Overall energy conversion efficiency of a photosynthetic vesicle

Sener, Melih; Strumpfer, Johan; Singharoy, Abhishek; Hunter, C. Neil; Schulten, Klaus

doi:10.7554/eLife.09541

Title: Overall energy conversion efficiency of a photosynthetic vesicle

Abstract

The chromatophore of purple bacteria is an intracellular spherical vesicle that exists in numerous copies in the cell and that efficiently converts sunlight into ATP synthesis, operating typically under low light conditions. Building on an atomic-level structural model of a low-light-adapted chromatophore vesicle from Rhodobacter sphaeroides, we investigate the cooperation between more than a hundred protein complexes in the vesicle. The steady-state ATP production rate as a function of incident light intensity is determined after identifying quinol turnover at the cytochrome $b c_{1}$ complex (cyt $b c_{1}$ ) as rate limiting and assuming that the quinone/quinol pool of about 900 molecules acts in a quasi-stationary state. For an illumination condition equivalent to 1% of full sunlight, the vesicle exhibits an ATP production rate of 82 ATP molecules/s. The energy conversion efficiency of ATP synthesis at illuminations corresponding to 1%–5% of full sunlight is calculated to be 0.12–0.04, respectively. The vesicle stoichiometry, evolutionarily adapted to the low light intensities in the habitat of purple bacteria, is suboptimal for steady-state ATP turnover for the benefit of protection against over-illumination.

Authors:

Sener, Melih ^[1]; Strumpfer, Johan ^[2]; Singharoy, Abhishek ^[3]; Hunter, C. Neil ^[4];

^[5]

Beckman Institute for Advanced Science and Technology, University of Illinois at Urbana-Champaign, Urbana, United States, Department of Physics, University of Illinois at Urbana-Champaign, Urbana, United States
Beckman Institute for Advanced Science and Technology, University of Illinois at Urbana-Champaign, Urbana, United States, Center for Biophysics and Computational Biology, University of Illinois at Urbana-Champaign, Urbana, United States
Beckman Institute for Advanced Science and Technology, University of Illinois at Urbana-Champaign, Urbana, United States
Department of Molecular Biology and Biotechnology, University of Sheffield, Sheffield, United Kingdom
Beckman Institute for Advanced Science and Technology, University of Illinois at Urbana-Champaign, Urbana, United States, Department of Physics, University of Illinois at Urbana-Champaign, Urbana, United States, Center for Biophysics and Computational Biology, University of Illinois at Urbana-Champaign, Urbana, United States

Publication Date:: Fri Aug 26 00:00:00 EDT 2016

Research Org.:: Oak Ridge National Laboratory (ORNL), Oak Ridge, TN (United States). Oak Ridge Leadership Computing Facility (OLCF); Energy Frontier Research Centers (EFRC) (United States). Photosynthetic Antenna Research Center (PARC)

Sponsoring Org.:: USDOE Office of Science (SC), Basic Energy Sciences (BES); National Science foundation (NSF); National Institutes of Health (NIH); European Research Council (ERC)

OSTI Identifier:: 1307791

Alternate Identifier(s):: OSTI ID: 1307792; OSTI ID: 1388788

Grant/Contract Number:: SC0001035; AC05-00OR22725; BB/M000265/1

Resource Type:: Published Article

Journal Name:: eLife

Additional Journal Information:: Journal Name: eLife Journal Volume: 5; Journal ID: ISSN 2050-084X

Publisher:: eLife Sciences Publications, Ltd.

Country of Publication:: United States

Language:: English

Subject:: 10 SYNTHETIC FUELS; 14 SOLAR ENERGY; solar (fuels); photosynthesis (natural and artificial); biofuels (including algae and biomass); bio-inspired; charge transport; membrane; synthesis (novel materials); synthesis (self-assembly)

Citation Formats


                    Sener, Melih, Strumpfer, Johan, Singharoy, Abhishek, Hunter, C. Neil, and Schulten, Klaus. Overall energy conversion efficiency of a photosynthetic vesicle.  United States: N. p., 2016. 
Web.  doi:10.7554/eLife.09541.

