{"id":278,"date":"2019-04-11T09:55:15","date_gmt":"2019-04-11T09:55:15","guid":{"rendered":"http:\/\/biocp04.epgpbooks.inflibnet.ac.in\/?post_type=chapter&#038;p=278"},"modified":"2019-04-11T09:59:08","modified_gmt":"2019-04-11T09:59:08","slug":"proton-motive-force","status":"publish","type":"chapter","link":"https:\/\/ebooks.inflibnet.ac.in\/biocp04\/chapter\/proton-motive-force\/","title":{"rendered":"Proton motive force"},"content":{"raw":"<div><span style=\"float: right\"><a href=\"https:\/\/youtu.be\/J660Uo1f0yU\" target=\"_blank\" rel=\"noopener\"><img src=\"http:\/\/epgpbooks.inflibnet.ac.in\/wp-content\/uploads\/2018\/11\/download.png\" alt=\"epgp books\" width=\"75px\" height=\"75px;\" \/><\/a>\r\n<\/span><\/div>\r\n<strong>\u00a0 \u00a0 <\/strong>\r\n\r\n&nbsp;\r\n\r\n&nbsp;\r\n\r\n&nbsp;\r\n\r\n<strong>PROTON MOTIVE FORCE<\/strong>\r\n\r\n<strong>Objectives<\/strong>\r\n<ol>\r\n \t<li>To understand the function of Proton Motive Force in production of ATP.<\/li>\r\n \t<li>To examine the importance of a proton gradients power the synthesis of ATP.<\/li>\r\n \t<li>To examine the importance of shuttles for aerobic oxidation of cytosolic NADH.<\/li>\r\n \t<li>To understand the cellular respiration which is regulated by the Need for ATP.<\/li>\r\n<\/ol>\r\n<strong>\u00a0 \u00a0 <\/strong>\r\n\r\n<strong>Introduction<\/strong>\r\n<ul>\r\n \t<li style=\"text-align: justify\">A proton-motive force is \"energy that is generated by the transfer of protons or electrons across an energy-transducing membrane and can be used for chemical, osmotic or mechanical work.\"<\/li>\r\n<\/ul>\r\n<ul>\r\n \t<li style=\"text-align: justify\">It can be \"generated by a variety of phenomena including the operation of an electron transport chain, illumination of a purple membrane, and the hydrolysis of ATP by a proton ATPase.\"<\/li>\r\n<\/ul>\r\n<img class=\"aligncenter size-full wp-image-279\" src=\"http:\/\/biocp04.epgpbooks.inflibnet.ac.in\/wp-content\/uploads\/sites\/215\/2019\/04\/1-130.png\" alt=\"\" width=\"470\" height=\"78\" \/>\r\n\r\n&nbsp;\r\n\r\n<strong>Proton Gradient<\/strong>\r\n\r\n&nbsp;\r\n\r\n&nbsp;\r\n\r\n<img class=\"aligncenter size-full wp-image-280\" src=\"http:\/\/biocp04.epgpbooks.inflibnet.ac.in\/wp-content\/uploads\/sites\/215\/2019\/04\/1-131.png\" alt=\"\" width=\"545\" height=\"341\" \/>\r\n<h2 style=\"text-align: center\"><strong>Figure-1. Proton motif force<\/strong><\/h2>\r\n<h2 style=\"text-align: center\"><strong>( Adapted from- http:\/\/classroom.sdmesa.edu\/eschmid\/Lectur46.gif)<\/strong><\/h2>\r\n&nbsp;\r\n\r\n<strong>Why Proton Gradients Are Necessary<\/strong>\r\n<ul>\r\n \t<li style=\"text-align: justify\">The reason proton gradients are required boils back down to chemistry. In other words, to convert carbon dioxide into organic molecules, life attaches hydrogen atoms to CO2<\/li>\r\n<\/ul>\r\n<ul style=\"text-align: justify\">\r\n \t<li>This pathway is exothermic (releasing energy that can be captured as ATP) right through to pyruvate, one of the central molecules in cell metabolism<\/li>\r\n<\/ul>\r\n<ul style=\"text-align: justify\">\r\n \t<li>But there's a problem, pointed out by William Martin in collaboration with Russell. All cells that use the acetyl-CoA pathway today depend on proton gradients. None of them can grow by fermentation<\/li>\r\n<\/ul>\r\n<ul>\r\n \t<li style=\"text-align: justify\">Why not? Because CO2 is a stable molecule and does not react easily, even with hydrogen - even when thermodynamics says it should react. CO2 is a bit like oxygen in this respect: Once it starts to react, it's not easily stopped. But a fire<span style=\"text-align: initial;font-size: 1em\"> needs a spark to get it going, and so, too, does CO2. Cells need the equivalent of a spark to get CO2 to react, and for cells, that spark is ATP<\/span><\/li>\r\n<\/ul>\r\n<ul>\r\n \t<li style=\"text-align: justify\">The problem is that the reaction of CO2 with H2 releases energy, but not a lot \u2014 only enough to make 1 ATP. That means cells have to spend 1 ATP to gain 1 ATP, so there's no net gain. If there's no gain, there's no growth; no growth, no life<\/li>\r\n<\/ul>\r\n<img class=\"aligncenter size-full wp-image-281\" src=\"http:\/\/biocp04.epgpbooks.inflibnet.ac.in\/wp-content\/uploads\/sites\/215\/2019\/04\/1-132.png\" alt=\"\" width=\"488\" height=\"161\" \/>\r\n<ul>\r\n \t<li style=\"text-align: justify\">Gradients break that cycle. It's not quite true to say that the reaction of CO 2 with H2 releases enough energy to make 1 ATP: it's actually enough to make 1.5 ATPs. But of course there's no such thing as 1.5 ATPs, at least not by stoichiometric chemistry; so the spare energy from the reaction, as chemistry, is lost<\/li>\r\n<\/ul>\r\n<ul style=\"text-align: justify\">\r\n \t<li>But that doesn't happen with a gradient. In principle, a reaction can be repeated over and over again, just to pump a proton over a membrane<\/li>\r\n<\/ul>\r\n<ul style=\"text-align: justify\">\r\n \t<li>When enough protons have accumulated, the proton motif force powers the formation of ATP<\/li>\r\n<\/ul>\r\n<ul>\r\n \t<li style=\"text-align: justify\">So a gradient allows cells to save up protons as \"loose change\", and that makes all the difference in the world \u2014 the difference between growth and no growth, life and no life.<\/li>\r\n<\/ul>\r\n<img class=\"aligncenter size-full wp-image-282\" src=\"http:\/\/biocp04.epgpbooks.inflibnet.ac.in\/wp-content\/uploads\/sites\/215\/2019\/04\/1-133.png\" alt=\"\" width=\"603\" height=\"234\" \/>\r\n\r\n&nbsp;\r\n\r\n<strong>A proton gradient powers the synthesis of ATP<\/strong>\r\n<ul>\r\n \t<li style=\"text-align: justify\">The overall process of ATP generation via the harnessing of a proton motif force is called chemiosmosis<\/li>\r\n \t<li style=\"text-align: justify\">The Chemiosmotic Theory Proposed by Peter Mitchell in the 1960\u2019s<\/li>\r\n \t<li style=\"text-align: justify\">The proton gradient generated by the oxidation of NADH and FADH2 is called the proton-motif force.The proton-motif force powers the synthesis of ATP<\/li>\r\n<\/ul>\r\n<strong>\u00a0 Coupled nature of respiration in mitochondria<\/strong>\r\n\r\n&nbsp;\r\n\r\n<img class=\"aligncenter size-full wp-image-283\" src=\"http:\/\/biocp04.epgpbooks.inflibnet.ac.in\/wp-content\/uploads\/sites\/215\/2019\/04\/1-134.png\" alt=\"\" width=\"538\" height=\"240\" \/>\r\n<p style=\"text-align: center\">(a) O2 consumed only with ADP, excess Pi.<\/p>\r\n<p style=\"text-align: center\">(b) (+) Uncoupler DNP (O2 consumed without ADP).<\/p>\r\n&nbsp;\r\n\r\n<strong>(A) <\/strong><strong>O<\/strong><strong>2<\/strong><strong> consumed only with ADP, excess Pi<\/strong>\r\n<ul>\r\n \t<li>Oxidation of substrates is coupled to the phosphorylation of ADP<\/li>\r\n \t<li>Respiration (consumption of oxygen) proceeds only when ADP is present<\/li>\r\n \t<li>The amount of O 2 consumed depends upon the amount of ADP added<\/li>\r\n<\/ul>\r\n&nbsp;\r\n\r\n<strong>(B) <\/strong><strong>(+) Uncoupler DNP (O 2 consumed without ADP)<\/strong>\r\n<ul>\r\n \t<li>Uncouplers stimulate the oxidation of substrates in the absence of ADP.<\/li>\r\n \t<li>Uncouplers are lipid-soluble weak acids.<\/li>\r\n \t<li>Both acidic and basic forms can cross the inner mitochondrial membrane.<\/li>\r\n \t<li>Uncouplers deplete any proton gradient by transporting protons across the membrane.