Electron transport chain - wikipedia Electron transport chain Jump to : navigation , search The electron transport chain in the mitochondrion is the site of oxidative phosphorylation in eukaryotes . The NADH and succinate generated in the citric acid cycle are oxidized , providing energy to power ATP synthase . Photosynthetic electron transport chain of the thylakoid membrane . An electron transport chain ( ETC ) is a series of complexes that transfer electrons from electron donors to electron acceptors via redox ( both reduction and oxidation occurring simultaneously ) reactions , and couples this electron transfer with the transfer of protons ( H ions ) across a membrane . This creates an electrochemical proton gradient that drives the synthesis of adenosine triphosphate ( ATP ) , a molecule that stores energy chemically in the form of highly strained bonds . The molecules of the chain include peptides , enzymes ( which are proteins or protein complexes ) , and others . The final acceptor of electrons in the electron transport chain during aerobic respiration is molecular oxygen although a variety of acceptors other than oxygen such as sulfate exist in anaerobic respiration . Electron transport chains are used for extracting energy via redox reactions from sunlight in photosynthesis or , such as in the case of the oxidation of sugars , cellular respiration . In eukaryotes , an important electron transport chain is found in the inner mitochondrial membrane where it serves as the site of oxidative phosphorylation through the action of ATP synthase . It is also found in the thylakoid membrane of the chloroplast in photosynthetic eukaryotes . In bacteria , the electron transport chain is located in their cell membrane . In chloroplasts , light drives the conversion of water to oxygen and NADP to NADPH with transfer of H ions across chloroplast membranes . In mitochondria , it is the conversion of oxygen to water , NADH to NAD and succinate to fumarate that are required to generate the proton gradient . Electron transport chains are major sites of premature electron leakage to oxygen , generating superoxide and potentially resulting in increased oxidative stress . Contents ( hide ) 1 Background 2 In mitochondria 2.1 Mitochondrial redox carriers 2.1. 1 Complex I 2.1. 2 Complex II 2.1. 3 Complex III 2.1. 4 Complex IV 2.2 Coupling with oxidative phosphorylation 2.3 Summary 3 In bacteria 3.1 Electron donors 3.2 Dehydrogenases 3.3 Quinone carriers 3.4 Proton pumps 3.5 Cytochrome electron carriers 3.6 Terminal oxidases and reductases 3.7 Electron acceptors 3.8 Summary 4 Photosynthetic 5 Summary 6 See also 7 References 8 External links Background ( edit ) The electron transport chain consists of a spatially separated series of redox reactions in which electrons are transferred from a donor molecule to an acceptor molecule . The underlying force driving these reactions is the Gibbs free energy of the reactants and products . The Gibbs free energy is the energy available ( `` free '' ) to do work . Any reaction that decreases the overall Gibbs free energy of a system is thermodynamically spontaneous . The function of the electron transport chain is to produce a transmembrane proton electrochemical gradient as a result of the redox reactions . If protons flow back through the membrane , they enable mechanical work , such as rotating bacterial flagella . ATP synthase , an enzyme highly conserved among all domains of life , converts this mechanical work into chemical energy by producing ATP , which powers most cellular reactions . A small amount of ATP is available from substrate - level phosphorylation , for example , in glycolysis . In most organisms the majority of ATP is generated in electron transport chains , while only some obtain ATP by fermentation . In mitochondria ( edit ) Most eukaryotic cells have mitochondria , which produce ATP from products of the citric acid cycle , fatty acid oxidation , and amino acid oxidation . At the mitochondrial inner membrane , electrons from NADH and FADH2 pass through the electron transport chain to oxygen , which is reduced to water . The electron transport chain comprises an enzymatic series of electron donors and acceptors . Each electron donor will pass electrons to a more electronegative acceptor , which in turn donates these electrons to another acceptor , a process that continues down the series until electrons are passed to oxygen , the most electronegative and terminal electron acceptor in the chain . Passage of electrons between donor and acceptor releases energy , which is used to generate a proton gradient across the mitochondrial membrane by actively `` pumping '' protons into the intermembrane space , producing a thermodynamic state that has the potential to do work . The entire process is called oxidative phosphorylation , since ADP is phosphorylated to ATP using the energy of hydrogen oxidation in many steps . A small percentage of electrons do not complete the whole series and instead directly leak to oxygen , resulting in the formation of the free - radical superoxide , a highly reactive molecule that contributes to oxidative stress and has been implicated in a number of diseases and aging . Mitochondrial redox carriers ( edit ) Energy obtained through the transfer of electrons down the ETC is used to pump protons from the mitochondrial matrix into the intermembrane space , creating an electrochemical proton gradient ( ΔpH ) across the inner mitochondrial membrane ( IMM ) . This proton gradient is largely but not exclusively responsible for the mitochondrial membrane potential ( ΔΨ ) . It allows ATP synthase to use the flow of H through the enzyme back into the matrix to generate ATP from adenosine diphosphate ( ADP ) and inorganic phosphate . Complex I ( NADH coenzyme Q reductase ; labeled I ) accepts electrons from the Krebs cycle electron carrier nicotinamide adenine dinucleotide ( NADH ) , and passes them to coenzyme Q ( ubiquinone ; labeled Q ) , which also receives electrons from complex II ( succinate dehydrogenase ; labeled II ) . Q passes electrons to complex III ( cytochrome bc complex ; labeled III ) , which passes them to cytochrome c ( cyt c ) . Cyt c passes electrons to Complex IV ( cytochrome c oxidase ; labeled IV ) , which uses the electrons and hydrogen ions to reduce molecular oxygen to water . Four membrane - bound complexes have been identified in mitochondria . Each is an extremely complex transmembrane structure that is embedded in the inner membrane . Three of them are proton pumps . The structures are electrically connected by lipid - soluble electron carriers and water - soluble electron carriers . The overall electron transport chain : NADH + H → Complex I → Q → Complex III → cytochrome c → Complex IV → H O ↑ Complex II ↑ Succinate Complex I ( edit ) In Complex I ( NADH : ubiquinone oxidoreductase , NADH - CoQ reductase , or NADH dehydrogenase ; EC 1.6. 5.3 ) , two electrons are removed from NADH and ultimately transferred to a lipid - soluble carrier , ubiquinone ( UQ ) . The reduced product , ubiquinol ( UQH ) , freely diffuses within the membrane , and Complex I translocates four protons ( H ) across the membrane , thus producing a proton gradient . Complex I is one of the main sites at which premature electron leakage to oxygen occurs , thus being one of the main sites of production of superoxide . The pathway of electrons is as follows : NADH is oxidized to NAD , by reducing Flavin mononucleotide to FMNH in one two - electron step . FMNH is then oxidized in two one - electron steps , through a semiquinone intermediate . Each electron thus transfers from the FMNH to an Fe - S cluster , from the Fe - S cluster to ubiquinone ( Q ) . Transfer of the first electron results in the free - radical ( semiquinone ) form of Q , and transfer of the second electron reduces the semiquinone form to the ubiquinol form , QH . During this process , four protons are translocated from the mitochondrial matrix to the intermembrane space . As the electrons become continuously oxidized and reduced throughout the complex an electron current is produced along the 180 Angstrom width of the complex within the membrane . This current powers the active transport of four protons to the intermembrane space per two electrons from NADH . This complex is inhibited by Alkylguanides ( Example : Guanethidine ) , Rotenone , Barbiturates , Chlorpromazine , Piericidin . Complex II ( edit ) In Complex II ( succinate dehydrogenase or succinate - CoQ reductase ; EC 1.3. 5.1 ) additional electrons are delivered into the quinone pool ( Q ) originating from succinate and transferred ( via flavin adenine dinucleotide ( FAD ) ) to Q. Complex II consists of four protein subunits : succinate dehydrogenase , ( SDHA ) ; succinate dehydrogenase ( ubiquinone ) iron - sulfur subunit , mitochondrial , ( SDHB ) ; succinate dehydrogenase complex subunit C , ( SDHC ) and succinate dehydrogenase complex , subunit D , ( SDHD ) . Other electron donors ( e.g. , fatty acids and glycerol 3 - phosphate ) also direct electrons into Q ( via FAD ) . Complex 2 is a parallel electron transport pathway to complex 1 , but unlike complex 1 , no protons are transported to the intermembrane space in this pathway . Therefore , the pathway through complex 2 contributes less energy to the overall electron transport chain process . This complex is inhibited by Carboxin . Complex III ( edit ) In Complex III ( cytochrome bc complex or CoQH - cytochrome c reductase ; EC 1.10. 2.2 ) , the Q - cycle contributes to the proton gradient by an asymmetric absorption / release of protons . Two electrons are removed from QH at the Q site and sequentially transferred to two molecules of cytochrome c , a water - soluble electron carrier located within the intermembrane space . The two other electrons sequentially pass across the protein to the Q site where the quinone part of ubiquinone is reduced to quinol . A proton gradient is formed by one quinol ( 2 H 2 + e − ( \ displaystyle ( \ ce ( 2H + 2e - ) ) ) ) oxidations at the Q site to form one quinone ( 2 H 2 + e − ( \ displaystyle ( \ ce ( 2H + 2e - ) ) ) ) at the Q site . ( in total four protons are translocated : two protons reduce quinone to quinol and two protons are released from two ubiquinol molecules ) . QH 2 + 2 cytochrome c ( Fe III ) + 2 H + ( in ) ⟶ Q + 2 cytochrome c ( Fe II ) + 4 H + ( out ) ( \ displaystyle ( \ ce ( QH2 + 2 cytochrome c ( Fe ^ ( III ) ) + 2 H+ ( in ) - > Q + 2 cytochrome c ( Fe ^ ( II ) ) + 4 H+ ( out ) ) ) ) When electron transfer is reduced ( by a high membrane potential or respiratory inhibitors such as antimycin A ) , Complex III may leak electrons to molecular oxygen , resulting in superoxide formation . This complex is inhibited by British Antilewisite ( BAL ) , Napthoquinone and Antimycin . Complex IV ( edit ) In Complex IV ( cytochrome c oxidase ; EC 1.9. 