1911 Encyclopædia Britannica/Evolution
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EVOLUTION. The modern doctrine of evolution or " evolving," as opposed to that of simple creation, has been defined by Prof. James Sully in the 9th edition of this encyclopaedia as a " natural history of the cosmos including organic beings, expressed in physical terms as a mechanical process." The following exposition of the historical development of the doctrine is taken from Sully's article, and for the most part is in his own words.
In the modern doctrine of evolution the cosmic system appears as a natural product of elementary matter and its laws. The various grades of life on our planet are the natural consequences of certain physical processes involved in the gradual transformations of the earth. Conscious life is viewed as conditioned by physical (organic and more especially nervous) processes, and as evolving itself in close correlation with organic evolution. Finally, human development, as exhibited in historical and prehistorical records, is regarded as the highest and most complex result of organic and physical evolution. This modern doctrine of evolution is but an expansion and completion of those physical theories (see below) which opened the history of speculation. It differs from them in being grounded on exact and verified research. As such, moreover, it is a much more limited theory of evolution than the ancient. It does not necessarily concern itself about the question of the infinitude of worlds in space and in time. It is content to explain the origin and course of development of the world, the solar or, at most, the sidereal system which falls under our own observation. It would be difficult to say what branches of science had done most towards the establishment of this doctrine. We must content ourselves by referring to the progress of physical (including chemical) theory, which has led to the great generalization of the conservation of energy; to the discovery of the fundamental chemical identity of the matter of our planet and of other celestial bodies, and of the chemical relations of organic and inorganic bodies; to the advance of astronomical speculation respecting the origin of the solar system, &c.; to the growth of the science of geology which has necessitated the conception of vast and unimaginable periods of time in the past history of our globe, and to the rapid march of the biological sciences which has made us familiar with the simplest types and elements of organism; finally, to the development of the science of anthropology (including comparative psychology, philology, &c.), and to the vast extension and improvement of all branches of historical study.
History of the Idea of Evolution
The doctrine of evolution in its finished and definite form is a modern product. It required for its formation an amount of scientific knowledge which could only be very gradually acquired. It is vain, therefore, to look for clearly defined and systematic presentations of the idea among ancient writers. On the other hand, nearly all systems of philosophy have discussed the underlying problems. Such questions as the origin of the cosmos as a whole, the production of organic beings and of conscious minds, and the meaning of the observable grades of creation, have from the dawn of speculation occupied men's minds; and the answers to these questions often imply a vague recognition of the idea of a gradual evolution of things. Accordingly, in tracing the antecedents of the modern philosophic doctrine we shall have to glance at most of the principal systems of cosmology, ancient and modern. Yet since in these systems inquiries into the esse and fieri of the world are rarely distinguished with any precision, it will be necessary to indicate very briefly the general outlines of the system so far as they are necessary for understanding their bearing on the problems of evolution.
The problem of the origin of the world was the first to engage man's speculative activity. Nor was this line of inquiry pursued simply as a step in the more practical problem of man's final destiny. The order of ideas observable in children suggests the reflection that man began to discuss the "whence " of existence before the "whither." At first, as in the case of the child, the problem of the genesis of things was conceived anthropomorphically: the question " How did the world arise?" first shaped itself to the human mind under the form " Who made the world?" As long as the problem was conceived in this simple manner there was, of course, no room for the idea of a necessary self-conditioned evolution. Yet the first indistinct germ of such an idea appears to emerge in combination with that of creation in some of the ancient systems of theogony. Thus, for example, in the myth of the ancient Parsees, the gods Ormuzd and Ahriman are said to evolve themselves out of a primordial matter. It may be supposed that these crude fancies embody a dim recognition of the physical forces and objects personified under the forms of deities, and a rude attempt to account for their genesis as a natural process. These first unscientific ideas of a genesis of the permanent objects of nature took as their pattern the process of organic reproduction and development, and this, not only because these objects were regarded as personalities, but also because this particular mode of becoming would most impress these early observers. This same way of looking at the origin of the material world is illustrated in the Egyptian notion of a cosmic egg out of which issues the god (Phta) who creates the world.
Passing from mythology to speculation properly so called, we find in the early systems of philosophy of India theories of emanation which approach in some respects the idea of evolution. Brahma is conceived as the eternal selfexistent being, which on its material side unfolds itself to the world by gradually condensing itself to material objects through the gradations of ether, fire, water, earth and the elements. At the same time this eternal being is conceived as the all-embracing world-soul from which emanates the hierarchy of individual souls. In the later system of emanation of Sankhya there is a more marked approach to a materialistic doctrine of evolution. If, we are told, we follow the chain of causes far enough back we reach unlimited eternal creative nature or matter. Out of this " principal thing " or " original nature " all material and spiritual existence issues, and into it will return. Yet this primordial creative nature is endowed with volition with regard to its own development. Its first emanation as plastic nature contains the original soul or deity out of which all individual souls issue.
Early Greek Physicists
Passing by Buddhism, which, though teaching the periodic destruction of our world by fire, &c., does not seek to determine the ultimate origin of the cosmos, we come to those early Greek physical philosophers who distinctly set themselves to eliminate the idea of divine interference with the world by representing its origin and changes as a natural process. The early Ionian physicists, including Thales, Anaximander and Anaximenes, seek to explain the world as generated out of a primordial matter (Gr. ὕλη; hence the name " Hylozoists "), which is at the same time the universal support of things. This substance is endowed with a generative or transmutative force by virtue of which it passes into a succession of forms. They thus resemble modern evolutionists, since they regard the world with its infinite variety of forms as issuing from a simple mode of matter. More especially the cosmology of Anaximander resembles the modern doctrine of evolution in its conception of the indeterminate ( τὸ ἄπειρον ) out of which the particular forms of the cosmos are differentiated. Again, Anaximander may be said to prepare the way for more modern conceptions of material evolution by regarding his primordial substance as eternal, and by looking on all generation as alternating with destruction, each step of the process being of course simply a transformation of the indestructible substance. Once more, the notion that this indeterminate body contains potentially in itself the fundamental contraries - hot, cold, &c. - by the excretion or evolution of which definite substances were generated, is clearly a forecasting of that antithesis of potentiality and actuality which from Aristotle downwards has been made the basis of so many theories of development. In conclusion, it is noteworthy that though resorting to utterly fanciful hypotheses respecting the order of the development of the world, Anaximander agrees with modern evolutionists in conceiving the heavenly bodies as arising out of an aggregation of diffused matter, and in assigning to organic life an origin in the inorganic materials of the primitive earth (pristine mud). The doctrine of Anaximenes, who unites the conceptions of a determinate and indeterminate original substance adopted by Thales and Anaximander in the hypothesis of a primordial and all-generating air, is a clear advance on these theories, inasmuch as it introduces the scientific idea of condensation and rarefaction as the great generating or transforming agencies. For the rest, his theory is chiefly important as emphasizing the vital character of the original substance. The primordial air is conceived as animated. Anaximenes seems to have inclined to a view of cosmic evolution as throughout involving a quasi-spiritual factor. This idea of the air as the original principle and source of life and intelligence is much more clearly expressed by a later writer, Diogenes of Apollonia. Diogenes made this conception of a vital and intelligent air the ground of a teleological view of climatic and atmospheric phenomena. It is noteworthy that he sought to establish the identity of organic and inorganic matter by help of the facts of vegetal and animal nutrition. Diogenes distinctly taught that the world is of finite duration, and will be renewed out of the primitive substance.
Heraclitus again deserves a prominent place in a history of the idea of evolution. Heraclitus conceives of the incessant process of flux in which all things are involved as consisting of two sides or moments - generation and decay - which are regarded as a confluence of opposite streams. In thus making transition or change, viewed as the identity of existence and non-existence, the leading idea of his system, Heraclitus anticipated in some measure Hegel's peculiar doctrine of evolution as a dialectic process.' At the same time we may find expressed in figurative language the germs of thoughts which enter into still newer doctrines of evolution. For example, the notion of conflict (πόλεμος) as the father of all things and of harmony as arising out of a union of discords, and again of an endeavour by individual things to maintain themselves in permanence against the universal process of destruction and renovation, cannot but remind one of certain fundamental ideas in Darwin's theory of evolution.
Empedocles took an important step in the direction of modern conceptions of physical evolution by teaching that all things arise, not by transformations of some primitive form of matter, but by various combinations of a number of permanent elements. Further, by maintaining that the elements are continually being combined and separated by the two forces love and hatred, which appear to represent in a figurative way the physical forces of attraction and repulsion, Empedocles may be said to have made a considerable advance in the construction of the idea of evolution as a strictly mechanical process. It may be observed, too, that the hypothesis of a primitive compact mass (sphaerus), in which love (attraction) is supreme, has some curious points of similarity to, and contrast with, that notion of a primitive nebulous matter with which the modern doctrine of cosmic evolution usually sets out. Empedocles tries to explain the genesis of organic beings, and, according to Lange, anticipates the idea of Darwin that adaptations abound, because it is their nature to perpetuate themselves. He further recognizes a progress in the production of vegetable and animal forms, though this part of his theory is essentially crude and unscientific. More important in relation to the modern problems of evolution is his thoroughly materialistic way of explaining the origin of sensation and knowledge by help of his peculiar hypothesis of effluvia and pores. The supposition that sensation thus rests on a material process of absorption from external bodies naturally led up to the idea that plants and even inorganic subtances are precipient, and so to an indistinct recognition of organic life as a scale of intelligence.
In the theory of Atomism taught by Leucippus and Democritus we have the basis of the modern mechanical conceptions of cosmic evolution. Here the endless harmonious diversity of our cosmos, as well as of other worlds supposed to coexist with our own, is said to arise through the various combination of indivisible material elements differing in figure and magnitude only. The force which brings the atoms together in the forms of objects is inherent in the elements, and all their motions are necessary. The origin of things, which is also their substance, is thus laid in the simplest and most homogeneous elements or principles. The real world thus arising consists only of diverse combinations of atoms, having the properties of magnitude, figure, weight and hardness, all other qualities being relative only to the sentient organism. The problem of the genesis of mind is practically solved by identifying the soul, 1 This is brought out by F. Lassalle, Die Philosophie Herakleitos, p. 126.
or vital principle, with heat or fire which pervades in unequal proportions, not only man and animals, but plants and nature as a whole, and through the agitation of which by incoming effluvia all sensation arises.
Aristotle is much nearer a conception of evolution than his master Plato. It is true he sets out with a transcendent Deity, and follows Plato in viewing the creation of the cosmos as a process of descent from the more to the less perfect according to the distance from the original self-moving agency. Yet on the whole Aristotle leans to a teleological theory of evolution, which he interprets dualistually by means of certain metaphysical distinctions. Thus even his idea of the relation of the divine activity to the world shows a tendency to a pantheistic notion of a divine thought which gradually realizes itself in the process of becoming. Aristotle's distinction of form and matter, and his conception of becoming as a transition from actuality to potentiality, provides a new ontological way of conceiving the process of material and organic evolution.' To Aristotle the whole of nature is instinct with a vital impulse towards some higher manifestation. Organic life presents itself to him as a progressive scale of complexity determined by its final end, namely, man.' In some respects Aristotle approaches the modern view of evolution. Thus, though he looked on species as fixed, being the realization of an unchanging formative principle (φύσις), he seems, as Ueberweg observes, to have inclined to entertain the possibility of a spontaneous generation in the case of the lowest organisms. Aristotle's teleological conception of organic evolution often approaches modern mechanical conceptions. Thus he says that nature fashions organs in the order of their necessity, the first being those essential to life. So, too, in his psychology he speaks of the several degrees of mind as arising according to a progressive necessity. 3 In his view of touch and taste, as the two fundamental and essential senses, he may remind one of Herbert Spencer's doctrine. At the same time Aristotle precludes the idea of a natural development of the mental series by the supposition that man contains, over and above a natural finite soul inseparable from the body, a substantial and eternal principle (νοῦς) which enters into the individual from without. Aristotle's brief suggestions respect ing the origin of society and governments in the Politics show a leaning to a naturalistic interpretation of human history as a development conditioned by growing necessities.
Of Aristotle's immediate successors one deserves to be noticed here, namely, Strato of Lampsacus, who developed his master's cosmology into a system of naturalism. Strato appears to reject Aristotle's idea of an original source of movement and life extraneous to the world in favour of an immanent principle. All parts of matter have an inward plastic life whereby they can fashion themselves to the best advantage, according to their capability, though not with consciousness.
