苦荞VQ基因家族的全基因组鉴定及其在叶斑病原与激素处理下的表达谱分析

doi:10.3864/j.issn.0578-1752.2021.19.002

中国农业科学 ›› 2021, Vol. 54 ›› Issue (19): 4048-4060.doi: 10.3864/j.issn.0578-1752.2021.19.002

• 作物遗传育种·种质资源·分子遗传学 • 上一篇下一篇

苦荞VQ基因家族的全基因组鉴定及其在叶斑病原与激素处理下的表达谱分析

郑逢盛¹(),王海华^1,²(),邬清韬¹,申权^1,²,田建红¹,彭喜旭^1,³,唐新科^1,³()

¹湖南科技大学生命科学学院,湖南湘潭 411201
²经济作物遗传改良与综合利用湖南省重点实验室,湖南湘潭 411201
³重金属污染生态土壤修复与安全利用湖南省高校重点实验室,湖南湘潭 411201

收稿日期:2021-02-25 接受日期:2021-06-07 出版日期:2021-10-01 发布日期:2021-10-12
通讯作者: 王海华,唐新科
作者简介:郑逢盛,E-mail： 13647321087@163.com。
基金资助:
国家重点研发计划(2017YFE0117600);湖南省教育厅重点科研项目(19A176)

Genome-Wide Identification of VQ Gene Family in Fagopyrum tataricum and Its Expression Profiles in Response to Leaf Spot Pathogens

ZHENG FengSheng¹(),WANG HaiHua^1,²(),WU QingTao¹,SHEN Quan^1,²,TIAN JianHong¹,PENG XiXu^1,³,TANG XinKe^1,³()

¹School of Life Science, Hunan University of Science and Technology, Xiangtan 411201, Hunan
²Key Laboratory of Genetic Improvement and Multiple Utilization of Economic Crops in Hunan Province, Xiangtan 411201, Hunan
³Key Laboratory of Ecological Remediation and Safe Utilization of Heavy Metal-polluted Soils, College of Hunan Province, Xiangtan 411201, Hunan

Received:2021-02-25 Accepted:2021-06-07 Online:2021-10-01 Published:2021-10-12
Contact: HaiHua WANG,XinKe TANG

摘要/Abstract

摘要：

【目的】VQ基因家族在植物生长、发育以及对生物或非生物胁迫反应中发挥重要功能。在全基因组尺度上,全面鉴定苦荞（Fagopyrum tataricum L. Gaertn.）VQ（FtVQ）基因家族,分析其在苦荞叶斑病原——互格链格孢（Alternaria alternata）和黑孢霉（Nigrospora osmanthi）侵染和防御相关激素——水杨酸（SA）、茉莉酸（JA）、乙烯（ET）处理下的表达模式,为深入解析苦荞VQ基因家族在植物抗病防御中的功能及机理奠定基础,同时为优良基因资源发掘及抗病品种改良提供线索。【方法】基于VQ保守结构域的隐马尔可夫文件（PF05678）,采用HMMER 3.0对苦荞平苦一号基因组数据库进行比对搜索,鉴定VQ基因;通过DNAMAN、MapInspect、MEGA、MEME、OrthoFinder、PLACE等生物信息学工具分析基因结构、染色体分布、启动子顺式元件、蛋白质理化性质、蛋白质保守基序、蛋白质亚细胞定位和蛋白质系统进化关系;采用实时荧光定量PCR（qPCR）方法分析苦荞叶VQ基因在病原侵染或激素处理下的表达模式。【结果】从苦荞基因组中鉴定获得28个VQ基因,大小为566—1 454 bp,均无内含子,不均一地分布在8条染色体上。根据它们在染色体上的物理位置,命名为FtVQ1—FtVQ28。每一个FtVQ蛋白含有1个VQ基序——FxxxVQx（L/F/I/V/A/Y）TG（x代表任意氨基酸）。亚细胞定位预测表明,21个FtVQ蛋白定位在细胞核中,其余定位在叶绿体或细胞质中。根据蛋白质氨基酸序列与保守结构基序,FtVQ蛋白归类于5个亚家族,亚家族内基因结构和蛋白质基序相对保守。基因重复分析表明,苦荞基因组中有8对VQ旁系同源基因,均为大片段重复基因,提示大片段基因重复在FtVQ基因家族数量扩张中发挥主要作用;它们的非同义突变和同义突变的比值（Ka/Ks）均小于1,提示重复基因在进化中经历了纯化选择。启动子顺式元件预测表明,所有FtVQ基因启动子含有BIHD1OS、CGTCA、ERELEA4、W-box和类W-box等病原或SA、JA、ET反应元件,尤其在FtVQ10、FtVQ14、FtVQ15、FtVQ22、FtVQ23、FtVQ27的启动子区域密集程度更高。qPCR分析显示,在可检测的20个FtVQ基因中,有55%—70%的基因为病原或激素处理下的差异表达基因（DEGs）,其中72.7%—85.7%的DEGs的表达显著上调。【结论】苦荞基因组拥有28个VQ基因成员,部分VQ基因可能参与了苦荞对叶斑病原的抗性反应。

