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  Not only do the iron-sulphur cells provide a continuous supply of energy, but they also act as miniature electrochemical reactors, catalysing fundamental biochemical reactions, and concentrating the reaction products. The basic building blocks of life, including RNA, ADP, simple amino acids, small peptides, and so on, could all have been formed by virtue of the catalytic properties of the iron-sulphur minerals, and perhaps sedimental clays, in the reactions described by Gunter Wächtershäuser, but with two great advantages—they are concentrated by the membranes (preventing them from diffusing away into the oceans), and they are powered by a natural source of energy, the proton gradient.

  Life itself

  Does all this sound improbable? In the previous chapter, I suggested that the origin of life was not as improbable as the evolution of the eukaryotes. Think about what is happening here. Such conditions could not have been rare on the early earth. Volcanic activity has been estimated to be fifteenfold greater than today. The crust was thinner, the oceans shallower, and the tectonic plates were only just forming. Volcanic seepage sites must have existed across much of the surface of the earth, to say nothing of more violent volcanic activity. The formation of many millions of tiny cells, bounded by iron-sulphur membranes, requires no more than a difference in redox state and acidity between the oceans and the volcanic fluids emanating from deep in the crust—a difference that certainly existed.

  The early earth, as envisaged by Russell, is a giant electrochemical cell, which depends on the power of the sun to oxidize the oceans. UV rays split water and oxidize iron. Hydrogen, released from water, is so light that it is not retained by gravity, and evaporates off into space. The ocean becomes gradually oxidized, relative to the more reduced conditions in the mantle. According to the basic rules of chemistry, the mixing zone inevitably forms natural cells, replete with their own chemiosmotic and redox gradients. Mixing would have been assisted by the high tidal range, drawn the tug of the newly formed moon, which was closer to the earth then than today. We can be almost certain that such cells would really have formed, perhaps on a massive scale. And of course we can see their remains in the geology of places like Tynagh. There is a long way to go from here to make even a bacterium, but these conditions are a good first step.

  Not only would the requisite conditions have been probable, but they would have been stable and continuous. They depend only on the power of the sun, without requiring problematic inventions like photosynthesis or fermentation. The sun is only needed to oxidize the oceans, as we know it must. Of all the forms of energy mooted by astrobiologists—meteorite impacts, volcanic heat, lightning—the power of the sun has often been curiously overlooked by scientists, if not by prehistoric mythologies. As the distinguished microbiologist Franklin Harold put it in his classic text, The Vital Force (the title of which I honour in the title of this Part): ‘One cannot help but suspect that the great stream of energy that passes across the earth plays a larger role in biology than our current philosophy knows: that perhaps the flood of power not only permitted life to evolve, but called it into being.’

  For hundreds of millions of years, the sun provided the constant source of energy needed to pay the debt to the second law of thermodynamics. It created chemical disequilibria, and promoted the formation of naturally chemiosmotic cells. The primordial conditions are still faithfully replicated in the fundamental properties of all cells today. Both organic and inorganic cells are bounded by a membrane, which physically contains the cell’s organic constituents, preventing them from diffusing away into the oceans. In both organic and inorganic cells, biochemical reactions are catalysed by minerals (today embedded as the prosthetic groups of enzymes). In both cases, the membrane is the barrier as well as carrier of energy. In both cases, energy is captured by a chemiosmotic gradient, with a positive charge and acidic conditions on the outside, and relatively negative, alkaline conditions on the inside. In both cases, redox reactions, electron transport and proton pumping regenerate the gradient. When the bacteria and archaea finally emerged from their nursery, to venture into the open oceans, they took with them an unmistakable seal of their origin. They parade it still today.

