learning Behavioral psychology problem sheet

Your Name: ___________________Answer Sheet for Problem Set 1
Q1.a Enter the letter corresponding to the curve on the graph (A, B, C, or D) that matches the
stimulus condition most likely to have produced it.
Condition
Curve
Same Intruder, Same Side
Same Intruder, Different Side
Different Intruder, Same Side
Different Intruder, Different Side
Q1.b In a few sentences, explain your answer.
Q2.a Does one of the two stimuli (mechanical or electrical) produce faster habituation than the
other? If so, which one?
Q2.b If a nematode is first exposed to a mechanical stimulus, then to an electrical stimulus,
characterize their response to the electrical stimulus.
Q2.c If a nematode is first exposed to an electrical stimulus, then to a mechanical stimulus,
characterize their response to the mechanical stimulus.
Q2.d Can you see evidence of dishabituation in any of the graphs above? Explain where/why or
why not.
Q3a. Add three lines to the graph: one illustrating the underlying habituation process, a second
one illustrating the underlying sensitization process, and the third one illustrating the observable
change in startle response. Use the empty box as a legend to label your curves.
Dual Process Model
120
% of Baseline Response
115
110
105
100
95
90
85
80
1
2
3
4
5
6
7
8
9
10
Trial Number
Q3.b What will be the observed response, based on the dual-process model (a word or phrase is
fine)?
Q4:Write the letters in the table below that correspond with the matching locations on the figure.
Term
Letter
Gill
Siphon
Sensory Neuron
Motor Neuron
Facilitating Interneuron
Location of Glutamate Release
Location of Serotonin Release
Q4.b When serotonin is released, what effect does it have on the postsynaptic neuron?
Q4.c When glutamate is released, what effect does it have on the postsynaptic neuron?
PSYC 332 – Learning and Behavior
Problem Set #1
Instructions: This sheet has the questions, and information needed to answer the questions. There is a
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Q1. Male three-spined stickleback are territorial fish. When one male invades the territory of
another male, the invader is attacked ferociously. However, the attacking response, which
consists mainly of biting, may habituate.
To study this phenomenon, a male stickleback was placed in a glass box. This fish is the
Intruder. The box was then placed into the left side of another male stickleback’s territory, the
Territory Owner. The biting response of the Territory Owner, which was restricted to strikes at
the glass box, was measured every two minutes. The results of a typical male stickleback are
presented in the left panel of the graph below.
In the next session, either the same or a different Intruder was placed in the glass box, and
then the box was presented either in the same (left) side of the territory or the different (right)
side. A typical Territory Owner’s responses to these four stimulus conditions — [1] same
Intruder presented in the same side of the territory, [2] same Intruder presented in the different
side, [3] different (new) Intruder presented in the same side, and [4] different (new) Intruder
presented in the different side — are displayed in the right panel. For simplicity, assume that the
Intruder’s behavior does not change across sessions.
Q1.a On the answer sheet, match the curves (labeled as A, B, C or D) to the most likely condition
that produced that curve.
Q1.b Explain your reasoning.
Q2: The graphs below show the response of four groups of nematodes (tiny soil worms called
Vorticella convallaria) to a mechanical or an electrical stimulus. In the left panel, the
Experimental group experienced several presentations of a mechanical stimulus during the first
300 seconds and then presentations of an electrical stimulus thereafter; the Control group
experienced only presentations of the electrical stimulus. In the right panel, another Experimental
group was presented first with the electrical stimulus and then the mechanical stimulus; the other
Control group in this case experienced only the mechanical stimulus.
Q2.a Does one of the two stimuli (mechanical or electrical) produce faster habituation than the
other? If so, which one?
Q2.b If a nematode is first exposed to a mechanical stimulus, then to an electrical stimulus,
characterize their response to the electrical stimulus.
Q2.c If a nematode is first exposed to an electrical stimulus, then to a mechanical stimulus,
characterize their response to the mechanical stimulus.
