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MOLLUSCA

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Originally appearing in Volume V18, Page 673 of the 1911 Encyclopedia Britannica.
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MOLLUSCA , one of the See also:

great " phyla," or sub-kingdoms, of the See also:animal See also:pedigree or See also:kingdom. The See also:shell-bearing forms belonging to this See also:group which were known to See also:Linnaeus were placed by him (in 1748) in the third See also:order of his class Vermes under the name " Testacea," whilst the Echinoderms, Hydroids and Annelids, with the naked Mollusca, formed his second order termed " Zoophyta." Ten years later he replaced the name " Zoophyta " by " Mollusca," which was thus in the first instance applied, not to the Mollusca at See also:present so termed, but to a group consisting chiefly of other organisms. Gradually, however, the See also:term Mollusca became used to include those Mollusca formerly placed among the " Testacea," as well as the naked Mollusca. It is important to observe that the term gaXaraa, of which Mollusca is merely a latinized See also:form, was used by See also:Aristotle to indicate a group consisting of the ctittle-fishes only. As now classified, the Mollusca consist of the following subdivisions:- Grade A.—Isopleura. Class I.—Amphineura (see See also:CHIToN). Grade B.—Prorhipidoglossomorpha. Class II.—See also:Gastropoda (q.v.). Class III.—See also:Scaphopoda (q.v.). Class IV.—See also:Lamellibranchia (q.v.). Grade C.—Siphonopoda. Class V.—See also:Cephalopoda (q.v.).

See also:

History of See also:Classification.—The definite erection of the Mollusca into the position of one of the great See also:primary See also:groups of the animal kingdom is due to See also:George See also:Cuvier (1788-1800), who largely occupied himself with the See also:dissection of representatives of this type.l* An See also:independent anatomical investigation of the Mollusca had been carried on by the remarkable Neapolitan naturalist Poli (1791), whose researches° were not published until after his See also:death (1817), and were followed by the beautiful See also:works of another Neapolitan zoologist, the illustrious Delle Chiaje.° The embranchement or sub-kingdom Mollusca, as defined by Cuvier, included the following classes of shellfish: (I) the cuttles or poulps, under the name CEPHALOPODA; (2) the snails. whelks and slugs, both terrestrial and marine, under the name GASTROPODA; (3) the See also:sea-butterflies or winged-snails, under the name See also:PTEROPODA; (4) the clams, mussels and oysters, under the name ACEPIALA; (5) the See also:lamp shells, under the name BRACHICPODA; (6) the sea-squirts or ascidians, under the name NUDA; and (7) the barnacles and sea-acorns, under the name CIRRIOPODA. The See also:main limitations of the sub-kingdom or phylum Mollusca. as laid down by Cuvier, and the See also:chief divisions thus recognized within its limits by him, hold See also:good to the present See also:day. At the same See also:time, three of the classes considered by him as Mollusca have been one by one removed from that association in consequence of improved knowledge, and one additional class, incorporated since his day with the Mollusca with See also:general approval, has, after more than See also:forty years, been again detached and assigned an independent position owing to newly acquired knowledge. The first ofCuvier's classes to be removed from the Mollusca was that of the Cirrhopoda. Their See also:affinities with the See also:lower See also:Crustacea were recognized by Cuvier and his contemporaries, but it was one of the brilliant discoveries of that remarkable and too-little-honoured naturalist, J. See also:Vaughan See also:Thompson, of See also:Cork, which decided their position as Crustacea. The metamorphoses of the Cirrhopoda were described and figured by him in 183o in a very See also:complete manner, and the legitimate conclusion as to their affinities was formulated by him.° Thus it is to Thompson (1830), and not to Burmeister (1834), as erroneously stated by Keferstein, that the merit of this See also:discovery belongs. The next class to be removed from Cuvier's * These figures refer to the Bibliography at the end of the See also:article. 670 Mollusca was that of the Nuda, better known as See also:Tunicata. In 1866 the See also:Russian embryologist Kowalewsky startled the zoological See also:world with a See also:minute See also:account of the developmental changes of Ascidia, one of the Tunicata,6 and it became evident that the affinities of that class were with the See also:Vertebrata, whilst their structural agreements with Mollusca were only superficial. The last class which has been removed from the Cuvierian Mollusca is that of the Lamp-shells or See also:Brachiopoda. The history of its See also:dissociation is connected with that of the class, viz. the See also:Polyzoa or Bryozoa, which has been both added to and again removed from the Mollusca between Cuvier's date and the present day.

