Three Substrains of the CyanobacteriumAnabaenasp. Strain PCC 7120 Display Divergence in Genomic Sequences andhetCFunction

ABSTRACTAnabaenasp. strain PCC 7120 is a model strain for molecular studies of cell differentiation and patterning in heterocyst-forming cyanobacteria. Subtle differences in heterocyst development have been noticed in different laboratories working on the same organism. In this study, 360 mutations, including single nucleotide polymorphisms (SNPs), small insertion/deletions (indels; 1 to 3 bp), fragment deletions, and transpositions, were identified in the genomes of three substrains. Heterogeneous/heterozygous bases were also identified due to the polyploidy nature of the genome and the multicellular morphology but could be completely segregated when plated after filament fragmentation by sonication.hetCis a gene upregulated in developing cells during heterocyst formation inAnabaenasp. strain PCC 7120 and found in approximately half of other heterocyst-forming cyanobacteria. Inactivation ofhetCin 3 substrains ofAnabaenasp. PCC 7120 led to different phenotypes: the formation of heterocysts, differentiating cells that keep dividing, or the presence of both heterocysts and dividing differentiating cells. The expression of PhetZ-gfpin thesehetCmutants also showed different patterns of green fluorescent protein (GFP) fluorescence. Thus, the function ofhetCis influenced by the genomic background and epistasis and constitutes an example of evolution under way.IMPORTANCEOur knowledge about the molecular genetics of heterocyst formation, an important cell differentiation process for global N2fixation, is mostly based on studies withAnabaenasp. strain PCC 7120. Here, we show that rapid microevolution is under way in this strain, leading to phenotypic variations for certain genes related to heterocyst development, such ashetC. This study provides an example for ongoing microevolution, marked by multiple heterogeneous/heterozygous single nucleotide polymorphisms (SNPs), in a multicellular multicopy-genome microorganism.

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Causal Role of Single Nucleotide Polymorphisms within themprFGene of Staphylococcus aureus in Daptomycin Resistance

Antimicrobial Agents and Chemotherapy ◽

10.1128/aac.01184-13 ◽

2013 ◽

Vol 57 (11) ◽

pp. 5658-5664 ◽

Cited By ~ 48

Author(s):

Soo-Jin Yang ◽

Nagendra N. Mishra ◽

Aileen Rubio ◽

Arnold S. Bayer

Keyword(s):

Staphylococcus Aureus ◽

Single Nucleotide Polymorphisms ◽

Point Mutations ◽

Nucleotide Polymorphisms ◽

Wild Type ◽

Single Nucleotide ◽

Content Type ◽

Reading Frame ◽

A Charge

ABSTRACTSingle nucleotide polymorphisms (SNPs) within themprFopen reading frame (ORF) have been commonly observed in daptomycin-resistant (DAPr)Staphylococcus aureusstrains. Such SNPs are usually associated with a gain-in-function phenotype, in terms of either increased synthesis or enhanced translocation (flipping) of lysyl-phosphatidylglycerol (L-PG). However, it is unclear if suchmprFSNPs are causal in DAPrstrains or are merely a biomarker for this phenotype. In this study, we used an isogenic set ofS. aureusstrains: (i) Newman, (ii) its isogenic ΔmprFmutant, and (iii) several intransplasmid complementation constructs, expressing either a wild-type or point-mutated form of themprFORF cloned from two isogenic DAP-susceptible (DAPs)-DAPrstrain pairs (616-701 and MRSA11/11-REF2145). Complementation of the ΔmprFstrain with singly point-mutatedmprFgenes (mprFS295LormprFT345A) revealed that (i) individual and distinct point mutations within themprFORF can recapitulate phenotypes observed in donor strains (i.e., changes in DAP MICs, positive surface charge, and cell membrane phospholipid profiles) and (ii) these gain-in-function SNPs (i.e., enhanced L-PG synthesis) likely promote reduced DAP binding toS. aureusby a charge repulsion mechanism. Thus, for these two DAPrstrains, the definedmprFSNPs appear to be causally related to this phenotype.

