The Development of the Head-Muscles in Gallus Domesticus, and the Morphology of the Head-Muscles in the Sauropsida
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HEAD-MUSCLES IN GALLUS AND OTHER SA.UROPS1DA. 511 The Development of the Head-muscles in Gallus domesticus, and the Morphology of the Head-muscles in the Sauropsida. F. H. Edgeworth, IU.IJ., D.Sc, Professor of Medicine, University College, Bristol (from the Anatomical Laboratory). With 39 Text-figures. A FEW papers have been published on the early stages of development of the head-muscles of Gallus and other birds, and on the insertion of the mandibular muscles into Meckel's cartilage and the lower jaw; but as yet there has been no description of the formation of all the muscles in any one bird which might serve as a basis for a discussion of their morpho- logy. The following is a contribution to the subject founded on an investigation of Gallus. I found it was impossible to understand these head-muscles without comparing them with those of Reptiles, but the diffi- culty of doing so was accentuated by the almost complete absence of knowledge of their development; for although Corning has described the early stages in Lacerta, the later ones are not known, and no other Reptiles have been investi- gated. Several friends, however, were good enough to lend me sections or embryos of the later stages of Chelone, Alliga- tor, Sphenodon, Agatna, and Chama^leon, and I was able to procure those of Tropidonotus natris, so that it was possible to follow the growth and changes in the muscles which take place and give each group its special charac- teristics. VOL. 5], PAET 4.—NEW SERIES. 39 512 F. H. EDGEWOBTH. The treatises of Gadow and Hoffman have been taken as guides for the adult anatomy of the head-muscles of the Sauropsida. Some changes in the nomenclature have been made in an attempt to apply identical names to homologous muscles, for examination of their development shows that different names have been given to homologous muscles,1 and the same name to muscles of different origins and morpho- logy.8 A list of the terms employed is given in an appendix. VISCERAL CARTIT-AG.ES. The development of the visceral cartilages in the various groups of the Sauropsida has already been described by TEXT-FIG. 1.—Transverse section through embryo of Gallus, with 11 trunk somites. (For explanation of lettering see p. 555.) Parker, Howes and Swinnerton, and by Broom ; and I have very little to add. An interesting point is, that in early stages of Tropidonotus there is a long pterygoid process pro- jecting forwards from the upper end of the quadrate (text-fig. 37, p. 543). By the time chondrification has taken place this pterygoid process has atrophied; and there is only its proximal 1 E.g. the anterior part of the ventral longitudinal muscles has been called " cerato-mandibularis" (Sphenodon and Lacertilia vera), " genio- hyoid" (Chelonia and Rhiptoglossa), " maxillo-hyoideus" (Ophidia), and " anterior belly of inaxillo-coracoideus " (Crocodilia). a E. g. the name " eerato-hyoideus " has been given to a muscle developed from the first branchial myotome of Sphenodon and Lacertilia vera, to the hyoglossus in Crocodilia and Rhiptoglossa, and to a muscle developed from the posterior mylohyoid in Birds. HBAD-MCJSOLBS IN GALLUS AND OTHER SAUROPSIDA. 513 end left—as a stump, which Parker (who, however, did not see the earlier stage) erroneously homologised with the " pedicle " in the frog. TEXT-FIG. 2.—Coronal section through embryo of Gallus, with 14 trunk somites ; outside the dorsal aorta, on either side, is seen the upper lateral part of the pharynx (cp. text-fig. 3), and outside this the upper part of the dorsal cephalic ccelom, which is con- tinuous behind with the first trunk somite. (For explanation of lettering see p. 555.) In all the Sauropsida, consequently, there is a long ptery- goid process to the quadrate in early stages of development. ftypobf. spf mes TEXT-FIG. 3.—Transverse section through embryo of Gallus, with 18 trunk somites. (For explanation of lettering see p. 555.) In Sphenodon, Lacertilia vera, Rhiptoglossa, and Croco- dilia this pterygoid process has a processus ascendens, or epi- pterygoid, which is developed very early in Sphenodon (vide Howes and Swinnerton), late in the Alligator (text-fig. 22, p. 527). 514 F. E. EDGEWOBTH. There are three median hyobranchial cartilages in Grallus. Parker called them the cartilago entoglossa, basihyal, and tbrmy TBXT-FIG. 4.—Sagittal section through embryo of Gallus, with 2 fully formed and 1 partially formed gill-pouch. (For explana- tion of lettering see p. 555.) urohyal, and stated that the first named was formed by lateral fusion of the lower ends of the ceratohyals. Geoffrey Smith TEXT-FIG. 5.—Sagittal section through embryo of Gallus, with 3 gill-pouches. From the anterior portion of the gut a hollow pouch projects laterally, the wall of which is cut by this section (the Anlage of the premandibular segment), (For explanation of lettering see p. 555.) HEAD-MUSCLES IN GALLTJS AND OTHER SAUROPSIDA. 