555:. Many believe this limits their applicability in clinical settings. In spite of these beliefs, there have been many ligands that have reported successful pre-clinical animal studies. Given that some bivalent ligands can have many advantages compared to their monovalent counterparts (such as tissue selectivity, increased binding affinity, and increased potency or efficacy), bivalents may offer some clinical advantages as well.
42:
388:
295:
317:, the concentration required to produce the half-maximal response). High-affinity ligand binding implies that a relatively low concentration of a ligand is adequate to maximally occupy a ligand-binding site and trigger a physiological response. Receptor affinity is measured by an inhibition constant or K
1247:
Matera, Carlo; Flammini, Lisa; Quadri, Marta; Vivo, Valentina; Ballabeni, Vigilio; Holzgrabe, Ulrike; Mohr, Klaus; De Amici, Marco; Barocelli, Elisabetta (2014-03-21). "Bis(ammonio)alkane-type agonists of muscarinic acetylcholine receptors: Synthesis, in vitro functional characterization, and in vivo
563:
Ligands of proteins can be characterized also by the number of protein chains they bind. "Monodesmic" ligands (μόνος: single, δεσμός: binding) are ligands that bind a single protein chain, while "polydesmic" ligands (πολοί: many) are frequent in protein complexes, and are ligands that bind more than
367:
level) implies that a relatively high concentration of a ligand is required before the binding site is maximally occupied and the maximum physiological response to the ligand is achieved. In the example shown to the right, two different ligands bind to the same receptor binding site. Only one of the
272:(lifetime of the receptor-ligand complex) does not correlate. High-affinity binding of ligands to receptors is often physiologically important when some of the binding energy can be used to cause a conformational change in the receptor, resulting in altered behavior for example of an associated
1211:
Matera, Carlo; Pucci, Luca; Fiorentini, Chiara; Fucile, Sergio; Missale, Cristina; Grazioso, Giovanni; Clementi, Francesco; Zoli, Michele; De Amici, Marco (2015-08-28). "Bifunctional compounds targeting both D2 and non-α7 nACh receptors: Design, synthesis and pharmacological characterization".
511:
Bivalent ligands consist of two drug-like molecules (pharmacophores or ligands) connected by an inert linker. There are various kinds of bivalent ligands and are often classified based on what the pharmacophores target. Homobivalent ligands target two of the same receptor types. Heterobivalent
572:
A privileged scaffold is a molecular framework or chemical moiety that is statistically recurrent among known drugs or among a specific array of biologically active compounds. These privileged elements can be used as a basis for designing new active biological compounds or compound libraries.
503:) determining the affinity is complicated by non-specific hydrophobic interactions. Non-specific hydrophobic interactions can be overcome when the affinity of the ligand is high. For example, PIP2 binds with high affinity to PIP2 gated ion channels.
415:(MP-SPR) can not only quantify the affinity from concentration based assays; but also from the kinetics of association and dissociation, and in the later cases, the conformational change induced upon binding. MP-SPR also enables measurements in high
1512:
Nimczick M, Pemp D, Darras FH, Chen X, Heilmann J, Decker M (August 2014). "Synthesis and biological evaluation of bivalent cannabinoid receptor ligands based on hCB₂R selective benzimidazoles reveal unexpected intrinsic properties".
564:
one protein chain, typically in or near protein interfaces. Recent research shows that the type of ligands and binding site structure has profound consequences for the evolution, function, allostery and folding of protein compexes.
2290:
Kombarov R, Altieri A, Genis D, Kirpichenok M, Kochubey V, Rakitina N, Titarenko Z (February 2010). "BioCores: identification of a drug/natural product-based privileged structural motif for small-molecule lead discovery".
2762:
1548:
Russo O, Berthouze M, Giner M, Soulier JL, Rivail L, Sicsic S, Lezoualc'h F, Jockers R, Berque-Bestel I (September 2007). "Synthesis of specific bivalent probes that functionally interact with 5-HT(4) receptor dimers".
512:
ligands target two different receptor types. Bitopic ligands target an orthosteric binding sites and allosteric binding sites on the same receptor. In scientific research, bivalent ligands have been used to study
262:
The interaction of ligands with their binding sites can be characterized in terms of a binding affinity. In general, high-affinity ligand binding results from greater attractive forces between the ligand and its
267:
while low-affinity ligand binding involves less attractive force. In general, high-affinity binding results in a higher occupancy of the receptor by its ligand than is the case for low-affinity binding; the
453:
Ligand efficacy refers to the ability of the ligand to produce a biological response upon binding to the target receptor and the quantitative magnitude of this response. This response may be as an
2755:
1283:
Erez M, Takemori AE, Portoghese PS (July 1982). "Narcotic antagonistic potency of bivalent ligands which contain beta-naltrexamine. Evidence for bridging between proximal recognition sites".
1682:"A Direct in Vivo Comparison of the Melanocortin Monovalent Agonist Ac-His-DPhe-Arg-Trp-NH2 versus the Bivalent Agonist Ac-His-DPhe-Arg-Trp-PEDG20-His-DPhe-Arg-Trp-NH2: A Bivalent Advantage"
2748:
643:
Other techniques include: fluorescence intensity, bimolecular fluorescence complementation, FRET (fluorescent resonance energy transfer) / FRET quenching surface plasmon resonance,
479:
Selective ligands have a tendency to bind to very limited kinds of receptor, whereas non-selective ligands bind to several types of receptors. This plays an important role in
1073:
647:, Coimmunopreciptation indirect ELISA, equilibrium dialysis, gel electrophoresis, far western blot, fluorescence polarization anisotropy, electron paramagnetic resonance,
423:(MST), an immobilization-free method was developed. This method allows the determination of the binding affinity without any limitation to the ligand's molecular weight.
328:
value from a competition binding experiment where the concentration of a ligand required to displace 50% of a fixed concentration of reference ligand is determined. The K
1629:
Busnelli M, Kleinau G, Muttenthaler M, Stoev S, Manning M, Bibic L, Howell LA, McCormick PJ, Di Lascio S, Braida D, Sala M, Rovati GE, Bellini T, Chini B (August 2016).
