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.