藜麦CqSGT1基因克隆、表达模式与DNA变异分析

doi:10.11686/cyxb2025063

摘要/Abstract

摘要：

藜麦的生长发育易受逆境胁迫，影响藜麦产业的发展。SGT1作为Skp1-4的抑制因子，通过分子伴侣或调控泛素化对植物抗逆反应进行调控，其中Skp1-4参与调控细胞周期、信号转导和基因表达等生物过程。鉴定藜麦SGT1基因，明确藜麦SGT1基因对生物及非生物胁迫的应答情况。利用生物信息学方法，鉴定出藜麦SGT1基因，并对其理化性质、序列特征、系统发育树、蛋白互作网络及表达模式等方面进行研究。从藜麦基因组鉴定出2个SGT1基因，分别命名为CqSGT1a和CqSGT1b，位于Chr06和Chr07上，藜麦SGT1蛋白主要富含碱性氨基酸，亚细胞定位在细胞核，无信号肽结构，二级结构以α-螺旋为主，属于疏水性蛋白，无跨膜结构，具有TPR-SGS-CS结构域。CqSGT1启动子区域存在藜麦生长发育和抵御逆境胁迫的光系统和激素等响应元件。CqSGT1基因与BvSGT1基因亲缘关系最近。qPCR技术分析表明，藜麦SGT1在花和籽粒中表达量较高，推测其表达与花和籽粒形成发育有关；低温初期表达水平上调，随之被抑制；水杨酸（SA）正向调控SGT1的表达，3 h时响应最显著。SGT1于2403（抗病材料）中响应霜霉病菌侵染，抗病材料中SGT1的表达量于接种后2 h显著上调，后期先降后升，在24 h时的响应最为强烈，表明SGT1a/b基因在藜麦霜霉病中发挥正调控作用。CqSGT1基因具有组织表达特异性，并且均响应低温、SA胁迫及霜霉病菌侵染，在藜麦器官生长发育和抗逆过程中发挥重要作用。

关键词: 藜麦, SGT1, 霜霉病菌, 生物信息学分析, 基因克隆

Abstract:

The growth and development of quinoa （Chenopodium quinoa） are negatively affected by adverse conditions， and that restricts the development of the quinoa industry. SGT1 （suppressor of the G2 allele of Skp1） participates in the plant stress resistance response by regulating molecular chaperones and ubiquitination. The SGT1 protein inhibits Skp1-4， which are components of ubiquitin ligase complexes that regulating biological processes such as the cell cycle， signal transduction， and gene expression. The aim of this study was to identify quinoa SGT1 genes and determine their transcriptional responses to biotic and abiotic stress conditions. The SGT1 genes of quinoa were identified using bioinformatics-based methods， and their sequence characteristics， phylogenetic relationships， and expression patterns were analyzed. In addition， the physical and chemical properties and protein interaction networks of their putative encoded proteins were determined. The quinoa genome was found to contain two SGT1 genes， CqSGT1a and CqSGT1b， located on chromosomes 6 and 7. The putative quinoa SGT1 proteins are rich in basic amino acids， lack signal peptides， and are dominated by α-helixes. They were predicted to localize to the nucleus. Both were predicted to be hydrophobic proteins without transmembrane structures， and both contained the characteristic TPR-SGS-CS domain. The CqSGT1 promoter regions contained cis-acting elements related to light and hormone responsiveness， suggesting that the expression of these genes is tightly regulated during growth， development， and stress responses. The CqSGT1 genes showed the closest relationship with BvSGT1 from Beta vulgaris. The results of qPCR analyses showed that the highest transcript levels of SGT1 in quinoa were in the flowers and grains， suggesting that their expression was related to the formation and development of these organs. Under low temperature stress， both SGT1 genes were initially up-regulated and then down-regulated. Treatment with salicylic acid induced the expression of SGT1， and the response was most significant at 3 h after inoculation. SGT1 responses were seen during downy mildew （Peronospora variabilis） infection in the resistant quinoa line 2403. In the resistant line， SGT1 transcript levels were significantly increased at 2 h after inoculation， then decreased， and then subsequently increased again. The strongest response was at 24 h， indicating that the SGT1a/b genes play a positive regulatory role in the response to quinoa downy mildew. Both CqSGT1 genes showed tissue-specific expression patterns and responded to low temperature， salicylic acid， and downy mildew infection. These results show that SGT1 plays an important role in the growth and development of quinoa， and in its responses to biotic and abiotic stress.

Key words: Chenopodium quinoa, SGT1, Peronospora variabilis, bioinformatics analysis, gene cloning

窦苗苗, 姜晓东, 孙慧琼, 徐宏申, 王锡亮, 杨博慧, 柴文婷, 赵珊珊, 张春来. 藜麦CqSGT1基因克隆、表达模式与DNA变异分析[J]. 草业学报, 2026, 35(1): 223-240.

