马尾松PmAOX基因克隆与不同逆境胁迫表达分析

doi:10.3969/j.issn.1000-2006.201911058

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） ›› 2020, Vol. 44 ›› Issue (4): 70-78.doi: 10.3969/j.issn.1000-2006.201911058

马尾松PmAOX基因克隆与不同逆境胁迫表达分析

孙晓波(), 陈佩珍, 吴晓刚, 吴帆, 季孔庶()

南京林业大学林木遗传与生物技术省部共建教育部重点实验室，南方现代林业协同创新中心, 江苏　南京　210037

收稿日期:2019-11-28 修回日期:2020-02-16 出版日期:2020-07-22 发布日期:2020-08-13
通讯作者: 季孔庶
作者简介:孙晓波（sun15336229768@163.com）
基金资助:
国家重点研发计划(2017YFD0600304);江苏高校优势学科建设工程资助项目

The cloning and expression analysis of PmAOX gene from Pinus massoniana under different stress

SUN Xiaobo(), CHEN Peizhen, WU Xiaogang, WU Fan, JI Kongshu()

Co -Innovation Center for the Sustainable Forestry in Southern China，Key Laboratory of Forest Genetics and Biotechnology，Ministry of Education，Nanjing Forestry University，Nanjing 210037，China

Received:2019-11-28 Revised:2020-02-16 Online:2020-07-22 Published:2020-08-13
Contact: JI Kongshu

摘要/Abstract

摘要： 目的

揭示马尾松（Pinus massoniana）抗氰呼吸途径的交替氧化酶（alternative oxidase，AOX）基因的功能。

方法

提取15年生马尾松不同组织RNA，运用RT-PCR及RACE技术克隆基因全长。以基因组DNA为模板,利用染色体步移法（chromosome walking，CK）克隆目的基因的启动子区域。通过qRT-PCR技术分析目的基因在不同组织及逆境胁迫下的表达。

结果

克隆获得基因全长为1 610 bp，开放阅读框（open reading finder, ORF）1 221 bp, 编码406个氨基酸，推测氨基酸含有与拟南芥AtAOX基因相同的双铁羧酸结构LET、 NERMHL、LEEA和RADE_H，将其命名为PmAOX。实时荧光定量PCR显示，PmAOX在马尾松花中表达量最高，根中最低；脱落酸（abscisic acid，ABA）（100 μmol/L）胁迫后表达量逐渐下降，持续胁迫至24 h时，表达量增加至最大；经高温（42 ℃）、H₂O₂、NaCl（200 mol/L）、CO₂（0.08% ~ 0.10%）、低温（4 ℃）及干旱（10% PEG₆₀₀₀）处理后，表达量均呈现先升高后降低的趋势；高温42 ℃ 处理至3 h时表达量最高；H₂O₂处理4 h时，表达量最高； NaCl胁迫至6 h时，表达量最高；CO₂、低温及干旱处理至12 h时，表达量最高。克隆得到PmAOX 起始密码子上游1 081bp的启动子区域，经PlantCARE分析显示PmAOX启动子区域具有CAAT-box 及TATA-box 的基本顺式作用元件和低温、干旱等多个胁迫诱导元件，同时还包括脱落酸、茉莉酸甲酯、赤霉素、水杨酸在内的激素调控元件和光及昼夜节律响应元件。

结论

PmAOX 可能与马尾松抗逆性相关，并受激素的诱导和调控。

关键词: 马尾松, PmAOX, 基因克隆, 逆境胁迫, 荧光定量, 启动子

Abstract: Objective

The objective was to understand the function of the cyanide-resistant alternate oxidase (AOX) gene on the cyanide-resistant respiration pathway in Pinus massoniana.

Method

RNA was extracted from 15-year-old P. massoniana, and the full length gene was cloned using RT-PCR and RACE. Using genomic DNA as a template, the PmAOX promoter region was cloned using chromosome walking (CK). The expression of target genes in different tissues under different kinds of stress was analyzed using qRT-PCR.

