CURRICULUM VITAE

JONATHAN ROGER BECKWITH

Born: December 25, 1935, Cambridge, Massachusetts

1953 Graduated Newton High School, Newton, Massachusetts 1957 A.B. in Chemistry, Harvard College, Cambridge, Massachusetts1961 Ph.D. in Biochemical Sciences, Harvard University, Cambridge, Massachusetts with Dr. Lowell Hager. 1960 Teaching Assistant, University of Illinois, Urbana,

Illinois 1961-65 NIH Postdoctoral Fellow with 1. Dr. Arthur Pardee, University of California, Berkeley, and Princeton

University, Princeton, New Jersey.2. Dr. William Hayes, MRC Microbial Genetic Research Unit,

Hammersmith Hospital, London, England. Jane Coffin Childs Foundation Postdoctoral Fellow with3. Dr. Sidney Brenner, MRC Microbial Genetic Research Unit,

Cambridge, England 4. Dr. Francois Jacob, Institut Pasteur, Paris, France

1965 Associate in Department of Bacteriology and Immunology, Harvard Medical School

1966 Assistant Professor in Department of Bacteriology and Immunology, Harvard Medical School1968 Associate Professor in Department of Bacteriology and Immunology,

Harvard Medical School 1969 Professor in Department of Bacteriology and Immunology, Harvard

Medical School; Professor in Department of Microbiology and Molecular Genetics, Harvard Medical School

1970 Recipient of the Eli Lilly Award for Outstanding Achievements in Microbiology 1970 Guggenheim Fellowship 1971-73 Chairman of the Department of Microbiology and Molecular Genetics1975-2000 Director of Genetics Training Grant, Harvard Medical School1980-American Cancer Society Research Professor 1980-85 Editorial Board. Annual Review of Genetics 1981-91 Member, Scientific Advisory Board, New England BioLabs, Beverly, MA. 1984-Member of the National Academy of Sciences 1985 Visiting Professor, University of California, Berkeley 1985-Fellow of the American Association for the Advancement of Science 1985- Advisory Board, Council for Responsible Genetics

1985- Advisory Board, Eritrean Relief Committee1986- Fellow of the American Academy of Arts and Science1986 Merit Award from the National Institutes of Health1986-90 Member, Scientific Advisory Board, International Institute of Genetics and Biophysics, Naples, Italy. 1986- Member of the Genetic Screening Study Group1987- 1999 Editorial Board, Genes and Development 1988 Jacques Monod Memorial Lecturer, Institut Pasteur, Paris 1989-Associate Member of the European Molecular Biology Organization1989-1995 Member of the Working Group on Ethical, Legal and Social Implications of the Human Genome Project for the National Center for Human Genome Research, NIH1990-92 Editorial Board, EMBO Journal1990-1993 President, Board of Directors, Science for the People1991 Samuel Rudin Visiting Professor-Columbia Univ. College of Physicians & Surgeons 1991 Lester O. Krampitz Lecturer in Microbiology, Case Western Reserve University1992-Fellow of the American Academy of Microbiology 1993 Awarded Genetics Society of America Medal1994-1998 Consultant to Genentech Corporation1997-1998 Member, Advisory Committee for "Genes, Environment and Behavior," a high school biology curriculum unit prepared by BSCS, Colorado Springs, CO.1998 Keynote Speaker at 99th Annual Meeting of the American Society of

Microbiology1999 Roger Stanier Memorial Lecturer, University of California, Berkeley1999-2002 Elected to Harvard Medical School Faculty Council2000-Editorial Board, ASM News2000-2002 Member of "Behavioral Genetics Working Group" of the Hastings Center for Bioethics and the American Assoc. for the Advancement of Science2000 Helen R. Whiteley Memorial Lecturer, University of Washington, Seattle2001 Frank and Joan Rothman Commencement Forum Lecturer, Brown University2001 Pan American Association for Biochemistry and Molecular Biology Award Lecturer for 2001 (presented in Argentina)2002-Member, Scientific Advisory Board, Center for Microbial Sciences, San Diego State University2002-Member, Advisory Board, Program in Science, Technology and Society Program, Kennedy School of Government, Harvard University2005 Recipient of Honorary Doctorate in Humane Letters from the Univ.of

Massachusetts, Lowell2005 Awarded Abbott Lifetime Achievement Award of the American Society

for Microbiology2006-2008 Member of ASM committee to select Abbott Lifetime Achievement Award2007 Elected Honorary Member of Phi Beta Kappa by the Alpha Iota Chapter of

Massachusetts at Harvard College June 5, 2007, in recognition of high attainments in liberal scholarship

2009 Awarded: 1)Selman Waksman Award in Microbiology from the National Academy of Sciences for “excellence in the field of microbiology” and

2)The Edinburgh Medal for professional achievements that have made a significant contribution to the understanding and well-being of humanity 3)Harvard Medical School Division of Medical Sciences Hooding Speaker “For inspired teaching and exceptional commitment and service to graduate education”.

2010 Brina Sheeman Shackleford Teaching Award from the PhD program in Biological and Biomedical Sciences, Harvard Medical School

2011 ASM Graduate Microbiology Teaching Award2012 Keynote Speaker at the Bacteria, Archaea and Phages Meeting at the Cold Spring Harbor

Laboratory2012 Keynote Speaker at Quest for Research Excellence Conference organized by the Office

of Research Integrity of the National Institutes of Health.2013 Keynote Speaker on Bioethics, University of Rochester Medical Center.

.

Addendum

I am American Cancer Society Professor of Microbiology and Molecular Genetics at Harvard Medical School. I teach genetics and I and my coworkers do research in bacterial genetics. Our work has been in the area of gene expression, the mechanism of protein secretion, the structure and function of membrane proteins, disulfide bond formation in proteins and cell division. I with my coworkers have published over 300 papers on these subjects. I have graduated 35 PhD students (as of 2009). I have also been active since 1969 in activities and public discussions of issues of the social impact of genetics. For many years I was active in Science for the People, an organization which worked to demystify scientific issues for the public and to describe the social and political influences on scientific research and its uses. More recently, as a member of the Genetic Screening Study Group, I have worked on critiques of biological determinist thinking. We have been critical of such areas as sociobiology, arguments that variation in human traits such as IQ and anti-social behavior are largely due to genetics, and claims that women are inferior to men in their mathematical ability for biological reasons. I have also been concerned with the social impact of genetic screening programs and of the Human Genome Project. As a result I was appointed to the first Working Group on Ethical, Legal and Social Implications of the Human Genome Project of the National

Center for Human Genome Research. In addition, our Study Group has published two reports on surveys that reveal "genetic discrimination" in such areas as health insurance resulting from the use of information from genetic tests. I am author of a memoir entitled “Making Genes, Making Waves: A Social Activist in Science” published by Harvard University Press in 2002.

Author of over 300 scientific publications and over 100 publications on science and society issues.

PUBLICATIONS:

BOOKS:Beckwith, J.R. and Zipser, D. editors "The Lac Operon." A book published by the Cold Spring Harbor Laboratory. (1970).

Beckwith, J., Davies, J. And Gallant, J.A. editors. Gene Function in Prokaryotes. Cold Spring Harbor Laboratory Press, Cold Spring Harbor. (1983).

Beckwith, J., and Silhavy, T. The Power of Bacterial Genetics: A Literature-Based Course. Cold Spring Harbor Laboratory Press, Cold Spring Harbor. 828 pages(1992).

Beckwith, J. Making Genes, Making Waves: A Social Activist in Science. Harvard University Press (2002). (Translated into Korean, 2009.)

Alper, J.S., Ard, C., Asch, A., Beckwith, J., Conrad, P., and Geller, L., eds. The Double-Edged Helix: Social Implications of Genetics in a Diverse Society. Johns Hopkins Univ. Press. Baltimore. (2002).

BOOKS AND SCIENTIFIC PUBLICATIONS: 1. Shaw, P.D., Beckwith, J., and Hager, L.P. Biological chlorination. II. The biosynthesis of delta-chlorolevulinic acid. J. Biol. Chem. 234:2560-2564 (1960).

2. Beckwith, J.R. and Hager, L.P. The synthesis of D, L-caldariomycin. J. Org. Chem. 26:5206-5208 (1961).

3. Pardee, A.B., and Beckwith, J.R. Genetic determination of constitutive enzyme levels. Biochim. Biophys. Acta. 60:452-454 (1962).

4. Beckwith, J.R., Pardee, A.B., Austrian, R. and Jacob, F. Co-ordination of the synthesis of the enzymes in the pyrimidine pathway in E. coli. J. Mol. Biol. 5:618-634 (1962).

5. Pardee, A.A., and Beckwith, J.R. Control of constitutive enzyme synthesis. In "Informational Macromolecules," Rutgers University, September, 1962. 255-269.

6. Beckwith, J.R., Clark, R. and Hager, L.P. Biological chlorination. VII. Studies on the biosynthesis of caldariomycin. J. Biol. Chem. 238:3086-3090 (1963).

7. Beckwith, J.R. and Hager, L.P. Biological chlorination. VIII. Later intermediates in the biosynthesis of caldariomycin. J. Biol. Chem. 238:3091-3094 (1963).

8. Beckwith, J.R. Restoration of operon activity by suppressors. Biochim. Biophys. Acta.76:162-164 (1963).

9. Beckwith, J.R. A deletion analysis of the lac operator region in E. coli. J. Mol. Biol.8:427-430 (1964).

10. Beckwith, J.R. Restoration of operon activity by suppressors, in Stubbe, H. (Ed.) Structure and Function of the Genetic Material (Erwin-Baur-Symposium III, 1963). Abhdlg. d. Dadb., Klasse f. Medizin, Mo. 1-119-124 (1964).

11. Broda, P., Beckwith, J.R. and Scaife, J. The characterization of a new type of F-prime factor in E. coli Kl2. Genetical Research 5:489-494 (1964).

12. Brenner, S. and Beckwith, J.R. Ochre mutants, a new class of suppressible nonsense mutants. J. Mol. Biol. 13:629-637 (1965).

