scholarly journals Competition between Metals for Binding to Methanobactin Enables Expression of Soluble Methane Monooxygenase in the Presence of Copper

2014 ◽  
Vol 81 (3) ◽  
pp. 1024-1031 ◽  
Author(s):  
Bhagyalakshmi Kalidass ◽  
Muhammad Farhan Ul-Haque ◽  
Bipin S. Baral ◽  
Alan A. DiSpirito ◽  
Jeremy D. Semrau
Keyword(s):  
Methane Monooxygenase ◽  
High Specificity ◽  
Copper Uptake ◽  
Content Type ◽  
Soluble Methane Monooxygenase ◽  
Sds Page ◽  
Genes Encoding ◽  
Key Factor ◽  
Membrane Bound

ABSTRACTIt is well known that copper is a key factor regulating expression of the two forms of methane monooxygenase found in proteobacterial methanotrophs. Of these forms, the cytoplasmic, or soluble, methane monooxygenase (sMMO) is expressed only at low copper concentrations. The membrane-bound, or particulate, methane monooxygenase (pMMO) is constitutively expressed with respect to copper, and such expression increases with increasing copper. Recent findings have shown that copper uptake is mediated by a modified polypeptide, or chalkophore, termed methanobactin. Although methanobactin has high specificity for copper, it can bind other metals, e.g., gold. Here we show that inMethylosinus trichosporiumOB3b, sMMO is expressed and active in the presence of copper if gold is also simultaneously present. Such expression appears to be due to gold binding to methanobactin produced byM. trichosporiumOB3b, thereby limiting copper uptake. Such expression and activity, however, was significantly reduced if methanobactin preloaded with copper was also added. Further, quantitative reverse transcriptase PCR (RT-qPCR) of transcripts of genes encoding polypeptides of both forms of MMO and SDS-PAGE results indicate that both sMMO and pMMO can be expressed when copper and gold are present, as gold effectively competes with copper for binding to methanobactin. Such findings suggest that under certain geochemical conditions, both forms of MMO may be expressed and activein situ. Finally, these findings also suggest strategies whereby field sites can be manipulated to enhance sMMO expression, i.e., through the addition of a metal that can compete with copper for binding to methanobactin.

scholarly journals A TonB-Dependent Transporter Is Responsible for Methanobactin Uptake by Methylosinus trichosporium OB3b

2016 ◽  
Vol 82 (6) ◽  
pp. 1917-1923 ◽  
Author(s):  
Wenyu Gu ◽  
Muhammad Farhan Ul Haque ◽  
Bipin S. Baral ◽  
Erick A. Turpin ◽  
Nathan L. Bandow ◽  
...  
Keyword(s):  
Methane Monooxygenase ◽  
Copper Uptake ◽  
Wild Type ◽  
Particulate Methane Monooxygenase ◽  
Methylosinus Trichosporium Ob3b ◽  
Gene Encoding ◽  
Methylosinus Trichosporium ◽  
Content Type ◽  
Soluble Methane Monooxygenase ◽  
Genes Encoding

ABSTRACTMethanobactin, a small modified polypeptide synthesized by methanotrophs for copper uptake, has been found to be chromosomally encoded. The gene encoding the polypeptide precursor of methanobactin,mbnA, is part of a gene cluster that also includes several genes encoding proteins of unknown function (but speculated to be involved in methanobactin formation) as well asmbnT, which encodes a TonB-dependent transporter hypothesized to be responsible for methanobactin uptake. To determine ifmbnTis truly responsible for methanobactin uptake, a knockout was constructed inMethylosinus trichosporiumOB3b using marker exchange mutagenesis. The resultingM. trichosporiummbnT::Gmrmutant was found to be able to produce methanobactin but was unable to internalize it. Further, if this mutant was grown in the presence of copper and exogenous methanobactin, copper uptake was significantly reduced. Expression ofmmoXandpmoA, encoding polypeptides of the soluble methane monooxygenase (sMMO) and particulate methane monooxygenase (pMMO), respectively, also changed significantly when methanobactin was added, which indicates that the mutant was unable to collect copper under these conditions. Copper uptake and gene expression, however, were not affected in wild-typeM. trichosporiumOB3b, indicating that the TonB-dependent transporter encoded bymbnTis responsible for methanobactin uptake and that methanobactin is a key mechanism used by methanotrophs for copper uptake. When thembnT::Gmrmutant was grown under a range of copper concentrations in the absence of methanobactin, however, the phenotype of the mutant was indistinguishable from that of wild-typeM. trichosporiumOB3b, indicating that this methanotroph has multiple mechanisms for copper uptake.

