Chapter 11 : Carbohydrate Catabolism: Pathways, Enzymes, Genetic Regulation, and Evolution

MyBook is a cheap paperback edition of the original book and will be sold at uniform, low price.

Preview this chapter:
Zoom in

Carbohydrate Catabolism: Pathways, Enzymes, Genetic Regulation, and Evolution, Page 1 of 2

| /docserver/preview/fulltext/10.1128/9781555818388/9781555810535_Chap11-1.gif /docserver/preview/fulltext/10.1128/9781555818388/9781555810535_Chap11-2.gif


Regulation of enzyme (levanase) and of an extra cellular fructosyltransferase (levansucrase [Lvs]) are reviewed in this chapter. Carbohydrate catabolism in other gram-positive bacteria is in most cases less well documented than that in . Catabolism of L-arabinose, D-xylose, sucrose, glycerol, and gluconate and the corresponding regulons are described in this chapter. Several gram-positive bacteria are able to catabolize lactose by means of diverse pathways. It has been known for decades that catabolic pathways are conserved elements in prokaryotes and more generally throughout all groups of living organisms. Three unique carbohydrate pathways in gram-positive bacteria have been highlighted here: pathways for glucitol and fructose utilization in and the tagatose-phosphate pathway for lactose-galactose degradation in and some lactic acid bacteria. The tagatose-phosphate pathway is a radical alternative to the otherwise ubiquitous Leloir galactose pathway. It is not known which of these pathways is t h e more recent innovation. species and have functional Leloir pathways , and other grampositive bacteria, including and , appear t o possess elements of this pathway. The diversity is such that it is impossible to define a typically gram-positive pattern of carbohydrate catabolism.

Citation: Steinmetz M. 1993. Carbohydrate Catabolism: Pathways, Enzymes, Genetic Regulation, and Evolution, p 157-170. In Sonenshein A, Hoch J, Losick R (ed), and Other Gram-Positive Bacteria. ASM Press, Washington, DC. doi: 10.1128/9781555818388.ch11

Key Concept Ranking

Transcription Start Site
Lactic Acid Bacteria
Highlighted Text: Show | Hide
Loading full text...

Full text loading...


Image of Figure 1
Figure 1

Catabolism of hexoses, linear hexitols, and sucrose in Transport and enzymatic steps are symbolized by arrows. The names of the structural genes (in some cases putative) of the relevant proteins are indicated in italics and ). All these genes are both mapped (except for phosphoglucoisomerase) and inducible (except and ). Various controls prevent (or reduce) simultaneous expression of alternative pathways for sucrose and fructose (see text). Transfructosylation from sucrose by Lvs results mainly in a mixture of glucose, fructose, and levan. encodes levanase. Fra, Glu, Gut, Md, and Suc: fructose, glucose, glucitol, mannitol, and sucrose, respectively; Suc-P, sucrose-phosphate (phosphorylated on position 6 of glucose moiety). Other sugar-phosphates are designated according to standard conventions; for example, Glu-6-P and Fru-l,6-dP designate glucose 6-phosphate and fructose 1,6-diphosphate.

Citation: Steinmetz M. 1993. Carbohydrate Catabolism: Pathways, Enzymes, Genetic Regulation, and Evolution, p 157-170. In Sonenshein A, Hoch J, Losick R (ed), and Other Gram-Positive Bacteria. ASM Press, Washington, DC. doi: 10.1128/9781555818388.ch11
Permissions and Reprints Request Permissions
Download as Powerpoint
Image of Figure 2
Figure 2

Organization of the gluconate operon and the xyloside-xylose regulon in The repressor of the operon is encoded by the first gene of the operon; and -Z very likely encode gluconate kinase, gluconate permease, and 6-phosphogluconate dehydrogenase, respectively; is transcribed from both the GntR-repressed promoter and a constitutive promoter within The repressor of the xylose-xyloside regulon (XylR) controls transcription of the and opérons; XynB is a xylosidase; XynC appears to be a membrane protein that might be a permease of xyloside and/or xylose; and and encode xylose isomerase and xylulose kinase, respectively. t, terminator.

Citation: Steinmetz M. 1993. Carbohydrate Catabolism: Pathways, Enzymes, Genetic Regulation, and Evolution, p 157-170. In Sonenshein A, Hoch J, Losick R (ed), and Other Gram-Positive Bacteria. ASM Press, Washington, DC. doi: 10.1128/9781555818388.ch11
Permissions and Reprints Request Permissions
Download as Powerpoint
Image of Figure 4
Figure 4

