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Effects of a corneal anesthetic on the extinction of the classically conditioned response in the rabbit

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Effects of a corneal anesthetic on the extinction of the classically conditioned response in the rabbit

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Content EFECTS OF A CORNEAL ANESTHETIC ON THE EXTINCTION OF THE
CLASSICALLY CONDITIONED RESPONSE IN THE RABBIT
Copyright 2003
by
Karla Robleto
A Thesis Presented to the
FACULTY OF THE GRADUATE SCHOOL
UNIVERSITY OF SOUTHERN CALIFORNIA
In Partial Fulfillment o f the
Requirements for the Degree
MASTER OF ARTS
(PSYCHOLOGY)
December 2003
Karla Robleto Student
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UMI Number: 1420396
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UNIVERSITY OF SOUTHERN CALIFORNIA
THE GRADUATE SCHOOL
UNIVERSITY PARK
LOS ANGELES, CALIFORNIA 90089-1695
V
This thesis,^written by
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under the direction o f h M thesis committee, and
approved by all its members, has been presented to and
accepted by the Director o f Graduate and Professional
Programs, in partial fulfillment o f the requirements fo r the
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DEDICATION
To my parents for their never ending support, patience and unconditional love.
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TABLE OF CONTENTS
DEDICATION ii
LIST OF FIGURES iv
ABSTRACT V
INTRODUCTION 1
METHOD 6
Subjects 6
Behavioral Training Procedures 6
Periorbital Stimulation Study 8
RESULTS 8
DISCUSSION 13
REFERENCES 17
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LIST OF FIGURES
FIGURE 1: Mean Percentage Conditioned Responses (CRs) During
each Treatment Condition
FIGURE 2 % Mean Percentage Conditioned Responses (CRs) During
Each day o f training
FIGURE 3: Mean Percentage Conditioned Response Amplitudes of
Each Extinction Session
FIGURE 4: Mean Amplitude of Unconditioned Responses (URs) to
Stimulation Thresholds
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ABSTRACT
Animals were presented with .seven daily sessions o f tone alone training. Before
the beginning o f each of the first four extinction sessions an artificial tear solution or
tetracaine hydrochloride (0.5%) was administered to the cornea of animals in the control
group (n=6) and experimental group (n=7) respectively. The percentage conditioned
responses, as determined by a repeated measures ANOVA, showed no difference in the
rate of responses between both the control and tetracaine group. However, the amplitude
o f the conditioned response was significantly reduced in the tetracaine group relative to
the control group early in extinction. The observed effects seem to suggest that while
motor output has been found to play an important role in extinction, sensory feedback
might not be necessary for this form o f learning.
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INTRODUCTION
Classical conditioning of the eyeblink nictitating .membrane (NM) response has
been widely used as a paradigm to investigate the neural basis o f learning and memory.
It is well established that the cerebellum and Its associated brainstem regions are
necessary for the establishment, maintenance and expression in this form o f associative
teaming, particularly in the standard delay procedure (McCormick & Thompson, 1984;
Thompson & Krupa, 1994; Chen, Bao & Locteard, 1996). In brief, lesions of the
hippocampus have teen found to not impair the acquisition o f the conditioned response
(CR) in simple delay eyeblink conditioning, where both the CS and US coterminate
(Kim, Clark & Thompson, 1995). In addition, lesion studies o f the cerebellum have
been found to abolish the ipsilateral CR while leaving the performance of the
unconditioned response (UR) unaffected (McCormick, Lavond, Clark, Kettner, Rising
& Thompson, 1981; McCormick, Clark, Lavond & Thompson, 1982). Also, lesions of
the cerebellar nuclei, specifically the interpositus (IP), completely prevent eyeblink
conditioning (Clark, McCormick, Lavond & Thompson, 1984); an effect that can also
be produced by very localized kainic acid lesions in this cerebellar region (Lavond,
Hembree & Thompson, 1985). Together, these results reveal the important role of the
cerebellum in eyeblink conditioning and suggest this brain structure to be the locus o f
memory for this type o f learning. Much of the understanding of classical conditioning,
however, has been gained in aspects related to acquisition o f the response. Extinction,
on the other hand, remains a relatively obscure phenomenon not only behaviorally but
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also at a neuronal level. Accordingly, what are the underlying processes that contribute
to behavioral extinction continues to be an important question in the study o f eyeblink
conditioning.
Since Pavlov first described the phenomenon, classical conditioning has
proven to bean, extremely useful model in the study of learning and memory. As one
example o f classical conditioning, eyeblink conditioning is arguably the best-understood
mechanism o f learning in vertebrates. Decades o f theoretical models and experimental
evidence by Thompson and colleagues have resulted in the detailed mapping o f the