  But this imprint, echoing the origin of life itself, was also life’s primary limitation. Why, we may ask, did bacteria never evolve beyond bacteria? Why did four billion years of bacterial evolution never succeed in producing a truly multicellular, intelligent bacterium? More specifically, why did the evolution of the eukaryotes require a union between an archaeon and a bacterium, rather than just the gradual accrual of complexity by a favoured line of bacteria or archaea? In Part 3, we’ll see that the answer to this long-standing riddle, and an explanation for the marvellous flowering of the eukaryotic line into plants and animals, lies in the fundamental nature of energy production by chemiosmotics across a bounding membrane.

  PART 3

  Insider Deal

  The Foundations of Complexity

  Bacteria ruled supreme on Earth for two billion years. They evolved almost unlimited biochemical versatility but never discovered the secrets of greater size or morphological complexity. Life on other planets may get stuck in the same rut. On Earth, large size and complexity only became possible once energy generation had been internalized in mitochondria. But why did bacteria never internalize their own energy generation? The answer lies in the tenacious survival of mitochondrial DNA, a two-billion-year-old paradox.

  A large cell with things inside—in eukaryotes, energy generation is internalized in mitochondria

  Here is a list of words to make an evolutionary biologist spill their beer: purpose, teleology, ramp of ascending complexity, non-Darwinian. All these terms are associated with a religious view of evolution—the sense that life was ‘programmed’ to evolve, to become more complex, to give rise to humanity on a smooth curve from the lowest animals to the angels, each approaching closer to God—the great ‘chain of being’. Such a view is popular not just with religious theorists, but nowadays with astrobiologists too. The idea that the laws of physics virtually summon life forth in the universe that we see around us is a comforting one, and evokes the idea that even human sentience may be an inevitable outcome of the workings of physics. I disagreed in Part 1, and we will consider the theme further in Part 3 by looking at the origin of biological complexity.

  In Part 1, we observed that all complex multicellular organisms on earth are composed of eukaryotic cells; in contrast, bacteria have remained resolutely bacterial for the best part of four billion years. There is a chasm between bacterial and eukaryotic cells, and life elsewhere in the universe might well get stuck in the bacterial rut. We have seen that the eukaryotic cell was first formed in an unusual union between a bacterium and an archaeon. The question we’ll look into now is the ‘seeding’ of complexity in eukaryotes: what exactly is it about the eukaryotic cell that seems to encourage the evolution of complexity? However misleading the impression may be, surveying the grand canvas of evolution after the appearance of the eukaryotic cell does engender a sense of purpose. The idea of a great chain of being, striving to approach closer to God, is not accidental, even if it is wrong. In Part 3, we’ll see that the seeds of complexity were sown by mitochondria, for once mitochondria existed, life was almost bound to become more complex. The drive towards greater complexity came from within, not from on high.

  In his celebrated book Chance and Necessity, the committed atheist and Nobel Prize-winning molecular biologist Jacques Monod tackled the theme of purpose. Plainly, he said, it is pointless to discuss the heart without mentioning that it is a pump, whose function is to pump blood around the body. But that is to ascribe purpose. Worse, if we were to say that the heart evolved to pump blood, we would be committing the ultimate sin of teleology—the assignment of a forward-looking purpose, a predetermined end-point to an evolutionary trajectory. But the heart could hardly have evolved ‘for’ anything else; if it didn’t evolve to pump blood, then it is truly a miracle that it happened to become so fine a pump.
Monod’s point was that biology is full of purpose and apparent trajectories, and it is perverse to pretend they don’t exist; rather, we must explain them. The question we must answer is this: how does the operation of blind chance, a random mechanism without foresight, bring about the exquisitely refined and purposeful biological machines that we see all around us?

  Darwin’s answer, of course, was natural selection. Blind chance serves only to generate random variation within a population. Selection is not blind, or at least not random: it selects for the overall fitness of an organism in its particular environment—the survival of the fittest. The survivors pass on their successful genetic constitution to their offspring. Thus any changes that improve the function of the heart at pumping blood will be passed on, while any that undermine it will be eliminated by selection. In each generation (in the wild) only a few per cent might survive to reproduce, and they will tend to be the luckiest or best adapted. Over many generations luck no doubt balances out, so natural selection tends to select the best adapted of the best adapted, inevitably refining function until other selective pressures balance out the tendency to change. Natural selection therefore works as a ratchet, which turns the operation of random variation into a trajectory. In retrospect this may well look like a ramp of ascending complexity.

