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Molecular Mechanism and Molecular Evolution of Glycosidases

Research Project

Project/Area Number 12460035
Research Category

Grant-in-Aid for Scientific Research (B)

Allocation TypeSingle-year Grants
Section一般
Research Field 応用微生物学・応用生物化学
Research InstitutionHOKKAIDO UNIVERSITY

Principal Investigator

MORI Haruhide (2001-2002)  Hokkaido Univ. Grad. School of Agr., Asso. Prof., 大学院・農学研究科, 助教授 (80241363)

千葉 誠哉 (2000)  北海道大学, 大学院・農学研究科, 教授 (30001449)

Co-Investigator(Kenkyū-buntansha) FUKUSHI Yukiharu  Hokkaido Univ. Grad. School of Agr., Lec., 大学院・農学研究科, 講師 (60218906)
KIMURA Atsuo  Hokkaido Univ. Grad. School of Agr., Prof., 大学院・農学研究科, 教授 (90186312)
森 春英  北海道大学, 大学院・農学研究科, 助手 (80241363)
Project Period (FY) 2000 – 2002
Project Status Completed (Fiscal Year 2002)
Budget Amount *help
¥14,400,000 (Direct Cost: ¥14,400,000)
Fiscal Year 2002: ¥2,800,000 (Direct Cost: ¥2,800,000)
Fiscal Year 2001: ¥2,800,000 (Direct Cost: ¥2,800,000)
Fiscal Year 2000: ¥8,800,000 (Direct Cost: ¥8,800,000)
Keywordsα-glucosidase / glycosidase / molecular evolution / グルコシダーゼ
Research Abstract

α-Glucosidase (EC 3.2.1.20), an exo-type glycosidase releasing α-glucose from substrate, is divided into two types of groups (family I and family II) that have different structures and substrate recognition. There are also the glycosidases, of which structures are homologous to each of family enzyme, such as α-amylase for family I and α-xylosidase for family II. The purpose of the project is to investigate the relationship of structure and function of two α-glucosidase families, and to understand the molecular mechanism of glycosidases. 1)The catalytic amino acid residues of two family enzymes were identified. Their functions were also analyzed. 2)The amino acids recognizing the glycon and aglycon moieties of substrate were identified. Two α-glucosidase groups showed the different recognition against the hydroxyl group in the glycon. 3)The crystal of enzyme was prepared. 4)The α-secondary kinetic isotope effects were observed, suggesting the formation of oxo-carbenium ion intermediate in the catalytic reaction. 5)The molecular analysis succeeded in elucidating the function and structure of two α-glucosidase families, and the findings are important to reveal the molecular mechanism of glycosidases homologous to family enzymes.

Report

(4 results)
  • 2002 Annual Research Report   Final Research Report Summary
  • 2001 Annual Research Report
  • 2000 Annual Research Report
  • Research Products

    (72 results)

All Other

All Publications (72 results)

  • [Publications] A.Kimura: "Molecular Anatomy of α-Glucosidase"Trends In Glycosci.Glycotechnol.. 12・68. 373-380 (2000)

    • Description
      「研究成果報告書概要(和文)」より
    • Related Report
      2002 Final Research Report Summary
  • [Publications] T.Takayanagi: "Evidence for A Single Acive Site on Isomalto-dextranase with Hydrolysis Activities of α-1,6-and α-1,4-Glucosidic Linkages"J.Appl.Glycosci.. 48・1. 55-61 (2001)

    • Description
      「研究成果報告書概要(和文)」より
    • Related Report
      2002 Final Research Report Summary
  • [Publications] M.Okuyama: "Carboxil Group of Asp-647 Residues as Possible Proton Donor in Catalytic Reaction of α-Glucosidase from Schizosaccharomyces pombe"Eur.J.Biochem.. 268・8. 2270-2280 (2001)

    • Description
      「研究成果報告書概要(和文)」より
    • Related Report
      2002 Final Research Report Summary
  • [Publications] M.Nishimoto: "Purification and Substrate Specificity of Honeybee, Apis mellifera L.,α-Glucosidase III"Biosci.Biotechnol.Biochem.. 65・7. 1610-1616 (2001)