neural circuits involved in conditioning o f the eyeblink response. Data obtained from
such studies, clearly show the essential plasticity for eyeblink conditioning to be located
within the cerebellum. Krupa and Thompson (1995), for instance, demonstrated that
injections of tetxodoxie (TTX) into the bracMum conjiBictivum, which Mock cerebellar
outflow, allowed for normal teaming even though no conditioned responses (CR’s) were
present during inactivation. Also, lesions o f the red nucleus, a major target for IP axons,
abolish expression of the CR yet do not interfere with learning o f eyeblink responses
leaving the learning related neuronal activity observed in the IP unaffected (Clark &
Lavond, 1993). Also physiological studies o f learning related-unit activity show that
inactivation of the IP removes the ‘signature1 ’ o f the engram everywhere else in the brain
(Clark, Gobi & Lavond, 1997; Clark & Lavond, 1996; Clark, Zhang & Lavond, 1997).
Further support for the involvement o f the cerebellum in eyeblink conditioning
comes from experimental observations that show that stimulation of two of its major
afferent fibers, specifically, mossy fibers and climbing fibers, can induce associative
2
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learning and can, in fact, substitute a peripheral CS and US respectively (Maiik,
Steinnaetz, Thompson, 1986; Steinmetz, Lavond, Thompson, 1989). Accordingly,
recording, lesion, stimulation and anatomical tracing studies have resulted in the
development o f a testable model which identifies in detail the pathways, nuclei and
possible sites o f plasticity involved in eyeblink conditioning (Thompson, 1986,1990).
According to the model, the CS is conveyed from the periphery via mossy fibers
originating from the pontine nuclei which send projections to afferent targets in the
cerebellum. in turn, information about the US is relayed thorough somatosensory nuclei
which sends projections to the inferior olive (IO) and is ultimately carried via climbing
fibers emanating from the 10. As it would be expected from a brain structure involved
in associative learning, information about the CS and the US converge in the
cerebellum, specifically the cortex and the IP, suggesting that both structures are
capable of supporting CS-US associations. In more detail, at the level of the IP, mossy
fibers coming from the pons and climbing fibers emanating from the 10 synapse with
the nuclei, relaying sensory and somatosensory information to this area. In the cerebellar
cortex, mossy fibers synapse with granule cells from which parallel fibers form synaptic
connections with Purlrinje neurons. The cortex also receives information from climbing
fibers coining from the 10 which wrap around Purkinje neurons and form a strong
synaptic input with them. Such evidence demonstrating convergence o f CS and US
within the cerebellum, implicates both the IP and cerebellar cortex as possible sites of
plasticity underlying eyeblink conditioning.
3
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Extinction can be defined as a reduction and/or non-performance of a
conditioned response (CR) following repeated CS alone presentations. In the NM
preparation, when the CR has been well conditioned to a tone CS with a corneal airpuff
unconditioned stimulus (US), subsequent tone alone presentations result in decrease and
eventual absence o f the CR. One of the many phenomena that challenge the idea that
excitatory associations are lost due to extinction is probably one o f the oldest described
since the inception o f classical conditioning. Spontaneous recovery, an effect seen after
extinction, was recognized by Pavlov since Ms early studies o f inhibitory learning. He
observed that CR’s would return after extinction if enough time is allowed to elapse
until the CS is tested again. The feet that the conditioned response can ‘recover’
between extinction trials seems to argue against the idea that the excitatory association
is lost or ‘fades away’ due to extinction training. TMs effect has teen tested and
consistently observed by researchers in a variety of conditioning protocols (Bouton,
1993; Schreurs, 1993; Napier, Macrae & Kehoe, 1992). There are several other
behavioral effects that indicate that extinction does not involve the destruction of the
original memory. One o f the most ‘straightforward’ methods that can be employed to
test the maintenance of the excitatory memory is that o f reacquisition where it is
observed that acquisition is fester when the original association is presented again after
extinction compared to normal acquisition rates seen in naive animals (Napier, Macrae,
& Kehoe, 1992).
Extinction studies that have used inactivation techniques have proven to be
useful in the study o f this phenomenon. It has previously been shown that muscimol
4
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infusions into the cranial motor nuclei, which do not block acquisition (Krupa, Weng &
Thompson, 1996), effectively prevent extinction o f the conditioned nictitating
membrane (MM) response in the rabbit (Krupa & Thompson, 1993). Further, lesions or
inactivation o f the cerebellum also prevent extinction in eyeblink conditioning (Perret &
Mauk, 1995; Ramnani & Yeo, 1996). In what is probably the ‘cleanest’ cerebellar lesion
