Epigenesis is the complex process by which
organisms are formed from a single cell. It is a continuation of
cosmic development to produce an individual living organism, which
is organised as a three-dimensional strucure of cells and organs
that are discrete from the environment only so long as it is alive.
When it dies it comes into chemical and energy equilibria with the
cosmos. This does not mean that while alive it is cut off from its
environment by an impenetrable barrier. On the contrary, it is very
much dependent on its environment for its continued existence and
the continually exchange matter and energy between the cosmos and
itself.
It is from its environment that a living organism
obtains those chemical substances needed for repair and the
replacement of worn-out parts, for growth and for
reproduction.
Some organisms — such as green plants
— are able to use, and to accumulate the energy of sunlight,
while others — such as all animals — rely on that
liberated during the chemical reactions involved with the breakdown
of food for all their energy needs. This energy, from whatever
source, is needed to drive the chemical reactions of building, as
well as for movement and the transport of chemicals within the
organism, and between it and its environment, across the
intervening boundaries of cells and the body.
The basic unit of which living organisms are
composed is the cell, a structure typically only micrometres in
diameter yet capable of carrying out the most elaborate chemical
processes. The simplest organisms living independently of others
are the bacteria, but their cells are less complex than those of
the simplest plants and animals.
The most complex plants and animals are composed
of many millions of cells and these are differentiated into a
number of cell types concerned with particular functions.
Living organisms generally respond to
environmental stresses in ways which prolong their survival. Thus,
for example, green plants grow towards the light, and animals move
away from extremes of heat and cold. In many cases these responses
are built into the nature of the organism; in others they are
acquired as the result of interactions with the environment itself
(nurture).
The constancy of the nature of different types of
organism is due to heredity, and is a consequence of the precise
replication of large chemical molecules —the nucleic acids -
which encode the instructions for the organism's development and
activities in the DNA chains comprising genes.
The nurture of an organism not only has a direct
effect on the expression of its heredity but also causes it to
become better adapted to its environment. Variation in the heredity
of related organisms results in some being better able to respond
to environmental stresses and these organisms are more likely to
survive and to become the parents of the next generation of
individuals (natural selection).
Progressive adaptation of the genes through
natural selection, with its concomitant elaboration of structure
and function, results in the evolution of new kinds of individuals.
It is in this way that the great diversity of living organisms have
originated from much more simply organised ancestors over the
millions of years that life has existed on this planet.
This theme may be developed by an account of the
chemical composition of living organisms, with descriptions of cell
structure and accounts of the simplest forms of free-living
organisms, the viruses, bacteria and blue-green algae. The
bioichemical aspects include an account of the synthesis of
proteins, and the formation of ATP, the energy currency of all
living organisms. This leads to a study of the nutrition in plants,
animals and other living organisms, and use to which the
assimilated food is put in the process of growth, reproduction and
development.
Development of an individual from a single cell
involves an increase in cell number and the subsequent
differentiation of the primary cell mass, that is, functional
specialisation of first cells. At first, the difference between the
cells is not great and regeneration of the whole embryo from a part
can occur, but, as development proceeds and the embryonic tissues
differentiate into adult tissues, this power of regeneration is
progressively lost. Thus, during the development of their offspring
the characteristics of the parents represented in the gametes by
replicates of their DNA molecules are progressively expressed. Our
knowledge of the processes involved in this cellular
differentiation is at present incomplete and speculative since it
is based on a small number of observations in a restricted number
of different organisms.
In complex organisms, an embryo develops from the
fertilised primary cell (zygote or spore) as the result of a series
of mitotic cell divisions. Although all the cells of an organism
formed in this way contain identical complements of genes, during
development morphological and physiological changes take place in
these clonal cells which result in the development of the many cell
types characteristic of plants and animals.
For example, the sea urchin zygote has a
characteristic band of pigmented granules in the cortex just below
the equator. The position of this definite band has enabled
embryologists to follow the distribution of the different regions
of the egg as the embryo develops. The first division of the
fertilised egg (cleavage) passes through both poles of, described
as the animal and vegetal poles, and divides the egg into equal
halves. The second cleavage is at right angles to the first and
also passes through the animal and vegetal poles and divides the
egg into equal quarters. The third is at right angles to the first
and the second in the horizontal plane just above the equator. The
embryo at this stage consists of four similar smaller cells called
mesomeres in the animal half and four similar larger macromeres in
the vegetal half. The inequality of the third cleavage is due to
the presence of more yolk in the vegetal half than in the animal
half. Yolk tends to impede cleavage. The greater amount of yolk in
an amphibian egg impedes cleavage in the vegetal region to an even
greater extent, and in reptilian and bird eggs, where the amount of
yolk is vast, there is no cleavage at all in the vegetal
region.
If a cell nucleus is planted into an egg from
which the nucleus has been removed (enucleate),of the same species,
normal embryos are produced, except when damage is caused to the
donor nucleus or recipient cytoplasm. In experimental work of any
kind, but particularly in biology where there are so many unknown
variables, it is absolutely essential to have controls which are
identical with the experimental individuals in every way except
that being investigated.
When a nucleus is taken from a hybrid frog embryo
formed in this way and put back, as it were, into an enucleate egg
of another species, an embryo is formed, but it is not viable. This
shows that the cytoplasm has had an effect on the nucleus. It is
also clear from such experiments that the cytoplasm is not inert
with regard to development. If it were inert, any nucleus would
function when in a foreign cytoplasm. If the cytoplasm of the egg
of a particular species were the same as that of a the donor egg
one would expect an egg containing a successfully transplanted
nucleus from another species would develop into a donor individual,
but it does not.