Q2.d Is there evidence of dishabituation in any of the graphs above? Explain where/why or why
not.
Q3: Dual Process Model of Habituation and Sensitization.
Imagine habituation in a rat across 10 exposures to an abrupt sound. The baseline startle response
to the sound should be 100 (100% of baseline response). Assume that the sound is of lowintensity, and there is little background arousal. The underlying habituation process should show
a reduction of response tendency of 15%, while the underlying sensitization process should show
an increase in response tendency of 5%.
Q3a. Add three lines to the graph: one illustrating the underlying habituation process, a second
one illustrating the underlying sensitization process, and the third one illustrating the observable
change in startle response. Use the empty box as a legend to label your curves.
Q3.b What will be the observed response, based on the dual-process model?
Dual Process Model
% of Baseline Response
120
115
110
105
100
95
90
85
80
1
2
3
4
5
6
Trial Number
7
8
9
10
Q4: Habituation and Dishabituation in Aplysia
Q4.a Match the terms in the list below to the locations on the diagram below, illustrating the
pathway of habituation of the gill-withdrawal reflex.
D
E
G
B
A
C
F
Terms:
Gill
Siphon
Sensory Neuron
Motor Neuron
Facilitating Interneuron
Location of Glutamate Release
Location of Serotonin Release
Q4.b When serotonin is released, what effect does it have on the postsynaptic neuron (i.e. on the
receiving side of the serotonergic synapse)?
Q4.c When glutamate is released, what effect does it have on the postsynaptic neuron (e.g. on the
receiving side of the glutamatergic synapse)?
Habituation and Dishabituation of the Gill-Withdrawal Reflex in Aplysia
Author(s): Harold Pinsker, Irving Kupfermann, Vincent Castellucci and Eric Kandel
Source: Science, New Series, Vol. 167, No. 3926 (Mar. 27, 1970), pp. 1740-1742
Published by: American Association for the Advancement of Science
Stable URL: http://www.jstor.org/stable/1728291 .
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rate was less than 3 pmole/hr. A1- Habitustionand Dishabituation
though HIOMT has been thought to of the Gill-WithdrawalReflex
be the rate-limitingenzyme in melatonin
in Aplysia
synthesis (1), these observations suggest that dibutyryl cyclic AMP may
Abstract. A behavioral reflex mecontrol melatonin production in this diated by identified motor neurons inn
experimental model through another the abdominal ganglion of Aplysia
mechanism. Dibutyryl cyclic AMP may undergoes two simple forms of shortprimarily stimulate the acetylation of term modification. When the gill-withserotonin and control melatonin pro- drawal reflex was repeatedly evoked by
duction through substrate availability a tactile stimulus to the siphon or
for HIOMT.
mantle shelf, the amplitude of the reDAVIDC. KLEIN sponse showed marked decrement (haGERALDR. BERG bituation). After a period of rest the
JOANWELLER,WALTER
GLINSMANN response showed spontaneous recovery.
Section on Physiological Controls,
The amplitude of a habituated reLaboratory of Biomedical Sciences,
sponse was facilitated by the presentaNational Institute of Child Health and tion of a strong tactile stimulus to
Human Development,
another part of the animal (dishabituaBethesda, Maryland 20014
tion). Many characteristicsof habituation and dishabituation in Aplysia are
References and Notes
similar to those in vertebrates.
1. R. J. Wurtman and J. Axelrod, Adlan. Pharmacol. 6A, 141 (1968); J. Axelrod and R. J.
Wurtman, ibid., p. 157.
2. J. Axelrod, P. D. Maclean, R. W. Albers,
H. Weissbach, in Regional Neurochemistry,
S. S. Kety and J. Elkes, Eds. (Macmillan,
New York, 1966), p. 307.
3. J. Axelrod, H. M. Shein, R. J. Wurtman,
Proc. Nat. Acad. Sci. U.S. 62, 544 (1969);
D. C. Klein, Fed. Proc. 28, 734 (1969).
4. D. C. Klein, in preparation.
5. B. Weiss and E. Costa, Science 156, 1750
(1967); J. Pharmacol. Exp. Therap. 161, 310
(1968).