581:
Main methods to study protein–ligand interactions are principal hydrodynamic and calorimetric techniques, and principal spectroscopic and structural methods such as
1804:"An in Vitro and in Vivo Investigation of Bivalent Ligands That Display Preferential Binding and Functional Activity for Different Melanocortin Receptor Homodimers"
192:
alters the conformation by affecting the three-dimensional shape orientation. The conformation of a receptor protein composes the functional state. Ligands include
2370:
658:
The dramatically increased computing power of supercomputers and personal computers has made it possible to study protein–ligand interactions also by means of
2808:
631:
431:
412:
1894:
Berque-Bestel I, Lezoualc'h F, Jockers R (December 2008). "Bivalent ligands as specific pharmacological tools for G protein-coupled receptor dimers".
1318:
Portoghese PS, Ronsisvalle G, Larson DL, Yim CB, Sayre LM, Takemori AE (1982). "Opioid agonist and antagonist bivalent ligands as receptor probes".
3004:
1584:
Soulier JL, Russo O, Giner M, Rivail L, Berthouze M, Ongeri S, Maigret B, Fischmeister R, Lezoualc'h F, Sicsic S, Berque-Bestel I (October 2005).
971:
Baaske P, Wienken CJ, Reineck P, Duhr S, Braun D (March 2010). "Optical thermophoresis for quantifying the buffer dependence of aptamer binding".
521:
228:
in solution. The solvent provides a chemical environment for the ligand and receptor to adapt, and thus accept or reject each other as partners.
2047:"Targeting putative mu opioid/metabotropic glutamate receptor-5 heteromers produces potent antinociception in a chronic murine bone cancer model"
3526:
1461:
Conn PM, Rogers DC, Stewart JM, Niedel J, Sheffield T (April 1982). "Conversion of a gonadotropin-releasing hormone antagonist to an agonist".
1139:
1108:
520:
and coworkers while studying the opioid receptor system. Bivalent ligands were also reported early on by
Micheal Conn and coworkers for the
2917:
1125:
403:
involve binding competition between a tagged ligand and an untagged ligand. Real-time based methods, which are often label-free, such as
3491:
2966:
2902:
2617:
3064:
3336:
2813:
2585:
3036:
2363:
376:. In this example, the concentration at which the full agonist (red curve) can half-maximally activate the receptor is about 5 x 10
3079:
3074:
302:
Agonist binding to a receptor can be characterized both in terms of how much physiological response can be triggered (that is, the
3293:
1980:"Opioid-induced tolerance and dependence in mice is modulated by the distance between pharmacophores in a bivalent ligand series"
1069:
626:
545:
408:
353:
3565:
2912:
551:
Bivalent ligands usually tend to be larger than their monovalent counterparts, and therefore, not 'drug-like' as in
Lipinski's
442:
Binding affinity data alone does not determine the overall potency of a drug or a naturally produced (biosynthesized) hormone.
954:
533:
2981:
2803:
1631:"Design and Characterization of Superpotent Bivalent Ligands Targeting Oxytocin Receptor Dimers via a Channel-Like Structure"
671:
585:
321:
value, the concentration required to occupy 50% of the receptor. Ligand affinities are most often measured indirectly as an
283:
A ligand that can bind to and alter the function of the receptor that triggers a physiological response is called a receptor
3470:
1931:"Inhibition of Inflammatory and Neuropathic Pain by Targeting a Mu Opioid Receptor/Chemokine Receptor5 Heteromer (MOR-CCR5)"
3321:
652:
2194:"Ligand-Binding-Site Structure Shapes Allosteric Signal Transduction and the Evolution of Allostery in Protein Complexes"
917:
2612:
2356:
247:
3288:
2986:
430:
in a quantitative study of the ligand-receptor binding affinity, see the comprehensive article on the configurational
662:. For example, a worldwide grid of well over a million ordinary PCs was harnessed for cancer research in the project
3480:
3346:
3178:
2431:
2414:
2397:
605:
525:
185:. In general, the interpretation of ligand is contextual with regards to what sort of binding has been observed.
3555:
3173:
2627:
2622:
2436:
1585:
648:
595:
420:
404:
357:
225:
216:, and this measurement typifies a tendency or strength of the effect. Binding affinity is actualized not only by
165:
bonding between a ligand and target molecule is atypical in biological systems. In contrast to the definition of
127:
1680:
Lensing CJ, Adank DN, Wilber SL, Freeman KT, Schnell SM, Speth RC, Zarth AT, Haskell-Luevano C (February 2017).
217:
2907:
2887:
2793:
2719:
2409:
807:
Teif VB, Rippe K (October 2010). "Statistical-mechanical lattice models for protein-DNA binding in chromatin".
644:
537:
500:
273:
205:
193:
158:
115:
1396:"Gonadotropin-releasing hormone stimulation of luteinizing hormone release: A ligand-receptor-effector model"
3550:
3414:
3351:
3326:
3209:
3129:
2788:
2547:
2519:
2446:
2426:
659:
615:
264:
189:
35:
1851:
Shonberg J, Scammells PJ, Capuano B (June 2011). "Design strategies for bivalent ligands targeting GPCRs".
3399:
3298:
3283:
3252:
3153:
2892:
2740:
2559:
2532:
675:
349:
201:
118:(conformation) of the target protein. In DNA-ligand binding studies, the ligand can be a small molecule,
3545:
3510:
3331:
3230:
3168:
3163:
3094:
3029:
2685:
2647:
2527:
2451:
2404:
1732:
Xu L, Josan JS, Vagner J, Caplan MR, Hruby VJ, Mash EA, Lynch RM, Morse DL, Gillies RJ (December 2012).
701:
529:
427:
341:
2096:"Ligand Binding Site Structure Shapes Folding, Assembly and Degradation of Homomeric Protein Complexes"
3419:
3262:
3257:
3139:
3134:
3121:
2656:
2652:
2604:
1991:
1745:
1470:
1407:
1033:
826:
763:
667:
636:
541:
517:
491:, because they bind to several other receptors in addition to the one generating the desired effect.
138:
445:
Potency is a result of the complex interplay of both the binding affinity and the ligand efficacy.