The name of J. Vaughan Thompson is again that which is primarily connected with the history of a Molluscan class. In 183o he pointed out that among the numerous kinds of "polyps" at that time associated by naturalists with the Hydroids, there were many which had a See also:

peculiar and more elaborate type of organization, and for these he proposed the name Polyzoa. Subsequently6 they were termed Bryozoa by See also:Ehrenberg (1831). See also:Henri Milne-See also:Edwards in 1844 demonstrated the affinities of the Polyzoa with the Molluscan class Brachiopoda, and proposed to See also:associate the three classes Brachiopoda, Polyzoa and Tunicata in a large group " Molluscoidea," co-See also:ordinate with the remaining classes of Cuvier's Mollusca, which formed a group retaining the name Mollusca. By subsequent writers the Polyzoa have in some cases been kept apart from the Mollusca and classed with the " Vermes ; whilst by others they have, together with the Brachiopoda, been regarded as true Mollusca. Increase of knowledge has now, however, established the conclusion that the agreement of structure supposed to obtain between Polyzoa and true Mollusca is delusive; and accordingly they, together with the Brachiopoda, were removed from the Molluscan phylum by Lankester in his article in the 9th edition of this See also:work (on the which present article is based). Further details in regard to this, the last revolution in Molluscan classification, will be found in the article POLYZOA. As thus purified by successive advances of embryological See also:research, the Mollusca were reduced to the Cuvierian classes of Cephalopoda, Pteropoda, Gastropoda and Acephala. Certain modifications in the disposition of these classes are naturally enough rendered necessary by the vast See also:accumulation of knowledge as to the See also:anatomy and See also:embryology of the forms comprised in them. Foremost among those who between 1840 and 188o laboured in this See also:field are the See also:French zoologists Henri Milne-Edwards' and Lacaze Duthiers,10 to the latter of whom we owe the most accurate dissections and beautiful illustrations of a number of different types. To See also:Kolliker," See also:Gegenbaur, a and more recently Spenger,13 amongst See also:German anatomists, we are indebted for See also:epoch-making researches of the same See also:kind.

In See also:

England, See also:Owen's anatomy of the pearly See also:nautilus," See also:Huxley's discussion of the general See also:morphology of the Mollusca," and Lankester's embryological investigations," have aided in advancing our knowledge of the group. Two remarkable works of a systematic See also:character dealing with the Mollusca deserve mention here-the See also:Manual of the Mollusca, by Dr S. P. See also:Woodward, a See also:model of clear systematic exposition, and the exhaustive See also:treatise on the Malacozoa or Weichthiere by See also:Professor Keferstein of See also:Gottingen, published as See also:part of See also:Bronn's Klassen and Ordnungen See also:des Thier-Reichs. The arrangement adopted by See also:Ray Lankester in the 9th edition, of the Ency. Brit. (See also:art. " Mollusca "; 1883) was as follows: Of the four Cuvierian classes mentioned above, the Pteropoda were See also:united with the Cephalopoda, on account of the apparent similarity of the cephalic tentacles in some of the former to the arms of the latter. An additional class was instituted for the reception of Dentalium and its few See also:allies, and for this class Bronn's name Scaphopoda was used. The Chitons and their allies were placed under the Gastropoda, as a distinct See also:branch called Isopleura, and for the Acephala de See also:Blainville's name Lamellibranchia was substituted. The latter were regarded as forming a distinct branch, See also:equivalent in See also:rank to the other three classes together, the latter all possessing the radula which is wanting in Lamellibranchs. Since the 9th edition of the Ency.

Brit. was published important advances have been made in our knowledge of the Mollusca, as the result of researches largely due to the See also:

