693:
1617:
1346:
31:
1285:
395:
2000:
2168:
1855:
1821:
919:
although external ions are attracted by the negative intracellular potential, entropy seeks to diffuse the ions already concentrated inside the cell. The converse phenomenon (osmosis supports transport, electric potential opposes it) can be achieved for Na in cells with abnormal transmembrane potentials: at
1357:
to drive the formation of proton gradients in chloroplasts, however, PSII utilizes vectorial redox chemistry to achieve this goal. Rather than physically transporting protons through the protein, reactions requiring the binding of protons will occur on the extracellular side while reactions requiring
918:
In the example of Na, both terms tend to support transport: the negative electric potential inside the cell attracts the positive ion and since Na is concentrated outside the cell, osmosis supports diffusion through the Na channel into the cell. In the case of K, the effect of osmosis is reversed:
2004:
Complex IV (CIV) catalyzes the transfer of two electrons from the cytochrome c reduced by CIII to one half of a full oxygen. Utilizing one full oxygen in oxidative phosphorylation requires the transfer of four electrons. The oxygen will then consume four protons from the matrix to form water while
2008:
1557:
719:
of ATP into ADP and an inorganic phosphate; for every molecule of ATP hydrolized, three Na are transported outside and two K are transported inside the cell. This makes the inside of the cell more negative than the outside and more specifically generates a membrane potential
160:
across that same membrane. In the former effect, the concentrated charge attracts charges of the opposite sign; in the latter, the concentrated species tends to diffuse across the membrane to an equalize concentrations. The combination of these two phenomena determines the
1659:
291:
1324:
forming the K state. This moves SB away from Asp85 and Asp212, causing H transfer from the SB to Asp85 forming the M1 state. The protein then shifts to the M2 state by separating Glu204 from Glu194 which releases a proton from Glu204 into the external medium. The SB is
885:
1995:{\displaystyle 2\underbrace {\text{cytochrome c}} _{\text{oxidized}}+{\ce {UQH_2}}+2\underbrace {{\ce {H^+}}} _{\text{matrix}}\longrightarrow 2\underbrace {\text{cytochrome c}} _{\text{reduced}}+{\ce {UQ}}+4\underbrace {{\ce {H^+}}} _{\text{IMS}}}
377:) correspond to turbines that convert the water's potential energy to other forms of physical or chemical energy, and the ions that pass through the membrane correspond to water traveling into the lower river. Conversely, energy can be used to
1304:. The proton pump relies on proton carriers to drive protons from the side of the membrane with a low H concentration to the side of the membrane with a high H concentration. In bacteriorhodopsin, the proton pump is activated by absorption of
81:
across a membrane. If there is an unequal distribution of charges across the membrane, then the difference in electric potential generates a force that drives ion diffusion until the charges are balanced on both sides of the membrane.
1404:
636:
of chloroplasts to drive the synthesis of ATP. The proton gradient can be generated through either noncyclic or cyclic photophosphorylation. Of the proteins that participate in noncyclic photophosphorylation,
529:. In a battery, an electrochemical potential arising from the movement of ions balances the reaction energy of the electrodes. The maximum voltage that a battery reaction can produce is sometimes called the
2163:{\displaystyle 2{\text{cytochrome c}}({\text{reduced}})+4{\ce {H+}}({\text{matrix}})+{\frac {1}{2}}{\ce {O2}}\longrightarrow 2{\text{cytochrome c}}({\text{oxidized}})+2{\ce {H+}}({\text{IMS}})+{\ce {H2O}}}
178:
73:
When there are unequal concentrations of an ion across a permeable membrane, the ion will move across the membrane from the area of higher concentration to the area of lower concentration through
756:
1337:. The protonation of Asp85 and Asp96 causes re-isomerization of the SB, forming the O state. Finally, bacteriorhodopsin returns to its resting state when Asp85 releases its proton to Glu204.
1816:{\displaystyle {\ce {NADH}}+{\ce {H^+}}+{\ce {UQ}}+4\underbrace {{\ce {H^+}}} _{\mathrm {matrix} }\longrightarrow {\ce {NAD^+}}+{\ce {UQH_2}}+4\underbrace {{\ce {H^+}}} _{\mathrm {IMS} }}
2377:
Shattock, Michael J.; Ottolia, Michela; Bers, Donald M.; Blaustein, Mordecai P.; Boguslavskyi, Andrii; Bossuyt, Julie; Bridge, John H. B.; Chen-Izu, Ye; Clancy, Colleen E. (2015-03-15).
2977:
Sun, Fei; Zhou, Qiangjun; Pang, Xiaoyun; Xu, Yingzhi; Rao, Zihe (2013-08-01). "Revealing various coupling of electron transfer and proton pumping in mitochondrial respiratory chain".
121:
and has industrial applications such as batteries and fuel cells. In biology, electrochemical gradients allow cells to control the direction ions move across membranes. In
2695:
Spitzer, Kenneth W.; Vaughan-Jones, Richard D. (2003), Karmazyn, Morris; Avkiran, Metin; Fliegel, Larry (eds.), "Regulation of
Intracellular pH in Mammalian Cells",
700:
Since the ions are charged, they cannot pass through cellular membranes via simple diffusion. Two different mechanisms can transport the ions across the membrane:
3055:
1581:
in two separate reactions. The process that occurs is similar to the Q-cycle in
Complex III of the electron transport chain. In the first reaction, PQH
3029:
Stephen T. Abedon, "Important words and concepts from
Chapter 8, Campbell & Reece, 2002 (1/14/2005)", for Biology 113 at the Ohio State University
416:
2925:"Photosynthetic complex stoichiometry dynamics in higher plants: biogenesis, function, and turnover of ATP synthase and the cytochrome b 6 f complex"
1264:
Proton gradients in particular are important in many types of cells as a form of energy storage. The gradient is usually used to drive ATP synthase,
1552:{\displaystyle 4h\nu +2{\ce {H2O}}+2{\ce {PQ}}+4{\ce {H+}}({\text{stroma}})\longrightarrow {\ce {O2}}+2{\ce {PQH2}}+4{\ce {H+}}({\text{lumen}})}
734:
An example of passive transport is ion fluxes through Na, K, Ca, and Cl channels. Unlike active transport, passive transport is powered by the
508:
3019:
2674:
2310:
3185:
1358:
the release of protons will occur on the intracellular side. Absorption of photons of 680nm wavelength is used to excite two electrons in
653:
directly contribute to generating the proton gradient. For each four photons absorbed by PSII, eight protons are pumped into the lumen.
