亚治疗剂量四环素饲喂对牛源微生物耐药基因的影响
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摘要
以低于治疗水平的氯四环素(CT),以及低于治疗水平的氯四环素和治疗水平的氧四环素组合(CT-OX)两种方式分别对肉牛进行抗生素处理,研究其对肠道大肠杆菌耐药基因型的影响。粪便样品采自商业肉牛饲养场,并有详实的抗生素使用历史记录,通过抗菌药物纸片法和稀释法敏感性试验测试分离出的大肠杆菌对四环素、氧四环素和氯四环素的敏感性。利用tet(A),tet(B)和tet(C)基因的引物进行多重PCR试验,检测对四环素耐药或中介的细菌样品(176个),结果发现全部样品均携带一种或者两种耐药基因,tet(A)在两组样品中的流行基本相同,但CT组中tet(B)的流行比例较CT-OX组小(P＜0.05),而tet(C)的流行比例则较大(P＜0.05)。同时,在对四环素表现为中介的样品(52个)检测结果中,发现其中92.3%呈现出携带tet(C)基因。另外,最小抑菌浓度值(MICs)结果表明,药物敏感性同时取决于四环素类别和耐药基因型两方面的因素。利用real-time PCR在转录水平上分析tet(C)基因,发现耐药型与中介型并非上游调控造成。但是,对tet(C)基因的测序分析发现耐药型的第1063处碱基由T突变为G。由上述数据可以看出,对肉牛的四环素饲喂种类可以影响到大肠杆菌的耐药基因流行。
将肉牛分为亚治疗剂量氯四环素的饲喂组及其对照组,在对其处理60天后进行粪便与咽部分泌物样品的采集,且对另一部分肉牛进行饲喂前后同一动物的样品采集,提取其宏基因组DNA,研究其中四环素耐药基因、磺胺耐药基因和红霉素耐药基因的浓度变化。结果表明,亚治疗剂量四环素的饲喂对tet(B)基因浓度的影响是显著的,四环素的存在的确对该基因的存在产生了选择压力；粪便中携带tet(C)基因的大肠杆菌数目较少,因此tet(C)基因在粪便中的浓度相对较低；咽部分泌物中tet(H)基因浓度较高；四环素处理同样对tet(M)、tet(O)和tet(W)基因的选择压力也是明显的,而粪便样品中这4类基因的浓度较高。对于磺胺耐药基因来说,sul2基因在粪便样品中表现出了较高的浓度。在红霉素耐药基因中,亚治疗剂量四环素的饲喂对erm(A)和erm(X)基因的影响是较为显著的。
通过对肉牛给予低于治疗水平的不同种类抗生素,研究其对废弃粪便微生物群落中四环素耐药基因数量与持久性的影响。试验肉牛被分成不同的抗生素处理组：氯四环素组,氯四环素和磺胺甲嘧啶组,以及对照组。将每个围栏中所有动物的新鲜粪便混匀为一份混合样品作为模式样品(每个处理组3份),分别在露天放置的第7、14、28、42、56、70、84、98、112、126和175天时采样并提取DNA,利用real-time PCR方法测定四环素耐药基因tet(B),tet(C),tet(L),tet(M),tet(W)以及16S-rRNA的浓度。结果显示,16S-rRNA的浓度在不同处理间相似,在56天内保持增加(P＜0.05)；总体上看,四环素组的耐药基因初始浓度较高(P＜0.05)；所有处理组的tet(B)和tet(C)浓度到56天时均增长了1～2个对数级,到175天时又降低到初始水平,而tet(M)与tet(W)的浓度与其他耐药基因相比较高。由此可见,四环素耐药基因可以在废弃粪便中持续存在超过175天,而某些基因的最初数量可能会导致错估其后的变化,其浓度的暂时改变并不能归因于微生物群落数量的变化。
The effect of administering feedlot cattle subtherapeutic levels of chlortetracycline (CT) or CT and therapeutic levels of oxytetracycline (CT-OX) on resistance genotype in Escherichia coli was investigated. Fecal samples were collected from cattle at commercial feedlots and had a documented history of antimicrobial use. Isolates were tested for susceptibility to tetracycline, oxytetracycline, and chlortetracycline using disk diffusion or broth microdilution. Detection of tet(A), tet(B), and tet(C) genes encoded by tetracycline-resistant isolates (N=176) was performed by multiplex PCR. All isolates encoded one or a combination of two resistance genes. Prevalence of tet(A) was similar between groups of E. coli, however prevalence of tet(B) was lower(P<0.5) and tet(C) was greater(P<0.5) in CT isolates. The nature of the tet determinants was further assessed in a group of intermediately tetracycline-resistant isolates (N=52). Minimum inhibitory concentrations showed that susceptibility was dependent on tetracycline analogue and the type of resistance determinant. The tet(C) gene was present in92%of these isolates. Copies of tet(C) transcripts, analyzed by real-time PCR, indicated that up-regulation did not occur in tetracycline-resistant isolates when compared to intermediately-resistant isolates. However, sequence analysis of the tet(C) gene revealed a T→G substitution at position1063in resistant isolates that may have affected phenotype. These data provide insight into the relationship between the type of tetracycline analogue administered to cattle and the prevalence of resistance genes in E. coli.
