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Howard C. BergHerchel Smith Professor of PhysicsProfessor of Molecular and Cellular Biology Member of the Rowland Institute at Harvard
Department of Molecular and Cellular Biology
617-495-0924
Curriculum VitaeEducationCalifornia Institute of Technology, Pasadena, California, 1952-1956, B.S. in Chemistry (1956), undergraduate research with Jerome Vinograd. Carlsberg Laboratory, Copenhagen, 1956-1957. Fulbright Fellow with Kai Linderstrøm-Lang. Harvard Medical School, Boston, Massachusetts, 1957-1959, preclinical study as a National Scholar. Harvard University, Cambridge, Massachusetts, 1959-1964, NIH Predoctoral Fellow. M.A. in Physics (1960), Ph.D. in Chemical Physics (1964), thesis with Norman Ramsey, "Spin exchange and surface relaxation in the atomic hydrogen maser". HonorsFulbright Fellow, 1956-1957National Scholar, Harvard Medical School, 1957-1959 NIH Predoctoral Fellow, 1959-1963 Junior Fellow, Harvard Society of Fellows, 1963-1966 NSF Science Faculty Professional Development Awardee, 1978-1979 Biological Physics Prize of the American Physical Society (with E.M Purcell), 1984 National Academy of Sciences, 1984 American Academy of Arts and Sciences, 1985 Fellow, American Association for the Advancement of Science, 1985 Fellow, American Physical Society, 1990 Fellow, American Academy of Microbiology, 1992 Beams Lecturer, Department of Physics, University of Virginia, 1992 Ida Beam Distinguished Visiting Professor, Department of Microbiology, University of Iowa, 1994 Annual Lecturer, Danish Academy of Technical Sciences, 1998 Guggenheim Fellow, 2000-2001 Chan Memorial Lecturer, Department of Chemistry, Berkeley, 2001 American Philosophical Society, 2002 Carlson Lectureship, Department of Biophysics, Johns Hopkins, 2002 Luria Lecturer, Department of Biology, MIT, 2002 Visiting Fellow Commoner, Trinity College Cambridge, 2005 2007 U.S. Genomics Award for Outstanding Investigator in the Field of Single Molecule Biology, awarded by the Biophysical Society Doctor of Science h. c., Faculty of Biology and Chemistry, University of Osnabrück, 2007 Professional ExperienceJunior Fellow, Harvard Society of Fellows, 1963-1966. Guest of Alec Bangham, Institute of Animal Physiology, Babraham, England, summer 1966. Assistant Professor of Biology and Chairman of the Board of Tutors in Biochemical Sciences, Harvard, 1966-1969. Associate Professor of Biochemistry and Molecular Biology and Chairman of the Board of Tutors in Biochemical Sciences, Harvard, 1969-1970. Guest of Julius Adler, University of Wisconsin, Madison, Wisconsin, fall 1970. Associate Professor of Molecular, Cellular and Developmental Biology, University of Colorado, 1970-1974. Professor of Molecular, Cellular and Developmental Biology, University of Colorado, 1974-1979, Associate Chairman, 1974-1975, Chairman, 1976-1977. Sabbatical with Roger Stanier, Pasteur Institute, Paris, and Jorgen Henrichsen, State Serum Institute, Copenhagen, 1978-1979. Professor of Biology, California Institute of Technology, 1979-1986. Student, Cold Spring Harbor course in Advanced Bacterial Genetics, summer 1985. Professor of Molecular & Cellular Biology, Harvard University, 1986-present. Member, The Rowland Institute for Science, 1986-present. Professor of Physics, Harvard University, 1997-present. Lorentz Professor, University of Leiden, spring 2001. Herchel Smith Professor of Physics, Harvard University, 2003-present Teaching experienceDeveloped courses at Harvard on membrane structure and function, at Colorado on the biology of sensory phenomena, statistical processes in molecular biology, and elements of biophysics, and at Caltech on motile behavior of cells and