657:
757:(OEC) within one PSII was provided by Pierre Joliot et al. They have shown that, if dark-adapted photosynthetic material (higher plants, algae, and cyanobacteria) is exposed to a series of single turnover flashes, oxygen evolution is detected with typical period-four damped oscillation with maxima on the third and the seventh flash and with minima on the first and the fifth flash (for review, see). Based on this experiment, Bessel Kok and co-workers introduced a cycle of five flash-induced transitions of the so-called
168:
734:
160:
35:
179:
in which the positive charge sitting on the chlorophyll dimer that undergoes the initial photoinduced charge separation is equally shared by the two monomers, in intact PSII the charge is mostly localized on one chlorophyll center (70−80%). Because of this, P680 is highly oxidizing and can take part
670:
The oxygen-evolving complex is the site of water oxidation. It is a metallo-oxo cluster comprising four manganese ions (in oxidation states ranging from +3 to +4) and one divalent calcium ion. When it oxidizes water, producing oxygen gas and protons, it sequentially delivers the four electrons from
686:. While obtaining these structures was in itself a great feat, they did not show the oxygen-evolving complex in full detail. In 2011 the OEC of PSII was resolved to a level of 1.9Å revealing five oxygen atoms serving as oxo bridges linking the five metal atoms and four water molecules bound to the
756:
The mechanism of water oxidation is understood in substantial detail. The oxidation of water to molecular oxygen requires extraction of four electrons and four protons from two molecules of water. The experimental evidence that oxygen is released through cyclic reaction of oxygen evolving complex
724:
in the USA. In 2014 the structure observed in 2011 was confirmed. Knowing the structure of
Photosystem II did not suffice to reveal how it works exactly. So now the race has started to solve the structure of Photosystem II at different stages in the mechanistic cycle (discussed below). Currently
725:
structures of the S1 state and the S3 state's have been published almost simultaneously from two different groups, showing the addition of an oxygen molecule designated O6 between Mn1 and Mn4, suggesting that this may be the site on the oxygen evolving complex, where oxygen is produced.
707:
cluster; more than 1,300 water molecules were found in each photosystem II monomer, some forming extensive hydrogen-bonding networks that may serve as channels for protons, water or oxygen molecules. At this stage, it is suggested that the structures obtained by
1884:
Kuntzleman T, Yocum CF (February 2005). "Reduction-induced inhibition and Mn(II) release from the photosystem II oxygen-evolving complex by hydroquinone or NH2OH are consistent with a Mn(III)/Mn(III)/Mn(IV)/Mn(IV) oxidation state for the dark-adapted enzyme".
999:
Okubo T, Tomo T, Sugiura M, Noguchi T (April 2007). "Perturbation of the structure of P680 and the charge distribution on its radical cation in isolated reaction center complexes of photosystem II as revealed by fourier transform infrared spectroscopy".
753:) is one of the most important reactions on the planet, since it is the source of nearly all the atmosphere's oxygen. Moreover, artificial photosynthetic water-splitting may contribute to the effective use of sunlight as an alternative energy-source.
187:
and green plants) is composed of around 20 subunits (depending on the organism) as well as other accessory, light-harvesting proteins. Each photosystem II contains at least 99 cofactors: 35 chlorophyll a, 12
671:
water to a tyrosine (D1-Y161) sidechain and then to P680 itself. It is composed of three protein subunits, OEE1 (PsbO), OEE2 (PsbP) and OEE3 (PsbQ); a fourth PsbR peptide is associated nearby.
1344:
Guskov A, Kern J, Gabdulkhakov A, Broser M, Zouni A, Saenger W (March 2009). "Cyanobacterial photosystem II at 2.9-A resolution and the role of quinones, lipids, channels and chloride".
1038:
Guskov A, Kern J, Gabdulkhakov A, Broser M, Zouni A, Saenger W (March 2009). "Cyanobacterial photosystem II at 2.9-A resolution and the role of quinones, lipids, channels and chloride".
357:
1185:
Zouni A, Witt HT, Kern J, Fromme P, Krauss N, Saenger W, Orth P (February 2001). "Crystal structure of photosystem II from
Synechococcus elongatus at 3.8 A resolution".
615:
2116:
893:
Loll B, Kern J, Saenger W, Zouni A, Biesiadka J (December 2005). "Towards complete cofactor arrangement in the 3.0 A resolution structure of photosystem II".
116:, photosystem II provides the electrons for all of photosynthesis to occur. The hydrogen ions (protons) generated by the oxidation of water help to create a
2657:
151:
is a chemical often used in laboratory settings to inhibit photosynthesis. When present, DCMU inhibits electron flow from photosystem II to plastoquinone.
469:
745:) shows the link between the electron transport (S-states advancement) and the relaxation process of the intermediate S-states (, n=0,1,2,3) formation
2549:
2160:
2109:
634:
316:
2142:
788:
In 2012, Renger expressed the idea of internal changes of water molecules into typical oxides in different S-states during water splitting.
175:
The core of PSII consists of a pseudo-symmetric heterodimer of two homologous proteins D1 and D2. Unlike the reaction centers of all other
716:
used, altering the observed OEC structure. This incentivized researchers to take their crystals to a different X-ray facilities, called
2178:
2201:
2150:
2102:
1841:
Kok B, Forbush B, McGloin M (June 1970). "Cooperation of charges in photosynthetic O2 evolution-I. A linear four step mechanism".
