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Viral evolution

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when an RNA virus replicates in its host cell, changes in their genes are occasionally introduced in error, some of which are lethal. One virus particle can produce millions of progeny viruses in just one cycle of replication, therefore the production of a few "dud" viruses is not a problem. Most mutations are "silent" and do not result in any obvious changes to the progeny viruses, but others confer advantages that increase the fitness of the viruses in the environment. These could be changes to the virus particles that disguise them so they are not identified by the cells of the
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synthesis. These processes likely evolved very early. The adaptive features of the enzymes catalyzing these early processes may have been maintained in the phage T4, bacterial, and eukaryotic lineages because they were established well-tested solutions to basic functional problems by the time these
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are similar suggesting that phage T4 shares ancestry with both bacteria and eukaryotes and has about equal similarity to each. Phage T4 may have diverged in evolution from a common ancestor of bacteria and eukaryotes or from an early evolved member of either lineage. Most of the phage genes showing
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for the origin of viruses was proposed in 2019. According to this hypothesis, the replication modules of viruses originated from the primordial genetic pool, although the long course of their subsequent evolution involved many displacements by replicative genes from their cellular hosts. By contrast,
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perspective, it has therefore been proposed that on celestial bodies such as Mars not only cells but also traces of former virions or viroids should be actively searched for: possible findings of traces of virions in the apparent absence of cells could provide support for the virus-first hypothesis.
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One of the problems for studying viral origins and evolution is the high rate of viral mutation, particularly the case in RNA retroviruses like HIV/AIDS. A recent study based on comparisons of viral protein folding structures, however, is offering some new evidence. Fold Super Families (FSFs) are
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Through these mechanisms new viruses are constantly emerging and present a continuing challenge in attempts to control the diseases they cause. Most species of viruses are now known to have common ancestors, and although the "virus first" hypothesis has yet to gain full acceptance, there is little
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are especially prone to mutations. In host cells there are mechanisms for correcting mistakes when DNA replicates and these kick in whenever cells divide. These important mechanisms prevent potentially lethal mutations from being passed on to offspring. But these mechanisms do not work for RNA and
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similar to viroids but has a protein coat derived from hepatitis B virus and cannot produce one of its own. It is, therefore, a defective virus. Although hepatitis delta virus genome may replicate independently once inside a host cell, it requires the help of hepatitis B virus to provide a protein
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are living cells that, like viruses, can reproduce only inside host cells. They lend support to this hypothesis, as their dependence on parasitism is likely to have caused the loss of genes that enabled them to survive outside a cell. This is also called the "degeneracy hypothesis", or "reduction
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allows viruses to quickly adapt to changes in their host environment. In addition, because viruses typically produce many copies in an infected host, mutated genes can be passed on to many offspring quickly. Although the chance of mutations and evolution can change depending on the type of virus
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Definitive exclusion of a hypothesis on the origin of viruses is difficult to make on Earth given the ubiquitous interactions between viruses and cells, and the lack of availability of rocks that are old enough to reveal traces of the earliest viruses on the planet. From an
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the genes encoding major structural proteins evolved from functionally diverse host proteins throughout the evolution of the virosphere. This scenario is distinct from each of the three traditional scenarios but combines features of the Virus-first and Escape hypotheses.
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Viruses have been able to continue their infectious existence due to evolution. Their rapid mutation rates and natural selection has given viruses the advantage to continue to spread. One way that viruses have been able to spread is with the evolution of virus
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The evolutionary history of viruses can to some extent be inferred from analysis of contemporary viral genomes. The mutation rates for many viruses have been measured, and application of a
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Viruses evolve through changes in their RNA (or DNA), some quite rapidly, and the best adapted mutants quickly outnumber their less fit counterparts. In this sense their evolution is
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are molecules of RNA that are not classified as viruses because they lack a protein coat. They have characteristics that are common to several viruses and are often called
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per round of replication). Although most viral mutations confer no benefit and often even prove deleterious to viruses, the rapid rate of viral mutation combined with
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Many viruses (for example, influenza A virus) can "shuffle" their genes with other viruses when two similar strains infect the same cell. This phenomenon is called
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There are three classical hypotheses on the origins of viruses and how they evolved: There are three main hypotheses that aim to explain the origins of viruses:
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La Scola B, Desnues C, Pagnier I, Robert C, Barrassi L, Fournous G, et al. (September 2008). "The virophage as a unique parasite of the giant mimivirus".
