Joan L. Slonczewski
Robert A. Oden, Jr. Professor of Biology
Department of Biology, Higley Hall
Kenyon College
202 North College Road
Gambier, Ohio 43022
Ultraphyte blog

Academic Positions
Administration and Service
Grant Awards
Research Publications
Science Fiction
Interdisciplinary Publicactions


Yale University,  Ph.D., Department of Molecular Biophysics and Biochemistry, 1982.
Dissertation:  "Regulation and Sensing of Bacterial pH in Escherichia coli." Advisor:  Robert M. Macnab.

Bryn Mawr College, A.B. in Biology, magna cum laude, 1977.


Kenyon College, Department of Biology.  Robert A. Oden, Jr. Professor, 2014-present; Professor, 2000-2014; Chair of Biology, 1993-1996; Associate Professor, 1991-2000; Assistant Professor, 1984-1991.
Research: pH stress response in E. coli, using GFP fluorimetry, H2 assays, microarrays, and proteomic 2-D gels.
Teaching: Microbiology, Molecular Biology, Microbial Ecology, and Virology.

U. Maryland at Baltimore, Department of Microbiology and Immunology, Visiting Associate Professor, 1998-1999.  Studied pH-regulated virulence factors in Proteus mirabilis and in Helicobacter pylori.

Princeton University, Molecular Biology Department, National Science Foundation Visiting Professorship for Women, 1990-1991.  Investigated the molecular biology of pH-regulated genes.   Created and taught a new course on microbiology.   Developed a mentoring program for women scientists.

University of Pennsylvania, Department of Biology, Postdoctoral Fellow, National Institutes of Health Research Service Award, 1982-1984.   Studied calcium flux in leukocyte chemotaxis.


Secretary to the Faculty, Executive Committee of Kenyon College, 2014-2016; 2009-2010; 2001-2002; 1995-1997.

Chair of the Faculty, 2006-2007.

American Society for Microbiology, Council Policy Committee, At-large District Member, 2007-2010; 2004-2007.

Member of Editorial Board, Applied and Environmental Microbiology, 2008-2010.

Director, Howard Hughes Medical Institute Undergraduate Biological Sciences Education Program Award, Science Division at Kenyon College, 2004-2008; 2000-2004; 1996-2000.  Leading the development of facilities and digital technology in five science departments, three pre-college enrichment programs, and summer science fellowships for minority and first-generation students.

Panelist, National Science Foundation, Prokaryotic Genetics Study Section, 2003-2006, 1990-1995.

Panelist, NASA Gravitation Ecology and Microbiology, 2002-2003.

Chair, Faculty Affairs Committee, 1999-2001.  Faculty Meeting, Kenyon College.

Chair, Biochemistry and Molecular Biology Program, 1999-2005.

Chair of Department of Biology, 1993-1996.  Implemented a new major program in Molecular Biology and Biochemistry.  Developed course program for non-majors, to complement our research-oriented program for biology majors.  Created a comprehensive pre-health advising program.

Chair, Facilities Subcommittee of the Campaign Planning Committee, 1996.   Developed campaign goal for a new science complex at Kenyon.

Director, Elementary School Science Month and Elementary School Scientists program.  Implemented hands-on science experiences for grades K-6 at fourteen elementary and middle schools in Mount Vernon City and Knox County, 1992-1994.

Chair, Division K. Microbial Physiology and Metabolism, American Society for Microbiology, 1996-1997.   Planned program for Annual Meeting.

Panelist, Howard Hughes Medical Institute Predoctoral Fellowship Program, Genetics & Molecular Biology, 1995-1997.


Faculty Service Award, Kenyon College, 2016.

Guest of Honor, International Conference on the Fantastic and the Arts 36, Orlando, March 2014.

John Campbell Memorial Award for Best Science Fiction Novel, The Highest Frontier, 2012.

Division I Lecturer, 109th General Meeting, American Society for Microbiology, Philadelphia, May 2009.

Robert Tomsich Award, for outstanding achievement in research in science, Kenyon College, 2001.

Vice-Chair, first bi-annual Gordon Conference on Microbial Stress Responses, Plymouth State College, 1994. Wrote and submitted the proposal, with Roberto Kolter. Led sessions or presented invited papers at subsequent meetings, 1996, 1998, 2000, 2002, 2014, 2016.

"Mover and Shaker" Award, Mount Vernon News, for the Elementary School Science Program in the Mount Vernon and Knox County elementary schools, 1993.

