亚麻生氰糖苷合成关键酶CYP79基因家族的鉴定及表达分析

doi:10.3724/SP.J.1006.2023.24042

作物学报 ›› 2023, Vol. 49 ›› Issue (3): 687-702.doi: 10.3724/SP.J.1006.2023.24042

• 作物遗传育种·种质资源·分子遗传学 • 上一篇下一篇

亚麻生氰糖苷合成关键酶CYP79基因家族的鉴定及表达分析

齐燕妮¹(), 李闻娟¹, 赵丽蓉², 李雯², 王利民¹, 谢亚萍¹, 赵玮¹, 党照¹, 张建平¹^,^*()

¹甘肃省农业科学院作物研究所, 甘肃兰州 730070
²甘肃农业大学农学院 / 甘肃省干旱生境作物重点实验室 / 甘肃省作物遗传改良与种质资源创新重点实验室, 甘肃兰州 730070

收稿日期:2022-02-21 接受日期:2022-07-21 出版日期:2023-03-12 网络出版日期:2022-08-19
通讯作者: 张建平
作者简介:E-mail: xbsdqyn@126.com
基金资助:
甘肃省农业科学院现代生物育种项目(2020GAAS08);甘肃省农业科学院现代生物育种项目(2022GAAS04);财政部和农业农村部国家现代农业产业技术体系建设专项(油料);国家自然科学基金项目(31760426);甘肃省知识产权计划项目(21ZSCQ026);甘肃省科技计划项目(21JR7RA722);甘肃省科技计划项目(21JR1RA354)

Identification and expression analysis of CYP79 gene family, a key enzyme for cyanogenic glycoside synthesis in flax

QI Yan-Ni¹(), LI Wen-Juan¹, ZHAO Li-Rong², LI Wen², WANG Li-Min¹, XIE Ya-Ping¹, ZHAO Wei¹, DANG Zhao¹, ZHANG Jian-Ping¹^,^*()

¹Institute of Crop Sciences, Gansu Academy of Agricultural Sciences, Lanzhou 730070, Gansu, China
²College of Agronomy, Gansu Agricultural University / Key Laboratory of Arid Land Crop Science in Gansu Province / Gansu Key Laboratory of Crop Improvement and Germplasm Enhancement, Lanzhou 730070, Gansu, China

Received:2022-02-21 Accepted:2022-07-21 Published:2023-03-12 Published online:2022-08-19
Contact: ZHANG Jian-Ping
Supported by:
Modern Biology Breeding Project of Gansu Academy of Agricultural Sciences(2020GAAS08);Modern Biology Breeding Project of Gansu Academy of Agricultural Sciences(2022GAAS04);China Agriculture Research System (Oil) of MOF and MARA;National Natural Science Foundation of China(31760426);Intellectual Property Planning project of Gansu Province(21ZSCQ026);Science and Technology Project of Gansu Province(21JR7RA722);Science and Technology Project of Gansu Province(21JR1RA354)

摘要/Abstract

摘要：

CYP79蛋白是生氰糖苷合成关键酶, 但关于亚麻CYP79基因的研究鲜有报道。本研究对包括亚麻在内的9种作物的CYP79基因家族进行了鉴定, 并分析了亚麻CYP79基因的序列特征、复制事件、共线性关系、系统进化、顺式作用元件及表达模式。结果表明, 在亚麻、白亚麻、毛果杨、木薯、芝麻、高粱、大豆、葡萄及水稻中分别鉴定到9、9、3、2、5、7、6、16和4个CYP79家族成员; 系统进化分析显示, CYP79基因的进化具有物种特异性; LuCYP79不均匀分布在4条染色体上, 具有1~3个外显子, 其启动子区含大量激素与逆境响应相关元件; 共克隆到8个亚麻CYP79基因的全长DNA序列及5个成员的全长cDNA序列; LuCYP79蛋白序列长度为282~565 aa, 等电点为5.84~9.14, 分子量为31.56~62.86 kD, 均为亲水性蛋白, 定位于内质网; 共有5对基因发生复制事件, 占全部基因的77.8%, 全部经历了强烈的纯化选择, 其中LuCYP79-1和LuCYP79-9在拟南芥和木薯中均具有同源基因。表达分析表明, LuCYP79家族成员具有组织特异性, 且各成员在不同遗传背景下表达模式不同, 其中LuCYP79-1、LuCYP79-7、LuCYP79-8和LuCYP79-9在4个品种中的表达差异显著。相关分析表明, 50 d时的LuCYP79-1/LuCYP79-7与成熟亚麻籽中生氰糖苷含量呈极显著正相关, 20 d时的LuCYP79-7/LuCYP79-8及(LuCYP79-7+LuCYP79-9)/LuCYP79-8分别与成熟亚麻籽中生氰糖苷含量呈极显著正相关, 初步推测其可能是亚麻籽生氰糖苷合成的关键基因。研究结果对进一步阐明亚麻CYP79蛋白的功能具有积极意义, 并为培育低生氰糖苷亚麻品种提供了理论参考。