3515:
2798:
2714:
2697:
2383:
474:
458:
377:
288:
174:
154:
150:
83:
3204:
2897:
2316:
2145:"Ligand Binding Site Structure Influences the Evolution of Protein Complex Function and Topology"
2009:
1876:
1763:
1494:
1425:
850:
816:
721:
600:
590:
243:
79:
287:. Ligands that bind to a receptor but fail to activate the physiological response are receptor
3560:
3465:
3449:
3245:
3240:
3149:
2564:
2308:
2272:
2223:
2174:
2125:
2076:
2027:
1960:
1911:
1868:
1833:
1781:
1711:
1662:
1608:
1566:
1530:
1486:
1443:
1376:
1335:
1300:
1265:
1229:
1190:
1135:
1104:
1051:
988:
899:
842:
789:
696:
610:
416:
103:
54:
1929:
Akgün E, Javed MI, Lunzer MM, Powers MD, Sham YY, Watanabe Y, Portoghese PS (November 2015).
3475:
3444:
3439:
3394:
3022:
2951:
2690:
2661:
2590:
2486:
2419:
2300:
2262:
2254:
2213:
2205:
2164:
2156:
2115:
2107:
2066:
2058:
2017:
1999:
1950:
1942:
1903:
1860:
1823:
1815:
1771:
1753:
1701:
1693:
1652:
1642:
1600:
1558:
1522:
1478:
1433:
1415:
1366:
1327:
1292:
1257:
1221:
1180:
1172:
1096:
1041:
980:
889:
881:
834:
779:
771:
209:
197:
3485:
3389:
3199:
2569:
958:
731:
488:
462:
30:
This article is about ligands in biochemistry. For ligands in coordination chemistry, see
1100:
838:
310:
of the agonist that is required to produce the physiological response (often measured as
1995:
1802:
Lensing CJ, Freeman KT, Schnell SM, Adank DN, Speth RC, Haskell-Luevano C (April 2016).
1749:
1474:
1411:
1037:
830:
767:
524:. Since these early reports, there have been many bivalent ligands reported for various
2702:
2509:
2379:
2348:
2267:
2242:
2218:
2193:
2169:
2144:
2120:
2095:
2071:
2046:
2022:
1979:
1955:
1930:
1828:
1803:
1776:
1733:
1706:
1681:
1586:"Design and synthesis of specific probes for human 5-HT4 receptor dimerization studies"
1185:
1156:
1131:
894:
869:
784:
751:
269:
221:
99:
1438:
1395:
666:, which ended in April 2007. Grid.org has been succeeded by similar projects such as
3539:
3453:
3367:
3235:
3069:
3046:
2780:
2504:
1978:
Daniels DJ, Lenard NR, Etienne CL, Law PY, Roerig SC, Portoghese PS (December 2005).
1331:
1090:
513:
372:. An agonist that can only partially activate the physiological response is called a
307:
146:
2320:
1880:
3278:
3099:
3089:
2729:
2494:
2490:
1498:
951:
711:
679:
620:
552:
480:
368:
agonists shown can maximally stimulate the receptor and, thus, can be defined as a
235:
170:
107:
71:
67:
854:
41:
2160:
2062:
1946:
1819:
1697:
1647:
1630:
1371:
1354:
1261:
1225:
1176:
1089:
Hammes, Stephen R.; Mendelson, Carole R. (2011). "Mechanisms of
Hormone Action".
3214:
3194:
2991:
2976:
2724:
2537:
775:
716:
706:
499:
For hydrophobic ligands (e.g. PIP2) in complex with a hydrophobic protein (e.g.
231:
130:. The relationship between ligand and binding partner is a function of charge,
87:
3014:
2258:
1984:
Proceedings of the
National Academy of Sciences of the United States of America
1907:
1738:
Proceedings of the
National Academy of Sciences of the United States of America
1400:
Proceedings of the
National Academy of Sciences of the United States of America
17:
3458:
3158:
3111:
2971:
2956:
2929:
2677:
2304:
2111:
1526:
1022:"Protein-binding assays in biological liquids using microscale thermophoresis"
396:
182:
142:
131:
2342:
3409:
3377:
3144:
3050:
2882:
2707:
2441:
2209:
2004:
1758:
1420:
1000:
726:
516:
and to investigate their properties. This class of ligands was pioneered by
94:, which means 'to bind'. In protein-ligand binding, the ligand is usually a
45:
2312:
2276:
2227:
2178:
2129:
2080:
2031:
1964:
1915:
1872:
1864:
1837:
1785:
1715:
1666:
1612:
1570:
1534:
1380:
1269:
1233:
1194:
1055:
992:
984:
903:
846:
793:
2045:
Smeester BA, Lunzer MM, Akgün E, Beitz AJ, Portoghese PS (November 2014).
1734:"Heterobivalent ligands target cell-surface receptor combinations in vivo"
1490:
1447:
1339:
1304:
177:, in biochemistry it is ambiguous whether the ligand generally binds at a
3372:
3341:
2924:
2499:
2456:
663:
303:
251:
162:
95:
1767:
1296:
387:
3104:
2481:
2013:
1657:
1046:
1021:
691:
454:
284:
239:
123:
111:
1604:
1562:
885:
294:
58:
3084:
2823:
1482:
1429:
277:
166:
31:
220:
interactions, but also by solvent effects that can play a dominant,
1020:
Wienken CJ, Baaske P, Rothbauer U, Braun D, Duhr S (October 2010).
2961:
2866:
2861:
2856:
2851:
2846:
2841:
2836:
2831:
2339:, a public database of measured protein-ligand binding affinities.
821:
484:
386:
381:
293:
178:
40:
3382:
2944:
2939:
2934:
337:
322:
311:
49:
3018:
2744:
2352:
528:(GPCR) systems including cannabinoid, serotonin, oxytocin, and
119:
90:
to serve a biological purpose. The etymology stems from Latin
2243:"Privileged scaffolds for library design and drug discovery"
2345:, a comprehensive database for ligand-protein interactions.
2336:
1355:"Heteromer Induction: An Approach to Unique Pharmacology?"
1155:
Cabanos, C; Wang, M; Han, X; Hansen, SB (8 August 2017).
961:, A complete guide to nonlinear regression, curvefit.com.
1127:
A pharmacology primer: theory, applications, and methods
340:. Ligand affinities can also be measured directly as a
419:
dissociation buffers thanks to a unique optical setup.
752:"Ligand-induced DNA condensation: choosing the model"
395:
Binding affinity is most commonly determined using a
391:
Two ligands with different receptor binding affinity.