3028:
467:
2184:
573:
439:
3254:
2712:
1653:
599:
486:
3048:
1608:
gets oxidized, adding an electron to another plastocyanin and PQ. Both reactions together transfer four protons into the lumen.
446:
286:{\displaystyle \nabla {\overline {\mu }}_{i}=\nabla \mu _{i}({\vec {r}})+z_{i}\mathrm {F} \nabla \varphi ({\vec {r}}){\text{,}}}
420:
2642:"The following table gives an idea of the intra and extra cellular ion concentrations in a squid axon and a mammalian cell"
3249:
3205:
594:, composed of four complexes embedded in the inner mitochondrial membrane. Complexes I, III, and IV pump protons from the
453:
3269:
3041:
1272:
transport. This section will focus on three processes that help establish proton gradients in their respective cells:
118:
2596:
435:
3259:
3153:
2729:"pH of the Cytoplasm and Periplasm of Escherichia coli: Rapid Measurement by Green Fluorescent Protein Fluorimetry"
656:
Several other transporters and ion channels play a role in generating a proton electrochemical gradient. One is TPK
370:
3264:
900:
626:
583:
172:
43:
2616:
Lodish, Harvey; Berk, Arnold; Zipursky, S. Lawrence; Matsudaira, Paul; Baltimore, David; Darnell, James (2000).
3148:
2214:
949:
591:
405:
142:
1586:
3274:
1394:
880:{\displaystyle \Delta G=RT\ln {\!\left({\frac {c_{\rm {in}}}{c_{\rm {out}}}}\right)}+(Fz)V_{\rm {membrane}}}
606:
entering the chain, ten protons translocate into the IMS. The result is an electric potential of more than
424:
409:
2785:
1566:
646:
138:
114:
3115:
3100:
2801:
619:
522:
460:
2209:
1036:
980:
595:
587:
569:
98:
90:
1329:
by Asp96 which forms the N state. It is important that the second proton comes from Asp96 since its
3074:
2487:
928:
692:
504:
130:
74:
1288:
Diagram of the conformational shift in retinal that initiates proton pumping in bacteriorhodopsin.
3094:
3064:
2526:
2272:
2219:
2194:
2179:
1066:
554:
546:
500:
343:
78:
1616:
3082:
3015:
2994:
2954:
2946:
2905:
2864:
2766:
2748:
2708:
2670:
2577:
2518:
2510:
2468:
2460:
2416:
2398:
2359:
2306:
2264:
2256:
1345:
1293:
1273:
1008:
750:
705:
661:
526:
518:
366:
2641:
350:
Sometimes, the term "electrochemical potential" is abused to describe the electric potential
3125:
3090:
2986:
2936:
2895:
2854:
2846:
2789:
2756:
2740:
2700:
2567:
2557:
2502:
2450:
2406:
2390:
2349:
2339:
2248:
2199:
2189:
954:
944:
912:
747:
701:
514:
333:
329:
157:
110:
86:
30:
2793:
2617:
2437:
Aperia, Anita; Akkuratov, Evgeny E.; Fontana, Jacopo Maria; Brismar, Hjalmar (2016-04-01).
1385:) which is released from PSII after gaining two protons from the stroma. The electrons in P
1284:
2814:
712:
633:
153:
66:
17:
680:) transports K into the thylakoid lumen and H into the stroma, which helps establish the
27:
Gradient of electrochemical potential, usually for an ion that can move across a membrane
2859:
2834:
2761:
2728:
2572:
2545:
2411:
2378:
2354:
2327:
743:
735:
669:
665:
638:
565:
378:
146:
94:
2923:
Schöttler, Mark Aurel; Tóth, Szilvia Z.; Boulouis, Alix; Kahlau, Sabine (2015-05-01).
3243:
3226:
3180:
2204:
1370:
1330:
1313:
642:
603:
580:
542:
541:
The generation of a transmembrane electrical potential through ion movement across a
169:
59:
2530:
3210:
3190:
2276:
1837:
1637:
1590:
1578:
1366:
892:
611:
122:
34:
Diagram of ion concentrations and charge across a semi-permeable cellular membrane.
2882:
Wickstrand, Cecilia; Dods, Robert; Royant, Antoine; Neutze, Richard (2015-03-01).
2239:
Nath, Sunil; Villadsen, John (2015-03-01). "Oxidative phosphorylation revisited".
2900:
2883:
2850:
2669:. Vol. 565 (Seventh ed.). Basel: Ciba-Geigy Limited. pp. 653–654.
2394:
1656:(NADH) which translocates four protons from the mitochondrial matrix to the IMS:
610:. The energy resulting from the flux of protons back into the matrix is used by
3200:
3195:
3172:
3110:
2704:
1825:
1326:
1317:
1301:
1148:
394:
152:
An electrochemical gradient has two components: a differential concentration of
126:
2544:
Höhner, Ricarda; Aboukila, Ali; Kunz, Hans-Henning; Venema, Kees (2016-01-01).
2455:
1593:
which then transfers it to plastocyanin. The second electron is transferred to
3220:
3143:
2990:
2546:"Proton Gradients and Proton-Dependent Transport Processes in the Chloroplast"
2506:
1585:
binds to the complex on the lumen side and one electron is transferred to the
1309:
1269:
716:
673:
2950:
2884:"Bacteriorhodopsin: Would the real structural intermediates please stand up?"
