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Home , Walter Gilbert

WALTER GILBERT Curriculum Vitae

Personal Data:

Born: March 1932, in , , U.S.A. Married: Celia Stone Gilbert, two children Website: http://wallygilbert.com

Education:

1953 B.A. , summa cum laude , and 1954 M.A. , Physics 1957 Ph.D. Cambridge University, Mathematics (: On Generalized Dispersion Relations for Pion-Nucleon Scattering)

Appointments:

1953-57 NSF predoctoral fellowships held at Harvard University and Cambridge University 1957-58 NSF postdoctoral fellowship held at Harvard in Physics 1958-59 Assistant to Professor Schwinger at Harvard and Lecturer in Physics 1959-64 Assistant Professor of Physics, Harvard University 1964-68 Associate Professor of Biophysics, Harvard University 1968-72 Professor of , Harvard University 1972-81 American Cancer Society Professor of Molecular , Harvard University 1978-83 Chair, Scientific Board of Directors, N.V. 1979-81 Co-Chair, Supervisory Board of Directors, Biogen N.V. 1981-84 Chair, Supervisory Board of Directors and Principal Executive Officer, Biogen N.V. 1985-86 Professor of Biology, Harvard University 1986-87 H. H. Timken Professor of Science, Dept. of Cellular and Developmental Biology, Harvard University 1987-93 Chair, Dept. of Cellular and Developmental Biology, Harvard University 1987-02 Carl M. Loeb University Professor, Dept. of Molecular and Cellular Biology, Harvard University 1992- Vice Chair, Board of Directors, Myriad Genetics, Inc. 1994-14 Member, Board of Scientific Governors, The Institute 1996-02 Chair, Board of Directors, NetGenics, Inc. 1996- 16 Chair, Board of Directors, Paratek Pharmaceuticals, Inc. 1998-10 Member, Board of Directors & Scientific Advisory Board, Memory Pharmaceuticals, Inc. 1999-04 Chair, Board of Directors and member of Scientific Advisory Board, Pintex Pharmaceuticals, Inc. 2000-04 Member, Board of Directors and Scientific Advisory Board, Transkaryotic Therapies, Inc. 2001-05 Member, Board of Directors, HospitalCareOnline.com, Inc. 2001- Managing Director, BioVentures Investors 2003-04 Carl M. Loeb University Research Professor, Dept. of Molecular and Cellular Biology, Harvard University 2004- Carl M. Loeb University Professor Emeritus, Harvard University 2015- Director, Amylyx, Inc.

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Awards:

United States Steel Foundation Award in of the National Academy of Sciences, 1968 Elected American Academy of Arts and Sciences, 1968 Guggenheim Fellowship, 1968-69, Paris Ledlie Prize, Harvard University, 1969 (with M. Ptashne) Elected National Academy of Sciences, 1976 V.D. Mattia Lectureship, Roche Institute of Molecular Biology, 1976 Smith Kline & French Lecturer, U.C. Berkeley, 1977 Warren Triennial Prize, Massachusetts General Hospital, 1977 (with S. Benzer) Louis and Bert Freedman Award, New York Academy of Sciences, 1977 Prix Charles-Leopold Mayer, Academie des Sciences, Institut de France, 1977 (with M. Ptashne and E. Witkin) Doctor of Science (Honorary), University of Chicago, 1978 Doctor of Science (Honorary), , 1978 Harrison Howe Award of the Rochester Branch of the American Chemical Society, 1978 Doctor of Science (Honorary), University of Rochester, 1979 Louisa Gross Horwitz Prize, Columbia University, 1979 (with F. Sanger) Gairdner Foundation Annual Award, The Gairdner Foundation, 1979 Albert Lasker Basic Medical Research Award, The Albert and Mary Lasker Foundation, 1979 (with F. Sanger) Prize Biochemical Analysis, The German Society for Clinical Chemistry, 1980 (with A. Maxam, F. Sanger, and A. Coulsen) Sober Award, The American Society of Biological , 1980 for Chemistry, 1980 (with F. Sanger and P. Berg) Doctor of Science (Honorary), Yeshiva University, 1981 Senior U.S. Scientist Humboldt Award, 1987 Foreign Member of the Royal Society, 1987 Honorary Fellow, Trinity College, Cambridge, England, 1991 New England Entrepreneur of the Year Award, 1991 Ninth National Biotechnology Ventures Award, 1997 The Fourth Annual Biotechnology Heritage Award, 2002 (with P. Sharp)

Publications:

In theoretical physics:

1. On generalized dispersion relations, II, A. Salam and W. Gilbert. Nuovo Cimento 3, 607 (1956). 2. Yang-Fermi ambiguity, W. Gilbert and G. Screaton. Phys. Rev. 104 , 1758 (1956). 3. New dispersion relations for pion-nucleon scattering, W. Gilbert. Phys. Rev. 108 , 1078 (1957). 4. Structure of the vertex function, S. Deser, W. Gilbert and E. Sudarshan. Phys. Rev. 115 , 731 (1959). 5. Structure of the forward scattering amplitude, S. Deser, W. Gilbert and E. Sudarshan. Phys. Rev. 117 , 266 (1960). 6. Integral representation of two-point functions, S. Deser, W. Gilbert and E. Sudarshan. Phys. Rev. 117 , 266 (1960). 7. Connection between gauge invariance and mass, D. Boulware and W. Gilbert. Phys. Rev. 126 , 1563 (1962). 8. Broken symmetries and massless particles, W. Gilbert. Phys. Rev. Lett . 12 , 713 (1964).