<\/li>\r\n<\/ul>\r\n<img class=\"aligncenter size-full wp-image-284\" src=\"http:\/\/biocp04.epgpbooks.inflibnet.ac.in\/wp-content\/uploads\/sites\/215\/2019\/04\/1-135.png\" alt=\"\" width=\"662\" height=\"175\" \/>\r\n<h2 style=\"text-align: center\"><strong>2,4-Dinitrophenol: an uncoupler<\/strong><\/h2>\r\n&nbsp;\r\n\r\n<strong>Mitchell\u2019s postulates for Chemiosmotic theory<\/strong>\r\n<ul>\r\n \t<li style=\"text-align: justify\">Intact inner mitochondrial membrane is required (to maintain a proton gradient)<\/li>\r\n \t<li style=\"text-align: justify\">Electron transport through the ETC generates a proton gradient (pumps H + from the matrix to the intermembrane space)<\/li>\r\n \t<li style=\"text-align: justify\">The membrane-spanning enzyme, ATP synthase, catalyzes the phosphorylation of ADP in a reaction driven by movement of H + across the inner membrane into the matrix<\/li>\r\n<\/ul>\r\n&nbsp;\r\n\r\n<strong>Shuttles are Necessary for Aerobic Oxidation of Cytosolic NADH<\/strong>\r\n<ul>\r\n \t<li style=\"text-align: justify\">Cytosolic NADH must enter the mitochondria to fuel oxidative phosphorylation, but NADH and NAD+ cannot diffuse across the inner mitochondrial membrane<\/li>\r\n \t<li style=\"text-align: justify\">Two shuttle systems for reducing equivalents:<\/li>\r\n<\/ul>\r\n<ol style=\"text-align: justify\">\r\n \t<li>Glycerol phosphate shuttle: insect flight muscles<\/li>\r\n \t<li>Malate-aspartate shuttle: predominant in liver and other mammalian tissues<\/li>\r\n<\/ol>\r\n<strong>1. Glycerol phosphate shuttle<\/strong>\r\n<ul>\r\n \t<li style=\"text-align: justify\">The glycerol-3-phosphate shuttle is a mechanism that regenerates NAD+ from NADH, a by-product of glycolysis. Its importance in transporting reducing equivalents is secondary to the malate-aspartate shuttle<\/li>\r\n<\/ul>\r\n<strong><em>\u00a0 \u00a0 \u00a0Mechanism:<\/em><\/strong>\r\n<ul>\r\n \t<li style=\"text-align: justify\">In this\u00a0 shuttle,\u00a0 the\u00a0 enzyme glycerol-3-phosphate\u00a0 dehydrogenase 1\u00a0 (GPDH-C) converts dihydroxyacetone phosphate <span style=\"text-align: initial;font-size: 1em\">(2) to <\/span>glycerol 3-phosphate <span style=\"text-align: initial;font-size: 1em\">(1) by oxidizing one molecule of NADH to NAD+<\/span><\/li>\r\n<\/ul>\r\n<ul style=\"text-align: justify\">\r\n \t<li>Glycerol-3-phosphate gets converted back to dihydroxyacetone phosphate by an inner membrane-bound mitochondrial glycerol-3-phosphate dehydrogenase 2 (GPDH-M), this time reducing one molecule of enzyme-bound flavin adenine dinucleotide <span style=\"text-align: initial;font-size: 1em\">(FAD) to FADH2. FADH2 then reduces <\/span>coenzyme Q <span style=\"text-align: initial;font-size: 1em\">(ubiquinone to ubiquinol) which enters into <\/span>oxidative phosphorylation. <span style=\"text-align: initial;font-size: 1em\">This reaction is irreversible.<\/span><\/li>\r\n<\/ul>\r\n<ul>\r\n \t<li style=\"text-align: justify\">The glycerol-3-phosphate shuttle allows the NADH synthesized in the cytosol by glycolysis to contribute to the oxidative phosphorylation <span style=\"text-align: initial;font-size: 1em\">pathway in the <\/span>mitochondria <span style=\"text-align: initial;font-size: 1em\">to generate ATP. It has been found in animals, fungi, and plants.<\/span><\/li>\r\n<\/ul>\r\n<img class=\"aligncenter size-full wp-image-285\" src=\"http:\/\/biocp04.epgpbooks.inflibnet.ac.in\/wp-content\/uploads\/sites\/215\/2019\/04\/1-136.png\" alt=\"\" width=\"523\" height=\"336\" \/>\r\n<h2 style=\"text-align: center\"><strong>Figure -2 Glycerol phosphate shuttle<\/strong><\/h2>\r\n<h2 style=\"text-align: center\"><strong>(Adapted from: http:\/\/oregonstate.edu\/instruct\/bb451\/winter14\/stryer7\/CH18\/figure_18_34.jpg))<\/strong><\/h2>\r\n&nbsp;\r\n\r\n<strong>Malate-aspartate shuttle<\/strong>\r\n<ul>\r\n \t<li style=\"text-align: justify\">The malate-aspartate shuttle is a biochemical system for translocating electrons\u00a0produced\u00a0during glycolysis across\u00a0the semipermeable inner\u00a0membrane of the mitochondrion for oxidative phosphorylation in eukaryotes.\u00a0These electrons\u00a0enter in\u00a0the electron transport\u00a0chain of\u00a0the mitochondria via reduction\u00a0equivalents to generate ATP.<\/li>\r\n<\/ul>\r\n&nbsp;\r\n\r\n&nbsp;\r\n\r\n<strong><em>Mechanism:<\/em><\/strong>\r\n<ul>\r\n \t<li style=\"text-align: justify\">The primary enzyme in the malate-aspartate shuttle is malate dehydrogenase.<\/li>\r\n<\/ul>\r\n<ul style=\"text-align: justify\">\r\n \t<li>First, in the cytosol, malate dehydrogenase catalyses the reaction of oxaloacetate and NADH to produce malate and NAD+.<\/li>\r\n<\/ul>\r\n<ul style=\"text-align: justify\">\r\n \t<li>Once malate is formed, the first antiporter (malate-alpha-ketoglutarate)imports the malate from the cytosol into the mitochondrial matrix and also exports alpha-ketoglutarate from the matrix into the cytosol simultaneously.<\/li>\r\n<\/ul>\r\n<ul style=\"text-align: justify\">\r\n \t<li>Oxaloacetate is then transformed into aspartate by mitochondrial aspartate aminotransferase. Since aspartate is an amino acid, an amino radical needs to be added to the oxaloacetate.<\/li>\r\n<\/ul>\r\n<ul>\r\n \t<li style=\"text-align: justify\">The second antiporter (the glutamate-aspartate antiporter) imports glutamate from the cytosol into the matrix and exports aspartate from the matrix to the cytosol.<\/li>\r\n<\/ul>\r\n<ul>\r\n \t<li style=\"text-align: justify\">The net effect of the malate-aspartate shuttle is purely redox: NADH in the cytosol is oxidized to NAD+, and NAD+ in the matrix is reduced to NADH.<\/li>\r\n<\/ul>\r\n<ul style=\"text-align: justify\">\r\n \t<li>Since the malate-aspartate shuttle regenerates NADH inside the mitochondrial matrix, it is capable of maximizing the number of ATPs produced in glycolysis (3\/NADH), ultimately resulting in a net gain of 38 ATP molecules per molecule of glucose metabolized.<\/li>\r\n<\/ul>\r\n<p style=\"text-align: justify\"><img class=\"aligncenter size-full wp-image-286\" src=\"http:\/\/biocp04.epgpbooks.inflibnet.ac.in\/wp-content\/uploads\/sites\/215\/2019\/04\/1-137.png\" alt=\"\" width=\"465\" height=\"244\" \/><\/p>\r\n\r\n<h2 style=\"text-align: center\"><strong>Figure- 3 Malate-aspartate shuttle<\/strong><\/h2>\r\n<h2 style=\"text-align: center\"><strong>( Adapted from- http:\/\/oregonstate.edu\/instruct\/bb451\/winter14\/stryer7\/CH18\/figure_18_35.jpg)<\/strong><\/h2>\r\n&nbsp;\r\n\r\n<strong>Regulation of Proton motif force<\/strong>\r\n<ul>\r\n \t<li>Protonmotif force ( D p) is the energy of the proton concentration gradient<\/li>\r\n<\/ul>\r\n<img class=\"aligncenter size-full wp-image-287\" src=\"http:\/\/biocp04.epgpbooks.inflibnet.ac.in\/wp-content\/uploads\/sites\/215\/2019\/04\/1-138.png\" alt=\"\" width=\"470\" height=\"297\" \/>\r\n<h2 style=\"text-align: center\"><strong>Figure \u2013 4. Proton concentration gradient in chemical and electrical condition.<\/strong><\/h2>\r\n<h2 style=\"text-align: center\"><strong>(Adapted from - http:\/\/www.ccrc.uga.edu\/~rcarlson\/bcmb3100\/Chap21)<\/strong><\/h2>\r\n&nbsp;\r\n\r\n<strong>ATP synthase<\/strong>\r\n<ul>\r\n \t<li style=\"text-align: justify\">There are 3 active sites, one in each b subunit<\/li>\r\n \t<li style=\"text-align: justify\">The c-e-g unit forms a \u201crotor\u201d<\/li>\r\n \t<li style=\"text-align: justify\">Rotation of the g subunit inside the a3b3 hexamer causes domain movements in the b-subunits, opening and closing the active sites<\/li>\r\n \t<li style=\"text-align: justify\">The a-b-g-a3b3 unit is the \u201cstator\u201d (the Fo channel is attached to a3b3 by the ab-d arm)<\/li>\r\n<\/ul>\r\n<img class=\"aligncenter size-full wp-image-288\" src=\"http:\/\/biocp04.epgpbooks.inflibnet.ac.in\/wp-content\/uploads\/sites\/215\/2019\/04\/1-139.png\" alt=\"\" width=\"375\" height=\"286\" \/>\r\n<h2 style=\"text-align: center\"><strong>Figure \u2013 5. ATP synthase.<\/strong><\/h2>\r\n<h2 style=\"text-align: center\"><strong>(Adapted<\/strong> <strong>from https:\/\/s3.amazonaws.com\/classconnection\/281\/flashcards\/2541281\/png\/session_15_ _oxidative_phosphorylation_-_google_slides_(1)-14C6815A20153228B19.png)<\/strong><\/h2>\r\n<h2 style=\"text-align: center\"><\/h2>\r\n<ul>\r\n \t<li>F0F1 ATP Synthase uses the proton gradient energy for the synthesis of ATP<\/li>\r\n \t<li>An F-type ATPase which generates ATP<\/li>\r\n \t<li>Composed of a \u201cknob-and-stalk\u201d structure<\/li>\r\n \t<li>F1 (knob) contains the catalytic subunits<\/li>\r\n \t<li>F0 (stalk) has a proton channel which spans the membrane.