587:
173:
1338:
HK. Hybrid kinases contain multiple phosphodonor and phosphoacceptor sites and use multi-step phospho-relay schemes instead of promoting a single phosphoryl transfer. In addition to the sensor domain and kinase core, they contain a CheY-like receiver domain and a His-containing phosphotransfer (HPt)
1466:
kinases; it has been speculated that the chemical instability of phosphoaspartate is responsible, and that increased stability is needed to transduce signals in the more complex eukaryotic cell. Notably, cross-talk between signaling mechanisms is very common in eukaryotic signaling systems but rare
1115:
activity against their cognate response regulators, so that their signaling output reflects a balance between their kinase and phosphatase activities. Many response regulators also auto-dephosphorylate, and the relatively labile phosphoaspartate can also be hydrolyzed non-enzymatically. The overall
1128:
Some histidine kinases are hybrids that contain an internal receiver domain. In these cases, a hybrid HK autophosphorylates and then transfers the phosphoryl group to its own internal receiver domain, rather than to a separate RR protein. The phosphoryl group is then shuttled to
1495:
of prokaryotic two-component systems called P2CS has been compiled to document and classify known examples, and in some cases to make predictions about the cognates of "orphan" histidine kinase or response regulator proteins that are genetically unlinked to a partner.
2574:
Tomomori C, Tanaka T, Dutta R, Park H, Saha SK, Zhu Y, Ishima R, Liu D, Tong KI, Kurokawa H, Qian H, Inouye M, Ikura M (Aug 1999). "Solution structure of the homodimeric core domain of
Escherichia coli histidine kinase EnvZ".
1063:
families that are not homodimers. Response regulators may consist only of a receiver domain, but usually are multi-domain proteins with a receiver domain and at least one effector or output domain, often involved in
1196:
has been estimated as around 30, or about 1–2% of a prokaryote's genome. A few bacteria have none at all – typically endosymbionts and pathogens – and others contain over 200. All such systems must be closely
1351:; bacteria that occupy niches with frequent environmental fluctuations possess more histidine kinases and response regulators. New two-component systems may arise by
904:
795:
686:
545:
423:
301:
131:
87:
75:
1454:
It is unclear why canonical two-component systems are rare in eukaryotes, with many similar functions having been taken over by signaling systems based on
2181:"A survey of HK, HPt, and RR domains and their organization in two-component systems and phosphorelay proteins of organisms with fully sequenced genomes"
1285:
may also be present. The kinase domain is responsible for the autophosphorylation of the histidine with ATP, the phosphotransfer from the kinase to an
237:
1359:, and the relative rates of each process vary dramatically across bacterial species. In most cases, response regulator genes are located in the same
1246:. The N-terminal domain of this protein forms part of the cytoplasmic region of the protein, which may be the sensor domain responsible for sensing
634:
249:
2457:"KdpD and KdpE, proteins that control expression of the kdpABC operon, are members of the two-component sensor-effector class of regulators"
1100:
that activates the RR's effector domain, which in turn produces the cellular response to the signal, usually by stimulating (or repressing)
2408:"The kdp system of Clostridium acetobutylicum: cloning, sequencing, and transcriptional regulation in response to potassium concentration"
1262:
are the key elements in two-component signal transduction systems. Examples of histidine kinases are EnvZ, which plays a central role in
2932:
Puthiyaveetil S, Kavanagh TA, Cain P, Sullivan JA, Newell CA, Gray JC, Robinson C, van der Giezen M, Rogers MB, Allen JF (Jul 2008).
2993:"Why chloroplasts and mitochondria retain their own genomes and genetic systems: Colocation for redox regulation of gene expression"
1417:. Two-component systems are well-integrated into developmental signaling pathways in plants, but the genes probably originated from
1987:"Two-component signal transduction pathways regulating growth and cell cycle progression in a bacterium: a system-level analysis"
2883:
Mavrianos J, Berkow EL, Desai C, Pandey A, Batish M, Rabadi MJ, Barker KS, Pain D, Rogers PD, Eugenin EA, Chauhan N (Jun 2013).
2504:
Perego M, Hoch JA (Mar 1996). "Protein aspartate phosphatases control the output of two-component signal transduction systems".
1363:
as their cognate histidine kinase; lateral gene transfers are more likely to preserve operon structure than gene duplications.
1326:
C-terminal kinase core. Members of this family, however, have an integral membrane sensor domain. Not all orthodox kinases are
992:; both histidine kinases and response regulators are among the largest gene families in bacteria. They are much less common in
1218:
924:
815:
706:
565:
443:
321:
151:
3328:
1130:
2667:
Vierstra RD, Davis SJ (Dec 2000). "Bacteriophytochromes: new tools for understanding phytochrome signal transduction".
2539:
West AH, Stock AM (Jun 2001). "Histidine kinases and response regulator proteins in two-component signaling systems".
1817:
West AH, Stock AM (Jun 2001). "Histidine kinases and response regulator proteins in two-component signaling systems".
1487:
analysis. (By contrast, eukaryotic kinases are typically easily identified, but they are not easily paired with their
2363:
Buckler DR, Anand GS, Stock AM (Apr 2000). "Response-regulator phosphorylation and activation: a two-way street?".
1347:
The number of two-component systems present in a bacterial genome is highly correlated with genome size as well as
1242:
2774:"Structural classification of bacterial response regulators: diversity of output domains and domain combinations"
2702:
Alex LA, Simon MI (Apr 1994). "Protein histidine kinases and signal transduction in prokaryotes and eukaryotes".
1483:
structure, many two-component systems – particularly histidine kinases – are relatively easy to identify through
1426:
952:
and respond to changes in many different environmental conditions. Two-component systems typically consist of a
3323:
1222:
912:
803:
694:
553:
431:
309:
139:
1304:
HKs can be roughly divided into two classes: orthodox and hybrid kinases. Most orthodox HKs, typified by the
1077:
985:
1770:"Phosphorylation site of NtrC, a protein phosphatase whose covalent intermediate activates transcription"
1322:
segment(s) that separate the protein into a periplasmic N-terminal sensing domain and a highly conserved
2279:
Shi, X; Wegener-Feldbrügge, S; Huntley, S; Hamann, N; Hedderich, R; Søgaard-Andersen, L (January 2008).
