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. 2020 Dec 30;22(1):341. doi: 10.3390/ijms22010341

Table 1.

Physiological and metabolic properties of Thermotogae species. YE: Yeast extract; BHI: Brain heart infusion; CMC: Carboxymethylcellulose; S0 = Elemental sulfur; Thio: Thiosulfate; Cys: Cysteine; AA: Acetic acid; LA: Lactic acid; ALA: Alanine; EPS: Exopolysaccharide; AABA: α-aminobutyrate; EtOH: Ethanol; AQDS: Anthraquinone-2,6-disulfonate; But: Butyrate; Val: Valerate; Glu: Glutamate; BuOH: Butanol; iBut: isobutyrate; iVal: isovalerate; PPA: Propionic Acid; Gly: Glycine; Pro: Proline; Fo: Formate; HPA: Hydroxyphenilacetate; PA: Phenylacetate; 3-IAA: Indole-3-acetate; 2-MeBu: 2-Methylbutyrate.

Genus Species Isolation Temp. Range/
Optimal (°C)		 pH
Range/
Optimal		 Cell
Dimension (Long by Wide) (µm) 		Growth Substrates NaCl Range/
Optimal
(%)		 Electron
Acceptor		 End
Products		 Ref.
Thermotoga Thermotoga petrophila Oil reservoir, Japan 47–88/
80		 5.2–9.0/
7.0		 2.0–7.0
by 0.7–1.0		 YE, peptone, glucose, fructose, ribose,
arabinose, sucrose, lactose, maltose,
starch, cellulose		 0.1–5.5/
1.0		 S0;
Thio		 AA, LA,
CO2, H2 		[72]
Thermotoga naphthophila Oil reservoir, Japan 48–86/
80		 5.4–9.0/
7.0		 2.0–7.0
by 0.8–1.2		 YE, peptone, glucose, galactose, fructose, mannitol, ribose, arabinose, sucrose,
lactose, maltose, starch		 0.1–6.0/
1.0		 S0;
Thio		 AA, LA,
CO2, H2 		[72]
Thermotoga maritima Geotermal
vent		 55–90/
80		 5.5–9.0/
6.5		 1.5–11.0
by 0.6		 ribose, xylose, glucose, sucrose, maltose,
lactose, galactose, starch, glycogen		 0.2–3.8/
2.7		 Fe (III)
S0;
Thio		 AA, LA, CO2, H2, ALA, EPS, AABA [3]
Thermotoga profunda Hot spring,
Japan		 50–72/
60		 6.0–8.6/
7.4		 0.8–2.1
by 0.4		 glucose, trehalose, cellobiose, arabinose,
xylose, ribose, pyruvate		 n. d S0;
Thio		 n. d [73]
Thermotoga caldifontis Hot spring,
Japan		 55–85/
70		 6.0–8.6/
7.4		 1.2–3.5
by 0.5		 glucose, maltose, trehalose, cellobiose,
arabinose, xylose, ribose, pyruvate, starch		 n. d Thio n. d [73]
Thermotoga neapolitana Submarine thermal vent 55–95/
77		 6.0–9.0/
7.5		 1.5–11.0
by 0.6		 fructose, fucose, galactose, mannose, rhamnose, pyruvate, glucosamine, lactulose, turanose, glycerol, dextrin, ribose, xylose, glucose, sucrose, maltose, lactose, starch, glycogen 0.2–6.0/
2.0		 S0 AA, ALA, CO2, H2 [74]
Pseudothermotoga Pseudothermotoga lettingae Thermophilic
bioreactor		 50–75/
65		 6.0–8.5/
7.0		 2.0–3.0
by 0.5–1.0		 glucose, EtOH, acetate, formate 0.0–2.8/
1.0		 S0; Thio; AQDS; Fe(III) AA, ALA, LA,
EtOH, AA, BA, CO2, H2 		[75]
Pseudothermotoga elfii Oil reservoir 50–72/
66		 5.5–7.5/
7.5		 2.0–3.0
by 0.5–1.0		 glucose, arabinose, fructose, lactose,
