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RECOMBINANT DNA TECHNIQUES and PROTEIN ENGINEERING

RECOMBINANT DNA TECHNIQUES and PROTEIN ENGINEERING. Introduction , historic milestones. Szilágyi László Ph.D. Dept. of Biochemistry (5.313.) szilagyl@elte.hu home page: www.szilagyl.web.elte.hu/GT-eng. List of lectures. Introduction; historic milestones Enzymes used in GT

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RECOMBINANT DNA TECHNIQUES and PROTEIN ENGINEERING

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  1. RECOMBINANT DNA TECHNIQUESandPROTEIN ENGINEERING Introduction, historicmilestones

  2. Szilágyi László Ph.D. • Dept. of Biochemistry (5.313.) • szilagyl@elte.hu • home page: www.szilagyl.web.elte.hu/GT-eng

  3. List of lectures • Introduction; historic milestones • Enzymes used in GT • Purification and analysis of nucleic acids • Polymerase chain reaction • Vectors & cloning strategies • Mutagenesis • Methods to study gene expression; next generation sequencing methods • Expression systems I. • Expression systems II.

  4. HISTORIC MILESTONES: 1869F. MIESCHER: DNA isolation 1886T. ESCHERICH: E. coli 1944O. AVERY: DNAis the genetic material 1953J. WATSON F. CRICK, M. WILKINS (Nobel prize, 1962) & R. FRANKLIN:DNA double helix 1957A. KORNBERG (1959):DNA polymerase 1961 J. MARMUR & P. DOTY: DNA renaturation and hybridization 1966 R. HOLEY, M. NIRENBERG, H.G. KHORANA, S. OCHOA(1968, 1959): genetic code 1962 W. ARBER (1978): bacterial restriction-modificaton 1967 M. GELLERT: DNA ligase 1970 H. SMITH (1978): first restriction endonuclease(HindII) 1972 D. NATHANS(1978):first restriction map(SV40) 1972 P. BERG(1980):first ligation 1973 S. COHEN & H. BOYER (1986): first recombinant DNA (E. coli pSC101 TetR + S. aureus pI250 AmpR) 1975 Asilomar conference - moratorium 1975E. SOUTHERN:DNA blotting -hybridization 1975G. KÖHLER & C. MILSTEIN (1984): monoclonal antibodies Miescher Friedrich. „Über die chemische Zusammensetzung der Eiterzellen." Hoppe-Seyler's medicinisch-chemische Untersuchungen4 (1871): 441-460

  5. HISTORIC MILESTONES: 1869F. MIESCHER: DNA isolation 1881 Ludwig Karl Martin Leonhard Albrecht KOSSEL(1910) DNA chemical structure 1886T. ESCHERICH: E. coli 1944O. AVERY: DNAis the genetic material 1953J. WATSON F. CRICK, M. WILKINS (Nobel prize, 1962) & R. FRANKLIN:DNA double helix 1957A. KORNBERG (1959):DNA polymerase 1961 J. MARMUR & P. DOTY: DNA renaturation and hybridization 1966 R. HOLEY, M. NIRENBERG, H.G. KHORANA, S. OCHOA(1968, 1959): genetic code 1962 W. ARBER (1978): bacterial restriction-modificaton 1967 M. GELLERT: DNA ligase 1970 H. SMITH (1978): first restriction endonuclease(HindII) 1972 D. NATHANS(1978):first restriction map(SV40) 1972 P. BERG(1980):first ligation 1973 S. COHEN & H. BOYER (1986): first recombinant DNA (E. coli pSC101 TetR + S. aureus pI250 AmpR) 1975 Asilomar conference - moratorium 1975E. SOUTHERN:DNA blotting -hybridization 1975G. KÖHLER & C. MILSTEIN (1984): monoclonal antibodies Untersuchungen über die Nukleine und ihre Spaltungsprodukte ("Investigations into the nucleins and their cleavage products", 1881)

