Preiss, L., D.B. Hicks, S. Suzuki, T. Meier, and T.A. Krulwich. 2015. Alkaliphilic bacteria with impact on industrial applications, concepts of early life forms, and bioenergetics of ATP synthesis. Front. Bioeng. Biotechnol. 3:75. Doi: 10.3389/bioe.s015.00075. PMCID:PMC4453477.

DeCaen, P.G., Y. Takahashi, T.A. Krulwich, M. Ito, and D.E. Clapham. 2014. Ionic selectivity and thermal adaptations within the voltage-gated sodium channel family of alkaliphilic Bacillus. Elife doi: 10.7554/eLife.04387. PMID:PMC4225499.

Attie, O., A. Jayaprakash, H. Shah, I.T. Paulsen, M. Morino, Y. Takahashi, I. Narummi, R. Sachidanandam, K. Satoh, M. Ito, and T.A. Krulwich. 2014. Draft genome sequence of Bacillus alcalophilus AV1934, a classic alkaliphile isolated from human feces in 1934. Genome Announc. 13:2. E01175-14, doi: 10.1128/genomeA.01175-14.

Preiss, L., J.D. Langer, D.B. Hicks, J. Liu, Ö. Yildiz, T.A. Krulwich and T. Meier. 2014. The structure of the c-ring ion binding site of the ATP synthase from Bacillus pseudofirmus OF4 is adapted to the alkaliphilic lifestyle. Molec. Microbiol. 92:973-984. PMCID:PMC4064006

Liu, J., S. Ryabichko, M. Bogdanov, O.J. Fackelmayer, W. Dowhan and T.A. Krulwich. 2014. Cardiolipin is dispensable for oxidative phosphorylation and non-fermentative growth of alkaliphilic Bacillus pseudofirmus OF4. J. Biol. Chem. 289:2960-2971. PMC3908427

Morino, M., T. Suzuki, M. Ito and T.A. Krulwich. 2014. Purification and functional reconstitution of a multi-subunit Mrp type Na+/H+ antiporter. J. Bacteriol. 196:28-35 PMC3911140

Price-Whelan, A., C.K. Poon, M.A. Benson, C.M. Roux, P.M. Dunman, V.J. Torres, and T.A. Krulwich. 2013. Transcriptional profiling of Staphylococcus aureus during growth in 2 M NaCl leads to clarification of physiological roles for Kdp and Ktr K+ uptake systems. mBio, vol 4 4e00407-13 PMC3747578

Krulwich, T. A. and M. Ito. 2013. Prokaryotic alkaliphiles. In: The Prokaryotes, Vol. 1, 4th ed. E. Rosenberg, E.F. DeLong, F. Thompson, S. Lory, E. Stackebrandt (eds) Springer New York, invited review.

Preiss, L., A.L. Klyszejko, D.B. Hicks, J. Liu, O. Fackelmayer, T.A. Krulwich, and T. Meier. 2013. The c-ring stoichiometry of ATP synthase is adapted to cell physiological requirements of alkaliphilic Bacillus pseudofirmus OF4. Proc. Natl. Acad. Sci. USA. PMC3651500

Liu, J., D.B. Hicks, and T.A. Krulwich. 2013. Roles of AtpI and two YidC-type proteins from alkaliphilic Bacillus pseudofirmus OF4 in ATP synthase assembly and non-fermentative growth. J. Bacteriol. PMC3553844.

Krulwich, T.A., G. Sachs, and E. Padan. 2011. Molecular aspects of bacterial pH sensing and homeostasis. Nature Microbiol. Rev. 9: 330-343. PMC3247762

Janto, B., A. Ahmed, M. Ito, J. Liu, D.B. Hicks, S. Pagni, O.J. Fackelmayer, T.A. Smith, J. Earl, L.D.H. Elbourne, K. Hassan, I.T. Paulsen, A.-B. Kolstø, N.J. Tourasse, G.D. Ehrlich, R. Boissy, D.M. Ivey, G. Li, Y. Xue, Y. Ma, F.Z. Hu and T.A. Krulwich. 2011. The genome of alkaliphilic Bacillus pseudofirmus OF4 reveals adaptations that support the ability to grow in an external pH range from 7.5 to 11.4. Environ. Microbiol. 13: 3289-3309. PMC3228905

Liu, J., O. Fackelmayer, D. B. Hicks, L. Preiss, T. Meier, E. Sobie and T. A. Krulwich. 2011. Mutations in a helix-1 motif of the ATP synthase c-subunit of Bacillus pseudofirmus OF4 cause functional deficits and changes in stability and mobility of the c-rotor on SDS-PAGE. Biochemistry 50: 5496-5506. PMC3115392

Krulwich, T. A., J. Liu, M. Morino, M. Fujisawa, M. Ito, and D. B. Hicks. 2010. Adaptive mechanisms of extreme alkaliphiles. In: Extremophiles Handbook. Horikoshi, K., Antranikian, G., Bull, A., Robb, F., Stetter, K. (eds), Springer, Heidelberg.

