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Electron transport chain
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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.[1] 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,[2] 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.[citation needed]
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 (ΔΨM). 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 bc1 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 → H2O ↑ 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 (UQH2), 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.[3]
The pathway of electrons is as follows:
NADH is oxidized to NAD+, by reducing Flavin mononucleotide to FMNH2 in one two-electron step. FMNH2 is then oxidized in two one-electron steps, through a semiquinone intermediate. Each electron thus transfers from the FMNH2 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, QH2. During this process, four protons are translocated from the mitochondrial matrix to the intermembrane space. [4] 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.[5]
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 bc1 complex or CoQH2-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 QH2 at the QO 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 Qi 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 Qo site to form one quinone ( 2 H 2 + e − {\displaystyle {\ce {2H+2e-}}} ) at the Qi 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 (O2), 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(H2S).
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 FOF1 ATP synthase complex to make ATP via oxidative phosphorylation. ATP synthase is sometimes described as Complex V of the electron transport chain.[6] The FO 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.[7] The number of c subunits it has determines how many protons it will require to make the FO turn one full revolution. For example, in humans, there are 8 c subunits, thus 8 protons are required.[8] After c subunits, protons finally enters matrix using a subunit channel that opens into the mitochondrial matrix.[7] 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 F1 component of the complex.[9]
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.[10] Synthetic uncouplers (e.g., 2,4-dinitrophenol, 2,4-dinitrocresol, CCCP) also exist, and can be lethal at high doses.[citation needed]
Summary[edit]
In the mitochondrial electron transport chain electrons move from an electron donor (NADH or QH2) to a terminal electron acceptor (O2) 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.[11]
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 → O2 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 → bc1 → 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, H2 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.[citation needed]
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 bc1 is a proton pump found in many, but not all, bacteria (it is not found in E. coli). As the name implies, bacterial bc1 is similar to mitochondrial bc1 (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 (O2) 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 bc1 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.[citation needed]
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 O2/H2O redox pair. NADH is the electron donor and O2 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 10−4 atm. The associated redox reaction, which is thermodynamically favorable in nature, is thermodynamic impossible under "standard" conditions.[citation needed]
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., O2) 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: a b 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
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History of the Ottoman Empire during World War I
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Part of a series on the
History of the
Ottoman Empire
Rise
(1299–1453)[show]
Gaza Thesis
Interregnum (1402–1413)
Fall of Constantinople
Classical Age
(1453–1566)[show]
Sultanate of Women
Transformation
(1566–1703)[show]
Köprülü Era (1656–1703)
Old Regime
(1703–1789)[show]
Tulip Era (1718–1730)
Decline and modernization
(1789–1908)[show]
Nizam-i Djedid
Tanzimat Era (1839–1876)
1st Constitutional Era (1876–1878)
Defeat and dissolution
(1908–1922)[show]
2nd Constitutional Era (1908–1920)
World War I (1914–1918)
Partitioning
Historiography
v
t
e
The Ottoman Empire participated in World War I as one of the Central Powers. The Ottoman Empire entered the war by carrying out a surprise attack on Russia's Black Sea coast on 29 October 1914, with Russia responding by declaring war on it on November 5th, 1914. Ottoman forces fought the Entente in the Balkans and the Middle Eastern theatre of World War I. The Ottoman Empire's defeat in the war in 1918 was crucial in the eventual dissolution of the empire in 1921.
Contents
[hide]
1 Military activities
1.1 War with Russia
1.2 War with Britain
1.3 Empire in home front
1.4 War in Eastern Europe
2 Political activities
2.1 1915
2.1.1 Capitulations and public debt, 1915
2.2 1916
2.3 1917
2.3.1 Diplomacy with new Russia, 1917
2.4 1918
2.4.1 Diplomacy with new states, 1918
2.4.2 Winding down toward the armistice, 1918
3 Genocide
4 Notes
5 References
6 Bibliography
Military activities[edit]
Main articles: Ottoman entry into World War I and Middle Eastern theatre of World War I
The Ottoman entry into World War I began on 29 October 1914 when it launched the Black Sea Raid against Russian ports. Following the attack, Russia and its allies (Britain and France) declared war on the Ottomans in November 1914. The Ottoman Empire started military action came after three months of formal neutrality, but it had signed a secret alliance with the Central Powers in August 1914.
The political reasons for the Ottoman Sultan's entry into the war are disputed.[1] and the Ottoman Empire was an agricultural state in an age of industrialized warfare,[2] Also, the economic resources of the empire were depleted by the cost of the Balkan Wars of 1912 and 1913.
The great land mass of Anatolia was between the Ottoman army's headquarters in Istanbul and many of the theatres of war. During Abdulhamit II's reign civilian communications had improved, but the road and rail network was not ready for war.[2] It took more than a month to reach Syria and nearly two months to reach Mesopotamia. To reach the border with Russia, the railway ran only 60 km east of Ankara, and from there, it was 35 days to Erzurum.[2] The Army used Trabzon port as logistical shortcut to east. It took less time to arrive at any of those fronts from London than from the Ottoman War Department because of the poor condition of Ottoman supply ships.