  6. HISTORIC MILESTONES: 1869F. MIESCHER: DNA isolation 1881 Ludwig Karl Martin Leonhard Albrecht KOSSEL(1910) DNA chemical structure 1886T. ESCHERICH: E. coli 1944O. AVERY: DNAis the genetic material 1953J. WATSON F. CRICK, M. WILKINS (Nobel prize, 1962) & R. FRANKLIN:DNA double helix 1957A. KORNBERG (1959):DNA polymerase 1961 J. MARMUR & P. DOTY: DNA renaturation and hybridization 1966 R. HOLEY, M. NIRENBERG, H.G. KHORANA, S. OCHOA(1968, 1959): genetic code 1962 W. ARBER (1978): bacterial restriction-modificaton 1967 M. GELLERT: DNA ligase 1970 H. SMITH (1978): first restriction endonuclease(HindII) 1972 D. NATHANS(1978):first restriction map(SV40) 1972 P. BERG(1980):first ligation 1973 S. COHEN & H. BOYER (1986): first recombinant DNA (E. coli pSC101 TetR + S. aureus pI250 AmpR) 1975 Asilomar conference - moratorium 1975E. SOUTHERN:DNA blotting -hybridization 1975G. KÖHLER & C. MILSTEIN (1984): monoclonal antibodies

  7. HISTORIC MILESTONES: 1869 F. MIESCHER: DNA isolation 1886 T. ESCHERICH: E. coli 1944 O. AVERY, C. MacLEOD, M. McCARTY: DNAis the genetic material 1953 J. WATSON F. CRICK, M. WILKINS (Nobel prize, 1962) & R. FRANKLIN:DNA double helix 1957 A. KORNBERG (1959):DNA polymerase 1961J. MARMUR & P. DOTY: DNA renaturation and hybridization 1966 R. HOLEY, M. NIRENBERG, H.G. KHORANA, S. OCHOA (1968, 1959): genetic code 1962 W. ARBER (1978): bacterial restriction-modificaton 1967 M. GELLERT: DNA ligase 1970 H. SMITH (1978): first restriction endonuclease(HindII) 1972 D. NATHANS(1978):first restriction map(SV40) 1972 P. BERG(1980):first ligation 1973 S. COHEN & H. BOYER (1986): first recombinant DNA (E. coli pSC101 TetR + S. aureus pI250 AmpR) 1975 Asilomar conference - moratorium 1975E. SOUTHERN:DNA blotting -hybridization 1975G. KÖHLER & C. MILSTEIN (1984): monoclonal antibodies Avery, Oswald T., Colin M. MacLeod, and Maclyn McCarty. Studies on the Chemical Nature of the Substance Inducing Transformation of Pneumococcal Types. Journal of Experimental Medicine 79, 2 (1 February 1944): 137-158.

  8. HISTORIC MILESTONES: 1869 F. MIESCHER: DNA isolation 1886 T. ESCHERICH: E. coli 1944O. AVERY, C. MacLEOD, M. McCARTY:DNAis the genetic material 1953 J. WATSON F. CRICK, M. WILKINS (Nobel prize, 1962) & R. FRANKLIN:DNA double helix 1957 A. KORNBERG (1959):DNA polymerase 1961J. MARMUR & P. DOTY: DNA renaturation and hybridization 1966 R. HOLEY, M. NIRENBERG, H.G. KHORANA, S. OCHOA (1968, 1959): genetic code 1962 W. ARBER (1978): bacterial restriction-modificaton 1967 M. GELLERT: DNA ligase 1970 H. SMITH (1978): first restriction endonuclease(HindII) 1972 D. NATHANS(1978):first restriction map(SV40) 1972 P. BERG(1980):first ligation 1973 S. COHEN & H. BOYER (1986): first recombinant DNA (E. coli pSC101 TetR + S. aureus pI250 AmpR) 1975 Asilomar conference - moratorium 1975E. SOUTHERN:DNA blotting -hybridization 1975G. KÖHLER & C. MILSTEIN (1984): monoclonal antibodies

  9. HISTORIC MILESTONES: 1869 F. MIESCHER: DNA isolation 1886 T. ESCHERICH: E. coli 1944O. AVERY, C. MacLEOD, M. McCARTY:DNAis the genetic material 1953 J. WATSON F. CRICK, M. WILKINS (Nobel prize, 1962) & R. FRANKLIN:DNA double helix 1957 A. KORNBERG (1959):DNA polymerase 1961J. MARMUR & P. DOTY: DNA renaturation and hybridization 1966 R. HOLEY, M. NIRENBERG, H.G. KHORANA, S. OCHOA (1968, 1959): genetic code 1962 W. ARBER (1978): bacterial restriction-modificaton 1967 M. GELLERT: DNA ligase 1970 H. SMITH (1978): first restriction endonuclease(HindII) 1972 D. NATHANS(1978):first restriction map(SV40) 1972 P. BERG(1980):first ligation 1973 S. COHEN & H. BOYER (1986): first recombinant DNA (E. coli pSC101 TetR + S. aureus pI250 AmpR) 1975 Asilomar conference - moratorium 1975E. SOUTHERN:DNA blotting -hybridization 1975G. KÖHLER & C. MILSTEIN (1984): monoclonal antibodies