Fujisawa, M., O. Fackelmayer, J. Liu, D. B. Hicks and T. A. Krulwich. 2010. The ATP Synthase a-subunit of extreme alkaliphiles is a distinct variant. J. Biol. Chem. 285: 32105-32115. PMC2952212

Morino, M., S. Natsui, T. Ono, T. H. Swartz, T. A. Krulwich and M. Ito. 2010. Single site mutations in the hetero-oligomeric Mrp antiporter from alkaliphilic Bacillus pseudofirmus OF4. J. Biol. Chem. 285: 30942-30950. PMC2945585

Hoffmann, J., L. Sokolova, L. Preiss, D.B. Hicks, T.A. Krulwich, N. Morgner, T. Meier, I. Wittig, H. Schägger and B. Brutschy. 2010. ATP synthases: cellular nanomotors characterized by LILBID mass spectrometry. Phys. Chem. Chem. Phys. 12: 13375-13382. PMC2955850

Preiss, L., Ö. Yildiz, D.B. Hicks, T.A. Krulwich and T. Meier. 2010. A new type of proton coordination in an F1F0-ATP synthase rotor ring. PLoS Biol. 8:e1000443. PMC2914638

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

Liu, J., M. Fujisawa, D. B. Hicks and T. A. Krulwich. 2009. Characterization of the functionally critical AXAXAXA and PXXEXXP motifs of the ATP synthase c-subunit from an alkaliphilic Bacillus. J. Biol. Chem. 284: 8714–8725.

Fujinami, S, N. Terahara, T. A. Krulwich and M. Ito. 2009. Motility and chemotaxis in alkaliphilic Bacillus species. Future Microbiol. 4: 1137-1149.

Terahara, N., T. A. Krulwich and M. Ito. 2009. Mutations alter the sodium versus proton use of a Bacillus clausii flagellar motor and confer dual ion use on Bacillus subtilis motors. Proc. Natl. Acad. Sci. USA 105:14359-14364.

Dzioba-Winogrodzki, J., O. Winogrodzki, T. A. Krulwich, M. A. Boin, C. C. Hase and P. Dibrov. 2009. The Vibrio cholerae Mrp system: cation/proton antiport properties and enhancement of bile salt-resistance in a heterologous host. J. Mol. Microbiol. Biotech 16: 176-186.

Morino, M., S. Natsui, T.H. Swartz, T.A. Krulwich and M. Ito. 2008. Single gene deletions of mrpA to mrpG and mrpE point mutations affect activity of the Mrp Na+/H+ antiporter of alkaliphilc Bacillus and formation of hetero-oligomeric Mrp complexes. J. Bacteriol. 190: 4162-4172.

 Krulwich, T. A., D. B. Hicks, T. Swartz and M. Ito. 2007. Bioenergetic adaptations that support alkaliphily. In: Physiology and Biochemistry of Extremophiles (C. Gerday and N. Glansdorff, eds.), 311-329, ASM Press, Washington, DC

Fujinami, S., T. Sato, J.S. Trimmer, B.W. Spiller, D.E. Clapham, T.A. Krulwich, I. Kawagishi and M. Ito. 2007. The voltage-gated Na+ channel NaVBP co-localizes with methyl-accepting chemotaxis proteins at cell poles of alkaliphilic Bacillus pseudofirmus OF4. Microbiology 153: 4027-4038].

Fujisawa, M., M. Ito and T. A. Krulwich. 2007. Three 2-component transporters with channel-like properties have monovalent cation/proton antiport activity. Proc. Natl. Acad. Sci. USA 104: 13289-13294.

Wei, Y., J. Liu, Y. Ma and T. A. Krulwich. 2007. Three putative cation/proton antiporters from the soda lake alkaliphile Alkalimonas amylolytica N10 complement an alkali-sensitive Escherichia coli mutant. Microbiology 153: 2168-2179.

Swartz, T. H., M. Ito, T. Ohira, S. Natsui, D. B. Hicks and T. A. Krulwich. 2007. Catalytic properties of Staphylococcus aureus and Bacillus members of the secondary Cation-Proton Antiporter-3 family are revealed by an optimized assay in an Escherichia coli host. J. Bacteriol. 189: 3081-3090.

