38. Susuki S, Kuenen JG, Schipper K, van der Velde S, Ishii S, Wu A et al (2014) Physiological and
genomic features of highly alkaliphilic hydrogen-utilizing Betaproteobacteria from a continental serpentinizing site. Nat Commun 5:3900
39. Krulwich TA, Hicks DB, Swartz TH, Ito M (2007) Bioenergetic adaptations that support
alkaliphily. In: Gerday C, Glansdorff N (eds) Physiology and biochemistry of extremophiles.
ASM Press, Washington, pp 311–329
40. Skulachev VP (1995) Membrane-linked energy transductions. Bioenergetic functions of
sodium H
+ is not unique as a coupling ion. FEBS J 151(2):199–208
41. Hicks DB, Krulwich TA (1995) The respiratory chain of alkaliphilic bacteria. Biochim Biophys
Acta 1229:303–314
42. Mulkidjanian AY, Galperin MY, Koonin V (2009) Co-evolution of primordial membranes and
membrane proteins. Trends Biochem Sci 34(4):206–215
43. Ventosa A, Nieto JJ, Oren A (1998) Biology of moderately halophilic aerobic bacteria.
Microbiol Mol Biol Rev 62(2):504–544
44. Ito M, Guffanti AA, Krulwich TA (2001) Mrp-dependent Na
+ /H
+ antiporters of Bacillus exhibit
characteristics that are unanticipated for completely secondary active transporters. FEBS Lett
496:117–120
45. Krulwich TA, Ito M, Guffanti AA (2001) The Na(+)-dependence of alkaliphily in Bacillus.
Biochim Biophys Acta 1505(1):158–168
46. Ito M, Xu H, Guffani AA, Wei Y, Zvi L, Clapham DE et al (2004) The voltage-gates 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
47. Horikoshi K (1999) Alkaliphiles: some applications of their products for biotechnology.
Microbiol Mol Biol Rev 63(4):735–750
48. Liu J, Xue Y, Wang Q, Wei Y, Swartz TH, Hicks DB et al (2005) The activity profile of the
NhaD-type Na
+ (Li
+ )/H
+ antiporter from the soda Lake Haloalkaliphile Alkalimonas amylolytica
is adaptive for the extreme environment. J Bacteriol 187(22):7589–7595
49. Janausch I, Zientz GE, Tran Q, Kroger HA, Unden G (2002) C4-dicarboxylate carriers and
sensors in bacteria. Biochim Biophys Acta 1553:39–56
50. Wutipraditkul N, Waditee R, Incharoensakdi A, Hibino T, Tanaka Y, Nakamura T et al (2005)
Halotolerant cyanobacterium Aphanothece halophytica contains NapA-Type Na
+ /H
+ antiporters
with novel ion specificity that are involved in salt tolerance at alkaline pH. Appl Environ
Microbiol 71(8):4176–4184
51. Krulwich TA (1985) Alkaliphiles: “basic” molecular problems of the pH tolerance and bioenergetics. Mol Microbiol 15(3):403–410
52. Hirota N, Kitada M, Imae Y (1981) Flagellar motors of alkalophilic Bacillus are powered by an
electrochemical potential gradient of Na
+ . FEBS Lett 132(2):278–280
53. Terahara N, Krulwich TA, Ito M (2008) 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 U S A 105(38):14359–14364
54. Fujinami S, Terahara N, Krulwich TA, Ito M (2009) Motility and chemotaxis in alkaliphilic
Bacillus species. Future Microbiol 4(9):1137–1149
55. Ren D, Navarro B, Xu H, Yue L, Shi Q, Clapham DE (2001) A prokaryotic voltage-gated
sodium channel. Science 294(5550):2372–2375
56. Olsson K, Keis S, Morgan HW, Dimroth P, Cook GM (2003) Bioenergetic properties of the
Thermoalkaliphilic Bacillus sp. strain TA2.A1. J Bacteriol 185(2):461–465
57. Guffanti AA, Finkelthal O, Hicks DB, Falk L, Sidhu A, Garro A (1986) Isolation and
characterization of new facultatively alkalophilic strains of Bacillus species. J Bacteriol
167(3):766–773
58. Kitada M, Lewis RJ, Krulwich TA (1983) Respiratory Chain of the alkalophilic bacterium
Bacillus firmus RAB and its non-alkalophilic mutant derivative. J Bacteriol 154(1):330–335
59. Krulwich TA, Ito M, Hicks DB, Gilmour R, Guffanti AA (1998) pH homeostasis and ATP
synthesis: studies of two processes that necessitate inward proton translocation in extremely
alkaliphilic Bacillus species. Extremophiles 2:217–222
154
P. H. Lebre and D. A. Cowan
genomic features of highly alkaliphilic hydrogen-utilizing Betaproteobacteria from a continental serpentinizing site. Nat Commun 5:3900
39. Krulwich TA, Hicks DB, Swartz TH, Ito M (2007) Bioenergetic adaptations that support
alkaliphily. In: Gerday C, Glansdorff N (eds) Physiology and biochemistry of extremophiles.