3.1 ) , sometimes called cytochrome AA3 , four electrons are removed from four molecules of cytochrome c and transferred to molecular oxygen ( O ) , producing two molecules of water . At the same time , eight protons are removed from the mitochondrial matrix ( although only four are translocated across the membrane ) , contributing to the proton gradient . The activity of cytochrome c oxidase is inhibited by cyanide , carbon monoxide , azide , hydrogen sulphide ( H S ) . Coupling with oxidative phosphorylation ( edit ) Depiction of ATP synthase , the site of oxidative phosphorylation to generate ATP . According to the chemiosmotic coupling hypothesis , proposed by Nobel Prize in Chemistry winner Peter D. Mitchell , the electron transport chain and oxidative phosphorylation are coupled by a proton gradient across the inner mitochondrial membrane . The efflux of protons from the mitochondrial matrix creates an electrochemical gradient ( proton gradient ) . This gradient is used by the F F ATP synthase complex to make ATP via oxidative phosphorylation . ATP synthase is sometimes described as Complex V of the electron transport chain . The F component of ATP synthase acts as an ion channel that provides for a proton flux back into the mitochondrial matrix . It is composed of a , b and c subunits . Protons in the inter-membranous space of mitochondria first enters the ATP synthase complex through a subunit channel . Then protons move to the c subunits . The number of c subunits it has determines how many protons it will require to make the F turn one full revolution . For example , in humans , there are 8 c subunits , thus 8 protons are required . After c subunits , protons finally enters matrix using a subunit channel that opens into the mitochondrial matrix . This reflux releases free energy produced during the generation of the oxidized forms of the electron carriers ( NAD and Q ) . The free energy is used to drive ATP synthesis , catalyzed by the F component of the complex . Coupling with oxidative phosphorylation is a key step for ATP production . However , in specific cases , uncoupling the two processes may be biologically useful . The uncoupling protein , thermogenin -- present in the inner mitochondrial membrane of brown adipose tissue -- provides for an alternative flow of protons back to the inner mitochondrial matrix . Thyroxine is also a natural uncoupler . This alternative flow results in thermogenesis rather than ATP production . Synthetic uncouplers ( e.g. , 2 , 4 - dinitrophenol , 2 , 4 - dinitrocresol , CCCP ) also exist , and can be lethal at high doses . Summary ( edit ) In the mitochondrial electron transport chain electrons move from an electron donor ( NADH or QH ) to a terminal electron acceptor ( O ) via a series of redox reactions . These reactions are coupled to the creation of a proton gradient across the mitochondrial inner membrane . There are three proton pumps : I , III , and IV . The resulting transmembrane proton gradient is used to make ATP via ATP synthase . The reactions catalyzed by Complex I and Complex III work roughly at equilibrium . This means that these reactions are readily reversible , by increasing the concentration of the products relative to the concentration of the reactants ( for example , by increasing the proton gradient ) . ATP synthase is also readily reversible . Thus ATP can be used to build a proton gradient , which in turn can be used to make NADH . This process of reverse electron transport is important in many prokaryotic electron transport chains . In bacteria ( edit ) In eukaryotes , NADH is the most important electron donor . The associated electron transport chain is NADH → Complex I → Q → Complex III → cytochrome c → Complex IV → O where Complexes I , III and IV are proton pumps , while Q and cytochrome c are mobile electron carriers . The electron acceptor is molecular oxygen . In prokaryotes ( bacteria and archaea ) the situation is more complicated , because there are several different electron donors and several different electron acceptors . The generalized electron transport chain in bacteria is : Donor Donor Donor ↓ ↓ ↓ dehydrogenase → quinone → bc → cytochrome ↓ ↓ oxidase ( reductase ) oxidase ( reductase ) ↓ ↓ Acceptor Acceptor Note that electrons can enter the chain at three levels : at the level of a dehydrogenase , at the level of the quinone pool , or at the level of a mobile cytochrome electron carrier . These levels correspond to successively more positive redox potentials , or to successively decreased potential differences relative to the terminal electron acceptor . In other words , they correspond to successively smaller Gibbs free energy changes for the overall redox reaction Donor → Acceptor . Individual bacteria use multiple electron transport chains , often simultaneously . Bacteria can use a number of different electron donors , a number of different dehydrogenases , a number of different oxidases and reductases , and a number of different electron acceptors . For example , E. coli ( when growing aerobically using glucose as an energy source ) uses two different NADH dehydrogenases and two different quinol oxidases , for a total of four different electron transport chains operating simultaneously . A common feature of all electron transport chains is the presence of a proton pump to create a transmembrane proton gradient . Bacterial electron transport chains may contain as many as three proton pumps , like mitochondria , or they may contain only one or two . They always contain at least one proton pump . Electron donors ( edit ) In the present day biosphere , the most common electron donors are organic molecules . Organisms that use organic molecules as an energy source are called organotrophs . Organotrophs ( animals , fungi , protists ) and phototrophs ( plants and algae ) constitute the vast majority of all familiar life forms . Some prokaryotes can use inorganic matter as an energy source . Such an organism is called a lithotroph ( `` rock - eater '' ) . Inorganic electron donors include hydrogen , carbon monoxide , ammonia , nitrite , sulfur , sulfide , manganese oxide , and ferrous iron . Lithotrophs have been found growing in rock formations thousands of meters below the surface of Earth . Because of their volume of distribution , lithotrophs may actually outnumber organotrophs and phototrophs in our biosphere . The use of inorganic electron donors as an energy source is of particular interest in the study of evolution . This type of metabolism must logically have preceded the use of organic molecules as an energy source . Dehydrogenases ( edit ) Bacteria can use a number of different electron donors . When organic matter is the energy source , the donor may be NADH or succinate , in which case electrons enter the electron transport chain via NADH dehydrogenase ( similar to Complex I in mitochondria ) or succinate dehydrogenase ( similar to Complex II ) . Other dehydrogenases may be used to process different energy sources : formate dehydrogenase , lactate dehydrogenase , glyceraldehyde - 3 - phosphate dehydrogenase , H dehydrogenase ( hydrogenase ) , etc . Some dehydrogenases are also proton pumps ; others funnel electrons into the quinone pool . Most dehydrogenases show induced expression in the bacterial cell in response to metabolic needs triggered by the environment in which the cells grow . Quinone carriers ( edit ) Quinones are mobile , lipid - soluble carriers that shuttle electrons ( and protons ) between large , relatively immobile macromolecular complexes embedded in the membrane . Bacteria use ubiquinone ( the same quinone that mitochondria use ) and related quinones such as menaquinone . Another name for ubiquinone is Coenzyme Q10 . Proton pumps ( edit ) A proton pump is any process that creates a proton gradient across a membrane . Protons can be physically moved across a membrane ; this is seen in mitochondrial Complexes I and IV . The same effect can be produced by moving electrons in the opposite direction . The result is the disappearance of a proton from the cytoplasm and the appearance of a proton in the periplasm . Mitochondrial Complex III uses this second type of proton pump , which is mediated by a quinone ( the Q cycle ) . Some dehydrogenases are proton pumps ; others are not . Most oxidases and reductases are proton pumps , but some are not . Cytochrome bc is a proton pump found in many , but not all , bacteria ( it is not found in E. coli ) . As the name implies , bacterial bc is similar to mitochondrial bc ( Complex III ) . Proton pumps are the heart of the electron transport process . They produce the transmembrane electrochemical gradient that enables ATP Synthase to synthesize ATP . Cytochrome electron carriers ( edit ) Cytochromes are pigments that contain iron . They are found in two very different environments . Some cytochromes are water - soluble carriers that shuttle electrons to and from large , immobile macromolecular structures imbedded in the membrane . The mobile cytochrome electron carrier in mitochondria is cytochrome c . Bacteria use a number of different mobile cytochrome electron carriers . Other cytochromes are found within macromolecules such as Complex III and Complex IV . They also function as electron carriers , but in a very different , intramolecular , solid - state environment . Electrons may enter an electron transport chain at the level of a mobile cytochrome or quinone carrier . For example , electrons from inorganic electron donors ( nitrite , ferrous iron , etc . ) enter the electron transport chain at the cytochrome level . When electrons enter at a redox level greater than NADH , the electron transport chain must operate in reverse to produce this necessary , higher - energy molecule . Terminal oxidases and reductases ( edit ) When bacteria grow in aerobic environments , the terminal electron acceptor ( O ) is reduced to water by an enzyme called an oxidase . When bacteria grow in anaerobic environments , the terminal electron acceptor is reduced by an enzyme called a reductase . In mitochondria the terminal membrane complex ( Complex IV ) is cytochrome oxidase . Aerobic bacteria use a number of different terminal oxidases . For example , E. coli does not have a cytochrome oxidase or a bc complex . Under aerobic conditions , it uses two different terminal quinol oxidases ( both proton pumps ) to reduce oxygen to water . Anaerobic bacteria , which do not use oxygen as a terminal electron acceptor , have terminal reductases individualized to their terminal acceptor . For example , E. coli can use fumarate reductase , nitrate reductase , nitrite reductase , DMSO reductase , or trimethylamine - N - oxide reductase , depending on the availability of these acceptors in the environment . Most terminal oxidases and reductases are inducible . They are synthesized by the organism as needed , in response to specific environmental conditions . Electron acceptors ( edit ) Just as there are a number of different electron donors ( organic matter in organotrophs , inorganic matter in lithotrophs ) , there are a number of different electron acceptors , both organic and inorganic . If oxygen is available , it is invariably used as the terminal electron acceptor , because it generates the greatest Gibbs free energy change and produces the most energy . In anaerobic environments , different electron acceptors are used , including nitrate , nitrite , ferric iron , sulfate , carbon dioxide , and small organic molecules such as fumarate . Since electron transport chains are redox processes , they can be described as the sum of two redox pairs . For example , the mitochondrial electron transport chain can be described as the sum of the NAD / NADH redox pair and the O / H O redox pair . NADH is the electron donor and O is the electron acceptor . Not every donor - acceptor combination is thermodynamically possible . The redox potential of the acceptor must be more positive than the redox potential of the donor . Furthermore , actual environmental conditions may be far different from standard conditions ( 1 molar concentrations , 1 atm partial pressures , pH = 7 ) , which apply to standard redox potentials . For example , hydrogen - evolving bacteria grow at an ambient partial pressure of hydrogen gas of 10 atm . The associated redox reaction , which is thermodynamically favorable in nature , is thermodynamic impossible under `` standard '' conditions . Summary ( edit ) Bacterial electron transport pathways are , in general , inducible . Depending on their environment , bacteria can synthesize different transmembrane complexes and produce different electron transport chains in their cell membranes . Bacteria select their electron transport chains from a DNA library containing multiple possible dehydrogenases , terminal oxidases and terminal reductases . The situation is often summarized by saying that electron transport chains in bacteria are branched , modular , and inducible . Photosynthetic ( edit ) In oxidative phosphorylation , electrons are transferred from a low - energy electron donor ( e.g. , NADH ) to an acceptor ( e.g. , O ) through an electron transport chain . In photophosphorylation , the energy of sunlight is used to create a high - energy electron donor and an electron acceptor . Electrons are then transferred from the donor to the acceptor through another electron transport chain . Photosynthetic electron transport chains have many similarities to the oxidative chains discussed above . They use mobile , lipid - soluble carriers ( quinones ) and mobile , water - soluble carriers ( cytochromes , etc . ) . They also contain a proton pump . It is remarkable that the proton pump in all photosynthetic chains resembles mitochondrial Complex III . Photosynthetic electron transport chains are discussed in greater detail in the articles Photophosphorylation , Photosynthesis , Photosynthetic reaction center and Light - dependent reaction . Summary ( edit ) Electron transport chains are redox reactions that transfer electrons from an electron donor to an electron acceptor . The transfer of electrons is coupled to the translocation of protons across a membrane , producing a proton gradient . The proton gradient is used to produce useful work . About 30 work units are produced per electron transport . See also ( edit ) CoRR hypothesis Electron equivalent Hydrogen hypothesis Respirasome References ( edit ) Jump up ^ Murray , Robert K. ; Daryl K. Granner ; Peter A. Mayes ; Victor W. Rodwell ( 2003 ) . Harper 's Illustrated Biochemistry . New York , NY : Lange Medical Books / MgGraw Hill . p. 96 . ISBN 0 - 07 - 121766 - 5 . Jump up ^ Karp , Gerald ( 2008 ) . Cell and Molecular Biology ( 5th ed . ) . Hoboken , NJ : John Wiley & Sons . p. 194 . ISBN 978 - 0 - 470 - 04217 - 5 . Jump up ^ Lauren , Biochemistry , Johnson / Cole , 2010 , pp 598 - 611 Jump up ^ Garrett & Grisham , Biochemistry , Brooks / Cole , 2010 , pp 598 - 611 Jump up ^ Garret and Grisham ( 2016 ) . biochemistry . University of Virginia . p. 687 . ISBN 978 - 1 - 305 - 57720 - 6 . Jump up ^ Jonckheere , An I. ; Smeitink , Jan A.M. ; Rodenburg , Richard J.T. ( 2017 - 03 - 10 ) . `` Mitochondrial ATP synthase : architecture , function and pathology '' . Journal of Inherited Metabolic Disease . 