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Alt, Christopher ; Antičić, Tome ; Baatar, B. ; Barna, Daniel ; Bartke, Jerzy ; Betev, Latchezar ; Białkowska, Helena ; Blume, Christoph ; Boimska, Bożena ; Botje, Michiel ; Bracinik, Juraj ; Bramm, Roland ; Bunčić, Predrag ; Cerny, Vladimír ; Christakoglou, Panagiotis ; Chvala, Ondrej ; Cramer, John G. ; Csató, Péter ; Dinkelaker, Peter ; Eckardt, Volker ; Flierl, Dominik Bernhard ; Fodor, Zoltán ; Foka, Panagiota ; Friese, Volker ; Gál, József ; Gaździcki, Marek ; Genchev, Vladimir ; Georgopoulos, George ; Gładysz-Dziaduś, Ewa ; Grebieszkow, Katarzyna ; Hegyi, Sándor ; Höhne, Claudia ; Kadija, Krešo ; Karev, Alexandre ; Kliemant, Michael ; Kniege, Stefan ; Kolesnikov, Vadim I. ; Kornas, Ewelina ; Korus, Robert ; Kowalski, Marek ; Kraus, Ingrid ; Kreps, Michal ; Leeuwen, Marco van ; Lévai, Péter ; Litov, Leandr ; Lungwitz, Benjamin ; Makariev, Martin ; Malakhov, Aleksandr I. ; Markert, Christina ; Mateev, Matej D. ; Melkumov, Georgy L. ; Mischke, André ; Mitrovski, Michael Kosta ; Molnár, József ; Mrówczyński, Stanislaw ; Nicolic, V. ; Pálla, Gabriella ; Panagiotou, Apostolos D. ; Panayotov, Dobromir ; Petridis, Athanasios ; Pikna, Miroslav ; Prindle, Duncan J. ; Pühlhofer, Falk ; Renfordt, Rainer Arno Ernst ; Roland, Christof ; Roland, Gunther ; Rybczyński, Maciej ; Rybicki, Andrzej ; Sandoval, Andres ; Schmitz, Norbert ; Schuster, Tim ; Seyboth, Peter ; Siklér, Ferenc ; Sitar, Branislav ; Skrzypczak, Ewa ; Stefanek, Grzegorz ; Stock, Reinhard ; Ströbele, Herbert ; Šuša, Tatjana ; Szentpétery, Imre ; Sziklai, János ; Szymański, Paweł ; Trubnikov, Victor ; Varga, Dezsö ; Vassiliou, Maria ; Veres, Gábor I. ; Vesztergombi, György ; Vranić, Danilo ; Wetzler, Alexander Albert ; Włodarczyk, Zbigniew ; Yoo, In-Kwon ; Zimányi, József
Results are presented from a search for the decays D0 -> K min pi plus and D0 bar -> K plus pi min in a sample of 3.8x10^6 central Pb-Pb events collected with a beam energy of 158A GeV by NA49 at the CERN SPS. No signal is observed. An upper limit on D0 production is derived and compared to predictions from several models.
Low energy beam transport (LEBT) for a future heavy ion driven inertial fusion (HIDIF [1]) facility is a crucial point using a Bi+ beam of 40 mA at 156 keV. High space charge forces (generalised perveance K=3.6*10-3) restrict the use of electrostatic focussing systems. On the other hand magnetic lenses using space charge compensation suffer from the low particle velocity. Additionally the emittance requirements are very high in order to avoid particle losses in the linac and at ring injection [2]. urthermore source noise and rise time of space charge compensation [3] might enhance particle losses and emittance. Gabor lenses [4] using a continuous space charge cloud for focussing could be a serious alternative to conventional LEBT systems. They combine strong cylinder symmetric focussing with partly space charge compensation and low emittance growth due to lower non linear fields. A high tolerance against source noise and current fluctuations and reduced investment costs are other possible advantages. The proof of principle has already been shown [5, 6]. To broaden the experiences an experimental program was started. Therefrom the first experimental results using a double Gabor lens (DGPL, see fig. 1 ) LEBT system for transporting an high perveance Xe+ beam will be presented and the results of numerical simulations will be shown.