Liu, X., X. Gong, D. B. Hicks, T. A. Krulwich, L. Yu and C.-A. Yu. 2007. Interaction between the cytochrome caa3 and F1F0-ATP synthase of alkaliphilic Bacillus pseudofirmus OF4 is demonstrated by saturation transfer electron paramagnetic resonance and differential scanning calorimetry assays. Biochemistry 46: 306-313.

Wei, Y., G. Deikus, B. Powers, V. Shelden, T. A. Krulwich and D. H. Bechhofer. 2006. Adaptive gene expression in Bacillus subtilis strains deleted for tetL. J. Bacteriol. 188: 7090-7100.

Terahara, N., M. Fujisawa, B. Powers, T. M Henkin, T. A. Krulwich and M. Ito. 2006. An intergenic stem-loop mutation in the Bacillus subtilis ccpA-motPS operon increases motPS transcription and the contribution of MotPS to motility. J. Bacteriol. 188: 2701-2705.

Padan, E., E. Bibi, M. Ito and T. A. Krulwich. 2005. Alkaline pH homeostasis in bacteria: new insights. Biochim. Biophys. Acta 1717: 67-88.

Swartz, T. H., S. Idewada, O. Ishikawa, M. Ito and T. A. Krulwich. 2005. The Mrp system: a giant among monovalent cation/proton antiporters? Extremophiles 9:345-354.

Krulwich, T. A., O. Lewinson, E. Padan and E. Bibi. 2005. Do physiological roles foster persistence of drug/multidrug-efflux pumps? A case study. Nature Rev. Microbiol. 3:566-572.

Liu, J., Y. Xue, Q. Wang, Y. Wei, T. Swartz, D. B. Hicks, Y. Ma and T. A. Krulwich. 2005. The activity profile of the NhaD-type Na+(Li+)/H+ antiporter from the soda lake alkaliphile Alkalimonas amylolytica is adaptive for its extreme environment. J. Bacteriol. 187: 7589-7595.

Ito, M., N. Terahara, S. Fujinami and T. A. Krulwich. 2005. Properties of motility in Bacillus subtilis powered by the H+-coupled MotAB flagellar stator, Na+-coupled MotPS or hybrid stators MotAS and MotPB. J. Mol. Biol. 352:396-408.

De Jesus, M., J. Jin, A. A. Guffanti and T. A. Krulwich. 2005. Importance of the GP dipeptide of the antiporter motif and other membrane-embedded proline and glycine residues in tetracycline efflux protein Tet(L). Biochemistry 44:12896-12904.

Swartz, T. H., M. Ito, D. B. Hicks, M. Nuqui, A. A. Guffanti and T. A. Krulwich. 2005. The Mrp Na+/H+ antiporter increases activity of the malate:quinone oxidoreductase of an Escherichia coli respiratory mutant. J. Bacteriol. 187:388-391.

Rivera-Torres, I. O., R. D. Krueger-Koplin, D. B. Hicks, S. M. Cahill, T. A. Krulwich and M. E. Girvin. 2004. pKa of the essential Glu54 and backbone conformation for subunit c from the H+-coupled F1F0 ATP synthase from an alkaliphilic Bacillus. FEBS Lett. 575:131-135.

Ito, M., H. Xu, A. A. Guffanti, Y. Wei, L. Zvi, D. E. Clapham and T. A. Krulwich. 2004. The voltage-gated Na+ channel NaVBP has a role in motility, chemotaxis and pH homeostasis of an alkaliphilic Bacillus. Proc. Natl. Acad. Sci. U. S. A. 101:10566-10571 .

Ito, M., D. B. Hicks, T. M. Henkin, A. A. Guffanti, B. Powers, K. Uematsu and T. A. Krulwich. 2004. MotPS is the stator-force generator for motility of alkaliphilic Bacillus and its homologue is a second functional Mot in Bacillus subtilis. Mol. Microbiol. 53:1035-1049.

Wang, Z., D. B. Hicks, A. A. Guffanti, K. Baldwin, and T. A. Krulwich. 2004. Replacement of amino acid sequence features of a- and c-subunits of ATP synthases of alkaliphilic Bacillus with the Bacillus consensus sequence results in defective oxidative phosphorylation and non-fermentative growth at pH 10.5. J. Biol. Chem. 279:26446-26554.

Kreuger-Koplin, R. D., P. L. Sorgen, S. T. Krueger-Koplin, I. O. Rivera-Torres, S.M. Cahill, D. B. Hicks, L. Grinius, T. A. Krulwich, and M. E. Girvin. 2004. An evaluation of detergents for NMR structural studies of membrane proteins. J. Biomol. NMR 17: 43-57.