interest excited in the subject by Lankester's article. See also:Attention has been especially directed to the investigation of the most See also:primitive forms in each group, and accordingly we can now form much more definite conceptions of the phylogeny and See also:evolution of the various classes. The most important and extensive contributions to this progress have been made by the Belgian zoologist, Dr See also:Paul Pelseneer, who has made the Mollusca his See also:special study. The Chitonidae and the Aplacophora are now separated from the Gastropoda and raised to the rank of a distinct class, under the name of Amphineura. On the other See also:hand, Boas and Pelseneer have shown that the Pteropoda have nothing to do with the Cephalopoda, but are Gastropoda modified for a pelagic See also:life; they are therefore now united with the Gastropoda. The Lamellibranchia are no longer regarded as a distinct branch in contrast to the remaining Mollusca; according to Pelseneer they are allied to the Gastropoda and Seaphopoda, all three classes being derived from a See also:common hypothetical ancestor, called Prorinpidoglossum. These three classes have therefore been united by Grobben into one branch or grade, the t' rorhipidoglossomorpha. General Characters of the Mollusca.—The forms comprised in the various groups, whilst exhibiting an extreme range of variety in shape, as may be seen on comparing an See also:oyster, a cuttle-See also:fish, and a sea-slug such as See also:Doris; whilst adapted, some to life on dry See also:land, others to the depths of the sea, others to rushing streams; whilst capable, some of See also:swimming, others of burrowing, crawling or See also:jumping, some, on the other hand, fixed and immobile; some amongst the most formidable of carnivores, others feeding on See also:vegetable mud, or on the minutest of microscopic organisms—yet all agree in possessing in common a very considerable number of structural details which are not possessed in common by any other animals. The structural features which the Mollusca do possess in common with other animals belonging to other great phyla of the animal kingdom arc those characteristic of the Coelomata, one of the two great grades (the other and lower being that of the See also:Coelentera) into which the higher animals, or Metazoa as distinguished from the See also:Protozoa, are divided. The Metazoa all commence their individual existence as a single See also:cell or plastid, which multiplies itself by transverse See also:division. Unlike the cells of Protozoa, these embryonic cells of the Metazoa do not remain each like its See also:neighbour and capable of independent life, but proceed to arrange themselves into two layers, taking the form of a See also:sac. The cavity of the two-cell-layered sac or diblastula thus formed is the primitive gut or See also:arch-enteron.

In the Coelentera, whatever subsequent changes of shape the little sac may undergo as it grows up to be See also:

polyp or jelly-fish, the See also:original arch-enteron remains as the one cavity pervading all regions of the See also:body. In the Coelomata, on the other hand, there is another cavity, dividing the body-See also:wall into two layers: an See also:internal layer surrounding the gut, and an See also:external layer. This cavity is excavated in a third See also:mass of cells distinct from the cells lining the gut, forming the endoderm, and the cells covering the See also:surface of the body, the ectoderm. This third mass of cells is the mesoderm. The Mollusca agree in being coelomate with the phyla Vertebrata, Platyhelmia (See also:flat-See also:worms), See also:Echinoderma, See also:Appendiculata (See also:insects, ringed-worms, &c.), and others—in fact, with all the Metazoa except the See also:sponges, See also:corals, polyps, and medusae. In common with all other Coelomata, the Mollusca are at one See also:period of life possessed of a prostomium or region in front of the mouth, which is the essential portion of the " See also:head," and is connected with the See also:property of forward locomotion in a definite direction and the steady See also:carriage of the body (as opposed to rotation of the body on its See also:long See also:axis). As a result, the Coelomata, and with them the Mollusca, present (in the first instance) the general See also:condition of body known as bilateral symmetry; the dorsal is differentiated from the ventral surface, whilst a right and a See also:left See also:side similarito, or rather the complements of, one another are permanently established. In common with all other Coelomata, the Mollusca have the mouth and first part of the alimentary See also:canal which leads into the met-enteron formed by a special invagination of the See also:outer layer of the primitive body-wall, not to be confounded with that which often, but not always, accompanies the antecedent formation of the archenteron; this invagination is termed the stomodaeum. Similarly an anal See also:aperture is formed in connexion with a special invagination which meets the hinder part of the met-enteron, and is termed the proctodaeum. The coelom is primarily and essentially the generative cavity: the reproductive cells arise from its walls, i.e. from the coelomic epithelium. True nephridia do not primarily open into the coelom, as was formerly taught, but are See also:intra-cellular ducts in the mesoderm. Such See also:organs are absent in Mollusca in the adult See also:state, but a pair of nephridia usually occurs in the larva.