Mischke, André ; Afanasiev, Sergey V. ; Antičić, Tome ; Baatar, B. ; Barna, Daniel ; Bartke, Jerzy ; Barton, Robert Allan ; Behler, Matthias ; Betev, Latchezar ; Białkowska, Helena ; Billmeier, Anja ; Blume, Christoph ; Blyth, Charles O. ; Boimska, Bożena ; Botje, Michiel ; Bracinik, Juraj ; Bramm, Roland ; Brun, René ; Bunčić, Predrag ; Cerny, Vladimír ; Chvala, Ondrej ; Cramer, John G. ; Csató, Péter ; Dinkelaker, Peter ; Eckardt, Volker ; Filip, Peter ; Fischer, Hans Gerhard ; Fodor, Zoltán ; Foka, Panagiota ; Freund, Peter ; Friese, Volker ; Gál, József ; Gaździcki, Marek ; Georgopoulos, George ; Gładysz-Dziaduś, Ewa ; Hegyi, Sándor ; Höhne, Claudia ; Igo, George J. ; Jones, Peter Graham ; Kadija, Krešo ; Karev, Alexandre ; Kolesnikov, Vadim I. ; Kollegger, Thorsten ; Kowalski, Marek ; Kraus, Ingrid ; Kreps, Michal ; Leeuwen, Marco van ; Lévai, Péter ; Malakhov, Aleksandr I. ; Margetis, Spyridon ; Markert, Christina ; Mayes, Bill W. ; Melkumov, Georgy L. ; Meurer, Christine ; Mischke, André ; Mitrovski, Michael Kosta ; Molnár, József ; Nelson, John M. ; Pálla, Gabriella ; Panagiotou, Apostolos D. ; Perl, Katarzyna ; Petridis, Athanasios ; Pikna, Miroslav ; Pinsky, Lawrence ; Pühlhofer, Falk ; Reid, Jeffrey G. ; Renfordt, Rainer Arno Ernst ; Retyk, Waldemar ; Roland, Christof ; Roland, Gunther ; Rybicki, Andrzej ; Sammer, Thomas ; Sandoval, Andres ; Sann, Hans ; Schmitz, Norbert ; Seyboth, Peter ; Siklér, Ferenc ; Sitar, Branislav ; Skrzypczak, Ewa ; Squier, Gordon Thomas Anderson ; Stock, Reinhard ; Ströbele, Herbert ; Šuša, Tatjana ; Szentpétery, Imre ; Sziklai, János ; Trainor, Thomas A. ; Varga, Dezsö ; Vassiliou, Maria ; Veres, Gábor I. ; Vesztergombi, György ; Vranić, Danilo ; Wenig, Siegfried ; Wetzler, Alexander Albert ; Whitten, Charles A. ; Yoo, In-Kwon ; Zaranek, Jacek ; Zimányi, József
Rapidity distributions for Lambda and anti-Lambda hyperons in central Pb-Pb collisions at 40, 80 and 158 AGeV and for K 0 s mesons at 158 AGeV are presented. The lambda multiplicities are studied as a function of collision energy together with AGS and RHIC measurements and compared to model predictions. A different energy dependence of the Lambda/pi and anti-Lambda/pi is observed. The anti-Lambda/Lambda ratio shows a steep increase with collision energy. Evidence for a anti-Lambda/anti-p ratio greater than 1 is found at 40 AGeV.
Die vorliegende Arbeit hat es sich zur Aufgabe gemacht, mit der Themenstellung "Netzaktivismus" eine relativ aktuelle Entwicklung aufzugreifen. Als nötig erschien mir hierbei von vornherein, den eigentlichen Kontext des Netzaktivismus, das Internet, einigermaßen genau in den Blick zu bekommen. Ursprünglich war also vorgesehen, zur Einleitung die militärgeschichtliche Herkunft des Internet zu klären und zu bewerten, um dann, anhand der Genese des Internet zu einem Massenmedium unter dem Stichpunkt "Gegenwärtige Nutzungsformen des Internet" der Frage nach der Faszination an diesem neuen Medium nachzugehen. Herausfinden wollte ich dabei, ob die Nutzer sich vornehmlich aktiv oder passiv an dem neuen Medium beteiligen und ob die "Aktivitäten", um die es schließlich in meiner Arbeit geht, überhaupt eine Relevanz für den Großteil der Internetnutzer hat. Dies stellte sich aber als schwieriger heraus, als erwartet. So kann auch in dieser Arbeit wenig darüber gesagt werden, warum denn das Internet in der Gegenwart eine so wichtige Rolle zu spielen scheint. An gegebener Stelle wird noch darauf zurückzukommen sein.