  10. HISTORIC MILESTONES: 1869 F. MIESCHER: DNA isolation 1886 T. ESCHERICH: E. coli 1944O. AVERY, C. MacLEOD, M. McCARTY:DNAis the genetic material 1953 J. WATSON F. CRICK, M. WILKINS (Nobel prize, 1962) & R. FRANKLIN:DNA double helix 1957 A. KORNBERG (1959):DNA polymerase 1961J. MARMUR & P. DOTY: DNA renaturation and hybridization 1966 R. HOLEY, M. NIRENBERG, H.G. KHORANA, S. OCHOA (1968, 1959): genetic code 1962 W. ARBER (1978): bacterial restriction-modificaton 1967 M. GELLERT: DNA ligase 1970 H. SMITH (1978): first restriction endonuclease(HindII) 1972 D. NATHANS(1978):first restriction map(SV40) 1972 P. BERG(1980):first ligation 1973 S. COHEN & H. BOYER (1986): first recombinant DNA (E. coli pSC101 TetR + S. aureus pI250 AmpR) 1975 Asilomar conference - moratorium 1975E. SOUTHERN:DNA blotting -hybridization 1975G. KÖHLER & C. MILSTEIN (1984): monoclonal antibodies hiperkróm effektus

  11. HISTORIC MILESTONES: 1869 F. MIESCHER: DNA isolation 1886 T. ESCHERICH: E. coli 1944O. AVERY, C. MacLEOD, M. McCARTY:DNAis the genetic material 1953 J. WATSON F. CRICK, M. WILKINS (Nobel prize, 1962) & R. FRANKLIN:DNA double helix 1957 A. KORNBERG (1959):DNA polymerase 1961J. MARMUR & P. DOTY: DNA renaturation and hybridization 1961R. HOLEY, M. NIRENBERG, H.G. KHORANA, S. OCHOA(1968, 1959): genetic code 1962 W. ARBER (1978): bacterial restriction-modificaton 1967 M. GELLERT: DNA ligase 1970 H. SMITH (1978): first restriction endonuclease(HindII) 1972 D. NATHANS(1978):first restriction map(SV40) 1972 P. BERG(1980):first ligation 1973 S. COHEN & H. BOYER (1986): first recombinant DNA (E. coli pSC101 TetR + S. aureus pI250 AmpR) 1975 Asilomar conference - moratorium 1975E. SOUTHERN:DNA blotting -hybridization 1975G. KÖHLER & C. MILSTEIN (1984): monoclonal antibodies

  12. HISTORIC MILESTONES: 1869 F. MIESCHER: DNA isolation 1886 T. ESCHERICH: E. coli 1944O. AVERY, C. MacLEOD, M. McCARTY:DNAis the genetic material 1953 J. WATSON F. CRICK, M. WILKINS (Nobel prize, 1962) & R. FRANKLIN:DNA double helix 1957 A. KORNBERG (1959):DNA polymerase 1961J. MARMUR & P. DOTY: DNA renaturation and hybridization 1961R. HOLEY, M. NIRENBERG, H.G. KHORANA, S. OCHOA(1968, 1959): genetic code 1962 W. ARBER (1978): bacterial restriction-modificaton 1967 M. GELLERT: DNA ligase 1970 H. SMITH (1978): first restriction endonuclease(HindII) 1972 D. NATHANS(1978):first restriction map(SV40) 1972 P. BERG(1980):first ligation 1973 S. COHEN & H. BOYER (1986): first recombinant DNA (E. coli pSC101 TetR + S. aureus pI250 AmpR) 1975 Asilomar conference - moratorium 1975E. SOUTHERN:DNA blotting -hybridization 1975G. KÖHLER & C. MILSTEIN (1984): monoclonal antibodies