scholarly journals Methanobactin from Methylocystis sp. Strain SB2 Affects Gene Expression and Methane Monooxygenase Activity in Methylosinus trichosporium OB3b

2015 ◽  
Vol 81 (7) ◽  
pp. 2466-2473 ◽  
Author(s):  
Muhammad Farhan Ul-Haque ◽  
Bhagyalakshmi Kalidass ◽  
Alexey Vorobev ◽  
Bipin S. Baral ◽  
Alan A. DiSpirito ◽  
...  
Keyword(s):  
Gene Expression ◽  
Methane Monooxygenase ◽  
Particulate Methane Monooxygenase ◽  
Methylosinus Trichosporium Ob3b ◽  
Methylosinus Trichosporium ◽  
Monooxygenase Activity ◽  
Content Type ◽  
Soluble Methane Monooxygenase ◽  
A Subunit ◽  
Membrane Bound

ABSTRACTMethanotrophs can express a cytoplasmic (soluble) methane monooxygenase (sMMO) or membrane-bound (particulate) methane monooxygenase (pMMO). Expression of these MMOs is strongly regulated by the availability of copper. Many methanotrophs have been found to synthesize a novel compound, methanobactin (Mb), that is responsible for the uptake of copper, and methanobactin produced byMethylosinus trichosporiumOB3b plays a key role in controlling expression of MMO genes in this strain. As all known forms of methanobactin are structurally similar, it was hypothesized that methanobactin from one methanotroph may alter gene expression in another. WhenMethylosinus trichosporiumOB3b was grown in the presence of 1 μM CuCl2, expression ofmmoX, encoding a subunit of the hydroxylase component of sMMO, was very low.mmoXexpression increased, however, when methanobactin fromMethylocystissp. strain SB2 (SB2-Mb) was added, as did whole-cell sMMO activity, but there was no significant change in the amount of copper associated withM. trichosporiumOB3b. IfM. trichosporiumOB3b was grown in the absence of CuCl2, themmoXexpression level was high but decreased by several orders of magnitude if copper prebound to SB2-Mb (Cu-SB2-Mb) was added, and biomass-associated copper was increased. Exposure ofMethylosinus trichosporiumOB3b to SB2-Mb had no effect on expression ofmbnA, encoding the polypeptide precursor of methanobactin in either the presence or absence of CuCl2.mbnAexpression, however, was reduced when Cu-SB2-Mb was added in both the absence and presence of CuCl2. These data suggest that methanobactin acts as a general signaling molecule in methanotrophs and that methanobactin “piracy” may be commonplace.

scholarly journals Insight from Molecular, Pathological, and Immunohistochemical Studies on Cellular and Humoral Mechanisms Responsible for Vaccine-Induced Protection of Rainbow Trout against Yersinia ruckeri

2013 ◽  
Vol 20 (10) ◽  
pp. 1623-1641 ◽  
Author(s):  
Sidhartha Deshmukh ◽  
Per W. Kania ◽  
Jiwan K. Chettri ◽  
Jakob Skov ◽  
Anders M. Bojesen ◽  
...  
Keyword(s):  
Rainbow Trout ◽  
Acute Phase Proteins ◽  
Head Kidney ◽  
Yersinia Ruckeri ◽  
Content Type ◽  
Immune Genes ◽  
Immunological Mechanisms ◽  
Genes Encoding ◽  
Immune Related Genes

