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Complete genome sequence of the model actinomycete Streptomyces coelicolor A3(2)

S.D.Bentley*,K.F.Chater?,A.-M.Cerden?o-Ta′rraga*,G.L.Challis??,N.R.Thomson*,K.D.James*,D.E.Harris*,M.A.Quail*,H.Kieser?, D.Harper*,A.Bateman*,S.Brown*,G.Chandra?,C.W.Chen§,M.Collins*,A.Cronin*,A.Fraser*,A.Goble*,J.Hidalgo*,T.Hornsby*, S.Howarth*,C.-H.Huang§,T.Kieser?,https://www.wendangku.net/doc/7a14067912.html,rke*,L.Murphy*,K.Oliver*,S.O’Neil*,E.Rabbinowitsch*,M.-A.Rajandream*,K.Rutherford*, S.Rutter*,K.Seeger*,D.Saunders*,S.Sharp*,R.Squares*,S.Squares*,K.Taylor*,T.Warren*,A.Wietzorrek?,J.Woodward*,B.G.Barrell*, J.Parkhill*&D.A.Hopwood?

*The Wellcome Trust Sanger Institute,Wellcome Trust Genome Campus,Hinxton,Cambridge CB101SA,UK

?John Innes Centre,Norwich Research Park,Colney,Norwich NR47UH,UK

?Department of Chemistry,University of Warwick,Coventry CV47AL,UK

§Institute of Genetics,National Yang-Ming University,Shih-Pai,Taipei112,Taiwan ........................................................................................................................................................................................................................... Streptomyces coelicolor is a representative of the group of soil-dwelling,?lamentous bacteria responsible for producing most natural antibiotics used in human and veterinary medicine.Here we report the8,667,507base pair linear chromosome of this organism,containing the largest number of genes so far discovered in a bacterium.The7,825predicted genes include more than 20clusters coding for known or predicted secondary metabolites.The genome contains an unprecedented proportion of regulatory genes,predominantly those likely to be involved in responses to external stimuli and stresses,and many duplicated gene sets that may represent‘tissue-speci?c’isoforms operating in different phases of colonial development,a unique situation for a bacterium. An ancient synteny was revealed between the central‘core’of the chromosome and the whole chromosome of pathogens Mycobacterium tuberculosis and Corynebacterium diphtheriae.The genome sequence will greatly increase our understanding of microbial life in the soil as well as aiding the generation of new drug candidates by genetic engineering.

Nutritionally,physically and biologically,soil is a particularly complex and variable environment.Streptomycetes are among the most numerous and ubiquitous soil bacteria1.They are crucial in this environment because of their broad range of metabolic pro-cesses and biotransformations.These include degradation of the insoluble remains of other organisms,such as lignocellulose and chitin(among the world’s most abundant biopolymers),making streptomycetes central organisms in carbon recycling.Unusually for bacteria,streptomycetes exhibit complex multicellular develop-ment,with differentiation of the organism into distinct‘tissues’:a branching,?lamentous vegetative growth gives rise to aerial hyphae bearing long chains of reproductive spores.The importance of streptomycetes to medicine results from their production of over two-thirds of naturally derived antibiotics in current use(and many other pharmaceuticals such as anti-tumour agents and immuno-suppressants),by means of complex‘secondary metabolic’path-ways.Furthermore,streptomycetes are members of the same taxonomic order(Actinomycetales)as the causative agents of tuberculosis and leprosy(Mycobacterium tuberculosis and M. leprae),the genomes of which have been sequenced2,3.Much should be learned about these pathogens from genome-level comparisons with harmless saprophytic relatives such as streptomycetes. Streptomyces coelicolor A3(2)is genetically the best known repre-sentative of the genus4.The single chromosome is linear with a centrally located origin of replication(oriC)and terminal inverted repeats(TIRs)carrying covalently bound protein molecules on the free50ends.Replication proceeds bidirectionally from oriC,leaving a terminal single-stranded gap on the discontinuous strand after removal of the last RNA primer.An unusual process of‘end-patching’by DNA synthesis primed from the terminal protein?lls the gap5.Studies of many streptomycetes,including most notably a close relative of the A3(2)strain,Streptomyces lividans66,estab-lished further novelties.More than a million base pairs(bp)of DNA at either end of the chromosomes can undergo extensive deletions and ampli?cations without compromising viability under labora-tory conditions6,and early comparisons of linkage maps established that most streptomycetes show conservation of gene order(synteny) in the core region7.Here,we report the use of an ordered cosmid library8to sequence the S.coelicolor genome.The strain used,M145, is a prototrophic derivative of strain A3(2)lacking its two plasmids (SCP1,linear,365kb,AL590463,AL590464;and SCP2,circular, 31kb,AL645771,which have been sequenced separately). Genome structure

General features of the chromosome sequence are shown in Table1 and Fig.1.At8,667,507bp it is the largest completely sequenced bacterial genome.The oriC and dnaA gene are about61kb left of the centre,at4,269,853–4,272,747bp.Like many other microbial genomes,there is a slight bias(55.5%)towards coding sequences on the leading strand.Although less pronounced than for most other eubacterial chromosomes,there is a discernible decrease in the GC bias around oriC,thought to be related to DNA replication9.In contrast to all other bacterial genomes studied to date,however,the Table1General features of the chromosome

Component of chromosome Property ............................................................................................................................................................................. Total size8,667,507bp Terminal inverted repeat21,653bp

GtC content72.12% Coding sequences7,825 ...of which pseudogenes55 Coding density88.9% Average gene length991bp Ribosomal RNAs6￡(16S–23S–5S) Transfer RNAs63

Other stable RNAs3 .............................................................................................................................................................................