关键词: 苦荞, VQ基因家族, 互格链格孢叶斑病, 黑孢霉叶斑病, 防御相关激素

Abstract:

【Objective】VQ gene family plays important roles in plant growth, development and responses to biotic or abiotic stress. The aim of this study is to comprehensively identify Fagopyrum tataricum L. Gaertn. VQ (FtVQ) gene family on genome-wide scale and analyze its expression profiles under challenge of leaf spot pathogens Alternaria. alternata and Nigrospora osmanthi, and treatment of defense-related hormones, such as salicylic acid (SA), jasmonic acid (JA) and ethylene (ET), thus providing a solid foundation not only for further elucidation possible roles of members of the VQ genes in defense response to leaf spot pathogens and underlying mechanisms in tartary buckwheat, but also for mining gene resources of breeding for crop disease resistance. 【Method】Based on the Hidden Markov Model profile of the conserved VQ domain (PF05678), HMMER 3.0 software was used to identify FtVQ genes from F. tataricum cv. Pinku1 genome database. Bioinformatic tools such as DNAMAN, MapInspect, MEGA, MEME, OrthoFinder and PLACE were used to analyze gene structure, chromosomal location of genes, cis-elements of gene promoters, physicochemical properties of proteins, conserved motifs of proteins, subcellular localization of proteins, and phylogenetic relationships. Quantitative real-time PCR (qPCR) was employed to analyze the expression profiles of leaf FtVQ genes of tartary buckwheat plants under infection of the pathogens and treatment of the hormones. 【Result】A total of 28 VQ genes were identified in the genomes of tartary buckwheat, with the gene size ranging from 566 to 1454 bp. The FtVQ genes contain no introns, and distribute unevenly on chromosomes 1-8. According to their physical locations on the chromosomes, the FtVQ genes were named from FtVQ1 to FtVQ28. Each of the FtVQ proteins has a highly conserved VQ motif FxxxVQx (L/F/I/V/A/Y) TG, where x represents any amino acid. Analysis of subcellular localization showed that 21 FtVQ proteins were predicted to the nucleus, and the others to the chloroplasts or cytoplasm. Based upon their amino acid sequence and presence of various conserved motifs, the FtVQ proteins were classified into five subfamilies (Ⅰ-Ⅴ). Each subfamily shared relatively conserved gene structures and protein motifs. The analysis of gene duplication revealed that F. tataricum genome had 8 pairs of paralogous pairs, all of which were segmental duplicated genes, suggesting that segmental duplication played major roles in FtVQ gene expansion. The ratio of nonsynonymous to synonymous substitutions (Ka/Ks) of paralogous pairs was less than 1, suggesting that they underwent purifying pressure during the evolution process. Prediction of cis-elements showed that pathogen-, SA-, JA-, or ET-responsive elements, such as BIHD1OS, CGTCA, ERELEA4, W-box and W-box-like sequences, were present within the promoters of all the FtVQ genes. Especially, FtVQ10, FtVQ14, FtVQ15, FtVQ22, FtVQ23 and FtVQ27 contained more elements in their promoter regions. 55% to 70% of the detectable FtVQ genes were differentially expressed genes (DEGs), and 72.7% to 85.7% of the DEGs were significantly induced on the level of transcription under the infection of leaf spot fungi A. alternata and N. osmanthi, or the treatment of SA, methyl jasmonate and ethephon. 【Conclusion】The tartary buckwheat genome contains 28 members of VQ gene family. Some FtVQ genes may be involved tartary buckwheat defense response to leaf spot pathogens.

Key words: tartary buckwheat (Fagopyrum tataricum (L.) Gaertn.), VQ gene family, Alternaria leaf spot, Nigrospora leaf spot, defense-related hormones

郑逢盛,王海华,邬清韬,申权,田建红,彭喜旭,唐新科. 苦荞VQ基因家族的全基因组鉴定及其在叶斑病原与激素处理下的表达谱分析[J]. 中国农业科学, 2021, 54(19): 4048-4060.