    • Description
      「研究成果報告書概要(和文)」より
    • Related Report
      2002 Final Research Report Summary
  • [Publications] T.Mizuno: "Isolation and Sequence of Puntative α-Glucosidase Gene from Brevibacterium fuscum var.dextranlyticum Strain 0407"J.Appl.Glycosci.. 48・3. 287-291 (2001)

    • Description
      「研究成果報告書概要(和文)」より
    • Related Report
      2002 Final Research Report Summary
  • [Publications] A.Kimura: "Reaction Mechanism of Glycosylase"Trands In Glycosci.Glycotechnol.. 13・Sup. SA-B01J (2001)

    • Description
      「研究成果報告書概要(和文)」より
    • Related Report
      2002 Final Research Report Summary
  • [Publications] J.-H.Lee: "Purification and Identification of Essential Ionizable Groups of Honeybee, Apis mellifera L., Trehalase"Biosci.Biotechnol.Biochem.. 65・12. 2657-2665 (2001)

    • Description
      「研究成果報告書概要(和文)」より
    • Related Report
      2002 Final Research Report Summary
  • [Publications] Y.-M.Kim: "Production of Maltopentaose and Biochemical Characterization of Maltopentaose-Forming Amylase"J.Microbiol.Biotechnol.. 11・4. 636-643 (2001)

    • Description
      「研究成果報告書概要(和文)」より
    • Related Report
      2002 Final Research Report Summary
  • [Publications] Y.-M.Kim: "Isolation of Bacillus sp.Air-5 Producing Maltopentaose Forming Amylase and Optimization of Maltopentaose Production"Kor.J.Biotechnol.Bioeng.. 16・3. 246-252 (2001)

    • Description
      「研究成果報告書概要(和文)」より
    • Related Report
      2002 Final Research Report Summary
  • [Publications] H.Mori: "Modulation of Activity and Substrate Binding Modes by Mutation of Single and Double Subsites+1/+2 and -5/-6 of Barley α-Amylase 1"Eur.J.Biochem.. 268・24. 6545-6558 (2001)

    • Description
      「研究成果報告書概要(和文)」より
    • Related Report
      2002 Final Research Report Summary
  • [Publications] M.Nishimoto: "Study on Three α-Glucosidase Isozymes from Honeybee, Apis mellifera L."J.Appl.Glycosci.. 49・2. 191-197 (2002)

    • Description
      「研究成果報告書概要(和文)」より
    • Related Report
      2002 Final Research Report Summary
  • [Publications] M.Okuyama: "Catalytic Amino Acid Residue Providing Proton Donar in α-Glucosidase Family II"J.Appl.Glycosci.. 49・2. 211-219 (2002)

    • Description
      「研究成果報告書概要(和文)」より
    • Related Report
      2002 Final Research Report Summary
  • [Publications] M.Okuyama: "Mutant Catalyzes α-Glucosynthase-type Reaction"Biosci.Biotechnol.Biochem.. 66・4. 928-933 (2002)

    • Description
      「研究成果報告書概要(和文)」より
    • Related Report
      2002 Final Research Report Summary
  • [Publications] T.Takayanagi: "Evaluation of Subsite Affinities of Isomalto-dextranase from Arthrobacter globiformis"J.Appl.Glycosci.. 49・2. 123-127 (2002)

    • Description
      「研究成果報告書概要(和文)」より
    • Related Report
      2002 Final Research Report Summary
  • [Publications] K.Fukuda: "Kinetic Studies on Substrate Specificity and Active Site of β-D-Glucosidase F1 from Streptomyces sp."J.Appl.Glycosci.. 49・3. 265-272 (2002)

    • Description
      「研究成果報告書概要(和文)」より
    • Related Report
      2002 Final Research Report Summary
  • [Publications] K.Fukuda: "Identification of Essential Ionizable Groups and Evaluation of Subsite Affinities in the Active Site of β-D-Glucosidase F1 from Streptomyces sp."Biosci.Biotechnol.Biochem.. 66・10. 2060-2067 (2002)

    • Description
      「研究成果報告書概要(和文)」より
    • Related Report
      2002 Final Research Report Summary
  • [Publications] T.Nishio: "Glycon Specificity Profiling of α-Glucosidase Using Monodeoxy and Mono-O-methyl Derivatives of p-Nitrophenyl α-D-Glucopyranoside"Carbohydr.Res.. 337・7. 629-634 (2002)

