833:, published a pioneering work in which he laid out his protocell theory on the origin of life. In his initial protocell model, Oparin took inspiration from Graham's description of colloids from 1861 as substances that usually give cloudy solutions and cannot pass through membranes. Oparin linked these properties to the protoplasm, and reasoned that precipitates of colloids form as clots or lumps of mucus or jelly, some of which have structural features that resemble the protoplasm. According to Oparin, protocells could therefore have formed by precipitation of colloids. In his later work, Oparin became more specific about his protocell model. He described the work of Bungenberg-de Jong on coacervates in his book from 1938, and postulated that the first protocells were coacervates.
818:
p. 82), most likely a description of coalescing coacervate droplets. Several years later, in 1929, Bungenberg-de Jong published a seminal paper with his PhD advisor, H. R. Kruyt, entitled âCoacervation. Partial miscibility in colloid systemsâ. In their paper, they give many more examples of colloid systems that flocculate into an emulsoid state, either by varying the temperature, by adding salts, co-solvents or by mixing together two oppositely charged polymer colloids, and illustrate their observations with the first microscope pictures of coacervate droplets. They term this phenomenon coacervation, derived from the prefix
669:
2599:
20:
577:
826:(heap), which relates to the dense liquid droplets. Coacervation is thus loosely translated as âto come together in a heapâ. Since then, Bungenberg-de Jong and his research group in Leiden published a range of papers on coacervates, including results on self-coacervation, salt effects, interfacial tension, multiphase coacervates and surfactant-based coacervates.
126:, leading to a dense phase in thermodynamic equilibrium with a dilute phase. The dispersed droplets of dense phase are also called coacervates, micro-coacervates or coacervate droplets. These structures draw a lot of interest because they form spontaneously from aqueous mixtures and provide stable compartmentalization without the need of a membraneâthey are
743:
show liquid-like properties, and was later confirmed by showing that liquid condensates can be reconstituted from purified protein and RNA in vitro. However, whether MLOs should be referred to as liquids, remains disputable. Even if initially they are liquid-like, over time some of them maturate into
869:
to refer to the dilute mixture of organic molecules that could have built up as a result of reactions between inorganic building blocks such as ammonia, carbon dioxide and water, in presence of UV light as an energy source. Oparin proposed that simple building blocks with increasing complexity could
845:
In the decades after that, until about 2000, the scientific interest in coacervates had faded. Oparin's theory on the role of coacervates in the origin of life had been replaced by interest in the RNA world hypothesis. Renewed interest in coacervates originated as scientists recognized the relevance
809:
could also be induced in certain polymer solutions by mixing them together. In particular, he reported the observation of opalescence (a turbid mixture) when equal volumes of acidified 0.5% âwashedâ gelatine solution, and 2% gum arabic solution were mixed. Tiebackx did not further analyse the nature
893:
scenarios have gained more attention since the 1980s with the work of
Morowitz, Luisi and Szostak. However, recently, there has been a rising interest in coacervates as protocells, resonating with current findings that reactions too slow or unlikely in aqueous solutions can be significantly favored
210:
H can be different from zero, and the process endothermic enough to overcome the entropic term and favor the de-mixed state (the blue curve shifts up). Low molecular-weight solutes will hardly reach such non-ideality, whereas for polymeric solutes, with increasing interactions sites N and therefore
836:
Other researchers followed, and in the 1930s and 1940s various examples of coacervation were reported, by
Bungenberg-de Jong, Oparin, Koets, Bank, Langmuir and others. In the 1950s and 1960s, focus shifted to a theoretical description of the phenomenon of (complex) coacervation. Voorn and Overbeek
804:
At the start of the 20th century, scientists had become interested in the stability of colloids, both the dispersions of solid particles and the solutions of polymeric molecules. It was known that salts and temperature could often be used to cause flocculation of a colloid. The German chemist F.W.
2600:
PeretĂł, Juli G., translator, writer of introduction. Inness, NatĂ lia, translator. Translation of: Oparin, A. I. (Aleksandr
Ivanovich), 1894-1980. Proiskhozhedenie zhizni. Translation of: Haldane, J. B. S. (John Burdon Sanderson), 1892-1964. Origin of life. Container of (expression): Oparin, A. I.
664:
of opposite charge. The Voorn-Overbeek approach applies the Debye-HĂŒckel approximation to the enthalpic term in the Flory-Huggins model, and considers two polyelectrolytes of the same length and at the same concentration. Complex coacervates are a subset of aqueous two-phase systems (ATPS),
182:
Associative LLPS is dominated by attractive interactions between macromolecules (such as electrostatic force between oppositely charged polymers), and segregative LLPS is driven by the minimization of repulsive interactions (such as hydrophobic effect on proteins containing a disordered region).
157:
A reawakening of coacervate research was seen in the 2000s, starting with the recognition in 2004 by scientists at the
University of California, Santa Barbara (UCSB) that some marine invertebrates (such as the sandcastle worm) exploit complex coacervation to produce water-resistant biological
1761:
Boulay, Gaylor; Sandoval, Gabriel J.; Riggi, Nicolo; Iyer, Sowmya; Buisson, RĂ©mi; Naigles, Beverly; Awad, Mary E.; Rengarajan, Shruthi; Volorio, Angela; McBride, Matthew J.; Broye, Liliane C. (2018-10-01). "Abstract PR09: Cancer-specific retargeting of BAF complexes by a prion-like domain".
817:
reported in his PhD thesis (Utrecht, 1921) two types of flocculation in agar solutions: one that leads to a suspensoid state, and one that leads to an emulsoid state. He observed the emulsoid state under the microscope and described small particles that merged into larger particles (Thesis,
178:; that is, the dense phase retains some of the original solvent â generally water â and does not collapse into solid aggregates, rather keeping a liquid property. Coacervates can be characterized as complex or simple based on the driving force for the LLPS: associative or segregative
324:
520:
The phase diagram of the mixture can be predicted by experimentally determining the two-phase boundary, or binodal curve. In a simplistic theoretical approach, the binodes are the compositions at which the free energy of de-mixing is minimal
655:
curve is defined. The conditions of the mixture in comparison to the two curves defines the phase separation mechanism: nucleation-growth of coacervate droplets (when the binodal region is crossed slowly) and spinodal decomposition.
649:
572:
133:
The term coacervate was coined in 1929 by Dutch chemist
Hendrik G. Bungenberg de Jong and Hugo R. Kruyt while studying lyophilic colloidal dispersions. The name is a reference to the clustering of colloidal particles, like
2257:
Wang, Ailin; Conicella, Alexander E; Schmidt, Hermann Broder; Martin, Erik W; Rhoads, Shannon N; Reeb, Ashley N; Nourse, Amanda; Ramirez
Montero, Daniel; Ryan, Veronica H; Rohatgi, Rajat; Shewmaker, Frank (2018-02-09).
841:
approximation. Veis and Aranyi suggested to extend this model with an electrostatic aggregation step in which charge-paired symmetrical soluble aggregates are formed, followed by phase separation into liquid droplets.
461:
1551:
Boeynaems, Steven; Alberti, Simon; Fawzi, Nicolas L.; Mittag, Tanja; Polymenidou, Magdalini; Rousseau, Frederic; Schymkowitz, Joost; Shorter, James; Wolozin, Benjamin; Van Den Bosch, Ludo; Tompa, Peter (June 2018).
659:
Associative LLPS is more complex to describe, as both solute polymers are present in the dilute and dense phase. Electrostatic-based complex coacervates are the most common, and in that case the solutes are two
2142:
Patel, Avinash; Lee, Hyun O.; Jawerth, Louise; Maharana, Shovamayee; Jahnel, Marcus; Hein, Marco Y.; Stoynov, Stoyno; Mahamid, Julia; Saha, Shambaditya; Franzmann, Titus M.; Pozniakovski, Andrej (August 2015).
516:
580:
Free energy of de-mixing according to Flory-Huggins approach. By determining the free-energy curve for different temperatures and taking the critical points, the phase diagram on the right can be constructed.
771:
structure), but in many cases transformation from liquid to solid results in the formation of pathological aggregates. Examples of both liquid-liquid phase separating and aggregation-prone proteins include
3913:
2601:(Aleksandr Ivanovich), 1894-1980. Proiskhozhedenie zhizni. Catalan. Container of (expression): Haldane, J. B. S. (John Burdon Sanderson), 1892-1964. Origin of life. Catalan. (28 November 2011).
837:
developed the first mean-field theory to describe coacervation. They estimated the total free energy of mixing as a sum of mixing entropy terms and mean-field electrostatic interactions in a
158:
adhesives. A few years later in 2009 the role of liquid-liquid phase separation was further recognized to be involved in the formation of certain membraneless organelles by the biophysicists
1846:
Li, Pilong; Banjade, Sudeep; Cheng, Hui-Chun; Kim, Soyeon; Chen, Baoyu; Guo, Liang; Llaguno, Marc; Hollingsworth, Javoris V.; King, David S.; Banani, Salman F.; Russo, Paul S. (March 2012).