The beef cattle were separated into subtherapeutic tetracycline treated group and non-treated group, and the fecal samples and pharyngeal secretion samples were collected at≥60DOF, and another animals were collected before and after tetracycline treat. The metagenomic DNA was extracted to quantify the tetracycline resistant gene, sulfanilamide resistant gene, and erythrocin resistant gene. The results showed that, the concerntration of tet(B) was affected by subtherapeutic tetracycline administering, and the administration put a selective pression on it. The concerntration of tet(C) gene is lower than others in fecal samples,because of the susceptibility in E.coli. The selective pression on tet(M), tet(O) and tet(W) is present, so that their concerntration in fecal samples showed a higher level. sul2gene show a higher concerntration in fecal samples. And for erythrocin resistant gene, the erm(A) an erm(X) gene were affected by the treat.
This study investigated the effects of administering beef cattle antimicrobials at subtherapeutic concentrations on the abundance and persistence of tetracycline resistance genes within the whole microbial community of fecal waste. Cattle were administered chlortetracycline, chlortetracycline plus sulfamethazine and no antimicrobials. Model fecal deposits (N=3) were prepared by mixing fresh feces from pens into a single composite sample. Real-time PCR was used to measure concentrations of tetracycline resistance genes tet(B), tet(C), tet(L), tef(M), tet(W) and16S-rRNA in DNA extracted from composite feces after7,14,28,42,56,70,84,98,112,126, and175days in the field. The concentrations of16S-rRNA in feces were similar across treatments and increased by day56(P<0.05). Generally, initial concentrations of tetracycline resistance genes were greater (P<0.05) in fecal pats from animals fed chlortetracycline. For all fecal treatments, tet(B) and tet(C) increased1～2log units by day56, and then decreased to the initial day7levels by day175(P>0.05). The concentrations of tet(M) and tet(W) were greater than other tetracycline resistance determinants. Tetracycline resistance genes can persist in fecal waste from cattle beyond175days and the initial load of some genes may underestimate concentrations at later time points. Temporal changes in the concentrations of resistance genes were likely due to shifts in microbial populations.