    • Description
      「研究成果報告書概要(和文)」より
    • Related Report
      2002 Final Research Report Summary
  • [Publications] H.Mori: "Barley α-amylase Met53 situated at the high-affinity subsite-2 belongs to a substrate binding motif in the α→β loop 2 of the catalytic (β/α)_8-barrel and is critical for activity and substrate specificity"Eur.J.Biochem.. 269・22. 5377-5390 (2002)

    • Description
      「研究成果報告書概要(和文)」より
    • Related Report
      2002 Final Research Report Summary
  • [Publications] B.Svensson: "Fascinating Facets of Function and Structure of Amylolytic Enzymes of Glycoside Hydrolase Family 13"Biologia. 57.Suppl 11. 5-19 (2002)

    • Description
      「研究成果報告書概要(和文)」より
    • Related Report
      2002 Final Research Report Summary
  • [Publications] M.Son: "Evidence of Intramolecular Transglucosylation Catalyzed by an α-Glucosidase"J.Appl.Glycosci.. 50・1. 41-44 (2003)

    • Description
      「研究成果報告書概要(和文)」より
    • Related Report
      2002 Final Research Report Summary
  • [Publications] San San Mar: "Purification, Characterization, and Sequence Analysis of Two α-Amylase Isoforms from Azuki Bean, Vigna angularis, Showing Different Affinity towards β-Cyclodextrin"Biosci.Biotechnol.Biochem.. 67(印刷中). (2003)

    • Description
      「研究成果報告書概要(和文)」より
    • Related Report
      2002 Final Research Report Summary
  • [Publications] B.Svensson: "Carbohydrate Bioengineering -(Gluco)amylases what have we lern so far?-"The Royal Society of Chemistry. 9 (2001)

    • Description
      「研究成果報告書概要(和文)」より
    • Related Report
      2002 Final Research Report Summary
  • [Publications] A. Kimura: "Molecular Anatomy of α-Glucosidase"Trends In Glycosci. Glycotechnol.. 12-68. 373-380 (2000)

    • Description
      「研究成果報告書概要(欧文)」より
    • Related Report
      2002 Final Research Report Summary
  • [Publications] T. Takayanagi: "Evidence for A Single Active Site on Isomalto-dextranase with Hydrolysis Activities of α-1, 6-and α-1, 4-Glucosidic Linkages"J. Appl. Glycosci. 48-1. 55-61 (2001)

    • Description
      「研究成果報告書概要(欧文)」より
    • Related Report
      2002 Final Research Report Summary
  • [Publications] M. Okuyama: "Carboxyl Group of Asp-647Residues as Possible Proton Donor in Catalytic Reaction of α-Glucosidase from Schizosaccharomyces pombe"Eur. J. Biochem.. 268-8. 2270-2280 (2001)

    • Description
      「研究成果報告書概要(欧文)」より
    • Related Report
      2002 Final Research Report Summary
  • [Publications] M. Nishimoto: "Purification and Substrate Specificity of Honeybee, Apis mellifera L. , α-Glucosidase III"Biosci. Biotechnol. Biochem.. 65-7. 1610-1616 (2001)

    • Description
      「研究成果報告書概要(欧文)」より
    • Related Report
      2002 Final Research Report Summary
  • [Publications] T. Mizumo: "Isolation and Sequence of Puntative α-Glucosidase Gene from Brevibacterium fuscum var. dextranlyticum Strain 0407"J. Appl. Glycosci.. 48-3. 287-291 (2001)

    • Description
      「研究成果報告書概要(欧文)」より
    • Related Report
      2002 Final Research Report Summary
  • [Publications] A. Kimura: "Reaction Mechanism of Glycosylase"Trends In Glycosci. Glycotechnol.. 13- (Supplement, SA-B01J). (2001)

    • Description
      「研究成果報告書概要(欧文)」より
    • Related Report
      2002 Final Research Report Summary
  • [Publications] J. -H. Lee: "Purification and Identification of Essential Ionizable Groups of Honeybee, Apis mellifera L. , Trehalase"Biosci. Biotechnol. Biochem.. 65-12. 2657-2665 (2001)

    • Description
      「研究成果報告書概要(欧文)」より
    • Related Report
      2002 Final Research Report Summary
  • [Publications] Y. -M. Kim: "Production of Maltopentaose and Biochemical Characterization of Maltopentaose-Forming Amylase"J. Microbiol. Biotechnol.. 11-4. 636-643 (2001)