870:
organize locally, or self-assemble, to form protocells with living properties. He performed experiments based on
Bungenberg de Jong's colloidal aggregates (coacervates) to encapsulate
2033:
Boke, Elvan; Ruer, Martine; WĂŒhr, Martin; Coughlin, Margaret; Lemaitre, Regis; Gygi, Steven P.; Alberti, Simon; Drechsel, David; Hyman, Anthony A.; Mitchison, Timothy J. (July 2016).
216:
3906:
3083:
3899:
814:
1658:
Riback, Joshua A.; Katanski, Christopher D.; Kear-Scott, Jamie L.; Pilipenko, Evgeny V.; Rojek, Alexandra E.; Sosnick, Tobin R.; Drummond, D. Allan (March 2017).
584:), across different temperatures (or other interaction parameter). Alternatively, by minimizing the change in free energy of de-mixing in regards to composition (
2800:
Drobot, Björn; Iglesias-Artola, Juan M.; Le Vay, Kristian; Mayr, Viktoria; Kar, Mrityunjoy; Kreysing, Moritz; Mutschler, Hannes; Tang, T-Y Dora (2018-09-07).
587:
2315:
Molliex, Amandine; Temirov, Jamshid; Lee, Jihun; Coughlin, Maura; Kanagaraj, Anderson P.; Kim, Hong Joo; Mittag, Tanja; Taylor, J. Paul (September 2015).
865:
independently hypothesized in the 1920s that the first cells in early Earth's oceans could be, in essence, coacervate droplets. Haldane used the term
846:
and versatility of the interactions that underlie complex coacervation in the natural fabrication of biological materials and in their self-assembly.
524:
3076:
763:) may lead to the formation of higher ordered structures and solidification of MLOs. Some MLOs serve their biological role as solid particles (e.g.
2688:
2647:
2692:
2651:
330:
724:). They have been shown to serve various functions: they can store and protect cellular material during stress conditions, they participate in
3069:
849:
Since 2009, coacervates have become linked to membraneless organelles and there has been a renewed interest in coacervates as protocells.
759:
in solution. IDRs can provide interactions responsible for LLPS, but over time conformational changes (sometimes promoted by mutations or
4469:
2900:
467:
1968:
Brangwynne, C. P.; Eckmann, C. R.; Courson, D. S.; Rybarska, A.; Hoege, C.; Gharakhani, J.; Julicher, F.; Hyman, A. A. (2009-05-21).
1083:
Brangwynne, C. P.; Eckmann, C. R.; Courson, D. S.; Rybarska, A.; Hoege, C.; Gharakhani, J.; Julicher, F.; Hyman, A. A. (2009-06-26).
2565:
Veis, Arthur; Aranyi, Catherine (September 1960). "Phase
Separation in Polyelectrolyte Systems. I. Complex Coacervates of Gelatin".
2883:
882:
further strengthened the theory that inorganic building blocks could increase in complexity and give rise to cell-like structures.
2429:"Remonstrantie der predikanten van Utrecht, overgelevert aen de Staten s'landts van Utrecht, raeckende het poinct van religie".
4036:
2765:
Dzieciol, Alicja J.; Mann, Stephen (2012-03-01). "ChemInform
Abstract: Designs for Life: Protocell Models in the Laboratory".
1482:
Overbeek, J. T. G.; Voorn, M. J. (May 1957). "Phase separation in polyelectrolyte solutions. Theory of complex coacervation".
2610:
945:
1424:"Simple Calculation of Phase Diagrams for LiquidâLiquid Phase Separation in Solutions of Two Macromolecular Solute Species"
4402:
2947:
1799:"Faculty Opinions recommendation of Phase separation of signaling molecules promotes T cell receptor signal transduction"
1209:"RNA-Based Coacervates as a Model for Membraneless Organelles: Formation, Properties, and Interfacial Liposome Assembly"
2202:"ALS Mutations Disrupt Phase Separation Mediated by α-Helical Structure in the TDP-43 Low-Complexity C-Terminal Domain"
4459:
2952:
748:
199:
187:
2317:"Phase Separation by Low Complexity Domains Promotes Stress Granule Assembly and Drives Pathological Fibrillization"
4464:
2980:
752:
764:
760:
740:
319:{\displaystyle \Delta _{mix}G={\frac {\phi }{N}}\ln \phi +(1-\phi )\ln(1-\phi )+\Delta _{\operatorname {mix} }H}
166:. Liquid organelles share features with coacervate droplets and fueled the study of coacervates for biomimicry.
4387:
3737:
3332:
3028:
1913:"Cajal Bodies, Nucleoli, and Speckles in the Xenopus Oocyte Nucleus Have a Low-Density, Sponge-like Structure"
4375:
3873:
2970:
789:
665:
which also include segregatively separated systems in which both phases are enriched in one type of polymer.
4529:
3521:
3202:
2876:
3207:
838:
744:
solids (gel-like or even crystalline, depending on the extent of spatial ordering within the condensate).
3441:
3197:
2975:
2505:
4534:
2942:
1611:"Considerations and Challenges in Studying Liquid-Liquid Phase Separation and Biomolecular Condensates"
1365:"Considerations and Challenges in Studying Liquid-Liquid Phase Separation and Biomolecular Condensates"
866:
705:. MLOs are mostly composed of proteins and nucleic acids, held together by weak intermolecular forces.
1258:"A review of the early development of the thermodynamics of the complex coacervation phase separation"
3891:
3742:
3023:
2664:
2260:"A single Nâterminal phosphomimic disrupts TDPâ43 polymerization, phase separation, and RNA splicing"
151:
2455:
1829:
1747:
735:
It is now widely believed that MLOs form through LLPS. This was first proposed after observing that
4446:
4392:
3589:
3490:
3407:
4539:
4481:
3616:
3392:
2869:
2434:
793:
682:
1207:
Aumiller, William M.; Pir Cakmak, Fatma; Davis, Bradley W.; Keating, Christine D. (2016-10-04).
3844:
3160:
2914:
2640:
1970:"Germline P Granules Are Liquid Droplets That Localize by Controlled Dissolution/Condensation"
1085:"Germline P Granules Are Liquid Droplets That Localize by Controlled Dissolution/Condensation"
3604:
3577:
3192:
2442:
2200:
Conicella, Alexander E.; Zerze, GĂŒl H.; Mittal, Jeetain; Fawzi, Nicolas L. (September 2016).
1816:
1734:
721:
3180:
1715:
Wheeler, Joshua R; Matheny, Tyler; Jain, Saumya; Abrisch, Robert; Parker, Roy (2016-08-15).
4491:
3548:
3543:
3419:
2813:
2715:
2145:"A Liquid-to-Solid Phase Transition of the ALS Protein FUS Accelerated by Disease Mutation"
1981:
1859:
1318:
1096:
686:
885:
The Oparin-Haldane hypothesis established the foundations of research on the chemistry of
8:
4066:
3784:
3626:
3582:
3538:
3276:
2985:
729:
159:
2817:
2719:
2111:
2094:
1985:
1863:
1322:
1100:
961:
644:{\displaystyle {\frac {\partial ^{2}\Delta _{\operatorname {mix} }G}{\partial x^{2}}}=0}
4360:
4282:
3834:
3002:
2842:
2801:
2747:
2682:
2628:
2547:
2411:
2364:
2351:
2316:
2292:
2259:
2234:
2201:
2182:
2067:
2034:
2015:
1888:
1847:
1779:
1692:
1659:
1635:
1610:
1586:
1553:
1456:
1423:
1404:
1391:
1364:
1342:
1282:
1257:
1184:
1149:
1130:
1065:
1016:
3061:
1945:
1912:
4338:
4254:
4156:
3926:
3829:
3533:
2997:
2937:
2847:
2829:
2782:
2739:
2731:
2670:
2616:
2606:
2582:
2539:
2486:
2403:
2356:
2338:
2297:
2279:
2239:
2221:
2174:
2166:
2124:
2116:
2072:
2054:
2007:
1999:
1950:
1932:
1893:
1875:
1697:
1679:
1640:
1591:
1573:
1530:
1507:
1499:
1461:
1443:
1396:
1334:
1287:
1238:
1230:
1189:
1171:
1122:
1114:
1057:
1008:
941:
785:
2522:
Just, Th.; Oparin, A. I.; Morgulis, Sergius (September 1938). "The Origin of Life".
2415:
2368:
2186:
2019:
1783:
1408:
1134:
3565:
3459:
3092:
2837:
2821:
2774:
2751:
2723:
2574:
2531:
2478:
2430:
2395:
2346:
2328:
2287:
2271:
2229:
2213:
2156:
2106:
2062:
2046:
1989:
1940:
1924:
1883:
1867:
1802:
1775:
1771:
1720:
1687:
1671:
1630:
1622:
1581:
1565:
1491:
1451:
1435:
1386:
1376:
1346:
1326:
1277:
1269:
1220:
1179:
1161:
1104:
1069:
1047:
1020:
998:
967:
933:
862:
858:
830:
202:
approach) is positive and the interaction enthalpies are all taken as equivalent (Î
163:
139:
71:
33:
2706:
FOX, SIDNEY W. (January 1965). "A Theory of
Macromolecular and Cellular Origins".