Die Dokumente enthalten jeweils die gleiche Arbeit, allerdings in drei unterschiedlichen Varianten, die sich in der Qualität der Bilder und damit in der Filegröße unterscheiden: * Bilder in voller Druckqualität (8,2 MB): DissWFOM1.pdf (Dokument1) * Photos in reduzierter Auflösung (3,1 MB): DissWFOM2.pdf (Dokument2) * Photos und Zeichnungen in red. Auflösung (1,4 MB): DissWFOM3.pdf (Dokument3)
Cold target recoil ion momentum spectroscopy (COLTRIM) has been employed to image the momentum distributions of continuum electrons liberated in the impact of slow He2+ on He and H2. The distributions were measured for fully determined motion of the nuclei that is as a function of the impact parameter and in a well de ned scattering plane The single ionization (SI) of H2 leading to H2+ recoil ions in nondissociative states (He2+ + H+ -> He2+ + H+ + e-) and the transfer ionization (TI) of H2 leading to H2 dissociation into two free protons (He2+ H2 -> He+ + H+ + H+ + e-) were investigated. Similar measurements have been carried out for He target, the corresponding atomic two electron system, i.e. the single ionization (SI) (He2+ + He -> He+ + He2+ e- and the transfer ionization (TI) (He2+ + He -> He+ + He2+ + e-). These measurements have been exploited to understand the results obtained for H target. In comparing the continuum electron momentum distributions for H2 with that for He, a high degree of similarity is observed. In the case of transfer ionization of H2, the electron momentum distributions generated for parallel and perpendicular molecular orientations revealed no orientation dependence. The in scattering plane electron momentum distributions for the transfer ionization of H2 by He2+ and for the transfer ionization of He by He2e showed that the salient feature of these distributions for both collisions systems consists in the appearance of two groups of electrons with difeerent structures. In addition to the group of the saddle electrons forming two jets separated by a valley along the projectile axis we nd a new group of electrons moving with a velocity higher than the projectile velocity These new fast forward electrons result from a narrow range of impact parameters and appear as image saddle in the projectile frame. In contrast to the transfer ionization of He, the fast forward electrons group disappears in the in scattering plane electron momentum distribution generated for the single ionization of He. Instead of this group another new group of electrons appear These electrons exhibit an amount of backscattering These backward elec trons appear as image saddle in the target frame The structures that the saddle electrons show are owing to the quasi molecular nature of the collision process For the TI of H2, the TI of He and the SI of He, a pi-orbital shape of the electron momentum distribution is observed This indicates the importance of the rotational coupling 2-p-theta -> 2p-pi in the initial promotion of the ground state followed by further promotions to the continuum The backward electrons as well as the fast forward electrons are not discussed in the theoretical literature at all. However, a number of obvious indications of the existence of the backward and fast forward electrons could be seen in the experimental works of Abdallah et al. as well as in the theoretical calculations of Sidky et al One might speculate that electrons which are promoted on the saddle for some time during the collision could finally swing around the He+ ion in the way out of the collision, i.e. either around the projectile in the forward direction as in the TI case forming the fast forward electrons or around the recoil ion in the backward direction as in the SI case forming the backward electrons. This might be a result of the strong gradient, and hence the large acceleration of the screened He+ potential.
The cytochrome bc1 complex is a cornerstone in bioenergetic electron transfer chains, where it carries out tasks as diverse as respiration, photosynthesis, and nitrogen fixation. This homodimeric multisubunit membrane protein has been studied extensively for several decades and the enzyme mechanism is described with the modified protonmotive Q cycle. Still, the molecular and kinetic description of the catalytic cycle is not complete and questions remain regarding the bifurcation of electron transfer at the quinol oxidation (Qo) site, substrate occupancy, pathways of proton conduction, and the nature of the Rieske protein domain movement. We used competitive inhibitors to study the molecular architecture at the Qo site with X-ray crystallography. The structure of the enzyme with the substrate analog 5-n-heptyl-6-hydroxy-4,7-dioxobenzothiazole (HHDBT) bound at the Qo site was determined at 2.5 Å resolution. Spectroscopic studies showed that HHDBT is negatively charged when bound at the active site. Mechanistic interpretations from inhibitor binding are in line with single occupancy model for quinol oxidation and structural analysis supports the proposed proton transfer pathway. For functional insight into the enzyme mechanism, redox-sensitive protonation changes were studied by Fourier transform infrared spectroscopy. The protein purification procedure was optimized for less delipidation and the isolated enzyme was more active. Furthermore, two new phospholipids were identified in the X-ray structures, including a cardiolipin. Strikingly, conserved lipid binding cavities were observed in structural comparison with homologous enzymes. The functional role of tightly bound phospholipids will be discussed. Finally, the Qo site is a target for various compounds of agricultural and pharmaceutical importance. Importantly, the X-ray structures permit detailed analysis of the molecular reasons for acquired resistance to and treatment failure of Qo site inhibitors, such as atovaquone, that is used to treat malaria and pneumonia, as discussed herein.