1488:
1418:
1396:
1356:
1274:
1097:
1048:
961:
1133:(HPT) and subsequently to a terminal RR, which can evoke the desired response. This system is called a
908:
799:
690:
549:
427:
305:
135:
2826:"The evolution of two-component systems in bacteria reveals different strategies for niche adaptation"
242:
3063:
3004:
2945:
2837:
2281:"Bioinformatics and experimental analysis of proteins of two-component systems in Myxococcus xanthus"
1431:
647:
262:
2934:"The ancestral symbiont sensor kinase CSK links photosynthesis with gene expression in chloroplasts"
1492:
1476:
1146:
1137:. Almost 25% of bacterial HKs are of the hybrid type, as are the large majority of eukaryotic HKs.
1069:
1065:
1055:
and an ATP binding domain, though there are reported examples of histidine kinases in the atypical
1029:
1726:"Identification of the site of phosphorylation of the chemotaxis response regulator protein, CheY"
2649:
2600:
2388:
2161:
2112:
1085:
1037:
965:
1435:, derived from chloroplasts but now integrated into the nuclear genome. CSK function provides a
1403:
organelles, and are typically of the hybrid kinase phosphorelay type. For example, in the yeast
1157:
conditions. These pathways have been adapted to respond to a wide variety of stimuli, including
2455:
Walderhaug MO, Polarek JW, Voelkner P, Daniel JM, Hesse JE, Altendorf K, Epstein W (Apr 1992).
3189:
3140:
3091:
3032:
2973:
2914:
2865:
2803:
2754:
2719:
2684:
2641:
2592:
2556:
2521:
2486:
2437:
2380:
2345:
2310:
2261:
2212:
2153:
2104:
2069:
2018:
1967:
1918:
1883:
1834:
1799:
1747:
1706:
1657:
1585:
1536:
1380:
1005:
937:
899:
790:
681:
540:
418:
296:
126:
3179:
3171:
3130:
3122:
3081:
3071:
3022:
3012:
2963:
2953:
2904:
2896:
2855:
2845:
2793:
2785:
2746:
2711:
2676:
2631:
2584:
2548:
2513:
2476:
2468:
2427:
2419:
2372:
2337:
2300:
2292:
2251:
2243:
2202:
2192:
2143:
2096:
2059:
2049:
2008:
1998:
1957:
1949:
1910:
1873:
1865:
1826:
1789:
1781:
1737:
1696:
1688:
1647:
1639:
1575:
1567:
1528:
1405:
1352:
1348:
1259:
1210:
1041:
957:
2341:
891:
782:
673:
532:
410:
288:
118:
1448:
1444:
1225:
1198:
1185:
1117:
1101:
1068:. Upon detecting a particular change in the extracellular environment, the HK performs an
1033:
977:
969:
2737:
Parkinson JS, Kofoid EC (1992). "Communication modules in bacterial signaling proteins".
3067:
3008:
2949:
2841:
2750:
2328:
Laub MT, Goulian M (2007). "Specificity in two-component signal transduction pathways".
1643:
3184:
3159:
3135:
3110:
3086:
3051:
3027:
2992:
2968:
2933:
2909:
2884:
2860:
2825:
2798:
2773:
2305:
2280:
2256:
2231:
2207:
2180:
2013:
1986:
1701:
1676:
1652:
1627:
1580:
1555:
1484:
1440:
1315:
1298:
1282:
1263:
1215:
1173:
1162:
1093:
1052:
80:
2636:
2619:
2552:
2481:
2456:
2432:
2407:
2376:
2064:
2037:
1914:
1878:
1853:
1830:
1794:
1769:
1742:
1725:
3317:
2715:
2517:
2423:
2054:
1962:
1953:
1937:
1901:
Varughese KI (Apr 2002). "Molecular recognition of bacterial phosphorelay proteins".
1785:
1532:
1410:
1400:
1319:
989:
953:
639:
254:
2653:
2604:
2472:
2392:
2165:
1451:, which aims to explain the retention of genes encoded by endosymbiotic organelles.
1289:
of the response regulator, and (with bifunctional enzymes) the phosphotransfer from
853:
744:
615:
481:
359:
205:
56:
17:
2116:
1869:
1414:
887:
778:
669:
528:
406:
284:
114:
2232:"Evolution and phyletic distribution of two-component signal transduction systems"
2087:
Attwood PV, Piggott MJ, Zu XL, Besant PG (Jan 2007). "Focus on phosphohistidine".
1297:. The kinase core has a unique fold, distinct from that of the Ser/Thr/Tyr kinase
494:
372:
218:
92:
3309:
3295:
3281:
3267:
3253:
3239:
3225:
2850:
2003:
865:
756:
627:
506:
384:
230:
68:
1724:
Sanders DA, Gillece-Castro BL, Stock AM, Burlingame AL, Koshland DE (Dec 1989).
1519:
Stock AM, Robinson VL, Goudreau PN (2000). "Two-component signal transduction".