430:. Phage T4 is an obligate intracellular parasite that reproduces within the host bacterial cell and its progeny are released when the host is destroyed by 339:. The way viruses reproduce in their host cells makes them particularly susceptible to the genetic changes that help to drive their evolution. The 553:(transmission between members of the same species that don't have a parent-child relationship) will usually evolve to have a higher virulence. 245:
proteins that show similar folding structures independent of the actual sequence of amino acids, and have been found to show evidence of viral
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Some viruses may have evolved from bits of DNA or RNA that "escaped" from the genes of a larger organism. The escaped DNA could have come from
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often appear within weeks or months after the beginning of a treatment. One of the main theoretical models applied to viral evolution is the
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Lam TT, Hon CC, Tang JW (February 2010). "Use of phylogenetics in the molecular epidemiology and evolutionary studies of viral infections".
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Viruses are ancient. Studies at the molecular level have revealed relationships between viruses infecting organisms from each of the
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are a valuable source of retrospective evidence about the evolutionary history of viruses, and have given birth to the science of
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This is also called the "virus-first hypothesis" and proposes that viruses may have evolved from complex molecules of protein and
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at the same time that cells first appeared on Earth and would have been dependent on cellular life for billions of years.
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Krupovic M, Dolja VV, Koonin EV (July 2019). "Origin of viruses: primordial replicators recruiting capsids from hosts".
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fragments forming sedimentary rocks that fossilize plants and animals. However, the genomes of many organisms contain
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doubt that the thousands of species of modern viruses have evolved from less numerous ancient ones. The
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may have new experimental evidence, with a long intermediary period in the evolution of cellular life.
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Waterborne transmission- leaving a host, infecting the water, and being consumed in a new host
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Based on the analyses of the evolution of the replicative and structural modules of viruses, a
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homology with bacteria and eukaryotes encode enzymes acting in the ubiquitous processes of
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in 1950. This is sometimes called the "vagrancy hypothesis", or the "escape hypothesis".
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larger cells. Over time, genes not required by their parasitism were lost. The bacteria
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Sit-and-wait-transmission- the virus is living outside a host for long periods of time
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Rinderpest and peste des petits ruminants: virus plagues of large and small ruminants
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Subfield of evolutionary biology and virology concerned with the evolution of viruses
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Proceedings of the National Academy of Sciences of the United States of America
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as a group of closely related viral strains competing within an environment.
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Tsagris EM, MartĂ­nez de Alba AE, Gozmanova M, Kalantidis K (November 2008).
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Miller ES, Kutter E, Mosig G, Arisaka F, Kunisawa T, RĂĽger W (March 2003).
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Fraile A, GarcĂ­a-Arenal F (2010). "The Coevolution of Plants and Viruses".
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Airborne transmission- passed on through the air (brought in by breathing)
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Boutwell CL, Rolland MM, Herbeck JT, Mullins JI, Allen TM (October 2010).
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Droplet transmission- passed on through body fluids (sneezing on someone)
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and could be the origin of some viruses. They were discovered in maize by
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species contain hundreds to thousands of sequences derived from ancient
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in the traditional sense, because they are much smaller than the finest
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coat so that it can be transmitted to new cells. In similar manner, the
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Vector transmission- picked up by a carrier and brought to a new host
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The coevolution of plants and viruses: resistance and pathogenicity
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Collier L, Balows A, Sussman M (1998). Mahy B, Collier LA (eds.).
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Sussman M, Topley W, Wilson GK, Collier L, Balows A (1998).
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less effective. Both of these changes occur frequently with
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Emerman M, Malik HS (February 2010). Virgin SW (ed.).
