Comp. Biochem. Physiol. Vol. 104B, No. 3, pp. 439-447, 1993 0305-0491/93 $6.00 + 0.00
Printed in Great Britain © 1993 Pergamon Press Ltd
MINI REVIEW
cDNA CLONES FROM FISH OPTIC NERVE
ILANACOHEN and MICHALSCHWARTZ*
Department of Neurobiology, The Weizmann Institute of Science, P.O. Box 26, 76100 Rehovot, Israel
(Fax 972-8-344-131)
(Received 3 August 1992; accepted 25 September 1992)
Abstract--l. The present review describes the results of a cloning that was successfully employed in the
study of optic nerve regeneration in fish.
2. Three intermediate filaments (IFs) expressed by the glial fish optic nerve were cloned.
3. By the use of this approach it was possible to resolve the controversial question of whether glial
fibrillary acidic protein (GFAP) is expressed in the fish optic nerve.
4. Moreover, as a result of the information that emerged from the cloning, it was possible to raise
well-characterized monospecific antibodies and successfully exploit them in order to determine glial cell
maturation, lineage and plasticity, monitor the glial cell response to injury of the fish optic nerve, and
compare it to that of a non-regenerative system.
THE FISH OPTIC NERVE AS A MODEL FOR CENTRAL
NERVOUS SYSTEM (CNS) REGENERATION
The visual system is part of the CNS that in fish can
regenerate spontaneously, unlike mammals (Sperry,
1948, Kiernan, 1979), and thus it serves as a useful
model for the study of nerve injury and regeneration
in the CNS (for reviews, see Graftstein, 1991; Sivron
T. and Schwartz M., submitted). In the fish, all optic
axons from each eye project to the contralateral lobe
of the optic tectum. This provides a readily accessible
pathway in which it is possible to produce lesions and
study the resulting changes in the ganglion cells,
axons and terminal fields. Axonal transport, cell body
changes, influences of non-neuronal cells and changes
in the synaptic target can be easily followed.
Studies carried out in an attempt to uncover the
reason for regeneration failure in the mammalian
CNS have demonstrated that non-neuronal cells play
a key role in determining the ability of a specific
neuron to regenerate. CNS neurons in mammals are
potentially capable of regeneration, but lack a per-
missive and/or supportive environment (Ram6n y
Cajal, 1959; Aguayo and David, 1981; Schwab and
Thoenen, 1985; Kierstead et al., 1989). Another line
of evidence comes from experiments which demon-
strate that rabbit optic nerve axons have the potential
to regenerate if their environment is suitably modified
by the introduction of soluble substances derived
from regenerating fish optic nerves (Schwartz et al.,
1985; Lavie et al., 1990). Accordingly, it is feasible
that fish CNS possesses the appropriate elements and
machinery to render the environment of its injured
*To whom correspondence should be addressed.
axons permissive to and/or supportive of axonal
regrowth.
The non-neuronal environment of CNS neurons in
vertebrates includes astrocytes, oligodendrocytes, res-
ident microglia and blood-borne macrophages (Ra-
m6n y Cajal, 1959; Stensaas, 1977; Murray, 1982).
One or more combinations of these cellular elements
and their response to axonal injury might contribute
to the ability or inability of CNS neurons to regener-
ate their injured axons. A comparison between the
non-neuronal CNS environments in fish and mam-
mals and their responses to axonal injury revealed
similarities in some aspects, but profound differences
in others. Attempts have been made to implicate these
differences in the species-dependent ability of CNS
neurons to regenerate.
Until recently, little was known about the glial cell
lineage in fish or the different subclasses of astrocytes
and oligodendrocytes in the fish optic nerve. The rat
optic nerve glial lineage is well characterized (Raft et
al., 1983), and a comparison between fish and rat
lineages might clarify the species-related differences in
plasticity and ability to support axonal regeneration.
The present review demonstrates the results, obtained
by the cloning approach, in the direction of uncover-
ing the glial cell lineage and plasticity during develop-
ment and after axonal injury in the fish optic nerve
model and comparing it to that of mammals.
IFs AS MARKERS OF GLIAL CELLS
Glial cells are characterized by their IFs. The type
of IF that they express signifies their state of
maturation. In addition, different cell types express
different IFs (Lazarides, 1982). The following six
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