Auris Nasus Larynx 28 (2001) 85 – 94

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Review
Functional anatomy of auditory brainstem nuclei: application to
the anatomical basis of brainstem auditory evoked potentials
Bernard Biacabe a,*, Jean Marc Chevallier b, Paul Avan c, Pierre Bonfils a
a
Laboratory of Research on the Physiology of the Hearing System, Formation Associée Claude Bernard and Formation CNRS UPRESA 7060,
Faculty of Medicine Necker Enfants-Malades, Uni6ersity Paris V, Boucicaut Hospital, 78 Rue de la Con6ention, 75015 Paris, France
b
Anatomy Institute of Paris, Uni6ersity Paris V, 45 Rue des Saints Pères, 75007 Paris, France
c
Laboratory of Biophysics, Faculty of Medicine Clermont Ferrand, Uni6ersity of Au6ergne1, BP 38, 63001 Clermont-Ferrand, France
Received 17 March 2000; received in revised form 18 May 2000; accepted 25 May 2000

Abstract

Brainstem auditory evoked potentials (BAEP) are used routinely in clinical practice to evaluate the normality of the lower
auditory system. The objective of this review is to describe the functional anatomy of the structures implicated in BAEP
generation (cochlear nerve and the auditory brainstem nuclei). Indications and results of BAEP in clinical practice are presented
and correlated with auditory structures, which generate each waveform of BAEP. © 2001 Elsevier Science Ireland Ltd. All rights
reserved.

Keywords: Brainstem auditory evoked potentials; Auditory nerve; Cochlear nucleus; Superior olivary complex

1. Introduction anatomy of the lower auditory pathways which, in

Brainstem auditory evoked potentials (BAEP) are
short-latency potentials recorded from the surface of
the head during a brief acoustic stimulation. These
potentials which consist of a series of positives and
negatives waves recorded within 10 ms of the stimulus
onset, are routinely used in clinical practice to evaluate
the normality of the lower auditory system [1]. Useful-
ness of BAEP can be extended by the knowledge of
central auditory structures from which originate the
different waveform components of BAEP.
The aim of this paper is to review the anatomical
structures identified as generators of BAEP, with em-
phasis on clinical applications. As auditory nerve and
auditory nuclei located in the brainstem (cochlear nu-
cleus, superior olivar complex) are the major structures
involved in BAEP generation, we first describe the
* Corresponding author. Tel.: + 33-1-53788103; fax: +33-1-
53788102.
E-mail address: bernard.biacabe@wanadoo.fr (B. Biacabe).
2000, is nothing else than a functional anatomy.
2. Functional anatomy of the auditory nerve
2.1. Cochlear tonotopy
The inner ear translates a physical input, the sound,
in bioelectrical outputs, action potentials in the audi-
tory nerve. The cochlea has a tonotopic organization
(i.e. high and low frequency sounds stimulate respec-
tively the basal and apical cells of the Corti organ)
which allow a first analysis of the frequency and inten-
sity characteristics of the acoustical stimulus. In the
auditory nerve, high frequencies are mainly mediated
by frequency-selective fibers (fibers which discharge for
a characteristic frequency or CF) whereas low frequen-
cies are coded by periodicity of discharge rate of the
auditory fibers [2]. Intensity is either related to the
discharge rate of frequency-selective fibers either or
transmitted by successive recruitment of neural
populations.

0385-8146/01/$ - see front matter © 2001 Elsevier Science Ireland Ltd. All rights reserved.
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86 B. Biacabe et al. / Auris Nasus Larynx 28 (2001) 85–94
2.2. Spiral ganglion and cochlear branch of the
cochleo-6estibular ner6e
Cells of the spiral ganglion are the somata of the first
neurons of the auditory pathways. The spiral ganglion
is located in the center of the modiulus (axis of the
cochlea) and includes 30 000 – 35 000 neurons in
human.