Regulons dependent on antiterminators and/or PTS control: comparison of regulation and genetic organization. The first three systems (a, b, and c) involve elements both structurally and functionally homologous: antiterminators, conditional terminators (ct), and enzymes II. (a) β-glucoside operon model. In the absence of β-glucoside inducer, transcription of the operon from the promoter is prevented by two conditional terminators bracketing the antiterminator gene. In these conditions, BglG function is inhibited by phosphorylation by the gene product (enzyme II). The presence of β-glucoside in the medium results in its transport and phosphorylation by enzyme II. This results in phosphate group rerouting. Dephosphorylated antiterminator then allows full transcription of the operon. BglB is the phospho-β-glucosidase ( ). (b) (Lvs) regulon. This regulon comprises the gene and the unlinked regulator operon. Both appear to be controlled by means of a regulatory cascade similar to that of the operon, i.e., positively by both sucrose and SacY antiterminator and negatively by SacX, a putative enzyme II. Several regulatory cross-talks with the regulon (see below) are not shown. The genes (DEG; see chapter 50) activate transcription from both and promoters ( ). (c) operon. Transcription of this operon is positively controlled by sucrose and the antiterminator encoded by the linked gene; appears not to play a role in this regulation ( ). The mechanism of SacT activation by the presence of sucrose is unknown (see text), (d) levanase operon. The first four genes encode an enzyme II (very poorly related to those mentioned above) controlling the LevR positive regulator. The fructose-dependent induction cascade appears similar to that of the operon but appears not to involve antitermination. LevR is a transcriptional activator binding upstream from the operon promoter and requiring σ as a cofactor ( ). (e) glycerol regulon. This regulon comprises at least two unlinked gene clusters and three promoters likely controlled by the gene product. This regulator (unrelated to BglG) appears to function as an antiterminator at conditional terminators just upstream from and ( ).

Citation: Steinmetz M. 1993. Carbohydrate Catabolism: Pathways, Enzymes, Genetic Regulation, and Evolution, p 157-170. In Sonenshein A, Hoch J, Losick R (ed), and Other Gram-Positive Bacteria. ASM Press, Washington, DC. doi: 10.1128/9781555818388.ch11
Permissions and Reprints Request Permissions
Download as Powerpoint
Image of Figure 3
Figure 3

Sucrose gene clusters in four bacteria. The comparison includes two gram-positive systems, from and and two gram-negative systems (sucrose operon in a close relative of enteric bacteria, and sucrose genes present on the pUR400 plasmid; the pUR400 sucrose regulon appears to be closely related to the chromosomal sucrose system). The four enzymes nsucrose, encoded by (in ) or genes, are structurally homologous. The ScrA protein contains a C-terminal extension containing a second phosphorylation site (enzyme III domain). The gene designated in or in and encodes related phosphosucrases (the pUR400 gene sequence is unknown). Expression of the sucrose genes is stimulated by the presence of different sugars in the medium (see text). The three other systems are specifically induced by sucrose via activation of the SacT antiterminator in or inactivation of ScrR repressors in the case of the pUR400 or systems. The two ScrR repressors are not strongly similar, but both belong to the LacI-GalR superfamily; the pUR400-encoded ScrR repressor binds fructose; the two gram-negative regulons are dependent on the cyclic-AMP receptor protein-cyclic-AMP complex. The and genes encode fructokinase and a sucrose porin (related to the LamB maltoporin), respectively, encodes unknown functions (probably not involved in sucrose metabolism) (see references , and and references therein). The sucrose genes were recendy cloned and sequenced; the gene products are homologous to those mentioned above. (encoding a short enzyme II) and are unlinked. Both are sucrose inducible via a negative control. Similar operatorlike sequences are found in both leader regions ( ).

Citation: Steinmetz M. 1993. Carbohydrate Catabolism: Pathways, Enzymes, Genetic Regulation, and Evolution, p 157-170. In Sonenshein A, Hoch J, Losick R (ed), and Other Gram-Positive Bacteria. ASM Press, Washington, DC. doi: 10.1128/9781555818388.ch11
Permissions and Reprints Request Permissions
Download as Powerpoint
Image of Figure 5
Figure 5

FTS-dependent lactose systems of and some lactic acid-producing bacteria: pathways and gene organization. (A) Transport and catabolism of lactose via the tagatose 6-phosphate pathway. Lactose is transported and phosphorylated by enzyme II (II) and then hydrolyzed into glucose (catabolized by the classical glycolysis pathway) and Gal-6-P, which is isomerized to tagatose 6-phosphate (Tag-6-P). Tag-6-P is converted, by means of a pathway similar to that of its isomer, fructose 6-phosphate (Fru-6-P), into two molecules of triose-phosphate (dihydroxyacetone-phosphate [DHA-P] and glyceraldehyde 3-phosphate [GA-3-P]) and then into phosphoenolpyru-vate (PEP). Dephosphorylation of PEP to pyruvate results partly in ATP production (not shown) and partly in phosphorylation of proteins of the cytoplasmic PTS cascade. This cascade ends with phosphorylation of the membrane enzyme II protein by the cytoplasmic enzyme III (III) protein. In lactic acid bacteria, lactate produced from pyruvate is excreted with protons; this-coexcretion contributes to energizing the proton motive force. A, H, I, and K, aldolases, hydrolases, isomerases, and kinases, respectively. The genes encoding the enzymes of the pathway are indicated in parentheses. (B) Organization of the opérons in and Arrows indicate the transcripts; t, t1, and t2 are transcriptional terminators. Three major features distinguish the operon from that of the orientation of the gene, the presence of a weak terminator (tl) downstream from and the presence of an eighth gene, of unknown function, at the 3′ end of the operon. The 5′ end of the operon has not yet been sequenced, but its transcript is similar in size to that of (Adapted from references , and )