1388:
2168:
864:
627:
337:
1445:
Suga M, Akita F, Hirata K, Ueno G, Murakami H, Nakajima Y, Shimizu T, Yamashita K, Yamamoto M, Ago H, Shen JR (January 2015).
1102:
594:
721:
2742:
2542:
2635:
284:
280:
276:
272:
268:
264:
260:
256:
252:
2258:
2083:
2018:
2700:
1612:
Vinyard DJ, Brudvig GW (May 2017). "Progress Toward a
Molecular Mechanism of Water Oxidation in Photosystem II".
588:
2705:
2677:
2650:
2628:
2613:
2578:
2535:
781:
state is largely considered to consist of manganese ions with oxidation states of Mn, Mn, Mn, Mn. Finally, the
570:
24:
2414:
575:
2623:
2042:
1657:"Current Understanding of the Mechanism of Water Oxidation in Photosystem II and Its Relation to XFEL Data"
1230:"Crystal structure of oxygen-evolving photosystem II from Thermosynechococcus vulcanus at 3.7-A resolution"
1138:"Current Understanding of the Mechanism of Water Oxidation in Photosystem II and Its Relation to XFEL Data"
1979:"Mechanism of light induced water splitting in Photosystem II of oxygen evolving photosynthetic organisms"
1922:"Evidence for intermediate S-states as initial phase in the process of oxygen-evolving complex oxidation"
1763:
Joliot P.; Barbieri G.; Chabaud R. (1969). "Un nouveau modele des centres photochimiques du systeme II".
785:
were proposed by
Jablonsky and Lazar as a regulatory mechanism and link between S-states and tyrosine Z.
639:
321:
Reaction center protein, binds
Chlorophyll P680, pheophytin, beta-carotene, quinone and manganese center
563:
2805:
1083:
X-Ray
Diffraction and Spectroscopy of Photosystem II at Room Temperature Using Femtosecond X-Ray Pulses
2515:
2502:
2489:
2476:
2463:
2450:
2437:
2399:
1798:
Joliot P (2003). "Period-four oscillations of the flash-induced oxygen formation in photosynthesis".
761:, describing the four redox states of OEC: When four oxidizing equivalents have been stored (at the S
717:
144:
54:
2800:
2727:
2409:
2363:
2306:
2133:
656:
591:
493:
515:
2795:
2759:
2737:
2717:
2311:
665:
94:
1081:
Junko Y, Kern J, Yachandra VK, Nilsson H, Koroidov S, Messinger J (2015). "Chapter 2, Section 3
2722:
2618:
2209:
1696:
Ibrahim M, Fransson T, Chatterjee R, Cheah MH, Hussein R, Lassalle L, et al. (June 2020).
712:
are biased, since there is evidence that the manganese atoms are reduced by the high-intensity
125:
2332:
2251:
2094:
1500:
Young ID, Ibrahim M, Chatterjee R, Gul S, Fuller F, Koroidov S, et al. (December 2016).
709:
675:
1559:"Light-induced structural changes and the site of O=O bond formation in PSII caught by XFEL"
1447:"Native structure of photosystem II at 1.95 Å resolution viewed by femtosecond X-ray pulses"
2404:
1933:
1717:
1633:
1621:
1570:
1513:
1458:
1403:
1300:
1241:
902:
551:
28:
1673:
1656:
1154:
1137:
8:
2774:
2764:
2732:
2368:
2225:
527:
1937:
1721:
1625:
1574:
1517:
1462:
1407:
1304:
1245:
1087:
Sustaining Life on Planet Earth: Metalloenzymes
Mastering Dioxygen and Other Chewy Gases
956:
Philosophical
Transactions of the Royal Society of London. Series B, Biological Sciences
906:
498:
378:
By convention, gene names are formed by Psb + subunit letter. For example, subunit O is
2810:
2769:
2301:
2075:
1954:
1921:
1866:
1854:
1823:
1780:
1776:
1740:
1697:
1594:
1534:
1501:
1482:
1427:
1369:
1326:
1210:
1167:
1113:
1063:
976:
951:
926:
682:
in 2001. Over the next years the resolution of the model was gradually increased to 2.9
62:
1264:
1229:
128:. The energized electrons transferred to plastoquinone are ultimately used to reduce
2079:
2000:
1959:
1902:
1858:
1815:
1745:
1678:
1637:
1598:
1586:
1557:
Suga M, Akita F, Sugahara M, Kubo M, Nakajima Y, Nakane T, et al. (March 2017).
