Untangling the Sequence of Events During the S2 → S3 Transition in Photosystem II and Implications for the Water Oxidation Mechanism

Total Page:16

File Type:pdf, Size:1020Kb

Untangling the Sequence of Events During the S2 → S3 Transition in Photosystem II and Implications for the Water Oxidation Mechanism Untangling the sequence of events during the S2 → S3 transition in photosystem II and implications for the water oxidation mechanism Mohamed Ibrahima,1, Thomas Franssonb,1, Ruchira Chatterjeec,1, Mun Hon Cheahd,1, Rana Husseina, Louise Lassallec, Kyle D. Sutherlinc, Iris D. Youngc,2, Franklin D. Fullere, Sheraz Gulc, In-Sik Kimc, Philipp S. Simonc, Casper de Lichtenbergd,f, Petko Chernevd, Isabel Bogaczc, Cindy C. Phamc, Allen M. Orvilleg,h, Nicholas Saicheki, Trent Northeni, Alexander Batyuke, Sergio Carbajoe, Roberto Alonso-Morie, Kensuke Tonoj,k, Shigeki Owadaj,k, Asmit Bhowmickc, Robert Bolotovskyc, Derek Mendezc, Nigel W. Moriartyc, James M. Holtonc,l,m, Holger Dobbeka, Aaron S. Brewsterc, Paul D. Adamsc,n, Nicholas K. Sauterc, Uwe Bergmanno, Athina Zounia,3, Johannes Messingerd,f,3, Jan Kernc, Vittal K. Yachandrac,3, and Junko Yanoc,3 aInstitut für Biologie, Humboldt-Universität zu Berlin, D-10115 Berlin, Germany; bInterdisciplinary Center for Scientific Computing, University of Heidelberg, 69120 Heidelberg, Germany; cMolecular Biophysics and Integrated Bioimaging Division, Lawrence Berkeley National Laboratory, Berkeley, CA 94720; dDepartment of Chemistry - Ångström, Molecular Biomimetics, Uppsala University, SE 75120 Uppsala, Sweden; eLinac Coherent Light Source, SLAC National Accelerator Laboratory, Menlo Park, CA 94025; fInstitutionen för Kemi, Kemiskt Biologiskt Centrum, Umeå Universitet, SE 90187 Umeå, Sweden; gDiamond Light Source Ltd, Harwell Science and Innovation Campus, OX11 0DE Didcot, United Kingdom; hResearch Complex at Harwell, Rutherford Appleton Laboratory, OX11 0FA Didcot, United Kingdom; iEnvironmental Genomics and Systems Biology Division, Lawrence Berkeley National Laboratory, Berkeley, CA 94720; jJapan Synchrotron Radiation Research Institute, Sayo-cho, Sayo-gun, 679-5198 Hyogo, Japan; kRIKEN SPring-8 Center, Sayo-cho, Sayo-gun, 679-5148 Hyogo, Japan; lStanford Synchrotron Radiation Lightsource, SLAC National Accelerator Laboratory, Menlo Park, CA 94025; mDepartment of Biochemistry and Biophysics, University of California, San Francisco, CA 94158; nDepartment of Bioengineering, University of California, Berkeley, CA 94720; and oStanford PULSE Institute, SLAC National Accelerator Laboratory, Menlo Park, CA 94025 Edited by Pierre Joliot, Institut de Biologie Physico-Chimique, Paris, France, and approved April 1, 2020 (received for review January 11, 2020) In oxygenic photosynthesis, light-driven oxidation of water to mo- chlorophyll a moiety P680 (most likely consisting of the four exci- lecular oxygen is carried out by the oxygen-evolving complex tonically coupled chlorophylls PD1,PD2,ChlD1,andChlD2)andthe BIOPHYSICS AND COMPUTATIONAL BIOLOGY (OEC) in photosystem II (PS II). Recently, we reported the room- neighboring pheophytin molecules PheoD1/D2 as well as the two temperature structures of PS II in the four (semi)stable S-states, S1, plastoquinones QA and QB. The one-electron photochemistry at S2,S3, and S0, showing that a water molecule is inserted during the → S2 S3 transition, as a new bridging O(H)-ligand between Mn1 Significance and Ca. To understand the sequence of events leading to the for- mation of this last stable intermediate state before O formation, 2 A new bridging oxygen ligand is incorporated between one of we recorded diffraction and Mn X-ray emission spectroscopy (XES) the Mn atoms and Ca in the Mn Ca cluster during the transition data at several time