关键词: 亚麻, CYP79基因家族, 生氰糖苷, 生物信息学, 表达分析

Abstract:

CYP79 is a key enzyme in the synthesis of cyanogenic glycoside. However, there is no systematical study of CYP79 genes in flax. In this study, we identified CYP79 gene family in 9 crops including flax, and focused on the sequence characteristics, duplication events, collinearity, evolution, cis-acting elements and expression patterns of LuCYP79 genes. The results revealed that a total of 9, 9, 3, 2, 5, 7, 6, 16, and 4 CYP79 family members were identified in flax, pale flax, poplar, cassava, sesame, sorghum, soybean, grape, and rice, respectively. Phylogenetic analysis showed that the evolution of CYP79 genes was species-specific. LuCYP79 were distributed unevenly on four chromosomes and had 1-3 exons. The promoter region of LuCYP79 contained lots of elements involved in response to hormone and stress. 8 full-length DNA sequences and 5 full-length cDNA sequences of flax LuCYP79 genes were cloned. LuCYP79 proteins contained 282-565 amino acid residues with molecular weight of 31.56-62.86 kD, and isoelectric point of 5.84-9.14. All LuCYP79 members were hydrophilic protein and located in endoplasmic reticulum. There were five pairs of LuCYP79 genes with duplication events, accounting for 77.8% of all genes, and all the duplication genes underwent strong purification selection. Both LuCYP79-1 and LuCYP79-9 had homologous genes in Arabidopsis and cassava. The relative expression levels showed that LuCYP79 family members were tissue-specific and had different expression patterns under different genetic backgrounds. There were significantly different in the relative expression of LuCYP79-1, LuCYP79-7, LuCYP79-8, and LuCYP79-9 in the four cultivars. Moreover, correlation analysis showed that LuCYP79-1/LuCYP79-7 in 50 days flaxseed was significantly positively correlated with the concentration of cyanogenic glycosides in mature flaxseed. LuCYP79-7/LuCYP79-8 and (LuCYP79-7+LuCYP79-9)/LuCYP79-8 in 20 days flaxseed were significantly positively correlated with the concentration of cyanogenic glycosides in mature flaxseed, respectively. It was preliminarily speculated that they might be the key genes in the synthesis of cyanogenic glycoside in flaxseed. These results have positive significance for further elucidating the function of CYP79 proteins in flax and provide theoretical references for breeding flax varieties with low cyanogenic glycoside.

Key words: flax, CYP79 gene family, cyanogenic glycoside, bioinformatics, the relative expression level

齐燕妮, 李闻娟, 赵丽蓉, 李雯, 王利民, 谢亚萍, 赵玮, 党照, 张建平. 亚麻生氰糖苷合成关键酶CYP79基因家族的鉴定及表达分析[J]. 作物学报, 2023, 49(3): 687-702.

QI Yan-Ni, LI Wen-Juan, ZHAO Li-Rong, LI Wen, WANG Li-Min, XIE Ya-Ping, ZHAO Wei, DANG Zhao, ZHANG Jian-Ping. Identification and expression analysis of CYP79 gene family, a key enzyme for cyanogenic glycoside synthesis in flax[J]. Acta Agronomica Sinica, 2023, 49(3): 687-702.

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表3

LuCYP79启动子顺式作用元件"