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Using Mycobacterium tuberculosis Single-Nucleotide Polymorphisms To Predict Fluoroquinolone Treatment Response

Antimicrobial Agents and Chemotherapy ◽

10.1128/aac.00076-19 ◽

2019 ◽

Vol 63 (7) ◽

Cited By ~ 3

Author(s):

Marva Seifert ◽

Edmund Capparelli ◽

Donald G. Catanzaro ◽

Timothy C. Rodwell

Keyword(s):

Mycobacterium Tuberculosis ◽

Single Nucleotide Polymorphisms ◽

Specific Resistance ◽

Therapeutic Targets ◽

World Health ◽

Population Pharmacokinetic ◽

Nucleotide Polymorphisms ◽

Single Nucleotide ◽

Content Type ◽

Level Increase

ABSTRACT Clinical phenotypic fluoroquinolone susceptibility testing of Mycobacterium tuberculosis is currently based on M. tuberculosis growth at a single critical concentration, which provides limited information for a nuanced clinical response. We propose using specific resistance-conferring M. tuberculosis mutations in gyrA together with population pharmacokinetic and pharmacodynamic modeling as a novel tool to better inform fluoroquinolone treatment decisions. We sequenced the gyrA resistance-determining region of 138 clinical M. tuberculosis isolates collected from India, Moldova, Philippines, and South Africa and then determined each strain’s MIC against ofloxacin, moxifloxacin, levofloxacin, and gatifloxacin. Strains with specific gyrA single-nucleotide polymorphisms (SNPs) were grouped into high or low drug-specific resistance categories based on their empirically measured MICs. Published population pharmacokinetic models were then used to explore the pharmacokinetics and pharmacodynamics of each fluoroquinolone relative to the empirical MIC distribution for each resistance category to make predictions about the likelihood of patients achieving defined therapeutic targets. In patients infected with M. tuberculosis isolates containing SNPs associated with a fluoroquinolone-specific low-level increase in MIC, models suggest increased fluoroquinolone dosing improved the probability of achieving therapeutic targets for gatifloxacin and moxifloxacin but not for levofloxacin and ofloxacin. In contrast, among patients with isolates harboring SNPs associated with a high-level increase in MIC, increased dosing of levofloxacin, moxifloxacin, gatifloxacin, or ofloxacin did not meaningfully improve the probability of therapeutic target attainment. We demonstrated that quantifiable fluoroquinolone drug resistance phenotypes could be predicted from rapidly detectable gyrA SNPs and used to support dosing decisions based on the likelihood of patients reaching therapeutic targets. Our findings provide further supporting evidence for the moxifloxacin clinical breakpoint recently established by the World Health Organization.

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Quantitative Analysis of Single-Nucleotide Polymorphism for Rapid Detection of TR34/L98H- and TR46/Y121F/T289A-Positive Aspergillus fumigatus Isolates Obtained from Patients in Iran from 2010 to 2014

Antimicrobial Agents and Chemotherapy ◽

10.1128/aac.02326-15 ◽

2015 ◽

Vol 60 (1) ◽

pp. 387-392 ◽

Cited By ~ 11

Author(s):

Faezeh Mohammadi ◽

Seyed Jamal Hashemi ◽

Jan Zoll ◽

Willem J. G. Melchers ◽

Haleh Rafati ◽

...

Keyword(s):

