Tn1 transposition in the course of natural transformation enables horizontal antibiotic resistance spread in Acinetobacter baylyi

Transposons are genetic elements that change their intracellular genomic position by transposition and are spread horizontally between bacteria when located on plasmids. It was recently discovered that transposition from fully heterologous DNA also occurs in the course of natural transformation. Here, we characterize the molecular details and constraints of this process using the replicative transposon Tn1 and the naturally competent bacterium Acinetobacter baylyi . We find that chromosomal insertion of Tn1 by transposition occurs at low but detectable frequencies and preferably around the A. baylyi terminus of replication. We show that Tn1 transposition is facilitated by transient expression of the transposase and resolvase encoded by the donor DNA. RecA protein is essential for the formation of a circular, double-stranded cytoplasmic intermediate from incoming donor DNA, and RecO is beneficial but not essential in this process. Absence of the recipient RecBCD nuclease stabilizes the double-stranded intermediate. Based on these results, we suggest a mechanistic model for transposition during natural transformation.

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Prevalence of Mycoplasma genitalium fluoroquinolone-resistance markers, and dual-class-resistance markers, in asymptomatic men who have sex with men

Journal of Medical Microbiology ◽

10.1099/jmm.0.001429 ◽

2021 ◽

Vol 70 (9) ◽

Author(s):

Teck-Phui Chua ◽

Kaveesha Bodiyabadu ◽

Dorothy A. Machalek ◽

Suzanne M. Garland ◽

Catriona S. Bradshaw ◽

...

Keyword(s):

Antibiotic Resistance ◽

Type Species ◽

Health Centre ◽

Mycoplasma Genitalium ◽

Fluoroquinolone Resistance ◽

Content Type ◽

Link Type ◽

Resistance Markers ◽

Sex With Men ◽

Dual Class

Introduction. Failure of fluoroquinolones, the principal treatment option for macrolide-resistant Mycoplasma genitalium infections, has recently emerged. This is of particular concern for men who have sex with men (MSM), who have high proportions of macrolide-resistant M. genitalium infections. Treatment failure with moxifloxacin is likely the result of single nucleotide polymorphisms (SNPs) in parC, whilst concurrent gyrA mutations may play a role. Gap Statement. The levels of fluoroquinolone resistance and dual-class (i.e. macrolide and fluoroquinolone) resistance in M. genitalium among asymptomatic MSM is unknown. Aim. To (i) determine the proportion of fluoroquinolone resistance and dual-class resistance in M. genitalium infections among asymptomatic MSM, (ii) explore any clinical and behavioural associations with fluoroquinolone resistance, and (iii) determine the distribution of antibiotic resistance among M. genitalium mgpB sequence types (STs). Methodology. M. genitalium positive samples (N=94) were obtained from 1001 asymptomatic MSM enrolled in a study at Melbourne Sexual Health Centre (Carlton, Australia) between August 2016 and September 2017. Sanger sequencing was performed to determine the proportion of M. genitalium infections with SNPs in parC that have previously been associated with failure of moxifloxacin (corresponding to amino changes S83I, D83R, D87Y and D87N) and in gyrA (corresponding to amino acid changes M95I, D99N, D99Y and D99G). Associations between clinical/behavioural factors and parC SNPs were examined. Strain typing was performed by sequencing a portion of the mgpB gene. Results. The proportion of MSM with infections harbouring parC and gyrA SNPs was 13.0 % [95 % confidence interval (CI): 6.8–23.2 %] and 4.7 % (95 % CI: 1.1–13.4 %), respectively; dual-class resistance was 13.0 %. No significant clinical/behavioural associations were found. Antibiotic resistance was not restricted to specific mgpB STs. Conclusion. One in eight (13 %) of asymptomatic MSM with M. genitalium had an infection with dual-class-resistance mutations. Typing by mgpB sequence suggested fluoroquinolone resistance is arising from independent mutation events. This study illustrates that asymptomatic MSM may act as a reservoir for antibiotic-resistant M. genitalium .

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bla OXA-48-like genome architecture among carbapenemase-producing Escherichia coli and Klebsiella pneumoniae in the Netherlands

Microbial Genomics ◽

10.1099/mgen.0.000512 ◽

2021 ◽

Vol 7 (5) ◽

Author(s):

Antoni P. A. Hendrickx ◽

Fabian Landman ◽

Angela de Haan ◽

Sandra Witteveen ◽

Marga G. van Santen-Verheuvel ◽

...

