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MORPHOLOGY (Gr. yopdsil, form)

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Originally appearing in Volume V18, Page 867 of the 1911 Encyclopedia Britannica.
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MORPHOLOGY (Gr. yopdsil, form), a term introduced by Goethe to denote in biology the study of the unity of type in organic form (for which the Linnaean term " Metamorphosis " had formerly been employed). It now usually covers the entire science of organic form. There are numerous restricted senses of the term in various sciences, but here we shall deal with it as a substantive side of zoology and botany. 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 begins with Aristotle. Founder of comparative anatomy and taxonomy, he established eight great divisions (to which are appended certain minor groups)—Viviparous Quadrupeds, Birds, Oviparous Quadrupeds and Apoda, Fishes, Malakia, Malacostraca, Entoma, and Ostracodermata—distinguishing the first four groups as Enaima (" with blood ") from the remaining four as Anaima (" bloodless "). In these two divisions we recognize the Vertebrata and Invertebrata of J. B. P. A. Lamarck, the first four groups corresponding with the Mammals, Birds, Reptiles, Fishes, whilst the others agree more loosely with the Cephalopods, Crustacea, Insecta, and Echinoderms with Mollusca other than the Cephalopods. Far from committing the mistake 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 special yfm'or beside the Viviparous Quadrupeds, more on account of their absence of limbs than of their aquatic habit. Not only is his method inductive, and his groups founded on the aggregate of known characters, but he foreshadows such generalizations as those of the correlation of organs, and ofthe progress of development from a general to a special form after-wards established by G. L. Cuvier and K. E. 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 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. Even after the reappearance of Aristotle's works in the 13th century, little can be recorded but revivals of his conclusions. Monographs on groups of plants and animals frequently appeared, those of P. Belon on Birds and G. 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 initiated by Vesalius, disappeared for centuries, unappreciated save by the surgecn Ambroise Pare. B. Palissy, like Leonardo da Vinci before him, discerned the true nature of fossils; and such flashes of 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 W. Harvey anticipated the generalizations of modern embryology by his researches on development and his theory of epigenesis. The encyclopaedic period of which Gesner is the highest representative was continued by Aldrovandi and others in the 17th century; but, aided by the Baconian movement, then influencing all scientific minds, it developed into one of genuinely systematic aim. At this stage of progress the most important part was taken by John Ray, whose classificatory labours among plants and animals were crowned with success. He first expelled the fabulous monsters and prodigies of which the encyclopaedists had handed on the tradition from medieval times, and succeeded, particularly among plants, in distinguishing many natural groups, for which his own terms sometimes survive—e.g. Dicotyledons and Monocotyledons, Umbelliferae and Leguminosae. The true precursor of Linnaeus, he introduced the idea of species in natural history, and reformed the practice of definition and terminology. Of the works which followed up Ray's systematic labours, none can be even named until we come to those of his great successor Linnaeus, whose grasp of logical method and 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, not only by the introduction of the binomial nomenclature and the renovation of descriptive terminology and method, but by the subordination of the species under the successive higher categories of genus, order and class, so reconciling the analytic and synthetic tendencies of his predecessors. Although the classification of plants by the number of their essential organs 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 extended those fragments of the natural system with which A. de Jussieu soon afterwards began to build. His classification of animals, too, was largely natural, and, though on the whole he lent his authority to maintain 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 investigator. While the artificial system was at the zenith of its usefulness, Bernard de Jussieu was arranging his gardens on the lines afforded by the fragmentary natural system of Linnaeus. His ideas were elabcrated by his nephew Antoine de Jussieu, who 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 Prodromus systematis naturalis (20 vols., 1818-1873), in which 8o,000 species were described and arranged. Mean-while the penetrative genius of Brown enabled him to unravel such structural complexities as those of Conifers and Cycads, Orchids and Proteaceae, thus demonstrating the possibility of ascertaining the systematic position of even the most highly modified floral types. Both Candolle and 3rown were thus no mere systematists, but genuine morphologists of the modern school. The labours of Bernard and Antoine de Jussieu initiated a parallel advance in zoology, the joint memoir on the classification of mammals with which Cuvier and Geoffroy St-Hilaire almost began their career receiving 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 A. von Haller and J. 