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From Two Cells To Eukarya
From Two Cells To Eukarya
From Two Cells To Eukarya
Wächterhäuser
MicroHypothesis
Fig. 1. Illustration of the divergence of the domains Bacteria, Archaea and Eukarya from stages PC-1, PC-2 and PC-3 in the second phase of
evolution (modified from Kandler, 1994a,b).
A. The overall process. (B is inset in Fig. 1A). The wavy line at the bottom signifies that the shown second phase of evolution continues an older
first phase of evolution beginning with the origin of life. The widths of the trunk and branches signify the diversity of subpopulation in different
habitats and the massiveness of parallel evolution. Pre-cells are symbolized by squares, the horizontal lines with thin and fat segments signifying
the linear intermediates of rolling circle plasmid replication. Archaea and Bacteria are symbolized by circles with circular chromosomes.
B. Symbiosis.
Fig. 2. Ancestral gene clusters (modified from Wächtershäuser, 1998), reconstructed for the bacterial common ancestor (BCA), the archaeal
common ancestor (ACA) and the pre-cells of stage PC-1. SecE, SecY, translocase units; NusG, NusA, anti-termination factors; L11, etc., large
subunit ribosomal proteins; S12, etc., small subunit ribosomal proteins; RpoB, etc., RNA polymerase units; EF-G, EF-Tu, elongation factors; Adk,
adenylate kinase; Map, methionine aminopeptidase; IF-1, initiation factor; Cmk, cytidylate kinase.
complex multicomponent genetic machinery, which cer- organisms with heterochiral hybrid membranes is selec-
tainly must have required containment inside a stable tively disfavoured. Therefore, a transformation of a bacte-
cellular structure. It was then argued that the pre-cells rial membrane into an archaeal membrane or vice versa
must have been bounded by a stable lipid membrane. This through a hybrid heterochiral membrane would have been
leads to the following key question: what were the lipids counter-selective. This contradicts all previous theories,
of this pre-cell membrane? which explicitly postulate or implicitly entail such transfor-
All extant species have lipids with basically the same mation (Cavalier-Smith, 1987; Sogin, 1991; Gupta and
structure: a glycerol-phosphate head group to which two Golding, 1996; Martin and Müller, 1998; Koga et al., 1998;
long-chain hydrophobic rests are attached. It is this struc- Horiike et al., 2001; Hartman and Fedorov 2002; Cavalier-
ture that gives the lipids the wedge-shape required for the Smith 2002). The problem of lipid membrane transforma-
formation of stable closed vesicles (Ackermann, 1992). It tion has been discussed by Forterre (2001). A proposal
therefore seems compelling that the stable membrane of by Zillig et al. (1992) avoids this transformation by postu-
the pre-cells had lipids with this kind of basic structure. lating that the bacterial lipid membrane and the archaeal
But then we come across a peculiar fact. All species of lipid membrane emerged independently from each other
the domains Bacteria and Eukarya have glycerol-3- by replacing an ancestral non-lipid cell membrane made
phosphate lipids, while all species of the domain Archaea of proteins. Incidentally, lipid incompatibilities in cases of
have glycerol-1-phosphate lipids (Kates, 1993; Nishihara chiral discrimination are well established (Sackmann,
et al., 1998). So far not a single exception has been 1982; Nassoy et al., 1995; Vollhardt et al., 1996; Kaganer
reported, nor a single case where both these two enanti- et al., 1999; Uragami et al., 2000).
omers of the chiral glycerol-phosphate lipids co-exist in a A solution of this problem is now proposed. Generally
hetero-chiral membrane. How do we account for this speaking, it is proposed that whereas evolution from a
situation. homochiral stable membrane through a less stable het-
Two possibilities come immediately to mind. The first erochiral membrane into the antipodal stable membrane
possibility is that the membrane of the pre-cells prior to would have been counter-selective and thus of low prob-
PC-1 consisted of lipids of the bacterial enantiomeric type ability, an evolution could have proceeded with much
and that these evolved into the archaeal enantiomeric higher probability from a heterochiral membrane of lesser
type. The second possibility is that the membrane of the stability separately into the two antipodal more stable het-
pre-cells prior to PC-1 consisted of lipids of the archaeal erochiral membranes. Specifically, according to this pro-
enantiomeric type and that these evolved into the bacterial posal the pre-cells before the divergence of the domain
enantiomeric type. Both possibilities require that evolution Bacteria did have a stable bilayer membrane of a race-
would have had to proceed through intermediate organ- mate of chiral lipids, which may well have been glycerol-
isms with heterochiral hybrid membranes of a mixture of phosphate lipids. These lipids were synthesized inside the
lipids of the bacterial and archaeal enantiomeric types. It pre-cells. The metabolism of the pre-cells was still in the
is here proposed that as a result of lipid incompatibility stage of a conversion of non-stereospecific reaction steps
such heterochiral hybrid membranes are less stable than to reaction steps catalysed by stereo-specific enzymes
the homochiral membranes of the Bacteria and Archaea. (Wächtershäuser, 1988; 1992). In this evolutionary phase
This means that a gradual evolution through intermediate the glycerol moiety of the lipids is assumed to have