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Studies on the Vertebrate Head. Part I. Fish. G. It. De Beer, M.A., B.Sc

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Studies on the Vertebrate Head. Part I. Fish. G. It. De Beer, M.A., B.Sc Studies on the Vertebrate Head. Part I. Fish. By G. It. de Beer, M.A., B.Sc, F.L.S., Fellow of Merton College, Demonstrator in Zoology and Comparative Anatomy, Oxford University. With 42 Text-figures. CONTENTS. PAGE 1. INTRODUCTION ......... 288 2. SELACHIANS ......... 289 (a) The Venae Capitis 289 (b) The Arteries 298 (i) Development 298 (ii) Relations of the Arteries ..... 301 3. TELEOSTOMES ......... 304 (a) Amiacalva. 304 (i) The Venae Capitis 304 (ii) The Relations of the Arteries . .309 (iii) Development of the Arteries .... 315 (Ji)Salmo trutta 316 (i) The Venae Capitis 316 (ii) The Arteries 318 (c) Cottus bubalis 319 The Venae Capitis 319 4. DIPNOI 320 Ceratodus forsteri ....... 320 (i) The Venae Capitis 320 (ii) The Arteries 323 5. CYCLOSTOMES ......... 324 Petromyzon ........ 324 (i) The Venae Capitis 324 (ii) The Arteries 327 288 G. E. DB BEER PAGE 6. CONSTANT RELATIONS ........ 328 7. INCONSTANT RELATIONS ....... 329 (a) The Mandibular Vessel 329 (b) The Hyomandibular 333 8. CONCLUSIONS . ... 337 9. SUMMARY 339 10. LIST OF LITERATURE CITED . ...... 339 1. INTRODUCTION. THE structure of the head of the vertebrate might appear to be more suitable as a subject for a text-book than for a paper. There exist innumerable and admirable descriptions of the structure of the head of various forms in books and papers, but it appeared to the writer that detailed comparisons between related forms belonging to one group not only constituted a less threadbare subject for investigation, but also might reveal some points either unknown, unexplained, or not well under- stood. Accordingly the present work aims at a description of few new structures, but at the comparison of the relations which these structures bear to one another in related forms. The present communication deals with the most lowly groups, the Fish and Cyclostomes. The forms investigated were: Scyllium canicula, Squalus acanthias, Pristiurus melanostomus, Heterodontus philippi, Torpedo ocellata, Amia calva, Salmo trutta, Cottus bubalis, Ceratodus forsteri, Petrornyzon planeri, P. fluviatilis. The relations of the eye-muscles and the oculomotor nerve have been dealt with elsewhere (14). Here the structures calling for consideration are the arteries and veins, and the relations which they bear to various nerves and cartilages. The work was done in the Department of Zoology and Comparative Anatomy, University Museum, Oxford. I desire to acknowledge my indebtedness to Professor Goodrich, P.B.S., for his encouragement during the work. VERTEBRATE HEAD 2S9 2. SELACHIANS. (a) The Venae Capitis. The external jugular veins of the adult dog-fish are, as is well known, not the primitive veins of the head region (see Hochstetter, 19 and 20 ; Babl, 26). The primitive vein, or vena capitis medialis as it is termed from its position, is the only cranial vein in ScyIlium, Stage I (Text-fig. 1). In TEXT-FIG, Selected transverse sections through Scylliuni oanicula, Stage I, in the regions of (a) trigeminal, (6) facial, (c) auditor nerve, (d) auditory vesicle, (e) glossopharyngeal, (/) vagus, (g) first postotic myotome. EXPLANATION OF LETTERING OF PIGS. 1-42. a, auricle; ah, abducens; aba, afferent branchial artery; abalt first branchial artery; ac, anterior cardinal; am, anterior cerebral artery ; acr, arteria centralis retinae ; aha, afferent hyoid artery; anteba, anterior efferent branchial artery; apa, afferent pseudobranchial artery; as, av, auditory sac ; on, auditory nerve ; auc, auditory capsule ; ba, branchial arch ; bas, third branchial arch ; baa, basilar artery ; hue, ramus buccalis facialis; ca, cerebral artery; cc, cross commissure between efferent branchial arteries; ch, ceratohyal; cv, connecting vessel between 290 G. R. DB BEER transverse sections of embryos it is seen to lie median to the' trunks of all the dorsal cranial nerves, i. e. trigeminal, facial, glossopharyngeal, and vagus ; and in the region of the auditory hypobranchial artery and ductus cuvieri; eim, connecting vessel between facial vein and vena capitis medialis ; da, dorsal aorta ; do, ductus cuvieri; e, eye; eb, epibranchial cartilage; ebai_h, first to fifth efferent branchial artery; ec, external carotid ; eha, efferent hyoid artery; ehv, hyoid vein; en, endostyle; epa, efferent pseudobranchial artery ; exr, external rectus ; exv, extra vein ; ,/', facial nerve ; fb, forebrain ; fgl, foramen for glossopharyngeal; fhmf, foramen for hyomandibular branch of facial; foe, foramen for oculomotor ; forba, foramen for orbital artery ; fv, facial vein ; cjl, glossopharyngeal nerve; ?