CHAPTER
10
THE
EVOLUTION OF EXPERIENCE
chapter
9
10.1.1.
When some of the débris orbiting a nascent sun coalesced to form
planet Earth the most experientially advanced organism upon it was a
small molecule. Today, some 4,500 million years later, it is a human
being. In general terms, how did the one evolve from the other? Our
current orthodoxy has its confident answer: it evolved via a long series
of small accretive changes, in which process nothing more was involved
than the laws and forces of physics and chemistry as conventionally
understood. At one end of the evolutionary process we have our own experience
- our perceptions, thoughts, feelings, volitions, imaginings, memories
etc.. At the other, the inanimate world as conceived by our current
mathematico-mechanistic physics. And according to orthodoxy the one
is no more than the other operating in ultra-complex situations that
it has itself built up over the aeons.
10.1.2.
Not surprisingly,
there have always existed biologists who have refused to accept this
answer, rightly regarding it as self-evidently absurd. However, the
great majority of these have no quarrel with orthodox physics; no less
than the materialists they accept this as a substantially true account
of the insentient, inanimate world. But a minority of them invoke some
other substance (mind) not at work in this world, and hence peculiar
to living things, to account for life. And they generally endow this
with some built-in nisus or teleological urge in order to account for
what, to us humans, seems the outstanding attribute of evolution: the
tendency of the biological world to give rise to ever more experientially
complex organisms with the passage of time. These biologists are the
dualists or vitalists, and I need hardly say that the irremediable shortcomings
of their theories have countless times shown them to be only marginally
less absurd than those of mechanistic orthodoxy. Which is only what
one might expect; once mechanistic physics (and in the rare instances
when it succeeds in saying anything physically meaningful, our mathematically
dominated 'new' physics is still purely mechanistic) has been accepted
as a substantially true account of the physical world, of necessity,
living organisms and their evolution present an insoluble problem.
10.1.3.
More intellectually
respectable are the organicists or Systems Theorists. These have proposed
no new physics, but in view of the obvious fact that mere mechanically
determined feedback is ludicrously incapable of accounting for living
organisms, to say nothing of their evolution, they claim that this gives
biologists the right to formulate their own basic laws, grounded on
the biological facts, independently of physical laws, and to leave to
future generations any ontological reconciliation of the two sets. The
essence of this organismic position is the claim that a whole, qua whole,
must somehow, as a superordinate entity in its own right, act back upon
the parts in such way as to bias their movements towards maintaining
itself as a viable system. In our present state of knowledge, the organicist
claims, one just has to accept this as an empirically established fact.
We have already (4.5.3.) provided one quote from Errol E. Harris on
this; here is another:
“It
[the cell] is no mere aggregate of chemical compounds or of synthetic
processes in equilibrium (though, of course, it is both), but is a
systematically structured, organised and temporally ordered web of
chemical interaction regulating itself for self-maintenance and displaying
an (as yet) scientifically inexplicable nisus towards order and elaboration
of coherent structure that runs counter to thermodynamic probability."1
There
are hundreds, if not thousands, of quotes saying essentially the same
thing in different words. We will content ourselves with two from Paul
Weiss2:
“Thus,
in contrast to the infinite number of possible interactions and combinations
among its constituent units which could take place in a mere complex,
in the living system only an extremely restricted selection from that
grab-bag of opportunities for chemical processes is being realized
at any one moment – a selection which can be understood solely
in its bearing on the concerted harmonious performance of a task by
the complex as a whole. This is the feature that distinguishes a living
system from a dead body ...”
“ ... the acknowledgement of field continua as ordering principles
in systems on the integral level is as valid and indispensable as
is the practical acceptance, on the differential level, of discrete
singularities within these continua, whether sub-atomic particles,
atoms, molecules, molecular assemblies, organelles, cells, or cell
assemblies ... I have yet to encounter any phenomena in the living
system which could be adequately described without reference to such
a dualistic scheme.”
