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      세균의 인산 항상성: 인산 수송 단백질들의 역할 = Bacterial Phosphate Homeostasis: Role of Phosphate Transporters

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      https://www.riss.kr/link?id=A101548014

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      국문 초록 (Abstract)

      인은 인지질, 탄수화물 및 핵산 등의 생분자 합성에 필요한 원소이다. 세균은 외부환경으로부터 인산이나 인산을 포함하는 영양소를 흡수하여 인을 얻고, 세포대사에 사용되고 남은 인산은 ...

      인은 인지질, 탄수화물 및 핵산 등의 생분자 합성에 필요한 원소이다. 세균은 외부환경으로부터 인산이나 인산을 포함하는 영양소를 흡수하여 인을 얻고, 세포대사에 사용되고 남은 인산은 polyphosphate 형태로 저장한다. 현재까지 알려진 다섯 개의 인산 수송 시스템 중, 인산에 특이적으로 높은 친화력을 갖는 Pst 시스템이 가장 중요한 역할을 하며, 그 발현은 세포외부 인산 농도에 반응하는 PhoB-PhoR two component 신호전달 시스템에 의해 조절된다. 반응 조절 단백질 PhoB는 인산 대사뿐 아니라 이와 관계없는 유전자들의 전사를 조절하는 것으로 알려졌으며, 따라서 PhoB의 활성이 조절되지 않으면 많은 종류의 다른 표현형이 나타난다. 본 총설은 각 인산 수송 시스템의 기능이 결여된 세균의 표현형에 대한 최근 연구 결과를 토대로 다음과 같은 내용을 기술하였다. 첫째, 세포 내부 인산의 적정 농도 유지를 위한 인산 수송 시스템들의 역할, 둘째, 인산뿐 아니라 여타 환경 신호와 관련된 수송 시스템의 다양한 표현형, 그리고 마지막으로, 수송 시스템들 간 혹은 그 조절자들 간의 표현형 중복을 분류하여 제시하였다. 이러한 내용은 결국 세균의 대사, 적응반응 및 병원성 발현에 미치는 인산 항상성의 중요성을 강조한다.

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      다국어 초록 (Multilingual Abstract)

      Phosphorous is an essential element for the synthesis of various biomolecules including phospholipids, carbohydrates and nucleic acids. Bacterial cells can uptake it as forms of phosphate and phosphate-containing nutrients from extracellular environme...

      Phosphorous is an essential element for the synthesis of various biomolecules including phospholipids, carbohydrates and nucleic acids. Bacterial cells can uptake it as forms of phosphate and phosphate-containing nutrients from extracellular environments, and reserve extra phosphate to polyphosphate inside the cell. Among five phosphate transport systems, Pst plays central roles in phosphate transport, and its expression is coordinated by the regulation of PhoB-PhoR two component signal transduction system in response to extracellular levels of phosphate. Genomic studies on the response regulator PhoB reveal many genes independent of phosphate metabolism. Based on recent findings on phenotypes of bacteria lacking proper function of each phosphate transport system, this review discusses roles of phosphate transporters in maintaining optimum intracellular phosphate levels, and presents diverse phenotypes of phosphate transporters related with other environmental signals as well as phosphate, then finally points out functional redundancy among phosphate transport systems or their regulators, which emphasize importance of phosphate homeostasis in governing metabolism, adaptation, and virulence of bacteria.

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      참고문헌 (Reference)

      1 Steed, P.M., "Use of the rep technique for allele replacement to construct mutants with deletions of the pstSCABphoUoperon: evidence of a new role for the PhoU protein in the phosphate regulon" 175 : 6797-6809, 1993

      2 van Veen, H.W., "Translocation of metal phosphate via the phosphate inorganic transport system of Escherichia coli" 33 : 1766-1770, 1994

      3 Aguena, M., "Transcriptional analysis of the pst operon of Escherichia coli" 268 : 518-524, 2002

      4 Fischer, R.J., "Transcription of the pst operon of Clostridium acetobutylicum is dependent on phosphate concentration and pH" 188 : 5469-5478, 2006

      5 Baek, J. H., "Transcript and protein level analyses of the interactions among PhoB, PhoR, PhoU and CreC in response to phosphate starvation in Escherichia coli" BLACKWELL PUBLISHING 277 : 254-259, 2007

      6 Kim, H. J., "Transcript accumulation from the rpoS gene encoding a stationary-phase sigma factor in Pseudomonas chlororaphis strain O6 is regulated by the polyphosphate kinase gene" SPRINGER-VERLAG 54 (54): 219-233, 2007

      7 Nezbedova, S., "The use of glass beads cultivation system to study the global effect of the ppk gene inactivation in Streptomyces lividans" 56 : 519-525, 2011