In the cosmology of the Stoics we have the germ of a monistic and pantheistic conception of evolution. All things are said to be developed out of an original being, which is at once material (fire) and spiritual (the Deity), and in turn they will dissolve back into this primordial source. At the same time the world as a developed whole is regarded as an organism which is permeated with the divine Spirit, and so we may say that the world-process is a self-realization of the divine Being. The formative principle or force of the world is said to contain the several rational germinal forms of things. Individual things are supposed to arise out of the original being, as animals and plants out of seeds. Individual souls are an efflux from the all-compassing world-soul. The necessity in the world's order is regarded by the Stoics as identical with the divine reason, and this idea is used as the basis of a teleological and optimistic view of nature. Very curious, in relation to modern evolutional ideas, is the Stoical doctrine that our world is but one of a series of exactly 1 Zeller says that through this distinction Aristotle first made possible the idea of development.
2 See this well brought out in G. H. Lewes's Aristotle, p. 187. Grote calls attention to the contrast between Plato's and Aristotle's way of conceiving the gradations of mind (Aristotle, ii. 171).
identical ones, all of which are destined to be burnt up and destroyed. The Epicurean
The Epicureans differed from the Stoics by adopting a purely mechanical view of the worldprocess. Their fundamental conception is that of Democritus; they seek to account for the formation of the cosmos, with its order and regularity, by setting out with the idea of an original (vertical) motion of the atoms, which somehow or other results in movements towards and from one another. Our world is but one of an infinite number of others, and all the harmonies and adaptations of the universe are regarded as a special case of the infinite possibilities of mechanical events. Lucretius regards the primitive atoms (first beginnings or first bodies) as seeds out of which individual things are developed. All living and sentient things are formed out of insentient atoms (e.g. worms spring out of dung). The peculiarity of organic and sentient bodies is due to the minuteness and shape of their particles, and to their special motions and combinations. So, too, mind consists but of extremely fine particles of matter, and dissolves into air when the body dies. Lucretius traces, in the fifth book of his poem, the progressive genesis of vegetal and animal forms out of the motherearth. He vaguely anticipates the modern idea of the world as a survival of the fittest when he says that many races may have lived and died out, and that those which still exist have been protected either by craft, courage or speed. Lucretius touches on the development of man out of a primitive, hardy, beast-like condition. Pregnant hints are given respecting a natural development of language which has its germs in sounds of quadrupeds and birds, of religious ideas out of dreams and waking hallucinations, and of the art of music by help of the suggestion of natural sounds. Lucretius thus recognizes the whole range of existence to which the doctrine of evolution may be applied.
In the doctrines of the Neoplatonists, of whom Plotinus is the most important, we have the worldprocess represented after the example of Plato as a series of descending steps, each being less perfect than its predecessors, since it is further removed from the first cause.' The system of Plotinus, Zellar remarks, is not strictly speaking one of emanation, since there is no communication of the divine essence to the created world; yet it resembles emanation inasmuch as the genesis of the world is conceived as a necessary physical effect, and not as the result of volition. In Proclus we find this conception of an emanation of the world out of the Deity, or the absolute, made more exact, the process being regarded as threefold-0) persistence of cause in effect, (2) the departure of effect from cause, and (3) the tendency of effect to revert to its cause.
The speculations of the fathers respecting the origin and course of the world seek to combine Christian ideas of the Deity with doctrines of Greek philosophy. The common idea of the origin of things is that of an absolute creation of matter and mind alike. The course of human history is regarded by those writers who are most concerned to refute Judaism as a progressive divine education. Among the Gnostics we meet with the hypothesis of emanation, as, for example, in the curious cosmic theory of Valentinus.
Early Schoolmen. - In the speculative writings of the middle ages, including those of the schoolmen, we find no progress towards a more accurate and scientific view of nature. The cosmology of this period consists for the most part of the Aristotelian teleological view of nature combined with the Christian idea of the Deity and His relation to the world. In certain writers, however, there appears a more elaborate transformation of the doctrine of creation into a system of emanation. According to John Scotus Erigena, the nothing out of which the world is created is the divine essence. Creation is the act by which God passes through the primordial causes, or universal ideas, into the region of particular things (processio), in order finally to return to himself (reversio). The transition from the 4 Zeller observes that this scale of decreasing perfection is a necessary consequence of the idea of a transcendent deity.
universal to the particular is of course conceived as a descent or degradation. A similar doctrine of emanation is to be found in the writings of Bernhard of Chartres, who conceives the process of the unfolding of the world as a movement in a circle from the most general to the individual, and from this back to the most general. This movement is said to go forth from God to the animated heaven, stars, visible world and man, which represent decreasing degrees of cognition.
Elaborate doctrines of emanation, largely based on Neoplatonic ideas, are also propounded by some of the Arabic philosophers, as by Farabi and Avicenna. The leading thought is that of a descending series of intelligences, each emanating from its predecessor, and having its appropriate region in the universe.
In the Jewish speculations of the middle ages may be found curious forms of the doctrine of emanations uniting the Biblical idea of creation with elements drawn from the Persians and the Greeks. In the later and developed form of the Kabbala, the origin of the world is represented as a gradually descending emanation of the lower out of the higher. Among the philosophic Jews, the Spanish Avicebron, in his Fons Vitae, expounds a curious doctrine of emanation. Here the divine will is viewed as an efflux from the divine wisdom, as the intermediate link between God, the first substance, and all things, and as the fountain out of which all forms emanate. At the same time all forms, including the higher intelligible ones, are said to have their existence only in matter. Matter is the one universal substance, body and mind being merely specifications of this. Thus Avicebron approaches, as Salomon Munk observes,' a pantheistic conception of the world, though he distinctly denies both matter and form to God.
Passing now to the later schoolmen, a bare mention must be made of Thomas Aquinas, who elaborately argues for the absolute creation of the world out of nothing, and of Albertus Magnus, who reasons against the Aristotelian idea of the past eternity of the world. More importance attaches to Duns Scotus, who brings prominently forward the idea of a progressive development in nature by means of a process of determination. The original substance of the world is the materia primo-prima, which is the immediate creation of the Deity. This serves Duns Scotus as the most universal basis of existence, all angels having material bodies. This matter is differentiated into particular things (which are not privations but perfections) through the addition of an individualizing principle (haecceitas) to the universal (quidditas). The whole world is represented by the figure of a tree, of which the seeds and roots are the first indeterminate matter, the leaves the accidents, the twigs and branches corruptible creatures, the blossoms the rational soul, and the fruit pure spirits or angels. It is also described as a bifurcation of two twigs, mental and bodily creation out of a common root. One might almost say that Duns Scotus recognizes the principle of a gradual physical evolution, only that he chooses to represent the mechanism by which the process is brought about by means of quaint scholastic fictions.
Revival of Learning
The period of the revival of learning, which was also that of a renewed study of nature, is marked by a considerable amount of speculation respecting the origin of the universe. In some of these we see a return to Greek theories, though the influence of physical discoveries, more especially those of Copernicus, Kepler and Galileo, is distinctly traceable.
An example of a return to early Greek speculation is to be met with in Bernardino Telesio. By this writer the world is explained as a product of three principles - dead matter, and two active forces, heat and cold. Terrestrial things arise through a confluence of heat, which issues from the heavens, and cold, which comes from the earth. Both principles have sensibility, and thus all products of their collision are sentient, that is, feel pleasure and pain. The superiority of animals to plants and metals in the possession of special organs of sense is connected with the greater complexity and heterogeneity of their structure.
1 Mélanges de philosophie juive et arabe, p. 225.
In the system of Giordano Bruno, who sought to construct a philosophy of nature on the basis of new scientific ideas, more particularly the doctrine of Copernicus, we find the outlines of a theory of cosmic evolution conceived as an essentially vital process. Matter and form are here identified, and the evolution of the world is presented as the unfolding of the world-spirit to its perfect forms according to the plastic substratum (matter) which is but one of its sides. This process of change is conceived as a transformation, in appearance only, of the real unchanging substance (matter and form) . All parts of matter are capable of developing into all forms; thus the materials of the table and chair may under proper circumstances be developed to the life of the plant or of the animal. The elementary parts of existence are the minima, or monads, which are at once material and mental. On their material side they are not absolutely unextended, but spherical. Bruno looked on our solar system as but one out of an infinite number of worlds. His theory of evolution is essentially pantheistic, and he does not employ his hypothesis of monads in order to work out a more mechanical conception.
A word must be given to one of Bruno's contemporary compatriots, namely Campanella, who gave poetic expression to that system of universal vitalism which Bruno developed. He argues, from the principle quicquid est in effectibus esse et in causis, that the elements and the whole world have sensation, and thus he appears to derive the organic part of nature out of the so-called " inorganic." Boehme. - Another writer of this transition period deserves a passing reference here, namely, Jacob Boehme the mystic, who by his conception of a process of inner diremption as the essential character of all mind, and so of God, prepared the way for later German theories of the origin of the world as the self-differentiation and self-externalization of the absolute spirit.
Hobbes and Gassendi
The influence of an advancing study of nature, which was stimulated if not guided by Bacon's writings, is seen in the more careful doctrines of materialism worked out almost simultaneously by Hobbes and Gassendi. These theories, however, contain little that bears directly on the hypothesis of a natural evolution of things. In the view of Hobbes, the difficulty of the genesis of conscious minds is solved by saying that sensation and thought are part of the reaction of the organism on external movement. Yet Hobbes appears (as Clarke points out) to have vaguely felt the difficulty; and in a passage of his Physics (chap. 25, sect. 5) he says that the universal existence of sensation in matter cannot be disproved, though he shows that when there are no organic arrangements the mental side of the movement (phantasma) is evanescent. The theory of the origin of society put forth by Hobbes, though directly opposed in most respects to modern ideas of social evolution, deserves mention here by reason of its enforcing that principle of struggle (bellum omnium contra omnes) which has played so conspicuous a part in the modern doctrine of evolution. Gassendi, with some deviations, follows Epicurus in his theory of the formation of the world. The world consists of a finite number of atoms, which have in their own nature a self-moving force or principle. These atoms, which are the seeds of all things, are, however, not eternal but created by God. Gassendi distinctly argues against the existence of a world-soul or a principle of life in nature.
In the philosophy of Descartes we meet with a dualism of mind and matter which does not easily lend itself to the conception of evolution. His doctrine that consciousness is confined to man, the lower animals being unconscious machines (automata), excludes all idea of a progressive development of mind. Yet Descartes, in his Principia Philosophiae, laid the foundation of the modern mechanical conception of nature and of physical evolution. In the third part of this work he inclines to a thoroughly natural hypothesis respecting the genesis of the physical world, and adds in the fourth part that the same kind of explanation might be applied to the nature and formation of plants and animals. He is indeed careful to keep right with the orthodox doctrine of creation by saying that he does not believe the world actually arose in this mechanical way out of the three kinds of elements which he here supposes, but that he simply puts out his hypothesis as a mode of conceiving how it might have arisen. Descartes's account of the mind and its passions is thoroughly materialistic, and to this extent he works in the direction of a materialistic explanation of the origin of mental life.
In Spinoza's pantheistic theory of the world, which regards thought and extension as but two sides of one substance, the problem of becoming is submerged in that of being. Although Spinoza's theory attributes a mental side to all physical events, he rejects all teleological conceptions and explains the order of things as the result of an inherent necessity. He recognizes gradations of things according to the degree of complexity of their movements and that of their conceptions. To Spinoza (as Kuno Fischer observes) man differs from the rest of nature in the degree only and not in the kind of his powers. So far Spinoza approaches the conception of evolution. He may be said to furnish a further contribution to a metaphysical conception of evolution in his view of all finite individual things as the infinite variety to which the unlimited productive power of the universal substance gives birth. Sir F. Pollock has taken pains to show how nearly Spinoza approaches certain ideas contained in the modern doctrine of evolution, as for example that of sell-preservation as the determining force in things.