Citation: Steinmetz M. 1993. Carbohydrate Catabolism: Pathways, Enzymes, Genetic Regulation, and Evolution, p 157-170. In Sonenshein A, Hoch J, Losick R (ed), and Other Gram-Positive Bacteria. ASM Press, Washington, DC. doi: 10.1128/9781555818388.ch11
Permissions and Reprints Request Permissions
Download as Powerpoint


1. Adams, C. V.,, J. A. Fornwald,, F. J. Schmidt,, M. Rosenberg,, and M. E. Brawner. 1988. Gene organization and structure of the Streptomyces lividans gal operon. J. Bacteriol. 170:203212.
2. Alpert, C. A.,, and B. M. Chassy. 1990. Molecular cloning and DNA sequence of lacE, the gene encoding the lactose-specific enzyme II of the phosphotransferase system of Lactobacillus casei. J. Biol. Chem. 265:2256122568.
3. Amster-Choder, O.,, F. Houman,, and A. Wright 1989. Protein phosphorylation regulates transcription of the β-glucoside utilization operon in E. coli. Cell 58:847855.
4. Asladinis, C.,, K. Schmid,, and K. Schmitt. 1989. Nucleotide sequences and operon structure of plasmid-borne genes mediating uptake and utilization of raffinose in Escherichia coli. J. Bacteriol. 171:67536763.
5. Aymerich, S.,, and M. Stelnmetz. 1992. Specificity determinants and structural features in the RNA target of the bacterial antiterminator proteins of the BglG/SacY family. Proc. Natl. Acad. Sci. USA 89:1041010414.
6. Bezzate, S.,, M. Steinmetz,, and S. Aymerlch. Unpublished data.
7. Blatch, G. L.,, and D. R. Woods. 1991. Nucleotide sequence and analysis of the Vibrio alginolyticus scrR repressor-encoding gene (scrR). Gene 101:1723.
8. Burchhardt, G.,, and H. Bahl. 1991. Cloning and analysis of the β-galactosidase-encoding gene from Clostridium thermosulfurogenes EMI. Gene 10.6:1319.
9. Cal, Y. 1991. Characterization of insertion sequences IS892 and related elements from the cyanobacterium Anabaena sp. strain PCC 7120. J. Bacteriol. 173:57715786.
10. Chamberí, R.,, and G. Gonzy-Tréboul. 1976. Levansucrase of B. subtilis: kinetic and thermodynamic aspects of the transfructosylation process. Eur. J. Biochem. 62:5564.
11. Charles, T. C.,, and T. M. Finan. 1991. Analysis of a 1600-kilobase Rhizobium meliloti megaplasmid using defined deletions generated in vivo. Genetics 127:520.
12. Chassy, B., 1983. Sucrose metabolism and glycosyltrans-ferase activity in oral streptococci, p. 310. In R. J. Doyle, and J. E. Ciardi (ed), Glucosyltransferases, Glucans, Sucrose, and Dental Caries. IRL Press, Washington, D.C..
13. Crutz, A. M.,, and M. Steinmetz. 1992. Transcription of the Bacillus subtilis sacX and sacY genes, encoding regulators of sucrose metabolism, is both inducible by sucrose and controlled by the DegS-DegU signalling system. J. Bacteriol. 174:60876095.
14. Crutz, A. M.,, M. Steinmetz,, S. Aymerlch,, R. Richter,, and D. Le Coq. 1990. Induction of levansucrase in Bacillus subtilis: an antitermination mechanism negatively controlled by the phosphotransferase system. J. Bacteriol. 172:10431050.
15. Débarbouillé, M.,, A. Fouet,, M. Arnaud,, A. Klier,, and G. Rapoport. 1990. The sacT gene regulating the sacPA operon in Bacillus subtilis shares strong homology with transcriptional antiteraiinators. J. Bacteriol. 172:39663973.
16. Débarbouillé, M.,, I. Martin-Verstraete,, A. Klier,, and G. Rapoport. 1991. The transcriptional regulator LevR of Bacillus subtilis has domains homologous to both σ54-and phosphotransferase system-dependent regulators. Proc. Natl. Acad. Sci. USA 88:22122216.
17. Débarbouillé, M.,, I. Martin-Verstraete,, A. Klier,, and G. Rapoport. 1991. The Bacillus subtilis sigL gene encodes an equivalent of o54 from gram-negative bacteria. Proc. Natl. Acad. Sci. USA 88:90929096.
18. Dekker, K.,, H. Yamagata,, K. Sakaguchl,, and S. Ukada. 1991. Xylose (glucose) isomerase gene from the thermo-phile Thermus thermophilus: cloning, sequencing, and comparison with other thermostable xylose isomerases. J. Bacteriol. 173:30783083.