S.coelicolor chromosome displays a downward rather than an upward shift,indicating a small bias towards C on the leading strand.

Coding density is largely uniform across the chromosome,with only a slight decrease in the distal regions.The distribution of different types of genes reveals,however,a central core comprising approximately half the chromosome and a pair of chromosome arms(Fig.1).Nearly all genes likely to be unconditionally essen-tial—such as those for cell division,DNA replication,transcription, translation and amino-acid biosynthesis—are located in the core (exceptions tend to be duplicate genes).In contrast,‘contingency’loci coding for probable non-essential functions,such as secondary metabolites,hydrolytic exoenzymes,the conservons(conserved operons)and‘gas vesicle’proteins(see below),lie in the arms. Curiously,this biphasic structure of the chromosome does not align with the position of oriC.The core appears to extend from around 1.5Mb to6.4Mb,giving uneven arm lengths of approximately 1.5Mb(left arm)and2.3Mb(right arm).The difference in arm lengths may re?ect some gross rearrangement or different rates of DNA accumulation in each arm.The fact that oriC is roughly central suggests some selective pressure for such positioning. Streptomyces coelicolor and M.tuberculosis are both

actinomycetes

scale is numbered anticlockwise(to correspond with the previously published map8)in megabases and indicates the core(dark blue)and arm(light blue)regions of the chromosome.Circles1and2(from the outside in),all genes(reverse and forward strand, respectively)colour-coded by function(black,energy metabolism;red,information transfer and secondary metabolism;dark green,surface associated;cyan,degradation of large molecules;magenta,degradation of small molecules;yellow,central or intermediary metabolism;pale blue,regulators;orange,conserved hypothetical;brown,pseudogenes;division,DNA replication,transcription,translation and amino-acid biosynthesis,colour coding as for circles1and2);circle4,selected‘contingency’genes(red,secondary metabolism;pale blue,exoenzymes;dark blue,conservon;green,gas vesicle proteins); circle5,mobile elements(brown,transposases;orange,putative laterally acquired genes);circle6,GtC content;circle7,GC bias((G2C/GtC),khaki indicates values .1,purple,1).The origin of replication(Ori)and terminal protein(blue circles)are also indicated.

but have very different lifestyles.Their genomes reveal much similarity at the level of individual gene sequences,and many similar gene clusters.Global comparison showed perceptible higher-order synteny as well,shown as a dot plot in Fig.2a.A prominent feature is the central broken diagonal cross pattern formed by the regions of synteny.This broken-X pattern is com-monly seen in comparisons of related bacteria and the breaks are attributed to multiple inversions centred on oriC 10.Normally,synteny extends over the whole of the compared chromosomes;however,for the comparison between S.coelicolor and M.tubercu-losis ,the broken-X pattern correlates only with the core of the S.coelicolor chromosome.Therefore this region and the whole M.tuberculosis chromosome must have had a common ancestor,with the chromosome arms of S.coelicolor consisting of subsequently acquired DNA.The syntenic regions mainly comprise genes con-cerned with primary cellular functions.The most strongly con-served is the gene cluster coding for the subunits of respiratory chain NADH dehydrogenase (systematic gene numbers SCO4562–4575).Functions/proteins coded for by other regions of synteny include the origin of replication (SCO3873–3892),urease activity (SCO1231–1236),pyrimidine biosynthesis (SCO1472–1488),argi-nine biosynthesis (SCO1570–1580),pentose phosphate pathway/tricarboxylic acid cycle (SCO1921–1953),histidine and tryptophan biosynthesis (SCO2034–2054),cell division (SCO2077–2092)and ribosomal proteins (SCO4701–4724).

The genome of the pathogenic actinomycete Corynebacterium diphtheriae has been sequenced recently (https://www.wendangku.net/doc/7a14067912.html,/

Projects/C_diphtheriae/).Comparison with the S.coelicolor chromosome gives a similar pattern to that for M.tuberculosis ,with the regions of synteny covering the entire C.diphtheriae chromosome and just the S.coelicolor core region (Fig.2b).The syntenic regions again correspond to genes coding for primary cellular functions and several of these regions are common to all three chromosomes.Mycobacterium tuberculosis and C.diphtheriae have more extensive synteny than either has with S.coelicolor (Fig.2c),re?ecting taxonomic groupings:C.diphtheriae and M.tuber-culosis are in the suborder Corynebacterineae of the actinomycetes,whereas S.coelicolor is in the Streptomycineae.

By investigating regions of unusual DNA content and/or genes with sequence similarity to those from known mobile genetic elements,we designated 14regions as potentially recently laterally acquired insertions (See Supplementary Information).By far the largest insertion contains 148genes and is located at a transfer RNA gene 11:as well as many hypothetical genes,it includes genes for heavy metal resistance (SCO6835–6837)and secondary metabolite production (SCO6827).Six other inserted regions have plasmid function genes in common with the integrative plasmid pSAM2of Streptomyces ambofaciens 12.Four of these pSAM2-like integrants appear to have inserted within a tRNA gene,including two that are adjacent to secondary metabolic clusters (calcium-dependent anti-biotic (CDA),SCO3250–3262;whiE ,SCO5327–5350).Notably,11of the 14insertions lie to the right of oriC ,correlating with the greater variation in DNA composition in the right half of the chromosome (Fig.1).