maltose, mannose, ribose, sucrose, xylose		 0.0–2.8/
1.0		 Thio AA, CO2, H2 [76]
Pseudothermotoga hypogea Oil reservoir, Africa 56–90/
70		 6.1–9.1/
7.3–7.4		 2.0–3.0
by 0.5–1.0		 fructose, galactose, glucose, lactose,
maltose, mannose, sucrose, xylose, xylan		 0.0–1.5/
0.2		 Thio AA, ALA, CO2, H2, EtOH [77]
Pseudothermotoga Pseudothermotoga subterranea Oil reservoir, Paris 50–75/
70		 6.0–8.5/
7.0		 3.0–10.0
by 0.5		 YE, peptone, tryptone, casein 0.0–2.4/
1.2		 Cys,
Thio		 n.d. [78]
Pseudothermotoga thermarum Hot spring,
Africa		 55–84/
70		 6.0–9.0/
7.0		 1.5–11.0
by 0.6		 starch, glucose, maltose 0.2–0.5/
0.35		 S0 n.d. [6]
Fervidobacterium Fervidobacterium nodosum Hot spring,
New Zealand		 40–80/
65–70		 6.0–8.0/
7.0		 1.0–2.5
by 0.5–0.55		 glucose, sucrose, starch and lactose n.d./<1.0 S0 AA, LA, CO2, H2, EtOH, But, Val [5]
Fervidobacterium pennavorans Hot spring,
Portugal		 50–80/
70		 5.5–8.0/
6.5		 2.0–20.0
by 0.5		 cellobiose, starch, glycogen, pullulan,
glucose, fructose, maltose, xylose,
native feathers		 0.0–4.0/
0.4		 S0;
Thio		 AA, CO2, ALA, Glu, EtOH, But, H2, BuOH [79]
Fervidobacterium islandicum Icelandic
Hot spring		 50–80/
65		 6.0–8.0/
7.2		 1.0–4.0
by 0.6		 pyruvate, ribose, glucose, maltose,
raffinose, starch, cellulose		 0.0–1.0/
0.2		 S0;
Thio		 LA, AA, H2, EtOH, CO2, iBut, iVal [80]
Fervidobacterium riparium Hot spring, Russia 46–80/
65		 5.7–7.9/
7.8		 1.0–3.0
by 0.4–0.5		 peptone, YE, pyruvate, glucose, xylose,
fructose, maltose, sucrose, cellobiose,
starch, xylan, CMC, cellulose, filter paper		 0.0–1.0/
0.0		 S0 H2, AA, CO2, PPA, iBut, But [81]
Fervidobacterium gondwanense Hot spring, Australia 45–80/
65–68		 5.5–8.5/
7.0		 4.0–40.0
by 0.5–0.6		 cellobiose, amylopectin, maltose, starch,
dextrin, xylose, glucose, pyruvate,
lactose, fructose, mannose, CMC, galactose		 0.0–0.6/
0.1		 S0 EtOH, AA, LA, CO2, H2 [82]
Fervidobacterium thailandese Hot spring, Thailand 60–88/
78–80		 6.5–8.5/
7.5		 1.1–2.5
by 0.5–0.6		 glucose, maltose, sucrose, fructose,
cellobiose, CMC, cellulose, starch		 <0.5/0.5 S0 n.d. [83]
Fervidobacterium changbaicum Hot spring, China 55–90/
75–80		 6.3–8.5/
7.5		 1.0–8.0
by 0.5–0.6		 glucose, lactose, fructose, sucrose, maltose, starch, sorbitol, cellobiose, trehalose,
galactose, melibiose, pyruvate, glycerin		 0.0–1.0/
0.0		 S0 n.d. [84]
Thermosipho Thermosipho
africanus 		Hot spring,
Africa		 53–77/
75		 6.0–8.0/
7.2		 3.0–4.0
by 0.5		 glucose, ribose, maltose, starch,
galactose, fructose, sucrose 		0.11–3.6 S0;
Thio		 AA, H2, CO2,
EtOH, LA		 [85]
Thermosipho
japonicus 		Hydrothermal vent, Japan 45–80/
72		 5.3–9.3/
7.2–7.6		 3.0–4.0