Single Nucleotide Polymorphism ◽

Quantitative Analysis ◽

Single Nucleotide Polymorphisms ◽

Aspergillus Fumigatus ◽

Tandem Repeat ◽

Promoter Region ◽

Nucleotide Polymorphisms ◽

Nucleotide Polymorphism ◽

Single Nucleotide ◽

Content Type

ABSTRACTWe employed an endpoint genotyping method to update the prevalence rate of positivity for the TR34/L98H mutation (a 34-bp tandem repeat mutation in the promoter region of thecyp51Agene in combination with a substitution at codon L98) and the TR46/Y121F/T289A mutation (a 46-bp tandem repeat mutation in the promoter region of thecyp51Agene in combination with substitutions at codons Y121 and T289) among clinicalAspergillus fumigatusisolates obtained from different regions of Iran over a recent 5-year period (2010 to 2014). The antifungal activities of itraconazole, voriconazole, and posaconazole against 172 clinicalA. fumigatusisolates were investigated using the European Committee on Antimicrobial Susceptibility Testing (EUCAST) broth microdilution method. For the isolates with an azole resistance phenotype, thecyp51Agene and its promoter were amplified and sequenced. In addition, using a LightCycler 480 real-time PCR system, a novel endpoint genotyping analysis method targeting single-nucleotide polymorphisms was evaluated to detect the L98H and Y121F mutations in thecyp51Agene of all isolates. Of the 172A. fumigatusisolates tested, the MIC values of itraconazole (≥16 mg/liter) and voriconazole (>4 mg/liter) were high for 6 (3.5%). Quantitative analysis of single-nucleotide polymorphisms showed the TR34/L98H mutation in thecyp51Agenes of six isolates. No isolates harboring the TR46/Y121F/T289A mutation were detected. DNA sequencing of thecyp51Agene confirmed the results of the novel endpoint genotyping method. By microsatellite typing, all of the azole-resistant isolates had genotypes different from those previously recovered from Iran and from the Dutch TR34/L98H controls. In conclusion, there was not a significant increase in the prevalence of azole-resistantA. fumigatusisolates harboring the TR34/L98H resistance mechanism among isolates recovered over a recent 5-year period (2010 to 2014) in Iran. A quantitative assay detecting a single-nucleotide polymorphism in thecyp51Agene ofA. fumigatusis a reliable tool for the rapid screening and monitoring of TR34/L98H- and TR46/Y121F/T289A-positive isolates and can easily be incorporated into clinical mycology algorithms.

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Identification and characterisation of single nucleotide polymorphisms in interferon regulatory factor-5 gene of Nigerian local chickens

Agricultura tropica et subtropica ◽

10.2478/ats-2021-0001 ◽

2021 ◽

Vol 54 (1) ◽

pp. 1-13

Author(s):

Samuel Olutunde Durosaro ◽

Michael Ohiokhuaobo Ozoje ◽

Ayotunde Olutumininu Adebambo ◽

Okanlawon Mohamed Onagbesan

Keyword(s):

Cell Differentiation ◽

Single Nucleotide Polymorphisms ◽

Interferon Regulatory Factor ◽

Type I ◽

Nucleotide Polymorphisms ◽

Regulatory Factor ◽

Single Nucleotide ◽

Interferon Regulatory Factor 5 ◽

Software Packages ◽

Local Chickens

Abstract The interferon regulatory factor gene family encodes transcription factors with multiple biological functions, which include reproduction, cell differentiation and immunity. Interferon regulatory factor-5 (IRF-5) gene is involved in immune defence against virus, stress response, activation of type I interferon genes, cell differentiation and growth. This experiment was conducted to identify and characterise single nucleotide polymorphisms in exons 3, 4, 5 and 7 of IRF-5 gene in Nigerian local chickens. Exons 3, 4, 5 and 7 of IRF-5 gene were amplified and sequenced. Single nucleotide polymorphisms (SNPs) present in exons 3, 4, 5 and 7 of IRF-5 gene were identified and analysed using Clustal W, DnaSp and SNAP2 software packages. Four SNPs, rs317511101, rs312902332, rs315149141 and rs739389464, were identified in exon 3 of IRF-5 gene in all the three genotypes. Exon 4 of the gene was conserved while three of the SNPs (rs736423928, 170C>T and rs740736761) identified in exon 7 were shared among the three genotypes. Linkage disequilibrium of 1.00 existed between rs317511101 and rs315149141 polymorphisms identified in exon 3 of normal feathered and frizzle feathered chickens. Mutation rs740736761 identified in exon 7 had the highest polymorphism information content obtainable for any biallelic marker. Most of the SNPs identified in exons 3, 5 and 7 were synonymous and singletons which could not be used for association study. The study concluded that only haplotypes in exons 3 and 7 of IRF-5 gene can be used in marker-assisted selection when improving Nigerian local chickens.

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Genome Stability in Engineered Strains of the Extremely Thermophilic Lignocellulose-Degrading Bacterium Caldicellulosiruptor bescii

Applied and Environmental Microbiology ◽

10.1128/aem.00444-17 ◽

2017 ◽

Vol 83 (14) ◽

Cited By ~ 12

Author(s):

Amanda M. Williams-Rhaesa ◽

Farris L. Poole ◽

Jessica T. Dinsmore ◽

Gina L. Lipscomb ◽

Gabriel M. Rubinstein ◽

...

