Identification of GRAS gene family members and their regulatory roles in reproductive development in Chenopodium quinoa

doi:10.3969/j.issn.1004-1524.20250081

Acta Agriculturae Zhejiangensis ›› 2026, Vol. 38 ›› Issue (1): 35-53.DOI: 10.3969/j.issn.1004-1524.20250081

• Crop Science • Previous Articles Next Articles

Identification of GRAS gene family members and their regulatory roles in reproductive development in Chenopodium quinoa

YANG Yang^a(), ZHANG Shuai^a, DONG Chenwenhua^a^,^b, ZENG Mengqiong^a, LIN Chun^a^,^b, MAO Zichao^a^,^b^,^*()

College of Agriculture and Biotechnology; b. Research Center of Featured Minor Crops, Yunnan Agricultural University, Kunming 650201, China

Received:2025-02-07 Online:2026-01-25 Published:2026-02-11

Abstract

Abstract:

The GRAS gene family plays an essential regulatory role in plant growth, development, and stress responses, but no relevant reports have been published on its function in the reproductive development of quinoa (Chenopodium quinoa). Based on the latest quinoa genome data, this study systematically identified and analyzed the quinoa GRAS family (CqGRAS) members, focusing on their gene structures, promoter cis-acting elements, as well as their expression patterns and regulatory mechanisms during vegetative and reproductive growth stages. Additionally, a comparative analysis was conducted between the CqGRAS members and the GRAS genes of Arabidopsis thaliana and the diploid ancestral species of quinoa, Chenopodium watsonii(A genome) and Chenopodium suecicum(B genome). A total of 51 CqGRAS genes were identified. These genes generally contained few introns and exhibited high homology with the GRAS genes of Arabidopsis and the diploid Chenopodium species. Promoter analysis revealed that these genes were enriched with cis-acting elements responsive to plant hormones (such as gibberellin, abscisic acid, ethylene, and jasmonic acid), growth and development, as well as stress signals. Phylogenetic analysis classified the CqGRAS family into 10 subfamilies, among which HAM (CqHAM01), PAT1 (CqPAT1-06/07/08), DELLA (CqDELLA01/02), DLT (CqDLT01/02), and SHR (CqSHR05/06) showed relatively high expression levels in inflorescences and developing seeds. Weighted gene co-expression network analysis (WGCNA) suggested that these CqGRAS genes associated with reproductive development might regulate the growth and development of quinoa flowers and seeds by integrating light signaling and hormone signaling pathways. This study provided novel insights into the role of the GRAS family in quinoa reproductive development and laid the foundation for further elucidation of its molecular mechanisms and breeding applications.

Key words: Chenopodium quinoa, GRAS gene family, reproductive regulation, weighted gene co-expression network analysis (WGCNA), reproductive development

CLC Number:

S519

YANG Yang, ZHANG Shuai, DONG Chenwenhua, ZENG Mengqiong, LIN Chun, MAO Zichao. Identification of GRAS gene family members and their regulatory roles in reproductive development in Chenopodium quinoa[J]. Acta Agriculturae Zhejiangensis, 2026, 38(1): 35-53.

Figures/Tables 10

References 75

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引物名称primer name	引物序列(5'→3') Primer sequence(5'→3')	产物大小/bp Product length/bp
1-CqDELLA01-F2	CGAAAGACACGAGCCACTAATAAA	169
1-CqDELLA01-R2	CCACCCTAGAGTCAAGCACCC
2-CqDELLA02-F2	ACTTCTAGTTCTCAATCTTTTGTTTTC	194
2-CqDELLA02-R2	AAAGTGTCTAGCACCCCCAG
3-CqPAT1-05-F2	CTTTCCCTTAGCAATGACAACA	105
3-CqPAT1-05-R2	AATGTTTAGGAAGGATGGTCGT
4-CqPAT1-06-F	AGGACTACTTCGGTGTTACTCTG	103
4-CqPAT1-06-R	CCAAGCTGAAGCCGAGATCAA
5-CqDLT01-F2	GGTATCAGGAGGGACGAAAG	166
5-CqDLT01-R2	CAGCAGCATCAACAGAACCC
6-CqDLT02-F2	CATGGATTAGTCTTTGATTTTCGAT	175
6-CqDLT02-R2	CATGCCATAAGTAGATCAAATAACAA
7-CqSHR05-F	GGCGTATAGTGAGGAGGTTTGTG	109
7-CqSHR05-R	TATTCCGGCAGTCGGAGAC
8-CqSHR06-F	CGGAGAGGAGAGAGAATGCC	137
8-CqSHR06-R	TGCCATCGACCACCCTTCT
CqActin-F2	TTGGAATCTCGCCGCCAAA	132
CqActin-R2	CCAGGTCCACCAACAATCCA

