1902 Encyclopedia > Morphology

Morphology




The term Morphology (_______, form), introduced by Goethe to denote the study of the unity of type in organic form (for which the Linnosan term METAMORPHOSIS (q.v.) had formerly been employed), now usually covers the entire science of organic form, and will be employed in this more comprehensive sense in the present article.

§ 1. Historical Outline.—If we disregard such vague likenesses as those expressed in the popular classifications of plants by size into herbs, shrubs, and trees, or of terrestrial animals by habit into beasts and creeping things, the history of morphology commences with Aris-totle. Founder of comparative anatomy and taxonomy, he established eight great divisions (to which are ap-pended certain minor groups) — Viviparous Quadrupeds, Birds, Oviparous Quadrupeds and Apoda, Fishes, Ma-lakia, Malacostraca, Entoma, and Ostracodermata—dis-tinguishing the first four groups as Enaima ("with blood") from the remaining four as Anaima (" blood-less "). In these two divisions we recognize the Ver-tebrata and Invertebrata of Lamarck, while the eight groups are identical with the Mammals, Birds, Reptiles, Fishes, the Cephalopods, Crustaceans, other Articulates, and Testaceans of recent zoology. Far, too, from com-mitting the mistake often attributed to him of reckoning Bats as Birds, or Cetaceans as Fishes, he discerned the true affinities of both, and erected the latter into a spe-cial ykvos beside the Viviparous Quadrupeds, far more on account of their absence of limbs than of their aquatic habit. Not only is his method inductive, and, as in modern systems, his groups natural, i.e., founded on the aggregate of known characters, but he foreshadows such generalizations as those of the correlation of organs, and of the progress of development from a general to a special form, long afterwards established by Cuvier and Von Baer respectively. In the correspondence he suggests between the scales of Fishes and the feathers of Birds, or in that hinted at between the fins of Fishes and the limbs of Quadrupeds, the idea of homology too is nascent; and from the compilation of his disciple Nicolaus of Damascus, who regards leaves as imperfectly-developed fruits, he seems almost to have anticipated the idea of the metamorphosis of plants. In short, we find a knowledge of structural facts and a comparative freedom from the errors induced by physiological resemblance, of which his successors such as Theophrastus and Pliny, generally mere classifiers by habit, show little trace, and which the moderns have but slowly regained. Little indeed can be recorded until the 13th century, when the reappearance of Aristotle's works gave a new impulse to the study of organic nature. Of the works of this period that of Albertus Magnus is far the most important; but they are all no more than re-vivals of Aristotle, marking the reappearance of scientific method and the reawakening of interest in and sympathy with nature. Meanwhile leech and apothecary, alchemist and witch, were accumulating considerable knowledge of plants, which, after the invention of printing, became collected and extended in the descriptive and well-illus-trated folios of Gesner and his successors, Fuchs, Lobel, and others, as well as by the establishment of botanic gardens and scientific academies, while, as Sachs expresses it, " in the sharpest contrast to the naive empiricism of the German fathers of botany came their Italian contemporary Ca?salpinus, as the thinker of the vegetable world." Both made systematic efforts,—the Germans vaguely seeking for natural affinities in mere similarities of habit, the Italian with no inconsiderable success striving towards an intel-lectual basis of classification. Monographs on groups of plants and animals frequently appeared, those of Belon on Birds and Rondelet on Fishes being among the earliest; and in the former of these (1555) we find a comparison of the skeletons of Bird and Man in the same posture and as nearly as possible bone for bone,—an idea which, despite the contemporaneous renaissance of human anatomy ini-tiated by Vesalius, disappeared for centuries, unappreciated save by the surgeon Ambroise Pare. Palissy, like Leonardo before him, discerned the true nature of fossils; and such flashes of morphological insight continued to appear from time to time during the 17th century. Thus, Joachim Jung recognized " the distinction between root and stem, the difference between leaves and foliaceous branches, the transition from the ordinary leaves to the folia floris," and Harvey anticipated the generalizations of modern embryo-logy by his researches on development and his theory of epigenesis.

The encyclopaedic period of which Gesner is the highest representative was continued by Aldrovandi, Jonston, and others in the 17th century, but, aided powerfully by the Baconian movement, then profoundly influencing all scientific minds, it developed rapidly into one of genuinely systematic aim. At this stage of progress by far the most important part was taken by John Bay, whose classificatory labours both among plants and animals were crowned with marvellous success. He first definitely expelled the fabulous monsters and prodigies of which the encyclopaedists had faithfully handed on the tradition from mediaeval times, and, like his predecessor Morison, classifying in a truly modem spirit by anatomical characters, he succeeded, particularly among plants, in distinguishing many natural groups, for which his very terms sometimes survive, e.g., Dicotyledons and Monocotyledons, Umbelliferae and Legu-minosae. The true precursor of Linnaeus, he introduced the idea of species in natural history, afterwards to become so rigid, and reformed the practice of definition and termino-logy. Of the many works which followed up Bay's systematic and monographic labours, though often, like those of Tournefort and Bivinus, Reaumur and Klein, of great importance, none can be even named until we come to those of his great successor Linnaeus, whose extraordinary grasp of logical method and unparalleled lucidity of thought and expression enabled him to reform and reorganize the whole labours of his predecessors into a compact and definite "systema naturae." The very genius of order, he established modern taxonomy (see BIOLOGY), not only by the introduction of the binomial nomenclature and the renovation of descriptive terminology and method, but by the subordination of the species—henceforth clearly defined—under the successive higher categories of genus, order, and class, so finally reconciling the analytic and synthetic tendencies of his predecessors. Although the classification of plants by the number of their essential organs (which vastly advanced not only the cultivation of botany but the knowledge of the flora of the globe, and by which he is popularly remembered) is highly artificial, it must be remembered that this artificiality is after all only a question of degree, and that he not only distinctly recognized its provisional character but collected and ex-tended those fragments of the natural system with which Jussieu soon afterwards commenced to build. His classi-fication of animals, too, was largely natural, and, though on the whole he unfortunately lent his authority to main-tain " that disastrous philosophic and scientific aberration " inherited from the alchemists through the last encyclopaedist of Gesner's school—the notion of three kingdoms of nature —he at least at one time discerned the fundamental unity of animals and vegetables, and united them in opposition to the non-living world as Organisata. At the same time he was still far more a scholastic naturalist than a modern in-vestigator, and his works represent little more than the full completion of the ancient era, and in the hands of fanatical followers served often to retard the commencement of the modern one. So, too, his excessive systematic and descriptive-precision, united as it was with comparative inattention to other than superficial characters, established a tendency, even yet not extinct, to rest contented with mere method and nomenclature instead of aiming at complete morpho-logical knowledge.

While the artificial system was at the zenith of its fame and usefulness, Bernard de Jussieu was arranging his garden on the lines afforded by the fragmentary natural system of Linnaeus. His ideas were elaborated by his nephew and successor Antoine de Jussieu, who for the first time published diagnoses of the natural orders, so giving the system its modern character. Its subsequent elaboration and definite establishment are due mainly to the labours of Pyrame de Candolle and Robert Brown. The former concentrated his own long life and that of his son upon a new " systema naturae," the colossal Prodrornus systematic naturalis (20 vols., 1818-1873), in which 80,000 species were described and arranged. Meanwhile the pene-trative genius of Brown enabled him to unravel such struc-tural complexities as those of Conifers and Cycads, Orchids and Proteaceae, thus demonstrating the possibility of ascer-taining the systematic position of even the most highly modified floral types. Both Candolle and Brown were thus no mere systematists, but genuine morphologists of the modern school. The former, as we shall afterwards see, established the theory of floral symmetry on grounds of pure comparative anatomy, and distinguished with greater success than hitherto between fundamental unity of struc-tural type and mere superficial similarity of physiological adaptation. The latter (Humboldt's "facile princeps botanicorum "), using the same ideas with even keener in-sight, made many memorable anatomical researches, such as those on the structure of the ovule and the seed, and indeed by his demonstration of the affinities of the gym-nosperms almost anticipated the discoveries of Hofmeister, who stands pre-eminent among his modern successors on account of his elucidation of the secret of phanerogamic reproduction.