Silver Medalist, National Professor of the Year program, Council for the Advancement and Support of Education, Washington DC, 1989.

Vector Laboratories Young Investigator Travel Award, for outstanding research presented at the American Society for Microbiology Annual Meeting, 1987.

Robert Tomsich Award, for science research, Kenyon College, 1997.

John Campbell Memorial Award for Best Science Fiction Novel, A Door into Ocean, 1987.


National Science Foundation RUI, $524,000. Acid and base stress in Escherichia coli, 2016-2019.

National Science Foundation RUI, $496,500. Acid and base stress in Escherichia coli, 2013-2016.

National Science Foundation RUI, $453,000. Acid and base stress in Escherichia coli, 2011-2013.

National Institutes of Health ARRA supplement, $68,118. pH Regulation in Bacillus subtilis, 2009-2010.

National Science Foundation RUI, $400,000. Acid and base stress in Escherichia coli, 2007-2010.

National Institutes of Health, $200,310.
pH regulation in Bacillus subtilis, 2007-2009.

Howard Hughes Medical Institute, Undergraduate Biological Sciences Competition, $1,500,000. Student research and teaching, curriculum development and pre-college outreach, 2004-2008.

National Science Foundation RUI, $396,000. Acid and base stress in Escherichia coli, 2003-2006.

Howard Hughes Medical Institute, Undergraduate Biological Sciences Competition, $700,000. Student research and teaching, curriculum development and pre-college outreach, 2000-2004.

National Science Foundation RUI, $370,000.  Acid and base stress in Escherichia coli, 2000-2003.

Howard Hughes Medical Institute, Undergraduate Biological Sciences Competition, $1,500,000. Student research and teaching, curriculum development and pre-college outreach, 1996-2000.

National Science Foundation RUI, $324,000. Acid stress genes in Escherichia coli, 1996-2000.

National Science Foundation RUI, $285,000. Acid resistance and stationary phase in Escherichia coli and Shigella flexneri, 1993-1996.

Dwight D. Eisenhower Mathematics and Science Education Program, Ohio Board of Regents, $44,200. Elementary Experimental Science Month: Scientists in the Classroom, 1992-1993.

Dwight D. Eisenhower Mathematics and Science Education Program, Ohio Board of Regents, $41,621, for Elementary School Scientists, 1993-1994.

National Science Foundation, Visiting Professorship for Women, at Princeton University, $77,137, 1990-1991.

National Science Foundation RUI, $255,000, pH-regulated genes in E. coli, 1990-1993.

National Institutes of Health, direct costs $166,000. "Genes induced by pH perturbation in Escherichia coli," 1987-1990.

National Institutes of Health, Academic Research Enhancement Award, $71,869, The genetics of pH Homeostasis in Escherichia coli," 1985-1987.

National Science Foundation College Science Instrumentation Program, $25,000, Equipment for Undergraduate Instruction in the Techniques of Subcellular Analysis by Ultracentrifugation, 1985-1987.

PUBLICATIONS.  *Undergraduate researchers

J. L. Slonczewski and John W. Foster, 2017, 2014, 2011, 2009. Microbiology: An Evolving Science. Core microbiology textbook for undergraduate science majors. W. W. Norton & Co., New York.

Primary Articles:
Creamer, K. E.,* F. S Ditmars,* P. J. Basting, K. S. Kunka,* I. N. Hamdallah,* S. P. Bush,* Z. Scott,* A. He,* S. R. Penix,* A. S. Gonzales,* E. K. Eder,* D. Camperchioli,* A. Berndt,* M. W. Clark, K. A. Rouhier, and J. L. Slonczewski. 2017. Benzoate and salicylate tolerant strains lose antibiotic resistance during laboratory evolution of Escherichia coli K-12. Appl. Env. Micro. 83:

Clark, M. W., A. M. Yie,* E. K. Eder,* R. G. Dennis,* P. J. Basting, K, A. Martinez II, Brian D. Jones, J. L. Slonczewski. 2015. Periplasmic acid stress increases cell division asymmetry (polar aging) of Escherichia coli. PLoS ONE 10:e0144650.

Harden, M. M.,* A. He,* K. Creamer,* M. W. Clark, I. Hamdallah,* K. A. Martinez 2nd, R. L. Kresslein,* S. P. Bush,* J. L. Slonczewski. 2015. Editor Selection. Acid-adapted strains of Escherichia coli K-12 obtained by experimental evolution. Appl. Env. Micro. 81:1932–1941.

