花生FAD2基因家族表达分析及其对低温胁迫的响应
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摘要
为了探究FAD2在花生低温响应中的作用,本研究从普通油酸花生中花16 (ZH16)和高油酸花生中花413 (ZH413)中克隆得到花生AhFAD2家族的全部基因,共7个。通过分析这些基因的表达模式发现,在ZH16和ZH413中各FAD2基因表达模式相似, AhFAD2-1A/B主要在花和发育的种子中表达, AhFAD2-3A/B主要在营养组织中表达, AhFAD2-4A/B主要在根和花中表达,表明AhFAD2基因在花生不同发育阶段和不同组织中发挥各自的生物学功能。在15℃下发芽6 d发现,ZH413的发芽率未显著下降,而ZH16的发芽率显著下降。种子萌发过程中,AhFAD2-1A/B和AhFAD2-4A/B均受低温诱导表达。在ZH16中AhFAD2-1A/B在低温诱导第6天开始显著上调表达,而在ZH413中第1天显著上调表达;在ZH16中AhFAD2-4A/B在低温诱导第3天出现显著上调表达,之后表达量下降,但在ZH413中第1天就显著上调表达,且始终维持在高水平表达。基于以上研究结果推测,高油酸花生在受到低温胁迫时,AhFAD2-1A/B编码蛋白失活,但AhFAD2-4A/B的高量表达在一定程度上弥补了这部分功能。同时也说明AhFAD2-1A/B功能的缺失并不是决定花生耐寒性的主要因素。本研究的开展为培育抗寒的高油酸花生品种奠定了理论基础,为高油酸花生在高纬度、高海拔地区推广提供了理论支持。
To explore the roles of FAD2s in response to cold stress in peanut, we cloned seven Ah FAD2 genes from normal oleate peanut ZH16 and high oleate peanut ZH413, respectively. The results of qRT-PCR showed that the expression patterns of FAD2 genes were similar in ZH16 and ZH413. AhFAD2-1 A/B was highly expressed in flower and developing seed, AhFAD2-3 A/B was mainly expressed in leaf and stem, and AhFAD2-4A/B was expressed specifically in root and flower, indicating that Ah FAD2 genes played their respective roles in different developmental stages and tissues of peanut. At 6 days after inducing under 15℃, the germination rate of ZH16 was significantly decreased while that of ZH413 was not significantly affected. The expression of both AhFAD2-1 A/B and AhFAD2-4 A/B were induced by cold stress. The expression of AhFAD2-1A/B was significantly up-regulated at 6 DAI in ZH16, while at 1 DAI in ZH413, suggesting that AhFAD2-1A/B was induced by cold more quickly in high oleate peanut. Furthermore, the expression of Ah FAD2-4A/B was significantly up-regulated at 3 DAI in ZH16 and then decreased, while it was increased immediately and maintained at high level for six days in ZH413. Based on these results, we speculate that up-regulation of AhFAD2-4 A/B may compensate the function of AhFAD2-1A/B that is deactivated under cold stress in high oleate peanut, and the deactivation of AhFAD2-1A/B is not the most important factors affecting peanut cold tolerance. This study provides a theoretical basis in breeding of high oleate peanut with high tolerance to cold stress, and the theoretical support for extension of high oleate peanut in both high latitude and high altitude regions.
To explore the roles of FAD2s in response to cold stress in peanut, we cloned seven Ah FAD2 genes from normal oleate peanut ZH16 and high oleate peanut ZH413, respectively. The results of qRT-PCR showed that the expression patterns of FAD2 genes were similar in ZH16 and ZH413. AhFAD2-1 A/B was highly expressed in flower and developing seed, AhFAD2-3 A/B was mainly expressed in leaf and stem, and AhFAD2-4A/B was expressed specifically in root and flower, indicating that Ah FAD2 genes played their respective roles in different developmental stages and tissues of peanut. At 6 days after inducing under 15℃, the germination rate of ZH16 was significantly decreased while that of ZH413 was not significantly affected. The expression of both AhFAD2-1 A/B and AhFAD2-4 A/B were induced by cold stress. The expression of AhFAD2-1A/B was significantly up-regulated at 6 DAI in ZH16, while at 1 DAI in ZH413, suggesting that AhFAD2-1A/B was induced by cold more quickly in high oleate peanut. Furthermore, the expression of Ah FAD2-4A/B was significantly up-regulated at 3 DAI in ZH16 and then decreased, while it was increased immediately and maintained at high level for six days in ZH413. Based on these results, we speculate that up-regulation of AhFAD2-4 A/B may compensate the function of AhFAD2-1A/B that is deactivated under cold stress in high oleate peanut, and the deactivation of AhFAD2-1A/B is not the most important factors affecting peanut cold tolerance. This study provides a theoretical basis in breeding of high oleate peanut with high tolerance to cold stress, and the theoretical support for extension of high oleate peanut in both high latitude and high altitude regions.