The labours of Bernard and Antoine de Jussieu initiated too a vast parallel advance in zoology, the joint memoir on the classification of mammals with which Cuvier and Geoffroy St-Hilaire almost commenced their career receiv-ing its dominant impulse from the "genera" of Antoine. Cuvier's works correspond in zoology to those of the whole period from the Jussieus to Brown, and epitomize the results of that line of advance. Although in some respects preceded by Haller and Hunter, who compared, though mainly with physiological aim, the same parts in different organisms, and much more distinctly by Vicq d'Azyr, the only real comparative anatomist of the 18th cen-' tury, he truly opens the era of detailed anatomical research united with exact comparison and clear generalization. The Regne Animal (1817) and the theory of types (verte-brate, molluscan, articulate, and radiate) are the results of this union of analysis and synthesis (although he himself, exasperated by the aberrations of the Naturphilosophie, was accustomed to proclaim the importance of detailed, empiricism alone), and mark the reconstitution of taxonomy on a new basis, henceforth to be no longer a matter of superficial description and nomenclature but a complete expression of structural resemblances and differences. More even than Linnaeus he is the founder of a great school, whose names and labours are imperishable. In Germany, Bojanus, Meckel, Von Siebold, and the illustrious Johannes Miiller, with his many living pupils, have carried on the work; in France, too, a succession of brilliant anatomists, such as De Quatrefages, Milne-Edwards, and Lacaze-Duthiers, are his intellectual heirs ; and in England he has been admirably represented by Owen.

The histological movement inaugurated by Bichat will be subsequently discussed ; the rise of embryology, how-ever, may be briefly noted, especially since it supplied the most obvious deficiency of the Cuvierian school. Here the principal figure is Von Baer, who established independently the four types of Cuvier on developmental grounds, so for the first time applying embryology as the touchstone of anatomical classifications, besides establishing his famous law of differentiation from a general towards a special form.

It is now necessary to return to Linnaeus, whose more speculative writings contain, though encumbered by fan-tastic hypotheses, the idea of floral metamorphosis (" Principium florum et foliorum idem est," <fcc). About the same time, and quite independently, C. F. Wolff, the embryo-logist, stated the same theory with greater clearness, for the first time distinctly reducing the plant to an axis bearing appendages—the vegetative leaves—which become meta-morphosed into bud-scales or floral parts through diminu-tion of vegetative force. Thirty years later the same view was again independently developed by Goethe in his now well-known pamphlet (Vermch die Metamorphose der Pflanzenzu erhldren, Gotha, 1790). In this brilliant essay the doctrine of the fundamental unity of floral and foliar parts is clearly enunciated, and supported by arguments from anatomy, development, and teratology. All the organs of a plant are thus modifications of one funda-mental organ—the leaf—and all plants are in like manner to be viewed as modifications of a common type—the JJrpfianze. The controversy as to the merits and import-ance of this essay, and of Goethe's morphological work in general, can scarcely be entered upon here. That Goethe discerned and proclaimed, and that more clearly than any of his predecessors or contemporaries, the fundamental idea of all morphology—the unity which underlies the multi-farious varieties of organic form—and that he systematically applied this idea to the interpretation of the most import-ant, most complex, and most varied animal and vegetable structures, is unquestionable. The difficulties arise when we seek to estimate the importance of his works in the chain of progress, and when we inquire whether, as some historians hold, his " urpflanze " was a mere ideal archetype, bringing forth as its fruit the innumerable metaphysical abstractions of the Naturphilosophie, and leading his countrymen, to their fall, into all the extravagances of that system ; or whether, as Haeckel maintains, it represented a concrete an-cestral form, soanticipatingthe viewof modern evolutionists. That to him Schelling was largely indebted for the founda-tion upon which he erected his philosophic edifice, as also that Goethe largely shared the same ideas, is unquestion-able ; but it must be remembered that he lived and made progress for forty years after the publication of this essay, that he was familiar with the whole scientific movement, and warmly sympathized with the evolutionary views of Lamarck and Geoffroy St-Hilaire ; it is not therefore to be wondered at that his writings should furnish evidence in favour of each and every interpretation of them. His other morphological labours must not be forgotten. Inde-pendently of Vicq d'Azyr, he discovered the human pre-maxillary bone ; independently of Oken, he proposed the vertebral theory of the skull ; and before Savigny, he dis-cerned that the jaws of insects were the limbs of the head.

In 1813 A. P. de Candolle published his Théorie Élé-mentaire de la Botanique, which he developed into the classic Organographie Végétale (1827). Although at first unac-quainted with Goethe's essay, and not clearly discerning the homology of leaves and floral parts, he established his theory of symmetry, reducing all flowers to " symmetrical" groupings of appendages on an axis and accounting for their various forms by cohesion and adhesion, by arrested or excessive development. The next great advance was the investigation by Schimper and Braun of phyllotaxis—the ascending spiral arrangement of foliar and floral organs—• thus further demonstrating their essential unity.

The term morphology was first introduced by Goethe in 1817, in a subsequent essay (Zur Naturwissenschaft iiherhaupit, besonders zur Morphologie). It did not come into use in botany until its popularization by Auguste de St-Hilaire in his admirable Morphologie Vegetóle (1841), and in zoology until later, although De Blainville, who also first employed the term type, had treated the external forms of animals under "morphologie." Though the Na-turphilosophie of Schelling and its countless modifications by his followers, its mystic theories of "polarization " and the like, its apparatus of assumption and abstraction, hy-pothesis and metaphor, cannot here be discussed, its un-doubted services must not be forgotten, since it not only stimulated innumerable reflective minds to the earnest study of natural science, but, by its incessant proclamation of the unity of nature and the free use of Platonic arche-types, gave a most powerful impulse to the study of com-parative anatomy, and nobly vindicated the claims of philo-sophic synthesis over those of merely analytic empiricism. Among its many adherents, some are of more distinctly theological type, others metaphysical, others mystical or poetic, others, again, more especially scientific; but its most typical and picturesque figure is Lorenz Oken, who epitomizes alike the best and the worst features of the school, and among whose innumerable pseudo-morpholo-gical dreams there occasionally occurred suggestions of the greatest fruitfulness,—notably, for instance, the independ-ent statement of the vertebral theory of the skull.

Although Lamarck shared in this movement, his great work (the Philosophie Zoologique, 1809), being setiolo-gical rather than morphological, scarcely claims discussion here. By far the most distinguished anatomist of the transcendental school is Geoffroy St-Hilaire, who being comparatively free from the extravagances of Oken, and uniting a depth of morphological insight scarcely inferior to that of Goethe with greater knowledge of facts and far wider influence and reputation in the scientific world (which affected to sneer at the poet as necessarily a mere amateur), had enormously greater influence on the progress of science than either. He started from the same studies of anatomi-cal detail as Cuvier, but, profoundly influenced by Buffon's view of unity of plan and by the evolutionary doctrines of Lamarck, he rapidly diverged into new lines, and again reached that idea of serial homology of which we have so frequently noted the independent origin. His greatest work, the Philosophie Anatomique (1818-1823), contains his principal doctrines. These are—(1) the theory of unity of organic composition, identical in spirit with that of Goethe ; (2) the theory of analogues, according to which the same parts, differing only in form and in degree of development, should occur in all animals; (3) the " principe des con-nexions," by which similar parts occur everywhere in similar relative positions; and (4) the " principe du balancement des organes," upon which he founded the study of tera-tology, and according to which the high development of one organ is allied to diminution of another. The advance in morphological theory is here obvious ; unfortunately, how-ever, in eager pursuit of often deceptive homologies, he wandered into the transcendentalism of the Naturphilo-sophie, and seems utterly to have failed to appreciate either the type theory of Cuvier or the discoveries of Von Baer. He earnestly defended Buffon's and Bonnet's earlier view of unity of plan in nature; and the controversy reached its climax in 1830, when he maintained the unity of structure in Cephalopods and Vertebrates against Cuvier before the Academy of Sciences. On the point of fact he was of course utterly defeated; the type theory was thenceforward fully accepted and the Naturphilosophie received its deathblow, while a "second emp-jric period" of exact anatomical and embryological research seemed for ever to replace it. Such was the popular view; only a few, like the aged Goethe, whose last literary effort was a masterly critique of the controversy, discerned that the very reverse interpretation was the deeper and essential one, that a veritable "scientific revolution" was in pro-gress, and that the supremacy of homological and synthetic over descriptive and analytic studies was thenceforward assured. The irreconcilable feud between the two leaders really involved a reconciliation for their followers ; theories of homological anatomy had thenceforward* to be strictly subjected to anatomical and embryological verification, while anatomy and embryology acquired a homological aim. This union of the solid matter and rigorous method of Cuvier with the generalizing spirit and philosophic aims of Geoffroy is well illustrated in the works of Owen; and, in short, the so-called Cuvierian school is in reality thenceforward also Geoffroyan.

