Chapter II: The Advance of Science, 1881-1906 (2)

The Giraffe-like animal called the Okapi, discovered by Sir Harry
Johnston in the Congo Forest. Photograph of the skin of a female
sent home by him in 1901, and now mounted and exhibited in the
Natural History Museum.]

Two “bandoliers” cut by the natives from the striped part of
the skin (the haunches) and at first supposed to be bits of the
hide of a new kind of Zebra. These were sent home by Sir Harry
Johnston in 1900.]

The main result of a good deal of such investigation is measured by our increased knowledge of the pedigree of organisms, what used to be called ‘classification.’ The anatomical study by the Australian professors, Hill and Wilson, of the teeth and the fœtus of the Australian group of pouched mammals--the marsupials--has entirely upset previous notions, to the effect that these are a primitive group, and has shown that their possession of only one replacing tooth is a retention of one out of many such teeth (the germs of which are present), as in placental mammals; and further that many of these marsupials have the nourishing outgrowth of the fœtus called the placenta fairly well developed, so that they must be regarded as a degenerate side-branch of the placental mammals, and not as primitive forerunners of that dominant series.

Photograph of the skull of a male Okapi--showing the paired boney
horn-cores--similar to those of the Giraffe, but connected with
the frontal bones and not with the parietals as the horn-cores of
Giraffes are.]

Drawings by Professor Grassi, of Rome, of the young of the common
Eel and its metamorphosis. All of the natural size. The uppermost
figure represents a transparent glass-like creature--which was
known as a rare “find” to marine naturalists, and received the
name _Leptocephalus_. Really it lives in vast numbers in great
depths of the sea--five hundred fathoms and more. It is hatched
here from the eggs of the common Eel which descends from the
ponds, lakes, and rivers of Europe in order to breed in these
great depths. The gradual change of the _Leptocephalus_ into a
young Eel or “Elver” is shown, and was discovered by Grassi. The
young Eels leave the great depth of the ocean and ascend the
rivers in immense shoals of many hundred thousand individuals,
and wriggle their way up banks and rocks into the small streams
and pools of the continent.

The above figures were published by Professor Grassi in November
1896, in the _Quarterly Journal of Microscopical Science_, edited
by E. Ray Lankester and published by Churchill & Sons.]

Speculations as to the ancestral connection of the great group of vertebrates with other great groups have been varied and ingenious; but most naturalists are now inclined to the view that it is a mistake to assume any such connection in the case of vertebrates of a more definite character than we admit in the case of starfishes, shell-fish, and insects. All these groups are ultimately connected by very simple, remote, and not by proximate ancestors, with one another and with the ancestors of vertebrates.

The unicellular parasite _Benedenia_, from the gut of the
common Poulp or Octopus. 1 is the normal male individual; 2 and
3 show stages in the production of spermatozoa on its surface
by budding; 4, 5 and 6 show a female parasite with spermatozoa
approaching it.]

Production of spermatozoa on the surface of the unicellular
parasite _Coccidium oviforme_, from the Rabbit’s intestines.]

The origin of the limbs of vertebrates is now generally agreed to be correctly indicated in the Thatcher-Mivart-Balfour theory to the effect that they are derived from a pair of continuous lateral fins, in fish-like ancestors, similar in every way to the continuous median dorsal fin of fishes.

Spermatozoa (often called “microgametes”) of the unicellular
parasite _Echinospora_ found in the gut of the small Centipede
_Lithobius mutabilis_.]

The discovery of the formation of true spermatozoa by simple unicellular animals of the group Protozoa is a startling thing, for it had always been supposed that these peculiar reproductive elements were only formed by multicellular organisms (figs. 21, 22, and 23). They have been discovered in some of the gregarina-like animalcules, the _Coccidia_, and also in the blood-parasites.

Among plants one of the most important discoveries relates to these same reproductive elements, the spermatozoa, which by botanists are called antherozoids. A great difference between the whole higher series of plants, the flowering plants or phanerogams, and the cryptogams or lower plants, including ferns, mosses, and algae, was held to be that the latter produce vibratile spermatozoa like those of animals which swim in liquid and fertilise the motionless egg-cell of the plant. Two Japanese botanists (and the origin of this discovery from Japan, from the University of Tokio, in itself marks an era in the history of science), Hirase and Ikeno, astonished the botanical world fifteen years ago by showing that motile antherozoids or spermatozoa are produced by two gymnosperms, the ging-ko tree (or _Salisburya_) and the cycads (fig. 24). The pollen-tube, which is the fertilising agent in all other phanerogams, develops in these cone-bearing trees, beautiful motile spermatozoa, which swim in a cup of liquid provided for them in connection with the ovules. Thus a great distinction between phanerogams and cryptogams was broken down, and the actual nature of the pollen-tube as a potential parent of spermatozoids demonstrated.

Spermatozoa (antherozoids) of _Cycas revoluta_, seen from the
side and from above. The spermatozoon is spherical, carrying a
spiral band of minute vibratile hairs (_cilia_) by which it is
propelled.]

When we come to the results of the digging out and study of extinct plants and animals, the most remarkable results of all in regard to the affinities and pedigree of organisms have been obtained. Among plants the transition between cryptogams and phanerogams has been practically bridged over by the discovery that certain fern-like plants of the Coal Measures--the _Cycadofilices_, supposed to be true ferns, are really seed-bearing plants and not ferns at all, but phanerogams of a primitive type, allied to the cycads and gymnosperms. They have been re-christened _Pteridosperms_ by Scott, who, together with F. Oliver and Seward, has been the chief discoverer in this most interesting field.