将肉牛分为亚治疗剂量氯四环素的饲喂组及其对照组,在对其处理60天后进行粪便与咽部分泌物样品的采集,且对另一部分肉牛进行饲喂前后同一动物的样品采集,提取其宏基因组DNA,研究其中四环素耐药基因、磺胺耐药基因和红霉素耐药基因的浓度变化。结果表明,亚治疗剂量四环素的饲喂对tet(B)基因浓度的影响是显著的,四环素的存在的确对该基因的存在产生了选择压力；粪便中携带tet(C)基因的大肠杆菌数目较少,因此tet(C)基因在粪便中的浓度相对较低；咽部分泌物中tet(H)基因浓度较高；四环素处理同样对tet(M)、tet(O)和tet(W)基因的选择压力也是明显的,而粪便样品中这4类基因的浓度较高。对于磺胺耐药基因来说,sul2基因在粪便样品中表现出了较高的浓度。在红霉素耐药基因中,亚治疗剂量四环素的饲喂对erm(A)和erm(X)基因的影响是较为显著的。
通过对肉牛给予低于治疗水平的不同种类抗生素,研究其对废弃粪便微生物群落中四环素耐药基因数量与持久性的影响。试验肉牛被分成不同的抗生素处理组：氯四环素组,氯四环素和磺胺甲嘧啶组,以及对照组。将每个围栏中所有动物的新鲜粪便混匀为一份混合样品作为模式样品(每个处理组3份),分别在露天放置的第7、14、28、42、56、70、84、98、112、126和175天时采样并提取DNA,利用real-time PCR方法测定四环素耐药基因tet(B),tet(C),tet(L),tet(M),tet(W)以及16S-rRNA的浓度。结果显示,16S-rRNA的浓度在不同处理间相似,在56天内保持增加(P＜0.05)；总体上看,四环素组的耐药基因初始浓度较高(P＜0.05)；所有处理组的tet(B)和tet(C)浓度到56天时均增长了1～2个对数级,到175天时又降低到初始水平,而tet(M)与tet(W)的浓度与其他耐药基因相比较高。由此可见,四环素耐药基因可以在废弃粪便中持续存在超过175天,而某些基因的最初数量可能会导致错估其后的变化,其浓度的暂时改变并不能归因于微生物群落数量的变化。
The effect of administering feedlot cattle subtherapeutic levels of chlortetracycline (CT) or CT and therapeutic levels of oxytetracycline (CT-OX) on resistance genotype in Escherichia coli was investigated. Fecal samples were collected from cattle at commercial feedlots and had a documented history of antimicrobial use. Isolates were tested for susceptibility to tetracycline, oxytetracycline, and chlortetracycline using disk diffusion or broth microdilution. Detection of tet(A), tet(B), and tet(C) genes encoded by tetracycline-resistant isolates (N=176) was performed by multiplex PCR. All isolates encoded one or a combination of two resistance genes. Prevalence of tet(A) was similar between groups of E. coli, however prevalence of tet(B) was lower(P<0.5) and tet(C) was greater(P<0.5) in CT isolates. The nature of the tet determinants was further assessed in a group of intermediately tetracycline-resistant isolates (N=52). Minimum inhibitory concentrations showed that susceptibility was dependent on tetracycline analogue and the type of resistance determinant. The tet(C) gene was present in92%of these isolates. Copies of tet(C) transcripts, analyzed by real-time PCR, indicated that up-regulation did not occur in tetracycline-resistant isolates when compared to intermediately-resistant isolates. However, sequence analysis of the tet(C) gene revealed a T→G substitution at position1063in resistant isolates that may have affected phenotype. These data provide insight into the relationship between the type of tetracycline analogue administered to cattle and the prevalence of resistance genes in E. coli.
The beef cattle were separated into subtherapeutic tetracycline treated group and non-treated group, and the fecal samples and pharyngeal secretion samples were collected at≥60DOF, and another animals were collected before and after tetracycline treat. The metagenomic DNA was extracted to quantify the tetracycline resistant gene, sulfanilamide resistant gene, and erythrocin resistant gene. The results showed that, the concerntration of tet(B) was affected by subtherapeutic tetracycline administering, and the administration put a selective pression on it. The concerntration of tet(C) gene is lower than others in fecal samples,because of the susceptibility in E.coli. The selective pression on tet(M), tet(O) and tet(W) is present, so that their concerntration in fecal samples showed a higher level. sul2gene show a higher concerntration in fecal samples. And for erythrocin resistant gene, the erm(A) an erm(X) gene were affected by the treat.