      8 Scanlan, D.J., "The response of the picoplanktonic marine cyanobacterium Synechococcus species WH7803 to phosphate starvation involves a protein homologous to the periplasmic phosphate-binding protein of Escherichia coli" 10 : 181-191, 1993

      9 Geiger, O., "The regulator gene phoB mediates phosphate stress-controlled synthesis of the membrane lipid diacylglyceryl-N,N,N-trimethylhomoserine in Rhizobium (Sinorhizobium) meliloti" 32 : 63-73, 1999

      10 Panhorst, M., "The pstSCAB operon for phosphate uptake is regulated by the global regulator GlxR in Corynebacterium glutamicum" 154 : 149-155, 2011

      1 Steed, P.M., "Use of the rep technique for allele replacement to construct mutants with deletions of the pstSCABphoUoperon: evidence of a new role for the PhoU protein in the phosphate regulon" 175 : 6797-6809, 1993

      2 van Veen, H.W., "Translocation of metal phosphate via the phosphate inorganic transport system of Escherichia coli" 33 : 1766-1770, 1994

      3 Aguena, M., "Transcriptional analysis of the pst operon of Escherichia coli" 268 : 518-524, 2002

      4 Fischer, R.J., "Transcription of the pst operon of Clostridium acetobutylicum is dependent on phosphate concentration and pH" 188 : 5469-5478, 2006

      5 Baek, J. H., "Transcript and protein level analyses of the interactions among PhoB, PhoR, PhoU and CreC in response to phosphate starvation in Escherichia coli" BLACKWELL PUBLISHING 277 : 254-259, 2007

      6 Kim, H. J., "Transcript accumulation from the rpoS gene encoding a stationary-phase sigma factor in Pseudomonas chlororaphis strain O6 is regulated by the polyphosphate kinase gene" SPRINGER-VERLAG 54 (54): 219-233, 2007

      7 Nezbedova, S., "The use of glass beads cultivation system to study the global effect of the ppk gene inactivation in Streptomyces lividans" 56 : 519-525, 2011

      8 Scanlan, D.J., "The response of the picoplanktonic marine cyanobacterium Synechococcus species WH7803 to phosphate starvation involves a protein homologous to the periplasmic phosphate-binding protein of Escherichia coli" 10 : 181-191, 1993

      9 Geiger, O., "The regulator gene phoB mediates phosphate stress-controlled synthesis of the membrane lipid diacylglyceryl-N,N,N-trimethylhomoserine in Rhizobium (Sinorhizobium) meliloti" 32 : 63-73, 1999

      10 Panhorst, M., "The pstSCAB operon for phosphate uptake is regulated by the global regulator GlxR in Corynebacterium glutamicum" 154 : 149-155, 2011

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      13 Gebhard, S., "The low-affinity phosphate transporter PitA is dispensable for in vitro growth of Mycobacterium smegmatis" 9 : 254-, 2009

      14 Diaz, M., "The high-affinity phosphate-binding protein PstS is accumulated under high fructose concentrations and mutation of the corresponding gene affects differentiation in Streptomyces lividans" 151 : 2583-2592, 2005

      15 Jacobsen, S.M., "The high-affinity phosphate transporter Pst is a virulence factor for Proteus mirabilis during complicated urinary tract infection" 52 : 180-193, 2008

      16 O'May, G.A., "The high-affinity phosphate transporter Pst in Proteus mirabilis HI4320 and its importance in biofilm formation" 155 : 1523-1535, 2009

      17 Castaneda-Garcia, A., "The glycerol-3-phosphate permease GlpT is the only fosfomycin transporter in Pseudomonas aeruginosa" 191 : 6968-6974, 2009

      18 Mudrak, B., "The Vibrio cholerae Pst2 phosphate transport system is upregulated in biofilms and contributes to biofilm-induced hyperinfectivity" 80 : 1794-1802, 2012

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      20 Luz, D.E., "The Pst system of Streptococcus mutans is important for phosphate transport and adhesion to abiotic surfaces" 27 : 172-181, 2012

      21 Gristwood, T., "The PhoBR two-component system regulates antibiotic biosynthesis in Serratia in response to phosphate" 9 : 112-, 2009

      22 Sultan, S.Z., "The PhoB regulatory system modulates biofilm formation and stress response in El Tor biotype Vibrio cholerae" 302 : 22-31, 2010

      23 Crepin, S., "The Pho regulon and the pathogenesis of Escherichia coli" 153 : 82-88, 2011

      24 Borsetti, F., "Tellurite uptake by cells of the facultative phototroph Rhodobacter capsulatus is a Delta pH-dependent process" 554 : 315-318, 2003

      25 Huang, Y., "Structure and mechanism of the glycerol-3-phosphate transporter from Escherichia coli" 301 : 616-620, 2003

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      32 Suda Tunpiboonsak, "Role of a Burkholderia pseudomallei Polyphosphate Kinase in an Oxidative Stress Response, Motilities, and Biofilm Formation" 한국미생물학회 48 (48): 63-70, 2010