Safferling, M., H. Griffith, J. Jin, J. Sharp, M. De Jesus, C. Ng, T. A. Krulwich and D.N. Wang. 2003. TetL tetracycline efflux protein from Bacillus subtilis is a dimer in the membrane and in detergent solution. Biochemistry 42:13969-13976.

Hicks, D. B., Z. Wang, Y. Wei, R. Kent, A. A. Guffanti, H. Banciu, D. H. Bechhofer and T. A. Krulwich. 2003. A newly discovered tenth atp gene and the conserved atpI gene of a Bacillus atp operon have a role in Mg2+ uptake. Proc. Natl. Acad. Sci. U.S.A. 100:10213-10218.

Wei, Y., T.W. Southworth, H. Kloster, M. Ito, A.A. Guffanti, A. Moir, and T.A. Krulwich. 2003. Mutational loss of a K+ and NH4+ transporter affects the growth and endospore formation of alkaliphilic Bacillus pseudofirmus OF4. J. Bacteriol. 185:5133-5147.

Guffanti, A.A., Y. Wei, S.V. Rood and T.A. Krulwich. 2002. An antiport mechanism for a member of the Cation Diffusion Facilitator Family: divalent cations efflux in exchange for K+ and H+. Mol. Microbiol. 45:145-153.

Jin, J., A.A. Guffanti, D.H. Bechhofer and T.A. Krulwich. 2002. Tet(L) and Tet(K) tetracycline-divalent metal/H+ antiporters: characterization of multiple catalytic modes and a mutagenesis approach to difference in their efflux substrate and coupling ion preferences. J. Bacteriol. 184:4722-4732.

Southworth, T.W., A.A. Guffanti, A. Moir and T.A. Krulwich. 2001. GerN: an endospore germination protein of Bacillus cereus is a Na+/H+-K+ antiporter. J. Bacteriol. 183:5896-5903.

Ito, M., A.A. Guffanti, and T.A. Krulwich. 2001. Mrp-dependent Na+/H+ antiporters of Bacillus exhibit characteristics that are unanticipated for completely secondary active transporters. FEBS Lett. 496:117-120.

Jin, J., A.A. Guffanti, C. Beck, and T.A. Krulwich. 2001. A 12-transmembrane segment (TMS) version (∆TMS VII-VIII) of the 14-TMS Tet(L) antibiotic-resistance protein retains monovalent cation transport modes but lacks tetracycline efflux capacity. J. Bacteriol. 183:2667-2671.

Gilmour, R, P. Messner, A.A. Guffanti, R. Kent, A. Scheberl, N. Kendrick, and T.A. Krulwich. 2000. Two-dimensional gel electrophoresis analyses of pH-dependent protein expression in facultatively alkaliphilic Bacillus pseudofirmus OF4 lead to characterization of an S-layer protein with a role in alkaliphily. J. Bacteriol. 182:5969-5981.

Ito, M., A.A. Guffanti, W. Wang, and T.A. Krulwich. 2000. Effects of non-polar mutations in each of the seven Bacillus subtilis mrp genes suggest complex interactions among the gene products in support of Na+– and alkali- but not cholate-resistance. J. Bacteriol. 182:5663-5670.

Wei, Y., A.A. Guffanti, M. Ito, and T.A. Krulwich. 2000. Bacillus subtilis YqkI is a novel Malic/NaLactate antiporter that enhances growth on malate at low protonmotive force. J. Biol. Chem. 275:30287-30292.

Wang.W., A.A. Guffanti, Y. Wei, M. Ito and T.A. Krulwich. 2000. Two types of Bacillus subtilis tetA(L) deletion strains reveal the physiological importance of TetA(L) in K+ acquisition as well as in Na+-, alkali-, and tetracycline resistance. J. Bacteriol. 182:2088-2095.

Takami, H. and T.A. Krulwich. 2000. Re-identification of facultatively alkaliphilic Bacillus firmus OF4 as Bacillus pseudofirmus OF4. Extremophiles 4:19-22.


Ito, M., A.A. Guffanti, B. Oudega, and T.A. Krulwich. 1999. mrp: a multigene, multifunctional locus in Bacillus subtilis with roles in resistance to cholate and to Na+, and in pH homeostasis. J. Bacteriol. 181:2394-2402.

Wei, Y., A.A. Guffanti and T.A. Krulwich. 1999. Sequence analysis and functional studies of a chromosomal region of alkaliphilic Bacillus firmus OF4 encoding an ABC-type transporter with similarity of sequence and Na+ exclusion capacity to the Bacillus subtilis NatAB transporter. Extremophiles 3:113-118.