The gigantic three-horned Reptile, _Triceratops_, as large as an
Elephant, found in Jurassic strata in North America. A model of
the skeleton may be seen in the Natural History Museum in London.]

By their fossil remains whole series of new genera of extinct mammals have been traced through the tertiary strata of North America and their genetic connections established; and from yet older strata of the same prolific source we have almost complete knowledge of several genera of huge extinct _Dinosauria_ of great variety of form and habit (fig. 25).

Photograph of the skeleton of a large carnivorous Reptile
from Triassic strata in North Russia, discovered by Professor
Amalitzky and named by him, _Inostransevia_. The head alone is
two feet in length.]

Photographs of completed models of the Devonian fish
_Drepanaspis_, from Devonian slates of North Germany, worked out
by Professor Traquair. The models are in the Natural History
Museum, London.]

The oldest fossil fish known--discovered in the Upper Silurian
strata of Scotland, and named _Birkenia_ by Professor Traquair.]

The discoveries by Seeley at the Cape, and by Amalitzky in North Russia of identical genera of Triassic reptiles, which in many respects resemble the Mammalia and constitute the group _Theromorpha_, is also a prominent feature in the palæontology of the past twenty-five years (fig. 26). Nor must we forget the extraordinary Devonian and Silurian fishes discovered and described by Professor Traquair (figs. 27 and 28). The most important discovery of the kind of late years has been that of the Upper Eocene and Miocene Mammals of the Egyptian Fayum, excavated by the Egyptian Geological Survey and by Dr. Andrews of the Natural History Museum, who has described and figured the remains. They include a huge four-horned animal as big as a rhinoceros, but quite peculiar in its characters--the _Arisinoïtherium_--and the ancestors of the elephants, a group which was abundant in Miocene and Pliocene times in Europe and Asia, and in still later times in America, and survives at the present day in its representatives the African and Indian elephant. One of the European extinct elephants--the _Tetrabelodon_--had, we have long known, an immensely long lower jaw with large chisel-shaped terminal teeth. It had been suggested by me that the modern elephant’s trunk must have been derived from the soft upper jaw and nasal area, which rested on this elongated lower jaw, by the shortening (in the course of natural selection and modification by descent) of this long lower jaw, to the present small dimensions of the elephant’s lower jaw, and the consequent down-dropping of the unshortened upper jaw and lips, which thus become the proboscis. Dr. Andrews has described from Egypt and placed in the Museum in London specimens of two new genera--one _Palæomastodon_, in which there is a long, powerful jaw, an elongated face, and an increased number of molar teeth (see figs. 29 and 30); the second, _Meritherium_ (fig. 31), an animal with a hippopotamus-like head, comparatively minute tusks, and a well-developed complement of incisor, canine, and molar teeth, like a typical ungulate mammal. Undoubtedly we have in these two forms the indications of the steps by which the elephants have been evolved from ordinary-looking pig-like creatures of moderate size, devoid of trunk or tusks. Other remains belonging to this great mid-African Eocene fauna indicate that not only the Elephants but the Sirenia (the Dugong and Manatee) took their origin in this area. Amongst them are also gigantic forms of Hyrax, like the little Syrian coney and many other new mammals and reptiles.

Photograph of a complete model of the skull and lower jaw of the
ancestral elephant, _Palæomastodon_, discovered by Dr. Andrews
in the Upper Eocene of the Fayum Desert, Egypt, and modelled
and restored under his direction in the Natural History Museum,
London. The comparatively short trunk or snout rested on the
broad front teeth of the long lower jaw. The face is elongated,
and the cheek-teeth are numerous.]

Photograph of the lower face of the skull of a specimen of
_Palæomastodon_ brought from Egypt in April, 1906, by Dr.
Andrews, and now in the Natural History Museum, London. The
six characteristic cheek-teeth on each side, and the pair of
sabre-like tusks in front, are well seen.]

Drawing of the skull and lower jaw of the _Meritherium_,
discovered by Dr. Andrews in the Upper Eocene of the Fayum
Desert. The shape of the skull and proportions of face and jaw
are like those of an ordinary hoofed mammal such as the pig;
but the cheek-teeth are similar to those of the _Mastodon_, and
whilst the full complement of teeth is present in the front of
the upper jaw, we can distinguish the big tusk-like incisor which
alone survives on each side in _Palæomastodon_, _Mastodon_, and
the elephants, as the great pair of tusks.]

Another great area of exploration and source of new things has been the southern part of Argentina and Patagonia, where Ameghino, Moreno, and Scott of Princeton have brought to light a wonderful series of extinct ant-eaters, armadilloes, huge sloths, and strange ungulates, reaching back into early Tertiary times. But most remarkable has been the discovery in this area of remains which indicate a former connection with the Australian land surface. This connection is suggested by the discovery in the Santa Cruz strata, considered to be of early Tertiary date, of remains of a huge horned tortoise which is generically identical with one found fossil in the Australian area of later date, and known as _Miolania_. In the same wonderful area we have the discovery in a cave of the fresh bones, hairy skin, and dung of animals supposed to be extinct, viz., the giant sloth, _Mylodon_, and the peculiar horse, _Onohippidium_. These remains seem to belong to survivors from the last submergence of this strangely mobile land-surface, and it is not improbable that some individuals of this ‘extinct’ fauna are still living in Patagonia. The region is still unexplored and those who set out to examine it have, by some strange fatality, hitherto failed to carry out the professed purpose of their expeditions.

