Conditioning Situation Versus Intermittent Stimulus
Conditioning Situation Versus Intermittent Stimulus
Conditioning Situation Versus Intermittent Stimulus
Acknowledgments
I also owe very best thanks to my daughter Joanna-Veronika Warwick for her
valuable editorial assistance.
W.W
Vll
Introduction
orienting reaction.
Another case, less dramatic than the first mentioned is an event observed
everyday in school. A bell sounding before class results in students entering
the classroom; the same sound at the end of the class results in students
leaving the room. Therefore, the same sound of a bell evokes two different
behaviors, depending on the
2 Conditioning
new immigrants to this country. One immigrant told me, “there is no life
here” and constantly returned to talk about his earlier times “there”
Chapter 5 consists of two parts. The first part presents results of studies on
the role of the experimental situation in instrumental conditioning; the
second part describes the studies on the single (intermittent) conditioned
stimulus tested in various situations.
Introduction 3
Rounding out the book, a general summary and conclusions once again
emphasize the effect of the situation on conditioning processes.
Classical Conditioned Reflexes
Ivan Pavlov’s historic discovery of the principle of the conditioned reflex was
an enormous break in the scientific approach to the understanding of brain
functions.
One of Pavlov’s first observations in this area was made during a study on the
secretion of saliva in dogs. Namely, Pavlov found that the animals salivated
not only when they were eating food offered by a caretaker, but also when
they saw the caretaker approaching or heard his footsteps. This simple
observation indicated that the inborn reaction of salivation to food (or other
objects) in the mouth may also appear to originally neutral stimuli which
accompanied or immediately preceded the food intake.
6 Conditioning
One of these topics was the process of inhibition of the previously acquired
behavior. Inhibition could be either external or internal. External inhibition
could be caused by an appearance of an extraneous stimulus, not involved in
the conditioned reflex arc. For instance, such extraneous stimulus could be a
sudden noise, not used previously, an unknown person who appeared
suddenly during the presentation of the conditioned stimulus, etc.
8 Conditioning
given 1-3 seconds after the beginning of the action of CS) the reinforcement
was delayed by an additional, e.g., 5 seconds, in each consecutive session,
until the delay reached several minutes. As a result of such procedure, the
conditioned salivation did not start immediately after the onset of the
conditioned stimulus, but only 30-60 seconds or longer after the beginning of
action of the CS. The same result could be achieved when after the
establishment of the conditioned reflex the delay in reinforcement was
prolonged to the desired length rapidly, without any gradual steps. However,
in this case, some complications in the dog’s behavior could appear during the
training period (Pavlov, p.89).
Generalization of Stimuli
other sensory systems (i.e., other analyzers). This phenomenon was called
generalization of stimuli (Pavlov 1960, p. 113). It is often observed both in the
laboratory and in natural life.
The ability of conditioning was also observed on the gall bladder; namely, the
contractions of the gall bladder increased when food was shown to a hungry
dog (experiments of Kurtsin and Gorshkova, described by Bykov 1957).
The extension of the research on conditioned reflexes also included some new
methods. An example of such novelty were experiments conducted by
W.H.Gantt who also was one of Pavlov’s pupils. Gantt chose the functions of
the heart and the circulatory system as the object of research and
demonstrated that the rules of conditioning also extended to this area (Gantt
1944, 1960, Gantt and Hoffman 1940).
All the above described experimental results, which represent only a small
part of the pioneer achievements of Pavlov and his pupils, were obtained with
the use of single intermittent stimuli. In their research there was no problem
of the situational background. However, Pavlov was aware of the existence of
a great number of various stimuli in the environment which could influence
the experimental results. In order to avoid interference of external stimuli
during experiments with conditioned reflexes, a special building (known also
as a “tower of silence ”) was constructed (see fig. 3 at the beginning of Lecture
3, Pavlov 1960).