«]_<, gill-slits 1 to 4; h, hypophysis ; ha, hypobranchial artery ; Kb, hind-brain ; hm, hyomandibular ; hmc, cut dorsal portion of hyomandibular ; hmf, ramus hyo- mandibularis facialis ; hmnis, hyomandibular muscle ; hv, hypo- physial vein; i, inf undibulum; ic, internal carotid; infr, inferior rectus; indb, inferior oblique; inos, inferior orbital sinus ; inr, internal rectus ; lev, lateral cutaneous vein ; ma, mandibular artery ; max, ramus maxillaris trigeinini; me, Meckel's cartilage ; md, ramus mandibularis trigemini; mdmx, maxillomandibular branch of the trigeminal; mexf, ramus mandibularis externus facialis ; minf, ramus mandibularis in- ternus facialis ; ms, mandibular sinus ; my, myotome; inyd, dorsal part of myotome ; vvyv, ventral part of myotome ; n, noto- chord ; nic, notch for internal carotid ; nrpf, notch for ramus palatinus facialis ; oc, oculomotor nerve ; ocs, occipital sinus ; ol, lateral occipital cartilage ; on, optic nerve ; op, operculum ; ophma, ophthalmica magna artery ; orba, orbital artery ; os, orbital sinus ; pa, pathetic nerve ; pant, pila antotica ; papa, primary afferent pseudobranchial artery ; pasc, processus ascen- dens ; pb, pharyngobranchial cartilage ; pc, parachordal; pun,, posterior cerebral artery ; pev, posterior cardinal vein ; peba, posterior efferent branchial artery ; po, processus oticus of the pterygoquadrate ; poc, polar cartilage ; pohm, processus oticus hyomandibularis ; ppb, processus palatobasalis ; ppq, palato- pterygoquadrate; pr, profundus nerve; ps, pseudobranch; r, rostrum ; re, ramus communicans between the facial and vagus ; rop, ramus ophthalmicus profundus; ros, rami ophthalmioi superiores ; ros VII, ramus ophthalmicus superficial facialis ; rpf, ramus palatinus facialis ; rvel, rudiment of the vena capitis lateralis; «,, 2> s» first, second, third somites; sh, stylohyal; si, sphenolateral cartilage ; sn, sinus venosus ; sp, spiracle ; sip, styloid process ; snob, superior oblique ; siios, superior orbital sinus ; sur, superior rectus ; sv, secondary vein ; te, trabecula cranii; tcom, trabecula communis ; th, thymus ; tp, trabecular plate ; Ir, trigeminal nerve ; v, vagus nerve ; va, ventral aorta ; vahm, vein accompanying the hyomandibular nerve ; zap, vein accompanying the profundus nerve; vea, velar artery; vev, velar vein; vl, lateralis branch of vagus; vn, ventricle; w,, ,JS 3, first ventral nerve-root, second and third ditto ; zs, rudiment of otic process of the hyomandibular. VEETEBRATE HEAD 291 sac it runs between the latter and the wall of the brain. These relations will be obvious from Text-fig. 1, and show that the term vena capitis medialis is accurately descriptive. Text-fig. 2 is a reconstruction of a slightly older embryo (Stage J) from the left side; the vena medialis can be seen passing on the inside of the nerves and of the auditory sac. Just behind the oculomotor nerve the veins of each side com- municate by the hypophysial vein. Between the glosso- TEXT-FIG. 2. Reconstruction of Stage J (Scyllium) from the left side. pharyngeal and vagus there appears the rudiment of a new vein, external to the medialis, which will form part of the vena capitis lateralis of the adult. In Text-fig. 3 (Stage K) the vena capitis lateralis is present external to the facial, but it has not yet joined with the posterior portion between the glossopharyn- geal and the vagus. This stage is shown in section in Text-fig. 4 ; there is only the medialis in the region of the glossopharyngeal. It is important to observe that in the region of the vagus the new vein is at this stage still internal to the nerve. For this reason it is here denoted ' secondary vein ' until it acquires its definitive external position. 292 G. R. DE BEER TEXT-FIG. 3. Reconstruction of Stage K (ScyIlium) from the dorsal side. TEXT-FIG. 4. Selected sections through Scyllium, Stage K, in the regions of (a) trigeminal, (6) facial, (c) auditory nerve, {d) auditory vesicle, (e) glossopharyngeal, •(/) slightly behind (e), (g) vagus, (7i) first postotic myotome. VERTEBRATE HEAD 293 TEXT-FIG. 5. Reconstruction of 21 mm. Heterodontus pliilippi from the left side. TEXT-FIG. 6. Selected sections through Stage 0, Scyllium, in the regions of (a) trigeminal, (6) facial, (c) and (d) auditory capsule, (e) glosso- pharyngeal, (/) vagus. 294 G. R. DE BEER Text-fig. 5 is taken from* a reconstruction of an embryo of Heterodontus in Stage L. The lateralis has joined the medialis just in front of the facial and in front of and behind the glossopharyngeal. By looping round, and the branches joining on the outer side, the first branch of the vagus has been TEXT-FIG. 7. Keconstruction of Scyllium from the dorsal side (the brain is removed), showing the vena capitis lateralis. enclosed by the lateralis. The medialis is still the principal vein. The secondary vein has become lateralis. Anteriorly the medialis forms the orbital sinuses. A more ventrally situated sinus (inferior orbital sinus) lies median to the oculomotor nerve
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