I cannot quit this immense field of gathered fact and growing generalisation without alluding to the study of animal embryology and the germ-layer theory, which has to some extent been superseded by the study of embryonic cell-lineage, so well pursued by some American microscopists. The great generalisation of the study of the germ-layers and their formation seems to be now firmly established--namely, that the earliest multicellular animals were possessed of one structural cavity, the enteron, surrounded by a double layer of cells, the ectoderm and endoderm. These _Enterocœla_ or _Cœlentera_ gave rise to forms having a second great body-cavity, the cœlom, which originated not as a split between the two layers, as was supposed twenty-five years ago by Haeckel and Gegenbaur and their pupils, but by a pouching of the enteron to form one or more cavities in which the reproductive cells should develop--pouchings which became nipped off from the cavity of their origin, and formed thus the independent cœlom. The animals so provided are the _Cœlomocœla_ (as opposed to the _Enterocœla_), and comprise all animals above the polyps, jelly-fish, corals, and sea-anemones. It has been established in these twenty-five years that the cœlom is a definite structural unit of the higher groups, and that outgrowths from it to the exterior (cœlomoducts) form the genital passages, and may become renal excretory organs also. The vascular system has not, as it was formerly supposed to have, any connection of origin with the cœlom, but is independent of it, in origin and development, as also are the primitive and superficial renal tubes known as nephridia. These general statements seem to me to cover the most important advance in the general morphology of animals which we owe to embryological research in the past quarter of a century.[16]

Before leaving the subject of animal morphology I must apologise for my inability to give space and time to a consideration of the growing and important science of anthropology, which ranges from the history of human institutions and language to the earliest prehistoric bones and implements. Let me therefore note here the discovery of the cranial dome of _Pithecanthropus_ in a river gravel in Java--undoubtedly the most ape-like of human remains, and of great age (see figs. 1 and 2); and, further, the Eoliths of Prestwich (see figs. 3 and 4), in the human authorship of which I am inclined to believe, though I should be sorry to say the same of all the broken flints to which the name ‘Eolith’ has been applied. The systematic investigation and record of savage races have taken on a new and scientific character. Such work as Baldwin Spencer’s and Haddon’s in Australasia furnish examples of what is being done in this way.

_Bacillus radicola_, the parasite which infests the roots of
leguminous plants and causes the growth of nodules whilst
assisting the plant in the assimilation of nitrogen: (_a_)
Nodule of the roots of the common Lupine, natural size; (_b_)
longitudinal section through a Lupine root and nodule; (_c_)
a single cell from a Lupine nodule showing the bacteria or
bacilli, as black particles in the protoplasm, magnified 600
diameters; (_d_) bacilli from the root nodule of the Lupine;
(_e_) triangular forms of the bacillus from the root nodules of
the Vetch; (_f_) oval forms from the root nodules of the Lupine;
(_d e f_) are magnified 1,500 diameters.]

_Physiology of Plants and Animals._--Since I have not space to do more than pick out the most important advances in each subject for brief mention, I must signalize in regard to the physiology of plants the better understanding of the function of leaf-green or chlorophyll due to Pringsheim and to the Russian Timiriaseff, the new facts as to the activity of stomata in transpiration discovered by Horace Brown, and the fixation of free nitrogen by living organisms in the soil and by organisms (_Bacillus radicola_) parasitic in the rootlets of leguminous plants (see fig. 32), which thus benefit by a supply of nitrogenous compounds which they can assimilate.

Great progress in the knowledge of the chemistry of the living cells or protoplasm of both plants and animals has been made by the discovery of the fact that ferments or enzymes are not only secreted externally by cells, but exist active and preformed _inside_ cells. Büchner’s final conquest of the secret of the yeast-cell by heroic mechanical methods--the actual grinding to powder of these already very minute bodies--first established this, and now successive discoveries of intracellular ferments have led to the conclusion that it is probable that the cell respires by means of a respiratory ‘oxydase,’ builds up new compounds and destroys existing ones, contracts and accomplishes its own internal life by ferments. Life thus (from the chemical point of view) becomes a chain of ferment actions. Another most significant advance in animal physiology has been the sequel (as it were) of Bernard’s discovery of the formation of glycogen in the liver, a substance not to be excreted, but to be taken up by the blood and lymph, and in many ways more important than the more obvious formation of bile which is thrown out of the gland into the alimentary canal. It has been discovered that many glands, such as the kidney and pancreas and the ductless glands, the suprarenals, thyroid, and others, secrete indispensable products into the blood and lymph. Hence myxœdema, exophthalmic goitre, Addison’s disease, and other disorders have been traced to a deficiency or excess of internal secretions from glands formerly regarded as interesting but unimportant vestigial structures. From these glands have in consequence been extracted remarkable substances on which their peculiar activity depends. From the suprarenals a substance has been extracted which causes activity of all those structures which the sympathetic nerve-system can excite to action; the thyroid yields a substance which influences the growth of the skin, hair, bones, &c.; the pituitary gland, an extract which is a specific urinary stimulant. Quite lately the mammalian ovary has been shown by Starling to yield a secretion which influences the state of nutrition of the uterus and mammæ. A great deal more might be said here on topics such as these--topics of almost infinite importance; but the fact is that the mere enumeration of the most important lines of progress in any one science would occupy many pages.

The continuity of the protoplasm of neighbouring vegetable cells,
by means of threads which perforate the cell-walls. Drawing
(_after_ Gardiner) of cells from the pulvinus of _Robinia_.]

Nerve-physiology has made immensely important advances. There is now good evidence that all excitation of one group of nerve-centres is accompanied by the _concurrent inhibition_ of a whole series of groups of other centres, whose activity might interfere with that of the group excited to action. In a simple reflex flexure of the knee the motor-neurones to the flexor muscles are excited, but concurrently the motor-neurones to the extensor muscles are thrown into a state of inhibition, and so equally with all the varied excitations of the nervous system controlling the movements and activities of the entire body.

