摘要
植物内生菌(Endophyte)是一类应用前景广阔的资源微生物,具有固氮、促进植物生长、增强宿主植物抗逆性等作用,可通过组织学方法或表面消毒等方法从植物组织中分离。重楼属植物是一类极具药用价值的植物,具有解毒、镇痛、抗菌、抗癌等功效,是许多中成药和新药的有效成分。随着医药产业对重楼的开发利用,野生重楼遭到了毁灭性的人为破坏,濒临枯竭。药用植物内生菌的研究表明,其内生菌具有合成与宿主植物相同或相似的活性成分的能力,有的内生菌还具有促进宿主植物合成活性成分的能力。本文选择对四川华重楼内生菌的研究,从分离药用植物的活性成分方面解决重楼资源不足的问题。
通过表面灭菌,从药用植物华重楼(Paris polyphylla var.chinensis )的地下根状茎中分离得到82株内生菌。通过滤纸片扩散法初筛和平板对峙法复筛,有多株菌有抑菌活性,本文对其中一株具有较好抑制作用的内生细菌PCE45进行了研究。根据形态学特征、生理生化特征及16SrDNA序列分析,确定PCE45为一株枯草芽孢杆菌(Bacillus subtilis)。
通过硫酸铵盐析、透析、丙酮低温沉淀,得到PCE45的粗提液,采用管碟法测得硫酸铵最佳饱和度为50%~70%。粗提液经SephadexG75柱、DE52纤维素柱和SephadexG25凝胶柱纯化得到一种抗菌肽,命名为PCP-1。稳定性测试表明,该抗菌肽对对蛋白酶不敏感,对高温、强酸、强碱有较好的耐受性,可造成稻瘟病菌菌丝畸形并抑制孢子萌发。抑菌谱表明该抗菌肽对玉米弯胞病菌等真菌和大肠杆菌等细菌有较强的抑菌效果。质谱测得其分子量为1058.3D。氨基酸组成分析表明该小肽主要由7种氨基酸组成。结合茚三酮反应呈阴性,酸水解后,茚三酮反应和双缩尿反应呈阳性的性质,推测PCP-1可能为低分子量的环状小肽。
Plant endophyte is a group of resource microorganism with broud perspective, which have ability of nitrogen fixation, could encourage plant growth and reinforce antireversion force of the plant . It can be separated from the inner part of plant by histology or disinfection. Paris L is a class of plant with variety of medicine value ,which have ability of detoxication, analgesia, antibiosis, anticancer et al. It is the major material of many prepared Chinese medicines. However, with the exploitation of medicine industrial, we are facing the crisis of it’s gradually drying up.
Base on the research results of medicine plants, it’s endophyte can synthetize the same active ingredient, some endophyte can encourage plant to synthetize active ingredient.We are study on endophyte in Paris polyphylla var.chinensis Hara which could produce active ingredient in order to resolve the crisis of wild paridis lacking.
Through surface sterilization ,82 strains endophytes were isolated and purified from the fresh hypogeal stems of Paris polyphylla var.chinensis . Many endophytes showed antimicrobial activity through preliminary antimicrobial screening against pathogenic bacterias and further antimicrobial screening against pathogenic fungi. The strain PCE45 with best antimicrobial activity was selected to the next series of studies.Based on the characteristics of morphology,physiology,and biochemistry as well as 16SrDNA analysis, the strain PCE45was classified into Bacillus subtilis.