The empire fell into disorder with the declaration of war along with Germany. On 11 November a conspiracy was discovered in Constantinople against Germans and the Committee of Union and Progress (CUP) in which some of the CUP leaders were shot. That followed the 12 November revolt in Adrianople against the German military mission. On 13 November, a bomb exploded in Enver Pasha's palace, which killed five German officers but missed the Enver Pasha. On 18 November were more anti-German plots. Committees formed around the country to rid the country of those who sided with Germany. Army and navy officers protested against the assumption of authority by Germans. On 4 December, widespread riots took place throughout the country. On 13 December was an anti-war demonstration by women in Konak (Izmir) and Erzurum. Throughout December, the CUP dealt with mutiny among soldiers in barracks and among naval crews. The head of the German Military Mission, Field Marshal von der Goltz, survived a conspiracy against his life.
The military power remained firmly in the hands of War Minister Enver Pasha, domestic issues (civil matters) on Interior Minister Talat Pasha, and, an interesting point, Cemal Pasha had sole control over Ottoman Syria.[3] Rest of the governance, provincial governors, ran their regions with differing degrees of autonomy.[3] An interesting case is Izmir; Rahmi Bey behaved almost as if his region was a neutral zone between the warring states.[4]
War with Russia[edit]
See also: Caucasus Campaign and Persian Campaign
Top: Destruction in the city of Erzurum; Left Upper: Russian forces; Left Lower: Wounded Muslim refugees; Right Upper:Ottoman forces; Right Lower: Armenian refuges
Ottoman's entrance into the war greatly increased the Triple Entente's military burdens. Russia had to fight on the Caucasus Campaign alone and in the Persian Campaign along with the United Kingdom. İsmail Enver Pasha set off for the Battle of Sarikamish with the intention of recapturing Batum and Kars, overrunning Georgia and occupying north-western Persia and the oil fields. Fighting the Russians in the Caucasus, however, the Ottomans lost ground, and over 100,000 soldiers, in a series of battles. 60,000 Ottoman soldiers died in the winter of 1916—17 on the Mus—Bitlis section of the front.[5] Ottomans preferred to keep the Caucasus militarily silent as they had to regroup reserves to retake Baghdad and Palestine from the British. 1917 and first half of 1918 was the time for negotiations. On 5 December 1917, the armistice of Erzincan (Erzincan Cease-fire Agreement) signed between the Russians and Ottomans in Erzincan that ended the armed conflicts between Russia and Ottoman Empire.[6] On 3 March, the Grand vizier Talat Pasha signed the Treaty of Brest-Litovsk with the Russian SFSR. It stipulated that Bolshevik Russia cede Batum, Kars, and Ardahan. In addition to these provisions, a secret clause was inserted which obligated the Russians to demobilize Armenian national forces.[7]
Between 14 March – April 1918 the Trabzon peace conference held among the Ottoman Empire and the delegation of the Transcaucasian Diet. Enver Pasha offered to surrender all ambitions in the Caucasus in return for recognition of the Ottoman reacquisition of the east Anatolian provinces at Brest-Litovsk at the end of the negotiations.[8] On 5 April, the head of the Transcaucasian delegation Akaki Chkhenkeli accepted the Treaty of Brest-Litovsk as a basis for more negotiations and wired the governing bodies urging them to accept this position.[9] The mood prevailing in Tiflis was very different. Tiflis acknowledge the existence of a state of war between themselves and the Ottoman Empire.[9]
In April 1918, the Ottoman 3rd Army finally went on the offensive in Armenia. Opposition from Armenian forces led to the Battle of Sardarapat, the Battle of Kara Killisse (1918), and the Battle of Bash Abaran. On 28 May 1918, the Armenian National Council based in Tiflis declared the Democratic Republic of Armenia. The new Republic of Armenia was forced to sign the Treaty of Batum.
In July 1918, Ottomans faced with the Centrocaspian Dictatorship at the Battle of Baku, with the goal of taking Britain Baku on the Caspian Sea.
War in the Caucasus and Persia
1st battalion of the Armenian volunteer unit. It was under the command of the Andranik during Persian Campaign.[10]
Terek Cossacks under Nikolai Baratov, Russian Caucasus Army
5th Infantry Regiment officers of the Battle of Sardarabad
War with Britain[edit]
See also: Mesopotamian campaign, Sinai and Palestine Campaign, and Battle of Gallipoli
February–April 1915, The Battle of Gallipoli
The British captured Basra in November 1914, and marched north into Iraq.[5] Initially Ahmed Djemal Pasha was ordered to gather an army in Palestine to threaten the Suez Canal. In response, the Allies—including the newly formed Australian and New Zealand Army Corps ("ANZACs")—opened another front with the Battle of Gallipoli. The army led by Ahmed Djemal Pasha (Fourth Army) to eject the British from Egypt was stopped at the Suez canal in February 1915, and again the next summer.[5] The canal was vital to the British war effort. The 1915 locust plague breaks out in the Palestine region, to be exact the Ottoman military hospitals record the period as March–October 1915:
The expected, and feared, British invasion came not through Cilicia or northern Syria, but through the straits.[3] The aim of the Dardanelles campaign was to support Russia. Most military observers recognized that the uneducated Ottoman soldier was lost without good leadership, and at Gallipoli Mustafa Kemal realized the capabilities of his man if their officers led from the front.[11] The war was something from a different era, as the agrarian Ottoman Empire faced two industrialized forces, at silent predawn attacks in which officers with drawn swords vent ahead of troops and only the troops to shout their battlecry of "Allahu Akbar!" when they reached the enemy’s trenches.[11]
The United Kingdom was obliged to defend India and the southern Persian oil territory by undertaking the Mesopotamian campaign. Britain also had to protect Egypt in the Sinai-Palestine-Syria Campaign. These campaigns strained Allied resources and relieved Germany.