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                    Sener, Melih, Strumpfer, Johan, Singharoy, Abhishek, Hunter, C. Neil, & Schulten, Klaus. Overall energy conversion efficiency of a photosynthetic vesicle.  United States.  https://doi.org/10.7554/eLife.09541

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                    Sener, Melih, Strumpfer, Johan, Singharoy, Abhishek, Hunter, C. Neil, and Schulten, Klaus. Fri .  
"Overall energy conversion efficiency of a photosynthetic vesicle".  United States.  https://doi.org/10.7554/eLife.09541.

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@article{osti_1307791,

  title        = {Overall energy conversion efficiency of a photosynthetic vesicle},

  author       = {Sener, Melih and Strumpfer, Johan and Singharoy, Abhishek and Hunter, C. Neil and Schulten, Klaus},

  abstractNote = {The chromatophore of purple bacteria is an intracellular spherical vesicle that exists in numerous copies in the cell and that efficiently converts sunlight into ATP synthesis, operating typically under low light conditions. Building on an atomic-level structural model of a low-light-adapted chromatophore vesicle from Rhodobacter sphaeroides, we investigate the cooperation between more than a hundred protein complexes in the vesicle. The steady-state ATP production rate as a function of incident light intensity is determined after identifying quinol turnover at the cytochrome b c 1 complex (cyt b ⁢ c 1 ) as rate limiting and assuming that the quinone/quinol pool of about 900 molecules acts in a quasi-stationary state. For an illumination condition equivalent to 1% of full sunlight, the vesicle exhibits an ATP production rate of 82 ATP molecules/s. The energy conversion efficiency of ATP synthesis at illuminations corresponding to 1%–5% of full sunlight is calculated to be 0.12–0.04, respectively. The vesicle stoichiometry, evolutionarily adapted to the low light intensities in the habitat of purple bacteria, is suboptimal for steady-state ATP turnover for the benefit of protection against over-illumination.},

  doi          = {10.7554/eLife.09541},

  journal      = {eLife},

  number       = ,

  volume       = 5,

  place        = {United States},

  year         = {Fri Aug 26 00:00:00 EDT 2016},

  month        = {Fri Aug 26 00:00:00 EDT 2016}

}

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Title: Overall energy conversion efficiency of a photosynthetic vesicle

Abstract

Citation Formats

Chromatophore Vesicles of Rhodobacter capsulatus Contain on Average One FOF1-ATP Synthase Each journal, March 2002

Molecular Architecture of the Thylakoid Membrane: Lipid Diffusion Space for Plastoquinone journal, April 2002

Atomic force microscopy of the bacterial photosynthetic apparatus: plain pictures of an elaborate machinery journal, March 2009

The theory of Forster-type migration between clusters of strongly interacting molecules: application to light-harvesting complexes of purple bacteria journal, November 1996

Adapting the Förster Theory of Energy Transfer for Modeling Dynamics in Aggregated Molecular Assemblies journal, March 2001

Supramolecular organization of the photosynthetic apparatus of Rhodobacter sphaeroides journal, February 1999

Adaptation of intracytoplasmic membranes to altered light intensity in Rhodobacter sphaeroides journal, September 2012

Turnover Number of Escherichia coli F0F1 ATP Synthase for ATP Synthesis in Membrane Vesicles journal, January 1997

Photosynthetic membrane development in Rhodopseudomonas sphaeroides. Spectral and kinetic characterization of redox components of light-driven electron flow in apparent photosynthetic membrane growth initiation sites. journal, March 1985

Theory on Rates of Excitation-Energy Transfer between Molecular Aggregates through Distributed Transition Dipoles with Application to the Antenna System in Bacterial Photosynthesis journal, January 1999

Theoretical examination of quantum coherence in a photosynthetic system at physiological temperature journal, October 2009

The optical cross section and absolute size of a photosynthetic unit journal, January 1986

Energy transfer in the inhomogeneously broadened core antenna of purple bacteria: a simultaneous fit of low-intensity picosecond absorption and fluorescence kinetics journal, January 1994

Atomic-level structural and functional model of a bacterial photosynthetic membrane vesicle journal, September 2007

The Accumulation of the Light-Harvesting 2 Complex during Remodeling of the Rhodobacter sphaeroides Intracytoplasmic Membrane Results in a Slowing of the Electron Transfer Turnover Rate of Photochemical Reaction Centers journal, June 2011

Relationship of proton motive force and the F0F1-ATPase with bio-hydrogen production activity of Rhodobacter sphaeroides: effects of diphenylene iodonium, hydrogenase inhibitor, and its solvent dimethylsulphoxide journal, June 2012