Putative transposase genes are found throughout the chromo-some in intact,truncated and frame-shifted forms.Many are associated with the multi-gene integrations described above.For the remainder,there is a particular concentration at the sub-TIR regions,35–95kb from the ends (Fig.1).This indicates a tolerance to insertion events in these regions and thus offers a possible route for chromosome expansion.Of the 78predicted transposase coding sequences,?ve are within transposons (one of which codes for a possible antibiotic resistance protein (SCO0107)),31form simple insertion elements and the remainder are not bounded by inverted repeats.Most fall into ?ve families,suggesting a degree of intra-chromosomal transposition.Such events offer a route for gene duplication.Two of the insertion elements mark the inner bound-aries of the TIRs,suggesting a possible role in their maintenance.

A plethora of proteins

With 7,825predicted genes,the S.coelicolor chromosome has an enormous coding potential.This ?gure compares with 4,289genes in the Gram-negative bacterium Escherichia coli ;4,099in the Gram-positive spore-former Bacillus subtilis ;6,203in the lower eukaryote Saccharomyces cerevisiae ;and a predicted 31,780in humans (https://www.wendangku.net/doc/7a14067912.html,/genomes/).The genome contains almost twice as many genes as that of M.tuberculosis .This large number of genes re?ects both a multiplicity of new protein families and an expansion within known families when compared with other bacteria (further information is available at https://www.wendangku.net/doc/7a14067912.html,/Projects/S_coe-licolor/).Many protein families that are signi?cantly expanded in S.coelicolor are involved in regulation,transport and degradation of extracellular nutrients (Table 2).

The genome shows a strong emphasis on regulation,with 965proteins (12.3%)predicted to have regulatory function.Discovery of so many regulators extends the observation that the proportion of regulatory genes increases with bacterial genome size 13.There is a clear preference for certain regulator groups.Sigma factors act by binding to and affecting the promoter speci?city of the RNA polymerase core enzyme,thus directing the selective transcription of gene sets.Streptomyces coelicolor codes for a remarkable 65sigma factors (the next highest number so far found is 23in Mesorhizo-bium loti ,with a genome size of 7.6Mb 14),of which 45are ‘ECF’(extra-cytoplasmic function)sigma factors (41from family

Figure 2Comparison of chromosome structure for S.coelicolor versus M.tuberculosis (a ),S.coelicolor versus C.diphtheriae (b )and M.tuberculosis versus C.diphtheriae (c ).Axes represent the proteins coded for in the order in which they occur on the

chromosomes.For each genome,DnaA is centrally located.Dots represent a reciprocal best match (by FASTA comparison 50)between protein sets.The bars above plots a and b indicate the core (solid,SCO1440–5869)and arm (hatched)regions of the S.coelicolor chromosome.

alone;Table2).Previously described ECF sigma factors(in S. coelicolor)respond to external stimuli and activate genes involved in disulphide stress,cell-wall homeostasis and aerial mycelium development15.Most of the other sigma factors fall into a single group(family54,Table2).Within this is a sub-group peculiar to Gram-positive bacteria,most of which have a single member; however,B.subtilis has three,controlling forespore development and the general stress response,and S.coelicolor has at least eight, many of them involved in responses to various stresses16.The numerous potentially stress-responsive sigma factors may account for the independent regulation of diverse stress response regulons in S.coelicolor17.Although widely distributed among bacteria,the atypical,enhancer-dependent sigma-54and its cognate activators18 are absent.

Streptomyces coelicolor also has abundant two-component regu-latory systems where typically,in response to an extracellular stimulus,an integral membrane sensor protein phosphorylates a response regulator,causing it to bind to speci?c promoter regions and thus activate or repress transcription.We identi?ed85sensor kinases and79response regulators,including53sensor–regulator pairs.The genome also codes for many members of previously described regulator families such as LysR,LacI,ROK,GntR,TetR, IclR,AraC,AsnC and MerR.The TetR family regulators in S. coelicolor form several subfamilies,often containing few or no members from the other genomes analysed.Furthermore,there is a group(family86,Table2)of25putative DNA-binding proteins that has no members from outside S.coelicolor and may constitute a new Streptomyces-speci?c family of regulators.Also notable is the presence of44putative serine/threonine protein kinases(family6.1, Table2).Examples of these typically eukaryotic regulators are now known to occur in many bacteria,but in much smaller numbers. Re?ecting its many interactions with the complex soil environ-ment,S.coelicolor has614proteins(7.8%)with predicted transport function.A large proportion of these are of the ABC transporter type,including81typical ABC permeases and141ATP-binding proteins(24of which are fused to membrane-spanning domains).Transporters for which the substrate is predictable include those for sugars,amino acids,peptides,metals and other ions.There are also several possible drug ef?ux proteins.Import of speci?c substrates would in part be facilitated by the75putative surface-anchored substrate-binding proteins of S.coelicolor.

The ability of S.coelicolor to exploit nutrients in the soil is abundantly demonstrated by our prediction of819potentially secreted proteins(10.5%).Secreted hydrolases are particularly numerous(for example,family7(Table2),which is over-rep-resented in S.coelicolor).They include60proteases/peptidases,13 chitinases/chitosanases,eight cellulases/endoglucanases,three amy-lases and two pectate lyases.As well as the complete Sec protein translocation system,S.coelicolor seems to contain the machinery and cognate signal sequences for the recently discovered TAT(twin arginine transport)system for exporting pre-folded proteins19(T. Palmer,personal communication).