In molecular biology:

9. Unstable ribonucleic acid revealed by pulse labelling of Escherichia coli , F. Gros, H. Hiatt, W. Gilbert, C.G. Kurland, R.W. Reisbrough and J.D. Watson. 190 , 581-585 (1961). 10. Molecular and biological characterization of messenger RNA, F. Gros, W. Gilbert, H.H. Hiatt, G. Attardi, P.F. Spahr and J.D. Watson. Cold Spring Harbor Symp . Quant. Biol. 26, 111-132 (1961). 11. Polypeptide synthesis in Escherichia coli . I. Ribosomes and the active complex, W. Gilbert. J. Mol. Biol. 6, 374-388 (1963).

2 12. Polypeptide synthesis in Escherichia coli . II. The polypeptide chain and S-RNA, W. Gilbert. J. Mol. Biol. 6, 389-403 (1963). 13. The binding of S-RNA by Escherichia coli ribosomes, M. Cannon, R. Krug and W. Gilbert. J. Mol. Biol. 7, 360-378 (1963). 14. Protein synthesis in Escherichia coli, W. Gilbert. Cold Spring Harbor Symp. Quant. Biol. 28 , 287-297 (1963). 15. Studies on methylated bases in transfer RNA, U.Z. Littauer, K. Muench, P. Berg, W. Gilbert and P.F. Spahr. Cold Spring Harbor Symp. Quant. Biol. 28 157-159 (1963). 16. Streptomycin, suppression and the code, J. Davies, W. Gilbert and L. Gorini. Proc. Natl. Acad. Sci. 51 , 883-890 (1964). 17. Isolation of the lac repressor, W. Gilbert and B. Müller-Hill. Proc. Natl. Acad. Sci. 56 , 1891-1898 (1966). 18. The lac operator is DNA, W. Gilbert and B. Müller-Hill. Proc. Natl. Acad. Sci. 58 , 2415-2421 (1967). 19. Mutants that make more lac repressor, B. Müller-Hill, L. Crapo and W. Gilbert. Proc. Natl. Acad. Sci. 59 , 1259-1264 (1968). 20. DNA replication: The rolling circle model, W. Gilbert and D. Dressler. Cold Spring Harbor Symp. Quant. Biol. 33 , 473-484 (1968). 21. Genetic repressors, M. Ptashne and W. Gilbert. Sci. Amer. 222(6) , 36-44 (1970). 22. The lactose repressor, W. Gilbert and B. Müller-Hill. In The Lactose Opero n (J. Beckwith and D. Zipser, eds.), Cold Spring Harbor Lab, New York, 93-109 (1970). 23. Repressors and genetic control, W. Gilbert. In The Neurosciences: Second Study Program (F.O Schmitt, ed.), The Rockefeller Univ. Press, New York, 946-954 (1970). 24. MSI and MSII made on ribosome in idling step of protein synthesis, W.A. Haseltine, R. Block, W. Gilbert and K. Weber. Nature 238 , 381-384 (1972). 25. The lac repressor and the lac operator, W. Gilbert. In CIBA Foundation Symp. 7: Polymerization in Biological Systems , Elsevier Publishing Co., Amsterdam, 245-259 (1972). 26. The sequence of the lac operator, W. Gilbert and A. Maxam. Proc. Natl. Acad. Sci. USA 70 , 3581-3584 (1973). 27. Sequences of controlling regions of the E. coli lactose operon, W. Gilbert, N. Maizels and A. Maxam. Genetics 79, 227 (1975). 28. Lactose operator sequences and the action of lac repressor, W. Gilbert, J. Gralla, J. Majors and A. Maxam. In Protein-Ligand Interactions (H. Sund and G. Blauer, eds.), Walter de Gruyter, Berlin, 193- 210 (1975). 29. Contacts between the lac repressor and DNA revealed by methylation, W. Gilbert, A. Maxam and A. Mirzabekov. In Control of Ribosome Synthesis (N. Kjeldgaard and O. Maaloe, eds.), Munksgaard, Copenhagen, 139-148 (1976). 30. Preferential protection of the minor groove of non-operator DNA by lac repressor against methylation by dimethyl sulphate, A.M. Kolchinsky, A.D. Mirzabekov, W. Gilbert and L. Li. Nucl. Acids Res. 3(1) , 11- 18 (1976). 31. Starting and stopping sequences for the RNA polymerase, W. Gilbert. In RNA Polymerase (R. Losick and M. Chamberlin, eds.), Cold Spring Harbor Lab, New York, 193-205 (1976). 32. How proteins recognize DNA sequences, W. Gilbert, J. Majors and A. Maxam. In Organization and Expression of Chromosomes , Dahlem Workshop Report, Berlin, 167-178 (1976). 33. Construction of plasmids carrying the cI gene of bacteriophage lambda, K. Backman, M. Ptashne and W. Gilbert. Proc. Natl. Acad. Sci. USA 73 , 4174-4178 (1976). 34. A new method for sequencing DNA, A.M. Maxam and W. Gilbert. Proc. Natl. Acad. Sci. USA 74 , 560-564 (1977). 35. Rous sarcoma virus genome is terminally redundant: The 5' sequence, W.A. Haseltine, A.M. Maxam and W. Gilbert. Proc. Natl. Acad. Sci. USA 74 , 989-993 (1977). 36. Promoter region for yeast 5S ribosomal RNA, A. Maxam, R. Tizard, K.G. Skryabin and W. Gilbert. Nature 267 , 643-645 (1977). 37. Mapping adenines, guanines and pyrimidines in RNA, H. Donis-Keller, A.M. Maxam and W. Gilbert. Nucl. Acids Res. 4(8) , 2527-2538 (1977). 38. A promoter region for yeast 5S RNA, W. Gilbert, A.M. Maxam, R. Tizard and K.G. Skryabin. In Eucaryotic Genetic System (J. Abelson and G. Wilcox, eds.), Academic Press, 15-23 (1977). 39. Contacts between the lac repressor and thymines in the lac operator, R. Ogata and W. Gilbert. Proc. Natl. Acad. Sci. USA 74 , 4973-4976 (1977). 40. Why genes in pieces? W. Gilbert. Nature 271 , 501 (1978).