The coelom opens to the exterior by ducts which are primarily genital ducts by which the ova or sperms are discharged. These ducts, however, as well as the coelomic epithelium, may assume excretory functions. In Mollusca the coelom is reduced and consists of two Tarts, the pericardial cavity which surrounds the See also:

heart, and the cavity of the gonads or generative organs. There is usually one pair of coelomic ducts leading from the pericardium to the exterior, and these are the excretory organs or kidneys, formerly known as the organs of Bojanus. The walls of the pericardium are also excretory in parts, these parts forming the pericardial glands. In the See also:majority of Mollusca the gonads are provided with a pair of ducts of their own. There are thus two pairs of coelomic ducts. This fact gives rise to the question whether the Mollusca are to be regarded as primitively segmented animals or not. In animals which exhibit typical segmentation or See also:metamerism, such as segmented worms (See also:Chaetopoda), each segment or metamere possesses its own coelomic cavity, a pair of coelomic ducts, and a pair of nephridia. The structure of the Mollusca in the greater number of cases agrees with the See also:hypothesis that the primitive form was unsegmented, and therefore had but one pair of coelomic ducts and one pair of nephridia. In existing forms the latter disappear in the adult. In the most primitive forms of several classes there are no distinct genital ducts, the gonads when mature discharging into or through the kidneys.

Among the Gastropoda, in the Aspidobranchia, there is no genital duct, and the gonad opens into the right See also:

kidney; in the more modified forms the left kidney alone is functional, the right has been converted into the genital duct. Among the Lamellibranchia again the kidneys serve as genital ducts in the Protobranchia and some Filibranchia. In the higher forms the opening of the gonad is shifted more and more towards the external aperture of each kidney until finally it is situated on the external surface, and thus the gonad secondarily acquires an independent aperture. In the Scaphopoda there is no distinct genital duct, the relations are as in Aspidobranchia. Among the Amphineura we find one pair of coelomic ducts in the Aplacophora, two pairs in the Chitons. In the former the genital coelom and the pericardial coelom are continuous and the reproductive cells See also:escape by the renal ducts. In the Chitons or Polyplacophora, on the other hand, the two cavities are See also:separate, and there are independent genital ducts. It is possible therefore to regard the latter condition as secondary, and to conclude that the separate genital ducts have been derived from the original single pair of coelomic ducts, as in Lamellibranchs. The Cephalopoda, however, do not harmonize so well with this view. The earliest forms of this class geologically are the Nautiloidea. Assuming that these ancestral forms resembled the existing Nautilus in their internal anatomy, they had two pairs of renal ducts and one pair of genital ducts, which would apparently indicate, not a single metamere or unsegmented body, but three metameres. There are however only two pairs of branchiae.

The Dibranchia, with only one pair of branchiae, one pair of renal organs, and one pair of genital ducts, are much more See also:

recent, not appearing till the end of the Secondary epoch, and therefore must be regarded as descended from the Tetrabranchia. The latter are represented in the Upper See also:Cambrian formations, together with Lamellibranchia and Gastropoda, and there are no earlier Molluscan fossils than these. Palaeontology therefore throws no See also:light on the question whether the metameric or the unsegmented Mollusca were the earlier. The development of the Cephalopoda affords at present no better See also:evidence that the metamerism is secondary. That of Nautilus, which would be most important in this inquiry, is unfortunately still unknown. In the Dibranchia true nephridia have not been detected in the embryo, nor has it been shown that the genital ducts are derived from the renal tubes. On the other hand, there is no evidence that the forms which show no metamerism, such as the Gastropoda, are descended from metameric ancestors. On the whole, then, the most probable conclusion is that the original ancestral form of the Mollusca was unsegmented, possessed one pair of true nephridia, and one pair of coelomic ducts whose See also:function was to conduct the generative products to the exterior. The chief types of Mollusca were already differentiated at the beginning of the See also:geological See also:record, and the metamerism which occurs in the Cephalopoda has been evolved within the limits of that class. External Characters.—The characteristic organs of Mollusca are the See also:mantle and shell, the See also:foot, the ctenidia and the radula,of which all but the last are external. The original form was bilaterally symmetrical, and this symmetry is retained in all the classes except the Gastropoda. At the anterior end the head is differentiated; it bears the sense-organs, and contains the See also:muscular pharynx within which is the radular apparatus.