6. G. A. Robison, R. W. Butcher, E. W. Sutherland, Annu. Rev. Biochem. 37, 149 (1968).
7. O. A. Trowell, Exp. Cell Res. 16, 118 (1959);
L. G. Raisz, J. Clin. Invest. 44, 103 (1965);
Nature 197, 1115 (1963).
8. L. G. Raisz and I. Nieman, Endocrinology
85, 446 (1969); Modified BGJb, Grand Island
Biological Co.
9. D. C. Klein, Anal. Biochem. 31, 480 (1969).
10. T. Posternak, E. W. Sutherland, W. F.
Henion, Biochim. Biophys. Acta 6S, 558
(1962).
11. J. Axelrod and H. Weissbach, J. Biol.
Chem. 236, 211 (1961); D. C. Klein and
S. Lines, Endocrinology 84, 1523 (1969). The
HIOMT reaction mixture includes S-adenosyl
methionine
(3 X 10-5M),
N-acetylserotonin
(10-4M), 100 nc of [14C-methyl] S-adenosyl
methionine (Amersham Searle), and 40 percent of the homogenate of one gland in a
total of 0.3 ml of 0.5M sodium phosphate
buffer (pH 7.9).
12. During review of this paper, Shein and Wurtman [Science 166, 519 (1969)] reported that
dibutyryl cyclic AMP increases the formation
of [14C]melatonin and [14C]serotonin from
[1&C]tryptophan. Our results agree with these
because to increase synthesis of melatonin
from tryptophan the synthesis of (the intermediate) serotonin may be increased. However, studies using the inhibitor of serotonin synthesis p-chlorophenylalanine (pCPA)
seem to indicate that the effect of dibutyryl
cyclic AMP in stimulating melatonin production is not dependent on prior elevation
of serotonin production. We have found that
whereas pCPA (1.0 rnM) inhibits the effect
of dibutyryl cyclic AMP on conversion of
[3H]tryptophan to [3H]melatonin to 35 | 20
percent of the normal stimulation, the effect
of dibutyryl cyclic AMP on [14C]serotonin
conversion to melatonin is not reduced by
pCPA, but is slightly enhanced (125 § 20
percent). This suggests that a specific site
of action of dibutyryl cyclic AMP involved
in stimulating melatonin production is at a
metabolic step that does not depend on the
new synthesis of serotonin, and will take
place when an exogenous source of serotonin
is provided, as in the experiments presented
here.
8 October 1969
The analysis of the neural mechanisms of learning and similar behavioral modifications requires an animal whose behavior is modifiable and
whose nervous system is accessible for
cellular analysis. In this and the subsequent two papers (1, 2) we have
applied a combined behavioral and
cellular neurophysiologicalapproach to
the marine mollusk Aplysia in order
to study a behavioral reflex that undergoes habituationand dishabituation.We
have progressively sumplifiedthe neural
circuit of this behavior so that the
action of individual neurons could be
related to the total reflex. As a result,
it is possible to analyze the locus and
the mechanisms of these behavioral
modifications. We now describe behavioral parameters of habituation and
dishabituation of the gill-withdrawal
reflex in Aplysia.
Habituation and dishabituation are
simple behavioral modifications often
considered to be the most elementary
forms of learning (3-5 ) . Habituation
is the decrement of a behavioral response that occurs when an initially
novel stimulus is repeatedly presented.
Spontaneous recovery of the decremented response occurs if the stimulus
is withheld for a period of time. Dishabituation, the restoration of a previously decremented response, occurs
following a change in the stimulus pattern, such as the presentation of another, stronger stimulus (4).
