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Holin

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Unlike class I and class II holins, which are composed of hydrophobic transmembrane helices, class III holins form a single highly hydrophilic TMD, with the N-terminus in the cytoplasm and the C-terminus in the periplasm. The first class III holin to be characterized was the
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such as the S protein of lambdoid phage 21 act by forming heptameric channels that depolarize the bacterial membrane. They are associated with SAR endolysins, which remain inactive in the periplasm prior to the depolarization of the membrane.
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Class II holins have two TMDs, with both the N- and the C-terminus in the cytoplasm. Their number of residues usually falls between 65 and 95. Examples include the S protein from lambdoid phage 21 and the Hol3626 protein from
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encode holins, making them the most diverse group of proteins with common function. Together with lysins, holins are being studied for their potential use as
476: 467: 144: 503: 45:, a component of bacterial cell walls. Holins have been shown to regulate the timing of lysis with great precision. Over 50 unrelated 1068:"Holins in Bacteria, Eukaryotes, and Archaea: Multifunctional Xenologues with Potential Biotechnological and Biomedical Applications" 708: 97:
in the periplasm and the C-terminus in the cytoplasm. They generally have over 95 residues. Examples of class I holins include the
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There are also several holin families that do not fall into the superfamilies designated above. These families include:
1224: 1111:"Characterization and determination of holin protein of Streptococcus suis bacteriophage SMP in heterologous host" 907:"Characterization and determination of holin protein of Streptococcus suis bacteriophage SMP in heterologous host" 422: 485: 539: 118: 240: 181: 161: 103: 90: 176: 166: 156: 98: 306: 204: 171: 151: 293: 135:-encoded t protein (T4 holin). Other examples include the holins of the ФCP39O and ФCP26F phage. 1109:
Shi, Yibo; Yan, Yaxian; Ji, Wenhui; Du, Bin; Meng, Xiangpeng; Wang, Hengan; Sun, Jianhe (2012).
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Ramanculov E, Young R (March 2001). "Genetic analysis of the T4 holin: timing and topology".
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Wang IN, Smith DL, Young R (2002). "Holins: the protein clocks of bacteriophage infections".
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Communicating Current Research and Educational Topics and Trends in Applied Microbiology
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Viruses that infect eukaryotic cells may use similar channel-forming proteins called
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Young, R.; Bläsi, U. (1995). "Holins: form and function in bacteriophage lysis".
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Veiga-Crespo P; Barros-Velázquez J; Villa T.G. (2007). Méndez-Vilas A (ed.).
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According to their structure there are three main classes of holins.
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Nieva, José Luis; Madan, Vanesa; Carrasco, Luis (2 July 2012).
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Shi Y, Yan Y, Ji W, Du B, Meng X, Wang H, Sun J (March 2012).
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in order to trigger and control the degradation of the host's
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Putative Transglycosylase-associated Holin (T-A Hol) Family
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The Putative Acholeplasma Phage L2 Holin (L2 Holin) Family
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The Mycobacterial Phage PBI1 Gp36 Holin (Gp36 Hol) Family
