ࡱ> %`%bjbjNN4,,@;( pT$p6p6p6P67|B<Zn=@==@=@4 KTO$hNhQr qru@"@[ssP== FMMMGt =r =MqrMMrT r =< `0p6Հ\k7\1PTyz1VDP M\8MdkXPPPdPPPqrqrqrqrdL$EL  Geographic distribution of the dominant strains of Staphylococcus aureus causing invasive infections in Europe. See list of authors attached Summary Background Staphylococcus aureus is one of the most important human pathogens and its methicillin-resistant variant (MRSA) is a major cause of hospital and community-acquired infection. We aimed to map the geographic distribution of the dominant strains that cause invasive infections on a continental scale in Europe. Methods In each country, staphylococcal reference laboratories secured the participation of an adequate number of laboratories with the aim to achieve national geo-demographic representation. Participating laboratories collected successive methicillin-susceptible (MSSA) and MRSA isolates per hospital from patients with invasive S. aureus infection using an agreed protocol in a systematic manner. All isolates were sent to the reference laboratories and characterised by quality controlled sequence typing of the variable region of the staphylococcal spa gene (spa typing) and data were uploaded to a central database. Pertinent demographic detail and genetic and phenotypic information were assembled for interactive interrogation at a purpose-build web-based mapping application. Findings Between September 2006 and February 2007, 357 laboratories serving 450 hospitals in 26 countries collected 2,890 MSSA and MRSA isolates from patients with invasive S. aureus infection. Spa types showed a wide geographical distribution with some spa types prevalent in all European countries. MSSA were more heteroclonal than MRSA. Genetic diversity of MRSA differed considerably between countries with dominant MRSA spa types forming distinctive geographical clusters. We provide evidence that a network approach consisting of decentralised typing and central data aggregation improve the understanding about the prevalence and spread of strains with particular public health importance. Interpretation In contrast to MSSA, MRSA spa types have a predominantly regional abundance indicative of the selection and spread of a limited number of clones within the community and health care networks. Funding All contributions of the staphylococcal reference laboratories were funded by national sources. Data collection, management and analysis were funded by the Dutch Ministry of Welfare and Sports and Directorate General for Health and Consumer Protection of the European Commission. Introduction Staphylococcus aureus is the main cause of purulent infection in humans [1]. S. aureus has the potential for local as well as disseminating infection and can cause lesions in all tissues and anatomical sites. Infections are mainly acquired in the community or in association with health care. The position as one of the most important human pathogens is largely due to the ubiquitous occurrence of S. aureus as a coloniser in humans, domestic animals and livestock [2]. Between 25-35% of healthy human individuals carry S. aureus on the skin or mucous membranes [3]. Any injury that compromises epithelial integrity, trauma, medical or surgical interventions, as well as viral infections, can lead to tissue invasion [4-6]. It is assumed that severity and outcome depend largely on the virulence of the introduced strain and the immune repertoire of the host [7, 8]. Occasionally, S. aureus acquires enhanced virulence and antimicrobial resistance through horizontal DNA transfer and maintains these mobile genetic elements in a predominantly clonal genomic background. Thus, clones of S. aureus are relatively stable and diversify predominantly by the accumulation of single nucleotide substitutions in the absence of frequent inter-strain recombination. It is therefore possible to discern different clonal lineages by simple genomic typing [9]. The ability to identify certain strains and their clonal background allows for several important observations on the evolution, epidemiology and spread of clones with particular public health importance, such as hospital- and community-, and livestock-associated methicillin-resistant S. aureus (MRSA). For MRSA, this is particularly important since it appears that certain lineages have disseminated over wide geographical regions and are currently a major threatening major to achievements in curative and public health medicine [10]. Geographical detail of this expansion on a continental scale has however been scant, due to the lack of appropriate surveys and agreement on a consistent application of standardized genomic typing approaches. At the same time little is known about the population structure and geographical abundance of methicillin-susceptible S. aureus (MSSA) which after all provide the genetic reservoir from which MRSA emerge. This investigation was designed to fill these knowledge gaps and provides (i) the first representative and contemporary snapshot of the genetic population structure of S. aureus that cause invasive infection in the European