13. Signer, E.R., Beckwith, J.R. and Brenner, S. Mapping of suppressor loci in E. coli. J. Mol. Biol. 14-153-166 (1965).

14. Newton, A., Beckwith J.R., Zipser, D. and Brenner, S. Nonsense mutants and polarity in the lac operon of E. coli. J. Mol. Biol. 14:290-296 (1965).

15. Beckwith, J.R., and Signer, E.R. Transposition of the lac operon of E. coli. I. Inversion of the lac operon and transduction of lac by ö80. J. Mol. Biol. 19:254-265 (1966).

16. Gorini, L. and Beckwith, J.R. Suppression. Ann. Rev. Microbiol. 20:401-422 (1966).

17. Fox, C.R., Beckwith J.R., Epstein, W. and Signer, E.R. Transpositon of the lac region of E. coli. II. On the role of thiogalactoside transacetylase in lactose metabolism. J. Mol. Biol. 19:576-579 (1966).

18. Signer, E.R. and Beckwith, J.R. Transposition of the lac region of E. coli. The mechanism of attachment of coliphage ö80 to the bacterial chromosome. J. Mol. Biol. 22:33-51 (1966).

19. Beckwith, J.R., Signer, E.R. and Epstein, W. Transposition of the lac region of E. coli. Cold Spr. Harb. Symp. Quant. Biol. 31:393-401 (1966).

20. Scaife, J.G. and Beckwith, J.R. Mutational alteration of the maximal level of lac operon expression. Cold Spr. Harb. Symp. Quant. Biol. 31:403-408 (1966).

21. Beckwith, J.R. and Signer, E.R. The effects of inversion, transposition and deletion on lac operon activity in E. coli. In "Regulation of Nucleic Acid and Protein Biosynthesis." Ed. Koningsberger Bosch, Elsevier, Amsterdam, 288-297 (1967).

22. Beckwith, J.R. The regulation of lac operon activity in E. coli. Science 156:597-604 (1967).

23. Arditti, R.R., Miller, J.H., Scaife, J.G. and Beckwith, J.R. The role of promoter sites in gene expression, in Molecular Genetics. A symposium of the GesellschaftDeutscher Naturforscher und Arzter, Ed. H.G. Wittman and H. Schuster. Springer-Verlag, Berlin, 213-222 (1967).

24. Ippen, K., Miller, J.H., Scaife, J.G. and Beckwith, J.R. New controlling element in the lac operon of E. coli. Nature 217:825-827 (1968).

25. Ohlsson, B.M., Strigini, P. and Beckwith, J.R. Allelic amber and ochre suppressors. J. Mol. Biol. 36:209 (1968).

26. Arditti, R.R., Scaife, J.G. and Beckwith, J.R. The nature of lac promoter mutants. J. Mol. Biol. 38:421 (1968).

27. Miller, J.H., Ippen, K., Scaife, J.G. and Beckwith, J.R. The promoter-operator region of the lac operon of E. coli. J. Mol. Biol. 38:413 (1968).

28. Miller, J.H., Beckwith, J.R., and Muller-Hill, B. The direction of reading of a regulatory gene in E. coli. Nature 220:1287 (1968).

29. Epstein, W. and Beckwith, J.R. Control of gene expression. Annual Reviews of Biochemistry 37:411-436 (1968).

30. Schwartz, D.O. and Beckwith, J.R. Mutagens which cause deletions in E. coli. Genetics 61:371 (1969).

31. Reznikoff, W., Miller, J.H., Scaife, J.G., and Beckwith, J.R. A mechanism for repressor action. J. Mol. Biol. 43:201 (1969).

32. Reznikoff, W., and Beckwith, J.R. Genetic evidence that the operator locus is distinct from the z gene in the lac operon of E. coli. J. Mol. Biol. 43:215 (1969).

33. Gottesman, S., and Beckwith, J.R. Directed transposition of the ara operon: A technique for the isolation of specialized transducing phages for any E. coli gene. J. Mol. Biol. 44:117-127 (1969).

34. Silverstone, A., Magasanik, B., Reznikoff, W.S., Miller, J.H. and Beckwith, J.R. Catabolite sensitive site of the lac operon. Nature 221-1012 (1969).

35. Shapiro, J., MacHattie, L., Eron, L., Ihler, G., Ippen, K., Beckwith, J., Arditti, R., Reznikoff, W., and MacGillivray, R. The isolation of pure lac operon DNA. Nature 224: 768-774 (1969).

36. Schwartz, D.O., and Beckwith, J.R. Mutants missing a factor necessary for the expression of catabolite-sensitive operons in E. coli in "The Lac Operon," 4l7-422 (1970).

37. Eron, L., Beckwith, J.R. and Jacob, F. Deletions of translation start signals in the lac operon. in "The Lac Operon," 353-358 (1970).

38. Zubay, G., Schwartz, D.O. and Beckwith, J.R. The mechanism of activation of catabolite-sensitive genes: a positive control system. Proc. Natl. Acad. Sci. 66:104-110 (1970).

39. Arditti, R.R., Eron, L., Zubay, G., Tocchini-Valentini, G., Conaway, S., and Beckwith, J. "In vitro" transcription of the lac genes. Cold Spr. Harb. Symp. Quant. Biol. 35:437-442 (1970).

40. Beckwith, J.R. and Zipser, D. editors "The Lac Operon." A book published by the Cold Spring Harbor Laboratory. (1970).

41. Eron, L., Arditti, R.R., Zubay, G., Connaway, S. and Beckwith, J. A cyclic-AMP binding protein which acts on the transcription process. Proc. Natl. Acad. Sci. 68:215-218 (1971).

42. Miller, J.H., Reznikoff, W.S., Silverstone, A.E., Ippen, K., Signer, E.R. and Beckwith, J.R. Fusions of the lac and trp regions of the E. coli chromosome. J. Bact. 104:1273-1279 (1970).

43. Beckwith, J.R. Lac: the genetic system. A review chapter in "The Lac Operon" 5-26 (1970).

46. Beckwith, J.R. Gene expression in bacteria and some concerns about the misuse of science. Bact. Revs. 34:222-227 (1970).

47. Eron, L., Morse, D., Reznikoff, W. and Beckwith, J. Fusions of the lac and trp regions of Escherichia coli: Covalently fused messenger RNA. J. Mol.

Biol. 60:203 (1971).

48. Zubay, G., Schwartz, D., and J. Beckwith. The mechanism of activation of catabolite-sensitive genes. Cold Spr. Harb. Sym. Quant. Biol. 35:433-435 (1970).

49. Ippen, K., Shapiro, J.A., and Beckwith, J. Transposition of the lac region to the gal region of the Escherichia coli chromosome: Isolation of λlac transducing phage. J. Bacteriol. 108:5-9 (1971).

50. Beckwith, J., Grodzicker, T., and Arditti, R. Evidence for two sites in the lac promoter region. J. Mol. Biol. 69-155-160 (1972).

51. Arditti, R., Grodzicker, T., and Beckwith, J.R. Cyclic adenosine monophosphate-independent mutants of the lactose operon of Escherichia coli. J. Bacteriol. 114:652-655 (1973).

52. Brickman, E., Soll, L., and Beckwith, J. Genetic characterization of mutations which affect catabolite sensitve operon in E. coli, including deletions of the gene for adenyl cyclase. J. Bacteriol. 116:582-587 (1973).

53. Beckwith, J., Rossow, P. Analysis of genetic regulatory mechanisms: the approaches and the pitfalls. Annu. Rev. Genetics 8: (1974).

54. Mitchell, D., Reznikoff, W., Beckwith, J. Genetic fusions defining trp and lac regulatory elements. J. Mol. Biol. 93:331 (1975).

55. Sabourin, D. and Beckwith, J. Deletion of the Escherichia coli gene crp. J. Bacteriol. 122:338-340 (1975).

56. Brickman, E., and Beckwith, J. Analysis of the regulation of E. coli alkaline phosphatase synthesis using deletions and o80 transducing phages. J. Mol. Biol. 96:307-316 (1975).

57. Mitchell, D., Reznikoff, W., Beckwith, J. Genetic fusions that help define a transcription termination region in E. coli. J. Mol. Biol. 101:441-457 (1976).

58. Inouye, H., Pratt, C., Beckwith, J. and Torriani, A.M. Alkaline phosphatase synthesis in a cell-free system using DNA and RNA templates. J. Mol. Biol. 110:75-87 (1977).

59. Silhavy, T., Casadaban, M.J., Shuman, H.A., and Beckwith, J.R. Conversion of â-galactosidase to a membrane-bound state by gene fusion. Proc. Nat'l. Acad. Sci. 73:3423-3427 (1976).

60. Casadaban, M.J., Silhavy, T.J., Berman, M., Shuman, H.A., Sarthy, A., and Beckwith, J.R. Construction and use of gene fusions using bacteriophage MU insertions. in DNA Insertion Elements, Plasmids and Episomes. Bukhari, A.I., Shapiro, J., and Adhya, S. Ed. (Cold Spring Harbor Laboratory, Cold Spring Harbor, N.Y. p. 531-536 (1977).

61. Guarente, L., Mitchell, D.H., and Beckwith, J. Transcription termination at the end of the trp operon of E. coli. J. Mol. Biol. 112:423-436 (1977)

62. Inouye, H., and Beckwith, J. Synthesis and processing of an E. coli alkaline phosphatase precursor in vitro. Proc. Natl. Acad. Sci. 74:1440-1444 (1977).

63. Silhavy, T., Shuman, H., Beckwith, J., and Schwartz, M. The use of gene fusions to study outer membrane protein localization in Escherichia coli. Proc. Natl. Acad. Sci. 74:5411-5415 (1977).

64. Guarente, L., and Beckwith. J. A mutant RNA polymerase of E. coli which terminates transcription in strains making defective rho factor. Proc. Natl Acad. Sci. 75:294-297 (1978).

65. Zabin, I., Fowler, A.V. and Beckwith, J.R. Position of the mutation in β-galactosidase ochre mutant Ull8. J. Bacteriol. 133:437-438 (1978).

