9.61	0.55	0.06	‒1.95	3.78
	BLUE	NI	7.85	8.95	6.35	10.00	8.76^*	0.82	0.09	‒0.86	0.08
		DS	5.85	7.30	4.30	9.60	7.08	1.18	0.17	0.00	‒0.78
GLA-22D	E1	NI	2.80	4.00^*	0.50	9.60	4.42^*	2.14	0.49	0.40	‒0.83
		DS	0.90	1.50^*	0.00	7.20	2.21	1.67	0.75	0.58	‒0.55
	E2	NI	8.50	10.00^*	6.10	10.00	9.08^*	0.86	0.09	‒1.06	0.84
		DS	7.80	9.70^*	4.00	9.90	7.83	1.31	0.17	‒0.53	‒0.44
	BLUE	NI	5.65	7.00	3.80	9.47	6.75^*	1.19	0.18	0.14	‒0.75
		DS	4.35	5.60	2.00	8.30	5.02	1.19	0.24	0.01	‒0.35
GLA-26D	E1	NI	0.50	1.70^*	0.00	7.60	1.93^*	1.44	0.75	1.27	2.04
		DS	0.00	0.90^*	0.00	2.80	0.41	0.62	1.54	1.80	2.94
	E2	NI	5.70	6.80^*	1.50	8.80	5.56^*	1.00	0.18	0.25	1.65
		DS	4.30	5.50^*	1.00	7.40	3.54	1.71	0.48	0.09	‒0.97
	BLUE	NI	3.10	4.25	1.80	7.07	3.75^*	0.99	0.26	0.73	0.52
		DS	2.15	3.20	0.50	4.75	1.97	1.00	0.51	0.23	‒0.69
GLA-30D	E2	NI	1.10	2.60^*	0.00	4.80	1.64^*	1.08	0.66	0.87	0.45
GLA-30D	E2	DS	0.90	1.20	0.00	4.60	0.82	0.64	0.78	1.48	6.11

Table S1

Fig. 1

Table 1

Statistical analysis of DH population and parental aging parameters"

性状 Trait	环境 Environ- ment	处理 Treatment	亲本 Parents		DH家系 DH line
性状 Trait	环境 Environ- ment	处理 Treatment	扬麦16 Yangmai 16	中麦895 Zhongmai 895	最小值 Min.	最大值 Max.	平均值 Mean	标准差 SD	变异系数 CV	偏度 Skew.	峰度 Kurt.
TMRS (d)	E1	NI	20.65	22.50^*	13.93	31.27	22.71^*	2.81	0.12	0.22	0.94
		DS	16.84	18.82^*	8.71	25.19	18.95	2.64	0.14	-0.42	0.74
	E2	NI	27.69	29.02^*	24.08	30.81	27.83^*	1.16	0.04	0.37	0.36
		DS	26.03	27.61^*	22.29	30.92	25.93	1.55	0.06	-0.05	-0.29
	BLUE	NI	24.17	25.76	20.08	30.10	25.27^*	1.67	0.07	0.28	0.48
		DS	21.43	23.21	16.86	27.35	22.45	1.76	0.08	-0.22	0.11
GLAD (d)	E1	NI	26.95	29.39^*	19.74	40.53	28.76^*	3.60	0.13	0.87	1.37
		DS	23.38	25.33^*	16.39	31.99	23.95	3.15	0.13	0.02	-0.80
	E2	NI	32.28	33.39	27.28	39.02	32.32^*	2.01	0.06	0.29	0.63
		DS	31.14	31.67	25.20	37.11	30.07	2.16	0.07	0.10	-0.14
	BLUE	NI	29.61	31.39	25.67	38.44	30.55^*	2.28	0.07	0.59	0.78
		DS	27.26	28.50	21.33	33.00	27.01	2.09	0.08	-0.02	-0.29
ARS (% d^-1)	E1	NI	-0.41	-0.38^*	-0.67	-0.27	-0.38^*	0.06	0.15	-1.35	4.70
		DS	-0.52	-0.45^*	-1.30	-0.32	-0.47	0.11	0.24	-3.99	22.73
	E2	NI	-0.31	-0.31	-0.37	-0.26	-0.32^*	0.02	0.06	-0.13	0.12
		DS	-0.32	-0.32	-0.41	-0.27	-0.34	0.03	0.07	-0.06	0.10
	BLUE	NI	-0.36	-0.34	-0.52	-0.28	-0.35^*	0.03	0.09	-1.08	4.13
		DS	-0.42	-0.39	-0.75	-0.32	-0.41	0.06	0.14	-3.16	15.50
T1 (d)	E1	NI	18.93	20.61^*	10.08	29.36	21.06^*	2.99	0.14	-0.23	1.22
		DS	15.05	17.04^*	4.59	23.91	17.58	2.84	0.16	-0.85	2.20
	E2	NI	26.43	27.82^*	22.97	29.75	26.60^*	1.06	0.04	0.35	1.25
		DS	24.62	26.50^*	20.94	29.23	24.79	1.49	0.06	-0.09	-0.39
	BLUE	NI	22.68	24.22	18.01	28.85	23.82^*	1.70	0.07	0.01	0.57
		DS	19.84	21.77	15.30	25.82	21.20	1.79	0.08	-0.40	0.43
T2 (d)	E1	NI	22.38	24.39^*	17.78	33.18	24.37^*	2.81	0.12	0.59	0.94
		DS	18.63	20.60^*	12.82	26.46	20.32	2.60	0.13	-0.10	-0.31
	E2	NI	28.95	30.22^*	25.20	32.32	29.06^*	1.33	0.05	0.33	-0.05
		DS	27.43	28.72^*	23.43	32.61	27.06	1.66	0.06	-0.01	-0.27
	BLUE	NI	25.66	27.30	22.15	32.39	26.71^*	1.74	0.07	0.46	0.47
		DS	23.03	24.66	18.41	28.90	23.70	1.78	0.08	-0.09	-0.09