Result

A full-length 1 610 bp gene was cloned to obtain a full-length open reading frame (ORF) of 1 221 bp encoding 406 amino acids. It was speculated that the amino acid sequence included the same ferrous carboxylic acid structure LET as the ArabidopsisAtAOX gene, NERMHL, LEEA, and RADE_H, and it was named PmAOX. The real-time fluorescence quantitative analysis showed that PmAOX expression was the highest in P. massoniana flowers, while it was the lowest in roots. ABA expression (100 μmol/L) decreased under ABA (100 μmol/L) stress and peaked 24 h after ABA (100 μmol/L) stress. After high temperature (42 °C), H₂O₂, NaCl (200 mmol/L), CO₂ (0.08% to 0.1%), low temperature (4 °C), and drought (10% PEG₆₀₀₀) stress, PmAOX expression increased before subsequently decreasing. The highest expression level was as follows: high temperature of 42 °C treatment at 3 h; H₂O₂ treatment at 4 h; NaCl stress at 6 h; CO₂, low temperature, and drought treatment at 12 h. A 1 081 bp promoter region upstream of the PmAOX start codon was cloned. Based on the PlantCARE analysis, we found that the PmAOX promoter region has basic cis-acting elements, including a CAAT-box, TATA-box, and multiple stress-inducing elements, such as low temperature and drought. It also includes hormonal regulatory elements, including abscisic acid (ABA), methyl jasmonate (MeJA), gibberellin (GA) and salicylic acid (SA), as well as light and circadian rhythm response elements.

Conclusion

PmAOX may be related to the resistance of P. massoniana, and it is induced and regulated by hormones.

Key words: Pinus massoniana, PmAOX, gene cloning, adversity stress, fluorescent quantification, promoter

中图分类号:

S722

孙晓波,陈佩珍,吴晓刚,等. 马尾松PmAOX基因克隆与不同逆境胁迫表达分析[J]. 南京林业大学学报（自然科学版）, 2020, 44(4): 70-78.

SUN Xiaobo, CHEN Peizhen, WU Xiaogang, WU Fan, JI Kongshu. The cloning and expression analysis of PmAOX gene from Pinus massoniana under different stress[J].Journal of Nanjing Forestry University (Natural Science Edition), 2020, 44(4): 70-78.DOI: 10.3969/j.issn.1000-2006.201911058.

图/表 4

图1

PmAOX 序列分析B图黑框勾选为编码序列的保守结构域，蓝框勾选为AHR芳香烃配合体；C图表示马尾松PmAOX 蛋白的三级结构；D图表示马尾松蛋白序列同其他物种聚类分析. Panel B. The black box is checked for the conserved domain of the coding sequence, and the blue box is checked for the AHR aromatic hydrocarbon complex. TKY0148.1. 密花豆 Spatholobus suberectus；XP_022028949.1. 向日葵Helianthus annuus；NP_001236166.2. 大豆Glycine max；XP_020886 713.1. 拟南芥Arabidopsis thaliana；PIN07923.1. 粉红钟花 Handroanthus impetiginosus；XP_018491746.1. 萝卜Raphanus sativus；ABK23336.1. 西加云杉Picea sitchensis；PmAOX. 马尾松Pinus massoniana. Panel C shows the tertiary structure of Pinus massoniana PmAOX protein. Panel D shows the cluster analysis of Pinus massoniana sequence with other species. AGZ61935.1. Nicotiana benthamiana; XP009804831.1. Nicotiana sylvestris; XP009618159.1. Nicotiana tomentosiformis; XP019262649.1. Nicotiana attenuate; XP027085311.1. Coffea Arabica; XP022028949.1. Helianthus annuus; GQ380559.1. Pinus pinea; PmAOX: P.massoniana; PON47568.1. Parasponia andersonii; AQM74365.1. Populus pruinosa; CAB64356.1. Populus tremula × Populus tremuloides; XP015897333.1. Ziziphus jujube; XP030946413.1. Quercus lobate; AAZ09196.1. Vigna unguiculata; XP009343192.1. Pyrus × bretschneideri; XP002317754.2. Populus trichocarpa; XP011001873.1. Populus euphratica; XP009145388.1. Brassica rapa; XP013647907.1. Brassica napus; VDD45298.1. Brassica oleracea; XP018491746.1. Raphanus sativus; AB003176.1. Arabidopsis thaliana; XP020886713.1. Arabidopsis lyrata subsp; XP007158132. Phaseolus vulgaris; XP004512369.1. Cicer arietinum; AGQ42774.1. Medicago sativa; XP027360032.1. Abrus precatorius; KHN42869.1. Glycine soja; NP001236166.2. Glycine max; XP025651389.1. Arachis hypogaea; KYP46143.1. Cajanus cajan; XP009343192.1. Pyrus × bretschneideri."

