Journal of Neuroscience, Vol 5, 1704-1716, Copyright © 1985 by Society for Neuroscience
Quantitative comparison of the structural features of slow and fast neuromuscular junctions in Manduca
MB Rheuben
The multiterminal slow and fast neuromuscular junctions of the moth Manduca
sexta were compared using scanning, thin-section, and freeze- fracture
techniques to see what structural features might underlie their functional
differences. Slow neuromuscular junctions, here formed on tonic muscle
fibers, produce a facilitating e.j.p. the amplitude of which is 1/5 to 1/3
the size of a fast excitatory junction potential (EJP) and the duration of
which is nearly four times longer. A slow junction consists of a single
terminal branch that is shorter in length than either of the pair of
branches that a fast junction forms close together on the muscle fiber.
Within the junction, slow nerve terminals exhibit longer, more frequent
constrictions and are very varicose compared with fast. Since fast larval
junctions on tonic muscle fibers are also varicose (Schaner and Rheuben,
1985), this is unlikely to represent an intrinsic property of the nerve.
However, calculations of the length constants of the varicose versus
nonvaricose shapes indicate that the effect of passive cable properties on
normal functioning may act to limit the length of the slow terminals more
than that of fast. Even though the varicose shape can be predicted to
prolong the time course of the EJP, calculations show that, at the measured
length, this would not explain the very long EJP that is observed. Within
the neuromuscular junctions, the synapses are characterized on the muscle
membrane by a patch of densely packed particles on the external leaflet and
on the nerve membrane by a single linear active zone. The total number of
synapses per slow junction is about 1/3 that of fast junctions. There is a
weak correlation between average area of the individual postsynaptic
particle patches and cross-sectional area of the muscle fibers that
transcends nerve and muscle fiber types. The average lengths of active
zones from the two types do not differ significantly. However, the number
of particles per active zone in slow junctions is about 55% of the number
in fast active zones. Chemically fixed slow nerve terminals have a greater
density of synaptic vesicles remaining than do fast. If a proportion of the
active zone particles represent structures directly involved in the
probability of transmitter release, such as Ca++ channels, then the latter
two characteristics may jointly reflect differences in capability to
release and mobilize transmitter that would partly explain the different
EJP amplitude and facilitation properties.