ATPase-like domain of the ASKHA (Acetate and Sugar Kinases/Hsc70/Actin) superfamily; The ASKHA superfamily, also known as actin-like ATPase domain superfamily, includes acetate and sugar kinases, heat-shock cognate 70 (Hsp70) and actin family proteins. They either function as conformational hydrolases (e.g. Hsp70, actin) that perform simple ATP hydrolysis, or as metabolite kinases (e.g. glycerol kinase) that catalyze the transfer of a phosphoryl group from ATP to their cognate substrates. Both activities depend on the presence of specific metal cations. ASKHA superfamily members share a common core fold that includes an actin-like ATPase domain consisting of two subdomains (denoted I _ II) with highly similar ribonuclease (RNase) H-like folds. The fold of each subdomain is characterized by a central five strand beta-sheet and flanking alpha-helices. The two subdomains form an active site cleft in which ATP binds at the bottom. Another common feature of ASKHA superfamily members is the coupling of phosphoryl-group transfer to conformational rearrangement, leading to domain closure. Substrate binding triggers protein motion.

                          10        20        30        40        50        60        70        80
                  ....*....|....*....|....*....|....*....|....*....|....*....|....*....|....*....|
gi 1958763153 194 GIAETLHYVLDRYPKAVQDTLVQNVFLTGGNVMYPGMKARVEKELLEMRPFQSSFQVQLASNPVLDAWYGARDWALDHlE 273
Cdd:cd10211   245 GLVETIEFVLKRYPAEQQDRLVQNVFLTGGNALFPGLKERLEKELRAIRPFGSPFNVVRAKDPVLDAWRGAAKWALDS-T 323

                          90       100
                  ....*....|....*....|..
gi 1958763153 274 DSGAWVTRKDYEEKGGEYLKEH 295
Cdd:cd10211   324 FEKVWITKQEYEEKGGEYLKEH 345

nucleotide-binding domain (NBD) of actin-related protein5 (Arp5) and similar proteins; Arp5, also called actin-like protein 5, may act as a core component of the chromatin remodeling INO80 complex which is involved in transcriptional regulation, DNA replication and probably DNA repair. It is involved in DNA double-strand break repair and UV-damage excision repair. Human Arp5 is encoded by the ACTR5 gene. Arabidopsis thaliana ARP5 (AtARp5) is a ubiquitously expressed nuclear protein involved in DNA repair and required for multicellular development of all organs. AtARp5 may be part of other chromatin remodeling machines in addition to INO80.

                          10        20        30        40        50        60        70        80
                  ....*....|....*....|....*....|....*....|....*....|....*....|....*....|....*....|
gi 1958763153 194 GIAETLHYVLDRYPKAVQDTLVQNVFLTGGNVMYPGMKARVEKELLEMRPFQSSFQVQLASNPVLDAWYGARDWALDHlE 273
Cdd:cd10211   245 GLVETIEFVLKRYPAEQQDRLVQNVFLTGGNALFPGLKERLEKELRAIRPFGSPFNVVRAKDPVLDAWRGAAKWALDS-T 323

                          90       100
                  ....*....|....*....|..
gi 1958763153 274 DSGAWVTRKDYEEKGGEYLKEH 295
Cdd:cd10211   324 FEKVWITKQEYEEKGGEYLKEH 345

Actin;

                          10        20        30        40        50        60        70        80
                  ....*....|....*....|....*....|....*....|....*....|....*....|....*....|....*....|
gi 1958763153 166 YHQLFvGTERIRAPEIIFQPSLIGEEQA--------GIAETLHYVLDRYPKAVQDTLVQNVFLTGGNVMYPGMKARVEKE 237
Cdd:pfam00022 267 STIIL-GAERFRVPEILFNPSLIGSESElpppqtavGIPELIVDAINACDVDLRPSLLANIVVTGGNSLFPGFTERLEKE 345

                          90       100       110       120       130
                  ....*....|....*....|....*....|....*....|....*....|....*..
gi 1958763153 238 LleMRPFQSSFQVQLASNPVLD-----AWYGARDWA-LDHLEDSgaWVTRKDYEEKG 288
Cdd:pfam00022 346 L--AQLAPPGVKVKIIAPGNTVerrysAWIGGSILAsLGTFQQM--WVSKQEYEEHG 398