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      38 Burall, L.S., "Proteus mirabilis genes that contribute to pathogenesis of urinary tract infection: identification of 25 signature-tagged mutants attenuated at least 100-fold" 72 : 2922-2938, 2004

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      40 Merkel, T.J., "Promoter elements required for positive control of transcription of the Escherichia coli uhpT gene" 174 : 2763-2770, 1992

      41 Jahid, I.K., "Polyphosphate stores enhance the ability of Vibrio cholerae to overcome environmental stresses in a low-phosphate environment" 72 : 7043-7049, 2006

      42 Price-Carter, M., "Polyphosphate kinase protects Salmonella enterica from weak organic acid stress" 187 : 3088-3099, 2005

      43 Sureka, K., "Polyphosphate kinase 2: a modulator of nucleoside diphosphate kinase activity in mycobacteria" 74 : 1187-1197, 2009

      44 Wanner, B.L., "Phosphorus assimilation and control of the phosphate regulon, In Escherichia coli and Salmonella typhimurium: Cellular and Molecular Biology" American Society for Microbiology Press 1357-1381, 1996

      45 Surin, B.P., "Phosphate-specific transport system of Escherichia coli: nucleotide sequence and gene-polypeptide relationships" 161 : 189-198, 1985

      46 van Veen, H.W., "Phosphate transport in prokaryotes: molecules, mediators and mechanisms" 72 : 299-315, 1997

      47 Yuan, Z.C., "Phosphate limitation induces catalase expression in Sinorhizobium meliloti, Pseudomonas aeruginosa and Agrobacterium tumefaciens" 58 : 877-894, 2005

      48 Rifat, D., "Phosphate depletion: a novel trigger for Mycobacterium tuberculosis persistence" 200 : 1126-1135, 2009

      49 Slater, H., "Phosphate availability regulates biosynthesis of two antibiotics, prodigiosin and carbapenem, in Serratia via both quorum-sensing-dependent and -independent pathways" 47 : 303-320, 2003

      50 Li, Y., "PhoU is a persistence switch involved in persister formation and tolerance to multiple antibiotics and stresses in Escherichia coli" 51 : 2092-2099, 2007

      51 Buckles, E.L., "PhoU enhances the ability of extraintestinal pathogenic Escherichia coli strain CFT073 to colonize the murine urinary tract" 152 : 153-160, 2006

      52 Pratt, J.T., "PhoB regulates motility, biofilms, and cyclic di-GMP in Vibrio cholerae" 191 : 6632-6642, 2009

      53 Pratt, J.T., "PhoB regulates both environmental and virulence gene expression in Vibrio cholerae" 77 : 1595-1605, 2010

      54 Lau, W. T., "Pho86p, an endoplasmic reticulum (ER) resident protein in Saccharomyces cerevisiae, is required for ER exit of the high-affinity phosphate transporter Pho84p" 97 : 1107-1112, 2000

      55 Motomura, K., "Overproduction of YjbB reduces the level of polyphosphate in Escherichia coli: a hypothetical role of YjbB in phosphate export and polyphosphate accumulation" 320 : 25-32, 2011

      56 Baek, J.H., "Novel gene members in the Pho regulon of Escherichia coli" BLACKWELL PUBLISHING 264 : 104-109, 2006

      57 Ault-Riche, D., "Novel assay reveals multiple pathways regulating stress-induced accumulations of inorganic polyphosphate in Escherichia coli" 180 : 1841-1847, 1998

      58 Kim, S.K., "Mutational analysis of the role of the first helix of region 4.2 of the sigma 70 subunit of Escherichia coli RNA polymerase in transcriptional activation by activator protein PhoB" 248 : 1-8, 1995

      59 Webb, D.C., "Mutational analysis of the Escherichia coli phosphate-specific transport system, a member of the traffic ATPase (or ABC) family of membrane transporters. A role for proline residues in transmembrane helices" 267 : 24661-24668, 1992

      60 Lamarche, M.G., "Modulation of hexa-acyl pyrophosphate lipid A population under Escherichia coli phosphate (Pho) regulon activation" 190 : 5256-5264, 2008

      61 Lamarche, M.G., "Membrane homeostasis requires intact pst in extraintestinal pathogenic Escherichia coli" 60 : 356-359, 2010

      62 Wu, H.J., "Manganese regulation of virulence factors and oxidative stress resistance in Neisseria gonorrhoeae" 73 : 899-916, 2010

      63 Fisher, S.L., "Kinetic comparison of the specificity of the vancomycin resistance VanS for two response regulators, VanR and PhoB" 35 : 4732-4740, 1996

      64 Metcalf, W.W., "Involvement of the Escherichia coli phn (psiD) gene cluster in assimilation of phosphorus in the form of phosphonates, phosphite, Pi esters, and Pi" 173 : 587-600, 1991