In Locke we find, with a retention of certain antievolutionist ideas, a marked tendency to this mode of viewing the world. To Locke the universe is the result of a direct act of creation, even matter being limited in duration and created. Even if matter were eternal it would, he thinks, be incapable of producing motion; and if motion is itself conceived as eternal, thought can never begin to be. The first eternal being is thus spiritual or " cogitative," and contains in itself all the perfections that can ever after exist. He repeatedly insists on the impossibility of senseless matter putting on sense.' Yet while thus placing himself at a point of view opposed to that of a gradual evolution of the organic world, Locke prepared the way for this doctrine in more ways than one. First of all, his genetic method as applied to the mind's ideas - which laid the foundations of English analytical psychology - was a step in the direction of a conception of mental life as a gradual evolution. Again he works towards the same end in his celebrated refutation of the scholastic theory of real specific essences. In this argument he emphasizes the vagueness of the boundaries which mark off organic species with a view to show that these do not correspond to absolutely fixed divisions in the objective world, that they are made by the mind, not by nature. 2 This idea of the continuity of species is developed more fully in a remarkable passage (Essay, bk. iii. ch. vi. § 12), where he is arguing in favour of the hypothesis, afterwards elaborated by Leibnitz, of a graduated series of minds (species of spirits) from the Deity down to the lowest animal intelligence. He here observes that " all quite down from us the descent is by easy steps, and a continued series of things, that in each remove differ very little from one another." Thus man approaches the beasts, and the animal kingdom is nearly joined with the vegetable, and so on down to the lowest and " most inorganical parts of matter." Finally, it is to be observed that Locke had a singularly clear view of organic arrangements (which of course he explained according to a theistic teleology) as an adaptation to the circumstances of the environment or to " the neighbourhood of the bodies that surround us." Thus he suggests that man has not eyes of a microscopic delicacy, because he would receive no great advantage from such acute organs, since though adding indefinitely to his speculative knowledge of the physical world they would 1 Yet he leaves open the question whether the Deity has annexed thought to matter as a faculty, or whether it rests on a distinct spiritual principle.
Locke half playfully touches on certain monsters, with respect to which it is difficult to determine whether they ought to be called men. (Essay, book iii. ch. vi. sect. 26, 27.) not practically benefit their possessor (e.g. by enabling him to avoid things at a convenient distance).3 Idea of Progress in History. - Before leaving the 17th century we must just refer to the writers who laid the foundations of the essentially modern conception of human history as a gradual upward progress. According to Flint, 4 there were four men who in this and the preceding century seized and made prominent this idea, namely, Bodin, Bacon, Descartes and Pascal. The former distinctly argues against the idea of a deterioration of man in the past. In this way we see that just as advancing natural science was preparing the way for a doctrine of physical evolution, so advancing historical research was leading to the application of a similar idea to the collective human life. English Writers of the 28th Centur
The theological discussions which make up so large a part of the English speculation of the 18th century cannot detain us here. There is, however, one writer who sets forth so clearly the alternative suppositions respecting the origin of the world that he claims a brief notice. We refer to David Hume. In his Dialogues concerning Natural Religion he puts forward tentatively, in the person of one of his interlocutors, the ancient hypothesis that since the world resembles an animal or vegetal organism rather than a machine, it might more easily be accounted for by a process of generation than by an act of creation. Later on he develops the materialistic view of Epicurus, only modifying it so far as to conceive of matter as finite. Since a finite number of particles is only susceptible of finite transpositions, it must happen (he says), in an eternal duration that every possible order or position will be tried an infinite number of times, and hence this world is to be regarded (as the Stoics maintained) as an exact reproduction of previous worlds. The speaker seeks to make intelligible the appearance of art and contrivance in the world as a result of a natural settlement of the universe (which passes through a succession of chaotic conditions) into a stable condition, having a constancy in its forms, yet without its several parts losing their motion and fluctuation.
French Writers of the 18th Century
Let us now pass to the French writers of the 18th century. Here we are first struck by the results of advancing physical speculation in their bearing on the conception of the world. Careful attempts, based on new scientific truths, an made to explain the genesis of the world as a natural process. Maupertuis, who, together with Voltaire, introduced the new idea of the universe as based on Newton's discoveries, sought to account for the origin of organic things by the hypothesis of sentient atoms. Buffon the naturalist speculated, not only on the structure and genesis of organic beings, but also on the course of formation of the earth and solar system, which he conceived after the analogy of the development of organic beings out of seed. Diderot, too, in his varied intellectual activity, found time to speculate on the genesis of sensation and thought out of a combination of matter endowed with an elementary kind of sentience. De la Mettrie worked out a materialistic doctrine of the origin of things, according to which sensation and consciousness are nothing but a development out of matter. He sought (L'Homme-machine) to connect man in his original condition with the lower animals, and emphasized (L'Homme-plante) the essential unity of plan of all living things. Helvetius, in his work on man, referred all differences between our species and the lower animals to certain peculiarities of organization, and so prepared the way for a conception of human development out of lower forms as a process of physical evolution. Charles Bonnet met the difficulty of the origin of conscious beings much in the same way as Leibnitz, by the supposition of eternal minute organic bodies to which are attached immortal souls. Yet though in this way opposing himself to the method of the modern doctrine of evolution, he aided the development of this doctrine by his view of the organic world as an ascending 3 A similar coincidence between the teleological and the modern evolutional way of viewing things is to be met with in Locke's account of the use of pain in relation to the preservation of our being (bk. ii. ch. vii. sect. 4).
4 Philosophy of History (1893), p. 103, where an interesting sketch of the growth of the idea of progress is to be found.
scale from the simple to the complex. Robinet, in his treatise De la nature, worked out the same conception of a gradation in organic existence, connecting this with a general view of nature as a progress from the lowest inorganic forms of matter up to man. The process is conceived as an infinite series of variations or specifications of one primitive and common type. Man is the chef-d'oeuvre of nature, which the gradual progression of beings was to have as its last term, and all lower creations are regarded as pre-conditions of man's existence, since nature " could only realize the human form by combining in all imaginable ways each of the traits which was to enter into it." The formative force in this process of evolution (or " metamorphosis ") is conceived as an intellectual principle (idee generatrice). Robinet thus laid the foundation of that view of the world as wholly vital, and as a progressive unfolding of a spiritual formative principle, which was afterwards worked out by Schelling. It is to be added that Robinet adopted a thorough-going materialistic view of the dependence of mind on body, going even to the length of assigning special nerve-fibres to the moral sense. The system of Holbach seeks to provide a consistent materialistic view of the world and its processes. Mental operations are identified with physical movements, the three conditions of physical movement, inertia, attraction and repulsion, being in the moral world self-love, love and hate. He left open the question whether the capability of sensation belongs to all matter, or is confined to the combinations of certain materials. He looked on the actions of the individual organism and of society as determined by the needs of self-preservation. He conceived of man as a product of nature that had gradually developed itself from a low condition, though he relinquished the problem of the exact mode of his first genesis and advance as not soluble by data of experience. Holbach thus worked out the basis of a rigorously materialistic conception of evolution.
The question of human development which Holbach touched on was one which occupied many minds both in and out of France during the 18th century, and more especially towards its close. The foundations of this theory of history as an upward progress of man out of a barbaric and animal condition were laid by Vico in his celebrated work Principii di scienza nuova. In France the doctrine was represented by Turgot and Condorcet.
German Writers of the 18th Centur
Leibnitz. - In Leibnitz we find, if not a doctrine of evolution in the strict sense, a theory of the world which is curiously related to the modern doctrine. The chief aim of Leibnitz is no doubt to account for the world in its static aspect as a co-existent whole, to conceive the ultimate reality of things in such a way as to solve the mystery of mind and matter. Yet by his very mode of solving the problem he is led on to consider the nature of the world-process. By placing substantial reality in an infinite number of monads whose essential nature is force or activity, which. is conceived as mental (representation), Leibnitz was carried on to the explanation of the successive order of the world. He prepares the way, too, for a doctrine of evolution by his monistic idea of the substantial similarity of all things, inorganic and organic, bodily and spiritual, and still more by his conception of a perfect gradation of existence from the lowest " inanimate " objects, whose essential activity is confused representation, up to the highest organized beingman - with his clear intelligence.' Turning now to Leibnitz's conception of the world as a process, we see first that he supplies, in his notion of the underlying reality as force which is represented as spiritual (quelque chose d'analogique au sentiment et a Tappan), both a mechanical and a teleological explanation of its order. More than this, Leibnitz supposes that the activity of the monads takes the form of a self-evolution. It is the following out of an inherent tendency or impulse to a series of changes, all of which were virtually pre-existent, and this process cannot be interfered with from without. As the individual monad, so the whole system which makes up the world is a gradual 1 G. H. Lewes points out that Leibnitz is inconsistent in his account of the intelligence of man in relation to that of lower animals, since when answering Locke he no longer regards these as differing in degree only.
development. In this case, however, we cannot say that each step goes out of the other as in that of individual development. Each monad is an original independent being, and is determined to take this particular point in the universe, this place in the scale of beings. We see how different this metaphysical conception is from that scientific notion of cosmic evolution in which the lower stages are the antecedents and conditions of the higher. It is probable that Leibnitz's notion of time and space, which approaches Kant's theory, led him to attach but little importance to the successive order of the world. Leibnitz, in fact, presents to us an infinite system of perfectly distinct though parallel developments, which on their mental side assume the aspect of a scale, not through any mutual action, but solely through the determination of the Deity. Even this idea, however, is incomplete, for Leibnitz fails to explain the physical aspect of development. Thus he does not account for the fact that organic beings - which have always existed as preformations (in the case of animals as animaux spermatiques) - come to be developed under given conditions. Yet Leibnitz prepared the way for a new conception of organic evolution. The modern monistic doctrine, that all material things consist of sentient elements, and that consciousness arises through a combination of these, was a natural transformation of Leibnitz's theory.2 Lessing. - Of Leibnitz's immediate followers we may mention Lessing, who in his Education of the Human Race brought out the truth of the process of gradual development underlying: human history, even though he expressed this in a form inconsistent with the idea of a spontaneous evolution.
Herder, on the other hand, Lessing's contemporary,. treated the subject of man's development in a thoroughly naturalistic spirit. In his Ideen zur Philosophic der Geschichte, Herder adopts Leibnitz's idea of a graduated scale of beings, at the same time conceiving of the lower stages as the conditions, of the higher. Thus man is said to be the highest product of nature, and as such to be dependent on all lower products. All material things are assimilated to one another as organic, the vitalizing principle being inherent in all matter. The development of man is explained in connexion with that of the earth, and in relation to climatic variations, &c. Man's mental faculties are viewed as related to his organization, and as developed under the pressure of the necessities of life.3 Kant. - Kant's relation to the doctrine of evolution is a many-sided one. In the first place, his peculiar system of subjective idealism, involving the idea that time is but a mental form to which there corresponds nothing in the sphere of noiimenal reality, serves to give a peculiar philosophical interpretation to every doctrine of cosmic evolution. Kant, like Leibnitz, seeks to reconcile the mechanical and teleological views of nature, only he assigns to these different spheres. The order of the inorganic world is explained by properly physical causes. In his Naturgeschichte des Himmels, in which he anticipated the nebular theory afterwards more fully developed by Laplace, Kant sought to explain the genesis of the cosmos as a product of physical forces and laws. The worlds, or systems of worlds, which fill infinite space are continually being formed and destroyed. Chaos passes by a process of evolution into a cosmos, and this again into chaos. So far as the evolution of the solar system is concerned, Kant held these mechanical causes as adequate. For the world as a whole, however, he postulated a beginning in time (whence his use of the word creation), and further supposed that the impulse of organization which was conveyed to chaotic matter by the Creator issued from a central point in the infinite space spreading gradually outwards. 4 While 2 Both Lewes and du Bois Reymond have brought out the points of contact between Leibnitz's theory of monads and modern biological speculations (Hist. of Phil. ii. 287, and Leibnitzsche Gedanken in der modernen Naturwissenschaft, p. 23 seq.).
For Herder's position in relation to the modern doctrine of evolution see F. von Barenbach's Herder als Vorgdnger Darwins, a work which tends to exaggerate the proximity of the two writers.