Actin-related protein [Cytoskeleton];

                          10        20        30        40        50        60        70        80
                  ....*....|....*....|....*....|....*....|....*....|....*....|....*....|....*....|
gi 1958763153 174 ERIRAPEIIFQPSLIGEE----------------------QAGIAETLHYVLDRYPKAVQDTLVQNVFLTGGNVMY---P 228
Cdd:COG5277   279 ERFLIGEILFNPNHEGFEsyiqqgrlriedavigdvvlygEMGLAEAIINSIMKCDVEIQDELYSNIILSGGAFNWsvpP 358

                          90       100       110       120       130       140
                  ....*....|....*....|....*....|....*....|....*....|....*....|....*..
gi 1958763153 229 GMK-------ARVEKELLEMRPfQSSFQVQLASNPVLDAWYGARDWALDhLEDSGAW--VTRKDYEE 286
Cdd:COG5277   359 GLEdvavdsvTRVQIELSELAP-ELKVNVRLVSDPQYSVWKGAIIYGYA-LPFSVKWswITKEGWYF 423

nucleotide-binding domain (NBD) of actin-related protein5 (Arp5) and similar proteins; Arp5, also called actin-like protein 5, may act as a core component of the chromatin remodeling INO80 complex which is involved in transcriptional regulation, DNA replication and probably DNA repair. It is involved in DNA double-strand break repair and UV-damage excision repair. Human Arp5 is encoded by the ACTR5 gene. Arabidopsis thaliana ARP5 (AtARp5) is a ubiquitously expressed nuclear protein involved in DNA repair and required for multicellular development of all organs. AtARp5 may be part of other chromatin remodeling machines in addition to INO80.

                          10        20        30        40        50        60        70        80
                  ....*....|....*....|....*....|....*....|....*....|....*....|....*....|....*....|
gi 1958763153 194 GIAETLHYVLDRYPKAVQDTLVQNVFLTGGNVMYPGMKARVEKELLEMRPFQSSFQVQLASNPVLDAWYGARDWALDHlE 273
Cdd:cd10211   245 GLVETIEFVLKRYPAEQQDRLVQNVFLTGGNALFPGLKERLEKELRAIRPFGSPFNVVRAKDPVLDAWRGAAKWALDS-T 323

                          90       100
                  ....*....|....*....|..
gi 1958763153 274 DSGAWVTRKDYEEKGGEYLKEH 295
Cdd:cd10211   324 FEKVWITKQEYEEKGGEYLKEH 345

Actin; ACTIN subfamily of ACTIN/mreB/sugarkinase/Hsp70 superfamily

                           10        20        30        40        50        60        70        80
                   ....*....|....*....|....*....|....*....|....*....|....*....|....*....|....*....|
gi 1958763153  161 FNLAAYHQLFVGTERIRAPEIIFQPSLIGEEQAGIAETLHYVLDRYPKAVQDTLVQNVFLTGGNVMYPGMKARVEKELLE 240
Cdd:smart00268 238 YELPDGNTIKVGNERFRIPEILFSPELIGLEQKGIHELVYESIQKCDIDVRKDLYENIVLSGGSTLIPGFGERLEKELKQ 317

                           90       100       110       120
                   ....*....|....*....|....*....|....*....|....*....
gi 1958763153  241 MRPFQSSFQVQLASNPVLDAWYGARDWA-LDHLEDSgaWVTRKDYEEKG 288
Cdd:smart00268 318 LAPKKLKVKVIAPPERKYSVWLGGSILAsLSTFEDM--WITKKEYEESG 364

Actin;

                          10        20        30        40        50        60        70        80
                  ....*....|....*....|....*....|....*....|....*....|....*....|....*....|....*....|
gi 1958763153 166 YHQLFvGTERIRAPEIIFQPSLIGEEQA--------GIAETLHYVLDRYPKAVQDTLVQNVFLTGGNVMYPGMKARVEKE 237
Cdd:pfam00022 267 STIIL-GAERFRVPEILFNPSLIGSESElpppqtavGIPELIVDAINACDVDLRPSLLANIVVTGGNSLFPGFTERLEKE 345

