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Development of Chemical Analysis for Metabolic Intermediates

Research Project

Project/Area Number 12206006
Research Category

Grant-in-Aid for Scientific Research on Priority Areas

Allocation TypeSingle-year Grants
Review Section Biological Sciences
Research InstitutionKyoto University

Principal Investigator

NISHIOKA Takaaki  Kyoto University, Graduate School of Agriculture, Professor, 農学研究科, 教授 (80026559)

Co-Investigator(Kenkyū-buntansha) TERABE Shigeru  University of Hyoto, Graduate School of Material Science, Professor, 物質理学研究科, 教授 (50115888)
Project Period (FY) 2000 – 2004
Project Status Completed (Fiscal Year 2004)
Budget Amount *help
¥89,200,000 (Direct Cost: ¥89,200,000)
Fiscal Year 2004: ¥21,600,000 (Direct Cost: ¥21,600,000)
Fiscal Year 2003: ¥21,600,000 (Direct Cost: ¥21,600,000)
Fiscal Year 2002: ¥21,000,000 (Direct Cost: ¥21,000,000)
Fiscal Year 2001: ¥25,000,000 (Direct Cost: ¥25,000,000)
Keywordsmetabolomics / metabolism / capillary electrophoresis / metabolite profile / homeostasis / CE / MS / ESI-IontrapMS / ESI-MS / TOFMS / 2次元分離 / LC / メタボローム解析 / 枯草菌 / 医療診断 / 創薬 / キャピラリー内濃縮 / pHジャンクション / ハイスループット化学分析 / レーザー励起蛍光 / アミノ酸 / 有機酸 / ゲノム / ネットワーク / 代謝物
Research Abstract

Most of the metabolites in living cells or tissues are polar and ionic substances that are not suitable for chemical analysis by using conventional methods such as GC-and LC-MS. We have successfully developed capillary electrophoresis coupled with mass spectrometry (CE-MS) as a method of metabolomics. We applied the method to analyze the metabolic intermediates of the central energy metabolism of the Bacillus subtilis cells and compared the metabolite profiles that schematically show the intercellular amount of the metabolites on the central metabolic pathways of the cells cultured under various environmental and genetic perturbations. The metabolite profiles were globally same when the rates of the cell division were the same under the different perturbations. This suggests that B. subtilis has a metabolite profile that is optimized for its cell division and that it maintain the profile against the environmental and genetic perturbations. At the same times when we analyzed the metabolites, we measured the gene expressions by using the DNA microarray of B. subtilis. Gene expression data were overlaid on the metabolite profiles. When perturbations were not so different from the control experiment, amount of the expressions of several enzyme genes were regulated. On the others where perturbations were quite different, B. subtilis changed not only gene expressions but also metabolic pathways; it used a quite different metabolic pathways from those of the control experiment. These results suggests that the metabolite profile is quantitatively and qualitatively regulated; this must be a way of "homeostasis". It is most likely that each living species has its own metabolite profile for cell division.

Report

(6 results)
  • 2004 Annual Research Report   Final Research Report Summary
  • 2003 Annual Research Report
  • 2002 Annual Research Report
  • 2001 Annual Research Report
  • 2000 Annual Research Report
  • Research Products

    (59 results)

All 2005 2004 2003 2002 2001 Other

All Journal Article (39 results) Book (6 results) Publications (14 results)

  • [Journal Article] Large-Scale Prediction of Cationic Metabolite Identity and Migration Time in Capillary Electrophoresis Mass Spectrometry Using Artificial Neural Networks2005

    • Author(s)
      Sugimoto M
    • Journal Title

      Anal Chem. 77

      Pages: 78-84

    • Description
      「研究成果報告書概要(和文)」より
    • Related Report
      2004 Final Research Report Summary
  • [Journal Article] メタポローム-ゲノム情報と環境の相互作用システム2005

    • Author(s)
      西岡孝明
    • Journal Title

      蛋白質・核酸・酵素 50

      Pages: 2198-2203

    • Description
      「研究成果報告書概要(和文)」より
    • Related Report
      2004 Final Research Report Summary
  • [Journal Article] Large-Scale Prediction of Cationic Metabolite Identity and Migration Time in Capillary Electrophoresis Mass Spectrometry Using Artificial Neural Networks.2005