It
is worth emphasising here that if the organicist contended that the
ultimate parts of organisms are much as the physicist conceives them,
he would be talking nonsense. It is just because he is contending that
physics has not yet attained to a true understanding of these parts
that the organicist can legitimately make the claims he does. As for
evolution, since the wholeness of any organism, however complex, is
just a natural attribute like any other, there is no reason why, in
conjunction with genetic mutation, replicative reproduction and Darwinian
selection, more complex wholes, if they possess some survival advantage
- as in some classes of organism they frequently do - should not evolve
on to increasingly elaborate levels of complexity.
10.1.4.
The next level in the ascent towards truth is defined by those organicists
who, from such attributes of living organisms as memory, habit, and
instinct, and the recapitulation of phylogeny in ontogeny, postulate
that the past is still in some way active, and exerts a force directly
on the present organism: that is, by definition of a force, causes the
material particles of which the organism is made to accelerate. Physical
orthodoxy acknowledges no such force - gravitation, electromagnetic
attractions and repulsions, and nuclear forces, strong and weak, making
up its whole inventory. By invoking a still active past, every thinker
of this persuasion - from Herbart, through von Hartmann, Bergson and
Geley to Rupert Sheldrake – has certainly taken a step in the
right direction; but because none has ever succeeded in constructing
a physics which coherently incorporates this active past, none has succeeded
in formulating a remotely satisfactory theory of how the past could
act within the present.
10.1.5.
We, of course, claim to have done precisely this; that is, as outlined
in the preceding chapters, developed this true or noumenal - as opposed
to merely phenomenal - physics: a physics in which, above all, the past
persists, and which, through sympathic association and mnemic causation,
is active in the physical present. From a physical world so conceived
the whole biological world from pre-cellular organisms to the advent
of homo sapiens, can be rationally derived. And it is such a derivation
- inevitably in the most general terms - that is attempted in this and
the three following chapters. Now, despite the fact that conventional
biological wisdom, as embodied in Neo-Darwinism, is unable to furnish
a rationally coherent account of even the simplest living organism,
this has proved no hindrance to its unearthing a vast, detailed, precisely
formulated and – albeit on a naively realistic conceptual level
- systematically organised body of observational and experimental fact.
So much is this the case, that what I am advancing as a true account
of experiential evolution may legitimately be viewed as a synthesis
of noumenal physics and Neo-Darwinism: that is, Neo-Darwinism incorporated
within an ontologically coherent conceptual schema. We may therefore
take the great truths of Neo-Darwinism as read and concentrate on everything
that conventional biology, grounded as it is on a radically false physics,
is incapable of explaining: basically, all those essential attributes
of living organisms and their evolution that derive from the preserved
past, sympathic association, mnemic causation, and paraphysical sequences.
This perfectly exemplifies what I wrote earlier (2.4.) about cosmology's
two methods - ontology (metaphysics) and science – complementing
one another: the strengths of each compensating for the other’s
deficiencies.
THE
COMPLEXIFICATION OF EXPERIENCE
10.2.1.
A point worth making at the outset is that complexity, as such, causes
nature no problems. As organisms grow more complex they show no sign
of an increasing tendency to break down. Complex organisms function
as naturally as simple - a cat no less efficiently than a cucumber.
All this is rooted in the fact that there is no limit to the number
of qualification sequences that sympathic association and mnemic causation
are capable of ordering.
10.2.2.
With this in mind, how, in broad terms, do we account for the evolving
complexity of experience from small molecules to human beings? We account
for it in terms of only five basic parameters.
Firstly, of course, there is all that arises as a consequence of our
noumenal physics: qualification sequences, a preserved past, its sympathic
association with the ongoing present, mnemic causation, and paraphysical
sequences.
Secondly, Darwinian selection. All successful species live by responding
to their environment – part inanimate, part composed of members
of their own and other species – in such way as to at least maintain
their numbers. That is, all species are obliged to create a viable ecological
niche, or perish.
Thirdly, breeding true. Organisms must be able to reproduce replicatively:
to produce offspring which closely resemble themselves in all respects.
Otherwise no changes, however advantageous, could be passed on down
the generations.
Fourthly, intra-specific variation. The members of a species, while
resembling one another in all essential respects, must yet exhibit within
these limits, considerable variation. Otherwise, there would be nothing
for selection to work on by way of adapting a species to changing environmental
conditions.