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An example would be how viral meningitis is passed on
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December 2007 1727: 1726: 1722: 1677: 1670: 1625: 1621: 1613: 1609: 1601: 1597: 1589: 1585: 1540: 1536: 1521: 1497: 1493: 1446: 1442: 1434: 1430: 1385: 1381: 1336: 1329: 1321: 1317: 1309: 1305: 1266: 1262: 1229:(2): e1000301. 1215: 1208: 1157: 1153: 1114:(8): e1500527. 1100: 1096: 1049: 1045: 1014:(11): 2168–79. 1000: 996: 941: 937: 930: 916: 905: 898: 884: 875: 868: 854: 843: 836: 822: 809: 769: 763: 759: 720: 711: 700: 696: 688: 684: 676: 672: 657: 635: 631: 623: 619: 613: 608: 559: 490: 469:DNA replication 444:model organisms 434:. The complete 405: 385:morbilliviruses 377:antigenic drift 365:morbilliviruses 350:antiviral drugs 330:molecular clock 276: 267:astrobiological 229: 192:subviral agents 121: 116: 96:antiviral drugs 72:influenza virus 20:Viral evolution 17: 12: 11: 5: 3452: 3442: 3441: 3436: 3431: 3416: 3415: 3403: 3391: 3368: 3367: 3365: 3364: 3354: 3343: 3340: 3339: 3337: 3336: 3331: 3326: 3321: 3316: 3315: 3314: 3304: 3299: 3294: 3289: 3284: 3283: 3282: 3277: 3272: 3262: 3257: 3252: 3246: 3244: 3240: 3239: 3237: 3236: 3231: 3230: 3229: 3224: 3219: 3218: 3217: 3207: 3202: 3197: 3192: 3187: 3177: 3171: 3169: 3165: 3164: 3162: 3161: 3156: 3151: 3146: 3141: 3136: 3131: 3126: 3121: 3116: 3115: 3114: 3105:Charles Darwin 3102: 3101: 3100: 3088: 3083: 3077: 3075: 3069: 3068: 3066: 3065: 3060: 3055: 3050: 3045: 3043:Non-ecological 3040: 3035: 3030: 3025: 3020: 3015: 3010: 3004: 3002: 2996: 2995: 2993: 2992: 2983: 2974: 2960: 2958: 2952: 2951: 2949: 2948: 2943: 2942: 2941: 2936: 2931: 2926: 2921: 2911: 2906: 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2249: 2244: 2239: 2234: 2229: 2224: 2219: 2214: 2212:Common descent 2209: 2208: 2207: 2197: 2192: 2190:Baldwin effect 2187: 2186: 2185: 2180: 2170: 2165: 2160: 2154: 2152: 2146: 2145: 2143: 2142: 2137: 2132: 2127: 2122: 2116: 2113: 2112: 2105: 2104: 2097: 2090: 2082: 2076: 2075: 2055: 2054:External links 2052: 2050: 2049: 2007: 2005: 2002: 2000: 1999: 1993: 1980: 1974: 1961: 1947: 1934: 1928: 1912: 1910: 1907: 1905: 1904: 1889: 1871: 1860:(3): 199–214. 1840: 1816: 1800:"Encephalitis" 1791: 1770: 1745: 1720: 1668: 1619: 1607: 1595: 1583: 1554:(3): 603–617. 1534: 1519: 1491: 1440: 1428: 1405:10.1086/655653 1379: 1350:(8): 859–864. 1327: 1315: 1303: 1260: 1206: 1151: 1094: 1043: 994: 935: 928: 903: 896: 873: 866: 841: 834: 807: 757: 730:(7): 449–458. 709: 694: 682: 670: 655: 629: 616: 612: 609: 607: 606: 601: 596: 591: 586: 581: 576: 571: 566: 560: 558: 555: 540: 539: 538: 537: 531: 530: 529: 523: 522: 521: 515: 514: 513: 507: 506: 505: 489: 486: 424:in the family 404: 401: 275: 272: 255:viral proteome 242: 241: 228: 225: 224: 223: 180: 177: 158: 155: 139: 120: 117: 115: 112: 48:point mutation 44:mutation rates 15: 9: 6: 4: 3: 2: 3451: 3440: 3437: 3435: 3432: 3430: 3427: 3426: 3424: 3414: 3409: 3404: 3402: 3397: 3392: 3390: 3385: 3380: 3379: 3376: 3363: 3359: 3355: 3353: 3345: 3344: 3341: 3335: 3332: 3330: 3327: 