Robustness and Optimality of Light Harvesting in Cyanobacterial Photosystem I journal, August 2002

Photosynthetic Excitons book, January 2000

Proton Translocation and ATP Synthesis by the FoF1-ATPase of Purple Bacteria book, January 2009

Three-Dimensional Structure of the Rhodobacter sphaeroides RC-LH1-PufX Complex: Dimerization and Quinone Channels Promoted by PufX journal, October 2013

Trapping Kinetics in Mutants of the Photosynthetic Purple Bacterium Rhodobacter sphaeroides: Influence of the Charge Separation Rate and Consequences for the Rate-Limiting Step in the Light-Harvesting Process journal, March 1994

The effect of different levels of the B800-850 light-harvesting complex on intracytoplasmic membrane development in Rhodobacter sphaeroides journal, April 1996

Spectral Dynamics of Individual Bacterial Light-Harvesting Complexes: Alternative Disorder Model journal, February 2008

The effect of correlated bath fluctuations on exciton transfer journal, March 2011

Membrane invagination in Rhodobacter sphaeroides is initiated at curved regions of the cytoplasmic membrane, then forms both budded and fully detached spherical vesicles: Membrane biogenesis in Rba. sphaeroides journal, April 2010

Energy transfer dynamics in an RC–LH1–PufX tubular photosynthetic membrane journal, August 2010

Projection structure of the photosynthetic reaction centre-antenna complex of Rhodospirillum rubrum at 8.5 A resolution journal, August 2002

Transfer of Energy from Bacteriochlorophyll to a Reaction Centre During Bacterial Photosynthesis journal, January 1963

Pigment Organization and Transfer of Electronic Excitation in the Photosynthetic Unit of Purple Bacteria journal, May 1997

The Light-Harvesting Apparatus in Purple Photosynthetic Bacteria: Introduction to a Quantum Biological Device book, August 2011

Identification of the First Steps in Charge Separation in Bacterial Photosynthetic Reaction Centers of Rhodobacter sphaeroides by Ultrafast Mid-Infrared Spectroscopy: Electron Transfer and Protein Dynamics journal, August 2008

Photosynthetic Vesicle Architecture and Constraints on Efficient Energy Harvesting journal, July 2010

Architecture and mechanism of the light-harvesting apparatus of purple bacteria journal, May 1998

Crystal structure of an integral membrane light-harvesting complex from photosynthetic bacteria journal, April 1995

Energy Transfer and Exciton Annihilation in the B800−850 Antenna Complex of the Photosynthetic Purple Bacterium Rhodopseudomonas acidophila (Strain 10050). A Femtosecond Transient Absorption Study journal, February 1997

Molecular Evidence for the Early Evolution of Photosynthesis journal, September 2000

Functional Consequences of the Organization of the Photosynthetic Apparatus in Rhodobacter sphaeroides: I. QUINONE DOMAINS AND EXCITATION TRANSFER IN CHROMATOPHORES AND REACTION CENTER{middle dot}ANTENNA COMPLEXES journal, January 2005

Structural and Functional Proteomics of Intracytoplasmic Membrane Assembly in <b><i>Rhodobacter sphaeroides</i></b> journal, January 2013

Excitonic polarons in quasi-one-dimensional LH1 and LH2 bacteriochlorophyll a antenna aggregates from photosynthetic bacteria: A wavelength-dependent selective spectroscopy study journal, February 2009

From high-resolution AFM topographs to atomic models of supramolecular assemblies journal, August 2007

The relation between membrane ionic current and ATP synthesis in chromatophores from Rhodopseudomonas capsulata journal, June 1983

A Spatial Model of the Chromatophore Vesicles of Rhodobacter sphaeroides and the Position of the Cytochrome bc1 Complex journal, August 2006

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Protein Shape and Crowding Drive Domain Formation and Curvature in Biological Membranes journal, January 2008

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Förster Energy Transfer Theory as Reflected in the Structures of Photosynthetic Light‐Harvesting Systems journal, February 2011

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Bacteriochlorophyll and Carotenoid Excitonic Couplings in the LH2 System of Purple Bacteria journal, October 2000

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General random matrix approach to account for the effect of static disorder on the spectral properties of light harvesting systems journal, March 2002