This study investigated the effects of administering beef cattle antimicrobials at subtherapeutic concentrations on the abundance and persistence of tetracycline resistance genes within the whole microbial community of fecal waste. Cattle were administered chlortetracycline, chlortetracycline plus sulfamethazine and no antimicrobials. Model fecal deposits (N=3) were prepared by mixing fresh feces from pens into a single composite sample. Real-time PCR was used to measure concentrations of tetracycline resistance genes tet(B), tet(C), tet(L), tef(M), tet(W) and16S-rRNA in DNA extracted from composite feces after7,14,28,42,56,70,84,98,112,126, and175days in the field. The concentrations of16S-rRNA in feces were similar across treatments and increased by day56(P<0.05). Generally, initial concentrations of tetracycline resistance genes were greater (P<0.05) in fecal pats from animals fed chlortetracycline. For all fecal treatments, tet(B) and tet(C) increased1～2log units by day56, and then decreased to the initial day7levels by day175(P>0.05). The concentrations of tet(M) and tet(W) were greater than other tetracycline resistance determinants. Tetracycline resistance genes can persist in fecal waste from cattle beyond175days and the initial load of some genes may underestimate concentrations at later time points. Temporal changes in the concentrations of resistance genes were likely due to shifts in microbial populations.
引文
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74 Kim S R, Nonaka L. Occurrence of tetracycline resistance genes tet (M) and tet(S) in bacteria frommarine acquaculture sites[J]. FEMS Microbiol. Lett,2004,237:147-156.
75 L-Abee-Lund TM, Sorum H. A global non-conjugativeTet C plasmid, pRAS3, from Aeromonas salmonicida [J]. Plasmid,2002,47:172-181.
76 Furushita M, Shiba T. Similarity of tetracycl ine resistance genes isolated from fish farmbacteria to those from clinical isolates [J]. Appl. Environ. Microbiol, 2003,69:5336-5342.
77 Lancaster H, Roberts AP. Characterization of Tn916S, a Tn916-like elementcontaining the tetracycline resistance determinant tet(S) [J]. J.Bacteriol,2004,186:4395-4398.
78 Del Grosso M, Abusco AC. Tn2009, a Tn916-like element containingmef (E) in Streptococcus pneumoniae. Antimicrob [J]. Agents Chemother,2004,48:2037-2042.
79 Giovanetti E, Brenciani A. The presence of the tet(O) gene in erythromycinand tetracycline-resistant strains of Streptococcus pyogenes[J]. Antimicrob. Agents Chemother,2003,47:2844-2849.
80 Brenciani A, Ojo KK. A new geneticelement, carrying tet(O) and mef (A) genes[J]. J. Antimicrob. Chemother,2004,54:991-998.
81 Giovanetti E, Brenciani A. The presence of the tet (0) gene in erythromycinand tetracycline-resistant strains of Streptococcus pyogenes[J]. Antimicrob. Agents Chemother,2003,47:2844-2849.
82 Brenciani A, Ojo KK. A new geneticelement, carrying tet(O) and mef(A) genes [J]. J. Antimicrob. Chemother,2004,54:991-998.
83 Melville CM, Brunel R. The Butyrivibrio fibrisolvens tet(W) gene is carried on the novel conjugative transposon TnB1230, which contains duplicatednitroreductase coding sequences [J]. J. Bacteriol,2004,186:3656-3659.
84 Available from:http://www.faculty.washington.edu/marilynr/(visited November 15, 2004).
85 Connell SR, Tracz DM. Ribosomal protection proteins and their mechanism of tetracycline resistance [J]. Antimicrob. Agents Chemother,2003,47:3675-3681.
86 Connell SR, Trieber CA. Mechanism of Tet (O), perturbs theconformation of the ribosomal decoding center [J]. Mol. Microbiol,2003,45:1463-1472.
