Part 15 (1/2)

FOR REFERENCE

Bragg, _Rays and Crystals_ (Ball & Sons).

FOOTNOTES:

[Footnote 70: Since this address was given, the results of the Eclipse Expedition to Brazil are considered to have confirmed in a satisfactory manner one of the most remarkable deductions made by Einstein from the principles which he maintains. The matter has roused so much interest that some of the leading exponents of the relativity principle have published careful accounts intended for students not familiar with it: it would therefore be superfluous to discuss the matter here.]

IX

PROGRESS IN BIOLOGY DURING THE LAST SIXTY YEARS

PROFESSOR LEONARD DONCASTER, F.R.S.

On November 24, 1859, _The Origin of Species_ was published, and this date marks the beginning of an epoch in every branch of biology. Before it, Biology had been almost entirely a descriptive science, but within a few years after the publication of the _Origin_ its effects began to colour all aspects of biological research. A co-ordinating and unifying principle had been found, and the leading idea of biologists ceased to be to describe living things as they are, and became transformed into the attempt to discover how they are related to one another. The first effect of this change of att.i.tude was chiefly to turn biologists towards the task of tracing phylogenetic or evolutionary relations.h.i.+ps between different groups of animals--the drawing up of probable or possible genealogical trees and the explanation of natural cla.s.sification on an evolutionary basis. When once, however, the notion of cause and effect, or more correctly of relations.h.i.+p, between the phenomena seen in living beings had become familiar to biologists, it spread far beyond the limits of tracing genealogical connexions between different animals and plants. It made possible the conception of a true Science of Life, in which every phenomenon seen in a living organism should fall into its true place in relation to the rest, and in which also the phenomena of life should be correlated with those discovered in the inorganic sciences of Chemistry and Physics.

The history of the various branches of biological science in the past sixty years reflects the general course of these tendencies. Until shortly after 1859, the study of morphology, or the comparative structure of animals (and of plants) was intimately related with that of physiology, that is, with the study of function. In the years following the appearance of the _Origin_, however, anatomists and morphologists were seized with a new interest. For the time at least, the chief aim in studying structure was no longer to explain function, but rather to explain how that structure had come into being in the course of evolution, and how it was related with h.o.m.ologous but different structures in other forms. The result was a tendency to a divorce between morphology and physiology, or at least between morphologists and physiologists, which led to the division into two more or less distinct sciences of what had hitherto been regarded as closely inter-related branches of one. The greater men of the early part of the period, such as Huxley, remained both morphologists and physiologists, but most of their followers fell inevitably into one or the other group, and in discussing the later phases of biological progress it will be necessary to keep them separate.

Apart from its effect on the systematic and anatomical side of Biology, the idea of Evolution, and especially of Darwin's theory of Natural Selection, had important consequences on that side of the science which may be described as Natural History. Before the appearance of Darwin's work, Natural History consisted chiefly in the observation and collection of facts about the habits and life-history of animals and plants, which as a rule had no unifying principle unless they were used, as in the Bridgewater Treatises, to ill.u.s.trate 'the power, wisdom, and goodness of G.o.d'. Now, however, a new motive was provided--that of discovering the uses to the organism of its various colours, structures, and habits, and the application of the principle of natural selection to show how these characters conduced to the preservation and further evolution of the species. And out of this interest in the theory of natural selection grew in the last twenty years of the nineteenth century the greatly increased attention to the facts and theories of heredity, which was stimulated by Darwin's hypothesis of Pangenesis and especially by Weismann's speculations about the nature and behaviour of the 'germ-plasm'. Before the appearance of Weismann's work, the germ-cells, which bear somehow or other the hereditary characters that appear in the offspring, were supposed to be produced directly from the body of the parent. Darwin provisionally suggested that every cell of every organism gives off minute particles which become congregated in the germ-cells, and that these cells thus contain representative portions of all parts of the parent's body. Weismann, on the basis of his work on the origin of the germ-cells in Medusae and Insects, maintained that these cells are not derived from the body, but only from pre-existing germ-cells stored within it--that, in fact, although an egg gives rise to a hen, a hen does not give rise to an egg, but only keeps inside her a store of embryonic eggs which mature and are laid as the time comes round. The theory had to be modified to suit the facts of regeneration and vegetative reproduction, but in essence it was accepted by the biological world and is the orthodox opinion (if such a word may be used in Science) at the present day. The difference between the two views is not only of theoretical interest, for it involves the whole question of whether characteristics acquired by an individual during its life in response to external conditions can or cannot be transmitted to offspring. If the germ-cells contain representatives of all parts of the body, modifications impressed on the body during its life may at least possibly be transmitted to offspring born after the modifications have taken place. If, however, the germ-cells are independent of the rest of the body, and only stored within it for safe-keeping like a deed-box in the vaults of a bank, it would seem impossible for any environmental influence, whether for good or ill, to take effect on the offspring.