35 ( 2 ) : 211 -- 225 . doi : 10.1007 / s10545 - 011 - 9382 - 9 . ISSN 0141 - 8955 . PMC 3278611 . PMID 21874297 . ^ Jump up to : Garrett , Reginald H. ; Grisham , Charles M. ( 2012 ) . Biochemistry ( 5th ed . ) . Cengage learning . p. 664 . ISBN 1 - 133 - 10629 - 3 . Jump up ^ Fillingame , Robert H ; Angevine , Christine M ; Dmitriev , Oleg Y ( 2003 - 11 - 27 ) . `` Mechanics of coupling proton movements to c - ring rotation in ATP synthase '' . FEBS Letters. 555 ( 1 ) : 29 -- 34 . doi : 10.1016 / S0014 - 5793 ( 03 ) 01101 - 3 . ISSN 1873 - 3468 . Jump up ^ Berg , Jeremy M. ; Tymoczko , John L. ; Stryer , Lubert ( 2002 - 01 - 01 ) . `` A Proton Gradient Powers the Synthesis of ATP '' . Jump up ^ Cannon , Barbara ; Nedergaard , Jan ( 2004 - 01 - 01 ) . `` Brown Adipose Tissue : Function and Physiological Significance '' . Physiological Reviews . 84 ( 1 ) : 277 -- 359 . doi : 10.1152 / physrev. 00015.2003 . ISSN 0031 - 9333 . PMID 14715917 . Jump up ^ Alberts , Bruce ; Johnson , Alexander ; Lewis , Julian ; Raff , Martin ; Roberts , Keith ; Walter , Peter ( 2002 - 01 - 01 ) . `` Electron - Transport Chains and Their Proton Pumps '' . Fenchel T ; King GM ; Blackburn TH ( September 2006 ) . Bacterial Biogeochemistry : The Ecophysiology of Mineral Cycling ( 2nd ed . ) . Elsevier . ISBN 978 - 0 - 12 - 103455 - 9 . Lengeler JW ( January 1999 ) . Drews G ; Schlegel HG , eds . Biology of the Prokaryotes . Blackwell Science . ISBN 978 - 0 - 632 - 05357 - 5 . Nelson DL ; Cox MM ( April 2005 ) . Lehninger Principles of Biochemistry ( 4th ed . ) . W.H. Freeman . ISBN 978 - 0 - 7167 - 4339 - 2 . Nicholls DG ; Ferguson SJ ( July 2002 ) . Bioenergetics 3 . Academic Press . ISBN 978 - 0 - 12 - 518121 - 1 . Stumm W ; Morgan JJ ( 1996 ) . Aquatic Chemistry ( 3rd ed . ) . John Wiley & Sons . ISBN 978 - 0 - 471 - 51185 - 4 . Thauer RK ; Jungermann K ; Decker K ( March 1977 ) . `` Energy conservation in chemotrophic anaerobic bacteria '' . Bacteriol Rev. 41 ( 1 ) : 100 -- 80 . PMC 413997 . PMID 860983 . White D. ( September 1999 ) . The Physiology and Biochemistry of Prokaryotes ( 2nd ed . ) . Oxford University Press . ISBN 978 - 0 - 19 - 512579 - 5 . Voet D ; Voet JG ( March 2004 ) . Biochemistry ( 3rd ed . ) . John Wiley & Sons . ISBN 978 - 0 - 471 - 58651 - 7 . Kim HS. ; Patel , K ; Muldoon - Jacobs , K ; Bisht , KS ; Aykin - Burns , N ; Pennington , JD ; Van Der Meer , R ; Nguyen , P ; et al. ( January 2010 ) . `` SIRT3 is a mitochondria - localized tumor suppressor required for maintenance of mitochondrial integrity and metabolism during stress '' . Cancer Cell. 17 ( 1 ) : 41 -- 52 . doi : 10.1016 / j. ccr. 2009.11. 023 . PMC 3711519 . PMID 20129246 . External links ( edit ) Electron Transport Chain Complex Proteins at the US National Library of Medicine Medical Subject Headings ( MeSH ) Khan Academy , video lecture UMich Orientation of Proteins in Membranes families / superfamily - 3 - Complexes with cytochrome b - like domains UMich Orientation of Proteins in Membranes families / superfamily - 4 - Bacterial and mitochondrial cytochrome c oxidases UMich Orientation of Proteins in Membranes families / superfamily - 2 - Photosynthetic reaction centers and photosystems UMich Orientation of Proteins in Membranes families / superfamily - 78 - Cytochrome PORN renodoxin reductase UMich Orientation of Proteins in Membranes families / superfamily - 130 - Electron transfer flavoproteins Metabolism , catabolism , anabolism General Metabolic pathway Metabolic network Primary nutritional groups Energy metabolism Aerobic respiration Glycolysis → Pyruvate decarboxylation → Citric acid cycle → Oxidative phosphorylation ( electron transport chain + ATP synthase ) Anaerobic respiration Electron acceptors are other than oxygen Fermentation Glycolysis → Substrate - level phosphorylation ABE Ethanol Lactic acid Specific paths Protein metabolism Protein synthesis Catabolism Carbohydrate metabolism ( carbohydrate catabolism and anabolism ) Human Glycolysis ⇄ Gluconeogenesis Glycogenolysis ⇄ Glycogenesis Pentose phosphate pathway Fructolysis Galactolysis Glycosylation N - linked O - linked Nonhuman Photosynthesis Anoxygenic photosynthesis Chemosynthesis Carbon fixation Xylose metabolism Radiotrophism Lipid metabolism ( lipolysis , lipogenesis ) Fatty acid metabolism Fatty acid degradation ( Beta oxidation ) Fatty acid synthesis Other Steroid metabolism Sphingolipid metabolism Eicosanoid metabolism Ketosis Reverse cholesterol transport Amino acid Amino acid synthesis Urea cycle Nucleotide metabolism Purine metabolism Nucleotide salvage Pyrimidine metabolism Other Metal metabolism Iron metabolism Ethanol metabolism Metabolism : Citric acid cycle enzymes Cycle Citrate synthase Aconitase Isocitrate dehydrogenase Oxoglutarate dehydrogenase Succinyl CoA synthetase Succinate dehydrogenase ( SDHA ) Fumarase Malate dehydrogenase and ETC Anaplerotic to acetyl - CoA Pyruvate dehydrogenase complex ( E1 , E2 , E3 ) ( regulated by Pyruvate dehydrogenase kinase and Pyruvate dehydrogenase phosphatase ) to α - ketoglutaric acid Glutamate dehydrogenase to succinyl - CoA Methylmalonyl - CoA mutase to oxaloacetate Pyruvate carboxylase Aspartate transaminase Mitochondrial electron transport chain / oxidative phosphorylation Primary Complex I / NADH dehydrogenase Complex II / Succinate dehydrogenase Coenzyme Q Complex III / Coenzyme Q - cytochrome c reductase Cytochrome c Complex IV / Cytochrome c oxidase Coenzyme Q10 synthesis : COQ2 COQ3 COQ4 COQ5 COQ6 COQ7 COQ9 COQ10A COQ10B PDSS1 PDSS2 Other Alternative oxidase Electron - transferring - flavoprotein dehydrogenase Retrieved from `` https://en.wikipedia.org/w/index.php?title=Electron_transport_chain&oldid=837323998 '' Categories : Cellular respiration Integral membrane proteins Hidden categories : All articles with unsourced statements Articles with unsourced statements from May 2010 Articles with unsourced statements from August 2012 Articles with unsourced statements from November 2011 Articles with unsourced statements from April 2015 Talk Contents About Wikipedia Български Bosanski Català Čeština Dansk Deutsch Español فارسی Français Galego 한국어 Bahasa Indonesia Italiano Magyar Bahasa Melayu 日本 語 Norsk Русский Simple English Српски / srpski Srpskohrvatski / српскохрватски Suomi Svenska ไทย Türkçe Українська Tiếng Việt 中文 20 more Edit links This page was last edited on 20 April 2018 , at 03 : 12 . 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Question: where does the electron transport chain get its electrons from


Four - wheel drive - wikipedia Four - wheel drive Jump to : navigation , search For other uses , see Four - wheel drive ( disambiguation ) . `` Four by four '' redirects here . For other uses , see Four by four ( disambiguation ) . The Jeep Wrangler is a 4WD vehicle with a transfer case to select low range or high range four - wheel drive Four - wheel drive , also called 4 × 4 ( `` four by four '' ) or 4WD , refers a two - axled vehicle with a drivetrain capable of providing torque to all its wheels simultaneously . It may be full - time or on - demand , and is typically linked via a transfer case providing an additional output drive - shaft and , in many instances , additional gear ranges . A four - wheeled vehicle with torque supplied to both axles is described as `` all - wheel drive '' ( AWD ) . However , `` four - wheel drive '' typically refers to a set of specific components and functions , and / or intended offroad application , which generally complies with modern use of the terminology . Contents ( hide ) 1 Definitions 1.1 4 × 4 1.2 4WD 1.3 AWD 1.4 IWD 1.5 SAE Recommended Practices 2 Design 2.1 Differentials 2.2 Limiting slippage 2.3 Operating Modes 3 History 3.1 Late 1800s 3.2 1900s - 1920s 3.3 1930s -- 1944 3.4 1945 - 1960s 3.5 1970s - 1990s 3.6 2000 - present 4 Uses 4.1 Road racing 4.2 In heavy trucks 4.3 In construction equipment 5 Terminology 6 Unusual systems 7 Introduction to off - roaders 8 Introduction to passenger cars 9 Systems by design type 9.1 Center differential with mechanical lock 9.2 Torsen center differential 9.3 Non-locking center differential 9.4 Multiple - clutch systems 9.5 Multi-plate clutch coupling 9.6 Part - time 10 See also 11 References 12 External links Definitions ( edit ) 4 × 4 / 4WD / AWD systems were developed in many different markets and used in many different vehicle platforms . There is no universally accepted set of terminology to describe the various architectures and functions . The terms used by various manufacturers often reflect marketing rather than engineering considerations or significant technical differences between systems . SAE International 's standard J1952 recommends only the term All - Wheel - Drive with additional sub classifications which cover all types of AWD / 4WD / 4x4 systems found on production vehicles . 4 × 4 ( edit ) Four - by - four ( 4 × 4 ) refers to the general class of vehicles . The first figure represents the total wheels ( more precisely , axle ends ) , and the second , the number that are powered . Syntactically , 4 × 2 means a four - wheel vehicle that transmits engine torque to only two axle - ends : the front two in front - wheel drive or the rear two in rear - wheel drive . Alternatively , a 6 × 4 vehicle has three axles , two of which provide torque to two wheel ends each . 