植物源细菌群体感应抑制因子的筛选及其对生物膜形成的影响
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摘要
长期以来,人们认为细菌之间彼此孤立,互不联系,但近几年的研究发现,细菌世界也和人类世界一样,它们也拥有自己的“语言”,这种信号分子被称为信号分子,细菌细胞产生信号分子并释放到环境中去,当环境中的信号分子达到一定浓度后开启细胞密度依赖的特定基因的表达,这一现象被称为“群体感应”(quorum sengsing,QS),它是细菌根据细胞密度变化进行基因表达调控的一种生理行为。研究发现,细菌通过群体感应系统调控致病过程的一些重要环节,如致病因子产生、生物膜形成等。
微生物是导致食品腐败和感染的主要因素,而且只有腐败细菌或致病细菌达到一定数量后才能导致食品的腐败和致病性的发生。随着抗菌药物和防腐剂的广泛应用,细菌的耐药性变得越来越严重,已成为全球关注的危害人类健康的重大难题。近年来,AHLs介导的群体感应现象如何影响食品的品质以及与食品腐败和致病菌致病性的关系成为食品防腐保鲜和致病菌防治领域的一个新的研究热点。因此,细菌群体感应系统干扰的研究对于新型防腐剂和抗菌药物的研究与开发具有重要意义。
本研究以革兰氏阴性菌的酰基高丝氨酸内酯介导的群体感应系统为靶标,从植物材料中筛选群体感应抑制因子,并研究其对大肠杆菌ATCC 8099产生物膜能力的抑制作用,为研究、开发新型抗菌药物和防腐剂提供理论依据和技术支持。具体研究如下：1植物源群体感应抑制因子的筛选
本研究利用产紫色素的菌株,Chromobacterium violaceum 026(CV026),进行群体感应抑制剂检测,其紫色素的产生受信号分子acylated lactone(AHL)的调控。采用平板检测法,研究了34种香辛料及花茶提取物对Chromobacterium violaceum 026(CV026)产紫色能力的抑制作用及生长的影响,从中筛选出薰衣草和玫瑰花提取物具有干扰Chromobacterium violaceum 026(CV026)群体感应系统的作用,且不影响Chromobacterium violaceum 026(CV026)的生长。
2薰衣草和玫瑰花提取物群体感应抑制效应的研究
本部分采用不同的提取方式,采用定性和定量两种方法研究薰衣草和玫瑰花提取物对Chromobacterium violaceum 026(CV026)群体感应系统的抑制作用,提取溶剂采用有机溶剂和水,结果表明薰衣草以乙酸乙酯作为提取溶剂,提取时间为24h效果最好。玫瑰花以水作为提取溶剂,提取方式为超声波处理1h效果最好。并研究其在不同浓度下对Chromobacterium violaceum 026(CV026)产紫色素能力的影响,发现抑制作用与提取物浓度呈明显的量效关系。
3薰衣草和玫瑰花提取物对大肠杆菌产生物膜能力的影响
本部分以产生物膜的菌株大肠杆菌ATCC 8099为研究基础,以薰衣草乙酸乙酯提取物和玫瑰花超声波处理1h的水提取液为研究对象,通过定性和定量两种方法检测两种提取物对大肠杆菌ATCC 8099产生物膜能力的影响。结果表明,两种提取物均可以明显抑制大肠杆菌ATCC 8099产生物膜。定性研究可以看出,添加提取物的实验组明显比对照组产生的膜厚度要薄,亮度更低,且可看见明显的孔洞。在不同浓度下研究薰衣草乙酸乙酯提取物对大肠杆菌ATCC 8099产生物膜能力的影响,发现抑制作用与提取物浓度呈明显的量效关系。
For a long time, bacteria was considered to be isolated and had no connection with each other. However, in recent years, studies have found that the world of bacteria was like the human world. They also have their own "language". This language was called signal molecular. Bacterial cells can produce signal molecules and release it into the environment. When the signal molecular reached a certain concentration in the environment, it made some specific genes which depended on the bacterial density express. This phenomenon was called "quorum sensing, QS"). Quorum sensing was known as a physiological behavior which can regulate gene expression according to the changes of cell density. The studies had found that bacterial quorum-sensing system can control the important steps in the pathogenic process, such as the production of pathogenic factors, biofilm formation and so on.