The repulse of British forces in Palestine in the spring of 1917 was followed by the loss of Jerusalem in December of the same year.[5] The Ottoman authorities deport the entire civilian population of Jaffa and Tel Aviv, The Tel Aviv and Jaffa deportation, pursuant to the order from Ahmed Jamal Pasha on 6 April 1917. The Muslim evacuees allowed to return before long, At the same period the Balfour Declaration was being negotiated (published on 2 November 1917) in which the British Government declares its support for the establishment of a Jewish national home in Palestine. Ahmed Jamal Pasha effectively separates these groups. The Jewish evacuees returned after the British conquest of Palestine.[12]
The Ottomans were eventually defeated due to key attacks by the British general Edmund Allenby.
War in Mesopotamia, Sinai and Palestine and Gallipoli
British troops entering Baghdad March 1917
6th Army field HQ
In 1917, Ottoman forces at the shores of the Dead Sea
Empire in home front[edit]
See also: Arab Revolt and Armenian resistance
The war tested to the limit the empire’s relations with its Arab population.[13] In February 1915 in Syria, Cemal Pasha exercised absolute power in both military and civil affairs.[14] Cemal Pasha was convinced that an uprising among local Arabs was imminent.[13] Leading Arabs were executed, and notable families deported to Anatolia.[13] Cemal’s policies did nothing to alleviate the famine that was gripping Syria; it was exacerbated by a British and French blockade of the coastal ports, the requisitioning of transports, profiteering and — strikingly — Cemal’s preference for spending scarce funds on public works and the restoration of historic monuments[13][15] During the war, Britain had been a major sponsor of Arab nationalist thought and ideology, primarily as a weapon to use against the power of the Empire. Sharif Hussein ibn Ali rebelled against the Ottoman rule during the Arab Revolt of 1916. In August he was replaced by Sharif Haydar, but in October he proclaimed himself king of Arabia and in December was recognized by the British as an independent ruler.[13] There was little the Empire could do to influence the course of events, other than try to prevent news of the uprising spreading, prevent it to demoralize the army or act as a propaganda for anti-Ottoman Arab factions.[13] On 3 October 1918 forces of the Arab Revolt enter Damascus accompanied by British troops, ending 400 years of Ottoman rule.
Home front
Lt. Col. Gerard Leachman disguised as a Bedouin.
Soldiers of the Sharif of Mecca carrying the Arab Flag during the Arab Revolt of 1916–1918
Resistance members from the Adapazari committee, 1915.
Murad of Sebastia and his comrades fought at Sivas during 1915
War in Eastern Europe[edit]
In order to support the other Central Powers, Enver Pasha sent 3 Army Corps or around 100.000 men to fight in Eastern Europe. [16]
VI Corps under command of Mustafa Hilmi Pasha participated in the Romanian Campaign between September 1916 and April 1918.
XV Corps under command of Yakup Şevki Subaşı and later Cevat Pasha fought in Galicia against the Russians between August 1916 and August 1917.
XX Corps under command of Abdul Kerim Pasha participated in the Salonika Campaign between December 1916 and May 1917.
The Rumeli Field Detachment (reinforced 177th Infantry Regiment) remained in Macedonia until May 1918.
Political activities[edit]
1915[edit]
The Constantinople Agreement on 18 March 1915 was a set of secret assurances, which Great Britain promised to give the Capital, and the Dardanelles to the Russians in the event of victory.[17] The city of Constantinople was intended to be a free port.
During 1915, British forces invalidated the Anglo-Ottoman Convention, declaring Kuwait to be an "independent sheikdom under British protectorate."[this quote needs a citation]
Capitulations and public debt, 1915[edit]
On 10 September 1915, Interior Minister Talat Pasha abolished the "Capitulations". On 10 September 1915 Grand Vizier Said Halim Pasha annulled (Vizer had the authority on annuls) the Capitulations, which ended the special privileges they granted to foreign nationals. The capitulation holders refused to recognize his action (unilateral action).[2] The American ambassador expressed the Great Power view:
The capitulary regime, as it exists in the Empire, is not an autonomous institution of the Empire, but the result of international treaties, of diplomatic agreements and of contractual acts of various sorts. The regime, consequently, cannot be modified in any of its parts and still less suppressed in its entirety by the Ottoman Government except in consequence of an understanding with the contracting Powers.[18]
—  Henry Morgenthau, Sr.