66. Bassford, P., Beckwith, J., Berman, J., Brickman, E., Casadaban, M., Guarente, L., Saint-Girons, I., Sarthy, A., Schwartz, M., Shuman, H., and Silhavy, T. Genetic fusions of the lac operon: a new approach to the study of biological processes in The Operon, ed. by J.H. Miller and W.S. Reznikoff. Cold Spring Harbor Laboratory. 245-261 (1978).

67. Beckwith, J., Silhavy, T., Inouye, H., Shuman, H., Schwartz, M., Emr, S., Bassford, P., and Brickman, E. The mechanism of localization of proteins in Escherichia coli in Dahlem Workshop on Transport of Macromolecules in cellular systems. Ed. by S. Silverstein. Dahlem, Berlin, 299-314 (1978).

68. Shuman, H.A., and Beckwith, J. Mutants of E. coli Kl2 that allow transport of maltose via the â-galactoside transport system. J. Bacteriol. 137:365-372 (1979).

69. Bassford, P.J., Silhavy, T., Shuman, H., Emr, S., Brickman, E., and Beckwith, J. The use of gene fusions to study protein localization in E. coli. in Microbiology 1979, ed. by D. Schlesinger. ASM Press, Washington. p. 30-33 (1979).

70. Bassford, P., and Beckwith, J. Mutants of Escherichia coli which accumulate the precursor of a secreted protein in the cytoplasm. Nature

277:538-541 (1979).

71. Berman, M., and Beckwith, J. Fusions of the lac operon to the transfer RNA gene tyrT of Escherichia coli. J. Mol. Biol. 130:285-301 (1979).

72. Berman, M., and Beckwith, J. Use of gene fusions to isolate promoter mutants in the transfer RNA gene, tyrT, of Escherichia coli. J. Mol. Biol. 130:305-315 (1979). 73. Silhavy, T.J., Bassford, P.J., and Beckwith, J.R. A genetic approach to the study of protein localization in E. coli in Bacterial Outer Membrane: Biosynthesis, Assembly and Functions. Ed. by M. Inouye. J. Wiley and Sons. 203-254 (1979).

74. Wanner, B., Sarthy, A., and Beckwith, J. Escherichia coli pleiotropic mutant that reduces amounts of several periplasmic and outer membrane proteins. J. Bacteriol. 140:229-239 (1979).

75. Brickman, E., Silhavy, T., Bassford, P.J., Shuman, H., and Beckwith. J. Sites within the lacZ gene for formation of hybrid active â-galactosidase molecules. J. Bacteriol. 139:13-18 (1979).

76. Bassford, P., Silhavy, T.J., and Beckwith, J. Use of gene fusion to study secretion by maltose-binding protein in Escherichia coli periplasm. J. Bacteriol. 139:19-31 (1979).

77. Silhavy, T.J., Brickman, E., Bassford, P.J., Casadaban, M.J., Shuman, H.A., Schwartz, V., Guarente, L., Schwartz, M., and Beckwith, J.R. Structure of the malB region in Escherichia coli K12. II. Genetic map of the malE,F,G operon. Mol. Gen. Genetics 174:249-259 (1979).

78. Guarente, L., Beckwith, J., Wu, A.M., and Platt, T. Localization of mutations in the transcription termination region of the tryptophan operon of E. coli. J. Mol. Biol. 133, 189-197 (1979).

79. Sarthy, A., Fowler, A., Zabin, I., and Beckwith, J. The use of gene fusions to determine a partial signal sequence of alkaline phosphatase. J. Bacteriol. 139, 932-939 (1979).

80. Shuman, H.A., Silhavy, T.J., and Beckwith, J. Labelling of protein with â-galactosidase by gene fusion: Identification of a cytoplasmic membrane component of the Escherichia coli maltose transport system. J. Biol. Chem. 255:168-174 (1980).

81. Silhavy, T.J., Emr, S., Schwartz, M., and Beckwith, J. l979. The use of gene 0fusions to study outer membrane protein localization in Limited proteolysis in microorganism.. ed. by G.N. Cohen and H. Holzer. DHEW

Publications No. (NIH) 79-1591. pp. 177-179.

82. Chang, C.N., Model, P., Inouye, H., and Beckwith, J. Alkaline phosphatase precursor is processed to the mature enzyme by an E. coli inner membrane preparation. J. Bacteriol.142:726-728 (1980).

83. Welply, J.K., Fowler, A.V., Beckwith, J.R., and Zabin, I. Map positions of early nonsense and deletion mutations in lacZ. J. Bacteriol. 142:732-734 (1980).

84. Bedouelle, H., Bassford, P.J., Jr., Fowler, A.V., Zabin, I., Beckwith, J., and Hofnung, M. The nature of mutational alterations in the signal sequence of the maltose binding protein of Escherichia coli. Nature 285:78-81 (1980).

85. Wu, A.M., Chapman, A.B., Platt, T., Guarente, L.P. and Beckwith, J. Deletions of distal sequences affect termination of transcription at the end of the tryptophan operon in Escherichia coli. Cell 19:829-836 (1980).

86. Sarthy, A., Michaelis, S., and Beckwith, J. Deletion map of the Escherichia coli structural gene for alkaline phosphatase. J. Bacteriol. 145:288-292 (1981).

87. Sarthy, A., Michaelis, S., and Beckwith, J. Use of gene fusions to determine the orientation of the phoA gene in the Escherichia coli chromosome. J. Bacteriol. 145:5145:293-298 (1981).

88. Ito, K., Bassford, P.J., Jr., and Beckwith, J. Protein localization in E. coli. Is there a common step in the secretion of periplasmic and outer membrane proteins? Cell 24:707-714 (1981).

89. Beckwith, J. A genetic approach to characterizing complex promoters in Escherichia coli. Cell 23:307-308 (1981).

90. Brown, S., Brickman, E.R., and Beckwith, J. Blue ghosts: A new method for isolating amber mutants in essential genes in Escherichia coli. J. Bacteriol. 146:422-425 (1981).

91. Inouye, H., Michaelis, S., Wright, A., and Beckwith, J. Cloning restriction map and generation of deletion mutants in vitro of the alkaline phosphatase structural gene (phoA) of E. coli. J. Bacteriol. 146:668-675 (1981).

92. Bassford, P.J., Emr, S.D., Silhavy, T.J., Beckwith, J., Bedouelle, H., Clement, J.-M., Hedpeth, J., and Hofnung, M. The genetics of protein secretion in Escherichia coli in Methods in Cell Biology. eds. A. Hand and C. Oliver. 23:28-33 (1981).

93. Oliver, D., and Beckwith, J. E. coli mutant pleiotropically defective in the secretion of E. coli proteins. Cell 25:765-772 (1981).

94. Ito, K., and Beckwith, J. Role of the mature protein sequence of maltose binding protein in its secretion across the E. coli cytoplasmic membrane. Cell 25:143-150 (1981).

95. Inouye, H., Barnes, W., and Beckwith, J. The signal sequence of alkaline phosphatase of E. coli. J. Bacteriol. 149:434-439 (1982).

96. Oliver, D.B., and Beckwith, J. Identification of a new gene (secA) and gene product involved in the secretion of envelope proteins in E. coli. J. Bacteriol. 150:686-691 (1982).

97. Garwin, J.L., and Beckwith, J. Secretion and processing of ribose binding protein Escherichia coli K-12. J. Bacteriol. 149:789-792 (1982).

98. Oliver, D., Kumamoto, C., Quinlan, M., and Beckwith, J. Pleiotropic mutants affecting the secretory apparatus of E. coli. Ann. Microbiol. (Inst. Pasteur) 133A:105-110 (1982).

99. Garwin, J.L., and Beckwith, J. Genetic approaches for studying protein localization. In Membranes and Transport, Vol. 1 (Ed. A.N. Martonosi) Plenum Publishing Corp, 1982

100. Michaelis, S., and Beckwith, J. Mechanism of incorporation of cell envelope proteins in Escherichia coli. Ann. Rev. Microbiol. 36:435-465 (1982).

101. Silhavy, T.J., and Beckwith, J. Isolation and characterization of mutants of Escherichia coli K12 affected in protein localization. in Methods in Enzymology 97:11 (1983).

102. Oliver, D., and Beckwith, J. Regulation of a membrane component required for protein secretion in Escherichia coli. Cell 130:311-3l9 (1982).

102a.Beckwith, J., Davies, J. And Gallant, J.A. editors. Gene Function in Prokaryotes. Cold Spring Harbor Laboratory Press, Cold Spring Harbor. (1983).

103. Beckwith, J., and Silhavy, T.J. Genetic analysis of protein export in Escherichia coli. Methods in Enzymology 97:3-11 (1983).

104. Michaelis, S., Guarente, L., and Beckwith, J. In vitro construction and characterization of phoA-lacZ gene fusions in Escherichia coli. J. Bacteriol.

154:356-365 (1983).

105. Michaelis, S., Inouye, H., Oliver, D., and Beckwith, J. Mutations that alter the signal sequence of alkaline phosphatase of Escherichia coli. J. Bacteriol. 154:366-374 (1983).

106. Kumamoto, C., and Beckwith, J . Mutations in a new gene, secB, cause defective protein localization in Escherichia coli. J. Bacteriol. 135:254-260 (1983).

107. Oliver, D., Kumamoto, C., Brickman, E., Ferro-Novick, S., Garwin, J., and Beckwith, J. Genetics of the secretory apparatus of E. coli. in Gene Expression. p. 371-383. UCLA Symposia on Molecular and Cellular Bioloy, New Series, eds. D. Hamer and M. Rosenberg (Alan R. Liss, Inc., N.Y. (1983).

108. Lopilato, J.E., Garwin, J.L., Emr, S.D., Silhavy, T.J., and Beckwith, J.R. D-ribose metabolism in Escherichia coli K12. Genetics, regulation and transport. J. Bacteriol. 158:665-673 (1984).

109. Brickman, E.R., Oliver, D.B., Garwin, J.L., Kumamoto, C. and Beckwith, J.R. The use of extragenic suppressors to define genes involved in protein export in Escherichia coli. Molecular and General Genetics. 196:24-27 (1984).

110. Froshauer, S. and Beckwith, J. The nucleotide sequence of the gene for malF protein, an inner membrane component of the maltose transport system of Escherichia coli. J. Biol. Chem. 259:10896-10903 (1984).