1807:
1798:
668:
4198:
3822:
3516:
3495:
3424:
3227:
2920:
1660:"Stress-Triggered Phase Separation Is an Adaptive, Evolutionarily Tuned Response"
1225:
1208:
908:
810:
of the flocs, but it is likely that this was an example of complex coacervation.
725:
661:
937:
567:{\displaystyle {\frac {\partial \Delta _{\operatorname {mix} }G}{\partial x}}=0}
4247:
4212:
4205:
4184:
4163:
4140:
4020:
3812:
3699:
3570:
3429:
3412:
3217:
3147:
2825:
2333:
2161:
2144:
2050:
1675:
1626:
1381:
875:
709:
2665:
Haldane, J. B. S. (John Burdon Sanderson), 1892-1964. Origin of life. (1929).
2217:
1569:
1273:
19:
4523:
4486:
4414:
4307:
4275:
4261:
4240:
4226:
4177:
4170:
4089:
3962:
3852:
3817:
3660:
3650:
3621:
3254:
3247:
2833:
2786:
2735:
2674:
2620:
2586:
2543:
2490:
2407:
2342:
2283:
2225:
2170:
2120:
2058:
2003:
1936:
1879:
1717:"Author response: Distinct stages in stress granule assembly and disassembly"
1683:
1577:
1534:
1503:
1447:
1439:
1338:
1234:
1175:
1166:
1118:
702:
690:
107:
2275:
1994:
1969:
1928:
1725:
1716:
1109:
1084:
4503:
4436:
4300:
4268:
4233:
4219:
4191:
4117:
4096:
3839:
3682:
3642:
3609:
3381:
3367:
3265:
3050:
3045:
2851:
2778:
2743:
2360:
2301:
2243:
2178:
2128:
2076:
2011:
1954:
1897:
1701:
1644:
1595:
1511:
1495:
1465:
1400:
1306:
1291:
1242:
1193:
1126:
1061:
1052:
1036:"Cement Proteins of the Tube-Building Polychaete Phragmatopoma californica"
1035:
1012:
971:
806:
694:
211:
decreasing entropic contribution, simple coacervation is much more likely.
119:
963:
The origin of life on the earth / Translated from the Russian by Ann Synge
874:
and enzymes within protocells. Work by chemists Sidney Fox, Kaoru Harada,
456:{\displaystyle \Delta _{mix}G=k_{B}T\left+\Delta _{\operatorname {mix} }H}
4365:
4110:
3863:
3729:
3677:
3672:
3185:
2892:
1911:
Handwerger, Korie E.; Cordero, Jason A.; Gall, Joseph G. (January 2005).
886:
879:
756:
576:
147:
2802:"Compartmentalised RNA catalysis in membrane-free coacervate protocells"
2578:
1871:
1305:
Brangwynne, Clifford P.; Tompa, Peter; Pappu, Rohit V. (November 2015).
190:(see equation). In ideal polymer solutions, the free-energy of mixing (Î
4496:
4443:
4030:
3943:
3771:
3719:
3712:
3707:
3692:
3687:
3528:
3463:
3212:
3154:
2908:
2551:
2482:
2399:
871:
773:
768:
736:
143:
3921:
1848:"Phase transitions in the assembly of multivalent signalling proteins"
1330:
1148:
Nakashima, Karina K.; Vibhute, Mahesh A.; Spruijt, Evan (2019-04-03).
1003:
986:
747:
Many proteins participating in the formation of MLO contain so-called
4103:
4082:
3984:
3776:
3761:
3748:
3500:
3288:
3119:
3007:
2992:
2962:
2727:
2469:
Jong, H. G. Bungenberg; Kruyt, H. R. (January 1930). "Koazervation".
903:
890:
717:
127:
2535:
2383:
4476:
4421:
4008:
3992:
3957:
3807:
3753:
3655:
3599:
3594:
3485:
3446:
3402:
3304:
3109:
928:
Booij, H. L.; Bungenberg de Jong, H. G. (1956), "Colloid Systems",
713:
698:
511:{\displaystyle \Delta _{\operatorname {mix} }H=\chi \phi (1-\phi )}
4508:
4426:
4407:
3967:
3952:
3789:
3667:
3451:
3436:
3139:
3114:
2861:
1770:(19_Supplement). American Association for Cancer Research: PR09.
1657:
781:
175:
115:
111:
1967:
1082:
852:
4454:
4397:
4382:
4332:
4062:
3880:
3858:
3555:
3478:
3473:
3238:
3129:
3124:
2799:
1609:
Alberti, Simon; Gladfelter, Amy; Mittag, Tanja (January 2019).
1363:
Alberti, Simon; Gladfelter, Amy; Mittag, Tanja (January 2019).
1150:"Biomolecular Chemistry in Liquid Phase Separated Compartments"
777:
751:(IDRs), parts of the polypeptide chain that can adopt multiple
1206:
89:
4431:
4058:
4002:
3997:
3362:
3354:
3172:
3134:
985:
Stewart, R.J.; Weaver, J.C.; Morse, D.E.; Waite, J.H (2004).
186:
The thermodynamics of segregative LLPS can be described by a
987:"The Tube Cement of Phragmatopoma californica: a solid foam"
51:
4370:
3974:
3922:
3868:
3560:
3101:
1550:
95:
77:
57:
42:
39:
2256:
2314:
2199:
1714:
83:
3091:
2141:
1760:
1608:
1554:"Protein Phase Separation: A New Phase in Cell Biology"
1524:
1362:
1147:
984:
927:
701:), MLOs are not separated from their surroundings by a
142:
to describe the proteinoid microspheres proposed to be
2035:"Amyloid-like Self-Assembly of a Cellular Compartment"
1034:
Zhao, H.; Sun, C.; Stewart, R.J.; Waite, J.H. (2005).
138:. The concept was later borrowed by Russian biologist
1910:
590:
527:
470:
333:
219:
86:
74:
48:
36:
2032:
1307:"Polymer physics of intracellular phase transitions"
1304:
150:. Coacervate-like protocells are at the core of the
98:
92:
60:
54:
1033:
784:. Aggregates of these proteins are associated with
681:Membraneless organelles (MLOs), also known as
80:
45:
2521:
643:
566:
510:
455:
318:
2388:Zeitschrift fĂŒr Chemie und Industrie der Kolloide
1845:
960:Oparin, Aleksandr Ivanovich; Synge, Ann. (1957).
689:. Unlike classic membrane-bound organelles (e.g.
4521:
2093:Alberti, Simon; Dormann, Dorothee (2019-12-03).
23:Coacervate droplets dispersed in a dilute phase
2558:
2092:
1484:Journal of Cellular and Comparative Physiology
3907:
3077:
2877:
1481:
853:Coacervates hypothesis for the origin of life
2764:
2687:: CS1 maint: multiple names: authors list (
2435:10.1163/2214-8264_dutchpamphlets-kb0-kb06696
194:G) is negative because the mixing entropy (Î
4470:Reverse transcriptase-related cellular gene
2384:"Gleichzeitige Ausflockung zweier Kolloide"
2095:"LiquidâLiquid Phase Separation in Disease"
959:
3914:
3900:
3084:
3070:
2884:
2870:
2691:) CS1 maint: numeric names: authors list (
2650:) CS1 maint: numeric names: authors list (
2564:
932:, Vienna: Springer Vienna, pp. 8â14,
2841:
2646:CS1 maint: multiple names: authors list (
2468:
2350:
2332:
2291:
2233:
2160:
2110:
2066:
1993:
1944:
1887:
1806:
1796:
1724:
1691:
1634:
1585:
1455:
1390:
1380:
1281:
1262:Advances in Colloid and Interface Science
1224:
1183:
1165:
1108:
1051:
1002:
676:
2381:
1764:Oral Presentations - Proffered Abstracts
728:and they are involved in the control of
708:MLOs are present in the cytoplasm (e.g.
667:
575:
18:
4451:Retroelements not elsewhere classified
4522:
2503:
1421:
206:H or Ï = 0). In non-ideal solutions, Î
3895:
3065:
2865:
2088:
2086:
1841:
1839:
1546:
1544:
1477:
1475:
1358:
1356:
124:liquid-liquid phase separation (LLPS)
1255:
174:Coacervates are a type of lyophilic
16:Aqueous phase rich in macromolecules
4403:Integrative and conjugative element
2948:Formamide-based prebiotic chemistry
2705:
2112:10.1146/annurev-genet-112618-043527
1428:The Journal of Physical Chemistry B
1040:The Journal of Biological Chemistry
991:The Journal of Experimental Biology
894:in such membraneless compartments.
13:
2891:
2083:
1836:
1541:
1472:
1353:
1154:Frontiers in Molecular Biosciences
930:Biocolloids and their Interactions
619:
605:
595:
549:
535:
531:
472:
441:
335:
304:
221:
188:Flory-Huggins polymer mixing model
14:
4551:
4460:Diversity-generating retroelement
2953:Alternative abiogenesis scenarios
2567:The Journal of Physical Chemistry
829:In the meantime, Russian chemist
169:
4465:Telomerase reverse transcriptase
4037:Microbes with highly unusual DNA
1525:Voorn, Michael Johannes (1956).