region, (ii) an insight into the geographical abundance of different clones among MSSA and MRSA on a continental scale, and (iii) a web-based mapping tool that provides interactive access and an intuitive illustration of the results generated by this large scale typing initiative. Methods Capacity building During three workshops organised for technical personnel from 28 European Staphylococcus aureus Reference Laboratories (SRLs), participants received hands-on training in spa sequence typing and spa sequence analysis according to a standard protocol using a purpose-designed software StaphType"! developed by Ridom"!, GmbH, Wuerzburg, Germany [11]. Proficiency testing was carried out by supplying each SRL with 5 isolates of well characterized S. aureus isolates and 5 sequence chromatograms (trace files) of known spa types as described previously [12, 13]. Only All participating laboratories, in the structured survey described here, that fulfilled quantifiable quality criteria which consisted of an unambiguous base-calling for all sequenced nucleotides for both forward and reverse sequencing runs of the test panel. were eligible to participate in the structured survey described here. Structured survey A protocol was drawn up and circulated for comment to all SRLs and agreed upon in April 2006. Following this protocol, European SRLs were asked to identify about 20 laboratories that serve hospitals, which are geographically and demographically representative for their country, and secure their participation. These laboratories were chosen form from those that regularly participate in the European Antimicrobial Resistance Surveillance System (EARSS). For six months, from September 2006 until February 2007, these participants were asked to submit the first 5 successive MSSA isolates and the first 5 successive MRSA isolates from individual patients with invasive infection. Isolates were dispatched by the participating laboratories to the SRLs accompanied by additional information, including the EARSS laboratory identifier, the sample number, the date of isolation, origin of clinical specimen, demographic detail, (such as age and gender), epidemiological context, defined as hospital-acquired when symptoms developed more than 48h after admission or as community onset otherwise, isoxacylpenicillin- or cefoxitin-resistance, all cause mortality 14 days after isolation of the first invasive isolate, and date when isolate was sent to the SRL. SRLs confirmed MRSA genotype by mecA PCR. Further genetic or phenotypic characterisation including multilocus sequence typing (MLST) the typing of staphylococcal cassette chromosome mec (SCCmec), identification of agr type and toxin genes such as lukF lukS gene indicative of Panton Valentine Leukocidin (PVL) expression was left to the SRLs own discretion. All SRLs preserved the isolates in their national strain collection and performed spa sequence typing according to the standard protocol, uploaded the sequence information and made this available by synchronisation with the central Ridom spa server curated by SeqNet.org at the Institute of Hygiene, University Hospital Muenster, Germany [13] [14] in order to apply a consistent spa nomenclature based on quality controlled sequence data. Epidemiological and typing data were communicated in parallel to a central SQL database at the National Institute for Public Health and the Environment of the Netherlands (RIVM). For each participating laboratory, SRLs also provided postal address or indicated their administrative region (such as region, province, state, department, or NUTS-2 level) if location of laboratories were to be aggregated on higher administrative geographical level (only Austria, Belgium, Czech Republic, and Poland). All data were anonymous and collected in accordance with the European Parliament and Council decision for the epidemiological surveillance and control of communicable disease in the European community [17, 18]. Ethical approval and informed consent were thus not obtained. Data analysis, geographical illustration and cluster identification All data were inspected for inconsistencies and analysed on a country by country basis and returned to SRLs for feedback, clarification of inconsistencies and final approval in February 2008. Additional molecular characteristics such as antibiotic susceptibilities and toxin profiles were still carried out and added to the databases until October 2008. After final approval, data were analysed using Stata"!, Version 8.0, (College Station, Texas, USA) and SAS"! version 9.1 (SAS Institute Inc., Cary, NC, USA) using conventional summary statistical approaches. Percentage diversity and 95% confidence intervals were calculated as described previously [15]. Postal address and location of all sampling laboratories were converted into decimal cartesian coordinates from the geocoding facility at  HYPERLINK "http://www.spatialepidemiology.net" www.spatialepidemiology.net [16]. Pairwise distances of laboratories that isolated MSSA and MRSA with identical spa types were calculated and distance matrices summarised