10 Conditioning
According to Pavlov (1951), the general rule of the conditioned reflex also
included the motor conditioned reflexes. When a movement is performed, it
produces a sensory input of impulses from the muscle. When this input is
each time followed by the reinforcement (e.g., food), the conditioned
salivation appears to the movement (such as a flexion of the dog’s leg) before
the food has been offered. This was demonstrated by Krasnogorski (1911; cit.
by Pavlov 1951), who used a special device to produce a passive lifting of the
dog’s leg. In this case, therefore, the motor conditioned stimulus was indeed
analogical to the acoustic, visual, or tactile conditioned stimuli (see a scheme
of this conditioned reflex in fig. 2.1).
11
12 Conditioning
FIGURE 2.1
(classical CR)
A diagram of motor conditioned reflex in experiments of Krasnogortski
according to the explanation of I.P. Pavlov (see Ch. 2).
The design of their first study was very simple. Using a mechanical device
which could be controlled by them from a distance, they produced a flexion of
the dog’s leg to an acoustic stimulus which was immediately followed by food.
After several repetitions of this procedure, the dog started to perform the leg
flexion actively (i.e., without any help of the mechanical device) to the
acoustic stimulus. Using various kinds of reinforcement (positive
reinforcement such as food, or negative reinforcement as a puff of air to the
ear, as well as the presence of the reinforcement in some cases and its
absence in other cases), they found that four
13
varieties of the motor conditioned reflexes exist. Let us briefly describe each
of these varieties.
First variety. When a specific movement (e.g., flexing the leg) is performed to
a conditioned stimulus (e.g., a tone) and is always followed by obtaining a
reward (e.g., food), then this movement will always spontaneously appear to
this stimulus.
Fourth variety. When a specific movement (e.g., leg flexion) appearing to the
conditioned stimulus (e.g., a tone), is always followed by a punishing
unconditioned stimulus (e.g., an air puff to the ear), then this movement will
cease to appear and an antagonistic movement (e.g., leg extension) will be
performed instead.
Konorski and Miller concluded that the motor conditioned reflexes differed
from the Pavlovian reflexes in that the reinforcement in the former was
dependent on the performance (or inhibition) of a specific movement,
whereas there is no such condition in the Pavlovian reflexes. Therefore, they
called the motor condi-
14 Conditioning
tioned reflexes “conditioned reflexes type II,” while reserving the name
“conditioned reflexes type I” for Pavlovian reflexes. Later the terms “classical
conditioning” for conditioned reflexes type I, and “instrumental conditioning”
for conditioned reflexes type II, were introduced by Hilgard and Marquis
(1940, Kimble 1961) and became commonly used by the behavioral scientists.
Konorski and Miller admitted that the prototype of the conditioned reflexes
type II was studied earlier by Thorndike (1911) who called this kind of
behavior “the trial-and-error learning.”
15
FIGURE 2.2
a) The dog performs the movement not only during the action of the
conditioned stimulus (CS) but also during the intervals between the
successive applications of the CS.
b) When food is offered exclusively in the presence of the CS, and never in its
absence, the movements appear only during the action of the CS.
1936, Konorski 1967) included the fact that the trained instrumental
movement appears not only in the presence of the conditioned stimulus but
also in its absence, i.e., in response to the experimental situation. As shown in
part a of fig. 2.2 (which is a copy of an original record taken during the early
period of the training), the instrumental movement (lifting the left foreleg)
appears not only during each application of the conditioned stimulus but also
in the intervals between the successive applications of CS. When the re-
16 Conditioning
FIGURE 2.3
« mil l) miimm ii
W l * --* ■ * — ul 1 1 Ml ...
From top to bottom: movements of the foreleg, pressing the lever, salivation.
Arrow denotes the increased portion of food.
inforcement (food) was given exclusively in the presence of the CS, the
instrumental movements started to be performed only during the presence of
CS and not in its absence (fig. 2.2, part b).