3 41. Sequence of a mouse germ-line gene for a variable region of an immunoglobulin light chain, S. Tonegawa, A.M. Maxam, R. Tizard, O. Bernard and W. Gilbert. Proc. Natl. Acad. Sci. USA 75 , 1485- 1489 (1978). 42. Immunoglobulin screening method to detect specific translation products, S. Broome and W. Gilbert. Proc. Natl. Acad. Sci. USA 75 , 2746-2749 (1978). 43. Molecular basis of base substitution hotspots in Escherichia coli, C. Coulondre, J.H. Miller, P.J. Farabaugh and W. Gilbert. Nature 274 , 775-780 (1978). 44. A bacterial clone synthesizing proinsulin, L. Villa-Komaroff, A. Efstratiadis, S. Broome, P. Lomedico, R. Tizard, S.P. Naber, W.L. Chick and W. Gilbert. Proc. Natl. Acad. Sci. USA 75 , 3727-3731 (1978). 45. An amino-terminal fragment of lac repressor binds specifically to lac operator, R.T. Ogata and W. Gilbert. Proc. Natl. Acad. Sci. USA 75 , 5851-5854 (1978). 46. DNA-binding site of lac repressor probed by dimethylsulfate methylation of lac operator, R.T. Ogata and W. Gilbert. J. Mol. Biol. 132 , 709-728 (1979). 47. and : Playgrounds of evolution, W. Gilbert. In Eucaryotic Gene Regulation (R. Axel, T. Maniatis and C.F. Fox, eds.), Academic Press, 1-12 (1979). 48. The structure and transcription of rat preproinsulin genes, A. Efstratiadis, P. Lomedico, N. Rosenthal, R. Kolodner, R. Tizard, F. Perler, L. Villa-Komaroff, S. Naber, W. Chick, S. Broome and W. Gilbert. In Eucaryotic Gene Regulation (R. Axel, T. Maniatis and C.F. Fox, eds.), Academic Press, 301-315 (1979). 49. The structure and evolution of the two nonallelic rat preproinsulin genes, P. Lomedico, N. Rosenthal, A. Efstratiadis, W. Gilbert, R. Kolodner and R. Tizard. Cell 18 , 545-558 (1979). 50. Pleiotropic mutations within two yeast mitochondrial cytochrome genes block mRNA processing, G.M. Church, P.P. Slonimski and W. Gilbert. Cell 18 , 1209-1215 (1979). 51. Hepatitis B virus genes and their expression in E. coli , M. Pasek, T. Goto, W. Gilbert, B. Zink, H. Schaller, P. MacKay, G. Leadbetter and K. Murray. Nature 282 , 575-579 (1979). 52. The structure of rat preproinsulin genes, P. Lomedico, N. Rosenthal, R. Kolodner, A. Efstratiadis and W. Gilbert. Ann. N.Y.Acad. Sci. , 425 (1980). 53. Contacts between Escherichia coli RNA polymerase and an early promoter of phage T7, U. Siebenlist and W. Gilbert. Proc. Natl. Acad. Sci. USA 77 , 122-126 (1980). 54. Useful proteins from recombinant bacteria, W. Gilbert and L. Villa-Komaroff. Sci. Am. 242(4), 74-84 (1980). 55. Sequencing end-labeled DNA with base-specific chemical cleavages, A.M. Maxam and W. Gilbert. In Methods in Enzymology , Vol. 65, Part I, 499-560 (K. Moldave and L. Grossman, ed.), Academic Press (1980). 56. E. coli RNA polymerase interacts homologously with two different promoters, U. Siebenlist, R.B. Simpson and W. Gilbert. Cell 20 , 269-281 (1980). 57. The evolution of genes: The chicken preproinsulin gene, F. Perler, A. Efstratiadis, P. Lomedico, W. Gilbert, R. Kolodner and J. Dodgson. Cell 20 , 555-566 (1980). 58. Yeast mitochondrial products required in trans for RNA splicing, G. Church and W. Gilbert. Miami Symp.: Mobilization and Reassembly of Genetic Information, 17, 379-394 (1980). 59. Eukaryotic signal sequence transports antigen in Escherichia coli , K. Talmadge, S. Stahl and W. Gilbert. Proc. Natl. Acad. Sci. USA 77 , 3369-3373 (1980). 60. The synthesis of insulin in bacteria: A model for the production of medically useful proteins in prokaryotic cells, L. Villa-Komaroff, S. Broome, S.P. Naber, A. Efstratiadis, P. Lomedico, R. Tizard, W.L. Chick and W. Gilbert. In Birth Defects: Original Article Series , Vol. XVI, No. 1, 53-68 (March of Dimes Birth Defects Found.) (1980) 61. Bacteria mature preproinsulin to proinsulin, K. Talmadge, J. Kaufman and W. Gilbert. Proc. Natl. Acad. Sci. USA 77 , 3988-3992 (1980). 62. Chemical probes for higher-order structure in RNA, D.A. Peattie and W. Gilbert. Proc. Natl. Acad. Sci. USA 77 , 4679-4682 (1980). 63. Construction of plasmid vectors with unique Pst I cloning sites in a signal sequence , K. Talmadge and W. Gilbert. Gene 12 , 235-241 (1980). 64. Tissue-specific exposure of chromatin structure at the 5' terminus of the rat preproinsulin II gene, C. Wu and W. Gilbert. Proc. Natl. Acad. Sci. USA 78 , 1577-1580 (1981). 65. Expression of active polypeptides in E. Coli , W. Gilbert. Recomb. DNA Tech. Bull. 4(1), 4-5 (1981). 66. DNA sequencing and gene structure, W. Gilbert. Science 214, 1305-1312 (1981).