1539:
1486:
1474:
1431:
1419:
1361:
1318:
1269:
1202:
1171:
1159:
1118:
1098:
1055:
1017:
981:
918:
804:
582:
248:
113:
1870:
1784:
1373:
1330:
1067:
2347:
2342:
2316:
2244:
2186:
2071:
1990:
1949:
1941:
1894:
1850:
1827:
1807:
1772:
1735:
1725:
1668:
1629:
1578:
1529:
1521:
1466:
1411:
1353:
1308:
1259:
1249:
1214:
1194:
1149:
1108:
1090:
1047:
1009:
971:
963:
930:
910:
839:
796:
750:
2562:
2527:
2394:
2378:
2291:
1995:
1978:
947:
874:
869:
441:
366:
117:
50:
1945:
1706:
transition in photosystem II and implications for the water oxidation mechanism"
1094:
167:
2710:
2570:
2432:
2373:
2125:
1710:
Proceedings of the
National Academy of Sciences of the United States of America
1234:
Proceedings of the
National Academy of Sciences of the United States of America
849:
800:
610:
247:
ions per monomer. There are several crystal structures of photosystem II. The
58:
1811:
1389:"Crystal structure of oxygen-evolving photosystem II at a resolution of 1.9 Å"
2789:
2605:
2588:
2337:
2296:
859:
737:
Water-splitting process: Electron transport and regulation. The first level (
683:
679:
459:
424:
197:
189:
184:
106:
98:
74:
20:
16:
First protein complex in light-dependent reactions of oxygenic photosynthesis
1730:
1313:
1288:
1254:
2286:
2004:
1963:
1906:
1819:
1749:
1682:
1641:
1590:
1543:
1478:
1423:
1365:
1322:
1273:
1206:
1163:
1122:
1059:
1021:
985:
967:
922:
823:
674:
The first structural model of the oxygen-evolving complex was solved using
121:
1862:
799:
of PSII are used as herbicides. There are two main chemical families, the
741:) shows the original Kok model of the S-states cycling, the second level (
733:
2583:
2510:
2445:
2281:
2129:
1558:
1446:
854:
414:
176:
1582:
1525:
1470:
1415:
914:
539:
2695:
2672:
1502:"Structure of photosystem II and substrate binding at room temperature"
1089:. Metal Ions in Life Sciences. Vol. 15. Springer. pp. 13–43.
449:
193:
1898:
1357:
1287:
Ferreira KN, Iverson TM, Maghlaoui K, Barber J, Iwata S (March 2004).
1051:
1013:
263:(3BZ1 and 3BZ2 are monomeric structures of the Photosystem II dimer),
159:
2484:
2458:
1198:
102:
90:
86:
812:
808:
361:
2191:
558:
82:
1136:
Cox, Nicholas; Pantazis, Dimitrios A.; Lubitz, Wolfgang (2020).
34:
2667:
2662:
2558:
2497:
2267:
827:
713:
622:
534:
522:
510:
434:
78:
66:
2124:
1762:
1695:
1286:
769:-state. In the absence of light, the OEC will "relax" to the S
2752:
2747:
2471:
819:
444:(PsbE–PsbF) as a secondary/protective electron carrier
140:
70:
1343:
1037:
2645:
2640:
1289:"Architecture of the photosynthetic oxygen-evolving center"
844:
816:
546:
201:
148:
2236:
1499:
1080:
101:
to plastoquinol. The energized electrons are replaced by
419:
Absorbs light energy and converts it to chemical energy
1386:
998:
112:
By replenishing lost electrons with electrons from the
19:"PSII" redirects here. For the video game console, see
777:
state is often described as being "dark-stable". The S
1556:
678:
from frozen protein crystals with a resolution of 3.8
2658:
Branched-chain alpha-keto acid dehydrogenase complex
2066:
Oettmeier W (1992). "Herbicides of photosystem II".
892:
225:
cluster (including two chloride ions), one non heme
2760:
Phosphoenolpyruvate sugar phosphotransferase system
1983:
Biochimica et Biophysica Acta (BBA) - Bioenergetics
1444:
1387:Umena Y, Kawakami K, Shen JR, Kamiya N (May 2011).
1184:
2557:
171:Schematic of PSII, highlighting electron transfer.
1840:
1654:
1135:
474:Also known as the oxygen evolving center, or OEC
464:Mobile intra-thylakoid membrane electron carrier
163:Cyanobacterial photosystem II, Monomer, PDB 2AXT.
2787:
946:
942:
940:
651:
2579:Photosynthetic reaction center complex proteins
1883:
1698:"Untangling the sequence of events during the S
89:that are then transferred through a variety of
1919:
1611:
2543:
2252:
2110:
937:
317:Photosynthetic reaction centre protein family
38:Cyanobacteria photosystem II, Dimer, PDB 2AXT
1380:
291:Protein Subunits (only with known function)
952:"Photosystem II: evolutionary perspectives"
2550:
2536:
2259:
2245:
2117:
2103:
1655:Cox N, Pantazis DA, Lubitz W (June 2020).
1227:
1033:
1031:
23:. For the Indonesian political party, see
2151:Trans-acenaphthene-1,2-diol dehydrogenase
2065:
1994:
1953:
1739:
1729:
1672:
1533:
1346:Nature Structural & Molecular Biology
1312:
1263:
1253:
1153:
1112:
1085:". In Kroneck PM, Sosa Torres ME (eds.).
1040:Nature Structural & Molecular Biology
975:
732:
655:
166:
158:
33:
1028:
865:Photosystem II light-harvesting protein
338:Photosystem II light-harvesting protein
154:
2788:
1976:
1797:
1634:10.1146/annurev-physchem-052516-044820
660:Proposed structure of Manganese Center
2531:
2240:
2098:
1674:10.1146/annurev-biochem-011520-104801
1155:10.1146/annurev-biochem-011520-104801
429:Quench excess photoexcitation energy
251:accession codes for this protein are
2743:Mitochondrial trifunctional protein
2169:Coenzyme Q - cytochrome c reductase
1920:Jablonsky J, Lazar D (April 2008).
1614:Annual Review of Physical Chemistry
815:are the most commonly used and the
765:-state), OEC returns to its basic S
749:Photosynthetic water splitting (or
13:
2076:10.1016/B978-0-444-89440-3.50018-7
1877:
1855:10.1111/j.1751-1097.1970.tb06017.x
1777:10.1111/j.1751-1097.1969.tb05696.x
1228:Kamiya N, Shen JR (January 2003).