87 Saphn CM, Blaha G. Localization of the ribosomalprotection protein Tet(O) on the ribosome-and the mechanismof tetracycline resistance[J]. Mol. Cell,2001,7:1037-1045.
88 Melville CM, Scott KP. Novel tetracycline resistance gene, tet (32), in the Clostridiumrelatedhuman colonic anaerobe K10 and its transmission in vitroto the rumen anaerobe Butyrivibrio fibrisolvens [J]. Antimicrob. Agents Chemother,2001,45: 3246-3249.
89 Whittle G, Whitehead TR. Identification of a newribosomal protection type of tetracycline resistance gene, tet (36), from swine manure pits [J]. Appl. Environ. Microbiol,2003,69:4151-4158.
90 Stanton TB, Humphrey SB. Isolation of tetracycline-resistant Megasphaera elsdenii strains with novel mosaic gene combinations of tet (0) andtet (W) from swine [J]. Appl. Environ. Microbiol,2003,69:3874-3882.
91 Stanton TB, McDowall JS. Diverse tetracycline-resistant genotypes of Megasphaera elsdeni i strains select ively cultured from swine feces [J]. Appl. Environ. Microbiol. 70,3754-3757.
92 Diaz-Torres ML, McNab R. Characterization of anovel tetracycline resistance determinate from the oral metage-nome[J]. Antimicrob. Agents Chemother, 2003,47:1430-1432.
93 Nonaka L, Suzuk S. New Mg2+-dependent oxytetracycline resistance determinant Tet34 in Vibrio isolates from marine intestinal contents.2002.
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76 Furushita M, Shiba T. Similarity of tetracycl ine resistance genes isolated from fish farmbacteria to those from clinical isolates [J]. Appl. Environ. Microbiol, 2003,69:5336-5342.
77 Lancaster H, Roberts AP. Characterization of Tn916S, a Tn916-like elementcontaining the tetracycline resistance determinant tet(S) [J]. J.Bacteriol,2004,186:4395-4398.
78 Del Grosso M, Abusco AC. Tn2009, a Tn916-like element containingmef (E) in Streptococcus pneumoniae. Antimicrob [J]. Agents Chemother,2004,48:2037-2042.
79 Giovanetti E, Brenciani A. The presence of the tet(O) gene in erythromycinand tetracycline-resistant strains of Streptococcus pyogenes[J]. Antimicrob. Agents Chemother,2003,47:2844-2849.
80 Brenciani A, Ojo KK. A new geneticelement, carrying tet(O) and mef (A) genes[J]. J. Antimicrob. Chemother,2004,54:991-998.
81 Giovanetti E, Brenciani A. The presence of the tet (0) gene in erythromycinand tetracycline-resistant strains of Streptococcus pyogenes[J]. Antimicrob. Agents Chemother,2003,47:2844-2849.
82 Brenciani A, Ojo KK. A new geneticelement, carrying tet(O) and mef(A) genes [J]. J. Antimicrob. Chemother,2004,54:991-998.
83 Melville CM, Brunel R. The Butyrivibrio fibrisolvens tet(W) gene is carried on the novel conjugative transposon TnB1230, which contains duplicatednitroreductase coding sequences [J]. J. Bacteriol,2004,186:3656-3659.
84 Available from:http://www.faculty.washington.edu/marilynr/(visited November 15, 2004).
85 Connell SR, Tracz DM. Ribosomal protection proteins and their mechanism of tetracycline resistance [J]. Antimicrob. Agents Chemother,2003,47:3675-3681.
86 Connell SR, Trieber CA. Mechanism of Tet (O), perturbs theconformation of the ribosomal decoding center [J]. Mol. Microbiol,2003,45:1463-1472.
87 Saphn CM, Blaha G. Localization of the ribosomalprotection protein Tet(O) on the ribosome-and the mechanismof tetracycline resistance[J]. Mol. Cell,2001,7:1037-1045.