We made the crude extract of PCE45 after the steps of ammonium sulfate salting-out, dialysis and acetone precipitation. Through the method of Oxford Plate,we find the best sulfate saturation is 50%~70% .After SephadexG75,DE52 and SephadexG25 column chromatography, we separated an antimicrobial peptide PCP-1 from the strain PCE45.The stability against high temperature and proteinase, and antimicrobial activity were also analyzed. The antimicrobial peptide PCP-1 was stable to proteinase and tolerated high temperature, strong acid and strong base. PCP-1 caused deformation of the hyphae of Pyricularia oryzae and prohibited the spore germination. It also inhibited fungi such as Curvularia lunata and bacteria such as Escherichia coli. Mass spectrogram measurement revealed its molecular weight of 1058.3 Da. The amino acid composition of the peptide composed of 7 amino acids. Ninhydrin reaction showed negative trait whereas after acid hydrolysis with positive ninhydrin reaction and biuret reaction. The ninhydrin reaction and biuret reaction imply that the peptide PCP-1 is a cyclic lipeptide.
引文
[1]邹文欣,谭仁祥.植物内生菌的研究新进展.植物学报,2001,43(9):881-892
[2]Stierle A ,Strobel G,Stierle D. Taxol and taxane production by Taxomyces andreanae ,an endophytic fungus of pacific yew(taxus brevifolia).Science,1993,260:214-216
[3]王建锋,吕华鹰,苏文金.植物内生真菌产紫杉醇的研究.微生物学通报,2000,27(1):58-60
[4]张玲琪,王海昆,邵华,等.美登木内生真菌产抗癌物质球毛壳甲素的分离及鉴定.中国药学杂志,2002,37(3):172-175
[5]李海燕,王志军,张玲琪.一种桃儿七内生真菌的分离初报.云南大学学报(自然科学版),1999,21(3):243
[6]曾松荣,邵华,张玲琪.从南方山荷叶分离出一株产鬼臼毒素类似物的内生真菌.微生物学杂志,2004,24(4):1221
[7]Gurney KA,Mantle PG.Biosynthesis of 1-N-methylal-bonoursin by an endophytic Streptomyces sp. J Nat Prod,1993,56:1194-1198
[8]Strobel GA,Miller RV,Miller C,et al.Cryptocandin,a potent antimycotic from the endophytic fungus Cryptosporiopsis of quercina. Microbiology,1999,14:1919-1926
[9]Brady SF,Clardy J, A new pentaketide antifungalagent isolated from an endophytic fungus. J Nat Prod, 2000, 63:1 447-1448
[10]Siegel MR,Latch GC,Bush LP,et al.Fungal endophyte2infected grasses:Alkaloid accumulation and aphid response.J Chem Ecol,1990,16:3301-3316
[11]Calhoun LA,Findrlay JA,Miller JD,et al.Metabolites toxic tospruce budwormfrom balsam fir needle endophytes. Mycol Res,1992,96:281-286
[12]Harpre J K,Ford E J ,strobel g a ,et al. 3- dihydroisobenzofuran from Pestalotiopsis microspora possessing antioxidant and antimycotic activities. Tetrahedron,2003,59: 2471-2476
[13]Huo Juan,Chen Shuang lin.A preliminary study on antioxadative activity of enophytic fungus Chactomella sp.from Eucmmia Ulmoieds.Journal of Nanchang University(Natural Science),2004,28(3):270-275
