94. McMillan DGG, Keis S, Dimroth P, Gregory M, Cook GM (2007) A specific adaptation in the
a-subunit of thermoalkaliphilic F 1 F O -ATP synthase enables ATP synthesis at high pH but not
at neutral pH values. J Biol Chem 282:17395–17404
95. Fujisawa F, Fackelmayer OJ, Liu J, Krulwich TA, Hicks DB (2010) The ATP synthase
a-subunit of extreme alkaliphiles is a distinct variant: mutations in the critical alkaliphilespecific residue Lys180 and other residues that support alkaliphile oxidative phosphorylation.
J Biol Chem 285:32105–32115
96. Cook GM, Keis S, Morgan HW, von Ballmoos C, Matthey U, Kaim G, Dimroth P (2003)
Purification and biochemical characterization of the F 1 F o -ATP synthase from
thermoalkaliphilic Bacillus sp. strain TA2.A1. J Bacteriol 85:4442–4449
97. Dimroth P, Cook GM (2004) Bacterial Na
+ - or H
+ -coupled ATP synthases operating at low
electrochemical potential. Adv Microb Physiol 49:175–218
98. Hicks DB, Krulwich TA (1990) Purification and reconstitution of the F 1 F O -ATP synthase from
alkaliphilic Bacillus firmus OF4. Evidence that the enzyme translocates H
+ but not Na
+ . J Biol
Chem 265:20547–20554
99. Hoffmann A, Dimroth P (1990) The ATPase of Bacillus alcalophilus. Purification and
properties of the enzyme. Eur J Biochem 194:423–430
100. Burne RA, Marquis RE (2000) Alkali production by oral bacteria and protection against dental
caries. FEMS Microbiol Lett 193:1–6
101. Yokaryo H, Tokiwa Y (2014) Isolation of alkaliphilic bacteria for production of high optically
pure L-(+)-lactic acid. J Gen Appl Microbiol 60:270–275
102. Wilks JC, Kitko RD, Cleeton SH, Lee GE, Ugwu CS, Jones BD, BonDurant SS, Slonczewski
JL (2009) Acid and base stress and transcriptomic responses in Bacillus subtilis. Appl Environ
Microbiol 75:981–990
103. Graham AF, Lund BM (1983) The effect of alkaline pH on growth and metabolic products of a
motile, yellow-pigmented Streptococcus sp. J Gen Microbiol 129:2429–2435
104. Hirota K, Aino K, Yumoto I (2013) Amphibacillus iburiensis sp. nov., an alkaliphile that
reduces an indigo dye. Int J Syst Evol Microbiol 63:4303–4308
105. Horikoshi K (2006) Alkaliphiles. Kodansha, New York
106. Aono R, Ito M, Joblin KN, Horikoshi K (1995) A high cell wall negative charge is necessary
for the growth of the alkaliphile Bacillus lentus C-125 at elevated pH. Microbiology
141:2955–2964
107. Hancock IC, Baddiley J (1985) Biosynthesis of the bacterial envelope polymers teichoic acid
and teichuronic acid. In: Martonosi NA (ed) The enzymes of biological membranes, vol 2. 2nd
edn. Plenum, New York, pp 279–307
108. Ward JB (1981) Teichoic and teichuronic acids: biosynthesis, assembly and location.
Microbiol Rev 45:211–243
109. Archibald AR, Baddiley J, Blumsom NL (1968) The teichoic acids. Adv Enzymol Relat Areas
Mol Biol 30:223–253
110. Archibald AR, Hancock IC, Harwood CR (1993) Cell wall structure, synthesis and turnover.
In: Sonenshein A, Hoch JA, Losick R (eds) Bacillus subtilis and other Gram-positive bacteria.
American Society for Microbiology, Washington, pp 381–410
111. Araki Y, Ito E (1989) Linkage units in cell walls of Gram-positive bacteria. CRC Crit Rev
Microbiol 17:121–135
112. Naumova IB, Shashkov AS (1997) Anionic polymers in cell walls of Gram-positive bacteria.
Biochemistry 62:809–840
113. Aono R, Horikoshi K (1983) Chemical composition of cell walls of alkalophilic strains of
Bacillus. J Gen Microbiol 129:1083–1087
114. Horikoshi K (1999) Alkaliphiles: some applications of their products for biotechnology.
Microbiol Mol Biol Rev 63:735–750
115. Koch AL (1986) The pH in the neighborhood of membranes generating a protonmotive force.
J Theor Biol 120:73–84
126
G. Mamo
a-subunit of thermoalkaliphilic F 1 F O -ATP synthase enables ATP synthesis at high pH but not
at neutral pH values. J Biol Chem 282:17395–17404
95. Fujisawa F, Fackelmayer OJ, Liu J, Krulwich TA, Hicks DB (2010) The ATP synthase
a-subunit of extreme alkaliphiles is a distinct variant: mutations in the critical alkaliphilespecific residue Lys180 and other residues that support alkaliphile oxidative phosphorylation.