  13. HISTORIC MILESTONES: 1869 F. MIESCHER: DNA isolation 1886 T. ESCHERICH: E. coli 1944O. AVERY, C. MacLEOD, M. McCARTY:DNAis the genetic material 1953 J. WATSON F. CRICK, M. WILKINS (Nobel prize, 1962) & R. FRANKLIN:DNA double helix 1957 A. KORNBERG (1959):DNA polymerase 1961J. MARMUR & P. DOTY: DNA renaturation and hybridization 1961R. HOLEY, M. NIRENBERG, H.G. KHORANA, S. OCHOA(1968, 1959): genetic code 1962 W. ARBER (1978): bacterial restriction-modificaton 1967 M. GELLERT: DNA ligase 1970 H. SMITH (1978): first restriction endonuclease(HindII) 1972 D. NATHANS(1978):first restriction map(SV40) 1972 P. BERG(1980):first ligation 1973 S. COHEN & H. BOYER (1986): first recombinant DNA (E. coli pSC101 TetR + S. aureus pI250 AmpR) 1975 Asilomar conference - moratorium 1975E. SOUTHERN:DNA blotting -hybridization 1975G. KÖHLER & C. MILSTEIN (1984): monoclonal antibodies

  14. HISTORIC MILESTONES: 1869 F. MIESCHER: DNA isolation 1886 T. ESCHERICH: E. coli 1944O. AVERY, C. MacLEOD, M. McCARTY:DNAis the genetic material 1953 J. WATSON F. CRICK, M. WILKINS (Nobel prize, 1962) & R. FRANKLIN:DNA double helix 1957 A. KORNBERG (1959):DNA polymerase 1961J. MARMUR & P. DOTY: DNA renaturation and hybridization 1961R. HOLEY, M. NIRENBERG, H.G. KHORANA, S. OCHOA(1968, 1959): genetic code 1962 W. ARBER (1978): bacterial restriction-modificaton 1967 M. GELLERT: DNA ligase 1970 H. SMITH (1978): first restriction endonuclease(HindII) 1972 D. NATHANS(1978):first restriction map(SV40) 1972 P. BERG(1980):first ligation 1973 S. COHEN & H. BOYER (1986): first recombinant DNA (E. coli pSC101 TetR + S. aureus pI250 AmpR) 1975 Asilomar conference - moratorium 1975E. SOUTHERN:DNA blotting -hybridization 1975G. KÖHLER & C. MILSTEIN (1984): monoclonal antibodies

  15. HISTORIC MILESTONES: 1869 F. MIESCHER: DNA isolation 1886 T. ESCHERICH: E. coli 1944O. AVERY, C. MacLEOD, M. McCARTY:DNAis the genetic material 1953 J. WATSON F. CRICK, M. WILKINS (Nobel prize, 1962) & R. FRANKLIN:DNA double helix 1957 A. KORNBERG (1959):DNA polymerase 1961J. MARMUR & P. DOTY: DNA renaturation and hybridization 1961R. HOLEY, M. NIRENBERG, H.G. KHORANA, S. OCHOA (1968, 1959): genetic code 1962 W. ARBER (1978): bacterial restriction-modificaton 1967 M. GELLERT: DNA ligase 1970 H. SMITH (1978): first restriction endonuclease(HindII) 1972 D. NATHANS(1978):first restriction map(SV40) 1972 P. BERG(1980):first ligation 1973 S. COHEN & H. BOYER (1986): first recombinant DNA (E. coli pSC101 TetR + S. aureus pI250 AmpR) 1975 Asilomar conference - moratorium 1975E. SOUTHERN:DNA blotting -hybridization 1975G. KÖHLER & C. MILSTEIN (1984): monoclonal antibodies

  16. HISTORIC MILESTONES: 1869 F. MIESCHER: DNA isolation 1886 T. ESCHERICH: E. coli 1944O. AVERY, C. MacLEOD, M. McCARTY:DNAis the genetic material 1953 J. WATSON F. CRICK, M. WILKINS (Nobel prize, 1962) & R. FRANKLIN:DNA double helix 1957 A. KORNBERG (1959):DNA polymerase 1961J. MARMUR & P. DOTY: DNA renaturation and hybridization 1961R. HOLEY, M. NIRENBERG, H.G. KHORANA, S. OCHOA (1968, 1959): genetic code 1962 W. ARBER (1978): bacterial restriction-modificaton 1967 M. GELLERT: DNA ligase 1970 H. SMITH (1978): first restriction endonuclease(HindII) 1972 D. NATHANS(1978):first restriction map(SV40) 1972 P. BERG(1980):first ligation 1973 S. COHEN & H. BOYER (1986): first recombinant DNA (E. coli pSC101 TetR + S. aureus pI250 AmpR) 1975 Asilomar conference - moratorium 1975E. SOUTHERN:DNA blotting -hybridization 1975G. KÖHLER & C. MILSTEIN (1984): monoclonal antibodies

