The Axatomy of the Autonomic Nervous System in the Dog1
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THE AXATOMY OF THE AUTONOMIC NERVOUS SYSTEM IN THE DOG1 NICHOLAS JAMES AIIZERES Ucpartnitiit of Aiintoiiry, Cnzvemtty of Xtclizgaii Scliool of ;2/cdicmc. Ann Arbor, Mtclatgan ELEVEN FIGURES INTRODUCTTOX I<iiowledgc clerivccl from cmprimeiits on tlie clog has not infrequently been applied to man without considering dif- fcmnccs in aiiatoniy. This is c~spcciallytrue of the autonomic nervous systeni, oiily parts of which have b:mi described in the adult clog. The cardiac ncli'vcs \\'ere described by Sc1iuran.- Iew ( '27) ant1 Soniclez ( '39), the gcw~alplan of the abrloniirial ancl sacral regions by Trumble ( '34), tlic lunibosacral trunk by Alehler, Fisclier and Alexander ( '52), the vagi lsy Hilsa- beck aid Hill ( '50) and BIcC'rca and D'hrcy ( '%), ant1 the urogenital plexuses by Schal~adasch( '26) aiicl Ncdowar ( '2.3. The present study was undertaken to fill gaps in previous clescriptions ancl to present an over-all view of the autonomic iierrous system in the (log, esclutliiig the cephalic region. In the pi-c~seiitiiivcstigation the XI< terminology has been used iiistpad of the usual RNA familiar in human anatomy loccause tlic study was based on the clog. Sincc a dog is a pronograde niamnial the terms cranial aiid caudal seeni nior(t appropriat r tliaii the ESA terms superior and inferior. Tiiis paper is n condensation of a clissei t:ition snbniittctl in paytial fulfillmc.~it of tlrc rcqiiireiiieuts for the drgrce of Doctor of Pliilosopliy in the University of Rlicliigaii. I wish to tlimik Dr. It. T. Wootlhuine for Ills interest and aclriec. For the supply of nlatciial tlie author vihlies to e\-pr('\s his appreciatioir to mcmbc~is of the 1)ep:irtiiiciits of P1iTsiolog.y :iiiil PIiar~u:~cologrof tlic Viiirersity of hliclrigan. 2 Present address: Dcpartniciit of Airntoiny, 1V:1pe Ultivcrsity School of JIcdieine, Detroit, Michigan. 283 286 NICHOLAS JAMES MIZERES MATERIAL AND METILOUS Ten adult clogs, 6 males and 4 fenialm, were injected with ernhalmiiig fluid sooii after death. In 4 speciniens rctl lead was added to facilitate dissectioii of small arteries. The cardiac region was dissected with the aid of a binocular dis- secting microscope. 1)ESCRIPTION The cerviccrl region Soiiidez’ (’31) study of this region ill the puppy is prob- ably the most dcltailecl work iii tlie clog. Both Xonidez (’31) and Finkelstein (1880) described the origin of the clepressor nerve in this area but I failed to find a iiervc which \vould fit their descriptions. It may possibly be very closely iii- corporatcd in the sheath of the vagus iierve near the nodose ganglion. 1. The crccszinl cerrictil gcrnglio.12. (fi%q. 1). The cranial cervical ganglion is it large fusiform niass lying adjacent to tlie origin of tlic iiiteriial carotid artc1.y antl tlccp to the lo~vcrmargin of the auditory 1)ulla. It is crossed ventrally by tlie nerve to the carotid sinus and the pharyngeal ramus of the vagus nerve. Its caudal portion lies dorsal to the auricular and occipital arteries antl the hifurcatioii of the coriirnon carotid artery. The riotlose gaiiglioii lies cauclo- lateral to it. I was able to o1)serve inconstant connections to the 9th’ 11th’ and 12th cranial nerves. A coniiectioii to the vagus nerve, just above the nodose ganglion, was ob- served in all 10 specimens. The internal carotid nerve (fig. 1) arises from the cranial pole of the cranial cervical ganglion and lies in close relation to the internal carotid artery as it enters its canal. Tlie artery is convoluted while in the canal and filaments from tlie iii- ternal carotid iierve were observed eiiteriiig the adventitia of the artery. A large nerve (fig. 1) emerges from the caudal pole of the cranial cervical ganglion and forms a clensp plexus n.liich follows the hraiiclics of the external carotid artery. X short thick branch was also observed joiniiig the pliaryngeal raiiiu:, of the vagus ncrvc. Xonidez ( ’31) regarded this liraiicli a:, eiitclriiig the “infei.ior pharyngeal ranius ” of the vagus iiervc’. In the spccirriciis I exarriiiiccl tlic superior and inferior phar- yiigeal ranii \wrc givcw off after this spnipathc~ticconnection entered a “conir1ioii” pharyiigcd ranius. h thin filament arose frorii the cciiti-a1 poi*tioiiof tlic cranial cervical gaiiglioii, coursed hehiiitl tho iiitcrnal carotid artery and nodose gaiiglioii arid eiitt.r(ld tlie first cervical iierv(i (fig. 1). In tlirccl specirrims tliis coniniunicati~igranius mit additiorial filici~to tlie sccoiid aiitl thiid ccrvical nerves ant1 to tlie clcsc(w1iiig cervical ~ier*ve,wlieii piweiit. In t1irc.e of the 10 cases this ramus was absent. C’audally tlic syrnpathctic trunk joiiis the vagus iic’rvc just bclow the nodose gaiiglioii (fig. 1). 