Fifthly, environmental variation. All over the planet, owing fundamentally
to plate tectonics, but subsidiarily to the activities of living organisms,
the inanimate environment, solid, liquid, and gaseous is in a constant
state of change, gradual or rapid, minor or major. Temperature and precipitation,
the chemical constitution of the atmosphere, the land/sea interface,
volcanic and seismic activity, mountain building and so forth - fluctuations
in these parameters give rise in course of time, at innumerable geographical
locations, to many significant changes. Species must respond adaptively
to such changes or perish, thereby producing yet further changes in
the environment of other species belonging to the same ecosystem.
10.2.3.
With these five parameters at work there is certainly no need whatever
to postulate some inbuilt orthogenetic nisus driving life on to ever
greater complexity. Certainly, we owe the very possibility of experiential
complexity to a physical world such as I am postulating. A physical
world as conceived by orthodoxy could not conceivably give rise to even
the simplest living organism. But this in no way implies that in our
system there is any inbuilt evolutionary drive towards greater complexity.
The opposite if anything, since the outstanding consequence of sympathic
association and mnemic causation is repetition not innovation –
as we see in such universal dispositions as instinct and habit. What
our noumenal physics does imply is that organisms can comfortably exist,
and consolidate themselves as a species, at any level of experiential
complexity. Inevitably, therefore, they are sometimes only a small step
away from an eco-niche requiring yet greater experiential complexity.
What causes this step to be taken?
10.2.4.
To answer this satisfactorily, we must bear in mind a number of basic
parameters germane to all such situations. Firstly, that the sole necessary
requirement for the establishment of a new species is its viability
in the face of Darwinian selection pressures. Secondly, that complexity
of experience is only one among innumerable biological attributes making
for such viability. Thirdly, that every new species arises as a result
of an accumulation of small changes to some already existing species.
From this it follows that new heights of experiential complexity must
arise as a consequence of small changes (mutations) to the genome of
a species whose viability is grounded on experiential complexity. The
basic cause of the arising of new species is environmental change of
one kind or another – topographical, climatic, or biological.
No environment is unchanging: there are only degrees of change, in terms
of magnitude, range or rapidity. But whatever the cause, to such changes
all species of the ecosystem affected must adapt or perish. And such
adaptation occurs as a result of Darwinian selection acting upon the
variations among the members of a species, these phenotypal variations
being, themselves, the consequence of small genotypal mutations. The
phenotypal variants within this gene pool offering the greatest survival
advantage under these changed conditions may well be significantly different
from those which offered it under the old. As a consequence they will
gradually replace these as the species’ norm. Sometimes, to such
an extent that the old species, as an interbreeding pool, splits into
two. That is, a new species is born. And it will sometimes be the case
that among species whose viability is grounded in complexity of experience,
there will arise in this way new species that are experiencing more
complexly than the old. Clearly, it is inherently probable that a wider
range of perception, a more varied behavioural repertoire, a richer
collaboration between perception and memory, more accurate anticipation,
and so on, may all offer better chances of survival for an organism
whose viability is predicated upon experiential complexity.
10.2.5.
Further, it is an attribute of experiential complexity that it tends
to feed on itself – that is, that a degree of positive feedback
is inherent in it. In an ecosystem, every species is part of the environment
of every other, changes in one species resulting in changes in others,
a whole wave of species changes tending to spread through the ecosystem
until it has settled into a new stability. And much of this interdependence
is of a competitive nature – in the struggle for available resources,
or even in the direct conflict of predator and prey. Any increase in
experiential complexity of one species opens up a new eco-niche for
any other species of the ecosystem sufficiently advanced experientially
to advance the further step needed to take advantage of this increase.
10.2.6.
When we say that experiential complexity evolves we mean only that,
as a general trend, the planet at any time will contain a species more
experientially complex than any existing at an earlier. These most experientially
complex species of their time constitute only a tiny minority of species.
The only essential attribute of any successful species is viability
– the ability to maintain its numbers. And there are innumerable
ways of doing that; experiential complexity being but one of innumerable
biological parameters constituting the functional core of an eco-niche.