  17. HISTORIC MILESTONES: 1869 F. MIESCHER: DNA isolation 1886 T. ESCHERICH: E. coli 1944O. AVERY, C. MacLEOD, M. McCARTY:DNAis the genetic material 1953 J. WATSON F. CRICK, M. WILKINS (Nobel prize, 1962) & R. FRANKLIN:DNA double helix 1957 A. KORNBERG (1959):DNA polymerase 1961J. MARMUR & P. DOTY: DNA renaturation and hybridization 1961R. HOLEY, M. NIRENBERG, H.G. KHORANA, S. OCHOA (1968, 1959): genetic code 1962 W. ARBER (1978): bacterial restriction-modificaton 1967 M. GELLERT: DNA ligase 1970 H. SMITH (1978): first restriction endonuclease(HindII) 1972 D. NATHANS(1978):first restriction map(SV40) 1972 P. BERG(1980):first ligation 1973 S. COHEN & H. BOYER (1986): first recombinant DNA (E. coli pSC101 TetR + S. aureus pI250 AmpR) 1975 Asilomar conference - moratorium 1975E. SOUTHERN:DNA blotting -hybridization 1975G. KÖHLER & C. MILSTEIN (1984): monoclonal antibodies

  18. HISTORIC MILESTONES: 1869 F. MIESCHER: DNA isolation 1886 T. ESCHERICH: E. coli 1944O. AVERY, C. MacLEOD, M. McCARTY:DNAis the genetic material 1953 J. WATSON F. CRICK, M. WILKINS (Nobel prize, 1962) & R. FRANKLIN:DNA double helix 1957 A. KORNBERG (1959):DNA polymerase 1961J. MARMUR & P. DOTY: DNA renaturation and hybridization 1961R. HOLEY, M. NIRENBERG, H.G. KHORANA, S. OCHOA (1968, 1959): genetic code 1962 W. ARBER (1978): bacterial restriction-modificaton 1967 M. GELLERT: DNA ligase 1970 H. SMITH (1978): first restriction endonuclease(HindII) 1972 D. NATHANS(1978):first restriction map(SV40) 1972 P. BERG(1980):first ligation 1973 S. COHEN & H. BOYER (1986): first recombinant DNA (E. coli pSC101 TetR + S. aureus pI250 AmpR) 1975 Asilomar conference - moratorium 1975E. SOUTHERN:DNA blotting -hybridization 1975G. KÖHLER & C. MILSTEIN (1984): monoclonal antibodies

  19. 1977 Foundation of Genentech 1977A. MAXAM, W. GILBERT, F. SANGER (1980): DNA sequencing 1978M. SMITH (1993): site-specific mutagenesis 1978 Protein mircosequencing 1978 First human genomic library 1978R. LETSINGER:automated oligonucleotide synthesis 1978 First industrial recombinant protein (Eli Lilly, humulin) 1982 BRINSTER & R. PALMITER:transgenic mouse 1982 Sequencing of lambda phage 1985 K. MULLIS (1993): polymerasechain reaction 1985 HUGO (a Human Genom Project) (3x109 bp) 1985 Sequence of first chromosome (yeast III., 315 kb) 1991 S. FODOR: DNA chip (Affymetrix) 1991 First complete procaryotic genom (Haemophylus influenzae, 1,9 Mb) 1991 First complete eucaryotic genom (S. cerevisiae, 12,068 Mb, 5885 gén) 1997 Escherichia coli complete genom (4,6 Mb, 4288) 1998Caenorhabditis elegans complete genom (97 Mb, 19099 gén) 1999 Human chromosome 22 is complete 2000 Draft sequence of human genome 2000 Human chromosome 21 is complete 2000 G. HADLACZKY: human satelite DNA based artificial chrom. (Chromos) 2001 Human chromosome 20 is complete 2002 Draft sequence of mouse genom 2003 Human chromosome 14 is complete