4 67. The production of immunologically active surface antigens of hepatitis B virus by Escherichia coli , P. Mackay, M. Pasek, M. Magazin, R.T. Kovacic, B. Allet, S. Stahl, W. Gilbert, H. Schaller, S.A. Bruce and K. Murray. Proc. Natl. Acad. Sci. USA 78 , 4510-4514 (1981). 68. Parabiosis as a model system for network interactions, K. Fischer Lindahl, W. Gilbert and K. Rajewsky. In The Immune System, Vol. 2, 24-32, Basel (1981). 69. Proinsulin from bacteria, K. Talmadge and W. Gilbert. In Proceedings of the International Plasmid Meeting (S. Levy, R. Clowes and E. Koenig, eds.), Plenum Publishing, 411-419 (1981). 70. An "internal" signal sequence directs secretion and processing of proinsulin in bacteria, K. Talmadge, J. Brosius and W. Gilbert. Nature 294 , 176-177 (1981). 71. Germ-line MuLV reintegrations in AKR/J mice, W. Herr and W. Gilbert. Nature 269 , 865-868 (1982). 72. Cellular location affects protein stability in Escherichia coli , K. Talmadge and W. Gilbert. Proc. Natl. Acad. Sci. USA 79 , 1830-1833 (1982). 73. Nucleotide sequence of the 3' half of AKV, W. Herr, V. Corbin and W. Gilbert. Nucl. Acids Res. 10(21) , 6931-6944 (1982). 74. Somatically acquired recombinant murine leukemia proviruses in thymic leukemias of AKR/J mice, W. Herr and W. Gilbert. J. Virology 46(1) , 70-82 (1983). 75. Isolation and mapping of cDNA hybridization probes specific for ecotropic and nonecotropic murine leukemia proviruses, W. Herr, D. Schwartz and W. Gilbert. Virology 125 , 139-154 (1983). 76. Nucleotide sequence of rous sarcoma virus, D.E. Schwartz, R. Tizard and W. Gilbert. Cell 32 , 853-869 (1983). 77. Comparison of the methylation patterns of the two rat insulin genes, R.L. Cate, W. Chick and W. Gilbert. J. Biol. Chem. 258(10) , 6645-6652 (1983). 78. Primary structure of chicken muscle pyruvate kinase mRNA, N. Lonberg and W. Gilbert. Proc. Natl. Acad. Sci. USA 80 , 3661-3665 (1983). 79. Monoclonal AKR/J thymic leukemias contain multiple JH immunoglobulin gene rearrangements, W. Herr, A.P. Perlmutter and W. Gilbert. Proc. Natl. Acad. Sci. USA 80 , 7433-7436 (1983). 80. Genomic sequencing, G.M. Church and W. Gilbert. Proc. Natl. Acad. Sci. USA 81 , 1991-1995 (1984). 81. Free and integrated recombinant murine leukemia virus DNAs appear in preleukemic thymuses of AKR/J mice, W. Herr and W. Gilbert. J. Virol 50(1), 155-162 (1984). 82. Antibodies of the secondary response can be expressed without switch recombination in normal mouse B cells, A.P. Perlmutter and W. Gilbert. Proc. Natl. Acad. Sci. USA 81 , 7189-7193 (1984). 83. B lineage-specific interactions of an immunoglobulin enhancer with cellular factors in vivo , A. Ephrussi, G.M. Church, S. Tonegawa and W. Gilbert. Science 227 , 134-140 (1985). 84. Intron/ structure of the chicken pyruvate kinase gene, N. Lonberg and W. Gilbert. Cell 40 , 81-90 (1985). 85. Detection in vivo of protein-DNA interactions within the lac operon of Escherichia coli , H. Nick and W. Gilbert. Nature 313 , 795-798 (1985). 86. Cell-type-specific contacts to immunoglobulin enhancers in nuclei, G.M. Church, A. Ephrussi, W. Gilbert and S. Tonegawa. Nature 313 , 798-801 (1985). 87. Chicken triosephosphate isomerase complements an Escherichia coli deficiency, D. Straus and W. Gilbert. Proc. Natl. Acad. Sci. USA 82 , 2014-2018 (1985). 88. Rous sarcoma virus encodes a transcriptional activator, S. Broome and W. Gilbert. Cell 40 , 537-546 (1985). 89. Active site of triosephosphate isomerase: In vitro mutagenesis and characterization of an altered enzyme, D. Straus, R. Raines, E. Kawashima, J.R. Knowles and W. Gilbert. Proc. Natl. Acad. Sci. USA 82 , 2272-2276 (1985). 90. Genes-in-pieces revisited, W. Gilbert. Science 228 , 823-824 (1985). 91. Genetic engineering in the Precambrian: Structure of the chicken triosephosphate isomerase gene, D. Straus and W. Gilbert. Mol. Cell. Biol. 5(12) , 3497-3506 (1985). 92. The genomic sequencing technique, G.M. Church and W. Gilbert. Prog. Clin. Biol. Res. 177, 17-21 (1985). 93. Detection of cytosine methylation in the maize alcohol dehydrogenase gene by genomic sequencing, H. Nick, B. Bowen, R.J. Ferl and W. Gilbert. Nature 319 , 243-346 (1986). 94. Origin of life: The RNA world, W. Gilbert. Nature 319, 618 (1986). 95. Evolution of antibodies: The road not taken, W. Gilbert. Nature 320 , 485-486 (1986). 96. Reaction energetics of a mutant triosephosphate isomerase in which the active-site glutamate has been changed to aspartate, R.T. Raines, E.L. Sutton, D.R. Straus, W. Gilbert and J.R. Knowles. Biochem. 