points during the S → S transition. At the 4 2 3 from the one-photon induced S intermediate state to the two- electron acceptor site, changes due to the two-electron redox 2 photon induced S state in the catalytic water oxidation re- chemistry at the quinones, Q and Q , are observed. At the donor 3 A B action in photosystem II. However, the sequence of events leading site, tyrosine Y and His190 H-bonded to it move by 50 μs after the Z to this change is not known. Here we report an X-ray crystallog- second flash, and Glu189 moves away from Ca. This is followed by raphy and spectroscopy study at room temperature using an X-ray Mn1 and Mn4 moving apart, and the insertion of O (H) at the X free electron laser to collect a “molecular movie” of the structural open coordination site of Mn1. This water, possibly a ligand of and oxidation state change steps leading to the insertion of this Ca, could be supplied via a “water wheel”-like arrangement of five new oxygen bridge, in the 50 μs to 200 ms time scales after waters next to the OEC that is connected by a large channel to the photon absorption, which triggers the S → S state transition. bulk solvent. XES spectra show that Mn oxidation (τ of ∼350 μs) 2 3 → during the S2 S3 transition mirrors the appearance of OX elec- Author contributions: U.B., A.Z., J.M., J.K., V.K.Y., and J.Y. designed research; M.I., T.F., tron density. This indicates that the oxidation state change and the R.C., M.H.C., R.H., L.L., K.D.S., I.D.Y., F.D.F., S.G., I.-S.K., P.S.S., C.d.L., P.C., I.B., C.C.P., insertion of water as a bridging atom between Mn1 and Ca are A.M.O., N.S., T.N., A. Batyuk, S.C., R.A.-M., K.T., S.O., A.B., U.B., A.Z., J.M., J.K., V.K.Y., highly correlated. and J.Y. performed research; T.F., R.C., M.H.C., R.H., N.S., T.N., J.M.H., A.S.B., P.D.A., N.K.S., A.Z., and J.M. contributed new reagents/analytic tools; M.I., T.F., R.C., M.H.C., L.L., K.D.S., A. Bhowmick, R.B., D.M., N.W.M., J.M.H., H.D., A.S.B., P.D.A., N.K.S., J.K., and J.Y. analyzed photosynthesis | photosystem II | water oxidation | data; and M.I., T.F., R.C., U.B., A.Z., J.M., J.K., V.K.Y., and J.Y. wrote the paper. oxygen-evolving complex | X-ray free electron laser The authors declare no competing interest. This article is a PNAS Direct Submission. ioxygen, which supports all aerobic life, is abundant in the This open access article is distributed under Creative Commons Attribution-NonCommercial- Datmosphere because of its constant regeneration via photo- NoDerivatives License 4.0 (CC BY-NC-ND). synthetic water oxidation in plants, algae, and cyanobacteria. The Data deposition: X-ray diffraction datasets and associated models have been deposited in water-splitting chemistry occurs in the oxygen-evolving complex the Research Collaboratory for Structural Bioinformatics (RCSB) Protein Data Bank (https://www.rcsb.org) under PDB ID codes 6W1O for the 0F, 6W1P for the 1F, 6W1Q (OEC) of photosystem II (PS II; Fig. 1A), which contains a het- for the 2F(50 μs), 6W1R for the 2F(150 μs), 6W1T for the 2F(250 μs), 6W1U for the eronuclear, oxo-bridged Mn4Ca cluster that catalyzes the reaction: 2F(400 μs), and 6W1V for the 2F(200 ms) data. 1M.I., T.F., R.C., and M.H.C. contributed equally to this work. → + − + + [1] 2 H2O O2 4 e 4 H . 