  20. 1977 Foundation of Genentech 1977A. MAXAM, W. GILBERT, F. SANGER (1980): DNA sequencing 1978M. SMITH (1993): site-specific mutagenesis 1978 Protein mircosequencing 1978 First human genomic library 1978R. LETSINGER:automated oligonucleotide synthesis 1978 First industrial recombinant protein (Eli Lilly, humulin) 1982 BRINSTER & R. PALMITER:transgenic mouse 1982 Sequencing of lambda phage 1985 K. MULLIS (1993): polymerasechain reaction 1985 HUGO (a Human Genom Project) (3x109 bp) 1985 Sequence of first chromosome (yeast III., 315 kb) 1991 S. FODOR: DNA chip (Affymetrix) 1991 First complete procaryotic genom (Haemophylus influenzae, 1,9 Mb) 1991 First complete eucaryotic genom (S. cerevisiae, 12,068 Mb, 5885 gén) 1997 Escherichia coli complete genom (4,6 Mb, 4288) 1998Caenorhabditis elegans complete genom (97 Mb, 19099 gén) 1999 Human chromosome 22 is complete 2000 Draft sequence of human genome 2000 Human chromosome 21 is complete 2000 G. HADLACZKY: human satelite DNA based artificial chrom. (Chromos) 2001 Human chromosome 20 is complete 2002 Draft sequence of mouse genom 2003 Human chromosome 14 is complete

  21. 1977 Foundation of Genentech 1977A. MAXAM, W. GILBERT, F. SANGER (1980): DNA sequencing 1978M. SMITH (1993): site-specific mutagenesis 1978 Protein microsequencing 1978 First human genomic library 1978R. LETSINGER ,M. CATUTHERS:automated oligonucleotide synthesis 1978 First industrial recombinant protein (Eli Lilly, humulin) 1982 BRINSTER & R. PALMITER:transgenic mouse 1982 Sequencing of lambda phage 1985 K. MULLIS (1993): polymerasechain reaction 1985 HUGO (a Human Genom Project) (3x109 bp) 1985 Sequence of first chromosome (yeast III., 315 kb) 1991 S. FODOR: DNA chip (Affymetrix) 1991 First complete procaryotic genom (Haemophylus influenzae, 1,9 Mb) 1991 First complete eucaryotic genom (S. cerevisiae, 12,068 Mb, 5885 gén) 1997 Escherichia coli complete genom (4,6 Mb, 4288) 1998Caenorhabditis elegans complete genom (97 Mb, 19099 gén) 1999 Human chromosome 22 is complete 2000 Draft sequence of human genome 2000 Human chromosome 21 is complete 2000 G. HADLACZKY: human satelite DNA based artificial chrom. (Chromos) 2001 Human chromosome 20 is complete 2002 Draft sequence of mouse genom 2003 Human chromosome 14 is complete

  22. 1977 Foundation of Genentech 1977A. MAXAM, W. GILBERT, F. SANGER (1980): DNA sequencing 1978M. SMITH (1993): site-specific mutagenesis 1978 Protein microsequencing 1978 First human genomic library 1978R. LETSINGER ,M. CATUTHERS:automated oligonucleotide synthesis 1978 First industrial recombinant protein (Eli Lilly, humulin) 1982 BRINSTER & R. PALMITER:transgenic mouse 1982 Sequencing of lambda phage 1985 K. MULLIS (1993): polymerasechain reaction 1985 HUGO (a Human Genome Project) (3x109 bp) 1985 Sequence of first chromosome (yeast III., 315 kb) 1991 S. FODOR: DNA chip (Affymetrix) 1991 First complete procaryotic genome(Haemophylus influenzae, 1,9 Mb) 1991 First complete eucaryotic genome(S. cerevisiae, 12,068 Mb, 5885 gén) 1997 Escherichia coli complete genome(4,6 Mb, 4288) 1998Caenorhabditis elegans complete genome(97 Mb, 19099 gén) 1999 Human chromosome 22 is complete 2000 Draft sequence of human genome 2000 Human chromosome 21 is complete 2000 G. HADLACZKY: human satelite DNA based artificial chrom. (Chromos) 2001 Human chromosome 20 is complete 2002 Draft sequence of mouse genome 2003 Human genom project completed

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