25 , 7142-7154 (1986). 5 97. The kinetic consequences of altering the catalytic residues of triosephosphate isomerase, R.T. Raines, D.R. Straus, W. Gilbert and J.R. Knowles. Phil. Trans. R. Soc. Lond. A 317 , 371-380 (1986). 98. The triosephosphate isomerase gene from maize: Introns antedate the plant-animal divergence, M. Marchionni and W. Gilbert. Cell 46 , 133-141 (1986). 99. On the antiquity of introns, W. Gilbert, M. Marchionni and G. McKnight. Cell 46 , 151-153 (1986). 100. The exon theory of genes, W. Gilbert. Cold Spring Harbor Symp. Quant. Biol. 52, 901-905 (1987). 101. Genome sequencing: Creating a new biology for the twenty-first century, W. Gilbert. Issues in Science & Technology III(3), 26-35 (1987). 102. Basic protein enhances the incorporation of DNA into lipid vesicles: Model for the formation of primordial cells, D.G. Jay and W. Gilbert. Proc. Natl. Acad. Sci. USA 84, 1978-1980 (1987) . 103. Formation of parallel four-stranded complexes by guanine-rich motifs in DNA and its implications for meiosis, D. Sen and W. Gilbert. Nature 334 , 364-366 (1988). 104. sequencing, W. Gilbert. Basic Life Sci. 46, 29-36 (1988). 105. Differential expression of acetylcholine receptor mRNA in nuclei of cultured muscle cells, S. Bursztajn, S.A. Berman and W. Gilbert. Proc. Natl. Acad. Sci. USA 86 , 2928-2932 (1989). 106. The monoclonal antibody B30 recognizes a specific neuronal cell surface antigen in the developing mesencephalic trigeminal nucleus of the mouse, D.Y. Stainier and W. Gilbert. J. Neurosci. 9(7) , 2468- 2485 (1989). 107. One-sided polymerase chain reaction: The amplification of cDNA, O. Ohara, R.L. Dorit and W. Gilbert. Proc. Natl. Acad. Sci. USA 86 , 5673-5677 (1989). 108. Direct genomic sequencing of bacterial DNA: The pyruvate kinase I gene of Escherichia coli , O. Ohara, R.L. Dorit and W. Gilbert. Proc. Natl. Acad. Sci. USA 86 , 6883-6887 (1989). 109. Detection of mutations and DNA polymorphisms using whole genome Southern Cross hybridization, G. Ruvkun, W. Gilbert and H.R. Horvitz. Nucl. Acids Res. 18(4) , 809-815 (1990). 110. DNA trapping electrophoresis, L. Ulanovsky, G. Drouin and W. Gilbert. Nature 343, 190-192 (1990). 111. Localization of an acetylcholine receptor intron to the nuclear membrane, S.A. Berman, S. Bursztajn, B. Bowen and W. Gilbert. Science 247 , 212-214 (1990). 112. Pioneer neurons in the mouse trigeminal sensory system, D.Y.R. Stainier and W. Gilbert. Proc. Natl. Acad. Sci. USA 87 , 923-927 (1990). 113. A sodium-potassium switch in the formation of four-stranded G4-DNA, D. Sen and W. Gilbert. Nature 344 , 410-414 (1990). 114. Factors released by ciliary neurons and spinal cord explants induce acetylcholine receptor mRNA expression in cultured muscle cells, S. Bursztajn, S.A. Berman and W. Gilbert. J. Neurobiol. 21 (3) , 387-399 (1990). 115. Simultaneous visualization of neuronal protein and receptor mRNA, S. Bursztajn, S.A. Berman and W. Gilbert. BioTechniques 9(4) , 440-449 (1990). 116. How big is the universe of exons? R.L. Dorit, L. Schoenbach and W. Gilbert. Science 250, 1377-1382 (1990). 117. Towards a paradigm shift in biology, W. Gilbert. Nature 349, 99 (1991). 118. Murine memory B cells are multi-isotype expressors, C.J. Wu, J.T. Karttunen, D.H.L. Chin, D. Sen and W. Gilbert. Immunol. 72 , 48-55 (1991). 119. The B30 ganglioside is a cell surface marker for neural crest derived neurons in the developing mouse, D.Y.R. Stainier, D.H. Bilder and W. Gilbert. Dev. Biol. 144 , 177-188 (1991). 120. Gene structure and evolutionary theory, W. Gilbert. In New Perspectives on Evolution , Wiley-Liss, Inc., 155-163 (1991). 121. Spatial domains in the developing forebrain: Developmental regulation of a restricted cell surface protein, D.Y.R. Stainier, D.H. Bilder and W. Gilbert. Dev. Biol. 147 , 22-31 (1991). 122. The structure of telomeric DNA: DNA quadriplex formation [Review], D. Sen and W. Gilbert. Curr. Opin. Struct. Biol. 1 , 35-438 (1991). 123. Neuronal differentiation and maturation in the mouse trigeminal sensory system, in vivo and in vitro , D.Y.R. Stainier and W. Gilbert. J. Comp. Neurol. 311, 300-312 (1991). 124. DNA sequencing, today and tomorrow, W. Gilbert. Hospital Practice 26(10) , 165-174 (1991). 125. The limited universe of exons [Review], R.L. Dorit and W. Gilbert. Curr. Opin. Struct. Biol. 1 , 973-977 (1991). 126. Zebrafish embryology and neural development [Review], C. Fulwiler and W. Gilbert. Curr. Opin. Cell Biol. 3, 988-991 (1991).