The See also:

rest of the body consists of the foot ventrally and the visceral mass dorsally. The foot is a muscular mass without cuticle or See also:skeleton, excepting certain cuticular structures such as the byssus of Lamellibranchs and the operculum of Gastropods, which do not aid in locomotion. The foot is usually the only See also:organ of locomotion. It corresponds to the ventral part of the body-wall in other animals. The muscular See also:tissue of the dorsal body-wall is much reduced and the integument here is thin and a.br. See also:lat)"\*'b' Fm. r.—Ctenidia of various Mollusca (original). A, Of Chiton: ft., fibrous tissue; a.b.v., afferent See also:blood-See also:vessel; e.b.v., efferent blood-vessel; g.l., laterally paired lamellae. B, Of See also:Sepia: letters as in A. C, Of Fissurella: letters as in A. D, Of Nucula: d, position of axis with blood-vessels; a, inner: b and c, outer See also:row of lamellae. E, Of Paludina: i, See also:intestine See also:running parallel with the axis of the ctenidium and ending in the anus a; be., rows of elongate processes corresponding to the two See also:series of lamellae of the upper figures. soft.

The external epithelium of the dorsal region secretes the shell. Between the edge of the shell and the foot there is a groove or cavity, chiefly See also:

developed laterally and posteriorly. The dorsal border of this groove is extended outwards and downwards as a See also:fold of the integument. There is some confusion of terms here: some writers See also:call the See also:free fold the mantle or See also:pallium, and this is the proper use of the term; but others apply the term to the whole of the dorsal integument, including both the projecting fold and the part covering the viscera. The shell extends to the edge of the mantle-fold, and the cavity between the mantle and the side of the body is the pallial chamber. This chamber serves two purposes: it is primarily the See also:respiratory cavity containing the gills, but it also serves to enclose the body so that the latter is surrounded by the shell, from which the head and foot can be protruded at the will of the animal. The shell consists of an organic basis the substance of which is called conchiolin, impregnated with carbonate of See also:lime, with a small proportion, 1–2 %, of phosphate of lime. On the outside of the shell is a non-calcified layer of conchiolin called the periostracum, secreted by the thickened edge of the mantle. The See also:zone of the external surface of the mantle within the edge secretes a layer formed of prisms of See also:calcite; the rest of the epithelium from this zone to the See also:apex secretes the inner layer of the shell, composed of successive laminae; this is the nacreous layer, and in certain See also:species has a commercial value as nacre or See also:mother-of-See also:pearl. Thus the growth of the shell in extent is due to additions to the prismatic layer at the edge, its growth in thickness to new layers of nacre deposited on its inner surface. In many cases in various classes the mantle is reflected over the edges of the shell, so as to See also:cover more or less completely its outer surface. When this covering is complete the shell is contained in a closed sac and is said to be " internal," but the sac is lined by ectoderm and the shell is always morphologically external.

In one or two cases the epithelium of the foot secretes a calcified shell, which is either free as in Argonaula or adherent as in Hipponyx. The ctenidia (fig. r) are the branchial organs of the Mollusca. In the primitive condition there is one on each side in the mantle cavity, towards the posterior end of the body. Each is an outgrowth of the body-wall at the side of the body, and consists of an axis containing two main vessels, an afferent and efferent, and bearing on either side a series of transverse plates whose blood-sinuses communicate with the vessels of the axis. The afferent vessel of the ctenidium receives blood from the vena cava or See also:

principal blood-sinus of the body, the efferent vessel opens into the See also:auricle of its own side. Near the See also:base of the ctenidium is a patch of sensory epithelium innervated from the branchial See also:nerve, forming a sense-organ called the osphradium, whose function is to test the See also:water entering the branchial cavity. The branchial current is maintained by the See also:cilia which cover the surface of the ctenidia, except in Cephalopoda, in which cilia are absent and the current is due to muscular See also:action. Thus in the primitive mollusc the mantle-cavity contains a symmetrical group of structures at the posterior end of the body, and this group of structures is called the pallial complex. It consists of the anus in the See also:middle, a renal organ and renal aperture on each side of this, and a ctenidium outside or anterior to the renal organ, an osphradium being situated at the base of the ctenidium. Internal Anatomy: See also:Digestive See also:Tube.—In primitive Mollusca the mouth and anus are the two extremities of the body, but the anus may be brought to an anterior position by a ventral flexure, complicated in Gastropoda by a lateral torsion. The alimentary tube consists of three regions: firstly, the anterior buccal mass with the See also:oesophagus, of ectodermic origin, and therefore bearing cuticular structures, namely the jaws and radula; secondly, the See also:mid-gut, of endodermic origin and including the See also:stomach and See also:liver; and, thirdly, the See also:hind-gut or intestine. The radula consists of a chitinous See also:band bearing See also:teeth, secreted by a ventral caecum of the pharnyx and moved by an apparatus of See also:cartilage and muscles.