microorganisms. Since returning to Harvard, have taught an introductory course on cell biology (with D. Branton), developed a Core course, “From DNA to Brain” (with J.E. Dowling), developed courses on statistical processes in cells and on introductory biophysics (the latter with J. Doyle, D. Nelson, and more recently, A. Samuel), and conducted graduate seminars on microbial development and behavior (once with R.M. Losick), motor molecules, and gliding motility. Professional societiesAmerican Academy of Arts and Sciences, American Academy for Microbiology, American Association for the Advancement of Science, American Physical Society (Chairperson, Division of Biological Physics, 1988-1989), American Society for Microbiology, Biophysical Society, National Academy of Sciences, American Philosophical Society. Committees and editorial boardsNIH Molecular Biology Study Section, 1974-1977; Biophysical Journal, 1978-1981; Cell Motility, 1980-1990; Modern Cell Biology, 1981-1989; Board on Physics and Astronomy, National Research Council, 1990-1993; American Institute of Physics: Biological Physics Series, 1994-, Advisory Committee on Physics Today, 1998-2003; Proceedings of the National Academy of Sciences, 2001-2002; Journal of Bacteriology, 2001-; NIH Prokaryotic Cell and Molecular Biology Study Section, 2005-. Current research interestsMotile behavior of bacteria: chemotaxis and flagellar rotation in Escherichia coli; flagellar rotation in a motile Streptococcus; swimming of a novel cyanobacterium; mechanisms of gliding motility. PublicationsBooks1. Berg, H.C. Random Walks in Biology. (Princeton, Princeton, 1983). Expanded edition, 1993. 2. Berg, H.C. E. coli in Motion. (Springer-Verlag, NY, 2004). ArticlesOn problems in atomic physics:On the separation of macromolecules according to mass: On the structure of cell membranes: On the motile behavior of microorganisms: On problems in atomic physics: 1. Berg, H.C. and Kleppner, D. Storage technique for atomic hydrogen. Rev. Sci. Instr. 33, 248-249 (1962). 2. Berg, H.C. Spin relaxation of atoms in molecular buffer gases. J. Chem. Phys. 43, 1851 (1965). 3. Berg, H.C. Amplitude and decay rate analysis of low level exponentially decaying radio-frequency signals. Rev. Sci. Instr. 36, 330-334 (1965). 4. Berg, H.C. Spin exchange and surface relaxation in the atomic hydrogen maser. Phys. Rev. 137, A1621-A1634 (1965). 5. Kleppner, D., Berg, H.C., Crampton, S.B., Ramsey, N.F., Vessot, R.F.C., Peters, H.E. and Vanier, J. Hydrogen maser principles and techniques. Phys. Rev. 138, A972-A983 (1965). 6. Crampton, S.B., Berg, H.C., Robinson, H.G. and Ramsey, N.F. Determination of the quadropole coupling constant in the N 14 atomic ground state. Phys. Rev. Lett. 24, 195-197 (1970). On the separation of macromolecules according to mass: 7. Berg, H.C. and Purcell, E.M. A method for separating according to mass a mixture of macromolecules or small particles suspended in a fluid, I. Theory. Proc. Natl. Acad. Sci., USA 58, 862-869 (1967). 8. Berg, H.C., Purcell, E.M. and Stewart, W.W. ..., II. Experiments in a gravitational field. ibid, pp. 1286-1291. 9. Berg, H.C. and Purcell, E.M. ..., III. Experiments in a centrifugal field. ibid, pp. 1821-1828. 10. Purcell, E.M. and Berg, H.C. Particle separator. U.S. Patent 3,523,610 (1970). 11. Barcilon, V. and Berg, H.C. Forced axial flow between rotating concentric cylinders. J. Fluid Mech. 47, 469-479 (1971). On the structure of cell membranes: 12. Berg, H.C., Diamond, J.M. and Marfey, P.S. Erythrocyte membrane: Chemical modification. Science 150, 64-67 (1965). 13. Berg, H.C. Membrane dipole potentials. Biophys. J. 8, 1051-1053 (1968). 