VMD: Visual molecular dynamics journal, February 1996

Membrane development in purple photosynthetic bacteria in response to alterations in light intensity and oxygen tension journal, May 2013

Electronic Excitations in Aggregates of Bacteriochlorophylls journal, September 1998

Membrane Curvature Induced by Aggregates of LH2s and Monomeric LH1s journal, December 2009

Ultrafast transient absorption spectroscopy: principles and application to photosynthetic systems journal, July 2009

Integration of energy and electron transfer processes in the photosynthetic membrane of Rhodobacter sphaeroides journal, October 2014

Natural photosystems from an engineer’s perspective: length, time, and energy scales of charge and energy transfer journal, October 2007

The architecture and function of the light-harvesting apparatus of purple bacteria: from single molecules to in vivo membranes journal, August 2006

The long-range organization of a native photosynthetic membrane journal, December 2004

Energy transfer between carotenoids and bacteriochlorophylls in light-harvesting complex II of purple bacteria journal, March 1999

Trapping, loss and annihilation of excitations in a photosynthetic system. I. Theoretical aspects journal, December 1983

Molecular stochastic simulations of chromatophore vesicles from Rhodobacter sphaeroides journal, April 2007

Open Quantum Dynamics Calculations with the Hierarchy Equations of Motion on Parallel Computers journal, July 2012

Unified treatment of quantum coherent and incoherent hopping dynamics in electronic energy transfer: Reduced hierarchy equation approach journal, June 2009

Chlorophyll Excitations in Photosystem I of Synechococcus elongatus journal, October 2002

Quantitative proteomic analysis of intracytoplasmic membrane development in Rhodobacter sphaeroides: Proteomics of membrane assembly in Rba. sphaeroides journal, May 2012

The 8.5Å Projection Structure of the Core RC–LH1–PufX Dimer of Rhodobacter sphaeroides journal, June 2005

Energy transfer and trapping in photosynthesis journal, August 1994

Three-Pulse Photon Echo Measurements on LH1 and LH2 Complexes of Rhodobacter sphaeroides : A Nonlinear Spectroscopic Probe of Energy Transfer journal, September 1997

Self-Assembly of Photosynthetic Membranes journal, April 2010

DOT2: Macromolecular docking with improved biophysical models journal, May 2013

Excitons and excitation transfer in the photosynthetic unit of purple bacteria journal, February 1998

Three-dimensional Reconstruction of a Membrane-bending Complex journal, March 2008

Crystal structure of the entire respiratory complex I journal, February 2013

Chromatophore Vesicles of Rhodobacter capsulatus Contain on Average One FOF1-ATP Synthase Each
journal, March 2002

Molecular Architecture of the Thylakoid Membrane: Lipid Diffusion Space for Plastoquinone
journal, April 2002

Atomic force microscopy of the bacterial photosynthetic apparatus: plain pictures of an elaborate machinery
journal, March 2009

The theory of Forster-type migration between clusters of strongly interacting molecules: application to light-harvesting complexes of purple bacteria
journal, November 1996

Adapting the Förster Theory of Energy Transfer for Modeling Dynamics in Aggregated Molecular Assemblies
journal, March 2001

Supramolecular organization of the photosynthetic apparatus of Rhodobacter sphaeroides
journal, February 1999

Adaptation of intracytoplasmic membranes to altered light intensity in Rhodobacter sphaeroides
journal, September 2012

Turnover Number of Escherichia coli F0F1 ATP Synthase for ATP Synthesis in Membrane Vesicles
journal, January 1997

Photosynthetic membrane development in Rhodopseudomonas sphaeroides. Spectral and kinetic characterization of redox components of light-driven electron flow in apparent photosynthetic membrane growth initiation sites.
journal, March 1985

Theory on Rates of Excitation-Energy Transfer between Molecular Aggregates through Distributed Transition Dipoles with Application to the Antenna System in Bacterial Photosynthesis
journal, January 1999

Theoretical examination of quantum coherence in a photosynthetic system at physiological temperature
journal, October 2009

The optical cross section and absolute size of a photosynthetic unit
journal, January 1986

Energy transfer in the inhomogeneously broadened core antenna of purple bacteria: a simultaneous fit of low-intensity picosecond absorption and fluorescence kinetics
journal, January 1994

