INTRODUCTION.
What is development? Does it imply preformation or epigenesis? This perplexing question of biology has reappeared recently as a problem of the day. Of late years there have been set forth contradictory doctrines, each seeking to explain the process by which the fertilised egg-cell, an apparently simple beginning, gives rise to the adult organism, which often is exceedingly complicated, and which has the capacity of producing new beginnings like that from which it itself arose.
The opposing views of to-day were in existence centuries ago, and they are known in the history of science as the theory of preformation or evolution, and the theory of epigenesis. That most of the great biologists of the seventeenth and eighteenth centuries were decided upholders of evolution was the natural result of the contemporary knowledge of facts. For they knew only the external signs of the process of development. All they saw was the embryo becoming adult, the bud growing out into a blossom, as the result of a process in which nutrition transformed smaller to greater parts. And so they regarded development as a simple process of growth resulting from nutrition. Their mental picture of the germ or beginning of an organism was an exceedingly reduced image of the organism, an image requiring for its development nothing but nutrition and growth. That the material eye failed to recognise the miniature they attributed to the imperfection of our senses, and to the extreme minuteness and resulting opacity of the object.
That it might satisfy our human craving for final causes, the theory of preformation had to be accompanied by a corresponding explanation of the origin of the miniatures. Biologists had already abandoned the error of such spontaneous generation as the origin of flies from decaying meat, and, in its place, had accepted the doctrine of the continuity of life, formulating it in the phrase, Omne vivum e vivo (Each life from a life), and in the similar phrase, Omne vivum ex ovo (Each life from an egg). One creature issued from another, within which it had lain as a germ, and the series was continuous. Thus, the theory of preformation gave rise to the conception that living things were a series of cases or wrappings, germ folded within germ. The origin of life was relegated to the beginning, at the creation of the world: it became the work of a supernatural Creator, who, when He formed the first creatures, formed with them, and placed within them, the germs of all subsequent creatures.
To reckon at their proper value the theory of preformation, and, still more, the doctrine of enfolded germs, the standard of appreciation must not be the present range of our knowledge. They must be viewed historically, in the light of the knowledge of these days.
Nowadays it is not so much pure reason as a wider empirical knowledge of nature, with its consequent transformation of ideas, that makes the doctrine of enfoldment difficult. Abstract thought sets no limit to smallness or greatness; for mathematics deals with the infinitely small and with the infinitely great. So long as actual observation had not determined the limits of minuteness in the cases in question, there were no logical difficulties in the doctrine of enfolded germs. The biology of earlier centuries had not our empirical standard. What appeared then to be a simple organic material we have resolved into millions of cells, themselves consisting of different chemical materials. The chemical materials have been analysed into their elements, and chemistry and physics have determined the dimensions of the ultimate molecules of these. It is only because the minute constitution of matter is no longer a secret to us that the theory of germ within germ now touches the absurd.
It was very different in earlier days; the acutest biologists and philosophers were evolutionists, and an epigenetic conception of the process of development could find no foothold alongside the apparent logical consistency of the theory of preformation.
Wolff's Theoria Generationis (1759) failed to convince his contemporaries, because he could bring against the closed system of the evolutionists only isolated observations, and these doubtful of interpretation; and because, in his time, on account of the rudimentary state of the methods of research in biology, men attached more importance to abstract reasoning than to observation. His effort was the more praiseworthy in that it was observation bearing witness against abstract and dogmatic conceptions. By means of actual observation he tried to expose the fallacy in preformation, to show that the organism was not fully formed in the germ, but that all development proceeded by new formation, or epigenesis; that the germ consisted of unorganised organic material, which became formed or organised only little by little in the course of its development, and that Nature really was able to produce an organism from an unorganised material simply by her inherent forces.
It is interesting to display the essential contrast between preformation and epigenesis in the poetical words of Wolff himself. 'You must remember,' so run his words in the second argument against the probability of preformation, 'that an evolution would be a phenomenon formed in its real essence by God at the Creation, but created in condition invisible, and so as to remain invisible for long before it would become visible. See, then, that a phenomenon of enfolding is a miracle, differing from ordinary miracles only in these: first, it was at the creation of the world that God produced it; second, it remained invisible for long before it became visible. In truth, therefore, all organic bodies would be miracles. Would not this change for us the presence of Nature? Would it not spoil her of her beauty? Hitherto we had a living Nature, displaying endless changes by her own forces. Now it would be a fabric displaying change in seeming only, in truth and essence remaining unchanged and as it was constructed, save that it gradually becomes more and more used up. Formerly it was a Nature destroying herself and creating herself anew, only that endless changes might become visible and new sides be brought to light. Now it would be a lifeless mass shedding off piece after piece until the stock should come to an end.'
None the less, who seeks in Wolff's 'Theoria Generationis' an account of the means or forces by which Nature builds up organic forms will seek in vain. The vis essentialis (inherent force) with which Wolff endowed his plastic organic material, or the nisus formativus (formative force), afterwards suggested to science by Blumenbach—what are they but empty words by which men seek to grasp in thought what has eluded them? Wolff's epigenesis was not a complete explanation—indeed, from its fundamental conception it could not possibly be such. For investigation of the natural forces by which development proceeds can advance only slowly and step by step, and for long will constitute the foremost task of biology. The prosecution of biological investigation will continuously endow the theory of epigenesis with a fuller and fuller meaning, but will never transform it into a solution final in the sense of the theory of preformation.