    • Author(s)
      Sugimoto M, Kikuchi S, Arita M, Soga T, Nishioka T, Tomita M.
    • Journal Title

      Anal Chem. 77(1)

      Pages: 78-84

    • Description
      「研究成果報告書概要(欧文)」より
    • Related Report
      2004 Final Research Report Summary
  • [Journal Article] Large-Scale Prediction of Cationic Metabolite Identity and Migration Time in Capillary Electrophoresis Mass Spectrometry Using Artificial Neural Networks.2005

    • Author(s)
      Sugimoto M, Kikuchi S, Arita M, Soga T, Nishioka T, Tomita M.
    • Journal Title

      Anal Chem. 77

      Pages: 78-84

    • Related Report
      2004 Annual Research Report
  • [Journal Article] Two- dimensional separation method for analysis of Bacillus subtilis metabolites via hyphenation of micro-liquid hromatography and capillary electrophoresis2004

    • Author(s)
      Jia, L.
    • Journal Title

      Anal. Chem. 76

      Pages: 1419-142

    • Description
      「研究成果報告書概要(和文)」より
    • Related Report
      2004 Final Research Report Summary
  • [Journal Article] Simultaneous determination of the main metabolites in rice leaves using capillary electrophoresis mass spectrometry and capillary electrophoresis diode array detection2004

    • Author(s)
      Sato, S
    • Journal Title

      Plant J 40

      Pages: 151-163

    • Description
      「研究成果報告書概要(和文)」より
    • Related Report
      2004 Final Research Report Summary
  • [Journal Article] Qualitative and quantitative analysis of amino acids by capillary electrophoresis electrospray ionization tandem mass spectrometry2004

    • Author(s)
      Soga, T.
    • Journal Title

      Electrophoresis 25

      Pages: 1964-197

    • Description
      「研究成果報告書概要(和文)」より
    • Related Report
      2004 Final Research Report Summary
  • [Journal Article] Two-dimensional separation method for analysis of Bacillus subtilis metabolites via hyphenation of micro-liquid chromatography and capillary electrophoresis2004

    • Author(s)
      Jia, L., Liu, B.-F., Terabe, S., Nishioka, T.
    • Journal Title

      Anal.Chem. 76

      Pages: 1419-1428

    • Description
      「研究成果報告書概要(欧文)」より
    • Related Report
      2004 Annual Research Report 2004 Final Research Report Summary
  • [Journal Article] Simultaneous determination of the main metabolites in rice leaves using capillary electrophoresis mass spectrometry and capillary electrophoresis diode array detection.2004

    • Author(s)
      Sato, S., Soga, T., Nishioka, T., Tomita, M.
    • Journal Title

      Plant J 40

      Pages: 151-163

    • Description
      「研究成果報告書概要(欧文)」より
    • Related Report
      2004 Annual Research Report 2004 Final Research Report Summary
  • [Journal Article] Qualitative and quantitative analysis of amino acids by capillary electrophoresis electrospray ionization tandem mass spectrometry2004

    • Author(s)
      Soga, T., Kakazu, Y., Robert, M., Tomita, M., Nishioka, T.
    • Journal Title

      Electrophoresis 25

      Pages: 1964-1972

    • Description
      「研究成果報告書概要(欧文)」より
    • Related Report
      2004 Annual Research Report 2004 Final Research Report Summary
  • [Journal Article] メタボローム解析分析法の開発から応用まで2004

    • Author(s)
      西岡孝明
    • Journal Title

      バイオサインスとインダストリー 62

      Pages: 229-233

    • NAID

      10013499958

    • Related Report
      2004 Annual Research Report
  • [Journal Article] 微生物のメタボーローム解析からわかること2004

    • Author(s)
      西岡孝明
    • Journal Title

      日本農芸化学会誌 78

      Pages: 568-571

    • NAID

      10013140487

    • Related Report
      2004 Annual Research Report
  • [Journal Article] Quantitative metabolome analysis using capillary electrophoresis mass spectrometry2003

    • Author(s)
      Soga, T.
    • Journal Title

      J. Proteome Res. 2

      Pages: 488-494

    • Description
      「研究成果報告書概要(和文)」より
    • Related Report
      2004 Final Research Report Summary
  • [Journal Article] Picomolar analysis of flavins in biological samples by dynamic pH junction-sweeping capillary electrophoresis with laser-induced fluorescence detection2003