Gronstad, A., E. Jaroszewicz, M. Ito, M.G. Sturr, T.A. Krulwich, and A.B. Kolsto. 1998. Physical map of alkaliphilic Bacillus firmus OF4 and detection of a large endogenous plasmid. Extremophiles 2:447-453.

Ito, M., A.A. Guffanti, J. Zemsky, D.M. Ivey, and T.A. Krulwich. 1997. The role of the nhaC-encoded Na+/H+ antiporter of alkaliphilic Bacillus firmus OF4. J. Bacteriol. 179:3851-3857.

Cheng, J., A.A. Guffanti, and T.A. Krulwich. 1997. A two gene ABC-type transport system involved in Na+ extrusion by Bacillus subtilis is induced by ethanol and protonophore. Mol. Microbiol. 23:1107-1120.

Gilmour, R., and T.A. Krulwich. 1997. Construction and characterization of a mutant of alkaliphilic Bacillus firmus OF4 with a disrupted cta operon and purification of a novel cytochrome bd. J. Bacteriol. 179:863-870.

Ito, M., B. Cooperberg, and T.A. Krulwich. 1997. Diverse genes of alkaliphilic Bacillus firmus OF4 that complement K+ uptake-deficient Escherichia coli include an ftsH homologue. Extremophiles 1:22-28.

Gilmour, R., and T.A. Krulwich. 1996. Purification and characterization of the succinate dehydrogenase complex and CO-reactive b-type cytochromes from the facultative alkaliphile Bacillus firmus OF4. Biochim. Biophys. Acta 1276:57-63.

Cheng, J., A.A. Guffanti, W. Wang, T.A. Krulwich, and D.H. Bechhofer. 1996. Chromosomal tetA(L) gene of Bacillus subtilis: regulation of expression and physiology of a tetA(L) deletion strain. J. Bacteriol. 178:2853-2860.

Cheng, J., K. Baldwin, A.A. Guffanti, and T.A. Krulwich. 1996. The Na+/H+ antiport activity conferred by the Bacillus subtilis tetA(L) gene, a 5′ truncation product of tetA(L), and related plasmid genes upon Escherichia coli. Antimicrob. Agents Chemother. 40:852-857.

Sturr, M.G., T.A. Krulwich, and D.B. Hicks. 1996. Purification of a cytochrome bd terminal oxidase encoded by the Escherichia coli app locus from a ΔcyoΔcyd strain complemented by genes from Bacillus firmus OF4. J. Bacteriol. 176:1742-1749.

Krulwich, T.A. 1995. Alkaliphiles: “basic” molecular problems of pH tolerance and bioenergetics. Molec. Microbiol. 15:403-410.

Guffanti, A.A. and T.A. Krulwich. 1995. Tetracycline/H+ antiport and Na+/H+ antiport catalyzed by the Bacillus subtilis Tet(L) transporter expressed in Escherichia coli. J. Bacteriol. 177:4557-4561.

Cheng, J., A.A. Guffanti, and T.A. Krulwich. 1994. The chromosomal tetracycline-resistance locus of Bacillus subtilis encodes a Na+/H+ antiporter that is physiologically important at elevated growth pH. J. Biol. Chem. 269:27365-27371

Guffanti, A.A. and T.A. Krulwich. 1994. Oxidative phosphorylation by ADP + Pi-loaded membrane vesicles from alkaliphilic Bacillus firmus OF4. J. Biol. Chem. 269:21576-21582.

Ivey, D.M., M.G. Sturr, T.A. Krulwich, and D.B. Hicks. 1994. The abundance of atp gene transcript and of the mem­brane F1Fo-ATPase as a function of the growth pH of alkaliphilic Bacil­lus firmus OF4. J. Bacteriol. 176:5167-5170.

Hicks, D.B., D. Cohen, and T.A. Krulwich. 1994. Reconstitu­tion of the energy-linked activities of the solubilized F1F0 ATP synthase from Ba­cillus subtilis. J. Bacteriol. 176:4192-4195.

Sturr, M.G., A.A. Guffanti, and T.A. Krulwich. 1994. Growth and bioenergetics of alkaliphilic Bacillus firmus OF4 in continuous culture at high pH. J. Bacteriol. 176:3111-3116.

Quirk, P.G., A.A. Guffanti, S. Clejan, J. Cheng, and T.A. Krulwich. 1994. Isolation of Tn917 insertional mutants of Bacillus subtilis that are resistant to the protonophore carbonyl cyanide m-chlorophenylhydrazone. Biochim. Biophys. Acta 1186:27-34.