111 4 speciniciiis the craiii:il cc>rvical gaiiglioii was obscrvetl scparatccl from the iiotloh~ ganglion bilatci*ally hut in 6 speciniciis t1it.y irere coiiiiectcti lvith short stiaaiicls iaangiiig froin multiple filarricliits to alnioht cornpletcl fusion. A thick mass of fiheix arising from tlic. caudal pole of the ci*aiiial ccrrical ganglion aiid lying 011 thcl hi furcation of thc coniiiioii carotid artery gives rise to iiuiii(~~*ousshort and loiig cloi.sal filanieiits. The short filaments ramify on the wall of the cai.otid sinus or hull) and coirirnoii carotid artery. Tlic long filaments coniprisc two sets. Oiic group courses oii tliv tlorsal wall of the common caieotitl artc1i.p aiid imnifies 011 all pal-ts of tlic niorc caudal portion of thcl ai*tc>ry.The sccoii(I gi-oul) cornpiks what Xoriidm ( ’31 ) calls tlic thpi*oitl iwi*v~ (fig. 1). Fusion of thew filarnciits is variable and in oiily 4 of my specinleiis \\’as a siiiglc iici~eol)serve(l. The thproitl 11e1~vcdescc~iids to\varc\ tlic thyi.oid artci-y as a plesiis (fig. 1 ) \vhich ~*eccivcsan aiiastorriotic filanicnt fi.orn the csteriial laryngeal nerve arid follo\vs tlic artcry tliiwtly into the tliy- roid gland. According to Xonidez ( ’31) the thyroid glaii(1 of the dog is innci~atcdby the cranial crlrrical ganglion ant1 tlic recurrelit laryngeal iierw. I was ahlo in all caws to fintl rccurrciit filwniciiis entcriiig the glaiid. 111 4 CRS;CS soiiic of the tcrniiiial himichrs of tlic recurrent larpiigc~al iicrvo niiiiglcvl with filaitieiits of tlic thyroid plexus. 2. ?’lie r.ccgos?/Inl,trflrefic. frzciik. In the cowse of dissec- tioii, it I\ as o1,servcd that incision of tlic coiiiicctivc tissue slicatli sui.iouiicliiig lmth vagus aiid symp:ithctic t ruiiks al- lo1vcltl tlioir scpai*atioii. By this “ slicath tlissection” it was INT. CAROTID N INT MAX A 8PLX. MARGIN OF AUDIT BULLA EXT CAROTIDA XI1 N PHARY NGE A RAMUS OF X NGEAL PLX. NT LARYNGEAL N. THYROID N. EXT LARYNGEAL N. VAGOSYMPATHETIC TK C CAROTIDA THYROID PLX. fon11tl that both trixiiks actually separate a few ceiitimcters cc~1~lialaclto the caudal crrvical ganglioii (fig. 2 E). hi the cixiiial portioii of the iiccli tlic vagus iiewe and sympathetic tlaiil; are coiiiplctcly fuscd. To iiiakc certain this scpara t'1011 aiitl fusioii \\-as iiot a11 ai-tifact of dissectioii, wrial scictioiis 290 SICHOLAS JAMES NIZERES of 6 cervical trunks were made and stained with liematosplin and eosin. It is readily seen that in the middle cervical region a thin comnioii perincurial septum (fig. 3 b) still sepai*ates tlic two trunks (fig. 3 a). In the upper cervical region this coni- nion perincurial septum disappears so that both vagal ailti synipatlietic fibers become interrniiiglecl. The vagosynipa- tlietic trunk sends bi.anclies to the trachea, esophagus, and common carotid artery. The thorcrcic region By incising tlic cpincurial sheath, the ~xteiit,location, ant1 variations of the raga1 and sympathetic branches forniing a cardiac nerve can be ascci*tained with coniparativc eascl. Tliih method of dissection has some application in nerve stirnula- tion. If, in the living animal, tlie caudal cervical ganglioxi can be separated carefully from the vagus nerve with a needle or scalpel, it might be stimulated so that its specific effect on the heart could he determined. The same could be done with the vagus nerve and its branches. The following tle- scription is liasccl on dissection of tlic sheath under a dissect- ing microscope. The figuiw are composite di~awingsant1 thc nerves represented arc those of most frequent occurrence. Because of differences in numher, course, and origin of cardiac nerves in dog and man, it is not possible to LISC iden- tical terminology. Cepl~aladto the caudal ccrvical gaiiglion there is no cardiac nerve in the dog, as was Formerly main- tained by Keng (1893), but later refuted by Sonidez ( ’39) and others. Thus, no superior (cranial) cautiiosymi)wtlicitic nerve can be itlentificd in this foi-m. Soniclez ncvcrthclcss named a superior cardiosympatlietic nerve, sincc ‘‘in sonic instances it (superior cardiosympathetic nerve) map also carry bundles of fibers from the middle cervical ganglion to the deep cardiac plexus or to the anterior wall of tlie atyia.” Although such an ai*i-angerneIitwas founcl in tlic majoiitp of specimens in the present study, it does not secin a valid basis for the namc ascrilwcl to it. So superficial cardiac plexus, comparal)lti to that in ~riaii, could br fomicl iii the clog. Tlicre is a plcsns at the tracheal bifurcation that iiiay be corripai.ct1 to the tlccp cardiac plcsus, lmt, as will be tlcscrihed, its components ai~iiot the same. Schui~awle~\-( ’2i) i~cgarcleclthis plexus as tlic pretracheal plcsus. He classifictl 6 cpicardial plexuses as visualized by the riiethylene blue nietliotl. Actually, only three of these can lw obsc>rvedwith tlie naked eye. These are his plexuses I ailti I1 (right arid left coronary), and IV, wliicli lies in the fold for the ligament of the left precaval vein in the dorsal wall of the left atriurii aiitl extends over the dorsal ventricular wall.