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The Neisserial Phage-associated Holin (NP-Holin) Family
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The Gene Transfer Agent-release Holin (GTA-Hol) Family
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Spore Morphogenesis and Germination Holin (BSH) Family
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The Putative Lactococcus lactis Holin (LLHol) Family
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The Putative Treponema 4 TMS Holin (Tre4Hol) Family
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The Mycobacterial 1 TMS Phage Holin (M1 Hol) Family
56:While canonical holins act by forming large pores, 1029:Biochimica et Biophysica Acta (BBA) - Biomembranes 549:The Putative Bacterial Archaeal Holin (BAH) Family 513:The Erwinia Phage Phi-Ea1h Holin (EPPE-Hol) Family 468:The Flp/Fap Pilin Putative Holin (FFPP-Hol) Family 329:The Actinobacterial 1 TMS Holin (A-1 Holin) Family 205:The Firmicute phage φU53 Holin (φU53 Holin) Family 147:, there are a total of seven holin superfamilies. 22:are a diverse group of small proteins produced by 849:Nieva, José; Carrasco, Luis (29 September 2015). 531:The Putative Archaeal 2 TMS Holin (A2-Hol) Family 459:The Putative Listeria Phage Holin (LP-Hol) Family 1023:Reddy, Bhaskara L.; Saier Jr, Milton H. (2013). 794:"Viroporins: structure and biological functions" 338:The 2 or 3 TMS Putative Holin (2/3 Holin) Family 276:The Bacterophase Dp-1 Holin (Dp-1 Holin) Family 387:The Putative Holin-like Toxin (Hol-Tox) Family 1066:Saier, Milton H.; Reddy, Bhaskara L. (2015). 619:Gründling A, Manson MD, Young R (July 2001). 8: 954: 952: 504:The Actinobacterial Phage Holin (APH) Family 414:The Xanthomonas Phage Holin (XanPHol) Family 214:The CidA/LrgA Holin (CidA/LrgA Holin) Family 432:The Caulobacter Phage Holin (CauHol) Family 1136: 1126: 1091: 1048: 932: 922: 876: 866: 825: 768: 646: 636: 369:phage P68 Putative Holin (P68 Hol) Family 675:"Bacteriophage Holins: Deadly Diversity" 668: 666: 441:The Enterobacterial Holin (EBHol) Family 1189: 576: 495:The Putative 3 TMS Holin (3-Hol) Family 423:The Prophage Hp1 Holin (Hp1Hol) Family 298:Phage Mu1/6 Holin (Mu1/6 Holin) Family 900: 898: 896: 7: 709:"What can bacteriophages do for us?" 486:The Brachyspira holin (B-Hol) Family 356:The Phage T1 Holin (T1 Holin) Family 1000:Transporter Classification Database 145:Transporter Classification Database 1168:10.1111/j.1574-6976.1995.tb00202.x 540:The Putative Holin-2 (PH-2) Family 320:The SPP1 Holin (SPP1 Holin) Family 285:The BhlA Holin (BhlA Holin) Family 245:Phage r1t Holin (r1t Holin) Family 232:The BlyA Holin (BlyA Holin) Family 223:The ArpQ Holin (ArpQ Holin) Family 14: 1192: 16:Diverse group of small proteins 743:Young, Ryland (1 March 2014). 598:10.1146/annurev.micro.54.1.799 196:The T4 Holin (T4 Holin) Family 1: 973:10.1016/S0378-1119(01)00365-1 682:J. Mol. Microbiol. Biotechnol 621:"Holins kill without warning" 1041:10.1016/j.bbamem.2013.07.004 625:Proc. Natl. Acad. Sci. U.S.A 101:S protein (λ holin) and the 798:Nature Reviews Microbiology 311:Holin (Vibrio Holin) Family 1241: 89:Class I holins have three 1156:FEMS Microbiology Reviews 761:10.1007/s12275-014-4087-z 718:: 885–893. Archived from 296:Streptomyces aureofaciens 107:phage P68 hol15 protein. 673:Young R (January 2002). 1072:Journal of Bacteriology 749:Journal of Microbiology 119:Clostridium perfringens 1128:10.1186/1743-422x-9-70 924:10.1186/1743-422X-9-70 638:10.1073/pnas.151247598 1220:Protein superfamilies 182:Holin superfamily VII 162:Holin superfamily III 122:bacteriophage Ф3626. 104:Staphylococcus aureus 91:transmembrane domains 41:to reach and degrade 177:Holin superfamily VI 167:Holin superfamily IV 157:Holin superfamily II 1084:10.1128/JB.02046-14 810:10.1038/nrmicro2820 172:Holin superfamily V 152:Holin superfamily I 996:"TC-Superfamilies" 586:Annu Rev Microbiol 243:Lactococcus lactis 33:at the end of the 1225:Membrane proteins 1035:(11): 2654–2671. 861:(10): 5169–5171. 