图1

图2

图3

表1

参考文献 38

1	徐飞, 袁澍, 梁厚果, 等. 交替氧化酶和解偶联蛋白在植物线粒体中的作用及其相互关系[J]. 植物生理学报, 2009, 45(2):105-110.
	XU F, YUAN W, LIANG H G, et al. The role of alternating oxidase and uncoupling proteins in plant mitochondria and their relationship[J]. Chinese Journal of Plant Physiology, 2009, 45(2): 105-110. DOI: 10.13592/j.cnki.ppj.2009.02.001.
2	吴强, 李红玉, 张立新, 等. 环境胁迫与植物抗氰呼吸[J]. 西北植物学报, 2003, 23(1):164-170.
	WU Q, LI H Y, ZHANG L X, et al. Environmental stress and plant cyanide⁃resistant respiration[J]. Journal of Northwest Botanical Sinica, 2003, 23(1): 164-170. DOI: CNKI:SUN:DNYX.0.2003-01-032.
3	VANLERBERGHE G C, MCINTOSH L, oxidase alternative: From gene to function[J]. Annu Rev Plant Physiol Plant Mol Biol, 1997， 48(48): 703-734. DOI: 10.1146/annurev.arplant. 48.1.703.
4	MILLENAAR, F.F, H. LAMBERS, The alternative oxidase: in vivo regulation and function[J]. Plant Biology, 2010， 5(1):2-15. DOI: 10. 1055/s-2003-37974.
5	CONSIDINE M, HOLTZAPFFEL R C, DAY D A, et al. Molecular distinction between alternative oxidase from monocots and dicots[J]. Plant Physiology, 2002， 129(3): 949-952. DOI: 10.1104/pp.004150.
6	NI Z, ZHOU P Y, XU M, et al. Development and characterization of chloroplast microsatellite markers for Pinus massoniana and their application in Pinus (Pinaceae) species[J]. Journal of Genetics, 2018, 97(2):1-7. DOI: 10.1007/s12041-018-0931-y.
7	XU C, WU X Q. Physiological and proteomic analysis of mycorrhizal Pinus massoniana inoculated with Lactarius insulsusunder drought stress[J]. Russian Journal of Plant Physiology, 2016, 63(5):709-717. DOI: 10.1134/S1021443716040178.
8	NI Z X, YE Y J, BAI T, XU M, et al. Complete chloroplast genome of Pinus massoniana (Pinaceae): Gene rearrangements, loss of ndh genes and short inverted repeats contraction expansion[J]. Molecules, 2017, 22(1):1528. DOI: 10.3390/molecules22091528.
9	季孔庶, 王章荣, 陈天华, 等. 马尾松插穗内源抑制物质的研究[J]. 林业科学, 1997, 33(2):142-151.
	JI K S, WANG Z R, CHEN T H, et al. Studies on endogenous inhibitory substances in Pinus massoniana cuttings[J]. Forestry Science, 1997, 33(2): 142-151.
10	季孔庶,王章荣. 马尾松插穗生根能力变异的研究[J]. 南京林业大学学报, 1998, 22（3）:66-70.
	JI K S, WANG Z R. Study on variation of rooting ability of Pinus massoniana cuttings[J]. Journal of Nanjing Forestry University, 1998, 22（3）:66-70. DOI: 10.3969/j.jssn.1000-2006.1998.03.015.
11	俞新妥, 卢建煌. 不同种源马尾松光合能力的比较研究[J]. 福建林学院学报, 1991,11(2):3-7.
	YU X T, LU J H. A comparative study on the photosynthetic capacity of Pinus massoniana from different provenances[J]. Journal of Fujian Forestry College, 1991,11(2): 3-7. DOI：10.13324/j.cnki.jfcf.1991.02.001.
12	阮维程，潘婷，季孔庶. 马尾松纤维素合成酶基因PmCesA1的克隆及其分析[J]. 分子植物育种, 2015,13(4): 861-870.
	RUAN W C,PAN T,JI K S. Cloning and analysis of cellulose synthase gene PmCesA1 from Masson pine[J]. Molecular Plant Breeding, 2015,13(4): 861-870. DOI: CNKI:SUN:FZZW.0.2015-04-035.
13	王猛. 马尾松抑制消减文库的构建及抗病性相关基因的克隆[D]. 长沙：中南林业科技大学, 2010.
	WANG M. Construction of a subtractive library of Pinus massoniana and cloning of disease resistance related genes [D]. Changsha: Central South University of Forestry and Technology, 2010.
14	蔡琼，丁贵杰，文晓鹏. 马尾松水通道蛋白PmPIP1基因克隆及在干旱胁迫下的表达分析[J]. 浙江农林大学学报, 2016,33(2): 191-200.
	CAI Q, DING G J, WEN X P. Cloning of aquaporin PmPIP1 gene from Pinus massoniana and its expression under drought stress[J]. Journal of Zhejiang Agriculture and Forestry University, 2016,33(2): 191-200. DOI: 10.11833/j.issn.2095-0756.2016.02.002.
15	王晓锋, 何卫龙, 蔡卫佳,等. 马尾松转录组测序和分析[J]. 分子植物育种, 2013, 11(3):385-392.
	WANG X F, HE W L, CAI W J, et al. Sequencing and analysis of transcriptome of Masson pine[J]. Molecular Plant Breeding, 2013, 11(3):385-392. DOI: CNKI:SUN:FZZW.0.2013-03-019.
16	MCDONALD A, VANLERBERGHEG. Branched mitochondrial electron transport in the Animalia: presence of alternative oxidase in several animal phyla[J]. Iubmb Life, 2004, 56(6): 333-341. DOI: 10.1080/1521-6540400000876.
17	NEIMANIS, KARINA, STAPLES, et al. Identification, expression and taxonomic distribution of alternative oxidases in non⁃angiosperm plants[J]. Gene, 2013, 26(6): 275-286. DOI: 10.1016/j.gene.2013.04.072.
18	CLIFTON R, LISTER R, PARKER K L, et al. Stress⁃induced co⁃expression of alternative respiratory chain components in Arabidopsis thaliana[J]. Plant Molecular Biology, 2005, 58(2):193-194. DOI: 10.1007/s11103-005-5514-7.
19	THIRKETTLE WATTS D, MCCABE T C, CLIFTON R, et al. Analysis of the alternative oxidase promoters from soybean[J]. Plant Physiology, 2003, 133(3):1158-1161. DOI: 10.1104/pp.103.028183.