J Biol Chem 285:32105–32115
96. Cook GM, Keis S, Morgan HW, von Ballmoos C, Matthey U, Kaim G, Dimroth P (2003)
Purification and biochemical characterization of the F 1 F o -ATP synthase from
thermoalkaliphilic Bacillus sp. strain TA2.A1. J Bacteriol 85:4442–4449
97. Dimroth P, Cook GM (2004) Bacterial Na
+ - or H
+ -coupled ATP synthases operating at low
electrochemical potential. Adv Microb Physiol 49:175–218
98. Hicks DB, Krulwich TA (1990) Purification and reconstitution of the F 1 F O -ATP synthase from
alkaliphilic Bacillus firmus OF4. Evidence that the enzyme translocates H
+ but not Na
+ . J Biol
Chem 265:20547–20554
99. Hoffmann A, Dimroth P (1990) The ATPase of Bacillus alcalophilus. Purification and
properties of the enzyme. Eur J Biochem 194:423–430
100. Burne RA, Marquis RE (2000) Alkali production by oral bacteria and protection against dental
caries. FEMS Microbiol Lett 193:1–6
101. Yokaryo H, Tokiwa Y (2014) Isolation of alkaliphilic bacteria for production of high optically
pure L-(+)-lactic acid. J Gen Appl Microbiol 60:270–275
102. Wilks JC, Kitko RD, Cleeton SH, Lee GE, Ugwu CS, Jones BD, BonDurant SS, Slonczewski
JL (2009) Acid and base stress and transcriptomic responses in Bacillus subtilis. Appl Environ
Microbiol 75:981–990
103. Graham AF, Lund BM (1983) The effect of alkaline pH on growth and metabolic products of a
motile, yellow-pigmented Streptococcus sp. J Gen Microbiol 129:2429–2435
104. Hirota K, Aino K, Yumoto I (2013) Amphibacillus iburiensis sp. nov., an alkaliphile that
reduces an indigo dye. Int J Syst Evol Microbiol 63:4303–4308
105. Horikoshi K (2006) Alkaliphiles. Kodansha, New York
106. Aono R, Ito M, Joblin KN, Horikoshi K (1995) A high cell wall negative charge is necessary
for the growth of the alkaliphile Bacillus lentus C-125 at elevated pH. Microbiology
141:2955–2964
107. Hancock IC, Baddiley J (1985) Biosynthesis of the bacterial envelope polymers teichoic acid
and teichuronic acid. In: Martonosi NA (ed) The enzymes of biological membranes, vol 2. 2nd
edn. Plenum, New York, pp 279–307
108. Ward JB (1981) Teichoic and teichuronic acids: biosynthesis, assembly and location.
Microbiol Rev 45:211–243
109. Archibald AR, Baddiley J, Blumsom NL (1968) The teichoic acids. Adv Enzymol Relat Areas
Mol Biol 30:223–253
110. Archibald AR, Hancock IC, Harwood CR (1993) Cell wall structure, synthesis and turnover.
In: Sonenshein A, Hoch JA, Losick R (eds) Bacillus subtilis and other Gram-positive bacteria.
American Society for Microbiology, Washington, pp 381–410
111. Araki Y, Ito E (1989) Linkage units in cell walls of Gram-positive bacteria. CRC Crit Rev
Microbiol 17:121–135
112. Naumova IB, Shashkov AS (1997) Anionic polymers in cell walls of Gram-positive bacteria.
Biochemistry 62:809–840
113. Aono R, Horikoshi K (1983) Chemical composition of cell walls of alkalophilic strains of
Bacillus. J Gen Microbiol 129:1083–1087
114. Horikoshi K (1999) Alkaliphiles: some applications of their products for biotechnology.
Microbiol Mol Biol Rev 63:735–750
115. Koch AL (1986) The pH in the neighborhood of membranes generating a protonmotive force.
J Theor Biol 120:73–84
126
G. Mamo