It was present in the ancestral mollusc, occurs in nearly all archaic types, and is only absent in the most specialized forms, in which it has evidently been lost; these forms are certain Neomeniomorpha, all the Lamellibranchia, various degenerate Gastropoda, and the Cirrhoteuthidae among Cephalopods. The teeth are secreted by a small number of cells at the closed end of the caecum, the basal membrane by a trans-See also:

verse row of cells in front of these. The teeth are disposed in trans-verse rows, and in each row they are arranged symmetrically on either side of a central tooth. In Polyplacophora there are eight on each side (8.I.8) ; in Scaphopoda two on each side (2.1.2); in almost all Cephalopoda three on each side (3.I.3) ; in Gastropoda the number varies very much in different subdivisions. Beneath the anterior parts of the radula where it emerges from the caecum are a pair of cartilages, and attached to these a number of special muscles by which the radula is moved backwards and forwards to See also:act as a rasp. The secretion of the radula at the closed end of the caecum is continuous, so that it is constantly growing forward as fast as its exposed anterior portion is worn away by use, just as a See also:finger-See also:nail is pushed forward by See also:constant growth at its posterior end,and is worn away or has to be cut See also:short from time to time at its outer end. Circulation.—The See also:system of blood-vessels is entirely separate from the coelomic cavities. It consists of See also:arteries, See also:veins and sinuses, but ramified capillaries are usually absent except in the integuments of Cephalopods. The arteries and veins have proper endothelial walls; they pass abruptly into the sinuses and in some cases communication is effected by orifices in the walls of the vessels, as for example in the vena cava of Nautilus. The heart is situated in the pericardium on the dorsal side of the intestine and at the posterior end of the animal. The pericardium never contains blood, as is well shown in those forms which have red corpuscles in their blood; these corpuscles are never found in the pericardium. The heart receives blood from the gills and mantle, and pumps it through arteries to the body.

It consists of a median ventricle with muscular walls and a cavity traversed by muscular strands. On either side of the ventricle, in the primitive condition, is a thin-walled auricle, opening into the ventricle by a valved opening. Each auricle forms the terminal enlargement of the efferent vein of the ctenidium of its own side. In Nautilus two pairs of auricles are present, corresponding with the two pairs of ctenidia. In the primitive form a single anterior aorta is given off from the ventricle, the two together representing the dorsal blood-vessel of Chaetopods. In more specialized forms a posterior aorta passes backwards from the ventricle, as in Gastropods and the majority of Lamellibranchs. The ramifications of the arteries convey the blood to all parts of the body, and it finally reaches the venous sinuses, the chief of which are the pedal, the pallial and the median-ventral. The last is between the pericardium and the foot; from it the blood passes through the renal organs to the ctenidia. Some blood, however, enters the auricles directly from the mantle, without passing through the ctenidia. In the majority of Gastropoda one gill and one auricle are lost. The blood is usually a colourless liquid containing amoeboid cells and sometimes other corpuscles called haematids. It may be coloured See also:

blue by haemocyanin, a respiratory See also:compound containing See also:copper.

In a few forms the blood contains haemoglobin, either in See also:

solution or in haematids (red blood-corpuscles). In the Gastropoda the muscular tissue of the buccal mass is coloured red by haemoglobin. See also:Nervous System.—The central nervous system may be described as consisting of a See also:collar surrounding the oesophagus, and two pairs of cords arising from the collar and passing backwards. The two pairs of cords arise from the same point of the collar. The ventral cords are the pedal, the dorso-lateral, the pleural, the former innervating the foot, the latter the mantle. The dorsal See also:half of the collar is the cerebral commissure, the ventral the labial commissure. The pedal cords are connected by commissures, and the pedal and pleural of each side are similarly connected. The pallial cords are united to one another posteriorly, dorsal to the rectum. This is the condition of the nervous system found in Chiton and the other Amphineura, but may not be in all respects the ancestral condition. Generally the system is differentiated into ganglia connected by nerve-cords consisting of nerve-See also:fibres only. At the point of the collar whence the nerve-cords arise are the cerebral ganglia; from these one pair of connectives passes to a pair of pedal ganglia, and another pair of connectives to a pair of pleural ganglia. Pedal and pleural on each side are connected by a pleuro-pedal connective Each pleural ganglion gives off a long nerve which supplies the viscera, and the two unite posteriorly below the intestine.