2Present address: Department of Bioengineering and Therapeutic Sciences, University of California, San Francisco, CA 94158. To allow this reaction to take place, the OEC accumulates four 3To whom correspondence may be addressed. Email: [email protected], oxidizing equivalents, stepping through intermediates that are [email protected], [email protected], or [email protected]. referred to as the S-states (Si,i= 0 through 4) (Fig. 1B) (1, 2). This article contains supporting information online at https://www.pnas.org/lookup/suppl/ These oxidation reactions are driven by the light-induced charge doi:10.1073/pnas.2000529117/-/DCSupplemental. separations in the reaction center of PS II that comprises the www.pnas.org/cgi/doi/10.1073/pnas.2000529117 PNAS Latest Articles | 1of12 Downloaded by guest on September 26, 2021 ABC Acceptor site Cytoplasm YZ - •− •− + Q Q E189 2H e A B Q Q H PQ A B 2 QB W4 e- S QB Q 4F QBH2 2 A W3 •− QAQB •+ PQH2 P680 P680 Ca Q Q + Dark State 1F A B W2 1 PheoD1 Yz• Yz 1 e- Mn4(III2,IV2) 5 Mn4(III3,IV) P680 PheoD2 0 1 •+ PD2 PD1 O2 S S Yz H+ e- •− 1 4 2 2 QA QB W1 3 e- + 3 ChlD2 Donor site • Yz P680 4 ChlD1 H+ H2O P +H2O 680 TyrT Z •− S4 H+ H2O S2 Mn4(III,IV3) + - Lumen QAQB - (H , e ) •+ H+ Yz YZ OEC e- Yz e- 2F E189 4 photons 3 Yz P680 + - •+ S W4 2HO O +4H +4e •+ + W3 2 2 P680 Yz Mn4(IV4) P680• e- S3 - X Q•−Q e P680 Ca A B Q•−Q•− A B W2 1 3F 5 1 QAQBH2 Q Q A B Q W1 4 2 B 2 3 QBH2 3 4 Fig. 1. The S2 → S3 transition in PS II. (A) The structure of PS II with the membrane-embedded helices and the large membrane extrinsic regions on the luminal side of PS II are shown as cartoon in gray and in color the main electron transfer components involved in charge transfer (P680, Pheo) and stabilization with the acceptor quinones, QA and QB, toward the cytoplasm, and YZ and OEC with catalytic Mn4Ca cluster on the donor side at the interface between the membrane and the lumen-bound part of PS II. (B) Kok cycle of the water oxidation reaction that is triggered by the absorption of photons (nanosecond light flashes, 1F to 4F) in the antenna and the reaction center P680. The white and gray areas show the redox changes of the OEC (S1 to S0) and YZ together with the charge separation reactions, while the outer blue circle shows the concomitant acceptor side chemistry of the quinones.
Recommended publications
  • Rail Transport in the World's Major Cities
    Feature Evolution of Urban Railways (part 2) Rail Transport in The World’s Major Cities Takao Okamoto and Norihisa Tadakoshi Many of the world’s large cities grew in development, particularly with regard to American city; and Hong Kong, Seoul and conjunction with railways, and today, the following: Tokyo representing Asian cities. large cities cannot depend only on motor • The correlation between railways and vehicles for transportation. With worries urban growth over global environmental issues, public • The location of terminals for intercity London transportation systems are increasingly and intra-city transport seen as an important way to expand and • Examination of various public transit Located in southeast England near the revitalize large cities, while consuming systems, including non-infrastructure mouth of the River Thames, London less energy and other resources. • Strategic planning of rail networks expanded during the Industrial Revolution This article looks at public transportation based on urban development trends (1760–1850) and secured its dominance systems in some major cities of the world and future models as the heart of the British Empire during and identifies similarities and differences The cities selected for this comparative the Victorian era (1837–1901). It’s in areas such as history of development, study are: London as the first city to adapt population grew from about 500,000 in railway networks, and method of rail technology to public transport; Paris, the 17th century to 4.5 million by the end operation. Our aim is to explore the future Berlin and Moscow as three major of the 19th century. The modern relationship between urban and transport European cities; New York as a North- metropolis of Greater London consists of The London Railway Network King’s Cross/St.Pancras Paddington LCY Victoria Waterloo LHR R.