Microbacteria was a major factor in the area of food spoilage and infection. Only the bacterial density reached a certain number can lead to the occurrence of food spoilage and pathogenicity. With antibiotics and preservatives widely used, bacterial resistance had become more and more serious. It had become a global problem, a major hazard to human health. In recent years, researches were on how AHLs-mediated quorum sensing affects food spoilage and pathogenicity of pathogenic bacteria. It was a new research focus in the field of the preservation of food and pathogen control. This kind of research can give us a promising approach in the field of food preservation and pathogen control.
In this study, we used the N-acylhomoserine lactone-mediated quorum-sensing system of gram-negative bacteria as a target to screen quorum sensing inhibitor from the plant materials and to study the influence on biofilm formation of Escherichia coli ATCC 8099. From this reaearch, we wanted to provide theoretical basis and technical support in the development of new antimicrobial agents. Specific studies are as follows:
1 Screeing for materials which can interrupt quorum sensing system
In this study, we use the bacteria, Chromobacterium violaceum 026 (CV026), which can pruduce purple pigment regulated by the signal molecule acylated lactone (AHL). We used the method of plate detection to study the capacity of inhibiting the production of purple pigment and the growth of CV026 by the extract of the 34 kinds of spices and tea. We screed extracts of lavender and roses had the capacity of QSI that can interrupt the quorum sensing system but can't interfer with the growth of CV026.
2 Research of the capacity of QSI of extracts of lavender and rose
In this part, we used different kinds of extraction methods to get the extraction. In order to check the capacity of inhibiting the quorum sensing system of Chromobacterium violaceum 026(CV026), we applied the quatitative and qualitative methods. We used organic solvents and water as extraction solvent. In order to get the best results, we used ethyl acetate ester as the extraction solvent of lavender and extracted for 24h quietly. Rose was different from lavender. We used water as the extraction solvent and extracted for 1h by ultrasonic treatment. Further, we studied the inhibition of production of purple pigment of CV026 by different concentrations of the extracts and found that inhibition with the concentration of the extracts showed significant dose-effect relationship.
3 Influence on biofilm formation of E.coli 8099 by lavender and rose extraction
In this part, we checked the influence on biofilm formation of E.coli ATCC 8099 by lavender ethyl acetate extraction and rose water extraction by ultrasonic treated for 1h. The studied qualitative and quantitative methods were applied in this part. The results showed that the two extracts can inhibit biofilm formation of E.coli ATCC 8099. From the qualitative results, we can see the biofilm was thinner and less brightter in the study group than in the control group. Also, we can see obvious holes in the study group. Furthermore, we studied the inhibition of biofilm formation of E. coli ATCC8099 by lavender extraction at different concentrations and found that inhibition with the concentration of the extracts showed significant dose-effect relationship.
微生物是导致食品腐败和感染的主要因素,而且只有腐败细菌或致病细菌达到一定数量后才能导致食品的腐败和致病性的发生。随着抗菌药物和防腐剂的广泛应用,细菌的耐药性变得越来越严重,已成为全球关注的危害人类健康的重大难题。近年来,AHLs介导的群体感应现象如何影响食品的品质以及与食品腐败和致病菌致病性的关系成为食品防腐保鲜和致病菌防治领域的一个新的研究热点。因此,细菌群体感应系统干扰的研究对于新型防腐剂和抗菌药物的研究与开发具有重要意义。
本研究以革兰氏阴性菌的酰基高丝氨酸内酯介导的群体感应系统为靶标,从植物材料中筛选群体感应抑制因子,并研究其对大肠杆菌ATCC 8099产生物膜能力的抑制作用,为研究、开发新型抗菌药物和防腐剂提供理论依据和技术支持。具体研究如下：1植物源群体感应抑制因子的筛选
本研究利用产紫色素的菌株,Chromobacterium violaceum 026(CV026),进行群体感应抑制剂检测,其紫色素的产生受信号分子acylated lactone(AHL)的调控。采用平板检测法,研究了34种香辛料及花茶提取物对Chromobacterium violaceum 026(CV026)产紫色能力的抑制作用及生长的影响,从中筛选出薰衣草和玫瑰花提取物具有干扰Chromobacterium violaceum 026(CV026)群体感应系统的作用,且不影响Chromobacterium violaceum 026(CV026)的生长。