20	晏婴才, 林宏辉, 梁厚果, 等. 不同低温胁迫对烟草愈伤组织抗氰交替途径诱导和交替氧化酶表达影响的比较[J]. 植物学报, 2004, 21(3):296-305.
	YAN Y C, LIN H H, LIANG H G, et al. Comparison of the effects of different low temperature stress on the induction of cyanide⁃resistant pathway and the expression of alternate oxidase in tobacco callus[J]. Chinese Journal of Plant Science, 2004, 21(3):296-305. DOI: CNKI:SUN:ZWXT.0.2004-03-006.
21	冯汉青, 马军, 李红玉,等. 黄化水稻幼苗转绿期AOX1基因家族的表达与功能分析[J]. 分子植物(英文版), 2006, 32(3):300-306.
	FENG H Q, MA J, LI H Y, et al. Expression and functional analysis of AOX1 gene family in yellowing rice seedlings during greening[J]. Molecular Plants, 2006, 32(3): 300-306. DOI: CNKI:SUN:ZWSI.0.2006-03-006.
22	WANG J, WANG X, LIU C, et al. The NgAOX1a gene cloned from Nicotianaglutinosa is implicated in the response to abiotic and biotic stresses[J]. Bioscience Reports, 2008, 28(5):259-266. DOI: 10.1042/BSR20080025.
23	SIEDOW J N, UMBACH A L. Plant mitochondrial electron transfer and molecular biology[J]. Plant Cell, 1995， 7(7): 821-827. DOI: 10.1105/tpc.7.7.821.
24	李严曼, 朱磊, 杨景华, 等. 西瓜交替氧化酶AOX2基因的克隆与分析[J]. 果树学报, 2011(5):171-176.
	LI Y M, ZHU L, YANG J H, et al. Cloning and analysis of watermelon oxidase AOX2 gene[J]. Journal of Fruit Science, 2011(5):171-176. DOI: CNKI:SUN:GSKK.0.2011-05-031.
25	CONSIDINE M J, DALEY D O, WHELAN J. The expression of alternative oxidase and uncoupling protein during fruit ripening in mango[J]. Plant Physiology, 2001, 126(5):1619-1629. DOI: 10.1104/pp.126.4.1619.
26	BARTOLI C G, GOMEZ F, GERGOFF G, et al. Up⁃regulation of the mitochondrial alternative oxidase pathway enhances photosynthetic electron transport under drought conditions[J]. Journal of Experimental Botany, 2005, 56(415):1269-1276. DOI: 10.1093/jxb/eri111.
27	VALYA V, LYUDMILA S S, KLIMENTINA D, et al. Variety⁃specific response of wheat (Triticum aestivum L.) leaf mitochondria to drought stress[J]. Journal of Plant Research, 2009, 122(4):445-454. DOI: 10.1007/s10265-009-0225-9.
28	WEN J, LIANG H. Effects of KCN and NaN₃ pretreatment on the cyanide⁃resistant respiration in tobacco callus[J]. Acta Bontanica Sinica, 1995, 37(9):711-717.
29	BEGCY K, MARIANO E D, MATTIELLO L, et al. An arabidopsis mitochondrial uncoupling protein confers tolerance to drought and salt stress in transgenic tobacco plants[J]. Plos One, 2011, 6(8):23776-23784. DOI: 10.1371/journal.pone.0023776.
30	李春荣. 水稻AOX1基因家族成员逆境下的表达分析及功能鉴定[D]. 合肥：安徽大学, 2013. LI C R. Expression analysis and functional identification of rice AOX1 gene family members under adverse conditions [D]. Hefei: Anhui University, 2013.
31	YAN Y C, LIN H H, LIANG H G, et al. Comparison of the effects of difterent low temperature stresses on the induction of the cyanide⁃resistant alternative pathway and the expression of alternative oxidase in tobacco callus[J].Chin Bull Bot, 2004, 21(3):296-305.
32	MOHANAPRIYA G, BHARADWAJ R, NOCEDA C, et al. Alternative oxidase (AOX) senses stress levels to coordinate auxin⁃induced reprogramming from seed germination to somatic embryogenesis-a role relevant for seed vigor prediction and plant robustness[J]. Frontiers in Plant Science, 2019, 12(2):1131-1134. DOI: 10.3389/fpls.2019.01134.
33	HILAL M. Saline stress alters the temporal patterns of xylem differentiation and alternative oxidase expression in developing soybean roots[J]. Plant Physiology, 1998, 117(3):695-701. DOI: 10.1104/pp.117.2.695.
34	ANNICCHIARICO P, NAZZICARI N, PECETTI L, et al. Pea genomic selection for Italian environments[J]. BMC Genomics, 2019(4):599-603. DOI: 10.1186/s12864-019-5920-x.
35	ANDJELKOVIĆ A, OLIVEIRA M T, CANNINO G, et al. Diiron centre mutations in Ciona intestinalis alternative oxidase abolish enzymatic activity and prevent rescue of cytochrome oxidase deficiency in flies[J]. Scientific Reports, 2015, 5(3):295-301. DOI: 10.1038/srep18295.
36	PURVIS A C, SHEWFELT R L. Does the alternative pathway ameliorate chilling injury in sensitive plant tissues?[J]. Physiologia Plantarum, 1993, 88(4):712-718. DOI: 10.1111/j.1399-3054.1993.tb01393.x.
37	SZABADOS L, CHARRIER B, KONDOROSI A, et al. New plant promoter and enhancer testing vectors[J]. Mol Breed, 1995, 1(4): 419–423. DOI: 10.1007/BF01248419.
38	李会宣, 董向峰, 高健. 毕赤酵母醇氧化酶基因AOX1启动子研究进展[J]. 生物技术, 2013, 1(4):87-91.
	LI H X, DONG X F, GAO J. Advances in the promoter of pichia pastoris alcohol oxidase gene AOX1[J]. Biotechnology, 2013, 1(4): 87-91. DOI: CNKI:SUN:SWJS.0.2013-04-021.