by 0.5		 YE, peptone, and tryptone, maltose, glucose, galactose, starch, sacharose, ribose, casein 0.7–7.9/
4.0		 S0;
Thio		 n.d. [86]
Thermosipho
geolei 		Oil reservoir, Russia 45–75/
70		 6.0–9.4/
7.5		 2.0–3.0
by 0.4–0.6		 Glucose, peptone, beef extract, YE 0.5–7.0/
2.0–3.0		 S0 H2, AA, ALA, CO2, iVal [87]
Thermosipho Thermosipho
affectus 		Hydrothermal vent,
Atlantic Ocean		 37–75/
70		 5.6–8.2/
6.6		 1.2–6.0
by 0.4–0.9		 YE, beef extract, glucose, maltose, sucrose, starch, dextrin, CMC, cellulose 1.0–5.5/
2.0		 S0 AA, H2,
CO2, EtOH		 [88]
Thermosipho
globiformans 		Hydrothermal vent 40–75/
68		 5.0–8.2/
6.8		 2.0–4.0
by 0.5		 YE, tryptone, starch 0.2–5.2/
2.5		 S0;
Fe2O3 		n.d. [89]
Thermosipho
melanesiensis 		Hydrothermal vent,
Pacific Ocean		 50–75/
70		 4.5–8.5/
6.5–7.5		 1.0–3.5
by 0.4–0.6		 BHI, malt extract, tryptone, sucrose, starch, glucose, maltose, lactose, cellobiose, galactose 1.0–6.0/
3.0		 S0 H2, AA,
ALA, CO2 		[90]
Thermosipho
activus 		Riftia sheath, Guaymas
Basin		 44–75/
65		 5.5–8.0/
6.0		 1.5–10.0
by 0.3–0.8		 glucose, maltose, cellobiose, cellulose,
filter paper, chitin, xylan, pectin, xanthan gum, YE, beef extract, tryptone, casein,
keratin, arabinose, xylose, gelatin		 0.3–6.0/
2.5		 S0,
Fe (III)		 AA, H2,
CO2 		[91]
Thermosipho
atlanticus 		Hydrothermal vent,
Atlantic Ocean		 45–80/
65		 5.0–9.0/
6.0		 1.0–2.6
by 0.2–0.6		 cellobiose, xylose, starch, LA, maltose,
mannose, trehalose, lactose, arabinose,
galactose, mannitol, peptone,
casamino acids, gelatin, BHI, YE, glucose		 1.5–4.6/
2.3		 S0,
Thio,
Cys		 AA, iVal, H2, Gly, ALA, Pro [92]
Geotoga Geotoga
subterranea 		Oilfields,
USA		 30–60/
45		 5.5–9.0/
6.5		 4.0– 7.5
by 0.5		 mannose, starch, maltodextrins, glucose,
lactose, sucrose, galactose, maltose		 0.5-10/
4.0		 S0 H2, CO2,
AA, EtOH		 [10]
Geotoga
petraea 		Oilfields,
USA		 30–55/
50		 5.5–9.0/
6.5		 3.0– 20.0
by 0.6		 mannose, starch, maltodextrins, glucose,
lactose, sucrose, galactose, maltose		 0.5–10/
3.0		 S0 H2, CO2,
AA, EtOH		 [10]
Petrotoga Petrotoga
miotherma 		Oilfields,
USA		 35–65/
55		 5.5–9.0/
6.5		 2.0– 7.5
by 0.6		 mannose, starch, maltodextrins, glucose,
lactose, sucrose, galactose, maltose,
maltodexstrins, xylose		 0.5–10/
2.0		 S0 H2, CO2,
AA, EtOH		 [10]
Petrotoga
olearia 		Oil reservoir, Russia 37–60/
55		 6.5–8.5/
7.5		 0.9–2.5
by 0.3–0.6		 arabinose, xylose, cellobiose, dextrin,
sucrose, glucose, fructose, maltose, ribose, trehalose, xylan, pyruvate, peptone, starch		 0.5–8.0/
2.0		 S0 H2, AA, LA, ALA, EtOH [93]
Petrotoga
sibirica 		Oil reservoir, Russia 37–55/
55		 6.5–9.4/
8.0		 0.9–2.5
by 0.3–0.6		 sucrose, glucose, fructose, maltose, ribose, trehalose, xylan, pyruvate, peptone, galactose 0.5–7.0/
