first group has its peak at 0.5 msec,
whereas the second has a peak at about
1.0 msec. Cell B is an example of a

cell having an action potential with a
short latency, which is about 0.5 msec;

cell C has a latency of approximately
0.9 msec.

All the cells in Fig. 1K could follow
repetitive activation ranging from 200
to 400 stimulations per second and
demonstrated a refractory period in the
range of 3 to 4 msec with relatively
small shift in latency for the second invasion near the refractory period (shift
less than 0.5 msec). Figure 1, D to F,
illustrates the refractory period of cell

potential were smaller than expected
(7, 12).
The orthedromic activation of Purkinje cells after stimulation of the
eighth nerve may be ascribed to at least
three pathways; two originate directly
from the vestibular bipolar cells (Fig.
1A), and the third is the vestibulo-

cerebellar projection through the vestibular nuclei. The two direct pathways end as climbing and mossy terminals in the cerebellar cortex (8). However, since activation of auricular Pur-

kinje cells by climbing fibers can be
recognized by its short latency and

characteristic burst activation (&), the
aforementioned
response
must
be

evoked by mossy terminals. It has not
been determined which of the two sys-

tems of mossy fibers, the direct vestibulo-cerebellar system or the disynaptic system through the vestibular nuclei, is responsible for the activation
in Fig. 1, G and J. Since the latencyis
fairly long, (7.5 and 5.0 msec) however,

these action potentials may be evoked
by disynaptic vestibulo-cerebellar fibers.
On the basis of these data, we con-

clude that ‘a number of Purkinje cells
from the auricular lobe project directly

C which was 3 msec for the antidromic
invasion. If the interval between the

‘two stimuli was increased to 3.2 msec
(Fig.

1F), a smaller action potential

was recorded (see Fig. 1D) which had
a larger initial segment-somadendritic

(IS-SD) separation (6).

The extracellular antidromic action
potentials evoked by stimulation of the
eighth nerve have characteristics very
similar to those already recorded by
Matthews et al. (12) who demonstrated
that, when a pair of stimuli are de-

livered at short intervals to the cerebel-

lar white matter, there is a delay in

the second response of the Purkinje
cell. We frequently observed this phenomenon during our experiments (Fig.
1F). Furthermore, there was, in many
instances, a large reduction of the amplitude of the second action potential,

suggesting that the cell had not completely recovered from the preceding
spike (Fig. 1F).
Figure 1, G to J, illustrates extracellular

and

intracellular

recordings

from other Purkinje ceils after stimulation of the eighth nerve. In Fig.
1G, an activation of the eighth nerve
with a stimulus 1.7 times greater than

the threshold stimulus evoked an antidromic

as

well

as

an

orthodromic

action potential. In Fig. 1, H to J, a

similar cell from the same electrode

tract was impaled intracellularly. As

the stimulus strength

increased from

1.2 to 1.5 times the threshold stimulus,
a subthreshold excitatory postsynaptic
potential was seen (Fig. 11). In Fig. 1J

an orthodromic activation was obtained

with a stimulus strength of 1.8 times

the threshold strength. In Fig. 1, I and
J, the stimulus artifact interfered with
the rising phases of the antidromic action potential. As in previous studies,
intracellular impalement of frog Purkinje cells proved very difficult, for
which reason both the resting potential
(approximately 40 mv) and the action
8 DECEMBER 1967

Fig. 2. Electron micrograph of retrograde changes in a Purkinje cell and degenerating
synaptic terminal on to vestibular receptorcells. (Top) Retrograde changes in a Purkinje
cell of frog auricular lobe 4 days after ipsilateral transection of the stato-acoustic
nerve extracranially (RPC). Note the dark ring formed around the cell nucleus by
the rearrangement of the endoplastic reticulum; note also the normal appearance of

‘the adjacent Purkinje cell (NPC) (x 5500). (Bottom) Degenerating synaptic terminal
3 days after ipsilateral removal of cerebellar cortex in the frog (DSB). Note the darkness of the terminal and the clustering of the mitochondria, RC, vestibular receptor
cell; SC. supporting cell (> 39000).
1329

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