Keyword(s):

Metabolic Engineering ◽

Single Nucleotide Polymorphisms ◽

Genetic Background ◽

Genome Stability ◽

Plant Biomass ◽

Nucleotide Polymorphisms ◽

Single Nucleotide ◽

Content Type ◽

Targeted Deletion ◽

Caldicellulosiruptor Bescii

ABSTRACT Caldicellulosiruptor bescii is the most thermophilic cellulose degrader known and is of great interest because of its ability to degrade nonpretreated plant biomass. For biotechnological applications, an efficient genetic system is required to engineer it to convert plant biomass into desired products. To date, two different genetically tractable lineages of C. bescii strains have been generated. The first (JWCB005) is based on a random deletion within the pyrimidine biosynthesis genes pyrFA, and the second (MACB1018) is based on the targeted deletion of pyrE, making use of a kanamycin resistance marker. Importantly, an active insertion element, ISCbe4, was discovered in C. bescii when it disrupted the gene for lactate dehydrogenase (ldh) in strain JWCB018, constructed in the JWCB005 background. Additional instances of ISCbe4 movement in other strains of this lineage are presented herein. These observations raise concerns about the genetic stability of such strains and their use as metabolic engineering platforms. In order to investigate genome stability in engineered strains of C. bescii from the two lineages, genome sequencing and Southern blot analyses were performed. The evidence presented shows a dramatic increase in the number of single nucleotide polymorphisms, insertions/deletions, and ISCbe4 elements within the genome of JWCB005, leading to massive genome rearrangements in its daughter strain, JWCB018. Such dramatic effects were not evident in the newer MACB1018 lineage, indicating that JWCB005 and its daughter strains are not suitable for metabolic engineering purposes in C. bescii. Furthermore, a facile approach for assessing genomic stability in C. bescii has been established. IMPORTANCE Caldicellulosiruptor bescii is a cellulolytic extremely thermophilic bacterium of great interest for metabolic engineering efforts geared toward lignocellulosic biofuel and bio-based chemical production. Genetic technology in C. bescii has led to the development of two uracil auxotrophic genetic background strains for metabolic engineering. We show that strains derived from the genetic background containing a random deletion in uracil biosynthesis genes (pyrFA) have a dramatic increase in the number of single nucleotide polymorphisms, insertions/deletions, and ISCbe4 insertion elements in their genomes compared to the wild type. At least one daughter strain of this lineage also contains large-scale genome rearrangements that are flanked by these ISCbe4 elements. In contrast, strains developed from the second background strain developed using a targeted deletion strategy of the uracil biosynthetic gene pyrE have a stable genome structure, making them preferable for future metabolic engineering studies.

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Genome-Wide Identification of Host-Segregating Single-Nucleotide Polymorphisms for Source Attribution of Clinical Campylobacter coli Isolates

Applied and Environmental Microbiology ◽

10.1128/aem.01787-20 ◽

2020 ◽

Vol 86 (24) ◽

Author(s):

Quentin Jehanne ◽

Ben Pascoe ◽

Lucie Bénéjat ◽

Astrid Ducournau ◽

Alice Buissonnière ◽

...

Keyword(s):

Single Nucleotide Polymorphisms ◽

Common Species ◽

Frequency Variation ◽

Common Source ◽

Campylobacter Coli ◽

Nucleotide Polymorphisms ◽

Source Attribution ◽

Single Nucleotide ◽

Content Type ◽

Genome Wide

ABSTRACT Campylobacter is among the most common causes of gastroenteritis worldwide. Campylobacter jejuni and Campylobacter coli are the most common species causing human disease. DNA sequence-based methods for strain characterization have focused largely on C. jejuni, responsible for 80 to 90% of infections, meaning that C. coli epidemiology has lagged behind. Here, we have analyzed the genome of 450 C. coli isolates to determine genetic markers that can discriminate isolates sampled from 3 major reservoir hosts (chickens, cattle, and pigs). These markers then were applied to identify the source of infection of 147 C. coli strains from French clinical cases. Using STRUCTURE software, 259 potential host-segregating markers were revealed by probabilistic characterization of single-nucleotide polymorphism (SNP) frequency variation in strain collections from three different hosts. These SNPs were found in 41 genes or intergenic regions, mostly coding for proteins involved in motility and membrane functions. Source attribution of clinical isolates based on the differential presence of these markers confirmed chickens as the most common source of C. coli infection in France. IMPORTANCE Genome-wide and source attribution studies based on Campylobacter species have shown their importance for the understanding of foodborne infections. Although the use of multilocus sequence typing based on 7 genes from C. jejuni is a powerful method to structure populations, when applied to C. coli, results have not clearly demonstrated its robustness. Therefore, we aim to provide more accurate data based on the identification of single-nucleotide polymorphisms. Results from this study reveal an important number of host-segregating SNPs, found in proteins involved in motility, membrane functions, or DNA repair systems. These findings offer new, interesting opportunities for further study of C. coli adaptation to its environment. Additionally, the results demonstrate that poultry is potentially the main reservoir of C. coli in France.