1422:
1335:
1181:
1177:
1154:
1112:
3056:
Proceedings of the
National Academy of Sciences of the United States of America
2997:
Proceedings of the
National Academy of Sciences of the United States of America
2938:
Proceedings of the
National Academy of Sciences of the United States of America
2247:
2148:
2132:"Two-component systems and their co-option for eukaryotic signal transduction"
2131:
2100:
2038:"Histidine protein kinases: key signal transducers outside the animal kingdom"
1692:
1384:
1278:
1271:
1267:
1169:
1073:
1009:
997:
1228:
proteins OmpF and OmpC. The KdpD sensor kinase proteins regulate the kdpFABC
1153:
to sense, respond, and adapt to a wide range of environments, stressors, and
3076:
3017:
2958:
1459:
1372:
1323:
1311:
1286:
1233:
1158:
1089:
1081:
1045:
3193:
3144:
3095:
3036:
2977:
2918:
2869:
2807:
2688:
2680:
2596:
2560:
2384:
2349:
2314:
2265:
2216:
2157:
2108:
2073:
2022:
1971:
1922:
1838:
1710:
1661:
1589:
1540:
1016:, two-component systems have been described as "conspicuously absent" from
3126:
2758:
2723:
2645:
2525:
2490:
2441:
1887:
1854:"Protein phosphorylation and regulation of adaptive responses in bacteria"
1803:
1751:
1571:
643:
586:
258:
172:
3305:
3291:
3277:
3263:
3249:
3235:
3221:
3175:
1463:
1331:
1327:
1290:
1150:
981:
945:
860:
751:
622:
501:
379:
225:
63:
2900:
2789:
2296:
3052:"Crosstalk and the evolution of specificity in two-component signaling"
2197:
1556:"Stimulus perception in bacterial signal-transducing histidine kinases"
1306:
993:
1768:
Sanders DA, Gillece-Castro BL, Burlingame AL, Koshland DE (Aug 1992).
1677:"Structure and function of HWE/HisKA2-family sensor histidine kinases"
1060:
1056:
3158:
Ortet P, Whitworth DE, Santaella C, Achouak W, Barakat M (Jan 2015).
1480:
1455:
1392:
1360:
1247:
1229:
1193:
1192:
and more. The average number of two-component systems in a bacterial
1017:
919:
810:
701:
560:
438:
316:
146:
2885:"Mitochondrial two-component signaling systems in Candida albicans"
2588:
1985:
Skerker JM, Prasol MS, Perchuk BS, Biondi EG, Laub MT (Oct 2005).
1436:
1388:
1376:
1294:
1165:
1013:
1001:
949:
3160:"P2CS: updates of the prokaryotic two-component systems database"
1447:; this observation has been described as a key prediction of the
968:
that mediates the cellular response, mostly through differential
3301:
3287:
3273:
3259:
3245:
3231:
3217:
1105:
973:
944:
serves as a basic stimulus-response coupling mechanism to allow
881:
848:
772:
739:
663:
610:
522:
488:
476:
400:
366:
354:
278:
212:
200:
108:
51:
1088:(RR) then catalyzes the transfer of the phosphoryl group to an
1938:"Keeping signals straight in phosphorelay signal transduction"
1395:. Two-component systems in eukaryotes likely originate from
1120:
of the response regulator ultimately controls its activity.
976:. Although two-component signaling systems are found in all
1409:, genes found in the nuclear genome likely originated from
1189:
2620:"Structure of CheA, a signal-transducing histidine kinase"
2179:
Salvado, B; Vilaprinyo, E; Sorribas, A; Alves, R (2015).
1391:, but have been described as "conspicuously absent" from
1628:"Evolution of two-component signal transduction systems"
177:
solved structure of the homodimeric domain of EnvZ from
3111:"P2CS: a database of prokaryotic two-component systems"
580:
Signal transducing histidine kinase, homodimeric domain
3300:
This article incorporates text from the public domain
3286:
This article incorporates text from the public domain
3272:
This article incorporates text from the public domain
3258:
This article incorporates text from the public domain
3244:
This article incorporates text from the public domain
3230:
This article incorporates text from the public domain
3216:
This article incorporates text from the public domain
3209:
2130:
Schaller, GE; Shiu, SH; Armitage, JP (10 May 2011).
2618:Bilwes AM, Alex LA, Crane BR, Simon MI (Jan 1999).
2230:Wuichet, K; Cantwell, BJ; Zhulin, IB (April 2010).
918:
898:
880:
875:
859:
847:
839:
834:
829:
809:
789:
771:
766:
750:
738:
730:
725:
720:
700:
680:
662:
657:
633:
621:
609:
601:
596:
591:
structure of CheA domain p4 in complex with TNP-ATP
579:
559:
539:
521:
516:
500:
487:
475:
467:
462:
457:
437:
417:
399:
394:
378:
365:
353:
345:
340:
335:
315:
295:
277:
272:
248:
236:
224:
211:
199:
191:
186:
165:
145:
125:
107:
102:
86:
74:
62:
50:
42:
37:
32:
830:Osmosensitive K channel His kinase sensor domain
27:Method of stimulus sensing and response in cells
3212:: The Prokaryotic 2-Component Systems Database
2036:Wolanin PM, Thomason PA, Stock JB (Sep 2002).
1763:
1761:
1266:, and CheA, which plays a central role in the
3109:Barakat M, Ortet P, Whitworth DE (Jan 2011).
2406:Treuner-Lange A, Kuhn A, Dürre P (Jul 1997).
1675:Herrou, J; Crosson, S; Fiebig, A (Feb 2017).
1092:residue on the response regulator's receiver
8:
2669:Seminars in Cell & Developmental Biology
1221:controls the differential expression of the
1554:Mascher T, Helmann JD, Unden G (Dec 2006).
1514:
1512:
1510:
1508:
1334:regulatory kinase NtrB (GlnL) is a soluble
1560:Microbiology and Molecular Biology Reviews
1270:system. Histidine kinases usually have an
872:
763:
654:
585:
513:
391:
269:
171:
99:
3183:
3134:
3085:
3075:
3026:
3016:
2967:
2957:
2908:
2859:
2849:
2797:
2635:
2480:
2431:
2304:
2255:
2206:
2196:
2147:
2063:
2053:
2012:
2002:
1961:
1877:
1852:Stock JB, Ninfa AJ, Stock AM (Dec 1989).
1793:
1741:
1700:
1651:
1579:
1621:
1619:
1617:
1615:
1613:
1611:
1609:
1607:
1605:
1603:
1601:
1599:
1504:
1051:containing a histidine phosphotransfer
2819:
2817:
2342:10.1146/annurev.genet.41.042007.170548
1439:-based regulatory system that couples
1111:Many HKs are bifunctional and possess
1044:(HK). Histidine kinases are typically
826:
717:
576:
454:
332:
166:His Kinase A (phospho-acceptor) domain
162:
29:
1201:to prevent cross-talk, which is rare
960:that senses a specific environmental
7:
2824:Alm E, Huang K, Arkin A (Nov 2006).