    • Author(s)
      Britz-McKibbin, P.
    • Journal Title

      Anal. Biochem. 313

      Pages: 89-96

    • Description
      「研究成果報告書概要(和文)」より
    • Related Report
      2004 Final Research Report Summary
  • [Journal Article] Sensitive and high-throughput analyses of purine metabolites by dynamic pH junction multiplexed capillary electrophoresis : A new tool for metabolomics studies2003

    • Author(s)
      Britz-Mckibbin, P.
    • Journal Title

      Anal. Sci 19

      Pages: 99-104

    • Description
      「研究成果報告書概要(和文)」より
    • Related Report
      2004 Final Research Report Summary
  • [Journal Article] Determination of pyridine and adenine nucleotide metabolites in Bacillus subtilis cell extract by sweeping borate complexation capillary electrophoresis2003

    • Author(s)
      Markuszewski, M.J.
    • Journal Title

      J. Chromatogr. A 989

      Pages: 293-301

    • Description
      「研究成果報告書概要(和文)」より
    • Related Report
      2004 Final Research Report Summary
  • [Journal Article] Analysis of carboxylic acid metabolites from the tricarboxylic acid cycle in Bacillus subtilis cell extract by capillary electrophoresis using an indirect photometric detection method2003

    • Author(s)
      Markuszewski, M.J.
    • Journal Title

      J. Chromatogr. A. 1010

      Pages: 113-121

    • Description
      「研究成果報告書概要(和文)」より
    • Related Report
      2004 Final Research Report Summary
  • [Journal Article] Mechanistic Study of Analyte Focusing by Dynamic pH Junction in Capillary Electrophoresis Using Computer Simulation2003

    • Author(s)
      Kim, J-B.
    • Journal Title

      Anal. Chem. 75

      Pages: 3986-3993

    • Description
      「研究成果報告書概要(和文)」より
    • Related Report
      2004 Final Research Report Summary
  • [Journal Article] Complementary On-line Preconcentration Strategies for Steroids by Capillary Electrophoresis.2003

    • Author(s)
      Britz-McKibbin, P
    • Journal Title

      J. Chromatogr. A 1013

      Pages: 65-76

    • Description
      「研究成果報告書概要(和文)」より
    • Related Report
      2004 Final Research Report Summary
  • [Journal Article] On-Line Sample Preconcentration of Cationic Analytes by Dynamic pH Junction in Capillary Electrophoresis.2003

    • Author(s)
      Kim, J-B
    • Journal Title

      J. Chromatogr. A 1018

      Pages: 251-256

    • Description
      「研究成果報告書概要(和文)」より
    • Related Report
      2004 Final Research Report Summary
  • [Journal Article] Subsecond Separa-tion of Cellular Flavin Coenzymes by Microchip Capillary Electrophoresis with Laser-Induced Fluorescence Detection.2003

    • Author(s)
      Liu, B.-F
    • Journal Title

      J. Chromatogr. A 1021

      Pages: 201-207

    • Description
      「研究成果報告書概要(和文)」より
    • Related Report
      2004 Final Research Report Summary
  • [Journal Article] ダイナミックな生命現象のメタボローム解析技術2003

    • Author(s)
      曽我朋義
    • Journal Title

      細胞工学 22

      Pages: 1337-1341

    • Description
      「研究成果報告書概要(和文)」より
    • Related Report
      2004 Final Research Report Summary
  • [Journal Article] Quantitative metabolome analysis using capillary electrophoresis mass spectrometry.2003

    • Author(s)
      Soga, T., Ohashi, Y., Ueno, Y., Naraoka, H., Tomita, M., Nishioka, T.
    • Journal Title

      J.Proteome Res. 2

      Pages: 488-494

    • Description
      「研究成果報告書概要(欧文)」より
    • Related Report
      2004 Final Research Report Summary
  • [Journal Article] Picomolar analysis of flavins in biological samples by dynamic pH junction-sweeping capillary electrophoresis with laser-induced fluorescence detection2003

    • Author(s)
      Britz-McKibbin, P., Markuszewski, M.J., Iyanahi, T., Matsuda, K., Nishioka, T., Terabe, S.
    • Journal Title