                          90       100       110       120       130
                  ....*....|....*....|....*....|....*....|....*....|....*..
gi 1958763153 238 LleMRPFQSSFQVQLASNPVLD-----AWYGARDWA-LDHLEDSgaWVTRKDYEEKG 288
Cdd:pfam00022 346 L--AQLAPPGVKVKIIAPGNTVerrysAWIGGSILAsLGTFQQM--WVSKQEYEEHG 398

nucleotide-binding domain (NBD) of actin and similar proteins; Actin is a ubiquitous protein involved in the formation of filaments that are major components of the cytoskeleton. It is a highly dynamic structural protein network involved in processes such as cell contraction, cell motility, vesicle trafficking, intracellular organization, cytokinesis, endocytosis and apoptosis. Actin is a monomeric globular protein (G-actin) that reversibly polymerizes to form filaments (F-actin). Each actin protomer binds one molecule of ATP and either calcium or magnesium ions. At low salt concentrations, actin exists as a monomer, and as the salt concentration rises F-actin forms, with the consequent hydrolysis of ATP. F-actin assembly is in constant flux with G-actin association occurring at the barbed end (+) and its disassociation at the pointed end (-). Actin monomers that have been released from the pointed end can be reused, if the ADP is exchanged for ATP. F-actin filaments can assemble into higher order structures, for example branched F-actin, and stress fibers. Actin binding proteins regulate actin filament dynamics by a range of functions including actin severing, depolymerizing, capping, stabilizing and de novo actin polymerization. Actins interaction with myosin is the basis of muscular contraction and many aspects of cell motility. In vertebrates there are three main groups of actin isoforms, alpha, beta and gamma. The alpha actins found in muscle tissues are a major constituent of the contractile apparatus. The beta and gamma actins co-exist in most cell types as components of the cytoskeleton and as mediators of internal cell motility. In plants there are many isoforms which are probably involved in a variety of functions such as cytoplasmic streaming, cell shape determination, tip growth, graviperception, cell wall deposition, etc.

                          10        20        30        40        50        60        70        80
                  ....*....|....*....|....*....|....*....|....*....|....*....|....*....|....*....|
gi 1958763153 147 EVEKPQVTTTVQPVFNLAAYHQLFVGTERIRAPEIIFQPSLIGEEQAGIAETLHYVLDRYPKAVQDTLVQNVFLTGGNVM 226
Cdd:cd10224   221 EMQTAASSSSLEKSYELPDGQVITIGNERFRCPEALFQPSFLGMEAAGIHETTYNSIMKCDVDIRKDLYANIVLSGGTTM 300

                          90       100       110       120       130       140
                  ....*....|....*....|....*....|....*....|....*....|....*....|....*
gi 1958763153 227 YPGMKARVEKELLEMRPfqSSFQVQLASNP--VLDAWYGARDWA-LDHLEDsgAWVTRKDYEEKG 288
Cdd:cd10224   301 FPGIADRMQKEITALAP--STMKIKIVAPPerKYSVWIGGSILAsLSTFQQ--MWISKQEYDESG 361

nucleotide-binding domain (NBD) of actin-related protein2 (Arp2) and similar proteins; Arp2, also called actin-like protein 2, is the ATP-binding component of the Arp2/3 complex, a multiprotein complex that mediates actin polymerization upon stimulation by nucleation-promoting factor (NPF). The Arp2/3 complex is comprised of 7 proteins (Arp2, Arp3, and five conserved proteins, ARPC1-5). It generates cytoplasmic branched filaments networks, by promoting nucleation of actin filaments as 70 degrees branches on the side of older filaments. It is activated, by simultaneously binding to a pre-existing filament and a nucleation promoting factor plus an actin monomer. Daughter branches subsequently detach/debranch from the mother filament. Its Arp2 and Arp3 subunits must be loaded with ATP for it to initiate the assembly of branched actin filaments. ATP hydrolysis may be required for branch initiation or debranching. The Arp2/3 complex is also found in the nucleus where it plays a role in promoting de novo actin polymerization and in RNA polymerase II-dependent transcription. This may in part be through regulating nuclear actin polymerization in a way like its function in the cytoplasm. Human Arp2 is encoded by the ACTR2 gene.