1467:in bacterial two-component systems.
2751:10.1146/annurev.ge.26.120192.000443
1730:The Journal of Biological Chemistry
1644:10.1146/annurev-micro-092611-150039
1936:Hoch JA, Varughese KI (Sep 2001).
1371:Two-component systems are rare in
25:
3050:Rowland MA, Deeds EJ (Apr 2014).
1028:Two-component systems accomplish
980:, they are most common by far in
2424:10.1128/jb.179.14.4501-4512.1997
2055:10.1186/gb-2002-3-10-reviews3013
1954:10.1128/jb.183.17.4941-4949.2001
1786:10.1128/jb.174.15.5117-5122.1992
1533:10.1146/annurev.biochem.69.1.183
1236:transport in bacteria including
2473:10.1128/jb.174.7.2152-2159.1992
2236:Current Opinion in Microbiology
1903:Current Opinion in Microbiology
1387:, and are relatively common in
942:two-component regulatory system
2541:Trends in Biochemical Sciences
1870:10.1128/MMBR.53.4.450-490.1989
1819:Trends in Biochemical Sciences
1314:membrane receptors and have a
1219:EnvZ/OmpR two-component system
1:
2637:10.1016/S0092-8674(00)80966-6
2553:10.1016/S0968-0004(01)01852-7
2377:10.1016/S0966-842X(00)01707-8
1915:10.1016/S1369-5274(02)00305-3
1831:10.1016/s0968-0004(01)01852-7
1743:10.1016/S0021-9258(20)88250-7
1632:Annual Review of Microbiology
1521:Annual Review of Biochemistry
1000:; although they do appear in
876:Available protein structures:
767:Available protein structures:
658:Available protein structures:
517:Available protein structures:
395:Available protein structures:
273:Available protein structures:
103:Available protein structures:
2851:10.1371/journal.pcbi.0020143
2716:10.1016/0168-9525(94)90215-1
2518:10.1016/0168-9525(96)81420-X
2004:10.1371/journal.pbio.0030334
1131:histidine phosphotransferase
1096:. This typically triggers a
3170:(Database issue): D536–41.
1413:and remain targeted to the
721:Histidine kinase N terminal
3345:
3299:
3285:
3271:
3257:
3243:
3229:
3215:
3121:(Database issue): D771–6.
2830:PLOS Computational Biology
1626:Capra EJ, Laub MT (2012).
1310:EnvZ protein, function as
1243:Clostridium acetobutylicum
2739:Annual Review of Genetics
2577:Nature Structural Biology
2330:Annual Review of Genetics
2248:10.1016/j.mib.2009.12.011
2149:10.1016/j.cub.2011.02.045
2101:10.1007/s00726-006-0443-6
1693:10.1016/j.mib.2017.01.008
1427:chloroplast sensor kinase
1281:kinase domain, but other
1072:reaction, transferring a
871:
762:
653:
584:
512:
390:
268:
181:by multi-dimensional NMR.
170:
98:
2772:Galperin MY (Jun 2006).
3077:10.1073/pnas.1317178111
3018:10.1073/pnas.1500012112
2959:10.1073/pnas.0803928105
2778:Journal of Bacteriology
2461:Journal of Bacteriology
2412:Journal of Bacteriology
2285:Journal of Bacteriology
1942:Journal of Bacteriology
1858:Microbiological Reviews
1774:Journal of Bacteriology
3164:Nucleic Acids Research
3115:Nucleic Acids Research
2681:10.1006/scdb.2000.0206
2365:Trends in Microbiology
1277:-binding domain and a
1078:adenosine triphosphate
1049:transmembrane proteins
964:, and a corresponding
2991:Allen JF (Aug 2015).
1681:Curr. Opin. Microbiol
1572:10.1128/MMBR.00020-06
1419:lateral gene transfer
1397:lateral gene transfer
1357:lateral gene transfer
1098:conformational change
1084:residue. The cognate
1432:Arabidopsis thaliana
1425:. An example is the
1080:(ATP) to a specific
1012:, and are common in
18:Two-component system
3329:Signal transduction
3210:http://www.p2cs.org
3127:10.1093/nar/gkq1023
3068:2014PNAS..111.5550R
3009:2015PNAS..11210231A
2950:2008PNAS..10510061P
2901:10.1128/EC.00048-13
2842:2006PLSCB...2..143A
2790:10.1128/JB.01887-05
2297:10.1128/jb.01502-07
2048:(10): REVIEWS3013.