ASM Press, Washington, pp 311–329
40. Skulachev VP (1995) Membrane-linked energy transductions. Bioenergetic functions of
sodium H
+ is not unique as a coupling ion. FEBS J 151(2):199–208
41. Hicks DB, Krulwich TA (1995) The respiratory chain of alkaliphilic bacteria. Biochim Biophys
Acta 1229:303–314
42. Mulkidjanian AY, Galperin MY, Koonin V (2009) Co-evolution of primordial membranes and
membrane proteins. Trends Biochem Sci 34(4):206–215
43. Ventosa A, Nieto JJ, Oren A (1998) Biology of moderately halophilic aerobic bacteria.
Microbiol Mol Biol Rev 62(2):504–544
44. Ito M, Guffanti AA, Krulwich TA (2001) Mrp-dependent Na
+ /H
+ antiporters of Bacillus exhibit
characteristics that are unanticipated for completely secondary active transporters. FEBS Lett
496:117–120
45. Krulwich TA, Ito M, Guffanti AA (2001) The Na(+)-dependence of alkaliphily in Bacillus.
Biochim Biophys Acta 1505(1):158–168
46. Ito M, Xu H, Guffani AA, Wei Y, Zvi L, Clapham DE et al (2004) The voltage-gates 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
47. Horikoshi K (1999) Alkaliphiles: some applications of their products for biotechnology.
Microbiol Mol Biol Rev 63(4):735–750
48. Liu J, Xue Y, Wang Q, Wei Y, Swartz TH, Hicks DB et al (2005) The activity profile of the
NhaD-type Na
+ (Li
+ )/H
+ antiporter from the soda Lake Haloalkaliphile Alkalimonas amylolytica
is adaptive for the extreme environment. J Bacteriol 187(22):7589–7595
49. Janausch I, Zientz GE, Tran Q, Kroger HA, Unden G (2002) C4-dicarboxylate carriers and
sensors in bacteria. Biochim Biophys Acta 1553:39–56
50. Wutipraditkul N, Waditee R, Incharoensakdi A, Hibino T, Tanaka Y, Nakamura T et al (2005)
Halotolerant cyanobacterium Aphanothece halophytica contains NapA-Type Na
+ /H
+ antiporters
with novel ion specificity that are involved in salt tolerance at alkaline pH. Appl Environ
Microbiol 71(8):4176–4184
51. Krulwich TA (1985) Alkaliphiles: “basic” molecular problems of the pH tolerance and bioenergetics. Mol Microbiol 15(3):403–410
52. Hirota N, Kitada M, Imae Y (1981) Flagellar motors of alkalophilic Bacillus are powered by an
electrochemical potential gradient of Na
+ . FEBS Lett 132(2):278–280
53. Terahara N, Krulwich TA, Ito M (2008) 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 U S A 105(38):14359–14364
54. Fujinami S, Terahara N, Krulwich TA, Ito M (2009) Motility and chemotaxis in alkaliphilic
Bacillus species. Future Microbiol 4(9):1137–1149
55. Ren D, Navarro B, Xu H, Yue L, Shi Q, Clapham DE (2001) A prokaryotic voltage-gated
sodium channel. Science 294(5550):2372–2375
56. Olsson K, Keis S, Morgan HW, Dimroth P, Cook GM (2003) Bioenergetic properties of the
Thermoalkaliphilic Bacillus sp. strain TA2.A1. J Bacteriol 185(2):461–465
57. Guffanti AA, Finkelthal O, Hicks DB, Falk L, Sidhu A, Garro A (1986) Isolation and
characterization of new facultatively alkalophilic strains of Bacillus species. J Bacteriol
167(3):766–773
58. Kitada M, Lewis RJ, Krulwich TA (1983) Respiratory Chain of the alkalophilic bacterium
Bacillus firmus RAB and its non-alkalophilic mutant derivative. J Bacteriol 154(1):330–335
59. Krulwich TA, Ito M, Hicks DB, Gilmour R, Guffanti AA (1998) pH homeostasis and ATP
synthesis: studies of two processes that necessitate inward proton translocation in extremely
alkaliphilic Bacillus species. Extremophiles 2:217–222
154
P. H. Lebre and D. A. Cowan