Epigenesis
in relation to evolutionary phenomena
According to
Piaget:
"...
biologists today, like Julian Huxley and Waddington with their "synthetic
theory" of species, are making phylogenesis depend in part on ontogenesis, and not
only the inverse. Indeed genes are not actually static elements but, rather factors
identical or analogous to enzymes, whose nature is revealed by their activity,
interdependently and subject to a whole set of regulations throughout the entire process
of embryogenetic growth in interaction with the environment. The result of this is that
the information supplied by the genotype is not only transmitted but also transformed in
the course of all this development, so that the essential system is no longer the genotype
in isolation but a total epigenetic system."7
Thus, even
on the genetical level, the notions of interactionism and constructivism - which Piaget
considers characteristic of epistemological activity in general - are thought to apply.
Instead of treating the genotype in isolation - which had been the tendency of classical
mutationsits - and as something mysteriously removed from the observable phenotypic
variations in a given race or species, modern genetics construed the collection of
genotypes as a relational totality or genetic system (known as a genome) in which there
were genes capable of performing different, but complementary, functions of a structural
and regulatory nature.
In other
words, some of the genes were of the traditional sort, so to speak, and conveyed or
transmitted structural information concerning such things as the physiological or
morphological properties of different biological dimensions of a given organism. On the
other hand, there were other genes which were concerned with regulating the organizational
properties of the genetic system as a whole - both in terms of the way structural genes
transmit their information, as well as in terms of the way in which the genetic system
responds to, and interacts with, the environment. Considered as a whole, structural and
regulatory genes represent a series of processes that encompass a range of qualities (such
as transmission, variation, deep structure and surface structures) said to be
characteristic of genetic phenomena.
In addition
to conceiving of the hereditary mechanism as a dynamic, interacting totality, Piaget also
describes how there is a growing tendency - to which he subscribes - in modern biology,
when considering the issue of the basic genetic unit, to switch the emphasis, or focus,
from the genome to the genetic pool or population.
Attendant to
this switch in focus, from the genotype to the genetic pool, there is also a change in
emphasis concerning the "reaction norm". The reaction norm refers to
the set of possible phenotypes that can be generated from a genotype - given variations in
the environment to which the genotype responds by way of altering the genetic system
(either in a structural or regulatory manner) such that phenotypic variations subsequently
appear. With the aforementioned change in emphasis and focus in certain facets of
biological investigation, the notion of "reaction norm" (while still appropriate
and meaningful in the context of individual genotypes) also came to be used as a way of
referring to the set of phenotypic possibilities represented by the mixture of genotypes
existing in any given population.
Seen from
the perspective of the interaction of genomes and population, evolution consists of the
complex series of transformations that occur on two levels - each of which affects the
other level. These levels are: (1) the responses of the individual genotype to the
surrounding environment - which itself represents a contribution to the genetic pool; (2)
the responses of the population as a whole to the surrounding environment, which includes
the influence of a variety of structural and regulatory possibilities generated internally
by individual genotypic reaction norms within the population.
Moreover,
just as there is a set of equilibration principles which co-ordinate the activity of the
genome or collection of genotypic possibilities within the hereditary mechanism of a given
organism, so too, according to Piaget, there is a set of organizational principles that
are auto-regulatory in nature with respect to the genetic pool as a whole. For Piaget,
this set of organizational principles represents both the source, as well as the result,
of evolution in general. (e.g., see his discussion on pages 278-284 in Biology and
Knowledge).