and compared by Student-t test. The Web Application SRL-Maps (http://www.spatialepidemiology.net/SRL-Maps) was developed to interrogate the data based on mapping of laboratory locations and centroids of administrative regions (when laboratory results were aggregated on the level of administrative region) A spatial scan statistic was used to identify geographical abundance of specific spa types at two levels, (i) country-specific frequencies that take into account all spa types within national boundaries and (ii) regional clusters of varying size independent of national boundaries. To identify spa types with higher than expected occurrence in any of the participating countries, the observed number of all spa types isolated within each country was compared with the number expected under the assumption of a European-wide random distribution. For the identification of regional clusters, circular windows around all laboratory locations were projected. For each location, the radius of the window was varied from zero to 1000 kilometres. This way a finite number of distinct windows was created. For each window, the observed number of isolates with a specific spa type was compared with the expected number under the assumption of a random distribution. Ten thousand random distributions were obtained by varying the composition of local spa types at all laboratory locations consistent with cumulative spa-specific frequencies using Monte Carlo Simulations. The test statistic was calculated by log likelihood ratio test, whereby countries or windows where the observed spa type frequencies differed from those obtained by simulation were considered to contain significant clusters with an alpha error of p<0.0001. SaTScan software was developed by Martin Kulldorff [19, 20]. Results Summary statistics Twenty-six SRLs from as many countries satisfactorily fulfilled the proficiency criteria for spa sequence typing and contributed data for final analysis. Between September 2006 and February 2007, 357 laboratories serving 450 hospitals collected 2,890 successive MSSA and MRSA isolates from patients with invasive S. aureus infection (2603 from blood cultures, 90.4%; 17 from CSF, 0.6%; 270 from puncture fluids of other normally sterile anatomical sites, 9.3%). At the SRLs, the spa sequences were determined and synchronised with the spa server at SeqNet.org. Submitted sequences were quality controlled by comparison with accompanying chromatograms (trace files) and all fulfilled the stringent quality criteria for excellent sequencing data [13]. Spa typing information combined with all additional data was submitted to the central database at RIVM. Final inspection of data revealed missing information for gender (28 isolates, 1%), age (54 isolates, 1.9%), sampling dates (74 isolates, 2.6%), epidemiological context (community onset or hospital acquisition, 568 isolates 19.7%), all cause mortality 14 days after S. aureus isolation (1052 isolates, 36.4%) and spa type (62 isolates, 2.1%).  HYPERLINK \l "Table1" Table 1 gives a summary overview of the number of participating laboratories, isolates and spa types submitted by country. Many laboratories were unable to collect 5 invasive MRSA isolates within the sampling period due to a low MRSA incidence in the hospitals they serve. Therefore, the combined collection consisted of two third MSSA (1923, 66.5%) and one third MRSA (967, 33.5%,  HYPERLINK \l "Table2" Table 2). Patients with invasive MRSA infections were older ( HYPERLINK \l "Figure1" figure 1) and had a median age of 69-years compared to a median age of 63-years in MSSA patients (p<0.001). Moreover, MRSA patients had a higher all cause mortality (20.8%) 14 days after the first isolation of S. aureus than MSSA patients (13.2%, p<0.0001). More males (1765/2862, 61.7%) than females had invasive S. aureus infections. The proportion of MRSA vs. MSSA did not differ between the sexes (p=0.2). Of the 231 MRSA that were reported as community onset (CO-MRSA), 226 (97.8%) were tested for the presence of PVL associated genes lukS, lukF and 11 (4.9%) were found to be positive. Of the 585 hospital-acquired MRSA (HA-MRSA), 551 (94.2%) were tested for PVL and 6 (1.1%) were found to be positive. This difference was significant (p=0.001). Spa types A total of 658 spa types were reported ( HYPERLINK \l "Table2" Table 2). Among all spa types, 564 and 154 were assigned to MSSA and MRSA respectively of which, 504 consisted of only MSSA and 94 of only MRSA. Sixty spa types contained both MSSA and MRSA. Thirty-six of the MSSA (1.9%) and 26 of the MRSA (2.7%) were non-typeable.  