      Anal.Biochem. 313

      Pages: 89-96

    • Description
      「研究成果報告書概要(欧文)」より
    • Related Report
      2004 Final Research Report Summary
  • [Journal Article] Sensitive and high-throughput analyses of purine metabolites by dynamic pH junction multiplexed capillary electrophoresis : A new tool for metabolomics studies2003

    • Author(s)
      Britz-Mckibbin, P., Nishioka, T., Terabe, S.
    • Journal Title

      Anal.Sci. 19

      Pages: 99-104

    • NAID

      10011102976

    • Description
      「研究成果報告書概要(欧文)」より
    • Related Report
      2004 Final Research Report Summary
  • [Journal Article] Determination of pyridine and adenine nucleotide metabolites in Bacillus subtilis cell extract by sweeping borate complexation capillary electrophoresis2003

    • Author(s)
      Markuszewski, M.J., Britz-McKibbin, P., Terabe, S., Matsuda, K., Nishioka, T.
    • Journal Title

      J.Chromatogr.A 989

      Pages: 293-301

    • NAID

      10011844131

    • Description
      「研究成果報告書概要(欧文)」より
    • Related Report
      2004 Final Research Report Summary
  • [Journal Article] Analysis of carboxylic acid metabolites from the tricarboxylic acid cycle in Bacillus subtilis cell extract by capillary electrophoresis using an indirect photometric detection method2003

    • Author(s)
      Markuszewski, M.J., Otsuka, K., Terabe, S., Matsuda, K, Nishioka, T.
    • Journal Title

      J.Chromatogr.A. 1010

      Pages: 113-121

    • Description
      「研究成果報告書概要(欧文)」より
    • Related Report
      2004 Final Research Report Summary
  • [Journal Article] Mechanistic Study of Analyte Focusing by Dynamic pH Junction in Capillary Electrophoresis Using Computer Simulation.2003

    • Author(s)
      Kim, J-B., Britz-McKibbin, P., Hirokawa, T., Terabe, S.
    • Journal Title

      Anal.Chem. 75

      Pages: 3986-3993

    • Description
      「研究成果報告書概要(欧文)」より
    • Related Report
      2004 Final Research Report Summary
  • [Journal Article] Complementary On-Line Preconcentration Strategies for Steroids by Capillary Electrophoresis.2003

    • Author(s)
      Britz-McKibbin, P., Ichihashi, T., Tsubota, Chen, D.D.Y., Terabe, S.
    • Journal Title

      J.Chromatogr.A 1013

      Pages: 65-76

    • Description
      「研究成果報告書概要(欧文)」より
    • Related Report
      2004 Final Research Report Summary
  • [Journal Article] On-Line Sample Preconcentration of Cationic Analytes by Dynamic pH Junction in Capillary Electrophoresis.2003

    • Author(s)
      Kim, J-B., Okamoto, Y., Terabe, S.
    • Journal Title

      J.Chromatogr.A 1018

      Pages: 251-256

    • Description
      「研究成果報告書概要(欧文)」より
    • Related Report
      2004 Final Research Report Summary
  • [Journal Article] Subsecond Separation of Cellular Flavin Coenzymes by Microchip Capillary Electrophoresis with Laser-Induced Fluorescence Detection.2003

    • Author(s)
      Liu, B.-F., Hisamoto, H., Terabe, S.
    • Journal Title

      J.Chromatogr.A 1021

      Pages: 201-207

    • Description
      「研究成果報告書概要(欧文)」より
    • Related Report
      2004 Final Research Report Summary
  • [Journal Article] Simultaneous determination of anionic intermediates for Bacillus subtilis metabolic pathways by capillary electrophoresis electrospray ionization mass spectrometry2002

    • Author(s)
      Soga, T
    • Journal Title

      Anal. Chem. 74

      Pages: 2233-2239

    • Description
      「研究成果報告書概要(和文)」より
    • Related Report
      2004 Final Research Report Summary
  • [Journal Article] Pressure-assisted capillary electro-phoresis electrospray ionization mass spectrometry for analysis of multivalent anions2002

    • Author(s)
      Soga, T
    • Journal Title

      Anal. Chem. 74

      Pages: 6224-6229

    • Description
      「研究成果報告書概要(和文)」より
    • Related Report
      2004 Final Research Report Summary
  • [Journal Article] On-line Focusing of Flavin Derivatives Using Dynamic pH Junction-Sweeping Capillary Electrophoresis with Laser-Induced Fluorescence Detection2002