Riggins, D.P.,* M. J. Narvaez,* K. A. Martinez, M. M. Harden,* and J. L. Slonczewski. 2013. Escherichia coli K-12 survives anaerobic exposure at pH 2 without RpoS, Gad, or hydrogenases, but shows sensitivity to autoclaved broth products. PLoS ONE 8:e56796.

Martinez, K. A., R.D. Kitko, J. Patrick Mershon,* H. E. Adcox,* K. A. Malek,* M. B. Berkmen, and J. L. Slonczewski. 2012. Cytoplasmic pH response to acid stress in individual cells of Escherichia coli and Bacillus subtilis observed by fluorescence ratio imaging microscopy. Appl. Env. Micro. 78:3706-3714

Deininger, K. N. W.,* A. Horikawa, R. D. Kitko, R. Tatsumi, J. L. Rosner, M. Wachi, and J. L. Slonczewski. 2011. A requirement of TolC and MDR efflux pumps for acid adaptation and GadAB induction in Escherichia coli. PLoS ONE 6: e18960.

Kitko, R. D., J. C. Wilks,* G. M. Garduque,* and J. L. Slonczewski. 2010. Osmolytes enhance pH homeostasis of Escherichia coli. PLoS ONE 5: e10078.

Noguchi, K.,* D. P. Riggins,* K. C. Eldahan,* R. D. Kitko, and J. L. Slonczewski. 2010. Hydrogenase-3 contributes to anaerobic acid resistance of Escherichia coli. PLoS ONE 5: e10132.

Slonczewski, J. L., J. A. Coker, and S. DasSarma. 2010. Microbial growth with multiple stressors. Microbe 5:110-116. Cover feature.

Slonczewski, J. L. M. Fujisawa, M. Dopson, and T. A. Krulwich. 2009. Cytoplasmic pH measurement and homeostasis in bacteria and archaea. Adv. Micro. Physiol. 55:1-79. Request PDF.

Kitko, R. D., R. L. Cleeton,* E. I. Armentrout,* G. E. Lee,* K. Noguchi,* M. B. Berkmen, B. D. Jones, and J. L. Slonczewski. 2009. Cytoplasmic Acidification and the benzoate transcriptome in Bacillus subtilis. PLoS ONE 4:e8255.

Wilks, J. C.,* R. D. Kitko, S. H. Cleeton,* G. Lee,* C. S. Ugwu,* B. D. Jones, S. S. BonDurant, and J. L. Slonczewski. 2009. Acid and base stress and transcriptomic responses in Bacillus subtilis. Appl. Env. Microbiol.75: 981-990.

Kannan, G.,* J. C. Wilks,* D. M. Fitzgerald,* B. D. Jones, S. S. BonDurant, and J. L. Slonczewski. 2008. Rapid acid treatment of Escherichia coli: transcriptomic response and recovery. BMC Microbiology 8:37.

Wilks, J. C. * and J. L. Slonczewski. 2007. pH of the cytoplasm and periplasm of Escherichia coli: rapid measurement by GFP fluorimetry. J. Bacteriol. 189:5601–5607.

Hayes, E. T.,* J.C. Wilks,* E. Yohannes, D. P. Tate,* M. Radmacher, S. S. BonDurant, and J. L. Slonczewski. 2006. pH and anaerobiosis coregulate catabolism, hydrogenases, ion and multidrug transporters, and envelope composition in Escherichia coli K-12. BMC Microbiology 6:89. BMC Research Highlight for 2006-2007.

Novak, J. P, S.-Y. Kim, J. Xu, O. Modlich, D. J. Volsky, D. Honys, J. L. Slonczewski, D. A. Bell, Fred R. Blattner, E. Blumwald, M. Boerma, M.Cosio, Z. Gatalica, M. Hajduch, J. Hidalgo, R. R. McInnes, M.C. Miller III, M. Penkowa, M. S. Rolph, J. Sottosanto, R. St-Arnaud, M. J Szego, D. Twell, and C. Wang. 2006. Generalization of DNA microarray dispersion properties: microarray equivalent of t-distribution. Biology Direct 1:27.

Yohannes, E., *A. E. Thurber, *J. C. Wilks, *D. P. Tate, and J. L. Slonczewski. 2005. Polyamine stress at high pH in Escherichia coli K-12. BMC Microbiology 5:59.

