伴矿景天WRKY基因家族鉴定及镉胁迫响应分析

doi:10.12302/j.issn.1000-2006.202201015

国家林草科技领军期刊
中国精品科技期刊
中国高校百佳科技期刊
江苏省新闻出版政府奖期刊奖
RCCSE林学权威期刊（A+）
CSCD核心期刊
Scopus数据库收录期刊
中文核心期刊
SCD核心期刊

作者加群：102861116

微信公众号：南京林业大学学报

高级检索

南京林业大学学报（自然科学版） ›› 2023, Vol. 47 ›› Issue (2): 49-60.doi: 10.12302/j.issn.1000-2006.202201015

伴矿景天WRKY基因家族鉴定及镉胁迫响应分析

王剑超¹^,²(), 邱文敏²(), 金康鸣², 陆铸畴², 韩小娇², 卓仁英², 刘晓光³^,^*(), 何正权¹^,^*()

1.三峡区域植物遗传与种质创新重点实验室(三峡大学),三峡大学生物与制药学院,湖北宜昌 443002
2.中国林业科学研究院亚热带林业研究所,浙江省林木育种技术研究重点实验室,浙江杭州 311400
3.唐山市农业科学研究院,河北唐山 063001

收稿日期:2022-01-12 修回日期:2022-06-25 出版日期:2023-03-30 发布日期:2023-03-28
通讯作者: * 何正权(zhq_he@163.com),教授,负责选题与实验方案设计;刘晓光(nyslxg@126.com),副研究员,负责数据分析与论文修改。
基金资助:
国家重点研发计划子课题(2016YFD080080104)

Comprehensive analysis of WRKY gene family in Sedum plumbizincicola responding to cadmium stress

WANG Jianchao¹^,²(), QIU Wenmin²(), JIN Kangming², LU Zhuchou², HAN Xiaojiao², ZHUO Renying², LIU Xiaoguang³^,^*(), HE Zhengquan¹^,^*()

1. Key Laboratory of Three Gorges Regional Plant Genetics & Germplasm Enhancement (CTGU),Biotechnology Research Center,China Three Gorges University, Yichang 443002, China
2. Key Laboratory of Tree Breeding of Zhejiang Province, The Research Institute of Subtropical Forestry, Chinese Academy of Forestry, Hangzhou 311400, China
3. Tangshan Academy of Agriculture Science, Tangshan 063001, China

Received:2022-01-12 Revised:2022-06-25 Online:2023-03-30 Published:2023-03-28

摘要/Abstract

摘要：

【目的】WRKY 转录因子在植物响应非生物胁迫过程中发挥着重要的调控作用,目前超积累植物伴矿景天(Sedum plumbizincicola) WRKY转录因子的研究较少。进行SpWRKY基因家族的全基因组鉴定及其在镉胁迫下的表达模式分析可为后续SpWRKYs 基因克隆和耐镉功能分析提供参考。【方法】对伴矿景天WRKY基因家族进行全基因组鉴定和生物信息学分析,利用转录组数据和qRT-PCR 技术检测WRKY基因在镉胁迫下的表达模式并进行转基因拟南芥耐镉性的分析。【结果】伴矿景天基因组中共鉴定出77个SpWRKYs,非均匀地分布在各条染色体上;系统进化树分析发现,伴矿景天WRKY成员被分成3大类群(Ⅰ—Ⅲ),第Ⅱ类群又被分成5个亚群(Ⅱa—e);共线性分析发现,伴矿景天WRKY基因家族和拟南芥(Arabidopsis thaliana) 之间存在7个共线基因对,和玉吊钟(Kalanchoe fedtschenkoi)之间存在53个共线基因对;SpWRKYs基因之间存在 19 个片段复制基因对;启动子序列分析发现,SpWRKYs 启动子有多种与激素和逆境响应等相关的顺式调控元件;镉胁迫表达分析发现,7 个SpWRKYs 受镉胁迫显著上调表达,整体表达趋势呈现先增后减的状态;在拟南芥中过表达SpWRKY69发现,SpWRKY69可以增强Cd离子向地上部的转运速率,对植株镉耐性起到负调控作用。【结论】伴矿景天WRKY基因家族与其他物种的WRKY家族结构相似,片段复制是其主要进化动力之一。镉胁迫条件下部分成员相对表达量显著变化,而SpWRKY69能增加Cd离子向地上部的转运速率,故其他WRKY成员可能也参与调控植株镉耐性。