[14]Dobereiner J,Baldani VLD,Reis VM. Endophytic occurrence of diazotropic bacteria in non-leguminous crops. In:Fendrik I,de Callo M,Vanderleyden J,eds. Azospirillum Viand Related Microorgnisms. Berlin:Springer2Verlag,1995. 3-14
[15]Hallmann J,Quadt-Hallmann A,Mahaffee W F,et al.Bacterial endophytes inagriculturalcrops. Can J.Microbiol,1997,43:895-914
[16]张集慧,王春兰,郭顺星,等.兰科药用植物5种内生真菌产生的植物激素.中国医学科学院学报,1999,21 (6):460-465
[17]Moy M,Li HM,Sullivan R,et al.Endophytic fungal beta - 1,6 -glucanase expression in the infected host grass.Plant Physiol,2002,Nov,130(3):1298-1308
[18]Luk’ianchuk VV,Reva ON,Polishchuk LV. Endophytic bacilli- producers of type II restriction endonucleases.Mikrobiologiia,2002,Jul– Aug,71(4):491-493
[19]Salituro GM,Pelaez F,Zhang B. Discovery of a small molecule insulin receptor activator. Recent Progress in Hormone Research,2001,56:107-126
[20]Zhang J,Wang C,Guo S,Chen J, et al.Studies on the plant hormones produced by 5 species of endophytic fungi isolated from medicinal plants. Zhongguo YiXue KeXueYuan XueBao,1999 Dec,21(6):460-465
[21]Lee J C,Lobkovsky E,Pliam N B,et al.Subglutinol A and B:immunosuppressive compounds from the endophytic funfus Fusarium subglutians. Org Chem,1995(60):7076-7077
[22]Strobel GA.Endophytes as sources of bioactive products.Microbes Infect,2003,5(6):535-544
[23]Baldani J I,Olivares FL,Hemerly,et al.Nitrogen 2 fixing endophytes:recent advances in the association with graminaceous plants grown in the tropics. In:Elemerich C,KordorosiA,Newton WE(eds).Biological Nitrogen Fixation for the 21st century. Netherlands:Kluwer Academic publishers,1997,203-206
[24]Doberieiner J,Day JM.Proceedings of the 1th international symposium on Nitrogen2fixation. 1974(2):518
[25]Baldani J,Carusol,Baldani VLD,et al.Recent advance in BNF with non2legume plants.Soil Biol Biochem,1997,29(5)911-922
[26]陈丽梅,樊妙姬,李玲.甘蔗固氮内生菌重氮营养醋杆菌的研究进展.微生物学通报, 2000,27 (1):63-66
[27]Cavalcante V,Dobereiner J.A new acid tolreant nitrogen fixing bacterium associated with sugar cane.Plant Soil,1988,108:23-31
[28]Boddey RM,Oliveira OC,Urquiaga S,et al.Biological nitrogen fixation associated with sugar cane and rice:contributions and prospects for improvement.PlantSoil,1995,174:195-209
[29]Clemence C,Eric G,Yves P,et al.Photosynthetic bradyrhizobia are natural endophytes of the African wild rice Oryza breviliguulata.Appl.Envi ron.Microbiol,2000(2):5437-5447