ZHENG FengSheng,WANG HaiHua,WU QingTao,SHEN Quan,TIAN JianHong,PENG XiXu,TANG XinKe. Genome-Wide Identification of VQ Gene Family in Fagopyrum tataricum and Its Expression Profiles in Response to Leaf Spot Pathogens[J]. Scientia Agricultura Sinica, 2021, 54(19): 4048-4060.

0
/ / 推荐

导出引用管理器 EndNote|Reference Manager|ProCite|BibTeX|RefWorks

链接本文: https://www.chinaagrisci.com/CN/10.3864/j.issn.0578-1752.2021.19.002

https://www.chinaagrisci.com/CN/Y2021/V54/I19/4048

图/表 9

表1

苦荞VQ基因及其序列特征"

基因名称 Gene name	序列登录号 Sequenced ID	染色体 Chromosome	内含子 Introns	正/反链 +/- strand	基因大小 Gene size (bp)	编码序列 Coding sequence	编码的蛋白质 Encoded protein
基因名称 Gene name	序列登录号 Sequenced ID	染色体 Chromosome	内含子 Introns	正/反链 +/- strand	基因大小 Gene size (bp)	编码序列 Coding sequence	氨基酸长度 aa length (aa)	分子量 Molecular length (kD)	等电点 pI	亚细胞定位 Subcellular localization
FtVQ1	FtPinG0005794900	1	0	+	772	549	182	20.00	8.71	细胞核Nucleus
FtVQ2	FtPinG0005409400	1	0	-	1059	588	195	20.90	10.11	细胞核Nucleus
FtVQ3	FtPinG0002545500	1	0	+	678	678	225	24.27	10.31	细胞核Nucleus
FtVQ4	FtPinG0002567300	1	0	-	654	651	216	23.52	7.07	细胞核Nucleus
FtVQ5	FtPinG0006482000	1	0	+	566	249	82	9.08	6.50	细胞核Nucleus
FtVQ6	FtPinG0001835400	2	0	+	688	405	134	15.21	11.09	叶绿体Chloroplast
FtVQ7	FtPinG0009771100	2	0	+	840	840	279	30.09	8.36	细胞核Nucleus
FtVQ8	FtPinG0004496500	2	0	-	938	297	98	10.54	9.83	细胞核Nucleus
FtVQ9	FtPinG0001284900	2	0	+	629	603	200	21.83	8.79	叶绿体Chloroplast
FtVQ10	FtPinG0001084300	3	0	+	895	507	168	18.02	9.82	细胞核Nucleus
FtVQ11	FtPinG0001089800	3	0	+	705	705	234	24.66	5.94	细胞核Nucleus
FtVQ12	FtPinG0002126400	3	0	+	780	729	242	25.78	10.20	细胞核Nucleus
FtVQ13	FtPinG0007726900	4	0	+	418	387	128	14.60	7.67	细胞核Nucleus
FtVQ14	FtPinG0001986300	4	0	+	357	354	116	13.24	9.52	细胞质Cytoplasm
FtVQ15	FtPinG0003123700	5	0	+	601	468	155	16.79	8.27	叶绿体Chloroplast
FtVQ16	FtPinG0007087100	5	0	+	453	429	142	15.71	10.54	细胞核Nucleus
FtVQ17	FtPinG0005585600	5	0	+	687	687	228	24.25	5.26	细胞核Nucleus
FtVQ18	FtPinG0002695900	6	0	+	850	603	200	21.09	5.20	叶绿体Chloroplast
FtVQ19	FtPinG0001480300	6	0	+	1454	1083	360	38.10	7.40	细胞核Nucleus
FtVQ20	FtPinG0006939600	6	0	+	899	477	158	17.28	5.56	细胞核Nucleus
FtVQ21	FtPinG0003079000	6	0	+	606	606	201	20.96	9.64	细胞核Nucleus
FtVQ22	FtPinG0009737600	6	0	+	797	594	197	21.25	6.08	细胞核Nucleus
FtVQ23	FtPinG0002065200	6	0	+	588	582	193	20.86	8.15	细胞核Nucleus
FtVQ24	FtPinG0002067700	6	0	+	468	444	147	16.01	4.48	细胞核Nucleus
FtVQ25	FtPinG0003595000	7	0	+	1343	882	293	30.96	10.77	细胞核Nucleus
FtVQ26	FtPinG0002292500	8	0	-	740	642	213	22.98	10.72	叶绿体Chloroplast
FtVQ27	FtPinG0003287200	8	0	+	699	630	209	22.81	6.62	细胞核Nucleus
FtVQ28	FtPinG0003376500	8	0	+	654	654	217	23.91	7.24	叶绿体Chloroplast