The spiral ganglion have two different neurons types
[3]. The major part is represented by type I spiral
ganglion cells (90 –95% of all the neurons) which are of
large size (12 –20 mm of diameter), bipolar and with a
bulky central nucleus. The cellular somata of these cells
are myelinized. Type I spiral ganglion cells contacts the
basis of the inner hair cells of the Corti’s organ. The
type II cells (5–10% of the spiral ganglion neurons) are
of small size (8–12 mm of diameter), unmyelinized and
contact the basis of the outer hair cells of the Corti’s
organ.
The dendritic extension of type I cells contacts only
one inner hair cell but one inner hair cell can be
contacted by many type I cells [4]. On the other hand,
dendrites of type II spiral ganglion cells contact ten to
20 outer hair cells.
The axons of the spiral ganglion cells project to the
central auditory structures through the cochlear branch
of the cochleo-vestibular nerve (auditory nerve). Fibers
coming from apical cells (low frequencies) and basal
cells (high frequencies) of the cochlea are located re-
spectively in the central and the peripheral parts of the
nerve [5]. The auditory nerve goes through the internal
auditory canal and the inner acoustical meatus and
enters the brainstem.
Fig. 1. Schematic transverse section of the medulla showing the location of the cochlear nucleus. From: Bonfils P, Chevaliar JM (1998). Anatomie
ORL. Medicine-Sciences Flammorian, Paris, p. 323 (with permission).
3. Functional anatomy of the brainstem auditory
structures
The major ascending auditory pathways are located
in the brainstem (cochlear nuclei, superior olivary com-
plex) (Figs. 1 and 2), in the midbrain tectum (lateral
lemniscus and inferior colliculus), in the thalamus (me-
dial geniculate body) and in the auditory cortex. Al-
though the knowledge of the central auditory centers is
still limited, the functional specificities of each structure
and relay begin to be outlined. These structures have in
common a cochleotopic organization, that is, the re-
gions of the different central auditory structures receive
neural inputs from frequency-related region of the
cochlea. As this review is devoted to auditory structures
involved in BAEP generations, we only describe the
auditory pathways located in the brainstem.
3.1. The cochlear nucleus (Fig. 1)
The cochlear nucleus is the first mandatory relay on
the ascending auditory pathway [6,7]. Indeed, it was
never found auditory nerve fibers which projected else-
where than in the cochlear nucleus. The cochlear nu-
cleus is located on the dorsolateral side of the
brainstem. Auditory nerve fibers, as they came in the
brainstem, quickly divide into two branches: one is
anterior or ascending, the second is posterior or de-
scending. A cochleotopy present in the cochlear nu-
cleus: low-frequency fibers project on ventral regions of
the nucleus whereas high frequency fibers project on
dorsal regions.
 B. Biacabe et al. / Auris Nasus Larynx 28 (2001) 85–94
Fig. 2. Schematic transverse section through the brainstem showing the location of the set of nuclei of the superior olivary complex. From: Bonfils
P, Chevaliar JM (1998). Anatomie ORL. Medicine-Sciences Flammorian, Paris, p. 323 (with permission).
The cellular structure of the cochlear nucleus is inho-
mogenous. Three main subdivisions have been de-
scribed, based on cytoarchitectonical characteristics and
auditory nerve projections [6,8 – 14]. The auditory nerve
ascending branch projects on the anteroventral cochlear
nucleus and it descending branch projects on the pos-
teroventral cochlear nucleus and the dorsal cochlear
nucleus. While specific cellular populations are confined
to particular regions of the cochlear nuclear, one cellu-
lar type, the multipolar or granule cells, is more widely
distributed. These cells which have a small cellular
somata (10 mm) with few extensions, are mainly present
in rodent species, whereas they are less abondant in
primates.
3.1.1. The antero6entral cochlear nucleus (AVCN)
The AVCN is the most prominent part of the coch-
lear nucleus. It is subdivided in anterior (AVCNa) and
posterior (AVCNp) regions [6]. The AVCNa and
AVCNp regions are mainly occupied by, respectively,
large spherical cells (20 – 30 mm of diameter) which have
a ‘bushy appearance’ and globular cells with long den-
dritic extensions [8,13]. Both of these cell types which
are intermingled with multipolar cells, are distributed
over the entire dorsoventral extent of the AVCN with a
tonotopic organization along a dorsoventral direction
[10].