213:
465:, depending on the physiological response produced.
3503:
3432:
3360:
3314:
3307:
3271:
3223:
3187:
3120:
3057:
2875:
2822:
2779:
2670:
2640:
2603:
2578:
2546:
2518:
2472:
2465:
2390:
1353:Portoghese PS, Akgün E, Lunzer MM (January 2017).
27:Substance that forms a complex with a biomolecule
2771:Intercellular signaling peptides and proteins /
2241:Welsch ME, Snyder SA, Stockwell BR (June 2010).
1157:"A Soluble Fluorescent Binding Assay Reveals PIP
1074:this article in the web archive on 2012 April 28
868:Baron R, Setny P, McCammon JA (September 2010).
114:. The binding typically results in a change of
1727:
1725:
1070:Configuration integral (statistical mechanics)
3030:
2756:
2364:
1797:
1795:
1624:
1622:
8:
2809:Transforming growth factor beta superfamily
3311:
3037:
3023:
3015:
2763:
2749:
2741:
2552:
2469:
2371:
2357:
2349:
632:Multi-parametric surface plasmon resonance
413:multi-parametric surface plasmon resonance
401:Homologous competitive binding experiments
298:Two agonists with similar binding affinity
2266:
2217:
2168:
2119:
2070:
2021:
2003:
1954:
1827:
1775:
1757:
1705:
1656:
1646:
1437:
1419:
1370:
1248:evaluation of their analgesic activity".
1184:
1045:
893:
820:
783:
487:that are non-selective tend to have more
874:Journal of the American Chemical Society
1250:European Journal of Medicinal Chemistry
1214:European Journal of Medicinal Chemistry
742:
522:gonadotropin-releasing hormone receptor
3527:Index of evolutionary biology articles
1206:
1204:
952:Homologous competitive binding curves
7:
2918:Nephroblastoma overexpressed protein
1515:Bioorganic & Medicinal Chemistry
1084:
1082:
1072:, 2008. this wiki site is down; see
870:"Water in cavity-ligand recognition"
809:Journal of Physics: Condensed Matter
2967:Parathyroid hormone-related protein
2903:CCN intercellular signaling protein
2618:Transcription preinitiation complex
2247:Current Opinion in Chemical Biology
1896:Current Drug Discovery Technologies
3337:Evolutionary developmental biology
2814:Vascular endothelial growth factor
2586:Signal transducing adaptor protein
1394:Blum JJ, Conn PM (December 1982).
1101:10.1093/oso/9780199744121.003.0006
399:ligand, known as a tagged ligand.
25:
449:Drug or hormone binding efficacy
3294:Evolution of sexual reproduction
2051:European Journal of Pharmacology
1092:Textbook of Endocrine Physiology
627:Dual polarisation interferometry
438:Drug or hormone binding potency
409:dual-polarization interferometry
354:isothermal titration calorimetry
258:Receptor/ligand binding affinity
212:. The rate of binding is called
2913:Connective tissue growth factor
2198:Molecular Biology and Evolution
157:is actually reversible through
3065:Genotype–phenotype distinction
3005:extracellular ligand disorders
2982:Tolloid-like metalloproteinase
2804:Platelet-derived growth factor
1935:Journal of Medicinal Chemistry
1808:Journal of Medicinal Chemistry
1635:Journal of Medicinal Chemistry
1593:Journal of Medicinal Chemistry
1551:Journal of Medicinal Chemistry
1285:Journal of Medicinal Chemistry
1161:Antagonism of TREK-1 Channels"
839:10.1088/0953-8984/22/41/414105
672:Human Proteome Folding Project
586:Fourier transform spectroscopy
332:value can be estimated from IC
1:
3322:Regulation of gene expression
639:and radioligand binding assay
577:Methods used to study binding
3492:Endless Forms Most Beautiful
3272:Evolution of genetic systems
3080:Gene–environment correlation
3075:Gene–environment interaction
2192:Abrusan G, Marsh JA (2019).
2161:10.1016/j.celrep.2018.02.085
2143:Abrusan G, Marsh JA (2018).
2100:Journal of Molecular Biology
2094:Abrusan G, Marsh JA (2019).
2063:10.1016/j.ejphar.2014.09.008
1947:10.1021/acs.jmedchem.5b01245
1820:10.1021/acs.jmedchem.5b01894
1698:10.1021/acschemneuro.6b00399
1648:10.1021/acs.jmedchem.6b00564
1372:10.1021/acschemneuro.7b00002
1332:10.1016/0024-3205(82)90362-9
1262:10.1016/j.ejmech.2014.01.032
1226:10.1016/j.ejmech.2015.06.039
1177:10.1016/j.celrep.2017.07.034
559:Mono- and polydesmic ligands
363:Low-affinity binding (high K
3471:Christiane Nüsslein-Volhard
1095:. Oxford University Press.
776:10.1529/biophysj.105.063909
469:Selective and non-selective
134:, and molecular structure.
3582:
3347:Hedgehog signaling pathway
3224:Developmental architecture
2259:10.1016/j.cbpa.2010.02.018
1908:10.2174/157016308786733591
606:Nuclear magnetic resonance
526:G protein-coupled receptor
472:
161:. Measurably irreversible
29:
3524:
3174:Transgressive segregation
3000:
2555:
2305:10.1007/s11030-009-9157-5
2112:10.1016/j.jmb.2019.07.014
1686:ACS Chemical Neuroscience
1527:10.1016/j.bmc.2014.06.008
1359:ACS Chemical Neuroscience
1001:"A hot road to new drugs"
918:"The difference between K
649:microscale thermophoresis
596:Fluorescence spectroscopy
421:Microscale thermophoresis
405:surface plasmon resonance
358:surface plasmon resonance
52:(orange) bound. Based on
2908:Cysteine-rich protein 61
2888:Agouti signaling protein
2794:Fibroblast growth factor
2698:Neuroendocrine signaling
750:Teif VB (October 2005).