In another experiment, the dog was trained to lift its right foreleg and place it
on a small board; each movement was rewarded with food. No intermittent
CS was used, therefore this instrumental movement was performed to the
situation only (fig. 2.3). When food reinforcement ceased to follow each
movement, the trained movement gradually disappeared. This supported the
view that
Early studies of Konorski and Miller (Miller and Konorski 1928, Konorski and
Miller 1936, Konorski 1939, 1967) included the phenomenon of “switching” or
“interchange” which consisted of the appearance of different reactions to the
same CS in specific circumstances. This problem will be discussed in the
following chapter.
All the studies mentioned above, as well as most studies conducted in his
laboratory by his pupils and associates, were described and analyzed by
Konorski in his last book entitled Integrative Activity of the Brain. An
interdisciplinary approach published, in 1967.
18 Conditioning
quired motor tasks for cat and dogs included pressing a lever, pulling a cord,
turning a knob, pushing the door, or in some special experiments, licking the
fur or scratching. Monkeys had to perform more complex movements, such
as manipulations with a wooden peg, hook, metal bar, or turning a knob by
270°, to get a reward. This kind of behavior, which was called by Thorndike
“trial-and-error learning,” corresponds to Konorski and Miller’s “conditioned
reflexes type II” (as mentioned earlier).
Skinner developed special techniques for studying the operant behavior. His
subjects were either pigeons or rats. The operant task for pigeons was to peck
at the visual stimulus and for rats to press the lever. The movements of the
subjects were automatically recorded enabling an immediate analysis of the
obtained data. This resulted in the compiling of a great amount of
experimental data related to such problems as discrimination of responses,
generalization of stimuli, delayed responses, inhibition (extinction), chained
schedules of conditioning, and others (Ferster and Skinner 1957). Skinner
also theorized on problems such as “drive” and “drive and conditioning,” and
discussed the effects of emotions and drugs on conditioning (Skinner 1938).
However, he did not relate the behavioral results of his studies to brain
functions.
One of the most important points of Skinner’s theoretical approach was the
nature of the stimulus. Skinner emphasized the fact of the presence of great
variability of stimuli in the environment and the necessity of the
differentiation of the chosen stimulus before its conditioning. He also
stressed that a chosen motor act to be conditioned is never completely new
but it could be performed
19
Because all motor activity of the organism is based on the inborn motor
reflexes, it is possible that the movement chosen for the training is not new
but could have been performed occasionally in the history of the subject.
Being aware of this, the investigator must be sure that the chosen movement
was never performed by the subject during the preliminary control sessions.
The appearance of that movement during the training sessions can be
considered proof of the establishment of this movement as an instrumental
conditioned reflex. The problem of the absolute novelty of the movement
should never be considered.
The above discussed points are only a part of Skinner’s hypotheses and
suggestions. The interested reader is directed to Skinner’s original
publications (Skinner 1938, Ferster and Skinner 1957).
The Phenomenon of Switching
21
22 Conditioning
tone) and the lifting of the left foreleg (CRII, the reaction trained to the bell
but never to the tone). According to Konorski’s interpretation, the obtained
result demonstrated that a CS connected by training with a definite defensive
CRII, may elicit this reaction to another CS related to a different defensive
reinforcement. Konorski also hypothesized that similar cases of switching
may also occur in conditioned reflexes with approach type of reinforcement
(Konorski 1939, p. 20). This supposition was verified by Fonberg (1961, 1967),
as described later in this chapter.
This type of experiment was performed by Konorski and Miller during their
visit to Pavlov’s laboratory (Konorski and Miller 1936, Konorski 1939). First,
an alimentary instrumental reaction of lifting the right hindleg to a specific
noise and a defensive (avoidance) reaction of lifting the right foreleg to the
sound of bubbling water were established in a dog. During each session a
band was fastened on the trained leg for recording purposes. It was found that
when the band was fastened on one leg only, the dog lifted this leg both to
alimentary and defensive stimuli. Only when the bands were fastened on both
trained legs, were the reactions correct, i.e., the alimentary CS produced the
alimentary CRII, and the defensive CS produced the defensive (avoidance)
CRII. However, when the alimentary CS was applied after a number of trials
with the defensive CS, the dog first performed a defensive CRII and then only
the correct alimentary CRII. And, vice versa, when the defensive CS was
preceded by a number of successive applications of alimentary CS, the
alimentary CRII appeared first and then only the correct defensive CRII was
performed. This case of switching (regarded by Konorski and Miller as a
result of the “inertia of excitation”) was not always observed in all dogs.