    • Description
      「研究成果報告書概要(欧文)」より
    • Related Report
      2002 Final Research Report Summary
  • [Publications] Y. -M. Kim: "Isolation of Bacillus sp. Air-5 Producing Maltopentaose Forming Amylase and Optimization of Maltopentaose Production"Kor. J. Biotechnol. Bioeng.. 16-3. 246-252 (2001)

    • Description
      「研究成果報告書概要(欧文)」より
    • Related Report
      2002 Final Research Report Summary
  • [Publications] H. Mori: "Modulation of Activity and Substrate Binding Modes by Mutation of Single and Double Subsites+1/+2 and -5/-6 of Barley α-Amylase 1"Eur. J. Biochem.. 268. 6545-6558 (2001)

    • Description
      「研究成果報告書概要(欧文)」より
    • Related Report
      2002 Final Research Report Summary
  • [Publications] B. Svensson: "(Gluco)amylases what have we lern so far?"Carbohydrate Bioengineering, The Royal Society of Chemistry. 67-75 (2001)

    • Description
      「研究成果報告書概要(欧文)」より
    • Related Report
      2002 Final Research Report Summary
  • [Publications] M. Nishimoto: "Study on Three α-Glucosidase Isozymes from Honeybee, Apis mellifera L."J. Appl. Glycosci.. 49-2. 191-197 (2002)

    • Description
      「研究成果報告書概要(欧文)」より
    • Related Report
      2002 Final Research Report Summary
  • [Publications] M. Okuyama: "Catalytic Amino Acid Residue Providing Proton Donar in α-Glucosidase Family II"J. Appl. Glycosci.. 49-2. 211-219 (2002)

    • Description
      「研究成果報告書概要(欧文)」より
    • Related Report
      2002 Final Research Report Summary
  • [Publications] M. Okuyama: "α-Glucosidase Mutant Catalyzes α-Glucosynthase-type Reaction"Biosci. Biotechnol. Biochem. 66-4. 928-933 (2002)

    • Description
      「研究成果報告書概要(欧文)」より
    • Related Report
      2002 Final Research Report Summary
  • [Publications] T. Takayanagi: "Evaluation of Subsite Affinities of Isomalto-dextranase from Arthrobacter globiformis"J. Appl. Glycosci.. 49-2. 123-127 (2002)

    • Description
      「研究成果報告書概要(欧文)」より
    • Related Report
      2002 Final Research Report Summary
  • [Publications] K. Fukuda: "Kinetic Studies on Substrate Specificity and Active Site of β-D-Glucosidase F1 from Streptomyces sp."J. Appl. Glycosci.. 49-3. 265-272 (2002)

    • Description
      「研究成果報告書概要(欧文)」より
    • Related Report
      2002 Final Research Report Summary
  • [Publications] K. Fukuda: "Identification of Essential Ionizable Groups and Evaluation of Subsite Affinities in the Active Site of β-D-Glucosidase F1 from Streptomyces sp."Biosci. Biotechnol. Biochem. 66-10. 2060-2067 (2002)

    • Description
      「研究成果報告書概要(欧文)」より
    • Related Report
      2002 Final Research Report Summary
  • [Publications] T. Nishio: "Glycon Specificity Profiling of α-Glucosidase Using Monodeoxy and Mono-O-methyl Derivatives of p-Nitrophenyl α-D-Glucopyranoside"Carbohydr. Res.. 337. 629-634 (2002)

    • Description
      「研究成果報告書概要(欧文)」より
    • Related Report
      2002 Final Research Report Summary
  • [Publications] H. Mori: "Barley α-Amylase Met53 Situated at the High-affinity Subsite-2 Belongs to a Substrate Binding Motif in the α→βLoop 2 of the Catalytic(β/α)8-Barrel and Is Critical for Activity and Substrate Specificity"Eur. J. Biochem. 269-22. 5377-5390 (2002)

    • Description
      「研究成果報告書概要(欧文)」より
    • Related Report
      2002 Final Research Report Summary
  • [Publications] B. Svensson: "Fascinating Facets of Function and Structure of Amylolytic Enzymes of Glycoside Hydrolase Family 13"Biologia. 57-Suppl 11. 5-19 (2002)