2752:
2514:
2464:
2402:
2344:
2260:
3138:
3133:
2562:
1641:
1629:
1625:
1265:
630:
615:
561:
374:
2998:
2958:
2909:
2868:
2770:
2581:
2522:
2488:"Regulation of the mitochondrial proton gradient by cytosolic Ca2+ signals"
2472:
2420:
2363:
2268:
2941:
2924:
2618:"Table 15-1, Typical Ion Concentrations in Invertebrates and Vertebrates"
1649:
1098:
959:
362:
51:
2744:
3163:
3105:
2005:
another four protons are pumped into the IMS, to give a total reaction
1829:
1334:
1321:
1297:
1123:
739:
664:
that is activated by Ca and conducts K from the thylakoid lumen to the
558:
381:, and chemical energy can be used to create electrochemical gradients.
2252:
3033:
1832:. The first step involving the transfer of two electrons from the UQH
1594:
1305:
939:
2835:"Molecular mechanisms for generating transmembrane proton gradients"
2438:
2833:
Gunner, M. R.; Amin, Muhamed; Zhu, Xuyu; Lu, Jianxun (2013-08-01).
2697:
The Sodium-Hydrogen
Exchanger: From Molecule to its Role in Disease
1604:
which then transfers it to PQ. In the second reaction, a second PQH
923:, the Na influx halts; at higher potentials, it becomes an efflux.
109:
Electrochemical energy is one of the many interchangeable forms of
1633:
1615:
1390:
1354:
1344:
1333:
state is unstable and rapidly reprotonated with a proton from the
1283:
1276:
and noncyclic photophosphorylation and oxidative phosphorylation.
691:
550:
29:
1620:
Detailed diagram of the electron transport chain in mitochondria.
1369:. These higher energy electrons are transferred to protein-bound
1359:
3037:
2890:. Structural biochemistry and biophysics of membrane proteins.
332:(the electrochemical potential is implicitly measured on a per-
388:
162:
47:
1844:
site. In the second step, two more electrons reduce UQ to UQH
85:
Electrochemical gradients are essential to the operation of
2597:"» What are the concentrations of different ions in cells?"
681:
168:
The combined effect can be quantified as a gradient in the
572:
to the extracellular region; a typical animal cell has an
521:
in the form of an applied voltage is used to modulate the
2439:"Na+-K+-ATPase, a new class of plasma membrane receptors"
2326:
Yang, Huanghe; Zhang, Guohui; Cui, Jianmin (2015-01-01).
2153:
2084:
1893:
1769:
1516:
1495:
1434:
1381:). This reduces plastoquinone (PQ) to plastoquinol (PQH
2888:
Biochimica et
Biophysica Acta (BBA) - General Subjects
2727:
Slonczewski, Joan L.; Wilks, Jessica C. (2007-08-01).
568:. By convention, physiological voltages are measured
361:
An electrochemical gradient is analogous to the water
156:
across a membrane and a differential concentration of
2011:
1858:
1662:
1407:
759:
181:
2786:"Rising Acidity in the Ocean: The Other CO2 Problem"
2432:
2430:
2328:"BK channels: multiple sensors, one activation gate"
3219:
3171:
3162:
3124:
3081:
2841:. Metals in Bioenergetics and Biomimetics Systems.
2839:
Biochimica et
Biophysica Acta (BBA) - Bioenergetics
77:. Ions also carry an electric charge that forms an
2379:"Na+/Ca2+ exchange and Na+/K+-ATPase in the heart"
2296:
2294:
2292:
2290:
2288:
2286:
2162:
1994:
1815:
1551:
879:
672:. On the other hand, the electro-neutral K efflux
285:
2972:
2970:
2968:
782:
2486:Poburko, Damon; Demaurex, Nicolas (2012-04-24).
2828:
2826:
2824:
2495:Pflügers Archiv: European Journal of Physiology
2443:American Journal of Physiology. Cell Physiology
753:change associated with successful transport is
1401:and H into the lumen, for a total reaction of
746:(the transmembrane potential). Formally, the
711:An example of active transport of ions is the
3049:
1573:, which then transfers two electrons from PQH
625:Similar to the electron transport chain, the
354:by an ionic concentration gradient; that is,
161:thermodynamically-preferred direction for an
8:
579:An electrochemical gradient is essential to
1836:reduced by CI to two molecules of oxidized
1349:Simplified diagram of photophosphorylation.
423:. Unsourced material may be challenged and
3168:
3073:Mechanisms for chemical transport through
3056:
3042:
3034:
576:of (−70)–(−50) mV.
305:the chemical potential of the ion species
101:, and certain other biological processes.
65:The electrical gradient, or difference in
2940:
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2760:
2571:
2561:
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2353:
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1510:
1505:
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1473:
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1433:
1428:
1423:
1406:
849:
848:
809:
808:
794:
793:
787:
781:
758:
487:Learn how and when to remove this message
379:pump water up into the lake above the dam
369:. Routes unblocked by the membrane (e.g.
278:
264:
263:
249:
243:
222:
221:
212:
196:
186:
180:
1377:) and then to unbound plastoquinone (PQ
1208:
1206:
1204:
1180:7.35 to 7.45 (normal arterial blood pH)
925:
629:of photosynthesis pump protons into the
58:The chemical gradient, or difference in
2231:
1222:
1220:
1218:
1200:
2810:
2799:
2794:10.1038/scientificamericanearth0908-22
2784:Brewer, Peter G. (September 1, 2008).
509:table of standard electrode potentials
54:. The gradient consists of two parts:
2979:Current Opinion in Structural Biology
2690:
2688:
2686:
7:
3186:Non-specific, adsorptive pinocytosis
2303:Lehninger Principles of Biochemistry
2301:Nelson, David; Cox, Michael (2013).
927:Common cellular ion concentrations (
421:adding citations to reliable sources
1561:After being released from PSII, PQH
1397:(OEC). This results in release of O
2185:Transmembrane potential difference
1807:
1804:
1801:
1734:
1731:
1728:
1725:
1722:
1719:
871:
868:
865:
862:
859:
856:
853:
850:
816:
813:
810:
798:
795:
760:
742:(a concentration gradient) and an
531:standard electrochemical potential
319:the charge per ion of the species
254:
250:
205:
182:
25:
1654:nicotinamide adenine dinucleotide
1624:In the electron transport chain,
1600:which then transfers it to heme b
165:'s movement across the membrane.