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Evolution and Single-Nucleotide Polymorphisms in Methicillin-Resistant Staphylococcus aureus Strains with Reduced Susceptibility to Vancomycin and Daptomycin, Based on Determination of the Complete Genome

Antimicrobial Agents and Chemotherapy ◽

10.1128/aac.05159-14 ◽

2015 ◽

Vol 59 (6) ◽

pp. 3585-3587 ◽

Cited By ~ 10

Author(s):

Tetsuo Yamaguchi ◽

Shingo Suzuki ◽

Sakiko Okamura ◽

Yuri Miura ◽

Ayaka Tsukimori ◽

...

Keyword(s):

Staphylococcus Aureus ◽

Single Nucleotide Polymorphisms ◽

Complete Genome ◽

Nucleotide Polymorphisms ◽

Single Nucleotide ◽

Methicillin Resistant ◽

Content Type ◽

Rifampin Resistance ◽

Generation Sequencing

ABSTRACTWe obtained a series of methicillin-resistantStaphylococcus aureusisolates, including both daptomycin-susceptible strain TD1 and daptomycin-resistant strain TD4, from a patient. We determined the complete genome sequences of TD1 and TD4 using next-generation sequencing, and only four single-nucleotide polymorphisms (SNPs) were identified, one each incapB(E58K),rpoB(H481Y),lytN(I16V), andmprF(V351E). We determined that these four SNPs were sufficient to cause the strains to develop daptomycin, vancomycin, and rifampin resistance.

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Novel Multiplex Single Nucleotide Polymorphism-Based Method for Identifying Epidemic Clones of Listeria monocytogenes

Applied and Environmental Microbiology ◽

10.1128/aem.00429-11 ◽

2011 ◽

Vol 77 (17) ◽

pp. 6290-6294 ◽

Cited By ~ 16

Author(s):

Sara Lomonaco ◽

Stephen J. Knabel ◽

Alessandra Dalmasso ◽

Tiziana Civera ◽

Maria Teresa Bottero

Keyword(s):

Single Nucleotide Polymorphism ◽

Listeria Monocytogenes ◽

Single Nucleotide Polymorphisms ◽

High Throughput ◽

High Throughput Screening ◽

Virulence Genes ◽

Nucleotide Polymorphisms ◽

Nucleotide Polymorphism ◽

Single Nucleotide ◽

Content Type

ABSTRACTA novel primer extension-based, multiplex minisequencing assay targeting six highly informative single nucleotide polymorphisms (SNPs) in four virulence genes correctly identified and differentiated all four epidemic clones (ECs) ofListeria monocytogenesand 9 other strains initially misclassified as non-ECs. This assay allows rapid, accurate, and high-throughput screening for all known ECs ofL. monocytogenes.

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Frequency and Distribution of Single-Nucleotide Polymorphisms withinmprFin Methicillin-Resistant Staphylococcus aureus Clinical Isolates and Their Role in Cross-Resistance to Daptomycin and Host Defense Antimicrobial Peptides

Antimicrobial Agents and Chemotherapy ◽

10.1128/aac.00970-15 ◽

2015 ◽

Vol 59 (8) ◽

pp. 4930-4937 ◽

Cited By ~ 46

Author(s):

Arnold S. Bayer ◽

Nagendra N. Mishra ◽

Liang Chen ◽

Barry N. Kreiswirth ◽

Aileen Rubio ◽

...