728:
204:, one bicarbonate, 20 lipids, the
47:water-plastoquinone oxidoreductase
14:
2822:
2701:Carbamoyl phosphate synthase II
2059:
2035:
2011:
1970:
1913:
1843:Photochemistry and Photobiology
1834:
1791:
1765:Photochemistry and Photobiology
1756:
1689:
1648:
1605:
1550:
1493:
1438:
1337:
2706:Aspartate carbamoyltransferase
2614:Pyruvate dehydrogenase complex
1280:
1221:
1178:
1129:
1074:
992:
886:
373:Manganese Stabilizing Protein
25:Indonesian Islamic Union Party
1:
2738:Glycine decarboxylase complex
2733:Fatty acid synthetase complex
1661:Annual Review of Biochemistry
1142:Annual Review of Biochemistry
880:
791:
652:Oxygen-evolving complex (OEC)
358:Manganese-stabilising protein
1996:10.1016/j.bbabio.2012.02.005
7:
2266:
2019:"Chlorotraizine herbicides"
1946:10.1529/biophysj.107.122861
1095:10.1007/978-3-319-12415-5_2
950:, Faller P (January 2003).
833:
180:in the splitting of water.
10:
2829:
2770:Sucrase-isomaltase complex
2636:Oxoglutarate dehydrogenase
718:X-ray Free Electron Lasers
663:
454:Primary electron acceptor
77:. Within the photosystem,
18:
2688:
2604:
2569:
2423:
2415:Michaelis–Menten kinetics
2387:
2356:
2325:
2274:
2218:
2200:
2177:
2159:
2141:
633:
621:
609:
604:
600:
581:
569:
557:
545:
533:
521:
509:
504:
492:
487:
482:
382:. The exceptions are D1 (
377:
336:
315:
55:light-dependent reactions
2728:Electron transport chain
2307:Diffusion-limited enzyme
1977:Renger G (August 2012).
329:Reaction center protein
145:non-cyclic electron flow
61:. It is located in the
2718:P450-containing systems
1812:10.1023/A:1024946610564
1800:Photosynthesis Research
1731:10.1073/pnas.2000529117
1314:10.1126/science.1093087
1255:10.1073/pnas.0135651100
666:oxygen-evolving complex
342:Binds manganese center
2723:Cytochrome b6f complex
2210:Cytochrome b6f complex
968:10.1098/rstb.2002.1186
746:
661:
172:
164:
109:and molecular oxygen.
39:
27:. For other uses, see
2563:multienzyme complexes
2400:Eadie–Hofstee diagram
2333:Allosteric regulation
783:intermediate S-states
736:
710:X-ray crystallography
676:X-ray crystallography
659:
170:
162:
85:of light to energize
37:
2410:Lineweaver–Burk plot
2070:. pp. 349–408.
397:Coenzymes/Cofactors
155:Structure of complex
29:PS2 (disambiguation)
2775:Tryptophan synthase
2765:Polyketide synthase
2226:Alternative oxidase
1938:2008BpJ....94.2725J
1926:Biophysical Journal
1722:2020PNAS..11712624I
1716:(23): 12624–12635.
1626:2017ARPC...68..101V
1583:10.1038/nature21400
1575:2017Natur.543..131S
1526:10.1038/nature20161
1518:2016Natur.540..453Y
1471:10.1038/nature13991
1463:2015Natur.517...99S
1416:10.1038/nature09913
1408:2011Natur.473...55U
1305:2004Sci...303.1831F
1246:2003PNAS..100...98K
915:10.1038/nature04224
907:2005Natur.438.1040L
398:
292:
183:Photosystem II (of
2369:Enzyme superfamily
2302:Enzyme promiscuity
747:
662:
396:
290:
173:
165:
114:splitting of water
63:thylakoid membrane
40:
2806:Manganese enzymes
2783:
2782:
2525:
2524:
2234:
2233:
2043:"Urea herbicides"
1899:10.1021/bi048460i
1569:(7643): 131–135.
1512:(7633): 453–457.
1358:10.1038/nsmb.1559
1104:978-3-319-12414-8
1052:10.1038/nsmb.1559
1014:10.1021/bi700157n
805:cyanuric chloride
649:
648:
645:
644:
564:metabolic pathway
478:
477:
394:
393:
236:and two putative
2818:
2552:
2545:
2538:
2529:
2528:
2405:Hanes–Woolf plot
2348:Enzyme activator
2343:Enzyme inhibitor
2317:Enzyme catalysis
2261:
2254:
2247:
2238:
2237:
2187:Catechol oxidase
2119:
2112:
2105:
2096:
2095:
2090:
2089:
2068:The Photosystems
2063:
2057:
2056:
2054:
2053:
2039:
2033:
2032:
2030:
2029:
2015:
2009:
2008:
1998:
1974:
1968:
1967:
1957:
1917:
1911:
1910:
1881:
1875:
1874:
1838:
1832:
1831:
1795:
1789:
1788:
1760:
1754:
1753:
1743:
1733:
1693:
1687:
1686:
1676:
1652:
1646:
1645:
1609:
1603:
1602:
1554:
1548:
1547:
1537:
1497:
1491:
1490:
1457:(7532): 99–103.
1442:
1436:
1435:
1393:
1384:
1378:
1377:
1341:
1335:
1334:
1316:
1299:(5665): 1831–8.
1284:
1278:
1277:
1267:
1257:
1225:
1219:
1218:
1199:10.1038/35055589
1193:(6821): 739–43.
1182:
1176:
1175:
1157:
1133:
1127:
1126:
1116:
1078:
1072:
1071:
1035:
1026:
1025:
996:
990:
989:
979:
962:(1429): 245–53.