Nor, if we traced the evolution of humans from small organic molecules,
would it be through these most experientially complex species of their
day. In many cases a certain evolutionary line is prevented from evolving
beyond a certain level of complexity. All species arise by gradual steps
from pre-existing species of the same anatomical and physiological groundplan;
and this may be such as to place a definite ceiling on their experiential
complexity. As we shall be seeing in some detail in the following three
chapters, the physical underpinning of ever more complex experience
is an increasingly complex nervous system. And one essential attribute
of such complexity is sheer number of nerve cells. But large nervous
systems require an internal skeleton to support and maintain them effectively.
Hence, invertebrate animals cannot evolve large nervous systems, the
large marine molluscs being by far the most experientially complex organisms
that nature can produce in this direction. With this minor exception,
all higher complexification of experience is confined to the vertebrate
phylum. And here, again, we shall find that there are many kinds of
eco-niche whose nature either puts an upper limit on experiential complexity,
or where greater experiential complexity has little if any survival
advantage. This last is particularly the case where selection pressures
are low. We have said that environmental change is the greatest stimulus
to evolution, but such change tends to be much greater, more frequent,
and more diverse on land than in the ocean. As a result, the evolution
of experiential complexity has been largely confined to the land dwelling
vertebrates.
EVOLUTION
OF REPLICATIVE REPRODUCTION
FIRST STAGE
10.3.1.
We have seen (10.2.2.) that the evolution of experience from molecule
to man has five basic causal parameters: mnemic causation, Darwinian
selection, intraspecific variation, replicative reproduction, and environmental
change. Now, the earliest self-reproducing cells are generally regarded
as having evolved by around 3800 m.y.b.p.1– that is,
some 700 million years after the formation of the Earth. According to
our theory, mnemic causation and environmental change would obviously
have been factors operating from the start. Also it seems natural enough
that as soon as individual metabolic complexes began to form there would
inevitably be some kind of passive competition among them for nutrients
and otherwise favourable locations. Likewise, that, although there were
no species, the various pre-biota composed of numerous individual metabolic
complexes would exhibit all manner of minute variations. But what of
replicative reproduction?
10.3.2.
Without replicative reproduction how can any change to an
organism be passed on to its progeny, generation by generation? We know
now that at the heart of replicative reproduction lies the genetic code,
based on a precise unvarying relationship between the constituent units
of proteins and nucleic acids. But such a code, or anything resembling
it, could not possibly have been incorporated into those elementary
metabolic systems with which, we are claiming, life on Earth began.
A first, primitive version of the genetic code finally evolved some
700 million years later. But since replicative reproduction is an essential
ingredient of organic evolution, and since it would seem to require,
in however crude a form, the genetic code, how could such an evolution
have possibly been achieved? There is only one answer worthy of consideration.
Although the genetic code is an essential ingredient of all replicative
reproduction above the most rudimentary level, it is not the only ingredient.
And it is owing to the operation of these other ingredients that a primitive
version of the genetic code was enabled to evolve. So we are claiming
that, like the other four evolutionary parameters, replicative reproduction,
if only as no more than a vague tendency, was operative right from the
formation of the Earth.
10.3.3.
Our five basic evolutionary parameters are far from being mutually exclusive.
They are simply five outstanding general features of the one evolutionary
process, and, as such, overlap in their effects. In this particular
case it is, above all, mnemic causation, present from the formation
of the Earth, which contributes so decisively to replicative reproduction.
As we saw in Chapter 9, mnemic causation, based upon sympathic association
between past and present experience, is, at bottom, no more than a question
of the physical forces which contributed to past rhythmic unities, modifying
present physical forces in such way as to reproduce these unities in
the present. Its whole general tendency is thus to reproduce past order
in the present. It is therefore essentially replicative in nature. It
is also essentially constructive, not, as we have pointed out, because
it contains some “urge to greater unities” but only because,
through its ability in the face of general dissipative tendencies to
maintain an established unity throughout a succession of individual
organisms, it provides a basis upon which slightly more complex unities
can arise – which, in turn perform the same role for unities yet
more complex, and so on indefinitely. This effect of mnemic causation,
at once both constructive and replicative, is present from the start
of life on Earth; of course, since, without it, life could never have
arisen. Before attempting to reconstruct the role mnemic causation must
have played in precellular evolution, we shall first glance at the central
feature of the role it plays in morphogenesis.