4WD ( edit ) Four wheel drive ( 4WD ) refers to vehicles with two axles providing torque to four wheel ends . In the North American market the term generally refers to a system that is optimized for off - road driving conditions . The term `` 4WD '' is typically designated for vehicles equipped with a transfercase which switches between 2WD and 4WD operating modes , either manually or automatically . AWD ( edit ) Main article : AWD ( vehicle ) All wheel drive ( AWD ) historically was synonymous with `` four - wheel drive '' on four - wheeled vehicles , and six - wheel drive on 6 × 6s , and so on , being used in that fashion at least as early as the 1920s . Today in North America the term is applied to both heavy vehicles as well as light passenger vehicles . When referring to heavy vehicles the term is increasingly applied to mean `` permanent multiple - wheel drive '' on 2 × 2 , 4 × 4 , 6 × 6 or 8 × 8 drive train systems that include a differential between the front and rear drive shafts . This is often coupled with some sort of anti-slip technology , increasingly hydraulic - based , that allows differentials to spin at different speeds but still be capable of transferring torque from a wheel with poor traction to one with better . Typical AWD systems work well on all surfaces , but are not intended for more extreme off - road use . When used to describe AWD systems in light passenger vehicles , it refers to a system that applies torque to all four wheels ( permanently or on demand ) and / or is targeted at improving on - road traction and performance ( particularly in inclement conditions ) , rather than for off - road applications . Some all wheel drive electric vehicles solve this challenge using one motor for each axle , thereby eliminating a mechanical differential between the front and rear axles . An example of this is the dual motor variant of the Tesla Model S , which on a millisecond scale can control the torque distribution electronically between its two motors . IWD ( edit ) Individual - wheel drive ( IWD ) was coined to identify those electric vehicles whereby each wheel is driven by its own individual electric motor . This system essentially has inherent characteristics that would be generally attributed to four - wheel drive systems like the distribution of the available torque to the wheels . However , because of the inherent characteristics of electric motors , torque can be negative , as seen in the Rimac Concept One and SLS AMG Electric . This can have drastic effects , as in better handling in tight corners . The term IWD can refer to a vehicle with any number of wheels . For example , the Mars rovers are 6 - wheel IWD . SAE recommended practices ( edit ) Per the SAE International standard J1952 AWD is the preferred term for all the systems described above . The standard subdivides AWD systems into three categories . Part - Time AWD systems require driver intervention to couple and decouple the secondary axle from the primarily driven axle and these systems do not have a center differential ( or similar device ) . The definition notes that part - time systems may have a low range . Full - Time AWD systems drive both front and rear axles at all times via a center ( inter-axle ) differential . The torque split of that differential may be fix or variable depending on the type of center differential . This system can be used on any surface at any speed . The definition does not address inclusion or exclusion of a low range gear . On - Demand AWD systems drive the secondary axle via an active or passive coupling device or `` by an independently powered drive system '' . The standard notes that in some cases the secondary drive system may also provide the primary vehicle propulsion . On - demand systems function primarily with only one powered axle until torque is required by the second axle . At that point either a passive or active coupling sends torque to the secondary axle . In addition to the above primary classifications the J1952 standard notes secondary classifications resulting in a total of eight system designations : Part - Time Non Synchro System Part - Time Synchro System Full - Time Fixed Torque System Full - Time Variable Torque Passive System Full - Time Variable Torque Active System On - Demand Synchro Variable Torque Passive System On - Demand Synchro Variable Torque Active System On - Demand Independently Powered Variable Torque Active System Design ( edit ) Differentials ( edit ) Main article : Differential The Lamborghini Murciélago is an AWD that powers the front via a viscous coupling unit if the rear slips . The HMMWV is a 4WD / AWD that powers all wheels evenly ( continuously ) via a manually lockable center differential , with Torsen differentials for both front and rear . Two wheels fixed to the same axle turn at the same speed as a vehicle goes around curves . This either forces one to slip , if possible , to balance the apparent distance covered , or creates uncomfortable and mechanically stressful wheel hop . To prevent this the wheels are allowed to turn at different speeds using a mechanical or hydraulic differential . This allows one driveshaft to independently drive two output shafts , axles that go from the differential to the wheel , at different speeds . The differential does this by distributing angular force ( in the form of torque ) evenly , while distributing angular velocity ( turning speed ) such that the average for the two output shafts is equal to that of the differential ring gear . When powered each axle requires a differential to distribute power between the left and right sides . When power is distributed to all four wheels a third or ' center ' differential can be used to distribute power between the front and rear axles . The described system handles extremely well , as it is able to accommodate various forces of movement and distribute power evenly and smoothly , making slippage unlikely . Once it does slip , however , recovery is difficult . If the left front wheel of a 4WD vehicle slips on an icy patch of road , for instance , the slipping wheel will spin faster than the other wheels due to the lower traction at that wheel . Since a differential applies equal torque to each half - shaft , power is reduced at the other wheels , even if they have good traction . This problem can happen in both 2WD and 4WD vehicles , whenever a driven wheel is placed on a surface with little traction or raised off the ground . The simplistic design works acceptably well for 2WD vehicles . It is much less acceptable for 4WD vehicles , because 4WD vehicles have twice as many wheels with which to lose traction , increasing the likelihood that it may happen . 4WD vehicles may also be more likely to drive on surfaces with reduced traction . However , since torque is divided amongst four wheels rather than two , each wheel receives approximately half the torque of a 2WD vehicle , reducing the potential for wheel slip . To prevent slippage from happening some vehicles have controls for independently locking center , front , and rear differentials Limiting slippage ( edit ) Main article : Limited - slip differential ( LSD ) Many differentials have no way of limiting the amount of engine power that gets sent to its attached output shafts . As a result , if a tire loses traction on acceleration , either because of a low - traction situation ( e.g. , - driving on gravel or ice ) or the engine power overcomes available traction , the tire that is not slipping receives little or no power from the engine . In very low traction situations , this can prevent the vehicle from moving at all . To overcome this , there are several designs of differentials that can either limit the amount of slip ( these are called ' limited - slip ' differentials ) or temporarily lock the two output shafts together to ensure that engine power reaches all driven wheels equally . Locking differentials work by temporarily locking together a differential 's output shafts , causing all wheels to turn at the same rate , providing torque in case of slippage . This is generally used for the center differential , which distributes power between the front and the rear axles . While a drivetrain that turns all wheels equally would normally fight the driver and cause handling problems , this is not a concern when wheels are slipping . The two most common factory - installed locking differentials use either a computer - controlled multi-plate clutch or viscous coupling unit to join the shafts , while other differentials more commonly used on off - road vehicles generally use manually operated locking devices . In the multi-plate clutch the vehicle 's computer senses slippage and locks the shafts , causing a small jolt when it activates , which can disturb the driver or cause additional traction loss . In the viscous coupling differentials the shear stress of high shaft speed differences causes a dilatant fluid in the differential to become solid , linking the two shafts . This design suffers from fluid degradation with age and from exponential locking behavior . Some designs use gearing to create a small rotational difference that hastens torque transfer . A third approach to limiting slippage is taken by a Torsen differential . A Torsen differential allows the output shafts to receive different amounts of torque . This design does not provide for traction when one wheel is spinning freely , where there is no torque , but provides excellent handling in less extreme situations . A typical Torsen II differential can deliver up to twice as much torque to the high traction side before traction is exceeded at the lower traction side . A fairly recent innovation in automobiles is electronic traction control . Traction control typically uses a vehicle 's braking system to slow a spinning wheel . This forced slowing emulates the function of a limited - slip differential , and , by using the brakes more aggressively to ensure wheels are being driven at the same speed , can also emulate a locking differential . It should be noted that this technique normally requires wheel sensors to detect when a wheel is slipping , and only activates when wheel slip is detected . Therefore , there is typically no mechanism to actively prevent wheel slip ( i.e. , you ca n't `` lock the differential '' in advance of wheel slip ) , rather the system is designed to expressly permit wheel slip to occur , and then attempt to send torque to the wheels with the best traction . If preventing all - wheel slip is a requirement , this is a limiting design . Operating modes ( edit ) The architecture of an AWD / 4WD system can be described by describing its possible operating modes . A single vehicle may have the ability to operate in multiple modes depending on driver selection . Mohan describes the modes as follows : Two Wheel Drive ( 2WD ) Mode - In this mode only one axle ( typically the rear axle ) is driven . The drive to the other axle is disconnected . The operating torque split ratio is 0 : 100 . Four Wheel Drive ( 4WD ) Mode - Here , depending on the nature of torque transfer to the axles , we can define three sub-modes ( below ) . Part - time Mode - The front and rear axle drives are rigidly coupled in the transfer case . Since the driveline does not permit any speed differentiation between the axles and would cause driveline wind - up , this mode is recommended only for ' part - time ' use in off - road or loose surface conditions where driveline wind - up is unlikely . Up to full torque could go to either axle depending on the road condition and the weight over the axles . Full - time Mode - Both axles are driven at all times but an inter-axle differential permits the axles to turn at different speeds as needed . This allows the vehicle to be driven ' full - time ' in this mode , regardless of the road surface , without fear of driveline wind - up . With standard bevel gear differentials the torque split is 50 : 50 . Planetary differentials can provide asymmetric torque splits as needed . A system that operates permanently in the full - time mode is sometimes called ' All - the - Time 4WD ' , ' All - Wheel - Drive ' or ' AWD ' . If the inter-axle differential is locked out then the mode reverts to a ' part - time mode ' . On - Demand Mode - In this mode the transfer case operates primarily in the 2WD mode . Torque is transferred to the secondary axle as needed by modulating the transfer clutch from ' open ' to a rigidly coupled state while avoiding any driveline wind - up . The torque modulation may be achieved by active electronic / hydraulic control systems , or by passive devices , based on wheel slip or wheel torque , as described in the section on traction control systems . In addition to these basic modes there could be implementations that combine these modes . The system could have a clutch across the center differential , for example , capable of modulating the front axle torque from a Full - time mode with the 30 : 70 torque split of the center differential to the 0 : 100 torque split of the 2WD mode History ( edit ) The Lohner - Porsche Mixte Hybrid was both the world 's first hybrid vehicle , and the first four - wheel drive without a steam - engine The 1903 Spyker 60 - HP was the world 's first 4WD , that was directly powered by an internal combustion engine , and the first 4WD race - car . Center transfer case sending power from the transmission to the rear axle ( right ) and front axle ( left ) Selection lever : 2H for two - wheel drive , 4H for high - range 4WD , 4L for low - range 4WD , and N for Neutral Late 1800s ( edit ) In 1893 , before the establishment of a modern automotive industry in Britain , English engineer Bramah Joseph Diplock patented a four - wheel - drive system for a steam - powered traction engine , including four - wheel steering and three differentials , which was subsequently built . The development also incorporated Bramah 's Pedrail wheel system in what was one of the first four - wheel - drive automobiles to display an intentional ability to travel on challenging road surfaces . It stemmed from Bramagh 's previous idea of developing an engine that would reduce the amount of damage to public roads . Ferdinand Porsche designed and built a four - wheel - driven electric vehicle for the k.u.k. Hofwagenfabrik Ludwig Lohner & Co. in Vienna in 1899 , presented to the public during the 1900 World Exhibition in Paris . The vehicle was a series hybrid car , that used an electric hub motor at each wheel , powered by batteries , which were in turn charged by a gasoline - engine generator . It was clumsily heavy and due to its unusual status the so - called Lohner - Porsche is not frequently given its credit as the first four - wheel - driven automobile . 