Ivey, D.M., A.A. Guffanti, J. Zemsky, E. Pinner, R. Karpel, S. Schuldiner, E. Padan, and T.A. Krulwich. 1993. Cloning and character­ization of a putative Ca+2/H+ antiporter gene from Escherichia coli upon func­tional complementation of Na+/H+ antiporter-deficient strains by the overexpressed gene. J. Biol. Chem. 268:11296-11303.

Quirk, P.G., E.A. Dunkley, Jr., P. Lee, and T.A. Krulwich. 1993. Identification of a putative Bacillus subtilis rho gene. J. Bacteriol. 175:647-654.

Quirk, P.G., D.B. Hicks, and T.A. Krulwich. 1993. Cloning of the cta operon from alkaliphilic Bacillus firmus OF4, and characteriza­tion of the pH-regulated cy­to­chrome caa3 oxidase it encodes. J. Biol. Chem. 268:678-685.

Ivey, D.M., J. Cheng, and T.A. Krulwich. 1992. A 1.6 kb region of Bacillus firmus OF4 DNA encodes a homolog of Escherichia coli and yeast DNA topoisomerases and may con­tain a translational read-through of UGA. Nucleic Acids Res. 20:4928.

Khan, S., D.M. Ivey, and T.A. Krulwich. 1992. Mem­brane ultrastructure of alkaliphilic Bacillus species stud­ied by rapid freeze electron microscopy. J. Bacteriol. 174:5123-5126.

Ivey, D.M., A.A. Guffanti, Z.H. Shen, N. Kudyan, and T.A. Krulwich. 1992. The cadC gene product of alkaliphilic Bacillus firmus OF4 partially restores Na+-resistance to an Escherichia coli strain lack­ing an Na+/H+ antiporter (NhaA). J. Bacteriol. 174:4878-4884.

Moody, A.J., R.B. Gennis, D.B., Hicks, W.J.., Ingledew, W.J., T.A. Krulwich, J.C. Rumbley, and P.R. Rich. 1992. Ligand binding properties of bacterial oxidases in rela­tion to cytochrome c oxidase. Biochem. Soc. Trans. 20:240S.

Guffanti, A.A. and T.A. Krulwich. 1992. Features of apparent non-chemiosmotic energization of oxidative phos­phorylation by alkaliphilic Bacillus firmus OF4. J. Biol. Chem. 267:9580-9588.

Ivey, D.M. and T.A. Krulwich. 1992. Two unrelated alkaliphilic Bacillus species possess identical de­vi­a­tions in sequence from those of conventional prokaryotes in re­gions of Fo genes implicated in proton transloca­tion through the ATP synthase. Res. Microbiol. 143:467-470.

Ivey, D.M., A.A. Guffanti, J.S. Bossewitch, E. Padan, and T.A. Krulwich. 1991. Molecular cloning and sequenc­ing of a gene from alkaliphilic Bacillus firmus OF4 that func­tional­ly complements an Escherichia coli strain carrying a dele­tion in the nhaA Na+/H+ antiporter gene. J. Biol. Chem. 266:23483-23489.

Ivey, D.M. and T.A. Krulwich. 1991. Structure and nu­cleotide sequence of the genes encoding the ATP syn­thase from alkaliphilic Bacillus firmus OF4. Mol. Gen. Genet. 229:292-300.

Quirk, P., A.A. Guffanti, R.J. Plass, S. Clejan, and T.A. Krulwich. 1991. Protonophore-resistance and cyto­chrome expression in mutant strains of the faculta­tive alkaliphile Bacillus firmus OF4. Biochim. Biophys. Acta 1058:131-140.

Ivey, D.M., D.B. Hicks, A.A. Guffanti, G. Sobel, and T.A. Krulwich. 1990. The problem of the electrochemical proton potential in alkaliphilic bacteria. Mosbach Collo­quium (Springer-Verlag, Heidelberg) 41:105-113.

Krulwich, T.A., P.G. Quirk, and A.A. Guffanti. 1990. Uncoupler-resistant mutants of bacteria. Microbiol. Revs. 54:52-65.

Krulwich, T.A. and D.M. Ivey. 1990. Bioenergetics in extreme environments. In: The Bacteria: A Treatise on Structure and Function, J. Sokatch and N. Ornston, eds., Volume 12, Bacterial Energetics, T. Krulwich, ed., Aca­dem­ic Press, Orlando, Florida.

Hicks, D.B. and T.A. Krulwich. 1990. Purification and reconstitution of the F1Fo-ATP synthase from alkaliphilic Bacillus firmus OF4: evidence that the enzyme translocates H+ but not Na+. J. Biol. Chem. 265:20547-20554.

Ivey, D.M. and T.A. Krulwich. 1990. Sequence of the gene encoding the ATP synthase beta subunit from alkaliphilic Bacillus firmus RAB. Nucl. Acids Res. 18:1296.