4 Kant held it probable that other planets besides our earth are inhabited, and that their inhabitants form a scale of beings, their perfection increasing with the distance of the planet which they inhabit from the sun.
in his cosmology Kant thus relies on mechanical conceptions, in his treatment of organic life his mind is, on the contrary, dominated by teleological ideas. An organism was to him something controlled by a formative organizing principle. It was natural, therefore, that he rejected the idea of a spontaneous generation of organisms (which was just then being advocated by his friend Forster), not only as unsupported by experience but as an inadequate hypothesis. Experience forbids our excluding organic activity from natural causes, also our excluding intelligence from purposeful (zwecktdtigen) causes; hence experience forbids our defining the fundamental force or first cause out of which living creatures arose.' Just as Kant thus sharply marks off the regions of the inorganic and the organic, so he sets man in strong opposition to the lower animals. His ascription to man of a unique faculty, free-will, forbade his conceiving our species as a link in a graduated series of organic developments. In his doctrine of human development he does indeed recognize an early stage of existence in which our species was dominated by sensuous enjoyment and instinct. He further conceives of this stage as itself a process of (natural) development, namely, of the natural disposition of the species to vary in the greatest possible manner so as to preserve its unity through a process of self-adaptation (Anarten) to climate. This, he says, must not be conceived as resulting from the action of external causes, but is due to a natural disposition (Anlage). From this capability of natural development (which already involves a teleological idea) Kant distinguishes the power of moral self-development or selfliberation from the dominion of nature, the gradual realization of which constitutes human history or progress. This moral development is regarded as a gradual approach to that rational, social and political state in which will be realized the greatest possible quantity of liberty. Thus Kant, though he appropriated and gave new form to the idea of human progress, conceived of this as wholly distinct from a natural (mechanical) process. In this particular, as in his view of organic actions, Kant distinctly opposed the idea of evolution as one universal process swaying alike the physical and the moral world. Schelling
In the earlier writings of Schelling, containing the philosophy of identity, existence is represented as a becoming, or process of evolution. Nature and mind (which are the two sides, or polar directions, of the one absolute) are each viewed as an activity advancing by an uninterrupted succession of stages. The side of this process which Schelling worked out most completely is the negative side, that is, nature. Nature is essentially a process of organic self-evolution. It can only be understood by subordinating the mechanical conception to the vital, by conceiving the world as one organism animated by a spiritual principle or intelligence (Weltseele). From this point of view the processes of nature from the inorganic up to the most complex of the organic become stages in the self-realization of nature. All organic forms are at bottom but one organization, and the inorganic world shows the same formative activity in various degrees or potences. Schelling conceives of the gradual self-evolution of nature in a succession of higher and higher forms as brought about by a limitation of her infinite productivity, showing itself in a series of points of arrest. The detailed exhibition of the organizing activity of nature in the several processes of the organic and inorganic world rests on a number of fanciful and unscientific ideas. Schelling's theory is a bold attempt to revitalize nature in the light of growing physical and physiological science, and by so doing to comprehend the unity of the world under the idea of one principle of organic development. His highly figurative language might leave us in doubt how far he conceived the higher stages of this evolution of nature as following the lower in time. In the introduction to his work Von der Weltseele, however, he argues in favour of the possibility of a transmutation of species in periods incommensurable with ours. The evolution of mind (the positive pole) proceeds by 1 Kant calls the doctrine of the transmutation of species " a hazardous fancy of the reason." Yet, as Strauss and others have shown, Kant's mind betrayed a decided leaning at times to a more mechanical conception of organic forms as related by descent.
way of three stages - theoretic, practical and aesthetical activity. Schelling's later theosophic speculations do not specially concern us here.
Followers of Schelling
Of the followers of Schelling a word or two must be said. Heinrich Steffens, in his Anthropologie, seeks to trace out the origin and history of man in connexion with a general theory of the development of the earth, and this again as related to the formation of the solar system. All these processes are regarded as a series of manifestations of a vital principle in higher and higher forms. Oken, again, who carries Schelling's ideas into the region of biological science, seeks to reconstruct the gradual evolution of the material world out of original matter, which is the first immediate appearance of God, or the absolute. This process is an upward one, through the formation of the solar system and of our earth with its inorganic bodies, up to the production of man. The process is essentially a polar linear action, or differentiation from a common centre. By means of this process the bodies of the solar system separate themselves, and the order of cosmic evolution is repeated in that of terrestrial evolution. The organic world (like the world as a whole) arises out of a primitive chaos, namely, the infusorial slime. A somewhat similar working out of Schelling's idea is to be found in H. C. Oersted's work entitled The Soul in Nature (Eng. trans.). Of later works based on Schelling's doctrine of evolution mention may be made of the volume entitled Natur and Idee, by G. F. Carus. According to this writer, existence is nothing but a becoming, and matter is simply the momentary product of the process of becoming, while force is this process constantly revealing itself in these products. Hegel
Like Schelling, Hegel conceives the problem of existence as one of becoming. He differs from him with respect to the ultimate motive of that process of gradual evolution which reveals itself alike in nature and in mind. With Hegel the absolute is itself a dialectic process which contains within itself a principle of progress from difference to difference and from unity to unity. " This process (W. Wallace remarks) knows nothing of the distinctions between past and future, because it implies an eternal present." This conception of an immanent spontaneous evolution is applied alike both to nature and to mind and history. Nature to Hegel is the idea in the form of hetereity; and finding itself here it has to remove this exteriority in a progressive evolution towards an existence for itself in life and mind. Nature (says Zeller) is to Hegel a system of gradations, of which one arises necessarily out of the other, and is the proximate truth of that out of which it results. There are three stadia, or moments, in this process of nature - (i) the mechanical moment, or matter devoid of individuality; (2) the physical moment, or matter which has particularized itself in bodies - the solar system; and (3) the organic moment, or organic beings, beginning with the geological organism - or the mineral kingdom, plants and animals. Yet this process of development is not to be conceived as if one stage is naturally produced out of the other, and not even as if the one followed the other in time. Only spirit has a history; in nature all forms are contemporaneous.2 Hegel's interpretation of mind and history as a process of evolution has more scientific interest than his conception of nature. His theory of the development of free-will (the objective spirit), which takes its start from Kant's conception of history, with its three stages of legal right, morality as determined by motive and instinctive goodness (Sittlichkeit), might almost as well be expressed in terms of a thoroughly naturalistic doctrine of human development. So, too, some of his conceptions respecting the development of art and religion (the absolute spirit) lend themselves to a similar interpretation. Yet while, in its application to history, Hegel's theory of evolution has points of resemblance with those doctrines which seek to explain the worldprocess as one unbroken progress occurring in time, it constitutes on the whole a theory apart and sui generis. It does not conceive of the organic as succeeding on the inorganic, or of conscious life 2 Hegel somewhere says that the question of the eternal duration of the world is unanswerable: time as well as space can be predicated of finitudes only.
as conditioned in time by lower forms. In this respect it resembles Leibnitz's idea of the world as a development; the idea of evolution is in each case a metaphysical as distinguished from a scientific one. Hegel gives a place in his metaphysical system to the mechanical and the teleological views; yet in his treatment of the world as an evolution the idea of end or purpose is the predominant one.
Of the followers of Hegel who have worked out his peculiar idea of evolution it is hardly necessary to speak. A bare reference may be made to J. K. F. Rosenkranz, who in his work Hegel's Naturphilosophie seeks to develop Hegel's idea of an earthorganism in the light of modern science, recognizing in crystallization the morphological element.
Of the other German philosophers immediately following Kant, there is only one who calls for notice here, namely, Arthur Schopenhauer. This writer, by his conception of the world as will which objectifies itself in a series of gradations from the lowest manifestations of matter up to conscious man, gives a slightly new shape to the evolutional view of Schelling, though he deprives this view of its optimistic character by denying any co-operation of intelligence in the world-process. In truth, Schopenhauer's conception of the world as the activity of a blind force is at bottom a materialistic and mechanical rather than a spiritualistic and teleological theory. Moreover, Schopenhauer's subjective idealism, and his view of time as something illusory, hindered him from viewing this process as a sequence of events in time. Thus he ascribes eternity of existence to species under the form of the " Platonic ideas." As Ludwig Noire observes, Schopenhauer has no feeling for the problem of the origin of organic beings. He says Lamarck's original animal is something metaphysical, not physical, namely, the will to live. " Every species (according to Schopenhauer) has of its own will, and according to the circumstances under which it would live, determined its form and organization, - yet not as something physical in time, but as something metaphysical out of time." Von Baer. - Be fore leaving the German speculation of the first half of the century, a word must be said of von Baer, to whose biological contributions we shall refer later in this article, who recognized in the law of development the law of the universe as a whole. In his Entwickelungsgeschichte der Thiere (p. 264) he distinctly tells us that the law of growing individuality is " the fundamental thought which goes through all forms and degrees of animal development and all single relations. It is the same thought which collected in the cosmic space the divided masses into spheres, and combined these to solar systems; the same which caused the weather-beaten dust on the surface of our metallic planet to spring forth into living forms." Von Baer thus prepared the way for Herbert Spencer's generalization of the law of organic evolution as the law of all evolution.
As we arrive at the r 9th century, though yet before the days of Darwin, biology is already beginning to affect the general aspect of thought. It might suffice to single out the influence of Auguste Comte, as the last great thinker who wrote before Darwinism began to permeate philosophic speculation. Though Comte did not actually contribute to a theory of cosmic organic evolution, he helped to lay the foundations of a scientific conception of human history as a natural process of development 'determined by general laws of human nature together with the accumulating influences of the past. Comte does not recognize that this process is aided by any increase of innate capacity; on the contrary, progress is to him the unfolding of fundamental faculties of human nature which always pre-existed in a latent condition; yet he may perhaps be said to have prepared the way for the new conception of human progress by his inclusion of mental laws under biology. Development of the Biological Doctrine
In the 19th century the doctrine of evolution received new biological contents and became transformed from a vague, partly metaphysical theory to the dominant modern conception. At this point it is convenient to leave the guidance of Professor J. Sully and to follow closely T. H. Huxley, who in the 9th edition of this encyclopaedia traced the history of the growth of the biological idea of evolution from its philosophical beginnings to its efflorescence in Charles Darwin.
In the earlier half of the 18th century the term " evolution " was introduced into biological writings in order to denote the mode in which some of the most eminent physiologists of that time conceived that the generation of living things took place; in opposition to the hypothesis advocated, in the preceding century, by W. Harvey in that remarkable work 1 which would give him a claim to rank among the founders of biological science, even had he not been the discoverer of the circulation of the blood.
One of Harvey's prime objects is to defend and establish, on the basis of direct observation, the opinion already held by Aristotle, that, in the higher animals at any rate, the formation of the new organism by the process of generation takes place, not suddenly, by simultaneous accretion of rudiments of all or the most important of the organs of the adult, nor by sudden metamorphosis of a formative substance into a miniature of the whole, which subsequently grows, but by epigenesis, or successive differentiation of a relatively homogeneous rudiment into the parts and structures which are characteristic of the adult.
" Et primo, quidem, quoniam per epigenesin sive partium superexorientium additamentum pullum fabricari certum est : quaenam pars ante alias omnes exstruatur, et quid de illa ejusque generandi modo observandum veniat, dispiciemus. Ratum sane est et in ovo manifeste apparet quod Aristoteles de perfectorum animalium generatione enuntiat : nimirum, non omnes partes simul fieri, sed ordine aliam post aliam; primumque existere particulam genitalem, cujus virtute postea (tanquam ex principio quodam) reliquae omnes partes prosiliant. Qualem in plantarum seminibus (fabis, puta, aut glandibus) gemmam sive apicem protuberantem cernimus, totius futurae arboris principium. Estque haec particula velut filius emancipatus seorsumque collocatus, et principium per se vivens; unde postea membrorum ordo describitur; et quaecunque ad absolvendum animal pertinent, disponuntur. 2 Quoniam enim nulla pars se ipsam general; sed postquam generata est, se ipsam jam auget; ideo eam primum oriri necesse est, quae principium augendi contineat (sive enim planta, sive animal est, aeque omnibus inest quod vim habeat vegetandi, sive nutriendi), 3 simulque reliquas omnes partes suo quamque ordine distinguat et formet; proindeque in eader primogenita particula anima primario inest, sensus, motusque, et totius vitae auctor et principium." (Exercitatio 51.) Harvey proceeds to contrast this view with that of the " Medici," or followers of Hippocrates and Galen, who, " badly philosophizing," imagined that the brain, the heart, and the liver were simultaneously first generated in the form of vesicles; and, at the same time, while expressing his agreement with Aristotle in the principle of epigenesis, he maintains that it is the blood which is the primal generative part, and not, as Aristotle thought, the heart.
In the latter part of the 17th century the doctrine of epigenesis thus advocated by Harvey was controverted on the ground of direct observation by M. Malpighi, who affirmed that the body of the chick is to be seen in the egg before the punctum sanguineum makes it appearance. But from this perfectly correct observation a conclusion which is by no means warranted was drawn, namely, that the chick as a whole really exists in the egg antecedently to incubation; and that what happens in the course of the latter process is no addition of new parts, " alias post alias natas," as Harvey puts it, but a simple expansion or unfolding of the organs which already exist, though they are too small and inconspicuous to be discovered. The weight of Malpighi's observations therefore fell into the scale of that doctrine which Harvey terms metamorphosis, in contradistinction to epigenesis. The views of Malpighi were warmly welcomed on philosophical grounds by Leibnitz, 4 who found in them a support to his 1 The Exercitationes de generatione animalium, which Dr George Ent extracted from him and published in 1651.
2 De generatione animalium, lib. ii. cap. x.
3 De generatione animalium, lib. ii. cap. iv.
4 " Cependant, pour revenir aux formes ordinaires ou aux Ames materielles, cette duree qu'il leur faut attribuer, a la place de celle qu'on avoit attribuee aux atomes pourroit faire douter si elles ne vont pas de corps en corps; ce qui seroit la metempsychose, a peu pros comme quelques philosophes ont cru la transmission du mouvement hypothesis of monads, and by Nicholas Malebranche; while, in the middle of the r8th century, not only speculative considerations, but a great number of new and interesting observations on the phenomena of generation, led the ingenious Charles Bonnet and A. von Haller, the first physiologist of the age, to adopt, advocate and extend them.