6 127. Cationic switches in the formation of DNA structures containing guanine-quartets, D. Sen and W. Gilbert. In Structure and Function, Vol. 1: Nucleic Acids , (R.H. Sarma & M.H. Sarma, eds.) Adenine Press, 43- 52 (1992). 128. Novel DNA superstructures formed by telomere-like oligomers, D. Sen and W. Gilbert. Biochem. 31 , 65- 70 (1992). 129. Guanine quartet structures, D. Sen and W. Gilbert. Methods in Enzymol . 211 , 191-199 (1992). 130. A fate map for the first cleavages of the zebrafish, D. Strehlow and W. Gilbert. Nature 361 , 451-453 (1993). 131. Identification and characterization of a nuclease activity specific for G4 tetrastranded DNA, Z. Liu, J.D. Frantz, W. Gilbert and B.K. Tye. Proc. Natl. Acad. Sci. USA 90 , 3157-3161 (1993). 132. RNA editing as a source of genetic variation, L.F. Landweber and W. Gilbert. Nature 363 , 179-182 (1993). 133. The boundaries of partially edited transcripts are not conserved in kinetoplastids: Implications for the guide RNA model of editing, L.F. Landweber, A.G. Fiks and W. Gilbert. Proc. Natl. Acad. Sci. USA 90, 9242-9246 (1993). 134. On the ancient nature of introns, W. Gilbert and M. Glynias. Gene 135, 137-144 (1993). 135. Phylogenetic analysis of RNA editing: A primitive genetic phenomenon, L.F. Landweber and W. Gilbert. Proc. Natl. Acad. Sci. USA 91 , 918-921 (1994). 136. The fates of the blastomeres of the 16-cell zebrafish embryo, D. Strehlow, G. Heinrich and W. Gilbert. Development 120, 1791-1798 (1994). 137. Large scale bacterial gene discovery by similarity search, K. Robison, W. Gilbert and G.M. Church. Nature Genetics 7, 205-214 (1994). 138. The genetic data environment an expandable GUI for multiple sequence analysis, S.W. Smith, R. Overbeek, C.R. Woese, W. Gilbert and P.M. Gillevet. CABIOS 10(6) 671-675 (1994). 139. The yeast KEM1 gene encodes a nuclease specific for G4 tetraplex DNA: Implication of in vivo functions for this novel DNA structure, Z. Liu and W. Gilbert. Cell 77 , 1083-1092 (1994). 140. Retinoic acid is necessary for development of the ventral retina in zebrafish, N. Marsh-Armstrong, P. McCaffery, W. Gilbert, J.E. Dowling and U.C. Drager. Proc. Natl. Acad. Sci. USA 91 , 7286-7290 (1994). 141. A yeast gene product, G4p2, with a specific affinity for quadruplex nucleic acids, J.D. Frantz and W. Gilbert. J. Biol. Chem. 270(16) , 9413-9419 (1995). 142. Absence of polymorphism at the ZFY locus on the human Y chromosome, R.L. Dorit, H. Akashi and W. Gilbert. Science 268 , 1183-1185 (1995). 143. Gene disruption of a G4-DNA dependent nuclease in yeast leads to cellular senescence and telomere shortening, Z. Liu, A. Lee and W. Gilbert. Proc. Natl. Acad. Sci. USA 92 , 6002-6006 (1995). 144. Large scale genomic sequencing: Optimization of genomic chemical sequencing reactions, M. Dolan, A. Ally, M.S. Purzycki, W. Gilbert and P.M. Gillevet. BioTechniques 19(2) , 264-273 (1995). 145. Exploring the Mycoplasma capricolum genome: A minimal cell reveals its physiology, P. Bork, C. Ouzounis, G. Casari, R. Schneider, C. Sander, M. Dolan, W. Gilbert and P.M. Gillevet. Mol. Microbiol. 16(5) , 955-967 (1995). 146. Tests of the exon theory of genes, W. Gilbert, M. Long, C. Rosenberg and M. Glynias. In Tracing Biological Evolution in Protein and Gene Structures, Proceedings of the 20th Taniguchi International Symp., Div. of Biophysics (M. Go and P. Schimmel, eds.), Elsevier Science B.V., 237-247 (1995). 147. A novel yeast gene product, G4p1, with a specific affinity for quadruplex nucleic acids, J.D. Frantz and W. Gilbert. J. Biol. Chem . 270(35) , 20692-20697 (1995). 148. Intron phase correlations and the evolution of the intron/exon structure of genes, M. Long, C. Rosenberg and W. Gilbert. Proc. Natl. Acad. Sci. USA 92, 12495-12499 (1995). 149. Evolution of the intron-exon structure of eukaryotic genes [Review], M. Long, S.J. de Souza and W. Gilbert. Curr. Opin. Genet. Devel. 5, 774-778 (1995). 150. More haemophilus and mycoplasma genes, K. Robison, W. Gilbert and G.M. Church. Science 271 , 1302-1303 (1996). 151. Discovery of amphibian Tc1-like transposon families, W.L. Lam, P. Seo, K. Robison, S. Virk and W. Gilbert. J. Mol. Biol. 257 , 359-366 (1996). 152. Exon shuffling and the origin of the mitochondrial targeting function in plant cytochrome c1 precursor, M. Long, S.J. de Souza, C. Rosenberg and W. Gilbert. Proc. Natl. Acad. Sci. USA 93, 7727-7731 (1996). 153. Introns and gene evolution, S.J. de Souza, M. Long and W. Gilbert. Genes to Cells 1, 493-505 (1996). 154. Active transposition in zebrafish, W.L. Lam, T.-S. Lee and W. Gilbert. Proc. Natl. Acad. Sci. USA 93, 10870-10875 (1996) . 