    [Show full text]
  • CAFR Includes the Dishiet's Basic Financial Statement Prepared in Accordance with Governrnental Acc'ounting Standards Board Statement 34
    SCHOOL DISTRICT OF BERLIN BOROUGH Berlin Borough Board of Education Berlin, New Jersey Comprehensive Annual Financial Report For the Fiscal Year Ended June 30, 2018 Comprehensive Annual Financial Report of the Berlin Borough Board of Education Berlin, New Jersey For the Fiscal Year Ended June 30, 2018 Prepared by Berlin Borough Board of Education Finance Department BERLIN BOROUGH SCHOOL DISTRICT INTRODUCTORY SECTION Page Letter of Transmittal 2 Organizational Chart 7 Roster of Officials 8 Consultants and Advisors 9 FINANCIAL SECTION Independent Auditor's Report 11 K-1 Report on Compliance and on Internal Control Over Financial Reporting Based on an Audit of Financial Statements Performed in Accordance with Government Auditing Standards 14 Required Supplementary Information - Part I Management's Discussion and Analysis 17 Basic Financial Statements A. District-wide Financial Statements: A-1 Statement of Net Position 27 A-2 Statement of Activities 28 B. Fund Financial Statements: Governmental Funds: B-1 Balance Sheet 30 B-2 Statement of Revenues, Expenditures, and Changes in Fund Balances 31 B-3 Reconciliation of the Statement of Revenues, Expenditures, and Changes in Fund Balances of Governmental Funds to the Statement of Activities 32 Proprietary Funds: B-4 Statement of Net Position 33 B-5 Statement of Revenues, Expenses, and Changes in Fund Net Position 34 B-6 Statement of Cash Flows 35 Fiduciary Funds: B-7 Statement of Fiduciary Net Position 36 B-8 Statement of Changes in Fiduciary Net Position 37 Notes to the Financial Statements 38 Page Required Supplementary Information - Part II C. Budgetary Comparison Schedules C-1 Budgetary Comparison Schedule - General Fund 72 C-1a Combining Schedule of Revenues, Expenditures and Changes in Fund Balance - Budget and Actual (if applicable) N/A C-2 Budgetary Comparison Schedule - Special Revenue Fund 78 Notes to the Required Supplementary Information C-3 Budget-to-GAAP Reconciliation 79 Required Supplementary Information - Part III L.
    [Show full text]
  • Stow Police Department Dispatch Log From: 01/01/2018 Thru: 01/07/2018 0000 - 2359
    Stow Police Department Dispatch Log From: 01/01/2018 Thru: 01/07/2018 0000 - 2359 For Date: 01/01/2018 - Monday Time Call Reason Action 0032 MOTOR VEHICLE STOP Citation/Warning Issued Location/Address: GREAT RD + COMMON RD Narrative: CBK on scene - S4 transporting operator and passenger to lobby 0106 MOTOR VEHICLE STOP VERBAL WARNING Location/Address: POMPOSITTICUT ST 0125 SUSPICIOUS ACTIVITY Services Rendered Location/Address: BOXBORO RD Narrative: Resident concerned about an individual in brown jacket and some type of bag in hand walking down street Narrative: S25 transporting individual to Bolton - making his way to Clinton - Bolton PD notified 0704 ALARM, BURGLAR Investigated Location/Address: KIRKLAND RD Narrative: ADT reporting residential panic alarm no answer when they attempted key holder. Narrative: s9 requesting fire no one coming to the door. s6 cancelling fire person at the door. Narrative: s6 reports all set, hearing problem she has trouble hearing phone or door. 0857 Town Beach Check AREA CHECKED Location/Address: [STW BEACH] TOWN BEACH - SUDBURY RD 0913 THREATS Investigated Location/Address: NORTH SHORE DR Narrative: Caller requesting police to respond to home for a domestic situation. Her daughter is receiving threatening texts from her boyfriend. Boyfriend not at home but may be in the area. s6 and s9 responding. Narrative: Attempting to obtain RO, Clinton PD given info on subject as his phone was pinged in Clinton. Narrative: RO granted. BOLO sent. 1249 ASSIST CITIZEN Services Rendered Location/Address: [STW 305GREAT] STOW POLICE DEPARTMENT - GREAT RD Narrative: Caller from Maynard requesting to talk with officer regarding what constitutes elder abuse.