    • Author(s)
      Britz-McKibbin, P
    • Journal Title

      Anal. Chem. 74

      Pages: 3736-3743

    • Description
      「研究成果報告書概要(和文)」より
    • Related Report
      2004 Final Research Report Summary
  • [Journal Article] Simultaneous determination of anionic intermediates for Bacillus subtilis metabolic pathways by capillary electrophoresis electrospray ionization mass spectrometry2002

    • Author(s)
      Soga, T., Ueno, Y., Naraoka, H., Ohashi, Y., Tomita, M., Nishioka, T.
    • Journal Title

      Anal.Chem. 74

      Pages: 2233-2239

    • Description
      「研究成果報告書概要(欧文)」より
    • Related Report
      2004 Final Research Report Summary
  • [Journal Article] Pressure-assisted capillary electro-phoresis electrospray ionization mass spectrometry for analysis of multivalent anions2002

    • Author(s)
      Soga, T., Ueno.Y., Naraoka, H., Matsuda, K., Tomita, M., Nishioka, T.
    • Journal Title

      Anal.Chem. 74

      Pages: 6224-6229

    • Description
      「研究成果報告書概要(欧文)」より
    • Related Report
      2004 Final Research Report Summary
  • [Journal Article] On-Line Focusing of Flavin Derivatives Using Dynamic pH Junction-Sweeping Capillary Electrophoresis with Laser-Induced Fluorescence Detection2002

    • Author(s)
      Britz-McKibbin, P., Otsuka, K., Terabe, S.
    • Journal Title

      Anal.Chem. 74

      Pages: 3736-3743

    • Description
      「研究成果報告書概要(欧文)」より
    • Related Report
      2004 Final Research Report Summary
  • [Journal Article] Capillary Electrophoretic Techniques toward the Metabolome Analysis2001

    • Author(s)
      Terabe, S.
    • Journal Title

      Pure Appl. Chem 73

      Pages: 1563-1572

    • Description
      「研究成果報告書概要(和文)」より
    • Related Report
      2004 Final Research Report Summary
  • [Journal Article] Capillary Electrophoretic Techniques toward the Metabolome Analysis2001

    • Author(s)
      Terabe, S., Markuszewski, M.J., Inoue, N., Otsuka, K., Nishioka, T.
    • Journal Title

      Pure Appl.Chem. 73

      Pages: 1563-1572

    • Description
      「研究成果報告書概要(欧文)」より
    • Related Report
      2004 Final Research Report Summary
  • [Book] Metabolomics. The Frontier of Systems Biolog2005

    • Author(s)
      Tomita, M., Nishioka, T.(Eds)
    • Total Pages
      256
    • Publisher
      Springer-Verlag, Tokyo
    • Description
      「研究成果報告書概要(和文)」より
    • Related Report
      2004 Final Research Report Summary
  • [Book] The Frontier of Metabolomics2005

    • Author(s)
      Tomita, M., Nishioka, T.(Eds)
    • Total Pages
      222
    • Publisher
      Springer-Verlag (Tokyo)
    • Description
      「研究成果報告書概要(欧文)」より
    • Related Report
      2004 Final Research Report Summary
  • [Book] Metabomics - a system of genome interacting with environment-2005

    • Author(s)
      Nishioka, T., Terabe, S., Soga, T., Matsuda, K., Fujita, Y.
    • Publisher
      Tanpakushitsu Kakusan Kouso
    • Description
      「研究成果報告書概要(欧文)」より
    • Related Report
      2004 Final Research Report Summary
  • [Book] Metabolomics. The Frontier of Systems Biology2005

    • Author(s)
      Tomita, M., Nishioka, T.(Eds)
    • Total Pages
      256
    • Publisher
      Springer-Verlag (Tokyo)
    • Description
      「研究成果報告書概要(欧文)」より
    • Related Report
      2004 Final Research Report Summary
  • [Book] メタボローム研究の最前線2003

    • Author(s)
      冨田 勝, 西岡孝明共編
    • Total Pages
      222
    • Publisher
      シュプリンガー・フェアラーク東京
    • Description
      「研究成果報告書概要(和文)」より
    • Related Report
      2004 Final Research Report Summary
  • [Book] Dynamic ascects of the life by using metabolomics (in Japanese)2003