The cytochrome bc1 complex or ubiquinol:cytochrome c oxidoreductase (QCR) catalyses electron transfer from ubiquinol to cytochrome c in respiration and photosynthesis coupled to a vectorial proton transport across the membrane, in which the enzyme resides. In both bacteria and eukaryotic organisms, QCR participates in supramolecular assembly of membrane proteins that comprise the respiratory or photosynthetic chain. In the present work, proton transfer pathways, substrate binding and the supramolecular assembly of the respiratory chain in yeast were probed by structure-based site-directed mutagenesis and characterization of the variants. Both active sites centre P, the place of quinol oxidation, and centre N, where quinone reduction takes place, lack direct access to the bulk solvent necessary for proton release and uptake. Based on the X-ray structure, proton transfer pathways were postulated. Analysis at centre P showed, that E272 and Y132 of cytochrome b are important for QCR catalysis as indicated by increased superoxide production and lowered Cyc1p reductase activity in these variants. Pre-steady state heme reduction kinetics in combination with stigmatellin resistance indicated that charge and length of the side chain at position 272 are crucial for efficient docking of the ISP to form the enzyme substrate complex and for electron bifurcation at centre P. Variants of Y312 and F129, both residues of cytochrome b, showed an increased Km indicating participation of these residues in coordination of ubiquinol or the possible intermediate semiquinone anion radical. F129 proved to be crucial for a functional Q-cycle as indicated by respiratory negative growth phenotype and a lowered H+/e- stoichiometry of F129 variants. At centre N, the postulated CL/K and E/R proton transfer pathways are located at opposite sites of the bound ubiquinone. Variants in the surface residues R218 (cytochrome b) and E52 (Qcr7) of the E/R pathway and E82 (Qcr7) of the CL/K pathway showed instability upon purification indicating an important role of these residues for QCR integrity. The slowed down centre N reduction kinetics in H85 (CL/K), R218 and N208 (both E/R) variant was attributed to a destabilised semiquinone anion consistent with the observed decreased sensitivity towards the site-specific inhibitor antimycin and an increased Km. Variants of residues of both pathway, E82Q and R218M, exhibited a decreased H+/e- stoichiometry indicating a crucial role of both residue for maintaining a working Q-cycle and supporting the proposed protonation of the substrate via the Cl/K and the E/R pathway. Long-range interaction between centre N and centre P were observed by altered reduction kinetics of the high potential chain and increased superoxide production in the centre N variants. The role of the cation-pi-interaction between F230 of Cyt1p and R19 of cytochrome c in binding of the redox carrier to QCR was analysed. In F230L hydrophobic interaction were partially lost as was deduced from the ionic strength dependence of Cyc1p reductase activity and Cycp1 binding, as detected by ionic strength sensitive Kd and Km for Cyc1p. The decreased enzymatic rate of F230W could be explained by a disturbed binding of Cyc1p to the variant enzyme. F230 may influence the heme mid point potential and thereby the electron transfer rate to Cyc1p. Reduction of Cobp via both centre P and centre N was disturbed suggesting an interaction between high and low potential chain. Supramolecular association between QCR and cytochrome c oxidase (COX) in yeast mitochondria was probed by affinity chromatography of a his-tagged QCR in the presence of the mild detergent digitonin. In comparison to purification with laurylmaltoside, the presence of both QCR and COX subunits was detected in the elution fractions by SDS-PAGE, Cyc1p reductase and TMPD oxidase activity assays and immunoblot analysis. The CL-dependent formation of the supercomplex between QCR and COX was analysed by replacement variants in the CL-binding site of QCR in CL containing and CL free environment. With an increasing number of replacements of the three lysines the CL-binding pocket supercomplex formation was not abolished, when CL is present as shown by BN-PAGE analysis. This was supported by the synergetic decrease in enzyme activity for both enzymes upon increased number of replacements. In the CL-free environment, no supracomplex formation was observed for a wildtype CL binding site. By replacements of two lysines in the CL-binding pocket, supercomplex formation could be recovered as revealed by BN-PAGE. This indicates, that CL may serve as a charge neutralizer for the lysines near the presumed interaction domain between complex III and complex IV. The obtained results for centre P provide new information of residues critical for stabilisation of ubiquinol and controlling electron short circuit reactions. The observations for centre N variants clearly support the proposed two proton transfer pathways and the role of the bound phospholipids in centre N kinetics. Variants in the Cyc1p binding site suggest a role for F230 both in Cyc1p binding and electron transfer. Clear interaction between the high and low potential chain in both Cyt1p and centre N variants strongly support long-range interactions in the complex. Studies on the supramolecular association of complex III and complex IV indicate a new role of Cl in stabilising a supracomplex.
