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Dr. C.L. (Conrad) Woldringh

Faculty of Science
Swammerdam Institute for Life Sciences
Photographer: onbekend

Visiting address
  • Science Park 904
Postal address
  • Postbus 94232
    1090 GE Amsterdam
Contact details
  • Profile

    Swammerdam Institute for Life Sciences (SILS)

    Faculty of Science,  University of Amsterdam

    Science Park 904  (room C3.106) 

    The Netherlands


    (Note new room number)


    Map of Science Park, Amsterdam

    Research interests

    • DNA organization in vitro and in vivo and the process of nucleoid segregation in Escherichia coli.
    • The bacterial cell cycle: relationships between DNA segregation and cell division and between position, size and shape of the nucleoid and cell shape.

    Molecular Cytology

    Cellular structures are built-up and modified by proteins. Besides their specific functions in metabolic processes, proteins also interact with each other and with other macromolecules present in the crowded cell in a non-specific way through thermal motion. These so-called excluded-volume interactions are determined by physical rules and can lead to self-assemblies, phase separations and self-organization. I am interested in how these physico-chemical rules contribute to and modulate the functioning and activity of specific proteins (polymerases, topoisomerses, translocases) involved in segregation and shape determination.


    Laboratory protocol for the isolation of nucleoids from E. coli spheroplasts. See:

    The bacterial cell cycle

    Considering its scale, the small bacterium may serve as a model system for understanding physical and structural principles underlying the self-organization of macromolecules in micro-compartments such as the eukaryotic chromosome. For instance, bacterial DNA segregation should be compared with the first step of eukaryotic segregation, the dissolution or disentanglement of the chromatids, rather than with the second step, the mitotic condensation, proteolysis and microtubular transport of the chromatids.
    For a further discussion see Syllabus "The Bacterial Cell Cycle" . See for cytological aspects of the cell cycle Section 2.1.

  • Collaborations



    Nanne Nanninga. Swammerdam Institute for Life Sciences.
    Norbert Vischer. Swammerdam Institute for Life Sciences.
    Cell cycle simulation program.
    Arieh Zaritsky (since 1974). Department of Life Sciences, Ben-Gurion University of the Negev, Be'er-Sheva, Israel.
    Norman Grover. Experimental Medicine and Cancer Research, Hadassah Medical School, Jerusalem, Israel.
    Theo Odijk. Section Theory of Complex Fluids, Kluyver Institute for Biotechnology, Delft University of Technology. Lorentz Institute for Theoretical Physics, Leiden.
    Suckjoon Jun . University of California at San Diego, U.S.A.
    Sharareh Tavaddod PhD in 2011 at Department of Physics, Zanjan,Iran.

    Flemming Hansen.  Department of Systems Biology Technical University of Denmark Matematiktorvet, Lyngby, Denmark