Using the
genome/genetic pool interactionist approach outlined above, Piaget plans to analyze what
he considers to be the three possible ways for theoretically accounting - in evolutionary
terms - for the relationship between organism and environment. These three ways are:
"(1)
environment takes control of the organism and molds it throughout its working existence,
affecting even its hereditary structures, which easily submit to its influence;
"or (2)
it is the organism that imposes certain independent hereditary structures on the
environment, the environment merely eliminating such structures as prove unsuitable or
nourishing those it finds congenial;
"or (3)
there are interactions between organism and environment such that both factors remain on
an equal footing of cooperation and importance."8
However,
since Piaget is primarily interested in establishing the biological underpinnings of
epistemological processes, he also needs to draw up a corresponding set of possibilities
with respect to the sorts of biological structures and functions (which, presumably, can
be accounted for by one, or more, of the three foregoing theoretical frameworks) in which
knowledge can be rooted. Thus:
"There
are, in effect, three possible kinds of knowledge: (1) the kind that is linked with
hereditary mechanisms (instinct, perception), which may or may not exist in man but which
correspond in biological terms to the sphere of characteristics transmitted by the genome;
(2) knowledge born of experience, which thus corresponds in biology to phenotypic
accommodation; and (3) the logico-mathematical kind of knowledge which is brought about by
operational co-ordinations (functions, etc.) and corresponds, in biology, to regulation
systems of any scale..."9
However,
before Piaget discusses these kinds of knowledge, he wants to lay the foundations for them
by demonstrating how an epigenetic approach can provide a tenable means of accounting for
evolutionary development . If successful, then, epigenetic principles also might serve as
a prototypic example, in general, for explaining a large variety of transformational
processes - including the transition from organic structures and functions to cognitive
systems.
When Piaget
mentions evolutionary theory in the context of the sort of organism/environment
relationships that can be construed in terms of the environment's taking control of the
organism, he, generally, has in mind: (a) Lamarck's (or a Lamarckian-like) theory in which
the exercise of various organs during the course of development influenced the direction
and nature of such development; (b) the way in which the changes brought about by the
exercising of organs during development were fixed in hereditary as acquired
characteristics; and, (c) the manner in which the environment - in terms of its physical
properties - forced or selected the exercising of particular organs that led to the fixing
of such hereditary characteristics.
For Lamarck,
the biological organism was, largely, a passive or malleable entity that was molded
according to the nature of the influences and pressures existing in the different features
of the immediate environment and the effect such forces had on various organs in a given
organism. Although Piaget does acknowledge the importance of the role played by the
environment in modifying the hereditary mechanism and credits Lamarck for having seen its
significance in the evolutionary context, he nonetheless criticizes Lamarck for, among
other things, having overestimated the passivity (or having underestimated the activity)
of the biological organism in bringing about evolutionary change - which is to say, that
even though the organism is influenced by external influences, such influences are
assimilated to various action schemata that exist prior to the organism's present
encounter with the environment.
Furthermore,
Piaget notes that on the basis of Lamarck's emphasis on the organism's supposed passivity,
one would tend to expect the organism to be malleable to an indefinite extent according to
the direction and nature of the environmental factor which was affecting the modification
in, say, a given organ's functioning. Yet, nowhere, according to Piaget, does one see
evidence of such malleability.
Instead, one
finds that, generally speaking, the limit situation with respect to malleability is a
function of the reaction norm that is, it is a function of the range of phenotypic
possibilities associated with both the individual genotype as well as the population
consisting of mixed genotypes. Consequently, as far as Piaget is concerned:
"...
what is lacking in the Lamarckian interpretation is the explicit recognition of the fact
that the effects of these exercises (i.e., of the organs) are always relative, not only to
the environment, but to the genotypic structure (pure or impure) of the lineages being
studied. To sum up, where Lamarck sees nothing but the effect of environment... there are
really interactions between external factors and the genome."10
At the
opposite extreme from the Lamarckian penchant for emphasizing the significance of the
environment in effecting evolutionary change, is the tendency of those theorists who play
up the importance of an organism's endogenous factors in bringing about such change. For
Piaget, the most notable of such theories is the mutationist school of thought.
According to
this approach to evolution, genotypes are generally considered to be static or invariable
hereditary structures except in instances in which variations (due to certain factors
internal to the organism) in these structures are introduced on a, supposedly, random
basis. The term for such variations is, naturally, mutation.
Moreover,
the classical mutationists contend that the role of environment as an evolutionary agent
is after the fact and, consequently, has no real evolutionary significance. In other
words, since evolutionary change is a function of random variations or mutations and the
role of the environment is restricted to a post facto selection of certain of these
variations which it favors, evolutionary transformation per se is, in the mutationist
conception, unrelated to environmental considerations and dependent only on the random
variations.