805:Tiebackx reported in 1911 that
761:post-translational modifications
749:intrinsically disordered regions
70:
32:
2793:
2758:
2699:
2658:
2593:
2515:
2497:
2462:
2422:
2375:
2308:
2250:
2193:
2135:
2026:
1961:
1904:
1797:Margulies, David (2016-05-17).
1790:
1754:
1708:
1651:
1602:
1518:
1422:Minton, Allen P. (2020-03-26).
1415:
1256:Veis, Arthur (September 2011).
672:Phase diagrams for coacervation
4388:Defective interfering particle
3738:Last universal common ancestor
3333:Defective interfering particle
3029:Last universal common ancestor
2382:Tiebackx, F. W. (April 1911).
1776:10.1158/1538-7445.pedca17-pr09
1298:
1249:
1200:
1141:
1076:
1027:
978:
953:
921:
505:
493:
429:
417:
408:
396:
297:
285:
276:
264:
106:) is an aqueous phase rich in
1:
4376:Clonally transmissible cancer
3874:Clonally transmissible cancer
3310:Satellite-like nucleic acids
1917:Molecular Biology of the Cell
1808:10.3410/f.726273110.793518440
914:
790:amyotrophic lateral sclerosis
2667:The rationalist annual, 1829
1226:10.1021/acs.langmuir.6b02499
966:. New York: Academic Press.
7:
3929:, and comparable structures
2605:. Universitat de ValĂšncia.
2524:American Midland Naturalist
938:10.1007/978-3-7091-5456-4_2
897:
716:) and in the nucleus (e.g.
10:
4556:
3430:Class II or DNA transposon
3425:Class I or retrotransposon
2826:10.1038/s41467-018-06072-w
2334:10.1016/j.cell.2015.09.015
2162:10.1016/j.cell.2015.07.047
2051:10.1016/j.cell.2016.06.051
1676:10.1016/j.cell.2017.02.027
1627:10.1016/j.cell.2018.12.035
1382:10.1016/j.cell.2018.12.035
889:, but the lipid-world and
799:
786:neurodegenerative diseases
4351:
4325:
4292:
4148:
4134:
4127:
4074:
4057:
4050:
4018:
3983:
3941:
3934:
3800:
3743:Earliest known life forms
3728:
3641:
3617:Repeated sequences in DNA
3509:
3391:
3380:
3353:
3325:
3275:
3264:
3237:
3226:
3171:
3100:
3038:
3024:Earliest known life forms
3016:
2961:
2930:
2899:
2218:10.1016/j.str.2016.07.007
2099:Annual Review of Genetics
1570:10.1016/j.tcb.2018.02.004
1274:10.1016/j.cis.2011.01.007
687:cell compartmentalization
152:Oparin-Haldane hypothesis
4393:Endogenous viral element
3590:Endogenous viral element
3408:Horizontal gene transfer
1440:10.1021/acs.jpcb.0c00402
1167:10.3389/fmolb.2019.00021
815:H. G. Bungenberg-de Jong
683:biomolecular condensates
198:S, combinatorial in the
3287:dsDNA satellite virus (
2276:10.15252/embj.201797452
1995:10.1126/science.1172046
1929:10.1091/mbc.e04-08-0742
1726:10.7554/elife.18413.018
1110:10.1126/science.1172046
794:frontotemporal dementia
3845:Helper dependent virus
3161:Biological dark matter
2943:MillerâUrey experiment
2915:Spontaneous generation
2779:10.1002/chin.201213265
2504:Oparin, A. I. (1924).
2450:Cite journal requires
1824:Cite journal requires
1742:Cite journal requires
1558:Trends in Cell Biology
1496:10.1002/jcp.1030490404
1053:10.1074/jbc.M508457200
972:10.5962/bhl.title.4528
861:and British biologist
677:Coacervates in biology
673:
645:
581:
568:
512:
457:
320:
24:
3605:Endogenous retrovirus
3578:Origin of replication
3294:ssDNA satellite virus
3284:ssRNA satellite virus
2806:Nature Communications
671:
646:
579:
569:
513:
458:
321:
22:
4492:Transposable element
4482:Spiegelman's Monster
3549:Secondary chromosome
3544:Extrachromosomal DNA
3420:Transposable element
2506:"The Origin of Life"
1670:(6): 1028â1040.e19.
1527:Complex coacervation
753:secondary structures
588:
525:
468:
331:
217:
4530:Colloidal chemistry
3785:Model lipid bilayer
3627:Interspersed repeat
2986:Primordial sandwich
2931:Prebiotic synthesis
2901:History of research
2818:2018NatCo...9.3643D
2720:1965Natur.205..328F
2603:L'origen de la vida
2579:10.1021/j100838a022
2471:Kolloid-Zeitschrift
1986:2009Sci...324.1729B
1980:(5935): 1729â1732.
1872:10.1038/nature10879
1864:2012Natur.483..336L
1323:2015NatPh..11..899B
1219:(39): 10042â10053.
1101:2009Sci...324.1729B
1095:(5935): 1729â1732.
857:Russian biochemist
822:and the Latin word
730:signal transduction
160:Clifford Brangwynne
140:Alexander I. Oparin
122:. It forms through
4361:Bio-like structure
4283:Tolecusatellitidae
3095:organic structures
3017:Earliest organisms
3003:Quasispecies model
2971:GADV-protein world
2639:has generic name (
2483:10.1007/bf01422833
2400:10.1007/bf01503532
674:
641:
582:
564:
508:
453:
316:
110:such as synthetic
25:
4535:Polymer chemistry
4517:
4516:
4347:
4346:
4321:
4320:
4317:
4316:
4255:Portogloboviridae
4157:Alphasatellitidae
4051:Non-cellular life
4046:
4045:
3927:non-cellular life
3889:
3888:
3830:Non-cellular life
3637:
3636:
3376:
3375:
3349:
3348:
3303:ssRNA satellite (
3059:
3058:
2998:Peptide-RNA world
2981:Ironâsulfur world
2938:Pseudo-panspermia
2714:(4969): 328â340.
2612:978-84-370-8607-1
1858:(7389): 336â340.
1434:(12): 2363â2370.
1331:10.1038/nphys3532
1004:10.1242/jeb.01330
947:978-3-211-80421-6
859:Aleksander Oparin
714:processing bodies
633:
556:
382:
250:
4547:
4132:
4131:
4072:
4071:
4055:
4054:
3939:
3938:
3916:
3909:
3902:
3893:
3892:
3566:Gene duplication
3389:
3388:
3385:self-replication
3273:
3272:
3235:
3234:
3093:Self-replicating
3086:
3079:
3072:
3063:
3062:
2917:(4th century BC)
2911:(5th century BC)
2886:
2879:
2872:
2863:
2862:
2856:
2855:
2845:
2797:
2791:
2790:
2762:
2756:
2755:
2728:10.1038/205328a0
2703:
2697:
2696:
2686:
2678:
2662:
2656:
2655:
2644:
2638:
2634:
2632:
2624:
2597:
2591:
2590:
2573:(9): 1203â1210.
2562:
2556:
2555:
2519:
2513:
2512:
2510:
2501:
2495:
2494:
2466:
2460:
2459:
2453:
2448:
2446:
2438:
2426:
2420:
2419:
2379:
2373:
2372:
2354:
2336:
2312:
2306:
2305:
2295:
2264:The EMBO Journal
2254:
2248:
2247:
2237:
2212:(9): 1537â1549.
2197:
2191:
2190:
2164:
2155:(5): 1066â1077.
2139:
2133:
2132:
2114:
2090:
2081:
2080:
2070:
2030:
2024:
2023:
1997:
1965:
1959:
1958:
1948:
1908:
1902:
1901:
1891:
1843:
1834:
1833:
1827:
1822:
1820:
1812:
1810:
1794:
1788:
1787:
1758:
1752:
1751:
1745:
1740:
1738:
1730:
1728:
1712:
1706:
1705:
1695:
1655:
1649:
1648:
1638:
1606:
1600:
1599:
1589:
1548:
1539:
1538:
1522:
1516:
1515:
1479:
1470:
1469:
1459:
1419:
1413:
1412:
1394:
1384:
1360:
1351:
1350:
1302:
1296:
1295:
1285:
1253:
1247:
1246:
1228:
1204:
1198:
1197:
1187:
1169:
1145:
1139:
1138:
1112:
1080:
1074:
1073:
1055:
1046:(52): 42938â44.