<\/li>\r\n \t<li>Passage of protons through the F0 (stalk) into the matrix is coupled to ATP formation<\/li>\r\n \t<li>Estimated passage of 3 H+ \/ ATP synthesized<\/li>\r\n<\/ul>\r\n<ul>\r\n \t<li style=\"text-align: justify\">Passage of protons through the F0 channel causes the rotor to spin in one direction and the stator to spin in the opposite direction<\/li>\r\n<\/ul>\r\n<img class=\"aligncenter size-full wp-image-289\" src=\"http:\/\/biocp04.epgpbooks.inflibnet.ac.in\/wp-content\/uploads\/sites\/215\/2019\/04\/1-140.png\" alt=\"\" width=\"485\" height=\"151\" \/>\r\n<p style=\"text-align: center\">Three protons = one complete rotation = one ATP<\/p>\r\n&nbsp;\r\n\r\n&nbsp;\r\n\r\n<img class=\"aligncenter size-full wp-image-290\" src=\"http:\/\/biocp04.epgpbooks.inflibnet.ac.in\/wp-content\/uploads\/sites\/215\/2019\/04\/1-141.png\" alt=\"\" width=\"475\" height=\"162\" \/>\r\n<h2 style=\"text-align: center\"><strong>Figure \u2013 6 Conformational changes in ATP synthase.<\/strong><\/h2>\r\n<h2 style=\"text-align: center\"><strong>\u00a0 1.\u00a0 ADP, Pi bind to an open site.<\/strong><\/h2>\r\n<h2 style=\"text-align: center\"><strong>2. Inward passage of protons, conformation change, ATP synthesis from ADP and Pi.<\/strong><\/h2>\r\n<h2 style=\"text-align: center\"><strong>3. ATP released from <\/strong>open<strong> site, ADP and Pi form ATP in the tight site.<\/strong><\/h2>\r\n<h2 style=\"text-align: center\"><strong>(Adapted from http:\/\/1.bp.blogspot.com\/_DZH2cmCoois\/R2hL3wt_O6I\/AAAAAAAAEHU\/7WNKoQjakDk\/s400\/tmp.jpg)<\/strong><\/h2>\r\n&nbsp;\r\n\r\n<strong>Proton flow around the \u201cc\u201d ring<\/strong>\r\n<ul>\r\n \t<li style=\"text-align: justify\">The number of c rings determines the number of protons required to synthesize a molecule of ATP.<\/li>\r\n \t<li style=\"text-align: justify\">The c ring of vertebrates consist of 8 subunits, making vertebrate ATP synthase the most efficient known.<\/li>\r\n<\/ul>\r\n<p style=\"text-align: justify\"><img class=\"aligncenter size-full wp-image-291\" src=\"http:\/\/biocp04.epgpbooks.inflibnet.ac.in\/wp-content\/uploads\/sites\/215\/2019\/04\/1-142.png\" alt=\"\" width=\"447\" height=\"295\" \/><\/p>\r\n\r\n<h2 style=\"text-align: center\"><strong>Figure \u2013 7 Proton flow around the C ring.<\/strong><\/h2>\r\n<h2 style=\"text-align: center\"><strong>(Adapted from-\u00a0<\/strong><strong>https:\/\/s3.amazonaws.com\/classconnection\/281\/flashcards\/2541281\/png\/session_15_- _oxidative_phosphorylation_-_google_slides_(5)-14C68FE9F127B4A4690.png)<\/strong><\/h2>\r\n&nbsp;\r\n\r\n<strong>SUMMARY<\/strong>\r\n<ul>\r\n \t<li style=\"text-align: justify\">It is generally accepted that the chemistry of water was the most crucial determinant in shaping life on earth. Among the more important chemical features of water is its dissociation into protons and hydroxyl ions.<\/li>\r\n<\/ul>\r\n<ul style=\"text-align: justify\">\r\n \t<li>The presence of relatively high proton concentrations in the ambient solution resulted in the evolution of proton pumps during the dawn of life on earth. These proton pumps maintained neutral pH inside the cells and generated electrochemical gradients of protons (proton-motive force) across their membranes.<\/li>\r\n<\/ul>\r\n<ul style=\"text-align: justify\">\r\n \t<li>The existence of proton-motif force enabled the evolution of porters driven by it that are most probably among the more primitive porters in the world. The directionality of the substrate transport by the porters could be to both sides of the membranes because they can serve as proton symporters or antiporters.<\/li>\r\n<\/ul>\r\n<ul>\r\n \t<li style=\"text-align: justify\">One of the most important subjects of this meeting is the mechanism by which proton-motif and other ion-motif forces drive the transport processes through porters.<\/li>\r\n<\/ul>\r\n&nbsp;\r\n\r\n<strong>Proton-motif force: free energy change during movement across a <em>proton<\/em> gradient<\/strong>\r\n<ul>\r\n \t<li style=\"text-align: justify\">\"Proton-motif force\" (Dp) is a Dy- or DE-like term (DE is <em>electromotive force<\/em> ) that combines the concentration and voltage effects of a proton gradient such that<\/li>\r\n<\/ul>\r\nDG = - n<em>F<\/em>Dp0' + n<em>F<\/em>Dp, and Dp0' = 0 (proton motive force = 0 under std conds), so\r\n\r\n&nbsp;\r\n\r\nDG = nF Dp\r\n\r\nDG can also be expressed as the sum of the DpH and Dym contributions:\r\n\r\n&nbsp;\r\n\r\nDG = - 2.303 RT DpH + n<em>F<\/em>Dym\r\n\r\nso nFDp = - 2.303 RT DpH + n<em>F<\/em>Dym\r\n\r\n&nbsp;\r\n\r\n&nbsp;\r\n\r\nor\r\n\r\nDp = (-2.303 RT DpH)\/n<em>F<\/em> + Dym\r\n\r\n&nbsp;\r\n<ul>\r\n \t<li style=\"text-align: justify\">This way of expressing the proton-motif force was probably adopted because of its elegant resemblance to the Nernst equation, but the clearest expression of the energy available from a proton gradient is probably DG = - 2.303 RT DpH + nFDym<\/li>\r\n<\/ul>\r\n<ul>\r\n \t<li style=\"text-align: justify\">Remember (this is a recording) that DpH = pHin - pHout and Dym = yin - yout because we started out by considering the movement of protons from the cytoplasm to the matrix:<\/li>\r\n<\/ul>\r\n<img class=\"aligncenter size-full wp-image-292\" src=\"http:\/\/biocp04.epgpbooks.inflibnet.ac.in\/wp-content\/uploads\/sites\/215\/2019\/04\/1-143.png\" alt=\"\" width=\"154\" height=\"39\" \/>\r\n\r\n&nbsp;\r\n<p style=\"text-align: justify\">The total free energy available from the movement of 1 mole of protons from the cytoplasm to the matrix under cellular conditions (DpH = 1.4, Dy = 0.14 V) is the sum of the free energy changes calculated in sections B and C:<\/p>\r\n&nbsp;\r\n\r\n<img class=\"aligncenter size-full wp-image-293\" src=\"http:\/\/biocp04.epgpbooks.inflibnet.ac.in\/wp-content\/uploads\/sites\/215\/2019\/04\/1-144.png\" alt=\"\" width=\"374\" height=\"53\" \/>\r\n\r\n&nbsp;\r\n\r\n<img class=\"aligncenter size-full wp-image-294\" src=\"http:\/\/biocp04.epgpbooks.inflibnet.ac.in\/wp-content\/uploads\/sites\/215\/2019\/04\/1-145.png\" alt=\"\" width=\"517\" height=\"252\" \/>\r\n<h2 style=\"text-align: center\"><strong>Figure \u2013 8. Transfer of Proton from the cytoplasm to mitochondria<\/strong><\/h2>\r\n<h2 style=\"text-align: center\"><strong>( Adapted from- http:\/\/www.ccrc.uga.edu\/~rcarlson\/bcmb3100\/Chap21)<\/strong><\/h2>\r\n<img class=\"aligncenter size-full wp-image-295\" src=\"http:\/\/biocp04.epgpbooks.inflibnet.ac.in\/wp-content\/uploads\/sites\/215\/2019\/04\/1-146.png\" alt=\"\" width=\"559\" height=\"376\" \/>\r\n<h2 style=\"text-align: center\"><strong>Figure- 9 Many Mitochondrial Transporters<\/strong><\/h2>\r\n<h2 style=\"text-align: center\"><strong>(Adapted from-http:\/\/www.ccrc.uga.edu\/~rcarlson\/bcmb3100\/Chap21)<\/strong><\/h2>\r\n<img class=\"aligncenter size-full wp-image-296\" src=\"http:\/\/biocp04.epgpbooks.inflibnet.ac.in\/wp-content\/uploads\/sites\/215\/2019\/04\/1-147.png\" alt=\"\" width=\"596\" height=\"379\" \/>\r\n\r\n&nbsp;\r\n\r\n<img class=\"aligncenter size-full wp-image-297\" src=\"http:\/\/biocp04.epgpbooks.inflibnet.ac.in\/wp-content\/uploads\/sites\/215\/2019\/04\/1-148.png\" alt=\"\" width=\"611\" height=\"349\" \/>\r\n\r\n&nbsp;\r\n\r\n<img class=\"aligncenter size-full wp-image-298\" src=\"http:\/\/biocp04.epgpbooks.inflibnet.ac.in\/wp-content\/uploads\/sites\/215\/2019\/04\/1-149.png\" alt=\"\" width=\"460\" height=\"335\" \/>\r\n\r\n&nbsp;\r\n\r\n&nbsp;\r\n<table>\r\n<tbody>\r\n<tr>\r\n<td><strong>you can view video on Proton motive force<\/strong><\/td>\r\n<td><a href=\"https:\/\/youtu.be\/J660Uo1f0yU\" target=\"_blank\" rel=\"noopener\"><img class=\"alignnone wp-image-120\" src=\"http:\/\/epgpbooks.inflibnet.ac.in\/wp-content\/uploads\/2018\/11\/download.png\" alt=\"\" width=\"36\" height=\"36\" \/><\/a><\/td>\r\n<\/tr>\r\n<\/tbody>\r\n<\/table>\r\n&nbsp;\r\n<div class=\"textbox learning-objectives\">\r\n<h3><strong>References<\/strong><\/h3>\r\n<strong>Web site<\/strong>\r\n<ul>\r\n \t<li><a href=\"http:\/\/www.nature.com\/scitable\/topicpage\/why-are-cells-powered-by-proton-gradients-14373960\">http:\/\/www.nature.com\/scitable\/topicpage\/why-are-cells-powered-by-proton-<\/a><a