基序类别 motif type	元件名称 name of element	位置 position	元件序列 sequence	预测功能注释 putative function annotation
基础元件 basic component	CAAT-box	-159,-215.-383,-439,-466,-522,+742,+802, +854,+917	CAAAT/CAAT	启动子、增强子区域常见的顺式作用元件; 转录起始位点-30 核心启动元件
	CCAAT-box	+756	CCAAT
	TATA-box	+44,+84,+141,+170,+227,+236,+287,+322,+334,+339,+357,+392,+475,+561,-920,-926,-994 +43,+83,+235,+333,+356,+391,+474 -139,-992 -169,-226,-321,-560,-925 -179 +206,+305	TATA/ ATATAA/ TATACA/TATAA/TACATAAA/TACAAAA
胁迫 coercion	LTR	+846	CCGAAA	低温响应元件
胁迫 coercion	MBS	+622,+879	CAACTG	干旱响应元件
光响应 light response	Box4	+220	ATTAAT	光响应保守区
	I-Box	+658	AGATAAGG	光响应元件
	chs-CMA2a	-1050	TCACTTGA	光响应元件
激素 hormone	CGTCA-motif	+750	CGTCA	茉莉酸甲酯响应元件
激素 hormone	TGACG-motif	-750	TGACG	茉莉酸甲酯响应元件
其他 other	CAT-box	+792	GCCACT	分生组织表达顺式调控元件
其他 other	CCAAT-box	-756	CAACGG	MYBHv1结合位点