Diagrammatic representation of the structures present in a
typical cell (_after_ Wilson). Note the two centrosomes,
sometimes single.]

The discovery of the continuity of the protoplasm through the walls of the vegetable cells by means of connecting canals and threads (see fig. 33) is one of the most startling facts discovered in connection with plant-structure, since it was held twenty years ago that a fundamental distinction between animal and vegetable structure consisted in the boxing-up or encasement of each vegetable cell-unit in a case of cellulose, whereas animal cells were not so imprisoned, but freely communicated with one another. It perhaps is on this account the less surprising that lately something like sense-organs have been discovered on the roots, stems, and leaves of plants, which, like the otocysts of some animals, appear to be really ‘statocytes,’ and to exert a varying pressure according to the relations of these parts of the plant to gravity. There is apparently something resembling a perception of the incidence of gravity in plants which reacts on irritable tissues, and is the explanation of the phenomena of geotropism. These results have grown out of the observations of Charles Darwin, followed by those of F. Darwin, Haberlandt, and Nemec.

FIG. 35.--THE NUMBER OF THE CHROMOSOMES: (_a_) Cell of the asexual
generation of the cryptogam _Pellia epiphylla_: the nucleus is
about to divide, a polar ray-formation is present at each end of
the spindle-shaped nucleus, the chromosomes have divided into two
horizontal groups each of sixteen pieces: _sixteen_ is the number
of the chromosomes of the ordinary tissue cells of _Pellia_.
(_b_) Cell of the sexual generation of the same plant (_Pellia_)
in the same phase of division, but with the _reduced_ number
of chromosomes--namely, _eight_ in each half of the dividing
nucleus. The completed cells of the sexual generation have only
_eight_ chromosomes. (_c_) Somatic or tissue cell of Salamander
showing _twenty-four_ =V=-shaped chromosomes, each of which is
becoming longitudinally split as a preliminary to division.
(_d_) Sperm-mother-cell from testis of Salamander, showing the
_reduced_ number of chromosomes of the sexual cells--namely,
_twelve_; each is split longitudinally. (From original drawings
by Prof. Farmer and Mr. Moore.)]

A few words must be said here as to the progress of our knowledge of cell-substance, and what used to be called the protoplasm question. We do not now regard protoplasm as a chemical expression, but, in accordance with von Mohl’s original use of the word, as a structure which holds in its meshes many and very varied chemical bodies of great complexity. Within these twenty-five years the ‘centrosome’ of the cell-protoplasm has been discovered (see fig. 34), and a great deal has been learnt as to the structure of the nucleus and its remarkable stain-taking bands, the chromosomes. We now know that these bands are of definite fixed number, varying in different species of plants and animals, and that they are halved in number in the reproductive elements--the spermatozoid and the ovum--so that on union of these two to form the fertilized ovum (the parent cell of all the tissues), the proper specific number is attained (see figs. 35 and 36). It has been pretty clearly made out by cutting up large living cells--unicellular animals--that the body of the cell alone, without the nucleus, can do very little but move and maintain for a time its chemical status. But it is the nucleus which directs and determines all definite growth, movement, secretion, and reproduction. The simple protoplasm, deprived of its nucleus, cannot form a new nucleus--in fact, can do very little but exhibit irritability. I am inclined to agree with those who hold that there is not sufficient evidence that any organism exists at the present time which has not both protoplasm and nucleus--in fact, that the simplest form of life at present existing is a highly complicated structure--a nucleated cell. That does not imply that simpler forms of living matter have not preceded those which we know. We must assume that something more simple and homogeneous than the cell, with its differentiated cell-body or protoplasm, and its cell kernel or nucleus, has at one time existed. But the various supposed instances of the survival to the present day of such simple living things--described by Haeckel and others--have one by one yielded to improved methods of microscopic examination and proved to be differentiated into nuclear and extra-nuclear substance.

Further stage in the division of the sexual cell drawn in Fig.
35(_e_), showing the _twelve_ chromosomes of the two nuclei of
the sperm-cells resulting from the division (_twelve_ instead of
_twenty-four_).]

The question of ‘spontaneous generation’ cannot be said to have been seriously revived within these twenty-five years. Our greater knowledge of minute forms of life, and the conditions under which they can survive, as well as our improved microscopes and methods of experiment and observation, have made an end of the arguments and instances of supposed abiogenesis. The accounts which have been published of ‘radiobes,’ minute bodies arising in fluids of organic origin when radium salts have been allowed to mix in minute quantities with such fluids, are wanting in precision and detail, but the microscopic particles which appear in the circumstances described seem to be of a nature identical with the minute bodies well known to microscopists and recognised as crystals modified by a colloid medium. They have been described by Rainey, Harting, and Ord, on different occasions, many years ago. They are not devoid of interest, but cannot be considered as having any new bearing on the origin of living matter.

