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Abstract:

This chapter on focuses on the need to draw attention to the progress in developing systems for genetic manipulation in actinomycetes. A significant fraction of this progress has depended on approaches first used in species. Interestingly, the largest open reading frame in IS117 is related to that of members of the IS110 family. There is some evidence that the efficiency of translation initiation may be affected by the nature of the first few codons; changes at these positions from rare codons to synonymous codons common in genes significantly increased expression of the genes in . Surface-grown colonies may be considered as multicellular organisms with several distinct cell types. Nevertheless, most of one's understanding of morphological differentiation comes from genetical work on and . In the course of this chapter, the author has mentioned the occurrence of multiple copies of genes encoding principal; σ-factor homologs and of two quite different genes for glutamine synthase. Major problems in the analysis of antibiotic biosynthetic pathways by traditional biochemical and chemical procedures have been caused by the low concentrations of the pathway enzymes.

Citation: Chater K, Hopwood D. 1993. , p 83-99. In Sonenshein A, Hoch J, Losick R (ed), and Other Gram-Positive Bacteria. ASM Press, Washington, DC. doi: 10.1128/9781555818388.ch6

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Hexose Monophosphate Pathway
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120. Jones, R. L.,, J. C. Jaskula,, and G. R. Janssen. 1992. In vivo translational start site selection on leaderless mRNA transcribed from the Streptomyces fradiae aph gene. J. Bacteriol. 174: 4753 4760.
121. Kataoka, M.,, T. Seki,, and T. Yoshida. 1991. Five genes involved in self-transmission of pSN22, a Streptomyces plasmid. J. Bacteriol. 173: 4220 4228.
122. Kataoka, M.,, T. Seki,, and T. Yoshida. 1991. Regulation and function of the Streptomyces plasmid pSN22 genes involved in pock formation and inviability. J. Bacteriol. 173: 7975 7981.
123. Kendall, K. J.,, and S. N. Cohen. 1988. Complete nucleotide sequence of the Streptomyces lividans plasmid pIJlOl and correlation of the sequence with genetic properties. J. Bacteriol. 170: 4634 4651.
124. Khokhlov, A. S.,, L. N. Anisova,, I. I. Tovarova,, I. Y. Kleiner,, I. V. Kovalenko,, O. I. Krasilnikova,, E. Y. Kornitskaya,, and S. A. Pliner. 1973. Effect of A-factor on the growth of asporogenous mutants of Streptomyces griseus, not producing this factor. Z. Allg. Mikrobiol. 13: 647 655.
125. Khosla, C.,, S. Ebert-Khosla,, and D. A. Hopwood. 1992. Targeted gene replacements in a Streptomyces polyketide synthase gene cluster: role for the acyl carrier protein. Mol. Microbiol. 6: 3237 3249.
126. Kieser, H. M.,, T. Kieser,, and D. A. Hopwood. 1992. A combined genetic and physical map of the Streptomyces coelicolor A3(2) chromosome. J. Bacteriol. 174: 5496 5507.
127. Kieser, T.,, and D. A. Hopwood. 1991. Genetic manipulation of Streptomyces: integrating vectors and gene replacement. Methods Enzymol. 204: 430 458.
128. Kinashi, H.,, and M. Shimaji. 1987. Detection of giant linear plasmids in antibiotic producing strains of Streptomyces by the OFAGE technique. J. Antibiot. 40: 913 916.
129. Kinashi, H.,, and M. Shimaji-Murayama. 1991. Physical characterization of SCP1, a giant linear plasmid from Streptomyces coelicolor. J. Bacteriol. 173: 1523 1529.
130. Kinashi, H.,, M. Shimaji-Murayama,, and T. Hanafusa. 1992. Integration of SCP1, a giant linear plasmid, into the Streptomyces coelicolor chromosome. Gene 115: 35 41.
131. King, A. A.,, and K. F. Chater. 1986. The expression of the Escherichia coli lacZ gene in Streptomyces. J. Gen. Microbiol. 132: 1739 1752.