Primary terminations of auditory axons on the
AVCN cells (peripheral afferent innervation) [15], have
various morphologic appearances. The most frequent is
87
the bud termination (94% of the endings) [12]. Less
frequent but more specific of the AVCN, is the cali-
ciform termination also called endbulbs of Held [15].
Additionally, the AVCN receives descending termina-
tions (central affering innervation) from the superior
olivar complex (lateral superior olive)[16], the contro-
lateral cochlear nucleus [17] and the ipsilateral cerebel-
lum [18]. Some central afferences might have an
inhibitory effect increasing the frequency selectivity of
the spherical cells.
The AVCN have bilateral extrinsic projections (Fig.
3) to the superior olivary complex (medial nucleus of
the trapezoid body), the nuclei of the lateral lemniscus
and the inferior colliculus [19].
3.1.2. The postero6entral cochlear nucleus (PVCN)
The PVCN is located posteriorly in the cochlear
nucleus, from the division of the auditory fibers to the
dorsal side of the nucleus. Two cellular types are found
in the PVCN [6]: the multipolar cells of Osen, very
similar to those of the AVCN, and the octopus cells
with a large somata (35 mm of diameter) and several
large dendritic extensions [9].
Peripheral innervation comes from the posterior or
descending branch of the auditory nerve. Terminations
of these primary auditory axons are various as in the
AVCN, except the endbulbs of Held. The PVCN re-
ceives central projections from the inferior colliculus
[20], the lateral lemniscus nuclei, the superior olivary
complex [21] and the controlateral cochlear nucleus
88 B. Biacabe et al. / Auris Nasus Larynx 28 (2001) 85–94

[17]. The intermediate acoustical stria gathers together The efferent fibers (Fig. 3) of the DCN project,
the extrinsic projections of the PVCN (Fig. 3), mainly to through the dorsal acoustical stria, to the contralateral
the contralateral nuclei of the lateral lemniscus and the nuclei of the lateral lemniscus and the inferior colliculus
inferior colliculus [22]. [19].

3.1.3. The dorsal cochlear nucleus (DCN)
The DCN covers the PVCN on its dorsal and poste-
rior parts. This nucleus is composed, from the periphery
to the center, of three cellular layers [6]: the molecular
layer, the pyramidal or fusiform cells layer and the
polymorphic cells layer. The fusiform cells have a pyra-
midal somata (15 –25 mm of diameter). Their dendritic
extensions project to the molecular layer as their axons
go, through the deep layer, to the dorsal acoustical stria
[23].
Together with an abondant peripheral innervation
from the auditory nerve, the DCN receives descending
projections from the inferior colliculus [20]: fibers from
the dorsal region of the inferior colliculus central nu-
cleus project bilaterally on the molecular and pyramidal
layers whereas fibers from the ventral region of the
inferior colliculus central nucleus project on the pyrami-
dal and polymorphic layers. These descending projec-
tions which respect the cochlear tonotopy, seem to have
an excitatory effect. Additionally, the DCN receives
projections from the superior olivary complex [17].
3.1.4. Neurotransmission in the cochlear nucleus
The neuromediator of the auditory nerve fibers is an
excitatory amino-acid, probably glutamate or aspartate
[24,25]. Neuromediators from the descending central
projections would be acetylcholine [26], gamma-
aminobutyric acid (GABA), glycine [27] and
noradrenalin.
In summary, the analysis of acoustical information in
the cochlear nucleus is complex. It results of many
interactions between the peripherical auditory system
and the upper auditory nuclei, as evidence by the
numerous descending projections from the inferior col-
liculus and the thalamus.
3.2. The superior oli6ary complex (Fig. 2)
The superior olivary complex (SOC) is a set of nuclei
located in the brainstem, near the trapezoid body, and
between the lateral lemniscus nuclei anteriorly and the
facial nucleus posteriorly. It is limited medially by the
medial lemniscus and laterally by the lateral corti-

Fig. 3. Cochlear nuclear efferences. Major projections are represented by dark lines and accessory efferences by thin lines. AVCNa, anterior part
of the anteroventral cochlear nucleus; DCN, dorsal cochlear nucleus; LSO, lateral superior olive; MSO, medial superior olive; MNTB, medial
nucleus of the trapezoid body; PON, periolivary nuclei; DAS, dorsal acoustical stria; IAS, intermediate acoustical stria; VAS, ventral acoustical
stria; LL, lateral lemniscus; IC, inferior colliculus.