ABSTRACTThe immunological mechanisms associated with protection of vaccinated rainbow trout,Oncorhynchus mykiss, against enteric redmouth disease (ERM), caused byYersinia ruckeri, were previously elucidated by the use of gene expression methodology and immunochemical methods. That approach pointed indirectly to both humoral and cellular elements being involved in protection. The present study correlates the level of protection in rainbow trout to cellular reactions in spleen and head kidney and visualizes the processes by applying histopathological, immunohistochemical, andin situhybridization techniques. It was shown that these cellular reactions, which were more prominent in spleen than in head kidney, were associated with the expression of immune-related genes, suggesting a Th2-like response.Y. ruckeri, as shown byin situhybridization (ISH), was eliminated within a few days in vaccinated fish, whereas nonprotected fish still harbored bacteria for a week after infection. Vaccinated fish reestablished normal organ structure within a few days, whereas nonprotected fish showed abnormalities up to 1 month postinfection. Protection in the early phase of infection was mainly associated with the expression of genes encoding innate factors (complement factors, lysozyme, and acute phase proteins), but in the later phase of infection, increased expression of adaptive immune genes dominated. The histological approach used has shown that the cellular changes correlated with protection of vaccinated fish. They comprised transformation of resident cells into macrophage-like cells and increased occurrence of CD8α and IgM cells, suggesting these cells as main players in protection. Future studies should investigate the causality between these factors and protection.

scholarly journals Small-Molecule Inhibition of Choline Catabolism in Pseudomonas aeruginosa and Other Aerobic Choline-Catabolizing Bacteria

2011 ◽  
Vol 77 (13) ◽  
pp. 4383-4389 ◽  
Author(s):  
Liam F. Fitzsimmons ◽  
Stevenson Flemer ◽  
A. Sandy Wurthmann ◽  
P. Bruce Deker ◽  
Indra Neil Sarkar ◽  
...  
Keyword(s):  
Pseudomonas Aeruginosa ◽  
Burkholderia Cepacia ◽  
Sinorhizobium Meliloti ◽  
Growth Inhibitor ◽  
Bioinformatic Analysis ◽  
Content Type ◽  
Genes Encoding ◽  
Dependent Growth

ABSTRACTCholine is abundant in association with eukaryotes and plays roles in osmoprotection, thermoprotection, and membrane biosynthesis in many bacteria. Aerobic catabolism of choline is widespread among soil proteobacteria, particularly those associated with eukaryotes. Catabolism of choline as a carbon, nitrogen, and/or energy source may play important roles in association with eukaryotes, including pathogenesis, symbioses, and nutrient cycling. We sought to generate choline analogues to study bacterial choline catabolismin vitroandin situ. Here we report the characterization of a choline analogue, propargylcholine, which inhibits choline catabolism at the level of Dgc enzyme-catalyzed dimethylglycine demethylation inPseudomonas aeruginosa. We used genetic analyses and13C nuclear magnetic resonance to demonstrate that propargylcholine is catabolized to its inhibitory form, propargylmethylglycine. Chemically synthesized propargylmethylglycine was also an inhibitor of growth on choline. Bioinformatic analysis suggests that there are genes encoding DgcA homologues in a variety of proteobacteria. We examined the broader utility of propargylcholine and propargylmethylglycine by assessing growth of other members of the proteobacteria that are known to grow on choline and possess putative DgcA homologues. Propargylcholine showed utility as a growth inhibitor inP. aeruginosabut did not inhibit growth in other proteobacteria tested. In contrast, propargylmethylglycine was able to inhibit choline-dependent growth in all tested proteobacteria, includingPseudomonas mendocina,Pseudomonas fluorescens,Pseudomonas putida,Burkholderia cepacia,Burkholderia ambifaria, andSinorhizobium meliloti. We predict that chemical inhibitors of choline catabolism will be useful for studying this pathway in clinical and environmental isolates and could be a useful tool to study proteobacterial choline catabolismin situ.

scholarly journals Efficient Production of 2,5-Diketo-d-Gluconate via Heterologous Expression of 2-Ketogluconate Dehydrogenase in Gluconobacter japonicus

2015 ◽  
Vol 81 (10) ◽  
pp. 3552-3560 ◽  
Author(s):  
Naoya Kataoka ◽  
Minenosuke Matsutani ◽  
Toshiharu Yakushi ◽  
Kazunobu Matsushita
Keyword(s):  
Mutant Strain ◽  
Gluconobacter Oxydans ◽  
Efficient Production ◽  
Broad Host Range ◽  
Putative Promoter Region ◽  
Content Type ◽  
Biotransformation Process ◽  
Genes Encoding ◽  
Membrane Bound ◽  
Almost All