132. Kobler, L.,, G. Schwertfirm,, H. Schmleger,, A. Bolotin,, and I. Sladkova. 1991. Construction and transduction of a shuttle vector bearing the cos site of Streptomyces phage 0C31 and determination of its cohesive ends. FEMS Microbiol. Lett. 78: 347 354.
133. Keller, K.-P.,, and G. Reiss. 1989. Heterologous expression of the α-amylase inhibitor gene cloned from an amplified genomic sequence of Streptomyces tendae. J. Bacteriol. 171: 4953 4957.
134. Kormanec, J.,, M. Farkasovsky,, and L. Potúeková. 1992. Four genes in Streptomyces aureofaciens containing a domain characteristic of principal sigma factors. Gene 122: 63 70.
135. Kretschmer, S. 1987. Nucleotide segregation pattern during branching in Streptomyces granaticolor mycelia. J. Basic Microbiol. 27: 203 206.
136. Kretschmer, S.,, and C. Kummer. 1987. Increase of nucleoid size with increasing age of hyphal region in vegetative mycelia of Streptomyces granaticolor. J. Basic Microbiol. 27: 23 27.
137. Kroening, T. A.,, and K. E. Kendrick. 1987. In vivo regulation of histidine ammonia-lyase activity from Streptomyces griseus. J. Bacteriol. 169: 823 829.
138. Kuhstoss, S.,, and R. N. Rao. 1991. Analysis of the integration function of the streptomycete bacteriophage øC31. J. Mol. Biol. 222: 897 908.
139. Kuhstoss, S.,, M. A. Richardson,, and R. N. Rao. 1991. Plasmid vectors that integrate site-specifically in Streptomyces spp. Gene 97: 143 146.
140. Kumada, Y.,, E. Takano,, K. Nagaoka,, and C. J. Thompson. 1990. Streptomyces hygroscopicus has two glutamine synthetase genes. J. Bacteriol. 172: 5343 5351.
141. Kunze, Z. M.,, S. Wall,, R. Appelberg,, M. T. Silva,, F. Portaels,, and J. J. McFadden. 1991. IS 901, a new member of a widespread class of atypical insertion sequences, is associated with pathogenicity in Mycobacterium avium. Mol. Microbiol. 5: 2265 2272.
142. Lawlor, E. J.,, H. A. Baylls,, and K. F. Chater. 1987. Pleiotropic morphological and antibiotic deficiencies result from mutations in a gene encoding a tRNA-like product in Streptomyces coelicolor A3(2). Genes Dev. 1: 1305 1310.
143. Leblond, P.,, P. Demuyter,, J. M. Simonet,, and B. Decaris. 1990. Genetic instability and hypervariability in Streptomyces ambofaciens: towards an understanding of a mechanism of genome plasticity. Mol. Microbiol. 4: 707 714.
144. Leblond, P.,, P. Demuyter,, J.-M. Simonet,, and B. Decaris. 1991. Genetic instability and associated genome plasticity in Streptomyces ambofaciens: pulsed-field gel electrophoresis evidence for large DNA alterations in a limited genomic region. J. Bacteriol. 173: 4229 4233.
145. Leblond, P.,, F. X. Francou,, J.-M. Simonet,, and B. Decaris. 1990. Pulsed-field gel electrophoresis analysis of the genome of Streptomyces ambofaciens strains. FEMS Microbiol. Lett. 72: 79 88.
146. Leskiw, B. K. Personal communication.
147. Leskiw, B. K.,, M. J. Bibb,, and K. F. Chater. 1991. The use of a rare codon specifically during development? Mol. Microbiol. 5: 2861 2867.
148. Leskiw, B. K.,, E. J. Lawlor,, J. M. Fernandez-Abalos,, and K. F. Chater. 1991. TTA codons in some genes prevent their expression in a class of developmental, antibiotic-negative Streptomyces mutants. Proc. Natl. Acad. Sci. USA 88: 2461 2465.