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 century, he opens the era of detailed anatomical research united with exact comparison and clear generalization. The Regne animal (1817) and the theory of types (vertebrate, molluscan, articulate, and radiate) are the results of this union of analysis and synthesis 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. In Germany, L. H. Bojanus, J. F. Meckel, C. T. E. von Siebold and Johannes Muller, with his many pupils, carried on the work; in France, too, a succession of brilliant anatomists, such as A. De Quatrefages, A. Milne-Edwards and H. de Lacaze-Duthiers, were his intellectual heirs; and in England he has been admirably represented by Sir R. Owen. It is now necessary to return to Linnaeus, whose more speculative writings contain, though encumbered by fantastic hypo-theses, the idea of floral metamorphosis. About the same time, and quite independently, C. F. Wolff, the embryologist, 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 metamorphosed into bud-scales or floral parts through diminution of vegetative force. Thirty years later the same view was again independently developed by Goethe in his now well-known pamphlet (Versuch die Metamorphose der Pflanzen zuerklaren, 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 fundamental organ—the leaf—and all plants are in like manner to be viewed as modifications of a common type—the Urpflanze. 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 into all the extravagances of that system; or whether, as E. H. Haeckel maintains, it represented a concrete ancestral form, so anticipating the view of modern evolutionists, it is certain that to him F. W. S. von Schelling was indebted for the foundation upon which he erected his philosophic edifice, as also that Goethe shared the same ideas. 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. Independently of Vicq d' Azyr, he discovered the human premaxillary bone; independently of L. Oken, he proposed the vertebral theory of the skull; and before S. C. Savigny, he discerned that the jaws of insects were the limbs of the head. In 18r3 A. P. de Candolle published his Theorie elementaire de la botanique, which he developed into the classic Organographie vegetale (1827). 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 advance was the investigation by W. P. Schimper and A. 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 uberhaupt, besonders zur Morphologic). It did not come into use in botany until its popularization by Auguste de St-Hilaire in his Morphologic vegetale (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 Naturphilosophie of Schelling and its countless modifications by his followers, its mystic theories of " polarization " and the like, its apparatus of assumption and abstraction, hypothesis and metaphor, cannot here be discussed, its undoubted services must not be forgotten, since it stimulated innumerable reflective minds to the earnest study of natural science, gave a powerful impulse to the study of comparative anatomy and vindicated the claims of philosophic synthesis over those of 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-morphological dreams there occasionally occurred suggestions of the greatest fruitfulness—notably, for instance, the independent statement of the vertebral theory of the skull. 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, had greater influence on the progress of science than either. He started from the same studies of anatomical detail as Cuvier, but, influenced by Buffon's view of unity of plan and by the evolutionary doctrines of Lamarck, 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 Philosophic 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 connexions," 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 teratology, 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, however, in eager pursuit of often deceptive homologies, he wandered into the transcendentalism of the Naturphilosophie, and seems utterly to have failed to appreciate either the type theory of Cuvier or the discoveries of Von Baer. He defended Buffon's and Bonnet's earlier view of unity of plan in nature; and the controversy reached its climax in 183o, 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 death-blow. 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 progress, 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. The further evolution of the idea of homology is sketched below, while the extent and rapidity of the subsequent progress of the knowledge of all the structural aspects of plants and animals alike make a historical survey impossible up to the appearance of the Origin of Species (1859). The needful 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 organisms, which in various and varying environments, have undergone differentiation into the vast multitude of existing 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 rudimentary 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 Linnaean 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 Haeckel's Generelle Morphologic (1866), while the valuable contemporaneous Principles of Biology of Herbert Spencer also gave special attention to the relation of morphology to physiology. 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 C. F. B. Mirbel, " Tout titre organise, complet dans ses parties, distinct et separe des autres titres, 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 decomposable 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 H. Cassini practically agreed with Mirbel in attempting to regard separate plants as individuals, the widest interpretation of the individual is that of G. Gallesio (1816), who proposed to regard as an individual the entire product of a single seed, alike whether this developed ihto 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 Linnaeus, " Gemmae totidem herbae," 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 xvIII. 