The further evolution of the idea of homology is sketched below (§ 7), while the extent and rapidity of the subsequent progress of the knowledge of all the structural aspects of plants and animals alike make an historical survey impos-sible up to the appearance of the Origin of Species (1859); however, no further qualitative advance was possible, since, as Sachs has best pointed out, morphology necessarily contains, under the Linnasan dogma of the constancy of species, the same two inconsistent and irreconcilable lines of thought which we saw represented by Csesalpinus and the early German botanists respectively,—on one side the want of strictly scientific classification, and on the other the vaguely-felt existence of a natural relationship. Strict classification of forms supposed constant excludes in fact any natural relationship. The type theory, the theory of unity of organic composition, and the like, are susceptible indeed of two explanations—they may be regarded as either ex-pressing a creative plan, or taken as purely Platonic and archetypal ideas. Both are tenable on theological and metaphysical grounds respectively, but the fact must not be disguised that of this unity of type no explanation in the least degree scientific, i.e., in terms of the pheno-mena of the natural world, does or can exist. The need-ful solution was effected by Darwin. The " urpflanze " of Goethe, the types of Cuvier, and the like, at once became intelligible as schematic representations of ancestral organ-isms, which, in various and varying environments, have undergone differentiation into the vast multitude of exist-ing forms. All the enigmas of structure become resolved ; "representative" and "aberrant," "progressive" and "degraded," "synthetic" and "isolated," "persistent" and "prophetic" types no longer baffle comprehension; conformity to type represented by differentiated or rudi-mentary organs in one organism is no longer contradicted by their entire disappearance in its near allies, while systematist and morphologist become related simply as specialist and generalizer, all through this escape from the Linnsean dogma of the fixity of species. The phenomena of individual development receive interpretation in terms of ancestral history; and embryology thus becomes divided into ontogeny and phylogeny, the latter, too, coming into intimate relation with palaeontology, while classification seeks henceforth the reconstruction of the genealogical tree. All these results were clearly developed in the most important work of the new period, HaeckePs Generelle Morphologie (1866), while the valuable contemporaneous Principles of Biology of Herbert Spencer also gave special attention to the relation of morphology to physiology.1
For bibliography see Cams, Geschichte der Zoologie ; Sachs, Ges-

§ 2. Results.—Though the preceding is but a meagre outline of the rise and progress of the science, no corre-sponding sketch of its results can be here attempted. A description of the refined applications of the doctrine of floral metamorphosis, an inquiry into the morphology of the Cryptogams, or an account of such beautiful homo-logies as those presented by the Arthropods or the Echino-derms is alike impossible; least of all can we consider the splendid simplification of the supremely complex prob-lem of vertebrate structure by the elaboration of a new theory of the skull, and by such luminous discoveries as those of the segmental organs, or of the origin and homo-logies of the spinal and cranial nerves. For these organo-logical conceptions the reader must study such articles as those on AMPHIBIA, BIRDS, HYDROZOA, MOLLUSCA, &C, and such works as those of Huxley, Gegenbaur and Haeckel, Balfour and Parker, Payer, Eichler, or Asa Gray, and (provided with the needful bibliographical equipment afforded by the various " Jahresberichte " and the kindred English publications) must indeed also plunge into the current literature of the science. And there too must be sought the innumerable attempts at taxonomic synthesis which such organological progress is constantly originating (see ANIMAL KINGDOM, BIOLOGY, vol. iii. p. 690 sq., and VEGETABLE KINGDOM). Embryological generalizations, such as Haeckel's " gastrsea theory," Lankester's rival " pla-nula theory," or the ingenious " ccelome theory " of Hert-wig, have been recently thoroughly criticized in Balfour's Embryology. The present article will be confined to a brief discussion of a few main problems, commencing with the cell theory and the problem of organic individuality —these being selected partly because of their special illus-trativeness and intrinsic importance, partly because they have somewhat less recently been summarized."

§ 3. Histology—Cell Theory.—Although the application of the simple microscope to the minute structure of plants and animals had been in progress since the end of the 17th century, the rise of modern histology really dates from the Anatomie Generate (1801) of Bichat, which analyses the organism into a series of simple tissues with definite structural characters. This new impulse, together with the improvement of optical appliances, led to much further research. "Fibres" and "globules," "laminae" and "nuclei," were described, and even "cells" by Mirbel in 1806, and in 1835 Johannes Midler pointed out the existence of cells resembling those of plants in the vertebrate notochord. The cellu-lar and nucleated structure of epidermis and other tissues was soon demonstrated, wdiile Robert Brown discovered the nucleus of the vegetable cell. In 1838 Schleiden referred all vegetable tissues to the cellular type, and traced back the plant embryo to a single nucleated cell, while in 1839 Schwann boldly extended this con-ception of plant structure and development to the animal world, and so fully constituted the "cell theory."

Schwann's cells were essentially nucleated vesicles with fluid contents which originated in an intracellular substance ; but this view was soon abandoned. Dujardin had discovered that the bodies of Foraminifera were composed of a viscous granular contractile sarcode, and Von Mohl described independently in similar terms the contents of the vegetable cell as protoplasm. This was identi-fied by Max Schultze as Dujardin's sarcode, the newer name sur-viving ; and this living matter, and not the membrane, he showed to be the essential constituent of the cell, since which his amended definition of the cell as a unit-mass of nucleated protoplasm has been generally accepted. Prevost and Dumas had noticed the segmentation of the ovum into masses as early as 1824, and these were naturally identified as cells immediately after the publication of Schwann's work. In 1846 Kolliker showed that all tissues arise from these segmentation masses, and that the multiplication of animal and vegetable cells takes place by a continuation of the same process,—that of transverse division already observed in the Protozoa.

These points gained, the attention of histologists was withdrawn for a considerable time from the scrutiny of the minute structure of the cell itself to be concentrated on the modes of origin of these unit-masses, and their subsequent differentiation and aggregation into tissues and organs. The minute structure and histogenesis of


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chichte d. Botanik; Cuvier, Hist. d.Sci.; Is. G. St-Hilaire, Hist.Nat. Gen.; Masters in Mid.-Chir. Rev., 1858, &c; also articles GOETHE, LINNAEUS, OKEN, &C. plants, as well as of at least

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the higher animals, have been studied with much and ever-increasing accuracy of detail. (See ANATOMY, HISTOLOGY, EMBRYOLOGY.) Both vegetable and animal tissues have been simply classified both according to their adult forms and according to the embryonic layers from which they respectively arise. This scrutiny of plant and animal structure over and above the special generalizations of the botanist and the zoologist has afforded much result to general histology. The improvement of technical methods has of late years largely aided the progress of discovery. A return from the study of the cell-aggregate to that of the cell has commenced, and the question of cell-structure may be said to be again paramount in histology. The process of transverse division has of late been much elucidated, and, although its complex details cannot here be entered upon, the result has been to establish a minute and thorough correspond-ence in cases so widely dissimilar as pollen-grains from a flower-bud, the epidermis of a tadpole, or the cells of a tumour—a result which obviously enhances the morphological completeness of the cell theory. Minor modes of cell-multiplication also are not without their morphological interest. Gemination, familiar in the yeast plant, occurs in other low and simple organisms, and may probably be identified with the formation of polar vesicles in ova as a modi-fication of transverse division. Schleiden had supposed all new cells to originate within pre-existing cells, and this process, known as free-cell-formation, may really be observed in various plant and animal tissues. The protoplasm groups itself round new nuclei, the new cells being in fact formed much as Schwann had in his turn supposed ; but these nuclei have repeatedly been shown to arise from segmentation of the original nucleus, and thus this pro-cess too seems a mere modification of the general one of transverse division. Conjugation, too—that coalescence of two similar cells wdiich may be observed in many Algaî, Fungi, and Protozoa—is to be considered as the undifferentiated form of that fertilization which occurs in higher animals and plants, the two apparently similar masses having become respectively differentiated into ovum and spermatozoon, or into egg-cell and antherozoid. An indefinite number of amoeboid cells sometimes flow together into a single mass,—a phenomenon regarded by some as multiple-conjugation, or perhaps more probably as an almost mechanical coalescence of exhausted cells, from which conjugation proper and finally fertili-zation may indeed have originated. The amoeboid cells of higher animals similarly unite when drawn, and this formation of Plas-modia, as these are termed, seems to be a deep-seated property of the amoeboid cell. Similarly, too, the process of rejuvenescence which occurs in many of the lowest plants and animals, such as Protococcus and Amceba, where the protoplasm passes from a rest-ing and encysted to a naked and mobile stage, has many analogues not only among the Protista but even in the tissues of higher animals, while the phases which the lowest organisms more or less exhibit—the encysted, the ciliated, the amoeboid, and the plasmo-dial—may be regarded as the fundamental forms of a "life-cycle," fully represented indeed only in such extremely low organisms as Protomyxa and Myxomycètes, yet nowhere completely suppressed. The very highest plants and animals may thus be considered as aggregates of more or less differentiated and variously arranged encysted, amoeboid, and ciliated cells, while their development and subsequent changes, their variations normal and pathological, in reality exhibit phases more or less distinct of the ancestral life-cycle.