2薰衣草和玫瑰花提取物群体感应抑制效应的研究
本部分采用不同的提取方式,采用定性和定量两种方法研究薰衣草和玫瑰花提取物对Chromobacterium violaceum 026(CV026)群体感应系统的抑制作用,提取溶剂采用有机溶剂和水,结果表明薰衣草以乙酸乙酯作为提取溶剂,提取时间为24h效果最好。玫瑰花以水作为提取溶剂,提取方式为超声波处理1h效果最好。并研究其在不同浓度下对Chromobacterium violaceum 026(CV026)产紫色素能力的影响,发现抑制作用与提取物浓度呈明显的量效关系。
3薰衣草和玫瑰花提取物对大肠杆菌产生物膜能力的影响
本部分以产生物膜的菌株大肠杆菌ATCC 8099为研究基础,以薰衣草乙酸乙酯提取物和玫瑰花超声波处理1h的水提取液为研究对象,通过定性和定量两种方法检测两种提取物对大肠杆菌ATCC 8099产生物膜能力的影响。结果表明,两种提取物均可以明显抑制大肠杆菌ATCC 8099产生物膜。定性研究可以看出,添加提取物的实验组明显比对照组产生的膜厚度要薄,亮度更低,且可看见明显的孔洞。在不同浓度下研究薰衣草乙酸乙酯提取物对大肠杆菌ATCC 8099产生物膜能力的影响,发现抑制作用与提取物浓度呈明显的量效关系。
For a long time, bacteria was considered to be isolated and had no connection with each other. However, in recent years, studies have found that the world of bacteria was like the human world. They also have their own "language". This language was called signal molecular. Bacterial cells can produce signal molecules and release it into the environment. When the signal molecular reached a certain concentration in the environment, it made some specific genes which depended on the bacterial density express. This phenomenon was called "quorum sensing, QS"). Quorum sensing was known as a physiological behavior which can regulate gene expression according to the changes of cell density. The studies had found that bacterial quorum-sensing system can control the important steps in the pathogenic process, such as the production of pathogenic factors, biofilm formation and so on.
Microbacteria was a major factor in the area of food spoilage and infection. Only the bacterial density reached a certain number can lead to the occurrence of food spoilage and pathogenicity. With antibiotics and preservatives widely used, bacterial resistance had become more and more serious. It had become a global problem, a major hazard to human health. In recent years, researches were on how AHLs-mediated quorum sensing affects food spoilage and pathogenicity of pathogenic bacteria. It was a new research focus in the field of the preservation of food and pathogen control. This kind of research can give us a promising approach in the field of food preservation and pathogen control.
In this study, we used the N-acylhomoserine lactone-mediated quorum-sensing system of gram-negative bacteria as a target to screen quorum sensing inhibitor from the plant materials and to study the influence on biofilm formation of Escherichia coli ATCC 8099. From this reaearch, we wanted to provide theoretical basis and technical support in the development of new antimicrobial agents. Specific studies are as follows:
1 Screeing for materials which can interrupt quorum sensing system
In this study, we use the bacteria, Chromobacterium violaceum 026 (CV026), which can pruduce purple pigment regulated by the signal molecule acylated lactone (AHL). We used the method of plate detection to study the capacity of inhibiting the production of purple pigment and the growth of CV026 by the extract of the 34 kinds of spices and tea. We screed extracts of lavender and roses had the capacity of QSI that can interrupt the quorum sensing system but can't interfer with the growth of CV026.
2 Research of the capacity of QSI of extracts of lavender and rose
In this part, we used different kinds of extraction methods to get the extraction. In order to check the capacity of inhibiting the quorum sensing system of Chromobacterium violaceum 026(CV026), we applied the quatitative and qualitative methods. We used organic solvents and water as extraction solvent. In order to get the best results, we used ethyl acetate ester as the extraction solvent of lavender and extracted for 24h quietly. Rose was different from lavender. We used water as the extraction solvent and extracted for 1h by ultrasonic treatment. Further, we studied the inhibition of production of purple pigment of CV026 by different concentrations of the extracts and found that inhibition with the concentration of the extracts showed significant dose-effect relationship.