3325: 3322: 3320: 3317: 3313: 3310: 3309: 3308: 3307:Phylogenetics 3305: 3303: 3300: 3298: 3295: 3293: 3290: 3288: 3285: 3281: 3278: 3276: 3273: 3271: 3268: 3267: 3266: 3263: 3261: 3258: 3256: 3253: 3251: 3248: 3247: 3245: 3241: 3235: 3232: 3228: 3225: 3223: 3220: 3216: 3213: 3212: 3211: 3210:Structuralism 3208: 3206: 3203: 3201: 3198: 3196: 3193: 3191: 3188: 3186: 3185:Catastrophism 3183: 3182: 3181: 3178: 3176: 3173: 3172: 3170: 3166: 3160: 3157: 3155: 3152: 3150: 3147: 3145: 3144:Neo-Darwinism 3142: 3140: 3137: 3135: 3132: 3130: 3127: 3125: 3122: 3120: 3117: 3113: 3112: 3108: 3107: 3106: 3103: 3099: 3098: 3094: 3093: 3092: 3089: 3087: 3084: 3082: 3079: 3078: 3076: 3074: 3070: 3064: 3061: 3059: 3058:Reinforcement 3056: 3054: 3051: 3049: 3046: 3044: 3041: 3039: 3036: 3034: 3031: 3029: 3026: 3024: 3021: 3019: 3016: 3014: 3011: 3009: 3006: 3005: 3003: 3001: 2997: 2991: 2990:Catastrophism 2987: 2984: 2982: 2981:Macromutation 2978: 2977:Micromutation 2975: 2973: 2969: 2965: 2962: 2961: 2959: 2957: 2953: 2947: 2944: 2940: 2937: 2935: 2932: 2930: 2927: 2925: 2922: 2920: 2917: 2916: 2915: 2912: 2910: 2907: 2905: 2902: 2900: 2897: 2895: 2892: 2890: 2887: 2885: 2884:Immune system 2882: 2880: 2877: 2875: 2872: 2870: 2867: 2865: 2862: 2858: 2855: 2854: 2853: 2850: 2848: 2845: 2843: 2840: 2838: 2835: 2831: 2828: 2826: 2823: 2822: 2821: 2818: 2817: 2815: 2813: 2808: 2800: 2797: 2795: 2792: 2790: 2787: 2785: 2782: 2780: 2777: 2776: 2774: 2770: 2767: 2765: 2762: 2760: 2757: 2755: 2752: 2750: 2747: 2745: 2744:symbiogenesis 2742: 2741: 2740: 2737: 2735: 2732: 2730: 2727: 2725: 2722: 2721: 2719: 2717: 2712: 2706: 2703: 2701: 2698: 2696: 2693: 2691: 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Blackwell. 1940: 1935: 1931: 1929:0-12-088385-6 1925: 1921: 1920: 1914: 1913: 1900: 1896: 1892: 1886: 1882: 1875: 1867: 1863: 1859: 1855: 1851: 1844: 1829: 1823: 1821: 1805: 1804:PubMed Health 1801: 1795: 1780: 1774: 1759: 1755: 1749: 1734: 1730: 1724: 1716: 1712: 1707: 1702: 1698: 1694: 1690: 1686: 1682: 1675: 1673: 1664: 1660: 1655: 1650: 1646: 1642: 1638: 1634: 1630: 1623: 1616: 1611: 1604: 1599: 1592: 1587: 1579: 1575: 1570: 1565: 1561: 1557: 1553: 1549: 1545: 1538: 1530: 1526: 1522: 1516: 1512: 1508: 1504: 1503: 1495: 1487: 1483: 1478: 1473: 1468: 1463: 1459: 1455: 1451: 1444: 1437: 1432: 1424: 1420: 1415: 1410: 1406: 1402: 1398: 1394: 1390: 1383: 1375: 1371: 1367: 1363: 1358: 1353: 1349: 1345: 1344:FASEB Journal 1341: 1334: 1332: 1325:, p. 272 1324: 1319: 1313:, p. 273 1312: 1307: 1299: 1295: 1291: 1287: 1283: 1279: 1275: 1271: 1264: 1256: 1252: 1247: 1242: 1237: 1232: 1228: 1224: 1220: 1213: 1211: 1202: 1198: 1194: 1190: 1186: 1182: 1178: 1174: 1170: 1166: 1162: 1155: 1147: 1143: 1138: 1133: 1129: 1125: 1121: 1117: 1113: 1109: 1105: 1098: 1090: 1086: 1082: 1078: 1074: 1070: 1066: 1062: 1058: 1054: 1047: 1039: 1035: 1031: 1027: 1022: 1017: 1013: 1009: 1005: 998: 990: 986: 981: 976: 971: 966: 962: 958: 955:(6): 344–55. 954: 950: 946: 939: 931: 925: 921: 914: 912: 910: 908: 899: 893: 889: 882: 880: 878: 869: 867:0-340-66316-2 863: 859: 852: 850: 848: 846: 837: 831: 827: 820: 818: 816: 814: 812: 803: 799: 795: 791: 787: 783: 780:(7): 449–58. 