1031:
1025:
1024:
1006:
982:
976:
975:
957:
951:
950:
925:
831:Alexander Oparin
722:nuclear speckles
685:, are a form of
662:polyelectrolytes
650:
648:
647:
642:
634:
632:
631:
630:
617:
613:
612:
603:
602:
592:
573:
571:
570:
565:
557:
555:
547:
543:
542:
529:
517:
515:
514:
509:
480:
479:
462:
460:
459:
454:
449:
448:
436:
432:
383:
375:
365:
364:
349:
348:
325:
323:
322:
317:
312:
311:
251:
243:
235:
234:
146:(protocells) on
105:
104:
101:
100:
97:
94:
91:
88:
85:
82:
79:
76:
67:
66:
63:
62:
59:
56:
53:
50:
47:
44:
41:
38:
4555:
4554:
4550:
4549:
4548:
4546:
4545:
4544:
4520:
4519:
4518:
4513:
4353:
4343:
4313:
4288:
4199:Finnlakeviridae
4144:
4123:
4065:
4061:
4042:
4014:
3979:
3930:
3920:
3890:
3885:
3835:Synthetic virus
3823:Artificial cell
3796:
3724:
3633:
3522:RNA replication
3517:DNA replication
3505:
3496:Group II intron
3394:
3384:
3372:
3363:Mammalian prion
3345:
3321:
3300:dsRNA satellite
3297:ssDNA satellite
3267:
3260:
3229:
3222:
3167:
3096:
3090:
3060:
3055:
3034:
3012:
2976:Clay hypothesis
2957:
2926:
2921:Primordial soup
2895:
2890:
2860:
2859:
2798:
2794:
2763:
2759:
2704:
2700:
2680:
2679:
2663:
2659:
2645:
2636:
2635:
2626:
2625:
2613:
2598:
2594:
2563:
2559:
2536:10.2307/2420646
2520:
2516:
2508:
2502:
2498:
2467:
2463:
2451:
2449:
2440:
2439:
2428:
2427:
2423:
2380:
2376:
2313:
2309:
2255:
2251:
2198:
2194:
2140:
2136:
2091:
2084:
2031:
2027:
1966:
1962:
1909:
1905:
1844:
1837:
1825:
1823:
1814:
1813:
1795:
1791:
1759:
1755:
1743:
1741:
1732:
1731:
1713:
1709:
1656:
1652:
1607:
1603:
1549:
1542:
1523:
1519:
1480:
1473:
1420:
1416:
1361:
1354:
1317:(11): 899â904.
1303:
1299:
1254:
1250:
1205:
1201:
1146:
1142:
1081:
1077:
1032:
1028:
997:(26): 4727â34.
983:
979:
958:
954:
948:
926:
922:
917:
909:Artificial cell
900:
867:primordial soup
855:
802:
726:gene expression
710:stress granules
679:
626:
622:
618:
608:
604:
598:
594:
593:
591:
589:
586:
585:
548:
538:
534:
530:
528:
526:
523:
522:
475:
471:
469:
466:
465:
444:
440:
374:
373:
369:
360:
356:
338:
334:
332:
329:
328:
307:
303:
242:
224:
220:
218:
215:
214:
209:
205:
197:
193:
172:
144:primitive cells
136:bees in a swarm
73:
69:
35:
31:
17:
12:
11:
5:
4553:
4543:
4542:
4540:Origin of life
4537:
4532:
4515:
4514:
4512:
4511:
4506:
4501:
4500:
4499:
4489:
4484:
4479:
4474:
4473:
4472:
4467:
4462:
4457:
4449:
4441:
4440:
4439:
4429:
4424:
4419:
4410:
4405:
4400:
4395:
4390:
4385:
4380:
4379:
4378:
4373:
4363:
4357:
4355:
4349:
4348:
4345:
4344:
4342:
4341:
4336:
4329:
4327:
4323:
4322:
4319:
4318:
4315:
4314:
4312:
4311:
4304:
4296:
4294:
4290:
4289:
4287:
4286:
4279:
4272:
4265:
4258:
4251:
4248:Polydnaviridae
4244:
4237:
4230:
4223:
4216:
4213:Globuloviridae
4209:
4206:Fuselloviridae
4202:
4195:
4188:
4185:Bicaudaviridae
4181:
4174:
4167:
4164:Ampullaviridae
4160:
4152:
4150:
4146:
4145:
4141:Naldaviricetes
4138:
4136:
4129:
4125:
4124:
4122:
4121:
4114:
4107:
4100:
4093:
4086:
4078:
4076:
4069:
4052:
4048:
4047:
4044:
4043:
4041:
4040:
4034:
4026:
4024:
4021:Incertae sedis
4016:
4015:
4013:
4012:
4005:
4000:
3995:
3989:
3987:
3981:
3980:
3978:
3977:
3972:
3971:
3970:
3965:
3955:
3949:
3947:
3936:
3932:
3931:
3919:
3918:
3911:
3904:
3896:
3887:
3886:
3884:
3883:
3878:
3877:
3876:
3871:
3861:
3855:
3849:
3848:
3847:
3842:
3832:
3827:
3826:
3825:
3820:
3810:
3804:
3802:
3798:
3797:
3795:
3794:
3793:
3792:
3787:
3779:
3774:
3769:
3764:
3758:
3757:
3756:
3745:
3740:
3734:
3732:
3726:
3725:
3723:
3722:
3717:
3716:
3715:
3710:
3702:
3700:Kappa organism
3697:
3696:
3695:
3690:
3685:
3680:
3675:
3665:
3664:
3663:
3658:
3647:
3645:
3639:
3638:
3635:
3634:
3632:
3631:
3630:
3629:
3624:
3614:
3613:
3612:
3607:
3602:
3597:
3587:
3586:
3585:
3575:
3574:
3573:
3571:Non-coding DNA
3568:
3563:
3553:
3552:
3551:
3546:
3541:
3536:
3526:
3525:
3524:
3513:
3511:
3507:
3506:
3504:
3503:
3498:
3493:
3491:Group I intron
3488:
3483:
3482:
3481:
3471:
3470:
3469:
3466:
3457:
3454:
3449:
3444:
3434:
3433:
3432:
3427:
3417:
3416:
3415:
3413:Genomic island
3410:
3399:
3397:
3393:Mobile genetic
3386:
3378:
3377:
3374:
3373:
3371:
3370:
3365:
3359:
3357:
3351:
3350:
3347:
3346:
3344:
3343:
3342:
3341:
3338:
3329:
3327:
3323:
3322:
3320:
3319:
3318:
3317:
3314:
3308:
3301:
3298:
3295:
3292:
3285:
3281:
3279:
3270:
3262:
3261:
3259:
3258:
3251:
3243:
3241:
3232:
3224:
3223:
3221:
3220:
3218:dsDNA-RT virus
3215:
3213:ssRNA-RT virus
3210:
3208:(â)ssRNA virus
3205:
3203:(+)ssRNA virus
3200:
3195:
3190:
3189:
3188:
3177:
3175:
3169:
3168:
3166:
3165:
3164:
3163:
3158:
3148:Incertae sedis
3144:
3143:
3142:
3137:
3132:
3127:
3117:
3112:
3106:
3104:
3098:
3097:
3089:
3088:
3081:
3074:
3066:
3057:
3056:
3054:
3053:
3048:
3042:
3040:
3036:
3035:
3033:
3032:
3026:
3020:
3018:
3014:
3013:
3011:
3010:
3005:
3000:
2995:
2990:
2989:
2988:
2978:
2973:
2967:
2965:
2959:
2958:
2956:
2955:
2950:
2945:
2940:
2934:
2932:
2928:
2927:
2925:
2924:
2923:(19th century)
2918:
2912:
2905:
2903:
2897:
2896:
2893:Origin of life
2889:
2888:
2881:
2874:
2866:
2858:
2857:
2792:
2757:
2698:
2657:
2611:
2592:
2557:
2514:
2496:
2461:
2452:|journal=
2421:
2394:(4): 198â201.
2374:
2327:(1): 123â133.
2307:
2249:
2192:
2134:
2105:(1): 171â194.
2082:
2045:(3): 637â650.
2025:
1960:
1923:(1): 202â211.
1903:
1835:
1826:|journal=
1789:
1753:
1744:|journal=
1707:
1650:
1621:(3): 419â434.
1601:
1564:(6): 420â435.
1540:
1517:
1471:
1414:
1375:(3): 419â434.