Beside the capitulations, there was another issue which evolved under the shadow of capitulations. The dept and financial control (revenue generation) of the empire was intertwined under single institution, which its board was constituted from Great Powers rather than Ottomans. There is no sovereignty in this design. The public debt could and did interfere in state affairs because it controlled (collected) one-quarter of state revenues.[18] The debt was administered by the Ottoman Public Debt Administration and its power extended to the Imperial Ottoman Bank (equates to modern central banks). Debt Administration controlled many of the important revenues of the empire. The council had power over every financial affairs. Its control extended to determine the tax on livestock in districts. Ottoman public debt was part of a larger scheme of political control, through which the commercial interests of the world had sought to gain advantages that may not be to Empire's interest. The immediate purpose of the abolition of capitulations and the cancellation of foreign debt repayments was to reduce the foreign stranglehold on the Ottoman economy; a second purpose — and one to which great political weight was attached – was to extirpate non—Muslims from the economy by transferring assets to Muslim Turks and encouraging their participation with government contracts and subsidies.[19]
1916[edit]
The French-Armenian Agreement of 27 October 1916, was reported to the interior minister, Talat Pasha, which agreement negotiations were performed with the leadership of Boghos Nubar the chairman of the Armenian National Assembly and one of the founder of the AGBU.
1917[edit]
In 1917 the Ottoman Cabinet considered maintaining relations with Washington after the United States had declared war on Germany on 6 April. But the views of the war party prevailed and they insisted on maintaining a common front with their allies. Thus, relations with America were broken on 20 April 1917.
Diplomacy with new Russia, 1917[edit]
The 1917 Russian revolution changed the realities. The war devastated not only Russian soldiers, also the Russian economy was breaking down under the heightened strain of wartime demand by the end of 1915. The tsarist regime’s advances for the security on its southern borders proved ruinous.[20] The tsarist regime's desire to control the Eastern Anatolia and the straits (perceived as an underbelly), in the end created the conditions that brought about Russia's own downfall. Unable to use Straits disrupted the Russian supply chain. Russia might survived without the Straits, but the strain was the tipping point for its war economy.[20] This question was left to Soviet historians: "whether a less aggressive policy toward the Ottoman Empire before the war would have caused Istanbul to maintain neutrality or whether Russia later might have induced Istanbul to leave the war,[a] the outcome of tsarist future would be different.[20] Nicholas's inept handling of his country and the war destroyed the Tsar and ended up costing him both his reign and his life.
Enver immediately instructed the Vehib Pasha, Third Army, to propose a ceasefire to Russia’s Caucasus Army.[21] Vehib cautioned withdrawing forces, as due to the politics in Russia — neither Russia’s Caucasus Army nor Caucasian civil authorities give assurance that an armistice would hold.[22] On 7 November 1917 the Bolshevik Party led by Vladimir Lenin over threw the Provisional Government in a violent coup plunged Russia into multitude of civil wars between ethnic groups. The slow dissolution of Russia’s Caucasus Army relieved one form of military threat from the east but brought another one. Russia was a long time threat, but at the same time kept the civil unrest in his land at bay without spreading to Ottomans in a violent. On 3 December the Ottoman foreign minister Ahmed Nesimi Bey informed the "Chamber of Deputies" about the prospects. Chamber discussed the possible outcomes and priorities. On 15 December Armistice between Russia and the Central Powers signed. On 18 December Armistice of Erzincan signed. The Bolsheviks’ anti-imperialist formula of peace with no annexations and no indemnities was close to Ottoman position. The Bolsheviks' position brought a conflict with the Germany's aim to preserve control over the East European lands it occupied and with Bulgaria’s claims on Dobruja and parts of Serbia. In December Enver informed the Quadruple Alliance that they would like to see the 1877 border (Russo-Turkish War (1877–1878)), pointing out that the only Ottomans lost territory and 1877 boarder was Ottoman territories inhabited by Muslims.[23] Ottomans did not pushed 1877 position too hard, scared to fall back to bilateral agreements. On the other hand, Germany, Austria-Hungary, and Bulgaria clearly stood behind on the pulling back the Ottoman and Russian forces from Iran.[24] Ottomans wanted Muslim Iran be under its own control. The ambassador to Berlin, Ibrahim Hakki Pasha, wrote: "Although Russia may be in a weakened state today, it is always an awesome enemy and it is probable that in a short time it will recover its former might and power.[23]
On 22 December 1917, the first meeting between Ottomans and the Bolsheviks, the temporary head Zeki Pasha, until Talat Pasha's arrival, requested of Lev Kamenev to put an end to atrocities being committed on Russian-occupied territory by Armenian partisans. Kamenev agreed and added "an international commission should be established to oversee the return of refugees (by own consent) and deportees (by forced relocation) to Eastern Anatolia. The battle of ideals, rhetoric, and material for the fate of Eastern Anatolia opened with this dialog .[23]
