High prevalence of methicillin resistant Staphylococcus aureus in pigs

Recently methicillin resistant Staphylococcus aureus (MRSA) was isolated from pigs and pig farmers in The Netherlands. In order to assess the dissemination of MRSA in the Dutch pig population, we screened 540 pigs in 9 slaughterhouses, where a representative portion of Dutch pigs (63%) was slaughtered in 2005. We found 209 (39%) of the pigs to carry MRSA in their nares. Forty-four of 54 groups of 10 consecutive pigs (81%), each group from a different farm, and all slaughterhouses were affected. All MRSA isolates belonged to 1 clonal group, showing Multi-Locus Sequence Type 398 and closely related spa types (mainly t011, t108 and t1254). Three types of the Staphylococcal Chromosome Cassette (SCCmec) were found: III (3%), IVa (39%) and V (57%). All 44 tested isolates (1 isolate per group) were resistant to tetracycline, reflecting the high and predominant use of tetracyclines in pig husbandry. Twenty-three percent of the isolates were resistant to both erythromycin and clindamycin and 36% to kanamycin, gentamicin and tobramycin but only a single isolate was resistant to co-trimoxazole and none to ciprofloxacin and several other antibiotics. The percentage of MRSA positive pigs was significantly different among slaughterhouses and among groups within slaughterhouses, indicating a high prevalence of MRSA in pigs delivered from the farms as well as cross contamination in the slaughterhouses.     

Retrospective detection of methicillin resistant and susceptible Staphylococcus aureus ST398 in Danish slaughter pigs

This retrospective study documents the first detection in Danish pigs of methicillin resistant and susceptible variants of the Staphylococcus aureus lineage ST398, which was previously described among pigs, pig farmers and veterinary staff in The Netherlands as well as in human patients and infected animals in Europe. Ten out of 100 pigs tested were shown to carry ST398 in the nasal cavity and two out of three farms tested were positive. The 10 porcine ST398 isolates had spa type t034 (n=9) and t1793 (n=1), and were resistant to erythromycin, clindamycin and tetracycline in addition to penicillin. The detection of ST398 in Danish pigs suggests that this new emerging zoonotic bacterium is rapidly spreading in the pig population in Europe.     

Methicillin resistant Staphylococcus aureus colonization in pigs and pig farmers

Methicillin resistant Staphylococcus aureus (MRSA) colonization has recently been identified in pigs and people that work with pigs, raising concerns about the role of pigs as reservoirs of MRSA for human infection. The objectives of this study were to evaluate the prevalence of MRSA colonization in pigs and pig farmers in Ontario, Canada and to characterize MRSA strains. Nasal and rectal swabs were collected from 285 pigs from three different age groups from 20 pig farms. Nasal swabs were collected from farm personnel and a brief questionnaire was also administered. The prevalence of MRSA colonization in farms was 45% (9/20) whereas the prevalence in pigs was 24.9% (71/285). There was no difference in MRSA colonization between age groups. The prevalence of MRSA colonization in pig farmers was 20% (5/25). There was a correlation between the presence of MRSA in pigs and humans on farms (P value=0.001). The results of spa typing revealed the predominant strain in pigs and humans was eGenomics spa type 539 (Ridom t034, clonal complex 398) which accounted for 59.2% of isolates and has been reported in pigs in Europe. A common human epidemic clone, CMRSA-2 (USA100, clonal complex 5) was also found in both pigs and pig personnel. Indistinguishable strains were found in pigs and pig personnel on all five farms with a colonized human. This study demonstrates that MRSA is common in pigs in Ontario, Canada, and provides further support to concerns about transmission of MRSA between pigs and humans.     

Study of methicillin resistant Staphylococcus aureus (MRSA) in Danish pigs at slaughter and in imported retail meat reveals a novel MRSA type in slaughter pigs