href=\"http:\/\/www.nature.com\/scitable\/topicpage\/why-are-cells-powered-by-proton-gradients-14373960\">gradients-14373960<\/a><\/li>\r\n \t<li><a href=\"http:\/\/www.ccrc.uga.edu\/~rcarlson\/bcmb3100\/Chap21\">http:\/\/www.ccrc.uga.edu\/~rcarlson\/bcmb3100\/Chap21<\/a><\/li>\r\n \t<li><a href=\"https:\/\/en.wikipedia.org\/wiki\/Chemiosmosis\">https:\/\/en.wikipedia.org\/wiki\/Chemiosmosis<\/a><\/li>\r\n \t<li><a href=\"http:\/\/www.biologyaspoetry.com\/terms\/proton_motive_force.html\">http:\/\/www.biologyaspoetry.com\/terms\/proton_motive_force.html<\/a><\/li>\r\n<\/ul>\r\n&nbsp;\r\n\r\n<strong>Books<\/strong>\r\n<ul>\r\n \t<li style=\"text-align: justify\">Harper\u2019sIllustrated Biochemistry by Robert K.. Murray, Daryl K. Granner, Peter A. Mayes, 26th Edition (2003)<\/li>\r\n \t<li style=\"text-align: justify\">Lehninger\u2019S Principle of Biochemistry David L. Nelson and Michael M. Cox,5th Edition (2008)<\/li>\r\n \t<li style=\"text-align: justify\">Instant Notes: Biochemistry, 2nd Edition, B.D. Hames &amp; N. M. Hooper (2005)<\/li>\r\n<\/ul>\r\n<p style=\"text-align: justify\"><\/p>\r\n\r\n<\/div>\r\n&nbsp;\r\n\r\n&nbsp;","rendered":"<div><span style=\"float: right\"><a href=\"https:\/\/youtu.be\/J660Uo1f0yU\" target=\"_blank\" rel=\"noopener\"><img decoding=\"async\" src=\"http:\/\/epgpbooks.inflibnet.ac.in\/wp-content\/uploads\/2018\/11\/download.png\" alt=\"epgp books\" width=\"75px\" height=\"75px;\" \/><\/a><br \/>\n<\/span><\/div>\n<p><strong>\u00a0 \u00a0 <\/strong><\/p>\n<p>&nbsp;<\/p>\n<p>&nbsp;<\/p>\n<p>&nbsp;<\/p>\n<p><strong>PROTON MOTIVE FORCE<\/strong><\/p>\n<p><strong>Objectives<\/strong><\/p>\n<ol>\n<li>To understand the function of Proton Motive Force in production of ATP.<\/li>\n<li>To examine the importance of a proton gradients power the synthesis of ATP.<\/li>\n<li>To examine the importance of shuttles for aerobic oxidation of cytosolic NADH.<\/li>\n<li>To understand the cellular respiration which is regulated by the Need for ATP.<\/li>\n<\/ol>\n<p><strong>\u00a0 \u00a0 <\/strong><\/p>\n<p><strong>Introduction<\/strong><\/p>\n<ul>\n<li style=\"text-align: justify\">A proton-motive force is &#8220;energy that is generated by the transfer of protons or electrons across an energy-transducing membrane and can be used for chemical, osmotic or mechanical work.&#8221;<\/li>\n<\/ul>\n<ul>\n<li style=\"text-align: justify\">It can be &#8220;generated by a variety of phenomena including the operation of an electron transport chain, illumination of a purple membrane, and the hydrolysis of ATP by a proton ATPase.&#8221;<\/li>\n<\/ul>\n<p><img loading=\"lazy\" decoding=\"async\" class=\"aligncenter size-full wp-image-279\" src=\"http:\/\/biocp04.epgpbooks.inflibnet.ac.in\/wp-content\/uploads\/sites\/215\/2019\/04\/1-130.png\" alt=\"\" width=\"470\" height=\"78\" srcset=\"https:\/\/ebooks.inflibnet.ac.in\/biocp04\/wp-content\/uploads\/sites\/215\/2019\/04\/1-130.png 470w, https:\/\/ebooks.inflibnet.ac.in\/biocp04\/wp-content\/uploads\/sites\/215\/2019\/04\/1-130-300x50.png 300w, https:\/\/ebooks.inflibnet.ac.in\/biocp04\/wp-content\/uploads\/sites\/215\/2019\/04\/1-130-65x11.png 65w, https:\/\/ebooks.inflibnet.ac.in\/biocp04\/wp-content\/uploads\/sites\/215\/2019\/04\/1-130-225x37.png 225w, https:\/\/ebooks.inflibnet.ac.in\/biocp04\/wp-content\/uploads\/sites\/215\/2019\/04\/1-130-350x58.png 350w\" sizes=\"auto, (max-width: 470px) 100vw, 470px\" \/><\/p>\n<p>&nbsp;<\/p>\n<p><strong>Proton Gradient<\/strong><\/p>\n<p>&nbsp;<\/p>\n<p>&nbsp;<\/p>\n<p><img loading=\"lazy\" decoding=\"async\" class=\"aligncenter size-full wp-image-280\" src=\"http:\/\/biocp04.epgpbooks.inflibnet.ac.in\/wp-content\/uploads\/sites\/215\/2019\/04\/1-131.png\" alt=\"\" width=\"545\" height=\"341\" srcset=\"https:\/\/ebooks.inflibnet.ac.in\/biocp04\/wp-content\/uploads\/sites\/215\/2019\/04\/1-131.png 545w, https:\/\/ebooks.inflibnet.ac.in\/biocp04\/wp-content\/uploads\/sites\/215\/2019\/04\/1-131-300x188.png 300w, https:\/\/ebooks.inflibnet.ac.in\/biocp04\/wp-content\/uploads\/sites\/215\/2019\/04\/1-131-65x41.png 65w, https:\/\/ebooks.inflibnet.ac.in\/biocp04\/wp-content\/uploads\/sites\/215\/2019\/04\/1-131-225x141.png 225w, https:\/\/ebooks.inflibnet.ac.in\/biocp04\/wp-content\/uploads\/sites\/215\/2019\/04\/1-131-350x219.png 350w\" sizes=\"auto, (max-width: 545px) 100vw, 545px\" \/><\/p>\n<h2 style=\"text-align: center\"><strong>Figure-1. Proton motif force<\/strong><\/h2>\n<h2 style=\"text-align: center\"><strong>( Adapted from- http:\/\/classroom.sdmesa.edu\/eschmid\/Lectur46.gif)<\/strong><\/h2>\n<p>&nbsp;<\/p>\n<p><strong>Why Proton Gradients Are Necessary<\/strong><\/p>\n<ul>\n<li style=\"text-align: justify\">The reason proton gradients are required boils back down to chemistry. In other words, to convert carbon dioxide into organic molecules, life attaches hydrogen atoms to CO2<\/li>\n<\/ul>\n<ul style=\"text-align: justify\">\n<li>This pathway is exothermic (releasing energy that can be captured as ATP) right through to pyruvate, one of the central molecules in cell metabolism<\/li>\n<\/ul>\n<ul style=\"text-align: justify\">\n<li>But there&#8217;s a problem, pointed out by William Martin in collaboration with Russell. All cells that use the acetyl-CoA pathway today depend on proton gradients. None of them can grow by fermentation<\/li>\n<\/ul>\n<ul>\n<li style=\"text-align: justify\">Why not? Because CO2 is a stable molecule and does not react easily, even with hydrogen &#8211; even when thermodynamics says it should react. CO2 is a bit like oxygen in this respect: Once it starts to react, it&#8217;s not easily stopped. But a fire<span style=\"text-align: initial;font-size: 1em\"> needs a spark to get it going, and so, too, does CO2. Cells need the equivalent of a spark to get CO2 to react, and for cells, that spark is ATP<\/span><\/li>\n<\/ul>\n<ul>\n<li style=\"text-align: justify\">The problem is that the reaction of CO2 with H2 releases energy, but not a lot \u2014 only enough to make 1 ATP. That means cells have to spend 1 ATP to gain 1 ATP, so there&#8217;s no net gain. If there&#8217;s no gain, there&#8217;s no growth; no growth, no life<\/li>\n<\/ul>\n<p><img loading=\"lazy\" decoding=\"async\" class=\"aligncenter size-full wp-image-281\" src=\"http:\/\/biocp04.epgpbooks.inflibnet.ac.in\/wp-content\/uploads\/sites\/215\/2019\/04\/1-132.png\" alt=\"\" width=\"488\" height=\"161\" srcset=\"https:\/\/ebooks.inflibnet.ac.in\/biocp04\/wp-content\/uploads\/sites\/215\/2019\/04\/1-132.png 488w, https:\/\/ebooks.inflibnet.ac.in\/biocp04\/wp-content\/uploads\/sites\/215\/2019\/04\/1-132-300x99.png 300w, https:\/\/ebooks.inflibnet.ac.in\/biocp04\/wp-content\/uploads\/sites\/215\/2019\/04\/1-132-65x21.png 65w, https:\/\/ebooks.inflibnet.ac.in\/biocp04\/wp-content\/uploads\/sites\/215\/2019\/04\/1-132-225x74.png 225w, https:\/\/ebooks.inflibnet.ac.in\/biocp04\/wp-content\/uploads\/sites\/215\/2019\/04\/1-132-350x115.png 350w\" sizes=\"auto, (max-width: 488px) 100vw, 488px\" \/><\/p>\n<ul>\n<li style=\"text-align: justify\">Gradients break that cycle. It&#8217;s not quite true to say that the reaction of CO 2 with H2 releases enough energy to make 1 ATP: it&#8217;s actually enough to make 1.5 ATPs. But of course there&#8217;s no such thing as 1.5 ATPs, at least not by stoichiometric chemistry; so the spare energy from the reaction, as chemistry, is lost<\/li>\n<\/ul>\n<ul style=\"text-align: justify\">\n<li>But that doesn&#8217;t happen with a gradient. In principle, a reaction can be repeated over and over again, just to pump a proton over a membrane<\/li>\n<\/ul>\n<ul style=\"text-align: justify\">\n<li>When enough protons have accumulated, the proton motif force powers the formation of ATP<\/li>\n<\/ul>\n<ul>\n<li style=\"text-align: justify\">So a gradient allows cells to save up protons as &#8220;loose change&#8221;, and that makes all the difference in the world \u2014 the difference between growth and no growth, life and no life.