HYPERLINK \l "Table3" Table 3 shows the rank order by frequencies of the most frequent spa types isolated during the survey. MRSA was less diverse than MSSA. Only, five spa types accounted for 50% of all MRSA isolates, whereas the same proportion of MSSA isolates was divided into 26 most frequent MSSA spa types. Estimates of the genetic diversity differed significantly with a diversity index for MSSA of 0.985 (CI95 0.983 0.987) and 0.940 (CI95 0.933 0.947) for MRSA. While MSSA diversity ranged between 1.0 in Iceland and 0.934 in Latvia (data not shown), MRSA diversity between countries was more heterogeneous ranging from 0.41 in Slovenia to 0.91 in Austria ( HYPERLINK \l "Figure2" Figure 2) indicative of the presence of few dominant MRSA spa types in several countries. MRSA showed a higher degree of geographical clustering as the average distance between laboratories that isolated the same spa type was significantly smaller than for MSSA ( HYPERLINK \l "Table2" Table 2), but we found no correlation (r = -0.09, p = 0.75) between genetic diversity of MRSA and overall proportion of MRSA among S. aureus blood stream infections at country level as reported to the EARSS database for 2007 [21]. Clustering of spa types at country and regional level In 18 countries, 22 different spa types were found with unexpected frequencies ( HYPERLINK \l "Table4" Table 4). Most (88.7%) of these isolates were MRSA. No more than two spa types clustered in any country. There was a frequent regional coincidence with respect to clusters in neighbouring countries sharing identical spa types. The Czech Republic and Germany shared spa type t003 (t003/ST225/SCCmecII), Bulgaria and Romania shared t030 (t030/ST239/ SCCmecIII), England and Ireland t032 (t032/ST22/ SCCmecIV), Italy and Croatia, although separated by the Adriatic Sea shared t041 (t041/ST228/ SCCmecIII) an iIberic strain t067 (t067/ST5&125/ SCCmecI&IV) first identified through this initiative dominates in Spain and is found in Southern France [22] . The notion of regional spread was supported by the cluster statistic that projected windows beyond national boundaries ( HYPERLINK \l "Table5" Table 5). These regional clusters differed considerably in the range between the observed frequencies of spa types and those expected under the assumption of an even European distribution. The degree of unexpectedness, which is an indication of the significance of each cluster, is expressed by the log likelihood ratio (LLR). The majority of regional clusters extended beyond national boundaries and consisted of MRSA (73.9%). The most significant cluster was identified in Spain and consisted of spa type t067 (t067/ST5,125/ SCCmecIV,  HYPERLINK \l "Figure3" Figure 3). A Northern Balkan/Adriatic cluster consisted of spa type t041 (t041/ST228/ SCCmecIII) and included Austria, Hungary, Slovenia, Croatia and Northern and Central Italy. Germany, the Czech Republic and Western-Poland were included in a large regional cluster of spa type t003 (t003/ST225/SCCmecII), which was geographically centred in Leipzig and had a radius of 389 km. The largest cluster in size (radius 930 km) as well as in numbers (115 isolates) consisted of spa type t008 (t008/ST8/SCCmecI&IV) and was centred in southern France. In the British Isles, t032(t032/ST22/SCCmecIV), known as epidemic MRSA (EMRSA) 15, was the dominant type and one of the most significant clusters. Two additional clusters of spa type t032 clusters, albeit less significant and much smaller, were located in the Brandenburg area around Berlin and in Central Hungary. A smaller cluster (but ranking on sixth position in terms of significance) was located in Flanders on the Belgium Dutch border and consisted of spa type t740(t740/ST45/SCCmecIV). The clusters with the smallest size included those submitted by single laboratories most likely reflecting local outbreaks. A single regional cluster consisted of MSSA only (spa type t230, ST45) was located in the east of Denmark, indicating that regional spread of S. aureus is not limited to MRSA alone. SRL-Maps The web application, SRL-Maps provides a community tool for the interrogation of the geographical distribution of different spa types. All laboratory and regional locations across Europe are represented as placemarks on a Google map. Clicking on a placemark displays all spa types identified at that location (and their frequency) along with the number (and number of locations) if found elsewhere. The European distribution of any spa type can then be displayed and placemarks on the map are colour coded based on whether the isolates at each location are all MSSA (green), all MRSA (red) or are a mix of MSSA & MRSA (yellow) showing the number of isolates inside the placemark. Graphical charts are displayed showing spa type-specific proportion of MRSA/MSSA, all cause mortality and the age distribution among cases ( HYPERLINK \l "Figure3" Figure 3). This method thus allows for the identification and mapping of strains with particular public health importance and further exploration by the scientific community is encouraged. Observations on specific spa types For both MSSA and MRSA isolates, there was no association between specific spa types and all cause mortality after 14 days, indicating that no spa type was associated with increased virulence. Spa type t041 (t041/ST228/ SCCmecIII) was the only spa type that showed a significant association with hospital acquisition (OR= 4.2, p<0.001). At the same time, it represented the third most frequent MRSA in the database after t032 (t032/ST22/ SCCmecIV) and t008 (t008/ST8/SCCmecI&IV,  HYPERLINK \l "Table3" Table 3,  HYPERLINK \l "Table5" Table 5) and formed the second most significant cluster. Of the 11 CO-MRSA isolates that were found to be PVL-positive, three could be assigned to spa type t044 (t044/ST80/SCCmecIV) the so called European community-acquired MRSA, and another three had spa type t008 (t008/ST8/SCCmecIV) and are indistinguishable form from the USA300 community-acquired MRSA. Two other PVL-positive CO-MRSA isolates had spa type t622, which is closely related to spa type t008 differing only by the loss of a single repeat unit. On three occasions MSSA spa type t171 (t171/ST123) was identified, twice in Ireland once in Sweden. Isolates with this spa type are usually often fusidic acid-resistant and frequently associated with outbreaks of skin infections [23]. MRSA belonging to the multilocus sequence type ST398 have recently emerged in several European countries and are regarded to be livestock-associated (LA-MRSA). Of all spa types associated with this clone only spa types t011, t034, t571, t1255, t2383 were identified on 12 occasions (1.3%) in 8 countries during this survey. None of these isolates displayed an MRSA phenotype or was found to harbour the mecA gene. Discussion The present survey represents the first cross sectional study set out to simultaneously identify the predominant strains causing invasive S. aureus infections in 26 countries of the European region. Except for Slovakia, Estonia, Lithuania and Luxemburg, all Member States of the European Union participated and achieved a country-wide coverage. In the run up to this study, a successful effort was undertaken to agree on a single molecular typing approach to scale up the typing capacity and improve quality assessment by introducing proficiency testing for staphylococcal reference typing laboratories which had the intention to participate in advanced S. aureus surveillance. This effort provided the basis for the execution of a mutually agreed protocol using a standardized sampling frame and a quality-controlled genetic typing approach [11, 13] based on the sequencing of the variable region of the S. aureus spa gene (spa typing) [24]. However, given the scope and ambition of this investigation it is not surprising that the study still suffered from several operational shortcomings. In order to keep the amount of work manageable for the participating SRLs, it was decided to include about twenty laboratories which should be demographically and geographically representative of each of the participating countries. Certainly, this number is arbitrary and cannot equally represent small and large countries in Europe alike. Laboratory enrolment based on population size would provide a more appropriate sampling strategy but would also impose a proportional and frequently unacceptable amount of work on SRLs in large countries if the sample size from small countries should remain meaningful. But even middle sized countries like Greece, Finland, Bulgaria, Romania, were unable to enrol the intended number of laboratories mainly for technical and logistic reasons. Naturally, the number of laboratories and isolates varied between countries and geo-demographical representativeness could still be improved. It was the intention to collect equal number of successive MSSA and MRSA in all laboratories during the six months sampling interval. This proved to be unrealistic, especially in countries where MRSA rates are low. As a result, Norway and Iceland could not provide any MRSA, whereas Sweden, Denmark and the Netherlands provided less then ten isolates each. Cyprus and Malta had only one participating laboratory, but since both provide the microbiological service for the totality of the respective island population (for Cyprus only The Republic of Cyprus), they were allowed to submit up to 20 isolates. But when taking these potential threats to validity into account, the simultaneous collection of 2890 isolates from patients with invasive S. aureus infection treated in 450 hospitals during a six-month study interval is unparalleled and remains unmatched by previous investigations, which have drawn their conclusions from convenience samples of predominantly MRSA collected by laboratories for different clinical or biological reasons. The current collection includes one third MRSA and thus over-represents the natural population of MRSA causing invasive disease which was 22% in the European Union in 2007 [21]. All isolates were collected from laboratories and hospitals participating in EARSS for which estimates of the overall catchment population is known. Thus, the present sample of hospitals catered for approximately 22 million people, totalling 4.4% of the citizens living in the European Union. Despite the above limitations, the sample provides a realistic insight into the epidemiology and genetic diversity of S. aureus strains causing invasive infection in Europe. The age distribution and mortality was consistent with the expected range [25, 26]. High frequencies were ascertained in the very young (infants and under 5-year-olds) and older age groups with males