 B. Biacabe et al. / Auris Nasus Larynx 28 (2001) 85–94 89

Fig. 4. Superior olivary complex efferences. Major projections are represented by dark lines and accessory efferences by thin lines. AVCNa,
anterior part of the anteroventral cochlear nucleus; AVCN, posterior part of the anteroventral cochlear nucleus; PVCN, posteroventral cochlear
nucleus; DCN, dorsal cochlear nucleus; LSO, lateral superior olive; MSO, medial superior olive; MNTB, medial nucleus of the trapezoid body;
PON, periolivary nuclei; OCB, olivo-cochlear bundle.

cospinal pathway. The topographic anatomy of the
SOC shows a great variability from a species to an-
other. However, in mammalians, three nuclei are
present: the lateral superior olive (LSO), the medial
superior olive (MSO) and the medial nucleus of the
trapezoid body (MNTB). Surrounding these nuclei,
groups of neurons make up the periolivary nuclei [28].
3.2.1. The medial nucleus of the trapezoid body
(MNTB)
The principal neurons of the MNTB have a large
spherical somata (20 mm of diameter) with an eccentric
nucleus [29]. Other cells types are described as elon-
gated neurons and multipolar cells. The principal neu-
rons receive projections via endbulbs of Held
axosomatic endings, from the globular and spherical
cells of the contralateral AVCN [30]. Additionally,
globular cells give terminations to the dorso-medial
periolivary neurons whereas spherical cells project to
the medial superior olive [29]. The neurotransmitter of
principal neurons of the MNTB seems to be glycine [24]
with an inhibitory effect on the other MNTB cells.
Axons of principal cells project (Fig. 4) on the other
SOC nuclei and especially on the lateral superior olive
with a cochleotopic organization, but never on the
inferior colliculus [31]. The MNTB has a cochleotopic
organization. Cells with high and low characteristic
frequencies are located respectively in the ventro-medial
and dorso-lateral regions of the nucleus [31]. Isofre-
quency cells range from a dorso-medial to a ventro-lat-
eral dimension according with the endbulbs of Held
distribution on principal neurons.
Data on MNTB functions are still limited: it could
inhibit the other nuclei of the SOC during a contralat-
eral stimulation and play a role in spatial location of
sound. Its size which varies from a species to another, is
especially bulky in whale and in bat (location of ultra-
sounds) [32].
3.2.2. Lateral superior oli6e (LSO)
The LSO is located near the trapezoid body, on the
medial edge of the facial nerve descending branch. It
has a ‘S’ shape with three subdivisions: the medial
region is more bulky than the two lateral regions.
Different cells types are present in the LSO but the
fusiform or multipolar cells (20 –25 mm of diameter) are
the largest and most characteristic cellular type of this
nucleus (80% of the cell population). The neurotrans-
mitter of these neurons is glycin. The LSO has a
tonotopic organization with the high and low frequen-
cies located laterally and medially, respectively, in the
nucleus [33].
90 B. Biacabe et al. / Auris Nasus Larynx 28 (2001) 85–94
The LSO receive projections from the ipsilateral and
contralateral AVCN: high and low frequencies region
of the AVCN contact respectively the LSO dorso-lat-
eral and dorso-medial regions [34]. Additionally, the
LSO gets glycinergic inputs from the ipsilateral MNTB
[35,36].
The fusiform cells of the LSO project (Fig. 4) on the
central nucleus of the inferior colliculus [37], the lateral
lemniscus and the other SOC nuclei but never on the
cochlea or the cochlear nuclei [38]. This projection,
which retains the tonotopic organization of the LSO, is
crossed for axons of the LSO medial region and un-
crossed for axons of the LSO lateral region [39]. The
neurotransmitter of these projections to the inferior
colliculus is glycine with inhibitory and excitatory ef-
fects for ipsilateral or contralateral projections,
respectively.
