why do chloroplasts and mitochondria have genomes?
TRANSCRIPT
Lectures by
John F. Allen
School of Biological and Chemical Sciences, Queen Mary, University of London
Cell Biology and Developmental Genetics
1jfallen.org
Cell Biology and Developmental Genetics
Lectures by John F. Allen
Endosymbiosis and the origin of bioenergetic organelles. Some history
Endosymbiosis and the origin of bioenergetic organelles. A modern view
Mitochondria as we know them and don't know them
Why do chloroplasts and mitochondria have genomes?
Co-location for Redox Regulation
Mitochondria, ageing, and sex – energy versus fidelity
Cell Biology and Developmental Genetics
Lectures by John F. Allen
Slides and supplementary information:
jfallen.org/lectures
School of Biological and Chemical Sciences Seminars
WEDNESDAYS AT 12 NOON IN G23, G. E. FOGG BUILDING
3 February 2010
Dr Nick LaneProvost’s Venture Research Fellow, University College LondonLife Ascending. The Ten Great Inventions of Evolution
Lecture 4
Why do chloroplasts and mitochondria have genomes?
I II III IV ATPase
Mitochondrial matrix
Inter-membrane space
Chloroplast stroma
Thylakoid lumen
Cyt b6-f Photosystem I ATPasePhotosystem II
RubisCO
Problem
Why Do Mitochondria and Chloroplasts Have Their Own Genetic Systems?
Why do mitochondria and chloroplasts require their own separate genetic systems when other organelles that share the same cytoplasm, such as peroxisomes
and lysosomes, do not? …. The reason for such a costly arrangement is not clear, and the hope that
the nucleotide sequences of mitochondrial and chloroplast genomes would provide the answer has proved unfounded. We cannot think of compelling
reasons why the proteins made in mitochondria and chloroplasts should be made there rather than in the
cytosol.Molecular Biology of the Cell
© 1994 Bruce Alberts, Dennis Bray, Julian Lewis, Martin Raff, Keith Roberts, and James D. WatsonMolecular Biology of the Cell, 3rd edn. Garland Publishing
Proposed solutions (hypotheses)
There is no reason. “That’s just how it is”. (Anon)
The “Lock-in” hypothesis. (Bogorad, 1975). In order for core components of multisubunit complexes to be synthesised, de novo, in the correct
compartment.
The evolutionary process of transfer of genes from organelle to nucleus is still incomplete.
E.g. Herrmann and Westhoff, 2001: The partite plant genome is not in a phylogenetic equilibrium. All available data suggest that the ultimate aim of genome restructuring in the plant cell, as in the eukaryotic cell in general,
is the elimination of genome compartmentation while retaining physiological compartmentation.
The frozen accident. The evolutionary process of gene transfer was underway when something happened that stopped it. E.g. von Heijne,
1986.
It’s all a question of hydrophobicity. The five-helix rule. (Anon)
Some proteins (with co-factors) cannot be imported. (Anon)
Co-location for Redox Regulation - CoRR (Allen 1993, 2003 et seq.)
Why Do Mitochondria and Chloroplasts Have Their Own Genetic Systems?
Proposed solution (hypothesis)
Why Mitochondria and Chloroplasts Have Their Own Genetic Systems
Allen, J. F. (1993) J. Theor. Biol. 165, 609-631
Allen, J. F. (2003) Phil. Trans. R. Soc. B458, 19-38
Co-location for Redox Regulation - CORR
Vectorial electron and proton transfer exerts regulatory control over expression of genes encoding proteins directly
involved in, or affecting, redox poise.
This regulatory coupling requires co-location of such genes with their gene products; is indispensable; and operated
continuously throughout the transition from prokaryote to eukaryotic organelle.
Organelles “make their own decisions” on the basis of environmental changes affecting redox state.