631:(16): 9348–9352. 143:According to the 1232: 1197: 1196: 1188: 1179: 1162:(1–2): 191–205. 1150: 1140: 1130: 1115:Virology Journal 1105: 1095: 1062: 1052: 1011: 1010: 1008: 1006: 991: 985: 984: 956: 947: 946: 936: 926: 911:Virology Journal 902: 891: 890: 880: 870: 868:10.3390/v7102866 846: 840: 839: 829: 789: 783: 782: 772: 740: 734: 733: 731: 730: 724: 713: 704: 698: 697: 679: 670: 661: 660: 650: 640: 616: 610: 609: 581: 133:bacteriophage T4 126:Class III holins 93:(TMDs) with the 1240: 1239: 1235: 1234: 1233: 1231: 1230: 1229: 1205: 1204: 1203: 1191: 1183: 1153: 1108: 1065: 1022: 1019: 1017:Further reading 1014: 1004: 1002: 993: 992: 988: 958: 957: 950: 904: 903: 894: 848: 847: 843: 791: 790: 786: 742: 741: 737: 728: 726: 722: 711: 706: 705: 701: 677: 672: 671: 664: 618: 617: 613: 583: 582: 578: 574: 557: 141: 128: 113: 111:Class II holins 99:bacteriophage λ 87: 79: 74: 17: 12: 11: 5: 1238: 1236: 1228: 1227: 1222: 1217: 1207: 1206: 1202: 1201: 1181: 1180: 1151: 1106: 1063: 1018: 1015: 1013: 1012: 986: 967:(1–2): 25–36. 948: 892: 841: 804:(8): 563–574. 784: 755:(3): 243–258. 735: 699: 662: 611: 575: 573: 570: 569: 568: 563: 556: 553: 552: 551: 542: 533: 524: 515: 506: 497: 488: 479: 470: 461: 452: 443: 434: 425: 416: 407: 398: 389: 380: 371: 367:Staphylococcus 358: 349: 340: 331: 322: 313: 300: 287: 278: 269: 256: 247: 234: 225: 216: 207: 198: 185: 184: 179: 174: 169: 164: 159: 154: 140: 137: 127: 124: 112: 109: 86: 85:Class I holins 83: 78: 75: 73: 72:Classification 70: 27:bacteriophages 15: 13: 10: 9: 6: 4: 3: 2: 1237: 1226: 1223: 1221: 1218: 1216: 1213: 1212: 1210: 1200: 1195: 1190: 1186: 1177: 1173: 1169: 1165: 1161: 1157: 1152: 1148: 1144: 1139: 1134: 1129: 1124: 1120: 1116: 1112: 1107: 1103: 1099: 1094: 1089: 1085: 1081: 1077: 1073: 1069: 1064: 1060: 1056: 1051: 1046: 1042: 1038: 1034: 1030: 1026: 1021: 1020: 1016: 1001: 997: 990: 987: 982: 978: 974: 970: 966: 962: 955: 953: 949: 944: 940: 935: 930: 925: 920: 916: 912: 908: 901: 899: 897: 893: 888: 884: 879: 874: 869: 864: 860: 856: 852: 845: 842: 837: 833: 828: 823: 819: 815: 811: 807: 803: 799: 795: 788: 785: 780: 776: 771: 766: 762: 758: 754: 750: 746: 739: 736: 725:on 2016-03-03 721: 717: 710: 703: 700: 695: 691: 687: 683: 676: 669: 667: 663: 658: 654: 649: 644: 639: 634: 630: 626: 622: 615: 612: 607: 603: 599: 595: 591: 587: 580: 577: 571: 567: 564: 562: 559: 558: 554: 550: 546: 543: 541: 537: 534: 532: 528: 525: 523: 519: 516: 514: 510: 507: 505: 501: 498: 496: 492: 489: 487: 483: 480: 478: 474: 471: 469: 465: 462: 460: 456: 453: 451: 447: 444: 442: 438: 435: 433: 429: 426: 424: 420: 417: 415: 411: 408: 406: 402: 399: 397: 393: 390: 388: 384: 381: 379: 375: 372: 370: 368: 362: 359: 357: 353: 350: 348: 344: 341: 339: 335: 332: 330: 326: 323: 321: 317: 314: 312: 310: 304: 301: 299: 297: 291: 288: 286: 282: 279: 277: 273: 270: 268: 266: 260: 257: 255: 251: 248: 246: 244: 238: 235: 233: 229: 226: 224: 220: 217: 215: 211: 208: 206: 202: 199: 197: 193: 190: 189: 188: 183: 180: 178: 175: 173: 170: 168: 165: 163: 160: 158: 155: 153: 150: 149: 148: 146: 139:Gene families 138: 136: 134: 125: 123: 121: 120: 110: 108: 106: 105: 100: 96: 92: 84: 82: 76: 71: 69: 67: 62: 59: 54: 52: 51:antibacterial 48: 47:gene families 44: 43:peptidoglycan 40: 36: 32: 28: 25: 21: 1159: 1155: 1118: 1114: 1075: 1071: 1032: 1028: 1003:. 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Index

dsDNA
bacteriophages
cell wall
lytic cycle
lysins
peptidoglycan
gene families
antibacterial
viroporins
transmembrane domains
N-terminus
bacteriophage λ
Staphylococcus aureus
Clostridium perfringens
bacteriophage T4
Transporter Classification Database
Holin superfamily I
Holin superfamily II
Holin superfamily III
Holin superfamily IV
Holin superfamily V
Holin superfamily VI
Holin superfamily VII
1.E.8
The T4 Holin (T4 Holin) Family
1.E.13
The Firmicute phage φU53 Holin (φU53 Holin) Family
1.E.14
The CidA/LrgA Holin (CidA/LrgA Holin) Family
1.E.15

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