ABSTRACT2,5-Diketo-d-gluconate (2,5DKG) is a compound that can be the intermediate ford-tartrate and also vitamin C production. AlthoughGluconobacter oxydansNBRC3293 produces 2,5DKG fromd-glucose viad-gluconate and 2-keto-d-gluconate (2KG), with accumulation of the product in the culture medium, the efficiency of 2,5DKG production is unsatisfactory because there is a large amount of residuald-gluconate at the end of the biotransformation process. Oxidation of 2KG to 2,5DKG is catalyzed by a membrane-bound flavoprotein-cytochromeccomplex: 2-keto-gluconate dehydrogenase (2KGDH). Here, we studied thekgdSLCgenes encoding 2KGDH inG. oxydansNBRC3293 to improve 2,5DKG production byGluconobacterspp. ThekgdS,kgdL, andkgdCgenes correspond to the small, large, and cytochrome subunits of 2KGDH, respectively. ThekgdSLCgenes were cloned into a broad-host-range vector carrying a DNA fragment of the putative promoter region of the membrane-bound alcohol dehydrogenase gene ofG. oxydansfor expression inGluconobacterspp. According to our results, 2KGDH that was purified from the recombinantGluconobactercells showed characteristics nearly the same as those reported previously. We also expressed thekgdSLCgenes in a mutant strain ofGluconobacter japonicusNBRC3271 (formerlyGluconobacter dioxyacetonicusIFO3271) engineered to produce 2KG efficiently from a mixture ofd-glucose andd-gluconate. This mutant strain consumed almost all of the starting materials (d-glucose andd-gluconate) to produce 2,5DKG quantitatively as a seemingly unique metabolite. To our knowledge, this is the first report of aGluconobacterstrain that produces 2,5DKG efficiently and homogeneously.

scholarly journals Response of a Rice Paddy Soil Methanogen to Syntrophic Growth as Revealed by Transcriptional Analyses

2014 ◽  
Vol 80 (15) ◽  
pp. 4668-4676 ◽  
Author(s):  
Pengfei Liu ◽  
Yanxiang Yang ◽  
Zhe Lü ◽  
Yahai Lu
Keyword(s):  
Paddy Field ◽  
Rice Paddy ◽  
Transcription Unit ◽  
Heterodisulfide Reductase ◽  
Content Type ◽  
Acetate Assimilation ◽  
Adaptive Mechanisms ◽  
Genes Encoding ◽  
Membrane Bound ◽  
Key Steps

ABSTRACTMembers ofMethanocellalesare widespread in paddy field soils and play the key role in methane production. These methanogens feature largely in these organisms' adaptation to low H2and syntrophic growth with anaerobic fatty acid oxidizers. The adaptive mechanisms, however, remain unknown. In the present study, we determined the transcripts of 21 genes involved in the key steps of methanogenesis and acetate assimilation ofMethanocella conradiiHZ254, a strain recently isolated from paddy field soil.M. conradiiwas grown in monoculture and syntrophically withPelotomaculum thermopropionicum(a propionate syntroph) orSyntrophothermus lipocalidus(a butyrate syntroph). Comparison of the relative transcript abundances showed that three hydrogenase-encoding genes and all methanogenesis-related genes tested were upregulated in cocultures relative to monoculture. The genes encoding formylmethanofuran dehydrogenase (Fwd), heterodisulfide reductase (Hdr), and the membrane-bound energy-converting hydrogenase (Ech) were the most upregulated among the evaluated genes. The expression of the formate dehydrogenase (Fdh)-encoding gene also was significantly upregulated. In contrast, an acetate assimilation gene was downregulated in cocultures. The genes coding for Fwd, Hdr, and the D subunit of F420-nonreducing hydrogenase (Mvh) form a large predicted transcription unit; therefore, the Mvh/Hdr/Fwd complex, capable of mediating the electron bifurcation and connecting the first and last steps of methanogenesis, was predicted to be formed inM. conradii. We propose thatMethanocellamethanogens cope with low H2and syntrophic growth by (i) stabilizing the Mvh/Hdr/Fwd complex and (ii) activating formate-dependent methanogenesis.

scholarly journals Marker Exchange Mutagenesis ofmxaF, Encoding the Large Subunit of the Mxa Methanol Dehydrogenase, in Methylosinus trichosporium OB3b