2241:Biotechnology and Bioengineering
393:
1296:generates a proton gradient in
1124:X (negatively charged proteins)
2929:Journal of Experimental Botany
2699:, Springer US, pp. 1–15,
2136:
2128:
2107:
2099:
2088:
2057:
2049:
2028:
2020:
1929:
1740:
1546:
1538:
1481:
1478:
1470:
841:
832:
513:The term typically applies in
275:
269:
260:
233:
227:
218:
149:quickly transmit information.
1:
3206:Receptor-mediated endocytosis
3014:. Pearson Benjamin Cummings.
3010:Campbell & Reece (2005).
2595:Philips, Ron Milo & Ron.
1852:site. The total reaction is:
1389:are replenished by oxidizing
715:(NKA). NKA is powered by the
574:internal electrical potential
2901:10.1016/j.bbagen.2014.05.021
2851:10.1016/j.bbabio.2013.03.001
2395:10.1113/jphysiol.2014.282319
668:, which helps establish the
191:
113:through which energy may be
2705:10.1007/978-1-4615-0427-6_1
1589:which then transfers it to
907:is the charge per ion, and
696:Diagram of the Na-K-ATPase.
119:electroanalytical chemistry
3291:
3154:Secondary active transport
2550:Frontiers in Plant Science
2456:10.1152/ajpcell.00359.2015
2305:. New York: W.H. Freeman.
1182:6.9 - 7.8 (overall range)
523:thermodynamic favorability
498:
436:"Electrochemical gradient"
371:membrane transport protein
18:Proton electromotive force
3071:
2991:10.1016/j.sbi.2013.06.013
2507:10.1007/s00424-012-1106-y
2383:The Journal of Physiology
1648:) by the transfer of two
1612:Oxidative phosphorylation
1114:
958:
953:
948:
943:
938:
935:
627:light-dependent reactions
584:oxidative phosphorylation
173:electrochemical potential
143:sodium-potassium gradient
44:electrochemical potential
3255:Electrochemical concepts
3149:Primary active transport
2345:10.3389/fphys.2015.00029
2215:Proton exchange membrane
592:electron transport chain
40:electrochemical gradient
2733:Journal of Bacteriology
2563:10.3389/fpls.2016.00218
2332:Frontiers in Physiology
1395:oxygen-evolving complex
50:that can move across a
2809:Cite journal requires
2164:
1996:
1817:
1621:
1553:
1350:
1289:
881:
697:
287:
135:chemiosmotic potential
35:
3101:Facilitated diffusion
2165:
1997:
1828:(CIII) catalyzes the
1818:
1619:
1554:
1348:
1287:
882:
695:
614:to combine inorganic
288:
91:electrochemical cells
33:
3250:Cellular respiration
3075:biological membranes
2622:www.ncbi.nlm.nih.gov
2210:Electrochemical cell
2009:
1856:
1660:
1405:
1353:PSII also relies on
1341:Photophosphorylation
901:absolute temperature
757:
588:cellular respiration
586:. The final step of
547:biological processes
417:improve this section
179:
99:cellular respiration
60:solute concentration
3270:Physical quantities
2745:10.1128/JB.00615-07
2155:
2086:
1895:
1771:
1518:
1497:
1436:
932:
600:intermembrane space
505:electrode potential
137:used to synthesize
3095:mediated transport
3065:Membrane transport
2942:10.1093/jxb/eru495
2665:Diem K, Lenter C.
2646:www.chm.bris.ac.uk
2220:Reversal potential
2180:Concentration cell
2160:
2143:
2074:
1992:
1991:
1984:
1951:
1944:
1928:
1921:
1883:
1878:
1871:
1813:
1812:
1797:
1759:
1739:
1715:
1622:
1587:iron-sulfur center
1549:
1506:
1485:
1424:
1351:
1290:
926:
877:
698:
555:muscle contraction
537:Biological context
533:of that reaction.
501:concentration cell
344:electric potential
283:
79:electric potential
69:across a membrane.
62:across a membrane.
36:
3260:Electrophysiology
3237:
3236:
3233:
3232:
3083:Passive transport
3021:978-0-8053-7146-8
2739:(15): 5601–5607.
2676:978-3-9801244-0-9
2667:Scientific Tables
2312:978-1-4292-3414-6
2253:10.1002/bit.25492
2158:
2146:
2134:
2120:
2105:
2097:
2077:
2071:
2055:
2041:
2026:
2018:
1989:
1973:
1968:
1966:
1958:
1949:
1940:
1937:
1935:
1926:
1910:
1905:
1903:
1886:
1876:
1867:
1864:
1862:
1786:
1781:
1779:
1762:
1747:
1704:
1699:
1697:
1689:
1675:
1666:
1544:
1530:
1509:
1488:
1476:
1462:
1450:
1439:
1427:
1312:, which leads to
1294:bacteriorhodopsin
1280:Bacteriorhodopsin
1274:bacteriorhodopsin
1198:
1197:
822:
751:Gibbs free energy
662:potassium channel
602:(IMS); for every
527:chemical reaction
519:electrical energy
497:
496:
489:
471:
367:hydroelectric dam
281:
272:
230:
194:
117:. It appears in
46:, usually for an
42:is a gradient of
16:(Redirected from
3282:
3265:Membrane biology
3169:
3126:Active transport
3091:Simple diffusion
3058:
3051:
3044:
3035:
3025:
3003:
3002:
2974:
2963:
2962:
2944:
2935:(9): 2373–2400.
2920:
2914:
2913:
2903:
2879:
2873:
2872:
2862:
2845:(8–9): 892–913.
2830:
2819:
2818:
2812:
2807:
2805:
2797:
2781:
2775:
2774:
2764:
2724:
2718:
2717:
2692:
2681:
2680:
2662:
2656:
2655:
2653:
2652:
2638:
2632:
2631:
2629:
2628:
2613:
2607:
2606:
2604:
2603:
2592:
2586:
2585:
2575:
2565:
2541:
2535:
2534:
2492:
2483:
2477:
2476:
2458:
2449:(7): C491–C495.