149. Leskiw, B. K.,, M. Mevarech,, L. S. Barritt,, S. E. Jensen,, D. J. Henderson,, D. A. Hopwood,, C. J. Bruton,, and K. F. Chater. 1990. Discovery of an insertion sequence, IS 116, from Streptomyces clavuligerus and its related-ness to other transposable elements from actinomycetes. J. Gen. Microbiol. 136: 1251 1258.
150. Lomovskaya, N. D. Personal communication.
151. Lydiate, D. J.,, H. Ikeda,, and D. A. Hopwood. 1986. A2.6 kb DNA sequence of Streptomyces coelicolor A3(2) which functions as a transposable element. Mol. Gen. Genet. 203: 79 88.
152. MacNeil, D. J. 1988. Characterization of a unique methyl-specific restriction system in Streptomyces avermitilis. J. Bacteriol. 170: 5607 5612.
153. MacNeil, D. J.,, J. L. Occi,, K. M. Gewain,, T. MacNeil,, P. H. Gibbons,, C. L. Ruby,, and S. J. Danis. 1992. Complex organization of the Streptomyces avermitilis genes encoding the avermectin polyketide synthase. Gene 115: 119 125.
154. Madon, J.,, and R. Hütter. 1991. Transformation system for Amycolatopsis (Nocardia) mediterranei: direct transformation of mycelium with plasmid DNA. J. Bacteriol. 173: 6325 6331.
155. Malpartlda, F. Personal communication.
156. Malpartlda, F.,, J. Niemi,, R. Navarrete,, and D. A. Hopwood. 1990. Cloning and expression in a heterologous host of the complete set of genes for biosynthesis of the Streptomyces coelicolor antibiotic undecylprodigi-osin. Gene 93: 91 99.
157. Martin, J. F.,, and A. L. Demain. 1980. Control of antibiotic synthesis. Microbiol. Rev. 44: 230 251.
158. Martin, J. F.,, and P. Liras. 1989. Organization and expression of genes involved in the biosynthesis of antibiotics and other secondary metabolites. Annu. Rev. Microbiol. 43: 173 206.
159. Martinez, E.,, M. A. Fernández-Moreno,, J. L. Caballero,, D. A. Hopwood,, and F. Malpartlda. Unpublished data.
160. Matsushima, P.,, K. L. Cox,, and R. H. Baltz. 1987. Highly transformable mutants of Streptomyces fradiae defective in several restriction systems. Mol. Gen. Genet. 206: 393 400.
161. Mazodier, P.,, G. Guglielmi,, J. Davles,, and C. J. Thompson. 1992. Characterization of the gro£Z.-like genes in Streptomyces albus. J. Bacteriol. 173: 7382 7386.
162. Mazodier, P.,, R. Petter,, and C. Thompson. 1989. Inter-generic conjugation between Escherichia coli and Streptomyces species. J. Bacteriol. 171: 3583 3585.
163. Mazodier, P.,, C. Thompson,, and F. Boccard. 1990. The chromosomal integration site of the Streptomyces element pSAM2 overlaps a putative tRNA gene conserved among actinomycetes. Mol. Gen. Genet. 222: 431 434.
164. McCarthy, A. J.,, and S. T. Williams. 1992. Actinomycetes as agents of biodégradation in the environment—a review. Gene 115: 189 192.
165. McCormick, J. Personal communication.
166. McCue, L. A.,, J. Kwak,, M. J. Babcock,, and K. E. Kendrick. 1992. Molecular analysis of sporulation in Streptomyces griseus. Gene 115: 173 179.
167. McDowall, K. J.,, D. Doyle,, M. J. Butler,, C. Blnnie,, M. Warren,, and I. S. Hunter,. 1991. Molecular genetics of oxytetracycline production by Streptomyces rimosus, p. 105 116. In D. Noack,, H. Kriigel,, and S. Baumberg (ed.), Genetics and Product Formation in Streptomyces. Plenum Press, New York.
