Coupling between information processing and redox chemistry.

Cellular co-location of energy transduction with genome function

John F. AllenSchool of Biological and Chemical Sciences, Queen Mary University of London

and Research Department of Genetics, Evolution and Environment, University College London

Coupling between information processing and redox chemistry.

Cellular co-location of energy transduction with genome function

John F. AllenSchool of Biological and Chemical Sciences, Queen Mary University of London

and Research Department of Genetics, Evolution and Environment, University College London

Horizontal Gene Transfer and the Last Universal Common Ancestor •Horizontal Gene Transfer and the Last Universal Common Ancestor •

The Open University • 5th and 6th September 2013The Open University • 5th and 6th September 2013

11

http://en.wikipedia.org/wiki/Geologic_time_scale

LUCALUCA here seems to mean: here seems to mean:

The Last Common Ancestor of all things alive today...The Last Common Ancestor of all things alive today...

...therefore Horizontal Gene Transfer could have occurred between LUCA ...therefore Horizontal Gene Transfer could have occurred between LUCA and things now no longer aliveand things now no longer alive

What do we mean by What do we mean by LUCALUCA...?...?

A. The Last Common Ancestor of all things alive A. The Last Common Ancestor of all things alive today (on Earth)...today (on Earth)...

B. The Last Common Ancestor of all things that B. The Last Common Ancestor of all things that have ever livedhave ever lived (on Earth)... (on Earth)...

This tree diagram, used to show the divergence of species, is the only illustration in the Origin of Species.

http://en.wikipedia.org/wiki/On_the_Origin_of_Species

LUCALUCA here must mean: here must mean:

B. The Last Common Ancestor B. The Last Common Ancestor of all things that of all things that have ever have ever

lived...lived...

...in which case... ...in which case...

• • With what did LUCA engage in With what did LUCA engage in Horizontal Gene Transfer...?Horizontal Gene Transfer...?

• • Can any sensible distinction be made Can any sensible distinction be made between LUCA’s Horizontal and Vertical between LUCA’s Horizontal and Vertical

Gene Transfer...?Gene Transfer...?

• • Can Horizontal Gene Transfer have Can Horizontal Gene Transfer have played any part in the Origin of Life?played any part in the Origin of Life?

Electron transport – chains of coupled oxidation Electron transport – chains of coupled oxidation and reduction reactionsand reduction reactions

Coupling between information Coupling between information processing and redox chemistry.processing and redox chemistry.

Homeostasis...Homeostasis...

...the maintenance of a constant internal ...the maintenance of a constant internal environment despite changes in the external environment despite changes in the external

environment. environment.

Coupling between information Coupling between information processing and redox chemistry.processing and redox chemistry.

http://en.wikipedia.org/wiki/Feedback

A block diagram for feedback. The feedback is negative if B < 0

Coupling between information Coupling between information processing and redox chemistry.processing and redox chemistry.

Redox Homeostasis...Redox Homeostasis...

...the maintenance of a constant electrochemical potential ...the maintenance of a constant electrochemical potential and ionic concentration gradient across a cellular boundary, and ionic concentration gradient across a cellular boundary, despite changes in the electrochemical potential of the despite changes in the electrochemical potential of the external environment and despite changing identities and external environment and despite changing identities and activities of electron donors and acceptors.activities of electron donors and acceptors.

Allen JF (2010) Journal of Cosmology, 10, 3362-3373Allen JF (2010) Journal of Cosmology, 10, 3362-3373

A feedback loop involving the elements of a redox repressor or activator incorporating an iron-sulphur centre in a polypeptide

Allen J. F. (2010) Journal of Cosmology, 10, 3362-3373

Allen J. F. (2010) Journal of Cosmology, 10, 3362-3373

A feedback loop involving the elements of a two-component redox regulatory system

Nick Lane, John F. Allen and William Martin (2010) How did LUCA make a living? Chemiosmosis in the origin of life. Bioessays 32: 271-280

A. The proton motive force across the boundary of LUCA.  The pmf is made by an alkaline (high pH) internal effluent from LUCA's founding hydrothermal vent and by an acidic (low pH) external environment of carbonic acid solution.

B. The proton motive force of living cells descended from LUCA.  The pmf is made by an alkaline (high pH) cytoplasm and by an acidic (low pH) extracellular environment.  The gradient is continuously replenished by electrons flowing across the membrane from donors to acceptors.

Steps from a porous inorganic metal-sulphide vesicle to a free-living cell.A. Hydrothermal fluid is alkaline, in contrast to the acidic ocean into which it flows through the boundary of a porous, catalytic, metal sulphide vesicle.