Parametricallysimilar forms of shortterm habituation, which last from several minutes to several hours, have been
demonstratedfor a variety of behavioral responses in all animals which have
clearly developed central nervous sys-
A
4.0
3.5
3.0
2.5
2.0
Fig. 1. (A3 Dorsal view of an intact animal showing a fully contracted gill. Normally
the parapodia and mantle shelf obscure the view of the gill, but they have been retracted to allow direct observation. The relaxed position of the gill is indicated by the
broken lines. The tactile receptive field for the gill-withdrawal reflex includes the
siphon and the edge of the mantle shelf. (B) The animal was immobilized in a small
aquarium containing cooled and aerated circulating seawater. The edge of the mantle
shelf was pinned to a substage, and a constant and quantifiable tactile stimulus consisting
of a brief jet of seawater was delivered by a Water Pik (a commercially available oral
hygiene apparatus). The stimuli were controlled by a Grass S-8 stimulator and were
usually 800 msec. The gill contractions were monitored with a photocell placed under
the gill. The output of the photocell was linearly related to the area uncovered as the
gill contracted and was recorded on a polygraph. (C) Gill responses to individual tactile
stimuli of different intensities. The stimuli were separated by very long intervals of
time. The intensity of the stimulus could be adjusted anywhere from a very light touch
( 1.0, arbitrary units) to a very intense pressure (5.0) . The weakest stimulus (2.0)
evoked only a small gill contraction which consisted of a simple, short-latency withdrawal. Stronger stimuli (2.5, 3.0) evoked bigger and longer lasting gill responses of
similar short latency but, if strong enough (3.5, 4.0), could bring in a second, longer
latency component. The latency for this second component was quite variable, and with
the strongest stimuli it sometimes merged with the first component.
SCIENCE, VOL. 167
1740
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tems (3-6). The behavioral similarity
across species suggests that there may
be common neuronal mechanisms of
short-term habituation.
We have examined habituation and
dishabituation of a behavioral reflex
controlled by the abdominal ganglion
of Aplysia. This ganglion offers a
number of advantages for the cellular
neurophysiologicalanalysis of behavioral mechanisms. It contains a small
number of nerve cells, all of which are
large enough to be penetrated with
microelectrodes for recording synaptic
potentials and for direct stimulation.
Many of these cells have been identified as unique individuals or as members of functional clusters (7). The
connections of some of the cells with
each other (8) and with peripheral
sensory and motor structures (9) have
also been specified.
The specific behavior that we have
chosen for analysis is a gill-withdrawal
reflex that occurs as part of a larger
defensive withdrawal response that is
triggered by a potentially noxious tactile stimulus (Fig. 1A). Analogous defensive escape and withdrawal responses are present in other invertebrates as well as in vertebrates. Probably because defensive reflexes must
be fast to be effective, the neural circuitry of these reflexes is usually relatively simple, often involving only a
few synaptic relays. In addition, defensive reflexes typically habituate quite
readily (6).
The gill system of Aplysia offers further advantages for a neural analysis
of behavior. First, most of the motor
neurons that control gill contractions
have been identified (9). Second, the
gill reflex can be effectively studied in
a restrained, but otherwise intact, animal. Third, reflex withdrawal of the
gill can be elicited from stimulation of
a receptive field that does not include
the gill itself, thereby minimizing the
contributionof local peripheralreflexes.
Finally, gill movements occur both
reflexly, as a result of sensory stimulation, and spontaneously, as a result of
the endogenous activity of neurons
within the ganglion (9). The gill system therefore also serves as a potential
model for a variety of more complex
behavioral processes that involve stimulus pairing or pairing of a spontaneous
response with a reinforcing stimulus.
A major problem in studying behavioral responses in Aplysia is to restrain
these soft-bodied animals with minimum damage and to apply reproducible
stimuli to their peripheral sensory re-
ceptors. We accomplished this in the
apparatus shown in Fig. IB. The animal was immobilized in a small seawater aquarium, and gill contractions
were monitored with a photocell placed
under the gill. The gill-withdrawalreflex can be evoked by a tactile stimulus
within a receptive field that is centered
on the siphon and mantle shelf and
falls off sharply in the surrounding
regions. The area along the dorsal edge
of the mantle shelf was pinned to a
substage, and a tactile stimulus was delivered by means of a brief jet of seawater (10).