There are usually three small ganglia on the course of this visceral dommissure, namely, the right and left visceral ganglia and the abdominal. The perioesophageal nerve-See also:

ring of Chaetopoda and See also:Arthropoda is represented, not by the collar first mentioned in the above description, but by the commissures connecting the cerebral and pedal ganglia. The labial commissure supplies only the buccal mass and the oesophagus and stomach. The special sense-organs are a pair of eyes on the head, a pair of otocysts or statocysts, and a pair of osphradia which have already been mentioned. In certain cases See also:accessory eyes are also present, e.g. the pallial eyes of Pecten and other Lamellibranchs, and of Chitons. The otocysts are invaginations of the epithelium of the foot, but are innervated from the cerebral ganglia, and the same innervation has been proved in some cases for the osphradia. See also:Reproduction and Development.—Molluscs are usually of separate sexes, but sexual dimorphism is seldom highly developed. Hermaphroditism is secondary, and occurs in one sub-class of Gastropoda, in some Lamellibranchs, and in one sub-order of Amphineura. In Cephalopods and the majority of Gastropods copulation occurs. As a See also:rule no parental care is exhibited, but See also:incubation of the developing ova within some part of the parental body, or receptacles attached to the See also:parent, occurs in some Lamellibranchs, some Gastropods, and in Argonauta among the Cephalopods. True viviparity, that is the development of the ova within the oviduct, is very rare, occurring only in one See also:case among the Amphineura and in some aquatic and pulmonate Gastropoda. The See also:egg-cell of Mollusca is either free from See also:food-material—a See also:simple protoplasmic corpuscle—or charged with food-material to a greater or less extent.

Those cases which appear to be most typical —i.e. which adhere to a See also:

procedure which was probably common at one time to all then existing Mollusca and has been departed from only in later and special lines of descent—show approximately the following history. By division .of the egg-cell a mulberry-mass of embryonic-cells is formed (morula), which dilates, forming a one-celllayered sac (blastula). By invagination one portion of this See also:sphere becomes tucked into the other—as in the preparation of a See also:woven See also:night-cap for the head. The orifice of invagination (blastopore) narrows, and we now have a two-cell-layered sac—the gastrula. The invaginated layer is the enteric cell-layer or endoderm ; the outer cell-layer is the dermic cell-layer or ectoderm. The cavity communicating with the blastopore and lined by the endoderm is the archenteron. The blastopore, together with the whole embryo, now elongates. The blastopore then closes along the middle portion of its extent, which corresponds with the later developed foot. At the same time the stomodaeum, or oral invagination, forms around the anterior remnant of the blastopore, and the proctodaeum, or anal invagination, forms around the posterior remnant of the blastopore. There are, however, See also:variations in regard to the relation of the blastopore to the mouth and to the anus which are probably modifications of the original See also:process described above. In eggs which contain a larger quantity of food-yolk, the process by which the endoderm is enveloped by the ectoderm is somewhat different. Segmentation in these is very unequal, and results in the formation of small cells called micromeres and large cells called megameres, as in fig.

4. As the micromeres become more numerous they gradually envelop the megameres until the latter are completely enclosed. The gastrula is in these cases said to be formed by epibole. Between ectoderm and endoderm a third intermediate cell-layer A~~t_ ,o . /2 I See also:

Pond-See also:Snail, Limnaeus stagnalis. A, First four cells resulting from the cleavage of the original egg-cell. B, Side-view of the same. C, Diblastula See also:stage showing the two cell-layers and the blastopore. D, E, F, Trochosphere stage, D older than E or F. G, Three-See also:quarter view of a Diblastula, to show the orifice of invagination of the endoderm or blastopore, (bl). H, I, Veliger stage later than D. is formed, which is called the mesoderm, and gives rise to the muscular and connective tissues to the vascular system, and to the excretory and generative organs.

The mesoderm arises for the most part from the endoderm. When the segmentation is unequal one of the megameres gives rise by successive divisions to two primary mesoderm cells called mesomeres; these See also:

divide to form two masses of cells called mesoblastic bands. The coelom is formed as a cavity or cavities in the interior of these cell-masses.

End of Article: MOLLUSCA

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