7 155. Novel G qα isoform is a candidate transducer of rhodopsin signaling in a Drosophila testes-autonomous pacemaker, C.E. Alvarez, K. Robison and W. Gilbert. Proc. Natl. Acad. Sci. USA 93, 12278-12282 (1996). 156. Intron positions correlate with module boundaries in ancient proteins, S.J. de Souza, M. Long, L. Schoenbach, S.W. Roy and W. Gilbert. Proc. Natl. Acad. Sci. USA 93, 14632-14636 (1996). 157. Jak1 kinase is required for cell migrations and anterior specification in zebrafish embryos, G. Conway, A. Margoliath, S. Wong-Madden, R.J. Roberts and W. Gilbert. Proc. Natl. Acad. Sci. USA 94, 3082-3087 (1997). 158. Delta-interacting protein A and the origin of hepatitis delta antigen, M. Long, S.J. de Souza and W. Gilbert. Science 276 , 824-825 (1997). 159. Dealing with database explosion: A cautionary note, U. Bhatia, K. Robison and W. Gilbert. Science 276 , 1724-1725 (1997). 160. Ligands for ErbB-family receptors encoded by a neuregulin-like gene, H. Chang, D.J. Riese II, W. Gilbert, D.F. Sterns and U.J. McMahan. Nature 387 , 509-512 (1997). 161. Origin of genes, W. Gilbert, S.J. de Souza and M. Long. Proc. Natl. Acad. Sci. USA 94, 7698-7703 (1997). 162. A novel zebrafish gene expressed specifically in the photoreceptor cells of the retina, H. Chang and W. Gilbert. Biochem. & Biophys. Res. Comm. 237, 84-89 (1997). 163. The yeast splice site revisited: New exon consensus from genomic analysis, M. Long, S.J. de Souza and W. Gilbert. Cell 91 , 739-740 (1997). 164. The correlation between introns and the three-dimensional structure of proteins, S.J. de Souza, M. Long, L. Schoenbach, S.W. Roy and W. Gilbert. Gene 205 , 141-144 (1997). 165. Relationship between "proto-splice sites" and intron phases: Evidence from dicodon analysis, M. Long, S.J. de Souza, C. Rosenberg and W. Gilbert. Proc. Natl. Acad. Sci. USA 95, 219-223 (1998). 166. Toward a resolution of the introns early/late debate: Only phase zero introns are correlated with the structure of ancient proteins, S.J. de Souza, M. Long, R.J. Klein, S. Roy, S. Lin, and W. Gilbert. Proc. Natl. Acad. Sci. USA 95, 5094-5099 (1998) . 167. Introns and the RNA world, W. Gilbert and S.J. de Souza. In The RNA World, Second Edition , Cold Spring Harbor Laboratory Press, 221-231 (1999). 168. Centripetal modules and ancient introns, S.W. Roy, M. Nosaka, S.J. de Souza and W. Gilbert. Gene 238 , 85-91 (1999). 169. EID: The exon-intron database--an exhaustive database of protein-coding intron-containing genes, S. Saxonov, I. Daizadeh, A. Fedorov and W. Gilbert. Nucl. Acids Res. 28(1) , 185-190 (2000). 170. The genomic structure of C14orf1 is conserved across eukarya, C. Ottolenghi, I. Daizadeh, A. Ju, S. Kossida, G. Renault, M. Jacquet, A. Fellous, W. Gilbert and R. Veitia. Mammalian Genome 11 , 786-788 (2000). 171. Footprints of primordial introns on the eukaryotic genome, S.W. Roy, B.P. Lewis, A. Fedorov and W. Gilbert. TIG 17(9) , 496-499 (2001). 172. Intron distribution difference for 276 ancient and 131 modern genes suggests the existence of ancient introns, A. Fedorov, X. Cao, S. Saxonov, S.J. de Souza, S.W. Roy and W. Gilbert. Proc. Natl. Acad. Sci. USA 98, 13177-13182 (2001). 173. Regularities of context-dependent codon bias in eukaryotic genes, A. Fedorov, S. Saxonov and W. Gilbert. Nucl. Acids Res . 30(5), 1192-1197 (2002). 174. Do introns favor or avoid regions of amino-acid conservation? T. Endo, A. Fedorov, S.J. de Souza and W. Gilbert. Mol. Biol. Evol. 19(4), 521-525 (2002). 175. The signal of ancient introns is obscured by intron density and homolog number, S.W. Roy, A. Fedorov and W. Gilbert. Proc. Natl. Acad. Sci. USA 99, 15513-15517 (2002). 176. Large-scale comparison of intron positions among animal, plant, and fungal genes, A. Fedorov, A.F. Merican and W. Gilbert. Proc. Natl. Acad. Sci. USA 99, 16128-16133 (2002). 177. Book review of Watson and DNA: Making a Scientific Revolution by V.K. McElheny, W. Gilbert. Nature 421 , 315-316 (2003). 178. Phylogenetically older introns strongly correlate with module boundaries in ancient proteins, A. Fedorov, S. Roy, X. Cao and W. Gilbert. Genome Res. 13. 1155-1157 (2003). 179. Large-scale comparison of intron positions in mammalian genes shows intron loss but no gain, S.W. Roy, A. Fedorov and W. Gilbert. Proc. Natl. Acad. Sci. USA 100 , 7158-7162 (2003). 180. The universe of exons revisited, S. Saxonov and W. Gilbert. Genetica 118 , 267-278 (2003). 181. Mystery of intron gain, A. Fedorov, S. Roy, L. Fedorova and W. Gilbert. Genome Res . 13 , 2236-2241, (2003). 8 182. The pattern of intron loss, S. W. Roy and W. Gilbert. Proc. Natl. Acad. Sci. USA 102 , 713-718 (2005).