Keyword(s):

Escherichia Coli ◽

Antibiotic Resistance ◽

The Netherlands ◽

Resistance Genes ◽

Type Species ◽

Antibiotic Resistance Genes ◽

Gene Content ◽

Content Type ◽

E Coli ◽

Link Type

Carbapenem-hydrolysing enzymes belonging to the OXA-48-like group are encoded by bla OXA-48-like alleles and are abundant among Enterobacterales in the Netherlands. Therefore, the objective here was to investigate the characteristics, gene content and diversity of the bla OXA-48-like carrying plasmids and chromosomes of Escherichia coli and Klebsiella pneumoniae collected in the Dutch national surveillance from 2014 to 2019 in comparison with genome sequences from 29 countries. A combination of short-read genome sequencing with long-read sequencing enabled the reconstruction of 47 and 132 complete bla OXA-48-like plasmids for E. coli and K. pneumoniae , respectively. Seven distinct plasmid groups designated as pOXA-48-1 to pOXA-48-5, pOXA-181 and pOXA-232 were identified in the Netherlands which were similar to internationally reported plasmids obtained from countries from North and South America, Europe, Asia and Oceania. The seven plasmid groups varied in size, G+C content, presence of antibiotic resistance genes, replicon family and gene content. The pOXA-48-1 to pOXA-48-5 plasmids were variable, and the pOXA-181 and pOXA-232 plasmids were conserved. The pOXA-48-1, pOXA-48-2, pOXA-48-3 and pOXA-48-5 groups contained a putative conjugation system, but this was absent in the pOXA-48-4, pOXA-181 and pOXA-232 plasmid groups. pOXA-48 plasmids contained the PemI antitoxin, while the pOXA-181 and pOXA-232 plasmids did not. Furthermore, the pOXA-181 plasmids carried a virB2-virB3-virB9-virB10-virB11 type IV secretion system, while the pOXA-48 plasmids and pOXA-232 lacked this system. A group of non-related pOXA-48 plasmids from the Netherlands contained different resistance genes, non-IncL-type replicons or no replicons. Whole genome multilocus sequence typing revealed that the bla OXA-48-like plasmids were found in a wide variety of genetic backgrounds in contrast to chromosomally encoded bla OXA-48-like alleles. Chromosomally localized bla OXA-48 and bla OXA-244 alleles were located on genetic elements of variable sizes and comprised regions of pOXA-48 plasmids. The bla OXA-48-like genetic element was flanked by a direct repeat upstream of IS1R, and was found at multiple locations in the chromosomes of E. coli . Lastly, K. pneumoniae isolates carrying bla OXA-48 or bla OXA-232 were mostly resistant for meropenem, whereas E. coli bla OXA-48, bla OXA-181 and chromosomal bla OXA-48 or bla OXA-244 isolates were mostly sensitive. In conclusion, the overall bla OXA-48-like plasmid population in the Netherlands is conserved and similar to that reported for other countries, confirming global dissemination of bla OXA-48-like plasmids. Variations in size, presence of antibiotic resistance genes and gene content impacted pOXA-48, pOXA-181 and pOXA-232 plasmid architecture.

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Phylum barrier and Escherichia coli intra-species phylogeny drive the acquisition of antibiotic-resistance genes

Microbial Genomics ◽

10.1099/mgen.0.000489 ◽

2021 ◽

Vol 7 (8) ◽

Author(s):

Marie Petitjean ◽

Bénédicte Condamine ◽

Charles Burdet ◽

Erick Denamur ◽

Etienne Ruppé

Keyword(s):

Escherichia Coli ◽

Antibiotic Resistance ◽

Resistance Genes ◽

Type Species ◽

Antibiotic Resistance Genes ◽

Resistance Pattern ◽

Content Type ◽

E Coli ◽

Link Type ◽

Specific Distribution

Escherichia coli is a ubiquitous bacterium that has been widely exposed to antibiotics over the last 70 years. It has adapted by acquiring different antibiotic-resistance genes (ARGs), the census of which we aim to characterize here. To do so, we analysed 70 301 E. coli genomes obtained from the EnteroBase database and detected 1 027 651 ARGs using the AMRFinder, Mustard and ResfinderFG ARG databases. We observed a strong phylogroup and clonal lineage specific distribution of some ARGs, supporting the argument for epistasis between ARGs and the strain genetic background. However, each phylogroup had ARGs conferring a similar antibiotic class resistance pattern, indicating phenotypic adaptive convergence. The G+C content or the type of ARG was not associated with the frequency of the ARG in the database. In addition, we identified ARGs from anaerobic, non- Proteobacteria bacteria in four genomes of E. coli , supporting the hypothesis that the transfer between anaerobic bacteria and E. coli can spontaneously occur but remains exceptional. In conclusion, we showed that phylum barrier and intra-species phylogenetic history are major drivers of the acquisition of a resistome in E. coli .

