102 Ii. n. Lindeman:

and somewhat anterior/yo In this medially-situated area, the sensory cells lie scattered amongst the supporting cells, and the crystal layer of the overlying statoconial membrane is extremely thin.

Differences in stnlCture of the sensory hairs are found in the different sensory regions. On the cristae, the hairs in the peripheral parts of the epithelium are longer than those in the central part. The stereocilia in the central areas of the cristae and in the striola of the maculae, are thicker and more club-shaped than the stereocilia at the periphery.

The course and ramifications of the vestibular nerve have been studied, and special attention has been paid to the innervation of the macula sacculi. This macula is innervated by myelinated fibres from both the saccular and Voit's nerve. In all species an overlapping of fibres from the two branches is seen. Furthermore, in the guinea pig bundles of thin myelinated fibres can be followed from Voit's nerve far into the main part of the macula sacculi.

The vestibular epithelia are innervated by myelinated nerve fibres of different calibre. The fibres which run to the central parts of the cristae and to the striola of the maculae, are thicker than those which run to the periphery. In the sensory epithelium a plexus of very fine fibres is seen. These fibres are provided with beads, which probably represent boutons. Each of the thin fibres appears to make contact with a large number of type II cells, with the nerve chalices of type I cells and with thicker intraepithelial fibres. Below the sensory epithelium, unmyelinated fibres are observed, some of which can be followed into the sensory epithelium. Moreover, thin unmyelinated fibres are seen in the walls of thc membranous labyrinth, just outside the sensory epithelia of the cristae and the maoula utriculi.

The statoconial membranes show regional differences in the size of the crystals, thickness of the crystal layer, and structure of the gelatinous substance enveloping the crystals. In the guinea pig, the crystal layer in the striola of the macula utriculi is thinner than that on either side, whereas on the macula sacculi, the crystal layer in the striola is thicker than that on either side. The size of the crystals varies from about II. fl. to about 30 fl. in the guinea pig. The largest crystals are found above the lateral part of the macula utriculi and the antero-inferior part of the macula sacculi. In the striola of both maculae, only small crystals are seen. The gelatinous substance shows a fibrillar appearance, much coarser in the striola than at the periphery.

The present study reveals a highly differentiated structural organization of the vestibular sensory regions. These observations indicate that the different areas are functionally dissimilar, a suggestion which is further supported by the findings of clear regional differences in the sensitivity of the vestibular sensory cells to ototoxic antibiotica.

Acknowledgement. The present investigation was carried out at the Ear, Nose and Throat Department, Sahlgrenska Hospital, University of Gothenburg.

I wish to express my gratitude to the Head of the Department, ProfeBBor Gosta Herberts, for his interest in the study and for providing good working facilities. Professor Hans Eng­strom introduced me to the sensory regions of the inner ear. I am greatly indebted to him for his inspiring interest in the work and for critical advice. Througho-ut the study Dr. Goran Bredberg has offered me great help, and I am most greatful to him for stimulating dis­cllssions and advice.

103

Illy sincere thanks are due to Profes"or Fred Walberg and Professor Alf Brodal at. th~ Anatomical Institute. University of Osla. far their constructive criticism and gencraus h"lp. particularly in the preparation of the manuscript.

I wish to expl'Cll8 my gra.titude to Dr .. John Balln'ltyne for oorrerting th~ En~liRh t('xt and for valu"ble advice. The skilful technical as"istancr and "d,.ire of Res. Enl!in~cr Anton Andersson. is greatfully acknowledged. For stimulating discuRsions I w"nt to thank Dr. B,·rn· hard Kcllerhal •• Dr. Itoy Naco""n. and my faUII". A.~o";ate Profcs"or Hcnrik H. Lind~m"n.

~·urthcrmarc. I ant indcbted to Asoocinte l'rof~"or llano Hikk~rt for &R"i"t"nre in t"king the X.raye; ta Dr. SUSAn Sehanche far translatian of the main part of the manuscript into English; and to Mrs. Nanti Ambarg far expert help in making the .ehematic drawingR .

. Thi. sturly WII.II supported in part b)' the Norwegian Research Counril for S"ience amI the Humaniti"s; in part by the Physiologirnl Psychology Branrh. Office of ]';"",,1 H,·scar .. h. Washington DC. under Contract. N 625584264 and l' 6105267 C OO!IO with n. I-:ngstri;m. This support is gratefully acknowledged.

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Subject Index

Ampulla, membranous 20 Anastomosis - cochleo-vestibular 24, 26 - f"cio-cochlear 25 -- facio-vestibular 25, 20 Antibiotics, ototoxic 39, 99 Aquaeductus vestibuli 17

Basal body 39,57,58,67,68 Basement membrane 3.5, 49 Bouton. (see also Nerve endings) - en passant 81 - terminal 81

Cisterna perilymphatiea 14, 23 Collapse figure 39, 54, 0.5 Cortilyml'h 5:1 Crista ampullaris 21,22,48 - anterior 21,49, 51 - - area of sensory epithelium of 49 - - morphological polarization of 57,64,

66,69 - - number of sensory cells of 49, 51 - central part-s of 48-52, 54, 64, 70, 71. 97 - innervation of sensory epithelium of

26, 29, 73-76, 79 - lateralis (horizontalis) 21, 49, 51 - - area of sensory epithelium of 49 - - morphological polarization of 57,

64,66,69 - - number of sensory cells of 49, 51 - peripheral parts of 4M-52, 54, 64, 70,

71,97 - posterior 21 - - area of sensory epithelium of 49 - - morphological polarization of 57,

64,60,69 - - number of sensory cells of 49, 51 - regional differcnces in structure of