111. Kumamoto, C.A., Oliver, D.B., and Beckwith, J.R Signal sequence mutations disrupt the coupling between secretion and translation in Escherichia coli. Nature 308:863-864 (1984).

112. Ebright, R., P.Cossart, B. Gicquel-Sanzey, and J. Beckwith. The molecular basis of DNA-sequence recognition by the catabolite gene activator protein (CAP): Detailed inferences from three mutations that alter DNA-sequence specificity. Proc. Natl. Acad. Sci. 81:7274-7278 (1984).

113. Ebright, R., P. Cossart, B. Gicquel-Sanzey and J. Beckwith. Mutations that alter the DNA sequence specificity of the catabolite gene activator protein (CAP) in E. coli. Nature 311:232-235 (1984).

114. Ferro-Novick, S., Honma, M. and Beckwith, J. The product of gene secC is involved in the synthesis of exported proteins in Escherichia coli. Cell 38:211-217 (1984).

115. Kumamoto, C., and Beckwith, J. Evidence for specificity at an early step in protein export in Escherichia coli. J. Bact. 163:267-274 (1985).

116. Lee, C.A., Fournier, M.J., and Beckwith, J. The 6S RNA of Escherichia coli is not essential for growth of protein secretion. J. Bacteriol. 161:1156-1161 (1985).

117. Beckwith, J.R., Galizzi, A., and Smith, G.R. The tools of Bacterial Genetics in Genetics of Bacteria ed. by J. Scaife, D. Leach and A. Galizzi. Academic Press p. 1-23, (1985).

118. Ebright, R., and Beckwith. The catabolite gene activator protein (CAP) is not required for indole-3-acetic acid to activate transcription of the araBAD operon of Escherichia coli K-12. Mol. Gen. Genet. 201:51-55. (1985).

119. Silhavy, T.J., and Beckwith, J. Uses of lac fusions for the study of biological problems. Microbiol. Revs. 49: 398-418 (1985).

120. Beckwith, J., and Ferro-Novick, S. Genetic studies on protein export in bacteria. In Current Topics in Microbiology and Immunology. Protein Secretion and Export in Bacteria. eds. H.C. Wu and P.C. Tai. Vol. 125, p. 5-28, Springer-Verlag, New York. (1986)

121. Beckwith, J. The Operon: an historical account. In Escherichia coli and Salmonella typhimurium. Cellular and Molecular Biology. eds. F. Neidhart et al. ASM Press. 1439-1443 (1987).

122. Beckwith, J. The lac operon. In Escherichia coli and Salmonella typhimurium. eds. F. Neidhart et al. Cellular and Molecular Biology. ASM Press. 1444-1452 (1987).

123. Manoil, C., and J. Beckwith. TnphoA: A transposon probe for protein export signals. Proc. Natl. Acad Sci. 82:8129-8133 (1985).

124. Strauch, K., C.A. Kumamoto and J. Beckwith. Does secA mediate coupling between secretion and translation in Escherichia coli? J. Bacteriol. 166:505-512 (1986).

125. Lee, C. and J. Beckwith. Suppression of growth and protein secretion defects in Escherichia coli secA mutants by decreasing protein synthesis. J. Bacteriol. 166:878-883 (1986).

126. Lee, C. and J. Beckwith. Cotranslational and posttranslational protein translocation in prokaryotic systems. Ann. Rev. Cell Biol. 2:315-336 (1986).

127. Michaelis, S., J. Hunt and J. Beckwith. Effects of signal sequence mutations on the kinetics of alkaline phosphatase export to the periplasm in Escherichia coli. J. Bacteriol. 167:160-167 (1986).

128. Lopilato, J., S. Bortner and J. Beckwith. Mutations in a new chromosomal gene of E. coli K-12,pcn, reduce plasmid copy number of pBR322 and its derivatives. Mol. Gen. Genetics 205:285-290 (1986).

129. Boyd, B., C.-D. Guan, S. Willard, W. Wright and J. Beckwith. The enzymatic activity of alkaline phosphatase precursor depends on its cellular location. in A.M. Torriani et al., ed. Phosphate Metabolism in Microorganisms. p. 89-93, (1987).

130. Manoil, C., D. Boyd and J. Beckwith. Using fusions to alkaline phosphatase to study protein insertion into the cytoplasmic membrane. in A.M. Torriani et al., ed. Phosphate Metabolism in Microorganisms, ASM Press, p. 94-98,1987. 13l. Manoil, C., and J. Beckwith. A genetic approach to analyzing membrane protein topology. Science 233: 1403-1408 (1986).

132. Gardel, C., S. Benson, J. Hunt, S. Michaelis and J. Beckwith. secD, a new gene involved in protein export. J. Bacteriol. 169-1286-1290 (1987).

133. Boquet, P., C. Manoil and J. Beckwith. Use of TnphoA to detect genes for exported proteins in Escherichia coli: Identification of the plasmid encoded gene for a periplasmic acid phosphatase. J. Bacteriol 169:1663-1669 (1987).

134. Gutierrez, C., J. Barondess, C. Manoil and J. Beckwith. The use of the transposon TnphoA to detect genes for cell envelope proteins subject to a common regulatory stimulus: The analysis of osmo-regulated genes in E. coli J. Mol. Biol. 195:289-297 (1987). 135. Ebright, R., Kolb, A., Buc, H., Kunkel, T.A., Krakow, J.S. and J. Beckwith. Role of Glul8l in DNA-sequence recognition by the catabolite gene activator protein (cap) of Escherichia coli: Altered DNA-sequence recognition properties of Vall8l and Leul81 CAP. Proc. Natl. Acad. Sci.84:6083-6087 (1987).

136. Boyd, D., C. Manoil and J. Beckwith. Determinants of Membrane protein topology. Proc. Natl. Acad. Sci. 84:8525-8529 (1987).

137. Beckwith, J. Genetics at the Institut Pasteur: Substance and Style. ASM News 54: #10, (1987) 551-557.

138. Froshauer, S., G.N. Green, D. Boyd, K. McGovern and J. Beckwith. Genetic analysis of the membrane insertion and topology of MalF, a cytoplasmic membrane protein of Escherichia coli. J. Mol. Biol.200:501-11 (1988).

139. Riggs, P , Derman, A., and J. Beckwith. A mutation affecting the regulation of a secA-lacZ fusion defines a new sec gene. Genetics 118:571-579 (1988).

140. Strauch, K., and J. Beckwith. An E. coli mutant preventing degradation of abnormal periplasmic proteins. Proc. Natl. Acad. Sci. 85:1576-1580 (1988).

141. Manoil, C., D.H. Boyd and J. Beckwith. Molecular genetic analysis of membrane protein topology. Trends in Genetic Science. 4:223-226, #8, August 1988.

142. Li, P., J. Beckwith, and H. Inouye. Alteration of the amino-terminus of the mature sequence of a periplasmic protein can severely affect protein export in Escherichia coli. Proc. Natl. Acad. Sci. 85:7685-7689 (1988).

143. Boyd, D., and J. Beckwith. Positively charged amino acids can act as topogenic determinants in membrane proteins. Proc. Natl. Acad. Sci. 86:9446-9450 (1989). 144. Strauch, K., K. Johnson and J. Beckwith. Characterization of degP, a gene required for proteolysis in the cell envelope and essential for growth of Escherichia coli at high temperature. J. Bacteriol. 171:2689-2696 (1989).

145. Lee, C., P. Li, H. Inouye, E. Brickman and J. Beckwith. Genetic studies on the inability of â-galactosidase to be translocated across the E. coli cytoplasmic membrane. J. Bacteriol. 171:4609-4616 (1989).

146. Schatz, P.J., P.D. Riggs, A. Jacq, M.J. Fath and J. Beckwith. The secE gene codes for an integral membrane protein required for protein export in E. coli. Genes and Development. 3:1035-1044 (1989).

147. San Millan, J.L., D. Boyd, R. Dalbey, W. Wickner and J. Beckwith. The use of phoA fusions to study the topology of the E. coli inner membrane protein, leader peptidase. J. Bacteriol. 171:5536-5541 (1989).

148. Manoil, C., J.J. Mekalanos and J. Beckwith. Alkaline phosphatase fusions; sensors of subcellular location. J. Bacteriol. 172: 515-518 (1990).

149. Boyd, D., C. Manoil, S. Froshauer, J.-L. San Millan, N.Green, K.

McGovern, C. Lee and J. Beckwith. The use of gene fusions to study membrane protein topology. in Protein Folding: Deciphering the Second Half of the Genetic Code. eds. J. King and L. Gierasch. AAAS. (1990). pp.314-322.

150. Barondess, J., and J. Beckwith. A bacterial virulence determinant encoded by lysogenic coliphage lambda. Nature. 346:871-874 (1990)

151. Gardel, C., K. Johnson, A. Jacq and J. Beckwith. The secD locus of Escherichia coli codes for two membrane proteins required for protein export. EMBO J. 9: 3209-3216 (1990).

152. Schatz, P., and J. Beckwith. The genetics of protein secretion in Escherichia coli.Ann. Rev. Genet.24: 215-248 (1990)

153. Ehrmann, M., D. Boyd and J. Beckwith. Genetic analysis of membrane protein topology by a sandwich gene fusion approach. Proc. Natl. Acad. Sci. 87: 7574-7578 (1990).

154. Boyd, D., and J. Beckwith. The role of charged amino acids in the localization of secreted and membrane proteins. Cell. 62: 1031-1033 (1990).

155. Baba, T., A. Jacq, E. Brickman, J. Beckwith, T. Taura, C. Ueguchi, Y. Akiyama and K. Ito. Characterization of cold-sensitive secY mutants of Escherichia coli. J. Bacteriol. 172:7005-7010 (1990).

156. Signer, E.R., and Beckwith, J. Cloning with ö80lac: The French Connection. BioEssays. 12:503-507 (1990).

157. Beckwith, J. The all-purpose gene fusion. in Perspectives on Cellular Regulation: From Bacteria to Cancer. ed. by M. Inouye et al. Wiley-Liss. (1991). pp. 85-94.

158. Beckwith, J. Strategies for finding mutants. in Methods in Enzymology, ed. J.H. Miller. Academic Press, San Diego. 204:3-18 (1991).