[30]Reinhold B,Hurek T,Niemann EG,et al..Close association of Azospirillum and diazotrophic rods with different root rones of Kallar grass.Appl.Environ.Microbiol,1986,52:520-526
[31]Lu H,Zou W X,Meng JC,et al.New bioactive metabolites produced by Colletot richum sp., an endophytic fungi in Artemisia annua.Plant Science,2000,151:67-73
[32]Hallmann J,Quadt-hallmann A,Mahaffee W F,et al.Bacterial endophytes in agricultural crops.Can J.Microbiol,1997,43:891-895
[33]Ravel F,Courty C,Coret A,Charmet G.Beneficial effects of Neotyphodium lolii on the growth and the water status in perennial ryegrass cultivated under nitrogen deficiency or drought stress. Agronomie,1997,17:173-181
[34]Malinowski DP,Belesky DP.Tall fescue aluminum torelance is affected by Neotyphodium coenophyialum endophyte. J Plant Nutr,1999,22:1335-1349
[35]Bacon CW,Siegel MR.Endophyte parasitism of tall fescue. J.Prod A gric, 1998 (1):45-55
[36]Hill NS,Belesky DP,Stringer WC.Competitiveness of tall fescue as influenced by Acremonium coenophial um.Crop Sci,1991(31):185-190
[37]姜广策,林永成,周世宁,等.中国南海红树内生真菌次级代谢物的研究.中山大学学报(自然科学版),2000,39(6):68-72
[38]Clay K,Cheplick GP,Wray SM.1989.Impact of the fungus Balansi n henni ngsisnn on the grass Panicum agrostoi des:frequency of infection ,plant growth and reproduction and resistance to pests.Oecologia,80:374-380
[39]Azevedo JL,Maccheroni JM,Pereira JO,et al.Edophytic microorganisms:a review on insect control and recent advances on tropical plants.J Biotech,2003,3(1):40-65
[40]Shi JingYing,Chen WeiXin,Ai Yuan.Advances in the study of endophytes and their effects on control of plant disease.Acta Ecologica Sinica,2006,26(7):2395-2401
[41]Yan MengHong,Cai ZhengQin,Han JiGang,et al.The applied research of endophytic bacteria in biological control of plant diease.Biotechnology Information,2004,(3):8-12
[42]Van Peer R,Niemann GJ. Schippers B.Induced resistance and phytoalexin accumulation in biological control of Fusarium wilt of carnation by Pseudomonas sp.strain WCS417. Phytopathology,1991,81:728-734
[43]Vanloon LC,Bakker Pahm,Pieterse CM J.Systemic resistance induced by rhizosphere bacteria.Annual Review of Phytopathology1998,36:453-483
[44]Sutherland BL, Hume DE,Tapper BA.Allelopathic effects of endophyte-infected perennial ryegrass extracts on white clover seedlings. New Zeal J Agr Res,1999,42:19-26
[45]Clay K, Holah J.Fungal endophyte symbiosis and plant diversity in successional fields. Science,1999,285:1742 -1745
[46]Wilson D,Carroll GC,Avoidance of high-endophyte space by gall-forming insects. Ecology,1997,78:2153-2163
[47]李恒.重楼植物.北京:北京科学出版社,1998.