1352:
1311:Nature Physics
1297:
1248:
1199:
1140:
1075:
1026:
977:
952:
946:
919:
918:
916:
913:
912:
911:
906:
899:
896:
876:Stanley Miller
863:J.B.S. Haldane
854:
851:
813:Dutch chemist
801:
798:
767:stabilised by
678:
675:
640:
637:
629:
625:
621:
616:
611:
607:
601:
597:
563:
560:
554:
551:
546:
541:
537:
533:
507:
504:
501:
498:
495:
492:
489:
486:
483:
478:
474:
452:
447:
443:
439:
435:
431:
428:
425:
422:
419:
416:
413:
410:
407:
404:
401:
398:
395:
392:
389:
386:
381:
378:
372:
368:
363:
359:
355:
352:
347:
344:
341:
337:
315:
310:
306:
302:
299:
296:
293:
290:
287:
284:
281:
278:
275:
272:
269:
266:
263:
260:
257:
254:
249:
246:
241:
238:
233:
230:
227:
223:
207:
203:
195:
191:
171:
170:Thermodynamics
168:
108:macromolecules
15:
9:
6:
4:
3:
2:
4552:
4541:
4538:
4536:
4533:
4531:
4528:
4527:
4525:
4510:
4507:
4505:
4502:
4498:
4495:
4494:
4493:
4490:
4488:
4487:Tandem repeat
4485:
4483:
4480:
4478:
4475:
4471:
4468:
4466:
4463:
4461:
4458:
4456:
4453:
4452:
4450:
4448:
4445:
4442:
4438:
4435:
4434:
4433:
4430:
4428:
4425:
4423:
4420:
4417:
4416:
4415:Nanobacterium
4411:
4409:
4406:
4404:
4401:
4399:
4396:
4394:
4391:
4389:
4386:
4384:
4381:
4377:
4374:
4372:
4369:
4368:
4367:
4364:
4362:
4359:
4358:
4356:
4350:
4340:
4337:
4334:
4331:
4330:
4328:
4324:
4310:
4309:
4308:Rhizidiovirus
4305:
4303:
4302:
4298:
4297:
4295:
4291:
4285:
4284:
4280:
4278:
4277:
4276:Thaspiviridae
4273:
4271:
4270:
4266:
4264:
4263:
4262:Pospiviroidae
4259:
4257:
4256:
4252:
4250:
4249:
4245:
4243:
4242:
4241:Plasmaviridae
4238:
4236:
4235:
4231:
4229:
4228:
4227:Halspiviridae
4224:
4222:
4221:
4217:
4215:
4214:
4210:
4208:
4207:
4203:
4201:
4200:
4196:
4194:
4193:
4189:
4187:
4186:
4182:
4180:
4179:
4178:Avsunviroidae
4175:
4173:
4172:
4171:Anelloviridae
4168:
4166:
4165:
4161:
4159:
4158:
4154:
4153:
4151:
4147:
4143:
4142:
4137:
4133:
4130:
4126:
4120:
4119:
4115:
4113:
4112:
4108:
4106:
4105:
4101:
4099:
4098:
4094:
4092:
4091:
4090:Duplodnaviria
4087:
4085:
4084:
4080:
4079:
4077:
4073:
4070:
4068:
4064:
4060:
4056:
4053:
4049:
4038:
4035:
4033:
4032:
4028:
4027:
4025:
4023:
4022:
4017:
4010:
4006:
4004:
4001:
3999:
3996:
3994:
3991:
3990:
3988:
3986:
3982:
3976:
3973:
3969:
3966:
3964:
3963:Mitochondrion
3961:
3960:
3959:
3956:
3954:
3951:
3950:
3948:
3945:
3940:
3937:
3935:Cellular life
3933:
3928:
3924:
3917:
3912:
3910:
3905:
3903:
3898:
3897:
3894:
3882:
3879:
3875:
3872:
3870:
3867:
3866:
3865:
3862:
3860:
3856:
3854:
3853:Nanobacterium
3850:
3846:
3843:
3841:
3838:
3837:
3836:
3833:
3831:
3828:
3824:
3821:
3819:
3818:Cell division
3816:
3815:
3814:
3811:
3809:
3806:
3805:
3803:
3799:
3791:
3788:
3786:
3783:
3782:
3780:
3778:
3775:
3773:
3770:
3768:
3765:
3763:
3759:
3755:
3752:
3751:
3750:
3746:
3744:
3741:
3739:
3736:
3735:
3733:
3731:
3727:
3721:
3718:
3714:
3711:
3709:
3706:
3705:
3703:
3701:
3698:
3694:
3691:
3689:
3686:
3684:
3681:
3679:
3676:
3674:
3671:
3670:
3669:
3666:
3662:
3661:Hydrogenosome
3659:
3657:
3654:
3653:
3652:
3651:Mitochondrion
3649:
3648:
3646:
3644:
3643:Endosymbiosis
3640:
3628:
3625:
3623:
3622:Tandem repeat
3620:
3619:
3618:
3615:
3611:
3608:
3606:
3603:
3601:
3598:
3596:
3593:
3592:
3591:
3588:
3584:
3581:
3580:
3579:
3576:
3572:
3569:
3567:
3564:
3562:
3559:
3558:
3557:
3554:
3550:
3547:
3545:
3542:
3540:
3537:
3535:
3532:
3531:
3530:
3527:
3523:
3520:
3519:
3518:
3515:
3514:
3512:
3510:Other aspects
3508:
3502:
3499:
3497:
3494:
3492:
3489:
3487:
3484:
3480:
3477:
3476:
3475:
3472:
3467:
3465:
3461:
3458:
3455:
3453:
3450:
3448:
3445:
3443:
3440:
3439:
3438:
3435:
3431:
3428:
3426:
3423:
3422:
3421:
3418:
3414:
3411:
3409:
3406:
3405:
3404:
3401:
3400:
3398:
3396:
3390:
3387:
3383:
3379:
3369:
3366:
3364:
3361:
3360:
3358:
3356:
3352:
3339:
3336:
3335:
3334:
3331:
3330:
3328:
3324:
3315:
3312:
3311:
3309:
3306:
3302:
3299:
3296:
3293:
3290:
3286:
3283:
3282:
3280:
3278:
3274:
3271:
3269:
3263:
3257:
3256:
3255:Avsunviroidae
3252:
3250:
3249:
3248:Pospiviroidae
3245:
3244:
3242:
3240:
3236:
3233:
3231:
3225:
3219:
3216:
3214:
3211:
3209:
3206:
3204:
3201:
3199:
3196:
3194:
3191:
3187:
3184:
3183:
3182:
3179:
3178:
3176:
3174:
3170:
3162:
3159:
3157:
3156:
3152:
3151:
3150:
3149:
3145:
3141:
3138:
3136:
3133:
3131:
3128:
3126:
3123:
3122:
3121:
3118:
3116:
3113:
3111:
3108:
3107:
3105:
3103:
3102:Cellular life
3099:
3094:
3087:
3082:
3080:
3075:
3073:
3068:
3067:
3064:
3052:
3049:
3047:
3044:
3043:
3041:
3037:
3030:
3027:
3025:
3022:
3021:
3019:
3015:
3009:
3006:
3004:
3001:
2999:
2996:
2994:
2991:
2987:
2984:
2983:
2982:
2979:
2977:
2974:
2972:
2969:
2968:
2966:
2964:
2960:
2954:
2951:
2949:
2946:
2944:
2941:
2939:
2936:
2935:
2933:
2929:
2922:
2919:
2916:
2913:
2910:
2907:
2906:
2904:
2902:
2898:
2894:
2887:
2882:
2880:
2875:
2873:
2868:
2867:
2864:
2853:
2849:
2844:
2839:
2835:
2831:
2827:
2823:
2819:
2815:
2811:
2807:
2803:
2796:
2788:
2784:
2780:
2776:
2772:
2768:
2761:
2753:
2749:
2745:
2741:
2737:
2733:
2729:
2725:
2721:
2717:
2713:
2709:
2702:
2694:
2690:
2684:
2676:
2672:
2668:
2661:
2653:
2649:
2642:
2630:
2622:
2618:
2614:
2608:
2604:
2596:
2588:
2584:
2580:
2576:
2572:
2568:
2561:
2553:
2549:
2545:
2541:
2537:
2533:
2529:
2525:
2518:
2507:
2500:
2492:
2488:
2484:
2480:
2476:
2472:
2465:
2457:
2444:
2436:
2432:
2425:
2417:
2413:
2409:
2405:
2401:
2397:
2393:
2389:
2385:
2378:
2370:
2366:
2362:
2358:
2353:
2348:
2344:
2340:
2335:
2330:
2326:
2322:
2318:
2311:
2303:
2299:
2294:
2289:
2285:
2281:
2277:
2273:
2269:
2265:
2261:
2253:
2245:
2241:
2236:
2231:
2227:
2223:
2219:
2215:
2211:
2207:
2203:
2196:
2188:
2184:
2180:
2176:
2172:
2168:
2163:
2158:
2154:
2150:
2146:
2138:
2130:
2126:
2122:
2118:
2113:
2108:
2104:
2100:
2096:
2089:
2087:
2078:
2074:
2069:
2064:
2060:
2056:
2052:
2048:
2044:
2040:
2036:
2029:
2021:
2017:
2013:
2009:
2005:
2001:
1996:
1991:
1987:
1983:
1979:
1975:
1971:
1964:
1956:
1952:
1947:
1942:
1938:
1934:
1930:
1926:
1922:
1918:
1914:
1907:
1899:
1895:
1890:
1885:
1881:
1877:
1873:
1869:
1865:
1861:
1857:
1853:
1849:
1842:
1840:
1831:
1818:
1809:
1804:
1800:
1793:
1785:
1781:
1777:
1773:
1769:
1765:
1757:
1749:
1736:
1727:
1722:
1718:
1711:
1703:
1699:
1694:
1689:
1685:
1681:
1677:
1673:
1669:
1665:
1661:
1654:
1646:
1642:
1637:
1632:
1628:
1624:
1620:
1616:
1612:
1605:
1597:
1593:
1588:
1583:
1579:
1575:
1571:
1567:
1563:
1559:
1555:
1547:
1545:
1536:
1532:
1528:
1521:
1513:
1509:
1505:
1501:
1497:
1493:
1489:
1485:
1478:
1476:
1467:
1463:
1458:
1453:
1449:
1445:
1441:
1437:
1433:
1429:
1425:
1418:
1410:
1406:
1402:
1398:
1393:
1388:
1383:
1378:
1374:
1370:
1366:
1359:
1357:
1348:
1344:
1340:
1336:
1332:
1328:
1324:
1320:
1316:
1312:
1308:
1301:
1293:
1289:
1284:
1279:
1275:
1271:
1268:(1â2): 2â11.