944:
935:
934:
901:(7070): 1040–4.
890:
840:Oxygen evolution
751:oxygen evolution
706:
705:
704:
696:
695:
602:
601:
480:
479:
470:Manganese center
399:
395:
293:
289:
246:
245:
244:
235:
234:
233:
224:
223:
222:
214:
213:
138:
137:
136:
120:that is used by
2828:
2827:
2821:
2820:
2819:
2817:
2816:
2815:
2801:Light reactions
2786:
2785:
2784:
2779:
2684:
2600:
2565:
2556:
2526:
2521:
2433:Oxidoreductases
2419:
2395:Enzyme kinetics
2383:
2379:List of enzymes
2352:
2321:
2292:Catalytic triad
2270:
2265:
2235:
2230:
2214:
2196:
2173:
2155:
2137:
2126:Oxidoreductases
2123:
2093:
2086:
2064:
2060:
2051:
2049:
2041:
2040:
2036:
2027:
2025:
2017:
2016:
2012:
1975:
1971:
1918:
1914:
1882:
1878:
1839:
1835:
1796:
1792:
1761:
1757:
1705:
1701:
1694:
1690:
1653:
1649:
1610:
1606:
1555:
1551:
1498:
1494:
1443:
1439:
1402:(7345): 55–60.
1391:
1385:
1381:
1342:
1338:
1285:
1281:
1226:
1222:
1183:
1179:
1134:
1130:
1105:
1079:
1075:
1036:
1029:
997:
993:
945:
938:
891:
887:
883:
875:Photoinhibition
870:Reaction Centre
836:
794:
780:
776:
772:
768:
764:
731:
729:Water splitting
703:
700:
699:
698:
694:
691:
690:
689:
687:
668:
654:
442:Cytochrome b559
243:
241:
240:
239:
237:
232:
230:
229:
228:
226:
221:
218:
217:
216:
212:
209:
208:
207:
205:
157:
143:or are used in
135:
133:
132:
131:
129:
118:proton gradient
51:protein complex
49:) is the first
32:
17:
12:
11:
5:
2826:
2825:
2814:
2813:
2808:
2803:
2798:
2796:Photosynthesis
2781:
2780:
2778:
2777:
2772:
2767:
2762:
2757:
2756:
2755:
2750:
2740:
2735:
2730:
2725:
2720:
2715:
2714:
2713:
2711:Dihydroorotase
2708:
2703:
2692:
2690:
2686:
2685:
2683:
2682:
2681:
2680:
2675:
2670:
2665:
2655:
2654:
2653:
2648:
2643:
2633:
2632:
2631:
2626:
2621:
2610:
2608:
2602:
2601:
2599:
2598:
2597:
2596:
2591:
2581:
2575:
2573:
2571:Photosynthesis
2567:
2566:
2555:
2554:
2547:
2540:
2532:
2523:
2522:
2520:
2519:
2506:
2493:
2480:
2467:
2454:
2441:
2427:
2425:
2421:
2420:
2418:
2417:
2412:
2407:
2402:
2397:
2391:
2389:
2385:
2384:
2382:
2381:
2376:
2371:
2366:
2360:
2358:
2357:Classification
2354:
2353:
2351:
2350:
2345:
2340:
2335:
2329:
2327:
2323:
2322:
2320:
2319:
2314:
2309:
2304:
2299:
2294:
2289:
2284:
2278:
2276:
2272:
2271:
2264:
2263:
2256:
2249:
2241:
2232:
2231:
2229:
2228:
2222:
2220:
2216:
2215:
2213:
2212:
2206:
2204:
2198:
2197:
2195:
2194:
2189:
2183:
2181:
2175:
2174:
2172:
2171:
2165:
2163:
2157:
2156:
2154:
2153:
2147:
2145:
2139:
2138:
2122:
2121:
2114:
2107:
2099:
2092:
2091:
2084:
2058:
2034:
2010:
1989:(8): 1164–76.
1969:
1932:(7): 2725–36.
1912:
1893:(6): 2129–42.
1876:
1833:
1806:(1–3): 65–72.
1790:
1771:(5): 309–329.
1755:
1703:
1699:
1688:
1667:(1): 795–820.
1647:
1620:(1): 101–116.
1604:
1549:
1492:
1437:
1379:
1336:
1279:
1220:
1177:
1128:
1103:
1073:
1027:
1008:(14): 4390–7.