  Ultimately, biological fitness is written in the sequence of the genes, for they alone are passed on to the next generation (well, almost alone: mitochondria are, too). Over evolutionary time, alterations in the genetic sequence, subjected to round after round of natural selection, build tiny refinement upon tiny refinement, until finally erecting the dizzying cathedral of biological complexity. Although Darwin knew nothing of genes, the genetic code at once suggests a mechanism for producing random variation in a population: mutations in the sequence of ‘letters’ in DNA can change the sequence of amino acids in proteins, which might have a positive, or a negative, or a neutral, effect on their function. Copying errors alone generate such variation. Each generation produces perhaps several hundred small changes in the DNA sequence (out of several billion letters), which may or may not affect fitness. Such small changes undoubtedly occur, and generate some of the raw material for the slow evolutionary change anticipated by Darwin. The gradual divergence in the sequence of genes of different species, over hundreds of millions of years, shows this process in action.

  But small mutations are not the only way to bring about change in the genome (the complete library of genes in one organism), and the more we learn about genomics (the study of genomes), the less important small mutations seem to be. At the least, greater complexity demands more genes—the small bacterial genome could hardly code for a whole human being, still less the myriad genetic differences between individuals. Surveying species, there is a general correlation between the degree of complexity and the number of genes, if not the total DNA content. So where do all these extra genes come from? The answer is duplications of existing genes, or whole genomes, or from the union of two or more different genomes, or from the spread of repetitive DNA sequences—apparently ‘selfish’ replicators, which copy themselves throughout the genome, but may later be co-opted to serve some useful function (useful, that is, to the organism as a whole).

  None of these processes is strictly Darwinian, in the sense of gradual, small refinements to an existing genome. Rather, they are large-scale, dramatic changes in the total DNA content—giant leaps across genetic space, transforming existing gene sequences at a single stroke—even if they generate the raw material for new genes, rather than the new genes themselves. Excepting these leaps across genetic space, the process is otherwise Darwinian. Changes to the genome are brought about in an essentially random manner, and then subjected to rounds of natural selection. Small changes hone the sequence of new genes to new tasks. So long as the big jumps in DNA content do not generate an unworkable monster, they can be tolerated. If there is no benefit in having twice as much DNA, then we can be sure that natural selection will jettison it again—but if complex organisms need a lot of genes, then the elimination of superfluous DNA surely puts a ceiling on the maximum possible complexity, for it eliminates the raw material needed to form new genes.

  This brings us back to the ramp of complexity. We have seen that there is a big discontinuity between bacteria and eukaryotes. It is remarkable that bacteria are still bacteria: while enormously varied and sophisticated in biochemical terms, they have resolutely failed to generate real morphological complexity in four billion years of evolution. In their size, shape, and appearance, they can hardly be said to have evolved in any direction at all. In contrast, in half the time open to bacteria, the eukaryotes unquestionably ascended a ramp of complexity—they developed elaborate internal membrane systems, specialized organelles, complex cell cycles (rather than simple cell division), sex, huge genomes, phagocytosis, predatory behaviour, multicellularity, differentiation, large size, and finally spectacular feats of mechanical engineering: flight, sight, hearing, echolocation, brains, sentience. Insofar as this progression happened over time, it can reasonably be plotted out as a ramp of ascending complexity. So we are faced with bacteria, which have nearly unlimited biochemical diversity but no drive towards complexity, and eukaryotes, which have little biochemical diversity, but a marvellous flowering in the realm of bodily design.