关键词: 伴矿景天, WRKY转录因子, 镉胁迫, 基因表达, 拟南芥转基因

Abstract:

【Objective】 WRKY transcription factors play important regulatory roles for a plant abiotic stress response. However, little information is available about WRKY transcription factors in the hyperaccumulating plant Sedum plumbizincicola. The identification of SpWRKY gene family members and analysis of their expression patterns under cadmium stress can provide a reference for molecular cloning of SpWRKYs gene and functional characterization of cadmium tolerance.【Method】 In this study, the genome-wide identification and bioinformatics analysis of WRKY gene family members were carried out. Expression patterns of WRKY genes under cadmium stress were derived from transcriptome data and qPCR, and the roles of SpWRKY69 in cadmium tolerance were assessed by the heterologous expression in Arabidopsis thaliana.【Result】 There were 77 SpWRKYs identified in S. plumbizincicola, unevenly distributed on the chromosomes. Based on the phylogenetic analysis, the SpWRKY proteins were classified into three groups (Ⅰ-Ⅲ), among which the second group was further divided into five subgroups (Ⅱa-e). The synteny analysis showed that there were seven collinear gene pairs in the S. plumbizincicola and A. thaliana WRKY gene family and 53 collinear gene pairs for Kalanchoe fedtschenkoi; 19 pairs of SpWRKYs were identified as the segmental duplication. Cis-regulatory elements related to stress and hormone responses were found in the promoters of SpWRKYs. The expression profiles showed that the expression levels of the seven SpWRKYs were significantly up-regulated under cadmium stress, exhibiting a pattern of increasing initially and then decreasing. Over-expression of SpWRKY69 in A. thaliana showed that SpWRKY69 can enhance the transport rate of Cd ions to the shoot and play a negative role in regulating plant cadmium tolerance.【Conclusion】 Structural features of WRKY gene family members in S. plumbizincicola are similar to those of other species, and the fragment duplication is one of the main evolutionary forces. The relative expression levels of some members change significantly under cadmium stress, and SpWRKY69 can increase the transport rate of Cd ions to the shoot, and so other WRKY members may also be involved in the regulation of plant cadmium tolerance. These results provide a foundation for further functional characterization of SpWRKYs related to Cd tolerance.

Key words: Sedum plumbizincicola, WRKY transcription factor, cadmium stress, gene expression, Arabidopsis transformation

中图分类号:

S718

王剑超,邱文敏,金康鸣,等. 伴矿景天WRKY基因家族鉴定及镉胁迫响应分析[J]. 南京林业大学学报（自然科学版）, 2023, 47(2): 49-60.

WANG Jianchao, QIU Wenmin, JIN Kangming, LU Zhuchou, HAN Xiaojiao, ZHUO Renying, LIU Xiaoguang, HE Zhengquan. Comprehensive analysis of WRKY gene family in Sedum plumbizincicola responding to cadmium stress[J].Journal of Nanjing Forestry University (Natural Science Edition), 2023, 47(2): 49-60.DOI: 10.12302/j.issn.1000-2006.202201015.