<\/li>\n<\/ul>\n<p><img loading=\"lazy\" decoding=\"async\" class=\"aligncenter size-full wp-image-282\" src=\"http:\/\/biocp04.epgpbooks.inflibnet.ac.in\/wp-content\/uploads\/sites\/215\/2019\/04\/1-133.png\" alt=\"\" width=\"603\" height=\"234\" srcset=\"https:\/\/ebooks.inflibnet.ac.in\/biocp04\/wp-content\/uploads\/sites\/215\/2019\/04\/1-133.png 603w, https:\/\/ebooks.inflibnet.ac.in\/biocp04\/wp-content\/uploads\/sites\/215\/2019\/04\/1-133-300x116.png 300w, https:\/\/ebooks.inflibnet.ac.in\/biocp04\/wp-content\/uploads\/sites\/215\/2019\/04\/1-133-65x25.png 65w, https:\/\/ebooks.inflibnet.ac.in\/biocp04\/wp-content\/uploads\/sites\/215\/2019\/04\/1-133-225x87.png 225w, https:\/\/ebooks.inflibnet.ac.in\/biocp04\/wp-content\/uploads\/sites\/215\/2019\/04\/1-133-350x136.png 350w\" sizes=\"auto, (max-width: 603px) 100vw, 603px\" \/><\/p>\n<p>&nbsp;<\/p>\n<p><strong>A proton gradient powers the synthesis of ATP<\/strong><\/p>\n<ul>\n<li style=\"text-align: justify\">The overall process of ATP generation via the harnessing of a proton motif force is called chemiosmosis<\/li>\n<li style=\"text-align: justify\">The Chemiosmotic Theory Proposed by Peter Mitchell in the 1960\u2019s<\/li>\n<li style=\"text-align: justify\">The proton gradient generated by the oxidation of NADH and FADH2 is called the proton-motif force.The proton-motif force powers the synthesis of ATP<\/li>\n<\/ul>\n<p><strong>\u00a0 Coupled nature of respiration in mitochondria<\/strong><\/p>\n<p>&nbsp;<\/p>\n<p><img loading=\"lazy\" decoding=\"async\" class=\"aligncenter size-full wp-image-283\" src=\"http:\/\/biocp04.epgpbooks.inflibnet.ac.in\/wp-content\/uploads\/sites\/215\/2019\/04\/1-134.png\" alt=\"\" width=\"538\" height=\"240\" srcset=\"https:\/\/ebooks.inflibnet.ac.in\/biocp04\/wp-content\/uploads\/sites\/215\/2019\/04\/1-134.png 538w, https:\/\/ebooks.inflibnet.ac.in\/biocp04\/wp-content\/uploads\/sites\/215\/2019\/04\/1-134-300x134.png 300w, https:\/\/ebooks.inflibnet.ac.in\/biocp04\/wp-content\/uploads\/sites\/215\/2019\/04\/1-134-65x29.png 65w, https:\/\/ebooks.inflibnet.ac.in\/biocp04\/wp-content\/uploads\/sites\/215\/2019\/04\/1-134-225x100.png 225w, https:\/\/ebooks.inflibnet.ac.in\/biocp04\/wp-content\/uploads\/sites\/215\/2019\/04\/1-134-350x156.png 350w\" sizes=\"auto, (max-width: 538px) 100vw, 538px\" \/><\/p>\n<p style=\"text-align: center\">(a) O2 consumed only with ADP, excess Pi.<\/p>\n<p style=\"text-align: center\">(b) (+) Uncoupler DNP (O2 consumed without ADP).<\/p>\n<p>&nbsp;<\/p>\n<p><strong>(A) <\/strong><strong>O<\/strong><strong>2<\/strong><strong> consumed only with ADP, excess Pi<\/strong><\/p>\n<ul>\n<li>Oxidation of substrates is coupled to the phosphorylation of ADP<\/li>\n<li>Respiration (consumption of oxygen) proceeds only when ADP is present<\/li>\n<li>The amount of O 2 consumed depends upon the amount of ADP added<\/li>\n<\/ul>\n<p>&nbsp;<\/p>\n<p><strong>(B) <\/strong><strong>(+) Uncoupler DNP (O 2 consumed without ADP)<\/strong><\/p>\n<ul>\n<li>Uncouplers stimulate the oxidation of substrates in the absence of ADP.<\/li>\n<li>Uncouplers are lipid-soluble weak acids.<\/li>\n<li>Both acidic and basic forms can cross the inner mitochondrial membrane.<\/li>\n<li>Uncouplers deplete any proton gradient by transporting protons across the membrane.<\/li>\n<\/ul>\n<p><img loading=\"lazy\" decoding=\"async\" class=\"aligncenter size-full wp-image-284\" src=\"http:\/\/biocp04.epgpbooks.inflibnet.ac.in\/wp-content\/uploads\/sites\/215\/2019\/04\/1-135.png\" alt=\"\" width=\"662\" height=\"175\" srcset=\"https:\/\/ebooks.inflibnet.ac.in\/biocp04\/wp-content\/uploads\/sites\/215\/2019\/04\/1-135.png 662w, https:\/\/ebooks.inflibnet.ac.in\/biocp04\/wp-content\/uploads\/sites\/215\/2019\/04\/1-135-300x79.png 300w, https:\/\/ebooks.inflibnet.ac.in\/biocp04\/wp-content\/uploads\/sites\/215\/2019\/04\/1-135-65x17.png 65w, https:\/\/ebooks.inflibnet.ac.in\/biocp04\/wp-content\/uploads\/sites\/215\/2019\/04\/1-135-225x59.png 225w, https:\/\/ebooks.inflibnet.ac.in\/biocp04\/wp-content\/uploads\/sites\/215\/2019\/04\/1-135-350x93.png 350w\" sizes=\"auto, (max-width: 662px) 100vw, 662px\" \/><\/p>\n<h2 style=\"text-align: center\"><strong>2,4-Dinitrophenol: an uncoupler<\/strong><\/h2>\n<p>&nbsp;<\/p>\n<p><strong>Mitchell\u2019s postulates for Chemiosmotic theory<\/strong><\/p>\n<ul>\n<li style=\"text-align: justify\">Intact inner mitochondrial membrane is required (to maintain a proton gradient)<\/li>\n<li style=\"text-align: justify\">Electron transport through the ETC generates a proton gradient (pumps H + from the matrix to the intermembrane space)<\/li>\n<li style=\"text-align: justify\">The membrane-spanning enzyme, ATP synthase, catalyzes the phosphorylation of ADP in a reaction driven by movement of H + across the inner membrane into the matrix<\/li>\n<\/ul>\n<p>&nbsp;<\/p>\n<p><strong>Shuttles are Necessary for Aerobic Oxidation of Cytosolic NADH<\/strong><\/p>\n<ul>\n<li style=\"text-align: justify\">Cytosolic NADH must enter the mitochondria to fuel oxidative phosphorylation, but NADH and NAD+ cannot diffuse across the inner mitochondrial membrane<\/li>\n<li style=\"text-align: justify\">Two shuttle systems for reducing equivalents:<\/li>\n<\/ul>\n<ol style=\"text-align: justify\">\n<li>Glycerol phosphate shuttle: insect flight muscles<\/li>\n<li>Malate-aspartate shuttle: predominant in liver and other mammalian tissues<\/li>\n<\/ol>\n<p><strong>1. Glycerol phosphate shuttle<\/strong><\/p>\n<ul>\n<li style=\"text-align: justify\">The glycerol-3-phosphate shuttle is a mechanism that regenerates NAD+ from NADH, a by-product of glycolysis. Its importance in transporting reducing equivalents is secondary to the malate-aspartate shuttle<\/li>\n<\/ul>\n<p><strong><em>\u00a0 \u00a0 \u00a0Mechanism:<\/em><\/strong><\/p>\n<ul>\n<li style=\"text-align: justify\">In this\u00a0 shuttle,\u00a0 the\u00a0 enzyme glycerol-3-phosphate\u00a0 dehydrogenase 1\u00a0 (GPDH-C) converts dihydroxyacetone phosphate <span style=\"text-align: initial;font-size: 1em\">(2) to <\/span>glycerol 3-phosphate <span style=\"text-align: initial;font-size: 1em\">(1) by oxidizing one molecule of NADH to NAD+<\/span><\/li>\n<\/ul>\n<ul style=\"text-align: justify\">\n<li>Glycerol-3-phosphate gets converted back to dihydroxyacetone phosphate by an inner membrane-bound mitochondrial glycerol-3-phosphate dehydrogenase 2 (GPDH-M), this time reducing one molecule of enzyme-bound flavin adenine dinucleotide <span style=\"text-align: initial;font-size: 1em\">(FAD) to FADH2. FADH2 then reduces <\/span>coenzyme Q <span style=\"text-align: initial;font-size: 1em\">(ubiquinone to ubiquinol) which enters into <\/span>oxidative phosphorylation. <span style=\"text-align: initial;font-size: 1em\">This reaction is irreversible.<\/span><\/li>\n<\/ul>\n<ul>\n<li style=\"text-align: justify\">The glycerol-3-phosphate shuttle allows the NADH synthesized in the cytosol by glycolysis to contribute to the oxidative phosphorylation <span style=\"text-align: initial;font-size: 1em\">pathway in the <\/span>mitochondria <span style=\"text-align: initial;font-size: 1em\">to generate ATP. It has been found in animals, fungi, and plants.<\/span><\/li>\n<\/ul>\n<p><img loading=\"lazy\" decoding=\"async\" class=\"aligncenter size-full wp-image-285\" src=\"http:\/\/biocp04.epgpbooks.inflibnet.ac.in\/wp-content\/uploads\/sites\/215\/2019\/04\/1-136.png\" alt=\"\" width=\"523\" height=\"336\" srcset=\"https:\/\/ebooks.inflibnet.ac.in\/biocp04\/wp-content\/uploads\/sites\/215\/2019\/04\/1-136.png 523w, https:\/\/ebooks.inflibnet.ac.in\/biocp04\/wp-content\/uploads\/sites\/215\/2019\/04\/1-136-300x193.png 300w, https:\/\/ebooks.inflibnet.ac.in\/biocp04\/wp-content\/uploads\/sites\/215\/2019\/04\/1-136-65x42.png 65w, https:\/\/ebooks.inflibnet.ac.in\/biocp04\/wp-content\/uploads\/sites\/215\/2019\/04\/1-136-225x145.png 225w, https:\/\/ebooks.inflibnet.ac.in\/biocp04\/wp-content\/uploads\/sites\/215\/2019\/04\/1-136-350x225.png 350w\" sizes=\"auto, (max-width: 523px) 100vw, 523px\" \/><\/p>\n<h2 style=\"text-align: center\"><strong>Figure -2 Glycerol phosphate shuttle<\/strong><\/h2>\n<h2 style=\"text-align: center\"><strong>(Adapted from: http:\/\/oregonstate.edu\/instruct\/bb451\/winter14\/stryer7\/CH18\/figure_18_34.jpg))<\/strong><\/h2>\n<p>&nbsp;<\/p>\n<p><strong>Malate-aspartate shuttle<\/strong><\/p>\n<ul>\n<li style=\"text-align: justify\">The malate-aspartate shuttle is a biochemical system for translocating electrons\u00a0produced\u00a0during glycolysis across\u00a0the semipermeable inner\u00a0membrane of the mitochondrion for oxidative phosphorylation in eukaryotes.