Steps from a porous inorganic metal-sulphide vesicle to a free-living cell. B. The porosity of the vesicle is decreased by the intrusion of products of autocatalytic carbon reduction, symbolized by the cycle of carbon. The boundary of the vesicle begins to sustain a small gradient of hydrogen ion concentration and electrical potential difference: positive (P-phase) outside; negative (N-phase) inside.

Steps from a porous inorganic metal-sulphide vesicle to a free-living cell.C. The porosity decreases further. A mechanism of negative feedback interrupts proton influx, and an active transport drives proton efflux.

Steps from a porous inorganic metal-sulphide vesicle to a free-living cell.D. The vesicle is now formed from the organic intrusion alone, which presents a sufficient barrier to proton influx for the active proton efflux to generate and sustain a proton motive force. The dependency on hydrothermal fluid is lost, along with the inorganic container, though remnants of the latter remain to support vectorial electron and proton translocation.

Steps from a porous inorganic metal-sulphide vesicle to a free-

living cell.

Allen JF (2010) Redox homeostasis in the emergence of life. On the constant internal environment of nascent living cells. Journal of Cosmology, 10, 3362-3373

Coupling between information processing and redox chemistry.

Cellular co-location of energy transduction with genome function

Coupling between information processing and redox chemistry.

Cellular co-location of energy transduction with genome function

Photosynthesis and respirationPhotosynthesis and respiration

P+ve

N–ve

P+ve

N–ve

P+ve

N–ve

After: David S. Goodsell (1993) The Machinery of Life. Springer

P+ve

N–ve

Allen, J. F. (1993) Control of Gene Expression by Redox Potential and the Requirement for Chloroplast and Mitochondrial Genomes. Journal of Theoretical Biology, 165, 609-631

P+ve

N–ve

Coupling between information

processing and redox chemistry. Cellular co-location of energy

transduction with genome function

Coupling between information processing and redox chemistry.

Cellular co-location of energy transduction with genome function

CoCo-location for -location for RRedox edox RRegulationegulation

Allen, J. F. (2003) The function of genomes in bioenergetic organelles. Philosophical Transactions of the Royal

Society of London Series B-Biological Sciences, 358, 19-37

jfallen.org/corr

Allen J. F., de Paula W. B. M., Puthiyaveetil S. & Nield J. (2011) Trends in Plant Science 16(12), 645-655

Coupling between information processing and redox chemistry.

Cellular co-location of energy transduction with genome function

Coupling between information processing and redox chemistry.

Cellular co-location of energy transduction with genome function

The Redox Switch Hypothesis for The Redox Switch Hypothesis for the first cyanobacterium and for the first cyanobacterium and for

the origin of oxygenic the origin of oxygenic photosynthesisphotosynthesis

Allen J. F. (2005) A redox switch hypothesis for the origin of two light reactions in photosynthesis. FEBS Letters 579: 963-968

Archean landscapeArchean landscape by Peter Sawyer of the Smithsonian Institution by Peter Sawyer of the Smithsonian Institution

Phanerozoic LandscapeStromatolites at Hamelin Pool, Western Australia

Photograph 4 October 2007 by Catherine Colas des Francs-Small, The University of Western Australia

Policeman Point, Coorong Lagoon, South AustraliaPhotograph 9 March 2009 by the author

Policeman Point, Coorong Lagoon, South AustraliaPhotograph 9 March 2009 by the author

Dietrich, L.E.P., Tice, M.M. and Newman, D.K. (2006) The co-evolution of life and Earth. Current Biology 16, R396-R400

http://en.wikipedia.org/wiki/Geologic_time_scale

Coupling between information processing and redox chemistry.

Cellular co-location of energy transduction with genome function

Coupling between information processing and redox chemistry.

Cellular co-location of energy transduction with genome function

Acknowledgements...Acknowledgements...

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William MartinHeinrich-Heine-Universität Düsseldorf

Michael RussellNASA-JPL, California Institute of Technology

Nick LaneUniversity College London

John RavenUniversity of Dundee

Liang Wang • John Allen • Noor Agip • Iskander Ibrahim • Wilson de PaulaLiang Wang • John Allen • Noor Agip • Iskander Ibrahim • Wilson de PaulaAugust 29, 2013August 29, 2013

(↑). The Earth–Moon system photographed from the Cassini orbiter, Saturn. July 19, 2013.(↑). The Earth–Moon system photographed from the Cassini orbiter, Saturn. July 19, 2013.NASA/JPL-Caltech/Space Science Institute. NASA/JPL-Caltech/Space Science Institute.

Thank you for your Thank you for your attention.attention.

http://jfallen.orghttp://jfallen.org

Horizontal Gene Transfer and the Last Universal Common Ancestor •Horizontal Gene Transfer and the Last Universal Common Ancestor •

The Open University • 5th and 6th September 2013The Open University • 5th and 6th September 2013