图/表 10

表1

图1

图2

图3

图4

图5

图6

图7

图8

图9

参考文献 48

[1]	ISHIGURO S, NAKAMURA K. Characterization of a cDNA encoding a novel DNA-binding protein,SPF1,that recognizes SP8 sequences in the 5’ upstream regions of genes coding for sporamin and β-amylase from sweet potato[J]. Molec Gen Genet, 1994, 244(6):563-571.DOI:10.1007/BF00282746.
[2]	EULGEM T, RUSHTON P J, ROBATZEK S, et al. The WRKY superfamily of plant transcription factors[J]. Trends Plant Sci, 2000, 5(5):199-206.DOI:10.1016/S1360-1385(00)01600-9.
[3]	WANG L N, ZHU W, FANG L C, et al. Genome-wide identification of WRKY family genes and their response to cold stress in Vitis vinifera[J]. BMC Plant Biol, 2014, 14:103.DOI:10.1186/1471-2229-14-103.
[4]	REN C M, ZHU Q, GAO B D, et al. Transcription factor WRKY70 displays important but no indispensable roles in jasmonate and salicylic acid signaling[J]. J Integr Plant Biol, 2008, 50(5):630-637.DOI:10.1111/j.1744-7909.2008.00653.x.
[5]	GUILLAUMIE S, MZID R, MÉCHIN V, et al. The grapevine transcription factor WRKY2 influences the lignin pathway and xylem development in tobacco[J]. Plant Mol Biol, 2010, 72(1):215.DOI:10.1007/s11103-009-9563-1.
[6]	JIANG C H, HUANG Z Y, XIE P, et al. Transcription factors WRKY70 and WRKY11 served as regulators in rhizobacterium Bacillus cereus AR156-induced systemic resistance to Pseudomonas syringae pv.tomato DC3000 in Arabidopsis[J]. J Exp Bot, 2015, 67(1):157-174.DOI:10.1093/jxb/erv445.
[7]	SHENG Y B, YAN X X, HUANG Y, et al. The WRKY transcription factor,WRKY13,activates PDR8 expression to positively regulate cadmium tolerance in Arabidopsis[J]. Plant Cell Environ, 2019, 42(3):891-903.DOI:10.1111/pce.13457.
[8]	TAO Z, KOU Y J, LIU H B, et al. OsWRKY45 alleles play different roles in abscisic acid signalling and salt stress tolerance but similar roles in drought and cold tolerance in rice[J]. J Exp Bot, 2011, 62(14):4863-4874.DOI:10.1093/jxb/err144.
[9]	CHU X Q, WANG C, CHEN X B, et al. The cotton WRKY gene GhWRKY41 positively regulates salt and drought stress tolerance in transgenic Nicotiana benthamiana[J]. PLoS One, 2015, 10(11):e0143022.DOI:10.1371/journal.pone.0143022.
[10]	LIU Z Q, FANG H H, PEI Y X, et al. WRKY transcription factors down-regulate the expression of H₂S-generating genes,LCD and DES in Arabidopsis thaliana[J]. Sci Bull, 2015, 60(11):995-1001.DOI:10.1007/s11434-015-0787-y.
[11]	SUN Y D, YU D Q. Activated expression of AtWRKY53 negatively regulates drought tolerance by mediating stomatal movement[J]. Plant Cell Rep, 2015, 34(8):1295-1306.DOI:10.1007/s00299-015-1787-8.
[12]	中国生态环境部. 2014全国土壤污染状况调查公报[EB/OL].(2014-04-17)[2021-10-12]. https://www.mee.gov.cn/gkml/sthjbgw/qt/201404/t20140417_270670.htm.
[13]	LANE T W, MOREL F M. A biological function for cadmium in marine diatoms[J]. Proc Natl Acad Sci USA, 2000, 97(9):4627-4631.DOI:10.1073/pnas.090091397.
[14]	ANDRESEN E, KÜPPER H. Cadmium toxicity in plants[J]. Met Ions Life Sci, 2013, 11:395-413.DOI:10.1007/978-94-007-5179-8_13.
[15]	RODRÍGUEZ-SERRANO M, ROMERO-PUERTAS M C, PAZMIÑO D M, et al. Cellular response of pea plants to cadmium toxicity:cross talk between reactive oxygen species,nitric oxide,and calcium[J]. Plant Physiol, 2009, 150(1):229-243.DOI:10.1104/pp.108.131524.
[16]	CHENG S P. Effects of heavy metals on plants and resistance mechanisms.A state-of-the-art report with special reference to literature published in Chinese journals[J]. Environ Sci Pollut Res Int, 2003, 10(4):256-264.DOI:10.1065/espr2002.11.141.2.
[17]	FAROON O, ASHIZAWA A, WRZAHJ S, et al. Toxicological profile for cadmium[R]. Cleveland: USA. US Department of Energy, 1989.
[18]	MUSZYNSKA E, HANUS-FAJERSKA E. Why are heavy metal hyperaccumulating plants so amazing?[J]. BioTechnologia, 2015, 4:265-271.DOI:10.5114/bta.2015.57730.
[19]	LI J T, GURAJALA H K, WU L H, et al. Hyperaccumulator plants from China:a synthesis of the current state of knowledge[J]. Environ Sci Technol, 2018, 52(21):11980-11994.DOI:10.1021/acs.est.8b01060.
[20]	PENG J S, WANG Y J, DING G, et al. A pivotal role of cell wall in cadmium accumulation in the Crassulaceae hyperaccumulator Sedum plumbizincicola[J]. Mol Plant, 2017, 10(5):771-774.DOI:10.1016/j.molp.2016.12.007.
[21]	XU D, LU Z C, JIN K M, et al. SPDE:a multi-functional software for sequence processing and data extraction[J]. Bioinformatics, 2021, 37(20):3686-3687.DOI:10.1093/bioinformatics/btab235.
[22]	EDGAR R C. MUSCLE:Multiple sequence alignment with high accuracy and high throughput[J]. Nucleic Acids Res, 2004, 32(5):1792-1797.DOI:10.1093/nar/gkh340.
[23]	WATERHOUSE A M, PROCTER J B, MARTIN D M A, et al. Jalview Version 2: a multiple sequence alignment editor and analysis workbench[J]. Bioinformatics, 2009, 25(9):1189-1191.DOI:10.1093/bioinformatics/btp033.
[24]	KUMAR S, STECHER G, LI M, et al. MEGA X:molecular evolutionary genetics analysis across computing platforms[J]. Mol Biol Evol, 2018, 35(6):1547-1549.DOI:10.1093/molbev/msy096.
[25]	ZHANG H K, GAO S H, LERCHER M J, et al. EvolView,an online tool for visualizing,annotating and managing phylogenetic trees[J]. Nucleic Acids Res, 2012, 40(W1):W569-W572.DOI:10.1093/nar/gks576.