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Horizontal Gene Transfer of Antibiotic Resistance from Acinetobacter baylyi to Escherichia coli on Lettuce and Subsequent Antibiotic Resistance Transmission to the Gut Microbiome

mSphere ◽

10.1128/msphere.00329-20 ◽

2020 ◽

Vol 5 (3) ◽

Cited By ~ 2

Author(s):

Marlène Maeusli ◽

Bosul Lee ◽

Sarah Miller ◽

Zeferino Reyna ◽

Peggy Lu ◽

...

Keyword(s):

Antibiotic Resistance ◽

Gene Transfer ◽

Horizontal Gene Transfer ◽

Resistance Genes ◽

Gut Microbiome ◽

Acinetobacter Baylyi ◽

Antibiotic Resistant ◽

Content Type ◽

Plant Foods ◽

Link Type

ABSTRACT Agricultural use of antibiotics is recognized by the U.S. Centers for Disease Control and Prevention as a major contributor to antibiotic-resistant infections. While most One Health attention has been on the potential for antibiotic resistance transmission from livestock and contaminated meat products to people, plant foods are fundamental to the food chain for meat eaters and vegetarians alike. We hypothesized that environmental bacteria that colonize plant foods may serve as platforms for the persistence of antibiotic-resistant bacteria and for horizontal gene transfer of antibiotic-resistant genes. Donor Acinetobacter baylyi and recipient Escherichia coli were cocultured in vitro, in planta on lettuce, and in vivo in BALB/c mice. We showed that nonpathogenic, environmental A. baylyi is capable of transferring plasmids conferring antibiotic resistance to E. coli clinical isolates on lettuce leaf discs. Furthermore, transformant E. coli from the in planta assay could then colonize the mouse gut microbiome. The target antibiotic resistance plasmid was identified in mouse feces up to 5 days postinfection. We specifically identified in vivo transfer of the plasmid to resident Klebsiella pneumoniae in the mouse gut. Our findings highlight the potential for environmental bacteria exposed to antibiotics to transmit resistance genes to mammalian pathogens during ingestion of leafy greens. IMPORTANCE Previous efforts have correlated antibiotic-fed livestock and meat products with respective antibiotic resistance genes, but virtually no research has been conducted on the transmission of antibiotic resistance from plant foods to the mammalian gut (C. S. Hölzel, J. L. Tetens, and K. Schwaiger, Pathog Dis 15:671–688, 2018, https://doi.org/10.1089/fpd.2018.2501; C. M. Liu et al., mBio 9:e00470-19, 2018, https://doi.org/10.1128/mBio.00470-18; B. Spellberg et al., NAM Perspectives, 2016, https://doi.org/10.31478/201606d; J. O’Neill, Antimicrobials in agriculture and the environment, 2015; Centers for Disease Control and Prevention, Antibiotic resistance threats in the United States, 2019). Here, we sought to determine if horizontal transmission of antibiotic resistance genes can occur between lettuce and the mammalian gut microbiome, using a mouse model. Furthermore, we have created a new model to study horizontal gene transfer on lettuce leaves using an antibiotic-resistant transformant of A. baylyi (AbzeoR).

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A pentaplex real-time PCR assay for rapid identification of major beta-lactamase genes KPC, NDM, CTX, CMY, and OXA-48 directly from bacteria in blood

Journal of Medical Microbiology ◽

10.1099/jmm.0.001465 ◽

2021 ◽

Vol 70 (12) ◽

Author(s):

Taalin R. Hoj ◽

Bradley McNeely ◽

Kylie Webber ◽

Evelyn Welling ◽

William G. Pitt ◽

...