1900s - 1920s ( edit ) The world 's first four - wheel - drive car directly powered by an internal combustion engine , commissioned for the Paris to Madrid race of 1903 , the Spyker 60 H.P. , was presented that year by Dutch brothers Jacobus and Hendrik - Jan Spijker of Amsterdam . The two - seat sports car featured permanent four - wheel drive and was also the first car ever equipped with a six - cylinder engine , as well as four - wheel braking . Later used as a hill - climb racer , it is now an exhibit in the Louwman Museum ( the former Nationaal Automobiel Museum ) in the Hague , the Netherlands . Designs for four - wheel drive in America came from the Twyford Company of Brookville , Pennsylvania in 1905 , six were made there around 1906 ; one still exists and is displayed annually . The second American four - wheel - drive vehicle was built in 1908 by ( what became ) the Four Wheel Drive Auto Company ( FWD ) of Wisconsin ( not to be confused with the term `` FWD '' as an acronym for front - wheel drive ) . FWD would later produce around 15,000 of its four - wheel - drive Model B trucks for the British and American armies during World War I . Approximately 11,500 of the Jeffery / Nash Quad trucks were built for similar use between 1913 and 1919 . The Quad not only came with four - wheel drive and four - wheel brakes , but also featured four - wheel steering . The Quad was one of the first successful four - wheel drive vehicles ever to be made , and its production continued for 15 years with a total of 41,674 units made by 1928 . The Reynolds - Alberta Museum has a four - wheel - drive vehicle , named `` Michigan '' , from 1905 in an unrestored storage . The American Marmon - Herrington Company was founded in 1931 to serve a growing market for moderately priced four - wheel - drive vehicles . Marmon - Herrington specialized in converting Ford trucks to four - wheel drive and got off to a successful start by procuring contracts for military aircraft refueling trucks , 4 × 4 chassis for towing light weaponry , commercial aircraft refueling trucks , and an order from the Iraqi Pipeline Company for what were the largest trucks ever built at the time . Daimler - Benz also has a history in four - wheel drive . In 1907 the Daimler Motoren Gesellschaft had built a four - wheel - driven vehicle called Dernburg - Wagen , also equipped with four - wheel steering , that was used by German colonial civil servant , Bernhard Dernburg , in Namibia . Mercedes and BMW , in 1926 , introduced a rather sophisticated four - wheel drive , the G1 , the G4 and G4 following . The 1937 Mercedes - Benz G5 and BMW 325 4 × 4 featured full - time four - wheel drive , four - wheel steering , three locking differentials , and fully independent suspension . They were produced because of a government demand for a four - wheel - drive passenger vehicle . The modern G - series / Wolf such as the G500 and G55 AMG still feature some of the attributes , with the exception of fully independent suspension since it can compromise ground clearance . The Unimog is another Mercedes truck . 1930s -- 1944 ( edit ) Starting in 1936 , Japanese company Tokyu Kurogane Kogyo built approximately 4,700 four - wheel - drive roadsters , called the Kurogane Type 95 reconnaissance car , used by the Imperial Japanese Army from 1937 until 1944 , during the Second Sino - Japanese War . Three different bodystyles were manufactured ; a 2 - door roadster , a 2 - door pickup truck and a 4 - door phaeton , all equipped with a transfer case that engaged the front wheels , powered by a 1.3 litre , 2 - cylinder , air - cooled OHV V - twin engine . The first Russian produced four - wheel - drive vehicle , also in part for civilian use , was the GAZ - 61 , developed in the Soviet Union in 1938 . `` Civilian use '' may be a bit of a misnomer , as most if not all were used by the Soviet government and military ( as command cars ) , but the GAZ - 61 - 73 version is the first four - wheel drive vehicle with a normal closed sedan body . Elements of the chassis were used in subsequent military vehicles such as the 1940 GAZ - 64 and the 1943 GAZ - 67 , as well as the post-war GAZ - 69 , and the properly civilian GAZ - M - 72 , based on the rear - wheel drive GAZ - 20 `` Victory '' and built from 1955 - 1958 . Soviet civilian life did not allow the proliferation of civilian products such as the Jeep in North America , but through the 1960s the technology of Soviet 4 × 4 vehicles stayed on par with British , German , and American models , even exceeding it in some aspects , and for military purposes just as actively developed , produced and used . It was not until `` go - anywhere '' vehicles were needed for the military on a large scale , that four - wheel drive found its place . The World War II Jeep , originally developed by American Bantam but mass - produced by Willys and Ford , became the best - known four - wheel - drive vehicle in the world during the war . 1945 - 1960s ( edit ) Willys introduced the model CJ - 2A in 1945 , the first full - production four - wheel - drive vehicle for sale in the general marketplace . Thanks to the ubiquitous World War II jeep 's success , its rugged utilitarianism set the pattern for many four - wheel drive vehicles to come . The similarly boxy , also inline - 4 powered Land Rover appeared at the Amsterdam Motor Show in 1948 . Originally conceived as a stop - gap product for the struggling Rover car company , despite chronic under - investment it succeeded far better than the passenger cars . Inspired by a Willys MB that was frequently run off - road on the farm belonging to chief engineer Maurice Wilks , Land Rover developed the more refined yet still off - road capable luxury 4WD Range Rover in the 1970s . With the acquisition of the `` Jeep '' name in 1950 Willys had cornered the brand . Its successor , Kaiser Jeep , introduced a revolutionary 4WD wagon called the Wagoneer in 1963 . Not only was it technically innovative , with independent front suspension and the first automatic transmission coupled to 4WD , it was equipped and finished as a regular passenger automobile . In effect , it was the ancestor of the modern SUV . The luxury Rambler or Buick V8s powered Super Wagoneer produced from 1966 to 1969 raised the bar even higher . Jensen applied the Formula Ferguson ( FF ) full - time all - wheel - drive system to 318 units of their Jensen FF built from 1966 to 1971 , marking the first time 4WD was used in a production GT sports car . While most 4WD systems split torque evenly , the Jensen split torque roughly 40 % front , 60 % rear by gearing the front and rear at different ratios . 1970s - 1990s ( edit ) American Motors ( AMC ) acquired Kaiser 's Jeep Division in 1970 and quickly upgraded and expanded the entire line of off - road 4WD vehicles . With its added roadworthiness , the top range full - size Grand Wagoneer continued to compete with traditional luxury cars . Partially hand - built , it was relatively unchanged during its production through 1991 , even after Chrysler 's buyout of AMC . Subaru introduced the category - expanding Leone in 1972 , an inexpensive compact station wagon with a light - duty , part - time four - wheel drive system that could not be engaged on dry pavement . In September AMC introduced Quadra Trac full - time AWD for the 1973 model year Jeep Cherokee and Wagoneer . Thanks to full - time AWD , which relieved the driver of getting out to lock hubs and having to manually select between 2WD and 4WD modes , it dominated all other makes in FIA rally competition . Gene Henderson and Ken Pogue won the Press - on - Regardless Rally FIA championship with a Quadra Trac equipped Jeep in 1972 . 1969 Jensen FF , world 's first 4WD in a production GT sports car 1987 AWD AMC Eagle wagon , the most popular model in the line 1981 AMC Eagle AWD convertible American Motors introduced the innovative Eagle for the 1980 model year . These were the first American mass production cars to use the complete front - engine , four - wheel drive system . The AMC Eagle was offered as a sedan , coupe , and station wagon with permanent automatic all - wheel - drive passenger models . The new Eagles combined Jeep technology with an existing and proven AMC passenger automobile platform . They ushered a whole new product category of `` sport - utility '' or crossover SUV . AMC 's Eagles came with the comfort and high - level appointments expected of regular passenger models and used the off - road technology for an extra margin of safety and traction . The Eagle 's thick viscous fluid center differential provided quiet and smooth transfer of power that was directed proportionally to the axle with the greatest traction . This was a true full - time system operating only in four - wheel drive without undue wear on suspension or driveline components . There was no low range in the transfer case . This became the forerunner of the designs that followed from other manufacturers . The automobile press at the time tested the traction of the Eagles and described it as far superior to the Subaru 's and that it could beat many so - called off - road vehicles . Four Wheeler magazine concluded that the AMC Eagle was `` The beginning of a new generation of cars . '' The Eagles were popular ( particularly in the snowbelt ) , had towing capacity , and came in several equipment levels including sport and luxury trims . Two additional models were added in 1981 , the sub-compact SX / 4 and Kammback . A manual transmission and a front axle - disconnect feature were also made available for greater fuel economy . During 1981 and 1982 a unique convertible was added to the line . The Eagle 's monocoque body was reinforced for the conversion and had a steel targa bar with a removable fiberglass roof section . The Eagle station wagon remained in production for one model year after Chrysler acquired AMC in 1987 . Audi also introduced a permanently all - wheel - driven road - going car , the Audi Quattro , in 1980 . Audi 's chassis engineer , Jörg Bensinger , had noticed in winter tests in Scandinavia that a vehicle used by the German Army , the Volkswagen Iltis , could beat any high - performance Audi . He proposed developing a four - wheel - drive car , that would also be used for rallying to improve Audi 's conservative image . The Audi Quattro system became a feature on production cars . In 1987 , Toyota also developed a car built for competition in rally campaigns . A limited number of road - going FIA Homologation Special Vehicle Celica GT - Four ( known as Toyota Celica All - Trac Turbo in North America ) were produced . The All - Trac system was later available on serial production Toyota Camry , Toyota Corolla , and Toyota Previa models . Some of the earliest mid-engined four - wheel - drive cars were the various road - legal rally cars made for Group B homologation , such as the Ford RS200 made from 1984 to 1986 . In 1989 , niche maker Panther Westwinds created a mid-engined four - wheel - drive , the Panther Solo 2 . 