14. Berg, H.C. Sulfanilic acid diazonium salt: A label for the outside of the human erythrocyte membrane. Biochim. Biophys. Acta 183, 65-78 (1969). 15. Bender, W.W., Garan, H. and Berg, H.C. Proteins of the human erythrocyte membrane as modified by pronase. J. Mol. Biol. 58, 783-797 (1971). 16. Whiteley, N.M. and Berg, H.C. Amidination of the outer and inner surfaces of the human erythrocyte membrane. J. Mol. Biol. 87, 541-561 (1974). 17. Berg, H.C. and Hirsh, D. Synthesis of diazotized ( 35 S)sulfanilic acid of high specific activity: A label for the outer surface of cell membranes. Anal. Biochem. 66, 629-631 (1975). 18. Wennogle, L.P. and Berg, H.C. Covalent attachment of polydeoxythymidylic acid to human erythrocytes. J. Mol. Biol. 123, 471-483 (1978). 19. Wennogle, L.P. and Berg, H.C. Isolation of cell surface proteins by hybridization. J. Mol. Biol. 124, 689-699 (1978). On the motile behavior of microorganisms: 1971 - 1975 20. Berg, H.C. How to track bacteria. Rev. Sci. Instr. 42, 868-871 (1971). 21. Futrelle, R.P. and Berg, H.C. Specification of gradients used for studies of chemotaxis. Nature 239, 517-518 (1972). 22. Berg, H.C. and Brown, D.A. Chemotaxis in Escherichia coli analysed by three-dimensional tracking. Nature 239, 500-504 (1972). 23. Berg, H.C. and Anderson, R.A. Bacteria swim by rotating their flagellar filaments. Nature 245, 380-384 (1973). 24. Brown, D.A. and Berg, H.C. Temporal stimulation of chemotaxis in Escherichia coli. Proc. Natl. Acad. Sci. USA 71, 1388-1392 (1974). 25. Berg, H.C. Dynamic properties of bacterial flagellar motors. Nature 249, 77-79 (1974). 26. Berg, H.C. and Brown, D.A. Chemotaxis in Escherichia coli analyzed by three-dimensional tracking. Addendum. Antibiotics and Chemotherapy 19, 55-78 (1974). 27. Berg, H.C. Flagellar control of bacterial motion. In Functional Linkage in Biomolecular Systems, ed. Schmitt, F.O., Schneider, D.M. and Crothers, D.M. (Raven Press, New York, 1975) pp. 290-296. 28. Berg, H.C. Flagellar rotation. Proceedings of the First Intersectional Congress of the IAMS, Vol. 1 (Science Council of Japan, Tokyo) 1975, pp. 665-673. 29. Berg, H.C. Bacterial behaviour. Nature 254, 389-392 (1975). 30. Berg, H.C. Chemotaxis in bacteria. Ann. Rev. Biophys. Bioeng. 4, 119-136 (1975). 31. Berg, H.C. How bacteria swim. Sci. Am. 233 (2), 36-44 (1975). 32. Berg, H.C. and Tedesco, P.M. Transient response to chemotactic stimuli in Escherichia coli. Proc. Natl. Acad. Sci., USA 72, 3235-3239 (1975). 33. Berg, H.C. Bacterial movement. In Swimming and Flying in Nature, Vol. 1, ed. Wu, T.Y.-T., Brokaw, C.J. and Brennen, C. (Plenum, New York, 1975) pp. 1-11. On the motile behavior of microorganisms: 1976 - 1980 34. Berg, H.C. How spirochetes may swim. J. Theor. Biol. 56, 269-273 (1976). 35. Berg, H.C. Does the flagellar rotary motor step? In Cold Spring Harbor Conferences on Cell Proliferation, Vol. 3: Cell Motility, ed. Goldman, R., Pollard, T. and Rosenbaum, J. (Cold Spring Harbor, 1976) pp. A47-A56. 36. Berg, H.C. Rotary engines in biology. In New Directions in Kinematics Research, ed. Roth, B. (Proceedings of NSF Workshop, Stanford, August 23, 1976) pp. 101-103. 37. Manson, M.D., Tedesco, P., Berg, H.C., Harold, F.M. and van der Drift, C. A protonmotive force drives bacterial flagella. Proc. Natl. Acad. Sci., USA 74, 3060-3064 (1977). 38. Berg, H.C. and Purcell, E.M. Physics of chemoreception. Biophys. J. 20, 193-219 (1977). 39. Berg, H.C., Bromley, D.B. and Charon, N.W. Leptospiral motility. Symp. Soc. Gen. Microbiol. 28, 285-294 (1978). 40. Berg, H.C. The tracking microscope. Adv. Opt. Elect. Microsc. 7, 1-15 (1978). 41. Berg, H.C. and Turner, L. Movement of microorganisms in viscous environments. Nature 278, 349-351 (1979). 