A marked example of multiple paralogues in S.coelicolor is a four-gene cluster that we named the conservon(for conserved operon). In the13such clusters(cvnA,B,C,D,1-13)there is unidirectional transcription and often overlap of translational start and stop codons,suggesting an operon structure.The only other known cvn cluster is present in M.tuberculosis.The protein products form distinct and exclusive families(Table2;families178,177,214,180; CvnA,B,C and D,respectively).The?rst gene codes for a probable membrane protein weakly resembling sensor kinases,the fourth codes for a possible ATP/GTP-binding protein,and the other two are of unknown function.In four of the clusters the immediate downstream gene codes for a predicted cytochrome P-450. Paralogous enzymes may sometimes represent isozymes active at different stages in the developmental cycle.One such example is the differential activities of duplicate gene clusters for glycogen syn-thesis in the vegetative and aerial mycelium20.Here we highlight a further?ve examples of paralogues for metabolic enzymes in S. coelicolor.(1)Two gene clusters code for enzymes of the pentose phosphate pathway(SCO1935–1939and SCO6657–6663).(2)Four loci for tryptophan biosynthesis(SCO2036–2043,SCO2117,

Table2Occurrence of a selection of protein families in six related genomes

Majority description*Sco Mtu Cdi Bsu Mlo Eco ................................................................................................................................................................................................................................................................................................................................................................... ECF sigma factor(13)41(0.52)10(0.25)7(0.29)7(0.17)16(0.23)1(0.02) Sigma factor(54)14(0.17)3(0.07)2(0.08)8(0.19)3(0.04)4(0.09) Two-component sensor kinase(1.3)27(0.34)8(0.20)5(0.20)12(0.29)41(0.60)20(0.46) Two-component sensor kinase(1.15)6(0.07)00000 Two-component regulator(1.6)50(0.63)2(0.05)5(0.20)9(0.21)5(0.07)7(0.16) Two-component regulator(1.5)24(0.30)11(0.28)6(0.24)13(0.31)21(0.31)14(0.32) Serine/threonine protein kinase(6.1)44(0.56)13(0.33)5(0.20)8(0.19)14(0.20)8(0.18) Regulator(LacI)(2.4)33(0.42)1(0.02)2(0.08)12(0.29)15(0.22)13(0.30) Regulator(ROK)(36)23(0.29)3(0.07)3(0.12)3(0.07)6(0.08)7(0.16) Regulator(TetR)(112)18(0.22)1(0.02)0000 Regulator(KorSA/GntR)(2.9)10(0.12)00000 Regulator(WhiB-like)8(0.10)4(0.10)3(0.12)000 DNA-binding(86)25(0.31)00000 ABC transport(ATP-binding)(2.1.3)27(0.34)4(0.10)8(0.33)6(0.14)3(0.04)3(0.06) Transport(permease)(2.3)36(0.45)4(0.10)1(0.04)8(0.19)25(0.37)3(0.06) Transport(sugar)(2.2)36(0.45)4(0.10)1(0.04)8(0.19)26(0.38)3(0.06) Transport(substrate-binding)(1.8)35(0.44)4(0.10)1(0.04)6(0.14)24(0.35)3(0.06) Integral membrane(59)14(0.17)14(0.35)3(0.12)2(0.04)00 Membrane ATPase(42)13(0.16)12(0.30)6(0.24)4(0.09)3(0.04)1(0.02) Secreted hydrolase(7)100(1.27)19(0.48)8(0.33)21(0.51)8(0.11)9(0.20) Secreted hydrolase(7.3)17(0.21)0001(0.01)0 Secreted chitinase(7.8)5(0.06)00000 Secreted protease(191)10(0.12)3(0.07)0000 Secreted protease(7.6)8(0.10)00000 Secreted cellulase(7.4)7(0.08)1(0.02)01(0.02)00 Secreted hypothetical(17)25(0.31)11(0.28)8(0.33)13(0.31)8(0.11)9(0.20) Hypothetical(63)25(0.31)006(0.14)00 Conservon(Cvn1–4;178,177,214,180)13(0.16)1(0.02)0000 Hypothetical(204)13(0.16)00000 Hypothetical(19)12(0.15)00000 ................................................................................................................................................................................................................................................................................................................................................................... Numbers indicate absolute number of proteins from each genome in each family and the percentage of the total proteins in that genome in parentheses.Family number is indicated in parentheses in the majority description column.The hierarchical numbering system re?ects use of higher BlastP thresholds to break large complex families into discrete https://www.wendangku.net/doc/7a14067912.html,plete data are available from http:// https://www.wendangku.net/doc/7a14067912.html,/Projects/S_coelicolor/.Sco,S.coelicolor;Mtu,M.tuberculosis;Cdi,C.diphtheriae;Bsu,B.subtilis;Mlo,M.loti;Eco,E.coli.

*Groupings are based on sequence similarity,so individual families do not necessarily include all representatives of each type of protein in each genome(see Methods).

SCO2147,SCO3211–3214)include two trpC,two trpD and three trpE genes.A trpCDGE locus is within the gene cluster for pro-duction of CDA21,a peptide antibiotic that contains tryptophan(in the‘unnatural’D form).The local cluster may ensure adequate tryptophan for CDA biosynthesis at the appropriate stage in the life cycle,independently of the needs of protein synthesis.(3)Five homologues of fabH code for a dedicated ketosynthase for the?rst step in fatty acid biosynthesis(condensation of acetyl-coenzyme A (CoA)with malonyl-CoA to yield acetoacetyl-CoA).One of the?ve (SCO2388)is in the main fatty acid biosynthetic operon and is essential22.Three of the other four fabH homologues(SCO5888, 3246,1271)are in gene clusters for secondary metabolism:the red and cda clusters,and a cluster of unknown product(see below).At least the?rst two clusters determine molecules with fatty acid components,and the presence of fabH paralogues makes it highly probable that some of the steps in their biosynthesis use dedicated enzymes,rather than sharing enzymes functioning in primary metabolism23.(4)Three clusters code for a typical four-subunit respiratory nitrate reductase(SCO0216–0219,SCO4947–4950, SCO6532–6535),indicating the importance of a capacity for micro-aerobic growth in what was classically regarded as an obligate aerobe.(5)Flexibility in respiration is further indicated by a second (partial)copy of the operon coding for subunits of the respiratory chain NADH dehydrogenase(SCO4599–4608). Unexpectedly,there are two gene clusters(SCO0649–0658, SCO6499–6508)similar in sequence and gene order to an operon from Halobacterium sp.that is involved in the production of gas vesicle proteins,including the eight genes essential for this pheno-type24.Many overtly water-living bacteria use gas vesicles as?o-tation devices,but the only previous occurrence of gas vesicle genes (but not so far of the vesicles themselves)in a soil organism is in Bacillus megaterium25.The bene?t of gas vesicles to Streptomyces is unknown,but perhaps such buoyancy devices would allow spores to remain at the oxygen-rich surface during dispersal and germination in waterlogged soil.