The examination of the precise modes of cell-division, particularly in the hands of botanists (see BIOLOGY, and summary in Sachs's Vorlcsungen iiber Pflamen Physiologie, 1883), are also constantly throwing the most interesting light upon the structure of the adult organism. Thus then, in our own day as in those of Bichat or Schwann, the labours of the histologist, when inspired by higher aims than that of the mere multiplication of descriptive detail, are of supreme morphological importance, and result in the demonstra-tion of a unity of organic structure deeper even than any which we owe to Linnseus or Cuvier, Goethe or Geoffroy.





§ 4. Individuality.—Probably no subject in the whole range of biology has been more extensively discussed than that of the nature of organic individuality. The history of the controversy is of interest, since besides leading up to solid results it serves, perhaps better than any other case, to illustrate the slow emergence of the natural sciences from the influence of scholastic thought. Starting from the obvious unity and indivisibleness of Man and other higher animals, and adopting some definition such as that of Mirbel (exceptionally unmetaphysieal, however), "Tout être organisé, complet dans ses parties, distinct et séparé des autres êtres, est un individu," it was attempted times without number to discover the same conception elsewhere in nature, or rather to impose it upon all other beings, plants and animals alike. The results of different inquirers were of course utterly discrepant. It seemed easy and natural to identify a tree or herb corresponding to the individual animal, yet difficulties at once arose. Many apparently distinct plants may arise from a common root, or a single plant may be decomposed into branches, twigs, shoots, buds, or even leaves, all often capable of separate existence. These, again, are decompos-able into tissues and cells, the cells into nucleus, &c, and ultimately into protoplasmic molecules, these finally into atoms,—the inquiry thus passing outside organic nature altogether and meeting the old dispute as to the ultimate divisibility of matter. In short, as Haeckel remarks, scarcely any part of the plant can be named which has not been taken by some one for the individual. It is necessary, therefore, briefly to notice some of the principal works on the subject, and these may conveniently be taken in descending order.

While Cassini practically agreed with Mirbel in attempting to regard separate plants as individuals, the widest interpretation of the individual is that of Gallesio (1816), who proposed to regard as an individual the entire product of a single seed, alike whether this developed into a uni-axial plant extended continuously like a Banyan, or multiplied asexually by natural or artificial means like the Weeping-willow or the Canadian Pondweed, of each of which, on this view, there is only a single individual in Britain, happily discontinuous.

At once the oldest and most frequently maintained view is that which regards the bud or shoot consisting of a single axis with appendages as the plant-individual, of which the tree represents a colony, like a branched hydroid Polyp. This conception, often attributed to Aristotle, but apparently without foundation, appears distinctly in the writings of Hippocrates and Theophrastus,—the latter saying, "The bud grows on the tree like a plant in the ground." The aphorism of Linnajus, "Gemma? totidem herbal," is well known; and in this view C. F. Wolff and Humboldt concurred, while Erasmus Darwin supported it by an appeal to the facts of anatomy and development. The most influential advocate of the bud theory during the first half of the present century was, however, Du Petit-Thouars, who, although starting much as usual with a "prkicipe unique d'existence," supported his theory on extensive though largely incorrect observations on stem structure and growth. For him the tree is a colony of phytons, each being a bud with its axillant leaf and fraction of the stem and root. Passing over numerous minor authors, we come to the central work of Alex. Braun (1853), in which, as Sachs has clearly pointed out, the illegitimate com-bination of Naturphilosophie with inductive morphology reaches its extreme. He reviews, however, all preceding theories, admits the difficulty of fixing upon any as final, since the plant, physio-logically considered, is rather a dividmim than an individuum, and proposes as a compromise, or indeed as a partial cutting of the knot, the adoption of the shoot as the morphological individual, comparable to an animal, especially because, unlike the cell, leaf, &e., it includes all the representative characters of the species. Darwin and Spencer on the whole also accept the bud or shoot as at any rate the most definite individual.

The theory of metamorphosis naturally led Goethe, Oken, and others to regard the leaf as the individual, while J ohannes Midler, Steenstrup, and others adopted the same view on various physio-logical grounds. Gaudichaud elaborated a theory intermediate between this view and that of Du Petit-Thouars, according to which the plant was built up of individuals, each consisting of a leaf with its subjacent internode of stem, which was regarded as the leaf-base, and this was supported by Edward Forbes and others, while the nominally converse view—that of the leaf as a mere outward ex-pansion of the stem-segment—was proposed by Hochstetter.

Though sundry attempts at identifying various tissues, such as the fibro-vascular bundles, as the constituent individuals may be passed over, those associated with the cell theory are of great importance. Schwann decided in favour of the cell and regarded the plant as a cell-community, in which the separate elements were like the bees of a swarm,—a view virtually concurred in in all essential respects by Schleiden, Virchow, and other founders of the cell theory. Yet, although the structure and functions of the plant are ultimately and exclusively cellular, it is impossible to ignore the fact that, save in the very lowest organisms, these are subordi-nated and differentiated iuto larger aggregates, and form virtually but the bricks of a building, and hence the later theories outlined above. Of attempts to find the individual in the nucleus or the protoplasm granules it is of course unnecessary to speak further.
So far the theories of absolute individuality. The conception of relative individuality is well traced by Fiseh upwards from the more or less vague suggestions in the writings of Goethe, Boeper, and the elder De Candolle to its clear expression in Alphonse de Candolle and Schleiden, both of whom take the cell, the shoot, and the multi-axial plant as forming three successive and subordinated categories. Wageli too recognized not only the necessity of establish-ing such a series (cell, organ, bud, leafy axis, multi-axial plant) but the distinction between morphological and physiological in-dividualities afterwards enunciated by Haeckel.

Passing over the difficulties which arise even among the Protozoa (see FOKAMINIFERA), we find that a similar controversy (fully chronicled in Haeckel's Kalkselvwanvme) has raged over the in-dividuality of Sponges. While the older observers were content to regard each sponge-mass as an individual, aviewin which Lieberkufm and other monographers substantially concurred, the application of the microscope led to the view suggested by James Clark, and still stoutly supported by Saville Kent, that the Sponge is a city of amceboid or mfusorian individuals. Carter looked upon the separate ampullaceous sacs as the true individuals, while Schmidt, defining the individual by the possession of a single exhalent aperture, dis-tinguishes Sponges into solitary and social. Later, however, he terms them Zoa impcrsonalia.

For the higher animals the problem, though perhaps really even more difficult, is less prominent. As Haeckel points out, the earlier discussions and even the comparatively late essay of Johannes Miiller take an almost purely psychological or at least a physiological point of view ; and the morphological aspect of the inquiry only came forward when the study of much lower forms, such as Cestoid Worms (see PLATYHELMINTHES) or Siphonophores (see HYDROZOA), had raised the difficulties with which botanists had so long been familiar. With the rapid progress of embryology, too, arose new problems ; and in 1842 Steenstrup introduced the conception of an '' alternation of generations" as a mode of origin of distinct individuals by two methods, for him fundamentally similar, the sexual from im-pregnated females and the asexual from unimpregnated "nurses,"— a view adopted by Edward Forbes and many other naturalists, but keenly criticized by Carpenter and Huxley. In Leuckart's remark-able essay on polymorphism (1853) the Siphonophora were analysed into colonies, and their varied organs shown to be morphologically equivalent, while the alternate generations of Steenstrup were reduced to a case of polymorphism in development. Leuckart further partly distinguished individuals of different orders, as well as between morphological and physiological individuals.