表1

图1

图2

图3

图4

表2

图5

图6

图7

参考文献 29

[1]	CHI Y, YANG Y, ZHOU Y, ZHOU J, FAN B, YU J Q, CHEN Z. Protein-protein interactions in the regulation of WRKY transcription factors. Molecular Plant, 2013, 6(2):287-300. doi: 10.1093/mp/sst026
[2]	JING Y, LIN R. The VQ motif-containing protein family of plant- specific transcriptional regulators. Plant Physiology, 2015, 169(1):371-378. doi: 10.1104/pp.15.00788
[3]	CHENG Y, ZHOU Y, YANG Y, CHI Y J, ZHOU J, CHEN J Y, WANG F, FAN B, SHI K, ZHOU Y H, YU J Q, CHEN Z. Structural and functional analysis of VQ motif-containing proteins in Arabidopsis as interacting proteins of WRKY transcription factors. Plant Physiology, 2012, 159(2):810-825. doi: 10.1104/pp.112.196816
[4]	WEYHE M, ESCHEN-LIPPOLD L, PECHER P, SCHEEL D, LEE J. Ménage à trois: The complex relationships between mitogen- activated protein kinases, WRKY transcription factors, and VQ-motif- containing proteins. Plant Signaling & Behavior, 2014, 9(8):29519.
[5]	ZHANG L, LI X, MA B, GAO Q, DU H, HAN Y, LI Y, CAO Y, QI M, ZHU Y, LU H, MA M, LIU L, ZHOU J, NAN C, QIN Y, WANG J, CUI L, LIU H, LIANG C, QIAO Z. The tartary buckwheat genome provides insights into rutin biosynthesis and abiotic stress tolerance. Molecular Plant, 2017, 10(9):1224-1237. doi: 10.1016/j.molp.2017.08.013
[6]	YANG J, DUAN G, LI C, LIU L, HAN G, ZHANG Y, WANG C. The crosstalks between jasmonic acid and other plant hormone signaling highlight the involvement of jasmonic acid as a core component in plant response to biotic and abiotic stresses. Frontiers in Plant Science, 2019, 10:1349. doi: 10.3389/fpls.2019.01349
[7]	KIM DY, KWON S I, CHOI C, LEE H, AHN I, PARK S R, BAE S C, LEE S C, HWANG D J. Expression analysis of rice VQ genes in response to biotic and abiotic stresses. Gene, 2013, 529(2):208-214. doi: 10.1016/j.gene.2013.08.023
[8]	LI N, LI X, XIAO J, WANG S. Comprehensive analysis of VQ motif-containing gene expression in rice defense responses to three pathogens. Plant Cell Report, 2014 33(9):1493-1505. doi: 10.1007/s00299-014-1633-4
[9]	ZHOU Y, YANG Y, ZHOU X, CHI Y, FAN B, CHEN Z. Structural and functional characterization of the VQ protein family and VQ protein variants from soybean. Scientific Report, 2016, 6:34663.
[10]	LIU C, LIU H, ZHOU C, TIMKO M P. Genome-wide identification of the VQ protein gene family of tobacco (Nicotiana tabacum L.) and analysis of its expression in response to phytohormones and abiotic and biotic stresses. Genes, 2020, 11(3):284. doi: 10.3390/genes11030284
[11]	LAI Z, LI Y, WANG F, CHENG Y, FAN B, YU J Q, CHEN Z. Arabidopsis sigma factor binding proteins are activators of the WRKY33 transcription factor in plant defense. The Plant Cell, 2011, 23(10):3824-3841. doi: 10.1105/tpc.111.090571
[12]	WANG H, HU Y, PAN J, YU D. Arabidopsis VQ motif-containing proteins VQ12 and VQ29 negatively modulate basal defense against Botrytis cinerea. Scientific Report, 2015, 5:14185.
[13]	ALI M R M, UEMURA T, RAMADAN A, ADACHI K, NEMOTO K, NOZAWA A, HOSHINO R, ABE H, SAWASAKI T, ARIMURA G I. The ring-type E3 ubiquitin ligase JUL1 targets the VQ-motif protein JAV1 to coordinate jasmonate signaling. Plant Physiology, 2019, 179(4):1273-1284. doi: 10.1104/pp.18.00715
[14]	CHU W Y, LIU B, WANG Y J, PAN F, CHEN Z, YAN H W, XIANG Y. Genome-wide analysis of poplar VQ gene family and expression profiling under PEG, NaCl, and SA treatments. Tree Genetics & Genomes, 2016, 12:124.