24 Conditioning
lifting the foreleg (an alimentary CRII, previously established to the bell).
These experiments showed that the alimentary CRII may appear to a
defensive CS in conditions others than inertia of preceding excitatory
processes.
The obtained results were interpreted by Konorski and Miller (1933, 1936;
Konorski 1939, 1967) in the following way. There are two kinds of stimuli: the
determining and the releasing stimuli. The determining stimuli, such as a
band on the dog’s leg, or overall experimental situation, do not elicit the
conditioned reaction but they determine which kind of reaction is to be
performed; the alimentary or defensive background produced by preceding
application of unconditioned stimulus could also be considered a determining
stimulus.
The releasing stimuli are usual intermittent stimuli that elicit the reaction. As
previously described, a partly inhibitory (differential) alimentary CRI could
elicit a defensive CRII trained to other CS; also, a deep extinction of an
alimentary CRI could result in the appearance of a defensive CRII established
to a different CS. An application of an aversive unconditioned stimulus could
also release a previously established alimentary CRII.
The boundary between the determining and the releasing stimuli is not
absolute, however; for instance, a continuous determining stimulus may be,
at the same time, a releasing stimulus, and vice versa, a sporadic stimulus
may both determine and release the reaction (Konorski 1967, p. 389).
Two different avoidance CRIIs were trained in four dogs in the usual
chamber. The conditioned stimuli were the whistle, the bell, the tone, and the
light. The training was conducted in separate
series to each CS. Each animal was trained to lift the right hindleg to the CS 1
to avoid a nociceptive reinforcing stimulus which was either an electric shock
or a puff-to-ear in various dogs. Then, in another series of sessions, the dog
was trained to place its left foreleg on a platform to stimulus CS2 to avoid a
nociceptive stimulus. After the establishment of the second CRII, a test was
performed in which CS1 and CS2 were applied in the same session. It was
found that both reflexes (lifting the right hindleg to CS 1 and placing the left
foreleg on the platform to avoid nociceptive stimulus) were usually
performed correctly and they did not interfere with each other when the
animal was calm. However, when the animal was excited, both avoidance
reactions appeared in the same trial (to either CS1 or CS2), often
simultaneously or almost simultaneously; these movements also were
observed in the intertrial intervals. This kind of switching was observed in
two dogs.
Approach Behavior
In another study (Fonberg 1967), two dogs were trained to press a lever to the
tone to obtain food. Then chronic electrodes were implanted in these dogs’
lateral hypothalamic “feeding” area (Anand and Brobeck 1951). Electrical
stimulation of the hypothalamic points corresponding to the tips of the
electrodes, produced eating in satiated animals. When the hypothalamic
stimulation was substituted for food reinforcement, the dogs began to
perform the previously trained alimentary CRII (lever pressing).
26 Conditioning
28 Conditioning
the dog’s cerebral cortex during the experiments on switching. In the morning
session, an auditory stimulus was paired with electric shock to the dog’s left
hindleg; in the afternoon session, the same auditory stimulus was paired with
an electric shock to the right hindleg. This training resulted in lifting the left
hindleg to the CS in the morning session and lifting the right hindleg to the
same CS in the afternoon session. At the same time, the EEG record showed
the presence of high amplitude waves in the leads from the boundary area
between the motor and parietal areas, ipsilateral to the trained legs, i.e., from
the left hemisphere in the morning sessions and from the right hemisphere
in the afternoon sessions. This EEG pattern appeared at the beginning of the
session, before the application of the CS, and persisted throughout the
session. This kind of EEG activity could reflect the action of the tonic
stimulus.
periments and Pavlovian studies was that the context studies were conducted
on human subjects whereas Pavlovian studies were conducted on dogs.