2000 - present ( edit ) In the United States , as of late 2013 , AWD vehicles comprised 32 per cent of new light vehicle sales , up five per cent since 2008 . This is in large part due to the popularity of the crossover . Most crossovers offer the popular technology , in spite of it increasing vehicle price and fuel consumption . Car manufacturers have inundated consumers with marketing proclaiming AWD as a safety feature , although the advantage of AWD over FWD is in accelerating , not braking or steering . Tests have shown that though AWD gives improved acceleration in wintery conditions , it does not help with braking . In 2008 , Nissan introduced the GT - R featuring a rear - mounted transaxle . The AWD system requires two drive shafts , one main shaft from the engine to the transaxle and differential and a second drive shaft from the transaxle to the front wheels . Uses ( edit ) Road racing ( edit ) Spyker is credited with building and racing the first ever four - wheel drive racing car , the Spyker 60 HP in 1903 . Bugatti created a total of three four - wheel - drive racers , the Type 53 , in 1932 , but the cars were notorious for having poor handling . Miller produced the first 4WD car to qualify for the Indianapolis 500 , the 1938 Miller Gulf Special . Ferguson Research Ltd. built the front - engine P99 Formula One car that actually won a non-World Championship race with Stirling Moss in 1961 . In 1968 , Team Lotus raced cars in the Indy 500 and three years later in Formula 1 with the Lotus 56 , that had both turbine engines and 4WD , as well as the 1969 4WD - Lotus 63 that had the standard 3 - litre V8 Ford Cosworth engine . Matra also raced a similar MS84 , and McLaren entered their M9A in the British Grand Prix , while engine manufacturers Ford - Cosworth produced their own version which was tested but never raced . All these F1 cars were considered inferior to their RWD counterparts , as the advent of aerodynamic downforce meant that adequate traction could be obtained in a lighter and more mechanically efficient manner , and the idea was discontinued , even though Lotus tried repeatedly . Nissan and Audi had success with all - wheel drive in road racing with the former 's advent of the Nissan Skyline GT - R in 1989 . So successful was the car that it dominated the Japanese circuit for the first years of production , going on to bigger and more impressive wins in Australia before weight penalties eventually levied a de facto ban on the car . Most controversially was the win pulled off at the 1990 Macau Grand Prix where the car led from start to finish . Audi 's dominance in the Trans - Am Series in 1988 was equally controversial as it led to a weight penalty mid season and to a rule revision banning all - AWD cars , its dominance in Super Touring eventually led to a FIA ban on AWD system in 1998 . New 2011 24 Hours of Le Mans regulations may revive AWD / 4WD in road racing , though such systems are only allowed in new hybrid - powered Le Mans Prototypes . One example is the Audi R18 e-tron quattro ( winner of 2012 race , the first ever hybrid / 4WD to win Le Mans ) , utilizing an electric motor in the front axle while combining the engine motor in the rear . In heavy trucks ( edit ) Medium - duty trucks and Heavy - duty trucks have recently adopted 4 × 4 drive - trains . 4 × 4 medium - duty trucks became common after Ford Motor Company began selling Ford Super Duty trucks . The Super Duty trucks shared many parts between the light duty and medium duty , reducing production costs . The Dana 60 front axle is used on both medium - and light - duty Super Duty trucks . Furthermore , the Big Three share / shared parts between the companies reducing costs . The Dana S 110 is currently being used for the rear drive , under Ford and Ram 's medium - duty trucks . The Dana 110 was also used on the General Motors 4 × 4s as well . Ram Trucks began selling medium - duty trucks , 4 × 4 and 4 × 2 , in 2008 . General Motors sold a 4 × 4 for model years 2005 - 2009 . GM 4 × 4 medium - duty trucks Heavy duty International Workstar Ford medium - duty 4 × 4 drive train In construction equipment ( edit ) A Case backhoe loader with 4WD Volvo introduced the Model 646 four - wheel drive backhoe loader in 1977 . Case Corporation followed suit in the U.S. in 1987 . Terminology ( edit ) In engineering terms , `` four - wheel drive '' designates a vehicle with power delivered to four wheel ends spread over at least two axles . The term 4 × 4 ( pronounced four by four ) was in use to describe North American military four - wheel - drive vehicles as early as the 1940s , with the first number indicating the number of wheel ends on a vehicle and the second indicating the number of driven wheels . Trucks with dual tires on the rear axle and two driven axles are designated as 4 × 4s despite having six wheels because the paired rear wheels behave as a single wheel for traction and classification purposes . True 6 × 6 vehicles which have three powered axles , and are classified as 6 × 6s regardless of how many wheels they have . Examples of these with two rear , one front axle are the 6 - wheeled Pinzgauer , which is popular with defense forces around the globe , and 10 - wheeled GMC CCKW made famous by the U.S. Army in World War II . 4 - wheeler ( or four - wheeler ) is a related term applying to all - terrain vehicles , and not to be confused with four - wheel drive . The `` four '' in the instance referring to the vehicle having four wheels , not necessarily all driven . Unusual systems ( edit ) Prompted by a perceived need for a simple , inexpensive all - terrain vehicle for oil exploration in North Africa , the French motor manufacturer Citroën developed the 2CV Sahara in 1958 . Unlike other 4 × 4 vehicles which use a conventional transfer case to drive the front and rear axle , the Sahara had two engines , each independently driving a separate axle , with the rear engine facing backwards . The two throttles , clutches and gear change mechanisms could be linked , so the two 12 hp ( 9 kW ) 425 cc ( 26 cu in ) engines could run together , or they could be split and the car driven solely by either engine . Combined with twin fuel tanks and twin batteries ( which could be set up to run either or both engines ) , the redundancy of two separate drive trains meant that they could make it back to civilization even after major mechanical failures . Only around 700 of these cars were built , and only 27 are known to exist today . BMC experimented with a twin - engine Mini Moke ( dubbed the `` Twini Moke '' ) in the mid-1960s , but never put it into production . This made advantage of the Mini 's ' power pack ' layout , with a transverse engine and the gearbox in the engine sump . Simply by fitting a second engine / gearbox unit across the rear , a rudimentary 4 × 4 system could be produced . Early prototypes had separate gear levers and clutch systems for each engine . Later versions sent for evaluation by the British Army had more user - friendly linked systems . In 1965 , A.J.M. Chadwick patented a 4WD system , GB 1113068 , that used hemispherical wheels for an all - terrain vehicle . Twenty years later , B.T.E. Warne , patented , GB 2172558 , an improvement on Chadwick 's design that did not use differential gear assemblies . By using near - spherical wheels with provision to tilt and turn each wheel co-ordinatively , the driven wheels maintain constant traction . Furthermore , all driven wheels steer and , as pairing of wheels is not necessary , vehicles with an odd number of wheels are possible without affecting the system 's integrity . Progressive deceleration is made possible by dynamically changing the front - to - rear effective wheel diameter ratios . Suzuki Motors introduced the Suzuki Escudo Pikes Peak Edition in 1996 . Earlier Suzuki versions were twin engined , from 1996 on the engine is a twin - turbocharged 2.0 L V6 , mated to a sequential 6 - speed manual transmission . Nissan Motors has developed a system called E-4WD . It is designed for cars that are normally front - wheel drive , however the rear wheels are powered by electric motors . This system was introduced in some variants of the Nissan Cube and Tiida . ( This is similar to the system used on the Ford Escape Hybrid AWD . ) Chrysler 's Jeep Division debuted the twin engine , 670 hp ( 500 kW ) Jeep Hurricane concept at the 2005 North American International Auto Show in Detroit . This vehicle has a unique `` crab crawl '' capability , which allows it to rotate 360 ° in place . This is accomplished by driving the left wheels as a pair and right wheels as a pair , as opposed to driving the front and rear pairs . A central gearbox allows one side to drive in the opposite direction to the other . It also has dual Hemi V8s . Some hybrid vehicles such as the Lexus RX400h provide power to an AWD system through a pair of electric motors , one to the front wheels and one to the rear . In the case of the AWD model version of the Lexus RX400h ( and its Toyota - branded counterpart , the Harrier hybrid ) , the front wheels can also receive drive power directly from the vehicle 's gasoline engine as well as via the electric motors , whereas the rear wheels derive power only from the second electric motor . Transfer of power is managed automatically by internal electronics based on traction conditions and need , making this an all - wheel - drive system . The 4RM system used in the Ferrari FF in 2011 is unique in that it has a rear transaxle with secondary front transaxle connected directly to the engine . The car operates primarily as a rear wheel drive vehicle . Clutches in the front transaxle engage when the rear wheels slip . Drive to the front wheels is transmitted through two infinitely - variable clutch packs which are allowed to ' slip ' to give the required road wheel speeds . The front transaxle has three gears , two forward , and reverse . The two forward gears of the front transmission match the lower four forward gears of the rear transmission . It is not used in higher gears . The connection between this gearbox and each front wheel is via independent haldex - type clutches , without a differential . Due to the difference in ratios the clutches continually slip and only transmit , at most , 20 % of the engine 's torque . Introduction to off - roaders ( edit ) Year Manufacturer Model / Series Mass Military 1941 GAZ GAZ - 64 x 1941 Willys - Overland Willys MA / MB x 1941 Volkswagen Typ 128 Schwimmwagen x 1945 Willys CJ - 2A x 1946 Dodge Power Wagon x 1947 Jeep Willys truck x 1948 Ford F - series x 1948 Rover Land Rover Series x 1951 Alfa Romeo Alfa Romeo Matta x 1951 Austin Motor Company Austin Champ x 1951 Fiat Fiat Campagnola x 1951 Nissan Nissan Patrol x 1954 Toyota Land Cruiser x 1956 Auto Union DKW Munga x 1963 Kaiser Jeep Wagoneer ( SJ ) x 1968 Suzuki Lj10 x 1970 British Leyland Range Rover x 1972 UAZ UAZ - 469 x 1976 Lada Lada Niva x 1979 Mercedes - Benz G - Class x 1982 Mitsubishi Pajero x Introduction to passenger cars ( edit ) Year Manufacturer Model / Series Mass Small series 1938 GAZ GAZ - 61 x ( 238 ) 1955 GAZ GAZ M - 72 x ( 4677 units ) 1966 Jensen Jensen FF x ( 320 units ) 1972 Subaru Subaru Leone x 1980 American Motors AMC Eagle x 1980 Audi Audi Quattro x 1982 Renault Renault 18 Combi 4 × 4 x Fiat Fiat Panda 4 × 4 x 1984 Alfa Romeo Alfa 33 4 × 4 x 1984 Mitsubishi Mitsubishi Cordia 4WD Turbo x 1984 Volkswagen Volkswagen Passat Syncro x 1984 Peugeot Peugeot 205 T16 x ( 200 units ) 1985 BMW BMW 325iX x 1985 Lancia Lancia Delta HF 4WD x 1985 Mazda Mazda 323 4WD x 1985 Suzuki Suzuki Samurai x 1986 Toyota Toyota Celica GT - Four All - Trac Turbo x 1986 Ford Ford RS200 x ( 220 units ) Mercedes - Benz Mercedes W124 4MATIC x Porsche Porsche 959 x ( 292 units ) 1988 Opel Opel Vectra 4 × 4 x 1988 Pontiac Pontiac 6000 STE AWD x Citroën Citroën BX x Chevrolet Chevrolet Astro Full - time AWD x 1991 Lamborghini Lamborghini Diablo VT x 1991 Bugatti Bugatti EB110 x ( ca . 300 units ) Fiat Tempra 4 × 4 x Volvo 850 AWD x Jaguar Jaguar X-Type x 2008 Saab Saab 9 - 3 Turbo X x Dacia Dacia Duster x 2011 Ferrari Ferrari FF x ( ca . 