645:bio-layer interferometry
501:lipid-gated ion channels
348:) using methods such as
181:site, as is the case in
116:conformational isomerism
3352:Notch signaling pathway
3327:Gene regulatory network
3210:Dual inheritance theory
2789:Epidermal growth factor
2005:10.1073/pnas.0506627102
1759:10.1073/pnas.1211762109
1421:10.1073/pnas.79.23.7307
660:computational chemistry
616:Atomic force microscope
238:labeled compounds used
48:(blue) with its ligand
36:Ligand (disambiguation)
3566:Ligands (biochemistry)
3400:cis-regulatory element
3308:Control of development
3188:Non-genetic influences
3154:evolutionary landscape
2893:Agouti-related protein
2560:cAMP-dependent pathway
1865:10.1002/cmdc.201100101
985:10.1002/anie.200903998
676:Compute Against Cancer
392:
350:fluorescence quenching
338:Cheng Prusoff equation
306:) and in terms of the
299:
63:
34:. For other uses, see
3511:Nature versus nurture
3415:Cell surface receptor
3332:Evo-devo gene toolkit
3231:Developmental biology
3169:Polygenic inheritance
3095:Quantitative genetics
2987:Tumor necrosis factor
2686:Synaptic transmission
2682:Neurocrine signaling
2605:Transcription factors
2452:PI3K/AKT/mTOR pathway
2210:10.1093/molbev/msz093
1026:Nature Communications
702:Allosteric regulation
530:melanocortin receptor
428:statistical mechanics
390:
342:dissociation constant
297:
153:. The association or
139:intermolecular forces
44:
3420:Transcription factor
3135:Genetic assimilation
3122:Genetic architecture
1326:(12–13): 1283–1286.
1007:. February 24, 2010.
668:World Community Grid
637:Ligand binding assay
518:Philip S. Portoghese
226:non-covalent binding
188:Ligand binding to a
151:Van der Waals forces
3516:Morphogenetic field
3433:Influential figures
2799:Nerve growth factor
2715:Mechanotransduction
2703:Exocrine signalling
2510:Signaling molecules
2384:Signal transduction
2293:Molecular Diversity
1996:2005PNAS..10219208D
1990:(52): 19208–19213.
1750:2012PNAS..10921295X
1744:(52): 21295–21300.
1475:1982Natur.296..653C
1412:1982PNAS...79.7307B
1297:10.1021/jm00349a016
1124:Kenakin TP (2006).
1038:2010NatCo...1..100W
940:. 31 December 2019.
880:(34): 12091–12097.
831:2010JPCM...22O4105T
768:2005BpJ....89.2574T
756:Biophysical Journal
568:Privileged scaffold
495:Hydrophobic ligands
475:Binding selectivity
175:inorganic chemistry
126:which binds to the
3205:Genomic imprinting
2898:Angiogenic protein
2725:Ion channel gating
2457:Integrin receptors
2391:Signaling pathways
1047:10.1038/ncomms1093
957:2007-12-19 at the
722:DNA binding ligand
601:Circular dichroism
591:Raman spectroscopy
432:partition function
393:
300:
224:role which drives
137:Binding occurs by
64:
3533:
3532:
3466:Eric F. Wieschaus
3428:
3427:
3246:Pattern formation
3150:Fitness landscape
3012:
3011:
2738:
2737:
2720:Phototransduction
2678:Intracrine action
2636:
2635:
2599:
2598:
2487:Neurotransmitters
2155:(12): 3265–3276.
2106:(19): 3871–3888.
1941:(21): 8647–8657.
1641:(15): 7152–7166.
1605:10.1021/jm050234z
1599:(20): 6220–6228.
1563:10.1021/jm070552t
1557:(18): 4482–4492.
1521:(15): 3938–3946.
1469:(5858): 653–655.
1406:(23): 7307–7311.
1141:978-0-12-370599-0
1110:978-0-19-974412-1
979:(12): 2238–2241.
973:Angewandte Chemie
938:The Science Snail
886:10.1021/ja1050082
697:Schild regression
611:Mass spectrometry
532:systems, and for
254:binding studies.
210:neurotransmitters
98:which produces a
16:(Redirected from
3573:
3556:Chemical bonding
3476:William McGinnis
3445:Richard Lewontin
3440:C. H. Waddington
3312:
3289:Neutral networks
3039:
3032:
3025:
3016:
2952:Hedgehog protein
2765:
2758:
2751:
2742:
2691:Chemical synapse
2591:Scaffold protein
2553:
2548:Second messenger
2474:Receptor ligands
2470:
2373:
2366:
2359:
2350:
2325:
2324:
2287:
2281:
2280:
2270:
2238:
2232:
2231:
2221:
2204:(8): 1711–1727.
2189:
2183:
2182:
2172:
2140:
2134:
2133:
2123:
2091:
2085:
2084:
2074:
2042:
2036:
2035:
2025:
2007:
1975:
1969:
1968:
1958:
1926:
1920:
1919:
1891:
1885:
1884:
1848:
1842:
1841:
1831:
1814:(7): 3112–3128.
1799:
1790:
1789:
1779:
1761:
1729:
1720:
1719:
1709:
1692:(6): 1262–1278.
1677:
1671:
1670:
1660:
1650:
1626:
1617:
1616:
1590:
1581:
1575:
1574:
1545:
1539:
1538:
1509:
1503:
1502:
1483:10.1038/296653a0
1458:
1452:
1451:
1441:
1423:
1391:
1385:
1384:
1374:
1350:
1344:
1343:
1315:
1309:
1308:
1280:
1274:
1273:
1244:
1238:
1237:
1208:
1199:
1198:
1188:
1171:(6): 1287–1294.
1152:
1146:
1145:
1121:
1115:
1114:
1086:
1077:
1066:
1060:
1059:
1049:
1017:
1011:
1008:
996:
968:
962:
948:
942:
941:
914:
908:
907:
897:
865:
859:
858:
824:
804:
798:
797:
787:
762:(4): 2574–2587.