159. Ehrmann, M. and J. Beckwith. Proper insertion of a complex membrane protein in the absence of its amino-terminal export signal. J. Biol. Chem. 266:16530-16533 (1991).

160. Schatz, P.J., K.L. Bieker, K.M. Ottemann, T.J. Silhavy and J. Beckwith. One of three transmembrane stretchs is sufficient for the functioning of the SecE protein, a membrane component of the E. coli secretion machinery. EMBO J. 10:1749-1757 (1991).

161. Carson, M.J., J. Barondess and J. Beckwith. The FtsQ protein of Escherichia coli: membrane topology, abundance, and cell division phenotypes due to overproduction and insertion. J. Bacteriol. 173: 2187-2195 (1991).

162. Barondess, J.J., Carson, M., Guzman-Verduzco, L.M., and Beckwith. J. Alkaline phosphatase fusions in the study of cell division genes. Res. Microbiol. 142: 295-299 (1991).

163. McGovern, K., Ehrmann, M., and Beckwith. J. Decoding signals for membrane protein assembly using alkaline phosphatase fusions. EMBO J. 10:2773-2782 (1991).

164. McGovern, K., and Beckwith, J. Membrane insertion of the MalF protein under conditions of reduced sec gene function. J. Biol. Chem. 266:20870-20876 (1991).

165. Derman, A.I., and Beckwith, J. Escherichia coli alkaline phosphatase fails to acquire disulfide bonds when retained in the cytoplasm. J. Bacteriol. 173:7719-7722 (1991).

166. Bardwell, J.C.A., McGovern, K., and Beckwith, J. Identification of a protein required for disulfide bond formation in vivo. Cell. 67:581-589 (1991).

167. Bassford, P., Beckwith, J., Ito, K., Kumamoto, C., Mizushima, S., Oliver, D., Randall, L., Silhavy, T., Tai, P.C., and Wickner, B. The primary exportpathway in E. coli. Cell 65:367-368 (1991).

168. Traxler, B., Lee, C., Boyd, D., and Beckwith, J. Dynamics of assembly of a cytoplasmic membrane protein in Escherichia coli. J. Biol. Chem. 267:5339-5345(1992).

169. Traxler, B., and Beckwith, J. Steps in the Assembly of a Cytoplasmic Membrane Protein- The MalF Component of the Maltose Transport Complex. in Membrane Biogenesis and Protein Targeting, eds. W. Neupert and R. Lill. (Series- New Comprehensive Biology), 49-61 (1992).

170. Guzman, L.-M., Barondess, J.J., and Beckwith, J. FtsL, an essential cytoplasmic membrane protein involved in cell division in E. coli. J. Bacteriol. 174:7716-7728 (1992).

171. Beckwith, J., and Silhavy, T. The Power of Bacterial Genetics: A Literature-Based Course. Cold Spring Harbor Laboratory Press, Cold Spring Harbor. 828 pages(1992).

172. Traxler, B., and Beckwith, J. The assembly of a hetero-oligomeric membrane protein complex. Proc. Natl. Acad. Sci. 89:10852-10856 (1992).

173. Johnson, K., Murphy, C.K., and Beckwith, J. Protein export in Escherichia coli. Current Opinion in Biotechnology 3:481-485 (1992).

174. Boyd, D., Traxler, B., and Beckwith, J. Analysis of the topology of a membrane protein using a minimum number of alkaline phosphatase fusions. J. Bacteriol.175:553-556 (1993).

175. Traxler, B., Boyd, D., and Beckwith, J. The topological analysis of integral cytoplasmic membrane proteins. J. Membr. Biol. 132:1-11 (1993).

176. Derman, A., Puziss, J., Bassford, P.J., Jr. and Beckwith, J. A signal sequence is not required for protein export in prlA mutants of Escherichia coli. EMBO J. 12:879-888 (1993).

177. Martin, J.L., Waksman, G., Bardwell, J.C.A., Beckwith, J., and Kuriyan, J. Crystallization of DsbA, an Escherichia coli protein required for disulfide bond formation in vivo. J. Mol. Biol. 230:1097-1100 (1993).

178. Pogliano, K.J., and Beckwith, J. The cs sec mutants of Escherichia coli reflect the cold sensitivity of protein export itself. Genetics 133:763-773 (1993).

179. Boyd, D., Traxler, B., Jander, G., Prinz, W., and Beckwith, J. Gene fusion approaches to membrane protein topology. in Molecular Biology and Function of Carrier Proteins, ed.s L. Reuss, J.M. Russell, Jr. and M. Jennings. Rockefeller Univ. Press., N.Y. pp. 23-37 (1993).

180. Bardwell, J.C.A., Lee, J.-O., Jander, G., Martin, N., Belin, D. and Beckwith, J. A pathway for disulfide bond formation in vivo. Proc. Natl. Acad. Sci. 90:1038-1042 (1993).

181. Derman, A., Prinz, W., Belin, D., and Beckwith, J. Mutants that allow disulfide bond formation in the cytoplasm of Escherichia coli. Science 262:1744-1746 (1993).

182. Bardwell, J.C.A., and Beckwith, J. The bonds that tie:catalyzed disulfide bond formation. Cell 74:769-771 (1993).

183. Pogliano, K.J., and Beckwith, J. Genetic and molecular characterization of the Escherichia coli secD operon and its products. J. Bacteriol. 176:804-814 (1994).

184. Beckwith, J. Early steps in secretion: conservation of mechanism across species. Forme. volume? 13-16 (1994).

185. Bardwell, J.C.A., Derman, A., Belin, D., Jander, G., Prinz, W., Martin, N., and Beckwith, J. Pathways of disulfide bond formation in proteins in vivo. in Molecular Biology of Phosphate in Microorganisms, eds. A.M. Torriani, E. Yagil and S. Silver. ASM Press, Washington. 270-275 (1994).

186. Murphy, C.K., and Beckwith, J. Residues essential for the function of SecE, a membrane component of the Escherichia coli secretion apparatus, are located in a conserved cytoplasmic region. Proc. Natl. Acad. Sci. USA. 91:2557-2561 (1994).

187. Prinz, W., and Beckwith, J. Gene fusion analysis of membrane protein topology: a direct comparison of alkaline phosphatase and â-lactamase fusions. J. Bacteriol.176:6410-6413 (1994).

188. Pogliano, J.A., and Beckwith, J. SecD and SecF facilitate protein export in Escherichia coli. EMBO J. 13: 554-561 (1994).

189. Jander, G., Martin, N., and Beckwith, J. Two cysteines in each periplasmic domain of the membrane protein DsbB are required for its function in protein disulfide bond formation. EMBO J. 13:5121-5128 (1994).

190. Barondess, J.J., and Beckwith, J. bor gene of phage , involved in serum resistance, encodes a widely conserved outer membrane lipoprotein. J. Bacteriol. 177:1247-1253 (1995).

191. Guzman, L.-M., Belin, D., Carson, M., and Beckwith, J. pBAD expression vectors containing the tightly regulated arabinose PBAD promoter. J. Bacteriol. 177:4121-4130 (1995).

192. Murphy, C.K., Stewart, E.J., and Beckwith, J. A double counter-selection system for the study of null alleles of essential genes in Escherichia coli. Gene. 155:1-7 and 163:, 165 (1995).

193. Derman, A.L., and Beckwith, J. Alkaline phosphatase localized to the Escherichia coli cytoplasm is slowly activated in cells whose growth has been suspended: a caution for gene fusion studies. J. Bacteriol. 177:3764-3770 (1995).

194. Murphy, C., Prinz, W., Pohlschroder, M., Derman, A., and Beckwith, J. Essential features of the pathway for protein translocation across the Escherichia coli cytoplasmic membrane. in Protein Kinesis. Cold Spring

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Harbor Symposium for Quantitative Biology. 60:277-283 (1995).

195. Guilhot, C., Jander, G., Martin, N., and Beckwith, J. Evidence that the pathway of disulfide bond formation in Escherichia coli involves interactions between the cysteines of DsbB and DsbA. Proc. Natl. Acad. Sci., USA, 92:9895-9899 (1995).

196. Economou, A., Pogliano, J., Beckwith, J., Oliver, D.B., and Wickner, W. SecA membrane cycling at SecYEG is driven by distinct ATP binding and hydrolysis events and is regulated by SecD and SecF. Cell 83:1171-1182 (1995).

197. Murphy, C.K., and Beckwith, J. Export of proteins to the cell envelope in Escherichia coli. In Escherichia coli and Salmonella typhimurium. Cellular and Molecular Biology. eds. F. Neidhart et al. ASM Press, Washington, pp. 967-978 (1996).

198. Jander, G., Cronan, J.E., and Beckwith, J. Biotinylation in vivo as a sensitive indicator of protein secretion and membrane protein insertion. J. Bacteriol. 178:3049-3058 (1996).

199. Beckwith, J. From physiology to DNA and back. Introduction to "Gene Regulation in Bacteria" eds. S. Lynch and E.C.C. Lin. Chapman and Hall, New York. pp.1-5 (1996).

200. Pohlschroder, M., Murphy, C., and Beckwith, J. In vivo analyses of interactions between SecE and SecY, core components of the Escherichia coli protein translocation machinery. J. Biol. Chem. 271:19908-19914 (1996).

201. Prinz, W.A., Spiess, C., Ehrmann, M., Schierle, C., and Beckwith, J. Targeting of signal sequenceless proteins for export in E. coli with altered protein translocase. EMBO J. 15:5209-5217 (1996).

202. Rietsch, A., Belin, D., Martin, N., and Beckwith, J. A pathway for protein disulfide bond isomerization in E. coli.Proc. Natl. Acad. Sci. 93:13048-13053 (1996).

203. Prinz, W.A., Åslund, F., Holmgren, A., and Beckwith, J. The role of the thioredoxin and glutaredoxin pathways in preventing disulfide bond formation in the cytoplasm. J. Biol. Chem. 272:15661-15667 (1997).