Many genes for secondary metabolism

Chromosomal gene clusters specifying the biosynthesis of the aromatic polyketide antibiotic actinorhodin,the so-called RED complex of red oligopyrrole prodiginine antibiotics,and the non-ribosomal peptide CDA had previously been analysed26,27,as had the whiE cluster of genes coding for a type II polyketide synthase for a grey spore pigment28.The genome sequence reveals a further18 clusters that would code for enzymes characteristic of secondary metabolism(Fig.3).These include type I modular and both type I and type II iterative polyketide synthases(PKSs),chalcone synthases,non-ribosomal peptide synthetases(NRPSs),terpene cyclases,and others.The distribution of the clusters on the chromo-some seems non-random,with some preponderance in the arms, but more especially in a region near the right-hand core–arm boundary(Fig.1).Comparison with similar clusters from other organisms and the application of recently developed sequence analysis tools have,in some cases,provided insight into the probable structure of the end products determined by these genes.For example,using predictive models for substrate amino-acid recog-nition29,30,the two NRPSs coded for by SCO0492and

SCO7681–

Figure3Secondary metabolites known or predicted to be made by S.coelicolor A3(2), grouped according to their putative function.These are:antibiotics(a),siderophores(b), pigments(c),lipids(d)and other molecules(e).The chromosomal locations of the gene clusters are:actinorhodin,SCO5071–5092;prodiginines(mixture of butyl-meta-cycloheptylprodiginine(shown)and undecylprodiginine),SCO5877–5898;CDA complex (CDA1,R?OPO3H2,R0?H;CDA2,R?OPO3H2,R0?Me;CDA3b,R?OH,R0?H; CDA4b,R?OH,R0?Me),SCO3210–3249;desferrioxamines(mixture of desferrioxamine G1(shown)and desferrioxamine E),SCO2782–2785;coelichelin, SCO0489–0499;coelibactin(structure is that predicted for a late intermediate attached to the PCP domain in the last module of the coelibactin NRPS;R?H/Me,the complete structure cannot be predicted as the regiospeci?city of several methyltransferases,a cytochrome P-450and an oxidoreductase coded for by genes in the cluster cannot be deduced),SCO7681–7691;TW95a(structure is the product obtained from heterologous expression of the whiE minimal PKS and the whiE-ORFIV genes;the structure of the grey spore pigment has not been elucidated),SCO5314–5320;tetrahydroxynaphthalene (predicted product of the chalcone synthase,which may be further modi?ed by enzymes coded for by other genes in the cluster),SCO1206–1208;isorenieratene,SCO0185–0191;hopanoids(mixture of aminotrihydroxybacteriohopane(shown)and hopene), SCO6759–6771;eicosapentaenoic acid,SCO0124–0129;geosmin,SCO6073; butyrolactones(believed to be assembled by the scbA gene product),SCO6266.The structures of the remaining putative secondary metabolites are unknown.The chromosomal location of these clusters and the type of secondary metabolic enzyme(s) coded for are:SCO6429–6438,NRPS;SCO6273–6288and SCO6826-6827,

type I polyketide synthases;SCO7669–7671and SCO7222,chalcone synthases; SCO5222–5223,sesquiterpene cyclase;SCO5799–5801,siderophore synthetase; SCO1265–1273,type II fatty acid synthase;SCO0381–0401,deoxysugar synthases/ glycosyl transferases.

7683were deduced to catalyse the biosynthesis of novel side-rophores named‘coelichelin’31and‘coelibactin’(G.L.C.,unpub-lished data),respectively.A third cluster,SCO2782–2785,probably directs the biosynthesis of two further siderophores,desferrioxa-mines G1and E32.Two large open reading frames(ORFs)(SCO0126 and0127)code for multi-enzymes with a domain organization very similar to a type I iterative PKS/FAS from a Gram-negative bacterium,Shewanella sp.,that catalyses biosynthesis of eicosapen-taenoic acid33.We therefore predict a role for this cluster in polyunsaturated fatty acid biosynthesis.Similarly,the cluster SCO6759–6771has been implicated in hopanoid biosynthesis34, and SCO1206–1208in tetrahydroxynaphthalene biosynthesis35.The sesquiterpene cyclase coded for by SCO6073is probably involved in geosmin biosynthesis(B.Gust,K.Fowler,T.K.,G.L.C.and K.F.C., personal communication)and SCO0185–0191probably directs biosynthesis of the carotenoid isoreneriatine36.