42. Manson, M.D., Tedesco, P.M. and Berg, H.C. Energetics of flagellar rotation in bacteria. J. Mol. Biol. 138, 541-561 (1980). 43. Berg, H.C. Dynamics and energetics of the bacterial rotary motor. In Protein dynamics and energy transduction, ed. Ishiwata, S.-I. (Taniguchi Foundation, 1980) pp. 312-344. 44. Berg, H.C. Chemotaxis in Bacteria: A beginner's guide to the literature. Lect. Notes. Biomath. 38, 377-378 (1980). On the motile behavior of microorganisms: 1981 - 1985 45. Berg, H.C., Manson, M.D. and Conley, M.P. Dynamics and energetics of flagellar rotation in bacteria. Symp. Soc. Exp. Biol. 35, 1-31 (1982). 46. Segall, J.E., Manson, M.D. and Berg, H.C. Signal processing times in bacterial chemotaxis. Nature 296, 855-857 (1982). 47. Lapidus, I.R. and Berg, H.C. Gliding motility of Cytophaga sp. strain U67. J. Bacteriol. 151, 384-398 (1982). 48. Block, S.M., Segall, J.E. and Berg, H.C. Impulse responses in bacterial chemotaxis. Cell 31, 215-226 (1982). 49. Smyth, R.D. and Berg, H.C. Change in flagellar beat frequency of Chlamydomonas in response to light. Cell Motility Suppl. 1, 211-215 (1982). 50. Khan, S. and Berg, H.C. Isotope and thermal effects in chemiosmotic coupling to the flagellar motor of Streptococcus. Cell 32, 913-919 (1983). 51. Khan, S. and Berg, H.C. Isotope and thermal effects in chemiosmotic coupling to the membrane ATPase of Streptococcus. J. Biol. Chem. 258, 6709-6712 (1983). 52. Block, S.M., Segall, J.E. and Berg, H.C. Adaptation kinetics in bacterial chemotaxis. J. Bacteriol. 154, 312-323 (1983). 53. Berg, H.C. and Khan, S. A model for the flagellar rotary motor. In Mobility and Recognition in Cell Biology, ed. Sund, H. and Veeger, C. (deGruyter, Berlin, 1983) pp. 485-497. 54. Ishihara, A., Segall, J.E., Block, S.M. and Berg, H.C. Coordination of flagella on filamentous cells of Escherichia coli. J. Bacteriol. 155, 228-237 (1983). 55. Block, S.M. and Berg, H.C. Successive incorporation of force-generating units in the bacterial rotary motor. Nature 309, 470-472 (1984). 56. Conley, M.P. and Berg, H.C. Chemical modification of Streptococcus flagellar motors. J. Bacteriol. 158, 832-843 (1984). 57. Berg, H.C. and Block, S.M. A miniature flow cell designed for rapid exchange of media under high-power microscope objectives. J. Gen. Microbiol. 130, 2915-2920 (1984). 58. Berg, H.C. Bacterial flagellar rotation and its chemotactic control. In Information and Energy Transduction in Biological Membranes, ed. Bolis, C.L., Helmreich, E.J.M. and Passow, H. (Liss, New York, 1984) pp. 215-219. 59. Berg, H.C. Bovine-like rhodopsin in algae. Nature 311, 702 (1984). 60. Segall, J.E, Ishihara, A. and Berg, H.C. Chemotactic signaling in filamentous cells of Escherichia coli. J. Bacteriol. 161, 51-59 (1985). 61. Krikos, A., Conley, M.P., Boyd, A., Berg, H.C. and Simon, M.I. Chimeric chemosensory transducers in Escherichia coli. Proc. Natl. Acad. Sci., USA 82, 1326-1330 (1985). 62. Berg, H. C. Physics of bacterial chemotaxis. In Sensory Perception and Transduction in Aneural Organisms, ed. Colombetti, G., Lenci, F. and Song, P.S. (Plenum, New York, 1985) pp. 19-30. 63. Berg, H.C. Flagellar Motility. In Roche Seminars on Bacteria: Number 1 in a Series. Properties, Problems and Therapeutic Progress, ed. Moellering, R.C. (Hoffman-LaRoche, Nutley, N.J., 1985), pp. 1-23. 64. Khan, S. Meister, M. and Berg, H.C. Constraints on flagellar rotation. J. Mol. Biol. 184, 645-656 (1985). On the motile behavior of microorganisms: 1986 - 1990 65. Berg, H.C. Chemotaxis gene unveiled. Nature 321, 200-201 (1986). 66. Segall, J.E., Block, S.M. and Berg, H.C. Temporal comparisons in bacterial chemotaxis. Proc. Natl. Acad. Sci., USA 83, 8987-8991 (1986). 