Atomic-level structural and functional model of a bacterial photosynthetic membrane vesicle
journal, September 2007

The Accumulation of the Light-Harvesting 2 Complex during Remodeling of the Rhodobacter sphaeroides Intracytoplasmic Membrane Results in a Slowing of the Electron Transfer Turnover Rate of Photochemical Reaction Centers
journal, June 2011

Relationship of proton motive force and the F0F1-ATPase with bio-hydrogen production activity of Rhodobacter sphaeroides: effects of diphenylene iodonium, hydrogenase inhibitor, and its solvent dimethylsulphoxide
journal, June 2012

Robustness and Optimality of Light Harvesting in Cyanobacterial Photosystem I
journal, August 2002

Photosynthetic Excitons
book, January 2000

Proton Translocation and ATP Synthesis by the F_oF₁-ATPase of Purple Bacteria
book, January 2009

Three-Dimensional Structure of the Rhodobacter sphaeroides RC-LH1-PufX Complex: Dimerization and Quinone Channels Promoted by PufX
journal, October 2013

Trapping Kinetics in Mutants of the Photosynthetic Purple Bacterium Rhodobacter sphaeroides: Influence of the Charge Separation Rate and Consequences for the Rate-Limiting Step in the Light-Harvesting Process
journal, March 1994

The effect of different levels of the B800-850 light-harvesting complex on intracytoplasmic membrane development in Rhodobacter sphaeroides
journal, April 1996

Spectral Dynamics of Individual Bacterial Light-Harvesting Complexes: Alternative Disorder Model
journal, February 2008

The effect of correlated bath fluctuations on exciton transfer
journal, March 2011

Membrane invagination in Rhodobacter sphaeroides is initiated at curved regions of the cytoplasmic membrane, then forms both budded and fully detached spherical vesicles: Membrane biogenesis in Rba. sphaeroides
journal, April 2010

Energy transfer dynamics in an RC–LH1–PufX tubular photosynthetic membrane
journal, August 2010

Projection structure of the photosynthetic reaction centre-antenna complex of Rhodospirillum rubrum at 8.5 A resolution
journal, August 2002

Transfer of Energy from Bacteriochlorophyll to a Reaction Centre During Bacterial Photosynthesis
journal, January 1963

Pigment Organization and Transfer of Electronic Excitation in the Photosynthetic Unit of Purple Bacteria
journal, May 1997

The Light-Harvesting Apparatus in Purple Photosynthetic Bacteria: Introduction to a Quantum Biological Device
book, August 2011

Identification of the First Steps in Charge Separation in Bacterial Photosynthetic Reaction Centers of Rhodobacter sphaeroides by Ultrafast Mid-Infrared Spectroscopy: Electron Transfer and Protein Dynamics
journal, August 2008

Photosynthetic Vesicle Architecture and Constraints on Efficient Energy Harvesting
journal, July 2010

Architecture and mechanism of the light-harvesting apparatus of purple bacteria
journal, May 1998

Crystal structure of an integral membrane light-harvesting complex from photosynthetic bacteria
journal, April 1995

Energy Transfer and Exciton Annihilation in the B800−850 Antenna Complex of the Photosynthetic Purple Bacterium Rhodopseudomonas acidophila (Strain 10050). A Femtosecond Transient Absorption Study
journal, February 1997

Molecular Evidence for the Early Evolution of Photosynthesis
journal, September 2000

Functional Consequences of the Organization of the Photosynthetic Apparatus in Rhodobacter sphaeroides: I. QUINONE DOMAINS AND EXCITATION TRANSFER IN CHROMATOPHORES AND REACTION CENTER{middle dot}ANTENNA COMPLEXES
journal, January 2005

Structural and Functional Proteomics of Intracytoplasmic Membrane Assembly in <b><i>Rhodobacter sphaeroides</i></b>
journal, January 2013

Excitonic polarons in quasi-one-dimensional LH1 and LH2 bacteriochlorophyll a antenna aggregates from photosynthetic bacteria: A wavelength-dependent selective spectroscopy study
journal, February 2009

From high-resolution AFM topographs to atomic models of supramolecular assemblies
journal, August 2007

The relation between membrane ionic current and ATP synthesis in chromatophores from Rhodopseudomonas capsulata
journal, June 1983

A Spatial Model of the Chromatophore Vesicles of Rhodobacter sphaeroides and the Position of the Cytochrome bc1 Complex
journal, August 2006