168. McHenney, M. A.,, and R. H. Baltz. 1988. Transduction of plasmid DNA in Streptomyces spp. and related genera by bacteriophage FP43. J. Bacteriol. 170: 2276 2282.
169. McLaughlin, J. R.,, C. L. Murray,, and J. C. Rabinowitz. 1981. Unique features of the ribosome binding site of the Gram-positive Staphylococcus aureus β-lactamase gene. J. Biol. Chem. 256: 11283 11291.
170. McVittie, A. M. 1974. Ultrastructural studies on sporulation in wild-type and white colony mutants of Streptomyces coelicolor. J. Gen. Microbiol. 81: 291 302.
171. Miyake, K.,, T. Kuzuyama,, S. Horinouchi,, and T. Beppu. 1990. The A-factor-binding protein of Streptomyces griseus negatively controls streptomycin production and sporulation. J. Bacteriol. 172: 3003 3008.
172. Morino, T.,, K. Takagi,, T. Nakamura,, T. Takita,, H. Saito,, and H. Takahashi. 1986. Studies of cosmid transduction in Streptomyces lividans and Streptomyces parvulus. Agrie. Biol. Chem. 50: 2493 2497.
173. Muth, G.,, B. Nussbaumer,, W. Wohlleben,, and A. Püh-ler. 1989. A vector system with temperature-sensitive replication for gene disruption and mutational cloning in streptomycetes. Mol. Gen. Genet. 219: 341 348.
174. Nagaso, H.,, S. Saito,, H. Saito,, and H. Takashashi. 1988. Nucleotide sequence and expression of a Streptomyces griseosporus proteinaceous alpha-amylase inhibitor (Haimll) gene. J. Bacteriol. 170: 4451 4457.
175. Neal, R. J.,, and K. F. Chater. 1991. Bidirectional promoter and terminator regions bracket mmr, a resistance gene embedded in the Streptomyces coelicolor A3(2) gene cluster encoding methylenomycin production. Gene 100: 75 83.
176. Novotna, J.,, and Z. Hostálék. 1985. Phosphorylation of hexoses in Streptomyces aureofaciens: evidence that the phosphoenolpyruvate:sugar phosphotransferase system is not operative. FEMS Microbiol. Lett. 28: 347 350.
177. Ochi, K. 1987. Metabolic initiation of differentiation and secondary metabolism by Streptomyces griseus: significance of the stringent response (ppGpp) and GTP content in relation to A-factor. J. Bacteriol. 169: 3608 3616.
178. Ogata, S.,, H. Suenaga,, Y. Koyama-Miyoshi,, and S. Hayashida. 1984. Cloning of the his gene of Streptomyces azureus in temperate phage SAtl. J. Gen. Appi. Microbiol 30: 405 409.
179. O'Hagan, D. 1991. The Polyketide Metabolites . Ellis Hor-wood, Chichester, United Kingdom.
180. Okamoto, S.,, T. Nlhlra,, H. Kataoka,, A. Suzuki,, and Y. Yamada. 1992. Purification and molecular cloning of a butyrolactone autoregulator receptor from Streptomyces Virginias. J. Biol. Chem. 267: 1093 1098.
181. Plaskitt, K. A. Personal communication".
182. Plohl, M.,, and V. Gamulin. 1990. Five transfer RNA genes lacking CCA termini are clustered in the chromosome of Streptomyces rimosus. Mol. Gen. Genet. 222: 129 134.
183. Puglia, A. M. Personal communication.
184. Rao, R. N.,, M. A. Richardson,, and S. Kuhstoss. 1987. Cosmid shuttle vectors for cloning and analysis of Streptomyces DNA. Methods Enzymol. 153: 166 198.
185. Reiter, W. D.,, P. Palm,, and S. Yeats. 1989. Transfer RNA genes frequently serve as integration sites for prokaryotic genetic elements. Nucleic Acids Res. 17: 1907 1914.