_Psychology._--I have given a special heading to this subject because its emergence as a definite line of experimental research seems to me one of the most important features in the progress of science in the past quarter of a century. Thirty-five years ago we were all delighted by Fechner’s psycho-physical law, and at Leipzig I, with others of my day, studied it experimentally in the physiological laboratory of that great teacher, Carl Ludwig. The physiological methods of measurement (which are the physical ones) have been more and more widely, and with guiding intelligence and ingenuity, applied since those days to the study of the activities of the complex organs of the nervous system which are concerned with ‘mind’ or psychic phenomena. Whilst some enthusiasts have been eagerly collecting ghost stories and records of human illusion and fancy, the serious experimental investigation of the human mind, and its forerunner the animal mind, has been quietly but steadily proceeding in truly scientific channels. The science is still in an early phase--that of the collection of accurate observations and measurements--awaiting the development of great guiding hypotheses and theories. But much has been done, and it is a matter of gratification to Oxford men that through the liberality of the distinguished electrician, Mr. Henry Wilde, F.R.S., a lectureship of Experimental Psychology has been founded in the University of Oxford, where the older studies of Mental and Moral Philosophy, Logic and Metaphysics have so strong a hold, and have so well prepared the ground for the new experimental development. The German investigators W. Wundt, G. E. Müller, C. Stumpf, Ebbinghaus, and Munsterberg have been prominent in introducing laboratory methods, and have determined such matters as the elementary laws of association and memory, and the perceptions of musical tones and their relations. The work of Goldschneider on ‘the muscular sense,’ of von Frey on the cutaneous sensations, are further examples of what is being done.

The difficult and extremely important line of investigation, first scientifically treated by Braid under the name ‘Hypnotism,’ has been greatly developed by the French school, especially by Charcot. The experimental investigation of ‘suggestion,’ and the pathology of dual consciousness and such exceptional conditions of the mind, has been greatly advanced by French observers.

The older work of Ferrier and Hitzig on the functions of the parts of the brain has been carried further by Goltz and Munk in Germany, and by Schäfer, Horsley, and Sherrington in England.

The most important general advance seems to be the recognition that the mind of the human adult is a social product; that it can only be understood in relation with the special environment in which it develops, and with which it is in perpetual interaction. Professor Baldwin, of Princeton, has done important work on this subject. Closely allied is the study of what is called ‘the psychology of groups,’ the laws of mental action of the individual as modified by his membership of some form of society. French authors have done valuable work here.

These two developments of psychology are destined to provide the indispensable psychological basis for Social Science, and for the anthropological investigation of mental phenomena.

Hereafter, the well-ascertained laws of experimental psychology will undoubtedly furnish the necessary scientific basis of the art of education, and psychology will hold the same relation to that art as physiology does to the art of medicine and hygiene.

There can be little doubt, moreover, of the valuable interaction of the study of physical psychology and the theories of the origin of structural character by natural selection. The relation of the human mind to the mind of animals, and the gradual development of both, form a subject full of rich stores of new material, yielding conclusions of the highest importance, which has not yet been satisfactorily approached.

I am glad to be able to give wider publicity here to some conclusions which I communicated to the Jubilee volume of the ‘Société de Biologie’ of Paris in 1899. I there discussed the significance of the great increase in the size of the cerebral hemispheres in recent, as compared with Eocene Mammals (see fig. 5), and in Man as compared with Apes, and came to the conclusion that ‘the power of building up appropriate cerebral mechanism in response to individual experience,’ or what may be called ‘educability,’ is the quality which characterizes the larger cerebrum, and is that which has led to its selection, survival, and further increase in volume. The bearing of this conception upon questions of fundamental importance in what has been called genetic psychology is sketched as follows:

‘The character which we describe as “educability” can be transmitted; it is a congenital character. But the _results_ of education can _not_ be transmitted. In each generation they have to be acquired afresh. With increased “educability” they are more readily acquired and a larger variety of them. On the other hand, the nerve-mechanisms of instinct are transmitted, and owe their inferiority as compared with the results of education to the very fact that they are _not_ acquired by the individual in relation to his particular needs, but have arisen by selection of congenital variation in a long series of preceding generations.’

‘To a large extent the two series of brain-mechanisms, the “instinctive” and the “individually acquired,” are in opposition to one another. Congenital brain-mechanisms may prevent the education of the brain and the development of new mechanisms specially fitted to the special conditions of life. To the educable animal the less there is of specialised mechanism transmitted by heredity, the better. The loss of instinct is what permits and necessitates the education of the receptive brain.’

‘We are thus led to the view that it is hardly possible for a theory to be further from the truth than that expressed by George H. Lewes and adopted by George Romanes, namely, that instincts are due to “lapsed” intelligence. The fact is that there is no community between the mechanisms of instinct and the mechanisms of intelligence, and that the latter are later in the history of the development of the brain than the former, and can only develop in proportion as the former become feeble and defective.’[17]