Although three of the S.coelicolor clusters specify antibiotics, most of the others are probably responsible for products with different functions.For example,hopanoids may protect against water loss through the plasma membrane in the aerial mycelium34, and eicosapentaenoic acid may help to maintain membrane?uidity at low temperature.It is notable that at least three clusters probably code for siderophore biosynthesis,implying that S.coelicolor is under strong selective pressure to scavenge iron in situations of low iron availability.Thus,products of some of these clusters might accurately be labelled‘stress metabolites’,predicted to combat stresses of a physical(desiccation,low temperature),chemical (low iron)or biological(competition)nature.

Cell and developmental biology

Escherichia coli and B.subtilis multiply by binary?ssion,whereas S. coelicolor grows as a non-dividing,many-branched mycelium, mainly by tip growth,with multiple copies of the genome in each hyphal compartment.Unigenomic dispersive exospores are borne as chains on specialized,little-branched aerial hyphae that probably extend by intercalary growth37.The genome sequence provides some new insights into this complex life cycle.

Initiation of DNA replication in S.coelicolor involves an oriC-linked dnaA gene,the product of which interacts with an unusually large number(17)of DnaA boxes at the replication origin38.In addition to its initiator function,DnaA is a transcription factor in a diverse range of bacteria39.It is therefore conspicuous that42(82%) out of the51‘strong’DnaA boxes of S.coelicolor(TT(G/ A)TCCACA38)lie in non-coding DNA upstream of genes.DnaA may conceivably coordinate the replication of multiple genomes in each hyphal compartment with cell-cycle-dependent gene expression.

Our limited understanding of bacterial chromosome partitioning is based largely on studies of low copy number plasmids of Gram-negative bacteria40.The parAB gene pair on many such plasmids codes for ParA,an ATPase of unclear function,and ParB,which binds one or more parS sites near parA and parB.Many bacteria (including S.coelicolor,but not E.coli)contain parAB genes near oriC,and in some cases parS target sites have been identi?ed41.In S. coelicolor,there is a high concentration of putative parS sites surrounding oriC42,with18‘perfect’sites(GTTTCACGTGAAAC) in a515-kb segment(4,174,551–4,689,985).Unlike DnaA boxes, nearly all of the parS sites are immediately downstream of genes, perhaps indicating selection for avoidance of effects on gene expression resulting from ParB–parS binding. Streptomycetes have at least three different kinds of septa43.It is therefore surprising that genes clearly homologous to conserved ‘divisome’(cell division)genes of other bacteria are generally present only once or(in the case of ftsA)not at all.Presumably the different kinds of cell division involve dedicated accessory proteins.This contrasts with genes coding for enzymes for pepti-doglycan synthesis and metabolism:there are eight ftsI/mrdA(class 2/3transpeptidase)and?ve mrcA/mrcB(peptidoglycan synthetase) homologues.

A principal difference between S.coelicolor and unicellular rods concerns septum placement.In rods,division involves a centrally located septum,with alternative division sites close to the cell poles usually being silent.This involves the minC,D and E genes in E.coli, and the minC,D and divIVA genes of B.subtilis44.In hyphae of Streptomyces,there is no centre point,and division events are usually far from hyphal‘poles’.Consistent with this,there are no minC,minE or divIVA-like genes in S.coelicolor.On the other hand, there is a large family of perceptibly minD-like genes(which, notably,reveal distant similarity to parA).These may control the use of potential division sites at various positions(for example, polar,sub-polar,between pre-existing septa,or at branch points). Discussion

The genome sequence of S.coelicolor has revealed much about the many adaptations of this model actinomycete to life in the highly competitive soil environment.Derived from an ancestor common to other actinomycetes,the chromosome has acquired the ability to replicate in a linear form and appears to have expanded by lateral acquisition and internal duplication of DNA.Chromosome expan-sion has provided a wealth of genes,allowing the organism a more complex life cycle,adapting to a wider range of environmental conditions and exploiting a greater variety of nutrient sources.This has coincided with an increase in regulatory systems,with a particular emphasis on detection of,and response to,extracellular stimuli.The preferential incorporation(and subsequent mainten-ance)of occasionally bene?cial sequences outside the ancestral core has created chromosome arms comprised mostly of‘non-essential’functions.The abundance of previously uncharacterized metabolic enzymes,particularly those likely to be involved in the production of natural products,is a resource of enormous potential value. Understanding of such enzymes will facilitate the genetic engineer-ing of pathways to produce new compounds with potential thera-peutic activity,including much needed antimicrobials45.The incomplete genome sequence of an industrial species,S.avermiti-lis46,appears to contain a different set of gene clusters for secondary metabolism from S.coelicolor.It may be that the arm regions of different streptomycete chromosomes have been accumulated separately,and therefore contain a largely different complement of contingency genes representing a huge pool of metabolic diversity.A Methods

Genome sequencing

We sequenced the genome of S.coelicolor A2(3)from325overlapping clones.Of these,305 were cosmids8,one was the terminal plasmid pLUS221and19were selected from a set of 3,456bacterial arti?cial chromosomes mapped to the sequences of?nished cosmid contigs by end sequencing.The methods for clone growth and isolation,sonication to produce 1.4–2-kb fragments,library preparation in either M13or pUC18vectors,and sequencing were as described previously47.Most of the clones were digested with Dra I,and insert puri?ed,before the fragmentation step in order to remove cloning vector.This was not done for those clones known to contain Dra I sites,and in these cases DNA from the cloning vector was greatly over-represented in the subclone libraries.The?nished325 clones formed a contiguous sequence extending from within the left TIR to the right end of the genome.The genome sequence was completed by extending the incomplete left TIR with a7-kb consensus sequence copied from the right TIR.The sequence was assembled,?nished and annotated as described previously2,using Artemis(https://www.wendangku.net/doc/7a14067912.html,/ Software/Artemis)to collate data and facilitate annotation.Protein families were constructed,independently of annotation,by performing an‘all-against-all’Blast(NCBI Blast version2)comparison48of proteins within a database containing all predicted protein products from six genomes(Table2),then single-linkage clustering using a Blast threshold of70bits.We checked composition of families using https://www.wendangku.net/doc/7a14067912.html,plex families were resolved by raising the Blast threshold to100,150or200bits,as re?ected in the hierarchical family numbering system(for example,family2.1.3was created using a Blast threshold of150on family2.1).