1477:sequence similarity
1258:Signal transducing
1147:signal transduction
1070:autophosphorylation
1030:signal transduction
3176:10.1093/nar/gku968
2704:Trends in Genetics
2506:Trends in Genetics
2198:10.7717/peerj.1183
1168:state, changes in
1086:response regulator
1038:response regulator
984:, particularly in
966:response regulator
1475:Because of their
1381:filamentous fungi
1375:. They appear in
1330:bound, e.g., the
1260:histidine kinases
1254:Histidine kinases
1006:filamentous fungi
938:molecular biology
934:
933:
930:
929:
925:structure summary
825:
824:
821:
820:
816:structure summary
716:
715:
712:
711:
707:structure summary
575:
574:
571:
570:
566:structure summary
453:
452:
449:
448:
444:structure summary
331:
330:
327:
326:
322:structure summary
161:
160:
157:
156:
152:structure summary
16:(Redirected from
3336:
3198:
3197:
3187:
3155:
3149:
3148:
3138:
3106:
3100:
3099:
3089:
3079:
3047:
3041:
3040:
3030:
3020:
2988:
2982:
2981:
2971:
2961:
2929:
2923:
2922:
2912:
2880:
2874:
2873:
2863:
2853:
2821:
2812:
2811:
2801:
2769:
2763:
2762:
2734:
2728:
2727:
2699:
2693:
2692:
2664:
2658:
2657:
2639:
2615:
2609:
2608:
2571:
2565:
2564:
2536:
2530:
2529:
2501:
2495:
2494:
2484:
2452:
2446:
2445:
2435:
2403:
2397:
2396:
2360:
2354:
2353:
2325:
2319:
2318:
2308:
2276:
2270:
2269:
2259:
2227:
2221:
2220:
2210:
2200:
2176:
2170:
2169:
2151:
2127:
2121:
2120:
2084:
2078:
2077:
2067:
2057:
2033:
2027:
2026:
2016:
2006:
1982:
1976:
1975:
1965:
1933:
1927:
1926:
1898:
1892:
1891:
1881:
1849:
1843:
1842:
1814:
1808:
1807:
1797:
1765:
1756:
1755:
1745:
1721:
1715:
1714:
1704:
1672:
1666:
1665:
1655:
1623:
1594:
1593:
1583:
1551:
1545:
1544:
1516:
1406:Candida albicans
1353:gene duplication
1349:ecological niche
1232:responsible for
1211:Escherichia coli
1186:chemoattractants
1042:histidine kinase
958:histidine kinase
873:
827:
764:
718:
655:
589:
577:
514:
458:Histidine kinase
455:
392:
336:Histidine kinase
333:
270:
179:Escherichia coli
175:
163:
100:
33:Histidine kinase
30:
21:
3344:
3343:
3339:
3338:
3337:
3335:
3334:
3333:
3324:Protein domains
3314:
3313:
3312:
3298:
3284:
3270:
3256:
3242:
3228:
3206:
3201:
3157:
3156:
3152:
3108:
3107:
3103:
3049:
3048:
3044:
3003:(33): 10231–8.
2990:
2989:
2985:
2944:(29): 10061–6.
2931:
2930:
2926:
2889:Eukaryotic Cell
2882:
2881:
2877:
2823:
2822:
2815:
2784:(12): 4169–82.
2771:
2770:
2766:
2736:
2735:
2731:
2701:
2700:
2696:
2666:
2665:
2661:
2617:
2616:
2612:
2573:
2572:
2568:
2538:
2537:
2533:
2503:
2502:
2498:
2454:
2453:
2449:
2418:(14): 4501–12.
2405:
2404:
2400:
2362:
2361:
2357:
2327:
2326:
2322:
2278:
2277:
2273:
2229:
2228:
2224:
2178:
2177:
2173:
2136:Current Biology
2129:
2128:
2124:
2086:
2085:
2081:
2035:
2034:
2030:
1984:
1983:
1979:
1935:
1934:
1930:
1900:
1899:
1895:
1851:
1850:
1846:
1816:
1815:
1811:
1780:(15): 5117–22.
1767:
1766:
1759:
1736:(36): 21770–8.
1723:
1722:
1718:
1674:
1673:
1669:
1625:
1624:
1597:
1553:
1552:
1548:
1518:
1517:
1506:
1502:
1473:
1449:CoRR hypothesis
1445:gene expression
1443:to chloroplast
1369:
1345:
1256:
1149:systems enable
1143:
1126:
1118:phosphorylation
1034:phosphorylation
1026:
978:domains of life
592:
182:
76:OPM superfamily
28:
23:
22:
15:
12:
11:
5:
3342:
3340:
3332:
3331:
3326:
3316:
3315:
3214:
3213:
3205:
3204:External links
3202:
3200:
3199:
3150:
3101:
3062:(15): 5550–5.
3042:
2983:
2924:
2875:
2813:
2764:
2729:
2694:
2659:
2610:
2566:
2531:
2496:
2447:
2398:
2355:
2320:
2271:
2222:
2171:
2142:(9): R320–30.
2122:
2079:
2042:Genome Biology
2028:
1977:
1948:(17): 4941–9.
1928:
1893:
1844:
1809:
1757:
1716:
1667:
1595:
1546:
1527:(1): 183–215.
1503:
1501:
1498:
1485:bioinformatics
1472:
1471:Bioinformatics
1469:
1441:photosynthesis
1429:(CSK) gene in
1368:
1365:
1344:
1341:
1316:signal peptide
1264:osmoregulation
1255:
1252:
1223:outer membrane
1216:osmoregulatory
1174:quorum signals
1145:Two-component
1142:
1139:
1125:
1122:
1025:
1022:
932:
931:
928:
927:
922:
916:
915:
902:
896:
895:
885:
878:
877:
869:
868:
863:
857:
856:
851:
845:
844:
841:
837:
836:
832:
831:
823:
822:
819:
818:
813:
807:
806:
793:
787:
786:
776:
769:
768:
760:
759:
754:
748:
747:
742:
736:
735:
732:
728:
727:
723:
722:
714:
713:
710:
709:
704:
698:
697:
684:
678:
677:
667:
660:
659:
651:
650:
637:
631:
630:
625:
619:
618:
613:
607:
606:
603:
599:
598:
594:
593:
590:
582:
581:
573:
572:
569:
568:
563:
557:
556:
543:
537:
536:
526:
519:
518:
510:
509:
504:
498:
497:
492:
485:
484:
479:
473:
472:
469:
465:
464:
460:
459:
451:
450:
447:
446:
441:
435:
434:
421:
415:
414:
404:
397:
396:
388:
387:
382:
376:
375:
370:
363:
362:
357:
351:
350:
347:
343:
342:
338:
337:
329:
328:
325:
324:
319:
313:
312:
299:
293:
292:
282:
275:
274:
266:
265:
252:
246:
245:
240:
234:
233:
228:
222:
221:
216:
209:
208:
203:
197:
196:
193:
189:
188:
184:
183:
176:
168:
167:
159:
158:
155:
154:
149:
143:
142:
129:
123:
122:
112:
105:
104:
96:
95:
90:
84:
83:
78:
72:
71:
66:
60:
59:
54:
48:
47:
44:
40:
39:
35:
34:
26:
24:
14:
13:
10:
9:
6:
4:
3:
2:
3341:
3330:
3327:
3325:
3322:
3321:
3319:
3311:
3307:
3303:
3297:
3293:
3289:
3283:
3279:
3275:
3269:
3265:
3261:
3255:
3251:
3247:
3241:
3237:
3233:
3227:
3223:
3219:
3211:
3208:
3207:
3203:
3195:
3191:
3186:
3181:
3177:
3173:
3169:
3165:
3161:
3154:
3151:
3146:
3142:
3137:
3132:
3128:
3124:
3120:
3116:
3112:
3105:
3102:
3097:
3093:
3088:
3083:
3078:
3073:
3069:
3065:
3061:
3057:
3053:
3046:
3043:
3038:
3034:
3029:
3024:
3019:
3014:
3010:
3006:
3002:
2998:
2994:
2987:
2984:
2979:
2975:
2970:
2965:
2960:
2955:
2951:
2947:
2943:
2939:
2935:
2928:
2925:
2920:
2916:
2911:
2906:
2902:
2898:
2895:(6): 913–22.