_Darwinism._--Under the title ‘Darwinism’ it is convenient to designate the various work of biologists tending to establish, develop, or modify Mr. Darwin’s great theory of the origin of species. In looking back over twenty-five years it seems to me that we must say that the conclusions of Darwin as to the origin of species by the survival of selected races in the struggle for existence are more firmly established than ever. And this because there have been many attempts to gravely tamper with essential parts of the fabric as he left it, and even to substitute conceptions for those which he endeavoured to establish, at variance with his conclusions. These attempts must, I think, be considered as having failed. A great deal of valuable work has been done in consequence; for honest criticism, based on observation and experiment, leads to further investigation, and is the legitimate and natural mode of increase of scientific knowledge. Amongst the attempts to seriously modify Darwin’s doctrine may be cited that to assign a great and leading importance to Lamarck’s theory as to the transmission by inheritance of newly ‘acquired’ characters, due chiefly to American palæontologists and to the venerated defender of such views, who has now closed his long life of great work, Mr. Herbert Spencer; that to attribute leading importance to the action of physiological congruity and incongruity in selective breeding, which was put forward by another able writer and naturalist who has now passed from among us, Dr. George Romanes; further, the views of de Vries as to the discontinuity in the origin of new species, supported by the valuable work of Mr. Bateson on discontinuous variation; and lastly, the attempt to assign a great and general importance to the facts ascertained many years ago by the Abbé Mendel as to the cross-breeding of varieties and the frequent production (in regard to certain characters in certain cases) of pure strains rather than of breeds combining the characters of both parents. On the other hand we have the splendid series of observations and writings of August Weismann, who has, in the opinion of the majority of those who study this subject, rendered the Lamarckian theory of the origin and transmission of new characters altogether untenable, and has, besides, furnished a most instructive, if not finally conclusive, theory or mechanical scheme of the phenomena of Heredity in his book ‘The Germ-plasm.’ Professor Karl Pearson and the late Professor Weldon--the latter so early in life and so recently lost to us--have, with the finest courage and enthusiasm in the face of an enormous and difficult task, determined to bring the facts of variation and heredity into the solid form of statistical statement, and have organised, and largely advanced in, this branch of investigation which they have termed ‘Biometrics.’ Many naturalists throughout the world have made it the main object of their collecting and breeding of insects, birds, and plants, to test Darwin’s generalisations and to expand the work of Wallace in the same direction. A delightful fact in this survey is that we find Mr. Alfred Russel Wallace (who fifty years ago conceived the same theory as that more fully stated by Darwin) actively working and publishing some of the most convincing and valuable works on Darwinism. He is still alive and not merely well, but pursuing his work with vigour and ability. It was chiefly through his researches on insects in South America and the Malay Islands that Mr. Wallace was led to the Darwinian theory; and there is no doubt that the study of insects, especially of butterflies, is still one of the most prolific fields in which new facts can be gathered in support of Darwin and new views on the subject tested. Prominent amongst naturalists in this line of research has been and is Edward Poulton of Oxford, who has handed on to the study of entomology throughout the world the impetus of the Darwinian theory. I must here also name a writer who, though unknown in our laboratories and museums, seems to me to have rendered very valuable service in later years to the testing of Darwin’s doctrines and to the bringing of a great class of organic phenomena within the cognisance of those naturalists who are especially occupied with the problems of Variation and Heredity. I mean Dr. Archdall Reid, who has with keen logic made use of the immense accumulation of material which is in the hands of medical men, and has pointed out the urgent importance of increased use by Darwinian investigators of the facts as to the variation and heredity of that unique animal, man, unique in his abundance, his reproductive activity, and his power of assisting his investigator by his own record. There are more observations about the variation and heredity of man and the conditions attendant upon individual instances than with regard to any other animal. Medical men need only to grasp clearly the questions at present under discussion in order to be able to furnish with ease data absolutely invaluable in quantity and quality. Dr. Archdall Reid has in two original books full of insight and new suggestions, the ‘Present Evolution of Man’ and ‘Principles of Heredity,’ shown a new path for investigators to follow.

The attempt to resuscitate Lamarck’s views on the inheritance of acquired[18] characters has been met not only by the demand for the production of experimental proof that such inheritance takes place, which has never been produced, but on Weismann’s part by a demonstration that the reproductive cells of organisms are developed and set aside from the rest of the tissues at so early a period that it is extremely improbable that changes brought about in those other tissues by unaccustomed incident forces can be communicated to the germ-cells so as to make their appearance in the offspring by heredity. Apart from this, I have drawn attention to the fact that Lamarck’s first and second laws (as he terms them) of heredity are contradictory the one of the other, and therefore may be dismissed. In 1894 I wrote:

‘Normal conditions of environment have for many thousands of generations moulded the individuals of a given species of organism, and determined as each individual developed and grew “responsive” quantities in its parts (characters): yet, as Lamarck tells us, and as we know, there is in every individual born a potentiality which has _not_ been extinguished. Change the normal conditions of the species in the case of a young individual taken to-day from the site where for thousands of generations its ancestors have responded in a perfectly defined way to the normal and defined conditions of environment; reduce the daily or the seasonal amount of solar radiation to which the individual is exposed; or remove the aqueous vapour from the atmosphere; or alter the chemical composition of the pabulum accessible; or force the individual to previously unaccustomed muscular effort or to new pressures and strains; and (as Lamarck bids us observe), in spite of all the long-continued response to the earlier normal specific conditions, the innate congenital potentiality shows itself. The individual under the new quantities of environing agencies shows _new_ responsive quantities in those parts of its structure concerned, new or _acquired_ characters.

‘So far, so good. What Lamarck next asks us to accept, as his “second law,” seems not only to lack the support of experimental proof, but to be inconsistent with what has just preceded it. The new character which is _ex hypothesi_, as was the old character (length, breadth, weight of a part) which it has replaced--a response to environment, a particular moulding or manipulation by incident forces of the potential congenital quality of the race--is, according to Lamarck, all of a sudden raised to extraordinary powers. The new or freshly acquired character is declared by Lamarck and his adherents to be capable of transmission by generation; that is to say, it alters the potential character of the species. It is no longer a merely responsive or reactive character, determined quantitatively by quantitative conditions of the environment, but becomes fixed and incorporated in the potential of the race, so as to persist when other quantitative external conditions are substituted for those which originally determined it. In opposition to Lamarck, one must urge, in the first place, that this thing has never been shown experimentally to occur; and in the second place, that there is no ground for holding its occurrence to be probable, but, on the contrary, strong reason for holding it to be improbable. Since the old character (length, breadth, weight) had not become fixed and congenital after many thousands of successive generations of individuals had developed it in response to environment, but gave place to a new character when new conditions operated on an individual (Lamarck’s first law), why should we suppose that the new character is likely to become fixed after a much shorter time of responsive existence, or to escape the operation of the first law? Clearly there is no reason (so far as Lamarck’s statement goes) for any such supposition, and the two so-called laws of Lamarck are at variance with one another.’