In 1852 Huxley proposed the view which he still substantially maintains (see BIOLOGY). Starting from such an undoubted homo-logy as that of the egg-producing process of Hydra with a free-swimming Medusoid, he points out that the title of individual, if applied to the latter, must logically be due to the former also, and avoids this confusion between organ and individual by defining the individual animal, as Gallesio had done the plant, as the entire product of an impregnated ovum,—the swarm of Aphides or free Medusa? which in this way might belong to a single individual being termed Zooids.

In Carus's System of Animal Morphology (1853) another theory was propounded, but the problem then seems to have fallen into abeyance until 1865, when it formed the subject of a prolonged and fruitful discussion in the Principles of Biology. Adopting the cell (defined as an aggregate of the lowest order, itself formed of physio-logical units) as the morphological unit, Spencer points out that these may either exist independently, or gradually exhibit unions into aggregates of the second order, like the lower Algae, of which the individuality may be more or less pronounced. The union of such secondary aggregates or compound units into individuals of a yet higher order is then traced through such intermediate forms as are represented by the higher seaweeds or the Liverworts, from the thallus of which the axes and appendages of Monocotyledons and Dicotyledons are ingeniously derived. The shoot of a flowering-plant is thus an aggregate of the third order ; it branches into an aggregate of the fourth or higher order, and finally as a tree "acquires a degree of composition too complex to be any longer defined." Proceeding to animals, the same method is applied. The Protozoa are aggregates of the first order. These, like plants, exhibit transitions, of which Radiolarians, Foraminifera, and Sponges are taken as examples, to such definite compound wholes as Hydra ; and such secondary aggregates multiply by gemmation into permanent aggregates of the third order, which may exhibit all degrees of integration up to that of the Siphonophora, where the individualities of the Polyps are almost lost in that of the aggregate form. The whole series of articulated animals are next interpreted as more or less integrated aggregates of the third order, of which the lower Annelids are the less developed forms, the Arthropods the more highlyintegrated and individualized. Molluscs and Vertebrates are regarded as aggregates of the second order.

In 1866 appeared the latest morphological classic, the Generelle Morphologic of Haeckel. Here pure morphology is distinguished into two sub-sciences,—the first purely structural, tectology, which regards the organism as composed of organic individuals of different orders ; the second essentially stereometric, promorphology. To tectology, defined as the science of organic individuality, a large section of the work is devoted. Dismissing the theory of absolute individuality as a metaphysical figment, and starting from the view of Schleiden, De Candolle, and Nageli of several successive categories of relative individuals, he distinguishes more clearly than heretofore the physiological individual (or Won), characterized by definiteness and independence of function, from the morphological individual (or morphon), characterized similarly by definiteness of form ; of the latter he establishes six categories, as follows :—

1. Plastides (cytodes and cells), or elementary organisms.
2. Organs (cell-stocks or cell-fusions), simple or homoplastic or-
gans (tissues), or heteroplastic organs. Organ-systems, organ-apparatuses.
3. Antimeres (opposite or symmetrical or homotypic parts), e.g.,
rays of radiate animals, '' halves of bilaterally symmetrical animals."
4. Metameres (successive or homodynamous parts), e.g., stem-
segments of Phanerogams, segments or zoonites of Annelids or Vertebrates.
5. Personse, shoots or buds of plants, polyps of Ccelenterates,
&c, " individuals " in the narrowest sense among the higher animals.
6. Corms (stocks or colonies), e.g., trees, chains of Salpa?, polyp-
stocks, &c.

In his subsequent monograph on calcareous Sponges, and in a final paper, he somewhat modifies these categories by substituting one category of extreme comprehensiveness, that of the idorgan, in place of the three separate orders of organs, antimeres, and meta-meres. The idorgan (of course clearly distinguished from the physiological organ or biorgan) is finally defined as a morphological unit consisting of two or more plastids, which does not possess the positive character of the person or stock. These are distinguished into homoplasts or homo-organs and alloplasts or alloe-organs, the former including, as subdivisions, plastid-aggregates and plastid-fusions, the latter idomeres, antimeres, and metameres. The former definition of the term antimere, as denoting at once each separate ray of a radiate, or the right and left halves of a bilaterally sym-metrical animal, is corrected by terming each ray a paramere, and its symmetrical halves the antimeres. Thus an ordinary Medusoid has four parameros and eight antimeres, a Star-fish five and ten. The con-ception of the persona is largely modified, not only by withdrawing the comparison of the animal with the vegetable shoot and by omit-ting the antimere and métamete as necessary constituents, but by taking the central embryonic form of all the Metazoa—the gastrula (fig. 1) and its assumed ancestral representative, the gastrrea—as the simplest and oldest form of per-sona. The different morphological stages to which it may attain are clas-sified into three series: (1) Monax-onial inarticulate persons, i.e., uni-axial and unsegmented without anti-meres or metameres, as in Sponges, or lowest Hydroids ; (2) Stauraxonial inarticulate persons with antimeres, but without metameres, e.g., Coral, Medusa, Turbellarian, Trematodc, Bry-ozoon ; (3) Stauraxonial articulate per-sons with antimeres and metameres, e. g., Annelids, Arthropods, Vertebrates. The colonies of Protozoa are mere idor- Fio. 1.—Gastrula in optical sec-gans. True corms composed of united ^^t^JO persons, occur only among Sponges, p0re and arch-enteron), as also Hydroids, Siphonophores, Corals, Bry- outer and inner layers, ectoderm ozoa, Tunicates, and Echinoderms, of and endoderm. (AíterHaeckel.) which the apparent parameros are regarded as highly centralized per-son* of a radially-budded worm colony; and these can be classified according to the morphological rank of their constituent persona?. They usuallyarisebygemmation from a single persona, yet in Sponges and Corals occasionally by fusion of several originally distinct persons or corms. The theory of successive subordinate orders of individuality being thus not only derived from historical criticism of previous theories but brought into conformity with the actual facts of development and descent,—various groups of organisms being referred to their several categories,—the remaining problem of tectology, that of the relation of the morphological to the physio-logical individuality, is finally discussed. Of the latter, three cate-gories are proposed:—(1) the '' actual bion or complete physiological individual," this being the completely developed organic form which has reached the highest grade of morphological individuality proper to it as a representative of, e.g., its species ; (2) the "virtual bion or potential physiological individual," including any incompletely developed form of the former from the ovum upwards ; and (3) the "partial bion or apparent physiological individual," such frag-ments of the actual or virtual bion as may possess temporary inde-pendence without reproducing the species—this latter category having, however, inferior importance.

Haeckel's theory, indeed in its earlier form, has been adopted by Gegenbaur and other morphologists, also in its later form by Jäger, who, however, rejects the category of idorgan on the ground of the general morphological principle that every natural body which carries on any chemical changes with its environment becomes differentiated into more or less concentric layers ; but the subject, especially as far as animals are concerned, is again recently dis-cussed in a large work by Perrier. Starting from the cell or plastid, he terms a permanent colony a méride, and these may remain isolated like Sagitta or Botifer, or may multiply by gemmation to form higher aggregates which he terms zoides. Such zoides may he irregular, radiate, or linear aggregates, of which the two former classes especially are termed denies. The organ—Haeckel's idorgan— is excluded, since tissues and organs result from division of labour in the anatomical elements of the merides, and so have only a secondary individuality, "carefully to be distinguished from the individuality of those parts whose direct grouping has formed the organism, and which live still, or have lived, isolated from one another." Perrier further points out that undifferentiated colonies are sessile, as Sponges and Corals, while a free state of existence is associated with the concentration and integration of the colony into an individual of a higher order.

So far the various theories of the subject; detailed criticism is impossible, but some synthesis and reconciliation must be attempted. Starting from the cell as the morphological unit, we find these forming homogeneous aggregates in some Protozoa and in the early development of the ovum. But integration into a whole, not merely aggregation into a mass, is essential to the idea of individu-ality ; the earliest secondary unit, therefore, is the gastrula or meride. This stage is permanently represented by an unbranched Hydra or Sponge or by a Planarian. These secondary units may, however, form aggregates either irregular as in most Sponges, in-definitely branched as in the Hydroids and Actinozoa, or linear as in .such Planarians as Catenula. Such aggregations, colonies, or denies, not being aggregated, do not fully reach individuality of the third order. This is attained, however, for the branched series by such forms as Siphonophores among Hydrozoa, or Benilla or Pennatula among Actinozoa; for linear aggregates again by the higher Worms, and still more fully by Arthropods and Vertebrates. Aggregates of a yet higher order may occur, though rarely. A longitudinally dividing Nais or laterally branched Syflis are obviously aggregates of these tertiary units, which, on Haeckel's view, become integrated in the Echinoderm, which would thus reach a complete indivi-duality of the fourth order. A chain of Salpse or a colony of Pyro-.soma exhibits an approximation to the same rank, which is more nearly obtained by a radiate group of Botryllus around their central cloaca, while the entire colony of such an Ascidian would represent the individual of the fifth order in its incipient and unintegrated state,—these and the preceding intermediate forms being, of course, readily intelligible, and indeed, as Spencer has shown, inevitable on the theory of evolution.