Keyword(s):

Antibiotic Resistance ◽

Klebsiella Pneumoniae ◽

Real Time ◽

Real Time Pcr ◽

Type Species ◽

New Delhi ◽

Beta Lactamase ◽

Content Type ◽

Link Type ◽

Carbapenem Resistant

Introduction. Antibiotic resistance, particularly in cases of sepsis, has emerged as a growing global public health concern and economic burden. Current methods of blood culture and antimicrobial susceptibility testing of agents involved in sepsis can take as long as 3–5 days. It is vital to rapidly identify which antimicrobials can be used to effectively treat sepsis cases on an individual basis. Here, we present a pentaplex, real-time PCR-based assay that can quickly identify the most common beta-lactamase genes ( Klebsiella pneumoniae carbapenemase (KPC); New Delhi metallo-beta-lactamase (NDM); cefotaximase-Munich (CTX-M); cephamycin AmpC beta-lactamases (CMY); and Oxacillinase-48 (OXA-48)) from pathogens derived directly from the blood of patients presenting with bacterial septicemia. Aim. To develop an assay which can rapidly identify the most common beta-lactamase genes in Carbapenem-resistant Enterobacteriaceae bacteria (CREs) from the United States. Hypothesis/Gap Statement. Septicemia caused by carbapenem-resistant bacteria has a death rate of 40–60 %. Rapid diagnosis of antibiotic susceptibility directly from bacteria in blood by identification of beta-lactamase genes will greatly improve survival rates. In this work, we develop an assay capable of concurrently identifying the five most common beta-lactamase and carbapenemase genes. Methodology. Primers and probes were created which can identify all subtypes of Klebsiella pneumoniae carbapenemase (KPC); New Delhi metallo-beta-lactamase (NDM); cefotaximase-Munich (CTX); cephamycin AmpC beta-lactamase (CMY); and oxacillinase-48 (OXA-48). The assay was validated using 13 isolates containing various PCR targets from the Centre for Disease Control Antimicrobial Resistance Isolate Bank Enterobacterales Carbapenemase Diversity Panel. Blood obtained from volunteers was spiked with CREs and bacteria were separated, lysed, and subjected to analysis via the pentaplex assay. Results. This pentaplex assay successfully identified beta-lactamase genes derived from bacteria separated from blood at concentrations of 4–8 c.f.u. ml−1. Conclusion. This assay will improve patient outcomes by supplying physicians with critical drug resistance information within 2 h of septicemia onset, allowing them to prescribe effective antimicrobials corresponding to the resistance gene(s) present in the pathogen. In addition, information supplied by this assay will lessen the inappropriate use of broad-spectrum antimicrobials and prevent the evolution of further antibiotic resistance.

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Prevalence and antibiotic resistance of Escherichia coli O157:H7 in beef at a commercial slaughterhouse in Moro, Kwara State, Nigeria

Access Microbiology ◽

10.1099/acmi.0.000289 ◽

2021 ◽

Vol 3 (11) ◽

Author(s):

Busayo I. Ajuwon ◽

Sola K. Babatunde ◽

Olatunji M. Kolawole ◽

Adeyinka E. Ajiboye ◽

Abosede H. Lawal

Keyword(s):