Charakterisierung der alternativen NADH-Ubichinon-Oxidoreduktase (NDH2) aus Yarrowia lipolytica
(2004)
Neben dem protonenpumpenden Komplex I (NDH-1) der Atmungskette besitzt die obligat aerobe Hefe Yarrowia lipolytica eine alternative NADH:Ubichinon Oxidoreduktase (NDH-2). Diese Enzyme, die in den Atmungsketten von Pflanzen, Pilzen und Bakterien vorkommen, bestehen aus nur einer Untereinheit, führen jedoch dieselbe Reaktion aus wie Komplex I, nämlich die Elektronenübertragung von NADH auf Ubichinon, wobei allerdings keine Protonen über die Membran transloziert werden. Nur peripher mit der Membran assoziiert, können alternative Dehydrogenasen entweder zur cytosolischen Seite (extern) oder zur Matrixseite (intern) orientiert sein. Y. lipolytica besitzt im Gegensatz zu anderen Ascomyceten nur eine einzige extern orientierte alternative Dehydrogenase mit einer vorhergesagten Masse von ca. 60 kD und einem nicht kovalent gebundenem Molekül FAD als Cofaktor. Durch Fusion des Leserasters mit der Präsequenz der 75 kD Untereinheit von Komplex I war die interne Expression des Enzyms (NDH2i) gelungen, die das Überleben von Komplex I Deletionsmutanten ermöglichte. Im Rahmen der vorliegenden Arbeit wurde die alternative Dehydrogenase von Y. lipolytica innerhalb ihrer natürlichen Membranumgebung charakterisiert. Das Enzym reagierte mit verschiedenen Chinonanaloga, wobei mit dem hydrophilen Q1 eine höhere katalytische Rate erzielt wurde als mit DBQ, das dem natürlich vorkommenden Q9 am ähnlichsten ist. Da hydrophobe Substrate fast ausschließlich in der Lipidphase der Membranen gelöst vorliegen, musste bei der Bestimmung von kinetischen Parametern (ebenso wie bei Komplex I) auf eine gleichbleibend große Membranphase im Messvolumen geachtet werden. Mit dem standardmäßig benutzten Substrat DBQ reagierte YLNDH2 nach einem Ping-Pong Reaktionsmechanismus. Dieser beschreibt eine abwechselnde Bindung der beiden Substrate, wobei das Enzym die Elektronen von NADH aufnimmt (E-FADH2) und an Ubichinon weitergibt (E-FAD); es existiert kein ternärer Enzym-Substrat Komplex. Gestützt durch Kristallstrukturen des analogen Enzyms QR1 mit NADPH bzw. mit Durochinon, liegt die Vermutung nahe, dass beide Substrate in ähnlicher Weise und sehr wahrscheinlich in der gleichen Bindungstasche binden. Ein Ping-Pong Reaktionsmechanismus wurde bereits für die NADH:DCPIP Oxidoreduktase Aktivität von zwei weiteren alternativen Enzymen postuliert, jedoch noch nie für ein physiologisches Substrat. Als bislang wirksamster Inhibitor für alternative Dehydrogenasen wurde 1-hydroxy-2-dodecyl-4(1H)chinolon (HDQ) entdeckt. HDQ hemmte NDH2 in Membranen aus Y. lipolytica mit einer I50 von 200 nM, was der 500fachen Hemmwirkung des gängig verwendeten Flavon auf das isolierte Enzym NDI1 von S. cerevisiae entspricht. Allerdings hemmte HDQ auch Komplex I mit einer I50 von 2 µM, ähnlich wie es bei Platanetin in Pflanzenmitochondrien der Fall war [Roberts et al., 1996]. Mit dem Ziel, ein polyklonales Antiserum gegen die native YLNDH2 zu generieren, wurde das Enzym in E. coli heterolog exprimiert. Die Expression führte zur Bildung von Einschlusskörpern, aus denen das