2434:
2425:
2424:
2414:
2389:(6): 1361–1382.
2374:
2368:
2367:
2357:
2347:
2323:
2317:
2316:
2298:
2281:
2280:
2236:
2200:Electrodiffusion
2190:Action potential
2169:
2167:
2166:
2161:
2159:
2156:
2154:
2151:
2144:
2135:
2132:
2127:
2126:
2125:
2118:
2106:
2103:
2098:
2095:
2087:
2085:
2082:
2075:
2072:
2064:
2056:
2053:
2048:
2047:
2046:
2039:
2027:
2024:
2019:
2016:
2001:
1999:
1998:
1993:
1990:
1987:
1985:
1980:
1979:
1978:
1971:
1959:
1956:
1950:
1947:
1945:
1938:
1927:
1924:
1922:
1917:
1916:
1915:
1908:
1896:
1894:
1891:
1884:
1877:
1874:
1872:
1865:
1822:
1820:
1819:
1814:
1811:
1810:
1798:
1793:
1792:
1791:
1784:
1772:
1770:
1767:
1760:
1754:
1753:
1752:
1745:
1738:
1737:
1716:
1711:
1710:
1709:
1702:
1690:
1687:
1682:
1681:
1680:
1673:
1667:
1664:
1558:
1556:
1555:
1550:
1545:
1542:
1537:
1536:
1535:
1528:
1519:
1517:
1514:
1507:
1498:
1496:
1493:
1486:
1477:
1474:
1469:
1468:
1467:
1460:
1451:
1448:
1440:
1437:
1435:
1432:
1425:
1260:Proton gradients
1254:
1253:Medium dependent
1251:
1245:
1242:
1236:
1233:
1227:
1224:
1213:
1210:
933:
922:
913:Faraday constant
910:
906:
898:
890:
886:
884:
883:
878:
876:
875:
874:
828:
827:
823:
821:
820:
819:
803:
802:
801:
788:
730:
609:
515:electrochemistry
492:
485:
481:
478:
472:
470:
429:
397:
389:
357:
341:
330:Faraday constant
327:
322:
318:
308:
304:
292:
290:
289:
284:
282:
279:
274:
273:
265:
253:
248:
247:
232:
231:
223:
217:
216:
201:
200:
195:
187:
158:chemical species
131:proton gradients
111:potential energy
75:simple diffusion
21:
3290:
3289:
3285:
3284:
3283:
3281:
3280:
3279:
3240:
3239:
3238:
3229:
3215:
3158:
3120:
3077:
3067:
3062:
3022:
3009:
3006:
2976:
2975:
2966:
2922:
2921:
2917:
2881:
2880:
2876:
2832:
2831:
2822:
2808:
2798:
2783:
2782:
2778:
2726:
2725:
2721:
2715:
2694:
2693:
2684:
2677:
2664:
2663:
2659:
2650:
2648:
2640:
2639:
2635:
2626:
2624:
2615:
2614:
2610:
2601:
2599:
2594:
2593:
2589:
2543:
2542:
2538:
2490:
2485:
2484:
2480:
2436:
2435:
2428:
2376:
2375:
2371:
2325:
2324:
2320:
2313:
2300:
2299:
2284:
2238:
2237:
2233:
2229:
2224:
2175:
2117:
2038:
2007:
2006:
1970:
1907:
1854:
1853:
1851:
1847:
1843:
1835:
1783:
1744:
1701:
1672:
1658:
1657:
1647:
1614:
1607:
1603:
1598:
1584:
1576:
1570:
1565:travels to the
1564:
1527:
1459:
1403:
1402:
1400:
1388:
1384:
1380:
1376:
1363:
1343:
1282:
1262:
1257:
1252:
1248:
1243:
1239:
1234:
1230:
1225:
1216:
1211:
1202:
1181:
1152:
1089:
1075:
1057:
1042:
920:
911:represents the
908:
904:
896:
891:represents the
888:
844:
804:
789:
783:
755:
754:
728:
726:
690:
679:
659:
650:
607:
539:
511:
493:
482:
476:
473:
430:
428:
414:
398:
387:
355:
339:
325:
320:
316:
311:
306:
303:
297:
239:
208:
185:
177:
176:
154:electric charge
147:neural synapses
107:
28:
23:
22:
15:
12:
11:
5:
3288:
3286:
3278:
3277:
3275:Thermodynamics
3272:
3267:
3262:
3257:
3252:
3242:
3241:
3235:
3234:
3231:
3230:
3225:
3223:
3217:
3216:
3214:
3213:
3208:
3203:
3198:
3193:
3188:
3183:
3177:
3175:
3166:
3160:
3159:
3157:
3156:
3151:
3146:
3141:
3136:
3130:
3128:
3122:
3121:
3119:
3118:
3113:
3108:
3103:
3098:
3087:
3085:
3079:
3078:
3072:
3069:
3068:
3063:
3061:
3060:
3053:
3046:
3038:
3032:
3031:
3026:
3020:
3005:
3004:
2985:(4): 526–538.
2964:
2915:
2894:(3): 536–553.
2874:
2820:
2811:|journal=
2776:
2719:
2713:
2682:
2675:
2657:
2633:
2608:
2587:
2536:
2478:
2426:
2369:
2318:
2311:
2282:
2247:(3): 429–437.