\u00a0These electrons\u00a0enter in\u00a0the electron transport\u00a0chain of\u00a0the mitochondria via reduction\u00a0equivalents to generate ATP.<\/li>\n<\/ul>\n<p>&nbsp;<\/p>\n<p>&nbsp;<\/p>\n<p><strong><em>Mechanism:<\/em><\/strong><\/p>\n<ul>\n<li style=\"text-align: justify\">The primary enzyme in the malate-aspartate shuttle is malate dehydrogenase.<\/li>\n<\/ul>\n<ul style=\"text-align: justify\">\n<li>First, in the cytosol, malate dehydrogenase catalyses the reaction of oxaloacetate and NADH to produce malate and NAD+.<\/li>\n<\/ul>\n<ul style=\"text-align: justify\">\n<li>Once malate is formed, the first antiporter (malate-alpha-ketoglutarate)imports the malate from the cytosol into the mitochondrial matrix and also exports alpha-ketoglutarate from the matrix into the cytosol simultaneously.<\/li>\n<\/ul>\n<ul style=\"text-align: justify\">\n<li>Oxaloacetate is then transformed into aspartate by mitochondrial aspartate aminotransferase. Since aspartate is an amino acid, an amino radical needs to be added to the oxaloacetate.<\/li>\n<\/ul>\n<ul>\n<li style=\"text-align: justify\">The second antiporter (the glutamate-aspartate antiporter) imports glutamate from the cytosol into the matrix and exports aspartate from the matrix to the cytosol.<\/li>\n<\/ul>\n<ul>\n<li style=\"text-align: justify\">The net effect of the malate-aspartate shuttle is purely redox: NADH in the cytosol is oxidized to NAD+, and NAD+ in the matrix is reduced to NADH.<\/li>\n<\/ul>\n<ul style=\"text-align: justify\">\n<li>Since the malate-aspartate shuttle regenerates NADH inside the mitochondrial matrix, it is capable of maximizing the number of ATPs produced in glycolysis (3\/NADH), ultimately resulting in a net gain of 38 ATP molecules per molecule of glucose metabolized.<\/li>\n<\/ul>\n<p style=\"text-align: justify\"><img loading=\"lazy\" decoding=\"async\" class=\"aligncenter size-full wp-image-286\" src=\"http:\/\/biocp04.epgpbooks.inflibnet.ac.in\/wp-content\/uploads\/sites\/215\/2019\/04\/1-137.png\" alt=\"\" width=\"465\" height=\"244\" srcset=\"https:\/\/ebooks.inflibnet.ac.in\/biocp04\/wp-content\/uploads\/sites\/215\/2019\/04\/1-137.png 465w, https:\/\/ebooks.inflibnet.ac.in\/biocp04\/wp-content\/uploads\/sites\/215\/2019\/04\/1-137-300x157.png 300w, https:\/\/ebooks.inflibnet.ac.in\/biocp04\/wp-content\/uploads\/sites\/215\/2019\/04\/1-137-65x34.png 65w, https:\/\/ebooks.inflibnet.ac.in\/biocp04\/wp-content\/uploads\/sites\/215\/2019\/04\/1-137-225x118.png 225w, https:\/\/ebooks.inflibnet.ac.in\/biocp04\/wp-content\/uploads\/sites\/215\/2019\/04\/1-137-350x184.png 350w\" sizes=\"auto, (max-width: 465px) 100vw, 465px\" \/><\/p>\n<h2 style=\"text-align: center\"><strong>Figure- 3 Malate-aspartate shuttle<\/strong><\/h2>\n<h2 style=\"text-align: center\"><strong>( Adapted from- http:\/\/oregonstate.edu\/instruct\/bb451\/winter14\/stryer7\/CH18\/figure_18_35.jpg)<\/strong><\/h2>\n<p>&nbsp;<\/p>\n<p><strong>Regulation of Proton motif force<\/strong><\/p>\n<ul>\n<li>Protonmotif force ( D p) is the energy of the proton concentration gradient<\/li>\n<\/ul>\n<p><img loading=\"lazy\" decoding=\"async\" class=\"aligncenter size-full wp-image-287\" src=\"http:\/\/biocp04.epgpbooks.inflibnet.ac.in\/wp-content\/uploads\/sites\/215\/2019\/04\/1-138.png\" alt=\"\" width=\"470\" height=\"297\" srcset=\"https:\/\/ebooks.inflibnet.ac.in\/biocp04\/wp-content\/uploads\/sites\/215\/2019\/04\/1-138.png 470w, https:\/\/ebooks.inflibnet.ac.in\/biocp04\/wp-content\/uploads\/sites\/215\/2019\/04\/1-138-300x190.png 300w, https:\/\/ebooks.inflibnet.ac.in\/biocp04\/wp-content\/uploads\/sites\/215\/2019\/04\/1-138-65x41.png 65w, https:\/\/ebooks.inflibnet.ac.in\/biocp04\/wp-content\/uploads\/sites\/215\/2019\/04\/1-138-225x142.png 225w, https:\/\/ebooks.inflibnet.ac.in\/biocp04\/wp-content\/uploads\/sites\/215\/2019\/04\/1-138-350x221.png 350w\" sizes=\"auto, (max-width: 470px) 100vw, 470px\" \/><\/p>\n<h2 style=\"text-align: center\"><strong>Figure \u2013 4. Proton concentration gradient in chemical and electrical condition.<\/strong><\/h2>\n<h2 style=\"text-align: center\"><strong>(Adapted from &#8211; http:\/\/www.ccrc.uga.edu\/~rcarlson\/bcmb3100\/Chap21)<\/strong><\/h2>\n<p>&nbsp;<\/p>\n<p><strong>ATP synthase<\/strong><\/p>\n<ul>\n<li style=\"text-align: justify\">There are 3 active sites, one in each b subunit<\/li>\n<li style=\"text-align: justify\">The c-e-g unit forms a \u201crotor\u201d<\/li>\n<li style=\"text-align: justify\">Rotation of the g subunit inside the a3b3 hexamer causes domain movements in the b-subunits, opening and closing the active sites<\/li>\n<li style=\"text-align: justify\">The a-b-g-a3b3 unit is the \u201cstator\u201d (the Fo channel is attached to a3b3 by the ab-d arm)<\/li>\n<\/ul>\n<p><img loading=\"lazy\" decoding=\"async\" class=\"aligncenter size-full wp-image-288\" src=\"http:\/\/biocp04.epgpbooks.inflibnet.ac.in\/wp-content\/uploads\/sites\/215\/2019\/04\/1-139.png\" alt=\"\" width=\"375\" height=\"286\" srcset=\"https:\/\/ebooks.inflibnet.ac.in\/biocp04\/wp-content\/uploads\/sites\/215\/2019\/04\/1-139.png 375w, https:\/\/ebooks.inflibnet.ac.in\/biocp04\/wp-content\/uploads\/sites\/215\/2019\/04\/1-139-300x229.png 300w, https:\/\/ebooks.inflibnet.ac.in\/biocp04\/wp-content\/uploads\/sites\/215\/2019\/04\/1-139-65x50.png 65w, https:\/\/ebooks.inflibnet.ac.in\/biocp04\/wp-content\/uploads\/sites\/215\/2019\/04\/1-139-225x172.png 225w, https:\/\/ebooks.inflibnet.ac.in\/biocp04\/wp-content\/uploads\/sites\/215\/2019\/04\/1-139-350x267.png 350w\" sizes=\"auto, (max-width: 375px) 100vw, 375px\" \/><\/p>\n<h2 style=\"text-align: center\"><strong>Figure \u2013 5. ATP synthase.<\/strong><\/h2>\n<h2 style=\"text-align: center\"><strong>(Adapted<\/strong> <strong>from https:\/\/s3.amazonaws.com\/classconnection\/281\/flashcards\/2541281\/png\/session_15_ _oxidative_phosphorylation_-_google_slides_(1)-14C6815A20153228B19.png)<\/strong><\/h2>\n<h2 style=\"text-align: center\"><\/h2>\n<ul>\n<li>F0F1 ATP Synthase uses the proton gradient energy for the synthesis of ATP<\/li>\n<li>An F-type ATPase which generates ATP<\/li>\n<li>Composed of a \u201cknob-and-stalk\u201d structure<\/li>\n<li>F1 (knob) contains the catalytic subunits<\/li>\n<li>F0 (stalk) has a proton channel which spans the membrane.<\/li>\n<li>Passage of protons through the F0 (stalk) into the matrix is coupled to ATP formation<\/li>\n<li>Estimated passage of 3 H+ \/ ATP synthesized<\/li>\n<\/ul>\n<ul>\n<li style=\"text-align: justify\">Passage of protons through the F0 channel causes the rotor to spin in one direction and the stator to spin in the opposite direction<\/li>\n<\/ul>\n<p><img loading=\"lazy\" decoding=\"async\" class=\"aligncenter size-full wp-image-289\" src=\"http:\/\/biocp04.epgpbooks.inflibnet.ac.in\/wp-content\/uploads\/sites\/215\/2019\/04\/1-140.png\" alt=\"\" width=\"485\" height=\"151\" srcset=\"https:\/\/ebooks.inflibnet.ac.in\/biocp04\/wp-content\/uploads\/sites\/215\/2019\/04\/1-140.png 485w, https:\/\/ebooks.inflibnet.ac.in\/biocp04\/wp-content\/uploads\/sites\/215\/2019\/04\/1-140-300x93.png 300w, https:\/\/ebooks.inflibnet.ac.in\/biocp04\/wp-content\/uploads\/sites\/215\/2019\/04\/1-140-65x20.png 65w, https:\/\/ebooks.inflibnet.ac.in\/biocp04\/wp-content\/uploads\/sites\/215\/2019\/04\/1-140-225x70.png 225w, https:\/\/ebooks.inflibnet.ac.in\/biocp04\/wp-content\/uploads\/sites\/215\/2019\/04\/1-140-350x109.png 350w\" sizes=\"auto, (max-width: 485px) 100vw, 485px\" \/><\/p>\n<p style=\"text-align: center\">Three protons = one complete rotation = one ATP<\/p>\n<p>&nbsp;<\/p>\n<p>&nbsp;<\/p>\n<p><img loading=\"lazy\" decoding=\"async\" class=\"aligncenter size-full wp-image-290\" src=\"http:\/\/biocp04.epgpbooks.inflibnet.ac.in\/wp-content\/uploads\/sites\/215\/2019\/04\/1-141.png\" alt=\"\" width=\"475\" height=\"162\" srcset=\"https:\/\/ebooks.inflibnet.ac.in\/biocp04\/wp-content\/uploads\/sites\/215\/2019\/04\/1-141.png 475w, https:\/\/ebooks.inflibnet.ac.in\/biocp04\/wp-content\/uploads\/sites\/215\/2019\/04\/1-141-300x102.png 300w, https:\/\/ebooks.inflibnet.ac.in\/biocp04\/wp-content\/uploads\/sites\/215\/2019\/04\/1-141-65x22.png 65w, https:\/\/ebooks.inflibnet.ac.in\/biocp04\/wp-content\/uploads\/sites\/215\/2019\/04\/1-141-225x77.png 225w, https:\/\/ebooks.inflibnet.ac.in\/biocp04\/wp-content\/uploads\/sites\/215\/2019\/04\/1-141-350x119.png 350w\" sizes=\"auto, (max-width: 475px) 100vw, 475px\" \/><\/p>\n<h2 style=\"text-align: center\"><strong>Figure \u2013 6 Conformational changes in ATP synthase.