747:
250:studies and for
206:signaling lipids
190:receptor protein
128:DNA double helix
61:
21:
3581:
3580:
3576:
3575:
3574:
3572:
3571:
3570:
3536:
3535:
3534:
3529:
3520:
3499:
3486:Sean B. Carroll
3424:
3356:
3303:
3267:
3219:
3200:Maternal effect
3183:
3116:
3053:
3043:
3013:
3008:
2996:
2871:
2818:
2775:
2769:
2739:
2734:
2666:
2632:
2595:
2574:
2570:Lipid signaling
2542:
2514:
2461:
2386:
2377:
2333:
2328:
2289:
2288:
2284:
2240:
2239:
2235:
2191:
2190:
2186:
2142:
2141:
2137:
2093:
2092:
2088:
2044:
2043:
2039:
1977:
1976:
1972:
1928:
1927:
1923:
1893:
1892:
1888:
1850:
1849:
1845:
1801:
1800:
1793:
1731:
1730:
1723:
1679:
1678:
1674:
1628:
1627:
1620:
1588:
1583:
1582:
1578:
1547:
1546:
1542:
1511:
1510:
1506:
1460:
1459:
1455:
1393:
1392:
1388:
1352:
1351:
1347:
1317:
1316:
1312:
1282:
1281:
1277:
1246:
1245:
1241:
1210:
1209:
1202:
1160:
1154:
1153:
1149:
1142:
1123:
1122:
1118:
1111:
1088:
1087:
1080:
1067:
1063:
1019:
1018:
1014:
999:
970:
969:
965:
959:Wayback Machine
949:
945:
933:
929:
925:
921:
916:
915:
911:
867:
866:
862:
806:
805:
801:
749:
748:
744:
740:
732:SAMPL Challenge
688:
579:
570:
561:
514:receptor dimers
509:
507:Bivalent ligand
497:
489:adverse effects
477:
471:
463:inverse agonist
451:
440:
426:For the use of
374:partial agonist
366:
347:
335:
331:
326:
320:
315:
260:
53:
39:
28:
23:
22:
18:Receptor ligand
15:
12:
11:
5:
3579:
3577:
3569:
3568:
3563:
3558:
3553:
3551:Cell signaling
3548:
3538:
3537:
3531:
3530:
3525:
3522:
3521:
3519:
3518:
3513:
3507:
3505:
3501:
3500:
3498:
3497:
3496:
3495:
3483:
3478:
3473:
3468:
3463:
3462:
3461:
3450:François Jacob
3447:
3442:
3436:
3434:
3430:
3429:
3426:
3425:
3423:
3422:
3417:
3412:
3407:
3402:
3397:
3392:
3387:
3386:
3385:
3375:
3370:
3364:
3362:
3358:
3357:
3355:
3354:
3349:
3344:
3339:
3334:
3329:
3324:
3318:
3316:
3309:
3305:
3304:
3302:
3301:
3296:
3291:
3286:
3281:
3275:
3273:
3269:
3268:
3266:
3265:
3260:
3255:
3250:
3249:
3248:
3243:
3233:
3227:
3225:
3221:
3220:
3218:
3217:
3212:
3207:
3202:
3197:
3191:
3189:
3185:
3184:
3182:
3181:
3179:Sequence space
3176:
3171:
3166:
3161:
3156:
3147:
3142:
3137:
3132:
3126:
3124:
3118:
3117:
3115:
3114:
3109:
3108:
3107:
3097:
3092:
3087:
3082:
3077:
3072:
3067:
3061:
3059:
3055:
3054:
3044:
3042:
3041:
3034:
3027:
3019:
3010:
3009:
3001:
2998:
2997:
2995:
2994:
2989:
2984:
2979:
2974:
2969:
2964:
2959:
2954:
2949:
2948:
2947:
2942:
2937:
2927:
2922:
2921:
2920:
2915:
2910:
2900:
2895:
2890:
2885:
2879:
2877:
2873:
2872:
2870:
2869:
2864:
2859:
2854:
2849:
2844:
2839:
2834:
2828:
2826:
2820:
2819:
2817:
2816:
2811:
2806:
2801:
2796:
2791:
2785:
2783:
2781:Growth factors
2777:
2776:
2770:
2768:
2767:
2760:
2753:
2745:
2736:
2735:
2733:
2732:
2727:
2722:
2717:
2712:
2711:
2710:
2700:
2695:
2694:
2693:
2688:
2680:
2674:
2672:
2671:Other concepts
2668:
2667:
2665:
2664:
2659:
2650:
2644:
2642:
2638:
2637:
2634:
2633:
2631:
2630:
2625:
2620:
2615:
2609:
2607:
2601:
2600:
2597:
2596:
2594:
2593:
2588:
2582:
2580:
2576:
2575:
2573:
2572:
2567:
2562:
2556:
2550:
2544:
2543:
2541:
2540:
2535:
2530:
2524:
2522:
2516:
2515:
2513:
2512:
2507:
2505:Growth factors
2502:
2497:
2484:
2478:
2476:
2467:
2463:
2462:
2460:
2459:
2454:
2449:
2444:
2439:
2434:
2429:
2424:
2423:
2422:
2417:
2407:
2402:
2400:
2394:
2392:
2388:
2387:
2380:Cell signaling
2378:
2376:
2375:
2368:
2361:
2353:
2347:
2346:
2340:
2332:
2331:External links
2329:
2327:
2326:
2299:(1): 193–200.
2282:
2233:
2184:
2135:
2086:
2037:
1970:
1921:
1902:(4): 312–318.
1886:
1859:(6): 963–974.
1843:
1791:
1721:
1672:
1618:
1576:
1540:
1504:
1453:
1386:
1365:(3): 426–428.
1345:
1310:
1291:(7): 847–849.
1275:
1239:
1200:
1158:
1147:
1140:
1134:. p. 79.
1132:Academic Press
1116:
1109:
1078:
1061:
1012:
1010:
1009:
963:
943:
931:
927:
923:
919:
909:
860:
815:(41): 414105.