    [Show full text]
  • Berlin Metro Map by Zuti
    Hohen Mühlenbeck Bernau Borgsdorf Neuendorf Bergfelde Schönfließ Mönchmühle Karow Röntgental Friedenstal Oranienburg Bernau Lehnitz Birkenwerder Hugenotten Navarrapl Buch Zepernick Guyotstr bei Bernau Rosenthal Nord Arnoux HAVEL Französisch Hauptstr Buchholz Kirche Frohnau Friedrich Engels 50 HAVEL Wiesenwinkel Blankenfelder Berlin Angerweg © Copyright Visual IT Ltd Nordendstr Rosenthaler ® Zuti and the Zuti logo are registered trademarks Hermsdorf www.zuti.co.uk Nordend Schillerstr Marienstr BERLIN WALL BERLIN Uhlandstr Pasewalker Blankenburg Hennigsdorf Waldemar Waidmannslust Pasewalker Platanenstr Heinrich Böll Blankenburger Weg Heiligensee Pankower Am Iderfenngraben Kuckhoffstr Pastor Niemöller Platz Schulzendorf Galenusstr Wittenau Hermann Hesse Grabbeallee Waldstr Pastor Niemöller Ahrensfelde REINICKENDORF Ahrensfelde Tschaikowskistr HAVEL Rathaus Würtzstr Wartenberg Reinickendorf Mendelstr Tegel Wilhelmsruh M1 Pankow Zingster Falkenberger Karl B Heinersdorf Prendener Welsestr Nerven Bürgerpark Stiftsweg Heinersdorf Falkenberg Barnimplatz Alt Tegel klinik Alt Reinickendorf Pankow Rathaus Zingster Ribnitzer Schönholz Pankow PANKOW Hohenschönhausen Eichborn Ahrenshooper Niemegker Borsigwerke damm Pankow Rothenbachstr Paracelsus Bad Kirche Prerower U8 Mehrower Holzhauser Lindauer Hansastr Malchower Wuhletalstr HAVEL Heinersdorf Kirche Otisstr Allee Wollankstr JUNGFERNHEIDE Residenzstr Pankow Feldtmannstr Rüdickenstr Max Hermann TEGELER SEE Am Wasserturm M5 Scharnweber Masurenstr M2 Pasedagplatz Berliner Allee Franz Neumann Am Steinberg
    [Show full text]
  • S25 (Gültig Ab 15.12.2019) S25
    S Teltow Stadt — S+U Friedrichstr. Bhf — S Hennigsdorf Bhf > S25 (gültig ab 15.12.2019) S25 S-Bahn Berlin GmbH Alle Züge 2. Klasse und f (Tarif des Verkehrsverbundes Berlin-Brandenburg [VBB]) montags bis freitags, nicht an Feiertagen Verkehrshinweise S Teltow Stadt ab 0 05 0 25 0 45 4 05 F20 23 45 S Lichterfelde Süd 0 08 0 28 0 48 4 08 23 48 S Osdorfer Str. 0 10 0 30 0 50 4 10 23 50 S Lichterfelde Ost Bhf 0 12 0 32 0 52 4 12 23 52 S Lankwitz 0 14 0 34 0 54 4 14 23 54 S Südende 0 16 0 36 0 56 4 16 23 56 S Priesterweg 0 19 0 39 0 59 4 19 23 59 S Südkreuz Bhf 0 21 0 41 1 01 4 21 0 01 S+U Yorckstr. S2 S25 S26 U7 0 24 0 44 1 04 4 24 0 04 S Anhalter Bahnhof 0 27 0 47 1 07 4 07 4 27 0 07 S+U Potsdamer Platz Bhf 0 29 0 49 1 09 4 09 4 29 0 09 S+U Brandenburger Tor 0 31 0 51 4 11 4 31 0 11 S+U Friedrichstr. Bhf O 0 33 0 53 4 13 4 33 0 13 S+U Friedrichstr. Bhf ab 0 33 0 53 4 13 4 33 0 13 S Oranienburger Str. 0 35 0 55 4 15 4 35 0 15 S Nordbahnhof 0 37 0 57 4 17 4 37 0 17 S Humboldthain 0 40 1 00 4 20 4 40 0 20 S+U Gesundbrunnen Bhf O 0 41 1 01 4 21 4 41 0 21 S+U Gesundbrunnen Bhf ab 0 42 1 02 4 22 4 42 0 22 S Bornholmer Str.