10.3.4.
The essential role of the genome in morphogenesis is to enable the cellular
– fundamentally proteinic – processes to produce a new protein
at exactly the time and place it is needed in the whole developmental
process. And what triggers any such instance is the metabolic context.
For orthodoxy, this context is, of course, purely physical. For us,
it is both physical and psychical, having in addition a past component
at least as important as the physical present. This past component,
as we have just seen, is essentially a tendency – embodied in
mnemic causation – to replicate the past. In the normal healthy
embryo, these two components operate, of course, in near-perfect constructive
collaboration. When the embryo is interfered with, these two operational
contexts are no longer harmonious, and then, what the two produce between
them may no longer make biological sense. But in the normal course of
nature such interference is reduced to a minimum, the whole morphogenetic
process taking place in an undisturbed, nutrient rich medium provided
by egg or womb.
10.3.5.
Here, we are concerned only with the evolution of the genetic code up
to the arrival of its first crude version in around 3800 m.y.b.p. The
morphogenetic process with which science is almost wholly concerned
is that of multicellular organisms. But the first multicellular organisms
did not appear on Earth until some 800 m.y.b.p. But stiil, the latter
can legitimately be viewed as a more elaborate version of the former,
the role of the genome being just the same: to provide proteins as and
when they are needed at that dual developmental context – composed
of both physical and mnemic forces. This general situation applies essentially
throughout, and we are concerned here only with its first, rudimentary
version. It is perhaps worth mentioning here the great steps taken along
the evolutionary pathway and the times at which they occurred.
Formation of planet Earth: c. 4600m.y.b.p.
Advent of the first primitive cell, complete with rudimentary genome:
c. 3800 m.y.b.p.
Advent of photosynthetic bacteria: c. 3200 m.y.b.p.
Development of aerobic photosynthesis: c. 2300 m.y.b.p.
Oxygenation of atmosphere: c. 2000 m.y.b.p.
Advent of unicellular eurkaryotes: c. 1400 m.y.b.p.
Development of sexual reproduction: c. 1200 m.y.b.p.
Advent of first multicellular organisms: c. 800 m.y.b.p.
Advent of first elaborate multicellular organisms: c. 600 m.y.b.p.
10.3.6.
Our immediate task, then, is to show how, on the primordial Earth, this
essential morphogenetic situation of a primitive genome evolved via
successively closer approximations. But in order to render such an account
intelligible we must first point out certain basic biochemical conditions
which had to be fulfilled in order that there might evolve ever more
complex syntheses of metabolic processes. Proteins are by far the most
important macromolecular constituents of living organisms. So that the
evolution of life centrally involved the emergence first, of amino acids;
then their polymerisation into chains of varying length, known as polypeptides,
the longest and most elaborate of which are the proteins. These polypeptides
evolved within increasingly elaborate self-sustaining systems of metabolic
processes of which they were (and still are) the principal macromolecular
constituents. All the other macromolecular constituents of metabolic
systems are intimately dependent upon polypeptides, more particularly
proteins, both for their advent, either by manufacture or incorporation,
and their subsequent functioning.
10.3.7.
Now, as is well known, this whole process of polymerisation is endergonic
(energy requiring). At the outset of organic evolution – the synthesis
of the monomers themselves - the energy source was physical: ultra violet
light, lightning, volcanic activity, etc. But this would be far too
crude and sporadic for elaborately organised biochemical systems. At
this level, the energy of polymerisation is obtained from coupled exergonic
(energy yielding) chemical reactions. Today, throughout virtually the
whole organic world, the energy of polymerisation and constructive metabolism
generally is obtained from the coupled breaking of one of the two terminal
“high-energy” pyrophosphate bonds of ATP (adenosine triphosphate).