[48]中国科学昆明植物所.云南植物志.北京:科学出版社,1997
[49]汤海峰,赵越平,蒋永培.重楼属植物的研究概况.中草药,1998 ,29 (12) :839-842
[50]何顺志.贵州省重楼属药用植物分类鉴定及地理分布的研究,中草药,1990,21(10):462
[51]王丽,李云飞,唐荣.18-26SrDNA在4种重楼属植物中的定位植物分类学报,2004,42(5):419-426
[52]唐荣华.毛丽.唐小为.十一种重楼属植物的RAPD分析.四川大学学报(自然科学版) ,2003,40(4):778-782
[53]张瑞,唐铭霞,翁周.重楼属植物AFLP实验体系的建立.四川大学学报(自然科学版), 2006,43(5):1105~1109
[54]张霄霖,刘月婵.重楼的研究与应用.中国中医药科技,2000,7 (5):346-347
[55]武珊珊,高文远,段宏泉,等.重楼化学成分和药理作用研究进展.中草药,2004,35(3):344-347
[56]Chen CX, Zhou J. Studies on chem ical constituents of P arispolyphy lla var. yunnanensis Acta Bot Yunnan,1981,3(1):89-93
[57]Chen CX, Zhou J.Two new steroid sapogenins of Paris polyphylla var. yunnanensis. Acta Bot Yunnan,1992,14(1):111-113
[58]Editorial Board of China Herbal,State A dministration of Traditional ChineseM edicine, China. China Herbal.Shanghai:Shanghai Scientific and Technical Publishers,1999
[59]Indresh K,Seshadri R,Seshadri T R.Constitution of pariphyllin-A and pariphyllin-B, the saponinsiso lated from the tubers of Paris polyphylla.Indian J Chem,1975,13(8):781
[60]Singh SB,Raghunath S. Furo stano l saponins from Paris polyphylla structures of polyphyllin G and H. Phy tochemistry,1982,21(8):2079
[61]Xu XM , Zhong ZC, Studies on chem ical constituents of Paris polyphylla var. ch inensis. Chin Tradit HerbDrugs,1988,19(6):242-249
[62]W ang Q,Yu XS.Determ ination of ecdysterone in 19 species of the resource p lants. China Pharm Univ, 1990,21 (3):229-231
[63]周安寰,黄永明,李勋.云木香及七叶一枝花中氨基酸的鉴定和含量测定.中草药,1984,15(11):l6-22
[64]陈昌祥,周俊,张玉童,等.滇产植物皂素成分的研究.云南植物研究,1983,5(2):219
[65]Chen CX, Zhang YT, Zhou J. The glyco sides of aerial parts of Paris polyphylla var. yunnanensis. Acta Bot Yunnan,1995,17(4):473-478
[66]王强,徐国钧,程永宝.中药七叶一枝花类的抑菌和止血作用的研究.中国药科大学学报,1989,20(4):251
[67]欧阳录明,黄晓敏.中草药体外抗白色念珠菌的实验研究.中国中医药信息杂志,2000,7(3):26
[68]成兴华.蚤休加抗痨药综台治疗淋巴结核溃疡30例.中国中西医结合杂志,1998,18(9):559
[69]舒哗.喉舒的抗炎作用.基层中药杂志,1997,11(4):15
[70]宋守荣,魏羽佳,曹煜,等.中药山宝皮宁治疗单纯疱疹临床及实验研究.贵阳医学院报.2004,29(3):226
[71]陈雪珍.综合治疗慢性盆腔炎80例.江苏中医,1999,20(5):22
[72]Zhang XL.Studies and application of Paris polyphy lla.Chin J Technol Tradit Med,2000,7 (5):346-347
[73]高学敏,肖艳萍.三七血伤宁胶囊的药理分析.首都医药,2004,(4):51
[74]Luo G,Wu TK,Zhou YL,et al.P relim inary studies on hemo stasia of Rhizoma Parid is saponin C.Pharm acol Clin Chin Mater Med,1988,4(2);37-39
[75]Takuo K,Masami Y,Kiyoshi S,et al.Studies on antitumor activities and antitumor principles of Chinese herbs I.Antitum or Activities Chin Herbs,1985,105(8):791-795
[76]Ji S,Zhou TS,Zhang JZ. Determ ination of antitumo rcytotoxic active substance gracillin in Rhizoma Paridis and Yunnan W hite.Chin Trad it Pat Med, 1992, 15(2):33-36