Received3December2001;accepted27March2002.

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Supplementary Information accompanies the paper on Nature’s website

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Acknowledgements

We would like to acknowledge the support of the Wellcome Trust Sanger Institute core sequencing and informatics groups.This work was funded by the Biotechnology and Biological Sciences Research Council and by the Wellcome Trust through its Beowulf Genomics Initiative.C.W.C.and C.-H.H.were supported by the R.O.C.National Science Council and Ministry of Education.

Competing interests statement

The authors declare that they have no competing?nancial interests.

Correspondence and requests for materials should be addressed to S.D.B.

(e-mail:sdb@https://www.wendangku.net/doc/7a14067912.html,)or D.A.H.(e-mail:david.hopwood@https://www.wendangku.net/doc/7a14067912.html,).The complete sequence is deposited in GenBank/EMBL under accession number AL645882.

建筑工程模板工程施工技术

建筑工程模板工程施工技术摘要：模板技术在建筑施工得到了广泛的推广和应用。在进行模板建筑时，要依据建筑施工的实际情况，选择适当的模板体系，才能保证工程的质量和施工的安全，使建筑工程的经济效益和社会效益最大化。本文首先介绍建筑工程模板工程施工的要求，再对模板工程的施工技术要点进行简要的探究。关键词：建筑工程；模板工程；施工技术前言模板施工是混凝土成型施工必不可少的环节之一，其施工目的是保证工程结构及构件的形状及尺寸，并确保相对位置的准确性，因此要求模板首先必须具有足够的强度及刚度，同时还要有足够的稳定性且便于安装和拆除。建筑工程模板施工要求较多，而且比较复杂，因此要对模板施工工作引起高度的重视。 1.工程概况某高层建筑，总建筑面积为14344M2，场地类型为三类土，建筑物框架抗震等级为二级，剪力墙抗震等级为一级，建筑结构安全为二级。地上共七层，建筑总高为35.00m，场地平整开阔。工程七层屋面梁（标高为28.80 米）处，①、 ⑧轴线外各有一超大雨棚，具体为：轴外雨棚大小尺寸为3200×30400mm。雨棚由悬臂板和悬臂梁组成。悬臂梁尺寸为350×700，檐口梁为200×700，板厚100， C30 砼。 2.建筑工程模板工程施工技术要点 2.1模式工程施工基本技术要求在建筑工程施工中，必须保证混凝土结构的工程质量以及施工安全，为了降低工程的成本，以及缩短施工进度，在模板工程施工中，必须满足四点技术要求（1）模板施工中必须充分保障混凝土的结构以及其他构件的尺寸和位置，也就是说模板的位置尺寸必须满足图纸设计要求；（2）模板必须具备一定强度和稳定度，能够承受混凝土的重量以及侧向的压力，而且在施工过程中，避免模板所承受的压力处于极限状态之下；（3）模板的构造尽可能简单，便于装拆，并符合混凝土浇筑和养护等要求；（4）模板的连接必须紧密，在接缝处必须采取加密措施，如果出现接缝不严密，必须及时采取措施，保证接缝处不出现漏浆等现象。 2.2模板配置技术模板的配置必须根据图纸的尺寸，对于结构形体相对简单的构件，其模板配可直接根据施工图纸来配置。对于模板、横档及楞木的断面以及它们之间的间距、支撑系统的配置，可查表或者按规范进行选择。而对于结构比较复杂的构件，比如楼梯等，配置模板通常采取放大洋的方法。所谓放大样，指的是根据结构图纸，在地面上画出结构构件的实体形状，然后测量出各个部分模板的准确尺寸，然后制定模板。 2.3模板施工技术一般而言，建筑工程模板施工工序为：垫层模板→基础梁模板→构造柱模板 →柱模板一墙板模板→圈梁模板→梁模板→楼板模板→楼梯模板。 2.3.1垫层模板对于垫层模板而言，其基础的高度不高，但是体积很大，安装模板前必须准确核实基础的中心线以及标高，事先弹出轴线和四周边线，根据边线尺寸将侧面模板对准，并且在校正模板垂直度以及标高之后，用支撑将模板固定，用以维持

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目录一、编制依据............................................................. 错误!未定义书签。二、工程概况............................................................. 错误!未定义书签。三、施工部署............................................................. 错误!未定义书签。四、施工准备............................................................. 错误!未定义书签。五、进度计划............................................................. 错误!未定义书签。六、施工工艺............................................................. 错误!未定义书签。七、质量标准............................................................. 错误!未定义书签。八、成品保护............................................................. 错误!未定义书签。九、应注意的质量问题 ............................................ 错误!未定义书签。十、安全注意事项 .................................................... 错误!未定义书签。十一、附图 ............................................................... 错误!未定义书签。

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建筑安装工程承包合同(2) 合同编号发包方: 承包方: 根据《中华人民共和国经济合同法》和《建筑安装工程承包合同条例》及有关规定，结合本工程的具体情况，经双方协商一致，签订本合同，以资共同遵守。第一条工程概况 1?工程名称: .工程地点: ?工程计划批准单位及文号: .工程范围和内容：全部工程建筑面积平方米。（各单项工程详见工程项目一览表）第二条工程期限

