~ Pergamon
0306-4522(94)00371-8
Neuroscience Vol. 64, No. 1, pp. 61 70, 1995
Elsevier Science Ltd
Copyright © 1994 IBRO
Printed in Great Britain. All rights reserved
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INTRACRANIAL TRANSPLANTATION AND SURVIVAL
OF TUBEROMAMMILLARY HISTAMINERGIC NEURONS
H. BERGMAN,*t~ J. I. NAGY§ and A. C. GRANHOLM:~
tDepartment of Cell Biology, University of Link6ping, Link6ping, Sweden
:~Department of Basic Science, University of Colorado School of Dentistry, Denver, CO 80262, U.S.A.
§Department of Physiology, University of Manitoba, Winnipeg, Manitoba, Canada
Abstract--Investigations were undertaken to determine whether fetal histaminergic neurons in the
tuberomammillary nucleus of the posterior hypothalamus survive intracranial transplantation to adult
hosts. Two methods of transplantation were utilized. Grafts were placed either into the delayed cavity
of a fimbria-fornix lesion or directly into the hippocampus using stereotaxic techniques. The tissue was
taken from rat fetuses at embryonic days 16-17 and grafted into adult rats of either the Sprague-Dawley
or the Fischer 344 strain. Routine histology and immunohistochemistry were used to evaluate the grafts.
All transplants to Sprague-Dawley rats showed signs of rejection, while no signs of rejection were seen
in any of the Fischer 344 rats. Transplants placed directly into the delayed fimbria-fornix cavity did not
grow as well or contain as many surviving neurons as the intraparenchymal grafts. The largest number
of surviving histamine-positive neurons was obtained with grafts of posterolateral blocks of hypothalamus
from fetal day 17 placed directly into the CAI region of the rostral hippocampal formation of Fischer
344 hosts. Histamine-immunoreactive cell bodies with neuritic outgrowth were found in all Fischer 344
rats that received hypothalamic grafts. Cell bodies exhibited histamine immunoreactivity evenly through-
out the cytoplasm and had morphological characteristics resembling histaminergic neurons in situ. Axonal
outgrowth extended throughout the grafted hypothalamic tissue, and was sometimes seen in the host
hippocampal tissue as well.
It is concluded that fetal histaminergic neurons survive transplantation to the adult hippocampal
formation, and that this allograft procedure can supplement current strategies to investigate the function
of histaminergic tuberomamillary neurons in the central nervous system.
The existence of magnocellular neurons in the tuberal
region of the hypothalamus has been known for
several decades) ° These neurons in the tubero-
mammillary nucleus (TM) were subsequently found
to be histaminergic, 43 and were described as consist-
ing of five subgroups, located medially, laterally and
ventrally. 2°'26,35'44 The target areas of histaminergic
neurons in the rodent brain have been mapped using
antibodies directed against conjugated histamine 27'37
or the histamine-synthesizing enzyme histidine decar-
boxylase. 12'22'41'42 The ontogenesis of histaminergic
neurons appears to be distinctly different from that of
the catecholaminergic neurons, 21 but closely related
to the development of serotonergic neurons.l'4° Their
neurogenesis in the rat begins at embryonic day (E)
13, peaks on El6 and is completed by El8. 31 Hista-
minergic neurons with well developed dendrites
and axons extending considerable distances from
cell bodies were shown to survive in hypothalamic
explants from neonatal rats for up to nine weeks
in vitro. 32 It has also been demonstrated that
*To whom correspondence should be addressed, at Denver.
Abbreviations: ADA, adenosine deaminase; BSA, bovine
serum albumin; E, embryonic day; GFAP, glial fibrillary
acidic protein; PB, phosphate buffer; PBS, phosphate-
buffered saline; TH, tyrosine hydroxylase; TM, tubero-
mammillary nucleus.
61
explanted histaminergic neurons can innervate co-
explanted hippocampal tissue in vitro. 19 However,
long-term studies of histaminergic plasticity and
axonal growth properties have not been performed to
date.
Biochemical and immunohistochemical data has
demonstrated that TM histaminergic neurons inner-
vate all areas of the hippocampal formation with
moderate densities of histamine-positive axons, and a
large proportion of the histidine decarboxylase was
reported to be concentrated in synaptosomal frac-
tions indicating localizations in axons and terminals
in this brain region. 2,29 Approximately 60% of these
histaminergic projections were suggested to enter the
hippocampal formation via a dorsal pathway consist-
ing of the fimbria-fornix and cingulum, and the
remainder via a ventral route through the amygdaloid
area. 2 The same routes of hippocampal innervation
have been described for other monoaminergic neur-
ons) 8 Since the anatomy and physiology of the
hippocampal formation are well known, and the
afferent and efferent connections can be relatively
easily manipulated by, for example, surgical fimbria-
fornix transection,~l this brain region has often served
as an innervation target for studies of neuronal
plasticity. Bilateral fimbria-fornix transection results
in a partial deafferentation of the hippocampal for-
mation, producing specific spatial memory impair-