204. Pogliano, J., Pogliano, K., Weiss, D., Losick, R., and Beckwith, J. Inactivation of FtsI inhibits constriction of the FtsZ cytokinetic ring and delays the assembly of FtsZ rings at potential division sites. Proc. Natl. Acad. Sci. 94:559-564 (1997).

205. Weiss, D.S., Pogliano, K., Carson, M., Guzman, L.-M., Fraipont, C., Nguyen-Distèche,M., Losick, R., and Beckwith, J. Localization of the Escherichia coli cell division protein FtsI (PBP3) to the division site and cell pole. Mol. Microbiol.25:671-681 (1997).

206. Guzman, L.M., Weiss, D.S., and Beckwith, J. Domain swapping analysis of FtsI, FtsL and FtsQ: bitopic membrane proteins essential for cell divsion in Escherichia coli. J. Bacteriol. 179:5094-5103 (1997).

207. Pogliano, J., Lynch, A.S., Belin, D., Lin, E.C.C., and Beckwith, J. Regulation of Escherichia coli cell envelope proteins involved in protein folding and degradation by the Cpx two-component system. Genes and Development 11:1169-1182 (1997).

208. Pohlschröder, M., Prinz, W., Hartmann, E., and Beckwith, J. Protein translocation in the three domains of life: variations on a theme. Cell 91:563-566 (1997).

209. Rietsch, A., Bessette, P., Georgiou, G., and Beckwith, J. Reduction of the periplasmic disulfide bond isomerase, DsbC, occurs by passage of electrons from cytoplasmic thioredoxin. J. Bacteriol. 179:6601-6608 (1997).

210. Prinz, W.A., Boyd, D.H., Ehrmann, M., and Beckwith, J. The protein translocation apparatus contributes to determining the topology of an integral membrane protein in Escherichia coli. J. Biol. Chem. 273:8419-8424 (1998).

211. Boyd, D., Schierle, C., and Beckwith, J. How many membrane proteins are there? Protein Sci. 7:201-205 (1998).

212. Leeds, J.A., and Beckwith, J. Lambda repressor N-terminal DNA binding domain as an assay for protein transmembrane segment interactions in vivo. J. Mol. Biol.280:799-810 (1998).

213. Debarbieux, L., and Beckwith, J. The reductive enzyme thioredoxin 1 acts as an oxidant when it is exported to the Escherichia coli periplasm. Proc. Natl. Acad. Sci., USA 95:10751-10756 (1998).

214. Stewart, E.J., Åslund, F. and Beckwith, J. Disulfide bond formation in the Escherichia coli cytoplasm: an in vivo role reversal for the thioredoxins. EMBO J. 17:5543-5550 (1998).

215. Rietsch, A., and Beckwith, J. The genetics of disulfide bond

metabolism. Annu. Rev. Genet. 32:163-184 (1998).

216. Beckwith, J. Applicazioni della genetica. Nell'Enciclopedia Italiana X (Enciclopedia del Novecento, Supplemento II. 680-699 (1998).

217. Ghigo, J.-M., Weiss, D.S., Chen, J.C., Yarrow, J.C., and Beckwith, J. Localization of FtsL to the Escherichia coli septal ring. Mol. Microbiol. 31:725-738 (1999).

218. Weiss, D.S., Chen, J.C., Ghigo, J.-M., Boyd, D., and Beckwith, J. Localization of FtsI (PBP3) to the septal ring requires its membrane anchor, the Z-ring, FtsA, FtsQ and FtsL J. Bacteriol. 181:508-520 (1999).

219. Chen, J.C., Weiss, D.S., Ghigo, J.-M., and Beckwith, J. Septal localization of FtsQ, an essential cell division protein in Escherichia coli. J. Bacteriol. 181:521-530 (1999).

220. Åslund, F., and Beckwith, J. The thioredoxin superfamily:specificity, redundancy and gray area genomics. J. Bacteriol. 181:1375-1379 (1999).

222. Mössner, D., Huber-Wunderlich, M., Rietsch, A., Beckwith, J., Glockshuber, R., and Åslund, F. Importance of redox potential for the in vivo function of the cytoplasmic disulfide reductant thioredoxin from Escherichia coli. J. Biol. Chem.274:25254-25259 (1999).

223. Åslund, F., Zheng, M., Beckwith, J., and Storz, G. Regulation of the OxyRtranscription factor by hydrogen peroxide and the cellular thiol-disulfide status Proc. Natl. Acad. Sci., USA 96:6161-6165 (1999).

224. Åslund, F., and Beckwith, J. Bridge over troubled waters: sensing stress by disulfide bond formation. Cell 96:751-753 (1999).

225. Debarbieux, L., and Beckwith, J. Electron avenue: pathways of disulfide bond formation and isomerization. Cell. 99:117-119 (1999).

226. Stewart, E.J., Katzen, F., and Beckwith,J. Six conserved cysteines of the membrane protein DsbD are required for the transfer of electrons from the cytoplasm to the periplasm of E. coli. EMBO J. 18:5963-5971 (1999).

227. Bessette, P.H., Åslund, F., Beckwith, J. and Georgiou, G. Efficient folding of proteins with multiple disulfide bonds in the Escherichia coli cytoplasm. Proc. Natl. Acad. Sci., USA 96:13703-13708 (1999).

228. Ghigo, J.-M., and Beckwith, J. Cell division in Escherichia coli: the role

of FtsL domains in septal localization, function and oligomerization. J. Bacteriol. 182:116-129 (2000).

229. Boyd, D., Weiss, D.S., Chen, J.C., and Beckwith, J. Towards single copy gene expression systems making gene cloning physiologically relevant: lambda InCh, a simple E. coli plasmid/chromosome shuttle system. J. Bacteriol. 182:842-847 (2000)

230. Ritz,D., Patel, H., Doan, B., Zheng, M., Åslund, F., Storz, G., and J. Beckwith. Thioredoxin 2 is involved in the oxidative stress reponse in Escherichia coli. J. Biol. Chem. 275:2505-2512 (2000).

231. Debarbieux, L., and Beckwith, J. On the functional interchangeability, oxidant vs. reductant, of members of the thioredoxin superfamily. J. Bacteriol. 182:723-727 (2000).

232. Tian, H.-P., Boyd, D., and Beckwith, J. A mutant hunt for defects in membrane protein assembly yields mutations in the bacterial SRP and Sec systems. Proc. Natl. Acad. Sci. 97:4730-4735 (2000).

233. Beckwith, J. The all-purpose gene fusion. Methods in Enzymology 326:3-7 (2000).

234. Leeds, J.A., and Beckwith, J. A gene fusion methods for assaying protein transmembrane segment interactions in vivo. Methods in Enzymology 327:165-175 (2000).

235. Kadokura, H., Bader, M., Tian, H.-P., Bardwell, J.C.A., and Beckwith, J. Roles of a conserved arginine residue of DsbB in linking protein disulfide bond formation pathway to the respiratory chain of Escherichia coli. Proc. Natl. Acad. Sci. 97:10884-10889 (2000).

236. Katzen, F., and Beckwith, J. Transmembrane electron transfer by the membrane protein DsbD occurs via a disulfide bond cascade. Cell 103:769-779 (2000).

237. Beckwith, J., Curtiss III, R., Hatfield, W., Ingraham, J., Low, B., Magasanik, B., Neidhart, F., Reznikoff, W., Riley, M., and Schaechter, M. Escherichia Coli and Salmonella 2000:the view from here. Microbial Mol. Biol. Revs. 65:119-130 (2001).

238. Goldstone, D., Haebel, P.W., Katzen, F., Bader, M.W., Bardwell, J.C., Beckwith, J., and Metcalf, P. DsbC activation by the N-terminal domain of DsbD. Proc. Natl. Acad. Sci., U.S.A. 98:9551-9556 (2001).

239. Ritz, D., Lim, J., Reynolds, C.M., Poole, L.B., and Beckwith, J. A triplet

repeat expansion resulting in the addition of a single amino acid converts the E. coli peroxiredoxin AhpC into a disulfide reductase. Science. 294:158-160 (2001).

240. Leeds, J., Boyd, D., Huber, D.R., Sonoda, G.K., Luu, H.T., Engelman, D.M., and Beckwith, J. Genetic selection for and molecular dynamic modeling of a protein transmembrane domain multimerization motif from a random Escherichia coli genomic library. J. Mol. Biol. 313:181-195 (2001).

241. Chen, J.C., and Beckwith, J. FtsQ, FtsL, and FtsI require FtsK but not FtsN for co-localization with FtsZ during Escherichia coli cell division. Mol. Microbiol. 42:395-413 (2001).

242. Kadokura, H., and Beckwith, J. The expanding world of oxidative folding. Nature Cell Biology. 3:E247-E250 (2001).

243. Ritz, D., and Beckwith, J. Roles of thiol-redox pathways in bacteria. Annual Reviews of Microbiology 55:21-48 (2001).

244. Katzen, F., and Beckwith, J. Disulfide bond formation in the periplasm of Escherichia coli. Methods in Enzymology 348:54-66 (2002).

245. Ritz, D., and Beckwith, J. Redox state of cytoplasmic thioredoxin. Methods in Enzymology 347:360-370 (2002).

246. Tian, H., and Beckwith, J. A genetic screen yields mutation in genes encoding all known components of the Escherichia coli signal recognition particle pathway. J. Bacteriol. J. Bacteriol 184:111-118 (2002).

247. Jurado, P., Ritz, D., Beckwith, J., de Lorenzo, V., and Fernandez, F.A. Production of functional single-chain antibodies in the cytoplasm of Escherichia coli. J. Mol. Biol. 320:1-10 (2002).

248. Chen, J.C., Minev, M,, and Beckwith, J. Analysis of ftsQ mutant alleles in Escherichia coli: complementation, septal localization, and recruitment of downstream cell division proteins. J. Bacteriol. 184:695-705 (2002).

249. Haebel, P.W., Goldstone, D., Katzen, F., Beckwith, J. and Metcalf, P. The disulfide bond isomerase DsbC is activated by an Ig-fold electron transporter: crystal structure of the DsbC-DsbDá complex. EMBO J. 21:4774-4784 (2002).