Keyword(s):

Staphylococcus Aureus ◽

Single Nucleotide Polymorphisms ◽

Surface Charge ◽

Host Defense ◽

Cross Resistance ◽

Nucleotide Polymorphisms ◽

Single Nucleotide ◽

Methicillin Resistant ◽

Content Type ◽

Positive Surface Charge

ABSTRACTMprF is responsible for the lysinylation of phosphatidylglycerol (PG) to synthesize the positively charged phospholipid (PL) species, lysyl-PG (L-PG). It has been proposed that the single-nucleotide polymorphisms (SNPs) within themprFopen reading frame (ORF) are associated with a gain-in-function phenotype in terms of daptomycin resistance inStaphylococcus aureus. (Note that although the official term is daptomycin nonsusceptibility, we use the term daptomycin resistance in this paper for ease of presentation.) Using 22 daptomycin-susceptible (DAPs)/daptomycin-resistant (DAPr) clinical methicillin-resistantS. aureus(MRSA) strain pairs, we assessed (i) the frequencies and distribution of putativemprFgain-in-function SNPs, (ii) the relationships of the SNPs to both daptomycin resistance and cross-resistance to the prototypical endovascular host defense peptide (HDP) thrombin-induced platelet microbicidal protein (tPMP), and (iii) the impact ofmprFSNPs on positive surface charge phenotype and modifications of membrane PL profiles. Most of themprFSNPs identified in our DAPrstrains were clustered within the two MprF loci, (i) the central bifunctional domain and (ii) the C-terminal synthase domain. Moreover, we were able to correlate the presence and location ofmprFSNPs in DAPrstrains with HDP cross-resistance, positive surface charge, and L-PG profiles. Although DAPrstrains withmprFSNPs in the bifunctional domain showed higher resistance to tPMPs than DAPrstrains with SNPs in the synthase domain, this relationship was not observed in positive surface charge assays. These results demonstrated that both charge-mediated and -unrelated mechanisms are involved in DAP resistance and HDP cross-resistance inS. aureus.

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Loss of Bacitracin Resistance Due to a Large Genomic Deletion amongBacillus anthracisStrains

mSystems ◽

10.1128/msystems.00182-18 ◽

2018 ◽

Vol 3 (5) ◽

Cited By ~ 2

Author(s):

Yoshikazu Furuta ◽

Hayato Harima ◽

Emiko Ito ◽

Fumito Maruyama ◽

Naomi Ohnishi ◽

...

Keyword(s):

Genetic Diversity ◽

Single Nucleotide Polymorphisms ◽

Bacillus Cereus ◽

Bacillus Anthracis ◽

Bacterial Species ◽

Nucleotide Polymorphisms ◽

Single Nucleotide ◽

Genomic Deletion ◽

Content Type ◽

Large Genomic Deletion

ABSTRACTBacillus anthracisis a Gram-positive endospore-forming bacterial species that causes anthrax in both humans and animals. In Zambia, anthrax cases are frequently reported in both livestock and wildlife, with occasional transmission to humans, causing serious public health problems in the country. To understand the genetic diversity ofB. anthracisstrains in Zambia, we sequenced and compared the genomic DNA ofB. anthracisstrains isolated across the country. Single nucleotide polymorphisms clustered these strains into three groups. Genome sequence comparisons revealed a large deletion in strains belonging to one of the groups, possibly due to unequal crossing over between a pair of rRNA operons. The deleted genomic region included genes conferring resistance to bacitracin, and the strains with the deletion were confirmed with loss of bacitracin resistance. Similar deletions between rRNA operons were also observed in a fewB. anthracisstrains phylogenetically distant from Zambian strains. The structure of bacitracin resistance genes flanked by rRNA operons was conserved only in members of theBacillus cereusgroup. The diversity and genomic characteristics ofB. anthracisstrains determined in this study would help in the development of genetic markers and treatment of anthrax in Zambia.IMPORTANCEAnthrax is caused byBacillus anthracis, an endospore-forming soil bacterium. The genetic diversity ofB. anthracisis known to be low compared with that ofBacillusspecies. In this study, we performed whole-genome sequencing of Zambian isolates ofB. anthracisto understand the genetic diversity between closely related strains. Comparison of genomic sequences revealed that closely related strains were separated into three groups based on single nucleotide polymorphisms distributed throughout the genome. A large genomic deletion was detected in the region containing a bacitracin resistance gene cluster flanked by rRNA operons, resulting in the loss of bacitracin resistance. The structure of the deleted region, which was also conserved among species of theBacillus cereusgroup, has the potential for both deletion and amplification and thus might be enabling the species to flexibly control the level of bacitracin resistance for adaptive evolution.

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