[328] In the "preliminary notice" of his Crayfish paper--_Isis_, pp 1093-1100, 1825.
[329] "On the Anatomy and the Affinities of the Family of the Medusae," _Phil. Trans._, 1849; _Sci. Memoirs_, i., pp. 9-32.
[330] _Phil. Trans._, cxliii., p. 368, 1853.
[331] The principle of achromatism was discovered (by Fraunhofer) and achromatic microscopes introduced in the early part of the 19th century. The use of chemical reagents, such as acetic acid, and various hardening fluids, came into fashion not long after. J. Muller seems to have been one of the first to realise their importance. Remak himself invented one or two fixing and hardening mixtures (pp. 87, 127, 1855), which enabled him to cut excellent hand sections. Section-cutting machines were not invented till later (V. Hensen, 1866, His, 1870).
[332] _Untersuchungen uber die Entwickelung der Wirbelthiere_, folio, pp. x.x.xvii + 195, 12 plates, Berlin, 1850-1855.
CHAPTER XIII
THE RELATION OF LAMARCK AND DARWIN TO MORPHOLOGY.
It is a remarkable fact that morphology took but a very little part in the formation of evolution-theory. When one remembers what powerful arguments for evolution can be drawn from such facts as the unity of plan and composition and the law of parallelism, one is astonished to find that it was not the morphologists at all who founded the theory of evolution.
It is true that the noticeable resemblances of animals to one another, the possibility of arranging them in a system, the vague perception of an all-pervading plan of structure, did suggest to many minds the thought that systematic affinities might be due to blood-relationship.
Thus Leibniz considered that the cat tribe might possibly be descended from a common ancestor,[333] and another great philosopher, Immanuel Kant, was led by his perception of the unity of type to suggest as possible the derivation of the whole organic realm from one parent form, or even ultimately from inorganic matter. In the course of his masterly discussion of mechanism and teleology,[334] he writes, "The agreement of so many genera of animals in a certain common schema, which appears to be fundamental not only in the structure of their bones, but also in the disposition of their remaining parts--so that with an admirable simplicity of original outline, a great variety of species has been produced by the shortening of one member and the lengthening of another, the involution of this part and the evolution of that--allows a ray of hope, however faint, to penetrate into our minds, that here something may be accomplished by the aid of the principle of the mechanism of Nature (without which there can be no natural science in general). This a.n.a.logy of forms, which with all their differences seem to have been produced according to a common original type, strengthens our suspicions of an actual relationship between them in their production from a common parent, through the gradual approximation of one animal-genus to another--from those in which the principle of purposes seems to be best authenticated, _i.e._, from man down to the polype, and again from this down to mosses and lichens, and finally to the lowest stage of Nature noticeable by us, viz., to crude matter."[335]
So, too, Buffon"s evolutionism was suggested by his study of the structural affinities of animals, and Erasmus Darwin in his _Zoonomia_ (1794) brought forward as one of the strongest proofs of evolution, "the essential unity of plan in all warm-blooded animals."[336]
But, as a matter of historical fact, no morphologist, not even Geoffroy, deduced from the facts of his science any comprehensive theory of evolution. The pre-Darwinian morphologists were comparatively little influenced by the evolution-theories current in their day, and it was in the anatomist Cuvier and the embryologist von Baer that the early evolutionists found their most uncompromising opponents.
Speaking generally, and excepting for the moment the theory of Lamarck, we may say that the evolution-theories of the 18th and 19th centuries arose in connection with the transcendental notion of the _ech.e.l.le des etres_, or scale of perfection. This notion, which plays so great a part in the philosophy of Leibniz, was very generally accepted about the middle of the 18th century, and received complete and even exaggerated expression from Bonnet and Robinet. Buffon also was influenced by it.
Towards the beginning of the 19th century the idea was taken up eagerly by the transcendental school and by them given, in their theories of the "one animal," a more morphological turn. Their recapitulation theory was part and parcel of the same general idea.
One understands how easily the notion of evolution could arise in minds filled with the thought of the ideal progression of the whole organic kingdom towards its crown and microcosm, man. Their theory of recapitulation led them to conceive evolution as the developmental history of the one great organism.[337] Many of them wavered between the conception of evolution as an ideal process, as a _Vorstellungsart_, and the conception of it as an historical process. Bonnet, Oken, and the majority of the transcendentalists seem to have chosen the former alternative; Robinet, Trevira.n.u.s, Tiedemann, Meckel, and a few others held evolution to be a real process.