779: 775: 768: 761: 753: 749: 745: 741: 737: 733: 729: 725: 718: 716: 714: 707: 703: 698: 691: 686: 679: 674: 666: 662: 658: 652: 648: 644: 641:. ASM Press. 640: 633: 626: 621: 617: 615: 605: 602: 600: 597: 595: 592: 590: 587: 585: 582: 580: 577: 575: 572: 570: 567: 565: 562: 561: 554: 552: 548: 544: 535: 534: 532: 527: 526: 524: 519: 518: 516: 511: 510: 508: 503: 502: 500: 499: 498: 496: 485: 482: 478: 477:recombination 474: 470: 465: 461: 457: 453: 449: 445: 441: 440:gene products 437: 433: 429: 428: 423: 419: 418: 414:that infects 413: 412:bacteriophage 409: 403:Bacteriophage 400: 398: 394: 390: 386: 380: 378: 374: 373:genetic shift 366: 361: 357: 355: 351: 347: 346:immune system 342: 338: 333: 331: 326: 324: 323:paleovirology 320: 316: 312: 308: 305: 301: 297: 293: 285: 282:Time-line of 280: 271: 268: 262: 260: 256: 252: 248: 238: 234: 231: 230: 221: 217: 215: 210: 206: 201: 197: 193: 189: 185: 181: 178: 175: 171: 167: 163: 159: 156: 152: 148: 144: 140: 137: 136: 135: 132: 130: 126: 111: 109: 105: 101: 97: 93: 89: 85: 81: 77: 73: 69: 65: 60: 57: 53: 49: 45: 41: 37: 33: 29: 25: 21: 3319:Polymorphism 3302:Astrobiology 3250:Biogeography 3205:Saltationism 3195:Orthogenesis 3180:Alternatives 3109: 3095: 3028:Cospeciation 3023:Cladogenesis 2972:Saltationism 2929:Mating types 2852:Color vision 2837:Avian flight 2759:mitochondria 2704: 2497:Canalisation 2375:Biodiversity 2120:Introduction 2064: 2042:. Retrieved 2023:(1): 101–6. 2020: 2016: 1984: 1965: 1938: 1922:. Elsevier. 1918: 1909:Bibliography 1880: 1874: 1857: 1853: 1843: 1832:. Retrieved 1808:. Retrieved 1803: 1794: 1783:. Retrieved 1773: 1762:. Retrieved 1757: 1748: 1737:. Retrieved 1732: 1723: 1688: 1684: 1636: 1632: 1622: 1610: 1605:, p. 16 1598: 1586: 1551: 1547: 1537: 1501: 1494: 1457: 1453: 1443: 1438:, p. 24 1431: 1396: 1392: 1382: 1347: 1343: 1318: 1306: 1273: 1269: 1263: 1226: 1223:PLOS Biology 1222: 1168: 1165:Astrobiology 1164: 1154: 1111: 1107: 1097: 1056: 1052: 1046: 1011: 1007: 997: 952: 948: 938: 919: 887: 857: 825: 777: 773: 760: 727: 723: 697: 692:, p. 26 685: 680:, p. 25 673: 638: 632: 627:, p. 24 620: 614: 541: 495:transmission 491: 488:Transmission 425: 415: 406: 381: 370: 334: 327: 315:retroviruses 289: 284:paleoviruses 263: 254: 243: 236: 232: 214:Acanthamoeba 212: 184:nucleic acid 154:hypothesis". 133: 122: 64:epidemiology 61: 50:or more per 19: 18: 3329:Systematics 3200:Mutationism 3018:Catagenesis 2946:Snake venom 2879:Eusociality 2857:in primates 2847:Cooperation 2775:In animals 2595:butterflies 2568:Cephalopods 2558:Brachiopods 2490:Development 2464:Mate choice 2217:Convergence 2200:Coevolution 2158:Abiogenesis 2038:11089/43149 1276:(1): 5–49. 341:RNA viruses 216:castellanii 166:transposons 143:parasitised 40:RNA viruses 3423:Categories 3190:Lamarckism 3168:Philosophy 3091:David Hume 3053:Peripatric 3048:Parapatric 3033:Ecological 3013:Anagenesis 3008:Allopatric 3000:Speciation 2964:Gradualism 2889:Metabolism 2749:chromosome 2739:Eukaryotes 2517:Modularity 2434:Population 2360:Population 2321:Speciation 2299:Panspermia 2252:Extinction 2247:Exaptation 2222:Divergence 2195:Cladistics 2183:Reciprocal 2163:Adaptation 2044:2023-05-11 1987:. Arnold. 