What is effectively a water molecule (H2O) is lost by the
two monomers – OH from one and H from the other; and in an essentially
two-step process involving a double-headed intermediate, the energy
for this endergonic condensation is supplied by the exergonic hydrolysis
of the high-energy pyrophosphate bond, the OH becoming attached to one
of the phosphate groups, and the H to the other. But this presupposes
an already well-advanced biochemistry. So that a gulf of some hundreds
of millions of years lies between the primordial stage where the energy
for biosynthesis was supplied by physical energy sources, and the evolutionary
level when it was first supplied by ATP. This implies a long period
when metabolism must have depended on more chemically elementary sources
of energy.
10.3.8.
By far the most outstanding candidate for this role is the thioester
bond: that between thiols (R' - SH) and carboxylic - more particularly
amino – acids (R-COOH). Between the reactants a ‘water molecule’
(H + OH) is lost, and a highly endergonic bond (S~C) is formed between
the sulphur and carbon atoms. Bacteria exist, even today, where peptides
are synthesised from amino acid thioesters. The main objection to the
thioester bond fulfilling this role is its highly endergonic nature.
However, in recent years, this objection has lost much of its cogency
with the discovery of deep-sea hydrothermal vents, situated as these
are in hot highly acidic, sulphur-rich waters closely surrounded by
much colder water rich in the molecular constituents of life2.
These vents are now seen as constituting one of the most favourable
kinds of birthplace for life on Earth. The main evolutionary role of
the thioesters was essentially the same as ATP at a later date: to supply,
in coupled reactions, by the hydrolytic breaking of the thioester bond,
the energy of polymerisation through condensation (effectively the loss
of a water molecule) between two amino acids. So we are suggesting that
the principal energy source for all the ongoing polymerisation of the
metabolic systems that constituted primordial life was the S~C thioester
bond.
10.3.9.
The principal components of these metabolic systems would have been
peptides of a wide range of polymerisation, though all short in comparison
with today’s proteins. We must never lose sight of the fact that
the basic nature of living systems is that they are not producing anything
but themselves: each part, qua part, is a means to an end, that end
being the maintainance of the whole system of processes in the face
of an environment whose constant tendency is to erode it. The role of
sympathic association and mnemic causation in all this is fundamental.
What are being sympathically associated are experiences – each
of which is a unity, and therefore something more than the merely arithmetical
sum of its parts. Mnemic causation, the force making for the reemergence
of the past in the present proceeds from such experiential unities.
It is the life force, since, without it there would be no life. Each
polypeptide fulfilled some vital function in the whole system. Preeminent
among such functions was that of organic catalyst (enzyme). All the
chemical reactions within the system were mediated by enzymes, the enzyme
attaching to both reactants in such way as to render their reaction
as smoothly efficient as possible. Moreover, as parts of the whole,
they responded mnemically to the dictates of the state of the whole
– effecting the reaction or not effecting it as the case might
be.
10.3.10.
We are now in a position to return to direct engagement with our theme
of replicative reproduction in the epoch before even a rudimentary version
of the genetic code had evolved. Our first concern must be with those
parameters, if any existed, which made for replicative reproduction,
and hence the preservation of systemic complexity, after the demise
of the individual systems which possessed them. Clearly, these must
have centred upon mnemic causation, whose very nature it is to preserve
or bebuild the past in the present. Now, in the necessary absence of
bacteria, the death of a particular metabolic system from whatever cause
must have been of a very different nature from the death of a higher
organism. Presumably, causes of death were the exhaustion of customary
sources of nutrients (individual hydrothermal vents possessed only a
limited life span); sudden cataclysms born of seismic activity or the
impact of large extraterrestrial bodies; poisoning by environmental
increase of destructive substances; sudden large changes in ambient
temperature; and, constituting a special case3, ingestion
by some other metabolic complex. Under such modes of assault the metabolic
system would have been radically disrupted, with disintegration of sorts
setting in. In a way strange to us, these fragments of disintegration
would also have played the role of germ cells. The fragments would have
consisted of complexes of polypeptides, and the more such fragments
resembled earlier stages of the disintegrated synthesis, the more likely,
if the environment were favourable, they would be to grow back, under
the influence of mnemic causation, to this original synthesis. For a
new metabolic system to develop it would not have been necessary to
begin all over again from scratch, at the monomeric level. Other things
being equal, the more rapidly, accurately and abundantly a metabolic
system, on disintegration, could grow back into what, in all structural
and functional interrelationships, is effectively itself, the greater
the chance for Darwinian selection to operate in favour of the multiplication
of instances of that particular system.