  • Publications


    • Pelletier, J., K. Halvorsen, B.-Y. Ha, R. Paparcone, S.J. Sandler, C.L. Woldringh, W.P. Wong and S. Jun (2012) Physical manipulation of the Escherichia coli chromosome reveals its soft nature. Proc. Nat. Ac. Sci. (USA) 109(40): E2649-E2656 (doi: 10.1073/pnas.1208689109)
    • Wegner, A.S., Alexeeva, S., Odijk, T. and Woldringh, C.L. (2012) Characterization of  Escherichia coli nucleoids released by osmotic shock.  J. Struct. Biol. 178: 260-269
    • Woldringh, C.L. 2009/2010. Nucleoid structure and segregation, in Bacterial Chromatin (R.T. Dame and Ch.J.  Dorman, eds.), Springer Biomedical/Life Science, The Netherlands.
    • Woldringh, C.L. and N. Nanninga. 2006. Structural and physical aspects of bacterial chromosome segregation. J. Struct. Biol. 156: 273-283.
    • Zaritsky, A., C.L. Woldringh, M. Einav and S. Alexeeva. 2006. Thymine limitation and thymine starvation to study bacterial physiology and cytology. J. Bacteriol. 188: 1667-1679.
    • Cunha, S., C.L. Woldringh, Th. Odijk. 2005. Restricted diffusion within the isolated Escherichia coli nucleoid. J. Struct. Biol. 150: 226-232.
    • Elmore, S., Müller, M., Vischer, N.O.E., Odijk, Th., Woldringh, C.L. 2005. Single-particle tracking of oriC-GFP fluorescent spots during chromosome segregation in Escherichia coli. J. Struct. Biol. 151: 275-287.
    • Woldringh, C.L. 2005. Is Escherichia coli getting old? BioEssays 27: 1-5.
    • Norris, V., C.L. Woldringh and E, Mileykovskaya. 2004. A hypothesis to explain division site selection in Escherichia coli by combining nucleoid occlusion and Min. FEBS Lett. 561: 3-10.
    • Zaritsky, A. and C.L. Woldringh. 2003. Localizing cell division in spherical Escherichia coli by nucleoid occlusion. FEMS Microbiol. Lett. 226: 209-214.
    • Gons, H.J., J. Ebert, H.L. Hoogveld, L. van der Hove, R. Pelk, W. Takkenberg and C.L. Woldringh. 2002. Observations on cyanobacterial population collapse in eutrophic lake water. Antonie van Leeuwenhoek 81: 319-326.
    • Nanninga, N., M. Roos and C.L. Woldringh. 2002. Models on stickiness of replicated Escherichia coli oriC. Microbiology 148: 3327-3328.
    • Stuger, R., C.L. Woldringh, C.C. van der Weijden, N.O.E. Vischer, B.M. Bakker, R.J.M. van Spanning, J.L. Snoep and H. Westerhoff. 2002. DNA supercoiling by gyrase is linked to nucleoid compaction. Molecular Biology Reports 29: 79-82.
    • Woldringh, C.L. 2002. The role of co-transcriptional translation and protein translocation (transertion) in bacterial chromosome segregation. Mol. Microbiol. 45: 17-29.
    • Cunha, S., Th. Odijk, E. Sueleymanoglu, C.L. Woldringh. 2001. Isolation of the Escherichia coli nucleoid. Biochimie 83: 149-154.
    • Cunha, S., C.L. Woldringh, Th. Odijk. 2001.  Polymer-mediated compaction and internal dynamics of isolated Escherichia coli nucleoids. J. Struct. Biol. 136: 53-66.
    • Grover, N.B. and C.L. Woldringh. 2001. Dimensional regulation of cell-cycle events in Escherichia coli during steady-state growth. Microbiology 147: 171-181.
    • Pas, E., M. Einav, C.L. Woldringh and A. Zaritsky. 2001. Perpendicular planes of FtsZ arcs in spheroidal Escherichia coli cells. Biochimie 83: 121-124.
    • Roos, M., A.B.M. van Geel, M.E.G. Aarsman, J.T.M. Veuskens, C.L. Woldringh, N. Nanninga. 2001. The replicated ftsQAZ and minB chromosomal regions of Escherichia coli segregate on average in line with nucleoid movement. Mol. Microbiol. 39: 633-640.
    • Roos, M.,R. Lingeman, C. L. Woldringh, and N. Nanninga.2001. Experiments on the movement of DNA regions in Escherichia coli evaluated by computer simulation. Biochimie 83: 67-75.