rekombinante Enzym unter denaturierenden Bedingungen gereinigt und zur Immunisierung eines Kaninchens verwendet werden konnte. Das Antiserum kreuzreagierte mit der nativen und der internen Version von YLNDH2 und zeigte nur wenige unspezifische Bindungen. Es wurde gezeigt, dass Y. lipolytica Stämme ohne NDH2 und Komplex I mit NDH2i als einziger Dehydrogenase erzeugt werden konnten. In N. crassa war der Versuch, NDE2 und Komplex I gleichzeitig zu deletieren, gescheitert, was zu der Schlussfolgerung führte, dass sich die beiden Enzyme in diesem Organismus möglicherweise kompensieren könnten. Dies war in Y. lipolytica ausgeschlossen worden, da wahrscheinlich kein (oder nur unzureichender) Austausch zwischen matrixständigem und cytosolischem NADH stattfindet. Die zielgerichtete Mutagenese hochkonservierter Bereiche im offenen Leserahmen von YLNDH2 lieferte das eindeutige Ergebnis, dass die zweite der beiden beta-alpha-beta-Bindungsdomänen NADH binden muß, da sich der KM Wert für NADH bei Mutation des essentiellen sauren Restes E320 dratisch erhöhte. Alle Mutationen, die die Dinukleotid Bindungsdomäne I betrafen, die danach folgerichtig den Cofaktor FAD binden muß, führten zu einem vollständigen Verlust von NDH2. Dieselbe Zuordnung der Bindungsstellen war bereits von Björklöf et al. [2000] vorgeschlagen worden. Eine Chinonbindungstelle konnte durch Mutagenese der beiden apolar/aromatischen Bereiche der Sequenz nicht identifiziert werden. Die Modifikation des C-Terminus von NDH2i führte zu nicht mehr messbarer Aktivität und stark verringerter Expression des Enzyms in mitochondrialen Membranen (siehe Anhang 7.5.1.1). Es kann daher vermutet werden, dass der C-Terminus für die korrekte Faltung eine wichtige Rolle spielt, möglicherweise sogar bei der Membranassoziation, wie bei Rasmusson [1999] vorgeschlagen wurde. Interessant war in diesem Zusammenhang, dass die C-terminal modifizierte NDH2i trotzdem das Überleben von Komplex I Deletionsmutanten bzw. das Wachstum auf DQA ermöglichte. In Membranen, die einen unterschiedlichen Gehalt an NDH2, jedoch die gleiche Gesamtmenge an Protein enthielten, wurde eine unerwartete lineare Abhängigkeit zwischen KM und Vmax Werten beobachtet. Dieses Phänomen wurde mit dem Modell der externen Diffusionslimitierung beschrieben, die in ähnlicher Weise auch bei immobilisierten Enzymen auftritt. Danach wird die Geschwindigkeit der enzymatischen Reaktion von YLNDH2 sowohl durch die kinetisch kontrollierte Rate, als auch durch die Transportrate des Ubichinons bestimmt, das aus der Membran heraus in die wässrige Umgebung des katalytischen Zentrums gelangen muss. Aus diesem Grund ist nicht nur die Maximalgeschwindigkeit, sondern auch die Michaelis Menten Konstante KM abhängig vom Gehalt des Enzyms in Membranen. Dies führte bei niedrig exprimierten mutanten Enzymen zur gleichzeitigen Abnahme von KM und Vmax. Eine externe Diffusionskontrolle der enzymatischen Reaktion wurde auch für Komplex I, dessen Reaktionszentrum im peripheren Arm vermutet werden kann, aber nicht für Komplex III aus S. cerevisiae, dessen Chinonbindungsstellen sich definitiv in hydrophober Umgebung befinden, beobachtet.