The Treaty of Brest-Litovsk represented an enormous success for the empire.[according to whom?] Minister of Foreign Affairs Halil Bey announced the achievement of peace to the Chamber of Deputies. He cheered the deputies further with his prediction of the imminent signing of a third peace treaty (the first Ukraine, second Russia, and with Romania). Halil Bey thought the Entente to cease hostilities and bring a rapid end to the war. The creation of an independent Ukraine promised to cripple Russia, and the recovery of Kars, Ardahan and Batum gave the CUP a tangible prize. Nationalism emerged at the center of the diplomatic struggle between the Central Powers and the Bolsheviks. Empire recognized that Russia’s Muslims, their co-religionists, are disorganized and dispersed to come out as an entity in the future battles of ideals, rhetoric, and material. Thus, the Ottomans mobilized the Caucasus Committee to make claims on behalf of the Muslims.[25] The Caucasus Committee had declined Ottoman earnest requests to break from Russia and embrace independence. The Caucasian Christians was far ahead in this new world concept. Helping the Caucasian Muslims to be free, like their neighbors, would be the Ottomans’ challenge.[25]
1918[edit]
In the overall war effort, the CUP was convinced that empire's contribution was essential. Ottoman armies had tied down large numbers of Allied troops on various fronts, keeping them away from theatres in Europe where they would have been used against German and Austrian forces. Moreover, they claimed that their success at Gallipoli had been an important factor in bringing about the collapse of Russia, resulting in the revolution of April 1917. They had turned the war in favor of Germany and her allies.[26] Hopes were initially high for the Ottomans that their losses in the Middle East might be compensated for by successes in the Caucasus Campaign. Enver Pasha maintained an optimistic stance, hid information that made the Ottoman position appear weak, and let most of the Ottoman elite believe that the war was still winnable.[27]
Diplomacy with new states, 1918[edit]
Ottoman policy toward the Caucasus evolved according to the changing demands of the diplomatic and geopolitical environment.[28] What was the Ottoman premise in involving with the Azerbaijan and the North Caucasus? The Empire’s leaders, in the parliament discussions throughout 1917, understood that Russia’s collapse presented a historic window of opportunity to redraw the map of the Caucasus. They were convinced, however, that soon enough Russia would recover and reemerge as the dominant power in the region and shut that window.
The principle of "self-determination" became the criterion, or at least in part, to give them a chance to stand on their feet.[29] The Bolsheviks did not regard national separatism in this region as a lasting force. Their expectation was whole region come under a "voluntary and honest union" [b] and this union bearing no resemblance to Lenin’s famous description of Russia as a "prison house of peoples."[30] Lenin's arrival to Russia was formally welcomed by Nikolay Chkheidze, the Menshevik Chairman of the Petrograd Soviet.
Ottoman's did not see a chance of these new states to stand against new Russia. These new Muslim states needed support to be emerged as viable independent states. In order to consolidate a buffer zone with Russia (both for the Empire and these new states), however, Ottomans needed to expel the Bolsheviks from Azerbaijan and the North Caucasus before the end of war.[31] Based on 1917 negotiations, Enver concluded that Empire should not to expect much military assistance from the Muslims of the Caucasus as they were the one in need. Enver also knew the importance of Kars—Julfa railroad and the adjacent areas for this support. Goal was set forward beginning from 1918 to end of the war.
The Empire duly recognized the Transcaucasian Democratic Federative Republic in February 1918. This preference to remain part of Russia led Caucasusian politics to the Trebizond Peace Conference to base their diplomacy on the incoherent assertion that they were an integral part of Russia but yet not bound[28] The representatives were Rauf Bey for the Empire, and Akaki Chkhenkeli from the Transcaucasian delegation.
On 11 May, a new peace conference opened at Batum. The Treaty of Batum was signed on 4 June 1918, in Batum between the Ottoman Empire and three Trans-Caucasus states: First Republic of Armenia, Azerbaijan Democratic Republic and Democratic Republic of Georgia.
The goal was to assist Azerbaijan Democratic Republic at Battle of Baku, then turn north to assist the embattled Mountainous Republic of the Northern Caucasus and then sweep southward to encircle the British in Mesopotamia and retake Baghdad.[29] The British in Mesopotamia already moving north, with forty vans (claimed to loaded with gold and silver for buying mercenary) accompanied with only a brigade, to establish a foothold. At the time Baku was under the control of the 26 Baku Commissars which were Bolshevik and Left Socialist Revolutionary (SR) members of the Baku Soviet Commune. The commune was established in the city of Baku. In this plan, they expected resistance from Bolshevik Russia and Britain, but also Germany, which opposed the extension of their influence into the Caucasus.[29] Ottoman's goal to side with Muslims of Azerbaijan and MRNC managed to get Bolsheviks of Russia, Britain and Germany on the same side of a conflict box at this brief point in the history.
Diplomacy with new states
Nikolay Chkheidze
Hovhannes Katchaznouni [c]
Mammad Amin Rasulzade
Leaders of the MRNC.
Azerbaijani National Council, 28 May 1918
Winding down toward the armistice, 1918[edit]
İkdam on 4 November 1918 announcing Enver, Talat, Cevdet left the country.