引物名称primer name	引物序列(5'→3') Primer sequence(5'→3')	产物大小/bp Product length/bp
1-CqDELLA01-F2	CGAAAGACACGAGCCACTAATAAA	169
1-CqDELLA01-R2	CCACCCTAGAGTCAAGCACCC
2-CqDELLA02-F2	ACTTCTAGTTCTCAATCTTTTGTTTTC	194
2-CqDELLA02-R2	AAAGTGTCTAGCACCCCCAG
3-CqPAT1-05-F2	CTTTCCCTTAGCAATGACAACA	105
3-CqPAT1-05-R2	AATGTTTAGGAAGGATGGTCGT
4-CqPAT1-06-F	AGGACTACTTCGGTGTTACTCTG	103
4-CqPAT1-06-R	CCAAGCTGAAGCCGAGATCAA
5-CqDLT01-F2	GGTATCAGGAGGGACGAAAG	166
5-CqDLT01-R2	CAGCAGCATCAACAGAACCC
6-CqDLT02-F2	CATGGATTAGTCTTTGATTTTCGAT	175
6-CqDLT02-R2	CATGCCATAAGTAGATCAAATAACAA
7-CqSHR05-F	GGCGTATAGTGAGGAGGTTTGTG	109
7-CqSHR05-R	TATTCCGGCAGTCGGAGAC
8-CqSHR06-F	CGGAGAGGAGAGAGAATGCC	137
8-CqSHR06-R	TGCCATCGACCACCCTTCT
CqActin-F2	TTGGAATCTCGCCGCCAAA	132
CqActin-R2	CCAGGTCCACCAACAATCCA