Comparison of Retinal-Based and Chlorophyll-Based Photosynthesis: A Biothermokinetic Description of Photochemical Reaction Centers
book, January 1993

Protein Shape and Crowding Drive Domain Formation and Curvature in Biological Membranes
journal, January 2008

The native architecture of a photosynthetic membrane
journal, August 2004

Protein-Induced Membrane Curvature Investigated through Molecular Dynamics Flexible Fitting
journal, July 2009

Interplay between excitation kinetics and reaction-center dynamics in purple bacteria
journal, September 2010

The Photosynthetic Apparatus of Rhodopseudomonas palustris: Structures and Organization
journal, April 2006

Förster Energy Transfer Theory as Reflected in the Structures of Photosynthetic Light‐Harvesting Systems
journal, February 2011

Modeling the electron transport chain of purple non‐sulfur bacteria
journal, January 2008

Bacteriochlorophyll and Carotenoid Excitonic Couplings in the LH2 System of Purple Bacteria
journal, October 2000

The effects of protein crowding in bacterial photosynthetic membranes on the flow of quinone redox species between the photochemical reaction center and the ubiquinol-cytochrome c2 oxidoreductase
journal, January 2011

General random matrix approach to account for the effect of static disorder on the spectral properties of light harvesting systems
journal, March 2002

VMD: Visual molecular dynamics
journal, February 1996

Membrane development in purple photosynthetic bacteria in response to alterations in light intensity and oxygen tension
journal, May 2013

Electronic Excitations in Aggregates of Bacteriochlorophylls
journal, September 1998

Membrane Curvature Induced by Aggregates of LH2s and Monomeric LH1s
journal, December 2009

Ultrafast transient absorption spectroscopy: principles and application to photosynthetic systems
journal, July 2009

Integration of energy and electron transfer processes in the photosynthetic membrane of Rhodobacter sphaeroides
journal, October 2014

Natural photosystems from an engineer’s perspective: length, time, and energy scales of charge and energy transfer
journal, October 2007

The architecture and function of the light-harvesting apparatus of purple bacteria: from single molecules to in vivo membranes
journal, August 2006

The long-range organization of a native photosynthetic membrane
journal, December 2004

Energy transfer between carotenoids and bacteriochlorophylls in light-harvesting complex II of purple bacteria
journal, March 1999

Trapping, loss and annihilation of excitations in a photosynthetic system. I. Theoretical aspects
journal, December 1983

Molecular stochastic simulations of chromatophore vesicles from Rhodobacter sphaeroides
journal, April 2007

Open Quantum Dynamics Calculations with the Hierarchy Equations of Motion on Parallel Computers
journal, July 2012

Unified treatment of quantum coherent and incoherent hopping dynamics in electronic energy transfer: Reduced hierarchy equation approach
journal, June 2009

Chlorophyll Excitations in Photosystem I of Synechococcus elongatus
journal, October 2002

Quantitative proteomic analysis of intracytoplasmic membrane development in Rhodobacter sphaeroides: Proteomics of membrane assembly in Rba. sphaeroides
journal, May 2012

The 8.5Å Projection Structure of the Core RC–LH1–PufX Dimer of Rhodobacter sphaeroides
journal, June 2005

Energy transfer and trapping in photosynthesis
journal, August 1994

Three-Pulse Photon Echo Measurements on LH1 and LH2 Complexes of Rhodobacter sphaeroides : A Nonlinear Spectroscopic Probe of Energy Transfer
journal, September 1997

Self-Assembly of Photosynthetic Membranes
journal, April 2010

DOT2: Macromolecular docking with improved biophysical models
journal, May 2013

Excitons and excitation transfer in the photosynthetic unit of purple bacteria
journal, February 1998

Three-dimensional Reconstruction of a Membrane-bending Complex
journal, March 2008

Crystal structure of the entire respiratory complex I
journal, February 2013

A Model for Photoreceptor-Based Magnetoreception in Birds
journal, February 2000

Excitation energy trapping by the reaction center ofRhodobacter Sphaeroides
journal, January 2000

Temporally and spectrally resolved subpicosecond energy transfer within the peripheral antenna complex (LH2) and from LH2 to the core antenna complex in photosynthetic purple bacteria.
journal, December 1995