  When confronted with the divide between bacteria and eukaryotes, the Darwinian might reply: ‘Ah, but the bacteria did generate complexity—they gave rise to the more complex eukaryotes, which in turn gave rise to many organisms of inordinately greater complexity.’ This is true, but only in a sense, and here is the rub. The mitochondria, I shall argue, could only be derived by endosymbiosis—a union of two genomes in the same cell, or a giant leap across genetic space—and without mitochondria, the complex eukaryotic cell simply could not evolve. This viewpoint stems from the idea that the eukaryotic cell itself was forged in the merger that gave rise to mitochondria, and that the possession of mitochondria is, or was in the past, a sine qua non of the eukaryotic condition. This picture differs from the mainstream view of the eukaryotic cell, so let’s remind ourselves quickly why it matters.

  In Part 1, we examined the origin of the eukaryotic cell, as surmised by Tom Cavalier-Smith, which best represents the mainstream view. To recapitulate, a prokaryotic cell (without a nucleus) lost its cell wall, perhaps through the action of an antibiotic produced by other bacteria, but survived the loss, as it already had an internal protein skeleton (cytoskeleton). The loss of the cell wall had profound consequences for the cell in terms of its lifestyle and manner of reproduction. It developed a nucleus and a complicated life cycle. Using its cytoskeleton to move around and change shape like an amoeba, it developed a new, predatory lifestyle, engulfing large particles of food such as whole bacteria by phagocytosis. In short, the first eukaryotic cell evolved its nucleus and its eukaryotic lifestyle by standard Darwinian evolution. At a relatively late stage, one such eukaryotic cell happened to engulf a purple bacterium, perhaps a parasite like Rickettsia. The internalized bacteria survived and eventually transmuted, by standard Darwinian evolution, into mitochondria.

  Notice two things about this line of reasoning: first, it exhibits what we might call a Darwinian bias, in that it limits the importance attributed to the union of two dissimilar genomes, a basically non-Darwinian mode of evolution; and second, it limits the importance of mitochondria in this process. Mitochondria are incorporated into a fully functional eukaryotic cell, and are readily lost again in many primitive lines such as Giardia. Mitochondria, in this view, are an efficient means of generating energy, but no more nor less than that. The new cell simply had a Porsche engine fitted, in place of its old-fashioned milk-cart motor. I think this view gives little real insight into why all complex cells possess mitochondria, or conversely, why mitochondria are needed for the evolution of complexity.

  Now consider the hydrogen hypothesis of Bill Martin and Miklos Müller, which we also discussed in Part 1. According to this rad
ical hypothesis, a mutual chemical dependency between two very different prokaryotic cells led to a close relationship between the two. Eventually one cell physically engulfed the other, combining two genomes within a single cell: a giant leap across genetic space to create a ‘hopeful monster’. This genetic leap, in turn, set up a series of Darwinian selection pressures on the new entity, leading to a transfer of genes from the guest to the host. The critical point of the hydrogen hypothesis is that there never was a primitive eukaryote, one that supposedly possessed a nucleus and had a predatory lifestyle, but did not have any mitochondria. Rather, the first eukaryote was born of the union between two prokaryotes, a fundamentally non-Darwinian process—there was no halfway house.

  Just look at Figure 9, a tree of life drawn in 1905 by the Russian biologist Konstantine Merezhkovskii, to see what an uncomfortable reversal of the standard branching tree of life this creates. There has been plenty of controversy over trees of life in the past, notably from Stephen Jay Gould, who claimed that the Cambrian explosion inverted the usual tree. The Cambrian explosion refers to the great, and geologically sudden, proliferation of life around 560 million years ago. Later on, most of the major branches were ruthlessly pollarded, as whole phyla fell extinct. Daniel Dennett, in Darwin’s Dangerous Idea, lambasts Gould’s apparently radical evolutionary trees for being the same as any other evolutionary tree, except with distorted axes—a low-lying scrub bush, throwing up a few scraggly shoots, rather than a lofty tree of life. But there is no danger of this in Merezhkovskii’s case. His evolutionary tree is a genuinely upside down variety. Here, the branches fuse, rather than bifurcate, to generate a new domain of life.