Methicillin-resistant Staphylococcus aureus (MRSA), especially CC398, have emerged in livestock worldwide. We investigated the occurrence of MRSA in pigs at slaughter and in retail meat. During 2009, nasal swabs (n=789) were taken from pigs at slaughter. Moreover, 866 meat samples [Danish: pork (153), broiler meat (121), beef (142) and; imported: pork (173), broiler meat (193), and beef (84)] were randomly collected in retail stores and outlets. MRSA was isolated from nasal swabs or from meat samples after preenrichment (Mueller Hinton broth with 6.5% NaCl), selective enrichment (tryptone soya broth with 4 mg/L cefoxitine and 75 mg/L aztreonam) and selective plating on Brilliance Chromogenic MRSA agar. The presence of mecA was confirmed by PCR and the MRSA isolates were spa typed. Novel MRSA spa types were characterized by MLST, PFGE and SCCmec typing. Thirteen percent (101/789) of the pigs had MRSA. Based on spa types 93% corresponded to CC398 (spa t011, t034, t1451, t2876, t2974), 4% to CC30 (t1333) and one isolate to CC1 (t0127). The spa type t1333 (CC30), which is common among methicillin susceptible S. aureus (MSSA) from pigs in Denmark, contained a SCCmec cassette type V and czrC zinc resistance gene. Imported broiler meat had the highest occurrence (18%) of MRSA, followed by imported pork (7.5%) and Danish pork (4.6%). MRSA ST398 was found for the first time in Danish beef (1.4%). The finding of MRSA CC30 (spa t1333) suggest possible spread of the SCCmec cassette normally associated with ST398 into another S. aureus lineage common in pigs.     

Epidemiology of methicillin resistant Staphylococcus pseudintermedius in guide dogs in Finland

Background

Methicillin resistant Staphylococcus pseudintermedius (MRSP) and Staphylococcus aureus (MRSA) are common multi-drug resistant (MDR) bacteria in dogs. In 2012–2013 three dogs of the Guide Dog School of the Finnish Federation of the Visually Impaired were found to be MRSP positive. Guide dogs have regular contact with each other during their first year of life and prolonged contact when in training. Since dogs are placed in different parts of Finland after training, there is a risk for national spread of MDR bacteria. In this study the prevalence of MRSP and MRSA, as well as the risk factors for MRSP were determined in the Finnish guide dog population. MRSP isolates were investigated using molecular methods and compared to the earlier isolates.

Results

Out of 132 tested dogs 4 were MRSP positive thus giving the prevalence estimate of 3% (95% CI: 1–8%) for MRSP in the target population. MRSA was not detected (prevalence estimate 0%, 95% CI: 0–3%). Risk factors associated with MRSP were being a breeding bitch (OR = 8.4; 95% CI: 1.1–64.1, P = 0.012), the number of veterinary visits (OR = 1.23; 95% CI: 1.0–1.5, P = 0.025) and number of antimicrobial courses (OR = 1.63; 95% CI: 1.0–2.55; P = 0.035). Identified MRSP isolates belonged to five different sequence types (ST45, 71, 402, 403 and 404). All ST71 isolates carried SCCmec II-III, while the SCCmec type of the ST45 and ST402 (a single locus variant of ST45) isolates were non-typeable with the method used.

Conclusions

MRSP and MRSA had low prevalence in the studied dog population despite the close contact between dogs, and the MRSP population was heterogenic. Antimicrobial therapy and veterinary visits are risk factors for MRSP even among a small case group.

Electronic supplementary material

The online version of this article (doi:10.1186/s13028-015-0129-8) contains supplementary material, which is available to authorized users.     

Occurrence and characterization of livestock-associated methicillin-resistant Staphylococcus aureus in pig industries of northern Thailand

This study was conducted to determine the prevalence of livestock-associated methicillin-resistant Staphylococcus aureus (LA-MRSA) in pigs, farm workers, and the environment in northern Thailand, and to assess LA-MRSA isolate phenotypic characteristics. One hundred and four pig farms were randomly selected from the 21,152 in Chiang Mai and Lamphun provinces in 2012. Nasal and skin swab samples were collected from pigs and farm workers. Environmental swabs (pig stable floor, faucet, and feeder) were also collected. MRSA was identified by conventional bacterial culture technique, with results confirmed by multiplex PCR and multi locus sequence typing (MLST). Herd prevalence of MRSA was 9.61% (10 of 104 farms). Among pigs, workers, and farm environments, prevalence was 0.68% (two of 292 samples), 2.53% (seven of 276 samples), and 1.28% (four of 312 samples), respectively. Thirteen MRSA isolates (seven from workers, four from environmental samples, and two from pigs) were identified as Staphylococcal chromosomal cassette mec IV sequences type 9. Antimicrobial sensitivity tests found 100% of the MRSA isolates resistant to clindamycin, oxytetracycline, and tetracycline, while 100% were susceptible to cloxacillin and vancomycin. All possessed a multidrug-resistant phenotype. This is the first evidence of an LA-MRSA interrelationship among pigs, workers, and the farm environment in Thailand.     