10.3.11.
Such being the case, there are a number of obvious ways in which this
relation between metabolic system and offspring might evolve. All of
these would be changes in the parent ‘organism’ such as
to favour accuracy, rapidity, and abundance of offspring. Perhaps the
first to evolve would be a parent metabolic system which tended to disintegrate
in such way as to include among the fragments some earlier stage or
stages of its own development. The whole tendency of mnemic causation
would then be to repeat the subsequent developmental stages, especially
if environmental conditions were similar – in effect, to reproduce
the parent complex. The next evolutionary step would be not to wait
for the fragmentation of ‘death’ but for the metabolic complex
to develop in such way that from time to time some part of it constituting
an earlier stage of development would ‘bud off’ from it
and so lead an independent life, governed as always by mnemic causation
with its tendency to reproduce past experience - an experience being
some particular ordered process. Finally, there would be no need for
a parent organism to restrict itself to one such offspring; a whole
type of metabolism could evolve which included such continuous production
of offspring as among its principal activities. We envisage that it
was at about this evolutionary stage when nucleoside base pairing, with
its potential for carrying replicative reproduction to an altogether
higher order of accuracy ubiquity and complexity entered the situation.
EVOLUTION
OF REPLICATIVE REPRODUCTION
SECOND STAGE
10.4.1.
The genetic code consists essentially of very precise relationships
between the monomeric constituents of proteins and those of nucleic
acids. Our metabolic complexes are essentially composed of polypeptides
– which may be regarded as the same as proteins in all respects
save only in the lesser number of amino acids composing them. But how
did nucleic acids enter these systems? We have already noted that virtually
all the energy of the living world is supplied by the hydrolysis of
one or more of the phosphate bonds of ATP. But ATP – adenosine
triphosphate – derives directly from adenosine diphosphate (ADP),
which, equally directly, derives from adenosine monophosphate (AMP).
The difference is just whether one, two, or three phosphate groups are
serially attached to the adenosine. And adenosine itself is composed
of the nucleobase adenine attached to a molecule of the sugar, ribose.
Furthermore, all this applies equally to the other three nucleobases
involved in the structure of ribonucleic acid (RNA): that is, the other
purine, guanine, and the two pyramidines, cytosine and uracil. And although
adenine is that one of the four most engaged in general metabolism,
the other three are also significantly involved; and indeed, might just
as well as adenine, have assumed the primary role in energy production.
Indeed, at the much earlier stage of evolution with which we are presently
concerned, the role of energy producer might well have been more equally
shared among them than it is today.
10.4.2.
So we see that there is much in common between the molecules most intimately
involved in energy production in living organisms, and those involved
in replicative reproduction. We suggest, then, that it was in the first
role, as providing an energy source superior to the thioester bond,
that the phosphates and the nucleobases first became an intimate part
of metabolic complexes, entering into all manner of constructive processes
with the major constituents of these complexes - the polypeptides. All
this provided an essential functional preliminary to the emergence of
their second and momentous role of raising replicative reproduction,
with its high responsiveness to Darwinian selection, to an altogether
higher order of complexity.
10.4.3.
One major difference between the polymerisation of nucleobases (RNA)
and that of amino acids (proteins) is that, whereas in the proteins
each amino acid unit is directly bonded to both its neighbours, this
is not the case for RNA. Here, the bonded units of the polymeric chain
are not the bases at all. Instead, the chain is composed of alternating
units of ribose and phosphate molecules. Each molecule of ribose is
bonded to a base. However, the double strand is formed by bonds between
bases, with each purine bonded to a pyramidine: adenine⇔uracil
and guanine⇔cytosine. This, of course, is the complementary base
pairing on which this whole vast contribution to replicative reproduction,
and hence to biological evolution, ultimately depends. It means that,
given a single strand of RNA, a complementary strand can form on it.