991:
936:
884:
882:
879:
878:
877:
872:
867:
862:
857:
852:
850:Photosynthesis
847:
842:
835:
832:
822:which include
793:
790:
778:
774:
770:
766:
762:
730:
727:
701:
692:
664:Main article:
653:
650:
647:
646:
643:
642:
637:
631:
630:
625:
619:
618:
613:
607:
606:
598:
597:
586:
579:
578:
573:
567:
566:
561:
555:
554:
549:
543:
542:
537:
531:
530:
525:
519:
518:
513:
507:
506:
502:
501:
496:
490:
489:
485:
484:
483:Photosystem II
476:
475:
472:
466:
465:
462:
456:
455:
452:
446:
445:
438:
431:
430:
427:
421:
420:
417:
411:
410:
405:
392:
391:
375:
374:
371:
355:
351:
350:
348:
344:
343:
340:
335:
331:
330:
327:
323:
322:
319:
314:
310:
309:
304:
299:
242:
231:
219:
210:
156:
153:
134:
105:water to form
59:photosynthesis
43:Photosystem II
15:
9:
6:
4:
3:
2:
2824:
2823:
2812:
2809:
2807:
2804:
2802:
2799:
2797:
2794:
2793:
2791:
2776:
2773:
2771:
2768:
2766:
2763:
2761:
2758:
2754:
2751:
2749:
2746:
2745:
2744:
2741:
2739:
2736:
2734:
2731:
2729:
2726:
2724:
2721:
2719:
2716:
2712:
2709:
2707:
2704:
2702:
2699:
2698:
2697:
2694:
2693:
2691:
2687:
2679:
2676:
2674:
2671:
2669:
2666:
2664:
2661:
2660:
2659:
2656:
2652:
2649:
2647:
2644:
2642:
2639:
2638:
2637:
2634:
2630:
2627:
2625:
2622:
2620:
2617:
2616:
2615:
2612:
2611:
2609:
2607:
2606:Dehydrogenase
2603:
2595:
2592:
2590:
2587:
2586:
2585:
2582:
2580:
2577:
2576:
2574:
2572:
2568:
2564:
2560:
2553:
2548:
2546:
2541:
2539:
2534:
2533:
2530:
2517:
2513:
2512:
2507:
2504:
2500:
2499:
2494:
2491:
2487:
2486:
2481:
2478:
2474:
2473:
2468:
2465:
2461:
2460:
2455:
2452:
2448:
2447:
2442:
2439:
2435:
2434:
2429:
2428:
2426:
2422:
2416:
2413:
2411:
2408:
2406:
2403:
2401:
2398:
2396:
2393:
2392:
2390:
2386:
2380:
2377:
2375:
2374:Enzyme family
2372:
2370:
2367:
2365:
2362:
2361:
2359:
2355:
2349:
2346:
2344:
2341:
2339:
2338:Cooperativity
2336:
2334:
2331:
2330:
2328:
2324:
2318:
2315:
2313:
2310:
2308:
2305:
2303:
2300:
2298:
2297:Oxyanion hole
2295:
2293:
2290:
2288:
2285:
2283:
2280:
2279:
2277:
2273:
2269:
2262:
2257:
2255:
2250:
2248:
2243:
2242:
2239:
2227:
2224:
2223:
2221:
2217:
2211:
2208:
2207:
2205:
2203:
2199:
2193:
2190:
2188:
2185:
2184:
2182:
2180:
2176:
2170:
2167:
2166:
2164:
2162:
2158:
2152:
2149:
2148:
2146:
2144:
2140:
2135:
2131:
2127:
2120:
2115:
2113:
2108:
2106:
2101:
2100:
2097:
2087:
2085:9780444894403
2081:
2077:
2073:
2069:
2062:
2048:
2044:
2038:
2024:
2020:
2014:
2006:
2002:
1997:
1992:
1988:
1984:
1980:
1973:
1965:
1961:
1956:
1951:
1947:
1943:
1939:
1935:
1931:
1927:
1923:
1916:
1908:
1904:
1900:
1896:
1892:
1888:
1880:
1872:
1868:
1864:
1860:
1856:
1852:
1849:(6): 457–75.
1848:
1844:
1837:
1829:
1825:
1821:
1817:
1813:
1809:
1805:
1801:
1794:
1786:
1782:
1778:
1774:
1770:
1766:
1759:
1751:
1747:
1742:
1737:
1732:
1727:
1723:
1719:
1715:
1711:
1707:
1692:
1684:
1680:
1675:
1670:
1666:
1662:
1658:
1651:
1643:
1639:
1635:
1631:
1627:
1623:
1619:
1615:
1608:
1600:
1596:
1592:
1588:
1584:
1580:
1576:
1572:
1568:
1564:
1560:
1553:
1545:
1541:
1536:
1531:
1527:
1523:
1519:
1515:
1511:
1507:
1503:
1496:
1488:
1484:
1480:
1476:
1472:
1468:
1464:
1460:
1456:
1452:
1448:
1441:
1433:
1429:
1425:
1421:
1417:
1413:
1409:
1405:
1401:
1397:
1390:
1383:
1375:
1371:
1367:
1363:
1359:
1355:
1352:(3): 334–42.
1351:
1347:
1340:
1332:
1328:
1324:
1320:
1315:
1310:
1306:
1302:
1298:
1294:
1290:
1283:
1275:
1271:
1266:
1261:
1256:
1251:
1247:
1243:
1240:(1): 98–103.
1239:
1235:
1231:
1224:
1216:
1212:
1208:
1204:
1200:
1196:
1192:
1188:
1181:
1173:
1169:
1165:
1161:
1156:
1151:
1147:
1143:
1139:
1132:
1124:
1120:
1115:
1110:
1106:
1100:
1096:
1092:
1088:
1084:
1077:
1069:
1065:
1061:
1057:
1053:
1049:
1046:(3): 334–42.