    [Show full text]
  • Activation Energies for Two Steps in the S2 → S3 Transition of Photosynthetic Water Oxidation from Time-Resolved Single-Frequency Infrared Spectroscopy
    Activation energies for two steps in the S2 S3 transition of photosynthetic water oxidation from time-resolved single- frequency infrared spectroscopy Cite as: J. Chem. Phys. 153, 215101 (2020); https://doi.org/10.1063/5.0027995 Submitted: 01 September 2020 . Accepted: 05 November 2020 . Published Online: 01 December 2020 Sarah M. Mäusle, Aiganym Abzaliyeva, Paul Greife, Philipp S. Simon, Rebeca Perez, Yvonne Zilliges, and Holger Dau COLLECTIONS Paper published as part of the special topic on 65 Years of Electron Transfer ARTICLES YOU MAY BE INTERESTED IN Reflections on electron transfer theory The Journal of Chemical Physics 153, 210401 (2020); https://doi.org/10.1063/5.0035434 Combining the best of two worlds: Stimulated Raman excited fluorescence The Journal of Chemical Physics 153, 210901 (2020); https://doi.org/10.1063/5.0030204 Removing artifacts in polarizable embedding calculations of one- and two-photon absorption spectra of fluorescent proteins The Journal of Chemical Physics 153, 215102 (2020); https://doi.org/10.1063/5.0023434 J. Chem. Phys. 153, 215101 (2020); https://doi.org/10.1063/5.0027995 153, 215101 © 2020 Author(s). The Journal ARTICLE of Chemical Physics scitation.org/journal/jcp Activation energies for two steps in the S2 → S3 transition of photosynthetic water oxidation from time-resolved single-frequency infrared spectroscopy Cite as: J. Chem. Phys. 153, 215101 (2020); doi: 10.1063/5.0027995 Submitted: 1 September 2020 • Accepted: 5 November 2020 • Published Online: 1 December 2020 Sarah M. Mäusle, Aiganym Abzaliyeva, Paul Greife, Philipp S. Simon, Rebeca Perez, Yvonne Zilliges, and Holger Daua) AFFILIATIONS Department of Physics, Freie Universität Berlin, Arnimallee 14, 14195 Berlin, Germany Note: This paper is part of the JCP Special Topic on 65 Years of Electron Transfer.
    [Show full text]
  • World Para Swimming World Records Long Course
    World Para Swimming World Records Long Course World Para Swimming World Records created by IPC Sport Data Management System Gender: Men | Course: Long Course Men's 50 m Freestyle Class Name NPC Birth Time Date City Country S1 Mamistvalov, Itzhak ISR 1979 01:03.80 2014-03-09 Esbjerg Denmark S2 Zou, Liankang CHN 1995 00:50.65 2016-09-11 Rio de Janeiro Brazil S3 Huang, Wenpan CHN 1995 00:38.81 2017-12-06 Mexico City Mexico S4 Leslie, Cameron NZL 1990 00:37.14 2019-09-13 London Great Britain S5 Fantin, Antonio ITA 2001 00:30.16 2019-06-01 Lignano Sabbiadoro Italy S6 Xu, Qing CHN 1992 00:28.57 2012-09-04 London Great Britain S7 Trusov, Andrii UKR 2000 00:27.07 2019-09-15 London Great Britain S8 Tarasov, Denis RUS 1993 00:25.32 2014-08-10 Eindhoven Netherlands S9 Barlaam, Simone ITA 2000 00:24.00 2019-09-15 London Great Britain S10 Brasil, Andre BRA 1984 00:23.16 2012-08-31 London Great Britain S11 Yang, Bozun CHN 1986 00:25.27 2012-09-01 London Great Britain S12 Veraksa, Maksym UKR 1984 00:22.99 2009-10-21 Reykjavik Iceland S13 Boki, Ihar BLR 1994 00:23.20 2015-07-13 Glasgow Great Britain IPC Sport Data Management System Page 1 of 22 29 September 2021 at 00:11:44 CEST World Para Swimming World Records Long Course Men's 100 m Freestyle Class Name NPC Birth Time Date City Country S1 Mamistvalov, Itzhak ISR 1979 02:15.83 2012-09-01 London Great Britain S2 Liu, Benying CHN 1996 01:46.63 2016-09-11 Rio de Janeiro Brazil S3 Lopez Diaz, Diego MEX 1994 01:32.69 2018-06-08 Berlin Germany S4 Dadaon, Ami Omer ISR 2000 01:19.77 2021-05-17 Funchal Portugal
    [Show full text]
  • Fahrplan-S25.Pdf