used in investigating cerebellar cortex involvement in eyeblink conditioning Thompson
and colleagues (1996) showed that Purkinje cell degeneration (pcd) mice, which lack
neural output from the cerebellar cortex, are not only able to team the eyeblink CR but
are also capable o f extinction indicating that extinction o f eyeblink CRs does not require
Purkinje cells or the cerebellar cortex for that matter. Collectively, these results suggest
that cerebellar pathways play an important rote in underlying processes of extinction
and are consistent with the widely accepted view that extinction is not simply a fading
away o f learning but rather involves other processes (e.g. Benton, 1991; Pavlov, 1927;
Rescorla, 1997; Rescoria & Wagner, 1972).
There are at least two possible and not exclusive explanations for this
inactivation-extinction result: 1) response-produced afferent feedback from the eye is
important for extinction, or 2) “reafterenee” feedback from the motor nuclei and higher
motor systems to other brain structures is important for extinction. It is clear that at least
response-produced afferent feedback plays no significant role in acquisition. We test
tins afferent feedback hypothesis here, giving extinction training with local
anesthetization o f the cornea.
5
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METHOD
Subjects
Mew Zealand albino rabbits (n-22), weighing 2.0 and 2.4 kg were used for the
experiment. Animals were individually boused with ad-lib food and water. Temperature
and a 12 hr light/ dark cycle were continuously controlled.
Behavioral Training Procedures
Rabbits were habituated to a Plexiglas restraint and the training chamber for two
daily sessions o f one hour in duration. At the end of the second day o f habituation, a
nylon suture was surgically placed in the apex of the aminals’ left nictitating membrane.
Training began on the following day.
Behavioral training consisted o f three phases. Phase 1 comprised the acquisition
phase in which animals were presented with four daily sessions o f tone-airpuff pairings.
Each acquisition session consisted of 100 trials divided into ten Hocks often trials. Each
block contained one tone alone trial, one airpuff atone trial and eight tone-akpuff
pairings. Intertriai intervals ranged randomly between 20 and 40 s. Tone-akpuff pairings
consisted o f a tone CS presented for 350 msec (1 kHz, 85 dB) with a coterminating
comeal airpuffUS (100 msec). Conditioned responses were defined as nictitating
membrane extensions o f at least 0.5mm occurring between 35 ms and 250 ms alter CS
onset. On tone alone trials the critical CR range was between 35 ms and 750 ms after CS
onset.
6
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In phase I, rabbits were required to meet the following criteria: 1) perform at
least eight conditioned responses (conditioned responses) within nine consecutive trials
by the end of the third session, and 2) give at least 80% conditioned responses during
the final acquisition session This ensured that all rabbits had reached the same level of
performance by the end o f the fourth session. At the end o f this phase animals were
randomly assigned to either a control or an experimental group.
Phase II consisted of four tone alone extinction sessions. Each session in phases
II and III consisted o f 100 trials o f CS alone presentations. The tone was presented for
350 msec with an intertrial interval that varied randomly between 20 and 40 s. Before
each session an artificial tear solution (Phoenix Pharmaceutical Inc.) was administered
to the cornea of animals in the control group. The experimental group was given
tetracaine Hydrochloride (0.5%), which effectively anesthetized the rabbits’ cornea for
more than an hour. The dosage each animal received before each session consisted of
two drops o f solution given in intervals o f five minutes during a period o f ten minutes (a
total of three applications). Phase III consisted o f three tone alone extinction sessions
with no solution administered to either group before each session.
Animals’ responses (NM extensions) were recorded using a minitorque
potentiometer attached to a nylon suture loop in the apex of the rabbits’ left nictitating
membrane. Data were saved on an IBM PC for later analysis.
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To control for the possibility that the local anesthetic might act directly on motor
nerves or nucleus, a group of 4 animals were implanted with stimulating electrodes
periorbitalfy. Four different current ranges were established according to the amplitude
of the eyeblink that resulted from the delivery o f the shock during a baseline test: High
range levels consisted o f shocks in the 3mA range which resulted in eyeblinks of more
than 10mm in amplitude. Mid range levels evoked eyeblinks that were 5mm in
amplitude. A shock of 2mA was used. Threshold levels included eyeblinks that were
between .5 and 1mm in amplitude and were generated by a shock o f 1mA. Subthreshold
levels consisted o f a shock of .5 naA which would not evoke any visible response.
All rabbits were presented with 4 daily sessions o f periorbital shock stimulation.
Animals were administered tetracaine or aline before the beginning o f each test session
and each treatment condition was counterbalanced for all 4 animals. A session consisted