Bonnet affirms that, before fecundation, the hen's egg contains an excessively minute but complete chick; and that fecundation and incubation simply cause this germ to absorb nutritious matters, which are deposited in the interstices of the elementary structures of which the miniature chick, or germ, is made up. The consequence of this intussusceptive growth is the " development " or " evolution " of the germ into the visible bird. Thus an organized individual (tout organise) " is a composite body consisting of the original, or elementary, parts and of the matters which have been associated with them by the aid of nutrition "; so that, if these matters could be extracted from the individual (tout), it would, so to speak, become concentrated in a point, and would thus be restored to its primitive condition of a germ; " just as, by extracting from a bone the calcareous substance which is the source of its hardness, it is reduced to its primitive state of gristle or membrane."2 " Evolution " and " development " are, for Bonnet, synonymous terms; and since by " evolution " he means simply the expansion of that which was invisible into visibility, he was naturally led to the conclusion, at which Leibnitz had arrived by a different line of reasoning, that no such thing as generation, in the proper sense of the word exists in nature. The growth of an organic being is simply a process of enlargement, as a particle of dry gelatine may be swelled up by the intussusception of water; its death is a shrinkage, such as the swelled jelly might undergo on desiccation. Nothing really new is produced in the living world, but the germs which develop have existed since the beginning of things; and nothing really dies, but, when what we call death takes place, the living thing shrinks back into its germ state.3 et celle des especes. Mais cette imagination est bien eloignee de la nature des choses. 11 n'y a point de tel passage; et c'est ici ou les transformations de Messieurs Swammerdam, Malpighi, et Leewenhoek, qui sont des plus excellens observateurs de notre tems, sont venues a mon secours et m'ont fait admettre plus aisement, que 1'animal, et toute autre substance organisee ne commence point lorsque nous le croyons, et que sa generation apparente n'est qu'un developpement et une espece d'augmentation. Aussi ai-je remarque que l'auteur de la Recherche de la verite, M. Regis, M. Hartseeker, et d'autres habiles hommes n'ont pas ete fort eloignes de ce sentiment." Leibnitz, Systeme nouveau de la nature (1695). The doctrine of " Emboitement " is contained in the Considerations sur le principe de vie (1705); the preface to the Theodicee (1710); and the Principes de la nature et de la grace (§ 6) (1718).
' " Il est vrai que la pensee la plus raisonnable et la plus conforme a l'experience sur cette question tres difficile de la formation du foetus; c'est que les enfans sont déjà presque tout formes avant meme l'action par laquelle ils sont conqus; et que leurs meres ne font que leur donner l'accroissement ordinaire dans le temps de la grossesse." De la recherche de la verite, livre ii. chap. vii. p. 334 (7th ed., 1721).
2 Considerations sur les corps organises, chap. x.
Bonnet had the courage of his opinions, and in the Palingenesie philosophique, part vi. chap. iv., he develops a hypothesis which he terms " evolution naturelle "; and which, making allowance for his peculiar views of the nature of generation, bears no small resemblance to what is understood by " evolution " at the present day : " Si la volonte divine a cree par un seul Acte l' Universalite des titres, d'ou venoient ces plantes et ces animaux dont Moyse nous decrit la Production au troisieme et au cinquieme jour du renouvellement de notre monde ?
" Abuserois-je de la liberte de conjectures si je disois, que les Plantes et les Animaux qui existentujourd'hui sont parvenus par une sorte d'evolution naturelle des Etres organises qui peuplaient ce premier Monde, sorti immediatement des Mains du Createur ? .. .
" Ne supposons que trois revolutions. La Terre vient de sortir des Mains du Createur. Des causes preparees par sa Sagesse font developper de toutes parts les Germes. Les Etres organises commencent a jouir de l'existence. Its etoient probablement alors bien differens de ce qu'ils sont aujourd'hui. Its l'etoient autant que ce premier Monde differoit de celui que nous habitons. Nous manquons de moyens pour juger de ces dissemblances, et peut-titre que le plus habile Naturaliste qui auroit ete place dans ce premier Monde y auroit entierement meconnu nos Plantes et nos Animaux." The two parts of Bonnet's hypothesis, namely, the doctrine that all living things proceed from pre-existing germs, and that these contain, one enclosed within the other, the germs of all future living things, which is the hypothesis of " emboitement," and the doctrine that every germ contains in miniature all the organs of the adult, which is the hypothesis of evolution or development, in the primary senses of these words, must be carefully distinguished. In fact, while holding firmly by the former, Bonnet more or less modified the latter in his later writings, and, at length, he admits that a " germ " need not be an actual miniature of the organism, hut that it may be merely an " original preformation " capable of producing the latter.4 But, thus defined, the germ is neither more nor less than the "particula genitalis" of Aristotle, or the "primordium vegetale" or " ovum " of Harvey; and the " evolution " of such a germ would not be distinguishable from " epigenesis." Supported by the great authority of Haller, the doctrine of evolution, or development, prevailed throughout the whole of the 18th century, and Cuvier appears to have substantially adopted Bonnet's later views, though probably he would not have gone all lengths in the direction of " emboitement." In a well-known note to Charles Leopold Laurillard's Eloge, prefixed to the last edition of the Ossemens fossiles, the " radical de l'etre " is much the same thing as Aristotle's " particula genitalis " and Harvey's " ovum." Bonnet's eminent contemporary, Buffon, held nearly the same views with respect to the nature of the germ, and expresses them even more confidently.
" Ceux qui ont cru que le coeur etoit le premier forme, se sont trompes; ceux qui disent que c'est le sang se trompent aussi : tout est forme en meme temps. Si l'on ne consulte que l'observation, le poulet se voit dans l'ceuf avant qu'il ait ete couve." J'ai ouvert une grande quantite d'ceufs a differens temps avant et apres l'incubation, et je me suis convaincu par mes yeux que le poulet existe en entier dans le milieu de la cicatrule au moment qu'il sort du corps de la poule." The " moule interieur " of Buffon is the aggregate of elementary parts which constitute the individual, and is thus the equivalent of Bonnet's germ, as defined in the passage cited above. But Buffon further imagined that innumerable "molecules organiques " are dispersed throughout the world, and that alimentation consists in the appropriation by the parts of an. organism of those molecules which are analogous to them. Growth, therefore, was, on this hypothesis, partly a process of simple evolution, and partly of what has been termed syngenesis. Buffon's opinion is, in fact, a sort of combination of views, essentially similar to those of Bonnet, with others, somewhat similar to those of the " Medici " whom Harvey condemns. The " molecules organiques " are physical equivalents of Leib nitz's " monads." It is a striking example of the difficulty of getting people to use their own powers of investigation accurately, that this form of the doctrine of evolution should have held its ground so long; for it was thoroughly and completely exploded, not long after its enunciation, by Caspar Frederick Wolff, who in his Theoria generationis, published in 1759, placed the opposite theory of epigenesis upon the secure foundation of fact, from which it has never been displaced. But Wolff had no immediate Ce mot (germe) ne designera pas seulement un corps organise reduit en petit; it designera encore toute espece de preformation originelle dont un Tout organique peut resulter comme de son principe immediat." - Palingenesie philosophique, part. x. chap. ii.
M. Cuvier considerant que tous les titres organises sont derives de parens, et ne voyant dans la nature aucune force capable de produire l'organisation, croyait a la pre-existence des germes; non pas a la pre-existence d'un titre tout forme, puisqu'il est bien evident que ce n'est que par des developpemens successifs que l'etre acquiert sa forme; mais, si l'on peut s'exprimer ainsi, a la pre-existence du radical de l'etre, radical qui existe avant que la serie des evolutions ne commence, et qui remonte certainement, suivant la belle observation de Bonnet, a plusieurs generations." - Laurillard, Eloge de Cuvier, note 12.
6 Histoire naturelle, tom. ii. ed. ii. (1750), p.
Ibid. p. 351. See particularly Buffon, l.c. p. 41.
successors. The school of Cuvier was lamentably deficient in embryologists; and it was only in the course of the first thirty years of the igth century that Prevost and Dumas in France, and, later on, Ddllinger, Pander, von Bar, Rathke, and Remak in Germany, founded modern embryology; and, at the same time, proved the utter incompatibility of the hypothesis of evolution as formulated by Bonnet and Haller with easily demonstrable facts.
Nevertheless, though the conceptions originally denoted by " evolution " and " development " were shown to be untenable, the words retained their application to the process by which the embryos of living beings gradually make their appearance; and the terms" development," " Entwickelung,"and " evolutio " are now indiscriminately used for the series of genetic changes exhibited by living beings, by writers who would emphatically deny that " development " or " Entwickelung " or " evolutio," in the sense in which these words were usually employed by Bonnet or Haller, ever occurs.
Evolution, or development, is, in fact, at present employed in biology as a general name for the history of the steps by which any living being has acquired the morphological and the physiological characters which distinguish it. As civil history may be divided into biography, which is the historyof individuals, and universal history, which is the history of the human race, so evolution falls naturally into two categories - the evolution of the individual (see Embryology) and the evolution of the sum of living beings. The Evolution of the Sum of Living Beings
The notion that all the kinds of animals and plants may have come into existence by the growth and modification of primordial germs is as old as speculative thought; but the modern scientific form of the doctrine can be traced historically to the influence of several converging lines of philosophical speculation and of physical observation, none of which go further back than the 17th century. These are: I. The enunciation by Descartes of the conception that the physical universe, whether living or not living, is a mechanism, and that, as such, it is explicable on physical principles.
2. The observation of the gradations of structure, from extreme simplicity to very great complexity, presented by living things, and of the relation of these graduated forms to one another.
3. The observation of the existence of an analogy between the series of gradations presented by the species which compose any great group of animals or plants, and the series of embryonic conditions of the highest members of that group.
4. The observation that large groups of species of widely different habits present the same fundamental plan of structure; and that parts of the same animal or plant, the functions of which are very different, likewise exhibit modifications of a common plan.
5. The observation of the existence of structures, in a rudimentary and apparently useless condition, in one species of a group, which are fully developed and have definite functions in other species of the same group.
6. The observation of the effects of varying conditions in modifying living organisms.
7. The observation of the facts of geographical distribution.
8. The observation of the facts of the geological succession of the forms of life.
1. Notwithstanding the elaborate disguise which fear of the powers that were led Descartes to throw over his real opinions, it is impossible to read the Principes de la philosophie without acquiring the conviction that this great philosopher held that the physical world and all things in it, whether living or not living, have originated by a process of evolution, due to the continuous operation of purely physical causes, out of a primitive relatively formless matter.' ' As Billion has well said : - " L'idee de ramener l'explication de tons les phenomenes a des principes mecaniques est assurement grande et belle,ce pas est le plus hardi qu'on peut faire en philosophic, et c'est Descartes qui l'a fait." - l.c. p. 50.
The following passage is especially instructive: " Et tant s'en Taut que je veuille que ion croie toutes les choses que j'ecrirai, que meme je pretends en proposer ici quelques-unes que je crois absolument titre fausses; A savoir, je ne doute point que le monde n'ait etc cree au commencement avec autant de perfection qu'il en a; en sorte que le soleil, la terre, la lune, et les etoiles ont etc des lors; et que la terre n'a pas eu seulement en soi les semences des plantes, mais que les plantes meme en ont couvert une partie; et qu'Adam et Eve n'ont pas etc crees enfans mais en Age d'hommes parfaits. La religion chretienne veut que nous le croyons ainsi, et la raison naturelle nous persuade entierement cette verite; car si nous considerons la toute puissance de Dieu, nous devons juger que tout ce qu'il a fait a eu des le commencement toute la perfection qu'il devoit avoir. Mais neanmoins, comme on connoitroit beaucoup mieux quelle a etc la nature d'Adam et celle des arbres de Paradis si on avoit examine comment les enfants se forment peu a peu dans le ventre de leurs mores et comment les plantes sortent de leurs semences, que si on avoit seulement considers quels ils ont etc quand Dieu les a crees : tout de meme, nous ferons mieux entendre quelle est generalement la nature de toutes les choses qui sont au monde si nous pouvons imaginer quelques principes qui soient fort intelligibles et fort simples, desquels nous puissions voir clairement que les astres et la terre et enfin tout ce monde visible auroit pu titre produit ainsi que de quelques semences (bien que nous sachions qu'il n'a pas ste produit en cette fawn) que si nous la decrivions seulement comme it est, ou bien comme nous croyons qu'il a ste cree. Et parceque je pense avoir trouvs des principes qui sont tels, je tacherai ici de les expliquer." 2 If we read between the lines of this singular exhibition of force of one kind and weakness of another, it is clear that Descartes believed that he had divined the mode in which the physical universe had been evolved; and the Traits de l'homme and the essay Sur les passions afford abundant additional evidence that he sought for, and thought he had found, an explanation of the phenomena of physical life by deduction from purely physical laws.