We first examined the responses of
the animal to individual stimuli that
were presented to the same spot on the
skin and differed only in intensity
(Fig. 1C). The weakest stimulus evoked
only a small gill contraction which
consisted of a simple, short-latency
withdrawal. Stronger stimuli evoked
bigger and longer lasting gill responses
of similar short latency, but, if strong
enough, these stimuli also brought in a
second component that usually had a
much longer latency.
In order to simplify the behavioral
analysis of habituation we adjusted the
stimulus intensity (Fig. 1C) to obtain
a short-latency component in the absence of a superimposed late component. In addition, we further restricted
A1
Rest
122 min
1
10
4
81
14
Total = 80 stimuli
ISI = 3 min
A2
A
I
I1
ISI = 1- min
120
0
80
00–
60
23
21
18 A
Dishabituation
Total = 20 stimuli
27
25
10 sec
B
– – – – – –
100
C
0L
0.
13
9
3′
–
–
–
–
–
–
–
–
–
.
–
–
–
-0
0
40
20
0I
II
0
I
20
I
I
40
I
I
60
1I
1I
80
1
100
I
120
1
.
140
Minutes to first recovery trial
Fig. 2. Habituation,spontaneousrecovery,and dishabituationof the gill-withdrawal
reflex. (A) Recordsfrom two responsehabituationsin a single preparation.The interval between stimuli (ISI) and total number of habituatorystimuli are indicated.
Part 1 shows decrementof the responsewith repetitionof the stimulus.Following a
122-minuterest the responsewas almost fully recovered.Part 2 shows a later experiment from the same preparation.After rehabituationof the responsea dishabituatory
stimulusconsistingof a strong and prolongedtactile stimulusto the neck region was
presented at the arrow. Successive responses were facilitated for several minutes.
(B) The time courseof recoverywas estimatedby habituatingindividualanimalswith
repeated stimuli and testing for the percent of recovery by presenting a single stimulus
after different intervals of rest. The curve is based on 44 separate habituation and re-
covery runs in 27 differentanimals.Each point is the averageof three measures(last
point based on only two) taken at roughly the same interval. In longer experiments,
later responses would often recover beyond the initial control level (for example,
compare the first response in Al with that in A2). For the purpose of this figure
all responses equal to or greater than the control response for that run were assigned
a value of 100 percent. The shortest time in which full recovery occurred was 10
minutes, whereas the longest time in which the response was not fully recovered
was 122 minutes.
27 MARCH 1970
1741
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dishabituation, we have
habituation of the
oStim = 2.5 ISI = 100 sec
dishabituatory stimulus with repeated
presentatioJls, (v) greater habituation
,>rwith short rather than long intera _J
stimulus intervals, and (vi) greater
with weak rather than
habituation
*
* :.
4Q
strong stimuli. Three other parametric
30
characteristics of habituation have
been noted but do not apsometimes
30
b
– t
pear to be characteristicof habituation
in Aplysia. These features are (vii)
20
o
20
.
greater habituation with. repeated pe
riods of habituationand recovery, (viii)
Q
generalizationof habituation to a stimulus ill another part of the receptive
3\
\
c
l l I
t
i i
o
O 10 20 30 40 50 60 7O
10
8
6
4
field, and (ix) prolongation of recovery
2
Consecutive minutes
Consecutive stimuli
additional stimulation after
following
20 sec
2 sec
Fig. 3. (A) Habituationwith weak and strongtactile stimuli.Responsesto the stronger the response has decremented to an
stimulus(filledcircles) are initialb largerand show less decrementwith repetitionthan asymptote. The existence of a satisresponsesto the weaker stimulus (open circles). (B) Spontaneouscontractionsand factory fit between many characteristics
reflex contractionsof gill. Part 1, spontaneousgill contractions(filled circles) remain of habituation in Aplysia md in verteconstantin amplitudewhile refles contractions(opencircles) producedby tactile stimuli
presentedat 5-minuteintervalsshowhabituationand then recoverywith a 30-minuterest. brates suggests that it may be of genPart 2, samplerecordsfrom this experimentto compareamplitudeof refles and spon- eral iulterestto analyze the underlying
taneous contractions(a) before reflex habituation,(b) during maximumhabituation, neuronal mechanismsSand this will be
and (c) following recoverywith rest. The spontaneouscontractionshown is the one the object of the following two papers
occurringclosest in time, either before or after the reflex contraction.Note the differ(1, 2).
ence in time calibration.