Most of the LSO neurons have a physiological activ-
ity in response to binaural stimulation (type EI: excita-
tory effect for an ipsilateral stimulation and inhibitory
effect for a contralateral stimulation) [33,40]. The
fusiform cells are sensible to interaural differences of
intensity and latency. Additionally, some LSO cells
with a characteristic frequency less than 1 kHz, react
exclusively to an ipsilateral stimulation (type EE) [33].
All the characteristics of the LSO cells suggest that the
LSO is implicated in the spatial location of sounds.
3.2.3. The medial superior oli6e (MSO)
The MSO is located between the LSO and the
MNTB. The main cellular type of this nucleus is repre-
sented by the fusiform cells (8 – 15 mm of diameter)[41]
which are tonotopically organized: high and low fre-
quencies cells are distributed in the ventral and dorsal
regions of the nucleus, respectively.
These cells receive projections from the spherical cells
of the ipsi and contralateral AVCN [42]. Two different
endings are present: the first type corresponds to a
glutamatergic termination with an excitatory effect
(type IA) [43] and the second is GABAergic or glyciner-
gic with an inhibitory effect (type IB) [44]. No other
region of the cochlear nucleus projects on the MSO
[42]. Fusiform cells get additionally inputs from the
principal cells of the MNTB and from the periolivary
nuclei [45].
The fusiform cells of the MSO give projections to the
central nucleus of the inferior colliculus (Fig. 4). This
projection which is essentially ipsilateral, respects the
tonotopic organization of the nucleus: the MSO dorsal
and ventral regions connect the dorso-lateral and the
ventro-medial regions, respectively, of the central nu-
cleus of the inferior colliculus [46].
Spatial location of sound is related to interaural
phase differencies for sound frequencies inferior to 2
kHz and interaural intensity differences for sound fre-
quencies superior to 2 kHz [47]. Most of the MSO cells
are binaural neurons but exhibit preferentially re-
sponses to unilateral stimulation (type EE). These cells
are insensitive to interaural difference of intensity. On
the other hand, low frequency cells are sensitive to
interaural difference of phase [47] which could be re-
lated to a spatial location function of the MSO. How-
ever, morphological and functional differences of the
MSO among species suggest that this nucleus might
have different functions according to the species. In cat,
the MSO could play as a detector of interaural differ-
ences whereas in bat, it could act as a monaural nu-
cleus, sensitive to frequency modulations [48].
3.2.4. Perioli6ary neurons (PON)
The PON surround the three main SOC nuclei. Most
of these neurons are located near the anterior and
ventral regions of the LSO and MSO nuclei and form
clusters called trapezoid body nucleus. Even there are
major topographic differences among species, some cel-
lular populations have similar morphologic, histochem-
ical and anatomical properties [28,49]. From these
neurons, originates the olivo-cochlear efferent system
(Fig. 4).
The main afference of the PON comes from the
octopus cells of the PVCN via the intermediate acousti-
cal stria [13,50] (Fig. 3). Additionally, PON receives
projections from other structures of the SOC, the lat-
eral lemniscus and the inferior colliculus.
3.2.5. Oli6o-cochlear efferent system (OCES)
The olivo-cochlear pathways [51], include two sys-
tems which originate from periolivary neurons of the
SOC: the medial and the lateral olivo-cochlear efferent
systems.
The medial olivo-cochlear system comes from the
medial part of the SOC and projects bilaterally to the
base of outer hair cells, predominantly to the contralat-
eral side [52,53]. Some of these neurons give collateral
branches to the cochlear nucleus [54]. The projection of
the medial olivo-cochlear system is tonotopically orga-
nized and predomines in the frequency cochlear regions
between 2 and 20 kHz in cat [55]. The axons of the
medial olivo-cochlear system are myelinated. The lat-
eral olivo-cochlear system originates from the lateral
part of the SOC and projects on the dendritic exten-
sions of the ipsilateral spiral ganglion cells. Fibers of
this system are unmyelinized.