of delivery o f all four current ranges presented in alternating order. Possible effects of
the local anesthetic on performance o f the response elicited by periorbital electrical
stimulation were subsequently determined.
RESULTS
Of the 22 rabbits used in the experiment, five did not reach learning criteria and
were thus excluded from further training and analysis. Three rabbits foiled to perform
eight conditioned responses out of nine consecutive trials by the end o f the third
8
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acquisition session and two reached less than 80% conditioned responses on the last day
of acquisition.
Corneal desensitization was effectively achieved in rabbits administered
tetracaine. Gentle corneal taps at the end o f each extinction session, showed that
tetracaine completely suppressed eyeblinks normally elicited by the presentation o f the
eye tap.
Figure 1 shows a summary o f the learning performance as measured by the
percentage of conditioned responses given by both groups during each treatment
condition (refer to Figure 2 for CR performance on each day).
70
60
50
o 40
| 30
20
10
0
■control
□ tetracaine
acttot edasg enodrug
Experimental phase
Figure 1 Each data point shows the overall learning performance for each phase in the
experiment. Acttot: Average CR performance during the four days o f acquisition.
Edrug: Average CR performance during the four days o f extinction with drug
administration. Enodrug: Average CR performance during the 3 days o f extinction
without drug administration. Totreacq: Average CR performance during reacquisition
9
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A repeated measures analyses of variance (MANOVA) showed that all animals
attained similar rates o f conditioned responses during the acquisition phase o f the
experiment , indicating that both groups reached the same level o f learning before being
given extinction training, (F (1,11)= 0.012, p < 0.91). In addition, no differences in
percent CR’s were found between the groups as a result o f drug administration, F
(1,11)= 0.31, p=0.58. Both the experimental and control groups gave similar extinction
response frequencies, which continued to decline in subsequent no drug extinction
sessions, F (1,11)=0.47, p < 0.83.
too - i
■ O — control
90 -|
■tetracaine
80 - I
70 1
I 6 0 -
o
c 50 -
2™
m
a .
40 H
v
30
20 i
CM
8 "
co
E
m
V .
I S T
Session
Figure 2 Percent of conditioned responses during each training session for both control
and experimental groups. No significant differences were found between the groups
throughout the course of training.
10
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Although there were no differences in the rate of conditioned responses
throughout extinction sessions between the groups, it is possible that comeal
desensitization could have produced significant changes in the amplitude o f these
conditioned responses while leaving CR rate intact. This was examined by normalizing
the amplitudes o f each extinction session with the average conditioned response
amplitude o f the last acquisition session. This yielded a measurement that reflected
changes in the strength o f the response between acquisition and extinction training
(Figure 3).
Figure 3 shows evidence of suppression in the amplitude o f the conditioned
responses of the tetracaine group during treatment days. On day two o f extinction
training, the tetracaine group presented significantly lower amplitudes compared to
controls, F (l,l 1)=L7, p=0.01. However, with the exception o f this day, conditioned
response amplitudes were not different between both groups during extinction training.
11
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- -■ o - - • control
— ■— tetracaine
2.5 -
2
13
1
ft
I
0.5 -
C O
f —
S5
to
Extinction day
Figure 3 Amplitudes o f each extinction day with those o f the last day of acquisition (see
Results section). A trend towards amplitude suppression can be seen in the tetracaine
group during treatment days. This effect was significant early in extinction.
In addition, tetracaine anesthetization of the cornea had no effect at all on the
amplitudes o f the NM responses elicited by periorbital electrical stimulation. Using a
just supratfareshold stimulus of intensity that varied between 1 and 3 mA, the mean NM
response amplitude for the control condition was 1.54 mm and for the tetracaine
condition was 1.44 mm, (t=31.8, p<0,99) (See Figure 4)
12
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14 ■ High range
□ Mid range
■Threshold
12
T
10
T
■ Sufathreshold
E
E
8
0
2
■
X
1
Tetracaine mean Control mean
Treatment
Figure 4 Mean amplitude of eyeblink responses for the four levels o f periorbital shock
stimulation administered throughout the four test sessions. High range: 3mA; Mid
range: 2mA; Threshold: 1mA; Subthreshold: .5mA.
The results o f this study indicate that blockage of corneal sensation does not
disrupt the extinction process. Animals in the tetracaine group show a gradual
decrement in CR performance that Is no different from controls even alter drug
administrations are stopped (Phase III). Further, the feet that the experimental group
extinguished in a manner that was no significantly different from the control group
seems to rule out the possibility that drag administrations may have also affected CR