                          10        20        30        40        50        60        70        80
                  ....*....|....*....|....*....|....*....|....*....|....*....|....*....|....*....|
gi 1958763153 171 VGTERIRAPEIIFQPSLIGEEQAGIAETLHYVLDRYPKAVQDTLVQNVFLTGGNVMYPGMKARVEKELLEM--------- 241
Cdd:cd10220   247 VGGERFEAPEALFQPHLIDVEGPGIAELLFNTIQAADIDTRPELYKHIVLSGGSTMYPGLPSRLEKEIKQLylervlkgd 326

                          90       100       110       120       130       140
                  ....*....|....*....|....*....|....*....|....*....|....*....|....
gi 1958763153 242 RPFQSSFQVQLASNP-----------VLDawygardwalDHLEDSGA-WVTRKDYEEKGGEYLK 293
Cdd:cd10220   327 TERLSKFKIRIEDPPrrkhmvflggaVLA----------DIMKDKDEfWITRQEYEEQGVRVLD 380

nucleotide-binding domain (NBD) of the actin-like protein 7 (ACTL7)-like family; The ACTL7-like family includes ACTL7A, ACTL7B and ACTL9 (also known as ACTL7C). In mammalian, ACTL7A is expressed in a wide variety of adult tissues, while the ACTL7B is expressed in spermatids through the elongation phase of spermatid development. ACTL7A, also called actin-like-7-alpha, or T-ACTIN-2 in mouse, may play an important role in formation and fusion of Golgi-derived vesicles during acrosome biogenesis. ACTL7B, also called actin-like-7-beta, acts as a key regulator of spermiogenesis that is required for male fertility. ACTL9 is a testis-specific protein that plays an important role in fusion of proacrosomal vesicles and perinuclear theca formation.

                          10        20        30        40        50        60        70        80
                  ....*....|....*....|....*....|....*....|....*....|....*....|....*....|....*....|
gi 1958763153 167 HQLFVGTERIRAPEIIFQPSLIGEEQAGIAETLHYVLDRYPKAVQDTLVQNVFLTGGNVMYPGMKARVEKELLEMRPfqs 246
Cdd:cd10214   241 HLITIGKERFRCPEMLFNPSLIGSKQPGLHTLTMNSLNKCDANLKKDLAKNILLCGGSTMFDGFPDRFQKELSKLCP--- 317

                          90       100       110       120       130
                  ....*....|....*....|....*....|....*....|....*....|....
gi 1958763153 247 sfqvqlASNPVLDA--------WYG----ARDWALDHLedsgaWVTRKDYEEKG 288
Cdd:cd10214   318 ------NDNPIVAAsperkysvWTGgsilASLKSFQQL-----WVRRREYEERG 360

nucleotide-binding domain (NBD) of actin-related protein 1 (Arp1) and similar proteins; Arp1, also called centractin, actin-like protein, alpha-centractin, actin-RPV, or centrosome-associated actin homolog, may be a component of a multi-subunit centrosomal complex involved in microtubule-based vesicle motility. In yeast, actin-related protein is essential for viability and is associated with the centrosome. In vertebrates, Arp1 is a core component of the dynactin complex which assists cytoplasmic dynein by increasing its processivity and by regulation of its cargo binding. The dynactin complex is required for the spindle translocation late in anaphase and is involved in a cell wall synthesis checkpoint. ARP1 forms the backbone filament of the dynactin rod structure and serves as the scaffold for the remaining subunits. It is required for proper orientation of the mitotic spindle. Arp1 is the only actin-related protein known to form actin-like filaments. Human Arp1/centractin is encoded by the ACTR1A gene.

                          10        20        30        40        50        60        70        80
                  ....*....|....*....|....*....|....*....|....*....|....*....|....*....|....*....|
gi 1958763153 143 EETPEVEKPQVTTtvqpvFNLAAYHQLFVGTERIRAPEIIFQPSLIGEEQAGIAETLHYVLDRYPKAVQDTLVQNVFLTG 222
Cdd:cd10216   223 EEKLEEEKTEKAQ-----YTLPDGSTIEIGPERFRAPEILFNPELIGLEYPGVHEVLVDSIQKSDLDLRKTLYSNIVLSG 297