clearly more at risk for invasive infections than females. Patients with MRSA were older than patients with MSSA and were 2.4 times more likely to have their infection attributed to hospital acquisition. Invasive MRSA carried a higher all cause mortality after 14 days. This finding is most likely confounded by a host of variables which distinguish MRSA patients from MSSA patients in fundamental ways. Mortality was independent of spa type, indicating that this study did not identify any single spa type that stands out with respect to invasiveness, virulence or other factors that would increase the risk of fatal outcome after 14 days. While a cross-sectional study has its limitations to control for many of the above mentioned confounders and has a restricted ability to detect subtle differences in virulence properties between different clonal lineages, this sample provides an unbiased estimate on the frequencies of specific spa types which have been reported to cause outbreaks and have attracted considerable public health attention. CO-MRSA were more likely to contain genes coding for PVL-toxin genes, but only 6 of the PVL-positive CO-MRSA isolates, which made up 0.5% of the overall sample, had spa types consistent with the community-acquired MRSA clones most frequently reported in Europe (t008/ST8/SCCmecIV and t044/ST80/SCCmecIV) [27]. These values when compared to the overall numbers of MRSA are small but still warrant attention since systemic infections with PVL-positive CA-MRSA are rare. Livestock-associated spa types representing multilocus sequence type ST398 [28] made up 0.4% of the overall sample. However, none were methicillin resistant. With the decision to utilize spa sequence typing as a common platform to address the geographical abundance and clone structure of S. aureus, various potential threats to meaningful observations need to be taken into account. Firstly, the evolution of a single gene that expresses a surface exposed protein that interacts with the host immune system may be constrained by the collective immune responses of the host population. It is thus prone to various degrees of homoplasy caused by convergent or parallel evolution brought about by a high mutability of this chromosomal region. [29]. Likewise, a single sequence of approximately 280 bps under immune selection may be a weak indicator for the genetic background of a genome that is approximately 10,000x larger, even in a predominantly clonally evolving species such as S. aureus. The chromosomal location of the spa gene in the proximity of the origin of replication makes it prone to occasional mobilisation during DNA strand replication. Large chromosomal replacements have been described that can transpose the spa gene into a heterologous genetic background [30] confounding the conclusions about the clonal decent of isolates based on spa typing alone [31]. Finally, spa typing seems not discriminatory enough to sufficiently discern between MRSA isolates given the fact that only 5 spa types accounted for 50% of all 967 MRSA isolated during this survey. Although, spa typing has been advocated for outbreak investigation in hospitals [32] spa typing may have a limited discriminatory ability for single hospital epidemiological purposes especially in regions were few spa types are endemic. In a wider epidemiological and geographical context, however, sub-optimal resolution is less of a threat. The findings of the present study provide a good indication that homoplasy and large chromosomal replacement are not sufficiently frequent to dilute the molecular signature of clonal dissemination on a contingent geographical scale. Moreover, when comparing agr type, SCCmec type, toxin and susceptibility profiles within and between different spa types, a significant within spa type consistency and between spa type discordance supports the notion that spa typing indeed provides a useful marker for major clonal lineages [33].  Predominant spa types showed a wide geographical distribution. The average distance between the locations from which the same spa types were sampled was smaller for MRSA isolates than MSSA isolates, suggesting a higher spatial clustering of MRSA. Moreover, genetic diversity of invasive? MRSA differed considerably between countries and, finally, cluster statistics corroborated a fundamental difference between MSSA and MRSA with respect to regional dissemination. The majority of isolates that formed regional clusters were MRSA and all 15 major MRSA spa types (defined as more than 10 isolates in the database) were typically hospital-acquired, of which 13 occurred in clusters and no more than three clusters overlapped in the same region. Conversely, of all 27 major MSSA spa types were more often from community onset invasive disease, only five were part of significant clusters and one (t230) consisted of MSSA alone. Thus invasive MRSA clones in Europe have a predominantly regional abundance and so far it is unclear if clusters are rather static and more or less confined to geographic areas or reflect multiple waves, of polycentric MRSA epidemics