The experimental data described in this chapter are strictly related to the
influence of the situational background on behavior. They also support the
hypothesis that conditioning of the situation does take place, and its effect on
behavior dominates over the effect of the single conditioned stimulus. This
problem will be further discussed later in this book.
Cases of Dependence of Conditioned Behavior on Situational Factors
32 Conditioning
However, this transfer was observed only when the general experimental
situation was unchanged. After placing the animal on the experimental table
and then rotating the table by 180°, the previously observed effect was no
longer observed. That is, stimulation of the left occipital cortex or left motor
cortex produced normal contralateral movements, whereas stimulation of the
right occipital cortex remained ineffective. On the other hand, stimulation of
the right motor cortex elicited contralateral reactions, i.e., head rotation to the
left and flexion of the left forepaw. As commented by Giurgea (1989) “even
such as ancient, phylogenetically and ontogenetically rooted physiological
response as motor contralaterality following efficient motor cortex
stimulation might be critically modulated by shortened conditional reflex in
the experimental situation.”
34 Conditioning
lus was applied several times in each experimental sessions at intervals of 3-5
minutes and was not reinforced with food. However, food was given during
the intervals, without any relation to the intermittent stimulus. Already on
the first session of this study, the latency of the instrumental reaction to the
CS, which was 4 seconds in the first trial, increased to 12 seconds in the 7th
trial. On the other hand, the salivary CR which was 35 points on the
manometer scale, in the first trial, decreased to 18 in the 7th trial although
the salivation during eating remained about the same up to the end of the
session. During the following sessions both salivary and instrumental
conditioned reactions to the CS gradually diminished and after 6-8 sessions
they completely disappeared. However, as soon as the reinforcing stimulus,
food, was given again to the extinguished stimulus (bubbling), both salivary
and instrumental conditioned reflexes gradually recovered.
On the other hand, when the dog was tested on another day, in the free-
behavior room, the same amount of chlorpromazine did not have any effect
on conditioned reaction. The same dog, after injection of this drug, taken just
in front of the Pavlovian room, showed a full neuroleptic effect, whereas
when placed in front of the free-behavior room, this dog appeared almost
normal as if he did not receive any drug. When taken back in front of the
Pavlovian room, this dog again showed the neuroleptic behavior whereas no
such behavior was observed when placed in front of the free-behavior room.
Such alternation in behavior could be induced 10-12 times during a period of
30-45 minutes, in various dogs.
This finding supports earlier observations of Kupalov (1961) that dogs were
more resistant to neurogenic procedures in the free-behavior situation than
in the Pavlovian room. The situational effects on behavior were also
demonstrated in studies of Zuckermann and Buffy (1960), and more recently
by Khananashvili (1983).
The tests for conditioning showed that rats in the morphine groups,
compared to the saline group, showed an anticipatory hypothermia in the pre-
injection environment; this was just opposite of the unconditioned
hyperthermia to morphine. On the other hand, the rats in the morphine
groups showed a conditioned hyperthermia when tested in the injection
environment after a period of abstinence. The results indicate that the
changes of body temperature depended on the environment which was
related to the morphine injection. In other words, it can be said that
conditioning to the environment took place.
38 Conditioning
environmental tolerance to the hypothermic effect of ethanol developed. The
environment-dependent tolerance was also seen when these mice were tested
with intraperitoneal injections of ethanol.
Two patients, a man 51 (patient 1) and a woman 52 years of age (patient 2),
were taken to a doctor’s office, a lecture room, a car, an apartment, and a gift
shop. It was found that the patients were excessively dependent on
environmental cues. Here are the excerpts of observations made in various
situations.
When in the doctor’s office some medical instruments were put on the desk,
the female patient immediately picked up the blood pressure gauge and
measured the doctor’s blood pressure. Then she took the tongue depressor
and examined the doctor’s throat.