19. De Vos, W. M., 1991. Disaccharide utilization in lactic acid bacteria, p. 447457. In H. Heslot,, J. Davies,, J. Florent,, L. Bobichon,, G. Durand,, and L. Penasse (ed.). Proceedings of the 6th International Symposium on the Genetics of Industrial Microorganisms. Société Française de Microbiologie, Paris.
20. Errington, J.,, and C. Vogt. 1990. Isolation and characterization of mutations in the gene encoding an endogenous Bacillus subtilis β-galactosidase and its regulator. J. Bacteriol. 172:488490.
21. Fischer, R.,, R. Eisermann,, B. Reiche,, and W. Hengstenberg. 1989. Cloning, sequencing, and overexpression of the mannitol-specific enzyme-III-encoding gene of Staphylococcus camosus. Gene 82:249257.
22. Fornwald, J. A.,, F. J. Schmidt,, C. W. Adams,, M. Rosenberg,, and M. E. Brawner. 1987. Two promoters, one inducible and one constitutive, control transcription of the Streptomyces lividans galactose operon. Proc. Natl. Acad. Sci. USA 84:21302134.
23. Fouet, A.,, M. Arnaud,, A. Klier,, and G. Rapoport. 1987. Bacillus subtilis sucrose specific enzyme II of the phosphotransferase system. Expression in Escherichia coli and homology to enzymes II from enteric bacteria. Proc. Natl. Acad. Sci. USA 84:87738777.
24. Fouet, A.,, A. Klier,, and G. Rapoport. 1986. Nucleotide sequence of the sucrase gene of Bacillus subtilis. Gene 43:221225.
25. Freeze, E.,, W. Klofat,, and E. Galliërs. 1970. Commitment to sporulation and induction of glucose-phos-phoenolpyruvate-transferase. Biochim. Biophys. Acta 22:265289.
26. Fujita, Y.,, and T. Fujita. 1983. Genetic analysis of a pleiotropic deletion mutation (Δigf) in Bacillus subtilis. J. Bacteriol. 154:864869.
27. Fujita, Y.,, and T. Fujita. 1986. Identification and nucleotide sequence of the promoter region of the Bacillus subtilis gluconate operon. Nucleic Acids Res. 14:12371252.
28. Fujita, Y.,, and T. Fujita. 1987. The gluconate operon gnt of Bacillus subtilis encodes its own transcriptional negative regulator. Proc. Natl. Acad. Sci. USA 84:45244528.
29. Fujita, Y.,, and T. Fujita. 1989. Effect of mutations causing gluconate kinase or gluconate permease deficiency on expression of the Bacillus subtilis gnt operon. J. Bacteriol. 171:17511754.
30. Fujita, Y.,, T. Fujita,, and Y. Miwa. 1990. Evidence for posttranscriptional regulation of synthesis of the Bacillus subtilis Gnt repressor. FEBS Lett. 267:7174.
31. Fujita, Y.,, T. Fujita,, Y. Miwa,, J. Nihashi,, and Y. Aratani. 1986. Organization and transcription of the gluconate operon, gnt, of Bacillus subtilis. J. Biol. Chem. 261:1374413753.
32. Fujita, Y.,, A. Ramaley,, and E. Freeze. 1977. Location and properties of glucose dehydrogenase in sporulating cells and spores of Bacillus subtilis. J. Bacteriol. 132: 282293.
32a.. Fujita, Y.,, K. Shlndo,, Y. Miwa,, and K. Yoshida. 1991. Bacillus subtilis inositol dehydrogenase-encoding gene (idh): sequence and expression in Escherichia coli. Gene 108:121125.
33. Gartner, D.,, M. Geissendörfer,, and W. Hillen. 1988. Expression of the Bacillus subtilis xyl operon is repressed at the level of transcription and is induced by xylose. 7. Bacteriol. 170:31023109.
34. Gay, P. 1979. Ph.D. thesis. Université Paris VI, Paris.
35. Gay, P.,, H. Chalumeau,, and M. Steinmetz. 1983. Chromosomal localization of gut, fruC, and pfk mutations affecting glucitol catabolism in Bacillus subtilis. J. Bacteriol. 153:11331137.
36. Gay, P.,, P. Cordier,, M. Marquet,, and A. Delobbe. 1973. Carbohydrate metabolism and transport in Bacillus subtilis. A study of ctr mutations. Mol. Gen. Genet. 121:355368.
37. Gay, P.,, and A. Delobbe. 1973. Fructose transport in Bacillus subtilis. Eur. J. Biochem. 79:363373.
38. Gay, P.,, D. Le Coq,, M. Steinmetz,, T. Berkelman,, and C. I. Kado. 1985. Positive selection procedure for entrapment of insertion sequence elements in gram-negative bacteria. J. Bacteriol. 164:918921.