Miao-miao DOU, Xiao-dong JIANG, Hui-qiong SUN, Hong-shen XU, Xi-liang WANG, Bo-hui YANG, Wen-ting CHAI, Shan-shan ZHAO, Chun-lai ZHANG. Cloning， expression profiling and DNA variation analysis of the disease-resistance gene CqSGT1 in quinoa （Chenopodium quinoa）[J]. Acta Prataculturae Sinica, 2026, 35(1): 223-240.

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基因名称 Gene name	正向序列 Forward primer （5′→3′）	反向序列 Reverse primer （5′→3′）	用途 Application
CqSGT1a	ACGGAAAGGAACTGCGTGTA	GGCTGCTTGTGCATTGGTAG	qRT-PCR， SemiRT-PCR
CqSGT1b	TTACCTACGAAAAGGGACCGC	TGCTTGTGCATTGGTAGGCT	qRT-PCR， SemiRT-PCR
CqEF1α	GACAAGCGTGTGATCGAGAG	TCGGCCTTAAGTTTGTCGAG	qRT-PCR， SemiRT-PCR
TI-CqSGT1a/b	ATGGCGACGGATCTCGAGACCAAG	TTAGTATTCCCATTTCTTCATCTC	基因克隆Gene clone
pCAMBIA1300-Cq SGT1a/b	CGCGGATCCAATGGCGACGGATCTCGAGACCAAG	CGCGTCGACTTAGTATTCCCATTT CTTCATCTC	瞬时表达Transient expression
M13	GTTGTAAAACGACGGCCAG	CAGGAAACAGCTATGAC	基因克隆和瞬时表达Gene clone and transient expression

基因名称 Gene name	正向序列 Forward primer （5′→3′）	反向序列 Reverse primer （5′→3′）	用途 Application
CqSGT1a	ACGGAAAGGAACTGCGTGTA	GGCTGCTTGTGCATTGGTAG	qRT-PCR， SemiRT-PCR
CqSGT1b	TTACCTACGAAAAGGGACCGC	TGCTTGTGCATTGGTAGGCT	qRT-PCR， SemiRT-PCR
CqEF1α	GACAAGCGTGTGATCGAGAG	TCGGCCTTAAGTTTGTCGAG	qRT-PCR， SemiRT-PCR
TI-CqSGT1a/b	ATGGCGACGGATCTCGAGACCAAG	TTAGTATTCCCATTTCTTCATCTC	基因克隆Gene clone
pCAMBIA1300-Cq SGT1a/b	CGCGGATCCAATGGCGACGGATCTCGAGACCAAG	CGCGTCGACTTAGTATTCCCATTT CTTCATCTC	瞬时表达Transient expression
M13	GTTGTAAAACGACGGCCAG	CAGGAAACAGCTATGAC	基因克隆和瞬时表达Gene clone and transient expression

元件名称 Element name	核心序列 Core sequence	元件功能 Element function	数量Quantity
元件名称 Element name	核心序列 Core sequence	元件功能 Element function	CqSGT1a	CqSGT1b
chs-CMA1a	TTACTTAA	部分光响应元件Part of light responsive element	1	-
3-AF1 binding site	TAAGAGAGGAA	光响应元件Light responsive element	1	-
HD-Zip 3	GTAAT（G/C）ATTAC	蛋白质结合位点Protein binding site	1	-
ARE	AAACCA	厌氧诱导必不可少的顺式作用调节元件Cis-acting regulatory element essential for the anaerobic induction	4	1
GT1-motif	GGTTAA	光响应元件Light responsive element	2	-
Box 4	ATTAAT	参与光响应的部分保守DNA模块Part of conserved DNA module involved in light responsiveness	3	4
G-box	CACGAC	参与光反应的顺式作用调节元件Cis-acting regulatory element involved in light responsiveness	1	-
GA-motif	ATAGATAA	部分光响应元件Part of light responsive element	1	-
TCT-motif	TCTTAC	部分光响应元件Part of light responsive element	1	-
P-box	CCTTTTG	赤霉素响应元件Gibberellin-responsive element	2	2
Gap-box	CAAATGAA（A/G）A	光响应元件Light responsive element	1	-
TCA-element	CCATCTTTTT	参与水杨酸反应的顺式作用调节件Cis-acting element involved in salicylic acid responsiveness	1	-
ATCT-motif	AATCTAATCC	参与光响应的部分保守DNA模块Part of conserved DNA module involved in light responsiveness	1	-
TATA-box	AATCTAATCC	转录起始-30核心启动子元件Core promoter element around-30 of transcription start	64	53
MBSI	AAAAAAC（G/C）GTTA	MYB结合位点参与类黄酮生物合成基因的调控元件MYB-binding site as a regulatory element involved in the control of flavonoid biosynthetic genes	-	1
CAAT-box	CAAAT	启动子和增强子区域调控元件Promoter and enhancer region regulatory elements	-	12
TCT-motif	TCTTAC	部分光响应元件Part of light responsive element	-	2
O₂-site	GTTGACGTGA	参与玉米醇溶蛋白代谢调控的顺式调节元件Cis-acting regulatory element involved in zein metabolism regulation	-	1
MRE	AACCTAA	MYB结合位点（参与光响应）MYB binding site involved in light responsiveness	-	1
Circadian	AACCTAA	参与昼夜节律控制的顺式作用调节元件Cis-acting regulatory element involved in circadian control	-	1