基因名称 Gene name	基因ID Gene ID	染色体 Chromosome	氨基酸长度/aa Amino acid length/aa	蛋白质分子质量/u Protein molecular weight/u	等电点 Isoelectric point	亚细胞定位 Subcellular localization
CqDELLA01	CQ012679	Cq8B	459	50 811.86	5.51	溶酶体Cytolysosome
CqDELLA02	CQ040570	Cq8A	474	52 509.87	5.66	溶酶体Cytolysosome
CqDELLA03	CQ001679	Cq6B	617	67 367.76	4.90	细胞核Nuclear
CqDELLA04	CQ026984	Cq6A	618	67 500.87	4.90	细胞核Nuclear
CqDLT01	CQ004440	Cq5B	648	73 031.86	5.77	叶绿体Chloroplast
CqDLT02	CQ029757	Cq5A	657	73 968.04	5.47	叶绿体Chloroplast
CqHAM01	CQ002321	Cq6B	610	67 095.88	6.46	细胞核Nuclear
CqHAM02	CQ027701	Cq6A	602	66 232.26	6.82	细胞核Nuclear
CqHAM03	CQ003474	Cq5B	785	85 801.61	6.15	细胞核Nuclear
CqHAM04	CQ028875	Cq5A	781	85 891.76	6.12	细胞核Nuclear
CqLAS01	CQ033480	Cq9B	490	55 803.58	7.02	细胞核Nuclear
CqLAS02	CQ051297	Cq9A	484	54 848.37	6.76	细胞核Nuclear
CqLISCL01	CQ032605	Cq9B	350	41 049.14	8.10	溶酶体Cytolysosome
CqLISCL02	CQ041202	Cq7A	725	82 353.95	4.95	细胞核Nuclear
CqLISCL03	CQ032598	Cq9B	689	78 590.85	5.18	细胞核Nuclear
CqLISCL04	CQ041204	Cq7A	619	70 272.91	6.29	细胞核Nuclear
CqLISCL05	CQ032601	Cq9B	679	76 337.79	5.60	细胞核Nuclear
CqLISCL06	CQ032604	Cq9B	548	62 438.96	7.94	溶酶体Cytolysosome
CqLISCL07	CQ041201	Cq7A	662	74 879.34	6.61	细胞核Nuclear
CqLISCL08	CQ032603	Cq9B	667	75 299.73	5.23	细胞核Nuclear
CqLISCL09	CQ041200	Cq7A	686	77 311.71	4.87	细胞核Nuclear
CqLISCL10	CQ030634	Cq5A	760	86 162.35	5.50	细胞核Nuclear
CqLISCL11	CQ055311	Cq2A	780	87 684.22	7.07	溶酶体Cytolysosome
CqLISCL12	CQ025002	Cq1B	818	91 640.62	7.49	细胞核Nuclear
CqLISCL13	CQ032609	Cq9B	756	84 422.57	5.55	细胞核Nuclear
CqLISCL14	CQ041197	Cq7A	756	84 533.61	5.81	细胞核Nuclear
CqPAT1-01	CQ024284	Cq1B	545	61 454.56	4.73	叶绿体Chloroplast
CqPAT1-02	CQ048458	Cq1A	426	48 153.43	5.27	溶酶体Cytolysosome
CqPAT1-03	CQ016853	Cq3B	639	70 122.51	5.46	细胞核Nuclear
CqPAT1-04	CQ044170	Cq3A	767	84 245.91	6.39	细胞核Nuclear
CqPAT1-05	CQ022180	Cq4B	635	70 854.40	8.16	叶绿体Chloroplast
CqPAT1-06	CQ037875	Cq4A	634	70 854.39	7.57	叶绿体Chloroplast
CqPAT1-07	CQ010139	Cq8B	544	61 079.89	6.33	细胞核Nuclear
CqPAT1-08	CQ038026	Cq8A	544	61 110.88	6.51	细胞核Nuclear
CqPAT1-09	CQ013243	Cq2B	466	52 280.60	5.82	叶绿体Chloroplast
CqPAT1-10	CQ034654	Cq9B	541	60 293.76	6.56	细胞核Nuclear
CqPAT1-11	CQ050106	Cq9A	588	65 884.16	6.69	叶绿体Chloroplast
CqSCL3L01	CQ019876	Cq4B	454	50 710.01	6.51	细胞核Nuclear
CqSCL3L02	CQ035148	Cq4A	454	50 837.30	6.65	细胞核Nuclear
CqSCL4L01	CQ003361	Cq6B	557	60 789.24	4.38	细胞核Nuclear
CqSCL4L02	CQ028754	Cq6A	610	67 157.51	4.74	细胞核Nuclear
CqSCR01	CQ052618	Cq9A	872	95 159.47	6.39	细胞核Nuclear
CqSCR02	CQ007129	Cq7B	861	93 906.00	6.40	细胞核Nuclear
CqSCR03	CQ014997	Cq2B	396	43 813.31	6.99	叶绿体Chloroplast
CqSCR04	CQ054542	Cq2A	396	43 859.22	7.32	细胞核Nuclear
CqSHR01	CQ003291	Cq6B	475	53 457.35	5.88	线粒体、细胞骨架 Mitochondrion, cytoskeleton
CqSHR02	CQ028690	Cq6A	389	43 994.11	5.05	细胞骨架Cytoskeleton
CqSHR03	CQ018831	Cq3B	439	50 575.84	6.38	叶绿体Chloroplast
CqSHR04	CQ044974	Cq3A	440	50 599.98	6.73	叶绿体Chloroplast
CqSHR05	CQ003315	Cq6B	472	53 064.48	5.94	细胞核Nuclear
CqSHR06	CQ028714	Cq6A	467	52 415.84	5.86	细胞核Nuclear