67. Lowe, G., Meister, M. and Berg, H.C. Rapid rotation of flagellar bundles in swimming bacteria. Nature 325, 637-640 (1987). 68. Shimada, K. and Berg, H.C. Response of the flagellar rotary motor to abrupt changes in extracellular pH. J. Mol. Biol. 193, 585-589 (1987). 69. Berg, H.C., Block, S.M., Conley, M.P., Nathan, A.R., Power, J.N. and Wolfe, A.J. Computerized video analysis of tethered bacteria. Rev. Sci. Instrum. 58, 418-423 (1987). 70. Wolfe, A.J., Conley, M.P., Kramer, T.J. and Berg, H.C. Reconstitution of signaling in bacterial chemotaxis. J. Bacteriol. 169, 1878-1885 (1987). 71. Meister, M., Lowe, G. and Berg, H.C. The proton flux through the bacterial flagellar motor. Cell 49, 643-650 (1987). 72. Meister, M. and Berg, H.C. The stall torque of the bacterial flagellar motor. Biophys. J. 52, 413-419 (1987). 73. Meyer, P.W., Matus, I.J. and Berg, H.C. Avoidance of Phycomyces in a controlled environment. Biophys. J. 51, 425-437 (1987). 74. Berg, H.C. A physicist looks at bacterial chemotaxis. Cold Spring Harbor Symp. Quant. Biol. 53, 1-9 (1988). 75. Wolfe, A.J., Conley, M.P. and Berg, H.C. Acetyladenylate plays a role in controlling the direction of flagellar rotation. Proc. Natl. Acad. Sci., USA 85, 6711-6715 (1988). 76. Blair, D.F. and Berg, H.C. Restoration of torque in defective flagellar motors. Science 242, 1678-1681 (1988). 77. Berg, H.C. Biological applications of scanning tunneling microscopy. Physics Today 41(1), S18-S19 (1989). 78. Block, S.M., Blair, D.F. and Berg, H.C. Compliance of bacterial flagella measured with optical tweezers. Nature 338, 514-517 (1989). 79. Meister, M., Caplan, S.R. and Berg, H.C. Dynamics of a tightly coupled mechanism for flagellar rotation. Biophys. J. 55, 905-914 (1989). 80. Conley, M.P., Wolfe, A.J., Blair, D.F. and Berg, H.C. Both CheA and CheW are required for reconstitution of chemotactic signaling in Escherichia coli. J. Bacteriol. 171, 5190-5193 (1989). 81. Wolfe, A.J. and Berg, H.C. Migration of bacteria in semisolid agar. Proc. Natl. Acad. Sci. USA 86, 6973-6977 (1989). 82. Blair, D.F. and Berg, H.C. The MotA protein of Escherichia coli is a proton-conducting component of the flagellar motor. Cell 60, 439-449 (1990). 83. Berg, H.C. Physical constraints on microbial behavior: How you act if you are very small. J. Chem. Ecol. 16, 119-120 (1990). 84. Berg, H.C. Bacterial microprocessing. Cold Spring Harbor Symp. Quant. Biol. 55, 539-545 (1990). 85. Berg, H.C. and Turner, L. Chemotaxis of bacteria in glass capillary arrays. Biophys J. 58, 919-930 (1990). 86. Schnitzer, M.J., Block, S.M., Berg, H.C. and Purcell, E.M. Strategies for chemotaxis. Symp. Soc. Gen. Microbiol. 46, 15-34 (1990). On the motile behavior of microorganisms: 1991- 1995 87. Block, S.M., Fahrner, K.A. and Berg, H.C. Visualization of bacterial flagella by video-enhanced light microscopy. J. Bacteriol. 173, 933-936 (1991). 88. Budrene, E.O. and Berg, H.C. Complex patterns formed by motile cells of Escherichia coli. Nature 349, 630-633 (1991). 89. Block, S.M., Blair, D.F. and Berg, H.C. Compliance of bacterial polyhooks measured with optical tweezers. Cytometry 12, 492-496 (1991). 90. Blair, D.F. and Berg, H.C. Mutants in the MotA protein of Escherichia coli reveal domains critical for proton conduction. J. Mol. Biol. 221, 1433-1442 (1991). 91. Blair, D.F., Kim, D.Y. and Berg, H.C. Mutant MotB proteins in Escherichia coli. J. Bacteriol. 173, 4049-4055 (1991). 92. Berg, H.C. Bacterial motility: Handedness and symmetry. Ciba Found. Symp. 162, 58-72 (1991). 93. Budrene, E.O. and Berg, H.C. Pattern formation by bacteria. Current Biology 1, 83 (1991). 94. Stolz, B. and Berg, H.C. Evidence for interactions between MotA and MotB, torque generating elements of the flagellar motor of Eschericia coli. J. Bacteriol. 