2230:
2228:
2225:
2223:
2222:
2217:
2212:
2207:
2202:
2197:
2195:Cell potential
2192:
2187:
2182:
2176:
2174:
2171:
2150:
2141:
2138:
2130:
2124:
2115:
2112:
2109:
2101:
2093:
2090:
2081:
2070:
2067:
2062:
2059:
2051:
2045:
2036:
2033:
2030:
2022:
2014:
1983:
1977:
1965:
1962:
1954:
1943:
1934:
1931:
1920:
1914:
1902:
1899:
1890:
1881:
1870:
1861:
1849:
1845:
1841:
1833:
1809:
1806:
1803:
1796:
1790:
1778:
1775:
1766:
1757:
1751:
1742:
1736:
1733:
1730:
1727:
1724:
1721:
1714:
1708:
1696:
1693:
1685:
1679:
1670:
1645:
1613:
1610:
1605:
1601:
1596:
1582:
1574:
1568:
1562:
1548:
1540:
1534:
1525:
1522:
1513:
1504:
1501:
1492:
1483:
1480:
1472:
1466:
1457:
1454:
1446:
1443:
1431:
1422:
1419:
1416:
1413:
1410:
1398:
1386:
1382:
1378:
1374:
1361:
1342:
1339:
1281:
1278:
1261:
1258:
1256:
1255:
1246:
1237:
1228:
1214:
1199:
1196:
1195:
1192:
1189:
1187:
1185:
1183:
1178:
1175:
1171:
1170:
1168:
1166:
1164:
1162:
1160:
1157:
1154:
1150:
1145:
1144:
1142:
1140:
1138:
1135:
1132:
1129:
1126:
1120:
1119:
1116:
1113:
1110:
1107:
1104:
1101:
1095:
1094:
1091:
1086:
1083:
1080:
1077:
1072:
1069:
1063:
1062:
1059:
1054:
1051:
1049:
1047:
1044:
1039:
1033:
1032:
1029:
1026:
1023:
1020:
1017:
1014:
1011:
1005:
1004:
1001:
998:
995:
992:
989:
986:
983:
977:
976:
973:
970:
967:
963:
962:
957:
952:
947:
942:
937:
873:
870:
867:
864:
861:
858:
855:
852:
847:
843:
840:
837:
834:
831:
826:
818:
815:
812:
807:
800:
797:
792:
786:
780:
777:
774:
771:
768:
765:
762:
744:electric field
736:arithmetic sum
724:
689:
686:
677:
670:electric field
657:
648:
639:photosystem II
538:
535:
495:
494:
401:
399:
392:
386:
383:
348:
347:
337:
323:
314:
309:
299:
277:
271:
268:
262:
259:
256:
252:
246:
242:
238:
235:
229:
226:
220:
215:
211:
207:
204:
199:
193:
190:
184:
106:
103:
95:photosynthesis
71:
70:
63:
26:
24:
14:
13:
10:
9:
6:
4:
3:
2:
3287:
3276:
3273:
3271:
3268:
3266:
3263:
3261:
3258:
3256:
3253:
3251:
3248:
3247:
3245:
3228:
3227:Degranulation
3224:
3222:
3218:
3212:
3209:
3207:
3204:
3202:
3199:
3197:
3194:
3192:
3189:
3187:
3184:
3182:
3181:Efferocytosis
3179:
3178:
3176:
3174:
3170:
3167:
3165:
3161:
3155:
3152:
3150:
3147:
3145:
3142:
3140:
3137:
3135:
3132:
3131:
3129:
3127:
3123:
3117:
3114:
3112:
3109:
3107:
3104:
3102:
3099:
3096:
3092:
3089:
3088:
3086:
3084:
3080:
3076:
3070:
3066:
3059:
3054:
3052:
3047:
3045:
3040:
3039:
3036:
3030:
3027:
3023:
3017:
3013:
3008:
3007:
3000:
2996:
2992:
2988:
2984:
2980:
2973:
2971:
2969:
2965:
2960:
2956:
2952:
2948:
2943:
2938:
2934:
2930:
2926:
2919:
2916:
2911:
2907:
2902:
2897:
2893:
2889:
2885:
2878:
2875:
2870:
2866:
2861:
2856:
2852:
2848:
2844:
2840:
2836:
2829:
2827:
2825:
2821:
2816:
2803:
2795:
2791:
2787:
2780:
2777:
2772:
2768:
2763:
2758:
2754:
2750:
2746:
2742:
2738:
2734:
2730:
2723:
2720:
2716:
2714:9781461504276
2710:
2706:
2702:
2698:
2691:
2689:
2687:
2683:
2678:
2672:
2668:
2661:
2658:
2647:
2643:
2637:
2634:
2623:
2619:
2612:
2609:
2598:
2591:
2588:
2583:
2579:
2574:
2569:
2564:
2559:
2555:
2551:
2547:
2540:
2537:
2532:
2528:
2524:
2520:
2516:
2512:
2508:
2504:
2500:
2496:
2489:
2482:
2479:
2474:
2470:
2466:
2462:
2457:
2452:
2448:
2444:
2440:
2433:
2431:
2427:
2422:
2418:
2413:
2408:
2404:
2400:
2396:
2392:
2388:
2384:
2380:
2373:
2370:
2365:
2361:
2356:
2351:
2346:
2341:
2337:
2333:
2329:
2322:
2319:
2314:
2308:
2304:
2297:
2295:
2293:
2291:
2289:
2287:
2283:
2278:
2274:
2270:
2266:
2262:
2258:
2254:
2250:
2246:
2242:
2235:
2232:
2226:
2221:
2218:
2216:
2213:
2211:
2208:
2206:
2205:Galvanic cell
2203:
2201:
2198:
2196:
2193:
2191:
2188:
2186:
2183:
2181:
2178:
2177:
2172:
2170:
2148:
2139:
2122:
2113:
2110:
2091:
2079:
2068:
2065:
2060:
2043:
2034:
2031:
2012:
2002:
1981:
1975:
1963:
1960:
1952:
1941:
1932:
1918:
1912:
1900:
1897:
1888:
1879:
1868:
1859:
1839:
1831:
1827:
1823:
1794:
1788:
1776:
1773:
1764:
1755:
1749:
1712:
1706:
1694:
1691:
1683:
1677:
1668:
1655:
1652:from reduced
1651:
1643:
1639:
1635:
1631:
1627:
1618:
1611:
1609:
1599:
1592:
1588:
1580:
1572:
1559:
1532:
1523:
1520:
1511:
1502:
1499:
1490:
1464:
1455:
1452:
1444:
1441:
1429:
1420:
1417:
1414:
1411:
1408:
1396:
1392:
1372:
1371:plastoquinone
1368:
1364:
1356:
1347:
1340:
1338:
1336:
1332:
1328:
1323:
1319:
1315:
1314:isomerization
1311:
1307:
1303:
1300:is through a
1299:
1295:
1286:
1279:
1277:
1275:
1271:
1268:rotation, or