Maurer, L. M.,* E. Yohannes, S. S. BonDurant, M. Radmacher, and J. L. Slonczewski. 2005. pH regulates genes for flagellar motility, catabolism, and oxidative stress in Escherichia coli K-12. J Bacteriol. 187:304-319.

Yohannes, E., D. M. Barnhart,* and J. L. Slonczewski. 2004. pH-Dependent Catabolic Protein Expression during Anaerobic Growth of Escherichia coli K-12. J. Bacteriol. 186:192-199. PDF

Stancik, L. M.,* D. M. Stancik,* B. Schmidt, D. M. Barnhart,* Y. N. Yoncheva,* and J. L. Slonczewski.  2002.  pH-dependent expression of periplasmic proteins and amino acid catabolism in Escherichia coli.  J. Bacteriol. 184:4246-4258. PDF

Slonczewski, J. L. and C. Kirkpatrick.  2002.  Proteomic analysis of pH-Dependent stress responses in Escherichia coli and Helicobacter pylori, using two-dimensional gel electrophoresis.  Methods in Enzymology, Bacterial Pathogenesis Part C, Academic Press.

Kirkpatrick, C., L. M. Maurer,* N. E. Oyelakin,* Y. Yontcheva,* R. Maurer, and J. L. Slonczewski.  2001.  Acetate and formate stress: opposite responses in the proteome of Escherichia coli.  J. Bacteriol. 183:6466-6477. PDF

Slonczewski, J. L.,  D. J. McGee, J. Phillips,* C. Kirkpatrick, and H. L. T. Mobley.   2000.  pH-Dependent protein profiles of Helicobacter pylori analyzed by two-dimensional gels.  Helicobacter 5:240-247.

Blankenhorn, D., Phillips, J.,* Slonczewski, J. L.  1999.  Acid- and base-induced proteins during aerobic and anaerobic growth of Escherichia coli revealed by two-dimensional gel electrophoresis. J. Bacteriol. 181: 2209-2216. PDF

Lambert, L.,* K. Abshire, D. Blankenhorn, and J. L. Slonczewski.  1997.  Global induction of proteins by benzoate at low pH in Escherichia coli.   J. Bacteriol. 179:7595-7599.

Hersh, B. M.,* F. T. Farooq,* D. N. Barstad,* D. L. Blankenhorn, and J. L. Slonczewski. 1996. A glutamate-dependent acid resistance gene in Escherichia coli. J. Bacteriol. 178:3978-3981.

Slonczewski, J. L. and J. W. Foster. 1996. Ch. 96. pH-regulated genes and survival at extreme pH. Escherichia coli and Salmonella typhimurium: Cellular and Molecular Biology, Frederick C. Neidhardt, Ed. in Chief, American Society for Microbiology Press, Washington D.C., pp. 1539-1549.

Lin, J., I. S. Lee, J. Frey,* J. L. Slonczewski, and J. W. Foster. 1995. Comparative analysis of extreme acid survival in Salmonella typhimurium, Shigella flexneri, and Escherichia coli. J. Bacteriol. 177:4097-4101.

Small, P., D. Blankenhorn, D. Welty, E. Zinser,* and J. L. Slonczewski. 1994. Acid and base resistance in Escherichia coli and Shigella flexneri: the role of rpoS and growth pH. J. Bacteriol. 176: 1729-1737.

Rosner, J. L., and Slonczewski, J. L. 1994. Dual regulation of inaA by the multiple antibiotic resistance (Mar) and superoxide (SoxRS) stress response systems of Escherichia coli. J. Bacteriol. 176:6262-6269

Lee, I. S., J. L. Slonczewski, and J. W. Foster. 1994. A low pH-inducible stationary phase acid tolerance response in S. typhimurium. J. Bacteriol. 176:1422-1426.

White, S., F. E. Tuttle,* D. Blankenhorn, D. C. Dosch, and J. L. Slonczewski. 1992. pH dependence and gene structure of inaA in Escherichia coli. J. Bacteriol. 174:1537-1543.

Watson, N., D. S. Dunyak, E. L. Rosey, J. L. Slonczewski, and E. R. Olson. 1992. Identification of elements involved in transcriptional regulation of the Escherichia coli cad operon by external pH. J. Bacteriol. 174:530-540.