Spinoza abounds in the same sense, and is as usual perfectly candid " Naturae leges et regulae, secundum quas omnia fiunt et ex unis formis in alias mutantur, sunt ubique et semper eadem." 3 Leibnitz's doctrine of continuity necessarily led him in the same direction; and, of the infinite multitude of monads with which he peopled the world, each is supposed to be the focus of an endless process of evolution and involution. In the Protogaea, xxvi., Leibnitz distinctly suggests the mutability of species " Alii mirantur in saxis passim species videri quas vel in orbe cognito, vel saltem in vicinis locis frustra quaeras. Ita Cornua Ammonis, quae ex nautilorum numero habeantur, passim et forma et magnitudine (nam et pedali diametro aliquando reperiuntur) ab omnibus illis naturis discrepare dicunt, quas praebet mare. Sed quis absconditos ejus recessus aut subterraneas abyssos pervestigavit ? quam multa nobis animalia antea ignota offert novus orbis ? Et credibile est per magnas illas conversiones etiam animalium species plurimum immutatas." Thus in the end of the 17th century the seed was sown which has at intervals brought forth recurrent crops of evolutional hypotheses, based, more or less completely, on general reasonings.
Among the earliest of these speculations is that put forward by Benoit de Maillet in his Telliained, which, though printed in 1735, was not published until twenty-three years later. Considering that this book was written before the time of Haller, or Bonnet, or Linnaeus, or Hutton, it surely deserves more respectful consideration than it usually receives. For De Maillet not only has a definite conception of the plasticity of living things, and of the production of existing species by the modification of their predecessors, but he clearly apprehends the cardinal maxim of modern geological science, that the explanation of the structure of the globe is to be sought in the deductive application to geological phenomena of the principles established inductively by the study of the present course of nature. Somewhat later, P. L. M. de Maupertuis 4 suggested a curious hypothesis as to the causes of variation, which he thinks may be sufficient to account for the origin of all animals 2 Principes de la philosophie, Troisieme partie, § 45.
3 Ethices, Pars tertia, Praefatio.
4 Systeme de la Nature. Essai sur la formation des corps organises, 1751, xiv.
from a single pair. Jean Baptiste Rene Robinet' followed out much the same line of thought as De Maillet, but less soberly; and Bonnet's speculations in the Palingenesie, which appeared in 1769, have already been mentioned. Buffon (1753-1778), at first a partisan of the absolute immutability of species, subsequently appears to have believed that larger or smaller groups of species have been produced by the modification of a primitive stock; but he contributed nothing to the general doctrine of evolution.
Erasmus Darwin (Zoonomia, 17 94 ), though a zealous evolutionist, can hardly be said to have made any real advance on his predecessors; and, notwithstanding the fact that Goethe had the advantage of a wide knowledge of morphological facts, and a true insight into their signification, while he threw all the power of a great poet into the expression of his conceptions, it may be questioned whether he supplied the doctrine of evolution with a firmer scientific basis than it already possessed. Moreover, whatever the value of Goethe's labours in that field, they were not published before 1820, long after evolutionism had taken a new departure from the works of Treviranus and Lamarck - the first of its advocates who were equipped for their task with the needful large and accurate knowledge of the phenomena of life as a whole. It is remarkable that each of these writers seems to have been led, independently and contemporaneously, to invent the same name of " biology " for the science of the phenomena of life; and thus, following Buffon, to have recognized the essential unity of these phenomena, and their contradistinction from those of inanimate nature. And it is hard to say whether Lamarck or Treviranus has the priority in propounding the main thesis of the doctrine of evolution; for though the first volume of Treviranus's Biologie appeared only in 1802, he says, in the preface to his later work, the Erscheinungen and Gesetze des organischen Lebens, dated 1831, that he wrote the first volume of the Biologie " nearly five-and-thirty years ago," or about 1796.
Now, in 1794, there is evidence that Lamarck held doctrines which present a striking contrast to those which are to be found in the Philosophic zoologique, as the following passages show: " 685. Quoique mon unique objet dans cet article n'ait etc que de traiter de la cause physique de l'entretien de la vie des titres organiques, malgre cela j'ai ose avancer en debutant, que l'existence de ces titres etonnants n'appartiennent nullement a la nature; que tout ce qu'on peut entendre par le mot nature, ne pouvoit donner la vie, c'est-a-dire, que toutes les qualites de la matiere, jointes a toutes les circonstances possibles, et meme a l'activite repandue dans l'univers, ne pouvaient point produire un titre muni du mouvement organique, capable de reproduire son semblable, et sujet a la mort.
" 686. Tous les individus de cette nature, qui existent, proviennent d'individus semblables qui tous ensemble constituent l'espece entiere. Or, je crois qu'il est aussi impossible a l'homme de connoitre la cause physique du premier individu de chaque espece, que d'assigner aussi physiquement la cause de l'existence de la matiere ou de l'univers entier. C'est au moins ce que le resultat de mes connaissances et de mes reflexions me portent a penser. S'il existe beaucoup de varietes produites par l'effet des circonstances, ces varietes ne denaturent point les especes ; mais on se trompe, sans doute souvent, en indiquant comme espece, ce qui n'est que variete; et alors je sens que cette erreur peut tirer a consequence dans les raisonnements que 1'on fait sur cette matiere." The first three volumes of Treviranus's Biologie, which contains his general views of evolution, appeared between 1802 and 1805. The Recherches sur l'organisation des corps vivants, which sketches out Lamarck's doctrines, was published in 1802; but the full development of his views in the Philosophic zoologique did not take place until 1809.
1 Considerations philosophiques sur la gradation naturelle des formes de l'etre; ou les essais de la nature qui apprend d faire l'homme (1768).
Recherches sur les causes des principaux faits physiques, par J. B. Lamarck. Paris. Seconde annee de la Republique. In the preface, Lamarck says that the work was written in 1776, and presented to the Academy in 1780; but it was not published before 2794, and at that time it presumably expressed Lamarck's mature views. It would be interesting to know what brought about the change of opinion manifested in the Recherches sur l'organisation des corps vivants, published only seven years later.
The Biologie and the Philosophic zoologique are both very remarkable productions, and are still worthy of attentive study, but they fell upon evil times. The vast authority of Cuvier was employed in support of the traditionally respectable hypotheses of special creation and of catastrophism; and the wild speculations of the Discours sur les revolutions de la surface du globe were held to be models of sound scientific thinking, while the really much more sober and philosophical hypotheses of the Hydrogeologie were scouted. For many years it was the fashion to speak of Lamarck with ridicule, while Treviranus was altogether ignored.
Nevertheless, the work had been done. The conception of evolution was henceforward irrespressible, and it incessantly reappears, in one shape or another, 2 up to the year 1858, when Charles Darwin and A. R. Wallace published their Theory of Natural Selection. The Origin of Species appeared in 1859; and thenceforward the doctrine of evolution assumed a position and acquired an importance which it never before possessed. In the Origin of Species, and in his other numerous and important contributions to the solution of the problem of biological evolution, Darwin confined himself to the discussion of the causes which have brought about the present condition of living matter, assuming such matter to have once come into existence. On the other hand, Spencer and E. Haeckel 5 dealt with the whole problem of evolution. The profound and vigorous. writings of Spencer embody the spirit of Descartes in the knowledge of our own day, and may be regarded as the Principes de la philosophic of the 19th century; while, whatever hesitation may not unfrequently be felt by less daring minds in following Haeckel in many of his speculations, his attempt to systematize the doctrine of evolution and to exhibit its influence as the central thought of modern biology, cannot fail to have a far-reaching influence on the progress of science.
If we seek for the reason of the difference between the scientific position of the doctrine of evolution in the days of Lamarck and that which it occupies now, we shall find it in the great accumulation of facts, the several classes of which have been enumerated above, under the second to the eighth heads. For those which are grouped under the second to the seventh of these classes, respectively, have a clear significance on the hypothesis of evolution, while they are unintelligible if that hypothesis be denied. And those of the eighth group are not only uninintelligible without the assumption of evolution, but can be proved never to be discordant with that hypothesis, while, in some cases, they are exactly such as the hypothesis requires. The demonstration of these assertions would require a volume, but the general nature of the evidence on which they rest may be briefly indicated.
2. The accurate investigation of the lowest forms of animal life, * ;commenced by Leeuwenhoek and Swammerdam, and continued by the remarkable labours of Reaumur, Abraham Trembley, Bonnet, and a host of other observers in the latter part of the 17th and the first half of the 18th centuries, drew the attention of biologists to the gradation in the complexity of organization which is presented by living beings, and culminated in the doctrine of the echelle des titres, so powerfully and clearly stated by Bonnet, and, before him, adumbrated by Locke and by Leibnitz. In the then state of knowledge, it appeared that all the species of animals and plants could be arranged in one series, in such a manner that, by insensible gradations, the mineral passed into the plant, the plant into the polype, the polype into the worm, and so, through gradually higher forms of life, to man, at the summit of the animated world.
But, as knowledge advanced, this conception ceased to be tenable in the crude form in which it was first put forward. Taking into account existing animals and plants alone, it became obvious that they fell into groups which were more or less sharply separated from one another; and, moreover, that even See the " Historical Sketch " prefixed to the last edition of the Origin of Species. ' First Principles and Principles of Biology (1860-1864). 6 Generelle Morphologie (1866).
the species of a genus can hardly ever be arranged in linear series. Their natural resemblances and differences are only to be expressed by disposing them as if they were branches springing from a common hypothetical centre.
Lamarck, while affirming the verbal proposition that animals form a single series, was forced by his vast acquaintance with the details of zoology to limit the assertion to such a series as may be formed out of the abstractions constituted by the common characters of each group.' Cuvier on anatomical, and Von Baer on embryological grounds, made the further step of proving that, even in this limited sense, animals cannot be arranged in a single series, but that there are several distinct plans of organization to be observed among them, no one of which, in its highest and most complicated modification, leads to any of the others.
The conclusions enunciated by Cuvier and Von Baer have been confirmed in principle by all subsequent research into the structure of animals and plants. But the effect of the adoption of these conclusions has been rather to substitute a new metaphor for that of Bonnet than to abolish the conception expressed by it. Instead of regarding living things as capable of arrangement in one series like the steps of a ladder, the results of modern investigation compel us to dispose them as if they were the twigs and branches of a tree. The ends of the twigs represent individuals, the ` smallest groups of twigs species, larger groups genera, and so on, until we arrive at the source of all these ramifications of the main branch, which is represented by a common plan of structure. At the present moment it is impossible to draw up any definition, based on broad anatomical or developmental characters, by which any one of Cuvier's great groups shall be separated from all the rest. On the contrary, the lower members of each tend to converge towards the lower members of all the others. The same may be said of the vegetable world. The apparently clear distinction between flowering and flowerless plants has been broken down by the series of gradations between the two exhibited by the Lycopodiaceae, Rhizocarpeae, and Gymnospermeae. The groups of Fungi, Licheneae and Algae have completely run into one another, and, when the lowest forms of each are alone considered, even the animal and vegetable kingdoms cease to have a definite frontier.
If it is permissible to speak of the relations of living forms to one another metaphorically, the similitude chosen must undoubtedly be that of a common root, whence two main trunks, one representing the vegetable and one the animal world, spring; and, each dividing into a few main branches, these subdivide into multitudes of branchlets and these into smaller groups of twigs.
As Lamarck has well said:-2 " 11 n'y a que ceux qui se sont longtemps et fortement occupes de la determination des especes, et qui ont consulte de riches collections, qui peuvent savoir jusqu'a quel point les especes, parmi les corps vivants, se fondent les unes dans les autres, et qui ont pu se convaincre que, dans les parties oft nous voyons des especes isolees, cela n'est ainsi que parcequ'il nous en manque d'autres qui en sont plus voisines et que nous n'avons pas encore recueillies.