186. Robinson, J. 1991. Polyketide synthase complexes: their structure and function in antibiotic biosynthesis. Proc. R. Soc. B 332: 107 114.
187. Rodicio, M. R.,, M. A. Alvarez,, and K. F. Chater. 1991. Isolation and genetic analysis of IS 112, an insertion sequence responsible for the inactivation of the Sail restriction-modification system of Streptomyces albus G. Mol. Gen. Genet. 225: 142 147.
188. Saier, M. H. 1985. Mechanism and Regulation of Carbohydrate Transport in Bacteria. Academic Press, Inc., Orlando, Fla..
189. Schauer, A. T.,, A. D. Nelson,, and J. B. Daniel. 1991. Tn4563 transposition in Streptomyces coelicolor and its application to isolation of new morphological mutants. J. Bacteriol. 173: 5060 5067.
190. Sedlmeier, R.,, and H. Schmieger,. 1991. tRNA genes in Streptomyces lividans: structure, organisation and construction of suppressor tRNA, p. 65 73. In S. Baum-berg,, H. Krügel,, and D. Noack (ed.), Genetics and Product Formation in Streptomyces. Plenum Press, New York.
191. Seno, E. T.,, and R. H. Baltz,. 1989. Structural organization and regulation of antibiotic biosynthesis and resistance genes in actinomycetes, p. 1 48. In S. Shapiro (ed.). Regulation of Secondary Metabolism in Actinomycetes. CRC Press, Inc., Boca Raton, Fla..
192. Seno, E. T.,, and K. F. Chater. 1983. Glycerol catabolism enzymes and their regulation in wild-type and mutant strains of Streptomyces coelicolor A3(2). J. Gen. Microbiol. 129: 1403 1413.
193. Sherman, D. H.,, F. Malpartida,, M. J. Bibb,, H. M. Kieser,, M. J. Bibb,, and D. A. Hopwood. 1989. Structure and deduced function of the granaticin-producing polyketide synthase gene cluster of Streptomyces violaceoruber Tü22. EMBOJ. 8: 2717 2725.
194. Shiina, T.,, K. Tanaka,, and H. Takahashi. 1991. Sequence of hrdB, an essential gene encoding sigma-like transcription factor of Streptomyces coelicolor A3 (2): homology to principal sigma factors. Gene 107: 145 148.
195. Siemieniak, D. R.,, J. L. Slightom,, and S. T. Chung. 1990. Nucleotide sequence of Streptomyces fradiae transposon TN 4556: a class II transposon related to Tn3. Gene 86: 1 9.
196. Simonet, J.-M.,, D. Schneider,, J.-N. Volff,, A. Darym,, and B. Decaris. 1992. Genetic instability in Streptomyces ambofaciens: Inducibility and associated genome plasticity. Gene 115: 49 54.
197. Smith, C. P.,, and K. F. Chater. 1988. Structure and regulation of controlling sequences for the Streptomyces coelicolor glycerol operon. J. Mol. Biol. 204: 569 580.
198. Smith, C. P.,, and K. F. Chater. 1988. Cloning and transcription analysis of the entire glycerol utilization (gylABX) operon of Streptomyces coelicolor A3 (2) and identification of a closely associated transcription unit. Mol. Gen. Genet. 211: 129 137.
199. Smith, D. J.,, M. K. R. Burnham,, J. H. Bull,, J. E. Hodgson,, J. M. Ward,, P. Browne,, J. Brown,, B. Barton,, A. J. Earl,, and G. W. Turner. 1992. β-Lactam antibiotic biosynthetic genes have been conserved in clusters in prokaryotes and eukaryotes. EMBO J. 9: 741 747.
200. Smith, M. C. M.,, C. J. Ingham,, C. E. Owen,, and N. T. Wood. 1992. Gene expression in the temperate Streptomyces phage (C31. Gene 115: 43 48.
201. Sohaskey, C. D.,, H. Im,, A. D. Nelson,, and A. T. Schauer. 1992. Tn .4556 and luciferase: synergistic tools for visualizing transcription in Streptomyces. Gene 115: 67 71.