Characteristics of methicillin resistant Staphylococcus aureus isolated from chicken meat and hospitalized dogs in Korea and their epidemiological relatedness

Methicillin resistant Staphylococcus aureus (MRSA) is one of the most important pathogens in human and veterinary hospitals. The isolation of MRSA from animals and foodstuffs has been reported with an increased incidence. However, methicillin (oxacillin) is not used in animal husbandry or in animal hospitals in Korea. In this study, three pre-MRSA and one silent mecA-carrying methicillin susceptible S. aureus (smMSSA) were isolated from retail chicken meat, and three MRSA were isolated from hospitalized dogs in Korea. The three pre-MRSA isolates were determined to have a staphylococcal cassette chromosome mec (SCCmec) type III, and the smMSSA isolate was not classified. The animal hospital isolates were found to contain SCCmec type II. Seven and 15 S. aureus isolated from hospitalized humans and bovine milk, respectively, were also examined in this study in order to determine the epidemiological origins of MRSA. Multilocus sequencing typing (MLST) revealed that the chicken meat and bovine milk isolates were closely related to the animal hospital isolates. The SCCmec characteristics and MLST analyses indicated the possibility of the human to animal transmission of MRSA. These results highlight the importance of identifying MRSA carriers as well as intercepting MRSA transmission because MRSA is becoming increasingly widespread without any plausible relationship with the use of methicillin (oxacillin).     

Prevalence of canine methicillin resistant Staphylococcus pseudintermedius in a veterinary diagnostic laboratory in Italy

The overall prevalence of methicillin-resistant Staphylococcus pseudintermedius (MRSP) was 2% (10/590) among 590 canine specimens submitted to an Italian veterinary diagnostic laboratory during a two-month period, and 21% (10/48) among Staphylococcus intermedius group (SIG) isolates. All methicillin-resistant strains exhibited additional resistance to fluoroquinolones, gentamicin, lincosamides, tetracyclines, and potentiated sulfonamides, belonged predominantly to spa type t02 and harboured SCCmec type II–III cassette.     

Prevalence of canine methicillin resistant Staphylococcus pseudintermedius in a veterinary diagnostic laboratory in Italy

The overall prevalence of methicillin-resistant Staphylococcus pseudintermedius (MRSP) was 2% (10/590) among 590 canine specimens submitted to an Italian veterinary diagnostic laboratory during a two-month period, and 21% (10/48) among Staphylococcus intermedius group (SIG) isolates. All methicillin-resistant strains exhibited additional resistance to fluoroquinolones, gentamicin, lincosamides, tetracyclines, and potentiated sulfonamides, belonged predominantly to spa type t02 and harboured SCCmec type II–III cassette.     

Methicillin‐resistant Staphylococcus aureus keratitis in a dog

The purpose of this study is to report a case of methicillin‐resistant Staphylococcus aureus (MRSA) keratitis in a dog. A 7‐year‐old intact male American cocker spaniel that had undergone removal of a nictitating gland was referred for severe ulcerative keratitis. Slit‐lamp examination showed swelling of the eyelid, mucopurulent discharge, conjunctival injection and chemosis, diffuse corneal edema and opacity, and a deep ulcer in central cornea. Gram staining of discharge from the eye demonstrated Gram‐positive cocci. Despite topical ofloxacin, oxytetracycline and polymyxin B ophthalmic solution and intravenous cefazolin, there was no improvement. Cultures revealed MRSA that was sensitive only to chloramphenicol, vancomycin, lincomycin, and clindamycin. The antibiotic regimen was changed to topical and systemic chloramphenicol. After 9 days of treatment, although inflammation started to be resolved, the dog developed nonregenerative anemia. The antimicrobial regimen was changed again to topical and systemic vancomycin. Inflammation continued to improve over the next week. MRSA should be considered a potential organism in infectious keratitis, especially when general antibiotics are not effective. Although topical and systemic chloramphenicol and/or vancomycin are effective for treating MRSA keratitis, vancomycin should only be used when culture and susceptibility results indicate it is appropriate and no other options are available. To our knowledge, this is the first detailed case report of MRSA keratitis in a dog.     