1267:
1263:
1259:
1252:
1244:
1240:
1236:
1232:
1227:
1222:
1218:
1214:
1210:
1203:
1195:
1191:
1186:
1181:
1177:
1173:
1168:
1163:
1159:
1155:
1151:
1144:
1136:
1132:
1128:
1124:
1120:
1116:
1111:
1106:
1102:
1098:
1094:
1090:
1086:
1079:
1071:
1067:
1063:
1059:
1054:
1049:
1045:
1041:
1037:
1030:
1022:
1018:
1014:
1010:
1005:
1000:
996:
992:
988:
981:
973:
969:
965:
964:
956:
949:
943:
939:
935:
931:
924:
920:
910:
907:
905:
902:
901:
895:
892:
888:
883:
881:
877:
873:
868:
864:
860:
850:
847:
843:
840:
834:
832:
827:
825:
821:
816:
811:
808:
797:
795:
791:
787:
783:
779:
775:
770:
766:
765:Balbiani body
762:
758:
754:
750:
745:
742:
738:
733:
731:
727:
723:
719:
715:
711:
706:
704:
703:lipid bilayer
700:
696:
692:
691:mitochondrion
688:
684:
670:
666:
663:
657:
654:
638:
635:
627:
623:
614:
609:
599:
578:
574:
561:
558:
552:
544:
539:
518:
502:
499:
496:
490:
487:
484:
481:
476:
463:
450:
445:
437:
433:
426:
423:
420:
414:
411:
405:
402:
399:
393:
390:
387:
384:
379:
376:
370:
366:
361:
357:
353:
350:
345:
342:
339:
326:
313:
308:
300:
294:
291:
288:
282:
279:
273:
270:
267:
261:
258:
255:
252:
247:
244:
239:
236:
231:
228:
225:
212:
201:
200:Flory-Huggins
189:
184:
181:
177:
167:
165:
161:
155:
153:
149:
145:
141:
137:
131:
129:
125:
121:
120:nucleic acids
117:
113:
109:
103:
65:
29:
21:
4504:Transpoviron
4437:Fungal prion
4413:
4306:
4301:Dinodnavirus
4299:
4281:
4274:
4269:Spiraviridae
4267:
4260:
4253:
4246:
4239:
4234:Ovaliviridae
4232:
4225:
4220:Guttaviridae
4218:
4211:
4204:
4197:
4192:Clavaviridae
4190:
4183:
4176:
4169:
4162:
4155:
4139:
4118:Varidnaviria
4116:
4109:
4102:
4097:Monodnaviria
4095:
4088:
4081:
4029:
4019:
3840:Viral vector
3766:
3683:Gerontoplast
3610:Transpoviron
3382:Nucleic acid
3368:Fungal prion
3266:Helper-virus
3253:
3246:
3153:
3146:
3051:Paleobiology
3046:Astrobiology
2809:
2805:
2795:
2770:
2766:
2760:
2711:
2707:
2701:
2666:
2660:
2602:
2595:
2570:
2566:
2560:
2527:
2523:
2517:
2499:
2477:(1): 39â48.
2474:
2470:
2464:
2443:cite journal
2424:
2391:
2387:
2377:
2324:
2320:
2310:
2267:
2263:
2252:
2209:
2205:
2195:
2152:
2148:
2137:
2102:
2098:
2042:
2038:
2028:
1977:
1973:
1963:
1920:
1916:
1906:
1855:
1851:
1817:cite journal
1792:
1767:
1763:
1756:
1735:cite journal
1710:
1667:
1663:
1653:
1618:
1614:
1604:
1561:
1557:
1526:
1520:
1490:(S1): 7â26.
1487:
1483:
1431:
1427:
1417:
1372:
1368:
1314:
1310:
1300:
1265:
1261:
1251:
1216:
1212:
1202:
1157:
1153:
1143:
1092:
1088:
1078:
1043:
1039:
1029:
994:
990:
980:
962:
955:
929:
923:
884:
856:
848:
844:
839:Debye-HĂŒckel
835:
828:
823:
819:
812:
807:flocculation
803:
757:random coils
746:
737:Cajal bodies
734:
707:
680:
658:
652:
583:
519:
464:
327:
213:
185:
179:
173:
156:
135:
132:
130:candidates.
123:
27:
26:
4447:microsphere
4366:Cancer cell
4111:Ribozyviria
3864:Cancer cell
3730:Abiogenesis
3678:Chromoplast
3673:Chloroplast
3456:Degradative
3198:dsRNA virus
3193:ssDNA virus
3186:Giant virus
3181:dsDNA virus
2812:(1): 3643.
2637:|last=
887:abiogenesis
880:Harold Urey
872:proteinoids
148:early Earth
4524:Categories
4497:Retroposon
4444:Proteinoid
4354:structures
4352:Comparable
4128:Unassigned
4031:Parakaryon
3944:Prokaryota
3772:Proteinoid
3767:Coacervate
3720:Nitroplast
3713:Trophosome
3708:Bacteriome
3693:Apicoplast
3688:Leucoplast
3529:Chromosome
3447:Resistance
3155:Parakaryon
2963:Protocells
2909:Panspermia
2773:(13): no.
2767:ChemInform
2530:(2): 472.
1529:. Centen.
915:References
741:P granules
164:Tony Hyman
28:Coacervate
4104:Riboviria
4083:Adnaviria
4067:Satellite
3985:Eukaryota
3781:Research
3762:Protocell
3501:Retrozyme
3460:Virulence
3442:Fertility
3289:Virophage
3277:Satellite
3268:dependent
3120:Eukaryota
3008:RNA world
2993:PAH world
2834:2041-1723
2787:0931-7597
2736:0028-0836
2683:cite book
2675:927006170
2629:cite book
2621:935643436
2587:0022-3654
2544:0003-0031
2491:0303-402X
2408:0372-820X
2343:0092-8674
2284:0261-4189
2226:0969-2126
2206:Structure
2171:0092-8674
2121:0066-4197
2059:0092-8674
2004:0036-8075
1937:1059-1524
1880:0028-0836
1684:0092-8674
1578:0962-8924
1535:901788902
1504:0095-9898
1448:1520-6106
1339:1745-2473
1235:0743-7463
1176:2296-889X
1119:0036-8075
904:Protocell
891:RNA-world
755:and form
718:nucleolus
620:∂
606:Δ
596:∂
550:∂
536:Δ
532:∂
503:ϕ
500:−
491:ϕ
488:χ
473:Δ
442:Δ
427:ϕ
424:−
415:
406:ϕ
403:−
391:ϕ
388:
377:ϕ
336:Δ
305:Δ
295:ϕ
292:−
283:
274:ϕ
271:−
259:ϕ
256:
245:ϕ
222:Δ
128:protocell
4477:Ribozyme
4422:Phagemid
4149:Families
4009:Protista
3993:Animalia
3958:Bacteria
3808:Organism
3801:See also
3777:Sulphobe
3754:Ribozyme
3749:RNA life
3656:Mitosome
3600:Prophage
3595:Provirus
3583:Replicon
3539:Circular
3486:Phagemid
3403:Mobilome
3395:elements
3305:Virusoid
3228:Subviral
3140:Protista
3125:Animalia
3110:Bacteria
3039:Research
2852:30194374
2744:14243409
2416:98519794
2369:18550463
2361:26406374
2302:29438978
2244:27545621
2187:14098476
2179:26317470
2129:31430179
2077:27471966
2020:42229928
2012:19460965
1955:15509651
1898:22398450
1784:86838379
1702:28283059
1645:30682370
1596:29602697
1512:13449108
1466:32118433
1409:59273868
1401:30682370
1292:21377640
1243:27599198
1213:Langmuir
1194:31001538
1135:42229928
1127:19460965
1062:16227622
1013:15579565
898:See also
699:lysosome
653:spinodal
116:proteins
112:polymers
4509:Xenobot
4427:Plasmid
4408:Jeewanu
4339:Obelisk
4135:Classes
4003:Plantae
3968:Plastid
3953:Archaea
3790:Jeewanu
3704:Organs
3668:Plastid
3468:Cryptic
3437:Plasmid
3135:Plantae
3115:Archaea
2843:6128941
2814:Bibcode
2752:7194753
2716:Bibcode
2552:2420646
2352:5149108
2293:5830921
2235:5014597
2068:5082712
1982:Bibcode
1974:Science
1889:3343696
1860:Bibcode
1693:5401687
1636:6445271
1587:6034118
1457:7104237
1392:6445271
1347:4643961
1319:Bibcode
1283:3476850
1185:6456709
1097:Bibcode
1089:Science
1070:7746883
1021:1104838
824:acervus
800:History
782:hnRNPA1
769:ÎČ-sheet
695:nucleus
651:), the
176:colloid
4455:Retron
4398:Fosmid
4383:Cosmid
4333:Nanobe
4293:Genera
4075:Realms
4063:Viroid
3881:Virome
3859:Nanobe
3556:Genome
3534:Linear
3479:Fosmid
3474:Cosmid
3239:Viroid
3230:agents
3031:(LUCA)
2850:
2840:
2832:
2785:
2750:
2742:
2734:
2708:Nature
2673:
2619:
2609:
2585:
2550:
2542:
2489:
2414:
2406:
2367:
2359:
2349:
2341:
2300:
2290:
2282:
2242:
2232:
2224:
2185:
2177:
2169:
2127:
2119:
2075:
2065:
2057:
2018:
2010:
2002:
1953:
1946:539164
1943:
1935:
1896:
1886:
1878:
1852:Nature
1782:
1700:
1690:
1682:
1643:
1633:
1594:
1584:
1576:
1533:
1510:
1502:
1464:
1454:
1446:
1407:
1399:
1389:
1345:
1337:
1290:
1280:
1241:
1233:
1192:
1182:
1174:
1160:: 21.