顺式元件Cis-element	典型序列 Typical sequence	特性 Characteristic	基因 Gene name
ARE	AAACCA	厌氧诱导必需的顺式作用元件 Cis-acting regulatory element essential for the anaerobic induction	LuCYP79-1, LuCYP79-2, LuCYP79-3, LuCYP79-4, LuCYP79-5, LuCYP79-6, LuCYP79-7, LuCYP79-8, LuCYP79-9
G-box	CACGTG/TACGTG	光响应顺式作用元件 Cis-acting regulatory element involved in light responsiveness	LuCYP79-1, LuCYP79-2, LuCYP79-3, LuCYP79-4, LuCYP79-5, LuCYP79-6, LuCYP79-7, LuCYP79-8, LuCYP79-9
GT1-motif	GGTTAAT/GGTTAA	光响应元件 Light responsive element	LuCYP79-1, LuCYP79-2, LuCYP79-3, LuCYP79-4, LuCYP79-5, LuCYP79-6, LuCYP79-7, LuCYP79-9
Box 4	ATTAAT	光响应保守模块的一部分 Part of a conserved DNA module involved in light responsiveness	LuCYP79-1, LuCYP79-2, LuCYP79-3, LuCYP79-5, LuCYP79-6, LuCYP79-7, LuCYP79-8, LuCYP79-9
GATA-motif	GATAGGG	光响应元件的一部分 Part of a light responsive element	LuCYP79-1, LuCYP79-3, LuCYP79-5, LuCYP79-7, LuCYP79-8, LuCYP79-9
ABRE	CGTACGTGCA/ CGCACGTGTC	脱落酸响应顺式元件 Cis-acting element involved in the abscisic acid responsiveness	LuCYP79-1, LuCYP79-2, LuCYP79-3, LuCYP79-4, LuCYP79-5, LuCYP79-6, LuCYP79-7, LuCYP79-8, LuCYP79-9
TGACG-motif	TGACG	MeJA响应顺式作用元件 Cis-acting regulatory element involved in the MeJA-responsiveness	LuCYP79-1, LuCYP79-2, LuCYP79-3, LuCYP79-4, LuCYP79-5, LuCYP79-6, LuCYP79-7, LuCYP79-8
CGTCA-motif	CGTCA	MeJA响应顺式作用元件 Cis-acting regulatory element involved in the MeJA-responsiveness	LuCYP79-1, LuCYP79-2, LuCYP79-3, LuCYP79-4, LuCYP79-5, LuCYP79-6, LuCYP79-7, LuCYP79-8
TGA-element	AACGAC	生长素响应元件 Auxin-responsive element	LuCYP79-1, LuCYP79-2, LuCYP79-5, LuCYP79-8
AuxRR-core	GGTCCAT	生长素响应顺式作用元件 Cis-acting regulatory element involved in auxin responsiveness	LuCYP79-4, LuCYP79-5
AuxRE	TGTCTCAATAAG	生长素响应元件的一部分 Part of an auxin-responsive element	LuCYP79-7
GARE-motif	TCTGTTG	赤霉素响应元件 Gibberellin-responsive element	LuCYP79-2, LuCYP79-9
P-box	CCTTTTG	赤霉素响应元件 Gibberellin-responsive element	LuCYP79-2
顺式元件Cis-element	典型序列 Typical sequence	特性 Characteristic	基因 Gene name
LTR	CCGAAA	低温响应顺式作用元件 Cis-acting element involved in low-temperature responsiveness	LuCYP79-1, LuCYP79-2, LuCYP79-4, LuCYP79-6, LuCYP79-7
MBS	CAACTG	参与干旱诱导的MYB结合位点 MYB binding site involved in drought-inducibility	LuCYP79-1, LuCYP79-2, LuCYP79-8, LuCYP79-9
TC-rich repeats	GTTTTCTTAC	参与防御和胁迫响应的顺式作用元件 Cis-acting element involved in defense and stress responsiveness	LuCYP79-2
RY-element	CATGCATG	参与种子特异性调控的顺式调控元件 Cis-acting regulatory element involved in seed-specific regulation	LuCYP79-4, LuCYP79-5, LuCYP79-7
CAT-box	GCCACT	分生组织相关的顺式调控元件 Cis-acting regulatory element related to meristem expression	LuCYP79-5, LuCYP79-6, LuCYP79-7
A-box	CCGTCC	顺式调控元件 Cis-acting regulatory element	LuCYP79-3, LuCYP79-7
GC-motif	CCCCCG	参与缺氧特异性诱导的类增强子元件 Enhancer-like element involved in anoxic specific inducibility	LuCYP79-1, LuCYP79-7
O2-site	GATGA(C/T)(A/G)TG (A/G)/GATGATGTGG	参与玉米醇溶蛋白新陈代谢调节的顺式调控元件 Cis-acting regulatory element involved in zein metabolism regulation	LuCYP79-1, LuCYP79-2, LuCYP79-6
MBSI	TTTTTACGGTTA	参与黄酮类合成基因调控的MYB结合位点 MYB binding site involved in flavonoid biosynthetic genes regulation	LuCYP79-3
HD-Zip 1	CAAT(A/T)ATTG	参与栅栏叶肉细胞分化的元件 Element involved in differentiation of the palisade mesophyll cells	LuCYP79-1