2894:
2890:
2886:
2879:
2876:
2871:
2867:
2862:
2857:
2852:
2847:
2843:
2839:
2835:
2831:
2827:
2820:
2818:
2814:
2809:
2805:
2800:
2795:
2791:
2787:
2783:
2779:
2775:
2768:
2765:
2760:
2756:
2752:
2748:
2744:
2740:
2733:
2730:
2725:
2721:
2717:
2713:
2709:
2705:
2698:
2695:
2690:
2686:
2682:
2678:
2675:(6): 511–21.
2674:
2670:
2663:
2660:
2655:
2651:
2647:
2643:
2638:
2633:
2630:(1): 131–41.
2629:
2625:
2621:
2614:
2611:
2606:
2602:
2598:
2594:
2590:
2589:10.1038/11495
2586:
2583:(8): 729–34.
2582:
2578:
2570:
2567:
2562:
2558:
2554:
2550:
2547:(6): 369–76.
2546:
2542:
2535:
2532:
2527:
2523:
2519:
2515:
2512:(3): 97–101.
2511:
2507:
2500:
2497:
2492:
2488:
2483:
2478:
2474:
2470:
2467:(7): 2152–9.
2466:
2462:
2458:
2451:
2448:
2443:
2439:
2434:
2429:
2425:
2421:
2417:
2413:
2409:
2402:
2399:
2394:
2390:
2386:
2382:
2378:
2374:
2370:
2366:
2359:
2356:
2351:
2347:
2343:
2339:
2335:
2331:
2324:
2321:
2316:
2312:
2307:
2302:
2298:
2294:
2291:(2): 613–24.
2290:
2286:
2282:
2275:
2272:
2267:
2263:
2258:
2253:
2249:
2245:
2242:(2): 219–25.
2241:
2237:
2233:
2226:
2223:
2218:
2214:
2209:
2204:
2199:
2194:
2190:
2186:
2182:
2175:
2172:
2167:
2163:
2159:
2155:
2150:
2145:
2141:
2137:
2133:
2126:
2123:
2118:
2114:
2110:
2106:
2102:
2098:
2095:(1): 145–56.
2094:
2090:
2083:
2080:
2075:
2071:
2066:
2061:
2056:
2051:
2047:
2043:
2039:
2032:
2029:
2024:
2020:
2015:
2010:
2005:
2000:
1996:
1992:
1988:
1981:
1978:
1973:
1969:
1964:
1959:
1955:
1951:
1947:
1943:
1939:
1932:
1929:
1924:
1920:
1916:
1912:
1908:
1904:
1897:
1894:
1889:
1885:
1880:
1875:
1871:
1867:
1864:(4): 450–90.
1863:
1859:
1855:
1848:
1845:
1840:
1836:
1832:
1828:
1825:(6): 369–76.
1824:
1820:
1813:
1810:
1805:
1801:
1796:
1791:
1787:
1783:
1779:
1775:
1771:
1764:
1762:
1758:
1753:
1749:
1744:
1739:
1735:
1731:
1727:
1720:
1717:
1712:
1708:
1703:
1698:
1694:
1690:
1686:
1682:
1678:
1671:
1668:
1663:
1659:
1654:
1649:
1645:
1641:
1637:
1633:
1629:
1622:
1620:
1618:
1616:
1614:
1612:
1610:
1608:
1606:
1604:
1602:
1600:
1596:
1591:
1587:
1582:
1577:
1573:
1569:
1566:(4): 910–38.