In its most condensed form my argument has been stated thus by Professor Poulton: Lamarck’s ‘first law assumes that a past history of indefinite duration is powerless to create a bias by which the present can be controlled; while the second assumes that the brief history of the present can readily raise a bias to control the future.’[19]

An important light is thrown on some facts which seem at first sight to favour the Lamarckian hypothesis by the consideration that, though an ‘acquired’ character is not transmitted to offspring as the consequence of the action of external agencies determining the ‘acquirement,’ yet the tendency to react exhibited by the parent _is_ transmitted, and if the tendency is exceptionally great a false suggestion of a Lamarckian inheritance can readily result. This inheritance of ‘variation in tendencies to react’ has a wide application, and has led me to coin the word ‘educability’ as mentioned in the section of this address on Psychology.

The principle of physiological selection advocated by Dr. Romanes does not seem to have caused much discussion, and has been unduly neglected by subsequent writers. It was ingenious, and was based on some interesting observations, but has failed to gain support.

The observations of de Vries--showing that in cultivated varieties of plants a new form will sometimes assert itself suddenly and attain a certain period of dominance, though not having been gradually brought into existence by a slow process of selection--have been considered by him, and by a good many other naturalists, as indicating the way in which new species arise in Nature. The suggestion is a valuable one if not very novel, but a great deal of observation will have to be made before it can be admitted as really having a wide bearing upon the origin of species. The same is true of those interesting observations which were first made by Mendel, and have been resuscitated and extended with great labour and ingenuity by recent workers, especially in this country by Bateson and his pupils. If it should prove to be true that varieties when crossed do not, in the course of eventual inter-breeding, produce intermediate forms as hybrids, but that characters are either dominant or recessive, and that breeds result having pure unmixed characters--we should, in proportion as the Mendelian law is shown to apply to all tissues and organs and to a majority of organisms, have before us a very important and determining principle in all that relates to heredity and variation. It remains, however, to be shown how far the Mendelian phenomenon is general. And it is, of course, admitted on all sides that, even were the Mendelian phenomenon general and raised to the rank of a law of heredity, it would not be subversive of Mr. Darwin’s generalisations, but probably tend to the more ready application of them to the explanation of many difficult cases of the structure and distribution of organisms.

Two general principles which Mr. Darwin fully recognised appear to me to deserve more consideration and more general application to the history of species than he had time to give to them, or than his followers have accorded to them. The first is the great principle of ‘correlation of variation,’ from which it follows that, whilst natural selection may be favouring some small and obscure change in an unseen group of cells--such as digestive, pigmentary or nervous cells, and that change a change of selective value--there may be, indeed often is, as we know, a correlated or accompanying change in a physiologically related part of far greater magnitude and prominence to the eye of the human onlooker. This accompanying or correlated character has no selective value, is not an adaptation--is, in fact, a necessary but useless by-product. A list of a few cases of this kind was given by Darwin, but it is most desirable that more should be established. For they enable us to understand how it is that specific characters, those seen and noted on the surface by systematists, are not in most cases adaptations of selective value. They also open a wide vista of incipient and useless developments which may suddenly, in their turn, be seized upon by ever-watchful natural selection and raised to a high pitch of growth and function.

The second, somewhat but by no means altogether neglected, principle is that a good deal of the important variation in both plants and animals is not the variation of a minute part or confined to one organ, but has really an inner physiological basis, and may be a variation of a whole organic system or of a whole tissue expressing itself at several points and in several shapes. In fact, we should perhaps more generally conceive of variation as not so much the accomplishment and presentation of one little mark or difference in weight, length, or colour, as the expression of a _tendency to vary_ in a given tissue or organ in a particular way. Thus we are prepared for the rapid extension and dominance of the variation if once it is favoured by selective breeding. It seems to me that such cases as the complete disappearance of scales from the integument of some osseous fishes, or the possible retention of three or four scales out of some hundreds present in nearly allied forms, favour this mode of conceiving of variation. So also does the marked tendency to produce membranous expansions of the integument in the bats, not only between the digits and from the axilla, but from the ears and different regions of the face. Of course, the alternative hairy or smooth condition of the integuments both in plants and animals is a familiar instance in which a tendency extending over a large area is recognised as that which constitutes the variation. In smooth or hairy varieties we do not postulate an individual development of hairs subjected one by one to selection and survival or repression.

_Disease._--The study of the physiology of unhealthy, injured, or diseased organisms is called pathology. It necessarily has an immense area of observation and is of transcending interest to mankind who do not accept their diseases unresistingly and die as animals do, so purifying their race, but incessantly combat and fight disease, producing new and terrible forms of it, by their wilful interference with the earlier rule of Nature.

Our knowledge of disease has been enormously advanced in the last quarter of a century, and in an important degree our power of arresting it, by two great lines of study going on side by side and originated, not by medical men nor physiologists in the narrow technical sense, but by naturalists, a botanist, and a zoologist. Ferdinand Cohn, Professor of Botany in Breslau, by his own researches and by personal training in his laboratory, gave to Robert Koch the start on his distinguished career as a bacteriologist. It is to Metschnikoff the zoologist and embryologist that we owe the doctrine of phagocytosis and the consequent theory of immunity now so widely accepted.