48-52, 54, 97 Crus commune 17,20 Crystals, see Statocouia Cupula 21, 65, 90--92, 94, 95

Directional sellsivity (see also Functional polarization) 57, 69

Dividing line of morphological polarization 43,58,61, 62, 78, 93

Du('t -- eoch lear I6 - definition of 20 - endolymphatic 17 -- rClIniC"ns 16 - ",,,'elliar Jfi, 18 - ~midrcular 20 - utricular 16,18

Endolymph 13, (;3 El'itl,,'lium, ,enRory :I.'i

Hanglion - W'nkulatc 25 -- "piral 24 -- vCfitihular 24, 25 - - infcrior 24, 29, 30 - --- subdivisions of 30 - ,- ,<mpcrior 24 j 30 Oclat inOUR ffllh,~tancc, sec Htatoconial

mcml;ranc:>, gelatinotiR :mbstanc('

Jlair!i, see Sensory hairn

Innervation (see also Nerve and Nerve fjbre~) - cristae ampulla.,," 26-29, 72, 73-77.

711 - macula sacculi 2(\-29,78, 79, HO - macllla utriculi 21i. 77, 78. 79 - type I cells 35, 72, 73, 74, 77, 78, 79 - type II cells 36, 72. 74, 80 Ist.hmll:9 gangIionaris 25, 30

KinocilillID, see Hensory hairs

IJimitinf! membrane 2:J

Macula -- npclceta 25 - Haceuli - - area of 45, 48 - - density of sensory ceJls of 45, 48 - - form of 14 - - innervation of 26--29, 7g, i9 - -- lIlorphological polarization of 58,

02.70 - - number of sensory cells of 45, 4R,

51,53

II:!

ct Index

Nerve - cochleo-se.ocule.r 24, 29 - crossing fibres in the me.cule. sacculi

27,29 - fe.cial 26 - intermedie.te 25 - saccule.r 116 - ste.toacoustic 24 - superior utricular 25 - vestibular 24 - Voit'. nerve 24,26,29,47 Nerve endings - nerve chalice 35, 72, 77, 79 - richly vesiculated 25, 36, 72, 76, 81 - sparsely vesiculated 36, 72, SO Nerve fibres - afferent 72, 78 - autonomic 73, 78, 82 - efferent 29, 72, 78, 81 - intraepithelial plexus of 72, 74-76,

78,81.82 - myeliusted - - density of 73,77,78,79 - -- diameters of 30, 72, 79 - - distribution of 72-74,77-80 - - numbers of 30, 56, 57 - unmyelinated 73,74,82 Nomencla.ture - canal 20 - duct 20 - ste.toconial membrane 92 - utricule.r duct 16 N uC\ei, vestibular 30

Oort, see Anastomosis, cochlea-vestibular Organ of Corti 11, 39, 55, 67, 78 Orientation, _ Morphological polarization Osmiophilic granules 36, 43, 47, 49, 54, 97 Otolith membrane, see Statoconial

membrane

Pars externa and interna, see Macula Perilymphatic space 13, 20, 23, 53 Pigment 20, 21, 23 Planum semilunatum 22 Plexus, intraepithelial 72, 74-76, 78,

SO,81 Polarization, _ Morphological polarization

Quantitative studies (_ Criste. ampullari., Macula, and lIIethods)

Reoeptors, see Sensory cells Recessus - e\lipticus 19 - "phericlis 14, 25

Subject Index 113

Recessus - utriculi 19,20 - between tbe non ampullary end of the

late .... 1 semicircular duct and tbe crus commune 17,20

Regional differences in structure of - cristae ampullares (see also Crista

ampullaris) 43-52, 54, 64, 73, 79, 97 - maculae (see also Macula) 39--48,

52-54,77,78,97 - - functional aspect of 69, 70, 98 Regional differences in sensitivity to ototoxic

antibiotics 99, 100 Reticular memb .... ne 31

Sac, endolympbatic 18 Saccule 14 Secretory epitbelium 20, 23, 78, 82 Semicircular ducts, see Duct, semicircular Sensory cells (see also Crista ampullaria

and Macula) - intermediate type 36, 72 - nuclei of 35, 40, 53 - type I cells 31, 35, 39 - - distribution of 45, 46, 51, 52 - - innervation of 35, 72-74, 77, 79, 80 - - ultrastructure of 35 - type II cells 31, 35, 39 - - distribution of 46,51,52 - - innervation of 36, 72, 74, 80 - - ult"" ructure of 35 Sensory hairs (see also lIIorphological

polarization) - a ..... ngement of 58 - kinocilium 32, 57-59, 64 - regional differences in structure of

61, 64, 70, 71, 97

Sensory hairs - stereocilia 32, 57, 58, 64 - ultraatructure of 58 Septum cruciatum 21 Spaces, intraepithelial 43,47, 53,78,97 Staining, see Methods Statoconia 83, 84, 88, 89, 92, 98 Sts toconial membrane 14, 20, 83, 84, 86,

92,97,98 - gelatinous substance of 87, 89, 1W-96,

97 - nomenclature of 92 - regional differences in structure of

84,86--89,93,94,98 - - functional aspects of 98 Stereocilia, see Sensory hairs Striola, see also Macula 32, 33, 39, 40, 41,

52, 53, 77, 78, 79, 97, 98, 99 Subepithelial tissue 10, 25, 73 Supporting cells 31, 3~, 49 Surface preparations, see Methods, micro­

disaection of the veetibular sensory regione

Traneitional epithelium 22 Type I cell, see Sensory cells Type II cell, see Sensory cells

Utricle 19 Utriculo,aaccular duct, see Duct, utricular

Valve, utriculo-endolymphatic 19 Vestibule 14, 64

Zone b 53,93