    S Teltow Stadt — S+U Friedrichstr. Bhf — S Hennigsdorf Bhf > S25 (gültig ab 13.12.2020) S25 S-Bahn Berlin GmbH Alle Züge 2. Klasse und f (Tarif des Verkehrsverbundes Berlin-Brandenburg [VBB]) montags bis freitags, nicht an Feiertagen Verkehrshinweise S Teltow Stadt ab 0 05 0 25 0 45 4 05 F20 23 45 S Lichterfelde Süd 0 08 0 28 0 48 4 08 23 48 S Osdorfer Str. 0 10 0 30 0 50 4 10 23 50 S Lichterfelde Ost Bhf 0 12 0 32 0 52 4 12 23 52 S Lankwitz 0 14 0 34 0 54 4 14 23 54 S Südende 0 16 0 36 0 56 4 16 23 56 S Priesterweg 0 19 0 39 0 59 4 19 23 59 S Südkreuz Bhf 0 21 0 41 1 01 4 21 0 01 S+U Yorckstr. S2 S25 S26 U7 0 24 0 44 1 04 4 24 0 04 S Anhalter Bahnhof 0 27 0 47 1 07 4 07 4 27 0 07 S+U Potsdamer Platz Bhf 0 29 0 49 1 09 4 09 4 29 0 09 S+U Brandenburger Tor 0 31 0 51 4 11 4 31 0 11 S+U Friedrichstr. Bhf O 0 33 0 53 4 13 4 33 0 13 S+U Friedrichstr. Bhf ab 0 33 0 53 4 13 4 33 0 13 S Oranienburger Str. 0 35 0 55 4 15 4 35 0 15 S Nordbahnhof 0 37 0 57 4 17 4 37 0 17 S Humboldthain 0 40 1 00 4 20 4 40 0 20 S+U Gesundbrunnen Bhf O 0 41 1 01 4 21 4 41 0 21 S+U Gesundbrunnen Bhf ab 0 42 1 02 4 22 4 42 0 22 S Bornholmer Str.
    [Show full text]
  • Train Simulator - Right Through Berlin
    Train Simulator - Right through Berlin © Historical S-Bahn Berlin e.V. 2020 Page 1 Train Simulator - Right through Berlin Manual "Mitten durch Berlin" - Operation Series 476 Table of contents Table of contents ....................................................................................................... 2 0. authors ................................................................................................................ 3 1. introduction .......................................................................................................... 3 2. system requirements .............................................................................................. 4 3. the driver's cab ..................................................................................................... 5 4. upgrading .......................................................................................................... 10 4.1 Train is cold (full upgrade) .............................................................................. 10 4.2 Transfer of trains from other Tf ......................................................................... 10 4.3 Train is fully upgraded .................................................................................... 10 4.4 Change of driver's cab ................................................................................... 11 5. door control ....................................................................................................... 11 6. driving and braking ...........................................................................................
    [Show full text]
  • Verkehrliche Voruntersuchung Und Standardisierte Bewertung Für Die Wiederinbetriebnahme Der Potsdamer Stammbahn
    Stammbahnsteig auf dem S-Bf Zehlendorf Bf Düppel, Bahnsteigkante Verkehrliche Voruntersuchung und Standardisierte Bewertung für die Wiederinbetriebnahme der Potsdamer Stammbahn Untersuchungsergebnisse Februar 2008 INTRAPLAN CONSULT GMBH • Orleansplatz 5a • 81667 München • Tel. 089 / 45 91 10 Fax 089 / 447 05 93 • e-mail: [email protected] Auftraggeber: Auftragnehmer: Senatsverwaltung für Stadtentwicklung Berlin Intraplan Consult GmbH Am Kölnischen Park 3 Orleansplatz 5a 10179 Berlin 81667 München Ministerium für Infrastruktur und Raumordnung des Landes Brandenburg Henning-von-Tresckow-Straße 2-8 14467 Potsdam Inhaltsverzeichnis 1 Aufgabenstellung und Zielsetzung 1 2 Projektstruktur 2 3 Grundlagen 5 3.1 Beschreibung des Investitionsvorhabens 5 3.2 Abgrenzung und Strukturierung des Untersuchungsraumes 10 3.3 Abbildung des Istzustandes 12 3.3.1 MIV-/ÖPNV-Angebot Istzustand 12 3.3.2 MIV-/ÖPNV-Nachfrage Istzustand 15 3.3.3 Umlegungsergebnisse ÖPNV Istzustand 19 4 Mengengerüst Ohnefall 21 4.1 Absehbare Strukturentwicklung 21 4.2 MIV-Maßnahmen 22 4.3 ÖPNV-Angebot Ohnefall 26 4.3.1 Regionalverkehrskonzept im Ohnefall 26 4.3.2 Betriebskonzept S-Bahn im Ohnefall 30 4.3.3 Bedienungskonzepte für die Betriebszweige U-Bahn und Straßen- bahn im Ohnefall 35 4.4 MIV-/ÖPNV-Nachfrage Ohnefall 35 4.4.1 Verflechtungsmatrix MIV/ÖPNV für den normalwerktäglichen Regel- verkehr 35 4.4.2 Verflechtungsmatrizen Ohnefall für den flughafenbezogenen Verkehr 36 4.4.3 Nahverkehrsrelevante Teilwege von SPFV-Kunden 38 4.5 Umlegung ÖPNV und Dimensionierungsnachweise 40