[15]	SONG W, ZHAO H, ZHANG X, LEI L, LAI J. Genome-wide identification of VQ motif-containing proteins and their expression profiles under abiotic stresses in maize. Frontiers in Plant Science, 2016, 6:1177.
[16]	ZHANG G, WANG F, LI J, DING Q, ZHANG Y, LI H, ZHANG J, GAO J. Genome-wide identification and analysis of the VQ motif-containing protein family in Chinese cabbage (Brassica rapa L. ssp. Pekinensis). International Journal of Molecular Sciences, 2015, 16(12):28683-28704. doi: 10.3390/ijms161226127
[17]	WANG M, VANNOZZI A, WANG G, ZHONG Y, CORSO M, CAVALLINI E, CHENG Z M. A comprehensive survey of the grapevine VQ gene family and its transcriptional correlation with WRKY proteins. Frontiers in Plant Science, 2015, 6:417.
[18]	CHEN P, WEI F, CHENG S, MA L, WANG H, ZHANG M, MAO G, LU J, HAO P, AHMAD A, GU L, MA Q, WU A, WEI H, YU S. A comprehensive analysis of cotton VQ gene superfamily reveals their potential and extensive roles in regulating cotton abiotic stress. BMC Genomics, 2020, 21(1):795. doi: 10.1186/s12864-020-07171-z
[19]	GARRIDO-GALA J, HIGUERA J J, MUÑOZ-BLANCO J, AMIL-RUIZ F, CABALLERO J L. The VQ motif-containing proteins in the diploid and octoploid strawberry. Scientific Report, 2019, 9(1):4942.
[20]	WANG Y, LIU H, ZHU D, GAO Y, YAN H, XIANG Y. Genome-wide analysis of VQ motif-containing proteins in Moso bamboo (Phyllostachys edulis). Planta, 2017, 246(1):165-181. doi: 10.1007/s00425-017-2693-9
[21]	LING L, QU Y, ZHU J, WANG D, GUO C. Genome-wide identification and expression analysis of the VQ gene family in Cicer arietinum and Medicago truncatula. PeerJ, 2020, 8:8471.
[22]	SHEN Q, PENG X X, HE F, LI S Q, XIAO Z Y, WANG H H, TANG X K, ZHOU M L. First report of Nigrospora osmanthi causing leaf spot on tartary buckwheat in China. Plant Disease, 2020. doi: 10.1094/PDIS-08-20-1773-PDN. doi: 10.1094/PDIS-08-20-1773-PDN
[23]	LYNCH M, CONERY J S. The evolutionary fate and consequences of duplicate genes. Science, 2000, 290(5494):1151-1155. doi: 10.1126/science.290.5494.1151
[24]	LI W H, GOJOBORI T, NEI M. Pseudogenes as a paradigm of neutral evolution. Nature, 1981, 292:237-239. doi: 10.1038/292237a0
[25]	STORZ J F. Genome evolution: Gene duplication and the resolution of adaptive conflict. Heredity, 2009, 102(2):99-100. doi: 10.1038/hdy.2008.114
[26]	ZHANG G, WEI B. Characterization of VQ motif-containing protein family and their expression patterns under phytohormones and abiotic stresses in melon (Cucumis melo L.). Plant Growth Regulation, 2019, 89(3):273-285. doi: 10.1007/s10725-019-00534-x
[27]	JIANG S Y, SEVUGAN M, RAMACHANDRAN S. Valine-glutamine (VQ) motif coding genes are ancient and non-plant-specific with comprehensive expression regulation by various biotic and abiotic stress. BMC Genomics, 2018, 19:342. doi: 10.1186/s12864-018-4733-7
[28]	UJI Y, KASHIHARA K, KIYAMA H, MOCHIZUKI S, AKIMITSU K, GOMI K. Jasmonic acid-induced VQ-motif-containing protein OsVQ13 influences the OsWRKY45 signaling pathway and grain size by associating with OsMPK6 in rice. International Journal of Molecular Science, 2019, 20:2917. doi: 10.3390/ijms20122917
[29]	EULGEM T, RUSHTON P J, ROBATZEK S, SOMSSICH I E. The WRKY superfamily of plant transcription factors. Trends in Plant Science, 2000, 5(5):199-206. doi: 10.1016/S1360-1385(00)01600-9