173, 7033-7037 (1991). 95. Berg, H.C. and Turner, L. Selection of motile nonchemotactic mutants of Escherichia coli by field-flow fractionation. Proc. Natl. Acad. Sci., USA 88, 8145-8148 (1991). 96. Berg, H.C. Studies of motile bacteria. In Physics News in 1991, ed. Schewe, P.F. (American Institute of Physics, New York, 1991). 97. Berg, H.C. Response of Escherichia coli to novel gradients. In Sensory Transduction, Proc. 45th Symp. Soc. Gen. Physiol., Chpt. 13, pp. 220-223 (Rockefeller Univ. Press, 1992). 98. Dailey, F.E. and Berg, H.C. Mutants in disulfide bond formation that disrupt flagellar assembly. Proc. Natl. Acad. Sci., USA 90, 1043-1047 (1993). 99. Dailey, F.E. and Berg, H.C. Change in direction of flagellar rotation in Escherichia coli mediated by acetate kinase. J. Bacteriol. 175, 3236-3239 (1993). 100. Hazelbauer, G.L., Berg, H.C. and Matsumura, P. Bacterial motility and signal transduction. Cell 73, 15-22 (1993). 101. Berg, H.C. and Turner, L. Torque generated by the flagellar motor of Escherichia coli. Biophys. J. 65, 2201-2216 (1993). 102. Frauenfelder, H. and Berg, H.C. Physics and Biology. Physics Today 47 (2), 20-21 (1994). 103. Fahrner, K.A., Block, S.M., Krishnaswamy, S., Parkinson, J.S. and Berg, H.C. A mutant hook-associated protein (HAP3) facilitates torsionally-induced transformations of the flagellar filament of Escherichia coli. J. Mol. Biol. 238, 173-186 (1994). 104. Conley, M.P., Berg, H.C., Tawa, P., Stewart, R.C., Ellefson, D.D. and Wolfe, A.J. pH dependence of CheA autophosphorylation in Escherichia coli. J. Bacteriol. 176, 3870-3877 (1994). 105. Berg, H.C. and Turner, L. Cells of Escherichia coli swim either end forward. Proc. Natl. Acad. Sci., USA 92, 477-479 (1995). 106. Berg, H.C. Torque generation by the flagellar rotary motor. Biophys. J. 68, 163s-167s (1995). 107. Samuel, A.D.T. and Berg, H.C. Fluctuation analysis of rotational speeds of the bacterial flagellar motor. Proc. Natl. Acad. Sci., USA 92, 3502-3506 (1995). 108. Woodward, D.E., Tyson, R., Myerscough, M.R., Murray, J.D., Budrene, E.O. and Berg, H.C. Spatio-temporal patterns generated by Salmonella typhimurium. Biophys. J. 68, 2181-2189 (1995). 109. Fung, D.C. and Berg, H.C. Powering the flagellar motor of Escherichia coli with an external voltage source. Nature 375, 809-812 (1995). 110. Frymier, P.D., Ford, R.M., Berg, H.C. and Cummings, P.T. Three-dimensional tracking of motile bacteria near a solid planar surface. Proc. Natl. Acad. Sci. USA 92, 6195-6199 (1995). 111. Berry, R.M., Turner, L. and Berg, H.C. Mechanical limits of flagellar motors probed by electrorotation. Biophys, J. 69, 280-286 (1995). 112. Budrene, E.O. and Berg, H.C. Dynamics of formation of symmetric patterns by chemotactic bacteria. Nature 376, 49-53 (1995). 113. Pitta, T.P. and Berg, H.C. Self-electrophoresis is not the mechanism for motility in swimming cyanobacteria. J. Bacteriol. 177, 5701-5703 (1995). On the motile behavior of microorganisms: 1996 - 114. Marykwas, D.L., Schmidt, S.A., and Berg, H.C. Interacting components of the flagellar motor of Escherichia coli revealed by the two-hybrid system in yeast. J. Mol. Biol. 256, 564-576 (1996). 115. Marykwas, D.L., and Berg, H.C. A mutational analysis of the interaction between FliG and FliM, two components of the flagellar motor of Escherichia coli. J. Bacteriol. 178, 1289-1294 (1996). 116. Berg, H.C. Touring machines. Curr. Biol. 6, 624 (1996). 117. Samuel, A.D.T. & Berg, H.C. Torque generating units of the bacterial flagellar motor step independently. Biophys. J. 71, 918-923 (1996). 118. Ehlers, K.M., Samuel, A.D.T. , Berg, H.C. & Montgomery, R. Do cyanobacteria swim using traveling surface waves? Proc. Natl. Acad. Sci. USA 93, 8340-8343 (1996). 