39. Giffard, P. M.,, C. L. Simpson,, C. P. Mihvard,, and N. A. Jacques. 1991. Molecular characterization of a cluster of at least two glucosyltransferase genes in Streptococcus salivar-ius ATCC 25975./ Gen. Microbiol. 137:25772593.
40. Glaser, P. (Institut Pasteur, Paris). 1991. Personal communication.
41. Goldman, M.,, and H. J. Blumenthal. 1963. Pathways of glucose in Bacillus subtilis. J. Bacteriol. 86:303311.
42. Gonzales-Candelas, L.,, D. Ramon,, and J. Polaina. 1990. Sequences and homology analysis of two genes encoding β-glucosidases from Bacillus polymyxa. Gene 95:3138.
43. Gonzy-Tréboul, G.,, J. H. De Vaard,, M. Zagorec,, and P. W. Postma. 1991. The glucose permease of the phosphotransferase system of Bacillus subtilis: evidence for IIGlc and IIIGlc domains. Mol. Microbiol. 5:12411249.
44. Hall, B. G.,, P. W. Betts,, and J. C. Wootton. 1989. DNA sequence analysis of artificially evolved ebg enzyme and ebg repressor genes. Genetics 123:635648.
45. Hancock, K. R.,, E. Rockman,, C. A. Young,, L. Pearce,, I. S. Maddox,, and D. B. Scott. 1991. Expression and nucleotide sequence of the Clostridium acetobutylicum β-galactosidase gene cloned in Escherichia coli. J. Bacteriol. 173:30843095.
46. Hastrup, S., 1988. Analysis of the Bacillus subtilis xylose regulon, p. 7983. In A. T. Ganesan, and J. A. Hoch (ed.), Genetics and Biotechnology of Bacilli, vol. 2 Academic Press, Inc., New York.
47. Hodgson, D., Primary metabolism-carbon catabolism. In E. M. Wellington (ed.), Streptomyces, in press. Plenum Biotechnology Handbooks, New York.
48. Holmberg, C.,, L. Beijer,, B. Rutberg,, and L. Rutberg. 1990. Glycerol catabolism in Bacillus subtilis: nucleotide sequence of the genes encoding glycerol kinase (glpK) and glycerol-3-phosphate dehydrogenase (glpD) J. Gen. Microbiol. 136:23672375.
49. Holmberg, C.,, and B. Rutberg. 1991. Expression of the gene encoding glycerol-3-phosphate dehydrogenase (glpD) in Bacillus subtilis is controlled by antitermina-tion. Mol. Microbiol. 5:28912900.
50. Holmberg, C.,, L. Rutberg,, and B. Rutberg (University of Lund). 1991. Personal communication.
51. Houman, F.,, M. R. Diaz-Torres,, and A. Wright. 1990. Transcriptional antitermination in the bgl operon of E. coli is modulated by a specific RNA binding protein. Cell 62:11531163.
51a.. Jäger, W.,, A. Schäfer,, A. Pühler,, G. Labes,, and W. Wohlleben. 1992. Expression of the Bacillus subtilis sacB gene leads to sucrose sensitivity in the gram-positive bacterium Corynebacterium glutamicum but not in Streptomyces lividans. J. Bacteriol. 174:54625465.
52. Kreuzer, P.,, D. Gartner,, R. Allmansberger,, and W. Hillen. 1989. Identification and sequence analysis of the Bacillus subtilis W23 xylR gene and xyl operator. J. Bacteriol. 171:38403845.
53. Kunst, F.,, M. Steinmetz,, J. A. Lepesant,, and R. De-donder. 1977. Presence of a third sucrose hydrolysing enzyme in Bacillus subtilis: constitutive levanase synthesis by mutants of Bacillus subtilis Marburg 168. Biochimie 59:287292.
54. Lawlis, V. B.,, M. S. Dennis,, E. Y. Chen,, D. H. Smith,, and D. J. Henner. 1984. Cloning and sequencing of the xylose isomerase and xylulose kinase genes of Escherichia coli. Appl. Environ. Microbiol. 47:1521.
55. Le Coq, D.,, A. M. Crutz,, R. Richter,, and M. Steinmetz. Unpublished data.
56. Leong-Morgenthaler, P.,, M. C. Zwahlen,, and H. Hot-tlnger. 1991. Lactose metabolism in Lactobacillus bulgaricus: analysis of the primary structure and expression of the genes involved. J. Bacteriol. 173:19511957.
57. Lepesant, J. A.,, F. Kunst,, M. Pascal,, J. Lepesant-Kej-zlarova,, M. Steinmetz,, and R. Dedonder,. 1976. Specific and pleiotropic regulatory mechanisms in the sucrose system of Bacillus subtilis 168, p. 5869. In D. Schles-singer (ed.), Microbiology—1976. American Society for Microbiology, Washington, D.C..