[26]	CHEN C J, CHEN H, ZHANG Y, et al. TBtools:an integrative toolkit developed for interactive analyses of big biological data[J]. Mol Plant, 2020, 13(8):1194-1202.DOI:10.1016/j.molp.2020.06.009.
[27]	LESCOT M, DÉHAIS P, THIJS G, et al. PlantCARE,a database of plant cis-acting regulatory elements and a portal to tools for in silico analysis of promoter sequences[J]. Nucleic Acids Res, 2002, 30(1):325-327.DOI:10.1093/nar/30.1.325.
[28]	WANG Y P, TANG H B, DEBARRY J D, et al. MCScanX:a toolkit for detection and evolutionary analysis of gene synteny and collinearity[J]. Nucleic Acids Res, 2012, 40(7):e49.DOI:10.1093/nar/gkr1293.
[29]	HAN X J, YIN H F, SONG X X, et al. Integration of small RNAs,degradome and transcriptome sequencing in hyperaccumulator Sedum alfredii uncovers a complex regulatory network and provides insights into cadmium phytoremediation[J]. Plant Biotechnol J, 2016, 14(6):1470-1483.DOI:10.1111/pbi.12512.
[30]	SHANNON P, MARKIEL A, OZIER O, et al. Cytoscape:a software environment for integrated models of biomolecular interaction networks[J]. Genome Res, 2003, 13(11):2498-2504.DOI:10.1101/gr.1239303.
[31]	CLOUGH S J, BENT A F. Floral dip:a simplified method for agrobacterium-mediated transformation of Arabidopsis thaliana[J]. Plant J, 1998, 16(6):735-743.DOI:10.1046/j.1365-313x.1998.00343.x.
[32]	SAMBROOK J, FRITSCH E, MANIATIS T. Molecular cloning:a laboratory manual[J]. Trends Biotechnol, 1991, 9(1): 213-214.
[33]	ALVAREZ M E, PENNELL R I, MEIJER P J, et al. Reactive oxygen intermediates mediate a systemic signal network in the establishment of plant immunity[J]. Cell, 1998, 92(6):773-784.DOI:10.1016/S0092-8674(00)81405-1.
[34]	WOHLGEMUTH H, MITTELSTRASS K, KSCHIESCHAN S, et al. Activation of an oxidative burst is a general feature of sensitive plants exposed to the air pollutant ozone[J]. Plant Cell Environ, 2002, 25(6):717-726.DOI:10.1046/j.1365-3040.2002.00859.x.
[35]	罗夏琳, 胡玉斐, 李攻科. 微波辅助消解-电感耦合等离子体原子发射光谱测定烟草中的重金属[J]. 分析科学学报, 2016, 32(2):249-252.DOI:10.13526/j.issn.1006-6144.2016.02.020.
	LUO X L, HU Y F, LI G K. Determination of heavy metals in tobaccos by microwave-assisted digestion-inductively coupled plasma-optical emission spectrometry[J]. J Anal Sci, 2016, 32(2):249-252.DOI:10.13526/j.issn.1006-6144.2016.02.020.
[36]	ÜLKER B, SOMSSICH I E. WRKY transcription factors:from DNA binding towards biological function[J]. Curr Opin Plant Biol, 2004, 7(5):491-498.DOI:10.1016/j.pbi.2004.07.012.
[37]	RUSHTON P J, SOMSSICH I E, RINGLER P, et al. WRKY transcription factors[J]. Trends Plant Sci, 2010, 15(5):247-258.DOI:10.1016/j.tplants.2010.02.006.
[38]	KIM C Y, ZHANG S Q. Activation of a mitogen-activated protein kinase cascade induces WRKY family of transcription factors and defense genes in tobacco[J]. Plant J, 2004, 38(1):142-151.DOI:10.1111/j.1365-313x.2004.02033.x.
[39]	XIE Z, ZHANG Z L, ZOU X L, et al. Annotations and functional analyses of the rice WRKY gene superfamily reveal positive and negative regulators of abscisic acid signaling in aleurone cells[J]. Plant Physiol, 2005, 137(1):176-189.DOI:10.1104/pp.104.054312.
[40]	GUPTA S, MISHRA V K, KUMARI S, et al. Deciphering genome-wide WRKY gene family of Triticum aestivum L[J]. Genes Genom, 2019, 41(1):79-94.DOI:10.1007/s13258-018-0742-9.
[41]	XIE T, CHEN C J, LI C H, et al. Genome-wide investigation of WRKY gene family in pineapple:evolution and expression profiles during development and stress[J]. BMC Genom, 2018, 19(1):490.DOI:10.1186/s12864-018-4880-x.
[42]	DONG J X, CHEN C H, CHEN Z X. Expression profiles of the Arabidopsis WRKY gene superfamily during plant defense response[J]. Plant Mol Biol, 2003, 51(1):21-37.DOI:10.1023/a:1020780022549.
[43]	CHRISTIAN A, YUE R, LIU Q, et al. The WRKY gene family in rice (Oryza sativa)[J]. J Integr Plant Biol, 2007(6):827-842.DOI:10.1111/j.1744-7909.2007.00504.x.
[44]	YANG B, JIANG Y Q, RAHMAN M H, et al. Identification and expression analysis of WRKY transcription factor genes in canola (Brassica napus L.) in response to fungal pathogens and hormone treatments[J]. BMC Plant Biol, 2009, 9:68.DOI:10.1186/1471-2229-9-68.
[45]	TIIKA R J, WEI J, MA R, et al. Identification and expression analysis of the WRKY gene family during different developmental stages in Lycium ruthenicum Murr.fruit[J]. Peer J, 2020, 8:e10207.DOI:10.7717/peerj.10207.
[46]	WANG D, CHEN Q Y, CHEN W W, et al. A WRKY transcription factor,EjWRKY17,from Eriobotrya japonica enhances drought tolerance in transgenic Arabidopsis[J]. Int J Mol Sci, 2021, 22(11):5593.DOI:10.3390/ijms22115593.
[47]	DANG F F, LIN J H, CHEN Y P, et al. A feedback loop between CaWRKY41 and H₂O₂ coordinates the response to Ralstonia solanacearum and excess cadmium in pepper[J]. J Exp Bot, 2019, 70(5):1581-1595.DOI:10.1093/jxb/erz006.
[48]	CAI Z D, XIAN P Q, WANG H, et al. Transcription factor GmWRKY142 confers cadmium resistance by up-regulating the cadmium tolerance 1-like genes[J]. Front Plant Sci, 2020, 11:724.DOI:10.3389/fpls.2020.00724.