Table 1

Table S2

Statistical analysis of DH population and parental SPAD values"

性状 Trait	环境 Environment	处理 Treatment	亲本 Parents		DH家系 DH Lines
性状 Trait	环境 Environment	处理 Treatment	扬麦16 Yangmai 16	中麦895 Zhongmai 895	最小值 Min.	最大值 Max.	平均值 Mean	标准差 SD	变异系数 CV	偏度 Skew.	峰度 Kurt.
SPAD-0D	E1	NI	59.21	59.97	54.92	67.10	61.43	2.45	0.04	‒0.45	0.26
	E1	DS	58.49	62.09^*	54.99	66.86	61.19	2.31	0.04	‒0.04	‒0.05
	E2	NI	59.66	60.42	53.36	64.38	59.33^*	1.96	0.03	‒0.09	0.32
	E2	DS	57.20	58.39	51.62	63.60	58.86	2.17	0.04	‒0.58	0.51
	BLUE	NI	59.43	60.19	54.92	63.99	60.39	1.91	0.03	‒0.60	0.20
	BLUE	DS	57.85	60.24^*	53.30	63.94	60.02	1.94	0.03	‒0.43	0.09
SPAD-10D	E1	NI	59.63	59.49	54.69	68.00	62.48	2.63	0.04	‒0.33	‒0.11
	E1	DS	61.59	67.49^*	55.17	70.06	62.72	2.76	0.04	0.03	‒0.09
	E2	NI	60.31	63.15^*	55.60	65.71	60.89^*	2.18	0.04	0.03	‒0.49
	E2	DS	58.14	62.35^*	55.70	65.29	60.26	2.00	0.03	‒0.05	‒0.37
	BLUE	NI	59.97	61.32	56.79	65.69	61.69^*	2.04	0.03	‒0.10	‒0.78
	BLUE	DS	59.87	64.92^*	56.73	67.52	61.48	2.02	0.03	0.19	0.06
SPAD-14D	E1	NI	57.24	62.31^*	52.54	70.12	61.76^*	3.27	0.05	‒0.18	0.11
	E1	DS	57.78	58.75	48.51	67.22	59.28	3.88	0.07	‒0.39	‒0.10
	E2	NI	59.64	62.21	54.21	65.83	60.26	2.46	0.04	‒0.19	‒0.37
	E2	DS	59.90	63.29^*	52.85	67.15	60.36	2.91	0.05	‒0.26	‒0.59
	BLUE	NI	58.44	62.26^*	54.96	67.06	61.01	2.54	0.04	‒0.14	‒0.37
	BLUE	DS	58.84	61.02	51.90	65.19	59.82	2.79	0.05	‒0.34	‒0.47
SPAD-18D	E1	NI	48.60	57.59^*	40.34	67.61	56.29^*	5.04	0.09	‒0.59	0.41
	E1	DS	28.86	43.55^*	7.68	65.41	48.49	9.49	0.20	‒0.81	1.33
	E2	NI	57.90	59.93^*	50.93	65.89	59.03^*	2.63	0.04	‒0.10	‒0.35
	E2	DS	56.68	60.49	50.61	64.82	58.18	2.55	0.04	‒0.27	‒0.22
	BLUE	NI	53.25	58.76^*	48.66	64.23	57.68^*	3.34	0.06	‒0.38	‒0.34
	BLUE	DS	42.77	52.02	35.59	63.45	53.38	5.22	0.10	‒0.54	0.20
SPAD-22D	E1	NI	32.36	43.49^*	16.52	65.24	44.34^*	9.65	0.22	‒0.43	‒0.09
	E1	DS	8.26	15.36^*	4.30	54.44	25.04	11.43	0.46	0.26	‒0.75
	E2	NI	49.19	54.03^*	37.76	63.07	53.53^*	4.82	0.09	‒0.66	0.37
	E2	DS	38.04	52.33^*	11.29	61.09	45.88	11.88	0.26	‒0.98	0.38
	BLUE	NI	40.77	48.76	30.56	62.94	48.97^*	6.26	0.13	‒0.39	‒0.07
	BLUE	DS	23.15	33.85	9.40	54.56	35.46	9.43	0.27	‒0.34	‒0.11
SPAD-26D	E1	NI	6.45	11.92^*	5.52	47.54	16.27^*	7.63	0.47	1.36	3.13
	E1	DS	3.84	5.38^*	2.97	25.72	8.30	3.39	0.41	1.57	4.09
	E2	NI	33.14	43.16^*	17.11	57.05	33.86^*	8.58	0.25	0.32	‒0.25
	E2	DS	27.06	35.05^*	6.24	56.54	18.99	13.07	0.69	1.15	0.27
	BLUE	NI	19.80	27.54	12.74	48.81	25.10^*	6.55	0.26	0.66	1.02
	BLUE	DS	15.45	20.22	3.12	35.17	13.63	7.21	0.53	1.12	0.33
SPAD-30D	E2	NI	6.20	11.81^*	3.27	44.51	11.29^*	6.44	0.57	1.94	5.22
SPAD-30D	E2	DS	4.27	8.09	1.90	31.35	7.61	3.95	0.52	3.10	14.24

Table S2

Fig. 2

Table S3

Table 2

Stabilizing QTL loci for senescence characterization parameters and SPAD values in different moisture environments"