Escherichia Coli ◽

Antibiotic Resistance ◽

Type Species ◽

Diffusion Method ◽

Foodborne Disease ◽

Production Chain ◽

Content Type ◽

E Coli ◽

Link Type ◽

Caecal Content

Background. Gastroenteritis due to foodborne disease is a leading cause of death in developing countries. In Nigeria, there is an increasing demand for beef. Yet, there is no surveillance for Escherichia coli O157:H7 contamination of raw beef and little is known about the carriage of this pathogen in Nigeria’s livestock. Methods. A total of 415 samples, including 180 cow carcass swabs, 180 caecal content samples, 16 water samples, 25 hand swabs and 14 knife swabs were collected at a large abattoir in the Moro region of Kwara State, Nigeria. The samples were enriched in modified tryptone broth containing novobiocine, and plated onto Sorbitol–MacConkey agar (Oxoid SR0172E) supplemented with 0.05 mg l−1 cefixime and 2.5 mg l−1 potassium tellurite (Oxoid) (CT-SMAC). Indole-producing isolates were confirmed serologically by serotyping with antisera specific for the O157 and H7 antigens. The E. coli O157:H7 isolates were further tested for their susceptibility to antibiotic agents using the disc diffusion method. Commercially available Gram-negative multi-discs (Oxoid) comprising nitrofurantoin (30 µg), ampicillin (5 µg), ceftazidime (30 µg), gentamicin (10 µg), ciprofloxacin (5 µg), augmentin (30 µg), ofloxacin (5 µg) and cefuroxime (30 µg) were tested. Results. Overall, 16 (3.9 %) samples were contaminated with E. coli O157:H7, of which 10 (5.6 %) were isolated from carcass swabs, 4 (2.2 %) from caecal content samples and 2 (12.5 %) from water. All isolates were multidrug-resistant (MDR), with resistance to ampicillin, ceftazidime and cefuroxime being the most common. Conclusion. This study provides evidence to suggest that E. coli O157:H7 exists in the beef production chain. The pathogen reveals a high frequency of multidrug resistance, suggesting that consumers and handlers of such meat are at risk of contracting antibiotic-resistant E. coli O157:H7-associated foodborne disease. Routine monitoring of antibiotic resistance is critical to uncovering novel therapeutic strategies that will help inform clinical practice guidelines.

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Reversal of heavy metal-induced antibiotic resistance by dandelion root extracts and taraxasterol

Journal of Medical Microbiology ◽

10.1099/jmm.0.001226 ◽

2020 ◽

Vol 69 (8) ◽

pp. 1049-1061

Author(s):

Kerry Yang ◽

Yanjie Zhang

Keyword(s):

Oxidative Stress ◽

Heavy Metals ◽

Heavy Metal ◽

Antibiotic Resistance ◽

Bacterial Growth ◽

Type Species ◽

Root Extract ◽

Content Type ◽

Root Extracts ◽

Link Type

Introduction. Metal exposure is an important factor for inducing antibiotic resistance in bacteria. Dandelion extracts have been used for centuries in traditional Chinese and Native American medicine. Aim. We assessed the effects of dandelion water extracts and taraxasterol on heavy metal-induced antibiotic resistance in Escherichia coli as well as the underlying mechanisms. Methodology. Dandelion extracts were obtained through 4 h of boiling in distilled water. Bacterial growth was monitored with a spectrophotometer. Biochemical assays were performed to assess the activities and gene transcriptions of β-lactamase and acetyltransferase. Oxidative stress was determined using an oxidation-sensitive probe, H2DCFDA. Results. The present study demonstrated that higher concentrations of nickel (>5 µg ml−1), cadmium (>0.1 µg ml−1), arsenic (>0.1 µg ml−1) and copper (>5 µg ml−1) significantly inhibited the growth of E. coli . Lower concentrations of nickel (0.5 µg ml−1), cadmium (0.05 µg ml−1) and arsenic (0.05 µg ml−1) had no effect on bacterial growth, but helped the bacteria become resistant to two antibiotics, kanamycin and ampicillin. The addition of dandelion root extracts and taraxasterol significantly reversed the antibiotic resistance induced by these heavy metals. The supplements of antibiotics and cadmium generated synergistic effects on the activities of β-lactamase and acetyltransferase (two antibiotic resistance-related proteins), which were significantly blocked by either dandelion root extract or taraxasterol. In contrast, oxidative stress was not involved in the preventative roles of dandelion root extracts and taraxasterol in heavy metal-induced antibiotic resistance. Conclusion. This study suggests that heavy metals induce bacterial antibiotic resistance and dandelion root extracts and taraxasterol could be used to help reverse bacterial resistance to antibiotics.

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Genome evolution drives transcriptomic and phenotypic adaptation in Pseudomonas aeruginosa during 20 years of infection

Microbial Genomics ◽

10.1099/mgen.0.000681 ◽

2021 ◽

Vol 7 (11) ◽

Author(s):

Samuel J.T. Wardell ◽

Jeff Gauthier ◽

Lois W. Martin ◽

Marianne Potvin ◽

Ben Brockway ◽

...