基因名 gene ID	序列 sequence	用途 application
SpWRKY42-RT-F	GTTGCCGCAAATGTTTAGCC	伴矿景天筛选基因 qRT-PCR引物与内参引物
SpWRKY42-RT-R	ATAGCTGCTGTGAAATGCGG
SpWRKY72-RT-F	AGGGTTCTTGTTCGGACTGA
SpWRKY72-RT-R	AGCGCTTCTCTGAACCTTCT
SpWRKY40-RT-F	AGTGTAGCTGCTGAATCCGA
SpWRKY40-RT-R	AAAAGTCCAACTCCGGCCTA
SpWRKY46-RT-F	GACAACCGGCACTTTTGACT
SpWRKY46-RT-R	TAGGCTGTGTTGGAGTTGGT
SpWRKY39-RT-F	AGTTTACCGGCGAAGATGGA
SpWRKY39-RT-R	TCCAAATCCGACGACTCTGT
SpWRKY69-RT-F	CCAAACCATAAAGCCACCGA
SpWRKY69-RT-R	TCGCCGGAACTCTTACAACT
SpWRKY29-RT-F	TCACGGCGGAGATATCGAAA
SpWRKY29-RT-R	AAGAGCATCGATACGTCCGT
UBC9-F	TGGCGTCGAAAAGGATTCTGA
UBC9-R	CCTTCGGTGGCTTGAATGGATA
SpWRKY69-F	ATGGCCGTCGACCTCGT	SpWRKY69全长基因克隆引物与拟南芥内参引物
SpWRKY69-R	TCACGACGACTCTAGAATGAGT
actin-F	GCACCCTGTTCTTCTTACCG
actin-R	AACCCTCGTAGATTGGCACA