位点 QTL	环境 Environment	处理 Treatment	左标记 Left marker	右标记 Right marker	置信区间 Confidence interval (cM)	位置 Position (Mb)	LOD值 LOD value	表型贡献率 PVE (%)	加性效应 Additive effect
QTMRS.xjau-4B	E1/BLUE	NI	AX-110194463	AX-111651924	120.5‒122.0	7.21‒19.03	4.72‒6.06	10.34‒11.46	0.53‒1.04
QTMRS.xjau-5D	E1/BLUE	NI	AX-111162962	AX-110717870	163.5‒168.5	459.07‒465.11	3.68‒4.28	7.26‒7.68	0.46‒0.83
QTMRS.xjau-4D	E2/BLUE	DS	AX-108944764	AX-94558069	52.5‒53.5	15.91‒16.59	7.35‒8.85	11.46‒11.93	‒0.65‒-0.59
QTMRS.xjau-7D	E2/BLUE	DS	AX-109167420	AX-111918348	109.5‒112.5	72.30‒77.23	3.17‒3.62	3.86‒5.31	0.37‒0.41
QGLAD.xjau-4D	E2/BLUE	DS	AX-108944764	Rht-D1	52.5‒55.5	15.91‒18.80	6.39‒10.61	6.97‒10.72	‒0.90‒-0.72
QT1.xjau-4B	E1/BLUE	NI	AX-86170717	AX-111651924	120.5‒122.0	7.21‒22.87	5.01‒6.11	10.13‒11.68	0.55‒1.10
QT1.xjau-5D	E1/BLUE	NI	AX-111162962	AX-110035093	161.5‒168.5	459.07‒464.13	4.75‒4.97	8.16‒9.32	0.53‒0.92
QT1.xjau-4D	E1/E2/BLUE	DS	AX-95659047	AX-94558069	49.5‒53.5	12.77‒16.59	7.47‒9.55	8.97‒14.11	-0.99‒-0.55
QT1.xjau-7D	E2/BLUE	DS	AX-109167420	AX-111918348	109.5‒112.5	72.30‒77.23	5.01‒5.51	6.56‒7.00	0.47‒0.52
QT2.xjau-4D	E2/BLUE	DS	AX-108944764	AX-94558069	52.5‒53.5	15.91‒16.59	9.38‒9.41	6.34‒12.58	-0.72‒-0.66
QT2.xjau-7B	E1/BLUE	DS	AX-111478329	AX-108883282	1.5‒2.5	1.21‒4.47	3.11‒5.80	5.94‒7.74	-0.65‒-0.56
QSPAD-0D.xjau-4D.1	E2/BLUE	NI	Rht-D1	AX-109445215	56.5‒60.5	18.80‒28.58	4.24‒5.10	3.64‒11.65	-0.73‒-0.59
QSPAD-0D.xjau-4D.2	E2/BLUE	DS	AX-89421921	Rht-D1	54.5‒55.5	18.80‒19.29	5.35‒7.80	9.37‒12.20	-0.90‒-0.63