1854:Apidologie 1834:2017-12-05 1828:"Smallpox" 1810:2017-12-05 1785:2017-12-05 1764:2017-12-05 1739:2017-11-27 704:, p.  611:References 481:nucleotide 473:DNA repair 464:eukaryotes 427:Myoviridae 397:rinderpest 311:vertebrate 220:satellites 147:rickettsia 3324:Protocell 3175:Darwinism 3063:Sympatric 2812:processes 2700:Tetrapods 2649:Kangaroos 2575:Dinosaurs 2512:Inversion 2481:Variation 2402:Gene flow 2395:Inclusive 2205:Mutualism 2150:Evolution 1899:794228026 1201:105299840 1193:1531-1074 1038:221581424 1004:"Viroids" 802:256744818 752:169035711 665:755638822 579:RNA virus 564:DNA virus 543:Virulence 422:DNA virus 337:Darwinian 319:sequences 317:. These 296:colloidal 274:Evolution 247:phylogeny 209:mimivirus 151:chlamydia 84:hepatitis 68:influenza 32:evolution 3434:Virology 3352:Category 3227:Vitalism 3222:Theistic 3215:Spandrel 2899:Morality 2894:Monogamy 2769:plastids 2734:Flagella 2690:Reptiles 2671:sea cows 2654:primates 2563:Molluscs 2541:Bacteria 2429:Mutation 2362:genetics 2338:Taxonomy 2284:Mismatch 2264:Homology 2178:Cheating 2173:Altruism 2071:Archived 2066:PBS Eons 1957:65207057 1663:12626685 1578:20674794 1529:20965070 1486:19284639 1423:20846038 1374:20865732 1298:35371362 1290:20367503 1255:20161719 1146:26601271 1081:18690211 1030:18764915 989:15430309 794:31142823 744:31142823 557:See also 460:bacteria 448:virology 307:germline 251:proteome 162:plasmids 92:vaccines 28:virology 3413:Viruses 3401:Science 3375:Portals 3243:Related 3073:History 2934:Meiosis 2869:Empathy 2864:Emotion 2764:nucleus 2705:Viruses 2695:Spiders 2607:Mammals 2590:Insects 2390:Fitness 2326:Species 2125:Outline 1715:2651395 1569:7127320 1477:2666653 1414:2945609 1366:8666162 1246:2817711 1173:Bibcode 1137:4643759 1116:Bibcode 1089:4422249 1061:Bibcode 980:1063197 957:Bibcode 389:measles 292:fossils 188:Viroids 114:Origins 82:), and 36:viruses 3362:Portal 3038:Hybrid 2874:Ethics 2716:organs 2678:Plants 2664:lemurs 2659:humans 2644:horses 2634:hyenas 2622:wolves 2617:canids 2551:origin 1991:  1972:  1955:  1945:  1926:  1897:  1887:  1713:  1706:209897 1703:  1661:  1654:150520 1651:  1576:  1566:  1527:  1517:  1484:  1474:  1460:: 30. 1421:  1411:  1372:  1364:  1296:  1288:  1253:  1243:  1199:  1191:  1144:  1134:  1087:  1079:  1053:Nature 1036:  1028:  987:  977:  926:  894:  864:  832:  800:  792:  750:  742:  663:  653:  436:genome 249:. 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Index

evolutionary biology
virology
evolution
viruses
RNA viruses
mutation rates
point mutation
genome
natural selection
epidemiology
influenza
influenza virus
AIDS
HIV
hepatitis
HCV
vaccines
antiviral drugs
resistant mutations
quasispecies model
viral quasispecies
three domains of life
last universal common ancestor
parasitised
rickettsia
chlamydia
plasmids
transposons
mobile genetic elements
Barbara McClintock

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