    • Wery, M., C.L. Woldringh and J. Rouviere-Yaniv. 2001. HU-GFP and DAPI co-localize on the Escherichia coli nucleoid. Biochimie 83: 193-201.
    • Zaritsky, A., C.L. Woldringh, R.H. Pritchard and I. Fishov. 2000. Surviving Escherichia coli in good shape: the many faces of bacillary bacteria. In J. Seckbach (ed.), Journey to diverse microbial worlds, pp. 347-364. Kluwer Academic Publishers, The Netherlands.
    • Fishov, I., and C.L. Woldringh. 1999. Visualization of membrane domains in Escherichia coli. Mol. Microbiol. 32: 789-794.
    • Huls, P. G., N.O.E. Vischer and C.L. Woldringh. 1999. Delayed nucleoid segregation in Escherichia coli. Mol. Microbiol. 33: 959-970.
    • Koppes, L.J.H., C.L. Woldringh and N. Nanninga. 1999. Escherichia coli contains a DNA replication compartment in the cell center. Biochimie 81: 803-810.
    • Roos, M., A.B.M. van Geel, M.E.G. Aarsman, J.T.M. Veuskens, C.L. Woldringh, N. Nanninga. 1999. Cellular localization of oriC during the cell cycle of Escherichia coli as analyzed by fluorescent in situ hybridization. Biochimie 81: 797-802.
    • Zaritsky, A., Van Geel, A., Fishov, I., Pas, E., Einav, M. & Woldringh, C.L. (1999). Visualizing multiple constrictions in spheroidal Escherichia coli cells. Biochimie 81: 897-900
    • Vischer, N.O.E., P.G. Huls, R.I. Ghauharali, G.J. Brakenhoff, N. Nanninga and C.L. Woldringh. 1999. Image cytometric method for quantifying the relative amount of DNA in bacterial nucleoids using Escherichia coli. J. Microsc. 196: 61-68.
    • Woldringh, C.L., and R. Van Driel. 1999. The Eukaryotic Perspective: Similarities and distinctions between pro- and eukaryotes. p. 77-90. In R.L. Charlebois (ed.), Organization of the prokaryotic genome. Chapter 5. American Society for Microbiology, Washington, D.C.
    • Woldringh, C.L., and T. Odijk. 1999. Structure of DNA withinthe bacterial cell: physics and physiology.p. 171-187. In R.L. Charlebois (ed.), Organization of the prokaryotic genome. Chapter 10. American Society for Microbiology, Washington, D.C.
    • Zaritsky, A., C.L. Woldringh, I. Fishov, N.O.E. Vischer and M. Einav. 1999. Varying division planes of secondary constrictions in spheroidal Escherichia coli cells. Microbiology 145: 1015-1022.
    • Drlica, K. and C.L. Woldringh. 1998. Chromosomal organization: nucleoids, chromosomal folding and DNA topology. In "Bacterial genomes: Physical structure and analysis." (eds.: F.J. de Bruijn, J.R. Lupski and G.M. Weinstock). Chapman and Hall, New York. Pp. 12-23.
    • van Helvoort, J. M. L. M., P.G. Huls, N.O.E. Vischer and C.L. Woldringh. 1998. Fused nucleoids resegregate faster than cell elongation in Escherichia coli pbpB(Ts) filaments after release from chloramphenicol inhibition. Microbiol. 144, 1339-1317.
    • Newman, E.B., L.I. Budman, E.C. Chan, R.C. Greene, R.T. Lin, C.L. Woldringh and R. D'Ari. 1998. Lack of S-adenosylmethionine results in a cell division defect in Escherichia coli. J. Bacteriol. 180: 3614-3619.
    • Gomes de Mesquita, D.S., J. Shaw, J.A. Grimbergen, M.A. Buys, L. Dewi, C.L. Woldringh. 1997. Vacuole segregation in the Saccharomyces cerevisiae vac2-1 mutant: structural and biochemical quantification of the segregation defect and formation of new vacuoles. Yeast 13: 999-1008.
    • Sillje, H.H., E.G. ter Schure, A.J. Rommens, P.G. Huls, C.L. Woldringh, A.J. Verkleij, J. Boonstra and C.T. Verrips. 1997. Effects of different carbon fluxes on G1 phase duration, cyclin expression and reserve carbohydrate metabolism in Saccharomyces cerevisiae. J. Bacteriol. 179: 6560-6565.