3 Influence on biofilm formation of E.coli 8099 by lavender and rose extraction
In this part, we checked the influence on biofilm formation of E.coli ATCC 8099 by lavender ethyl acetate extraction and rose water extraction by ultrasonic treated for 1h. The studied qualitative and quantitative methods were applied in this part. The results showed that the two extracts can inhibit biofilm formation of E.coli ATCC 8099. From the qualitative results, we can see the biofilm was thinner and less brightter in the study group than in the control group. Also, we can see obvious holes in the study group. Furthermore, we studied the inhibition of biofilm formation of E. coli ATCC8099 by lavender extraction at different concentrations and found that inhibition with the concentration of the extracts showed significant dose-effect relationship.
引文
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[1]綦国红,董明盛.细菌胞间的分子通讯与食品保藏新策略[J].广西农业生物科学,2005,24(3)：259-263.
[2]Fuqua W C, Winans S C. Quorum sensing in bacteria:the LuxR-Lux Ⅰ family of cell density-responsive transcriptional regulator[J]. Journal of Bacteriology,1994,176:269-275.
[3]Nealson K H, Platt T, Hastings J W. Cellular control of the synthesis and activity of the bacterial luminescent system[J]. Journal of Bacteriology,1970,104:313-322.
[4]郭嘉亮,陈卫民.细菌群体感应信号分子与抑制剂研究进展[J].生命科学,2007,19(2)：224-232.
[5]黄远帅,尹一兵.细菌密度感应系统的信号干扰及其应用[J].生命的化学,2005,25(2)：86-88.
[6]杨荣华,林家莲.香辛料的抗菌性[J].中国调味品,1999,12：2-4.
[7]D. A. Vattem, K. Mihalik, S. H. Crixell, et al. Dietary phytocheicals as quorum sensing inhibitors. Fitoterapia,2007,78:302-310.
[8]J.H. Choo, Y. Rukayadi, J.-K. Hwang. Inhibition of bacterial quorum sensing by vanilla extract. Microbiology,2006,42:637-641.
[9]H. Zhu, S. J. Sun. Inhibition of Bacterial quorum sensing-regulated behaviors by Tremella fuciformis Extract. Current Microbiology,2008:DOI10.1007/s 00284-008-9215-8.
[10]Gulgun Bosgelmez-tinaz, Seyhan Ulusoy, Aysel Ugur, et al. Inhibition of Quorum sensing-Regulated Behaviors by Scorzonera sandrasica. Current Microbiology,2007,55:114-118.
[11]Allison Adonizio, Kelsey Downum, Bradley C. Bennett, et al. Anti-quorum sensing activity of medicinal plants in southern Florida. Journal of Ethnopharmacology,2006,105:427-435.
[12]Robert J.C. McLean, Leland S. Pierson Ⅲ, Clay Fuqua. A simple screening protocol for the identification of quorum signal antagonists[J]. Journal of Microbiological Methods,2004(58): 351-360.
[13]P. Truchado, M. I. Gil, A. Allende, et al. Quorum sensing inhibitory and antimicrobial activities of honeys and the relationship with indivisdual phenolics[J]. Food Chemistry,2009,115:1337-1344.
[14]Basavaraj Girenha L. Cepeda, Kamlesh A. Soni, et al. Grapefruit juice and its furocoumarins inhibits autoinducer signaling and biofilm formation in bacteria. International Journal of Food Microbiology, 2008,125:204-208.
[1]吴霞,刘净,于志斌,等.薰衣草化学成分的研究[J].化学学报,2007,65(16)：1649-1653.
[2]胡喜兰,韩照祥,刘玉芬,等.薰衣草挥发油的抗氧化活性测定和成分比较[J].食品科技,2006,9：115-117.
[3]王玉芹,孙亚军,施献儿.薰衣草精油的化学成分与药理活性[J].国外医药,2004,19(1)：5-8.
[4]马猛华,崔波,于海峰,等.玫瑰花的研究进展[J].山东轻工业学院学报,2008,22(4)：38-42.
[5]李明,彭艳丽,韩莉,等.不同品种玫瑰花挥发油化学成分的GC-MS分析[J].中成药,2008,30(5)：726-730.
[6]McClean K H, Winson M K, Fish L, et al. Quorum sensing and Chromobacterium violaceum: exploitation of violacein production and inhibition for the detection of N-acylhomoserine lactone[J]. Microbiology,1997,143:3703-3711.
[7]綦国红,董明盛.细菌胞间的群体感应系统及其新型的抗菌策略[J].中国抗生素杂志,2006,31(3)：135-139.