PINSKER
HAROLD
KUPEERMANN
TRVING
VINCENTCASTELLUCCI
our analysis to the earliest peak of the sponse was facilitated, and successive
ERIC KANDEL
several
for
elevIated
response which is most closely asso- responses remained
Physiology and
Departments-of
dishabituatory
a
occasion,
On
minutes.
monothe
of
activlation
ciated with
University
synap!tic pathway to be analyzed in stimulus facilitatedthe decrementedre- Psychiatry, New York
Health
Pablic
and
School,
Medicczl
sponse to an amplitude greater than
neurophys.iologicalexperiments (2).
City
the
of
Institate
ReseGtrch
recontrol
With repetition of the tactile stim- the initial (unhabituated)
of New York, New York
ulus at intervals that ranged from 30 spoJlse.
As described above, a contraction
seconds to S minutes, the gill responses
habituated to an average of 25 percent similar to that evoked by tactile stimScience,
of control amplitude (S to S percent) ulation also occursl spontaneously. We
to
contractiou
spontaneolus
this
used
ibid.,
sometimes
(Fig. 2A). Hlabituationhas
Learning and Instinct in
been seen with intervals as long as examine whether reflex habituatioll reAnimals
20 minutes between stimuli. Typically, sults from fatigue of the gill musculathe major part of the decrement was ture. We found that spontaneous gill
Psychol. Rev. 173,
Psychol. Bull. 40,
produced by 1he first 5 to 10 s.timuli contractions that occurred before the
Ann. Phyonset of reflex habituatioue during
in a series (see Fig. 3, A and B).
siol. Rev. 32,
Periods of rest that ranged from 10 maximum response decrement, and afJ. Neurophysiot.
minutes to more than 2 hours were ter recovery of the reflex, were of
requiredfor full recovery from habi.tua- similar amplitude (Fig. 3B), which
ibid., p.
Science 15S,
tion To obtain a more quantitative indicated that gill fatigue is not a
Science
measure of the rate of recovery we factor in habituation This inference is
that
iding
the
by
supperted
further
plotted percent recovery in 44 separate
response habituations with a single an extrastimulus can dishabituate a
stimulus given after difEerentintervals habituated response and that strong
of rest (Fig. 2B). The data suggest stimuli (which are more likely to prothat there is a rapid plhase lasting 10 duce fatigue) produx less habituation
to 20 minutes that accounts for about than weak stimuli (Fig. 3A).
Thompson and Spencer (4) de75 to 85 percent of recovery, followed
by a slow and highly variable return scribed nine parametric characteristics
to the -originalresponse level and often of behaviorblhabituationin vertebraltes.
Six of these characteristics hfave conbeyond.
After habituation of ffie response, sistently been found in Aplysta. In adthe illustrations. Supported by PES grant
MH 15980, .career scientist award K 5-ME
a single strong tactile sti.mulus pre- dition to (i) response decrement,usual18558 to E.R.K., career development Waward
sented to another part of the animal ly a negative exponential function of
K1-ME 12240 to I.K., and a Canadian Medical Research Council Fellowship to V.C.
prolduceddishabituation (Fig. 2A, part the number of stimulus presentations,
2). The previously decremented re- (ii) spontaneous restoration with rest, 21 November 1969
contraction
A *Stim = 3.5 iSI = 100sec B | oSpontaneous
Reflex contraction J
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27 MARCH 1970
–
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1745