Developments in Southeast Europe squashed the Ottoman government's hopes. In September 1918, the Allied forces under the command of Louis Franchet d'Espèrey mounted a sudden offensive at the Macedonian Front, which proved quite successful. Bulgaria was forced to sue for peace in the Armistice of Salonica. This development undermined both the German and Ottoman cause simultaneously - the Germans had no troops to spare to defend Austria-Hungary from the newly formed vulnerability in Southeast Europe after the losses it had suffered in France, and the Ottomans suddenly faced having to defend Istanbul against an overland European siege without help from the Bulgarians.[27]
Grand Vizier Talaat Pasha visited both Berlin, and Sofia, in September 1918, and came away with the understanding that the war was no longer winnable. With Germany likely seeking a separate peace, the Ottomans would be forced to as well. Grand Vizier Talaat convinced the other members of the ruling party that they must resign, as the Allies would impose far harsher terms if they thought the people who started the war were still in power. He also sought out the United States to see if he could surrender to them and gain the benefits of the Fourteen Points despite the Ottoman Empire and the United States not being at war; however, the Americans never responded, as they were waiting on British advice as to how to respond which never came. On 13 October, Talaat and the rest of his ministry resigned. Ahmed Izzet Pasha replaced Talaat as Grand Vizier.
Two days after taking office, Ahmed Izzet Pasha sent the captured British General Charles Vere Ferrers Townshend to the Allies to seek terms on an armistice.[27] The British Cabinet were eager to negotiate a deal. British government interpreted that not only should Britain conduct the negotiations, but should conduct them alone. There may be a desire to cut the French out of territorial "spoils" promised to them in the Sykes-Picot agreement. Talaat (before resigning) had sent an emissary to the French as well, but that emissary had been slower to respond back. The British cabinet empowered Admiral Calthorpe to conduct the negotiations, and to explicitly exclude the French from them.[27] The negotiations began on Sunday, 27 October on the HMS Agamemnon, a British battleship. The British refused to admit French Vice-Admiral Jean Amet, the senior French naval officer in the area, despite his desire to join; the Ottoman delegation, headed by Minister of Marine Affairs Rauf Bey.[27]
Unknown to both sides, both sides were actually quite eager to sign a deal and willing to give up their objectives to do so. The British delegation had been given a list of 24 demands, but were told to concede on any of them except allowing the occupation of the forts on the Dardanelles as well as free passage through the Bosphorus; the British desired access to the Black Sea for the Rumanian front. Prime Minister David Lloyd George also desired to make a deal quickly before the United States could step in; according to the diary of Maurice Hankey:
[Lloyd George] was also very contemptuous of President Wilson and anxious to arrange the division of Empire between France, Italy, and G.B. before speaking to America. He also thought it would attract less attention to our enormous gains during the war if we swallowed our share of Empire now, and the German colonies later.[27]
The Ottomans, for their part, believed the war to be lost and would have accepted almost any demands placed on them. As a result, the initial draft prepared by the British was accepted largely unchanged; the Ottomans did not know they could have pushed back on most of the clauses, and the British did not know they could have demanded even more. The Ottomans ceded the rights to the Allies to occupy "in case of disorder" any Ottoman territory, a vague and broad clause.[27] The French were displeased with the precedent; French Premier Clemenceau disliked the British making unilateral decisions in so important a matter. Lloyd George countered that the French had concluded a similar armistice on short notice in the Armistice of Salonica which had been negotiated by French General d'Esperey, and that Great Britain (and Czarist Russia) had committed the vast majority of troops to the campaign against the Ottomans. The French agreed to accept the matter as closed.
On 30 October 1918, the Armistice of Mudros was signed, ending Ottoman involvement in World War 1. The Ottoman public, however, was given misleadingly positive impressions of the severity of the terms of the Armistice. They thought its terms were considerably more lenient than they actually were, a source of discontent later that the Allies had betrayed the offered terms.[27]
Genocide[edit]
Main articles: Armenian Genocide, Assyrian genocide, and Greek genocide
During WWI the Ottoman Empire engaged in a genocide against local ethnicities in its territory. The Armenian Genocide,[32] also known as the Armenian Holocaust,[33] was the Ottoman government's systematic extermination of 1.5 million Christian Armenians, mostly Ottoman citizens within the Ottoman Empire and its successor state, the Republic of Turkey.[34][35] The starting date is conventionally held to be 24 April 1915, the day that Ottoman authorities rounded up, arrested, and deported 235 to 270 Armenian intellectuals and community leaders from Constantinople to Ankara, the majority of whom were eventually murdered. The genocide was carried out during and after World War I and implemented in two phases: the wholesale killing of the able-bodied male population through massacre and subjection of army conscripts to forced labour, followed by the deportation of women, children, the elderly, and the infirm on death marches leading to the Syrian desert. Driven forward by military escorts, the deportees were deprived of food and water and subjected to periodic robbery, rape, and massacre.[36][37][38] Other indigenous and Christian ethnic groups such as the Assyrians and the Ottoman Greeks were similarly targeted for extermination by the Ottoman government in the Assyrian genocide and the Greek genocide, and their treatment is considered by some historians to be part of the same genocidal policy.[39][40] Most Armenian diaspora communities around the world came into being as a direct result of the genocide.[41]
Raphael Lemkin was explicitly moved by the Armenian annihilation to define systematic and premeditated exterminations within legal parameters and to coin the word genocide in 1943.[42] The Armenian Genocide is acknowledged to have been one of the first modern genocides,[43][44][45] because scholars point to the organized manner in which the killings were carried out in order to eliminate the Armenians, and it is the second most-studied case of genocide after the Holocaust.[46]
Turkey, the successor state of the Ottoman Empire, denies the word genocide as an accurate term for the mass killings of Armenians that began under Ottoman rule in 1915. It has in recent years been faced with repeated calls to recognize them as genocide.[47] To date, 29 countries have officially recognized the mass killings as genocide,[48] as have most genocide scholars and historians.[49][50][51]
Notes[edit]
Jump up ^ Sazonov’s commitment to a two-front war and disregard for Yudenich’s warnings to seek peace with the Ottomans had overstretched Russia.[20]
Jump up ^ The reference "voluntary and honest union" realized with the Soviet Union, as the 11th Red Army had its virtually unopposed advance to the region on 29 November 1920.