    • Gomes de Mesquita, D.S., H.B. van den Hazel, J. Bouwman, C.L. Woldringh. 1996. Characterization of new vacuolar segregation mutants, isolated by screening for loss of proteinase B self-activation. Eur. J. Cell Biol. 71: 237-247.
    • van Helvoort, J. M. L. M., J. Kool and C. L. Woldringh. 1996. Chloramphenicol causes fusion of separated nucleoids in Escherichia coli K-12 cells and filaments. J. Bacteriol. 178: 4289-4293.
    • Wang, Y.X., H. Zhao, T.M. Harding, D.S. Gomes de Mesquita, C.L. Woldringh, D,J. Klionsky, A.L. Munn and L.S. Weisman. 1996. Multiple classes of yeast mutants are defective in vacuole partitioning yet target vacuole proteins correctly. Mol. Biol. Cell 7: 1375-1389.
    • Woldringh, C.L., K. Fluiter and P.G. Huls. 1995. Production of senescent cells of Saccharomyces cerevisiae by centrifugal elutriation. Yeast 11: 361-369.
    • Woldringh, C. L., P. R. Jensen, and H. V. Westerhoff. 1995. Structure and partitioning of bacterial DNA: determined by a balance of compaction and expansion forces? FEMS Micribiol. Lett. 131: 235-242.
    • Gasset, G., R. Tixador, B. Eche, L. Lapchine, N. Moatti, P. Toorop and C.L. Woldringh. 1994. Growth and division of Escherichia coli under microgravity conditions. Res. Microbiol. 145: 111-120.
    • van Helvoort, J. M. L. M. and C. L. Woldringh. 1994. Nucleoid partitioning in Escherichia coli during steady state growth and upon recovery from chloramphenicol treatment.Mol. Microbiol. 13: 577-583.
    • Koch, A.L. and C.L. Woldringh. 1994. The metabolic inertness of the pole wall of a Gram-negative rod. J. theor. Biol. 171: 415-425.
    • Tixador, R., G. Gasset, B. Eche, N. Moatti, L. Lapchine, C.L. Woldringh, P. Toorop, J.P. Moatti, F. Delmotte and G. Tap. 1994. Behavior of bacteria and antibiotics under space conditions. Aviat. Space Environ. Med. 65: 551-556.
    • Vischer, N.O.E., P.G. Huls and C.L. Woldringh. 1994. Object-image: an interactive image analysis program using structured point collection. Binary. 6: 160-166.
    • Woldringh, C.L. 1994. Significance of plasmolysis spaces as markers for periseptal annuli and adhesion sites. Molec. Microbiol.  14: 597-607.
    • Woldringh, C. L., A. Zaritsky, and N. B. Grover. 1994. Nucleoid partitioning and the division plane in Escherichia coli. J. Bacteriol. 176: 6030-6038.
    • Mulder, E.and C.L. Woldringh. 1993. Plasmolysis bayes in Escherichia coli: are they related to development and positioning of division sites? J. Bacteriol. 175: 2241-2247.
    • Woldringh, C. L., P.G. Huls, and N.O.E. Vischer. 1993. Volume growth of daughter and parent cells during the cell cycle of Saccharomyces cerevisiae a/alpha as determined by image cytometry. J.Bacteriol. 175:3174-3181.
    • Zaritsky, A., C.L. Woldringh, Ch.E. Helmstetter and N.B. Grover. 1993. Dimensional rearrangement of Escherichia coli B/r cells during a nutritional shift-down. J. Gen. Microbiol. 139: 2711-2714.
    • Goede, A.P., G.J. Brakenhoff, C.L. Woldringh, J.W. Aalders, J.P. Imhof, P. van Kralingen, W.A. Mels, P. Schreinemakers and A. Zegers. 1992. Confocal microscopy in microgravity research. Adv. Space Res. 12: 109-112.
    • Huls, P. G., N. Nanninga, E. A. van Spronsen, J. A. C. Valkenburg, N. O. E. Vischer, and C. L. Woldringh. 1992. A computer-aided measuring system for the characterization of yeast populations combining 2D-image analysis, electronic particle counter, and flow cytometry. Biotechnol. Bioeng. 39: 343-250.