Zinc resistance of Staphylococcus aureus of animal origin is strongly associated with methicillin resistance

This study was conducted to determine the occurrence of zinc and copper resistances in methicillin-resistant Staphylococcus aureus (MRSA) from swine and veal calves in a global strain collection. The test population consisted of 476 porcine MRSA isolates from ten European countries, 18 porcine MRSA isolates from Canada and seven MRSA from China, 92 MRSA and 60 methicillin-susceptible S. aureus (MSSA) isolates from veal calves in the Netherlands and 88 porcine MSSA isolates from four European countries. Most porcine MRSA (n=454) and all bovine MRSA belonged to clonal complex (CC) 398 whereas 37 of the pig MRSA from Europe and the seven Chinese isolates belonged to other CCs and 3 isolates were not classified into a CC. All isolates were tested for susceptibility to zinc chloride and copper sulphate using agar dilution and tested by PCR for the czrC gene encoding zinc resistance. Phenotypic zinc resistance (MIC>2mM) was observed in 74% (n=324) and 42% (n=39) of European MRSA CC398 from pigs and veal calves, respectively, and in 44% of the Canadian isolates (n=8), but not among the Chinese isolates. Almost all (99%) zinc-resistant MRSA carried czrC. Of the 37 European non-CC398 MRSA, 62% were resistant to zinc, but only 46% of them carried czrC. The MICs of the MSSA isolates to zinc chloride ranged from 1 to 4mM and none carried czrC. The MICs of copper sulphate were associated neither with methicillin resistance nor with the detection of czrC. This study showed that zinc resistance and the czrC gene are widespread among CC398 MRSA isolates. This suggests that the use of zinc in feed might have contributed to the emergence of MRSA.     

Genetic diversity and methicillin resistance of Staphylococcus aureus originating from buffaloes with mastitis in Iran

The aims of the present study were to investigate the genetic diversity and methicillin resistance in S. aureus isolates recovered from mastitis-affected buffaloes. Five hundred seventy-eight milk samples were obtained from buffaloes with mastitis in three provinces, Iran. Ninety-one of the 578 tested samples contained S. aureus (15.74%), in two cases were methicillin resistant S. aureus (MRSA). Isolates were typed by spa typing, followed by MLST on some representative isolates and SCCmec typing for MRSA strains. The presence of genes encoding Panton–Valentine leukocidin (PVL) was also tested by PCR. Eight spa types were identified, with t3576 (n = 18), t7311 (n = 18) and t937 (n = 17) were the most common, followed by t304 (n = 11), t7308 (n = 9), t521 (n = 7), t267 (n = 6), and t527 (n = 5). MLST revealed four different sequence types (STs) including ST97 (related to t521 and t527 spa types), ST352 (related to t267), ST291 (related to t304 and t937) and ST522 (related to t7338, t7311 and t3576). Two MRSA were identified as t304-ST291-SCCmecIV and t7311-ST522-SCCmecIV. No PVL-positive S. aureus were found. A significant difference in geographical distribution of genotypes was observed, with some types being prevalent in all studied provinces (P < 0.001). The results demonstrated genetic diversity among the S. aureus strains involved in mastitis in buffaloes. This study also provides evidence of the presence of MRSA belonging to genotypes which have been earlier reported in human infections, emphasizing the need for their epidemiological monitoring.     

Quantification of within- and between-pen transmission of Foot-and-Mouth disease virus in pigs

Quantified transmission parameters of Foot-and-Mouth Disease Virus (FMDV) are needed for epidemic models used for control and surveillance. In this study, we quantified the within- and between-pen transmission of FMDV in groups of pigs by estimating the daily transmission rate beta, i.e. the number of secondary infections caused by one infectious pig during one day, using an SIR (susceptible-infectious-removed) model. Within-pen transmission was studied in four groups of ten pigs in which 5 infected and 5 susceptible pigs had direct contact; between-pen transmission was studied in one group of ten pigs in which 5 infected and 5 susceptible pigs had indirect contact. Daily results of virus isolation of oropharyngeal fluid were used to quantify the transmission rate beta, using Generalised Linear Modelling (GLM) and a maximum likelihood method. In addition, we estimated the expected time to infection of the first pig within a pen T(w) and in the indirect-contact pen T(b). The between-pen transmission rate beta(b) was estimated to be 0.59 (0.083-4.18) per day, which was significantly lower than the within-pen transmission rate beta(w) of 6.14 (3.75-10.06). T(w) was 1.6 h, and T(b) was 16 h. Our results show that the transmission rate is influenced by contact structure between pigs.     