1267:
1259:
1250:
1247:
1241:
1238:
1232:
1229:
1223:
1221:
1219:
1215:
1209:
1207:
1205:
1201:
1193:
1190:
1188:
1186:
1184:
1179:
1176:
1173:
1172:
1169:
1167:
1165:
1163:
1161:
1158:
1155:
1153:
1147:
1146:
1143:
1141:
1139:
1136:
1133:
1130:
1127:
1125:
1122:
1121:
1117:
1111:
1108:
1105:
1102:
1100:
1097:
1096:
1092:
1087:
1084:
1081:
1078:
1073:
1070:
1068:
1065:
1064:
1060:
1055:
1052:
1050:
1048:
1045:
1040:
1038:
1035:
1034:
1030:
1027:
1024:
1021:
1018:
1015:
1012:
1010:
1007:
1006:
1002:
999:
996:
993:
990:
987:
984:
982:
979:
978:
974:
971:
968:
965:
964:
961:
956:
951:
950:S. cerevisiae
946:
941:
934:
930:
924:
916:
914:
902:
894:
845:
838:
835:
829:
824:
805:
790:
784:
778:
775:
772:
769:
766:
763:
752:
749:
745:
741:
737:
732:
723:
718:
714:
709:
707:
703:
694:
688:Ion gradients
687:
685:
683:
675:
671:
667:
663:
654:
652:
644:
643:plastiquinone
640:
635:
632:
628:
623:
621:
617:
613:
605:
604:electron pair
601:
597:
593:
589:
585:
582:
581:mitochondrial
577:
575:
571:
567:
563:
560:
556:
552:
548:
544:
543:cell membrane
536:
534:
532:
528:
524:
520:
516:
510:
506:
502:
491:
488:
480:
477:December 2023
469:
466:
462:
459:
455:
452:
448:
445:
441:
438: –
437:
433:
432:Find sources:
426:
422:
418:
412:
411:
407:
402:This section
400:
396:
391:
390:
384:
382:
380:
376:
372:
368:
364:
359:
353:
345:
338:
335:
331:
324:
317:
310:
302:
296:
295:
294:
266:
257:
244:
240:
236:
224:
213:
209:
202:
197:
188:
174:
171:
170:thermodynamic
166:
164:
159:
155:
150:
148:
144:
140:
136:
132:
128:
124:
120:
116:
112:
104:
102:
100:
96:
92:
88:
83:
80:
76:
68:
64:
61:
57:
56:
55:
53:
49:
45:
41:
32:
19:
3211:Transcytosis
3191:Phagocytosis
3011:
2982:
2978:
2932:
2928:
2918:
2891:
2887:
2877:
2842:
2838:
2802:cite journal
2779:
2736:
2732:
2722:
2696:
2666:
2660:
2649:. Retrieved
2645:
2636:
2625:. Retrieved
2621:
2611:
2600:. Retrieved
2590:
2553:
2549:
2539:
2501:(1): 19–26.
2498:
2494:
2481:
2446:
2442:
2386:
2382:
2372:
2335:
2331:
2321:
2302:
2244:
2240:
2234:
2096:cytochrome c
2017:cytochrome c
2003:
1939:cytochrome c
1866:cytochrome c
1838:cytochrome c
1824:
1623:
1591:cytochrome f
1579:plastocyanin
1567:cytochrome b
1560:
1393:through the
1367:energy level
1365:to a higher
1352:
1331:deprotonated
1327:reprotonated
1291:
1263:
1249:
1240:
1231:
917:
893:gas constant
733:
721:
710:
699:
655:
647:cytochrome b
624:
612:ATP synthase
578:
553:conduction,
540:
530:
512:
483:
474:
464:
457:
450:
443:
431:
415:Please help
403:
360:
351:
349:
342:, the local
312:
300:
167:
151:
134:
127:chloroplasts
123:mitochondria
108:
84:
72:
39:
37:
3201:Potocytosis
3196:Pinocytosis
3173:Endocytosis
1826:Complex III
1318:Schiff base
1302:proton pump
921:+70 mV
899:represents
729:−60 mV
713:Na-K-ATPase
708:transport.
608:200 mV
133:generate a
3244:Categories
3221:Exocytosis
3144:Antiporter
2651:2019-06-07
2627:2019-06-07
2602:2019-06-07
2227:References
1638:ubiquinone
1310:wavelength
1270:metabolite
1194:8.1 - 8.2
1191:7.2 - 7.8
1076:1.3 - 1.5
1074:2.2 - 2.6
1043:0.5 - 0.8
945:Squid axon
929:millimolar
717:hydrolysis
684:gradient.
674:antiporter
499:See also:
447:newspapers
375:electrodes
141:, and the
89:and other
3139:Symporter
3134:Uniporter
2951:0022-0957
2753:0021-9193
2515:0031-6768
2465:0363-6143
2403:1469-7793
2261:1097-0290
2089:⟶
1982:⏟
1942:⏟
1930:⟶
1919:⏟
1869:⏟
1795:⏟
1741:⟶
1713:⏟
1650:electrons
1642:ubiquinol
1634:reduction
1630:catalyzes
1626:complex I
1571:f complex
1482:⟶
1415:ν
1308:of 568nm
1266:flagellar
1177:7.1 - 7.3
1134:300 - 400
1115:10 - 200
1079:10 - 3×10
1058:0.01 - 1
1056:30 - 100
985:100 - 140
960:Sea water
779:
761:Δ
727:of about
651:f complex
631:thylakoid
616:phosphate
566:sensation
562:secretion
404:does not
385:Chemistry
365:across a
352:generated
270:→
258:φ
255:∇
228:→
210:μ
206:∇
192:¯
189:μ
183:∇
115:conserved
87:batteries
3116:Carriers
3111:Channels
3093:(or non-
2999:23867107
2959:25540437
2910:24918316
2869:23507617
2771:17545292
2582:26973667
2531:18133149
2523:22526460
2473:26791490
2421:25772291
2364:25705194
2269:25384602
2173:See also
2104:oxidized
1875:oxidized
1848:at the Q
1840:at the Q
1640:(UQ) to
1320:(SB) in
1292:The way
1109:40 - 150
1000:30 - 300
725:membrane
641:(PSII),
570:relative
363:pressure
105:Overview
52:membrane
3164:Cytosis
3106:Osmosis
3012:Biology
2860:3714358
2762:1951819
2573:4770017
2556:: 218.