Jump up ^ Hovhannes Katchaznouni was in city of Van until 1914. He was on the Armenian delegation that conducted peace talks at Trabzon and Batoum negotiations with the Empire.
Includes most of to daysader nation of Europe
References[edit]
Jump up ^ Nicolle 2008, p. 167
^ Jump up to: a b c d Finkel 2007, p. 529
^ Jump up to: a b c Nicolle 2008, p. 174
Jump up ^ Nicolle 2008, p. 178
^ Jump up to: a b c d Finkel 2007, p. 530
Jump up ^ Tadeusz Swietochowski, Russian Azerbaijan 1905–1920, page 119.
Jump up ^ Hovannisian. "Armenia's Road to Independence", pp. 288–289. ISBN 1-4039-6422-X.
Jump up ^ (Shaw 1977, p. 326)
^ Jump up to: a b Richard Hovannisian "The Armenian people from ancient to modern times" Pages 292–293
Jump up ^ Aram, "Why Armenia Should be Free", page 22
^ Jump up to: a b Nicolle 2008, p. 176
Jump up ^ Friedman, Isaiah (1971). German Intervention on Behalf of the "Yishuv", 1917 , Jewish Social Studies, Vol. 33, pp. 23–43.
^ Jump up to: a b c d e f Finkel 2007, p. 537
Jump up ^ Finkel 2007, p. 531
Jump up ^ Melanie S. Tanielian. The Charity of War: Famine, Humanitarian Aid, and World War I in the Middle East (Stanford, CA.: Stanford University Press, 2017) http://www.sup.org/books/title/?id=28143
Jump up ^ [1] turkeyswar, Campaigns, Eastern Europe.
Jump up ^ The Greenwood Encyclopedia of International Relations: A-E, Ed. Cathal J. Nolan, (Greenwood Publishing Group, 2002), 350.
^ Jump up to: a b Kent 1996, p. 19
Jump up ^ Finkel 2007, pp. 536
^ Jump up to: a b c d Reynolds 2011, p. 253
Jump up ^ Reynolds 2011, p. 170
Jump up ^ Reynolds 2011, p. 171
^ Jump up to: a b c Reynolds 2011, p. 172
Jump up ^ Reynolds 2011, p. 173
^ Jump up to: a b Reynolds 2011, p. 190
Jump up ^ Kent 1996, p. 16
^ Jump up to: a b c d e f g h (Fromkin 2009, pp. 360–373)
^ Jump up to: a b Reynolds 2011, p. 217
^ Jump up to: a b c Reynolds 2011, p. 221
Jump up ^ Reynolds 2011, p. 192
Jump up ^ Reynolds 2011, p. 218
Jump up ^ Armenian Genocide (affirmation), The International Association of Genocide Scholars, That this assembly of the Association of Genocide Scholars in its conference held in Montreal, June 11–3, 1997, reaffirms that the mass murder of Armenians in Turkey in 1915 is a case of genocide which conforms to the statutes of the United Nations Convention on the Prevention and Punishment of Genocide. It further condemns the denial of the Armenian Genocide by the Turkish government and its official and unofficial agents and supporters.
Jump up ^ Fisk, Robert (14 October 2006). "Let me denounce genocide from the dock". The Independent. Archived from the original on 24 January 2014. Retrieved 31 August 2016.
Jump up ^ "8 facts about the Armenian genocide 100 years ago - CNN.com". CNN. Retrieved 2015-12-13.
Jump up ^ "100 Years Ago, 1.5 Million Armenians Were Systematically Killed. Today, It's Still Not A 'Genocide.'". The Huffington Post. Retrieved 2015-12-13.