    • Mulder, E., C.L. Woldringh, F. Tetart and J.-P. Bouche. 1992. New minC matations suggest different interactions of the same region of division inhibitor MinC with proteins specific for minD and dicB coinhibition pathways. J. Bacteriol. 174: 35-39.
    • van Bakel, M. N. Vischer, R. Tixador, G. Gasset and C.L. Woldringh. 1991. Influence of low-temperature storage and glucose starvation on growth recovery in Escherichia coli relA and relA+ strains. J. Biotech. 19: 159-172.
    • Gomes de Mesquita, D.S., R. ten Hoopen and C.L. Woldringh. 1991. Vacuolar segregation to the bud of Saccharomyces cerevisiae: an analysis of morphology and timing in the cell cycle. J. Gen. Microbiol. 137: 2447-2454.
    • Mulder, E., and C.L. Woldringh. 1991. Autoradiographic analysis of diaminopimelic acid incorporation in filamentous cells of Escherichia coli: repression of peptidoglycan synthesis around the nucleoid. J.Bacteriol. 173: 4751-4756.
    • Wientjes,F.B., C.L. Woldringh and N. Nanninga. 1991. Amount of peptidoglycan in cell walls of Gram-negative bacteria. J. Bacteriol. 173: 7684-7691.
    • Woldringh, C. L., E. Mulder, P. G. Huls, and N. Vischer. 1991. Toporegulation of bacterial division according to the nucleoid occlusion model. Res. Microbiol. 142: 309-320.
    • Mulder, E., M. El'Bouhali, E. Pas and C. L. Woldringh. 1990. The Escherichia coli minB mutation resembles gyrB in defective nucleoid segregation and decreased negative supercoiling of plasmids. Mol. Gen Genet. 221: 87-93.
    • Nanninga, N., F.B. Wientjes, B.L. de Jonge and C.L. Woldringh. 1990. Polar cap formation during cell division in Escherichia coli. Res. Microbiol. 141: 103-118.
    • Rutgers, C.A., P.J. Schaap, J. van 't Riet, C.L. Woldringh and H.A Raue. 1990. In vivo and in vitro analysis of structure-function relationships in ribosomal protein L25 from Saccharomyces cerevisiae. Biochim. Biophys. Acta 1050: 74-79.
    • Woldringh, C. L., E. Mulder, J. A. C. Valkenburg, F. B. Wientjes, A. Zaritsky, and N. Nanninga. 1990. Role of nucleoid in toporegulation of division. Res. Microbiol. 141: 39-49.
    • Mulder, E., and C.L. Woldringh. 1989. Actively replicating nucleoids influence the positioning of division sites in DNA-less cell forming filaments of Escherichia coli. J. Bacteriol. 171: 4303-4314.
    • Baldwin, W.W., M.J.-T.  Sheu, P.W. Bankston, and C.L. Woldringh. 1988. Changes in buoyant density and cell size of Escherichia coli in response to osmotic stress. J. Bacteriol. 170: 452-455.
    • Taschner, P.E.M., N. Ypenburg, B.G. Spratt and C.L. Woldringh. 1988. An amino acid substitution in penicillin-binding protein 3 creates pointed polar caps in Escherichia coli. J. Bacteriol. 170: 4828-4837.
    • Taschner, P.E., P.G. Huls, E. Pas and C.L. Woldringh. 1988. Division behavior and shape changes in isogenic ftsZ, ftsQ, ftsA, pbpB and ftsE cell division mutants of Escherichia coli during tempersture shift experiments. J, Bacteriol. 170: 1533-1540.
    • Woldringh, C. L., P. Huls, N. Nanninga, E. Pas, P. E. M. Taschner, and F. B. Wientjes. 1988. Autoradiographic analysis of peptidoglycan synthesis in shape and cell division mutants ofEscherichia coli MC4100. In: Antibiotic Inhibition of Bacterial Cell-Surface Assembly and function. P. Actor, L. Daneo-Moore, M. L. Higgins, M. R. J. Salton, and G. D. Shockman (eds.), pp. 66-78. American Society for Microbiology, Washington, D.C.
    • Buchnik, D. C.L. Woldringh and A. Zaritsky. 1987. Effects of temperature inactivation of penicillin-binding protein 2 on envelope growth in Escherichia coli. Ann. Inst. Pasteur/Microbiol. 138: 537-547.