58. Lin, E. C. C., 1987. Dissimilatory pathways for sugars, polyols, and carboxylates, p. 244284. In F. C. Neidhart,, J. L. Ingraham,, K. B. Low,, B. Magasanik,, M. Schaechter,, and H. E. Umbarger (ed.), Escherichia coli and Salmonella typhimurium: Cellular and Molecular Biology, vol. 1. American Society for Microbiology, Washington, D.C..
59. Lin, E. C. C.,, A. J. Hacking,, and J. Aguilar. 1976. Experimental models of acquisitive evolution. BioScience 26:548555.
60. Lindgren, V. 1978. Mapping of a genetic locus that affects glycerol 3-phosphate in Bacillus subtilis. J. Bacteriol. 133:667670.
61. Lindgren, V.,, and L. Rutberg. 1976. Genetic control of the glp system in Bacillus subtilis. J. Bacteriol. 127: 10471057.
62. Loesche, W. L. 1986. Role of Streptococcus mutans in human dental decay. Microbiol. Rev. 50:353380.
62a.. Losick, R. (Harvard University). 1992. Personal communication.
63. Macrina, F. L.,, K. R. Jones,, C. A. Alpert,, B. M. Chassy,, and S. M. Michalek. 1991. Repeated DNA sequence involved in mutations affecting transport of sucrose in Streptococcus mutans V403 via the phosphoenolpyru-vate phosphotransferase system. Infect. Immun. 59: 15351543.
64. Mahadevan, S.,, and A. Wright. 1987. A bacterial gene involved in transcription antitermination: regulation at a Rho-independent terminator in the bgl operon of E. coli. Cell 50:485494.
65. Martin, I.,, M. Debarbouillé,, E. Ferrari,, A. Klier,, and G. Rapoport. 1987. Characterization of the levanase gene of Bacillus subtilis which shows homology to yeast invertase. Mol. Gen. Genet. 208:177184.
66. Martin, I.,, M. Debarbouillé,, A. Klier,, and G. Rapoport. 1989. Induction and metabolite regulation of levanase synthesis in Bacillus subtilis. J. Bacteriol. 171:18851892.
67. Martin-Verstraete, I.,, M. Debarbouillé,, A. Klier,, and G. Rapoport. 1990. Levanase operon of Bacillus subtilis includes a fructose-specific phosphotransferase system regulating the expression of the operon. J. Mol. Biol. 214:657671.
68. Minton, N. P.,, S. P. Chambers,, W. J. Mitchell,, and J. K. Brehm. 1991. Program Abstr. 6th Int. Conf. Bacilli, abstr. Ml..
69. Mollet, B.,, and N. Pllloud. 1991. Galactose utilization in Lactobacillus helveticus: isolation and characterization of the galactokinase (galK) and galactose-1-phosphate uridyl transferase igalT) genes. J. Bacteriol. 173: 44644473.
70. Morse, M. L.,, K. L. Hill,, J. B. Egan,, and W. Hengstenberg. 1968. Metabolism of lactose by Staphylococcus aureus and its genetic basis. J. Bacteriol. 95:22702274.
71. Nègre, D.,, J.-C. Cortay,, I. G. Old,, A. Galinier,, C. Rich-aud,, I. Saint-Girons,, and A. J. Cozzone. 1991. Overproduction and characterization of the iclR gene product of Escherichia coli Kl 2 and comparison with that of Salmonella typhimurium LT2. Gene 97:2937.
72. Oskouian, B.,, and G. C. Stewart. 1990. Repression and catabolite repression of the lactose operon of Staphylococcus aureus. J. Bacteriol. 172:38043812.
73. Parker, L. L.,, and B. G. Hall. 1990. Characterization and nucleotide sequence of the cryptic eel operon of Escherichia coli K12. Genetics 124:455471.
74. Poolman, B.,, T. J. Royer,, S. E. Mainzer,, and B. F. Schmidt. 1989. Lactose transport system of Streptococcus thermophilus: a hybrid protein with homology to the melibiose carrier and enzyme III of phosphoenol-pyruvate-dependent phosphotransferase systems. J. Bacteriol. 171:244253.
75. Poolman, B.,, T. J. Royer,, S. E. Mainzer,, and B. F. Schmidt. 1990. Carbohydrate utilization in Streptococcus thermophilus: characterization of the genes for aldose 1-epimerase (mutarotase) and UDPglucose 4-epi-merase. J. Bacteriol. 172:40374047.
76. Rather, P. N.,, and C. P. Moran, Jr. 1988. Compartment-specific transcription in Bacillus subtilis: identification of the promoter for gdh. J. Bacteriol. 170:50865092.
77. Reizer, A.,, J. Deutscher,, M. H. Saier, Jr.,, and J. Reizer. 1991. Analysis of the gluconate (gnt) operon Bacillus subtilis. Mol. Microbiol. 5:10811089.