[77]李延.8种中草药的抗肿瘤作用实验.时珍国医国药,2001,l2(7):587-588
[78]詹锋,李质怀,刘和强.部分抗癌中药对HeLa细胞的抑制实验.兰州医学院学报,2003,29(4):l5-l6
[79]韦金育,李延,韦涛,等.50种广西常用中草药、壮药抗肿瘤作用的筛选研究.广西中医学院学报,2003,6(4):3-7
[80]金炜东,陈孝平,蔡红娇.重楼提取物对HepG2细胞的毒性作用.华中科技大学学报(医学版),2006,35(1):103-106
[81]Ravikumar P R,Hammesfah r P,Charles J.Cyto toxic saponins from the Chinese herbal drug Yunnan Baiyao. JPharm Sci,1979,68(7):900
[82]石小枫,杜德极,谢定成,等.重楼总皂苷对H22动物抑制性肿瘤的影响.中药材,1992,15(2):33-36
[83]程剑华,李以镔.抗癌植物药及其验方.南昌:江西科技出版社.1998:1l-13
[84]王建,黎海蒂,徐建伟,等.重楼皂甙对急性吗啡耐受大鼠痛反应及海马ACTH和G-EP含量的影响.第三军医大学学报,2000,22(12):1142
[85]Chiang HC,Wang JJ,wu RT.Immunomodulating effects of the hydrolysis products of formosanin C and beta-ecdysone from Paris formosana Hayata.Anticancer Res.1992,12(5):1475-1478
[86]李广勋.中药药理毒理与临床.天津:天津科技翻译出版公司,1992:415
[87]Hisashi M,Yutana,Toshio M,et a1.Protective Effects of Steroid Saponins from Paris polyphylla var. yunnanensis on Ethanol-or Indom-ethacin-Induced Gastric Mucosal Lesions in Rats:Structure Requirement for Activity and Mode of Action.Bioorganic& Medicinal Chemistry Letters,2003,13:l101-1106
[88]张继峰,田青,汤健等.从抗蛇毒药探寻内皮素拮抗剂的初步报告.南京医科大学学报(自然科学版),1993,13(3):228-229
[89]曹霖.七叶一枝花(Ⅱ)等4种化合物抑精子活性的研究.中草药,1987,18(10):19
[90]Ye YP, Hu L.200 Cases of women’s chlamydia infection treated with Zaoxiu powder. Shanxi J Trad it Chin Med,2000,21(8):352
[91]万近福,刘传东.昆明师范高等专科学校学报大孔吸附树脂分离重楼中偏诺类皂苷的研究,2005,27(4):7-8
[92]韦建荣,董汛.重楼中薯蓣皂甙元的反相高效液相色谱测定.色谱.1999,9(17):498-499
[93]Xu HaiWei , Li HaiDi , Wang Jian,et al.Rhizoma Paridis saponins reverses the decline of ACTH in hypothalamus of morphine tolerant arthritic rats.Chin J Neurosci.2001,17(3):259-263
[94]Zhou LiGang,Cao XiaoDong,Yang ChengZong,et,al.Endophytic fungi on Paris polyphyliavar.yunnna ensis and steroid analysis in the fungi. Natural Product Research and Development.2004,16(3):198-200
[95]黄芸,崔力剑,王强,等.南重楼Paris vietnamensis活性物质的分离与鉴定.药学学报,2006,41(4):361-364
[96]李群,葛方兰,王丽.重楼属植物初代培养过程中无菌培养物的建立.四川师范大学学报(自然科学版),2003,26(6):120-122
[97]侯玉平,胡晓立,沙莎,等.重楼的繁殖技术研究进展.云南大学学报(自然科学版),2004,26(6):60-62
[98]孟繁蕴,汪丽娅,张文生,等.滇重楼种胚休眠和发育过程中内源激素变化的研究.中药研究2006,34(4):36-38
[99]兰洁,杨明,马家骅.重楼提取工艺的研究.中草药,2006,37(6):871-872
[100]陈翠,杨丽云,吕丽芬,等.云南重楼根状茎切段苗繁育技术研究.西南农业学报,2007,20(4):706-710
[101]张玲琪,郭波,李海燕,等.长春花内生真菌的分离及其发酵产生药用成分的初步研究.中草药,2000,31(11):805-807
[102]张玲琪,谷苏,邵华,等.发酵产鸢尾酮真菌的分离鉴定及生香特性的初步研究.菌物系统,199,18(1):49-54
[103]Wang C,Wu J,Mei X. Enhancement of Taxol prodution and excretion in Taxus chinensis cell culture by fungal elicitation and medium renewal.Appl Microbiol Biotechnol,2001,55 (4):404-410