2015 ◽  
Vol 82 (5) ◽  
pp. 1549-1555 ◽  
Author(s):  
Muhammad Farhan Ul Haque ◽  
Wenyu Gu ◽  
Alan A. DiSpirito ◽  
Jeremy D. Semrau
Keyword(s):  
Large Subunit ◽  
Methane Monooxygenase ◽  
Methanol Dehydrogenase ◽  
Methylosinus Trichosporium Ob3b ◽  
Initial Oxidation ◽  
Methylosinus Trichosporium ◽  
Content Type ◽  
Marker Exchange ◽  
Oxidation Of Methane ◽  
Soluble Methane Monooxygenase

ABSTRACTMethanotrophs have remarkable redundancy in multiple steps of the central pathway of methane oxidation to carbon dioxide. For example, it has been known for over 30 years that two forms of methane monooxygenase, responsible for oxidizing methane to methanol, exist in methanotrophs, i.e., soluble methane monooxygenase (sMMO) and particulate methane monooxygenase (pMMO), and that expression of these two forms is controlled by the availability of copper. Specifically, sMMO expression occurs in the absence of copper, while pMMO expression increases with increasing copper concentrations. More recently, it was discovered that multiple forms of methanol dehydrogenase (MeDH), Mxa MeDH and Xox MeDH, also exist in methanotrophs and that the expression of these alternative forms is regulated by the availability of cerium. That is, expression of Xox MeDH increases in the presence of cerium, while Mxa MeDH expression decreases in the presence of cerium. As it had been earlier concluded that pMMO and Mxa MeDH form a supercomplex in which electrons from Mxa MeDH are back donated to pMMO to drive the initial oxidation of methane, we speculated that Mxa MeDH could be rendered inactive through marker-exchange mutagenesis but growth on methane could still be possible if cerium was added to increase the expression of Xox MeDH under sMMO-expressing conditions. Here we report thatmxaF, encoding the large subunit of Mxa MeDH, could indeed be knocked out inMethylosinus trichosporiumOB3b, yet growth on methane was still possible, so long as cerium was added. Interestingly, growth of this mutant occurred in both the presence and the absence of copper, suggesting that Xox MeDH can replace Mxa MeDH regardless of the form of MMO expressed.

scholarly journals The origin of aerobic methanotrophy within the Proteobacteria

2019 ◽  
Vol 366 (9) ◽  
Author(s):  
Christina S Kang ◽  
Peter F Dunfield ◽  
Jeremy D Semrau
Keyword(s):  
Gene Transfer ◽  
Lateral Gene Transfer ◽  
Methane Monooxygenase ◽  
Copper Uptake ◽  
Particulate Methane Monooxygenase ◽  
Conversion Of Methane ◽  
Genes Encoding ◽  
Aerobic Methanotrophs ◽  
Encoding Genes ◽  
Global Carbon

ABSTRACT Aerobic methanotrophs play critical roles in the global carbon cycle, but despite their environmental ubiquity, they are phylogenetically restricted. Via bioinformatic analyses, it is shown that methanotrophy likely arose from methylotrophy from the lateral gene transfer of either of the two known forms of methane monooxygenase (particulate and soluble methane monooxygenases). Moreover, it appears that both known forms of pyrroloquinoline quinone-dependent methanol dehydrogenase (MeDH) found in methanotrophs—the calcium-containing Mxa-MeDH and the rare earth element-containing Xox-MeDH—were likely encoded in the genomes before the acquisition of the methane monooxygenases (MMOs), but that some methanotrophs subsequently received an additional copy of Xox-MeDH-encoding genes via lateral gene transfer. Further, data are presented that indicate the evolution of methanotrophy from methylotrophy not only required lateral transfer of genes encoding for methane monooxygenases, but also likely the pre-existence of a means of collecting copper. Given the emerging interest in valorizing methane via biological platforms, it is recommended that future strategies for heterologous expression of methane monooxygenase for conversion of methane to methanol also include cloning of genes encoding mechanism(s) of copper uptake, especially for expression of particulate methane monooxygenase.

scholarly journals Gammaproteobacterial Methanotrophs Dominate Cold Methane Seeps in Floodplains of West Siberian Rivers

2014 ◽  
Vol 80 (19) ◽  
pp. 5944-5954 ◽  
Author(s):  
Igor Y. Oshkin ◽  
Carl-Eric Wegner ◽  
Claudia Lüke ◽  
Mikhail V. Glagolev ◽  
Illiya V. Filippov ◽  
...  
Keyword(s):  
West Siberia ◽  
Dry Weight ◽  
Small Rivers ◽  
Type I ◽  
Content Type ◽  
Cell Counts ◽  
Genes Encoding ◽  
Irtysh Basin ◽  
Methane Fluxes