799:
741:
739:
736:
735:
734:
729:
724:
719:
714:
709:
704:
699:
694:
687:
684:
641:
640:
634:
629:
624:
618:
613:
608:
603:
598:
593:
588:
578:
575:
569:
566:
560:
557:
546:nACh receptors
508:
505:
496:
493:
473:Main article:
470:
467:
450:
447:
439:
436:
364:
345:
333:
329:
324:
318:
313:
270:residence time
259:
256:
147:hydrogen bonds
132:hydrophobicity
26:
24:
14:
13:
10:
9:
6:
4:
3:
2:
3578:
3567:
3564:
3562:
3559:
3557:
3554:
3552:
3549:
3547:
3544:
3543:
3541:
3528:
3523:
3517:
3514:
3512:
3509:
3508:
3506:
3502:
3494:
3493:
3489:
3488:
3487:
3484:
3482:
3479:
3477:
3474:
3472:
3469:
3467:
3464:
3460:
3457:
3456:
3455:
3454:Jacques Monod
3451:
3448:
3446:
3443:
3441:
3438:
3437:
3435:
3431:
3421:
3418:
3416:
3413:
3411:
3408:
3406:
3403:
3401:
3398:
3396:
3393:
3391:
3388:
3384:
3381:
3380:
3379:
3376:
3374:
3371:
3369:
3368:Homeotic gene
3366:
3365:
3363:
3359:
3353:
3350:
3348:
3345:
3343:
3340:
3338:
3335:
3333:
3330:
3328:
3325:
3323:
3320:
3319:
3317:
3313:
3310:
3306:
3300:
3297:
3295:
3292:
3290:
3287:
3285:
3282:
3280:
3277:
3276:
3274:
3270:
3264:
3261:
3259:
3256:
3254:
3251:
3247:
3244:
3242:
3239:
3238:
3237:
3236:Morphogenesis
3234:
3232:
3229:
3228:
3226:
3222:
3216:
3213:
3211:
3208:
3206:
3203:
3201:
3198:
3196:
3193:
3192:
3190:
3186:
3180:
3177:
3175:
3172:
3170:
3167:
3165:
3162:
3160:
3157:
3155:
3151:
3148:
3146:
3143:
3141:
3138:
3136:
3133:
3131:
3128:
3127:
3125:
3123:
3119:
3113:
3110:
3106:
3103:
3102:
3101:
3098:
3096:
3093:
3091:
3088:
3086:
3083:
3081:
3078:
3076:
3073:
3071:
3070:Reaction norm
3068:
3066:
3063:
3062:
3060:
3056:
3052:
3048:
3040:
3035:
3033:
3028:
3026:
3021:
3020:
3017:
3007:
3006:
2999:
2993:
2990:
2988:
2985:
2983:
2980:
2978:
2975:
2973:
2970:
2968:
2965:
2963:
2960:
2958:
2955:
2953:
2950:
2946:
2943:
2941:
2938:
2936:
2933:
2932:
2931:
2928:
2926:
2923:
2919:
2916:
2914:
2911:
2909:
2906:
2905:
2904:
2901:
2899:
2896:
2894:
2891:
2889:
2886:
2884:
2881:
2880:
2878:
2874:
2868:
2865:
2863:
2860:
2858:
2855:
2853:
2850:
2848:
2845:
2843:
2840:
2838:
2835:
2833:
2830:
2829:
2827:
2825:
2821:
2815:
2812:
2810:
2807:
2805:
2802:
2800:
2797:
2795:
2792:
2790:
2787:
2786:
2784:
2782:
2778:
2774:
2766:
2761:
2759:
2754:
2752:
2747:
2746:
2743:
2731:
2728:
2726:
2723:
2721:
2718:
2716:
2713:
2709:
2706:
2705:
2704:
2701:
2699:
2696:
2692:
2689:
2687:
2684:
2683:
2681:
2679:
2676:
2675:
2673:
2669:
2663:
2660:
2658:
2654:
2651:
2649:
2646:
2645:
2643:
2639:
2629:
2626:
2624:
2621:
2619:
2616:
2614:
2611:
2610:
2608:
2606:
2602:
2592:
2589:
2587:
2584:
2583:
2581:
2577:
2571:
2568:
2566:
2563:
2561:
2558:
2557:
2554:
2551:
2549:
2545:
2539:
2536:
2534:
2533:Intracellular
2531:
2529:
2526:
2525:
2523:
2521:
2517:
2511:
2508:
2506:
2503:
2501:
2498:
2496:
2495:Neurohormones
2492:
2491:Neuropeptides
2488:
2485:
2483:
2480:
2479:
2477:
2475:
2471:
2468:
2464:
2458:
2455:
2453:
2450:
2448:
2445:
2443:
2442:Fas apoptosis
2440:
2438:
2435:
2433:
2430:
2428:
2425:
2421:
2418:
2416:
2413:
2412:
2411:
2408:
2406:
2403:
2401:
2399:
2396:
2395:
2393:
2389:
2385:
2381:
2374:
2369:
2367:
2362:
2360:
2355:
2354:
2351:
2344:
2341:
2338:
2335:
2334:
2330:
2322:
2318:
2314:
2310:
2306:
2302:
2298:
2294:
2286:
2283:
2278:
2274:
2269:
2264:
2260:
2256:
2253:(3): 347–61.
2252:
2248:
2244:
2237:
2234:
2229:
2225:
2220:
2215:
2211:
2207:
2203:
2199:
2195:
2188:
2185:
2180:
2176:
2171:
2166:
2162:
2158:
2154:
2150:
2146:
2139:
2136:
2131:
2127:
2122:
2117:
2113:
2109:
2105:
2101:
2097:
2090:
2087:
2082:
2078:
2073:
2068:
2064:
2060:
2056:
2052:
2048:
2041:
2038:
2033:
2029:
2024:
2019:
2015:
2011:
2006:
2001:
1997:
1993:
1989:
1985:
1981:
1974:
1971:
1966:
1962:
1957:
1952:
1948:
1944:
1940:
1936:
1932:
1925:
1922:
1917:
1913:
1909:
1905:
1901:
1897:
1890:
1887:
1882:
1878:
1874:
1870:
1866:
1862:
1858:
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1320:Life Sciences
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1068:Vu-Quoc, L.,
1065:
1062:
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1048:
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1027:
1023:
1016:
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308:concentration
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97:
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82:that forms a
81:
77:
73:
69:
60:
56:
51:
47:
43:
37:
33:
19:
3546:Biomolecules
3490:
3404:
3383:eyeless gene
3279:Evolvability
3253:Segmentation
3130:Canalisation
3100:Heterochrony
3090:Heritability
3058:Key concepts
3002:
2772:
2730:Gap junction
2565:Ca signaling
2528:Cell surface
2473:
2296:
2292:
2285:
2250:
2246:
2236:
2201:
2197:
2187:
2152:
2149:Cell Reports
2148:
2138:
2103:
2099:
2089:
2054:
2050:
2040:
1987:
1983:
1973:
1938:
1934:
1924:
1899:
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1889:
1856:
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1811:
1807:
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1389:
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1323:
1319:
1313:
1288:
1284:
1278:
1253:
1249:
1242:
1217:
1213:
1168:
1165:Cell Reports
1164:
1150:
1126:
1119:
1091:
1064:
1029:
1025:
1015:
1004:
976:
972:
966:
946:
937:
912:
877:
873:
863:
812:
808:
802:
759:
755:
745:
712:Docking@Home
680:Folding@Home
657:
642:
621:Paramagnetic
580:
571:
562:
553:rule of five
550:
510:
498:
481:pharmacology
478:
452:
444:
441:
425:
400:
397:radiolabeled
394:
373:
370:full agonist
369:
362:
336:through the
301:
282:
261:
236:radioisotope
232:Radioligands
230:
187:
171:metalorganic
159:dissociation
136:
110:on a target
91:
75:
72:pharmacology
68:biochemistry
65:
3481:Mike Levine
3390:Distal-less
3215:Polyphenism
3195:Epigenetics
3047:development
2992:Wnt protein
2977:Somatomedin
2641:By distance
2579:Assistants:
2538:Co-receptor
1853:ChemMedChem
1658:2434/430357
1256:: 222–232.