引文
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[22]张照华,王志慧,淮东欣,谭家壮,陈剑洪,晏立英,王晓军,万丽云,陈傲,康彦平,姜慧芳,雷永,廖伯寿.利用回交和标记辅助选择快速培育高油酸花生品种及其评价.中国农业科学,2018,51:1641-1652.Zhang Z H,Wang Z H,Huai D X,Tan J Z,Chen J H,Yan L Y,Wang X J,Wan L Y,Chen A,Kang Y P,Jiang H F,Lei Y,Liao BS.Fast development of high oleate peanut cultivars by using maker-assisted backcrossing and their evaluation.Sci Agric Sin,2018,51:1641-1652(in Chinese with English abstract).
[23]Jung S,Swift D,Sengoku E,Patel M,Teule F,Powell G,Moore K,Abbott A.The high oleate trait in the cultivated peanut(Arachis hypogaea L.).Isolation and characterization of two genes encoding microsomal oleoyl-PC desaturases.Mol Gen Genet,2000,263:796-805.
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[25]周丽侠,唐桂英,陈高,毕玉平,单雷.花生AhFAD2基因的多态性及其与籽粒油酸/亚油酸比值间的相关性.作物学报,2011,37:415-423.Zhou L Y,Tang G Y,Chen G,Bi Y P,Shan L.Correlation between AhFAD2 polymorphism and oleic acid/linoleic acid ratio in peanut seeds.Acta Agron Sin,2011,37:415-423(in Chinese with English abstract).
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[2]门爱军,庞兴国,胡东青,陆福军,李少骞.中国花生出口面临的困境及应对措施安徽农业科学,2016,44:257-258.Men A J,Pang X G,Hu D Q,Lu F J,Li S Q.Difficulties and countermeasures for China’s peanut export.J Anhui Agric Sci,2016,44:257-258(in Chinese without English abstract).
[3]刘娟,汤丰收,张俊,臧秀旺,董文召,易明林,郝西.国内花生生产技术现状及发展趋势研究.中国农学通报,2017,33(22):13-18.Liu J,Tang F S,Zhang J,Zang X W,Dong W Z,Yi M L,Hao X.Current status and development trends of peanut production technology in China.Chin Agric Sci Bull,2017,33(22):13-18(in Chinese with English abstract).
[4]Moore K M,Knauft D A.The inheritance of high oleic acid in peanut.Heredity,1989,80:8-10.
[5]王传堂,张建成,唐月异,于树涛,王强,刘峰,李秋.中国高油酸花生育种现状与展望.山东农业科学,2018,50(6):171-176.Wang C T,Zhang J C,Tang Y Y,Yu S T,Wang Q,Liu F,Li Q.Current situation and future directions of high oleic peanut breeding in China.Shandong Agric Sci,2018,50(6):171-176(in Chinese with English abstract).
[6]迟晓元,郝翠翠,陈明娜,潘丽娟,陈娜,王通,王冕,杨珍,梁成伟,禹山林.花生AhFAD2-1基因与油酸/亚油酸比值的关系.花生学报,2016,45(4):20-24.Chi X Y,Hao C C,Chen M N,Pan L J,Chen N,Wang T,Wang M,Yang Z,Liang C W,Yu S L.Correlation between Ah FAD2-1 and oleic acid/linoleic acid ratio in different peanut varieties.J Peanut Sci,2016,45(4):20-24(in Chinese with English abstract).
[7]Matos A R,Hourton-Cabassa C,Cicek D,Arrabaca J D,Zachowski A,Moreau F.Alternative oxidase involvement in cold stress response of Arabidopsis thaliana fad2 and fad3+cell suspensions altered in membrane lipid composition.Plant Cell Physiol,2007,48:856-865.
[8]Miquel M,James D,Dooner H,Browse A J.Arabidopsis requires polyunsaturated lipids for low-temperature survival.Proc Natl Acad Sci USA,1993,90:6208-6212.
[9]Watanabe K,Oura T,Sakai H,Kajiwara S.YeastΔ12 fatty acid desaturase:gene cloning,expression,and function.Biosci Biotechnol Biochem,2004,68:721-727.
[10]Kargiotidou A,Deli D,Galanopoulou D,Tsaftaris A,Farmaki T.Low temperature and light regulate delta 12 fatty acid desaturases(FAD2)at a transcriptional level in cotton(Gossypium hirsutum).J Exp Bot,2008,59:2043-2056.
[11]阮建,单雷,李新国,郭峰,孟静静,万书波,彭振英.花生FAD基因家族的全基因组鉴定与表达模式分析.山东农业科学,2018,50(6):1-9.Ruan J,Shan L,Li X G,Guo F,Meng J J,Wan S B,Peng Z Y.Genome-wide identification and expression pattern analysis of peanut FAD gene family.Shandong Agric Sci,2018,50(6):1-9(in Chinese with English abstract).
[12]Singh S C,Sinha R P,Haèder D P.Role of lipids and fatty acids in stress tolerance in cyanobacteria.Acta Protozoo,2002,41:297-308.
