(For part 1 of this series on the first two chapters, click
here.)
Having outlined the problem in Part I of The Vital Question, Nick Lane narrows
down the conditions for the origin of life in Part II. Chapter 3 is titled “Energy
at life’s origin” and Chapter 4 is “The emergence of cells”.
In Chapter 3, Lane cuts straight to the chase by outlining
six requirements for a cell, assumed to be the smallest “unit” of a living
organism.
·
A supply of reactive carbon
·
Free energy to drive some sort of metabolism
·
Catalysts (to overcome the “right” kinetic
barriers)
·
A method to excrete “waste” products
·
Compartmentalization
·
Informational material that is heritable
He skips past the details I really care about (the
formation of primitive metabolic cycles involving a range of small organics),
but no fault of his as we simply don’t know at this stage what might have
constituted these cycles. That’s why it is an area of active research I’m
engaged in. If the answers were known, this would be less interesting.
Lane’s goal is to quickly narrow the cradle of life to just
one viable option (that we are aware of) – alkaline hydrothermal vents. The
argument is made by quickly and effectively dismissing other possibilities
suggested in the literature. First, he rules out an open primordial ocean/soup.
This makes sense because concentrations of suitable organic molecules would be
miniscule, and no interesting chemistry will take place. He then rules out
freezing environments. Although useful for concentrating organics, a mechanism
for continuous supply of the building blocks is lacking. That’s a problem.
What might the source of carbon be? Lane thinks CO2
is a good candidate (CO being too low in concentration), and there is some
evidence that the amount of CO2 was significantly higher on the
Hadean earth compared to today. His candidate reaction to form organics such as
formate, formaldehyde, methanol, methane (this list represents increasingly
reduced C1 molecules), is the reaction of CO2 and H2.
The problem is that this reaction is not favorable within the same pH
environment, however if there was a pH gradient then this might work. Lane
calculates that the redox reaction is favorable if the oxidation of H2
took place at pH 10, and CO2 reduction to formaldehyde was at pH 6.
His solution: An iron-sulfide semiconducting barrier that you might find in an
alkaline hydrothermal vent. The outside of the vent has acidic waters (pH 5-7
has been suggested; I’ve even published a paper on this). The inside of the
vent is alkaline.
There’s a nice vignette about the discovery of the Lost City
alkaline hydrothermal vents of white smokers vindicating the predictions made
by Martin Russell years before. Lane effectively argues against the black
smokers (much hotter and more acidic) as the cradle of life. The higher
temperatures and much more acidic conditions favor hydrolysis – it is hard to
maintain any semblance of a short polymer (proteins, starches, DNA are all
polymers) in this environment. Lane also enumerates characteristics of the milder
white smokers that may allow both the concentration (via thermophoresis) and
incubation of small organics. This could be the route to more complex
molecules. He doesn’t specify details, and his group is building physical
models to simulate these conditions to test his hypothesis. (That’s how science
should work!) His model also ties in nicely with the proton pumps mentioned in
Part 1.
Chapter 4 opens with the problem of lateral gene transfer in
prokaryotes “erasing” the history that might otherwise be reconstructed through
phylogenetics. (Bill Martin’s ‘amazing disappearing tree’ is used as an
example.) Lane then enumerates some of the key differences between archaea and bacteria.
What they share was likely present in the Last Common Universal Ancestor (LUCA).
These include proteins and ribosome translational machinery, DNA and some form
of transcription, and an ATP synthase – i.e. the ability to pump protons.
However they differ in many other ways ranging from membrane composition to the
DNA replication apparatus. Lane uses these odd similarities and dissimilarities
to construct a hypothesis.
First, he narrows down fixing carbon to what may be the
simplest and most ancestral pathway – the acetyl-CoA pathway – except that a
simpler functional group could have initially substituted for CoA. Essentially
he invokes the fundamental use of thioester chemistry in fixing CO2
and H2 into the bevy of organics required in a primitive
biochemistry. He openly admits not knowing what this chemistry might look like
in detail and he handwaves the formation of DNA (a gaping hole that may not be
easy to solve given what we know from prebiotic syntheses). However, his
picture is intriguing at the very least. The iron-sulfide cores of the
hydrogenase and ferredoxin in methanogens suggest some continuity with the inorganic
network that constitutes the hydrothermal vents.
LUCA may have started out lodged in the inorganic membrane
between the acidic ocean and the alkaline hydrothermal liquid. The natural
proton gradient results in an influx of protons from the acidic ocean into the
cell. To maintain the influx requires relatively quick neutralization from the
alkaline fluid, either by protons diffusing out the other side or from an
influx of hydroxide. In any case, the membrane has to be leaky for this to work
– this may be the case for simple fatty acids that are candidates for primitive
cell membranes. (Many experiments suggest their synthesis is straightforward,
and they allow for both growth and division of cell-like structures.) The
problem is that LUCA is stuck where it is. Evolving a less permeable membrane
kills the proton motive force to drive a primitive metabolism.
Lane thinks there might be an ingenious solution that kills
two birds with one stone. The evolution of Na/H antiporters as a
‘preadaptation’ may have led to the further evolution of active pumping
proteins, which would co-evolve with increasing membrane permeability. These
could allow the cells increasing freedom from being stuck in the inorganic layer
allowing them to ‘colonize’ other areas, be it in the vent system or possibly
further afield. The bacteria and archaea may have come from two different evolutionary
routes in this process. It may (in broad sweeping terms) explain the
similarities and differences between the two domains of life. It certainly
avoids some of the other problems that attempt to find the root to the tree of
life (Lane enumerates the different possibilities, which he then argues
against.)
The factoid that jumped out at me while reading this section
was the amount of ‘waste’ generated compared to biomass synthesized. The ratio
is 40 to 1. Methanogens spend most of their energy budget generating methane
(and water) to pump protons. That’s the price to be paid in a free-living cell.
In the leaky membrane of a cell attached to the wall of a hydrothermal vent, more
energy may have been available in a natural proton gradient to drive a primitive
(and evolving) carbon metabolism synthesizing the building blocks of proteins
and nucleic acids. LUCA was a creature of the vent. Could we find LUCA now?
Unfortunately not, as any other vent organisms now would devour metabolites and
outcompete anything as primitive as LUCA. Could we build a LUCA under
artificial conditions? Possibly. It won’t be easy, but it will be very
interesting if some research group succeeds!
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