    • Description
      「研究成果報告書概要(欧文)」より
    • Related Report
      2002 Final Research Report Summary
  • [Publications] M. Son: "Evidence of Intramolecular Transglucosylation Catalyzed by an α-Glucosidase"J. Appl. Glycosci.. 50-1. 41-44 (2003)

    • Description
      「研究成果報告書概要(欧文)」より
    • Related Report
      2002 Final Research Report Summary
  • [Publications] San San Mar: "Purification, Characterization, and Sequence Analysis of Two α-Amylase Isoforms from Azuki Bean, Vigna angularis, Showing Different Affinity towards β-Cyclodextrin"Biosci. Biotechnol. Biochem.. 67. in press (2003)

    • Description
      「研究成果報告書概要(欧文)」より
    • Related Report
      2002 Final Research Report Summary
  • [Publications] M.Okuyama: "Catalytic Amino Acid Residue Providing Proton Donar in α-Glucosidase Family II"J. Appl. Glycosci.. 49・2. 211-219 (2002)

    • Related Report
      2002 Annual Research Report
  • [Publications] M.Nishimoto: "Study on Three α-Glucosidase Isozymes from Honeybee, Apis mellifera L"J. Appl. Glycosci.. 49・2. 191-197 (2002)

    • Related Report
      2002 Annual Research Report
  • [Publications] M.Okuyama: "Mutant Catalyzes α-Glucosynthase-type Reaction"Biosci. Biotechnol. Biochem.. 66・4. 928-933 (2002)

    • Related Report
      2002 Annual Research Report
  • [Publications] T.Takayanagi: "Evaluation of Subsite Affinities of Isomalto-dextranase from Arthrobacter globiformis"J. Appl. Glycosci.. 49・2. 123-127 (2002)

    • Related Report
      2002 Annual Research Report
  • [Publications] K.Fukuda: "Kinetic Studies on Substrate Specificity and Active Site of β-D-Glucosidase F1 from Streptomyces sp"J. Appl. Glycosci.. 49・3. 265-272 (2002)

    • Related Report
      2002 Annual Research Report
  • [Publications] K.Fukuda: "Identification of Essential Ionizable Groups and Evaluation of Subsite Affinities in the Active Site of j3-D-Glucosidase F1 from Streptomyces sp"Biosci. Biotechnol. Biochem.. 66・10. 2060-2067 (2002)

    • Related Report
      2002 Annual Research Report
  • [Publications] T.Nishio: "Glycon Specificity Profiling of α-Glucosidase Using Monodeoxy and Mono-O-methyl Derivatives of p-Nitrophenyl α-D-Glucopyranoside"Carbohydr. Res.. 337・7. 629-634 (2002)

    • Related Report
      2002 Annual Research Report
  • [Publications] H.Mori: "Barley α-Amylase Met53 Situated at the High-affinity Subsite -2 Belongs to a Subsrate Binding Motif in the α→β Loop 2 of the Catalytic (β/α)_8-Barre and Is Critical for Activity and Substrate Specificity"Eur. J. Biochem.. 269・22. 5377-5390 (2002)

    • Related Report
      2002 Annual Research Report
  • [Publications] B.Svensson: "Fascinating Facets of Function and Structure of Amylolytic Enzymes of Glycoside Hydrolase Family 13"Biologia. 57・Suppl 11. 5-19 (2002)

    • Related Report
      2002 Annual Research Report
  • [Publications] M.Son: "Evidence of Intramolecular Transglucosylation Catalyzed by an α-Glucosidase"J. Appl. Glycosci.. 50・1. 41-44 (2003)

    • Related Report
      2002 Annual Research Report
  • [Publications] San San Mar: "Purification, Characterization, and Sequence Analysis of Two α-Amylase Isoforms from Azuki Bean, Vigna angularis, Showing Different Affinity towards β-Cyclodextrin"Biosci. Biotechnol. Biochem.. 67(印刷中). (2003)

    • Related Report
      2002 Annual Research Report
  • [Publications] M.Okuyama: "Carboxyl Group of Asp-647 Residues as Possible Proton Donorin Catalytic Activities of α-1,6- and α-1,4-Glucosidic Linkages"Eur.J.Biochem.. 268・8. 2270-2280 (2001)