The exclusion of tissues and organs from rank in this series is thus seen to necessarily follow. Ectoderm and endoderm cannot exist alone ; they and the organs into which they differentiate arise merely, as Jäger expresses it, from that concentric lamination, or, with Perrier, from that polymorphism of the members of the colony, which is associated with organic and social existence. The idea of the antimere is omitted, as being essentially a promorpho-logical conception (for a Medusoid or a Star-fish, though of widely distinct order of individuality, are equally so divisible) ; that of the metamere is convenient to denote the secondary units of a linear tertiary individual ; the term persona, however, seems un-likely to survive, not only on account of its inseparable psycho-logical connotations, but because it has been somewhat vaguely applied alike to aggregates of the second and third order ; and the term colony, conn, or deme may indifferently be applied to those aggregates of primary, secondary, tertiary, or quaternary order which are not, however, integrated into a whole, and do not reach the full individuality of the next higher order. The term zooid is also objectionable as involving the idea of individualized organs, a view natural while the medusoid gonophores of a Hydrozoon were looked at as evolved of its homologue in Hydra, whereas the latter is really a degenerate form of the former. Passing to the vegetable world, here as before the cell is the unit of the first order, while aggregates representing almost every stage in the insensible evolu-tion of a secondary unit are far more abundant than among animals. Complete unity of the second order can hardly he allowed to the thallus, which Spencer proposes to compound and integrate into tertiary aggregates—the higher plants ; as in animals the embryo-logical method is preferable, both as avoiding gratuitous hypothesis and as leading to direct results. Such a unit is clearly presented by the embryo of higher plants in which the cell-aggregate is at once differentiated into parts and integrated into a whole. Such an embryo possesses axis and appendages as when fully developed (fig. 2).^ The latter, however, being as organs meTe lateral expan-sions of the concentric layers into which the plant embryo, like the animal, is differentiated, and so neither stages of evolution nor capable of separate existence, are not entitled to individual rank. The embryo, the bud, shoot, or uni-axial plant, all thus belong to the second order of individuality, like the Hydroid they resemble. Like the lower Ccelenterates, too, aggregates of such axes are formed by branching out from their low degree of integration. Such colonies can hardly be termed individuals of the third, much less of higher order, at least without somewhat abandoning that unity of treatment of plants and animals without which philosophi-cal biology disappears. Individuality of the second order is most fully reached by the flower,—the most highly differentiated and integrated form of axes and appendages. Such a simple inflores-cence as a raceme or umbel approximates to unity of the third order, to which a composite flower-head must be admitted to have attained, while a compound inflorescence is on the way to a yet higher stage.

If, as seems probable, a nomencla-ture be indispensable for clear ex-pression, it may be simply arranged in conformity with this view. Start-ing from the unit of the first order, the plastid or monad, and terming any undilferentiated aggregate a deme, we have a monad-demc integrating into a secondary unit or dyad, this ^ÄYltÄ rising through dyad-demes into a axis and appendages, as also triad, this forming triad-demes, and the three concentric embryonic these when differentiated becoming layers.

tetrads, the Botryllus-colony with which the evolution of compound individuality terminates being a tetrad-deme. The separate living form, whether monad, dyad, triad, or tetrad, requires also some dis-tinguishing name, for wdiich persona will probably ultimately be found most appropriate, since such usage is most in harmony with its inevi-table physiological and psychological connotations, while the genea-logical individual of Gallesio and Huxley, common also to all the cate-gories, may be designated with Haeckel the ovum-product or ovum-cycle, the complete series of forms needed to represent the species being the species-cycle (though this coincides with the former save in cases where the sexes are separate, or polymorphism occurs). For such a peculiar case as Diplozoon paradoxum, where two separate forms of the same species coalesce, and still more for such heterogeneous individuality as that of a Lichen, where a composite unit arises from the union of two altogether distinct forms—Fungus and Alga,—yet additional categories and terms are required.

§ 5. Promorphology.—Just as the physiologist constantly seeks to interpret the phenomena of function in terms of mechanical, physical, and chemical laws, so the morphologist is tempted to inquire whether organic as well as mineral forms are not alike reducible to simple mathematical law. And just as the crystallo-grapher constructs an ideally perfect mathematical form from an imperfect or fragmentary crystal, so the morphologist has frequently attempted to reduce the complex-curved surfaces of organic beings to definite mathematical expression. Canon Moseley (Phil. Trans., 1838) succeeded in showing, by a combination of measurement and mathematical analysis, that the curved surface of any turbinated or discoid shell might be considered as generated by the revolution, about the axis of the shell, of a curve, which continually varied its dimensions according to the law of the logarithmic spiral. For (ioodsir this logarithmic spiral, now carved on his tomb, seemed a fundamental expression of organic curvature and the dawn of a new epoch in natural science—that of the mathematical investiga-tion of organic form—and his own elaborate measurements of the body, its organs, and even its component cells seemed to yield, now the triangle, and again the tetrahedron, as the fundamental form. But such supposed results, savouring more of the Natur-philosophie than of sober mathematics, could only serve to dis-courage further inquiry and interest in that direction. Thus we find that even the best treatises on botany and zoology abandon the subject, satisfied with merely contrasting the simple geometrical ground-forms of crystals with the highly curved and hopelessly complicated lines and surfaces of the organism.

But there are other considerations which lead up to a mathe-matical conception of organic form, those namely of symmetry and regularity. These, however, are usually but little developed, botanists since Schleiden contenting themselves with throwing organisms into three groups—first, absolute or regular ; second, regular and radiate ; third, symmetrical bilaterally or zygomorphic —the last being capable of division into two halves only in a single plane, the second in two or more planes, the first in none at all. Burmeister, and more fully Bronn, introduced the fundamental improvement of defining the mathematical forms they sought not by the surfaces but by axes and their poles ; and Haeckel has developed the subject with an elaborateness of detail and nomen-clature which seems unfortunately to have impeded its study and acceptance, but of which the main results may, with slight varia-tions chiefly due to Jäger (Lehrb. d. Zool., i. 283), be briefly out-lined.

A. ANAXONIA—forms destitute of axes, and consequently
wholly irregular in form, e.g., Amoebae and many Sponges.
B. AXONIA—forms with definite axes.

I. HOMAXONIA—all axes equal.

(a) Spheres, where an indefinite number of equal axes can be drawn through the middle point, e.g., Sphcerozoum. (J) Polyhedra, with a definite number of like axes. Of these aconsiderable number occur in nature, for example, many Radiolarians (fig. 3), pollen-grains, &c, and they are again classifiable by the number and regularity of their faces.
II. PKOTAXONIA, where all the parts are arranged round a main axis, and of these we distinguish—
1. Monaxonia, with not more than one
definite axis. Here are distinguished
(as) those with similar poles, spheroid
(Coccodiscus) and cylinder (Pyrosoma)
and (b) those with dissimilar poles,
cone (Conulina).
2. Stauraxonia, where, besides the
„ . Fio. 3.—Radiolanan (Ethmo-
mam axes, a definite number of second- sphsera), an irregular endo-
ary axes are placed at right angles, and spharic polyhedron with equi-
the stereometric ground-form becomes angular faces. Type of Hom-
a pyramid. Here, again, may be distin- axoma-
guished (a) those with polos similar, Stauraxonia homopola, where
the stereometric form is the double pyramid (fig. 4), and (b) those with
poles dissimilar, Stauraxonia hetero-
pola, where the stereometric form is the /A.

single pyramid, and where we distin-guish a basal, usually oral, pole from an apical, aboral, or anal pole. The bases of these may be either regular or irre-gular polygons, and thus a new classi-fication into Homostaura and Iletero-staura naturally arises.
?ro.4.— Pollen of Passion Flower, as example of Stauraxonia ho-mopola. Ground-form a regu-lar double pyramid of six sides.