    • Grover, N.B., C.L. Woldringh and L.J. Koppes. 1987. Elongation and surface extension of individual cells of Escherichia coli B/r: comparison of theoretical and experimental size distributions. J. theor. Biol. 129: 337-348.
    • Hiemstra, H., N. Nanninga, C.L. Woldringh, M. Inouye and B. Witholt. 1987. Distribution of newly synthesized lipoprotein over the outer membrane and the peptidoglycan sacculus of an Escherichia coli lac-lpp strain. J. Bacteriol. 169: 5434-5444.
    • Koppes, L.J., C.L. Woldringh and N.B. Grover. 1987. Predicted steady-state cell size distributions for various growth models. J. theor. Biol. 129: 325-335.
    • Taschner, P.E., J.G. Verest and C.L. Woldringh. 1987.  Genetic and morphological characterization of ftsB and nrdB mutants of Escherichia coli. J. Bacteriol. 169: 19-25.
    • Woldringh, C. L., P. Huls, E. Pas, G. J. Brakenhoff, and N. Nanninga. 1987. Topography of peptidoglycan synthesis during elongation and polar cap formation in a cell division mutant of Escherichia coli MC4100. J. Gen. Microbiol. 133: 575-586.
    • Nanninga, N. and C.L. Woldringh. 1985. Cell growth, genome duplication and cell division. In "Molecular cytology of Escherichia coli" (ed.: N. Nanninga). Academic press, London. pp. 259-319.
    • Valkenburg, J.A.C., C.L. Woldringh, G.J. Brakenhoff, H.T.M. van derVoort, and N. Nanninga. 1985. Confocal scanning light microscopy of the Escherichia coli nucleoid: comparison with phase-contrast and electron microscope images. J. Bacteriol. 161: 478-483.
    • Wientjes, F.B., E. Pas, P.E. Taschner and C.L. Woldringh. 1985. Kinetics of uptake and incorporation of meso-diaminopimelic acid in different Escherichia coli strains. J. Bacteriol. 164: 331-337.
    • Woldringh C.L. and N. Nanninga. 1985.  Structure of nucleoid and cytoplasm in the intact cell. In "Molecular cytology of Escherichia coli" (ed.: N. Nanninga). pp. 161-197. Academic Press, London, New York.
    • Woldringh, C.L., J.A.C. Valkenburg, E. Pas, P.E.M. Taschner, P. Huls and F.B. Wientjes. 1985. Physiological and geometrical conditions for cell division in Escherichia coli. Ann. Inst. Pasteur/Microbiol. 136A: 131-138.
    • Nanninga, N., G.J. Brakenhoff, M. Meijer and C.L. Woldringh. 1984. Bacterial anatomy in retrospect and prospect. Ant. v. Leeuwenhoek 50: 433-460.
    • Valkenburg, J.A.C., and C.L. Woldringh. 1984. Phase separation between nucleoid and cytoplasm in Escherichia coli as defined by immersive refractometry. J. Bacteriol. 160:1151-1157.
    • Kubitschek, H.E. and C.L. Woldringh. 1983. Cell elongation and division probability during the  Escherichia coli growth cycle. J. Bacteriol. 153: 1379-1387.
    • Mullin, D.A., C.L. Woldringh, J.M. Henson and J.R. Watson. 1983. Cloning of the Escherichia coli dnaZX region and identification of its products. Mol. Gen. Genet. 192: 73-79.
    • Binnerts, J.S., C.L. Woldringh and G.J. Brakenhoff. 1982. Visualization of the nucleoid in living bacteria on poly-lusine coated surfaces by the immersion technique. J. Microscopy 125: 359-363.
    • Grossman, N., E.Z. Ron and C.L. Woldringh. 1982. Changes in cell dimensions during amino acid starvation of Escherichia coli. J. Bacteriol. 152: 35-41.
    • Nanninga, N., C.L. Woldringh and L.J.H. Koppes. 1982. Growth and division of Escherichia coli. In Cell growth (ed.: C. Nicolini.) PlenumPubl.Corp. pp. 225-271.
    • Trueba, F.J., O.M. Neijssel and C.L. Woldringh. 1982. Generality of the growth kinetics of the average individual cell in different bacterial populations. J. Bacteriol. 150: 1048-1055.