    • Related Report
      2001 Annual Research Report
  • [Publications] M.Nishimoto: "Purification and Substrate Specificity of Honeybee, Apismellifera L., α-Glucosidase III"Biosci.Biotechnol.Biochem.. 65・7. 1610-1616 (2001)

    • Related Report
      2001 Annual Research Report
  • [Publications] T.Mizuno: "Isolation and Sequence of α-Glucosidase Gene from Brevibacterium fuscum var. dextranlyticum Strain 0407"J.Appl.Glycosci. 48・3. 287-291 (2001)

    • Related Report
      2001 Annual Research Report
  • [Publications] A.Kimura: "Reaction Mechanism of Glycosylase"Trends In Glycosci.Glycotechnol.. 13. SA-BO1J-SA-B01J (2001)

    • Related Report
      2001 Annual Research Report
  • [Publications] J.-H.Lee: "Purification and Identification of Essential Ionizable Groups of Honeybee, Apismellifera L., Trehalase"Biosci.Biotechnol.Biochem.. 65・12. 2657-2665 (2001)

    • Related Report
      2001 Annual Research Report
  • [Publications] Y.-M.Kim: "Production of Maltopentaose and Biochemical Characterization of Maltopentaose-Forming Amylase"J.Microbiol.Biotechnol.. 11・4. 636-643 (2001)

    • Related Report
      2001 Annual Research Report
  • [Publications] H.Mori: "Modulation of Activity and Substrate Binding Modes by Mutation of Single and Double Subsites +1/+2 and -5/-6 of Barley α-Amylase 1"Eur.J.Biochem.. 268・24. 6545-6558 (2001)

    • Related Report
      2001 Annual Research Report
  • [Publications] M.Okuyama: "Catalytic Amino Acid Residue Providing Proton Donar in α-Glucosidase Family II"J.Appl.Glycosci.. (印刷中). (2002)

    • Related Report
      2001 Annual Research Report
  • [Publications] M.Okuyama: "α-Glucosidase Mutant Catalyzes α-Glucosynthase-type Reaction"Biosci.Biotechnol.Biochem.. (印刷中). (2002)

    • Related Report
      2001 Annual Research Report
  • [Publications] T.Takayanagi: "Evaluation of Subsite Affinities of Isomalto-dextranase from Arthrobacter globiformis"J.Appl.Glycosci.. (印刷中). (2002)

    • Related Report
      2001 Annual Research Report
  • [Publications] K.Fukuda: "Kinetic Studies on Substrate Specificity and Active Site of β-D-Glucosidase F_I from Streptomyces sp."J.Appl.Glycosci.. (印刷中). (2002)

    • Related Report
      2001 Annual Research Report
  • [Publications] M.Nishimoto: "Study on Three α-Glucosidase Isozymes from Honeybee,Apis mellifera L."J.Appl.Glycosci.. (印刷中). (2002)

    • Related Report
      2001 Annual Research Report
  • [Publications] A.Kimura: "Molecular Anatomy of α-Glucosidase"Trends In Glycosci.Glycotechnol.. 12・68. 373-380 (2000)

    • Related Report
      2000 Annual Research Report
  • [Publications] T.Takayanagi: "Evidence for A Single Active Site on Isomalto-dextranase with Hydrolysis Activities of α-1, 6-and α-1, 4-Glucosidic Linkages"J.Appl.Glycosci.. 48・1. 55-61 (2001)

    • Related Report
      2000 Annual Research Report
  • [Publications] M.Okuyama: "Carboxyl Group of Asp-647 Residues as Possible Proton Donorin Catalytic Reaction of α-Glucosidase from Schizosaccharomycespombe"Eur.J.Biochem.. (印刷中). (2001)

    • Related Report
      2000 Annual Research Report
  • [Publications] M.Nishimoto: "Purification and Substrate Specificity of Honeybee, Apismellifera L., α-Glucosidase III "Biosci.Biotechnol.Biochem.. (印刷中). (2001)

    • Related Report
      2000 Annual Research Report
  • [Publications] T.Mizuno: "Isolation and Sequence of α-Glucosidase Gene from Brevibacterium fuscum var. dextranlyticum Strain 0407"J.Appl.Glycosci.. (印刷中). (2001)

    • Related Report
      2000 Annual Research Report

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Published: 2000-04-01   Modified: 2016-04-21  

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