The simpler group, the Homostaura, may have either an even or an odd number of sides, and thus among the Homostaura we have even-sided and odd-sided, single and double pyramids. In those Homostaura with an even number of sides, such as Medusa?, the radial and inter-radial axes have simi-lar poles ; but in the series with an odd number of sides, like most Echi-noderms, each of the transverse axes is half radial and half semi-radial (fig. 5). Of the group of regular double pyra-mids the twelve-sided pollen-grain of Passifiora (fig. 4) may be taken as an example, having the ground-form of the hexagonal system, the hexagonal dodecahedron. Of the equal even-sided single pyramids (Heteropola homostaura), Alcyonium, Geryonia, Aurelia may be taken as ex-amples of the eight-sided, six-sided, and four-sided pyramids, while those with an odd number of sides may be illustrated by Ophiura or Primula with five sides, and the flower of Lily or Rush with three sides.

In the highest and most complicated group, the Heterostaura, the basal poly-gon is no longer regular but amphithect (áiMpídwKTOs = double-edged). Such a polygon has an even number of sides and Pla 6._starns, an e le can be divided into symmetrical halves of Heteropola homostaura. by each of two planes intersecting at right Ground-form a regular single angles in the middle point, and thus divid- P>'™nid of five sides, ing the whole figure into four congruent polygons. The longer of these axes may be termed lateral, the shorter the equatorial or dorso-ventral ; and these two axes, along with the main axes, always define the three dimensions of space. Ctenophores (fig. 6) furnish examples of eight-sided amphithect pyramids, some Madrepore Corals of six-sided, Crucifers, some Medusae, and Cestodes of four-sided amphi-thect pyramids.

In these forms the poles of the dorso-ventral and lateral axes are similar, and, as in the preceding Monaxonia and Stauraxonia, the centre of the body is defined by a line; and they are therefore termed Centraxonia, while the Protaxonia, which are defined by their central point, are called Centrostigma. There are, however, other forms, and these the most complicated, in which the poles of at least the dorso-ventral axis are unlike, and in which the body is thus defined not with reference to a line but to a median plane, and these have accordingly received the name of Centropipeda. Their ground-form is a polygon with an even number of sides, which can only be divided into two symmetrical halves by the one median plane. It can be obtained by halving an amphithect pyramid of double the number of sides, and is consequently termed a half amphithect pyramid (fig. 7). The whole amphithect pyramid may be most con-

veniently obtained by the reduplication of the ground-form as if in a-mirror. Of half amphithect pyramids there are again two forms, termed by Haeckel Amphipleura and Zygopleura, the former in-cluding the "bilaterally symmetrical" or irregularly radiate forms of previous authors, such as Spatangus, Viola, Orchis, while the Zygopleura include forms bilaterally symmetrical in the strictest sense, in which not more than two radial planes, and these at right

Fig. 6. Fig. 7.
FIG. 6.—Ctenophore (Eucharis). Ground-form an eight-sided double amphithect pyramid.

FIG. 7.—Spatangus. Ground-form a five-sided half amphithect pyramid, angles to each other, are present. The stereometric ground-form is a half rhombic pyramid. Haeckel again divides these, according to the number of antimeres, into Tetrapleura and Dipleura.

Promorphology has thus shown that the reigning dogma of the fundamental difference of organic and mineral forms is false, and that a crystallography of organic forms is possible,—the form of the cell or the cell-aggregate differing from the crystal merely by its more or less viscous state of aggregation, its inherited peculi-arities, and its greater adaptability to the environment. The classification into bilateral and radiate forms which usually does duty for more precise promorphological conceptions must be aban-doned as hopelessly confusing essentially different forms, or at least must be rigidly restricted, —the term radial to regular and double pyramids, the term bilateral to the Centropipeda if not indeed to dipleural forms. Similarly, the topographical and relative terms, anterior and posterior, upper and under, horizontal and vertical,, must be superseded by the tenus above applied to the axes and their poles, oral and aboral, dorsal and ventral, right and left.

§ 6. Natihre of Morphological Changes.—The main forms of organic structure being analysed and classified and their stage of individu-ality being ascertained, the question next arises, by what morpho-logical changes have they arisen, and into what categories can these modes of differentiation be grouped ? They at first sight seem innumerable, yet in reality are few. Goethe somewhat vaguely generalized them for the flower as ascending and descending meta-morphosis, expansion and contraction of organs, &c. ; but the first attempt at careful enumeration seems to be that of Auguste de St-Hilaire, who recognized defects of development, adhérences, excesses of production or "dédoublements," metamorphosis and displace-ment of organs. Subsequent authors have variously treated the subject ; thus Asa Gray enumerates as modifications of the flower— coalescence, adnation, irregularity, abortion, non-alternation or anteposition, multiplication, enation, unusual development of the axis, and other morphological modifications connected with fertili-zation. These are obviously too numerous, as may best be shown by a single comparison with the view of an animal morphologist. Thus Huxley, in discussing the arrangement of the Vertebrata, recognizes only three processes of modification, not only in the ancestral evolution of the Equidae, but in the individual develop-ment of animals generally ; these are "(1) excess of development of some parts in relation to others, (2) partial or complete suppres-sion of certain parts, (3) coalescence of parts originally distinct."' It is probable that this ' ' threefold law of evolution " may include all observed cases of change, even in the flower ; thus Chorisis and Peloria maybe regarded as peculiar forms of excess, wdiile displace-ment is probably in all cases only apparent, and really due tc-adhesion or coalescence (see BIOLOGY, vol. iii. p. 681 sq.).

§ 7. Nature of Morphological Correspondence — Categories of Homology.—To indicate all the steps by which the idea of mor-phological has been distinguished from that of physiological resemblance would be to examine the whole history of morphology; it must suffice to discuss the terminology of the subject which has, as ever, served not only as an index but as an engine of progress. For these two distinct forms of resemblance the terms homology and analogy gradually became specialized, and were filially estab-lished and clearly defined by Owen in 1843,—"the former as the same organ in different animals under every variety of form and function (e.g., fore-limbs of Draco volans and wings of Bird) ; the-second as a part or organ in one animal which has the same function as another part or organ in a different animal (e.g., parachute of Draco and wings of Bird)." He further distinguishes three kinds of homology :—(1) special, being "that above defined, namely, the correspondence of a part or organ determined by its relative position and connexions with a part or organ in a different animal, the _determination of which homology indicates that such animals are constituted on a common type," e.g., basilar process of human occipital with basi-occipital of fish; (2) general, that "higher relation in which a part or series of parts stands to the fundamental or general type, involving a knowledge of the type on which the group in question is constituted," e.g., the same human bone and •centrum of the last cranial vertebra ; (3) serial homology, "repre-sentative or repetitive relation in the segments of the same skeleton " (demonstrated when general and special homologies have been determined) ; thus usually the basi-occipital and basi-sphenoid are "homotypes." These terms were henceforth accepted by naturalists ; but the criterion of analogy and homology became for Agassiz and other embryologists developmental as well as comparative, reference to the ideal archetype becoming less and less frequent. Passing over the discussions of Agassiz and Bronn, of which the latter is criticized and partly incorporated by Haeckel, we find the last-named (1) placing serial under general homology ; (2) erecting categories of homology partially corre-sponding to those of individuality,—(a) homotypy (of antimeres), hence distinct from that of Owen, (b) homodynamy (of metameres), _(c) homonomy (of parts arranged on transverse axes) ; (3) defining special homology in terms of identity of embryonic origin. In 1870 this latter point was more fully insisted upon by Ray Lan-kester, who, decomposing it into two others, proposed to supersede the term homology by homogeny, being the correspondence of common descent, and homoplasy, denoting any superinduced correspondence of position and structure in parts embryonically distinct. Thus, the fore-limb of a mammal is homogenous with that of a bird, but the right and left ventricles of the heart in both are only homoplastic, these having arisen independently since the divergence of both groups from a uni-ventrieulate ancestor in relation to similarity of physiological needs. Mivart next pro-posed to retain homology as a generic term, with homogeny and homoplasy as two species under it, and carried the analysis into great detail, distinguishing at first twenty-five, hut later fifteen, kinds of correspondence :—(1) parts similar in function only, e.g., legs of Lizard and Lobster ; (2) parts similar both in function and relative position, wings of Bat and Bird ; (3) parts of common descent, fore-limb of Horse and Rhinoceros; (4) parts of similar embryonic origin, whatever be their racial genetic relations, e.g., occipitals of Panther and Perch ; (5) parts of dissimilar embryonic origin, whatever be their racial genetic relations, e.g., legs of Diptera; (6, 7, 8, 9, 10) laterally, vertically, serially, antero-posteriorly, and radially homologous parts; (11) subordinate serial homologues, e.g., joints of antenna; (12 and 13) secondary and tertiary subordinate serial homologues ; (14 and 15) special and general homologies (in Owen's sense). In his Kalkschwammc Haeckel proposed to term homophyly the truly phylogenetic homology in opposition to homomorphy, to which genealogic basis is wanting ; and finally Von Jhering has published a repetition of Lankester's view.