[104]江东福,马萍,张玲琪,等.龙血树真菌群及其对血竭形成的影响.云南植物研究,1995,17 (1):79
[105]陈晓梅,郭顺星,王春兰.四种内生真菌对金线莲无菌苗生长及多糖含量的影响.中国药学杂志,2005,40(1):13-16
[106]陈小静,冯定胜,赵明,等.四种华重楼内生细菌的初步研究.四川大学学报(自然科学版),2005,42(4):827-830
[107]孙桂丽,陈有为,夏国兴,等.云南重楼植物内生真菌的分离及抗菌活性筛选.微生物学杂志,2005,25(6):59-62
[108]张晓洁,查岭生,陈小静,等.一株产重楼皂甙内生细菌的分离与鉴定.微生物学通报,2006,33(5):84-88
[109]任智,张晓喻,祝凯,等.产薯蓣皂甙华重楼内生菌的筛选与鉴定.微生物学杂志,2007,27(2):6-9
[110]祝凯,张晓喻,任智,等.内芽孢杆菌属细菌SS02抗菌蛋白的分离纯化和其性质的研究.生物工程学报,2007,23(4):682-685
[113]东秀珠,蔡妙英.常用细菌系统鉴定手册.北京:科学出版社,2001.2
[114]张纪忠.微生物分类学.上海:复旦大学出版社,1990.12
[115]沈萍,范秀容,李广武.微生物学实验.北京:高等教育出版社,2003
[116]中国科学院微生物研究所细菌分类组.一般细菌常用鉴定方法.北京:科学出版社,1978.11
[117]申屠旭萍,陈宵峰,俞晓平.雷公藤内生真菌的分离及活性菌株的筛选.浙江农业学报,2006,18(5):308-312
[118]鲁素云.植物病害生物防治学.北京:北京农业大学出版社,1993,243-254
[119]中国科学院微生物研究所细菌分类组.一般细菌常用鉴定方法.北京:科学出版社,1978.11
[120]陆健,等.蛋白质纯化技术及应用.北京:化学工业出版社,2005.9
[121]何钟效.生物化学实验技术.北京:化学工业出版社,2004.3
[122]Gabriela S.Dveksler Carl W. Dieffenbach.PCR Primer:A Laboratory Manual.北京:科学出版社.
[123]刘志恒.现代微生物学.北京:科学出版社,2002,62-63
[122]边洪荣,李小娜,王会敏.重楼的研究及应用进展.中药材,2002,25(3):218-220
[123]杨海莲,孙晓璐,宋未.植物内生细菌的研究.微生物学通报,1998,25(4):224-227
[124]Johannes Haumann.Biotic interactions in plant-pathogen associations.London:CAB International,2001,90-91
[125] Li Haiyan, Qing Chen, Zhang Yanli,et al.Screening for endophytic fungi with antitumour and antifungal activities from Chinese medicinal plants.World Journal of Microbiology & Biotechnology.2005,21:1515-1519
[126]David M.W.Biologicol control of soilborne plant pathogens in the Rhizosphere with Bacteria.Ann.Rev.Phytopathol,1998,26:379-407
[127]郭亚辉,郭坚华,李斌.根据16SrRNA序列对假单胞菌属分类学的研究进展.微生物学杂志,2004,24(2):38-41
[128]王振雄,徐毅,周培瑾.嗜盐碱古生菌新种的系统分类学研究.微生物学报,2000,40(2):115-120
[129]黄海婵,裘娟萍.枯草芽孢杆菌防治植物病害的研究进展.浙江农业科学.2005,3:213-219
[130]徐积恩,朱明珍.抗生素.北京:科学出版社,1982,152
[131]黄国洋.农药实验技术与评价方法.北京:中国农业出版社,2000
[132]张兴,王兴林.植物化学保护实验指导.杨凌:西北农林科技大学出版社,2000:66-67
[133]Boman HG,Nilsson-Faye I,Paul K, et al.Characteristics of an inducible cell-free antibacterial reaction in hemolymph of Samia cynthia pupae.Infect Immunity,1974,10(1):136-145
[134]Vanittanakom N,Loeffler W.J Antibiotics ,1986 ,39(7):888-901
[135]Peypoux F,Guinand M,Michel G et al.Biochem,1978,17(19):3992-3996
[136]Peypoux F,Besson F,Michel G.J Antibiotics,1980,33(10):1146-1149
[137]Mhammedi A,Peypoux F,Besson F et al.J Antibiotics,1982,35(3):306-311
[138]Hansen JN.Antibiotics synthesized by posttranslational modification.Annual Review Microbiology,1993,47:535-564.
[139]Katz E,Demain AL.The peptide antibiotics of Bacillus:chemistry,biogenesis and possible functions. Bacteria Review,1997,41:449-474