2412:4376416
2355:4319557
2277:2598635
2025:reduced
1948:reduced
1830:Q-cycle
1335:cytosol
1322:retinal
1316:of the
1306:photons
1298:Archaea
1244:Ionised
1046:1 - 1.5
994:10 - 20
955:E. coli
740:osmosis
706:passive
598:to the
590:is the
559:hormone
545:drives
517:, when
461:scholar
425:removed
410:sources
3018:
2997:
2957:
2949:
2908:
2867:
2857:
2769:
2759:
2751:
2711:
2673:
2580:
2570:
2529:
2521:
2513:
2471:
2463:
2419:
2409:
2401:
2362:
2352:
2338:: 29.
2309:
2275:
2267:
2259:
2054:matrix
1925:matrix
1595:heme b
1475:stroma
975:Blood
969:Blood
940:Mammal
887:where
702:active
666:stroma
645:, and
596:matrix
564:, and
507:, and
463:
456:
449:
442:
434:
336:basis)
145:helps
67:charge
2527:S2CID
2491:(PDF)
2273:S2CID
1628:(CI)
1543:lumen
1391:water
1355:light
1235:Total
1212:Bound
972:Cell
966:Cell
748:molar
634:lumen
551:nerve
549:like
525:of a
468:JSTOR
454:books
293:with
3016:ISBN
2995:PMID
2955:PMID
2947:ISSN
2906:PMID
2892:1850
2865:PMID
2843:1827
2815:help
2767:PMID
2749:ISSN
2709:ISBN
2671:ISBN
2578:PMID
2519:PMID
2511:ISSN
2469:PMID
2461:ISSN
2417:PMID
2399:ISSN
2360:PMID
2307:ISBN
2265:PMID
2257:ISSN
1665:NADH
1644:(UQH
1632:the
1226:Free
1137:5-10
1118:500
1112:560
1031:500
1013:5-15
936:Ion
676:(KEA
660:, a
618:and
440:news
408:any
406:cite
334:mole
125:and
97:and
2987:doi
2937:doi
2896:doi
2855:PMC
2847:doi
2790:doi
2757:PMC
2741:doi
2737:189
2701:doi
2568:PMC
2558:doi
2503:doi
2499:464
2451:doi
2447:310
2407:PMC
2391:doi
2387:593
2350:PMC
2340:doi
2249:doi
2245:112
2133:IMS
1988:IMS
1885:UQH
1761:UQH
1746:NAD
1636:of
1577:to
1508:PQH
1387:680
1373:(PQ
1362:680
1149:HCO
1128:138
1106:110
1093:10
1090:10
1061:50
1041:10
1022:440
1016:145
1003:10
997:300
991:400
988:4-5
738:of
704:or
620:ADP
419:by
373:or
358:.
163:ion
139:ATP
48:ion
38:An
3246::
2993:.
2983:23
2981:.
2967:^
2953:.
2945:.
2933:66
2931:.
2927:.
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2886:.
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2804:}}
2800:{{
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2271:.
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2255:.
2243:.
1957:UQ
1688:UQ
1449:PQ
1217:^
1203:^
1174:pH
1159:29
1156:12
1099:Cl
1088:3
1082:10
1071:10
1067:Ca
1053:50
1037:Mg
1028:10
1025:30
1019:50
1009:Na
931:)
915:.
903:,
895:,
776:ln
731:.
682:pH
622:.
557:,
503:,
328:,
129:,
93:,
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3050:t
3043:v
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3001:.
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2100:(
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2080:2
2076:O
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2040:H
2035:4
2032:+
2029:)
2021:(
2013:2
1976:+
1972:H
1964:4
1961:+
1953:+
1933:2
1913:+
1909:H
1901:2
1898:+
1889:2
1880:+
1860:2
1850:i
1846:2
1842:o
1834:2
1808:S
1805:M
1802:I
1789:+
1785:H
1777:4
1774:+
1765:2
1756:+
1750:+
1735:x
1732:i
1729:r
1726:t
1723:a
1720:m
1707:+
1703:H
1695:4
1692:+
1684:+
1678:+
1674:H
1669:+
1646:2
1606:2
1602:H
1597:L
1583:2
1575:2
1569:6
1563:2
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1539:(
1533:+
1529:H
1524:4
1521:+
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1500:+
1491:2
1487:O
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1471:(
1465:+
1461:H
1456:4
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1438:O
1430:2
1426:H
1421:2
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1412:h
1409:4
1399:2
1383:2
1379:B
1375:A
1360:P
1151:3
1131:9
1103:4
1085:2
981:K
909:F
905:z
897:T
889:R
872:e
869:n
866:a
863:r
860:b
857:m
854:e
851:m
846:V
842:)
839:z
836:F
833:(
830:+
825:)
817:t
814:u
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806:c
799:n
796:i
791:c
785:(
773:T
770:R
767:=
764:G
722:V
678:3
658:3
649:6
490:)
484:(
479:)
475:(
465:·
458:·
451:·
444:·
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413:.
356:φ
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340:φ
326:F
321:i
315:i
313:z
307:i
301:i
298:μ
280:,
276:)
267:r
261:(
251:F
245:i
241:z
237:+
234:)
225:r
219:(
214:i
203:=
198:i
175::
20:)
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