78. Reizer, J.,, A. Reizer,, and M. H. Saier, Jr. 1990. The cellobiose permease of Escherichia coli consists of three proteins and is homologous to the lactose permease of Staphylococcus aureus. Res. Microbiol. 141:10611067.
79. Reynolds, A. E.,, J. Felton,, and A. Wright. 1981. Insertion of DNA activates the cryptic bgl operon in Escherichia coli K12. Nature (London) 293:625629.
80. Romantschuk, M.,, G. Y. Richter,, P. Mukhopadhyay,, and D. Mills. 1991. IS801, an insertion sequence element isolated from Pseudomonas syringae pathovar. Mol. Microbiol. 5:617622.
81. Roncero, M. I. G. 1983. Genes controlling xylan utilization by Bacillus subtilis. J. Bacteriol. 156:257263.
82. Rosey, E. L.,, B. Oskouian,, and G. C. Stewart. 1991. Lactose metabolism by Staphylococcus aureus: characterization of lacABCD, the structural genes of the tagatose 6-phosphate pathway. J. Bacteriol. 173:59925998.
83. Saheb, S. A. 1972. Etude de deux mutants du métabolisme du glycerol chez Bacillus subtilis. Can. J. Microbiol. 18:13151325.
84. Sa-Nogueira, I.,, and H. de Lancastre. 1991. Program Abstr. 6th Int. Conf. Bacilli., abstr. T9.
85. Sa-Nogueira, I.,, H. Paveia,, and H. de Lancastre. 1988. Isolation of constitutive mutants for L-arabinose utilization in Bacillus subtilis. J. Bacteriol. 170:28552857.
86. Sato, Y.,, F. Poy,, G. R. Jacobson,, and H. K. Kuramitsu. 1989. Characterization and sequence analysis of the scrA gene encoding enzyme IIScr of the Streptococcus mutans phosphoenolpyruvate-dependent sucrose phosphotransferase system. J. Bacteriol. 171:263271.
87. Sato, Y.,, Y. Yamamoto,, R. Suzuki,, H. Klzaki,, and H. K. Kuramitsu. 1991. Construction of scrA: :lacZ gene fusion to investigate regulation of the sucrose PTS of Streptococcus mutans. FEMS Microbiol. Lett. 79:339346.
88. Scheler, A.,, T. Rygus,, R. Allmansberger,, and W. Hillen. 1991. Molecular cloning, structure, promoters and regulatory elements for transcription of the Bacillus licheniformis encoded regulon for xylose utilization. Arch. Microbiol. 155:526534.
89. Schmid, K.,, R. Ebner,, K. Jahreis,, J. W. Lengeler,, and F. Tigemeyer. 1991. The sugar-specific porin, ScrY, is involved in sucrose uptake in enteric bacteria. Mol. Microbiol. 5:941950.
90. Schmidt, B. F.,, R. M. Adams,, C. Requadt,, S. Power,, and S. E. Malnzer. 1989. Expression and nucleotide sequence of the Lactobacillus bulgaricus β-galactosidase gene cloned in Escherichia coli. J. Bacteriol. 171:625635.
91. Schnetz, K.,, and B. Rak. 1988. Regulation of the bgl operon of Escherichia coli by transcriptional antitermi-nation. EMBO J. 7:32713278.
92. Scholzen, T.,, and E. Arndt. 1991. Organization and nucleotide sequence of ten ribosomal protein genes from the region equivalent to the spectinomycin operon in the archaebacterium Halobacterium marismortui. Mol. Gen. Genet. 228:7080.
93. Schörgendorfer, K.,, H. Scharb,, and R. M. Laflerty. 1988. Molecular characterization of Bacillus subtilis levanase and a C-terminal deleted derivative. J. Biotechnol. 7:247258.
94. Schroeder, C. J.,, C. Robert,, G. Lenzen,, L. L. McKay,, and A. Mercenier. 1991. Analysis of the lacZ sequence from two Streptococcus thermophilus strains: comparison with the Escherichia colt and Lactococcus bulgaricus β-galactosidase sequences. J. Gen. Microbiol. 137:369380.
95. Schroeder, V.,, S. M. Michalek,, and F. L. Macrina. 1989. Biochemical characterization and evaluation of virulence of a fructosyltransferase-deficient mutant of Streptococcus mutans V403. Infect. Immun. 57:35603569.
96. Shamanna, D. K.,, and K. E. Sanderson. 1979. Genetics and regulation of the D-xylose utilization in Salmonella typhimurium LT2. J. Bacteriol. 139:7179.
97. Shazand, K.,, P. Hwang,, J. Tucker,, J. C. Rabinowltz,, T. Leighton,, and M. Grunberg-Manago. 1990. Program. Abstr. Conf. Bacillus subtilis Genome, Paris, abstr. P.66.
98. Shimotsu, H.,, and D. Henner. 1986. Modulation of Bacillus subtilis levansucrase gene expression by suerose and regulation of the steady-state mRNA level by sacU and sacQ genes. J. Bacteriol. 168:380388.