250. Buddelmeijer, N., Judson, N., Boyd, D., Mekalanos, J.J., and Beckwith, J. YgbQ, a cell division protein in Escherichia coli and Vibrio cholerae, localizes in co-dependent fashion with FtsL to the division site. Proc. Natl. Acad. Sci. 99:6316-6321 (2002).

251. Kadokura, H., and Beckwith, J. Four cysteines of the membrane protein DsbB act in concert to oxidize its substrate DsbA. EMBO J. 21:2354-2363 (2002).

252. Katzen, F., Desmukh, M., Daldal, F., and Beckwith, J. Evolutionary domain fusion expanded the substrate specificity of the transmembrane electron transporter DsbD. EMBO J. 21:3960-3969 (2002).

253. Buddelmeijer, N., and Beckwith, J. Assembly of cell division proteins at the E. coli cell center. Curr. Opin. Microbiol. 5:553-557 (2002).

254. Toutain, C.M., Clarke, D.J., Leeds, J.A., Kuhn, J., Beckwith, J., Holland, I.B., and Jacq, A. The transmembrane domain of the DnaJ-like protein DjlA is a dimerisation domain. Mol. Gen. Genetics. 268:761-770 (2003).

255. Brandon, L.D., Goehring, N., Janakiraman, A., Yan, A.W., Wu, T., Beckwith, J., and Goldberg, M.B. IcsA, a polarly localized autotransporter with an atypical signal peptide, uses the Sec apparatus for secretion, although the Sec apparatus is circumferentially distributed. Molecular Microbiology 50:45-60 (2003)

256. Kadokura, H., Katzen, F., and Beckwith, J. Protein disulfide bond formation in prokaryotes. Annu. Rev. Biochem. 72:111-35 (2003).

257. Schierle, C.F., Berkmen, M., Huber, D., Kumamoto, C., Boyd, D., and Beckwith, J. The DsbA signal sequence directs efficient, co-translational export of passenger protein to the E. coli periplasm via the SRP pathway. J. Bacteriol. 185:5706-5713 (2003).

258. Katzen, F., and Beckwith, J. Role and location of the unusual redox-active cysteines in the hydrophobic domain of the transmembrane electron transporter DsbD. Proc. Natl. Acad. Sci., USA 100:10471-10476 (2003).

259. Ortenberg, R., and Beckwith, J. Functions of thiol-disulfide oxidoreductases in E. coli: redox myths, realities and practicalities. Antioxidants and Redox Signalling. 4:403-11 (2003).

260. Beckwith, J. Disulfide bond formation in the periplasm and cytoplasm of Escherichia coli. in C. Gitler and A. Danon. Cellular Implications of Redox Signalling. Imperial College Press, London pp. 213-232 (2003).

261. Buddelmeijer, N., and Beckwith, J. A complex of the Escherichia coli cell division proteins FtsL, FtsB and FtsQ forms independently of its localization to the septal region. Mol. Microbiol. 52:1315-1327 (2004).

262. Belin, D., Guzman, L.-G., Bost, S., Konakova, M., Silva, F., and Beckwith, J. Functional activity of eukaryotic signal sequences in Escherichia coli: the ovalbumin family of serine protease inhibitors. J. Mol. Biol. 335:437-453 (2004).

263, Kadokura, H., Tian, H., Zander, T., Bardwell, J.C.A. and Beckwith, J. Snapshots of DsbA in action: detection of proteins in the process of oxidative folding. Science 303:534-537 (2004).

264. Ortenberg, R., Gon, S., Porat, A., and Beckwith, J. Interactions of glutaredoxins, ribonucleotide reductase and components of the DNA replication system of E. coli. Proc. Natl. Acad. Sci., USA 101:7439-7444 (2004).

265. Porat, A., Cho, S.-H., and Beckwith, J. The unusual transmembrane transporter DsbD and its homologues: a bacterial family of disulfide reductases. Res. Microbiol. 155:617-622 (2004).

266. Goehring, N.W., Gueiros-Filho, F., and Beckwith, J. Premature targeting of a cell division protein to midcell allows dissection of divisome assembly in Escherichia coli. Genes Dev. 19:127-37 (2005).

267. Berkmen, M., Boyd, D., and Beckwith, J. The non-consecutive disulfide bond of Escherichia coli phytase (AppA) renders it dependent on the protein disulfide isomerase, DsbC. J. Biol. Chem. 280:11387-94 (2005).

268. Kadokura, H., Nichols, L., and Beckwith J. Mutational alterations of a key cis proline residue results in the trapping of enzymatic reaction intermediates of DsbA, a member of the thioredoxin superfamily. J. Bacteriol. 187:1519-1522 (2005).

269. Huber, D., Boyd, D., Xia, Y., Olma, M.H., Gerstein, M., and Beckwith, J. Use of thioredoxin as a reporter to identify a subset of Escherichia coli signal sequences that promote signal recognition particle-dependent translocation. J. Bacteriol. 187:2983-2991 (2005).

270. Or, E., Boyd, D., Gon, S., Beckwith, J., and Rapoport, T. The bacterial ATPase SecA functions as a monomer in protein translocation. J. Biol. Chem. 280:9097-9105 (2005).

271. Sevier, C.S., Kadokura, H., Tam, V.C., Beckwith, J., Fass, D., and Kaiser, C.A. The prokaryotic enzyme DsbB may share structural features with eukaryotic disulfide bond forming oxidoreductases. Protein Science 14:1630-1642 (2005).

272. Huber, D., Cha, M.-I., Debarbieux, L., Planson, A.-G., López, G., Tasayco, M.L., Chaffotte, A., Beckwith, J. A selection for mutants that interfere with folding of E. coli thioredoxin-1 in vivo. Proc. Natl. Acad. Sci., U.S.A 102: 18872-18877 (2005).

273. Goehring, N., and Beckwith, J. Diverse paths to midcell: assembly of the bacterial cell division machinery. Curr. Biol. 15:R514-R526 (2005).

274. Gon, S., Camara, J., Klungsøyr, H.K., Crooke, E., Skarstad K., and Beckwith J. Mutations in dnaA and dnaN reveal a novel regulatory mechanism that couples deoxyribonucleotide synthesis and DNA replication during the cell cycle in E. coli. EMBO J. 25:1137-47 (2006).

275. Segatori, L., Murphy, L., Arredondo, S., Kadokura, H., Gilbert, H., Beckwith, J., and Georgiou, G. Conserved role of the linker α-helix of the bacterial disulfide isomerase DsbC in the avoidance of misoxidation by DsbB. J. Biol. Chem. 281:4911-9 (2006).

276. Gon, S., Faulkner, M.J, and Beckwith, J. In vivo requirement for glutaredoxins and thioredoxins in the reduction of the ribonucleotide reductases of E. coli. Antioxidants and Redox Signalling 8:735-42 (2006).

277. Gon,S. and Beckwith, J. Ribonucleotide reductases : Influence of environment on synthesis and activity. Antioxidants and Redox Signalling 8:773-80 (2006).

278. Goehring, N., Gonzalez, M.D., and Beckwith, J. Premature targeting of cell division proteins to midcell reveals hierarchies of protein interactions involved in divisome assembly. Mol. Microbiol. 61:33-45 (2006).

279. Cho, S.-H., and Beckwith, J. Mutations of the membrane-bound disulfide reductase DsbD that block electron transfer steps from cytoplasm to periplasm in Escherichia coli. J. Bacteriol. 188:5066-76 (2006).

280. Huber, D., and Beckwith, J. Phage display extends its reach. Nat. Biotechnol. 7:793-4 (2006).

281. Berkmen, M., Boyd, D., and Beckwith, J. Disulfide bond formation in the periplasm. in The Periplasm, Ed. M. Ehrmann. ASM Press, Washington, D.C. pp 122-140 (2006).

282. Goehring, N.W., Petrovska, I., Boyd, D., and Beckwith, J. Mutants, suppressors, and wrinkled colonies: Mutant alleles of the cell division gene ftsQ point to functional domains in FtsQ and a role for domain 1C of FtsA in divisome assembly. J Bacteriol. 189:633-45 (2007). 283. Goehring, N.W., Robichon, C. and Beckwith, J. A role for the non-essential N-terminus of FtsN in divisome assembly. J. Bacteriol. 189:646-649 (2007).

284. Porat, A., Lillig, C.H., Johansson,C., Fernandes, A.P, Nilsson, L., Holmgren, A., and Beckwith, J. The reducing activity of glutaredoxin 3 towards cytoplasmic substrate proteins is restricted by methionine 43. Biochemistry 20: 3366-3377 (2007).

285. Li, W., Schulman, S., Boyd, D. Erlandson, K., Beckwith, J., and Rapoport, T.A. The plug domain of the SecY protein stabilizes the closed state of the translocation channel and maintains a membrane seal. Mol. Cell 26:511-521 (2007).

286. Cho, S.-H., Porat, A., Ye, J., and Beckwith, J. Redox-active cysteines of a membrane electron transporter DsbD show dual compartment accessibility. EMBO J. 8:3509-3520 (2007).

287. Ye. J., Cho, S.-H., Fuselier, J., Li, W., Beckwith, J. and Rapoport, T.A. Crystal structure of an unusual thioredoxin protein with a zinc finger domain. J. Biol. Chem. 282:34945-51 (2007).

288. Beckwith, J. What Lies Beyond Uranus? Preconceptions, Ignorance, Serendipity and Suppressors in the Search for Biology's Secrets. Genetics 176:733-740 (2007).

289. Scheffers, D.-J., Robichon, C., Haan, G. J., den Blauwen, T., Koningstein, G., van Bloois, E., Beckwith, J., and Luirink, K. Contribution of the FtsQ transmembrane segment to localization to the division site. J. Bacteriol. 189:7273-7280 (2007).

290. Kadokura, H., and Beckwith, J. Disulfide bond formation in bacteria. in Redox Biochemistry, ed. R. Banerjee. pp. 113-120. John Wiley and Sons, Hoboken. (2008).

291. Yamamoto, Y. Ritz, D., Planson, A-.G., Jonsson, T., Faulkner, M.J., Boyd, D., Beckwith, J. and Poole, L.B. Mutant AhpC peroxiredoxins suppress thiol-disulfide redox deficiencies and acquire deglutathionylating activity. Molecular Cell 29:36-45 (2008).