1133:
1125:
1117:
1068:
1060:
1019:
1011:
944:
788:(e.g.
778:TDP-43
4432:Prion
4326:Other
4059:Virus
3998:Fungi
3355:Prion
3326:Other
3173:Virus
3130:Fungi
2748:S2CID
2548:JSTOR
2509:(PDF)
2412:S2CID
2365:S2CID
2270:(5).
2183:S2CID
2016:S2CID
1780:S2CID
1405:S2CID
1343:S2CID
1131:S2CID
1066:S2CID
1017:S2CID
792:, or
4371:HeLa
3975:LUCA
3923:Life
3869:HeLa
3813:Cell
3561:Gene
2848:PMID
2830:ISSN
2783:ISSN
2740:PMID
2732:ISSN
2693:link
2689:link
2671:OCLC
2669:. .
2652:link
2648:link
2641:help
2617:OCLC
2607:ISBN
2583:ISSN
2540:ISSN
2487:ISSN
2456:help
2404:ISSN
2357:PMID
2339:ISSN
2321:Cell
2298:PMID
2280:ISSN
2240:PMID
2222:ISSN
2175:PMID
2167:ISSN
2149:Cell
2125:PMID
2117:ISSN
2073:PMID
2055:ISSN
2039:Cell
2008:PMID
2000:ISSN
1951:PMID
1933:ISSN
1894:PMID
1876:ISSN
1830:help
1748:help
1698:PMID
1680:ISSN
1664:Cell
1641:PMID
1615:Cell
1592:PMID
1574:ISSN
1531:OCLC
1508:PMID
1500:ISSN
1462:PMID
1444:ISSN
1397:PMID
1369:Cell
1335:ISSN
1288:PMID
1239:PMID
1231:ISSN
1190:PMID
1172:ISSN
1123:PMID
1115:ISSN
1058:PMID
1009:PMID
942:ISBN
878:and
780:and
739:and
162:and
4335:(?)
4039:(?)
3452:Col
3340:DNA
3337:RNA
3316:DNA
3313:RNA
2838:PMC
2822:doi
2775:doi
2724:doi
2712:205
2575:doi
2532:doi
2479:doi
2431:doi
2396:doi
2347:PMC
2329:doi
2325:163
2288:PMC
2272:doi
2230:PMC
2214:doi
2157:doi
2153:162
2107:doi
2063:PMC
2047:doi
2043:166
1990:doi
1978:324
1941:PMC
1925:doi
1884:PMC
1868:doi
1856:483
1803:doi
1772:doi
1721:doi
1688:PMC
1672:doi
1668:168
1631:PMC
1623:doi
1619:176
1582:PMC
1566:doi
1492:doi
1452:PMC
1436:doi
1432:124
1387:PMC
1377:doi
1373:176
1327:doi
1278:PMC
1270:doi
1266:167
1221:doi
1180:PMC
1162:doi
1105:doi
1093:324
1048:doi
1044:280
999:doi
995:207
968:doi
934:doi
796:).
774:FUS
697:or
610:mix
540:mix
477:mix
446:mix
309:mix
208:mix
204:mix
196:mix
192:mix
118:or
68:or
52:ÉËr
4526::
3925:,
3464:Ti
2846:.
2836:.
2828:.
2820:.
2808:.
2804:.
2781:.
2771:43
2769:.
2746:.
2738:.
2730:.
2722:.
2710:.
2685:}}
2681:{{
2633::
2631:}}
2627:{{
2615:.
2581:.
2571:64
2569:.
2546:.
2538:.
2528:20
2526:.
2485:.
2475:50
2473:.
2447::
2445:}}
2441:{{
2410:.
2402:.
2390:.
2386:.
2363:.
2355:.
2345:.
2337:.
2323:.
2319:.
2296:.
2286:.
2278:.
2268:37
2266:.
2262:.
2238:.
2228:.
2220:.
2210:24
2208:.
2204:.
2181:.
2173:.
2165:.
2151:.
2147:.
2123:.
2115:.
2103:53
2101:.
2097:.
2085:^
2071:.
2061:.
2053:.
2041:.
2037:.
2014:.
2006:.
1998:.
1988:.
1976:.
1972:.
1949:.
1939:.
1931:.
1921:16
1919:.
1915:.
1892:.
1882:.
1874:.
1866:.
1854:.
1850:.
1838:^
1821::
1819:}}
1815:{{
1801:.
1778:.
1768:78
1766:.
1739::
1737:}}
1733:{{
1719:.
1696:.
1686:.
1678:.
1666:.
1662:.
1639:.
1629:.
1617:.
1613:.
1590:.
1580:.
1572:.
1562:28
1560:.
1556:.
1543:^
1506:.
1498:.
1488:49
1486:.
1474:^
1460:.
1450:.
1442:.
1430:.
1426:.
1403:.
1395:.
1385:.
1371:.
1367:.
1355:^
1341:.
1333:.
1325:.
1315:11
1313:.
1309:.
1286:.
1276:.
1264:.
1260:.
1237:.
1229:.
1217:32
1215:.
1211:.
1188:.
1178:.
1170:.
1156:.
1152:.
1129:.
1121:.
1113:.
1103:.
1091:.
1087:.
1064:.
1056:.
1042:.
1038:.
1015:.
1007:.
993:.
989:.
940:,
820:co
776:,
732:.
720:,
712:,
693:,
412:ln
385:ln
280:ln
253:ln
154:.
114:,
96:eÉȘ
90:Ér
78:oÊ
40:oÊ
4418:"
4412:"
4011:'
4007:'
3946:"
3942:"
3915:e
3908:t
3901:v
3857:?
3851:?
3760:â
3747:?
3462:/
3307:)
3291:)
3085:e
3078:t
3071:v
2885:e
2878:t
2871:v
2854:.
2824::
2816::
2810:9
2789:.
2777::
2754:.
2726::
2718::
2695:)
2677:.
2654:)
2643:)
2623:.
2589:.
2577::
2554:.
2534::
2511:.
2493:.
2481::
2458:)
2454:(
2437:.
2433::
2418:.
2398::
2392:8
2371:.
2331::
2304:.
2274::
2246:.
2216::
2189:.
2159::
2131:.
2109::
2079:.
2049::
2022:.
1992::
1984::
1957:.
1927::
1900:.
1870::
1862::
1832:)
1828:(
1811:.
1805::
1786:.
1774::
1750:)
1746:(
1729:.
1723::
1704:.
1674::
1647:.
1625::
1598:.
1568::
1537:.
1514:.
1494::
1468:.
1438::
1411:.
1379::
1349:.
1329::
1321::
1294:.
1272::
1245:.
1223::
1196:.
1164::
1158:6
1137:.
1107::
1099::
1072:.
1050::
1023:.
1001::
974:.
970::
936::
639:0
636:=
628:2
624:x
615:G
600:2
562:0
559:=
553:x
545:G
521:(
506:)
497:1
494:(
485:=
482:H
451:H
438:+
434:]
430:)
421:1
418:(
409:)
400:1
397:(
394:+
380:N
371:[
367:T
362:B
358:k
354:=
351:G
346:x
343:i
340:m
314:H
301:+
298:)
289:1
286:(
277:)
268:1
265:(
262:+
248:N
240:=
237:G
232:x
229:i
226:m
180:.
102:/
99:t
93:v
87:s
84:ĂŠ
81:Ë
75:k
72:/
64:/
61:t
58:É
55:v
49:s
46:Ë
43:É
37:k
34:/
30:(
Text is available under the Creative Commons Attribution-ShareAlike License. Additional terms may apply.