Krulwich, T.A. and A.A. Guffanti. 1989. Alkalophilic bacteria. Ann. Rev. Microbiol. 43:435-463.

Rohde, M., F. Mayer, D.B. Hicks, and T.A. Krulwich. 1989. Immunoelectron microscopic localization of the F1FoATPase (ATP synthase) on the cytoplasmic mem­brane of alkalophilic Bacillus firmus RAB. Biochim. Biophys. Acta 985:233-235.

Clejan, S., A.A. Guffanti, M.A. Cohen, and T.A. Krulwich. 1989. Mutation of Bacillus firmus OF4 to duramycin resistance results in substan­tial replacement of membrane lipid phosphatidylethanolamine by its plasmal­ogen form. J. Bacteriol. 171:1744-1746.

Clejan, S., and T.A. Krulwich. 1988. Permeability studies of lipid vesicles from alkalophilic Bacillus firmus showing opposing effects of mem­brane iso­pren­oid and diacylglycerol fractions and suggest­ing a pos­sible basis for obligate alkalophily. Biochim. Biophys. Acta 946: 40-48.

Guffanti, A.A. and Krulwich, T.A. 1988. ATP synthesis is driven by an imposed ΔpH or ΔμH+ but not by an im­posed ΔpNa+ or ΔμNa+ in alkalophilic Bacillus firmus OF4 at high pH. J. Biol. Chem. 263:14748-14752.

Dunkley, E.A., Jr., S. Clejan, A.A. Guffanti, and T.A. Krulwich. 1988. Large decreases in membrane phosphatidyl­ethanolamine and diphosphatidylglycerol upon mutation to duramycin resistance do not change the protonophore resis­tance of Bacillus subtilis. Biochim. Biophys. Acta 943:13-18.

Davidson, M.W., K.A. Gray, D.B. Knaff, and T.A. Krulwich. 1988. Purification and characterization of two soluble cytochromes from the alkalophile Bacillus firmus RAB. Biochim. Biophys. Acta 933:470-477.

Clejan, S., A.A. Guffanti, L.H. Falk, and T.A. Krulwich. 1988. The protonophore resistance of Bacillus megaterium is correlated with elevated ra­tios of sat­u­rat­ed to unsatu­rated fatty acids in membrane phospholipids. Biochim. Biophys. Acta 932:43-51.

Hicks, D.B. and T.A. Krulwich. 1987. Studies of the F1 ATPase from Bacillus subtilis and its uncoupler-resistant mutant derivatives. J. Bacteriol. 169:4743-4749.

Krulwich, T.A., S. Clejan, L. Falk, and A.A. Guffanti. 1987. Incorporation of specific exoge­nous fatty acids into membrane lipids modulates protonophore resistance in Ba­cillus subtilis. J. Bacteriol. 169:4479-4485.

Guffanti, A.A., S. Clejan, L.H. Falk, D.B. Hicks, and T.A. Krulwich. 1987. Isolation and characterization of uncoupler-resistant mutants of Bacillus subtilis. J. Bacteriol. 169:4469-4478.

Clejan, S., T.A. Krulwich, K.R. Mondrus, and D. Seto-Young. 1986. Membrane lipid composition of obligately and facultatively alkalophilic strains of Bacillus. J. Bacteriol. 168:334-340.

Hicks, D.B. and T.A. Krulwich. 1986. The membrane ATP­ase of alkalophilic Bacillus firmus RAB is an F1-type ATPase. J. Biol. Chem. 261:12896-12902.

Guffanti, A.A., O. Finkelthal, D.B. Hicks, L. Falk, A. Sidhu, A. Garro, and T.A. Krulwich. 1986. Isolation and characterization of new facultatively alkalophilic strains of Bacillus. J. Bacteriol. 167:766-773.

Krulwich, T.A., A.A. Guffanti, M.Y. Fong, L. Falk, and D.B. Hicks. 1986. Alkalophilic Bacillus firmus RAB gen­er­ates variants which can grow at lower Na+ concentra­tions than the parental strain. J. Bacteriol. 165:884-889.

Seto-Young, D., M.L. Garcia, and T.A. Krulwich. 1985. Reconstitution of a bacterial Na+/H+ antiporter. J. Biol. Chem. 260:11393-11395.

Krulwich, T.A., R. Agus, M. Schneier, and A.A. Guffanti. 1985. The buffering capacity of bacilli that grow in different ranges of pH. J. Bacteriol. 162:768-772.