1.0		 S0 H2, AA, LA, ALA, EtOH [93]
Petrotoga Petrotoga
mobilis 		Oilfield,
North Sea		 40–65/
58–60		 5.5–8.5/
6.5–7.0		 1.0–50.0
by 0.5–1.5		 starch, xylan, maltodextrin, maltose,
cellobiose, sucrose, lactose, glucose, galactose, fructose, arabinose, xylose, ribose, rhamnose		 0.5–9.0/
3.0–4.0		 S0,
Thio		 H2, CO2,
AA, EtOH		 [94]
Petrotoga
halophila 		Offshore oil,
Africa		 45–65/
60		 5.6–7.8/
6.7–7.2		 2.0–45.0
by 0.5–0.7		 arabinose, cellobiose, fructose, galactose,
glucose, lactose, maltose, rhamnose, ribose, starch, sucrose, xylose, xylan, pyruvate		 0.5–9.0/
4.0–6.0		 S0 AA, LA,
ALA, H2, CO2 		[95]
Petrotoga
mexicana 		Offshore oil,
Africa		 25–65/
55		 5.8–8.5/
6.6		 1.0–30.0
by 0.5–0.7		 arabinose, cellobiose, fructose, galactose,
glucose, lactose, maltose, mannose, raffinose, rhamnose, ribose, starch, sucrose, xylose,
xylan, pyruvate.		 1.0–20.0/
3.0		 S0,
Thio,
Sulfite		 AA, LA, H2, CO2, ALA [96]
Petrotoga
japonica 		Oil reservoir, Japan 40–65/
60		 6.0–9.0/
7.5		 2.5–7.0
by 0.25–0.75		 starch, xylan, maltose, cellobiose, sucrose, lactose, glucose, galactose, fructose, casamino acids, mannose, arabinose, xylose, ribose 0.5–9.0/
0.5–1.0		 S0,
Thio		 AA, H2,
CO2, ALA		 [97]
Marinitoga Marinitoga
piezophila 		Hydrothermal chimney,
Pacific Ocean		 45–70/
65		 5.0–8.0/
6.0		 1.0–1.5
by 0.5		 starch, fructose, glucose, galactose,
maltose, cellobiose, ribose, acetate		 1.0–5.0/
3.0		 S0,
Thio,
Cys		 n.d. [98]
Marinitoga
litoralis 		Hot spring,
Indian Ocean		 45–70/
65		 5.5–7.5/
6.0		 1.0–7.0
by 0.8–1.0		 cellobiose, galactose, glucose, glycogen,
lactose, maltose, ribose, starch, BHI, casamino acids, casein, peptone, pyruvate, tryptone, YE		 0.8–4.6/
2.6		 S0 n.d. [99]
Marinitoga
okinawensis 		Hydrothermal field, Okinawa 30–70/
55–60		 5.5–7.4/
5.5–5.8		 1.5–5.0
by 0.5–0.8		 YE, tryptone, peptone, starch, glucose,
glycerol		 1.0–5.5/
3.0–3.5		 S0,
Cys		 n.d. [100]
Marinitoga
hydrogenitolerans 		Hydrothermal chimney,
Atlantic Ocean		 35–65/
60		 4.5–8.5/
6.0		 1.5–5.0
by 0.5–0.8		 glucose, starch, glycogen, chitin, YE,
BHI, peptone, casein, pyruvate, maltose		 1.0–6.5/
3.0–4.0		 S0,
Thio,
Cys		 AA, EtOH, Fo, H2, CO2 [101]
Marinitoga
artica 		Hydrothermal chimney,
Norwegian		 45–70/
65		 5.0–7.5/
5.5		 1.0–5.0
by 0.5–0.8		 glucose, trehalose, maltose, sucrose,
maltodextrin, starch, pectin, meat extract, tryptone, YE, pyruvate, fructose,
mannose, cellobiose, cellulose, peptone		 1.5–5.5/
2.5		 S0,
Cys		 n.d. [102]
Marinitoga
camini 		Hydrothermal chimney,
Atlantic Ridge		 25–65/
55		 5.0–9.0/
7.0		 2.0–3.0
by 0.5–1.0		 BHI, gluten, peptone, tryptone, pyruvate, glucose, fructose, maltose, cellobiose,
sucrose, starch, cellulose, CMC, pectin, chitin		 1.0–4.5/