292. Robichon, C., Goehring, N., King, G. and Beckwith J Artificial septal targeting of Bacillus subtilis cell division proteins in Escherichia coli: an interspecies approach to the study of protein-protein interactions in multi-protein complexes. J. Bacteriol. 190:6048-6059 (2008).

293. Faulkner, M.J., Veeravalli, K., Gon, S., Georgiou, G., and Beckwith, J. Functional plasticity of a peroxidase allows evolution of diverse disulfide-reducing pathways. Proc. Natl. Acad. Sci. U.S.A. 105:6735-40 (2008).

294. Dutton, R.J., Boyd, D., Berkmen. M., and Beckwith, J. Bacterial species exhibit diversity in their mechanisms and capacity for protein disulfide bond formation. Proc. Natl. Acad. Sci., U.S.A. 105:11933-11938 (2008).

295. Zhou, Y., Cierpicki, T., Jimenez, T.H.F., Lukasik, S.M., Ellena, J.F., Cafiso, D.S., Kadokura, H., Beckwith, J., and Bushweller, J.H. NMR Solution Structure of the Integral Membrane Enzyme DsbB – Functional Insights into DsbB Catalyzed Disulfide Bond Formation. Mol. Cell 31:896-908 (2008).

296. Eser, M., Masip, L., Kadokura, H., Georgiou, G. and Beckwith J. Disulfide Bond Formation by Exported Glutaredoxin Indicates Glutathione’s Presence in the E. coli Periplasm. Proc. Natl. Acad. Sci. U.S.A. 106:1572-1577 (2009). PMCID 2635786 297. Gonzalez, M., and Beckwith, J. Divisome under construction:  Distinct domains of the small membrane protein FtsB are necessary for interaction with multiple cell division proteins. J. Bacteriol. 191:2815-25 (2009).

298. Kadokura, H., and Beckwith, J. Detecting folding intermediates of a protein as it passes through the bacterial translocon. Cell 138:1164-1173 (2009).

299. Perez-Jimenez, R., Li, J., Kosuri, P., Sanchez-Romero, I., Wiita, A.P., Rodriguez-Larrea, D., Chueca, A., Holmgren, A., Miranda-Vizuete, A., Becker, K., Cho, S.-H., Beckwith, J., Gelhaye, E., Jacquot, J.P., Gaucher, E., Sanchez-Ruiz, J.M., Berne, B., and Fernandez, J.M. Diversity of Chemical Mechanisms in Thioredoxin Catalysis. Nature Structural and Molecular Biology 16:890-6 (2009)

300. Cho, S.-H., and Beckwith, J. Two snapshots of electron transport across the membrane insights in to the structure and function of DsbD J. Biol. Chem. 284:11416-11424 (2009). PMCID: PMC2670147

301. Beckwith, J. Genetic Suppressors and Recovery of Repressed Biochemical Memory. J. Biol. Chem. 284:12585-92 (2009)

302. Chung, H.S., Yao, Z., Goehring, N.W., Kishony, R., Beckwith, J.R., and Kahne, D.E. Rapid {beta}-lactam-induced lysis requires successful assembly of the cell division machinery. Proc. Natl. Acad. Sci., U.S.A. 106:21872-21877 (2009).

303. Shouldice, S.R., Cho, S.-H., Boyd, D. Heras, B., Eser, M., Beckwith, J., Riggs, P., Martin, J.L., and Berkmen, M. In vivo oxidative protein folding can be facilitated by oxidation-reduction cycling. Mol. Microbiol. 75:13-28 (2010). 304. Dutton, R.J., Wayman, A., Wei, J.-R., Rubin, E.J., Beckwith, J., and Boyd, D. Inhibition of bacterial disulfide bond formation by the anti-coagulant warfarin. Proc. Natl. Acad. Sci.107:297-301 (2010).

305. Li, W., Schulman, S., Dutton, R.J., Boyd, D., Beckwith, J., and Rapoport, T.A. Structure of a bacterial homolog of vitamin K epoxide reductase. Nature 463:507-512 (2010).

306. Gonzalez, M.D., Akbay, E.A., Boyd, D., and Beckwith, J. Multiple interaction domains in FtsL, a protein component of the widely conserved FtsLBQ cell division complex. J. Bacteriol. 192:2757-2768 (2010).

307. Huber, D. Chaffotte, A., Eser, M., Planson, AG, and Beckwith, J.. Amino acid residues important for folding of thioredoxin are revealed only by studying of the physiologically relevant reduced form of the protein. Biochemistry 49:8922-8 (2010).

308. Kadokura, H., and Beckwith, J. Mechanisms of oxidative protein folding in the bacterial cell envelope. Antioxidants and Redox Signalling. 13:1231-1246 (2010).

309. Rowland, S.L., Wadsworth, K.D., Robson, S.A., Robichon, C., Beckwith, J. and King, G.F. Residues in the extracytoplasmic β domain of DivIB mediate

its interaction with the divisomal transpeptidase PBP 2B: evidence from artificial septal targeting and site-directed mutagenesis. J. Bacteriol. 192:6116-25 (2010).

310. Elgán, T.H., Planson, A.-G., Beckwith, J., Güntert, B., and Berndt, K. Determinants of Activity in Glutaredoxins: An in vitro evolved Grx1-like Variant of E. coli Grx3. Biochemical J. 432:487-495 (2010).

311. Veeravalli, K., Boyd, D., Iverson, B.L., Beckwith, J., and Georgiou, G. Evolution of de novo biosynthetic pathways for γ-glutamyl cysteine and unnatural amino-thiol analogues. Nat. Chem.Biol.7:101-5. (2011).

312. Wang, X., Dutton, R., Beckwith, J., and Boyd, D. Membrane topology and mutational analysis of Mycobacterium tuberculosis VKOR, a protein involved in disulfide bond formation and a homologue of human vitamin K epoxide reductase. Antioxidants and Redox Signalling. 14:1413-1420 (2011). 312.Beckwith, J. The Operon as Paradigm: Normal Science and the Beginning of Biological Complexity. J. Mol. Biol. 409: 7-13 (2011).

313.Feeney, M., Gon, S., Veeravalli, K., Faulkner, M.J., Georgiou, G. and Beckwith, J. Repurposing lipoic acid changes electron flow in two important metabolic pathways of Escherichia coli. Proc. Natl. Acad. Sci. 108:7991-7996 (2011).

314. Robichon, C., Karimova, G., Beckwith, J., and Ladant, D. Role of leucine zipper in the association of the Escherichia coli cell division proteins FtsL and FtsB, J. Bacteriol. 193:4988-4922 (2011).

315. Cho, S.-H., Parsonage, D., Dutton, R., Thurston, C., Poole, L., Collet, J.-F., and Beckwith, J.. A new family of membrane electron transporters and its substrates, including the first cell-envelope peroxiredoxin, reveal a broadened reductive capacity of the oxidative bacterial cell envelope. MBio 3:1-11 (2012).

316. Feeney, M., Ke, N., and Beckwith, J. Mutations at several loci cause increased expression of ribonucleotide reductase in Escherichia coli. J. Bacteriol. 194:1515-1522 (2012).

317. Chng, S.-S., Dutton, R.J., Denoncin, K., Vertommen, D., Collet, I.-F., Kadokura, H. and Beckwith, J. Overexpression of the rhodanese PspE, a single cysteine-containing protein, restores disulfide bond formation to an Escherichia coli strain lacking DsbA. Mol. Microbiol. 85:996-1006 (2012).

318. Chng, S.-S., Xue, M., Garner, R.A., Kadokura, H., Boyd, D., Beckwith, J. and Kahne, D. Disulfide Rearrangement Triggered by Translocon Assembly Controls Lipopolysaccharide Export. Science 337:1665-1668 (2012).

319. Beckwith, J., The Sec-dependent pathway, Res. Microbiol. (2013), http://dx.doi.org/10.1016/j.resmic.2013.03.007

320. Beckwith, J. 50 years Fused to Lac. Ann. Rev. Microbiol. 67:1-19 (2013).

321. Beckwith, J. Clever Mutant Isolation: Defining the Components of Cell BiologicalProcesses. In: Stanley Maloy and Kelly Hughes, editors. Brenner’s Encyclopedia ofGenetics 2nd edition, Vol 2. San Diego: Academic Press; 2013. p. 33–37.

322. Hatahet, F., Boyd, D., and Beckwith, J. Disulfide bond formation in prokaryotes: History, diversity and design. BBA- Proteins and Proteomics. 1844:1402–1414 (2014). PMC4048783 http://dx.doi.org/10.1016/j.bbapap.2014.02.014

323. Dwyer, R.S., Malinverni, J.C., Boyd, D., Beckwith, J., and Silhavy, T.J. Folding LacZ in the periplasm of Escherichia coli. J Bacteriol. 196:3343-50 (2014). doi: 10.1128/JB.01843-14. Epub 2014 Jul 7.

324. Beckwith, J. Mission possible: Getting to yes with François Jacob. Res. Microbiol. 165:348-350 (2014).

325. Landeta, C., Blazyk, J., Hatahet, F., Meehan, B., Eser, M., Myrick, A., Bronstein, M., Miami, S., Arnold, H., Rubin, E.J., Furie, B., Furie, B., Beckwith, J., Dutton, R. and Boyd, D.Compounds targeting disulfide bond forming enzyme DsbB of Gram-negative bacteria. Nat Chem Biol. 4:292-8. (2015). doi: 10.1038/nchembio.1752.

326. Williamson, J., Cho, S., Ye, J., Collet, J.F., Beckwith,J., and Chou, J. Structure and Multi-State Function of the Transmembrane Electron Transporter CcdA. Nature Structural and Molecular Biology 22: 809-814 (2015). doi: 10.1038/nsmb.3099.327. Hatahet, F., Blazyk, J.L., Martineau, E., Mandela, E., Zhao, Y., Campbell, R.E., Beckwith, J., and Boyd, D. Altered Escherichia coli membrane protein assembly machinery allows proper membrane assembly of eukaryotic protein vitamin K epoxide reductase. Proc. Natl. Acad. Sci. U. S. A.(2015) Nov 23. pii: 201521260.