                          90       100       110       120       130       140
                  ....*....|....*....|....*....|....*....|....*....|....*....|....*..
gi 1958763153 223 GNVMYPGMKARVEKELLEMRPFQSSFQVQLASNPVLDAWYGARDWA-LDHLEDsgAWVTRKDYEEKG 288
Cdd:cd10216   298 GSTLFKGFGDRLLSEVKKLAPKDVKIRISAPPERLYSTWIGGSILAsLSTFKK--MWVSKKEYEEDG 362

nucleotide-binding domain (NBD) of the actin-related protein 4 (Arp4)-like subfamily; The Arp4-like subfamily includes Arp4, also called actin-like protein 4, from fungi and plants. Saccharomyces cerevisiae Arp4 acts synergistically with Arp8 to depolymerize F-actin; it binds ATP, but unlike conventional actin, does not form filaments. It is a component of the NuA4 histone acetyltransferase complex, the chromatin-remodeling INO80 complex and the SWR1 chromatin remodeling complex. Arabidopsis thaliana Arp4 is involved in several developmental processes including organization of plant organs, flowering time, anther development, flower senescence and fertility, probably by regulating the chromatin structure. This family also includes human homologs of yeast and plant, which are actin-like protein 6A (encoded by the ACTL6A gene; also known as ArpNbeta, 53 kDa BRG1-associated factor A/BRG1-associated factor 53A/BAF35A, and INO80 complex subunit K/INO80K) and actin-like protein 6B (encoded by the ACTL6B gene; also known as ArpNalpha, 53 kDa BRG1-associated factor B/BRG1-associated factor 53B/BAF35B). ACTL6A and ACTL6B are involved in transcriptional activation and repression of select genes by chromatin remodeling (alteration of DNA-nucleosome topology). They are components of numerous complexes with chromatin remodeling and histone acetyltransferase activity. ACTL6A is also a putative core component of the chromatin remodeling INO80 complex which is involved in transcriptional regulation, DNA replication and probably DNA repair. Schizosaccharomyces pombe actin-related protein 42 (Arp42) is also included in this family. It is also a component of SWI/SNF and RSC complexes.

                          10        20        30        40        50        60        70        80
                  ....*....|....*....|....*....|....*....|....*....|....*....|....*....|....*....|
gi 1958763153 166 YHQLFvGTERIRAPEIIFQPSLIGEEQAGIAETLHY------VLDrypkAVQ--D-----TLVQNVFLTGGNVMYPGMKA 232
Cdd:cd13395   281 YNIEF-GAERFKIPELLFDPSLVKGIPAPPSEGNELlglpqlVYT----SIGscDvdirpELYGNVVLTGGNSLLPGFTD 355

                          90       100       110       120       130       140       150
                  ....*....|....*....|....*....|....*....|....*....|....*....|....*....|.
gi 1958763153 233 RVEKELLEMRPFQSSFQVqLASNPVLD----AWYGardwaldhledsG-----------AWVTRKDYEEKG 288
Cdd:cd13395   356 RLNRELSEKAPGSLKLKI-LASGNTVErrfsSWIG------------GsilaslgsfqqMWISKQEYEEHG 413

nucleotide-binding domain (NBD) of Saccharomyces cerevisiae actin-related protein7 (Arp7) and similar proteins; Saccharomyces cerevisiae Arp7, also called actin-like protein 7, is a component of the chromatin structure remodeling complex (RSC), which is involved in transcription regulation and nucleosome positioning. It is also part of the SWI/SNF complex, an ATP-dependent chromatin remodeling complex, which is required for the positive and negative regulation of gene expression of many genes. Arp7 forms a stable heterodimer with Arp9 protein in both the RSC and SWI/SNF chromatin-remodeling complexes. It has been suggested that this dimer functions as a module with DNA bending proteins, to achieve correct architecture and facilitate complex-complex interactions. Fission yeast SWI/SNF and RSC complexes do not contain Arp7 and Arp8, but instead contain Arp9 and Arp42.