One
objection Piaget has to the mutationist position is that such a theory tends to treat the
hereditary mechanism in an atomistic fashion. Thus, genes are considered to be so many
separate boxes that open up only on those occasions when it releases a new mutation which
its internal processes have mysteriously produced and immediately thereafter seals itself
off from external influences.
This
atomistic conception of the gene effectively isolates the gene from all possibility of
interaction with the environment. As a result, one neglects the considerable evidence
compiled by Waddington, Julian Huxley and others concerning the complex relationship
between genome and environment that is said, by such investigators, to be responsible for
evolutionary change.
Moreover,
the mutationist view is in considerable contrast to the cybernetic features which Piaget
has argued are characteristic of biological phenomena on every level - from the hereditary
to the cognitive. Therefore, for Piaget, the mutationist position appears strangely
inconsistent with what seems to be a persistent feature of biological activity in general.
In addition
to the foregoing sorts of objections to the mutationist position, Piaget cites another
kind of argument. much later in Biology and Knowledge during a discussion of
instinctual phenomena. Piaget argues:
"But if
it is not considered in any way incompatible with survival that evolution should wait some
thousands of centuries to endow a horse with a tail and a mare made of hairs and not
feathers, it becomes rather difficult to envisage how long it would take to ingrain the
instincts of reproduction, nest-building, and so on among species whose very existence
depends on cognitive precision in relation to those instinctive mechanisms. We need only
take one example the eye in vertebrates'.11This is not indispensable as a means
of acquiring knowledge but it is indubitably useful. Bleuler's calculations showed that if
the mutations necessary for the formation of this organ had been brought about
simultaneously or co-jointly, they would have had a probability of only 1 in 1012,
in other words, practically none. On the other hand, if it had been a question of
successive mutations, in which new ones were simply added to preceding mutations so that a
cumulative effect was achieved, then it would have taken as many generations as would
correspond to the age of the world or even exceed it."13
Of course,
Piaget acknowledges that such calculations are dependent on a number of variables (such as
the age of the' earth, rates of mutation, population sizes, and so on) that have14been
assigned numerical values based on a certain amount of approximation and guess work.
However, even if one were to dramatically readjust the calculations in the
direction of estimations more favorable to the mutationist position'' (which may not be at
all justifiable except in a logically heuristic sense) and, thereby, reduced the
probability to, say, 1 in 1015, one is still talking about a very, very small
probability.
While one
might be willing to admit the logical, and even empirical, possibility of such a small
probability of occurrence becoming a reality on occasion, the boundaries of credulity are
distorted beyond recognition or reasonableness at the suggestion that numerous events of
such small probability should have happened with such frequency. Indeed, seemingly, one is
forced to seriously consider the possibility that such a combination of happy coincidences
might be the result of operative factors other than random mutations.
Any position
that arrives at a conclusion contrary to the foregoing appears to be demanding that the
individual take a great deal on faith. To believe that such a series of random
coincidences occurred on a regular basis appears to be a rather far-fetched exploitation
of the notion of logical possibility, and, as such, is far removed from any kind of
empirical evidence capable of pointing to its 'likelihood' or plausibility.
In my
opinion, however, the whole population genetics movement is seriously misguided to the
extent it attempts to supplant the individual in favor of the population. The
possibilities inherent in the population are little more than what is given to it by the
individuals of the population in both a negative and positive sense - "negative"
referring to the limitations which are inherent in a given species and
"positive" referring to the flexibility and scope of possibilities inherent in
the species.
In either
case, it remains to be seen how the population, per se, can generate anything new that was
not already inherent in the contributions of the various individuals to the genetic pool.
Mutations may cause the appearance of something new but mutations appear, on the one hand,
independent of the population concept and, on the other hand, mutations only can bring
about a change in terms of acting upon the givens and, therefore, the change is going to
be functionally related to, and not independent of, what it acts upon.
In short,
the population cannot produce something totally outside the structures on which it works.
Population is limited to the possibilities of modification at hand which are defined by
the nature of the potential inherent in the organisms which give expression to the
population.[Back to Text]
As such,
natural selection does not represent a systematic way of exploiting mutations. What works,
remains, and what does not work, disappears.[Back to Text]