28and development. The most influential advocate of the bud theory during the first half of the 19th century was, however, Du Petit-Thouars, who, although starting much as usual with a " principe 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 combination 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, physiologically considered, is rather a dividuum 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, &c., 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, ()ken and others to regard the leaf as the individual, while Johannes Muller, J. J. S. Steenstrup and others adopted the same view on various physiological grounds. C. Gaudichaud elaborated a theory inter-mediate 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 expansion of the stem-segment—was proposed by C. F. 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. T. 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 M. Schleiden, R. Virchow and other founders of the cell theory. Yet, although the structure and functions of the plant are ultimately and specially cellular, it is impossible to ignore the fact that, save in the very lowest organisms, these are subordinated and differentiated into 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 unnecessary to speak further. So far the theories of absolute individuality. The conception of relative individuality was first clearly expressed by 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. K. W. von Nageli too recognized not only the necessity of establishing such a series (cell, organ, bud, leafy axis, multi-axial plant) but the distinction between morphological and physiological individualities afterwards enunciated by Haeckel. Passing over the difficulties which arise even among the Protozoa we find that a similar controversy (fully chronicled in Haeckel's Kalkschwdmme) has raged over the individuality of sponges. While the older observers were content to regard each sponge-mass as an individual, a view in which J. N. Lieberkiihn and other monographers substantially concurred, the application of the microscope led to the view suggested by James Clark, and stoutly supported by Saville Kent, that the sponge is a city of amoeboid or infusorian individuals. H. J. Carter looked upon the separate ampullaceous sacs as the true individuals, while others, defining the individual by the possession of a single exhalent aperture, distinguish sponges into solitary and social. 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 Muller 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 PLATYELMIA) 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 Streenstrup 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 impregnated females and the asexual from unimpregnated " nurses " —a view adopted by Edward Forbes and many other naturalists, but keenly criticized by W. B. Carpenter and T. H. Huxley. In R. Leuckart's remarkable 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, starting from such an undoubted homology as that of the egg-producing process of Hydra with a free-swimming Medusoid, pointed out that the title of individual, if applied to the latter, must logically be due to the former also, and avoided this confusion between organ and individual by defining the individual 11 animal, as Gallesio had done the plant, as the entire product of an impregnated ovum—the swarm of Aphides or free Medusae 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 physiological units) as the morphological unit, H. 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 highly integrated and individualized. Molluscs and vertebrates are regarded as aggregates of the second order. In 1866 appeared a morphological classic, the Generelle Morphologie 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 bion), 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: I. Plastides (cytodes and cells), or elementary organisms. 2. Organs (cell-stocks or cell-fusions), simple or homoplastic organs (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. Personae, shoots or buds of plants, polyps of Coelenterates, &c., " individuals " in the narrowest sense among the higher animals. 6. Corms (stocks or colonies), e.g. trees, chains of Salpae, 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 metameres. 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 symmetrical animal, is corrected by terming each ray a paramere, and its symmetrical halves the antimeres. Thus an ordinary Medusoid has four parameres and eight antimeres, a starfish five and ten. The conception of the persona is largely modified, not only by with-drawing the comparison of the animal with the vegetable shoot and by omitting the antimere and metamere as necessary constituents, but by taking the central embryonic form of all the Metazoa—the gastrula (fig. I) and its assumed ancestral representative, the gastraea—as the simplest and oldest form of persona. The different morphological stages to which it may attain are classified into three series: (I) Monaxonial inarticulate personae, i.e. uniaxial and unsegmented without antimeres or metameres, as in sponges or lowest Hydroids; (2) Stauraxonial inarticulate personae with antimeres, but without metameres, e.g. coral, medusa, turbellarian, trematode, bryozoon; (3) Stauraxonial articulate personae with antimeres andmetameres, e.g. annelids, arthropods, vertebrates. The colonies of protozoa are mere idorgans. True corms, composed of united personae, occur only among sponges, hydroids, siphonophores, corals, bryozoa, tunicates and echinoderms, of which the apparent parameres are- regarded as highly centralized personae of a radially budded worm colony; and these can be classified according to the morphological rank of their constituent personae. They usually arise by gemmation from a single persona, yet in sponges and corals occasion-ally 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 j VGo~o.