Keyword(s):

Pseudomonas Aeruginosa ◽

Antibiotic Resistance ◽

Chronic Infection ◽

Type Species ◽

Average Rate ◽

Opportunistic Pathogen ◽

Altered Expression ◽

Content Type ◽

Link Type ◽

Long Read

The opportunistic pathogen Pseudomonas aeruginosa chronically infects the lungs of patients with cystic fibrosis (CF). During infection the bacteria evolve and adapt to the lung environment. Here we use genomic, transcriptomic and phenotypic approaches to compare multiple isolates of P. aeruginosa collected more than 20 years apart during a chronic infection in a CF patient. Complete genome sequencing of the isolates, using short- and long-read technologies, showed that a genetic bottleneck occurred during infection and was followed by diversification of the bacteria. A 125 kb deletion, an 0.9 Mb inversion and hundreds of smaller mutations occurred during evolution of the bacteria in the lung, with an average rate of 17 mutations per year. Many of the mutated genes are associated with infection or antibiotic resistance. RNA sequencing was used to compare the transcriptomes of an earlier and a later isolate. Substantial reprogramming of the transcriptional network had occurred, affecting multiple genes that contribute to continuing infection. Changes included greatly reduced expression of flagellar machinery and increased expression of genes for nutrient acquisition and biofilm formation, as well as altered expression of a large number of genes of unknown function. Phenotypic studies showed that most later isolates had increased cell adherence and antibiotic resistance, reduced motility, and reduced production of pyoverdine (an iron-scavenging siderophore), consistent with genomic and transcriptomic data. The approach of integrating genomic, transcriptomic and phenotypic analyses reveals, and helps to explain, the plethora of changes that P. aeruginosa undergoes to enable it to adapt to the environment of the CF lung during a chronic infection.

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Pathogenicity and drug resistance of animal streptococci responsible for human infections

Journal of Medical Microbiology ◽

10.1099/jmm.0.001339 ◽

2021 ◽

Vol 70 (3) ◽

Author(s):

Paulina Glajzner ◽

Eligia Maria Szewczyk ◽

Magdalena Szemraj

Keyword(s):

Antibiotic Resistance ◽

Type Species ◽

Streptococcus Bovis ◽

Content Type ◽

Streptococcus Species ◽

Link Type ◽

Animal Pathogens ◽

Streptococcal Species ◽

New Hosts ◽

Human Infections

Bacteria of the genus Streptococcus , earlier considered typically animal, currently have also been causing infections in humans. It is necessary to make clinicians aware of the emergence of new species that may cause the development of human diseases. There is an increasing frequency of isolation of streptococci such as S. suis , S. dysgalactiae , S. iniae and S. equi from people. Isolation of Streptococcus bovis/Streptococcus equinus complex bacteria has also been reported. The streptococcal species described in this review are gaining new properties and virulence factors by which they can thrive in new environments. It shows the potential of these bacteria to changes in the genome and the settlement of new hosts. Information is presented on clinical cases that concern streptococcus species belonging to the groups Bovis, Pyogenic and Suis. We also present the antibiotic resistance profiles of these bacteria. The emerging resistance to β-lactams has been reported. In this review, the classification, clinical characteristics and antibiotic resistance of groups and species of streptococci considered as animal pathogens are summarized.

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Pseudomonas aeruginosa is capable of natural transformation in biofilms

Microbiology ◽

10.1099/mic.0.000956 ◽

2020 ◽

Vol 166 (10) ◽

pp. 995-1003 ◽

Cited By ~ 1

Author(s):

Laura M. Nolan ◽

Lynne Turnbull ◽

Marilyn Katrib ◽

Sarah R. Osvath ◽

Davide Losa ◽

...

Keyword(s):

Pseudomonas Aeruginosa ◽

Type Species ◽

Natural Transformation ◽

Bacterial Species ◽

Extracellular Dna ◽

Virulence Determinants ◽

Bacterial Populations ◽

Exogenous Dna ◽

Content Type ◽

Link Type

Natural transformation is a mechanism that enables competent bacteria to acquire naked, exogenous DNA from the environment. It is a key process that facilitates the dissemination of antibiotic resistance and virulence determinants throughout bacterial populations. Pseudomonas aeruginosa is an opportunistic Gram-negative pathogen that produces large quantities of extracellular DNA (eDNA) that is required for biofilm formation. P. aeruginosa has a remarkable level of genome plasticity and diversity that suggests a high degree of horizontal gene transfer and recombination but is thought to be incapable of natural transformation. Here we show that P. aeruginosa possesses homologues of all proteins known to be involved in natural transformation in other bacterial species. We found that P. aeruginosa in biofilms is competent for natural transformation of both genomic and plasmid DNA. Furthermore, we demonstrate that type-IV pili (T4P) facilitate but are not absolutely essential for natural transformation in P. aeruginosa .

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