2.0		 S0,
Cys		 AA, iBut, iVal, H2, 3-IAA, LA CO2, HPA, PA [11]
Oceanotoga Oceanotoga
teriensis 		Offshore oil,
India		 25–70/
55– 58		 5.5–9.0/
7.5		 1.5–1.7 by 0.5–0.7 glucose, fructose, cellobiose, arabinose,
raffinose, rhamnose, sucrose, xylose,
ribose, starch, EtOH, formate, acetate,
BHI, YE, bio–trypticase		 0.0–12/
4.3		 S0,
Thio		 AA, H2,
CO2, EtOH		 [12]
Defluviitoga Defluviitog
tunisiensis 		Mesothermic digester 37–65/
55		 6.7–7.9/
6.9		 3.0–30.0
by 1.0		 arabinose, cellobiose, fructose, galactose,
glucose, lactose, maltose, mannose, raffinose, ribose, sucrose, xylose, cellulose, xylan		 0.2–3.0/
0.5		 S0,
Thio		 AA, H2,
CO2 		[9]
Mesotoga Mesotoga
infera 		Deep aquifer, France 30–50/
45		 6.2–7.9/
7.4		 2.0–4.0
by 1.0–2.0		 arabinose, cellobiose, fructose, galactose,
glucose, lactose, LA, mannose, maltose,
raffinose, ribose, sucrose, xylose		 0.0–1.5/
0.2		 S0 AA, CO2 [26]
Mesotoga
prima 		Sediment, USA 20–50/
37		 6.5–8.0/
7.5		 1.0 by 0.2 xylose, fructose, ribose, sucrose, mannose, galactose, maltose, lactose, peptone,
tryptone, casamino acids, glucose,
arabinose, cellobiose, casein, pyruvate		 2.0–6.0/
4.0		 S0,
Thio,
Sulfite		 AA, But, iBut, iVal, 2–MeBu [8]
Kosmotoga Kosmotoga
arenicorallina 		Hot spring,
Japan		 50–65/
60		 6.2–8.0/
7.1		 1.1–2.7
by 1.1–1.9		 xylose, maltose, glycerol 1.0–6.0/
3.0		 S0,
Cys		 n.d. [103]
Kosmotoga
pacifica 		Hydrothermal field, Pacific Ocean 33–78/
70		 6.2–8.0/
7.1		 1.0 by 0.6 maltose, YE, peptone, BHI, glycerol,
tryptone, xylose, glucose, fructose, cellobiose, trehalose, LA, propionate, glutamate		 0.5–6.0/
n.d.		 S0,
Cys		 n.d. [104]
Kosmotoga
olearia 		Fluid,
North Sea		 20–80/
65		 5.5–8.0/
6.8		 0.8–1.2
by 0.4–0.7		 maltose, ribose, sucrose, starch,
casamino acids, tryptone, pyruvate		 1.0–6.0/
2.5–3.0		 Thio H2, CO2, AA, EtOH, PPA [7]
Kosmotoga
shengliensis 		Oilfield,
China		 45–75/
65		 6.0–8.0/
7.0		 0.7–0.9 glucose, acetate, mEtOH, galactose, fructose, xylose, sucrose, maltose, sorbitol, lactose,
xylan, arabinose, formate, rhamnose,
glycerol, pyruvate, starch, LA		 0.0–4.0/
1.5		 S0,
Thio,
Sulfate		 AA, LA,
ALA, CO2,
H2 		[15]
Athalassatoga Athalassatoga
saccharophila 		Hot spring,
Japan		 30–60/
55		 4.5–7.5/
5.5–6.0		 0.8–2.0
by 0.7–0.8		 arabinose, fructose, glucose, lactose,
maltose, mannose, ribose, sucrose,
xylose, starch, glycogen, peptone, YE		 <1/0.0 Fe (III), Thio,
Cys		 AA, iBut,
iVal		 [14]
Mesoaciditoga Mesoaciditoga
lauensis 		Hydrothermal vent,
Pacific Ocean		 45–65/
57–60		 4.1–6.0/
5.5–5.7		 0.8–1.0
by 0.4		 YE, peptone, maltose, sucrose, glucose, xylose, ribose, starch, tryptone 0.5–6.0/
3.0		 S0;
Thio,
Cys		 n.d. [13]