515 subsequently stated that in a five-day-old chick each cerato- hyal is continuous below with a " branchial blastema." Transverse sections show that whereas the lower ends of premandsegan TEXT-FIG. 6.—Transverse section through an embryo of Galhis, showing fully formed Anlage of pi-emandibular segment. (For explanation of lettering see p. 555.) the first branchial bars meet each other in the middle line, the lower ends of the ceratohyals are separated from one another TEXT-FIG. 7.—Transverse section through an embryo of Gallus, with 3 fully formed gill-pouches, in hyoid and first branchial seg- ments. (For explanation of lettering see p. 555.) by a median procartilaginous structure (text-fig. 12, p. 519), which is continuous with a median structure in front and behind. Chondrification takes place, the middle portion of 516 P. H. EDGEWOB.TH. the ceratohyal disappears,1 and the lower end forms a short cornu to the cart, entoglossa, which is, therefore, a median basihyal or glossohyal with ceratohyal cornua. The middle piece (Copula i, of Gaupp) is a first basibranchial, and the urohyal (Copula ii) a second basibranchial.1 This conclusion is supported by comparison with the condi- tions found in Reptiles. In Sphenodon, Lacertilia vera, Ehiptoglossa, Chelonia, and Crocodilia there is a continuous basihyobranchial cartilage, and in the first four groups there are ceratohyal cornua homologous with those in Gallus. In TEXT-FIG. 8.—Sagittal section through an einhryo of Gallus, with 4 gill-pouches. (For explanation of lettering see p. 555.) Sphenodon these ceratohyal cornua are formed by ceratolij'al bars which are continuous above with the hyomandibulars, in Agama, Chamasleon, and Chelone they are the lower ends of ceratohyal bars which, in embryonic stages, were continuous with the hyomandibulars. In the Crocodilia there are no ceratohyal cornua to the basihyobranchial, and the single 1 The small bit of cartilage which is described by Geoffrey Smith as separating from the lower end of the infrastapedial on the eighth day is probably homologous with that called "stylohyal" by Parker in Uhelidon urbica. HEAD-MUSCLES IN GALLUS AND OTHER SAUROPSIDA. 517 embryo I had was too far advanced in development (chondri- fication was complete) for one to see this stage—of a cerato- hyal continuous from the hyomandibular above to the basi- hyobranchial below. A first branchial bar is present in all groups of the Saurop- sida, a second branchial in Chelonia1 (vide text-fig. 19, p. 525), Sphenodon, and Lacertilia vera, but is absent in Birds, Croco- dilia, Bhiptoglossa, and Ophidia. The main differences, consequently, between Birds and TEXT-FIG. 9.—Sagittal section through an embryo of Gallus, with 4 gill-pouches. It shows (partially) the formation of the ventral longitudinal muscles, which originate from the first (half) royo- torne and the four succeeding ones. Owing to the spiral twist of the embryo no single section shows all the five downgvowths. (For explanation of lettering see p. 555.) Reptiles in the state of the hyobranchial skeleton, are that in the former a second basibranchial is developed2 and the median cartilages become jointed. There are correlated 1 The suggestions of Gaupp in regard to the nature of the cart, entoglossa in Birds and the existence of a second branchial bar in Chelone are thus confirmed. 2 It is absent in some few Birds, e. g. Rhea, Platalea. 518 1<\ H. EDGEWOK.TH. differences in certain muscles of the head which will be described later on. Parker, who did not investigate any embryonic stages, R.SUfl TEXT-FIGS. 10 AND 11.—Sagittal sections through an embryo ot Gallus at the end of the fifth day. 10 is the more external. (For explanation of lettering see p. 555.) described the columella of the adult Ohamfeleon as consisting of stapedial, mediostapedial, extrastapedial, and supra- HEAD-MUSCLES IN GALLUS AND OTHER SAUROPS1DA. 519 stapedial portions, the extra- and supra-stapedial portions lying internal to the quadrate. Study of the development, however, shows that his " extrastapedial" is the persisting upper end of the ceratohyal which lies at first posterior to the quadrate (text-figs. 32 and 33, p. 536) in the hyoid segment. A true extrastapedial is absent. The hyoid bar of Tropidonotus similarly lies at first in the hyoid segment (text-fig. 37, p. 543). The " tuberculuni " (of Parker) attached to it is probably a suprastapedial element; it is developed as an outwards and upwards projecting process of TEXT-FIG. 12.—Transverse section through an embryo of Gallus at the beginning of the sixth day. (For explanation of lettering see p. 555.) the hyoid bar (at a stage later than that shown in text-fig. 37, p. 543), and only subsequently rotates forward. Parker stated that the "tuberculum" became adherent to the stapedial plate, but so far as my observations go it simply disappears.