1565:
1561:
1557:
1550:
1547:
1542:
1538:
1534:
1530:
1526:
1522:
1515:
1513:
1511:
1509:
1505:
1499:
1497:
1494:
1490:
1486:
1482:
1478:
1470:
1468:
1465:
1461:
1457:
1452:
1450:
1446:
1442:
1438:
1434:
1433:
1428:
1424:
1420:
1416:
1412:
1411:endosymbiosis
1408:
1407:
1402:
1401:endosymbiotic
1399:, often from
1398:
1394:
1390:
1386:
1382:
1378:
1374:
1367:In eukaryotes
1366:
1364:
1362:
1358:
1354:
1350:
1342:
1340:
1337:
1333:
1329:
1325:
1321:
1320:transmembrane
1317:
1313:
1309:
1308:
1302:
1300:
1296:
1293:phosphate to
1292:
1288:
1284:
1280:
1276:
1273:
1269:
1265:
1261:
1253:
1251:
1249:
1245:
1244:
1239:
1235:
1231:
1227:
1224:
1220:
1217:
1213:
1212:
1206:
1204:
1200:
1195:
1191:
1187:
1183:
1179:
1175:
1171:
1167:
1164:
1160:
1156:
1152:
1148:
1140:
1138:
1136:
1132:
1124:Phosphorelays
1123:
1121:
1119:
1114:
1109:
1107:
1103:
1099:
1095:
1091:
1087:
1083:
1079:
1075:
1071:
1067:
1062:
1058:
1054:
1050:
1047:
1043:
1039:
1035:
1031:
1023:
1021:
1019:
1015:
1011:
1007:
1003:
999:
995:
991:
990:cyanobacteria
987:
986:Gram-negative
983:
979:
975:
971:
967:
963:
959:
955:
951:
947:
943:
939:
926:
923:
921:
917:
914:
910:
906:
903:
901:
897:
893:
889:
886:
883:
879:
874:
870:
867:
864:
862:
858:
855:
852:
850:
846:
842:
838:
833:
828:
817:
814:
812:
808:
805:
801:
797:
794:
792:
788:
784:
780:
777:
774:
770:
765:
761:
758:
755:
753:
749:
746:
743:
741:
737:
733:
729:
724:
719:
708:
705:
703:
699:
696:
692:
688:
685:
683:
679:
675:
671:
668:
665:
661:
656:
652:
649:
645:
641:
638:
636:
632:
629:
626:
624:
620:
617:
614:
612:
608:
604:
600:
595:
588:
583:
578:
567:
564:
562:
558:
555:
551:
547:
544:
542:
538:
534:
530:
527:
524:
520:
515:
511:
508:
505:
503:
499:
496:
493:
490:
486:
483:
480:
478:
474:
470:
466:
461:
456:
445:
442:
440:
436:
433:
429:
425:
422:
420:
416:
412:
408:
405:
402:
398:
393:
389:
386:
383:
381:
377:
374:
371:
368:
364:
361:
358:
356:
352:
348:
344:
339:
334:
323:
320:
318:
314:
311:
307:
303:
300:
298:
294:
290:
286:
283:
280:
276:
271:
267:
264:
260:
256:
253:
251:
247:
244:
241:
239:
235:
232:
229:
227:
223:
220:
217:
214:
210:
207:
204:
202:
198:
194:
190:
185:
180:
174:
169:
164:
153:
150:
148:
144:
141:
137:
133:
130:
128:
124:
120:
116:
113:
110:
106:
101:
97:
94:
91:
89:
85:
82:
79:
77:
73:
70:
67:
65:
61:
58:
55:
53:
49:
45:
41:
36:
31:
19:
3167:
3163:
3153:
3118:
3114:
3104:
3059:
3055:
3045:
3000:
2996:
2986:
2941:
2937:
2927:
2892:
2888:
2878:
2836:(11): e143.
2833:
2829:
2781:
2777:
2767:
2742:
2738:
2732:
2710:(4): 133–8.
2707:
2703:
2697:
2672:
2668:
2662:
2627:
2623:
2613:
2580:
2576:
2569:
2544:
2540:
2534:
2509:
2505:
2499:
2464:
2460:
2450:
2415:
2411:
2401:
2371:(4): 153–6.
2368:
2364:
2358:
2333:
2329:
2323:
2288:
2284:
2274:
2239:
2235:
2225:
2188:
2184:
2174:
2139:
2135:
2125:
2092:
2088:
2082:
2045:
2041:
2031:
1997:(10): e334.
1994:
1991:PLOS Biology
1990:
1980:
1945:
1941:
1931:
1909:(2): 142–8.
1906:
1902:
1896:
1861:
1857:
1847:
1822:
1818:
1812:
1777:
1773:
1733:
1729:
1719:
1684:
1680:
1670:
1635:
1631:
1563:
1559:
1549:
1524:
1520:
1474:
1453:
1430:
1423:chloroplasts
1415:mitochondria
1404:
1370:
1346:
1305:
1303:
1257:
1241:
1237:
1209:
1207:
1202:
1144:
1135:phosphorelay
1134:
1127:
1110:
1032:through the
1027:
941:
935:
605:H-kinase_dim
178:
2089:Amino Acids
1385:slime molds
1336:cytoplasmic
1324:cytoplasmic
1312:periplasmic
1299:superfamily
1182:temperature
1178:antibiotics
1113:phosphatase
1076:group from
1066:DNA binding
1046:homodimeric
1010:slime molds
835:Identifiers
726:Identifiers
597:Identifiers
463:Identifiers
341:Identifiers
187:Identifiers
88:OPM protein
38:Identifiers
3318:Categories
2745:: 71–112.
2336:: 121–45.
1638:: 325–47.
1500:References
1489:substrates
1373:eukaryotes
1279:C-terminal
1272:N-terminal
1268:chemotaxis
1250:pressure.
1170:osmolarity
1104:of target
1102:expression
1074:phosphoryl
1040:(RR) by a
998:eukaryotes
972:of target
970:expression
888:structures
779:structures
670:structures
529:structures
407:structures
285:structures
115:structures
46:His_kinase
3310:IPR003852
3296:IPR011126
3282:IPR004105
3268:IPR011495
3254:IPR003661
3240:IPR010559
3226:IPR011712
2191:: e1183.
1687:: 47–54.
1460:threonine
1343:Evolution
1287:aspartate
1234:potassium
1199:regulated
1159:nutrients
1116:level of
1090:aspartate
1082:histidine
1024:Mechanism
946:organisms
866:IPR003852
757:IPR018984
628:IPR004105
507:IPR011712
385:IPR011495
231:IPR003661
69:IPR016380
3306:InterPro
3292:InterPro
3278:InterPro
3264:InterPro
3250:InterPro
3236:InterPro
3222:InterPro
3194:25324303
3145:21051349
3096:24706803
3037:26286985
2978:18632566
2919:23584995
2870:17083272
2808:16740923
2689:11145881
2654:16842653
2605:23334643
2597:10426948
2561:11406410
2393:39589537
2385:10754569
2350:18076326
2315:17993514
2266:20133179
2217:26339559
2166:18423129
2158:21549954
2109:17103118
2074:12372152
2023:16176121
1972:11489844
1923:11934609
1839:11406410
1711:28193573
1662:22746333
1590:17158704
1541:10966457
1493:database
1464:tyrosine
1339:domain.