1045:
1041:
1034:
1032:
1023:
1019:
1015:
1011:
1007:
1003:
995:
987:
983:
978:
973:
969:
965:
961:
957:
953:
949:
948:Rutherford AW
943:
941:
932:
928:
924:
920:
916:
912:
908:
904:
900:
896:
889:
885:
876:
873:
871:
868:
866:
863:
861:
860:Photosystem I
858:
856:
853:
851:
848:
846:
843:
841:
838:
837:
831:
829:
825:
821:
818:
814:
810:
806:
803:derived from
802:
798:
789:
786:
784:
760:
754:
752:
744:
740:
735:
726:
723:
719:
715:
711:
685:
681:
677:
672:
667:
658:
641:
638:
636:
632:
629:
626:
624:
620:
617:
614:
612:
608:
603:
599:
596:
593:
590:
587:
584:
580:
577:
574:
572:
568:
565:
562:
560:
556:
553:
550:
548:
544:
541:
540:NiceZyme view
538:
536:
532:
529:
526:
524:
520:
517:
514:
512:
508:
503:
500:
497:
495:
491:
486:
481:
473:
471:
468:
467:
463:
461:
460:Plastoquinone
458:
457:
453:
451:
448:
447:
443:
439:
436:
433:
432:
428:
426:
425:Beta-carotene
423:
422:
418:
416:
413:
412:
409:
406:
404:
401:
400:
389:
385:
381:
376:
372:
369:
368:
363:
359:
356:
353:
352:
349:
346:
345:
341:
339:
333:
332:
328:
325:
324:
320:
318:
312:
311:
308:
305:
303:
300:
298:
295:
294:
288:
286:
282:
278:
274:
270:
266:
262:
258:
254:
250:
203:
199:
198:plastoquinone
195:
191:
190:beta-carotene
186:
185:cyanobacteria
181:
178:
169:
161:
152:
150:
146:
142:
127:
123:
119:
115:
110:
108:
107:hydrogen ions
104:
100:
99:plastoquinone
96:
92:
88:
84:
80:
76:
75:cyanobacteria
72:
68:
64:
60:
56:
52:
48:
44:
36:
30:
26:
22:
21:PlayStation 2
2593:
2511:Translocases
2508:
2495:
2482:
2469:
2456:
2446:Transferases
2443:
2430:
2287:Binding site
2067:
2061:
2050:. Retrieved
2047:alanwood.net
2046:
2037:
2026:. Retrieved
2023:alanwood.net
2022:
2013:
1986:
1982:
1972:
1929:
1925:
1915:
1890:
1887:Biochemistry
1886:
1879:
1846:
1842:
1836:
1803:
1799:
1793:
1768:
1764:
1758:
1713:
1709:
1691:
1664:
1660:
1650:
1617:
1613:
1607:
1566:
1562:
1552:
1509:
1505:
1495:
1454:
1450:
1440:
1399:
1395:
1382:
1349:
1345:
1339:
1296:
1292:
1282:
1237:
1233:
1223:
1190:
1186:
1180:
1145:
1141:
1131:
1086:
1082:
1076:
1043:
1039:
1005:
1002:Biochemistry
1001:
994:
959:
955:
898:
894:
888:
824:chlortoluron
795:
787:
782:
773:state; the S
758:
755:
748:
742:
738:
673:
669:
528:BRENDA entry
407:
402:
387:
383:
379:
365:
347:CP47 (PsbB)
334:CP43 (PsbC)
306:
301:
296:
182:
177:photosystems
174:
124:to generate
122:ATP synthase
111:
57:of oxygenic
46:
42:
41:
2584:Photosystem
2282:Active site
1148:: 795–820.
855:Photosystem
516:IntEnz view
488:Identifiers
415:Chlorophyll
2790:Categories
2485:Isomerases
2459:Hydrolases
2326:Regulation
2052:2021-03-26
2028:2021-03-26
881:References
797:Inhibitors
792:Inhibitors
720:, such as
585:structures
552:KEGG entry
450:Pheophytin
386:) and D2 (
326:D2 (PsbD)
313:D1 (PsbA)
194:pheophytin
97:to reduce
2811:EC 1.10.3
2364:EC number
1599:205254025
1487:205241611
1432:205224374
1172:214645936
807:of which
801:triazines
505:Databases
440:Bound to
367:IPR002628
103:oxidizing
95:cofactors
91:coenzymes
87:electrons
2388:Kinetics
2312:Cofactor
2275:Activity
2132:family (
2130:diphenol
2005:22353626
1964:18178650
1907:15697239
1871:31914925
1820:16228566
1785:96744015
1750:32434915
1683:32208765
1642:28226223
1591:28219079
1544:27871088
1479:25470056
1424:21499260
1374:23034289
1366:19219048
1331:31521054
1323:14764885
1274:12518057
1207:11217865
1164:32208765
1123:25707465
1068:23034289
1060:19219048
1022:17371054
986:12594932
923:16355230
834:See also
830:(DCMU).
813:simazine
809:atrazine
759:S-states
640:proteins
628:articles
616:articles
589:RCSB PDB
499:1.10.3.9
408:Function
403:Cofactor
362:InterPro
307:Function
81:capture
2559:Enzymes
2498:Ligases
2268:Enzymes
2202:1.10.99
2192:Laccase
1955:2267143
1934:Bibcode
1863:5456273
1828:8742213
1741:7293653
1718:Bibcode
1622:Bibcode
1571:Bibcode
1535:5201176
1514:Bibcode
1459:Bibcode
1404:Bibcode
1301:Bibcode
1293:Science
1242:Bibcode
1215:4344756
1114:4688042
977:1693113
931:4394735
903:Bibcode
576:profile
559:MetaCyc
364::
297:Subunit
83:photons
79:enzymes
53:in the
2668:BCKDHB
2663:BCKDHA
2472:Lyases
2179:1.10.3
2161:1.10.2
2143:1.10.1
2082:
2003:
1962:
1952:
1905:
1869:
1861:
1826:
1818:
1783:
1748:
1738:
1681:
1640:
1597:
1589:
1563:Nature
1542:
1532:
1506:Nature
1485:
1477:
1451:Nature
1430:
1422:
1396:Nature
1372:
1364:
1329:
1321:
1272:
1265:140893
1262:
1213:
1205:
1187:Nature
1170:
1162:
1121:
1111:
1101:
1066:
1058:
1020:
984:
974:
929:
921:
895:Nature
828:diuron
714:X-rays
623:PubMed
605:Search
595:PDBsum
535:ExPASy
523:BRENDA
511:IntEnz
494:EC no.