    • Trueba, F.J., E.A. van Spronsen, J. Traas and C.L. Woldringh. 1982. Effects of temperature on the size and shape of Escherichia coli cells. Arch. Microbiol. 131: 235-240.
    • Zaritsky, A., C. L. Woldringh, N. B. Grover, J. Naaman, and R. F. Rosenberger. 1982. Growth and form in bacteria. Comments Molec. Cell. Biophys. 1: 237-260.
    • Nanninga, N. and C.L. Woldringh. 1981. The interpretation of chemically fixed and freeze-fractured bacterial nucleoplasm. Acta histochem., Suppl. Band XXIII, S. 39-53.
    • Woldringh, C.L., J.S. Binnerts and A. Mans. 1981. Variation in Escherichia coli buoyant density measured in Percoll gradients. J. Bacteriol. 148: 58-63.
    • Trueba, F. J., and C. L. Woldringh. 1980. Changes in cell diameter during the division cycle of Escherichia coli. J. Bacteriol. 142: 869-878.
    • Woldringh, C. L., N. B. Grover, R. F. Rosenberger, and A. Zaritsky. 1980. Dimensional rearrangement of rod-shaped bacteria following nutritional shift-up. II. Experiments with Escherichia coli B/r. J. Theor. Biol. 86: 441-454.
    • Zaritsky, A., C.L. Woldringh and D. Mirelman. 1979. Constant peptidoglycan density in the sacculus of Escherichia coli B/r growing at different rates. FEBS Lett. 98: 29-32.
    • Rosenberger, R.F., Grover, N.B., Zaritsky, A. & Woldringh, C.L. (1978). Control of microbial surface growth by density. Nature (Lond.) 271: 244-245
    • Koppes, L.H., C.L. Woldringh and N. Nanninga. 1978. Size variations and correlation of different cell cycle events in slow-growing Escherichia coli. J. Bacteriol. 134: 423-433.
    • Zaritsky. A., and C. L. Woldringh. 1978. Chromosome replication rate and cell shape in Escherichia coli: lack of coupling. J. Bacteriol. 135: 581-587.
    • Grover, N.B., C.L. Woldringh, A. Zaritsky, R.F. Rosenberger. 1977. Elongation of rod-shaped bacteria. J. theor. Biol. 67: 181-193.
    • Woldringh, C. L., M. A. de Jong, W. van den Berg, and L. Koppes. 1977. Morphological analysis of the division cycle of two Escherichia coli  substrains during slow growth. J. Bacteriol. 131: 270-279.
    • Meyer, M., M.A. de Jong, C.L. Woldringh and N. Nanninga. 1976. Factors affecting the release of folded chromosomes from Escherichia coli. Eur. J. Biochem. 63: 469-475.
    • Meyer, M., M.A. de Jong, C.L. Woldringh and N. Nanninga. 1976. Significance of folded chromosomes released from amino acid-starved Escherichia coli cells. Eur. J. Biochem. 65: 409-414.
    • Staugaard, P., F.M. van den Berg, C.L. Woldringh and N. Nanninga. 1976. Localization of ampicillin-sensitive sites in Escherichia coli by electron microscopy. J. Bacteriol. 127: 1376-1381.
    • Woldringh, C. L. 1976. Morphological analysis of nuclear separation and cell division during the life cycle of Escherichia coli. J. Bacteriol. 125: 248-257.
    • Woldringh C.L. and N. Nanninga. 1976 Organization of the nucleoplasm in Escherichia coli visualized by phase-contrast light microscopy, freeze fracturing, and thin sectioning. J. Bacteriol. 127: 1455-1464.
    • Woldringh, C.L. 1973. Effect of cations on the organization of the nucleoplasm in Escherichia coli prefixed with osmium tetroxide or glutaraldehyde. Cytobiologie 8: 97-111.
    • Woldringh, C.L. 1973. Effects of toluene and phenetyl alcohol on the ultrastructure of Escherichia coli. J. Bacteriol. 114: 1359-1361.
    • Woldringh, C. L. and W. van Iterson. 1972. Effects of treatment with sodium dodecyl sulfate on the ultrastructure of Escherichia coli. J. Bacteriol. 111: 801-813.
    • Woldringh, C.L. 1970. Lysis of the cell membrane of Escherichia coli K12 by ionic detergents. Biochim. Biophys. Acta. 224: 288-290.
  • Curriculum vitae