202. Solenberg, P. J.,, and R. H. Baltz. 1991. Transposition of Tn 5096 and other IS 493 derivatives in Streptomyces griseofuscus. J. Bacteriol. 173: 1096 1104.
203. Soliveri, J.,, K. L. Brown,, M. J. Buttner,, and K. F. Chater. 1992. Two promoters for the whiB sporulation gene of Streptomyces coelicolor A3(2) and their activities in relation to development. J. Bacteriol. 174: 6215 6220.
204. Sosio, M.,, J. Madon,, and R. Hütter. 1989. Excision of pIJ408 from the chromosome of Streptomyces glaucescens and its transfer to Streptomyces lividans. Mol. Gen. Genet. 218: 169 176.
205. Stein, D.,, and S. N. Cohen. 1989. A cloned regulatory gene of Streptomyces lividans can suppress the pigment deficiency phenotype of different developmental mutants. J. Bacteriol. 171: 2258 2261.
206. Stein, D. S.,, K. J. Kendall,, and S. N. Cohen. 1989. Identification and analysis of transcriptional regulatory signals for the kil and kor loci of Streptomyces plasmid pIJl0l. J. Bacteriol. 171: 5768 5775.
207. Stover, C. K.,, V. F. dela Cruz,, T. R. Fuerst,, J. E. Burlein,, L. A. Benson,, L. T. Bennett,, G. P. Bansal,, J. F. Young,, M. H. Lee,, G. F. Hatfull,, S. B. Snapper,, R. G. Garletta,, W. R. Jacobs, Jr.,, and B. R. Bloom. 1991. New use of BCG for recombinant vaccines. Nature 351(London): 456 460.
208. Strauch, E.,, E. Takano,, H. Baylis,, and M. J. Bibb. 1991. The stringent response in Streptomyces coelicolor A3(2). Mol. Microbiol. 5: 289 298.
209. Strickler, J. E.,, T. M. Berka,, J. Gorniak,, J. Fornwald,, R. Keys,, J. J. Rowland,, M. Rosenberg,, and D. P. Taylor. 1992. Two novel Streptomyces protein protease inhibitors: purification, activity, cloning and expression. J. Biol. Chem. 267: 3236 3241.
210. Strohl, W. R. 1992. Compilation and analysis of DNA sequences associated with apparent streptomycete promoters. Nucleic Acids Res. 5: 961 974.
211. Stuttard, C., 1989. Generalized transduction in Streptomyces species, p. 157 162. In C. L. Hershberger,, S. W. Queener,, and G. Hegeman (ed.), Genetics and Molecular Biology of Industrial Microorganisms. American Society for Microbiology, Washington, D.C..
212. Stuttard, C. Personal communication.
213. Summers, R. G.,, E. Wendt-Pienkowski,, H. Motamedi,, and C. R. Hutchinson. 1992. Nucleotide sequence of the tcmII-tcmIV region of the tetracenomycin ? biosynthetic gene cluster of Streptomyces glaucescens and evidence that the tcmN gene encodes a multifunctional cyclase-dehydratase- O-methyl transferase. J. Bacteriol. 174: 1810 1820.
214. Taguchi, S. 1992. Streptomyces subtilisin inhibitor: genetical characterization and its application. Actinomycetologica 6: 9 20.
215. Takahashi, H.,, T. Isogai,, T. Morino,, H. Kojima,, and H. Saito,. 1982. Development of phage vector systems in Streptomyces, p. 61 65. In Y. Ikeda, and T. Beppu (ed.), Proceedings of the IVth International Symposium on Genetics of Industrial Microorganisms, 1982. Kodansha, Tokyo.
216. Tan, H. 1991. Molecular genetics of developmentally regulated promoters in Streptomyces coelicolor A3(2). Ph.D. thesis, University of East Anglia, Norwich, United Kingdom.