performance due to motor nerve damage. One concern was whether tetracaine would
have any effects on motor performance that could account for any differences observed
DISCUSSION
13
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between the groups. To further test this, we examined eyeblink responses to periorbital
stimulation at different current thresholds. No differences were found in magnitude of
the UR to stimulation thresholds between control and tetracaine animals suggesting that
drug administration did not induce motor deficits specific to eyeblink conditioning.
However, an analysis o f response amplitudes showed that blockage o f corneal sensation
selectively affected CR performance without disrupting learning. Furthermore, there is
a general trend in amplitudes observed during treatment days where animals in the
tetracaine group seem to show more response suppression than the control group.
Overall, data from the present study suggest that proprioception related to the
cornea is not critical during extinction o f the NM. Furthermore, cutaneous
deaflerentation, achieved by a cornea! anesthetic, seems to leave UR performance intact.
While these findings seem to indicate that blockage of sensory feedback does not
interfere with extinction, a different effect has previously been reported during
acquisition. Kettlewel & Papsdorf (1971) found that, although UR amplitude and
latency remained unaffected, anesthetization o f the orbital region does have a
detrimental effect on acquisition o f the eye-blink response. The different effects seen in
acquisition and extinction due to blockage o f sensory feedback could be interpreted in
various ways. For example, it could be argued that during acquisition, cutaneous
deaflerentation would decrease the aversive or intensive value o f the US resulting in
poor learning o f the CS-US association. Extinction, on the other hand, would not be
affected by this manipulation given the nonreinfoicing nature o f the process itself. That
is, blockage o f sensory feedback during CS alone presentations would still permit the
14
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formation of what has been previously described as an inhibitory CS-n© US relation
(Pearce, 1987; Wagner, 1981; Bouton, 1991) thus allowing extinction to occur
uninterrupted.
Moreover, assuming that sensory feedback from the cornea was effectively
blocked, the disruption o f reafferenee during extinction training remains possible. As
extinction was not disrupted by this effect it nay be plausible that the performance o f
the response during tone alone presentations may have been sufficient for extinction to
occur. Although, the role of sensory input o f motor movements in eye-blink extinction
has not been previously studied, various experiments have focused and attempted to
explain the effects o f responding during extinction training. Most o f these observations
have found a positive relationship between the frequency o f responding during
extinction and the consequent deterioration o f such nonrelnforcement (Holland &
Rescoria, 1975; Rescorla & Skucy, 1969). Together with our data, these results may
indicate that expression o f the response may be an important component in the
formation o f inhibitory associations even in the absence o f sensory reafferenee.
It could be argued, however, that by administering tetracaine into the animals’
cornea we were not completely blocking sensory feedback. Indeed, peripheral areas that
may contribute to sensory relay in NM conditioning, such as the eyelids, may have not
been inactivated during extinction training. One possibility is that proprioception from
these areas may have been enough to activate areas such as the motor nuclei and
surrounding regions allowing extinction to occur uninterrupted.
15
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It has been suggested that the critical brain regions involved in extinction are
different from those required in acquisition. With growing evidence indieating that
extinction does not involve forgetting (Bouton, 1993,2002) the underlying neural
mechanisms mediating the suppression o f the behavior are stli vaguely understood. It
has previously been shown that muscimol infusions into the cranial motor nuclei which
do not block acquisition (Krupa, Weng & Thompson, 1996), effectively prevent
extinction o f the conditioned NM response in the rabbit (Krupa & THhompson, 1993).
Lesions or inactivation o f the cerebellum also prevent extinction in eyeblink
conditioning (Ferret & Mauk, 1995; Hardiman, Ramnani & Yeo, 1996). However, to the
extent that CR’s were prevented from occurring in these studies, results can be
reinterpreted simply as the consequence o f non-responding, independent o f necessary
cerebellar involvement
16
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Creator Robleto, Karla (author)

Core Title Effects of a corneal anesthetic on the extinction of the classically conditioned response in the rabbit

School Graduate School

Degree Master of Arts

Degree Program Psychology

Publisher University of Southern California (original), University of Southern California. Libraries (digital)

Tag biology, neuroscience,OAI-PMH Harvest,psychology, experimental

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