88 Melville CM, Scott KP. Novel tetracycline resistance gene, tet (32), in the Clostridiumrelatedhuman colonic anaerobe K10 and its transmission in vitroto the rumen anaerobe Butyrivibrio fibrisolvens [J]. Antimicrob. Agents Chemother,2001,45: 3246-3249.
89 Whittle G, Whitehead TR. Identification of a newribosomal protection type of tetracycline resistance gene, tet (36), from swine manure pits [J]. Appl. Environ. Microbiol,2003,69:4151-4158.
90 Stanton TB, Humphrey SB. Isolation of tetracycline-resistant Megasphaera elsdenii strains with novel mosaic gene combinations of tet (0) andtet (W) from swine [J]. Appl. Environ. Microbiol,2003,69:3874-3882.
91 Stanton TB, McDowall JS. Diverse tetracycline-resistant genotypes of Megasphaera elsdeni i strains select ively cultured from swine feces [J]. Appl. Environ. Microbiol. 70,3754-3757.
92 Diaz-Torres ML, McNab R. Characterization of anovel tetracycline resistance determinate from the oral metage-nome[J]. Antimicrob. Agents Chemother, 2003,47:1430-1432.
93 Nonaka L, Suzuk S. New Mg2+-dependent oxytetracycline resistance determinant Tet34 in Vibrio isolates from marine intestinal contents.2002.
94 Weaver KE, Rice LB. Plasmids and transposons In:The Enterococci:Pathogenesis, Molecular Biology and Antibiotic Resistance [J]. Gi lmore, M.S., Ed.,2002:219-263.
95 Avrain L, Vernozy-Rozand C. Evidence for natural horizontal transfer of tetO gene between Campylobacter jejuni strains in chickens [J]. J. Appl. Microbiol,2004, 97:134-140.
96 Luna VA, Roberts MC. The presence of the tetO gene in a variety of tetracycline resistant Streptococcus pneumoniae serotypes fromWashington State[J]. J. Antimicrob. Chemother,1998,42:613-619.
97 Weigel LM, Musser K. Genetic Analysis of a Vancomycin-resistant Staphylococcus aureus (VRSA) Isolated from New York. In:Abstracts of the 44th Interscience Conference on Antimicrobial Agents and Chemotherapy,2004,69:C1-941.
98 Dugan J, Rockey DD. Tetracycline resistance in Chlamydia suis mediated by genomic isolated inserted into the chlamydial inv-like gene[J]. Antimicrob. Agents Chemother, 2004,48:3989-3995.
99 Jones RB, Van der Pol B. Partial characterization of Chlamydia trachmatis isolates resistant to multiple antibiotics [J]. J. Infect. Dis.1990,162:1309-1315.
100 Somani J, Bhullar VB. Multiple drug-resistant Chlamydia trachomatis associated with clinical treatment failure[J]. J. Infect. Dis,2000,181:1421-1427.
101 Lefevre FC, Lepargneur JP. Tetracycline-resistant Chlamydia trachomatis in Toulouse, France [J]. Path. Biol,1997,45:376-378.
102 Moulder JM. Chlamydiaceae[J]. Bergys Manual of Systematic Bacteriology,1984, Vol. 1:729.
103 Sengelov G, Halling S. Susceptibility of Escherichia coli and Enterococcus faecium isolated from pigs and broiler chickens to tetracycline degradation products and distribution of tetracycline resistance determinants in E. coli from food animals.
104 Bryan A, Shapir N. Frequency and distribution of tetracycline resistance genes in genetically diverse, nonselected, nonclinical Escherichia coli strains isolated from diverse human and animal sources [J]. Appl. Environ. Microbiol,2004,70:2503-2507.
105 Pasquali F, De Cesare A. Phage types, ribotypes and tetracycline resistance genes ofSalmonella enterica subsp. Enterica serovar Typhimurium strains isolated from different origins in Italy [J]. Vet. Microbiol,2004,103:71-76.
106 Recchia GD, Hall RM. Gene cassettes:a new class of mobile element [J]. Microbiology,1995,141:3015-3027.
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