Bingham, R. J.,* K. S. Hall, and J. L. Slonczewski. 1990. Alkaline induction of a novel gene locus alx-1 in Escherichia coli. J. Bacteriol. 172:2184-2186.

Aliabadi, Z., Y. K. Park, J. L. Slonczewski, and J. W. Foster. 1988. Novel regulatory loci controlling oxygen- and pH-regulated gene expression in Salmonella typhimurium. J. Bacteriol. 170:842-851.

Zigmond, S. H., J. L. Slonczewski, M. W. Wilde, and M. Carson.  1988.  Polymorphonuclear leukocyte locomotion is insensitive to lowered cytoplasmic calcium levels. Cell Motility and the Cytoskeleton 9:184-189.

Slonczewski, J. L., T. N. Gonzalez,* F. M. Bartholomew, and N. J. Holt.  1987. Mud-directed lacZ fusions regulated by acid pH in Escherichia coli. J. Bacteriol. 169:3001-3006.

Slonczewski, J. L., M. W. Wilde, and S. H. Zigmond.  1985. Phosphorylase a activity as an indicator of neutrophil activation by chemotactic peptides. J. Cell Biol. 101:1191-1197.

Slonczewski, J. L., R. M. Macnab, J. R. Alger, and A. Castle.  1982. Effects of pH and repellent tactic stimuli on protein methylation levels in Escherichia coli. J. Bacteriol. 152:384-399.

Slonczewski, J. L., B. P. Rosen, J. R. Alger, and R. M. Macnab.  1981.  pH homeostasis in Escherichia coli:  Measurement by P31 nuclear magnetic resonance of methylphosphonate and phosphate. Proc. Natl. Acad. Sci. USA 78:6271-6275.


The Highest Frontier. College out in space—Invaded by undocumented aliens. Tor/Macmillan, September, 2011.

Brain Plague.  Intelligent microbes invade human brains, offering limitless powers--at a price.  Tor Books, August, 2000; Science Fiction Book of the Month Club, Alternate Selection.

The Children Star.  A planet with biochemistry so alien that only children can be genetically engineered to survive there.  And what unique alien intelligence is watching in secret?  Analog serial, April, 1998; Tor Books, September, 1998.

Daughter of Elysium. In the far future, biologists engineer humans to live for thousands of years, then face a revolt by the machines that made it possible. Avon, 1993; Easton Press signed first edition, 1993; Avon pbk, 1994.

The Wall around Eden. Quaker teen-agers face the environmental consequences of nuclear war. William Morrow, 1989; Avon pbk, 1990; Italian translation, Editrice Nord, 1991.

A Door into Ocean. Women biologists genetically engineer fantastic creatures on a planet covered entirely by ocean.  Science Fiction Book of the Month Club Main Selection, 1986; Arbor House, 1986; Avon pbk, 1987; Italian translation, Editrice Nord, 1988.

Still Forms on Foxfield. Quakers colonize a planet with bizarre alien inhabitants. DelRey Books, 1980.


J. L. Slonczewski and R. Marusak. 2004. A response to BIO 2010: Transforming Undergraduate Education for Future Research Biologists, from the perspective of the Biochemistry and Molecular Biology Major Program at Kenyon College. Biochem. Mol. Biol. Ed. 32:151-155.

J. L. Slonczewski and M. Levy. 2003. “Chapter 12: Science Fiction and the Life Sciences,” in The Cambridge Companion to Science Fiction, edited by E. James and F. Mendlesohn, Cambridge University Press, Cambridge, UK.

“Joan Slonczewski: Stranger than fiction.,” 2003. Profile in Nature 426:501.

J. L. Slonczewski. 2002. “Stem cells and human cloning,” Analog Science Fiction and Science Fact, November, pp. 26-29.

J. L. Slonczewski. 2000. “Tuberculosis bacteria join UN,” Nature 405:1001.

J. L. Slonczewski, S. Lindow, M. Levy, C. Jacobsen, and K. Hellekson. 2000. “Approaching The Children Star,” conference report with questions for writing and discussion, Science Fiction Research Association Review, July/August pp. 17-27.

J. L. Slonczewski. 1994. “Science in science fiction: making it work,” The Writer, pp 14-17.

J. L. Slonczewski. 1986. “A tale of two handmaids,” review of A Handmaid’s Tale by Margaret Atwood and The Magnificent Spinster by May Sarton, The Kenyon Review, New Series 8:120-124.