Jump up ^ Kieser, Hans-Lukas; Schaller, Dominik J. (2002), Der Völkermord an den Armeniern und die Shoah [The Armenian genocide and the Shoah] (in German), Chronos, p. 114, ISBN 3-0340-0561-X
Jump up ^ Walker, Christopher J. (1980), Armenia: The Survival of A Nation, London: Croom Helm, pp. 200–3
Jump up ^ Bryce, Viscount James; Toynbee, Arnold (2000), Sarafian, Ara, ed., The Treatment of Armenians in the Ottoman Empire, 1915–1916: Documents Presented to Viscount Grey of Falloden (uncensored ed.), Princeton, NJ: Gomidas Institute, pp. 635–49, ISBN 0-9535191-5-5
Jump up ^ Schaller, Dominik J; Zimmerer, Jürgen (2008). "Late Ottoman genocides: the dissolution of the Ottoman Empire and Young Turkish population and extermination policies – introduction". Journal of Genocide Research. 10 (1): 7–14. doi:10.1080/14623520801950820.
Jump up ^ Jones, Adam (2010). Genocide: A Comprehensive Introduction. Taylor & Francis. pp. 171–172. ISBN 978-0-203-84696-4. A resolution was placed before the IAGS membership to recognize the Greek and Assyrian/Chaldean components of the Ottoman genocide against Christians, alongside the Armenian strand of the genocide (which the IAGS has already formally acknowledged). The result, passed emphatically in December 2007 despite not inconsiderable opposition, was a resolution which I co-drafted, reading as follows:...
Jump up ^ "The Many Armenian Diasporas, Then and Now". GeoCurrents. Retrieved 13 December 2015.
Jump up ^ Totally Unofficial: The Autobiography of Raphael Lemkin. New Haven and London: Yale University Press. 2013. pp. 19–20.
Jump up ^ "Council of Europe Parliamentary Assembly Resolution". Armenian genocide. Retrieved 17 June 2016.
Jump up ^ Ferguson, Niall (2006). The War of the World: Twentieth-Century Conflict and the Descent of the West. New York: Penguin Press. p. 177. ISBN 1-59420-100-5.
Jump up ^ "A Letter from The International Association of Genocide Scholars" (PDF). Genocide Watch. 13 June 2005.
Jump up ^ Rummel, RJ (1 April 1998), "The Holocaust in Comparative and Historical Perspective", The Journal of Social Issues, 3 (2)
Jump up ^ "For Turkey, denying an Armenian genocide is a question of identity". america.aljazeera.com. Retrieved 2015-12-13.
Jump up ^ "National Assembly of the Republic of Armenia - Official Web Site - parliament.am". parliament.am. Archived from the original on 27 June 2015.
Jump up ^ Renwick Monroe, Kristen (2012). Ethics in an Age of Terror and Genocide: Identity and Moral Choice. Princeton University Press. p. 13. ISBN 0-691-15143-1. Retrieved 17 June 2016.
Jump up ^ Loytomaki, Stiina (2014). Law and the Politics of Memory: Confronting the Past. Routledge. p. 31. ISBN 978-1-136-00736-1. To date, more than 20 countries in the world have officially recognized the events as genocide and most historians and genocide scholars accept this view.
Jump up ^ Frey, Rebecca Joyce (2009). Genocide and International Justice. Infobase Publishing. p. 83. ISBN 978-0-8160-7310-8. Retrieved 15 April 2016.
Bibliography[edit]
Erickson, Edward J. (2001). Ordered to Die: A History of the Ottoman Army in the First World War. Westport, CT: Greenwood.
Fromkin, David (2009). A Peace to End All Peace: The Fall of the Ottoman Empire and the Creation of the Modern Middle East. Macmillan. ISBN 978-0-8050-8809-0.
Finkel, Caroline (2007). Osman's Dream: The History of the Ottoman Empire. Basic Books.
Kent, Marian (1996). The Great Powers and the End of the Ottoman Empire. Routledge. ISBN 0714641545.
Nicolle, David (2008). The Ottomans: Empire of Faith. Thalamus Publishing. ISBN 1902886119.
Francesco Pongiluppi (2015). The Energetic Issue as a Key Factor of the Fall of the Ottoman Empire. in "The First World War: Analysis and Interpretation" (ed. by Biagini and Motta), Vol. 2., Newcastle, Cambridge Scholars Publishing, pp. 453–464.
Reynolds, Michael A. (2011). Shattering Empires: The Clash and Collapse of the Ottoman and Russian Empires 1908–1918. Cambridge University Press. p. 324. ISBN 0521149169.
Shaw, Ezel Kural (1977). History of the Ottoman Empire and Modern Turkey. New York: Cambridge University Press. ISBN 0-521-21280-4.
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value: Osmanlı imperiyası Birinci dünya müharibəsində – Azerbaijani

value: Асманская імперыя ў Першай сусветнай вайне – Belarusian

value: Imperio otomano durante la Primera Guerra Mundial – Spanish

value: Empire ottoman dans la Première Guerre mondiale – French

value: Օսմանյան կայսրությունն Առաջին համաշխարհային պատերազմում – Armenian

value: Османская империя в Первой мировой войне – Russian

value: Osmanska riket under första världskriget – Swedish

value: Османська імперія в Першій світовій війні – Ukrainian

value: Lịch sử đế quốc Ottoman trong Đệ nhất thế chiến – Vietnamese

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