      65 Auesukaree, C., "Intracellular phosphate serves as a signal for the regulation of the PHO pathway in Saccharomyces cerevisiae" 279 : 17289-17294, 2004

      66 Rao, N.N., "Inorganic polyphosphate: essential for growth and survival" 78 : 605-647, 2009

      67 Rao, N.N., "Inorganic polyphosphate regulates responses of Escherichia coli to nutritional stringencies, environmental stresses and survival in the stationary phase" 23 : 183-195, 1999

      68 Ogawa, N., "Inorganic polyphosphate in Vibrio cholerae: genetic, biochemical, and physiologic features" 182 : 6687-6693, 2000

      69 Rao, N.N., "Inorganic polyphosphate in Escherichia coli: the phosphate regulon and the stringent response" 180 : 2186-2193, 1998

      70 Shi, X., "Inorganic polyphosphate in Bacillus cereus: motility, biofilm formation, and sporulation" 101 : 17061-17065, 2004

      71 Moberly, J.G., "Influence of pH and inorganic phosphate on toxicity of zinc to Arthrobacter sp. isolated from heavy-metal-contaminated sediments" 44 : 7302-7308, 2010

      72 Richards, G.R., "Induction of the Pho regulon suppresses the growth defect of an Escherichia coli sgrS mutant, connecting phosphate metabolism to the glucose-phosphate stress response" 194 : 2520-2530, 2012

      73 Critzer, F.J., "Increased transcription of the phosphate-specific transport system of Escherichia coli O157:H7 after exposure to sodium benzoate" 73 : 819-824, 2010

      74 Cesselin, B., "Inactivation of the Lactococcus lactis high-affinity phosphate transporter confers oxygen and thiol resistance and alters metal homeostasis" 155 : 2274-2281, 2009

      75 Gangaiah, D., "Importance of polyphosphate kinase 1 for Campylobacter jejuni viable-butnonculturable cell formation, natural transformation, and antimicrobial resistance" 75 : 7838-7849, 2009

      76 Reid, A.N., "Identification of Campylobacter jejuni genes contributing to acid adaptation by transcriptional profiling and genome-wide mutagenesis" 74 : 1598-1612, 2008

      77 Kuroda, A., "Guanosine tetra- and pentaphosphate promote accumulation of inorganic polyphosphate in Escherichia coli" 272 : 21240-21243, 1997

      78 Lemieux, M.J., "Glycerol-3-phosphate transporter of Escherichia coli: structure, function and regulation" 155 : 623-629, 2004

      79 Moraleda-Munoz, A., "Glycerol 3-phosphate inhibits swarming and aggregation of Myxococcus xanthus" 183 : 6135-6139, 2001

      80 Hsieh, Y.J., "Global regulation by the seven-component Pi signaling system" 13 : 198-203, 2010

      81 VanBogelen, R.A., "Global analysis of proteins synthesized during phosphorus restriction in Escherichia coli" 178 : 4344-4366, 1996

      82 Rodriguez-Garcia, A., "Genome-wide transcriptomic and proteomic analysis of the primary response to phosphate limitation in Streptomyces coelicolor M145 and in a DeltaphoP mutant" 7 : 2410-2429, 2007

      83 Yuan, Z.C., "Genome prediction of PhoB regulated promoters in Sinorhizobium meliloti and twelve proteobacteria" 34 : 2686-2697, 2006

      84 Schurdell, M.S., "Genetic evidence suggests that the intergenic region between pstA and pstB plays a role in the regulation of rpoS translation during phosphate limitation" 189 : 1150-1153, 2007

      85 Carmany, D.O., "Genetic and biochemical studies of phosphatase activity of PhoR" 185 : 1112-1115, 2003

      86 Esteban, A., "Expression of the pstS gene of Streptomyces lividans is regulated by the carbon source and is partially independent of the PhoP regulator" 8 : 201-, 2008

      87 Hoffer, S.M., "Expression of the pho regulon interferes with induction of the uhpT gene in Escherichia coli K-12" 176 : 370-376, 2001

      88 Jackson, R.J., "Expression of the PitA phosphate/metal transporter of Escherichia coli is responsive to zinc and inorganic phosphate levels" 289 : 219-224, 2008

      89 Monds, R.D., "Expression of the Pho regulon negatively regulates biofilm formation by Pseudomonas aureofaciens PA147-2" 42 : 415-426, 2001

      90 Bhatt, K., "Evidence that phosphate specific transporter is amplified in a fluoroquinolone resistant Mycobacterium smegmatis" 267 : 4028-4032, 2000

      91 Beard, S.J., "Evidence for the transport of zinc(II) ions via the pit inorganic phosphate transport system in Escherichia coli" 184 : 231-235, 2000

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