    Curriculum vitae

    I was born in 1940 in Batavia (Indonesia). After two years primary school in Switzerland, I finished school in Hilversum, studied biology at the University of Amsterdam (MSc. 1968) and got my PhD at the Laboratory for Electron Microscopy in 1974. Apart from a short term visit to Israel in 1974 and a sabbatical leave at the University of Texas at Austin in 1982, I remained associated with the University of Amsterdam, from where I had to retire in 2005.

    From Electron microscopy to cell cycle analysis

    • Although I started my research of the bacterium Escherichia coli at a time in whichapplication ofelectron microscope techniques (negative staining, thin sectioning, freeze fracturing) appeared promising, I became sceptical about the maintenance of bacterial ultrastructure during the necessary fixation and dehydration procedures. At the same time, my colleagues Nanne Nanninga (initiator) and Fred Brakenhoff (reinventor) started the development of the confocal scanning light microscope (CSLM).
    • With the development of new (fluorescent) microscope techniques (NSOM, SIM, SSIM, STORM, PALM) that allowed the study of living cells, protocols for transmission electron microscopy became out of use. However, e.m. techniques persist (like cryo-em) and knowledge of fixation and dehydration can still be useful. Therefore, see the link ("Cell preparation for (electron)microscopy") for an introduction to fixation and dehydration. See also the link ("Erfgoed electronenmicroscopie Amsterdam") for a description in Dutch of the history of electron microscopy in Amsterdam.
    • Inspired by the book of Maalø and Kjeldgaard, 1966, on "The control of macromolecular synthesis" and by the so-called Cooper-Helmstetter model (1968), I initiated the study of the E. coli cell cycle. This required the development of cytometric methods (agar filtration, image analysis) to study changes in structure and cell shape during the cell cycle (see collaborations with Arieh Zaritsky and Norman Grover).
    • To understand how the bacterial chromosome is segregated during the cell cycle, the structure of the DNA within the nucleoid had to be understood in more physical terms. In collaboration with Theo Odijk (since 1998) a model has been developed for the physical organization of the DNA based on isolated nucleoids (See link "Physical model nucleoid" from Wegner et al., 2012). Other groups (e.g. Suckjoon Jun) are now continuing the study ofthe physical properties of the bacterial nucleoid.

    A fluorescence microscope for Salatiga

    For my retirement in 2005 colleagues and friends contributed to "A microscope for Salatiga". We finally brought the microscope in October 2008 to the University of Salatiga. See link below for a blog-report of that journey.

  • Publications


    • Pelletier, J., Halvorsen, K., Ha, B-Y., Paparcone, R., Sandler, S. J., Woldringh, C. L., ... Jun, S. (2012). Physical manipulation of the Escherichia coli chromosome reveals its soft nature. Proceedings of the National Academy of Sciences of the United States of America, 109(40), E2649-E2656. [details]
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  • Ancillary activities
    No known ancillary activities