435:Heme B
302:Family
200:, two
196:, two
192:, two
73:, and
67:plants
2753:HADHB
2748:HADHA
2689:Other
2424:Types
2219:Other
2136:1.10)
1867:S2CID
1824:S2CID
1781:S2CID
1595:S2CID
1483:S2CID
1428:S2CID
1392:(PDF)
1370:S2CID
1327:S2CID
1211:S2CID
1168:S2CID
1064:S2CID
927:S2CID
820:ureas
571:PRIAM
141:NADPH
71:algae
2646:DLST
2641:OGDH
2516:list
2509:EC7
2503:list
2496:EC6
2490:list
2483:EC5
2477:list
2470:EC4
2464:list
2457:EC3
2451:list
2444:EC2
2438:list
2431:EC1
2080:ISBN
2001:PMID
1987:1817
1960:PMID
1903:PMID
1859:PMID
1816:PMID
1746:PMID
1679:PMID
1638:PMID
1587:PMID
1540:PMID
1475:PMID
1420:PMID
1362:PMID
1319:PMID
1270:PMID
1203:PMID
1160:PMID
1119:PMID
1099:ISBN
1056:PMID
1018:PMID
982:PMID
919:PMID
845:P680
826:and
817:aryl
811:and
722:SLAC
635:NCBI
592:PDBe
547:KEGG
437:559
388:PsbD
384:PsbA
380:PsbO
285:1IZL
281:1FE1
277:1ILX
273:1W5C
269:1S5L
265:2AXT
261:3BZ2
257:3BZ1
253:3WU2
202:heme
149:DCMU
130:NADP
93:and
45:(or
2696:CAD
2678:DLD
2673:DBT
2651:DLD
2072:doi
1991:doi
1950:PMC
1942:doi
1895:doi
1851:doi
1808:doi
1773:doi
1736:PMC
1726:doi
1714:117
1702:→ S
1669:doi
1630:doi
1579:doi
1567:543
1530:PMC
1522:doi
1510:540
1467:doi
1455:517
1412:doi
1400:473
1354:doi
1309:doi
1297:303
1260:PMC
1250:doi
1238:100
1195:doi
1191:409
1150:doi
1109:PMC
1091:doi
1048:doi
1010:doi
972:PMC
964:doi
960:358
911:doi
899:438
697:CaO
611:PMC
583:PDB
390:).
287:.
249:PDB
215:CaO
139:to
126:ATP
65:of
2792::
2629:E3
2624:E2
2619:E1
2594:II
2561::
2134:EC
2128::
2078:.
2045:.
2021:.
1999:.
1985:.
1981:.
1958:.
1948:.
1940:.
1930:94
1928:.
1924:.
1901:.
1891:44
1889:.
1865:.
1857:.
1847:11
1845:.
1822:.
1814:.
1804:76
1802:.
1779:.
1769:10
1767:.
1744:.
1734:.
1724:.
1712:.
1708:.
1677:.
1665:89
1663:.
1659:.
1636:.
1628:.
1618:68
1616:.
1593:.
1585:.
1577:.
1565:.
1561:.
1538:.
1528:.
1520:.
1508:.
1504:.
1481:.
1473:.
1465:.
1453:.
1449:.
1426:.
1418:.
1410:.
1398:.
1394:.
1368:.
1360:.
1350:16
1348:.
1325:.
1317:.
1307:.
1295:.
1291:.
1268:.
1258:.
1248:.
1236:.
1232:.
1209:.
1201:.
1189:.
1166:.
1158:.
1146:89
1144:.
1140:.
1117:.
1107:.
1097:.
1062:.
1054:.
1044:16
1042:.
1030:^
1016:.
1006:46
1004:.
980:.
970:.
958:.
954:.
939:^
925:.
917:.
909:.
897:.
688:Mn
370:)
354:O
283:,
279:,
275:,
271:,
267:,
259:,
255:,
238:Ca
227:Fe
206:Mn
147:.
69:,
2589:I
2551:e
2544:t
2537:v
2518:)
2514:(
2505:)
2501:(
2492:)
2488:(
2479:)
2475:(
2466:)
2462:(
2453:)
2449:(
2440:)
2436:(
2260:e
2253:t
2246:v
2118:e
2111:t
2104:v
2088:.
2074::
2055:.
2031:.
2007:.
1993::
1966:.
1944::
1936::
1909:.
1897::
1873:.
1853::
1830:.
1810::
1787:.
1775::
1752:.
1728::
1720::
1704:3
1700:2
1685:.
1671::
1644:.
1632::
1624::
1601:.
1581::
1573::
1546:.
1524::
1516::
1489:.
1469::
1461::
1434:.
1414::
1406::
1376:.
1356::
1333:.
1311::
1303::
1276:.
1252::
1244::
1217:.
1197::
1174:.
1152::
1125:.
1093::
1070:.
1050::
1024:.
1012::
988:.
966::
933:.
913::
905::
779:1
775:1
771:1
767:0
763:4
743:B
739:A
702:5
693:4
684:Å
680:Å
360:(
220:5
211:4
31:.
Text is available under the Creative Commons Attribution-ShareAlike License. Additional terms may apply.