Krulwich, T.A., J.G. Federbush, and A.A. Guffanti. 1985. Presence of a non-metabolizable solute that is translocated with Na+ enhances Na+-dependent pH homeosta­sis in an alkalophilic Bacillus. J. Biol. Chem. 260:4055-4058.

Guffanti, A.A., E. Chiu, and T.A. Krulwich. 1985. Failure of an alkalophilic bacterium to synthe­size ATP in re­sponse to a valinomycin-induced potassium diffu­sion po­tential at high pH. Arch. Biochem. Biophys. (is­sue hon­or­ing B.L. Horecker) 239:327-333.

Guffanti, A.A., M. Mann, T.L. Sherman, and T.A. Krulwich. 1984. Patterns of electrochemical proton gradi­ent formation by membrane vesi­cles from an obligately aci­do­philic bacterium. J. Bacteriol. 159:448-452.

Kitada, M. and T.A. Krulwich. 1984. Purification and characterization of the cytochrome oxidase from alkalophilic Bacillus firmus RAB. J. Bacteriol. 158:963-967.

Guffanti, A.A., R.T. Fuchs, M. Schneier, E. Chiu, and T.A. Krulwich. 1984. A ΔΨ-generated by respiration is not equivalent to a diffusion potential of the same magni­tude for ATP synthesis by Bacillus firmus RAB. J. Biol. Chem. 259: 2971-2975.

Garcia, M.L., A.A. Guffanti, and T.A. Krulwich. 1983. Characterization of the Na+/H+ antiporter of alkalophilic bacilli in vivo: ΔΨ-dependent 22Na+ efflux from starved cells. J. Bacteriol. 156:1151-1157.

Krulwich, T.A. 1983. Na+/H+ antiporters. Biochim. Biophys. Acta 726:245-264.

Kitada, M., A.A. Guffanti, and T.A. Krulwich. 1982. Bioenergetic properties and viability of the alkalophilic Bacillus firmus RAB as a function of pH and Na+ content of the medium. J. Bacteriol. 152:1096-1104.

Bonner, S., M. Mann, A.A. Guffanti, and T.A. Krulwich. 1982. Na+/solute symport in membrane vesicles from Bacil­lus alcalophilus. Biochim. Biophys. Acta 679:315-322.

Lewis, R.J., R. Prince, P.L. Dutton, D. Knaff, and T.A. Krulwich. 1981. The respiratory chain of Bacillus alcalophilus and its non-alkalophilic mu­tant de­riv­a­tive. J. Biol. Chem. 256:10543-10549.

Guffanti, A.A., H. Blumenfeld, and T.A. Krulwich. 1981. ATP synthesis by an uncoupler_resistant mutant of Bacil­lus megaterium. J. Biol. Chem. 256:8416-8421.

Guffanti, A.A., R.F. Bornstein, and T.A. Krulwich. 1981. Oxidative phosphorylation by membrane vesicles from Bacillus alcalophilus. Biochim. Biophys. Acta 635:619-630.

Mandel, K.G., A.A. Guffanti, and T.A. Krulwich. 1980. Monovalent cation/proton antiporters in membrane vesi­cles from Bacillus alcalophilus. J. Biol. Chem. 225:7391-7396.

Krulwich, T.A., K.G. Mandel, R.F. Bornstein, and A.A. Guffanti, 1979. A non-alkalophilic mutant of Bacillus alcalophilus lacks the Na+/H+ antiporter. Biochem. Biophys. Res. Comm. 91: 58-62.

Krulwich, T.A. and N. Pelliccione. 1979. Catabolic pathways of coryneforms, nocardia and mycobacteria. Ann. Rev. Microbiol. 33:95-111.

Pelliccione, N., B. Jaffin, M.E. Sobel and T.A. Krulwich. 1979. Induction of the phosphoenolpyruvate:hexose phosphotransferase system associated with relative anaerobiosis in an obligate aerobe. Eur. J. Biochem. 95: 69-75.

Note: Apart from the research efforts detailed above Terry Krulwich has been engaged in outreach efforts to enhance diversity in academic science and elsewhere; currently specific efforts include Program Directorship of Mount Sinai PREP (Post-baccalaureate Research Education Program; which is sponsored by NIGMS and in its 15th year). Several publications related to diversity efforts are cited below.

Butts, G.C., Y. Hurd, A.G. Palermo, A.G., D. Delbrune, S. Saran, C. Zony, and T.A. Krulwich. 2012. Role of institutional climate in fostering diversity in biomedical research workforce: a case study. Mt. Sinai J. Med. 79:498-511.

McGee, R., Jr., Saran, S., and T.A. Krulwich. 2012. Diversity in the biomedical research workforce: developing talent. Mt. Sinai J. Med. 79: 397-411.