" Je ne veux pas dire pour cela que les animaux qui existent forment une serie tres-simple et partout egalement nuancee; mais je dis qu'ils forment une serie rameuse, irregulierement graduee et qui n'a point de discontinuite dans ses parties, ou qui, du moms, n'en a toujours pas eu, s'il est vrai que, par suite de quelques especes perdues, it s'en trouve quelque part. 11 en resulte que les especes qui terminent chaque rameau de la serie generale tiennent, au moms d'un cote, a d'autres especes voisines qui se nuancent avec elles. Voila ce que l'etat bien connu des choses me met maintenant a portee de demontrer. Je n'ai besoin d'aucune hypothese ni d'aucune supposition pour cela : j 'en atteste tous les naturalistes observateurs." 3. In a remarkable essay 3 Meckel remarks: ' ` There is no good physiologist who has not been struck by the observation that the original form of all organisms is one and the 1 " 11 s'agit donc de prouver que la serie qui constitute l'echelle animale reside essentiellement dans la distribution des masses principales qui la composent et non dans celle des especes ni meme touj ours dans celle des genres." - Phil. zoologique, chap. v.
Philosophie zoologique, premiere partie, chap. iii.
" Entwurf einer Darstellung der zwischen dem Embryozustande der hoheren Thiere and dem permanenten der niederen stattfindenden Parallele," Beytrage zur vergleichenden Anatomie, Bd. ii. 1811.
same, and that out of this one form, all, the lowest as well as the highest, are developed in such a manner that the latter pass through the permanent forms of the former as transitory stages. Aristotle, Haller, Harvey, Kielmeyer, Autenrieth, and many others have either made this observation incidentally, or, especially the latter, have drawn particular attention to it, and drawn therefrom results of permanent importance for physiology." Meckel proceeds to exemplify the thesis, that the lower forms of animals represent stages in the course of the development of the higher, with a large series of illustrations.
After comparing the salamanders and the perenni-branchiate Urodela with the tadpoles and the frogs, and enunciating the law that the more highly any animal is organized the more quickly does it pass through the lower stages, Meckel goes on to say : " From these lowest Vertebrata to the highest, and to the highest forms among these, the comparison between the embryonic conditions of the higher animals and the adult states of the lower can be more completely and thoroughly instituted than if the survey is extended to the Invertebrata, inasmuch as the latter are in many respects constructed upon an altogether too dissimilar type; indeed they often differ from one another far more than the lowest vertebrate does from the highest mammal; yet the following pages will show that the comparison may be also extended to them with interest. In fact, there is a period when, as Aristotle long ago said, the embryo of the highest animal has the form of a mere worm, and, devoid of internal and external organization, is merely an almost structureless lump of polype-substance. Notwithstanding the origin of organs, it still for a certain time, by reason of its want of an internal bony skeleton, remains worm and mollusk, and only later enters into the series of the Vertebrata, although traces of the vertebral column even in the earliest periods testify its claim to a place in that series." - op. cit. pp. 4, 5.
If Meckel's proposition is so far qualified, that the comparison of adult with embryonic forms is restricted within the limits of one type of organization; and if it is further recollected, that the resemblance between the permanent lower form and the embryonic stage of a higher form is not special but general, it is in entire accordance with modern embryology; although there is no branch of biology which has grown so largely, and improved its methods so much since Meckel's time, as this. In its original form, the doctrine of " arrest of development," as advocated by Geoffroy Saint-Hilaire and Serres, was no doubt an over-statement of the case. It is not true, for example, that a fish is a reptile arrested in its development, or that a reptile was ever a fish; but it is true that the reptile embryo, at one stage of its development, is an organism which, if it had an independent existence, must be classified among fishes; and all the organs of the reptile pass, in the course of their development, through conditions which are closely analogous to those which are permanent in some fishes.
4. That branch of biology which is termed morphology is a commentary upon, and expansion of, the proposition that widely different animals or plants, and widely different parts of animals or plants, are constructed upon the same plan. From the rough comparison of the skeleton of a bird with that of a man by Pierre Delon, in the 16th century (to go no further back), down to the theory of the limbs and the theory of the skull at the present day; or, from the first demonstration of the homologies of the parts of a flower by C. F. Wolff, to the present elaborate analysis of the floral organs, morphology exhibits a continual advance towards the demonstration of a fundamental unity among the seeming diversities of living structures. And this demonstration has been completed by the final establishment of the cell theory (see Cytology), which involves the admission of a primitive conformity, not only of all the elementary structures in animals and plants respectively, but of those in the one of these great divisions of living things with those in the other. No a priori difficulty can be said to stand in the way of evolution, when it can be shown that all animals and all plants proceed by modes of development, which are similar in principle, from a fundamental protoplasmic material.
5. The innumerable cases of structures, which are rudimentary and apparently useless, in species, the close allies of which possess well-developed and functionally important homologous structures, are readily intelligible on the theory of evolution, while it is hard to conceive their raison . d'etre on any other hypothesis. However, a cautious reasoner will probably rather explain such cases deductively from the doctrine of evolution than endeavour to support the doctrine of evolution by them. For it is almost impossible to prove that any structure, however rudimentary, is useless - that is to say, that it plays no part whatever in the economy; and, if it is in the slightest degree useful, there is no reason why, on the hypothesis of direct creation, it should not have been created. Nevertheless, doubleedged as is the argument from rudimentary organs, there is probably none which has produced a greater effect in promoting the general acceptance of the theory of evolution.
6. The older advocates of evolution sought for the causes of the process exclusively in the influence of varying conditions, such as climate and station, or hybridization, upon living forms. Even Treviranus has got no further than this point. Lamarck introduced the conception of the action of an animal on itself as a factor in producing modification. Starting from the wellknown fact that the habitual use of a limb tends to develop the muscles of the limb, and to produce a greater and greater facility in using it, he made the general assumption that the effort of an animal to exert an organ in a given direction tends to develop the organ in that direction. But a little consideration showed that, though Lamarck had seized what, as far as it goes, is a true cause of modification, it is a cause the actual effects of which are wholly inadequate to account for any considerable modification in animals, and which can have no influence at all in the vegetable world; and probably nothing contributed so much to discredit evolution, in the early part of the 29th century, as the floods of easy ridicule which were poured upon this part of Lamarck's speculation. The theory of natural selection, or survival of the fittest, was suggested by William Charles Wells in 1813, and further elaborated by Patrick Matthew in 1831. But the pregnant suggestions of these writers remained practically unnoticed and forgotten, until the theory was independently devised and promulgated by Charles Robert Darwin and Alfred Russell Wallace in 1858, and the effect of its publication was immediate and profound.
Those who were unwilling to accept evolution, without better grounds than such as are offered by Lamarck, and who therefore preferred to suspend their judgment on the question, found in the principle of selective breeding, pursued in all its applications with marvellous knowledge and skill by Darwin, a valid explanation of the occurrence of varieties and races; and they saw clearly that, if the explanation would apply to species, it would not only solve the problem of their evolution, but that it would account for the facts of teleology, as well as for those of morphology; and for the persistence of some forms of life unchanged through long epochs of time, while others undergo comparatively rapid metamorphosis.
How far " natural selection " suffices for the production of species remains to be seen. Few can doubt that, if not the whole cause, it is a very important factor in that operation; and that it must play a great part in the sorting out of varieties into those which are transitory and those which are permanent.
But the causes and conditions of variation have yet to be thoroughly explored; and the importance of natural selection will not be impaired, even if further inquiries should prove that variability is definite, and is determined in certain directions rather than in others, by conditions inherent in that which varies. It is quite conceivable that every species tends to produce varieties of a limited number and kind, and that the effect of natural selection is to favour the development of some of these, while it opposes the development of others along their predetermined lines of modification.
7. No truths brought to light by biological investigation were better calculated to inspire distrust of the dogmas intruded upon science in the name of theology than those which relate to the distribution of animals and plants on the surface of the earth. Very skilful accommodation was needful, if the limitation of sloths to South America, and of the Ornithorhynchus to Australia, was to be reconciled with the literal interpretation of the history of the Deluge; and, with the establishment of the existence of distinct provinces of distribution, any serious belief in the peopling of the world by migration from Mount Ararat came to an end.
Under these circumstances, only one alternative was left for those who denied the occurrence of evolution; namely, the supposition that the characteristic animals and plants of each great province were created, as such, within the limits in which we find them. And as the hypothesis of " specific centres," thus formulated, was heterodox from the theological point of view, and unintelligible under its scientific aspect, it may be passed over without further notice, as a phase of transition from the creational to the evolutional hypothesis.
8. In fact, the strongest and most conclusive arguments in favour of evolution are those which are based upon the facts of geographical, taken in conjunction with those of geological, distribution.
Both Darwin and Wallace lay great stress on the close relation which obtains between the existing fauna of any region and that of the immediately antecedent geological epoch in the same region; and rightly, for it is in truth inconceivable that there should be no genetic connexion between the two. It is possible to put into words the proposition, that all the animals and plants of each geological epoch were annihilated, and that a new set of very similar forms was created for the next epoch, but it may be doubted if any one who ever tried to form a distinct mental image of this process of spontaneous generation on the grandest scale ever really succeeded in realizing it.
In later years the attention of the best palaeontologists has been withdrawn from the hodman's work of making " new species " of fossils, to the scientific task of completing our knowledge of individual species, and tracing out the succession of the forms presented by any given type in time.
Evolution at the Beginning of the 20th century
Since Huxley and Sully wrote their masterly essays in the 9th edition of this encyclopaedia, the doctrine of evolution has outgrown the trammels of controversy and has been accepted as a fundamental principle. Writers on biological subjects no longer have to waste space in weighing evolution against this or that philosophical theory or religious tradition; philosophical writers have frankly accepted it, and the supporters of religious tradition have made broad their phylacteries to write on them the new words. A closer scrutiny of the writers of all ages who preceded Charles Darwin, and, in particular, the light thrown back from Darwin on the earlier writings of Herbert Spencer, have made plain that without Darwin the world by this time might have come to a. general acceptance of evolution; but it seems established as a historical fact that the world has come to accept evolution, first, because of Darwin's theory of natural selection, and second, because of Darwin's exposition of the evidence for the actual occurrence of organic evolution. The evidence as set out by Darwin has been added to enormously; new knowledge has in many cases altered our conceptions of the mode of the actual process of evolution, and from time to time a varying stress has been laid on what are known as the purely Darwinian factors in the theory. The balance of these tendencies has been against the attachment of great importance to sexual selection, and in favour of attaching a great importance to natural selection; but the dominant feature in the recent history of the theory has been its universal acceptance and the recognition that this general acceptance has come from the stimulus given by Darwin.
A change has taken place in the use of the word evolution. Huxley, following historical custom, devoted one section of his article to the " Evolution of the Individual." The facts and theories respecting this are now discussed under such headings as Embryology; Heredity; Variation And Selection; under these headings must be sought information on the important recent modifications with regard to the theory of the relation between the development of the individual and the development of the race, the part played by the environment on the individual, and the modern developments of the old quarrel between evolution and epigenesis. The most striking general change has been against seeing in the facts of ontogeny any direct evidence as to phylogeny. The general proposition as to a parallelism between individual and ancestral development is no doubt indisputable, but extended knowledge of the very different ontogenetic histories of closely allied forms has led us to a much fuller conception of the mode in which stages in embryonic and larval history have been modified in relation to their surroundings, and to a consequent reluctance to attach detailed importance to the embryological argument for evolution. The vast bulk of botanical and zoological work on living and extinct forms published during the last quarter of the 19th . century increased almost beyond all expectation the PhyJogeny ev i dence for the fact of evolution. The discovery of a single fossil creature in a geological stratum of a wrong period, the detection of a single anatomical or physiological fact irreconcilable with origin by descent with modification, would have been destructive of the theory and would have made the reputation of the observer. But in the prodigious number of supporting discoveries that have been made no single negative factor has appeared, and the evolution from their predecessors of the forms of life existing now or at any other period must be taken as proved. It is necessary to notice, however, that although the general course of the stream of life is certain, there is not the same certainty as to the actual individual pedigrees of the existing forms. In the attempts to place existing creatures in approximately phylogenetic order, a striking change, due to a more logical consideration of the process of evolution, has become established and is already resolving many of the earlier difficulties and banishing from the more recent tables the numerous hypothetical intermediate forms so familiar in the older phylogenetic trees. The older method was to attempt the comparison between the highest member of a lower group and the lowest member of a higher group - to suppose, for example, that the gorilla and the chimpanzee, the highest members of the apes, were the existing representatives of the ancestors of man and to compare these forms with the lowest members of the human race. Such a comparison is necessarily illogical, as the existing apes are separated from the common ancestor by at least as large a number of generations as separate it from any of the forms of existing man. In the natural process of growth, the gap must necessarily be wider between the summits of the twigs than lower down, and, instead of imagining " missing links," it is necessary to trace each separate branch as low down as possible, and to institute the comparisons between the lowest points that can be reached. The method is simply the logical result of the fact that every existing form of life stands at the summit of a long branch of the whole tree of life. A due consideration of it leads to the curious paradox that if any two animals be compared, the zoologically lower will be separated from the common ancestor by a larger number of generations, since, on the average, sexual maturity is reached more quickly by the lower form. Naturally very many other factors have to be considered, but this alone is a sufficient reason to restrain attempts to place existing forms in linear phylogenetic series. In embryology the method finds its expression in the limitation of comparisons to the corresponding stages of low and high forms and the exclusion of the comparisons between the adult stages of low forms and the embryonic stages of higher forms. Another expression of the same method, due to Cope, and specially valuable to the taxonomist, is that when the relationship between orders is being considered, characters of subordinal rank must be neglected. It must not be supposed that earlier writers all neglected this method, or still less that all writers now employ it, but merely that formerly it was frequently overlooked by the best writers, and now is neglected only by the worst. The result is, on the one hand, a clearing away of much fantastic phylogeny, on the other, an enormous reduction of the supposed gaps between groups.