[8]McClean K H, Pierson L S P, Fuqua C, et al. A simple screening protocol for the identification of quorum signal antagonists[J]. Journal of Microbiology,2004,58:351-360.
[9]Robert J.C. McLean, Leland S. Pierson III, Clay Fuqua. A simple screening protocol for the identification of quorum signal antagonists[J]. Journal of Microbiological Methods,2004(58): 351-360.
[10]Gulgun Bosgelmez-tinaz, Seyhan Ulusoy, Aysel Ugur, et al. Inhibition of Quorum sensing-Regulated Behaviors by Scorzonera sandrasica[J]. Current Microbiology,2007,55:114-118.
[11]Allison Adonizio, Kelsey Downum, Bradley C. Bennett, et al. Anti-quorum sensing activity of medicinal plants in southern Florida[J]. Journal of Ethnopharmacology,2006(105):427-435.
[12]H. Zhu, S. J. Sun. Inhibition of Bacterial quorum sensing-regulated behaviors by Tremella fuciformis Extract[J]. Current Microbiology,2008:DOI10.1007/s 00284-008-9215-8.
[13]J.H. Choo, Y. Rukayadi, J.-K. Hwang. Inhibition of bacterial quorum sensing by vanilla extract[J]. Microbiology,2006,42:637-641.
[14]D.A. Vattem, K. Mihalik, S.H. Crixell, et al. Dietary phytochemical as quorum sensing inhibitors[J]. Fitoterapia,2007(78):302-310.
[15]P.,Truchado, M. I. Gil, A. Allende, et al. Quorum sensing inhibitory and antimicrobial activities of honeys and the relationship with indivisdual phenolics [J]. Food Chemistry,2009,115:1337-1344.
[16]Renee S. Blosser, Kendall M. Gray. Extraction of violacein from Chromobacterium violaceum provides a new quantitative bioassay for N- acyl homoserine lactone autoinducers[J]. Journal Microbiological Methods,2000(40):47-55.
[17]郭嘉亮,陈卫民.细菌群体感应信号分子与抑制剂研究进展[J].生命科学,2007,19(2)：224-232.
[18]黄远帅,尹一兵.细菌密度感应系统的信号干扰及其应用[J].生命的化学,2005,25(2)：86-88.
[19]Basavaraj Girenha L. Cepeda, Kamlesh A. Soni, et al. Grapefruit juice and its furocoumarins inhibits autoinducer signaling and biofilm formation in bacteria[J]. International Journal of Food Microbiology,2008,125:204-208.
[1]干霞芳,李蒙英.生物膜和生物膜形成菌的研究[J].安徽大学学报,2007,30(6)：91-93.
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[1]綦国红,董明盛.细菌胞间的分子通讯与食品保藏新策略[J].广西农业生物科学,2005,24(3)：259-263.
[2]Fuqua W C, Winans S C. Quorum sensing in bacteria:the LuxR-Lux Ⅰ family of cell density-responsive transcriptional regulator[J]. Journal of Bacteriology,1994,176:269-275.
[3]Nealson K H, Platt T, Hastings J W. Cellular control of the synthesis and activity of the bacterial luminescent system[J]. Journal of Bacteriology,1970,104:313-322.
[4]郭嘉亮,陈卫民.细菌群体感应信号分子与抑制剂研究进展[J].生命科学,2007,19(2)：224-232.
[5]黄远帅,尹一兵.细菌密度感应系统的信号干扰及其应用[J].生命的化学,2005,25(2)：86-88.
[6]杨荣华,林家莲.香辛料的抗菌性[J].中国调味品,1999,12：2-4.
[7]D. A. Vattem, K. Mihalik, S. H. Crixell, et al. Dietary phytocheicals as quorum sensing inhibitors. Fitoterapia,2007,78:302-310.
[8]J.H. Choo, Y. Rukayadi, J.-K. Hwang. Inhibition of bacterial quorum sensing by vanilla extract. Microbiology,2006,42:637-641.
[9]H. Zhu, S. J. Sun. Inhibition of Bacterial quorum sensing-regulated behaviors by Tremella fuciformis Extract. Current Microbiology,2008:DOI10.1007/s 00284-008-9215-8.