119. Berg, H.C. Symmetries in bacterial motility. Proc. Natl. Acad. Sci. USA 93, 14225-14228 (1996). 120. Turner, L., Caplan, S.R. & Berg, H.C. Temperature induced switching of the bacterial flagellar motor. Biophys. J. 71, 2227-2233 (1996). 121. Berry, R.M. & Berg, H.C. Torque generated by the bacterial flagellar motor close to stall. Biophys. J. 71, 3501-3510 (1996). 122. Pitta, T.P., Sherwood, E.E., Kobel, A.M. & Berg, H.C. Calcium is required for swimming by the nonflagellated cyanobacterim Synechococcus strain WH8113. J. Bacteriol. 179, 2524-2528 (1997). 123. Samuel, A.D.T. & Berg, H.C. Statistical kinetics of the bacterial flagellar motor. Phys. Rev. E 55, 7801-7804 (1997). 124. Berry, R.M. & Berg, H.C. Absence of a barrier to backwards rotation of the bacterial flagellar motor demonstrated with optical tweezers. Proc. Natl. Acad. Sci. USA 94, 14433-14437 (1997). 125. Scharf, B.E., Fahrner, K.A., Turner, L. & Berg, H.C. Control of direction of flagellar rotation in bacterial chemotaxis. Proc. Natl. Acad. Sci. USA 95, 201-206 (1998). 126. Berg, H.C. Comments on 'The use of flash photolysis for ... analysis of bacterial chemotactic behaviour ...' Mol Microbiol 25, 295-302 (1997). Mol. Microbiol. 27, 507-508 (1998). 127. Scharf, B.E., Fahrner, K.A. and Berg, H.C. CheZ has no effect on flagellar motors activated by CheY13DK106YW. J. Bacteriol. 180, 5123-5128 (1998). 128. Berg, H.C. Keeping up with the F1-ATPase. Nature 394, 324-325 (1998). 129. Berry, R.M. and Berg, H.C. Torque generated by the flagellar motor of Escherichia coli while driven backward. Biophys. J. 76, 580-587 (1999). 130. Turner, L., Samuel, A.D.T., Stern, A.S., and Berg, H.C. Temperature dependence of switching of the bacterial flagellar motor by the protein CheY13DK106YW. Biophys. J. 77, 597-603 (1999). 131. Samuel, A.D.T., Pitta, T.P., Ryu, W.S., Danese, P.N., Leung, E.C.W., and Berg, H.C. Flagellar determinants of bacterial sensitivity to c-phage. Proc. Natl. Acad. Sci. USA 96, 9863-9866 (1999). 132. Berg, H.C. Motile behavior of bacteria. Physics Today 53 (1), 24-29 (2000).. 133. Berg, H.C. Constraints on models for the flagellar rotary motor. Phil. Trans. Roy. Soc. B 355, 491-501 (2000). 134. Ryu, W.S., Berry, R.M. and Berg, H.C. Torque-generating units of the flagellar motor of Escherichia coli have a high duty ratio. Nature 403, 444-447 (2000). 135. Chen, X. and Berg, H.C. Torque-speed relationship of the flagellar rotary motor. Biophys J. 78, 1036-1041 (2000). 136. Turner, L., Ryu, W.S. and Berg, H.C. Real-time imaging of fluorescent flagellar filaments. J. Bacteriol. 182, 2793-2801 (2000). 137. Chen, X. and Berg, H.C. Solvent-isotope and pH effects on flagellar rotation in Escherichia coli. Biophys. J. 78, 2280-2284 (2000). 138. Sourjik, V., and Berg, H.C. Location of components of the chemotaxis machinery of Escherichia coli using fluorescent-protein fusions. Molec. Microbiol. 37, 740-751 (2000). 139. Skerker, J.M. and Berg, H.C. “Direct observation of extension and retraction of type IV pili.” Proc Natl Acad Sci U S A., 98, 6901-6904 (2001). 140. Miyata, M., Ryu, W.S. and Berg, H.C. Force and velocity of Mycoplasma mobile gliding. J. Bacteriol. 184, 1827-1831 (2002). 141. Sourjik, V., and Berg, H.C. Receptor sensitivity in bacterial chemotaxis. Proc. Natl. Acad. Sci. USA 99, 123-127 (2002). 142. Berg, H.C. How Spiroplasma might swim. J. Bacteriol 184, 2063-2064 (2002). 143. Sourjik, V. and Berg, H.C. Binding of the Escherichia coli response regulator CheY to its target measured in vivo by fluorescence resonance energy transfer. Proc. Natl. Acad. Sci. USA 99, 12669-12674 (2002). 144. Berg, H.C. The rotary motor of bacterial flagella. Annu. Rev. Biochem. 