This controversy on the heritability of 'acquired characters' was one of the most important towards the end of last century, and although the majority of biologists now follow Weismann in so far as they deny that 'acquired' characters are transmissible, the question is not yet completely settled; all that can be said is that, in spite of many attempts to prove the contrary, there is no satisfactory evidence of the transmission to offspring of effects impressed on the body of the parent, unless the germ-cells themselves have been affected by the same cause--as for example in some cases of long-continued poisoning by alcohol or similar drugs.

While the problem of the transmission of acquired characters, and of the cause of variation and its relation to evolution, was occupying much of the attention of biologists, the whole problem entered upon a new phase in the year 1900 with the re-discovery of Mendel's work on heredity.

Mendel worked with plants, and published his results in 1865, but at that time the biological world was too much occupied with the fierce controversy which raged over _The Origin of Species_ to take much notice of a paper the bearing of which upon it was not appreciated. Mendel's discovery never came to the notice of Darwin, was buried in an obscure periodical, and remained unknown until many years after the death of its author. In 1900 it was unearthed, and, largely owing to the work of Bateson, it rapidly became known as one of the most important contributions to Biology made during the period under review.

This is not the place to describe in detail the nature of Mendel's theory. Its essence is, firstly, that the various characteristics of an organism are in general inherited quite independently of one another; and, secondly, that the germ-cells of a hybrid are pure in respect of any one character, that is to say, that any one germ-cell can only transmit any unit character as it was received from one parent or the other, and not a combination of the two. This leads to a conception of the organism as something like a mosaic, in which each piece of the pattern is transmitted in inheritance independently of the rest, and in which any piece cannot be modified by a.s.sociation with a different but corresponding piece derived from another ancestor. It is impossible to say as yet whether this conception at all completely represents the nature of the living organism, but it is one which is exercising considerable influence in biological thought, and if established it will mark a revolution in Biology hardly inferior to that brought about in Physics and Chemistry by the discovery of radio-activity.

An important consequence of the advance in our knowledge of heredity a.s.sociated with the work of Mendel and his successors is a tendency to doubt whether natural selection is of such fundamental importance in shaping the course of evolution as was supposed in the years of the first enthusiasm which followed the publication of the _Origin_.

Darwin based his theory of Natural Selection on the belief which he derived from breeders of plants and animals, that the kind of variation used by them to produce new breeds was the small and apparently unimportant differences which distinguish a 'fine' from a 'poor'

specimen. He supposed that the skilled breeder picked out as parents of his stock those individuals which were slightly superior in one feature or another, and that by the acc.u.mulative effect of these successive selections not only was the breed steadily improved, but also, by divergent selection, new breeds were produced. Experience shows, however, that although this method is used to keep breeds up to the required standard, it is rarely, if ever, the means by which new breeds arise. New breeds commonly come into existence either by a 'sport' or mutation, or by crossing two already distinct races, and by selecting from among the heterogeneous descendants of the cross those individuals which show the required combination of characters. And it is further found that most of the distinguis.h.i.+ng features of various breeds of domestic animals and plants are inherited according to Mendel's Law, suggesting that each of these characters is a unit, like one piece of a mosaic, independent of the rest. Now it is easy to see how the selection of small, continuously varying characters could take place in Nature by the destruction of all those individuals which failed to reach a certain standard, but it is much more difficult to understand how natural selection could act on comparatively large, sporadic, unco-ordinated 'sports'. There is thus a distinct tendency at present to regard natural selection as less omnipotent in directing the course of evolution than was formerly supposed, but it must be admitted that no very satisfactory alternative hypothesis has been suggested. Some have supposed that there is a kind of organic momentum which causes evolution to continue in those directions in which it has already proceeded, while others have postulated, like Bergson, an _elan vital_ as a kind of directive agency.