<\/strong><\/h2>\n<h2 style=\"text-align: center\"><strong>\u00a0 1.\u00a0 ADP, Pi bind to an open site.<\/strong><\/h2>\n<h2 style=\"text-align: center\"><strong>2. Inward passage of protons, conformation change, ATP synthesis from ADP and Pi.<\/strong><\/h2>\n<h2 style=\"text-align: center\"><strong>3. ATP released from <\/strong>open<strong> site, ADP and Pi form ATP in the tight site.<\/strong><\/h2>\n<h2 style=\"text-align: center\"><strong>(Adapted from http:\/\/1.bp.blogspot.com\/_DZH2cmCoois\/R2hL3wt_O6I\/AAAAAAAAEHU\/7WNKoQjakDk\/s400\/tmp.jpg)<\/strong><\/h2>\n<p>&nbsp;<\/p>\n<p><strong>Proton flow around the \u201cc\u201d ring<\/strong><\/p>\n<ul>\n<li style=\"text-align: justify\">The number of c rings determines the number of protons required to synthesize a molecule of ATP.<\/li>\n<li style=\"text-align: justify\">The c ring of vertebrates consist of 8 subunits, making vertebrate ATP synthase the most efficient known.<\/li>\n<\/ul>\n<p style=\"text-align: justify\"><img loading=\"lazy\" decoding=\"async\" class=\"aligncenter size-full wp-image-291\" src=\"http:\/\/biocp04.epgpbooks.inflibnet.ac.in\/wp-content\/uploads\/sites\/215\/2019\/04\/1-142.png\" alt=\"\" width=\"447\" height=\"295\" srcset=\"https:\/\/ebooks.inflibnet.ac.in\/biocp04\/wp-content\/uploads\/sites\/215\/2019\/04\/1-142.png 447w, https:\/\/ebooks.inflibnet.ac.in\/biocp04\/wp-content\/uploads\/sites\/215\/2019\/04\/1-142-300x198.png 300w, https:\/\/ebooks.inflibnet.ac.in\/biocp04\/wp-content\/uploads\/sites\/215\/2019\/04\/1-142-65x43.png 65w, https:\/\/ebooks.inflibnet.ac.in\/biocp04\/wp-content\/uploads\/sites\/215\/2019\/04\/1-142-225x148.png 225w, https:\/\/ebooks.inflibnet.ac.in\/biocp04\/wp-content\/uploads\/sites\/215\/2019\/04\/1-142-350x231.png 350w\" sizes=\"auto, (max-width: 447px) 100vw, 447px\" \/><\/p>\n<h2 style=\"text-align: center\"><strong>Figure \u2013 7 Proton flow around the C ring.<\/strong><\/h2>\n<h2 style=\"text-align: center\"><strong>(Adapted from-\u00a0<\/strong><strong>https:\/\/s3.amazonaws.com\/classconnection\/281\/flashcards\/2541281\/png\/session_15_- _oxidative_phosphorylation_-_google_slides_(5)-14C68FE9F127B4A4690.png)<\/strong><\/h2>\n<p>&nbsp;<\/p>\n<p><strong>SUMMARY<\/strong><\/p>\n<ul>\n<li style=\"text-align: justify\">It is generally accepted that the chemistry of water was the most crucial determinant in shaping life on earth. Among the more important chemical features of water is its dissociation into protons and hydroxyl ions.<\/li>\n<\/ul>\n<ul style=\"text-align: justify\">\n<li>The presence of relatively high proton concentrations in the ambient solution resulted in the evolution of proton pumps during the dawn of life on earth. These proton pumps maintained neutral pH inside the cells and generated electrochemical gradients of protons (proton-motive force) across their membranes.<\/li>\n<\/ul>\n<ul style=\"text-align: justify\">\n<li>The existence of proton-motif force enabled the evolution of porters driven by it that are most probably among the more primitive porters in the world. The directionality of the substrate transport by the porters could be to both sides of the membranes because they can serve as proton symporters or antiporters.<\/li>\n<\/ul>\n<ul>\n<li style=\"text-align: justify\">One of the most important subjects of this meeting is the mechanism by which proton-motif and other ion-motif forces drive the transport processes through porters.<\/li>\n<\/ul>\n<p>&nbsp;<\/p>\n<p><strong>Proton-motif force: free energy change during movement across a <em>proton<\/em> gradient<\/strong><\/p>\n<ul>\n<li style=\"text-align: justify\">&#8220;Proton-motif force&#8221; (Dp) is a Dy- or DE-like term (DE is <em>electromotive force<\/em> ) that combines the concentration and voltage effects of a proton gradient such that<\/li>\n<\/ul>\n<p>DG = &#8211; n<em>F<\/em>Dp0&#8242; + n<em>F<\/em>Dp, and Dp0&#8242; = 0 (proton motive force = 0 under std conds), so<\/p>\n<p>&nbsp;<\/p>\n<p>DG = nF Dp<\/p>\n<p>DG can also be expressed as the sum of the DpH and Dym contributions:<\/p>\n<p>&nbsp;<\/p>\n<p>DG = &#8211; 2.303 RT DpH + n<em>F<\/em>Dym<\/p>\n<p>so nFDp = &#8211; 2.303 RT DpH + n<em>F<\/em>Dym<\/p>\n<p>&nbsp;<\/p>\n<p>&nbsp;<\/p>\n<p>or<\/p>\n<p>Dp = (-2.303 RT DpH)\/n<em>F<\/em> + Dym<\/p>\n<p>&nbsp;<\/p>\n<ul>\n<li style=\"text-align: justify\">This way of expressing the proton-motif force was probably adopted because of its elegant resemblance to the Nernst equation, but the clearest expression of the energy available from a proton gradient is probably DG = &#8211; 2.303 RT DpH + nFDym<\/li>\n<\/ul>\n<ul>\n<li style=\"text-align: justify\">Remember (this is a recording) that DpH = pHin &#8211; pHout and Dym = yin &#8211; yout because we started out by considering the movement of protons from the cytoplasm to the matrix:<\/li>\n<\/ul>\n<p><img loading=\"lazy\" decoding=\"async\" class=\"aligncenter size-full wp-image-292\" src=\"http:\/\/biocp04.epgpbooks.inflibnet.ac.in\/wp-content\/uploads\/sites\/215\/2019\/04\/1-143.png\" alt=\"\" width=\"154\" height=\"39\" srcset=\"https:\/\/ebooks.inflibnet.ac.in\/biocp04\/wp-content\/uploads\/sites\/215\/2019\/04\/1-143.png 154w, https:\/\/ebooks.inflibnet.ac.in\/biocp04\/wp-content\/uploads\/sites\/215\/2019\/04\/1-143-150x39.png 150w, https:\/\/ebooks.inflibnet.ac.in\/biocp04\/wp-content\/uploads\/sites\/215\/2019\/04\/1-143-65x16.png 65w\" sizes=\"auto, (max-width: 154px) 100vw, 154px\" \/><\/p>\n<p>&nbsp;<\/p>\n<p style=\"text-align: justify\">The total free energy available from the movement of 1 mole of protons from the cytoplasm to the matrix under cellular conditions (DpH = 1.4, Dy = 0.14 V) is the sum of the free energy changes calculated in sections B and C:<\/p>\n<p>&nbsp;<\/p>\n<p><img loading=\"lazy\" decoding=\"async\" class=\"aligncenter size-full wp-image-293\" src=\"http:\/\/biocp04.epgpbooks.inflibnet.ac.in\/wp-content\/uploads\/sites\/215\/2019\/04\/1-144.png\" alt=\"\" width=\"374\" height=\"53\" srcset=\"https:\/\/ebooks.inflibnet.ac.in\/biocp04\/wp-content\/uploads\/sites\/215\/2019\/04\/1-144.png 374w, https:\/\/ebooks.inflibnet.ac.in\/biocp04\/wp-content\/uploads\/sites\/215\/2019\/04\/1-144-300x43.png 300w, https:\/\/ebooks.inflibnet.ac.in\/biocp04\/wp-content\/uploads\/sites\/215\/2019\/04\/1-144-65x9.png 65w, https:\/\/ebooks.inflibnet.ac.in\/biocp04\/wp-content\/uploads\/sites\/215\/2019\/04\/1-144-225x32.png 225w, https:\/\/ebooks.inflibnet.ac.in\/biocp04\/wp-content\/uploads\/sites\/215\/2019\/04\/1-144-350x50.png 350w\" sizes=\"auto, (max-width: 374px) 100vw, 374px\" \/><\/p>\n<p>&nbsp;<\/p>\n<p><img loading=\"lazy\" decoding=\"async\" class=\"aligncenter size-full wp-image-294\" src=\"http:\/\/biocp04.epgpbooks.inflibnet.ac.in\/wp-content\/uploads\/sites\/215\/2019\/04\/1-145.png\" alt=\"\" width=\"517\" height=\"252\" srcset=\"https:\/\/ebooks.inflibnet.ac.in\/biocp04\/wp-content\/uploads\/sites\/215\/2019\/04\/1-145.png 517w, https:\/\/ebooks.inflibnet.ac.in\/biocp04\/wp-content\/uploads\/sites\/215\/2019\/04\/1-145-300x146.png 