A large number of the failures are known to be
due to faulty experimental technique, and to imperfect replication
of the donor nucleus. The proportion of nuclei giving normal
embryos decreases as the embryos from which they are taken become
more differentiated. This shows there is a progressive loss of
potency of cells to take over the control of normal development.
These experiments also show there is no selective permanent
inactivation of genes as differentiation throughout development
proceeds. Specialisation involves the differential activity of
genes present in every cell of the organism.
Further discussion of cellular differentiation
requires some information about the mechanism of gene action
because not all genes are active at the same time in different
cells, and consequently, the characteristics of different tissues
are due to the activity of certain genes or groups of genes which
are switched on.
The switching on of genes has been extensively
studied in microorganisms. A culture of the bacterium E.
coli adapted to a medium containing glucose as its carbon
source will grow rapidly if the glucose and other substances
required for growth are present in optimal amounts. If the
bacterial cells are separated from the medium, to avoid damaging
the cells, and the nutrient medium is replaced by one containing
lactose instead of glucose, there will result an immediate
cessation of population growth because the cells are unable to
utilise the new substrate. However, after a short interval, growth
will resume. During the delay period genes are switched on to form
the mechanisms to utilise lactose.
In more complex organisms cellular specialisation
and embryonic differentiation are very precise processes.
Development proceeds along definite pathways, with forks controlled
by switch mechanisms, at which differentiation occurs. The
presumptive areas of the embryo differentiate into the basic
tissues which then undergo further and further differentiation
until the adult tissues are formed.
Regulation of the formation of organs systems
that define a species requires that the genotype should remain
unchanged from generation to generation. In biochemical terms, we
mean that the proteins determined by the genotype's constituent
codes should have an invariable amino acid composition. This is
only possible if the coding mechanism is essentially conservative
and resistant to environmentally induced changes.
Nevertheless, in some instances it is
advantageous for an organism to conform to a stress. Galactosidase
induction in Escherichia coli is an example of this. When an
organism does conform it is better that it should do so in an
optimal way rather than in a variable one. When this bacterium is
placed in an environment containing a new food medium it is
advantageous that it should produce the appropriate enzyme in the
correct quantity for optimum substrate utilisation. Those variant
bacteria which can do this best will be more likely to survive,
grow and multiply than the others. Should a genetic change in the
mechanism controlling enzyme activity resulting in optimal
substrate usage occur in these selected bacteria, it will be
immediately assimilated into the genetic system of the population
because of its advantage to the individuals possessing it.
In higher organisms, genetic adaptations to
changes environment through natural selection begin by variations
in embryonic development that may result, for example, in a
slightly larger beak, a faster or slower rate of growth.
Ageing is a continuation of the processes of
development. These changes result from the accumulation of errors
in the workings of genes that eventually result in the organism
being unable to adapt to environmental stress.
The cellular controls of epigenesis are expressed
at the level of size and shape of cell populations. In this sense
evolution is limited by what has already been selected in terms of
the forms and organ structures generated during the process of
epigenesis. Emergent order is generated by distinctive types of
dynamic process in which genes play a significant but limited role.
Epigenesis as a process is the source of emergent evolutionary
properties, and it is the absence of a theory of organisms that
includes this basic generative process that has resulted in the
failure to account for the origin of the emergent characteristics
that identify species.
Many people have recognized this limitation of
Darwin's vision. It was the achievement of D'Arcy Thompson in his
volumes On Growth and Form (1917) in which he
single-handedly defines the problem of biological form in
mathematical terms and re- establishes the organism as the dynamic
vehicle of biological emergence. Once this is included in an
extended view of the living process, the focus shifts from
inheritance and natural selection to what Brian Goodwin has
described as creative emergence of high level structures as
the central quality of the evolutionary process. And, since
organisms are primary loci of this distinctive quality of life,
they become again the fundamental units of life, as they were for
Darwin. Inheritance and natural selection continue to play
significant roles in this expanded biology, but they become parts
of a more comprehensive dynamical theory of life which is focused
on the dynamics of emergent processes.
According to Goodwin, the consequences of this
altered perspective are considerable, particularly in relation to
the status of organisms, their creative potential, and the
qualities of life. Organisms cease to be simply survival machines
and assume intrinsic value, having worth in and of themselves, like
works of art. Such a realization arises from an altered
understanding of the nature of organisms as centres of autonomous
action and creativity, connected with a causal agency that cannot
be described as mechanical. It is relational order between
components that matters more than material composition in living
processes, so that emergent qualities predominate over quantities.
This consequence extends to social structure, where relationships,
creativity and values are of primary significance. As a result,
values enter fundamentally into the appreciation of the nature of
life and biology takes on the properties of a science of qualities.
This is not in conflict with the predominant science - of
quantities - but it does have a different focus and emphasis.
Darwinism sees the living process in terms that
emphasize competition, inheritance, selfishness, and survival as
the driving forces of evolution. These are certainly aspects of the
remarkable drama that includes our own history as a species. But it
is a very incomplete and limited story, both scientifically and
metaphorically, based upon an inadequate view of organisms; and it
invites us to act in a limited way as an evolved species in
relation to our environment, which includes other cultures and
species. These limitations have contributed to some of the
difficulties we now face, such as the crises of environmental
deterioration, pollution, decreasing standards of health and
quality of life, and loss of communal values. But Darwinism
short-changes us as regards our biological natures. We are every
bit as co-operative as we are competitive; as altruistic as we are
selfish; as creative and playful as we are destructive and
repetitive. And we are biologically grounded in relationships which
operate at all the different levels of our beings as the basis of
our natures as agents of creative evolutionary emergence, a
property we share with all other species. These are not romantic
yearnings and Utopian ideals. They arise from a rethinking of our
biological natures that is emerging from the sciences of complexity
and is leading towards a science of qualities which may help in our
efforts to reach a more balanced relationship with the other
members of our planetary society.