We have already in previous chapters[338] briefly noticed the relation of one or two of the transcendental evolution-theories to morphology, and there is little more to be said about them here. They had as good as no influence upon morphological theory, nor indeed upon biology in general.[339] It is different with the theory of Lamarck, which, although it had little influence upon biological thought during and for long after the lifetime of its author, is still at the present day a living and developing doctrine.
Lamarck"s affinity with the transcendentalists was in many ways a close one, but he differed essentially in being before all a systematist. Nor is the direct influence of the German transcendentalists traceable in his work--his spiritual ancestors are the men of his own race, the materialists Condillac and Cabanis, and Buffon, whose friend he was. The idea of a gradation of all animals from the lowest to the highest was always present in Lamarck"s mind, and links him up, perhaps through Buffon, with the school of Bonnet. The idea of the _ech.e.l.le des etres_ had for him much less a morphological orientation than it had even for the transcendentalists, for he was lacking almost completely in the sense for morphology. Lamarck"s scientific, as distinguished from his speculative work, was exclusively systematic, and it was systematics of a very high order. He introduced many reforms into the general cla.s.sification of animals. He was the first clearly to separate Crustacea (1799), and a little later (1800) Arachnids, from insects. He reduced to a certain orderliness the neglected tribes of the Invertebrates, and wrote what was for long the standard work on their systematics--the _Histoire naturelle des Animaux sans Vertebres_ (1816-22). His speculative work on biology is contained in three publications, the small book ent.i.tled _Considerations sur l"organisation des corps vivants_ (1802), the larger work of 1809, the _Philosophie zoologique_, and the introductory matter to his _Animaux sans Vertebres_ (vol. i., 1816).
It is no easy matter to give in short compa.s.s an account of Lamarck"s biological philosophy. He is an obscure writer, and often self-contradictory.
In the first part of the _Philosophie zoologique_ Lamarck is largely pre-occupied with the problem of whether species are really distinct, or do not rather grade insensibly into one another. As a systematist of vast experience Lamarck knew how difficult it is in practice to distinguish species from varieties. "The more," he writes, "we collect the productions of Nature, the richer our collections become, the more do we see almost all the gaps filled up and the lines of separation effaced. We find ourselves reduced to an arbitrary determination, which sometimes leads us to seize upon the slightest differences of varieties, and form from them the distinctive character of what we call a species, and at other times leads us to consider as a variety of a certain species individuals a little bit different, which others regard as forming a separate species."[340]
For Lamarck, as for Darwin later, the chief problem was not the evolution and differentiation of types of structure, but the mode of origin of species.
Lamarck is at great pains to show how arbitrary are our determinations of species, and how artificial the cla.s.sificatory groups which we distinguish in Nature. Strictly speaking, there are in Nature only individuals, "... this is certain, that among her products Nature has in reality formed neither cla.s.ses, nor orders, nor families, nor genera, nor constant species, but only individuals which succeed one another and resemble those that produced them. Now, these individuals belong to infinitely diversified races, which shade into one another under all the forms and in all the degrees of organisation, and each of which maintains itself without change, so long as no cause of change acts upon it" (p. 41).
But there is a natural order in the animal kingdom, a progression from the simpler to the more complex organisations, a natural _ech.e.l.le des etres_.
This order is shown by the relation to one another of the large cla.s.sificatory groups, for they can be arranged in series from the simplest to the most complex, somewhat as follows:--
1. Infusoria.
2. Polyps.
3. Radiates.
4. Worms.
5. Insects.
6. Arachnids.
7. Crustacea.
8. Annelids.
9. Cirripedes.
10. Molluscs.
11. Fishes.
12. Reptiles.
13. Birds.
14. Mammals.
But the order of Nature is essentially continuous, and the limits of even the best defined of these cla.s.ses are in reality artificial--"if the order of Nature were perfectly known in a kingdom, the cla.s.ses which we should be forced to establish in it would always const.i.tute entirely artificial sections" (p. 45).
In the same way the lesser cla.s.sificatory groups represent smaller sections of the one unique order of Nature. Note that Lamarck"s _ech.e.l.le_ is in no way a morphological one, and was not intended to be such. It is a scale of increasing physiological differentiation, and the stages of it are marked by the acquirement of this or that new organ (_cf._ Oken). "Observation of their state convinces one that in order to produce them successively Nature has proceeded gradually from the simpler to the more complex. Now Nature, having had in mind the realisation of a plan of organisation which would permit of the greatest perfecting (that of the Vertebrates), a plan very different from those which she has been obliged to form as a preliminary to reaching it, one understands that, among the mult.i.tude of animals, one must necessarily come across not a single system of organisation which has become progressively perfected, but diverse very distinct systems, each of which has come into existence at the moment when each primary organ first put in its appearance" (p. 171).