1332:nitrogen
1328:membrane
1291:aspartyl
1163:cellular
1151:bacteria
1141:Function
982:bacteria
962:stimulus
954:membrane
905:RCSB PDB
861:InterPro
796:RCSB PDB
752:InterPro
687:RCSB PDB
623:InterPro
546:RCSB PDB
502:InterPro
424:RCSB PDB
380:InterPro
302:RCSB PDB
226:InterPro
132:RCSB PDB
64:InterPro
3185:4384028
3136:3013651
3087:3992699
3064:Bibcode
3028:4547249
3005:Bibcode
2969:2474565
2946:Bibcode
2910:3675996
2861:1630713
2838:Bibcode
2799:1482966
2759:1482126
2724:8029829
2646:9989504
2526:8868347
2491:1532388
2442:9226259
2306:2223698
2257:3391504
2208:4558063
2117:6912202
2014:1233412
1888:2556636
1804:1321122
1752:2689446
1702:5534388
1653:4097194
1581:1698512
1393:animals
1307:E. coli
1283:domains
1238:E. coli
1203:in vivo
1018:animals
994:archaea
956:-bound
854:PF02702
745:PF09385
616:PF02895
482:PF07730
471:HisKA_3
360:PF07568
349:HisKA_2
206:PF00512
57:PF06580
3192:
3182:
3143:
3133:
3094:
3084:
3035:
3025:
2976:
2966:
2917:
2907:
2868:
2858:
2806:
2796:
2757:
2722:
2687:
2652:
2644:
2603:
2595:
2559:
2524:
2489:
2482:205833
2479:
2440:
2433:179285
2430:
2391:
2383:
2348:
2313:
2303:
2264:
2254:
2215:
2205:
2164:
2156:
2115:
2107:
2072:
2065:244915
2062:
2021:
2011:
1970:
1960:
1921:
1886:
1879:372749
1876:
1837:
1802:
1795:206329
1792:
1750:
1709:
1699:
1660:
1650:
1588:
1578:
1539:
1481:operon
1456:serine
1389:plants
1383:, and
1377:yeasts
1361:operon
1355:or by
1275:ligand
1248:turgor
1230:operon
1214:, the
1194:genome
1155:growth
1094:domain
1061:HisKA2
1053:domain
1014:plants
1008:, and
1002:yeasts
920:PDBsum
894:
884:
840:Symbol
811:PDBsum
785:
775:
734:HisK_N
731:Symbol
702:PDBsum
676:
666:
648:SUPFAM
602:Symbol
561:PDBsum
535:
525:
495:CL0025
468:Symbol
439:PDBsum
413:
403:
373:CL0025
346:Symbol
317:PDBsum
291:
281:
263:SUPFAM
219:CL0025
192:Symbol
147:PDBsum
121:
111:
43:Symbol
2650:S2CID
2601:S2CID
2389:S2CID
2185:PeerJ
2162:S2CID
2113:S2CID
1963:95367
1491:.) A
1462:, or
1437:redox
1421:from
1295:water
1226:porin
1166:redox
1106:genes
1036:of a
974:genes
950:sense
644:SCOPe
635:SCOP2
259:SCOPe
250:SCOP2
243:HisKA
238:SMART
195:HisKA
3304:and
3302:Pfam
3290:and
3288:Pfam
3276:and
3274:Pfam
3262:and
3260:Pfam
3248:and
3246:Pfam
3234:and
3232:Pfam
3220:and
3218:Pfam
3190:PMID
3141:PMID
3092:PMID
3033:PMID
2974:PMID
2915:PMID
2866:PMID
2804:PMID
2755:PMID
2720:PMID
2685:PMID
2642:PMID
2624:Cell
2593:PMID
2557:PMID
2522:PMID
2487:PMID
2438:PMID
2381:PMID
2346:PMID
2311:PMID
2262:PMID
2213:PMID
2154:PMID
2105:PMID
2070:PMID
2019:PMID
1968:PMID
1919:PMID
1884:PMID
1835:PMID
1800:PMID
1748:PMID
1707:PMID
1658:PMID
1586:PMID
1537:PMID
1479:and
1318:and
1240:and
1059:and
996:and
988:and
940:, a
913:PDBj
909:PDBe
892:ECOD
882:Pfam
849:Pfam
843:KdpD
804:PDBj
800:PDBe
783:ECOD
773:Pfam
740:Pfam
695:PDBj
691:PDBe
674:ECOD
664:Pfam
640:1b3q
611:Pfam
554:PDBj
550:PDBe
533:ECOD
523:Pfam
491:clan
489:Pfam
477:Pfam
432:PDBj
428:PDBe
411:ECOD
401:Pfam
369:clan
367:Pfam
355:Pfam
310:PDBj
306:PDBe
289:ECOD
279:Pfam
255:1b3q
215:clan
213:Pfam
201:Pfam
140:PDBj
136:PDBe
119:ECOD
109:Pfam
93:5iji
52:Pfam
3180:PMC
3172:doi
3131:PMC
3123:doi
3082:PMC
3072:doi
3060:111
3023:PMC
3013:doi
3001:112
2964:PMC
2954:doi
2942:105
2905:PMC
2897:doi
2856:PMC
2846:doi
2794:PMC
2786:doi
2782:188
2747:doi
2712:doi
2677:doi
2632:doi
2585:doi
2549:doi
2514:doi
2477:PMC
2469:doi
2465:174
2428:PMC
2420:doi
2416:179
2373:doi
2338:doi
2301:PMC
2293:doi
2289:190
2252:PMC
2244:doi
2203:PMC
2193:doi
2144:doi
2097:doi
2060:PMC
2050:doi
2009:PMC
1999:doi
1958:PMC
1950:doi
1946:183
1911:doi
1874:PMC
1866:doi
1827:doi
1790:PMC
1782:doi
1778:174
1738:doi
1734:264
1697:PMC
1689:doi
1648:PMC
1640:doi
1576:PMC
1568:doi
1529:doi
1208:In
1057:HWE
948:to
936:In
900:PDB
791:PDB
682:PDB
541:PDB
419:PDB
297:PDB
127:PDB
81:281
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