that are spreading across the continent. To unravel the dynamic evolution of epidemic MRSA clearly requires repeated observations. Irrespective of the dynamics, there are predictable forces that determine the differential clonal abundance of MRSA vs. MSSA. The emergence of MRSA occurs by chance and is a rare event although new findings suggest that it is more frequent then originally anticipated [34, 29]. The limited number of clones that acquired the methicillin resistance determinant (SCCmec) are more likely to expand in hospitals where exposure to antibiotics is higher and transmission more frequent than in the community. At the same time, geographic diffusion of MRSA clones will be facilitated by repeated hospital admissions [35] and referrals of MRSA carriers who typically belong to an older and frailer segment of the population. MSSA clones prevail in less structured populations and are thus not equally exposed to the same extent of selection and genetic drift. The observed levels of diversity are thus more easily maintained by MSSA, which are disseminated by hosts with different travel patterns than MRSA carriers. In order to explore the geographical distribution of different spa types, and the genetic and phenotypic detail of isolates in the European sample, the reader is encouraged to consult the purpose-designed interactive web application, SRL-Maps, at http://www.spatialepidemiology.net/SRL-Maps/. 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Country No. LabsNo. HospitalsNo. isolatesMSSAMRSA*No. spa types MSSANo. spa types MRSANo. not typedProp. not typed (%)Austria184817412054701910.6Belgium222219510788652510.5Bulgaria88542925231100.0Croatia1111885038271366.8Cyprus1116978500.0Czech2020145945164900.0Denmark1430112108470232.7Finland552215714700.0France2323225114111752700.0Germany27271809882562010.6Greece33352015126617.1Hungary1013110664435921.8Iceland115505000.0Ireland22221698584552600.0Italy19191478067531500.0Latvia11124338520100.0Malta111531225320.0Netherlands1821204195998962.9Norway11205555037023.6Poland232317913247421400.0Portugal1212884840361300.0Romania101036251118300.0Slovenia111258481029323.4Spain212120411391571921.0Sweden2047200195590552.5UK1518131716050152216.8Total35745028901923967563153622.1 Table 1: Summary overview of participating laboratories, hospitals, number of invasive isolates MSSA and MRSA and spa types by country. * Note that number and MRSA do not reflect a prevalence or occurrence in particular countries as the protocol asked for submission of the first five isolates of each phenotype. n*MSSAMRSATotal/Overallp-valueFrequency (%)28901923 (66.5)967 (33.4)2890 (100%)-Median age (IQR)283663 (46 75)69 (55 78)66 (49 76)<0.0001Male gender (%)28621159 (60.8)606 (63.3)1765 (61.7)0.2Mortality after 14 days (%)1839153 (13.2)141(20.8)294 (16.0)<0.0001Hospital acquisition (%)2322 777 (51.6)585 (71.7)1362 (58.7)<0.0001No. spa types2828564154658**-No. not typable282836 (1.9)26 (2.7)62 (2.2)-Index of diversity (CI95)28280.985 (0.983 0.987)0.940 (0.933 0.947)0.983 (0.982 0.984)<0.05***Mean distance in kilometres between laboratories which isolated identical spa types (CI95)1614****1088.5 (1081.9 1095.1)886.6 (875.0 898.2)-<0.05*** Table 2: Summary statistics of S. aureus isolated in 26 European countries. * Number of isolates for which data were available. ** Total number of spa types includes 60 spa types that are both MSSA and MRSA. *** Deduced from non-overlapping 95% confidence intervals. **** Includes only MRSA and MSSA with more than 10 isolates per spa type. Figure 1: Age distribution and mortality of invasive S. aureus infections in Europe. Age divided in infants (-1) and bands of five years  MSSAMRSASpatypeST*FreqPercentCum. %SpatypeST*FreqPercentCum. %1t002ST-5, ST-231934.8%4.8%1t032ST-2214014.5%14.4%2t084ST-15, ST-18894.6%9.5%2t008ST-8, ST-247, ST-250, ST-25412012.4%26.8%3t015ST-45844.4%13.8%3t041ST-111, ST-228727.4%34.2%4t091ST-7824.3%18.1%4t003ST-5, ST-225717.3%41.6%5t012ST-30774.0%22.1%5t002ST-5, ST-231626.4%48.1%6t127ST-1573.0%25.1%6t067ST-5, ST-125505.2%53.3%7t008ST-8, ST-247, ST-250, ST-254552.9%27.9%7t001ST-5, ST-222, ST-228303.1%56.4%8t021ST-30, ST-33, ST-55492.5%30.5%8t037ST-239, ST-240, ST-241272.8%59.2%9t005ST-22, ST-23, ST-60422.2%32.7%9t030ST-239, ST-246202.1%61.2%10t026ST-45, ST-47271.4%34.1%10t024ST-8141.4%62.7%11t065ST-45, ST-46261.4%35.4%11t190ST-8141.4%64.1%12t160ST-12, ST-13261.4%36.8%12t515ST-22121.3%65.5%13t056ST-101251.3%38.1%13t038ST-45121.2%66.7%14t050ST-45211.1%39.2%14t022ST-22111.1%67.8%15t078ST-26211.1%40.2%15t740ST-45111.1%69.0%16t164ST-20191.0%41.2%16t012ST-3090.9%69.9%17t346ST-15, ST-620 180.9%42.2%17t015ST-4590.9%70.8%18t024ST-8170.9%43.1%18t044ST-8090.9%71.8%19t230ST-45170.9%43.9%19t045ST-5, ST-22580.8%72.6%20t166ST-34160.8%44.8%20t127ST-180.8%73.4%other106255.2%100.0%other25826.6%100.0%Total1923100Total967100 Table 3: Twenty most frequent spa types and their sequence types (ST) among MSSA and MRSA isolated in 26 European countries. * ST = Multilocus sequence type as determined by SRLs (bold face) or derived from the SeqNet.org Spa typing data base.  