We must not forget that in this same period much of the immortal work of Pasteur on hydrophobia, of Behring and Roux on diphtheria, and of Ehrlich and many others to whom the eternal gratitude of mankind is due, has been going on. It is only some fifteen years since Calmette showed that if cobra poison were introduced into the blood of a horse in less quantity than would cause death, the horse would tolerate with little disturbance after ten days a full dose, and then day after day an increasing dose, until the horse without any inconvenience received an injection of cobra poison large enough to kill thirty horses of its size. Some of the horse’s blood being now withdrawn was found to contain a very active antidote to cobra poison--what is called an antitoxin. The procedure in the preparation of the antitoxin is practically the same as that previously adopted by Behring in the preparation of the antitoxin of diphtheria poison. Animals treated with injections of these antitoxins are immune to the poison itself when subsequently injected with it, or, if already suffering from the poison (as, for instance, by snake-bite), are readily shown by experiment to be rapidly cured by the injection of the appropriate antitoxin. This is, as all will admit, an intensely interesting bit of biology. The explanation of the formation of the antitoxin in the blood and its mode of antagonising the poison is not easy. It seems that the antitoxin is undoubtedly formed from the corresponding toxin or poison, and that the antagonism can be best understood as a chemical reaction by which the complex molecule of the poison is upset, or effectively modified.

The remarkable development of Metschnikoff’s doctrine of phagocytosis during the past quarter of a century is certainly one of the characteristic features of the activity of biological science in that period. At first ridiculed as ‘Metschnikoffism,’ it has now won the support of its former adversaries.

For a long time the ideal of hygienists has been to preserve man from all contact with the germs of infection, to destroy them and destroy the animals conveying them, such as rats, mosquitoes, and other flies. But it has now been borne in upon us that, useful as such attempts are, and great as is the improvement in human conditions which can thus be effected, yet we cannot hope for any really complete or satisfactory realisation of the ideal of escape from contact with infective germs. The task is beyond human powers. The conviction has now been arrived at that, whilst we must take every precaution to diminish infection, yet our ultimate safety must come from within--namely, from the activity, the trained, stimulated, and carefully guarded activity, of those wonderful colourless amœba-like corpuscles whose use was so long unrecognized, but has now been made clear by the patiently continued experiments and arguments of Metschnikoff, who has named them ‘phagocytes.’ The doctrine of the activity and immense importance of these corpuscles of the living body which form part of the all-pervading connective tissues and float also in the blood, is in its nature and inception opposed to what are called the ‘humoral’ and ‘vitalistic’ theories of resistance to infection. Of this kind were the beliefs that the _liquids_ of the living body have an inherent and somewhat vague power of resisting infective germs, and even that the mere living quality of the tissues was in some unknown way antagonistic to foreign intrusive disease-germs.

FIG. 37. FIG. 38. FIG. 39.

Fig. 37.--Phagocyte or colourless corpuscle of a guinea-pig
in the act of engulphing two Spirilla or parasitic vegetable
microbes of a spiral shape. Fig. 38.--The same half an hour
later, one of the Spirilla is nearly completely engulphed. Fig.
39.--The same ten minutes later still, one of the Spirilla is
completely absorbed into the substance (protoplasm) of the
phagocyte. (_From_ Metschnikoff’s book, “Immunity,” kindly
supplied by the Cambridge University Press.)]

The first eighteen years of Metschnikoff’s career, after his undergraduate course, were devoted to zoological and embryological investigations. He discovered many important facts, such as the alternation of generations in the parasitic worm of the frog’s lung--_Ascaris nigrovenosa_--and the history of the growth from the egg of sponges and medusæ. In these latter researches he came into contact with the wonderfully active cells, or living corpuscles, which in many low forms of life can be seen by transparency in the living animal. He saw that these corpuscles (as was indeed already known) resemble the well-known amœba, and can take into their soft substance (protoplasm) at all parts of their surface any minute particles and digest them, thus destroying them. In a transparent water-flea Metschnikoff saw these amœba-like, colourless, floating blood-corpuscles swallowing and digesting the spores of a parasitic fungus which had attacked the water-fleas and was causing their death. He came to the conclusion that this is the chief, if not the whole, value of these corpuscles in higher as well as lower animals, in all of which they are very abundant. It was known that when a wound bringing in foreign matter is inflicted on a vertebrate animal the blood-vessels became gorged in the neighbourhood and the colourless corpuscles escape through the walls of the vessels in crowds. Their business in so doing, Metschnikoff showed, is to eat up the foreign matter, and also to eat up and remove the dead, wounded tissue. He therefore called these white or colourless corpuscles ‘phagocytes,’ the eater-cells, and in his beautiful book on Inflammation, published twenty years ago, proved the extreme importance of their activity. At the same time he had shown that they eat up intrusive bacteria and other germs (see figs. 37 to 43); and his work for the last twenty years has mainly consisted in demonstrating that they are the chief, and probably the only, agents at work in either ridding the human body of an attack of disease-causing germs or in warding off even the commencement of an attack, so that the man or animal in which they are fully efficient is ‘immune’--that is to say, cannot be effectively attacked by disease-germs.

FIG. 40. FIG. 41. FIG. 42.

Fig. 40.--Phagocyte of a guinea-pig in the course of engulphing
a very mobile undulating spirillum. Fig. 41.--The same, forty
minutes later. Fig. 42.--The same taken half an hour after Fig.
41. (_From_ Metschnikoff’s “Immunity.”)]

A large kind of phagocyte of the guinea-pig, killed and stained
for microscopic examination. It shows the large spherical nucleus
and three specimens of the spirillum of relapsing-fever which
have been engulphed, and are lying within its protoplasm. They
would have been slowly digested--that is to say, dissolved by the
digestive juices within the phagocyte. (_From_ Metschnikoff’s
“Immunity.”)]