There has been a renewed activity in the study of existing forms from the point of view of obtaining evidence as to the nature and origin of species. Comparative anatomists have been learning to refrain from basing the diagnosis of a species, or the description of the condition of an organ, on the evidence of a single specimen. Naturalists who deal specially with museum collections have been compelled, it is true, for other reasons to attach an increasing importance to what is called the type specimen, but they find that this insistence on the individual, although invaluable from the point of view of recording species, is unsatisfactory from the point of view of scientific zoology; and propositions for the amelioration of this condition of affairs range from a refusal of Linnaean nomenclature in such cases, to the institution of a division between master species for such species as have been properly revised by the comparative morphologist, and provisional species for such species as have been provisionally registered by those working at collections. Those who work with living forms of which it is possible to obtain a large number of specimens, and those who make revisions of the provisional species of palaeontologists, are slowly coming to some such conception as that a species is the abstract central point around which a group of variations oscillate, and that the peripheral oscillations of one species may even overlap those of an allied species. It is plain that we have moved far from the connotation and denotation of the word species at the time when Darwin began to discuss the origin of species, and that the movement, on the one hand, tends to simplify the problem philosophically, and, on the other, to make it difficult for the amateur theorist.
The conception of evolution is being applied more rigidly to the comparative anatomy of organs and systems of organs. When a series of the modifications of an anatomical structure has been sufficiently examined, it is frequently possible to decide that one particular condition is primitive, ancestral or central, and that the other conditions have been derived from it. Such a condition has been termed, with regard to the group of animals or plants the organs of which are being studied, archecentric. The possession of the character in the archecentric condition in (say) two of the members of the group does not indicate that these two members are more nearly related to one another than they are to other members of the group; the archecentric condition is part of the common heritage of all the members of the group, and may be retained by any. On the other hand, when the ancestral condition is modified, it may be regarded as having moved outwards along some radius from the archecentric condition. Such modified conditions have been termed apocentric. It is obvious that the mere apocentricity of a character can be no guide to the affinities of its possessor. It is necessary to determine if the modification be a simple change that might have occurred in independent cases, in fact if it be a multiradial apocentricity, or if it involved intricate and precisely combined anatomical changes that we could not expect to occur twice independently; that is to say, if it be a uniradial apocentricity. Multiradial apocentricities lie at the root of many of the phenomena that have been grouped under the designation convergence. Especially in the case of manifest adaptations, organs possessed by creatures far apart genealogically may be moulded into conditions that are extremely alike. Sir E. Ray Lankester's term, homoplasy, has passed into currency as designating such cases where different genetic material has been pressed by similar conditions into similar moulds. These may be called heterogeneous homoplasies, but it is necessary to recognize the existence of homogeneous homoplasies, here called multiradial apocentricities. A complex apocentric modification of a kind which we cannot imagine to have been repeated independently, and which is to be designated as uniradial, frequently forms a new centre around which new diverging modifications are produced. With reference to any particular group of forms such a new centre of modification may be termed a metacentre, and it is plain that the archecentre of the whole group is a metacentre of the larger group cf which the group under consideration is a branch. Thus, for instance, the archecentric condition of any Avian structure is a metacentre of the Sauropsidan stem. A form of apocentricity extremely common and often perplexing may be termed pseudocentric; in such a condition there is an apparent simplicity that tive anatomy. reveals its secondary nature by some small and apparently meaningless complexity.
Another group of investigations that seems to play an important part in the future development of the theory of evolution relates to the study of what is known as organic symmetry. The differentiations of structure that char acterize animals and plants are being shown to be orderly and definite in many respects; the relations of the various parts to one another and to the whole, the modes of repetition of parts, and the series of changes that occur in groups of repeated parts, appear to be to a certain extent inevitable, to depend on the nature of the living material itself and on the necessary conditions of its growth. Closely allied to the study of symmetry is the study of the direct effect of the circumambient media on embryonic young and adult stages of living beings (see Embryology: Physiology; Heredity; and Variation And Selection), and a still larger number of observers have added to our knowledge of these. It is impossible here to give even a list of the names of the many observers who in recent times have made empirical study of the effects of growth-forces and of the symmetrical limitations and definitions of growth. It is to be noticed, however, that, even after such phenomena have been properly grouped and designated under Greek names as laws of organic growth, they have not become explanations of the series of facts they correlate. Their importance in the theory of evolution is none the less very great. In the first place, they lessen the number of separate facts to be explained; in the second, they limit the field within which explanation must be sought, since, for instance, if a particular mode of repetition of parts occur in mosses, in flowering-plants, in beetles and in elephants, the seeker of ultimate explanations may exclude from the field of his inquiry all the conditions individual to these different organic forms, and confine himself only to what is common to all of them; that is to say, practically only the living material and its environment. The prosecution of such inquiries is beginning to make unnecessary much ingenious speculation of a kind that was prominent from r880 to 'goo; much futile effort has been wasted in the endeavour to find on Darwinian principles special " selection-values " for phenomena the universality of which places them outside the possibility of having relations with the particular conditions of particular organisms. On the other hand, many of those who have been specially successful in grouping diverse phenomena under empirical generalizations have erred logically in posing their generalizations against such a vera causa as the preservation of favoured individuals and races. The thirty years which followed the publication of the Origin of Species were characterized chiefly by anatomical and embryological work ; since then there has been no diminution in anatomical and embryological enthusiasm, but many of the continually increasing body of investigators have turned again to bionomical work. Inasmuch as Lamarck attempted to frame a theory of evolution in which the principle of natural selection had no part, the interpretation placed on their work by many bionomical investigators recalls the theories of Lamarck, and the name Neo-Lamarckism has been used of such a school of biologists, particularly active in America. The weakness of the NeoLamarckian view lies in its interpretation of heredity; its strength lies in its zealous study of the living world and the detection therein of proximate empirical laws, a strength shared by very many bionomical investigations, the authors of which would prefer to call themselves Darwinians, or to leave themselves without sectarian designation.
Statistical inquiry into the facts of life has long been employed, and in particular Francis Galton, within the Darwinian period, has advocated its employment and developed its methods. BM- Within quite recent years, however, a special school q Y P has arisen with the main object of treating the processes of evolution quantitatively. Here it is right to speak of Karl Pearson as a pioneer of notable importance. It has been the habit of biologists to use the terms variation, selection, elimination, correlation and so forth, vaguely; the new school, which has been strongly reinforced from the side of physical science, insists on quantitative measurements of the terms. When the anatomist says that one race is characterized by long heads, another by round heads, the biometricist demands numbers and percentages. When an organ is stated to be variable, the biometricist demands statistics to show the range of the variations and the mode of their distribution. When a character is said to be favoured by natural selection, the biometricist demands investigation of the death-rate of individuals with or without the character. When a character is said to be transmitted, or to be correlated with another character, the biometricist declares the statement valueless without numerical estimations of the inheritance or correlation. The subject is still so new, and its technical methods (see Variation And Selection) have as yet spread so little beyond the group which is formulating and defining them, that it is difficult to do more than guess at the importance of the results likely to be gained. Enough, however, has already been done to show the vast importance of the method in grouping and codifying the empirical facts of life, and in so preparing the way for the investigation of ultimate " causes." The chief pitfall appears to be the tendency to attach more meaning to the results than from their nature they can bear. The ultimate value of numerical inquiries must depend on the equivalence of the units on which they are based. Many of the characters that up to the present have been dealt with by biometrical inquiry are obviously composite. The height or length of the arm of a human being, for instance, is the result of many factors, some inherent, some due to environment, and until these have been sifted out, numerical laws of inheritance or of correlation can have no more than an empirical value. The analysis of composite characters into their indivisible units and statistical inquiry into the behaviour of the units would seem to be a necessary part of biometric investigation, and one to which much further attention will have to be paid.
It is well known that Darwin was deeply impressed by differences in flora and fauna, which seemed to be functions of locality, and not the result of obvious dissimilarities of environment. A. R. Wallace's studies of island life, and the work of many different observers on local races of animals and plants, marine, fluviatile and terrestrial, have brought about a conception of segregation as apart from differences of environment as being one of the factors in the differentiation of living forms. The segregation may be geographical, or may be the result of preferential mating, or of seasonal mating, and its effects plainly can be made no more of than proximate or empirical laws of differentiation, of great importance in codifying and simplifying the facts to be explained. The minute attention paid by modern systematists to the exact localities of subspecies and races is bringing together a vast store of facts which will throw further light on the problem of segregation, but the difficulty of utilizing these facts is increased by an unfortunate tendency to make locality itself one of the diagnostic characters.
Consideration of phylogenetic series, especially from the palaeontological side, has led many writers to the conception that there is something of the nature of a growth-force inherent in organisms and tending inevitably towards divergent evolution. It is suggested that even in the absence of modification produced by any possible Darwinian or Lamarckian factors, that even in a neutral environment, divergent evolution of some kind would have occurred. The conception is necessarily somewhat hazy, but the words bathmism and bathmic Evolution have been employed by a number of writers for some such conception. Closely connected with it, and probably underlying many of the facts which have led to it, is a more definite group of ideas that may be brought together under the phrase " phylogenetic limitation of variation." In its simplest form, this phrase implies such an obvious fact as that whatever be the future development of, say, existing cockroaches, it will be on lines determined by the present structure of these creatures. In a more general way, the phrase implies that at each successive branching of the tree of life, the branches become more specialized, more defined, and, in a sense, more limited. The full implications of the group of ideas require, and are likely to receive, much attention in the immediate future of biological investigation, but it is enough at present to point out that until the more obvious lines of inquiry have been opened out much more fully, we cannot be in a position to guess at the existence of a residuum, for which such a metaphysical conception as bathmism would serve even as a convenient disguise for ignorance.
Almost every side of zoology has contributed to the theory of evolution, but of special importance are the facts and theories associated with the names of Gregor Mendel, A. Weismann and Hugo de Vries. These are discussed under the headings Heredity; Mendelism; and Variation And Selection. It has been a feature of great promise in recent contributions to the theory of evolution, that such contributions have received attention almost directly in proportion to the new methods of observation and the new series of facts with which they have come. Those have found little favour who brought to the debate only formal criticisms or amplifications of the Darwinian arguments, or re-marshallings of the Darwinian facts, however ably conducted. The time has not yet come for the attempt to synthesize the results of the many different and often apparently antagonistic groups of workers. The great work that is going on is the simplification of the facts to be explained by grouping them under empirical laws; and the most general statement relating to these that can yet be made is that no single one of these laws has as yet shown signs of taking rank as a vera causa comparable with the Darwinian principle of natural selection.
For evolution in relation to society see Sociology.
References. - Practically, every botanical and zoological publication of recent date has its bearing on evolution. The following are a few of the more general works: Bateson, Materials for the Study of Variation; Bunge, Vitalismus and Mechanismus; Cope, Origin of the Fittest, Primary Factors of Organic Evolution, Darwin's Life and Letters; H. de Vries, Species and Varieties and their Origin by Mutation; Eimer, Organic Evolution; Gulick, " Divergent Evolution through Cumulative Segregation," Jour. Linn. Soc. xx.; Haacke, Schopfung des Menschen; Mitchell, " Valuation of Zoological Characters," Trans. Linn. Soc. viii. pt. 7; Pearson, Grammar of Science; Romanes, Darwin and after Darwin; Sedgwick, Presidential Address to Section Zoology, Brit. Ass. Rep. 1899; Wallace, Darwinism; Weismann, The Germ-Plasm. Further references of great value will be found in the works of Bateson and Pearson referred to above, and in the annual volumes of the Zoological Record, particularly under the head " General Subject." (P. C. M.)