伴矿景天WRKY基因家族鉴定及镉胁迫响应分析

Comprehensive analysis of WRKY gene family in Sedum plumbizincicola responding to cadmium stress

RichHTML

PDF

可视化

摘要/Abstract

引用本文

使用本文

图/表 10

参考文献 48

相关文章 15

编辑推荐

Metrics

本文评价

作者

读者

审稿

[1]	却枫, 刘庆楠, 查若飞, 魏强. 孝顺竹中笋箨衰老相关WRKY转录因子的鉴定与分析[J]. 南京林业大学学报（自然科学版）, 2023, 47(6): 113-123.
[2]	黄碧芸, 卓仁英, 乔桂荣. 毛竹不同类型愈伤组织比较分析[J]. 南京林业大学学报（自然科学版）, 2023, 47(6): 141-149.
[3]	张婷, 柏杨, 亓希武, 于盱, 房海灵, 李莉, 刘冬梅, 梁呈元, 李维林. 薄荷MhWRKY57基因克隆及响应茉莉酸信号的表达分析[J]. 南京林业大学学报（自然科学版）, 2022, 46(6): 279-287.
[4]	王占军, 吴子琦, 王朝霞, 欧祖兰, 李杰, 蔡倩文, 徐忠东, 张照亮. 3个茶树品种WOX基因家族的进化及密码子偏好性比较[J]. 南京林业大学学报（自然科学版）, 2022, 46(2): 71-80.
[5]	纳晓莹, 刘刚, 刘桂丰, 王秀伟. 4种基因表达量和光合参数差异对白桦无性系幼苗生长的影响[J]. 南京林业大学学报（自然科学版）, 2022, 46(2): 88-94.
[6]	林莉莉, 胡安琪, 陈钢, 张霁月, 曹光球, 曹世江. 杉木ClWRKY44基因克隆及其表达特性分析[J]. 南京林业大学学报（自然科学版）, 2022, 46(1): 203-209.
[7]	王培龙, 杨妮, 张傲然, 唐努尔·塞力克, 李爽, 高彩球. 刚毛柽柳ThPCS1基因克隆与镉胁迫应答分析[J]. 南京林业大学学报（自然科学版）, 2021, 45(3): 71-78.
[8]	黎梦娟, 朱礼明, 霍俊男, 张景波, 施季森, 成铁龙. 唐古特白刺NtCBL1、NtCBL2基因克隆及表达分析[J]. 南京林业大学学报（自然科学版）, 2021, 45(3): 93-99.
[9]	王敏, 席东, 莫正海, 陈于, 赵玉强, 朱灿灿. 薄壳山核桃CiAGL6基因的克隆、亚细胞定位及表达[J]. 南京林业大学学报（自然科学版）, 2020, 44(4): 63-69.
[10]	田雪瑶, 周洁, 王保松, 何开跃, 何旭东. 柳树NAC基因的克隆与表达模式分析[J]. 南京林业大学学报（自然科学版）, 2020, 44(1): 119-124.
[11]	张庆, 魏树和, 代惠萍, 贾根良. 硒对茶树镉毒害的缓解作用研究[J]. 南京林业大学学报（自然科学版）, 2020, 44(1): 200-204.
[12]	王影,邱文敏,李鹤,贺雪莲,刘明英,韩小娇,曲同宝,卓仁英. 东南景天SaWRKY7基因对镉胁迫的响应研究[J]. 南京林业大学学报（自然科学版）, 2019, 62(03): 59-66.
[13]	陈圆,徐传红,韩建刚. Cd胁迫对湿地沉积物反硝化与氨化相对重要性的影响[J]. 南京林业大学学报（自然科学版）, 2019, 62(02): 64-72.
[14]	张春红,熊振豪,吴文龙,李维林. 黑莓果实发育成熟期木质素合成CAD酶及其基因表达[J]. 南京林业大学学报（自然科学版）, 2019, 62(01): 141-148.
[15]	吴凡,吴小芹. 细胞自噬对松材线虫繁殖存活的影响及线虫与松树互作早期自噬相关基因的表达[J]. 南京林业大学学报（自然科学版）, 2018, 61(05): 59-64.