Figure 2: Estimates of country-specific MRSA genetic diversity expressed by Simpsons index of diversity (diamonds) and 95% confidence intervals (bars) Country cluster No.Countryspa typeST*No. labs reporting clustered spa typeNo. labs particpating in surveyProp. labs reporting clustered spa type (percent)No. isolates observed No. isolates expected **No. MRSA among observed isolatesProp. MRSA among observed isolates (percent)1Austriat190871838.9110.91091.02Belgiumt7404562227.3151.011173.33Belgiumt0384582236.4120.8112100.04Bulgariat0302394850.0100.41990.05Croatiat04122871163.6142.1914100.06Czecht003225162080.0343.7634100.07Czecht13032015.050.45100.08Denmarkt2304571450.080.700.09Francet0088222395.76113.625488.510Francet777562326.190.7888.911Germanyt003225102737.0244.672395.812Germanyt0322292733.3298.8429100.013Greecet0448033100.050.13480.014Hungaryt062521020.060.346100.015Hungaryt2165971070.070.537100.016Irelandt03222182281.8388.338100.017Irelandt5152272231.880.78100.018Italyt041228131968.4233.6923100.019Italyt00122881942.191.549100.020Latviat43542741136.480.218100.021Latviat42536831127.350.075100.022Polandt037239112347.8212.171466.723Polandt127162326.1173.9617100.024Romaniat03023931030.060.276100.025Spaint0675, 125182185.7534.454381.126Spaint0025152171.42810.941450.027United Kingdomt03222121580.0286.432796.4total23547549.549481.7443888.7 Table 4: Overabundance of spa types at country level. *As determined by SRLs. **Average number of isolates with this spa type expected in country after 10,000 simulations on the basis of European-wide cumulative frequency. Regional cluster No.spa type ST*Window centerWindow radius (km)No. labs within window reporting clustered spa typeNo. isolates observedNo. isolates expected **Log likelihood ratioNo. MRSA among observed isolatesProp. MRSA among observed isolates (percent)1t0675, 125Alicante, Spain716.2219554.82124.894683.62t041228Split, Croatia521.93255910.5684.7659100.03t003225Leipzig, Germany386.08225810.0982.285493.14t0088Perpignan, France930.645311945.4672.8510588.25t03222Letterkenny, Ireland712.75216614.7465.226598.56t74045Goes, Netherlands81.146150.5650.361173.37t030239Lovetch, Bulgaria362.984160.6545.251593.88t037239Plock, Poland329.511212.1736.871885.79t03845Antwerpen, Belgium94.239120.8132.7112100.010t1908Krems, Austria62.855100.3329.63990.011t001228Sibenik, Coratia884.57152910.2227.5329100.012t435427Daugavpils, Latvia188.62380.1925.8200.013t03222Berlin, Germany128.184192.8523.5619100.014t425368Riga, Latvia0150.0622.765100.015t1271Inowroclaw, Poland131.324120.9122.6500.016t01545Lomza, Poland581.138351121.43720.017t777Laval, France277.14470.2520.617100.018t0917Ried, Austria770.67517042.2820.1445.719t51522Mullingar, Ireland196.22590.7218.949100.020t08125Makw Mazowiecki, Poland0140.0218.601375.021t0025, 231Coimbra, Portugal484.11143310.6218.372060.622t23045Hilleroed, Denmark234.559111.3617.1900.023t04480Nicosia, Cyprus916.16260.1917.096100.024t2054Clamart, France233.97450.1816.65360.025t03222Kecskemt, Hungary82.064163.0515.7816100.026t0625Szolnok, Hungary 144.91260.2915.246100.027t0025, 231Reims, France186.814268.2614.391350.0total350732182.6454173.9 Table 5: Cluster of spa types at regional level. *As determined by SRLs. **Average number of isolates with this spa type expected in window after 10,000 simulations on the basis of European-wide cumulative frequency Figure 3: Location of laboratories isolating S. aureus spa type t067. Based on the display of isolates from LAB ES056 in Madrid, Spain, the distribution of all other t067 isolates are shown (n=62). Each placemark indicates whether isolates are MSSA (green), MRSA(red) or a mixture (yellow) and the selected laboratory is blue. The number of isolates are indicated inside the placemark. The pie charts on the right show proportion MRSA/MSSA, all cause mortality after 14 days and the bar chart displays patient age distribution.      PAGE  PAGE 19 PAGE  PAGE 8 Is it discussed to what extent this was achieved PAGE \# "'Side: '#' '" I do not believe that our paper has data on spread of MRSA in the community. Robert: I agree with this view Suggest: relative few Commas around however? and? With? As the isolates both harbour SCCmec I and IV it makes it less likely that it is one cluster this is one example underlining the comments by Wolfgang Looking at the distribution of the isolates to the geography in Denmark again underline the points from Wolgang In my opinion isolates with different SCCmec types should not a priori be considered representing the same cluster Is itt really of common interest are these fusidic acid resistant? I read this as if it was surprising that not more different types were found I actually find it impressing that 8 different types were found among 12 cases The distribution as not especially representative for at least Italy and possibly also other countries I suggest it is modified se also line 22-25 This seems too long Not typed or non-tapable or a mixture of both? PAGE \# "'Side: '#' '" This should be rounded to whole numbers. Either you have an isolate or you dont. 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