[1]	武燕, 黄青, 刘讯, 郑睿, 岑佳宝, 丁波, 张运林, 符裕红. 西南喀斯特地区马尾松人工林林龄对土壤理化性质的影响[J]. 南京林业大学学报（自然科学版）, 2024, 48(3): 99-107.
[2]	范明阳, 胡萌, 杨园, 方炎明. 中国东部地区马尾松与黄山松群落分类及群落结构和物种多样性特征[J]. 南京林业大学学报（自然科学版）, 2024, 48(1): 47-58.
[3]	王章荣, 季孔庶, 徐立安, 邹秉章, 林能庆, 林景泉. 马尾松实生种子园营建技术、现实增益及多世代低成本经营新模式探讨[J]. 南京林业大学学报（自然科学版）, 2023, 47(6): 9-16.
[4]	王宇, 易艳灵, 刘海, 文晓晨, 李天一, 尹海锋, 李贤伟, 范川. 两种采伐方式对马尾松人工林林分空间结构的影响[J]. 南京林业大学学报（自然科学版）, 2023, 47(5): 138-146.
[5]	刘增才, 王淑婷, 佟鑫宇, 邹莉. 暴马桑黄GPS基因克隆及响应茉莉酸甲酯诱导表达研究[J]. 南京林业大学学报（自然科学版）, 2023, 47(4): 123-130.
[6]	魏祯祯, 宋程威, 郭丽丽, 郭琪, 侯小改. ‘凤丹’PoERF4基因的克隆及表达分析[J]. 南京林业大学学报（自然科学版）, 2023, 47(3): 56-62.
[7]	竹磊, 徐军亮, 章异平, 罗鹏飞, 师志强, 候佳玉, 翟乐鑫. 河南洛阳马尾松树干液流昼夜变化特征及其影响因子分析[J]. 南京林业大学学报（自然科学版）, 2023, 47(1): 92-100.
[8]	孙薇, 王斌, 楚秀丽, 王秀花, 张东北, 吴小林, 周志春. 马尾松容器苗生长和养分性状对磷添加和接种菌根菌的响应及关联[J]. 南京林业大学学报（自然科学版）, 2023, 47(1): 226-233.
[9]	季孔庶, 徐立安, 王登宝, 倪州献, 王章荣. 中国马尾松遗传改良研究历程与成就[J]. 南京林业大学学报（自然科学版）, 2022, 46(6): 10-22.
[10]	贾展慧, 贾晓东, 许梦洋, 莫正海, 翟敏, 宣继萍, 张计育, 王刚, 王涛, 郭忠仁. 薄壳山核桃原花青素合成关键酶基因的克隆与表达分析[J]. 南京林业大学学报（自然科学版）, 2022, 46(5): 49-57.
[11]	王立超, 苏胜荣, 陈凤毛, 董晓燕, 田成连, 王洋. 黄山马尾松林天牛及携带线虫种类初步调查[J]. 南京林业大学学报（自然科学版）, 2022, 46(4): 29-35.
[12]	吴佳雯, 尹艳楠, 谈家金, 郝德君. 蜡样芽孢杆菌NJSZ-13菌株诱导马尾松抗松材线虫病研究[J]. 南京林业大学学报（自然科学版）, 2022, 46(4): 53-58.
[13]	胡兴峰, 吴帆, 孙晓波, 陈厚平, 殷安政, 季孔庶. 38年生马尾松种源生长及材性联合分析[J]. 南京林业大学学报（自然科学版）, 2022, 46(3): 203-212.
[14]	吴帆, 朱沛煌, 季孔庶. 马尾松分布格局对未来气候变化的响应[J]. 南京林业大学学报（自然科学版）, 2022, 46(2): 196-204.
[15]	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.

马尾松PmAOX基因克隆与不同逆境胁迫表达分析

The cloning and expression analysis of PmAOX gene from Pinus massoniana under different stress

RichHTML

PDF

可视化

摘要/Abstract

引用本文

使用本文

图/表 4

参考文献 38

相关文章 15

编辑推荐

Metrics

本文评价

作者

读者

审稿