    [Show full text]
  • Berlin 2018 World Para Swimming World Series PI Classification Schedule Version 1.2
    Berlin 2018 World Para Swimming World Series PI Classification Schedule Version 1.2 Athletes: Must present to the Classification 15 minutes before the allocated time on the classification schedule. Must bring a passport or another official photo form of identification to classification. Will be required to read and sign a classification consent form prior to presenting to the classification panel. Must be accompanied by one support staff from their team. Must bring their own interpreter if they do not speak English. 4-Jun-18 Time SDMS ID NPC Family Name Given Name S SB SM Status 9:00 29994 TUR Inci Omer Onur S4 New 9:00 19482 ISR Gordiychuk Yuliya S9 SB8 SM9 Review 9:00 4201 BLR Shavel Natalia S5 SB4 SM5 Review 9:00 38962 ESP Cortes Melgar Jaume S5 SB4 SM5 New 10:30 33328 TUR Dogan Ismail Can S5 SB4 SM5 Review 10:30 23573 KAZ Yeldekenov Yerlan S9 SB8 New 10:30 22473 ISR Malyar Ariel S5 SB4 SM5 Review 10:30 35895 CZE Fryba Petr S8 New 12:15 15071 ESP Rolo Marichal Judit S7 SB7 SM7 Review 12:15 5338 AUT Onea Andreas S8 SB8 SM8 Review 12:15 26226 BLR Shchalkanau Yahor S10 SB9 SM10 Review Lunch Break 13:15 14:15 35154 KAZ Pronkina Viktoriya S7 New 14:15 19472 ESP Rodriguez Martinez Ines Rocio S4 SB3 SM3 New 14:15 22955 CZE Andrysek Petr S6 SB5 SM6 Review 14:15 26225 BLR Musarau Artsiom S7 SB6 SM7 Review 16:00 11498 EST Tilk Brenda S7 N/A N/A Review 16:00 20050 EST Topkin Matz S5 SB4 SM5 Review 16:00 17934 CZE Duskova Vendula S8 SB7 SM8 Review 16:00 22511 BLR Svadkouskaya Aliaksandra S10 SB9 SM10 Review 17:30 5942 ESP Gascon Sarai S9 SB9 SM9
    [Show full text]
  • Adding Transit to an Agent-Based Transportation Simulation: Concepts and Implementation
    Adding Transit to an Agent-Based Transportation Simulation Concepts and Implementation vorgelegt von Dipl. Inf.-Ing. ETH Marcel Rieser aus Affeltrangen (TG), Schweiz von der Fakultät V, Verkehrs- und Maschinensysteme der Technischen Universität Berlin zur Erlangung des akademischen Grades eines Doktor-Ingenieurs (Dr.-Ing.) genehmigte Dissertation Promotionsausschuss: Vorsitzende: Prof. Dr.-Ing. Chr. Ahrend Gutachter: Prof. Dr. rer. nat. Kai Nagel Gutachter: Prof. Dr.-Ing. Kay W. Axhausen Tag der wissenschaftlichen Aussprache: 10. Juni 2010 Berlin 2010 D 83 Contents Abstract ix Kurzfassung xi 1 Introduction 1 2 Public Transportation Systems 5 2.1 Definitions ............................... 5 2.2 Public Transportation in Developed Countries ................. 6 2.2.1 Overview ............................. 6 2.2.2 Data Requirements ........................ 7 2.3 Public Transportation in Developing Countries ................ 9 2.3.1 Overview ............................. 9 2.3.2 Data Requirements ........................ 9 3 Related Work 11 3.1 Transportation Simulation in General .................... 11 3.2 Multi-Modal and Transit Simulation ..................... 13 3.2.1 Transit Assignment ........................ 13 3.2.2 Multimodal Route Choice ...................... 14 3.2.3 Operational Simulation ....................... 15 3.2.4 Multi-Modal Simulation ....................... 16 4 Agent-Based Transportation Simulation 19 4.1 Overview ................................ 19 4.2 Controler ................................ 22 4.3 Initial Demand
    [Show full text]