QSPAD-10D.xjau-4D.1	E1/BLUE	NI	AX-89421921	Rht-D1	54.5‒55.5	18.80‒19.29	6.15‒10.69	9.06‒14.86	-0.95‒-0.91
QSPAD-10D.xjau-1A	E1/BLUE	DS	AX-94629887	AX-109406391	0.0‒0.5	8.64‒20.09	4.30‒5.82	4.86‒8.37	0.56‒0.94
QSPAD-10D.xjau-4B	E1/E2/BLUE	DS	AX-95685381	AX-94912823	43.5‒46.5	171.62‒172.72	5.59‒10.27	8.79‒14.06	0.86‒0.89
QSPAD-10D.xjau-4D.2	E1/BLUE	DS	AX-89421921	Rht-D1	54.5‒55.5	18.80‒19.29	7.13‒13.05	10.72‒17.18	-0.98‒-0.98
QSPAD-14D.xjau-4B	E2/BLUE	NI	AX-95685381	AX-108888649	43.5‒44.5	171.62‒263.38	8.39‒11.51	12.81‒17.04	1.22‒1.39
QSPAD-14D.xjau-4D	E2/BLUE	NI	AX-111071805	AX-111662392	61.5‒63.5	27.15‒33.06	6.03‒10.24	9.64‒15.73	-1.12‒-0.80
QSPAD-18D.xjau-4B	E1/E2/BLUE	NI	AX-95685381	AX-108888649	43.5‒44.5	171.62‒263.38	6.27‒8.18	9.75‒13.49	1.13‒1.69
QSPAD-18D.xjau-4D	E1/E2/BLUE	NI	AX-95659047	AX-108944764	47.5‒52.5	12.77‒15.91	6.75‒13.27	10.92‒17.45	-2.18‒-1.01
QSPAD-18D.xjau-7B	E1/BLUE	DS	AX-111581943	AX-110629026	0.0‒1.5	6.07‒7.00	3.33‒4.66	2.99‒7.05	‒2.36‒‒0.90
QSPAD-22D.xjau-4D	E1/E2/BLUE	DS	AX-108944764	AX-109478820	52.5‒57.5	15.91‒30.66	5.26‒9.05	7.21‒12.81	-4.45‒-3.75
QSPAD-22D.xjau-7D	E2/BLUE	DS	AX-110941147	AX-111918348	111.5‒112.5	76.62‒77.23	4.50‒6.25	5.02‒5.94	2.57‒3.78
QSPAD-26D.xjau-7B	E1/E2/BLUE	NI	AX-111581943	AX-110629026	0.0‒1.5	6.07‒7.00	3.42‒8.74	6.85‒13.27	-2.70‒-2.50
QSPAD-26D.xjau-1B	E2/BLUE	DS	AX-110534755	AX-111590092	128.5‒129.5	653.94‒665.05	2.90‒4.64	6.60‒10.30	-3.05‒-2.39
QSPAD-26D.xjau-4D	E2/BLUE	DS	AX-89421921	Rht-D1	54.5‒57.5	18.80‒19.29	4.21‒5.48	8.06‒9.76	-3.41‒-2.35