For Lamarck this order of Nature was not merely ideal--Nature had actually formed the cla.s.ses successively, proceeding from the simpler to the more complex; she had brought about this evolution by transforming the primitive species of animals, raising them to higher degrees of organisation, and modifying them in relation to the environment in which they found themselves.
Lamarck"s theory of evolution is worked out in great detail in his _Philosophie zoologique_, but the exposition is diffuse and disconnected; it is better in giving an account of it to follow the more concise, mature and general exposition which he gives in the Introduction to his _Histoire naturelle des Animaux sans Vertebres_.[341]
Near the beginning of the Introduction Lamarck gives us in a few short "Fundamental Principles" the main lines of his general philosophy. He is a confirmed materialist. Every fact and phenomenon is essentially physical and owes its existence or production entirely to material bodies or to relations between them. All change and all movement is in the last resort due to mechanical causes. Every fact or phenomenon observed in a living body is at once a physical fact or phenomenon and a product of organisation (p. 19). Life, thought and sensation are not properties of matter, but result from particular material combinations.
His thorough-going materialism is most clearly shown in its relation to living things in the first three of the "Zoological Principles and Axioms," which are developed further on in the book.
These are as follows:--"1. No kind or particle of matter can have in itself the power of moving, living, feeling, thinking, nor of having ideas; and if, outside of man, we observe bodies endowed with all or one of these faculties, we ought to consider these faculties as physical phenomena which Nature has been able to produce, not by employing some particular kind of matter which itself possesses one or other of these faculties, but by the order and state of things which she has const.i.tuted in each organisation and in each particular system of organs.
"2. Every animal faculty, of whatever nature it may be, is an organic phenomenon, and results from a system of organs or an organ-apparatus which gives rise to it and upon which it is necessarily dependent.
"3. The more highly a faculty is developed the more complex is the system of organs which produces it, and the higher the general organisation; the more difficult also does it become to grasp its mechanism. But the faculty is none the less a phenomenon of organisation, and for that reason purely physical" (p. 104).
According to these "axioms" function is a direct and mechanical effect of structure.
The curious thing is that in spite of his avowed materialism, Lamarck"s conception of life and evolution is profoundly psychological, and from the conflict of his materialism and his vitalism (of which he was himself hardly conscious), arise most of the obscurities and the irreductible self-contradiction of his theory.
Lamarck divided animals (psychologically!) into three great groups--apathetic or insensitive animals, animals endowed with sensation, and intelligent animals. The first group, which comprise all the lower Invertebrates, are distinguished from other animals by the fact that their actions are directly and mechanically due to the excitations of the environment; they have no principle of reaction to external influences, but pa.s.sively prolong into action the excitations they receive from without. They are _irritable_ merely. The second group are distinguished from the first by their possessing, in addition to irritability, a power which Lamarck calls the _sentiment interieur_. He has some difficulty in defining exactly what he means by it:--"I have no term to express this internal power possessed not only by intelligent animals but also by those that are endowed merely with the faculty of sensation; it is a power which, when set in action by the feeling of a need, causes the individual to act at once, _i.e._, in the very moment of the sensation it experiences; and if the individual is of those that are endowed with intelligence it nevertheless acts in such a case entirely without premeditation and before any mental operation has brought its _will_ into play" (p. 24).
It is the power we call instinct in animals (p. 25), and it implies neither consciousness nor will. It acts by transforming external into internal excitations.
To this second group of animals, possessing the _sentiment interieur_, belong the higher Invertebrates, notably insects and molluscs. Only animals possessed of a more or less centralised nervous system can manifest this _sentiment_, or principle of (unconscious) reaction to external stimuli.
The higher animals, or the four Vertebrate cla.s.ses, form the group of "intelligent animals." In virtue of their more complex organisation they possess in addition to the _sentiment interieur_ the faculties of intelligence and will.
Now, broadly put, Lamarck"s theory of evolution is that new organs are formed in direct reaction to needs (_besoins_) experienced by the _sentiment interieur_. The _sentiment interieur_ is therefore the cause not only of instinctive action but also of all morphogenetic processes.
Will and intelligence (which are confined to a relatively small number of animals) have little or nothing to do directly with evolution.
To understand the working-out of Lamarck"s evolution-theory we must revert to his conception of the _ech.e.l.le des etres_. What he wrote in the _Philosophie zoologique_ is here repeated in the work of 1816 with little modification.
There is a real progression from the simpler to the more complex organisations; Nature has gradually complicated her creatures by giving them new organs and therefore new faculties.
It is interesting to note that Lamarck expressly refers to Bonnet (p.