[13]张建军,闫世江,王浩,司龙亭,马志国,杨佳明．黄瓜种子脂肪酸含量与耐低温性关系的研究.安徽农业科学,2008,36:4859-4861.Zhang J J,Yan S J,Wang H,Si L T,Ma Z G,Yang J M.Study on the relationship between the contents of fatty acids and low temperature tolerance in cucumber seeds.J Anhui Agric Sci,2008,36:4859-4861(in Chinese with English abstract).
[14]Ishizaki-nishizawa O,Fujii T,Ohtani T,Toguri T.Lowtemperature resistance of higher plants is significantly enhanced by a nonspecific cyanobacterial desaturase.Nat Biotechnol,1996,14:1003-1006.
[15]黄锐之,刘智宏,郎春秀,胡张华,陈锦清.植物中多不饱和脂肪酸生物合成的基因工程.植物生理学通讯,2001,37:547-550.Huang R Z,Liu Z H,Lang C X,Hu Z H,Chen J Q.Genetic engineering of polyunsaturated fatty acid biosynthesis in plants.Plant Physiol Commun,2001,37:547-550(in Chinese with English abstract).
[16]陈儒钢,巩振辉,逯明辉,李大伟,黄炜.植物抗寒基因工程研究进展.西北植物学报,2008,28:1274-1280.Chen R G,Gong Z H,Lu M H,Li D W,Huang W.Research advance in the cold-resistance genetic engineering in plants.Acta Bot Boreali-Occident Sin,2008,28:1274-1280(in Chinese with English abstract).
[17]王洪春.植物抗逆性与生物膜结构功能研究的进展.植物生理学通讯,1985,(1):60-66.Wang H C.Advances in studies on plant stress resistance and biofilm structure.Plant Physiol Commun,1985,(1):60-66(in Chinese with English abstract).
[18]杨玉珍,罗青,彭方仁.不同种源香椿叶片脂肪酸组分含量与抗寒性关系.林业科技开发,2011,25(5):21-25.Yang Y Z,Luo Q,Peng F R.A relationship between cold resistance and content of fatty acid composition in leaves of Toona sinensis from different provenances.China For Sci Technol,2011,25(5):22-25(in Chinese with English abstract).
[19]戴晓峰,肖玲,武玉花,吴刚,卢长明.植物脂肪酸去饱和酶及其编码基因研究进展.植物学通报,2007,24:105-113.Dai X F,Xiao L,Wu Y H,Wu G,Lu C M.An overview of plant fatty acid desaturases and the coding genes.Chin Bull Bot,2007,24:105-113(in Chinese with English abstract).
[20]Yang Q,Fan C,Guo Z,Qin J,Wu J,Li Q,Fu T,Zhou Y.Identification of FAD2 and FAD3 genes in Brassica napus genome and development of allele-specific markers for high oleic and low linolenic acid contents.Theor Appl Genet,2012,125:715-729.
[21]Cao S J,Zhou X R,Wood C C,Green G A,Singh S P,Liu L X,Liu Q.A large and functionally diverse family of Fad2 genes in safflower(Carthamus tinctorius L.).BMC Plant Biol,2013,13:5.doi:10.1186/1471-2229-13-5.
[22]张照华,王志慧,淮东欣,谭家壮,陈剑洪,晏立英,王晓军,万丽云,陈傲,康彦平,姜慧芳,雷永,廖伯寿.利用回交和标记辅助选择快速培育高油酸花生品种及其评价.中国农业科学,2018,51:1641-1652.Zhang Z H,Wang Z H,Huai D X,Tan J Z,Chen J H,Yan L Y,Wang X J,Wan L Y,Chen A,Kang Y P,Jiang H F,Lei Y,Liao BS.Fast development of high oleate peanut cultivars by using maker-assisted backcrossing and their evaluation.Sci Agric Sin,2018,51:1641-1652(in Chinese with English abstract).
[23]Jung S,Swift D,Sengoku E,Patel M,Teule F,Powell G,Moore K,Abbott A.The high oleate trait in the cultivated peanut(Arachis hypogaea L.).Isolation and characterization of two genes encoding microsomal oleoyl-PC desaturases.Mol Gen Genet,2000,263:796-805.
[24]Wang Y,Zhang X G,Zhao Y L,Prakash C S,He G H,Yin D M.Insights into the novel members of the FAD2 gene family involved in high-oleate fluxes in peanut.NRC Res Press,2015,58:1-9.
[25]周丽侠,唐桂英,陈高,毕玉平,单雷.花生AhFAD2基因的多态性及其与籽粒油酸/亚油酸比值间的相关性.作物学报,2011,37:415-423.Zhou L Y,Tang G Y,Chen G,Bi Y P,Shan L.Correlation between AhFAD2 polymorphism and oleic acid/linoleic acid ratio in peanut seeds.Acta Agron Sin,2011,37:415-423(in Chinese with English abstract).
[26]D’Angeli S,Falasca G,Matteucci M M,Altamura M M.Cold perception and gene expression differ in Olea europaea seed coat and embryo during drupe cold acclimation.New Phytol,2013,197:123-138.
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