J Physiolo<w (Paris) (1995) 89, 137-145 137
©Elsevier. Paris
Overexpression of choline acetyltransferase reconstitutes
discrete acetylcholine release in some but not all synapse
formation-defective neuroblastoma cells
ZG Zhong a, H Misawa b, S Furuya c, Y Kimura a, M Noda a, S Yokoyama a, H Higashida a*
aDepartmenl q( Biophysics. Kanazawa Unieetwity School ql" Medicine, 13 I Takara-machi, Kamtz, wa 920: hDepartment of Neurology,
Tokyo Metropolitan Institute for Neuroscience, Tokyo 183: CNational hzslitute .[br Physiological Sciences, Okazaki 444, Japan
(Received 29 December 1994: accepted 2 February 1995)
Summary -- Secretion of acetylcholine (ACh) in neuroblastoma cells overexpressingcholine acctyltransferase(CHAT)was exalnined. With
transient transfection of ChAT eDNA, neuroblastoma cells, which have no endogenous ChAT and either adhere to myotubes or not. failed
to form functional synapses, and thus no evidence for release of ACh was detected. Stable neurohlastoma cell lines overexpressing ChAT
accumulated ACh inside the cell. and slowly released ACh to the outside of the cell in a calcium independent fashion. However, after
co-culturing them with rat muscle cells, these transformed cells adhered to myotubes and ACh was secreted in a discrete fashion into the
synaptic cleft efficiently in some neuroblastomacell lines but rather inefficientlyin another cell line. The results show that the latent secretion
machinery of ChAT overexpressing neuroblastoma cells either is competent or possess defect(s) in ACh release.
synapse formation / neuromuscularjunction / vesicular acetylcholine secretion / eDNA
Introduction
Cholinergic neurons which secrete acetylcholine (ACh)
play an important role in muscle contraction (Katz, 1966)
as well as in learning, memory and sleep (Bartus et al,
1982; Hasselmo and Bower, 1993; Sarter and Bruno,
1994). Choline acetyltransferase (CHAT; EC 2.3.1.6) is
the enzyme that is responsible tbr the synthesis of ACh
(Wu and Hersh. 1994). Full-length complementary DNAs
(cDNAs) encoding mammalian ChAT have been cloned
and sequenced (see table I of Wu and Hersh, 1994). Regu-
lation of this gene has been studied (Hersh el al, 1993;
Misawa el d, 1992, 1993, 1994; Quirin-Stricker et aJ.
1994; see also Wu and Hersh, 1994). Furthermore, re-
cently, possible co-ordinate expression of ChAT with ve-
sicular ACh transporter has been reported (Bejanin el a/.
1994; Efickson et al, 1994). ChAT cDNAs have been
expressed in CHO cells (Ishii el a/, 1990; Oda e/al, 1992),
insect cells (Habert e¢ d, 1992), fibroblast cells (Schin-
stine e¢ a/, 1992; Misawa et al, 1994) and neuroblastoma
cells (Ishii et al, 1990; Kimum e¢ a/. 1992; Misawa et
al, 1994; Zhong e/d, 1995). These genetically engineered
cells, specially neuroblastoma cells overexpressing CHAT,
would be useful for the study of cholinergic phenotype
expression, such as ACh release into the synaptic cleft.
To monitor discrete ACh output from cells, skeletal
muscle cells which possess nicotinic ACh receptors are
frequently used as a biological and chemical sensor, since
the time resolution is in the millisecond range (Sakmann,
*Correspondence and reprints
1992). There are two ways to detect ACh released from
presynaptic cells in the in vitro culture condition by elec-
trical recording of nicotinic ACh responses in muscle
cells. One is Nelson's method of co-culturing cells with
fused myotubes to facilitate synaptic contacts (Nelson el
al, [976; Nirenberg et d, 1983b). The other is to approach
the cell closely with an excised outside-out patch of muscle
membrane at the tip of a patch pipette to the surtitce of
secreting cells (Hume et al, 1983; Yong and Poo, 1983).
The former synapse formation experiments in co-culture
may have one disadwmtage in that synaptic contact is
greatly influenced by the factor of extracellular matrix
(Chow, 1990). However, it may have an advantage in that
ACh release can be exanlined under a more physiological
condition. Furthermore, the co-culture method provides to
study developmental changes of cell-cell interaction (Fu-
mya and Fumya, 1983: Fumya et d, 1983).
Here, we examined whether ACh can be secreted
into the synaptic cleft from neuroblastoma cells tran-
siently and stably transfected with CHAT. We demon-
strate two typical results in recording synaptic
responses in co-cultured myotubes. By transient trans-
fection of neuroblastoma cells with CHAT, transformed
cells are unable to produce miniature end-plate poten-
tials (MEPPs) in myotubes, in stably transformed cells
up-regulation of MEPP frequency is efficiently or less
efficiently obse~,ed in a cell line-specific manner, un-
veiling latent structure of secetou machinery utilized
in neuroblastoma cells. Three factors, synthesis and re-
lease of ACh and adherence to myotubes, are used
for dissecting synapse formation processes in neu-
roblastoma cells.