表3

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基因名称 Gene name	正向引物序列 Forward sequence (5'-3')	反向引物序列 Reverse sequence (5'-3')
LuCYP79-1	ATGAACATCGACGGCCAGGAGAAG	TCAGTTGGTAATCTTGGGGTACAAGTG
LuCYP79-2	ATGACCATGAAAACCTCCGACG	TCAGTTGGTAACCAAAGGGTACAAG
LuCYP79-3	ATGCCCATGAACACCTCCGACCGAT	TCAGTTGGTAATCTTGGGGTACAAGTGG
LuCYP79-4	ATGAAGTCATCTCCGATCAACACTATTAC	TCATTTGGTAGAGCGGCTGGGTAC
LuCYP79-5	ATGGCAAATAACCTCCCTCCGAG	TCATTTGGCTAAGGCAGGGTACAAG
LuCYP79-6	ATGGAAACCATGATCCTCCTCGTC	TCAAGAGAGTGTAGGGTATAAATGAGG
LuCYP79-7	ATGAACACCTTCACAATCATCCTCC	TCATTTGACGACAACGACAGGGTAC
LuCYP79-8	ATGGCGGCCATGAACACCTCC	TCATTTGGCTAAGGCAGGGTACAAG
LuCYP79-9	ATGGCCATGAACACCTCCGAC	TCAGTTGGTAATCTTAGGGTACAAGTGGG

基因名称 Gene name	正向引物序列 Forward sequence (5'-3')	反向引物序列 Reverse sequence (5'-3')
LuCYP79-1	ACCGCTGCTGTCACTACCACG	CTGCTCCTTCTGATGCGTTGG
LuCYP79-2	GGGAAGGTGATTAAGGAGGCTAATAAG	CCACTGCGTTTGACGGGTTG
LuCYP79-3	GCAAGCGAATAAGACGATGAGGG	GGTTGGTGATTTCATTGCGGGAG
LuCYP79-4	CTCCGACCGATGTCTATCCACCAC	CCAGTCATAATGCTACCTGCTCCT
LuCYP79-5	CGGCACAATGACATCTACCACCTC	CTGCAATAAGCCTACCTGCTCTTC
LuCYP79-6	TAAGGAGTCGAATAAGGCGTTGC	CACTGCGTTGGATGGATTGTCTA
LuCYP79-7	CACCGCCGTCATAATCCTCCTC	GCCGCTGGCTTCTTCCTCGTT
LuCYP79-8	CGGACCGATGGAGATTGACC	TCCCTCGCACATGCCTTGAC
LuCYP79-9	TCTCACCGACGTTCTTATTACCCTT	GGACTAGCCTATCTTTACCGACCAC

基因名称 Gene name	基因号 Gene ID	染色体位置 Chromosome location	外显子数量 No. of exons	蛋白序列长度 Protein length (aa)	分子量 Molecular weight (kD)	等电点 pI	亚细胞定位 Subcellular location
LuCYP79-1	L.us.o.m.scaffold38.16	Chr2:4491546-4492641(+)	2	282	31.56	5.84	ER
LuCYP79-2	L.us.o.m.scaffold38.15	Chr2:4495863-4497662(+)	2	554	62.17	9.14	ER
LuCYP79-3	L.us.o.m.scaffold38.14	Chr2:4499744-4501641(+)	2	558	62.57	9.01	ER
LuCYP79-4	L.us.o.m.scaffold137.29	Chr6:18190484-18192183(+)	2	537	60.32	8.97	ER
LuCYP79-5	L.us.o.m.scaffold137.31	Chr6:18195750-18196883(+)	1	377	41.69	8.11	ER
LuCYP79-6	L.us.o.m.scaffold96.139	Chr12:3278067-3280521(-)	3	521	58.68	6.97	ER
LuCYP79-7	L.us.o.m.scaffold13.272	Chr12:15779026-15781011(-)	2	525	59.10	8.12	ER
LuCYP79-8	L.us.o.m.scaffold13.263	Chr12:15823701-15825571(+)	2	549	60.89	7.26	ER
LuCYP79-9	L.us.o.m.scaffold28.185	Chr15:6106894-6109113(-)	2	565	62.86	9.03	ER