In this discussion, as in that of individuality, it is evident that we are dealing with numerous logical cross-divisions largely corre-sponding, no doubt, to the complex web of inter-relations presented by nature, yet remaining in need of disentanglement. Though we must set aside analogies of functional activity, the resemblances in external shape or geometric ground-form which correspond to these, e.g., Hydrozoa and Bryozoa, Fishes and Cetaceans, mimetic organisms, are nevertheless, as our historic survey showed, the first which attract attention ; and these homoplastic or homomor-phic forms, as Haeckel has shown, come as fairly within the province of the promorphologist as do isomorphic crystals within that of his an-organological colleague the crystallographer. Here, too, would he considered "radial," "vertical," "lateral" homology, " homotypy of antimeres," and all questions of symmetry, for which Haeckel's nomenclature of homaxonial, homopolic, &c, is distinctly preferable. Entering the field of tectology or morphology in the ordinary sense, we may next consider whether two organisms com-pared are of the same category of individuality—are homocategoric ; and under this serial homology, for instance, would come as a minor division, the correspondence between the units or parts of units of a linear dyad-deme or triad. From a third point of view, that of the embryologist, we trace the development of each multi-cellular organism (1) from the embryonic layers and systems into which the secondary unit (gastrula or plant embryo) differentiates, (2) from a unit-deme or unit of the inferior order or orders of individuality. The parts and units thus recognized by ontogenetic research., respectively or successively homodermic, homosystemic, and Iwmodemie, may then conveniently be termed (indifferently save for considerations of priority) either " specially homologous," " homogenous," " homophylic," or "homogenetic," in the language of phylogenetic theory. These three great classes of morphological correspondence — promorphologica], tectological, and embryological — may or may not coincide. But the completest homology, in which all forms of resemblance unite and from which they differentiate, is that expressed in the cell theory, or rather in that ovum theory which underlies it, and which Agassiz therefore not unjustly regarded as " the greatest discovery in the natural sciences of modern times."

§ 8. Results to Taxonomy.—The advance and modification of classifications which follow each morphological advance have been pointed out above, and taxonomy thus never quite reaches a level with morphological knowledge. That it requires much reform at present is obvious. Although the dogma of the constancy of species is no longer maintained, its results survive, and perhaps a majority of groups have still to be remonographed in the generalizing spirit with which Haeckel has treated the calcareous Sponges, or Car-penter, Parker, and Brady the Foraniinifera. The union of the Protophyta and Protozoa into the Protista (a generalization which research is constantly confirming) involves a final abandonment of the mediaeval figment of three kingdoms of nature, and a revival of the Organisata of Linnaeus. Physiological prejudices, too, are not completely expelled ; hence, for instance, the constant attempts to separate Animalia and Vegetabilia by physiological character-istics, which would be irrelevant even if in themselves valid. A strictly morphological standard must be applied to the construction of classifications and the pruning of genealogical trees ; organisms are " higher "or " lower " not according to their stage of evolution in beauty or intelligence but (as Huxley has most clearly pointed out in the essay referred to under § 6) to the degree of morphological differentiation by excess, suppression, or coalescence which they exhibit. Thus the supreme position of Man in classification must be abandoned, since the Primates are simply one of the less special-ized, i. e., lower orders of Mammals, and the Mammals themselves are on the whole distinctly less specialized than the Birds, or per-haps even some of the higher Reptiles. The morphological import-ance of the '' vegetable kingdom " sinks when tested by such a standard. The Cormophytes are all nothing more than an axis with appendages, and as such may fairly be compared, not with the entire animal assemblage, but merely to that group which is homomorphic (or rather isomorphic) with them as reducible to axis and appendages too. Such a group we find in the Hydrome-dusai, which we can easily model in imagination into all the special-izations of the floral world, a single genus like Clava or Tubularia affording a starting-point for countless '' natural orders."

§ 9. Relation of Morpliology to Physiology.—Although the pure morphologist investigates laws of structure only, and rightly elimi-nates the conceptions of life, environment, and function, yet if kept permanently apart from physiological considerations his labours would be incomplete and his results inexplicable, if not indeed almost illusory. For, however deeply one penetrates through super-ficial and adaptive characters to an apparently permanent and fundamental morphological type, this is itself but an earlier adapta-tion, showing the fading traces of an earlier adaptation still. And, conversely, the most superficial of adaptive characters, if trans-mitted to numerous varying descendants, may attain high morpho-logical importance. The morphological aspect of an organism is merely statical, and, like that of an eddy or a vortex-ring, becomes only truly intelligible when viewed in its dynamic aspect; and thus, though the demonstration of the structural unity of the organic world is in itself a great result, yet the desire of a deeper explanation of form as determined by function and environment is thereby rendered all the more pressing. An example may be taken from botany. Thus Airy beautifully explains the pheno-mena of phyllotaxis as adaptations to bud-life. Or again, in a common flower, say the Dead-nettle, all the details of form are in-deed described by the systematist with equal minuteness (a pro-ceeding which, except in so far as serving for specific identification, is of no further scientific value), but receive separate interpretation from the two distinct standpoints of the morphologist and physio-logist. The latter, to whom form is important merely so far as explanatory of function, shows how the tough persistent calyx is protective against various dangers, how the corolla serves to lure the fertilizing bees, which find in its lip a landing stage and in each lateral process a hold-fast, while its hood at once protects the pollen against rain and determines the curvature of the stamens,— this curvature, as well as their didynamous arrangement, median position, and linearly arranged anther-lobes being all adaptations through the medium of the bee's hairy back to meet the similarly placed stigma of another flower,—and so on. The morphologist, on the other hand, analyses the calyx into its five constituent sepals, reduces the corolla to a regular pentamerous type, ascertains the position of the four stamens, and asserts the loss of a fifth posterior one, finds the ovary to be primitively two-celled, and thus reaches a schematic conception of a not archetypal but ancestral form. This ground-form itself, however, suggests a new train of considera-tions both morphological and physiological respecting the origin of this primeval flower from a somewhat fern-like Cryptogam, of which the foliage-leaves, the envelopes of the spore-bearing leaves, the micro- and macrosporangiosphores had become permanently differ-entiated in ascending order ; of which the microspores, doubtless through the intervention of a spore-eating insect, had come to ger-minate upon the macrosporangium instead of upon the ground ; and in which this variation (evidently advantageous, since making ferti-lization at once more certain and more economical) was aided to per-petuate itself by the contemporaneous evolution of those floral colours which are nascent even among the Thallophytes. And thus the mor-phologist, though excluding teleological and functional considera-tions from his anatomical researches, has yet a physiological ideal, and enters sooner or later upon a new series of inquiries—those of the interdependence of structure and function. Milne-Edwards's law of the physiological division of labour, Dohrn's principle of functional change, the speculations of Claude Bernard, Spencer, and Haeckel, experimental inquiries such as those of Semper, where organisms are subjected to special modifications of their environment, and the like, are all contributions to this newest and evolutionary department of morphology. Such ideas are even applied to the study of cellular morphology. Thus, Spencer points out the relation of the shapes of cells to their environments; James ingeniously explains the occurrence of cell-division by the rapid increase of hulk over surface which the growth of a solid involves, and the corresponding increase of difficulty of nutrition ; and the writer has attempted to explain the forms of free and united cells as. specializations of a (protomyxoid) cycle in which variations of func-tional activity are accompanied by the assumption of corresponding forms, the whole series of changes depending upon the properties of protoplasm under the variations in the supply of energy from the environment. Rauber, His, and others have even attempted to explain embryological phenomena in terms of the simplest cellular mechanics, but as yet such speculations are somewhat crude.

§ 10. Orientation and Subdivisions of Morphology.—The position of morphology in the classification of the sciences and the proper mode of subdividing it cannot be discussed within these limits, although the latter is especially the subject of much disagreement. The position above assumed, that of including under morphology the whole statical aspects of the organic world, is that of Haeckel, Spencer, Huxley, and most recent animal morphologists ; botanists frequently, however, still use the term under its earlier and more limited significance.- (P. GE.)



The above article was written by: Patrick Geddes, F. R. S. E..



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