The first detection of methicillin-resistant Staphylococcus aureus ST398 in pigs in Korea

In order to determine the presence of methicillin-resistant Staphylococcus aureus (MRSA) in pigs in Korea, a total of 657 nasal swabs were collected from pigs on 66 different pig farms nationwide during February 2008-May 2009. The prevalences of MRSA positive samples in pigs and farms were 3.2% (21/657) and 22.7% (15/66), respectively. Two different types were found among the 21 MRSA isolates: 17 strains of livestock-associated type (LA; ST398 or ST541/spa t034) and 4 strains of human-associated type (HA; ST72/spa t664 or t2461). The most prevalent type of MRSA strain was ST398/t034 (12/21, 57%), followed by ST541/t034 (5/21, 23.8%). The rest of the isolates were ST72/t664 (n=2) and ST72/t2461 (n=2), respectively. Our data provide evidence for the existence of not only LA types (ST398 and ST541) but also HA type (ST72) MRSA in pigs in Korea. This survey provides the first evidence of LA type MRSA in animals in Korea. In addition, the presence of human MRSA clones in pigs observed in this study suggests an additional reservoir for human MRSA infection, and vice versa.     

Multidrug- and methicillin resistant Staphylococcus pseudintermedius as a cause of canine pyoderma: a case report

A case of a dog with a long-term inflammatory skin disorder due to infection with methicillin-resistant Staphylococcus pseudintermedius (MRSP) is described. After initial diagnostics of MRSP, follow-up swabs of the dog (nose, skin) were taken twice after four and seven weeks. MRSP was constantly isolated from the skin and once from the nose. Since infected humans might be a source of reinfection, the owners of the dog were screened (nasal) three times during their pet's therapy. Thereby, the male owner was found to be colonized with MRSP once in the first sampling round. Comparative typing of all MRSP-isolates by pulsed-field gel electrophoresis (PFGE), SCCmec typing, multilocus sequence typing (MLST), spa typing, PCR-detection of the leukotoxin encoding operon (LukI) and the Staphylococcus intermedius-exfoliative toxin (SIET) as well as antimicrobial resistance profiling by broth microdilution revealed that all five MRSP isolates from the dog and the single isolate from the owner were indistinguishable by any of the applied methods. All isolates were assigned to a certain strain, a multidrug-resistant MRSP belonging to sequence type (ST) 71, spa type (t)05, harbouring SCCmecIII as well as the genes encoding LukI and SIET. In this case, a number of reasons might have contributed to therapy failure and re-infection, respectively (e. g. contact to other MRSP-colonized dogs, contact to MRSP-colonized humans, refusal to clip the dog's fur). In addition, MRSP-contaminated objects or surfaces in the household, which were difficult to disinfect or simply not considered as a potential source of MRSP, might have served as a source of re-infection. These results envision the possibility of a dog-to-human transmission of MRSP and the relevance of this aspect as a potential source of re-infection in cases of bacterial-supported long-term skin disorders in canine patients. First cases of MRSP infections in humans have been described only recently. However, the general pathogenic potential of multidrug resistant MRSP in humans is unknown so far and needs further investigation.     