300w, https:\/\/ebooks.inflibnet.ac.in\/biocp04\/wp-content\/uploads\/sites\/215\/2019\/04\/1-145-65x32.png 65w, https:\/\/ebooks.inflibnet.ac.in\/biocp04\/wp-content\/uploads\/sites\/215\/2019\/04\/1-145-225x110.png 225w, https:\/\/ebooks.inflibnet.ac.in\/biocp04\/wp-content\/uploads\/sites\/215\/2019\/04\/1-145-350x171.png 350w\" sizes=\"auto, (max-width: 517px) 100vw, 517px\" \/><\/p>\n<h2 style=\"text-align: center\"><strong>Figure \u2013 8. Transfer of Proton from the cytoplasm to mitochondria<\/strong><\/h2>\n<h2 style=\"text-align: center\"><strong>( Adapted from- http:\/\/www.ccrc.uga.edu\/~rcarlson\/bcmb3100\/Chap21)<\/strong><\/h2>\n<p><img loading=\"lazy\" decoding=\"async\" class=\"aligncenter size-full wp-image-295\" src=\"http:\/\/biocp04.epgpbooks.inflibnet.ac.in\/wp-content\/uploads\/sites\/215\/2019\/04\/1-146.png\" alt=\"\" width=\"559\" height=\"376\" srcset=\"https:\/\/ebooks.inflibnet.ac.in\/biocp04\/wp-content\/uploads\/sites\/215\/2019\/04\/1-146.png 559w, https:\/\/ebooks.inflibnet.ac.in\/biocp04\/wp-content\/uploads\/sites\/215\/2019\/04\/1-146-300x202.png 300w, https:\/\/ebooks.inflibnet.ac.in\/biocp04\/wp-content\/uploads\/sites\/215\/2019\/04\/1-146-65x44.png 65w, https:\/\/ebooks.inflibnet.ac.in\/biocp04\/wp-content\/uploads\/sites\/215\/2019\/04\/1-146-225x151.png 225w, https:\/\/ebooks.inflibnet.ac.in\/biocp04\/wp-content\/uploads\/sites\/215\/2019\/04\/1-146-350x235.png 350w\" sizes=\"auto, (max-width: 559px) 100vw, 559px\" \/><\/p>\n<h2 style=\"text-align: center\"><strong>Figure- 9 Many Mitochondrial Transporters<\/strong><\/h2>\n<h2 style=\"text-align: center\"><strong>(Adapted from-http:\/\/www.ccrc.uga.edu\/~rcarlson\/bcmb3100\/Chap21)<\/strong><\/h2>\n<p><img loading=\"lazy\" decoding=\"async\" class=\"aligncenter size-full wp-image-296\" src=\"http:\/\/biocp04.epgpbooks.inflibnet.ac.in\/wp-content\/uploads\/sites\/215\/2019\/04\/1-147.png\" alt=\"\" width=\"596\" height=\"379\" srcset=\"https:\/\/ebooks.inflibnet.ac.in\/biocp04\/wp-content\/uploads\/sites\/215\/2019\/04\/1-147.png 596w, https:\/\/ebooks.inflibnet.ac.in\/biocp04\/wp-content\/uploads\/sites\/215\/2019\/04\/1-147-300x191.png 300w, https:\/\/ebooks.inflibnet.ac.in\/biocp04\/wp-content\/uploads\/sites\/215\/2019\/04\/1-147-65x41.png 65w, https:\/\/ebooks.inflibnet.ac.in\/biocp04\/wp-content\/uploads\/sites\/215\/2019\/04\/1-147-225x143.png 225w, https:\/\/ebooks.inflibnet.ac.in\/biocp04\/wp-content\/uploads\/sites\/215\/2019\/04\/1-147-350x223.png 350w\" sizes=\"auto, (max-width: 596px) 100vw, 596px\" \/><\/p>\n<p>&nbsp;<\/p>\n<p><img loading=\"lazy\" decoding=\"async\" class=\"aligncenter size-full wp-image-297\" src=\"http:\/\/biocp04.epgpbooks.inflibnet.ac.in\/wp-content\/uploads\/sites\/215\/2019\/04\/1-148.png\" alt=\"\" width=\"611\" height=\"349\" srcset=\"https:\/\/ebooks.inflibnet.ac.in\/biocp04\/wp-content\/uploads\/sites\/215\/2019\/04\/1-148.png 611w, https:\/\/ebooks.inflibnet.ac.in\/biocp04\/wp-content\/uploads\/sites\/215\/2019\/04\/1-148-300x171.png 300w, https:\/\/ebooks.inflibnet.ac.in\/biocp04\/wp-content\/uploads\/sites\/215\/2019\/04\/1-148-65x37.png 65w, https:\/\/ebooks.inflibnet.ac.in\/biocp04\/wp-content\/uploads\/sites\/215\/2019\/04\/1-148-225x129.png 225w, https:\/\/ebooks.inflibnet.ac.in\/biocp04\/wp-content\/uploads\/sites\/215\/2019\/04\/1-148-350x200.png 350w\" sizes=\"auto, (max-width: 611px) 100vw, 611px\" \/><\/p>\n<p>&nbsp;<\/p>\n<p><img loading=\"lazy\" decoding=\"async\" class=\"aligncenter size-full wp-image-298\" src=\"http:\/\/biocp04.epgpbooks.inflibnet.ac.in\/wp-content\/uploads\/sites\/215\/2019\/04\/1-149.png\" alt=\"\" width=\"460\" height=\"335\" srcset=\"https:\/\/ebooks.inflibnet.ac.in\/biocp04\/wp-content\/uploads\/sites\/215\/2019\/04\/1-149.png 460w, https:\/\/ebooks.inflibnet.ac.in\/biocp04\/wp-content\/uploads\/sites\/215\/2019\/04\/1-149-300x218.png 300w, https:\/\/ebooks.inflibnet.ac.in\/biocp04\/wp-content\/uploads\/sites\/215\/2019\/04\/1-149-65x47.png 65w, https:\/\/ebooks.inflibnet.ac.in\/biocp04\/wp-content\/uploads\/sites\/215\/2019\/04\/1-149-225x164.png 225w, https:\/\/ebooks.inflibnet.ac.in\/biocp04\/wp-content\/uploads\/sites\/215\/2019\/04\/1-149-350x255.png 350w\" sizes=\"auto, (max-width: 460px) 100vw, 460px\" \/><\/p>\n<p>&nbsp;<\/p>\n<p>&nbsp;<\/p>\n<table>\n<tbody>\n<tr>\n<td><strong>you can view video on Proton motive force<\/strong><\/td>\n<td><a href=\"https:\/\/youtu.be\/J660Uo1f0yU\" target=\"_blank\" rel=\"noopener\"><img loading=\"lazy\" decoding=\"async\" class=\"alignnone wp-image-120\" src=\"http:\/\/epgpbooks.inflibnet.ac.in\/wp-content\/uploads\/2018\/11\/download.png\" alt=\"\" width=\"36\" height=\"36\" \/><\/a><\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<p>&nbsp;<\/p>\n<div class=\"textbox learning-objectives\">\n<h3><strong>References<\/strong><\/h3>\n<p><strong>Web site<\/strong><\/p>\n<ul>\n<li><a href=\"http:\/\/www.nature.com\/scitable\/topicpage\/why-are-cells-powered-by-proton-gradients-14373960\">http:\/\/www.nature.com\/scitable\/topicpage\/why-are-cells-powered-by-proton-<\/a><a href=\"http:\/\/www.nature.com\/scitable\/topicpage\/why-are-cells-powered-by-proton-gradients-14373960\">gradients-14373960<\/a><\/li>\n<li><a href=\"http:\/\/www.ccrc.uga.edu\/~rcarlson\/bcmb3100\/Chap21\">http:\/\/www.ccrc.uga.edu\/~rcarlson\/bcmb3100\/Chap21<\/a><\/li>\n<li><a href=\"https:\/\/en.wikipedia.org\/wiki\/Chemiosmosis\">https:\/\/en.wikipedia.org\/wiki\/Chemiosmosis<\/a><\/li>\n<li><a href=\"http:\/\/www.biologyaspoetry.com\/terms\/proton_motive_force.html\">http:\/\/www.biologyaspoetry.com\/terms\/proton_motive_force.html<\/a><\/li>\n<\/ul>\n<p>&nbsp;<\/p>\n<p><strong>Books<\/strong><\/p>\n<ul>\n<li style=\"text-align: justify\">Harper\u2019sIllustrated Biochemistry by Robert K.. Murray, Daryl K. Granner, Peter A. Mayes, 26th Edition (2003)<\/li>\n<li style=\"text-align: justify\">Lehninger\u2019S Principle of Biochemistry David L. Nelson and Michael M. Cox,5th Edition (2008)<\/li>\n<li style=\"text-align: justify\">Instant Notes: Biochemistry, 2nd Edition, B.D. Hames &amp; N. M. Hooper (2005)<\/li>\n<\/ul>\n<p style=\"text-align: justify\">\n<\/div>\n<p>&nbsp;<\/p>\n<p>&nbsp;<\/p>\n","protected":false},"author":3,"menu_order":18,"template":"","meta":{"pb_show_title":"on","pb_short_title":"","pb_subtitle":"","pb_authors":["dr-padma-ambalam"],"pb_section_license":""},"chapter-type":[],"contributor":[60],"license":[],"class_list":["post-278","chapter","type-chapter","status-publish","hentry","contributor-dr-padma-ambalam"],"part":3,"_links":{"self":[{"href":"https:\/\/ebooks.inflibnet.ac.in\/biocp04\/wp-json\/pressbooks\/v2\/chapters\/278","targetHints":{"allow":["GET"]}}],"collection":[{"href":"https:\/\/ebooks.inflibnet.ac.in\/biocp04\/wp-json\/pressbooks\/v2\/chapters"}],"about":[{"href":"https:\/\/ebooks.inflibnet.ac.in\/biocp04\/wp-json\/wp\/v2\/types\/chapter"}],"author":[{"embeddable":true,"href":"https:\/\/ebooks.inflibnet.ac.in\/biocp04\/wp-json\/wp\/v2\/users\/3"}],"version-history":[{"count":6,"href":"https:\/\/ebooks.inflibnet.ac.in\/biocp04\/wp-json\/pressbooks\/v2\/chapters\/278\/revisions"}],"predecessor-version":[{"id":304,"href":"https:\/\/ebooks.inflibnet.ac.in\/biocp04\/wp-json\/pressbooks\/v2\/chapters\/278\/revisions\/304"}],"part":[{"href":"https:\/\/ebooks.inflibnet.ac.in\/biocp04\/wp-json\/pressbooks\/v2\/parts\/3"}],"metadata":[{"href":"https:\/\/ebooks.inflibnet.ac.in\/biocp04\/wp-json\/pressbooks\/v2\/chapters\/278\/metadata\/"}],"wp:attachment":[{"href":"https:\/\/ebooks.inflibnet.ac.in\/biocp04\/wp-json\/wp\/v2\/media?parent=278"}],"wp:term":[{"taxonomy":"chapter-type","embeddable":true,"href":"https:\/\/ebooks.inflibnet.ac.in\/biocp04\/wp-json\/pressbooks\/v2\/chapter-type?post=278"},{"taxonomy":"contributor","embeddable":true,"href":"https:\/\/ebooks.inflibnet.ac.in\/biocp04\/wp-json\/wp\/v2\/contributor?post=278"},{"taxonomy":"license","embeddable":true,"href":"https:\/\/ebooks.inflibnet.ac.in\/biocp04\/wp-json\/wp\/v2\/license?post=278"}],"curies":[{"name":"wp","href":"https:\/\/api.w.org\/{rel}","templated":true}]}}