4.A.2 The PTS Fructose-Mannitol (Fru) Family
The Fru family is a large and complex family which includes several sequenced fructose, mannose and mannitol-specific porters as well as several putative PTS porters of unknown specificities. The fructose porters of this family phosphorylate fructose on the 1-position. Those of family 4.6 phosphorylate fructose on the 6-position. As is true of other members of the PTS-GFL superfamily, the IIC domains of these permeases probably have a uniform 10 TMS topology (Vastermark and Saier 2016; McCoy et al. 2016; Cao et al. 2011).
The IIA, IIB and IIC domains of the fructose- and mannitol-specific porters are demonstrably homologous. The IIB and IIC domains of the fructose porters are only distantly related to the corresponding domains of the mannitol porters. The IIB and IIC domains of these porters are homologous to those of the Glc family (TC #4.A.1) (Chang et al., 2004). However, the structure of the IIA domain of the mannitol porter of E. coli has been determined, and it proved to possess an α2β2α3 secondary structure, a structure which is very different from the β-sandwich structure of IIAGlc. Further, the IIC domains of the mannitol and fructose porters are almost as dissimilar from each other as they are from the glucose (TC #4.A.1) or lactose (TC #4.A.3) families.
This family belongs to the PTS-GFL Superfamily.
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|4.A.2.1.1||Fructose porter (FruAB) (fructose-1-P forming) ||Bacteria ||Fructose IIB'BC-IIAMH complex of E. coli |
The FrwABCD putative transporter of unknown function. FruA is a 3-domain multiphosphoryl transfer protein: EIAni-HPr-IIAFru (Reizer et al., 1995).
FrwABCD of E. coli
FrwA (IIA) (P32670)
FrwB (IIB) (P69816)
FrwC (IIC) (P32672)
FrwD (IIB) (P32676)
The FryABC putative transporter of unknown function. FryA is a 3-domain multiphosphoryl transfer protein: EI-HPr-IIAFru (Reizer et al., 1995).
FryABC of E. coli
FryA (IIA) (P77439)
FryB (IIB) (P69808)
FryC (IIC) (P77579)
The mannitol/glucitol transporter, MtlA (IICBAMtl) (Kumar et al., 2011)
MtlA (IICBA) of Vibrio cholerae (Q9KKQ7)
The fructose-specific PTS Enzyme IIABC FruA (Araki et al., 2011).
FruA of Rhodococcus jostii (Q0S1N2)
Fructose Enzyme II complex (IIAFru - IIBFru - IICFru) (based on homology)
IIABCFru of Haloterrigena turkmenica
Fructose-specific PTS, PtfABC (functions with 8.A.7.1.4 and 8.A.8.1.4; Pickl et al., 2012). The transcriptional regulation of the fructose PTS in the very similar organism, Haloferax
mediterranei, has shown that GlpR is a transcriptional activator (Cai et al. 2014).
Fructose Enzyme II, PtfABC (IIABC) complex of Haloferax volcanii
PtfA (IIA) (D4GYE1)
PtfB (IIB) (D4GYE4)
PtfC (IIC) (D4GYE5)
Fructose-specific Enzyme IIABC (Gaurivaud et al. 2000).
Fructose IIABC of Spiroplasma citri
Fructose-specific PTS permease, FruIIBC/FruI-HPr-IIA (Johnson et al. 2008).
FruIIBC/FruI-HPr-IIA of Pseudomonas aeruginosa
Fructose PTS Enzyme IIBC, FruA. The gene encoding the IIA protein is an inactive pseudogene, and fructose appears to be phosphorylated by an ATP-dependent mechanism (Lee et al. 2012).
FruA of Deinococcus radiodurans
The tagatose-specific PTS transporter/kinase, TagIIA-TPr/TagIIB'BC (tagatose-1-P forming) (Shakeri-Garakani et al. 2004). TagIIA-TPr is a fusion of a IIA domain fused N-terminal to an HPr domain. TagIIB'BC has and inactive IIB' domain fused N-terminal to the active C-terminal IIBC domains. This arrangement resembles that for the E. coli fructose Enzyme II complex.
Tag PTS of Klebsiella pneumoniae
Mannitol porter (MtlA) (mannitol-1-P forming), the mannitol IICBA complex. The enzyme-transporter has been alterred genetically, sequenced, purified, reconstituted and characterized (Jacobson et al. 1983, Leonard and Saier 1983, Lee and Saier 1983, Manayan et al. 1988). Intramolecular phosphoryl transfer between the A and B domains of IIMtl is rate-limited by chemistry and not by the rate of formation or dissociation of a stereospecific complex in
which the active sites are optimally apposed (Suh et al. 2007). Substrates, in addition to D-mannitol, include D-glucitol (D-sorbitol), D-2-amino-2-deoxymannitol, D-2-deoxymannitol and D-arabitol (D-arabinitol) (Jacobson et al. 1983).
Mannitol IICBA complex of E. coli
Chromosomal fructose Enzyme IIABC (Fru1) of 654 aas; in an operon with fructose-1-P kinase (Patron et al. 2015).