质粒介导的喹诺酮耐药基因在动物源金黄色葡萄球菌的流行分布

为了解广东省动物源金黄色葡萄球菌耐药现状、分析耐喹诺酮类金黄色葡萄球菌耐药分子特征,本研究从2010—2011年广东33个养殖场的不同动物来源的样本中分离鉴定出46株金黄色葡萄球菌,采用二倍琼脂稀释法测定对14种抗菌药物的敏感性;采用聚合酶链式反应(PCR)检测耐喹诺酮金黄色葡萄球菌grlA、grlB、gyrA、gyrB基因在喹诺酮耐药决定区(QRDR)的突变特征、质粒介导喹诺酮类耐药(PMQR)基因(qnr、aac6′-Ib-cr、oqxA和qepA)的流行分布特征,同时还检测了β-内酰胺酶编码基因(blaCTX-M、blaCMY和blaSHV)、质粒介导金黄色葡萄球菌耐氟苯尼考的基因(cfr和fexA)和万古霉素耐药基因(vanA、vanB和vanC)。结果表明,46株金黄色葡萄球菌对克林霉素耐药率最高为67%,对环丙沙星和氟苯尼考耐药率为52%,对青霉素、氨苄西林、庆大霉素、红霉素、四环素和复方新诺明的耐药率在40%⁵0%之间,对苯唑西林和氯霉素耐药率为28%,对妥布霉素耐药率为13%,对头孢噻肟的耐药率在10%左右,有2株耐万古霉素。20株耐环丙沙星金黄色葡萄球菌的grlA和gyrA基因的QRDR发生了碱基突变并导致编码的氨基酸发生改变,GrlA氨基酸只有1种突变模式,Ser80→Phe,而GyrA氨基酸存在3种突变模式:Ser84→Leu,Ala或Phe。PMQR基因中,oqxA检出率最高(80%),其次是aac6′-Ib-cr占13%,qnrS1检出3株、qnrD检出2株。检测到2株菌株携带超广谱β-内酰胺酶编码基因blaSHV-12;23株氟苯尼考耐药菌株全部携带fexA基因,其中3株同时携带cfr基因;2株耐万古霉素菌株未检测到相关耐药基因。广东省动物源金黄色葡萄球菌多重耐药现象较为普遍,本研究首次在动物源金黄色葡萄球菌中检测到超广谱质粒编码的喹诺酮耐药基因流行分布广泛,大多数菌株同时携带2种以上质粒编码的耐药基因,提示养殖过程或治疗过程中使用任何一种药物(尤其是动物促生长剂)都可以筛选出耐药菌株。     

Transmission of methicillin resistant Staphylococcus aureus among pigs during transportation from farm to abattoir

The prevalence of methicillin resistant Staphylococcus aureus (MRSA) in pigs at abattoirs is higher than in pigs sampled on farms. This study investigated whether MRSA negative pigs can become MRSA positive during transportation from the farm to the abattoir after exposure to other pigs and environmental sources of MRSA. Nasal swabs were collected from four batches of pigs during loading at the farm, on arrival at the abattoir and after stunning. Environmental wipes were taken from lorries after transporting pigs and from lairages after holding pigs. All pigs (n = 117) tested MRSA negative before transportation. On arrival at the abattoir, 12/117 (10.3%) pigs in two batches tested MRSA positive. In lorries that tested positive after transportation, the prevalence of MRSA positive pigs was 21.1%, whereas no MRSA was detected in pigs that had been transported in lorries that tested negative after transportation. At stunning, all batches and 70/117 (59.8%) pigs tested MRSA positive. Pigs can become MRSA positive in the short period of time during transportation from the farm to stunning at the abattoir.     

Quantification of the spread of Mycoplasma hyopneumoniae in nursery pigs using transmission experiments

Mycoplasma hyopneumoniae (M. hyopneumoniae) is present in almost all swine herds worldwide, but transmission of the pathogen through herds is not yet fully clarified. The aim of this study, performed in 2003, was to investigate and to quantify the transmission of M. hyopneumoniae under experimental conditions by means of an adjusted reproduction ratio (R_n). This R_n-value, calculated according to the final size method, expresses the mean number of secondary infections due to one typical infectious piglet during the nursery period. The period lasted from 4 to 10 weeks of age, corresponding with the nursery period used in most European production systems. Additionally, a comparison was made between transmissions of highly virulent and low virulent isolates.

Forty-eight weaned piglets, free of M. hyopneumoniae, were housed in six separate pens. During 6 weeks, two animals experimentally infected with M. hyopneumoniae were housed together with six susceptible piglets. At the end of the study, the number of contact-infected animals was determined by the use of nPCR on bronchoalveolar lavage fluid. The R_n-values of the highly virulent and the low virulent isolates were estimated to be 1.47 (0.68–5.38) and 0.85 (0.33–3.39), respectively. No significant difference between the groups was found (P = 0.53). The overall R_n was estimated to be 1.16 (0.94–4.08). Under the present experimental conditions, the transmission of M. hyopneumoniae, assessed for the first time by a reproduction ratio, shows that one piglet infected before weaning will infect on average one penmate during the nursery phase.