In summary, functions of the SOC are complex: it is
involved in spatial location of sounds, in the acoustical
reflex and in physiology of the efferent olivo-cochlear
system. The functional properties of the olivo-cochlear
efferent pathway remain, in a large part, unknown. It
could be implicated in protection and dynamics and
protection of the inner ear during sound exposure [41].
 B. Biacabe et al. / Auris Nasus Larynx 28 (2001) 85–94
4. Anatomical basis of brainstem auditory evoked
potentials generators and clinical applications
4.1. BAEP generators
BAEP waveforms include a series of fluctuant far-
field potentials occurring during the first 10 ms follow-
ing a transient acoustical stimulation. They reflect
synchronous activity of auditory cells populations. The
BAEP waveforms appearance is dependant of the
recording electrodes locations (Fig. 5). Each waveform
can be referred to its polarity (P, positivity or N,
negativity) and order of occurrence (1 – 5). In human
BAEP, roman numerals are typically used (I – V). The
BAEP threshold is defined as the lowest intensity of
stimulation, which allows identification of the different
BAEP waveforms. In clinical practice, the 2 and 4 kHz
frequencies are usually tested.
From an historical point of view, auditory structures
involved in BAEP generation have been studied exten-
sively in human and animals. Experimental approaches
included simultaneous recordings of intracranial and
surface potentials [56,57] and effects of brainstem le-
sions following surgery [58 – 60] or injection of toxin
[61 – 63]. In human, correlations between brainstem
structures and BAEP waveforms were studied following
stroke, trauma or degenerative lesions [64,65], or from
peroperative recordings during surgery of the posterior
cranial fossa [66]. The aim of these studies was to
correlate each BAEP waveform to a specific anatomical
structure. This approach allowed to identify generators
of the first BAEP waveforms in human and animals (I,
N1) [66]. In the other hand, anatomical and functionnal
complexity of brainstem auditory structures can not
Fig. 5. (A) BAEP recorded from electrodes located on the vertex (vx)
and on the ipsilateral ear (ie). (B) Recording with electrodes on the
vertex and the contralateral ear (ce).
91
allow to directly connect anatomical structure to later
BAEP waveforms (III –V). Many brainstem auditory
structures are simultaneously actived by an acoustical
stimulus and BAEP waveform appearance depends of
the amount — positive or negative — of all these
activities. Additionally, some auditory structures with
asynchronous activity may not be detected during
BAEP recordings even though these structures are acti-
vated by the acoustical stimulus.
Recently, Melcher and Kiang [67 –69] reported a
‘cellular’ approach of BAEP generators with correlating
BAEP variations with neurotoxin-induced lesions of the
brainstem in cat. These authors studied the contribu-
tion of specific cellular populations to BAEP generation
(Fig. 6) and suggested a functional rather than anatom-
ical organization of brainstem auditory structures.
4.1.1. Generators of P1 and N1 wa6eforms
Structures which generate P1 and N1 are spiral gan-
glion cells of the cochlea [68]. These waveforms are
unchanged after cutting the auditory nerve or aspirat-
ing the cochlear nucleus [59,70]. The spiral ganglion
cells with high CF (above 2 kHz) mainly contribute to
the generation of P1 [69].
4.1.2. Generators of P2 wa6eforms
P2 is generated by the cochlear nucleus cells [62,68].
After surgical lesions on the medial edge of the CN (i.e.
disruption of the CN connections with upper central
auditory structures), amplitude of P2 waveform is unaf-
fected [71]. The globular cells of the posterior part of
the AVCN and of the anterior part of the PVCN are
the main cell population involved in P2 generation. The
CF of these cells is above 2 kHz. Additionally, octopus
cells of the posterior part of the PVCN (PVCNp) and
of the DCN contribute to P2 generation but with a
minor role [69]. It could be explained by the number of
octopus cells which are small as compared with the
number of globular cells [11]. Moreover, surgical cuts
of DCN or PVCNp efferences have no effect on P2
generation [58,59] which is consistent with the overall
minor participation of DCN and PVCNp in BAEP
generations.