2.本工程开工日期日。 3 ?如遇下列情况，经发包方现场监理工程师或工程师代表签证后，工期作相应顺延，并用书面形式确定顺延期限。道路、施工用水，或电源未按规定接通，影响承包方进场施工者。 (2) 明确由发包方负责供应的材料、设备、成品或半成品等未能按双方认定的时间进场，或进场的材料、设备、成品或半成品等向承包方交验时发现有缺陷, 需要修配、改、代、换而耽误施工进度者。 (3) 不属包干系数范围内的重大设计变更；提供的工程地质资料不准，使基础超深；施工方法与设计规定不符而增加工程量影响进度者。 (4) 在施工中因停水、停电连续影响8小时以上者。 (5) 发包方现场监理工程师或工程师代表无故拖延办理签证手续而影响下一工序施工者。 1.本工程合同总工期为天(日历天从开工之日算起)。日，竣工日期 (1) 发包方在合同规定开工日期前天，不能交承包方施工场地、进场

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xxx（xx）食品有限公司新建工场模板工程施工方案编制人：审核人： xx—xx建筑有限责任公司 2005-01-18

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三、施工准备 3.1技术准备模板施工前工程技术人员应熟悉图纸，对工人交底透彻。通过理论计算，确定模板的支撑方案。 3.2物质资源准备模板施工前必须保证材料的及时供应，本工程外管线以及厂房中部分梁柱为清水砼，故对模板的要求比较高，应选用平整度高、表面光滑、厚度一致的模板，对机械设备及时检修及购买。 3.3安全技术准备安全交底清楚，进入现场必须佩带安全帽扣好扣带，现场严禁抽烟、大小便，高空作业时必须系好扣好安全带，安全员应检查机械设备、临时用电有无安全隐患，必须作好各项安全防护措施。四、施工方法根据设计图纸放好大样，校对尺寸，然后做好标准样板，根据标准样板划线下料，划好后拉通对角线，检查对角线长度，以保证四角方正。本工程采用厚度为2cm厚的胶合九夹板，每块模板均在标准样板上下料，加工完毕后，都必须按照设计图纸要求以及质量验收标准进行严格检查，质量不合格者应返修。 4.1柱模安装：根据楼地面弹出的柱头方盘线，四周焊接短钢筋，柱边线以内剔毛，沿线外粘好后海棉，先粘后支模，严禁边粘边支模，矩形柱子采用九夹板和二四方料木条组合模板(图一)，用Φ12螺栓对拉固定模板，用短钢管制作柱箍筋，柱箍筋从混凝土面上200mm起每隔400mm 开始设置(图二)。在柱模的中间弹出中线，以检查柱的垂直度。定型模板的启口处要直、光、平，定装时要粘好海棉条后方可安装。柱模上部开设与梁模板连接的梁缺口，底部开设有清理孔，以便清除杂物，当柱子

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建筑工程模板工程施工方案

1.编制依据：1.编制依据 2.工程概况2.1设计概况

2.2结构平面图（详见设计图） 3.施工安排 3.1施工部位及工期要求 3.1.1本工程分为主楼和地下车库两部分，施工按后浇带划分流水段，进行流水施工。 3.1.2工期要求：本工程计划工期2010年6月13日至2012年12月30日，总工期932日历天，由于征地问题迟迟未得到解决，计划2012年8月30日结构封顶。

3.2劳动组织及职责分工 3.2.1模板工程负责人： 3.2.2根据工程特点和工期要求，模板工程中地下室墙、柱、顶板、梁模板需要投入大量的人力，安排具体管理和施工人员如下： 3.2.3模板工程施工人员在技术、生产负责人进行完详细的交底之后才能开始正式施工。模板工程负责人应提前熟悉图纸，明确各结构部位的特点和模板施工的要求，然后组织工人进行下料和拼装。木工应在专业负责人的指导下，按要求搭设模板支撑系统并支设模板，在操作过程中应注意控制各项偏差。普工则负责配合木工完成模板的加工和安装工作，其主要任务是配合木工以保证工序的连续性。 4.施工准备 4.1技术准备 4.1.1相关人员熟悉图纸及设计变更洽商，分析各结构构件的特点以确定使用模板材料和支撑系统，在此过程中注意施工难点和较特殊部位对模板支设的要求。 4.1.2编制施工方案和技术交底等技术资料，用以作为工人施工的依据和质量控制的标准。

4.1.3组织有关管理人员和劳务人员进行培训，组织各级施工人员学习有关技术规范、操作规程、工艺标准、质量验评标准及相关新技术、新材料、新工艺，并组织相应的技术培训。 4.1.4在模板设计过程中着重对梁柱接头、高柱、超高顶板等部位的考虑并制定相应技术措施。 4.2生产准备 4.2.1机具准备施工前准备模板工程所需的各种机械并组织进场和调试以确保正常使用。主要机械列表如下： 4.2.2材料准备 4.2.2.1各种模板材料提前做计划并提交材料部门进行采购和组织进场。 4.2.2.2木方进场要对其规格、材质、外观、含水率等指标进行验收，不合格的不允许进场。 4.2.2.3多层板进场时检查出厂合格证、规格尺寸等内容，以确保与施工要求相一致。 4.2.2.4碗扣架子和钢管扣件、可调顶托、支座等及时租赁进场备用。 4.2.2.5模板、支撑、龙骨、隔离剂采购和进场安排。模板、支撑、龙骨的需要数量见下页表，工程部提供计划,技术质量部控制质量。