                          10        20        30        40        50        60        70        80
                  ....*....|....*....|....*....|....*....|....*....|....*....|....*....|....*....|
gi 1958763153  64 MDQFHKALIELNMDSPEELQSYIQKLTLAVEQAKQKILQAEANLEVDVVDSKPETPdleplepsLEDVENINdFEPLfse 143
Cdd:cd10212   226 IQQFKSTMLQVSEKDLFELERYYKEQADIYAKQQEQLKQMDQQLQYTALTGSPNNP--------LVQKKNFL-FKPL--- 293

                          90       100       110       120       130       140       150       160
                  ....*....|....*....|....*....|....*....|....*....|....*....|....*....|....*....|
gi 1958763153 144 etpeveKPQVTTTVQPVFNLAAYhqlfvgterirapeiIFQPSLIGEE---QAGIAETLHYVLDRYPKAVQDTLVQNVFL 220
Cdd:cd10212   294 ------NKTLTLDLKECYQFAEY---------------LFKPQLISDKfspEDGLGPLMAKSVKKAPEQVYSLLLTNVII 352

                         170       180       190       200       210       220
                  ....*....|....*....|....*....|....*....|....*....|....*....|....*..
gi 1958763153 221 TGGNVMYPGMKARVEKELLEMRPFQ--SSFQVQLASNPVLDAWYGARDWALDHLEDSGAWVTRKDYE 285
Cdd:cd10212   353 TGSTSLIEGMEQRIIKELSIRFPQYklTTFANQVMMDRKIQGWLGALTMANLPSWSLGKWYSKEDYE 419

Actin-related protein 3; Provisional

                          10        20        30        40        50        60        70        80
                  ....*....|....*....|....*....|....*....|....*....|....*....|....*....|....*....|
gi 1958763153 171 VGTERIRAPEIIFQPSLIGEE-QAGIAETLHYVLDRYPKAVQDTLVQNVFLTGGNVMYPGMKARVEK------------- 236
Cdd:PTZ00280  268 VGYERFLGPEMFFHPEIFSSEwTTPLPEVVDDAIQSCPIDCRRPLYKNIVLSGGSTMFKGFDKRLQRdvrkrvdrrlkka 347

                          90       100       110       120       130
                  ....*....|....*....|....*....|....*....|....*....|....*
gi 1958763153 237 -ELLEMRPFQSSFQVQLASNPV--LDAWYGARDWALDHlEDSGAWVTRKDYEEKG 288
Cdd:PTZ00280  348 eELSGGKLKPIPIDVNVVSHPRqrYAVWYGGSMLASSP-EFEKVCHTKAEYDEYG 401

nucleotide-binding domain (NBD) of actin-related protein3 (Arp3) and similar proteins; Arp3, also called actin-like protein 3, is the ATP-binding component of the Arp2/3 complex, a multiprotein complex that mediates actin polymerization upon stimulation by nucleation-promoting factor (NPF). The Arp2/3 complex is comprised of 7 proteins (Arp2, Arp3, and five conserved proteins, ARPC1-5). It generates cytoplasmic branched filaments networks, by promoting nucleation of actin filaments as 70 degrees branches on the side of older filaments. It is activated, by simultaneously binding to a pre-existing filament and a nucleation promoting factor plus an actin monomer. Daughter branches subsequently detach/debranch from the mother filament. Its Arp2 and Arp3 subunits must be loaded with ATP for it to initiate the assembly of branched actin filaments. ATP hydrolysis may be required for branch initiation or debranching. The Arp2/3 complex is also found in the nucleus where it plays a role in promoting de novo actin polymerization and in RNA polymerase II-dependent transcription. This may in part be through regulating nuclear actin polymerization in a way like its function in the cytoplasm. Human Arp3 and Arp3B are encoded by the ACTR3 and ACTR3B genes respectively. Arp3B is also known as actin-related protein Arp4.

                          10        20        30        40        50        60        70        80
                  ....*....|....*....|....*....|....*....|....*....|....*....|....*....|....*....|
gi 1958763153 171 VGTERIRAPEIIFQPSLIGEE-QAGIAETLHYVLDRYPKAVQDTLVQNVFLTGGNVMYPG--------MKARVEK----- 236
Cdd:cd10221   265 VGYERFLAPEIFFNPEIASSDfTTPLPEVVDQVIQSCPIDTRRGLYKNIVLSGGSTMFKDfgrrlqrdVKRIVDArlkas 344

                          90       100       110       120       130
                  ....*....|....*....|....*....|....*....|....*....|....*...
gi 1958763153 237 -ELLEMRPFQSSFQVQLASNP-----VldaWYGARDWALDHlEDSGAWVTRKDYEEKG 288
Cdd:cd10221   345 eELSGGKLKPKPIDVNVISHPmqryaV---WFGGSMLASTP-EFYTVCHTKAEYEEYG 398