1220:: 367–383.
717:GPUGRID.net
707:Ki Database
653:switchSENSE
289:antagonists
274:ion channel
143:ionic bonds
88:biomolecule
3540:Categories
3459:Lac operon
3284:Robustness
3263:Modularity
3258:Metamerism
3164:Plasticity
3159:Pleiotropy
3112:Heterotopy
2972:Semaphorin
2957:Interferon
2930:Endothelin
2708:Pheromones
2648:Juxtacrine
1032:(7): 100.
738:References
459:antagonist
218:host–guest
202:activators
198:inhibitors
194:substrates
183:hemoglobin
141:, such as
3410:Morphogen
3395:Engrailed
3378:Pax genes
3299:Tinkering
3145:Epistasis
3140:Dominance
3051:phenotype
3003:see also
2883:Adipokine
2662:Endocrine
2657:Paracrine
2653:Autocrine
2520:Receptors
2500:Cytokines
2337:BindingDB
2057:: 48–52.
822:1004.5514
727:BindingDB
540:systems (
382:nanomolar
80:substance
46:Myoglobin
3561:Proteins
3373:Hox gene
3361:Elements
3342:Homeobox
2925:Cytokine
2482:Hormones
2432:JAK-STAT
2420:MAPK/ERK
2415:TGF beta
2321:23331540
2313:19468851
2277:20303320
2228:31004156
2179:29562182
2130:31306664
2081:25239072
2032:16365317
1965:26451468
1916:19075611
1881:10561038
1873:21520422
1838:26959173
1786:23236171
1768:42553664
1716:28128928
1667:27420737
1613:16190749
1571:17676726
1535:24984935
1381:28139906
1270:24534538
1234:26164842
1195:28793254
1056:20981028
1005:Phys.org
993:20186894
955:Archived
930:, and EC
904:20695475
847:21386588
794:16085765
686:See also
664:grid.org
483:, where
304:efficacy
265:receptor
252:in vitro
214:affinity
163:covalent
96:molecule
62:
3504:Debates
3315:Systems
3241:Eyespot
3105:Neoteny
2773:ligands
2613:General
2437:Akt/PKB
2268:2908274
2219:6657754
2170:5873459
2121:6739599
2072:4259840
2023:1323165
2014:4152590
1992:Bibcode
1956:5055304
1829:5679017
1777:3535626
1746:Bibcode
1707:5679024
1499:4303982
1491:6280058
1471:Bibcode
1448:6296828
1408:Bibcode
1340:6292615
1305:7108900
1186:5586213
1034:Bibcode
934:values"
895:2933114
827:Bibcode
785:1366757
764:Bibcode
692:Agonist
455:agonist
285:agonist
244:tracers
240:in vivo
155:docking
124:protein
112:protein
104:binding
86:with a
84:complex
3405:Ligand
3085:Operon
2824:Ephrin
2466:Agents
2343:BioLiP
2319:
2311:
2275:
2265:
2226:
2216:
2177:
2167:
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2118:
2079:
2069:
2030:
2020:
2012:
1963:
1953:
1914:
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1463:Nature
1446:
1439:347328
1436:
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1338:
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1183:
1138:
1107:
1054:
991:
902:
892:
855:103345
853:
845:
792:
782:
623:probes
417:saline
380:(nM =
278:enzyme
222:steric
208:, and
167:ligand
100:signal
92:ligare
76:ligand
32:Ligand
2962:Kinin
2876:Other
2867:EFNB3
2862:EFNB2
2857:EFNB1
2852:EFNA5
2847:EFNA4
2842:EFNA3
2837:EFNA2
2832:EFNA1
2628:TFIIH
2623:TFIID
2447:Hippo
2427:Notch
2317:S2CID
2010:JSTOR
1877:S2CID
1764:JSTOR
1589:(PDF)
1495:S2CID
1430:13076
1426:JSTOR
851:S2CID
817:arXiv
485:drugs
461:, or
378:Molar
179:metal
122:, or
106:to a
78:is a
3045:The
2945:EDN3
2940:EDN2
2935:EDN1
2398:GPCR
2309:PMID
2273:PMID
2224:PMID
2175:PMID
2126:PMID
2077:PMID
2028:PMID
1961:PMID
1912:PMID
1869:PMID
1834:PMID
1782:PMID
1712:PMID
1663:PMID
1609:PMID
1567:PMID
1531:PMID
1487:PMID
1444:PMID
1377:PMID
1336:PMID
1301:PMID
1266:PMID
1230:PMID
1191:PMID
1136:ISBN
1105:ISBN
1052:PMID
989:PMID
950:See
926:, IC
900:PMID
843:PMID
790:PMID
678:and
544:and
534:GPCR
411:and
234:are
173:and
149:and
108:site
74:, a
70:and
59:1MBO
50:heme
3049:of
2410:RTK
2405:Wnt
2301:doi
2263:PMC
2255:doi
2214:PMC
2206:doi
2165:PMC
2157:doi
2116:PMC
2108:doi
2104:431
2067:PMC
2059:doi
2055:743
2018:PMC
2000:doi
1988:102
1951:PMC
1943:doi
1904:doi
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1754:doi
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1702:PMC
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1653:hdl
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