Fru1 of Streptococcus agalactiae
Putative fructose Enzyme II complex, Fru3; IIA (148 aas)/IIBC (464 aas) (Richards et al. 2011).
IIA/IIBC of Streptococcus agalactiae
D-allose/D-ribose transporting Enzyme II complex (Fru2; IIA/IIB/IIC) (Patron et al. 2017). This system is similar to Frz of E. coli (TC#4.A.2.1.9) which is involved in environmental sensing, host adaptation and virulence (Patron et al. 2015). The regulatory mechanism has been studied (Patron et al. 2017).
D-Ribose and D-allose transporting Enzyme II complex of Streptococcus agalactiae
IIA, 149 aas
IIB, 103 aas
IIC, 367 aas
The tagatose-1-P-forming tagatose phosphorylating Enzyme IIA/IIBC, TagM/L (Van der Heiden et al. 2015). The product is phosphorylated by tagatose-1-P kinase (TagK), and then cleaved by tagatose-1,6-bisphosphate aldolase (GatY).
TagLK of Bacillus licheniformis
TagL (IIBCTag), 466 aas
TagM (IIATag), 152 aas
Cryptic mannitol permease, CmtA (IICB; 462 aas; 9 - 10 TMSs) - CmtB (IIA; 147 aas.)
CmtA-CmtB of E. coli
Fructose-like PTS Enzyme II complex, FrvA (IIA of 148 aas) - FrvB (IIBC of 483 aas and 9 predicted TMSs) (Reizer et al. 1994).
FrvIIa/IIBC of E. coli
Fructose-specific Enzyme I-HPr-Enzyme IIABC complex, all encoded within a single operon with genes in the order: ptsC (IIC), ptsA (IIA), ptsH (HPr), ptsI (Enzyme I) and ptsB (IIB) (Comas et al. 2008).
Fructose Enzyme II complex including EI and HPr of Haloarcula marismortui
The 2-O-α-mannosyl D-glycerate porter (2-O-α-mannosyl D-glycerate-6-P forming), MngA (HrsA) (Sampaio et al., 2004). The phosphorylated product is hydrolyzed to manose-6-P and glycerate by MngB, an α-mannosidase.
2-O-α-mannosyl D-glycerate IIABC complex of E. coli
The fructose porter, FruA (fructose-1-P forming IIABC) (Delobbe et al. 1975) FruA is 39% identical to 4.A.2.1.1). fructose can be metabolized to Fru-1-P via this system as well as Fru-6-P by another PTS system (Gay and Delobbe 1977).
Fructose IIABC of Bacillus subtilis (gi2633811)
The mannitol porter (MtlA) (mannitol-1-P forming), MtlAF. The system is encoded by the mannitol catabolic operon, mtlAFD, and is regulated by the transcription factor, MtlR (Joyet et al. 2015). MtlR contains an N-terminal helix-turn-helix motif followed by an Mga-like domain, two PTS regulatory domains (PRDs), an Enzyme IIBGat-like domain and an Enzyme IIAMtl-like domain, the last four of which can be phosphorylated by the PTS. MtlR proteins are also found in Geobacillus stearothermophilus and Lactobacillus casei, but the mechanisms of their action are different (Joyet et al. 2015). The dephosphorlyated form of the protein activates transcription (Heravi and Altenbuchner 2014).
Mannitol IICB/A (MtlA/F) of Bacillus subtilis (P42956)
The mannose porter (ManP) (37% identical to 4.A.2.1.1). It is encoded in an operon with 3 genes: manP-manA-yjdD, where manP codes for the IIBCA mannose transporter, manA codes for a mannose-6-P isomerase and YjdD codes for a 5-formyltetrahydrofolate cyclo-ligase, characterized in B. anthrasis. Expression of the operon is regulated by ManP and ManR, an activator, in response to external mannose (Sun and Altenbuchner 2010).
Mannose IIBCA of Bacillus subtilis (gi2633555)
|4.A.2.1.7||The fructose inducible fructose/xylitol porter, FruI (Benchabane et al., 2002; Tanzer et al., 2006; Wen et al., 2001)||Bacteria||FruI (IIABC) of Streptococcus mutans (DAA01814)|
|4.A.2.1.8||The constitutive fructose porter FruC/FruD (Benchabane et al., 2002; Tanzer et al., 2006; Wen et al., 2001)||Bacteria||FruC/D (IIBC/IIA) of Streptococcus mutans |
FruC (IIBC) (AAN57895)
FruD (IIA) (DAA01808)
The FrzABC PTS putative transporter (promotes bacterial fitness under stress conditions and promotes fimbrial (fim) gene expression indirectly (Rouquet et al., 2009). Might transport D-tagatose, D-psicose and/or D-sorbose, or a disaccharide of these (Rouquet et al. 2009); involved in environmental sensing, host adaptation and virulence (Patron et al. 2015).
FrzABC of E. coli
FrzA (IIA) (Q1R4S9)
FrzB (IIB) (Q8FC73)
FrzC (IIC) (Q1R4T1)