Then, when this is unravelled into two single strands, a complementary
strand can form on both of these, giving two versions of the original.
And so on, doubling in number at each generation. This is the first
root process of precise replicative reproduction, but its great importance
lies not in itself but only in its providing a necessary basis for a
second process. This is the rigidly unvarying attachment of amino acids
to small groups of the nucleobases composing RNA, such that the precise
replication of these also entails the precise replication of the amino
acids attached to them. So that monomer by monomer the polypeptides
can be as precisely replicated as the RNA strands themselves. Hence,
the replicative properties of RNA are supremely important for life,
not because they replicate RNA, but only because, through simple association,
they allow of the replication of polypeptides - ultimately, of proteins.
10.4.4.
Needless to say, although this process could hardly be simpler in its
general nature, in detailed feature it is far from simple. But most
of its details are now well known – in so far, that is, as anything
conceived in mechanistic terms can be said to be well known –
and it is no part of this work to present them in noumenal terms in
systematic detail. It is enough to understand that mnemic causation
is operative throughout, maintaining orderly process on a level of complexity
utterly beyond the powers of physical forces as conventionally understood,
to create and sustain. Because of the strong selective pressures to
which the replicative reproduction of any species of organism must be
subject, there is every reason why this RNA – polypeptide relationship
should evolve in terms of precision, rapidity, abundance and complexity,
from its first crude beginnings some 3800 m.y.b.p. to the DNA –
protein relationship constituting the genetic code as we have it today.
Again, we shall make no attempt to trace this evolution, which is best
left to the specialist. We simply contend that any empirical facts unearthed
by the scientists can only be grasped as contributions to a rationally
ordered system of processes within a context of sympathic association
and mnemic causation. However, before we leave this topic, there are
one or two general observations which should perhaps be made.
10.4.5.
In the polypeptide-RNA relationship it is overwhelmingly the polypeptide
which is the active partner, and the RNA the passive. The whole replicative
process is organised and monitored by enzymes. And if our contention
is correct that all the constituents of RNA gradually infiltrated a
metabolic system all of whose catalysts were polypeptides and many of
whose substrates contained amino acids as prominent constituents, this
is just what we would expect. In any case it is precisely the unchanging
nature of RNA which, second only to its base pairing propensities, makes
it an ideal material basis for replicative reproduction. The assembly
of RNA from its constituent ribose, phosphate, purine, and pyrimidine
molecules is effected entirely by enzymes. Most interesting of all,
each step in the translation of RNA into protein is initiated by an
ezyme known as tRNA synthetase. The substrate to which this binds is
one of the twenty out of hundreds of amino acids that are constituents
of protein. It then attaches this amino acid to a relatively short length
of RNA, known as a transfer RNA (tRNA) molecule, containing the complement
of one of the amino acid’s codons (anticodon). This engages with
the codon, whilst simultaneously the amino acid is being bonded into
its corresponding place in the growing peptide chain. But there is some
hard evidence4 that it is the more detailed nature of this
process which determines why only twenty – not necessarily the
most common – of the hundreds of amino acids that exist, are constituents
of protein. It seems that the tRNA under the influence of the tRNA synthetase
folds up in such a way that the anticodon, the fourth nucleotide from
the 3’ end (acceptor stem) of the tRNA molecule, and the tRNA
synthetase form a pocket into which the substrate amino acid fits. Subsequent
research has produced some evidence that no amino acid which is not
a constituent of protein will fit into such a pocket. Since twenty amino
acids are sufficient to give rise to a human being, life doesn’t
seem to have been seriously affected by this limitation.
NOTES
1 m.y.b.p. stands for million years before present.
2 A significant proportion of these dissolved nutrients would have come
from outer space as ingredients of comets and meteoroids.
3. Because sometimes, far from dying – that is, disintegrating
- the ingested complex begins a new and viable life as an endosymbiont
within the ingesting organism.
4. See: Shimizu M. Molecular
basis for the genetic code. Journal of Molecular Evolution 18:297-303;
1982
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