Obcells as Proto-Organisms: Membrane Heredity, Lithophosphorylation, and the Origins of the Genetic Code, the First Cells, and Photosynthesis |
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Authors: | Thomas Cavalier-Smith |
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Institution: | (1) Department of Zoology, University of Oxford, South Parks Road, Oxford, OX1 3PS, United Kingdom, GB |
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Abstract: | I attempt to sketch a unified picture of the origin of living organisms in their genetic, bioenergetic, and structural aspects.
Only selection at a higher level than for individual selfish genes could power the cooperative macromolecular coevolution
required for evolving the genetic code. The protein synthesis machinery is too complex to have evolved before membranes. Therefore
a symbiosis of membranes, replicators, and catalysts probably mediated the origin of the code and the transition from a nucleic
acid world of independent molecular replicators to a nucleic acid/protein/lipid world of reproducing organisms. Membranes
initially functioned as supramolecular structures to which different replicators attached and were selected as a higher-level
reproductive unit: the proto-organism. I discuss the roles of stereochemistry, gene divergence, codon capture, and selection
in the code's origin. I argue that proteins were primarily structural not enzymatic and that the first biological membranes
consisted of amphipathic peptidyl-tRNAs and prebiotic mixed lipids. The peptidyl-tRNAs functioned as genetically-specified
lipid analogues with hydrophobic tails (ancestral signal peptides) and hydrophilic polynucleotide heads. Protoribosomes arose
from two cooperating RNAs: peptidyl transferase (large subunit) and mRNA-binder (small subunit). Early proteins had a second
key role: coupling energy flow to the phosphorylation of gene and peptide precursors, probably by lithophosphorylation by
membrane-anchored kinases scavenging geothermal polyphosphate stocks. These key evolutionary steps probably occurred on the
outer surface of an `inside out-cell' or obcell, which evolved an unambiguous hydrophobic code with four prebiotic amino acids
and proline, and initiation by isoleucine anticodon CAU; early proteins and nucleozymes were all membrane-attached. To improve
replication, translation, and lithophosphorylation, hydrophilic substrate-binding and catalytic domains were later added to
signal peptides, yielding a ten-acid doublet code. A primitive proto-ecology of molecular scavenging, parasitism, and predation
evolved among obcells. I propose a new theory for the origin of the first cell: fusion of two cup-shaped obcells, or hemicells,
to make a protocell with double envelope, internal genome and ribosomes, protocytosol, and periplasm. Only then did water-soluble
enzymes, amino acid biosynthesis, and intermediary metabolism evolve in a concentrated autocatalytic internal cytosolic soup,
causing 12 new amino acid assignments, termination, and rapid freezing of the 22-acid code. Anticodons were recruited sequentially:
GNN, CNN, INN, and *UNN. CO2 fixation, photoreduction, and lipid synthesis probably evolved in the protocell before photophosphorylation. Signal recognition
particles, chaperones, compartmented proteases, and peptidoglycan arose prior to the last common ancestor of life, a complex
autotrophic, anaerobic green bacterium.
Received: 19 February 2001 / Accepted: 9 April 2001 |
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Keywords: | : Obcell — Genetic code — Polyphosphate bioenergetics — Membrane heredity — Protocell — Origin of photosynthesis — Cenancestor |
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