ABSTRACTA complex system of muddy fluid-discharging and methane (CH4)-releasing seeps was discovered in a valley of the river Mukhrinskaya, one of the small rivers of the Irtysh Basin, West Siberia. CH4flux from most (90%) of these gas ebullition sites did not exceed 1.45 g CH4h−1, while some seeps emitted up to 5.54 g CH4h−1. The δ13C value of methane released from these seeps varied between −71.1 and −71.3‰, suggesting its biogenic origin. Although the seeps were characterized by lowin situtemperatures (3.5 to 5°C), relatively high rates of methane oxidation (15.5 to 15.9 nmol CH4ml−1day−1) were measured in mud samples. Fluorescencein situhybridization detected 107methanotrophic bacteria (MB) per g of mud (dry weight), which accounted for up to 20.5% of total bacterial cell counts. Most (95.8 to 99.3%) methanotroph cells were type I (gammaproteobacterial) MB. The diversity of methanotrophs in this habitat was further assessed by pyrosequencing ofpmoAgenes, encoding particulate methane monooxygenase. A total of 53,828pmoAgene sequences of seep-inhabiting methanotrophs were retrieved and analyzed. Nearly all of these sequences affiliated with type I MB, including theMethylobacter-Methylovulum-Methylosomagroup, lake cluster 2, and several as-yet-uncharacterized methanotroph clades. Apparently, microbial communities attenuating methane fluxes from these local but strong CH4sources in floodplains of high-latitude rivers have a large proportion of potentially novel, psychrotolerant methanotrophs, thereby providing a challenge for future isolation studies.

scholarly journals Activity of a Holin-Endolysin System in the Insecticidal Pathogenicity Island ofYersinia enterocolitica

2018 ◽  
Vol 200 (16) ◽  
Author(s):  
Katharina Springer ◽  
Sandra Reuter ◽  
Mandy Knüpfer ◽  
Lukas Schmauder ◽  
Philipp-Albert Sänger ◽  
...  
Keyword(s):  
Insecticidal Activity ◽  
High Specificity ◽  
Pathogenicity Island ◽  
Open Reading Frames ◽  
Content Type ◽  
Toxin Complex ◽  
Genes Encoding ◽  
Conserved Genes ◽  
Nematocidal Activity ◽  
Lysis Cassette

ABSTRACTYersinia enterocoliticais a pathogen that causes gastroenteritis in humans. Because of its low-temperature-dependent insecticidal activity, it can oscillate between invertebrates and mammals as host organisms. The insecticidal activity of strain W22703 is associated with a pathogenicity island of 19 kb (Tc-PAIYe), which carries regulators and genes encoding the toxin complex (Tc). The island also harbors four phage-related and highly conserved genes of unknown functions, which are polycistronically transcribed. Two open reading frames showed significant homologies to holins and endolysins and exhibited lytic activity inEscherichia colicells upon overexpression. When a set ofYersiniastrains was tested in an equivalent manner, highly diverse susceptibilities to lysis were observed, and some strains were resistant to lysis. If cell lysis occurred (as demonstrated by membrane staining), it was more pronounced when two accessory elements of the cassette coding for an i-spanin and an o-spanin were included in the overexpression construct. The pore-forming function of the putative holin, HolY, was demonstrated by complementation of the lysis defect of a phage λ S holin mutant. In experiments performed with membrane preparations, ElyY exhibited high specificity for W22703 peptidoglycan, with a cleavage activity resembling that of lysozyme. Although the functionality of the lysis cassette from Tc-PAIYewas demonstrated in this study, its biological role remains to be elucidated.IMPORTANCEThe knowledge of how pathogens survive in the environment is pivotal for our understanding of bacterial virulence. The insecticidal and nematocidal activity ofYersiniaspp., by which the bacteria gain access to nutrients and thus improve their environmental fitness, is conferred by the toxin complex (Tc) encoded on a highly conserved pathogenicity island termed Tc-PAIYe. While the regulators and the toxin subunits of the island had been characterized in some detail, the role of phage-related genes within the island remained to be elucidated. Here, we demonstrate that this cassette encodes a holin, an endolysin, and two spanins that, at least upon overexpression, lyseYersiniastrains.

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