Table 2

Fig. 3

Fig. 4

Table 3

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性状 Trait	均方值Mean Square
性状 Trait	基因型 Genotype (G)	环境 Environment (E)	G×E互作 G×E	H²
%GLA-10D	0.22^***	5.07^***	0.11^***	0.60
%GLA-14D	1.56^***	131.19^***	0.86^***	0.60
%GLA-18D	6.49^***	2017.33^***	3.15^***	0.60
%GLA-22D	8.65^***	3441.3^***	3.51^***	0.66
%GLA-26D	5.39^***	1698.58^***	2.36^***	0.64
%GLA-30D	2.04^***	115.07^***	1.09^***	0.57
SPAD-0D	25.05^***	575.43^***	4.81	0.81
SPAD-10D	26.64^***	494.54^***	6.52^***	0.78
SPAD-14D	47.01^***	353.41^***	10.88^***	0.79
SPAD-18D	126.52^***	7855.07^***	40.73^***	0.74
SPAD-22D	372.19^***	51022.18^***	129.11^***	0.73
SPAD-26D	266.73^***	39912.29^***	116.02^***	0.68
SPAD-30D	62.44^***	2345.09^***	50.92^***	0.50
TMRS	18.13^***	5199.24^***	5.87^***	0.72
GLAD	24.98^***	4316.17^***	12.16^***	0.57
ARS	0.00^***	1.58^***	0.01^***	0.56
T1	18.79^***	5521.38^***	6.65^***	0.69
T2	18.56^***	4912.08^***	6.16^***	0.71

QTL mapping of stay-green-related traits in wheat under drought condition

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位点 QTL	左标记 Left marker	右标记 Right marker	物理位置 Position (Mb)	候选基因 Gene ID	功能注释 Functional annotation
QTMRS.xjau-4B	AX-110194463	AX-111651924	15.18	TraesCS4B01G021200	基本螺旋-环-螺旋(BHLH) DNA结合超家族蛋白 Basic helix-loop-helix (BHLH) DNA-binding superfamily protein
QTMRS.xjau-4B	AX-110194463	AX-111651924	13.30	TraesCS4B01G018400	衰老相关家族蛋白, 推测(DUF581) Senescence-associated family protein, putative (DUF581)
QT1.xjau-4D	AX-95659047	AX-94558069	16.30	TraesCS4D01G037000	半胱氨酸蛋白酶, 推测 Cysteine protease, putative
QSPAD-10D.xjau-4B	AX-95685381	AX-94912823	171.62	TraesCS4B01G131400	蛋白质锌诱导的促进因子1 Protein zinc induced facilitator-like 1
QSPAD-14D.xjau-4D	AX-111071805	AX-111662392	30.00	TraesCS4D01G054000	GRAS转录因子 GRAS transcription factor
QSPAD-18D.xjau-4D	AX-95659047	AX-108944764	15.52	TraesCS4D01G033900	锌指家族蛋白 Zinc finger family protein
QSPAD-26D.xjau-1B	AX-110534755	AX-111590092	658.91	TraesCS1B01G434300	WRKY家族转录因子 WRKY family transcription factor
QSPAD-26D.xjau-7B	AX-111581943	AX-110629026	6.40	TraesCS7B01G010200	叶绿体POR1的蛋白伴侣样蛋白Protein chaperone-like protein of POR1, chloroplastic