4.1.3. Generators of P3 wa6eforms
P3 waveform originates both from cochlear nucleus
and contralateral superior olivar complex cells [68]. In
the CN, spherical cells of the anterior part of the
AVCN generate a part of P3 whereas in the contralat-
eral SOC, principal cells of MNTB contribute to P3
generation [69]. Globular cells of the posterior part of
the AVCN are not cellular generators of P3, however
they indirectly participate to its generation as they drive
MNTB principal cells [30]. Additionally, neurons of the
LSO are also implicated [62]. All these cells have a CF
above 2 kHz.
92 B. Biacabe et al. / Auris Nasus Larynx 28 (2001) 85–94
Fig. 6. Contribution of different brainstem auditory to structures to BAEP generations. See text for details. Adapted from Melcher and Kiang
[59].
4.1.4. Generators of P4 wa6eforms
Ipsi and contralateral cells of the SOC participate in
P4 generation [63,68] with MSO principal cells iden-
tified as P4 generators [69]. Additionally, spherical cells
of the anterior part of the AVCN mainly contribute to
P4 waveform [68], even they are not cellular generators
of this waveform [69], by driving MSO principal cells
[42,72]. By the same way, MNTB modulate the P4
generators [68] as MNTB principal cells have an in-
hibitory effect on MSO principal cells [17].
4.1.5. Generators of P5 wa6eforms
Cellular generators of P5 are located in the lateral
lemniscus [59,70] and/or the inferior colliculus [69,73].
However, cells of the anterior part of the AVCN and
MSO principal cells contribute to P5 generation [68] as
the MSO principal cells project to the lateral lemniscus
and the inferior colliculus [46]. These cell populations
are not cellular generators of P5 as they are involved in
the generation of BAEP waveforms occurring before
P5. CF of these cells are below 10 kHz [69].
In summary, relations between BAEP waveforms and
brainstem auditory structures or cell types, were estab-
lished in animal species in most cases. However, to a
certain extent, these same relations can be transposed
to man when it is postulated that globular cell system
have a minor role in auditory pathways [32,74]. First,
cat and man have both a spherical cell system. Second,
the first three positivities of BAEP in cat and man
(P1 –P3 in cat and I–III man) have similar characteris-
tics [60]. Third, P4 in cat which is predominantly gener-
ating by MSO, have similar generators than V in man
[64,65].
Meltcher and Kiang [69] conclude: ‘BAEP in cats
reflects cellular activity in two parallel pathways, one
originating with globular cells and the other with spher-
ical cells. Since the globular cell pathway is poorly
represented in humans, we suggest that human BAEP is
largely generated by brainstem cells in the spherical cell
pathway’. This conclusion supports the hypothesis that
BAEP recordings in human may allow to derive infor-
mation from specific auditory pathways or specific au-
ditory cell populations, even though neuronal activity
can not be directly assessed.
 B. Biacabe et al. / Auris Nasus Larynx 28 (2001) 85–94
4.2. Clinical applications of BAEP
There are three major clinical uses of BAEP:
(1) Evaluate objectively the lower auditory system:
screening infants at risk for hearing loss allows to begin
earlier an auditory rehabilitation, when BAEP record-
ings show an increase of auditory thresholds or an
absence of waveforms at high intensities of stimulation.
(2) Diagnosis of acoustic neurinomas: the tumor
stretches or compresses the auditory nerve, producing a
delay in the response latency of the later BAEP wave-
forms or a loss of synchronous auditory cells activity.
Typically, the latency of I remains normal when the
latency of V response increases. The BAEP characteris-
tics commonly measured, are interaural latency differ-
ence of V, interwaveform latencies (I – III, I – V, III –V)
and waveform morphology. BAEP are considered ab-
normal when the I to V interpeak latency is longer than
4.4 ms and the interaural latency difference for V is
longer than 0.3 ms. Recently, intraoperative monitoring
of BAEP have been purposed during tumor resections
at the cerebellopontine angle, in order to help auditory
preservation [41].
(3) Diagnosis of lesions of the brain stem: involve-
ment of brainstem auditory structures by demyelinating
diseases produces loss of later BAEP waveforms, due to
loss of synchronous cells activity. For example, in
multiple sclerosis, V can be absent, despite the presence
of normal subjective audiometric thresholds [65].
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