In his photographic art, he has had many one-person shows: Selected exhibitions:

“From Science to Art,” City Gallery, San Diego, CA 2019 “Retrospective,” LabCentral, Cambridge, MA 2018 “Towers,” Viridian Gallery, Chelsea, NYC 2017 “Doors to Nowhere,” Salon R, Cambridge, MA 2017 “Broken City,” Khaki Gallery, Boston, MA 2016 “Journeying,” Permanent exhibition, AGH University, Krakow, Poland 2016 “Broken City” Viridian Gallery, Chelsea, NYC 2016 “Patterns & Recognition,” Seoul National University Bundang Hospital, curated by Chang and Jae Kim 2015-2016 “Transformations,” Viridian Artists, Chelsea, NYC 2014 “Patterns & Recognition,” The Howard Hughes Medical Institute, Janelia Farm, VA 2014 “Wally Gilbert,” CJ Gallery, Art San Diego 2013, San Diego, CA 2013 “Wally Gilbert: A Room of Light,” Milton Art Museum, Canton, MA 2013 “Wally Gilbert: Black & White,” Khaki Gallery, Boston, MA 2013 “Digital Constellations,” Lindau City Museum, Lindau, Germany 2013 “Wally Gilbert: New Black and White Images,” Viridian Artists, Chelsea, NYC 2013 “Wally Gilbert”, CJ Gallery, Art San Diego 2012, San Diego, CA 2012 “En-Lighten,” Khaki Gallery, Boston, MA 2012 “Journeying,” The Artemis Gallery, Krakow, Poland, curated by Wieslawa Piotrowska-Sowadska 2012 “Pattern & Recognition,” The Art Gallery, Antelope Valley College, Lancaster, CA 2012 “Squares, Triangles, and Lines,” Galerie im Einstein, Berlin 2011 “Projekt Norblin,” New Art Wet Music Foundation, Bydgoszcz, Poland 2011 “Squares and Triangles,” Viridian Artists, Chelsea, NYC 2011 “Vanishing,” CJ Gallery, San Diego, CA 2010 “Vanishing Profiles,” Khaki Gallery, Boston, MA 2010 “The Norblin Project and Other Images,” CJ Gallery and OCIO DESIGN GROUP, San Diego,CA 2010 “Wally@Wainwright,” Wainwright Bank, Cambridge, MA 2010 “Vanishing,” BAAK Gallery, Cambridge, MA 2009 Norblin Installation, Poznan, Poland, curated by Jan Kubasiewicz and Zuk Piwkowski 2009 “The Norblin Project and other Images,” CJ Art Gallery, San Diego, CA 2009 “IN COLOR & BEYOND,” Khaki Gallery, Boston, MA 2009 “Fresh Fruit,” Mayyim Hayyim Gallery, Newton, MA 2009 “Stillness and Motion,” Viridian Artists, Chelsea, NYC 2008 “LEEKS & CHAINS,” Khaki Gallery, Wellesley, MA 2008 “The Norblin Project and other Images,” CJ Art Gallery, San Diego, CA 2007 BAAK Gallery, Cambridge, MA 2007 Norblin Installation, Galeria PATIO, Lodz, Poland, curated by Zuk Piwkowski, Jan Kubasiewicz, and Aurelia Mandziuk 2007 Norblin Site Installation, Warsaw, Poland, curated by Jan Kubasiewicz and Zuk Piwkowski 2007 “The Norblin Project: Images of Decay,” American Center for Physics, College Park, MD 2007 “IN COLOR,” Khaki Gallery, Wellesley, MA 2007 “The Norblin Project: Images of Decay,” LACDA, Los Angeles, CA 2006 “The Norblin Project: Images of Decay,” Viridian Artists, Chelsea, NYC 2006 Jock Colville Hall, Churchill College, , Cambridge, UK 2006 Ann Janss Gallery, Los Angeles, CA 2005 Doran Gallery, Massachusetts College of Art, Boston, MA, curated by Jan Kubasiewicz 2004

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