[10]Gulgun Bosgelmez-tinaz, Seyhan Ulusoy, Aysel Ugur, et al. Inhibition of Quorum sensing-Regulated Behaviors by Scorzonera sandrasica. Current Microbiology,2007,55:114-118.
[11]Allison Adonizio, Kelsey Downum, Bradley C. Bennett, et al. Anti-quorum sensing activity of medicinal plants in southern Florida. Journal of Ethnopharmacology,2006,105:427-435.
[12]Robert J.C. McLean, Leland S. Pierson Ⅲ, Clay Fuqua. A simple screening protocol for the identification of quorum signal antagonists[J]. Journal of Microbiological Methods,2004(58): 351-360.
[13]P. Truchado, M. I. Gil, A. Allende, et al. Quorum sensing inhibitory and antimicrobial activities of honeys and the relationship with indivisdual phenolics[J]. Food Chemistry,2009,115:1337-1344.
[14]Basavaraj Girenha L. Cepeda, Kamlesh A. Soni, et al. Grapefruit juice and its furocoumarins inhibits autoinducer signaling and biofilm formation in bacteria. International Journal of Food Microbiology, 2008,125:204-208.
[1]吴霞,刘净,于志斌,等.薰衣草化学成分的研究[J].化学学报,2007,65(16)：1649-1653.
[2]胡喜兰,韩照祥,刘玉芬,等.薰衣草挥发油的抗氧化活性测定和成分比较[J].食品科技,2006,9：115-117.
[3]王玉芹,孙亚军,施献儿.薰衣草精油的化学成分与药理活性[J].国外医药,2004,19(1)：5-8.
[4]马猛华,崔波,于海峰,等.玫瑰花的研究进展[J].山东轻工业学院学报,2008,22(4)：38-42.
[5]李明,彭艳丽,韩莉,等.不同品种玫瑰花挥发油化学成分的GC-MS分析[J].中成药,2008,30(5)：726-730.
[6]McClean K H, Winson M K, Fish L, et al. Quorum sensing and Chromobacterium violaceum: exploitation of violacein production and inhibition for the detection of N-acylhomoserine lactone[J]. Microbiology,1997,143:3703-3711.
[7]綦国红,董明盛.细菌胞间的群体感应系统及其新型的抗菌策略[J].中国抗生素杂志,2006,31(3)：135-139.
[8]McClean K H, Pierson L S P, Fuqua C, et al. A simple screening protocol for the identification of quorum signal antagonists[J]. Journal of Microbiology,2004,58:351-360.
[9]Robert J.C. McLean, Leland S. Pierson III, Clay Fuqua. A simple screening protocol for the identification of quorum signal antagonists[J]. Journal of Microbiological Methods,2004(58): 351-360.
[10]Gulgun Bosgelmez-tinaz, Seyhan Ulusoy, Aysel Ugur, et al. Inhibition of Quorum sensing-Regulated Behaviors by Scorzonera sandrasica[J]. Current Microbiology,2007,55:114-118.
[11]Allison Adonizio, Kelsey Downum, Bradley C. Bennett, et al. Anti-quorum sensing activity of medicinal plants in southern Florida[J]. Journal of Ethnopharmacology,2006(105):427-435.
[12]H. Zhu, S. J. Sun. Inhibition of Bacterial quorum sensing-regulated behaviors by Tremella fuciformis Extract[J]. Current Microbiology,2008:DOI10.1007/s 00284-008-9215-8.
[13]J.H. Choo, Y. Rukayadi, J.-K. Hwang. Inhibition of bacterial quorum sensing by vanilla extract[J]. Microbiology,2006,42:637-641.
[14]D.A. Vattem, K. Mihalik, S.H. Crixell, et al. Dietary phytochemical as quorum sensing inhibitors[J]. Fitoterapia,2007(78):302-310.
[15]P.,Truchado, M. I. Gil, A. Allende, et al. Quorum sensing inhibitory and antimicrobial activities of honeys and the relationship with indivisdual phenolics [J]. Food Chemistry,2009,115:1337-1344.
[16]Renee S. Blosser, Kendall M. Gray. Extraction of violacein from Chromobacterium violaceum provides a new quantitative bioassay for N- acyl homoserine lactone autoinducers[J]. Journal Microbiological Methods,2000(40):47-55.
[17]郭嘉亮,陈卫民.细菌群体感应信号分子与抑制剂研究进展[J].生命科学,2007,19(2)：224-232.
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