    • Author(s)
      Soga, T., Sato, S, Nishioka, T., Tomita, M.
    • Publisher
      Saibou Kogaku
    • Description
      「研究成果報告書概要(欧文)」より
    • Related Report
      2004 Final Research Report Summary
  • [Publications] Britz-McKibbin, P, Nishioka, T., Terabe, S.: "Picomolar analysis of flavins in biological samples by dynamic pH junction-sweeping capillary electrophoresis with laser-induced fluorescence detection"Analytical Biochemistry. 313. 89-96 (2003)

    • Related Report
      2003 Annual Research Report
  • [Publications] Britz-Mckibbin, P., Nishioka, T., Terabe, S.: "Sensitive and high-throughput analyses of purine metabolites by dynamic pH junction multiplexed capillary electrophoresis : A new tool for metabolomics studies"Analytical Science. 19. 99-104 (2003)

    • Related Report
      2003 Annual Research Report
  • [Publications] Markuszewski, M.J., Terabe, S., Nishioka, T.: "Determination of pyridine and adenine nucleotide metabolites in Bacillus subtilis cell extract by sweeping borate complexation capillary electrophoresis"Journal of Chromatography. A,989. 293-301 (2003)

    • Related Report
      2003 Annual Research Report
  • [Publications] Soga, T., Nishioka, T.: "Quantitative metabolome analysis using capillary electrophoresis mass spectrometry"Journal of Proteome Research. 2. 488-494 (2003)

    • Related Report
      2003 Annual Research Report
  • [Publications] Markuszewski, M.J., Nishioka, T.: "Analysis of carboxylic acid metabolites from the tricarboxylic acid cycle in Bacillus subtilis cell extract by capillary electrophoresis using an indirect photometric detection method"Journal of Chromatography. A,1010. 113-121 (2003)

    • Related Report
      2003 Annual Research Report
  • [Publications] 冨田 勝, 西岡 孝明(共編): "メタボローム研究の最前線"シュプリンガー・フェアラーク東京. 222 (2003)

    • Related Report
      2003 Annual Research Report
  • [Publications] Soga, T.: "Simultaneous determination of anionic intermediates for Bacillus subtilis metabolic pathways by capillary electrophoresis electrospray ionization mass spectrometry"Analytical Chemistry. 74. 2233-2239 (2002)

    • Related Report
      2002 Annual Research Report
  • [Publications] Soga, T.: "Pressure-assisted capillary electrophoresis electrospray ionization mass spectrometry for analysis of multivalent anions"Analytical Chemistry. 74. 6224-6229 (2002)

    • Related Report
      2002 Annual Research Report
  • [Publications] Britz-McKibbin, P.: "Picomolar analysis of flavins in biological samples by dynamic pH junction-sweeping capillary electroplioresis with laser-induced fluorescence detection"Analytical Biochemistry. 313. 89-96 (2003)

    • Related Report
      2002 Annual Research Report
  • [Publications] Britz-McKibbin, P.: "Sensitive and high-throughput analyses of purine metabolites by dynamic pH junction multiplexed capillary electrophoresis : A new tool for metabolomics studies"Analytical Science. 19. 99-104 (2003)

    • Related Report
      2002 Annual Research Report
  • [Publications] Markuszewski, M.J: "Determination of pyridine and adenine nucleotide metabolites in Bacillus subtilis cell extract by sweeping borate complexation capillary electrophoresis"Journal of Chromatography, A. 989. 293-301 (2003)

    • Related Report
      2002 Annual Research Report
  • [Publications] S.Terabe.3名, T.Nishioka: "Capillary Electrophoretic Techniques toward the Metabolome Analysis"Pure & Appl. Chem.. 73. 1563-1572 (2001)

    • Related Report
      2001 Annual Research Report
  • [Publications] 寺部 茂, 4名, 西岡孝明: "細胞内代謝中間産物分析のためのオンライン試料濃縮キャピラリー電気泳動"生物物理化学. 45. 117-121 (2001)

    • Related Report
      2001 Annual Research Report
  • [Publications] Terabe,S: "Comprehensive analysis of metabolic intermediates by CE"Proceedings of HPCE 2001. 19-19 (2001)

    • Related Report
      2000 Annual Research Report

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Published: 2001-04-01   Modified: 2018-03-28  

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