物种 Species	基因号 Gene ID	物种 Species	基因号 Gene ID	物种 Species	基因号 Gene ID
白亚麻 Linum bienne	L.bie.m.scaffold143.40	芝麻 Sesamum indicum	rna5687	葡萄 Vitis vinifera	VIT_206s0009g01780.1
	L.bie.m.scaffold143.49		rna5689		VIT_206s0009g02820.1
	L.bie.m.scaffold349.34		rna5691		VIT_206s0009g02843.1
	L.bie.m.scaffold349.35		rna5692		VIT_206s0009g02850.1
	L.bie.m.scaffold349.36		rna9755		VIT_206s0009g02895.1
	L.bie.m.scaffold593.11	高粱 Sorghum bicolor	Sobic.001G012300.1.p		VIT_206s0009g02930.1
	L.bie.m.scaffold593.12		Sobic.001G185900.2.p		VIT_206s0009g02980.1
	L.bie.m.scaffold593.6		Sobic.001G185900.2.p		VIT_206s0009g03120.1
	L.bie.m.scaffold852.13		Sobic.001G187500.1.p		VIT_206s0009g03130.1
毛果杨 Populus trichocarpa	Potri.004G055200.1.p		Sobic.001G187600.1.p		VIT_213s0067g02720.1
	Potri.013G157200.1.p		Sobic.007G090457.1.p		VIT_213s0067g02730.1
	Potri.013G157400.1.p		Sobic.010G172200.1.p		VIT_213s0067g02780.1
木薯 Manihot esculenta	Manes.12G133500.1.p	大豆 Glycine max	Glyma.11G197300.1.p		VIT_213s0067g02790.1
木薯 Manihot esculenta	Manes.13G094200.1.p		Glyma.11G197400.1.p		VIT_213s0067g02830.1
水稻 Oryza sativa	LOC_Os03g37290.1		Glyma.13G051600.1.p		VIT_213s0106g00280.1
	LOC_Os04g08824.1		Glyma.18G052200.1.p		VIT_218s0001g13760.1
	LOC_Os04g08828.1		Glyma.20G065000.1.p
	LOC_Os04g09430.1		Glyma.20G065100.1.p

蛋白名称Protein name	α-螺旋 Alpha helix	β-转角 Beta turn	延伸链 Extended strand	无规则卷曲 Random coil
LuCYP79-9	120 (42.55%)	7 (2.48%)	35 (12.41%)	120 (42.55%)
LuCYP79-5	255 (46.03%)	22 (3.97%)	58 (10.47%)	219 (39.53%)
LuCYP79-4	240 (43.01%)	32 (5.73%)	86 (15.41%)	200 (35.84%)
LuCYP79-7	231 (43.02%)	23 (4.28%)	71 (13.22%)	212 (39.48%)
LuCYP79-8	177 (46.95%)	9 (2.39%)	48 (12.73%)	143 (37.93%)
LuCYP79-6	240 (46.07%)	21 (4.03%)	64 (12.28%)	196 (37.62%)
LuCYP79-3	234 (44.57%)	19 (3.62%)	68 (12.95%)	204 (38.86%)
LuCYP79-2	243 (44.26%)	21 (3.83%)	76 (13.84%)	209 (38.07%)
LuCYP79-1	247 (43.72%)	25 ( 4.42%)	76 (13.45%)	217 (38.41%)

亚麻生氰糖苷合成关键酶CYP79基因家族的鉴定及表达分析

Identification and expression analysis of CYP79 gene family, a key enzyme for cyanogenic glycoside synthesis in flax

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基因对 Gene pairs	复制事件 Duplication event	非同义替换率 Ka	同义替换率 Ks	Ka/Ks	选择类型 Selection type	分歧时间 Divergence time (MYa)
LuCYP79-1/LuCYP79-2	串联复制 Tandem duplication	0.0722	0.6753	0.1069	纯化选择 Purifying selection	55.3525
LuCYP79-2/LuCYP79-3	串联复制 Tandem duplication	0.0678	0.6603	0.1027	纯化选择 Purifying selection	54.1230
LuCYP79-4/LuCYP79-5	串联复制 Tandem duplication	0.1213	1.1045	0.1098	纯化选择 Purifying selection	90.5328
LuCYP79-1/LuCYP79-9	片段复制 Segmental duplication	0.0055	0.0164	0.3354	纯化选择 Purifying selection	1.3443
LuCYP79-4/LuCYP79-8	片段复制 Segmental duplication	0.1277	1.0731	0.1190	纯化选择 Purifying selection	87.9590

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