BacteriumEndosymbiontBioenergetic organelle
1. As now generally agreed, bioenergetic organelles evolved from free-living bacteria. 2. Gene transfer between the symbiont or organelle and the nucleus may occur in
either direction and is not selective for particular genes.3. There is no barrier to the successful import of any precursor protein, nor to its
processing and assembly into a functional, mature form. 4. Direct redox control of expression of certain genes was present in the bacterial
progenitors of chloroplasts and mitochondria, and was vital for cell function before, during, and after the transition from bacterium to organelle. The mechanisms of this control have been conserved.
5. For each gene under redox control, it is selectively advantageous for that gene to be retained and expressed only within the organelle.
6. For each bacterial gene that survives and is not under redox control, it is selectively advantageous for that gene to be located in the nucleus and expressed only in the nucleus and cytosol. If the mature gene product functions in chloroplasts or mitochondria, the gene is first expressed in the form of a precursor for import.
7. For any species, the distribution of genes between organelle and nucleus is the result of selective forces that continue to operate.
8. Those genes for which redox control is always vital to cell function have gene products involved in, or closely connected with, primary electron transfer. These genes are always contained within the organelle.
9. Genes whose products contribute to the organelle genetic system itself, or whose products are associated with secondary events in energy transduction, may be contained in the organelle in one group of organisms, but not in another.
10. Components of the redox-signalling pathways upon which co-location for redox regulation depends are themselves not involved in primary electron transfer, and so their genes have been relocated to the nucleus.
Co-location for Redox Regulation - CoRRTen assumptions, axioms, principles jfallen.org/corr
Co-location for Redox Regulation - CoRR
Prediction: Explanation of previous knowledge
Distribution of genes for components of oxidative phosphorylation between mitochondria
and the cell nucleus
Prediction: Experimental results
Redox control of mitochondrial and chloroplast gene expression
Prediction: Experimental results
Persistence of “bacterial” redox signalling components in chloroplasts and mitochondria
Co-location for Redox Regulation - CoRR
Prediction
Explanation of previous knowledge
Distribution of genes for components of oxidative phosphorylation between mitochondria and the cell nucleus
Redox regulation
Redox regulation
Nucleus Cytosol
N-phaseMitochondrial matrix
O2
H2O
I II III IV ATPase
Mitochondrial matrix
Inter-membrane space
I II III IV ATPase
Mitochondrial matrix
Inter-membrane space
H+ H+ H+
H+
NADH O2
ATP
ADP
H2O
NAD+ succinate fumarate
Redox regulation
Nucleus Cytosol
N-phaseMitochondrial matrix
O2
H2O
Allen JF (2003) The function of genomes in bioenergetic organellesPhilosophical Transactions of the Royal Society of London Series B-Biological Sciences 358: 19-37
Co-location for Redox Regulation - CORR
Prediction
Explanation of previous knowledge
Distribution of genes for components of photosynthetic phosphorylation between
chloroplasts and the cell nucleus
Redox regulation
Redox regulation
Light Light
Nucleus Cytosol
N-phaseChloroplast stroma
CO2
CH2O
Chloroplast stroma
Thylakoid lumen
Cyt b6-f Photosystem I ATPasePhotosystem II
RubisCO
Cyt b6-f Photosystem I ATPase
Chloroplast stroma
Thylakoid lumen
Photosystem II
RubisCO
H+H+ H+
H+
ATP
NADP+
O2
H2O
H+
H+
H+
ADP
NADPH
ATP
ADP
H2O O2
NADP+
NADPH
Redox regulation
Light Light
Nucleus Cytosol
N-phaseChloroplast stroma
CO2
CH2O
Allen JF (2003) The function of genomes in bioenergetic organellesPhilosophical Transactions of the Royal Society of London Series B-Biological Sciences 358: 19-37
Co-location for Redox Regulation - CoRR
Prediction: Explanation of previous knowledge
Distribution of genes for components of oxidative phosphorylation between mitochondria
and the cell nucleus
Prediction: Experimental results
Redox control of mitochondrial and chloroplast gene expression
Prediction: Experimental results
Persistence of “bacterial” redox signalling components in chloroplasts and mitochondria
Lecture 5
Co-location for Redox Regulation