元件名称 Element name	核心序列 Core sequence	元件功能 Element function	数量Quantity
元件名称 Element name	核心序列 Core sequence	元件功能 Element function	CqSGT1a	CqSGT1b
chs-CMA1a	TTACTTAA	部分光响应元件Part of light responsive element	1	-
3-AF1 binding site	TAAGAGAGGAA	光响应元件Light responsive element	1	-
HD-Zip 3	GTAAT（G/C）ATTAC	蛋白质结合位点Protein binding site	1	-
ARE	AAACCA	厌氧诱导必不可少的顺式作用调节元件Cis-acting regulatory element essential for the anaerobic induction	4	1
GT1-motif	GGTTAA	光响应元件Light responsive element	2	-
Box 4	ATTAAT	参与光响应的部分保守DNA模块Part of conserved DNA module involved in light responsiveness	3	4
G-box	CACGAC	参与光反应的顺式作用调节元件Cis-acting regulatory element involved in light responsiveness	1	-
GA-motif	ATAGATAA	部分光响应元件Part of light responsive element	1	-
TCT-motif	TCTTAC	部分光响应元件Part of light responsive element	1	-
P-box	CCTTTTG	赤霉素响应元件Gibberellin-responsive element	2	2
Gap-box	CAAATGAA（A/G）A	光响应元件Light responsive element	1	-
TCA-element	CCATCTTTTT	参与水杨酸反应的顺式作用调节件Cis-acting element involved in salicylic acid responsiveness	1	-
ATCT-motif	AATCTAATCC	参与光响应的部分保守DNA模块Part of conserved DNA module involved in light responsiveness	1	-
TATA-box	AATCTAATCC	转录起始-30核心启动子元件Core promoter element around-30 of transcription start	64	53
MBSI	AAAAAAC（G/C）GTTA	MYB结合位点参与类黄酮生物合成基因的调控元件MYB-binding site as a regulatory element involved in the control of flavonoid biosynthetic genes	-	1
CAAT-box	CAAAT	启动子和增强子区域调控元件Promoter and enhancer region regulatory elements	-	12
TCT-motif	TCTTAC	部分光响应元件Part of light responsive element	-	2
O₂-site	GTTGACGTGA	参与玉米醇溶蛋白代谢调控的顺式调节元件Cis-acting regulatory element involved in zein metabolism regulation	-	1
MRE	AACCTAA	MYB结合位点（参与光响应）MYB binding site involved in light responsiveness	-	1
Circadian	AACCTAA	参与昼夜节律控制的顺式作用调节元件Cis-acting regulatory element involved in circadian control	-	1

基因 Gene	染色体位置Chromosome location	Ref	Alt	JQ3	JQ6	Qingheili-8	Qingbaili-9	影响Effect（其他突变系Other mutant lines）	改变的密码子 The changed codon
CqSGT1a	Chr06：34577463	GA	G	GA， GA	GA， GA	GA， GA	G，G	内含子Intron	GTa/Gat
CqSGT1b	Chr07：83675475	A	AT	N	A， AT	A， AT	N	上游Upstream	aAt/aAT
CqSGT1b	Chr07：83677523	A	AC	AC， AC	AC， AC	AC， AC	AC， AC	内含子Intron	aAc/aAC
CqSGT1b	Chr07：83679761	CA	C	C， C	C， C	C， C	CA， C	内含子Intron	cCA/cCa
CqSGT1b	Chr07：83679830	TTTGTTGTTGTTG	T	TTTGTTGTTGTTG， TTTGTTGTTGTTG	N	N	TTTGTTGTTGTTG， T	内含子Intron	cTTTGTTGTTGTTGT/cTt
CqSGT1b	Chr07：83689890	A	ATAGCTGGCACAAACGGTGCCTTGT	A， A	A， ATAGCTGGCACAAACGGTGCCTTGT	A， ATAGCTGGCACAAACGGTGCCTTGT	A， ATAGCTGGCA CAAACGGTGCCTTGT	下游Downstream	aAt/aATAGCTGGCACAAACGGTGCCTTGTt
CqSGT1b	Chr07：83690874	A	AT	AT， AT	A， A	A， A	N	下游Downstream	tAt/tAT
CqSGT1b	Chr07：83691319	G	GA	GA， GA	GA， GA	GA， GA	GA， GA	下游Downstream	acGaaa/acGAaa