800 units ) Systems by design type ( edit ) Center differential with mechanical lock ( edit ) Alfa Romeo 164 Q4 ( central viscous coupling , epicyclic unit and Torsen rear differential ) Alfa Romeo 155 Q4 ( central epicyclic unit , Ferguson viscous coupling and Torsen rear differential ) AMC Eagle ( central viscous coupling ) Audi - Quattro Coupé , 80 , 90 , 100 & 200 ( locking center and rear differentials ) - up to 1987 Audi Q7 - double pinion 50 / 50 with lockup clutch pack BMW 3 series and 5 series in the 1980s - planetary center differential with a 37 - 63 ( front - back ) torque split and viscous lock ( also in rear differential but not front differential ) Chevrolet Rounded - Line K Fleetside , K Stepside , K Blazer , and K Suburban - permanent four - wheel drive ( 1973 - 1979 ) two - speed New Process 203 transfer case , center differential with 50 : 50 torque split and lock . An Eaton Automatic Differential Lock was optional for the rear hypoid differential . Ford - Escort ( RS 2000 16v 4 × 4 models and RS Cosworth ) , Sierra Cosworth , Sierra and Granada 4 × 4 models , Dodge Power Wagon - permanent four - wheel drive ( 1974 - 1979 ) two - speed New Process 203 transfer case , center differential with 50 : 50 torque split and lock . Ford Expedition ( 1997 -- present ) and Expedition EL / Max ( 2007 -- present ) - automatic ControlTrac four - wheel drive with two - speed dual range BorgWarner transfer case and intelligent locking center multi-disc differential Ford Explorer ( 1995 -- 2010 ) - automatic ControlTrac four - wheel drive with two - speed dual range BorgWarner transfer case and intelligent locking center multi-disc differential Ford F - Series - permanent four - wheel drive ( 1974 - 1979 ) two - speed New Process 203 transfer case , center differential with 50 : 50 torque split and lock . GMC Rounded - Line K Wideside , K Fenderside , K Jimmy , and K Suburban - permanent four - wheel drive ( 1973 - 1979 ) two - speed New Process 203 transfer case , center planetary differential with 50 : 50 torque split and lock . An Eaton Automatic Differential Lock was optional for the rear hypoid differential . H1 & Humvee NVG 242HD AMG open center differential , locked center differential , Neutral , low range locked . Also Torsen1 differential at the front and rear axle , The H1 moved to Torsen2 when ABS was added . The H1 Alpha had optional locking differentials in place of torsens Hummer H2 , H3 40 / 60 planetary with lock Jeep Grand Cherokee , Commander ( except models equipped with Quadra - Trac I ) Jeep Liberty , Jeep Cherokee ( XJ ) , Dodge Durango ( Select - Trac ) - NV 242 transfer case - rear drive , open center differential , locked center differential , Neutral , low range Full size Jeeps with Borg Warner QuadraTrac : limited slip center differential , 50 / 50 locked center differential . Low range could be used in locked or unlocked mode , allowing for use of low range on pavement . Land Rover Defender ( and Series III V8 models ) Land Rover Discovery / LR3 Land Rover Freelander Lada Niva ( VAZ - 2121 ) - full - time 4WD using open center differential . Transfer case with high / low range and manual central diff lock . Low range selectable in locked or unlocked mode , allowing use on pavement . Lexus RX300 - viscous coupling across the otherwise open center differential . Lincoln Navigator ( 1998 -- 2006 ) - automatic ControlTrac four - wheel drive with two - speed dual range BorgWarner transfer case and intelligent locking center multi-disc differential Navigator and Navigator L ( 2007 -- present ) use one - speed single range transfer case , no reduction gearing Mercedes - Benz Unimog ( locking center and rear with up to 10 low range gears ) . Mercedes - Benz G - Class ( locking center and lockers on both front - and rear axle ) Mercedes - Benz GL - Class - 4Matic all - wheel - drive system Mitsubishi Pajero ( also known as Montero or Shogun ) Porsche Cayenne - 38 / 62 planetary with lockup clutch pack Range Rover Classic 1970 -- 1995 all full - time 4WD either plate LSD , manual lock or Ferguson viscous centre differential . Range Rover 2nd Gen. 1994 -- 2002 full - time 4WD Ferguson viscous centre differential Suzuki Grand Vitara / Escudo ( 2005 and later models , excepting the XL - 7 ) - full - time 4WD using limited - slip center differential , off - road 4WD with selectable center differential lock and low range transfer case 4 mode ( 4h , 4h lock , 4l n ) , traction control and electronic stability control Subaru - manual transmissions come with 50 / 50 viscous - type center differential ; performance models include a planetary differential with computer regulated lockup ; automatic transmission models have an electronically controlled variable transfer clutch . Toyota Land Cruiser Toyota Sequoia ( Multi-mode ) Volkswagen Touareg - double pinion 50 / 50 with lockup clutch pack Torsen center differential ( edit ) Alfa Romeo Q4s - with ( Torsen T - 3 ) : 156 Crosswagon and Sportwagon 159 Brera , Spider Audis with quattro - various iterations of Torsen , the T - 3 starting from the 2007 B7 RS4 80 , 90 & Coupé ( Typ 89 ) 100 & 200 A4 , S4 , RS4 A5 & S5 A6 , S6 , RS6 A8 , S8 Q5 , Q7 Bentley Continental GT , Bentley Continental Flying Spur ( 2005 ) initially Torsen T - 2 , current have T - 3 Chevrolet Trailblazer SS Torsen T - 3 Lexus GX 470 , Toyota Land Cruiser Prado 120 Torsen T - 3 Range Rover 3rd Gen. 2002 -- 2009 Toyota 4Runner ( only Limited V8 model & 2010 Limited V6 model ) Torsen T - 3 with lock Toyota FJ Cruiser ( only manual models ) Torsen T - 3 with lock Toyota Hilux Surf Torsen T - 3 with lock Toyota Land Cruiser 200 / 2008 / V8 Torsen T - 3 with lock Toyota Sequoia ( North America ) ( only 2005 - 07 Models ) Volkswagen Passenger Cars with 4motion : Volkswagen Passat Torsen T - 2 ( B5. 5 model , not latest B6 model with transverse engine ) Volkswagen Phaeton Torsen T - 2 Non-locking center differential ( edit ) BMW 3 - series and X5 between 2001 and xDrive - planetary center differential with permanent 38 - 62 ( front - back ) torque split # Cadillac Escalade , STS AWD , SRX AWD ( The first two generations had a viscous clutch on the center differential ) # Chrysler 300C AWD # Dodge Ramcharger 1974 -- 1981 - NP203 FullTime 4WD Transfer Case Dodge Magnum , Charger AWD # GMC Yukon Denali , XL Denali , Sierra Denali # Mercedes 4MATIC cars , R class , and ML class ( note some MLs had low range ) # Plymouth Trailduster 1974 -- 1981 - NP203 FullTime 4WD Transfer Case Toyota Highlander # Toyota Sienna AWD ( - 2010 only ) # The above systems ending with `` # '' function by selectively using the traction control system ( via ABS ) to brake a slipping wheel . Multiple - clutch systems ( edit ) Acura RL , RDX ( SH - AWD ) Right and left axle shaft Acura MDX SH - AWD & VTM4 Ford Explorer - Ford 's full - time shift - on - the - fly Intelligent 4WD System ( I - 4WD ) on the 2011 Explorer with Terrain Management System and RSC ( Roll Stability Control ) , Curve Control functionality , HDC ( Hill Descent Control ) and HAA ( Hill Ascent Assist ) . Honda Ridgeline Honda Pilot Infiniti FX ( ATTESA E-TS ) Mercedes - Benz 1st generation 4MATIC ( normally rear - drive , automatic clutch in transfer case engages 4WD on demand ) Mitsubishi GTO MR / 3000GT VR - 4 Mitsubishi Lancer Evolution Series S - AWC 2010 Mitsubishi Outlander GT S - AWC Mitsubishi Outlander ( 2003 -- 2006 ) independent front and rear axle coupling , and Active Center Differential . Nissan GT - R ( ATTESA E-TS ) Nissan Skyline GT - R ( ATTESA E-TS and ATTESA E-TS - PRO ) front axle coupling , rear differential locking Nissan Skyline GTS4 ( ATTESA E-TS ) Nissan A31 Cefiro SE4 ( ATTESA E-TS ) Porsche 959 PSK front axle coupling , rear differential locking Saab 9 - 3 , Saab 9 - 5 , Saab 9 - 4X ( Saab XWD ) . Multi-plate clutch coupling ( edit ) Audi A3 quattro , Audi S3 , Audi TT quattro , Audi R8 ( with Haldex Traction ) BMW xDrive : latest 3 Series , latest 5 series , X3 , latest X5 series Chevrolet Equinox ( GMPCA ) Chrysler Pacifica ( BorgWarner ITM3e ) ( on 2007 model ) Dodge Nitro ( Quadra - Trac 1 ) Dodge Caliber Ford : Escape , Freestyle , Edge , Fusion , Five Hundred ( Freestyle , FiveHundred Haldex Traction based ) ( Escape Control Trac II , based ) Honda CR - V , HR - V , Element Hyundai Santa Fe , Hyundai Tucson Borg - Warner ITM 3e magnetic multi-plate clutch coupling Hyundai Veracruz IMJ magnetic multi-plate clutch coupling Infiniti : G35x , M35x Jeep Compass ( Freedom Drive ) Jeep Grand Cherokee and SRT8 NVG 249 , 247 Land Rover Freelander 2 / LR2 ( also Haldex Traction ) Lamborghini : AWD variants VT series ( viscous traction ) Lincoln : MKS , MKZ Mazdaspeed6 ( a power takeoff unit linked to clutch pack with torque sensitive rear differential ) Mazda : Tribute , CX - 7 , CX - 9 ( tribute Control Trac II , based ) Mercury : Milan , Montego , Mariner ( Montego Haldex Traction - based ) Mitsubishi Outlander ( current generation ) Nissan Murano automatic with manual lockup switch Porsche 911 AWD variants ( a version of BorgWarner ITM3e ) -- excluding the 964 - series Porsche 911 Carrera 4 31 / 69 planetary center differential Pontiac Torrent ( GMPCA ) Subaru low powered automatic transmission models Subaru Legacy , Outback , Impreza , Forester , Tribeca automatic transmission models : mechanical front drive , clutch coupled rear axle . Suzuki : SX4 , XL7 , Aerio , Swift / Cultus based Subaru Justy . ( viscous clutch ) Toyota RAV4 - from 2005 ( third generation only ) Toyota Sienna AWD ( 2011 and newer only ) Volkswagen Golf 4motion , Volkswagen Jetta 4motion , Volkswagen Tiguan 4motion , Volkswagen Passat ( B6 ) 4motion ( initially viscous coupling , later with Haldex Traction ) Volvo : S40 , S60 , S80 , V50 , V70 , XC70 , XC90 ( Visco system until 2003 ; then all Haldex Traction - based ) Note : the above all function like 2WD when multi-plate clutch coupling is not engaged ( with exception of Subaru models ) , and like 4WD highrange in a part - time 4WD system when the clutch is engaged ( usually by computer although some allow manual control ) . Some in this category have varying degrees of control in the torque distribution between front and rear by allowing some of the clutches in a multi-plate clutch coupling to engage and slip varying amounts . An example of a system like this is the BorgWarner i - Trac ( TM ) system . Note : the Haldex Traction - based car list was created from the list on Haldex Traction corporate web site : Haldex Cars . A version of the BorgWarner ITM3e system is used on 2006 and up Porsche 911TT 's . The Borg - Warner ITM 3e is also used in the 2006 - now Hyundai Santa Fe and the Hyundai Tucson . In the Hyundais , the ITM 3e acts like a full - time AWD with 95 : 5 normal torque split . In extreme conditions , the system can be locked in a 50 : 50 split via the 4WD LOCK button . Part - time ( edit ) These are vehicles that have no center differential . Since there is no center differential to allow for speed differences between the front and rear wheels when turning , a small amount of tire slippage must occur during turns . When used on slick surfaces , this is not a problem , but when turning on dry pavement , the tires grip , then are forced to slip , then grip again , and so on , until the turn is completed . This causes the vehicle to exhibit a ' hopping ' sensation . Using an engaged part - time 4WD system on a hard surface is not recommended , as damage to the drive - line eventually occurs . Chevrolet Rounded - Line K Fleetside , K Stepside , K Blazer , and K Suburban - conventional four - wheel drive ( 1973 - 1987 ) or shift - on - the - move four - wheel drive ( 1981 - 1987 ) two - speed New Process 205 or 208 transfer case . 0 : 100 torque split in Two High . 50 : 50 torque split lock in Four High and Four Low . An Eaton Automatic Differential Lock was optional for the rear hypoid differential . Note Rounded - Line `` K '' Pickups and Utilities were temporarily renamed to `` V '' for 1987 Chevrolet Tahoe , Trailblazer ( LT1 and LT3 models only ) , Tracker , Suburban , Silverado , Avalanche , Colorado , S - 10 series , K5 Blazer Dodge Power Wagon ( a Ram version with front and rear differential locks ) Dodge Ram , Dakota Dodge Nitro ( Quadra - Trac 2 ) Ford F series Ford Explorer ( 1991 - 1994 ) & Sport Trac Ford Ranger Geo Tracker GMC Rounded - Line K Wideside , K Fenderside , K Jimmy , and K Suburban - conventional four - wheel drive ( 1973 - 1987 ) or shift - on - the - move four - wheel drive ( 1981 - 1987 ) two - speed New Process 205 or 208 transfer case . 