旁系同源基因对 Paralogous pairs	Ka值 Ka value	Ks值 Ks value	Ka/Ks	重复年代（百万年） Duplication date (Mya)
FtVQ2-FtVQ10	0.5833	1.9208	0.3037	105.54
FtVQ7-FtVQ19	0.7035	3.3733	0.2085	185.35
FtVQ23-FtVQ27	0.4747	2.8328	0.1680	155.65
FtVQ15-FtVQ28	0.5945	3.4488	0.1724	189.50
FtVQ1-FtVQ12	0.4369	0.8331	0.5244	45.77
FtVQ11-FtVQ17	0.2927	1.3880	0.2110	76.27
FtVQ14-FtVQ24	1.0473	1.3114	0.7986	72.05
FtVQ4-FtVQ9	0.1333	5.0748	0.0260	278.83

苦荞VQ基因家族的全基因组鉴定及其在叶斑病原与激素处理下的表达谱分析

Genome-Wide Identification of VQ Gene Family in Fagopyrum tataricum and Its Expression Profiles in Response to Leaf Spot Pathogens

RichHTML

PDF

赞

可视化

摘要/Abstract

引用本文

使用本文

图/表 9

参考文献 29

相关文章 13

Metrics

本文评价

推荐阅读 0

[1]	赵海霞,肖欣,董玘鑫,吴花拉,李成磊,吴琦. 苦荞愈伤遗传转化体系的优化及用于FtCHS1的过表达分析[J]. 中国农业科学, 2022, 55(9): 1723-1734.
[2]	侯思宇,王欣芳,杜伟,冯晋华,韩渊怀,李红英,刘龙龙,孙朝霞. 苦荞WOX家族全基因组鉴定及响应愈伤诱导率表达分析[J]. 中国农业科学, 2021, 54(17): 3573-3586.
[3]	郝彦蓉,杜伟,侯思宇,王东航,冯红梅,韩渊怀,周美亮,张凯旋,刘龙龙,王俊珍,李红英,孙朝霞. 苦荞ARF基因家族的鉴定及生长素诱导下的表达模式[J]. 中国农业科学, 2020, 53(23): 4738-4749.
[4]	吴曹阳,梁诗涵,邱军,高金锋,高小丽,王鹏科,冯佰利,杨璞. 基于连续12年国家苦荞区域试验的中国苦荞品种选育现状分析[J]. 中国农业科学, 2020, 53(19): 3878-3892.
[5]	孙朝霞，侯思宇，令狐斌，刘荣华，王丽，杨武德，韩渊怀. 苦荞全生育期芦丁积累与其生物合成途径相关基因表达分析[J]. 中国农业科学, 2017, 50(18): 3473-3481.
[6]	路之娟，张永清，张楚，刘丽琴，杨春婷. 不同基因型苦荞苗期抗旱性综合评价及指标筛选[J]. 中国农业科学, 2017, 50(17): 3311-3322.
[7]	屈洋，周瑜，王钊，王鹏科，高金锋，高小丽，冯佰利. 苦荞产区种质资源遗传多样性和遗传结构分析[J]. 中国农业科学, 2016, 49(11): 2049-2062.
[8]	刘琴, 张薇娜, 朱媛媛, 胡秋辉. 不同产地苦荞籽粒中多酚的组成、分布及抗氧化性比较[J]. 中国农业科学, 2014, 47(14): 2840-2852.
[9]	高帆, 张宗文, 吴斌. 中国苦荞SSR分子标记体系构建及其在遗传多样性分析中的应用[J]. 中国农业科学, 2012, 45(6): 1042-1053.
[10]	孙朝霞, 侯思宇, 杨武德. 苯丙氨酸与UV-C对苦荞芦丁含量影响及相关基因表达分析[J]. 中国农业科学, 2011, 44(23): 4772-4780.
[11]	张强, 李艳琴. 基于矿质元素的苦荞产地判别研究[J]. 中国农业科学, 2011, 44(22): 4653-4659.
[12]	侯雅君,张宗文,吴斌,李艳琴 . 苦荞种质资源AFLP标记遗传多样性分析[J]. 中国农业科学, 2009, 42(12): 4166-4174 .
[13]	魏益民,张国权,Sietz,W.. 同源四倍体荞麦籽粒品质性状研究[J]. 中国农业科学, 1995, 28(增刊): 34-40 .