~° theories but brought into conformity c + i with the actual facts of development (After Haeckel.) and descent—various groups of organ- doderm. __ logical individual," this being the com- pletely 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 fragments of the actual or virtual bion as may possess temporary independence without reproducing the species—this latter category having, however, inferior importance. Haeckel's theory, indeed in its earlier form, has been adopted by C. Gegenbaur and other morphologists, also in its later form by G. Jager, 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 be-comes differentiated into more or less concentric layers; but the subject, especially as far as animals are concerned, was again discussed in a large work by E. Perrier. Starting from the cell or plastid, he terms a permanent colony a meride, and these may re-main isolated like Sagitta or Rotifer, or may multiply by gemmation to form higher aggregates which he terms zoides. Such zoides may be irregular, radiate or linear aggregates, of which the two former classes especially are termed demes. 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 the 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 individuality; the earliest secondary unit, therefore, is the gastrula or meride. This stage is permanently represented by an unbranched hydroid 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 demes, 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 Renilla 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 Syllis are obviously aggregates of these tertiary units, which, on Haeckel's view, become integrated in the Echinoderm, which would thus reach a complete individuality of the fourth order. A chain of Salps or a colony of Pyrosoma 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 Jager 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 promorphological conception (for a medusoid or a starfish, though of widely distinct order of individuality, is equally so divisible) ; that of the metamere is convenient to denote the secondary units of a linear tertiary individual; the term persona, however, seems unlikely to survive, not only on account of its inseparable psychological connotations, but because it has been somewhat vaguely applied alike to aggregates of the second and third order; and the term colony. corm or deme may indifferently be applied to those aggregates of primary, secondary, tertiary or quaternary order which are not, however, integrated intq 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 locked at as evolved of its homologue in Hydra, whereas the latter may be 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 evolution of a secondary unit are far more abundant than among animals. Complete unity of the second order can hardly be 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 mere lateral expansions 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 uniaxial plant, all thus belong to the second order of individuality, like the hydroid they resemble. Like the lower coelenterates, 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 philosophical 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 inflorescence 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 nomenclature be indispensable for clear expression, it may be simply arranged in conformity with this view. Starting from the unit (After Sachs.) of the first order, the plastid or monad, FIG. 2._ Embryo of and terming any undifferentiated ag- Dicotyledon, showing inci- gregate a deme, we have a monad-deme axi asealso the three concentric dyad integrating this rising through secondary denies, embryonic layers. into a triad, this forming triad-demes, and these when differentiated becoming 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 distinguishing name, for which persona will probably ultimately be found most appropriate, since such usage is most in harmony with its inevitable physiological and psychological connotations, while the genealogical individual of Gallesio and Huxley, common also to all the categories, 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 paredoxum, 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. 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 crystallographer 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 dimensions867 according to the law of the logarithmic spiral. For Goodsir 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 investigation of organic form—and his cwn 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 Naturphilosophie than of sober mathematics, could only serve to discourage 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 mathematical 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. H. C. C. Burmeister, and more fully H. G. 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 nomenclature which seems unfortunately to have impeded its study and acceptance, but of which the main results may, with slight variations chiefly due to Jager (Lehrb. d. Zool. i. 283), be briefly outlined.
End of Article: MORPHOLOGY (Gr. yopdsil, form)
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PAUL CHARLES MORPHY (1837–1884)

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