Others again have reverted towards the older belief in the inherited effects of environment--a belief which, in spite of the arguments of Weismann and his followers, has never been without its supporters. The present condition of this part of biology, as of many others, is one of open-mindedness approaching agnosticism. There is dissatisfaction with the beliefs which satisfied the preceding generation, and which were held up almost as dogmas, but there is no clear vision of the direction in which a truer view may be sought.

Before leaving this side of the subject, reference must be made to one important aspect of modern work on heredity--that of the inheritance of 'mental and moral' characteristics. As a result of the work of the biometric school founded by Galton and Pearson, it has been shown that the so-called mental and moral characteristics of man are inherited in the same manner and to the same extent as his physical features. Of the theoretical importance of this demonstration this is not the place to speak; its practical value is unquestionable, and may in the future have important effects on sociological problems.

Another notable line of advance, entirely belonging to the period under review, and chiefly the product of the present century, is seen in the science of Cytology--the investigation of the microscopic structure of the cells of which the body is composed. The marvellous phenomena of cell and nuclear division have revealed much of the formerly unsuspected complexity of living things, while the universality of the processes shows how fundamentally alike is life in all its forms. In recent years great progress has been made in correlating the phenomena of heredity and of the determination of s.e.x with the visible structural features of the germ-cells. Weismann attempted a beginning of this over thirty years ago, but the detailed knowledge of the facts was then insufficient.

Since the discovery of Mendel's Law, a great amount of work has been done, chiefly in America, by E.B. Wilson and T.H. Morgan and their pupils, on tracing the actual physical basis of hereditary transmission.

Although the matter is far from being completely known, the results obtained make it almost indubitable that inherited characters are in some way borne by the _chromosomes_ in the nuclei of the germ-cells.

The work of Morgan and his school has shown that the actual order in which these inherited 'factors' are arranged in the chromosomes can almost certainly be demonstrated, and his results go far to support the conception of the organism, referred to above, as a combination or mosaic of independently inherited features.

It was said at the beginning of this sketch that most of the more notable lines of advance in Biology could be traced back to the impetus given by the acceptance of the theory of Evolution, and the desire to test and prove that theory in every biological field. It is most convenient, therefore, to take this root-idea as a starting-point, and to see how the various branches of study have diverged from it and have themselves branched out in various ways, and how these branches have often again become intertwined and united in the later development of the science.

Perhaps the most obvious method of testing the theory of evolution is by the study of fossil forms, and our knowledge of these has progressed enormously during the period under review. Not only have a number of new and strange types of ancient life come to light, but in some cases, e.g.

in that of the horse and elephant, a very complete series of evolutionary stages has been discovered. In this branch, however, as in almost all others, the results have not exactly fulfilled the expectations of the early enthusiasts. On the one hand, evolution has been shown to be a much more complex thing than at first seemed probable; and on the other, many of the gaps which it was most hoped to fill still remain. A number of most remarkable 'missing links' have been discovered, such as, for example, _Archaeopteryx_, the stepping-stone between the Reptiles and the Birds, and the faith of the palaeontologist in the truth of evolution is everywhere confirmed. But the hope of finding all the stages, especially in the ancestry of Man, has not been realized, and it has been found that what at one time were regarded as direct ancestors are collaterals, and that the problem of human evolution is much less simple than was once supposed.

A second important piece of evidence in favour of evolution is provided by the study of the geographical distribution of animals, on which much work was done in the earlier part of the period under review. And in this connexion mention must be made of the science of Oceanography, for our whole knowledge of life in the abysses of the ocean, and almost all that we know of the conditions of life in the sea in general, has been gained in the last fifty years.