99. Shiroza, T.,, and H. K. Kuramitsu. 1988. Sequence analysis of the Streptococcus mutans fructosyltransferase gene and flanking regions. J. Bacteriol. 170:810816.
100. Slzemore, C.,, E. Buchner,, T. Rygus,, C. Witke,, F. Görz,, and W. Hillen. 1991. Organization, promoter analysis and transcriptional regulation of the Staphylococcus xylosus xylose utilization operon. Mol. Gen. Genet. 227:377384.
101. Smith, C. P. (University of Manchester). 1991. Personal communication.
102. Smith, C. P.,, and K. F. Chater. 1988. Structure and regulation of controlling sequences for the Streptomyces coelicolor glycerol operon. J. Mol. Biol. 204:569580.
103. Steinmetz, M. Unpublished data.
104. Steinmetz, M.,, and S. Aymerich,. 1990. The Bacillus subtilis sac-deg system: how and why?, p. 303311. In M. Zukowski,, A. T. Ganesan,, and J. A. Hoch (ed.), Genetics and Biotechnology of Bacilli, vol. 3. Academic Press, Inc., New York.
105. Steinmetz, M.,, D. Le Coq,, and S. Aymerich. 1989. Induction by sucrose of saccharolytic enzymes in Bacillus subtilis: evidence for two partially interchangeable regulatory pathways. J. Bacteriol. 171:15191523.
106. Steinmetz, M.,, D. Le Coq,, S. Aymerich,, G. Gonzy-Tréboul,, and P. Gay. 1985. The DNA sequence of the gene for the secreted Bacillus subtilis enzyme Ievansu-crase. Mol. Gen. Genet. 200:220228.
107. Steinmetz, M.,, D. Le Coq,, H. Ben Djemia,, and P. Gay. 1983. Analyse génétique de sacB, gène de structure d'une enzyme sécrétée, la lévane-saccharase de Bacillus subtilis. Mol. Gen. Genet. 191:138144.
108. Strauss, N. 1983. Role of glucose dehydrogenase in germination of Bacillus subtilis spores. FEMS Microbiol. Lett. 20:379384.
109. Sutrina, S. L.,, P. Reddy,, M. H. Saier, Jr.,, and J. Reizer. 1990. The glucose permease of Bacillus subtilis is a single polypeptide chain that functions to energize the sucrose permease. J. Biol. Chem. 265:1858118589.
110. Thomas, T. D.,, and V. L. Crow. 1984. Selection of galactose-fermenting Streptococcus thermophilus in lactose-limited chemostat cultures. Appl. Environ. Microbiol. 48:186191.
111. Thompson, J. 1987. Regulation of sugar transport and metabolism in lactic acid bacteria. FEMS Microbiol. Rev. 46:221232.
112. Ueda, S.,, T. Shiroza,, and H. K. Kuramitsu. 1988. Sequence analysis of the gtfC gene from Streptococcus mutans GC5. Gene 69:101109.
113. Van Rooijen, R. J.,, and W. M. De Vos. 1990. Molecular cloning, transcription analysis, and nucleotide sequence of lacR, a gene encoding the repressor of the lactose phosphotransferase system of Lactococcus lactis. J. Biol. Chem. 265:1849918503.
114. Van Rooijen, R. J.,, S. Vanschalkwijk,, and W. M. De Vos. 1991. Molecular cloning, characterization, and nucleotide sequence of the tagatose 6-phosphate pathway gene cluster of the lactose operon of Lactococcus lactis. J. Biol. Chem. 266:71767182.
114a.. Wagner, E.,, and R. Bruckner (University of Tubingen). 1992. Personal communication.
115. Westphellng, J.,, and M. Brawner. 1989. Two transcribing activities are involved in expression of the Streptomyces galactose operon. J. Bacteriol. 171:13551361.
116. Wong, H. C.,, Y. Ting,, H.-C. Lin,, F. Reichert,, K. Myambo,, K. W. K. Watt,, P. T. Toy,, and R. J. Drummond. 1991. Genetic organization and regulation of the xylose degradation genes in Streptomyces rubiginosus. J. Bacteriol. 173:68496858.
117. Zagorec, M.,, and M. Steinmetz. 1991. Construction of a derivative of Tn917 containing an outward directed promoter and its use in Bacillus subtilis. J. Gen. Microbiol. 137:107112.

This is a required field
Please enter a valid email address
Please check the format of the address you have entered.
Approval was a Success
Invalid data
An Error Occurred
Approval was partially successful, following selected items could not be processed due to error