基因名称 Gene name	基因ID Gene ID	染色体 Chromosome	氨基酸长度/aa Amino acid length/aa	蛋白质分子质量/u Protein molecular weight/u	等电点 Isoelectric point	亚细胞定位 Subcellular localization
CqDELLA01	CQ012679	Cq8B	459	50 811.86	5.51	溶酶体Cytolysosome
CqDELLA02	CQ040570	Cq8A	474	52 509.87	5.66	溶酶体Cytolysosome
CqDELLA03	CQ001679	Cq6B	617	67 367.76	4.90	细胞核Nuclear
CqDELLA04	CQ026984	Cq6A	618	67 500.87	4.90	细胞核Nuclear
CqDLT01	CQ004440	Cq5B	648	73 031.86	5.77	叶绿体Chloroplast
CqDLT02	CQ029757	Cq5A	657	73 968.04	5.47	叶绿体Chloroplast
CqHAM01	CQ002321	Cq6B	610	67 095.88	6.46	细胞核Nuclear
CqHAM02	CQ027701	Cq6A	602	66 232.26	6.82	细胞核Nuclear
CqHAM03	CQ003474	Cq5B	785	85 801.61	6.15	细胞核Nuclear
CqHAM04	CQ028875	Cq5A	781	85 891.76	6.12	细胞核Nuclear
CqLAS01	CQ033480	Cq9B	490	55 803.58	7.02	细胞核Nuclear
CqLAS02	CQ051297	Cq9A	484	54 848.37	6.76	细胞核Nuclear
CqLISCL01	CQ032605	Cq9B	350	41 049.14	8.10	溶酶体Cytolysosome
CqLISCL02	CQ041202	Cq7A	725	82 353.95	4.95	细胞核Nuclear
CqLISCL03	CQ032598	Cq9B	689	78 590.85	5.18	细胞核Nuclear
CqLISCL04	CQ041204	Cq7A	619	70 272.91	6.29	细胞核Nuclear
CqLISCL05	CQ032601	Cq9B	679	76 337.79	5.60	细胞核Nuclear
CqLISCL06	CQ032604	Cq9B	548	62 438.96	7.94	溶酶体Cytolysosome
CqLISCL07	CQ041201	Cq7A	662	74 879.34	6.61	细胞核Nuclear
CqLISCL08	CQ032603	Cq9B	667	75 299.73	5.23	细胞核Nuclear
CqLISCL09	CQ041200	Cq7A	686	77 311.71	4.87	细胞核Nuclear
CqLISCL10	CQ030634	Cq5A	760	86 162.35	5.50	细胞核Nuclear
CqLISCL11	CQ055311	Cq2A	780	87 684.22	7.07	溶酶体Cytolysosome
CqLISCL12	CQ025002	Cq1B	818	91 640.62	7.49	细胞核Nuclear
CqLISCL13	CQ032609	Cq9B	756	84 422.57	5.55	细胞核Nuclear
CqLISCL14	CQ041197	Cq7A	756	84 533.61	5.81	细胞核Nuclear
CqPAT1-01	CQ024284	Cq1B	545	61 454.56	4.73	叶绿体Chloroplast
CqPAT1-02	CQ048458	Cq1A	426	48 153.43	5.27	溶酶体Cytolysosome
CqPAT1-03	CQ016853	Cq3B	639	70 122.51	5.46	细胞核Nuclear
CqPAT1-04	CQ044170	Cq3A	767	84 245.91	6.39	细胞核Nuclear
CqPAT1-05	CQ022180	Cq4B	635	70 854.40	8.16	叶绿体Chloroplast
CqPAT1-06	CQ037875	Cq4A	634	70 854.39	7.57	叶绿体Chloroplast
CqPAT1-07	CQ010139	Cq8B	544	61 079.89	6.33	细胞核Nuclear
CqPAT1-08	CQ038026	Cq8A	544	61 110.88	6.51	细胞核Nuclear
CqPAT1-09	CQ013243	Cq2B	466	52 280.60	5.82	叶绿体Chloroplast
CqPAT1-10	CQ034654	Cq9B	541	60 293.76	6.56	细胞核Nuclear
CqPAT1-11	CQ050106	Cq9A	588	65 884.16	6.69	叶绿体Chloroplast
CqSCL3L01	CQ019876	Cq4B	454	50 710.01	6.51	细胞核Nuclear
CqSCL3L02	CQ035148	Cq4A	454	50 837.30	6.65	细胞核Nuclear
CqSCL4L01	CQ003361	Cq6B	557	60 789.24	4.38	细胞核Nuclear
CqSCL4L02	CQ028754	Cq6A	610	67 157.51	4.74	细胞核Nuclear
CqSCR01	CQ052618	Cq9A	872	95 159.47	6.39	细胞核Nuclear
CqSCR02	CQ007129	Cq7B	861	93 906.00	6.40	细胞核Nuclear
CqSCR03	CQ014997	Cq2B	396	43 813.31	6.99	叶绿体Chloroplast
CqSCR04	CQ054542	Cq2A	396	43 859.22	7.32	细胞核Nuclear
CqSHR01	CQ003291	Cq6B	475	53 457.35	5.88	线粒体、细胞骨架 Mitochondrion, cytoskeleton
CqSHR02	CQ028690	Cq6A	389	43 994.11	5.05	细胞骨架Cytoskeleton
CqSHR03	CQ018831	Cq3B	439	50 575.84	6.38	叶绿体Chloroplast
CqSHR04	CQ044974	Cq3A	440	50 599.98	6.73	叶绿体Chloroplast
CqSHR05	CQ003315	Cq6B	472	53 064.48	5.94	细胞核Nuclear
CqSHR06	CQ028714	Cq6A	467	52 415.84	5.86	细胞核Nuclear

Identification of GRAS gene family members and their regulatory roles in reproductive development in Chenopodium quinoa

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