0 : 100 torque split in Two High . 50 : 50 torque split lock in Four High and Four Low . An Eaton Automatic Differential Lock was optional for the rear hypoid differential . Note Rounded - Line `` K '' Pickups and Utilities were temporarily renamed to `` V '' for 1987 GMC Envoy , Yukon , Sierra , Jimmy , Sonoma Infiniti QX56 ( All - mode 4WD ) Auto - engages 4WD with slip Isuzu i - series , Isuzu Wizard Jeep Cherokee ( Quadra - Trac 2 ) Jeep Cherokee ( XJ ) , Jeep Comanche , Jeep Grand Cherokee ( ZJ ) , Jeep Liberty ( Command - Trac ) Jeep Wrangler ( Rubicon model has locking front and rear differentials ) Kia Sorento ( some 2002 - 2009 models with 2WD / 4HI / 4LO - mostly LX ) Land Rover Series I , II & III ( except V8 models ) Lincoln Mark LT Mazda B - series Mercedes - Benz G - Class Mitsubishi Raider Nissan Patrol Nissan Terrano II Nissan Armada , Pathfinder ( All - mode 4WD ) Auto - engages 4WD with slip Nissan Titan , Xterra , Frontier ( rear locker an option ) Subaru Loyale , GL / DL , BRAT Front / 4wd / 4wd lo , Justy 4WD Suzuki Sidekick ( pre-2005 models , and XL - 7 ) , Jimny , Vitara Toyota Hilux Toyota Tacoma ( rear locking differential optional ) Toyota Tundra TRD Toyota FJ Cruiser ( automatic transmission models ) ( also locking rear differential ) Toyota 4Runner ( only SR5 and pre 2010 Limited V6 models , 2010 Trail edition V6 models ) ( also locking rear differential on 2010 V6 ) See also ( edit ) Category : All - wheel - drive vehicles Four - wheel drive in Formula One Limited slip differential Off - road vehicle Sport utility vehicle Dune bashing Rock crawling 4WS Transfer case References ( edit ) ^ Jump up to : Mohan , Sankar ( 12 June 2000 ) . `` All - Wheel Drive / Four - Wheel Drive Systems and Strategies '' ( PDF ) . Seoul 2000 FISITA World Automotive Congres . Jump up ^ Andreev , Alexandr F. ; Kabanau , Viachaslau ; Vantsevich , Vladimir ( 2010 ) . Driveline Systems of Ground Vehicles : Theory and Design . CRC Press . p. 34 . Jump up ^ Dykes , Alex . `` Alphabet Soup : 4 × 4 vs 4WD vs AWD Where 's the Differential ? '' . thetruthaboutcars.com . Retrieved 18 December 2015 . ^ Jump up to : `` Surface Vehicle Recommended Practice - J1952 - All - Wheel Drive System Classification '' . SAE International . October 2013 . Jump up ^ Walczak , Jim . `` 4WD vs 2WD : The Differences Between 4 × 4 And 4 × 2 '' . about com . Retrieved 7 August 2010 . ^ Jump up to : Collard , Chris . `` 2WD vs AWD vs 4WD '' . ConsumerReports.com . Consumer Reports Magazine . Retrieved 15 December 2015 . Jump up ^ Williams , Mark . `` 4WD vs AWD : What 's the Diff '' . MotorTrend.com . Motor Trend Magazine . Retrieved 15 December 2015 . Jump up ^ Allisons.org Automotive History ( 1 ) `` 1929 : AEC started to build AWD trucks in conjunction with FWD ( UK ) '' Jump up ^ MEYER , DONALD E. THE FIRST CENTURY OF GMC TRUCK HISTORY ( 2 ) `` By December ( 1939 ) , GMC had orders for more than 4,400 all - wheel - drive military trucks . '' ^ Jump up to : Sheppard , Tom ( 1 September 2005 ) . `` Jeep Grand Cherokee 4WD System Breakthrough - 4x ... For ? '' . Four Wheeler Network . Retrieved 27 May 2014 . Jump up ^ Davies , Alex ( 10 October 2014 ) . `` The Model D Is Tesla 's Most Powerful Car Ever , Plus Autopilot '' . Wired.com . Retrieved 11 October 2014 . Musk said the added efficiency is thanks to the electronic system that will shift power between the front and rear motors from one millisecond to the next , so each is always operating at its most efficient point Jump up ^ http://www.carscoops.com/2013/04/harris-helps-us-understand-negative.html Jump up ^ The History of 4 Wheel Drive -- Haley Dodge Jump up ^ `` History of Hybrid Vehicles '' . HybridCars.com. 2006 - 03 - 27 . Archived from the original on 2009 - 09 - 04 . Retrieved 2011 - 10 - 25 . See year 1898 . Jump up ^ `` Lohner - Porsche Mixte Voiturette '' . Ultimate Car Page . 2007 - 11 - 19 . Retrieved 2011 - 10 - 25 . Jump up ^ History of 4WD Sport -- OF4WD ^ Jump up to : `` 1903 Spyker 60 HP '' . Retrieved 6 May 2011 . Jump up ^ Spyker 60 - hp Four - wheel Drive Racing Car -- Louwman Museum Jump up ^ `` The Novak Guide to the Dana Spicer Model 18 Transfer Case '' . Novak Conversions . Retrieved 16 September 2012 . ^ Jump up to : Eckermann , Erik ( 2001 ) . World history of the automobile . Society of Automotive Engineers . p. 76 . ISBN 978 - 0 - 7680 - 0800 - 5 . Retrieved 21 January 2013 . Jump up ^ Redgap , Curtis ; Watson , Bill ( 2010 ) . `` The Jefferys Quad and Nash Quad -- 4x4 Ancestor to the Willys Jeep '' . Allpar . Retrieved 6 December 2014 . Jump up ^ Nylund , Jimmy ( February 2009 ) . `` 27 Trailer Towing Tips '' . Four Wheeler . Retrieved 16 September 2012 . Jump up ^ The world 's first mass - produced compact 4WD car , restored -- Japanese Nostalgic Car Jump up ^ Editors of Air Force Journal of Logistics ( 2000 ) . The Logistics of War - a historical perspective . The Air Force Logistics Management Agency . p. 282 . ISBN 9781428993785 . Retrieved 21 January 2013 . It became a famous vehicle around the world and is mentioned in a majority of the histories . CS1 maint : Uses authors parameter ( link ) Jump up ^ Nunney , MJJ ( 2012 ) . Light and Heavy Vehicle Technology ( Fourth ed . ) . Routledge . p. 431 . ISBN 9780750680370 . Retrieved 21 January 2013 . Jump up ^ Foster , Patrick ( January 2008 ) . `` 1963 Jeep Wagoneer - Landmark Vehicle '' . 4 Wheel Drive & Sport Utility Magazine . Retrieved 21 January 2013 . Jump up ^ Auto Editors of Consumer Guide ( 25 July 2007 ) . `` 1966 -- 1971 Jensen FF '' . howstuffworks com . Retrieved 7 August 2010 . Jump up ^ Auto Editors of Consumer Guide ( 29 August 2007 ) . `` 1978 - 1979 Jeep Wagoneer Limited '' . HowStuffWorks.com . Retrieved 5 January 2011 . Jump up ^ Auto Editors of Consumer Guide ( 29 August 2007 ) . `` 1967 -- 1977 Jeep Wagoneer '' . howstuffworks com . Retrieved 7 August 2010 . Jump up ^ Calvin , Jean ( 1974 ) . Rallying to win : a complete guide to North American rallying . W.W. Norton . p. 150 . ISBN 978 - 0 - 87880 - 017 - 9 . Retrieved 5 January 2011 . Jump up ^ Sherman , Don ( February 2001 ) . `` All - Wheel - Drive Revisited : AMC 's 1980 Eagle pioneered the cross-over SUV '' . Automotive Industries . Retrieved 21 January 2013 . Jump up ^ Rettie , John ( August 1987 ) . `` Four - Wheeling Into Your Future '' . Popular Mechanics. 164 ( 8 ) : 58 . Retrieved 5 January 2011 . Jump up ^ Sass , Rob ( 9 March 2008 ) . `` A Breed of 4 - by - 4 Hatched on the Fly '' . The New York Times . Retrieved 21 January 2013 . Jump up ^ Padgett , Martin ( 2004 ) . Hummer . Zenith . p. 26 . ISBN 978 - 0 - 7603 - 1863 - 8 . Retrieved 21 January 2013 . Jump up ^ Foster , Patrick R. `` 1980 - 88 AMC Eagle 4wd '' . Autoweek ( archived at amxfiles ) . Archived from the original on 30 April 2008 . Retrieved 21 January 2013 . Jump up ^ Consumer 's Guide , ed. ( 2005 ) . History of the American Auto . Publications International . p. 532 . ISBN 978 - 0 - 7853 - 9874 - 5 . Jump up ^ `` Celica All Trac and Gt - Four FAQ '' . Alltrac.net . Retrieved 7 August 2010 . ^ Jump up to : Kranz , Rick . `` All - Wheel - Drive Vehicles Grow in Popularity With Car Shoppers '' . Edmunds . Retrieved 3 August 2017 . Jump up ^ Shunk , Chris . `` Thirty - percent of new car sales will be AWD by 2015 '' . autoblog . Retrieved 3 August 2017 . Jump up ^ `` Do You Really Need AWD in the Snow ? '' . Consumer Reports . Retrieved 3 August 2017 . Jump up ^ Peter , Cheney . `` The myth of all - wheel - drive and why it can be a deadly trap '' . The Gobe and Mail . Retrieved 3 August 2017 . Jump up ^ Loh , Edward ( December 2007 ) . `` First Look : 2008 Nissan GT - R '' . Retrieved 5 January 2011 . Jump up ^ Henry , Allan ( 1975 ) . The 4 - Wheel Drives : Racing 's Formula for Failure ? . Macmillan . Jump up ^ `` The 2011 Le Mans 24 Hours Regulations '' . lemans.org. 6 October 2010 . Archived from the original on 13 June 2010 . Retrieved 21 January 2013 . Jump up ^ `` Hybrid Tech Dominates The 24 Hours Of Le Mans Endurance Race ( VIDEO ) '' . Autoblog.com . Jump up ^ `` Volvo BM 646 Backhoe / Loader '' ( PDF ) . Volvo AB . Jump up ^ `` Instruction Book : Driving , Maintenance , Repair . Department of National Defence , Canada '' . bcoy1cpb.pacdat.net. 1940 . Retrieved 21 January 2013 . Jump up ^ Melissen , Wouter ( 1 December 2004 ) . `` Citroën 2CV Sahara '' . Ultimatecarpage.com . Retrieved 21 January 2013 . Jump up ^ `` 2009 Ford Escape Hybrid Review and Specs '' . Edmunds.com . Archived from the original on 1 March 2009 . Retrieved 1 August 2009 . Jump up ^ Horrell , Paul ( 25 January 2011 ) . `` Ferrari FF The inside scoop on Ferrari 's new FF '' . Topgear.com . Retrieved 6 May 2011 . Jump up ^ Abuelsamid , Sam ( 26 July 2010 ) . `` 2011 Ford Explorer puts four - cylinder EcoBoost engine above V6 in pecking order '' . Autoblog.com . Retrieved 26 September 2010 . Jump up ^ `` Ford - Cars , SUVs , Trucks & Crossovers '' ( in Spanish ) . FordVehicles.com . Archived from the original on 30 September 2010 . Retrieved 26 September 2010 . Jump up ^ according to Car and Driver Vol52No8 Feb 07 page 110 External links ( edit ) Car design Car classification By size Microcar City car Kei Subcompact Supermini Family car Compact Mid-size Full - size Custom Hot rod Lead sled Lowrider Street rod T - bucket Luxury Compact executive Executive Personal luxury car Minivan / Multi-purpose vehicle ( MPV ) Compact MPV Mini MPV Sport utility vehicle ( SUV ) Compact SUV Crossover SUV Mini SUV Sports car Grand tourer Hot hatch Muscle Pony Sport compact Supercar Antique Classic Economy Leisure activity vehicle Ute Van Voiturette Body styles 2 + 2 Baquet Barchetta Berlinetta Brougham Cabrio coach Cabriolet / Convertible Coupé Coupé de Ville Coupé utility Drophead coupe ( Convertible ) Fastback Hardtop Hatchback Landaulet Liftback Limousine Multi-stop truck Notchback Panel van Phaeton Pickup truck Quad coupé Retractable hardtop Roadster Runabout Saloon / Sedan Sedan delivery Sedanca de Ville ( Coupé de Ville ) Shooting - brake Spider / Spyder ( Roadster ) Station wagon Targa top Torpedo Touring car Town car ( Coupé de Ville ) T - top Vis - à - vis Specialized vehicles Amphibious Driverless ( autonomous ) Hearse Gyrocar Roadable aircraft Taxicab Tow truck Propulsion Alternative fuel Autogas Biodiesel Diesel Electric ( battery NEV ) Ethanol ( E85 ) Fuel cell Gasoline / petrol ( direct injection ) Homogeneous charge compression ignition Hybrid ( plug - in ) Hydrogen Internal combustion Liquid nitrogen Steam Drive wheels Front - wheel Rear - wheel Two - wheel Four - wheel Six - wheel Eight - wheel Twelve - wheel Engine position Front Mid Rear Layout ( engine / drive ) Front / front Front mid / front Rear / front Front / rear Rear mid / rear Rear / rear Front / four - wheel Mid / four - wheel Rear / four - wheel Engine configuration ( internal combustion ) Boxer Flat Four - stroke H - block Reciprocating Single - cylinder Straight Two - stroke V ( Vee ) W engine Wankel Portal Retrieved from `` https://en.wikipedia.org/w/index.php?title=Four-wheel_drive&oldid=822182990 '' Categories : Off - road vehicles Automobile layouts Automotive technologies Four - wheel drive layout Hidden categories : All articles with dead external links Articles with dead external links from October 2017 Articles with permanently dead external links CS1 maint : Uses authors parameter CS1 Spanish - language sources ( es ) Use dmy dates from August 2011 All articles with unsourced statements Articles with unsourced statements from April 2012 Articles with unsourced statements from July 2015 Talk Contents About Wikipedia Afrikaans Català Čeština Cymraeg Dansk Español Esperanto فارسی Français 한국어 Bahasa Indonesia Italiano Nederlands Norsk Norsk nynorsk Português Simple English Slovenčina Suomi Türkçe Tiếng Việt 中文 Edit links This page was last edited on 24 January 2018 , at 21 : 49 . 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Question: what is the meaning of 4wd in cars

refers a two-axled vehicle with a drivetrain capable of providing torque to all its wheels simultaneously