Comp. Biochem. Physiol. Vol. 95B, No. 2, pp. 375 379, 1990 0305-0491/90 $3.00 + 0.00
Printed in Great Britain © 1990 Pergamon Press plc
COMPARATIVE STUDIES ON GLUTAMATE
DECARBOXYLASE AND CHOLINE ACETYLTRANSFERASE
ACTIVITIES IN THE VERTEBRATE VESTIBULE
IVAN L6PEZ and GRACIELA MEZA*
Departamento de Neurociencias, Instituto de Fisiologia Celular, UNAM, Apdo Postal 70-600,
04510 M6xico (Tel: (905) 550-5215)
(Received 17 May 1989)
Abstract--1. Vestibular putative neurotransmitters GABA and acetylcholine synthesizing enzymes were
quantified in four vertebrate species to find a correlation between all-vertebrate vestibular hair cell II
(HCII) and synaptic contacts and appearance of hair cell I (HCI) and related synapses in terrestrial species.
2. Glutamate decarboxylase (GAD) and choline acetyltransferase (CHAT) values were: 3.76; 15.38;
21.68; 27.78 and 9.44; 450; 720; 970 n(pico)mol/mg protein/hr (min) in, respectively, frogs, guinea pigs,
rats and chicks.
3. GAD and ChAT omnipresence may indicate constant GABAergic HCII and its cholinergic efferent
synapses, their raised content, appearance of GABA-containing HCI and related cholinergic boutons in
higher vertebrates.
INTRODUCTION
Scarcely modified in the course of evolution the main
features of the inner ear can be recognized in all
vertebrates (Vinnikov, 1982). It consists of a membra-
nous labyrinth encased in a bony analogue located
generally in the petrous part of the temporal bone or
its analogue. In amphibians, birds and mammals the
membranous labyrinth shows the following charac-
teristics: three well defined semicircular canals, each
with its own sensory surface (cristae); two or three
distinct otolithic organs with separated sensory per-
iphery areas (maculae) and one to three other organs
lacking calciferous masses, the papillae, including the
papilla neglecta and the organ of Corti. The apparent
homologies among these organs in different species
have led anatomists to assess a common name to each
type of organ across species, even though there are
substantial morphological and functional variations.
The membranous labyrinth is generally divided into
a superior and an inferior division. The superior
division, which exhibits fewer evolutionary changes
than the inferior one, includes the semicircular canal,
the utricule and the papilla neglecta, which often
shares a labyrinthine cavity with one of the other
organs. Functionally, the organs of the superior
division tend to be vestibular, i.e. equilibrial, with
some exceptions. The inferior division, which changes
dramatically in both form and function in the course
of evolution, includes the saccule and its appendages:
the lagena, the cochlea and the recesses containing
the basilar and the amphibian papilae (Lewis et al.,
1985).
The labyrinthine wall is lined with a layer of
epithelial cells which are specialized in the end organs
to form the sensory epithelia. Each sensory epi-
thelium in the inner ear of vertebrates contains
two basic cell types (hair cells or sensory cells and
*Author to whom correspondence should be addressed.
supporting cells) as well as the nerve fibres inner-
vating the hair cells (Lewis et al., 1985).
The hair cell is the mechano-electric transducer
element which is vital to all functional properties of
vestibular end organs (Wers/ill, 1956). This type of
sensory cell is found exclusively in vertebrates and
has been described as a secondary cell of epithelial
origin (Lowenstein, 1974). The hair cells are very
similar in the macula of the utricule and of the
sacule and in the cristae. There is, however, a great
difference in the structure of the covering material
(Engstrom and Engstrom, 1981).
Wers~ill (1956) described two different kinds of
sensory elements: type I and type II cells. Type I cells
are flask-shaped with a rounded lower end. These
cells are surrounded to a great extent by a nerve
chalice which continues as an afferent nerve fibre
through the basement membrane. The type II cell is
more cylindrical but it also has a rounded lower end.
The type II cell has contacts with many nerve endings
which form afferent (centripetal) synapses along the
plasma membrane at different levels both below and
above the nucleus. This kind of cell is thought to be
phylogenetically the oldest, i.e. it exists in the sensory
periphery of all vertebrate classes whereas type I
appears in terrestrial vertebrates (Wers/ill, 1956). The
presence of synaptic endings filled with vesicles at the
base of a type II hair cell and outside the calix of the
type I hair cell of higher vertebrates supports the idea
that hair cells are also contacted by terminals of
efferent (centrifugal) origin (Lowenstein, 1974).
There is ample evidence that communication, both
centripetal and centrifugal, is chemical in nature
(Furukawa, 1978; Schessel and Highstein, 1981). The
identity of the neurotransmitters involved is presently
a matter of controversy, although gamma-amino-
butyric acid (GABA) and acetylcholine (Ach) have
been implicated as afferent and efferent neuro-
mediators, respectively (Gisselsson, 1960; Flock
and Lam, 1974; Rossi et al., 1977, 1980; Felix and
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