72, 19-54 (2003). 145. Berg, H.C. The bacterial rotary motor. In The Enzymes, 3rd Ed., Vol. 23 (Academic, 2003) pp. 143-202. 146. Fahrner, K.A., Ryu, W.S., and Berg, H.C. Bacterial flagellar switching under load. Nature 423, 938 (2003) 147. Gabel, C.V., and Berg, H.C. The speed of the flagellar rotary motor of Escherichia coli varies linearly with protonmotive force. Proc. Natl. Acad. Sci. USA 100, 8748-8751 (2003). 148. Jaffe, J.D., Berg, H.C., and Church, G.M. Proteogenomic mapping as a complementary method to perform genome annotation. Proteomics 4, 59-77 (2004) 149. Berg H.C., and Sourjik, V., “Functional interactions between receptors in bacterial chemotaxis.” Nature 428, 437 - 441 (2004) 150. Darnton, N., Turner, L., Breuer, K., and Berg, H.C., “Moving fluid with bacterial carpets,” Biophys J 86, 1863-1870 (2004) 151. Jaffe, J.D., Miyata, M., and Berg, H.C., “Energetics of gliding motility in Mycoplasma mobile.” Journal of Bacteriology 186, 4254-4261 (2004) 152. Jaffe J.D., Stange-Thomann N., Smith C., DeCaprio D., Fisher S., Butler J., Calvo S., Elkins T., FitzGerald M.G., Hafez N., Kodira C.D., Major J., Wang S., Wilkinson J., Nicol R., Nusbaum C., Birren B., Berg H.C., Church G.M., “The complete genome and proteome of Mycoplasma mobile.” Genome Res., 14, 1447-1461 (2004). 153. Liberman, L., Berg, H.C. and Sourjik, V. , “Effect of Chemoreceptor Modification on Assembly and Activity of the Receptor-Kinase Complex in Escherichia coli”, Journal of Bacteriology, 186, 6643-6646 (2004) 154. Vaknin, A. and Berg, H.C. “Single-cell FRET imaging of phosphatase activity in the Escherichia coli chemotaxis system.” Proc. Natl. Acad. Sci. USA, 101, 17072-17077 (2004). 155. DiLuzio, W.R., Turner, L., Mayer, M., Garstecki, P., Weibel, D.B., Berg, H.C., and Whitesides, G.M. “Escherichia coli swim on the right-hand side.”, Nature, 435, 1271-1274 (2005). 156. Bates, D., Epstein, J., Boyle, E., Fahrner, K., Berg, H., and Kleckner, N. “The Escherichia coli baby cell column: a novel cell synchronization method provides new insight into the bacterial cell cycle.”, Molec. Microbiol., 57, 380-391 (2005). 157. Berg, H.C. Q&A Howard Berg. Curr. Biol., 15, R189-R190 (2005). 158. Berg, H.C. “Swarming motility: it better be wet”, Curr. Biol., 15, R599-R560 (2005). 159. Vaknin, A. and Berg, H.C. “Osmotic stress mechanically perturbs chemoreceptors in Escherichia coli&148;, Proc. Natl. Acad. Sci. USA 103, 592-596 (2005). 160. Shimizu, T.S., Delalez, N., Pichler, K. and Berg, H.C. “Monitoring bacterial chemotaxis by using bioluminescence resonance energy transfer: absence of feedback from the flagellar motors&148;, Proc. Natl. Acad. Sci. USA 103, 2093-2097 (2005). 161. Berg, H.C. “Marvels of bacterial behavior.&148; Proc. Am. Philos. Soc., 150, 428-442 (2006). 162. Vaknin, A. and Berg, H.C. “Physical responses of bacterial chemoreceptors&148;, J. Mol. Biol. 366, 1416-1423 (2007). 163. Darnton, N.C. and Berg, H.C. “Force-extension measurements on bacterial flagella: triggering polymorphic transformations.&148; Biophys. J. 92, 2230-2236 (2007). 164. Berg, H.C. “Navigation on a micron scale.&148; Lect. Notes Phys. 711, 1-13 (2007). 165. Darnton, N.C., Turner, L., Rojevsky, S. and Berg, H.C. “On torque and tumbling in swimming Escherichia coli,&148; J. Bacteriol. 189, 1756-1764 (2007). 166. Murphy, G.E., Matson, E.G., Leadbetter, J.R., Berg, H.C. and Jensen, G.J. “Ultrastructure of Treponema primitia by electron cryotomography.&148; Mol. Microbiol., in press. 167. Shimizu, T.S., Schulmeister, S., Berg, H.C. and Sourjik, V. “Sensitivity as a function of receptor modification in bacterial chemotaxis,&148; J. Mol. Biol., submitted.
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