217. Tanaka, K.,, T. Shilna,, and H. Takahashi. 1988. Multiple principal sigma factor homologs in eubacteria: identification of the "rpoD box." Science 242: 1040 1042.
218. Tanaka, K.,, T. Shiina,, and H. Takahashi. 1991. Nucleotide sequence of genes hrdA, hrdC and hrdD from Streptomyces coelicolor A3(2) having similarity to rpoD genes. Mol. Gen. Genet. 229: 234 240.
219. Urabe, H.,, and H. Ogawara. 1992. Nucleotide sequence and transcriptional analysis of activator-regulator proteins for β-lactamase in Streptomyces cacaoi. J. Bacteriol. 174: 2834 2842.
220. Vats-Mehta, S.,, P. Bouvrette,, F. Shareck,, R. Morosoli,, and D. Kluepfel. 1990. Cloning of a second xylanase-encoding gene of Streptomyces lividans 66. Gene 86: 119 122.
221. Virolle, M. J.,, and M. J. Bibb. 1988. Cloning, characterization and regulation of an α-amylase gene from Streptomyces limosus. Mol. Microbiol. 2: 197 208.
222. Vujaklija, D.,, K. Ueda,, S.-K. Hong,, T. Beppu,, and S. Horinouchi. 1991. Identification of an A-factor-dependent promoter in the streptomycin biosynthetic cluster of Streptomyces griseus. Mol. Gen. Genet. 229: 119 128.
223. Wang, P.,, S. S. Harvey,, P. F. G. Sims,, and P. Broda. The construction of Streptomyces cyaneus genomic libraries in Escherichia coli is dependent upon the use of Mcr-deficient strains. Gene 119: 127 129.
224. Weber, J. M.,, J. O. Leung,, S. J. Swanson,, K. B. Idler,, and J. B. McAlpine. 1991. An erythromycin derivative produced by targeted gene disruption in Saccharopolys-pora erythraea. Science 252: 114 117.
225. Weber, J. M.,, and R. Losick. 1988. The use of a chromosome integration vector to map erythromycin resistance and production genes in Saccharopolyspora erythraea (Streptomyces erythreus). Gene 68: 173 180.
226. Westpheling, J.,, and M. Brawner. 1989. Two transcribing activities are involved in expression of the Streptomyces galactose operon. J. Bacteriol. 171: 1355 1361.
227. Westpheling, J.,, M. Ranes,, and R. Losick. 1985. RNA polymerase heterogeneity in Streptomyces coelicolor. Nature (London) 313: 22 27.
228. Wildermuth, H.,, and D. A. Hopwood. 1970. Septation during sporulation in Streptomyces coelicolor. J. Gen. Microbiol. 60: 51 59.
229. Willey, J. Personal communication.
230. Willey, J.,, R. Santamaria,, J. Guijarro,, M. Geistlich,, and R. Losick. 1991. Extracellular complementation of a developmental mutation implicates a small sporulation protein in aerial mycelium formation by Streptomyces coelicolor. Cell 65: 641 650.
231. Woudt, L. P.,, K. Rietveld,, M. Verdurmen,, J. van Haarlem,, G. P. van Wezel,, E. Vijgenboom,, and L. Bosch. Three fu/Hike) genes in Streptomyces ramocissimus. Submitted for publication.
232. Wright, F.,, and M. J. Bibb. 1992. Codon usage in the G+C-rich Streptomyces genome. Gene 113: 55 65.
233. Wu, X.,, and K. Roy. 1993. Complete nucleotide sequence of a linear plasmid from Streptomyces clavuligerus and characterization of its RNA transcripts. J. Bacteriol. 175: 37 52.
234. Yagi, Y. 1990. Transposition of Tn 4560 in Streptomyces avermitilis. J. Antibiot. 43: 1204 1206.
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237. Zakrzewska-Czerwinska, J.,, and H. Schrempf. 1992. Characterization of an autonomously replicating region from the Streptomyces lividans chromosome. J. Bacteriol. 174: 2688 2693.
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