Both patients were taken by the doctor to a lecture room with a buffet of food
and beverages. Patient 1 helped himself to the food
Conditioned Behavior on Situational Factors 39
and the orange juice and behaved like a guest. In contrast, patient 2 behaved
like a hostess. When she saw some stacks of chairs, she proceeded to set the
chairs side by side. Then she offered some foods on various plates and a glass
of orange juice to the doctor. She did not serve herself.
The doctor took patient 1 and his female friend to his apartment and said that
it was a “museum.” Patient 1 immediately started to examine the paintings as
if he were in a real museum. His behavior was appropriate for any usual
museum visitor. Moreover, when he saw that one painting was down on the
floor, he took the hammer and nails (which had been left next to the
painting), hammered a nail into the wall and hung the painting. Then the
doctor and patient walked around the apartment and also entered the
bedroom. When the patient saw the bed without the bedspread, he
immediately started to undress and then got into bed and prepared to sleep.
The behavior of patient 2 in the bedroom was slightly different. She did not
undress but lay down immediately. Then she accompanied the doctor to a
table with various items used for intramuscular injections. When the doctor
showed her a syringe, she took it and gave the injection to the doctor.
All studies described in this chapter show the decisive role of the
environmental situation on the behavior. From the point of view of
conditioning, the behavior became associated (conditioned) to the situation
due to the repetitive application of the reinforcing stimulus in this situation.
This problem will be discussed from the theoretical point of view in a later
chapter.
Studies on the Role of the Situation and the Intermittent Stimulus in
Conditioning
In a usual Pavlovian chamber, each dog was trained to perform two reactions:
one was lifting the right hindleg (CR1) to an acous-
41
42 Conditioning
tic stimulus, the sound of a rattle (CS1), and the other was placing the right
forepaw on the feeder’s platform (CR2) to a visual stimulus, a rotating disc
(CS2). Each reaction was trained in a separate series of sessions. A two- week
series of daily sessions with CS1-CR1 alternated with a fortnight series of
sessions with CS2- CR2. It was observed that at the beginning of the next
series each CS evoked a “wrong” reaction, i.e., the reaction trained during the
preceding series (for instance CR2 to CS1, or CR1 to CS2). Only when the
wrong reaction was not reinforced by food did the correct reaction appear. It
was also found that during further sessions of a series, the dogs often
performed both reactions to each CS (i.e., both CR1 and CR2 to either CS1 or
CS2).
In this experimental series, each dog was trained in two situations different
from each other. One of these situations was a usual Pavlovian chamber
(situation A) in which the dog was trained to perform reaction CR2 (placing
the right forepaw on the platform)
to a visual stimulus CS2. Another situation was a large empty pen (without
any stand or a platform) arranged on the floor of a room located in a separate
building (situation C). In this situation the dogs were trained to perform
another instrumental reaction CR3 (lifting the left foreleg in one dog, and
assuming a “begging” posture in the other dog) to a stimulus called “noise.”
44 Conditioning
his study, dogs were trained to perform a definite instrumental reaction to get
a food reinforcement. However, when the experimental procedure was
changed, namely the food reinforcement was given immediately after the
start of application of a definite conditioned stimulus, the dog began to eat at
once, without first performing the instrumental reaction. After training (in
which the delay in offering food was gradually extended) it was possible to
obtain a period up to 10 seconds of waiting for food, without the instrumental
reaction. In this case, the situation included a procedure different from the
standard procedure used with other conditioned stimuli; such a change in the
experimental procedure was also a change in the situation and this resulted in
obtaining a different kind of behavior. This study further supports the idea
that changes in the situation may result in changes in the conditioned
reaction.
Considering the obtained data, a question may be asked— what is the role of a
single stimulus in the process of instrumental conditioning? Can it retain its
acquired property of evoking the instrumental reaction when it is used in an
environment different than that in which it was originally trained? In order to
answer this question, another study was undertaken, described as follows.
(aCR) to stimulus aCS (the sound of a flute) for food reinforcement. In the
same chamber, each dog in the defensive group was trained to lift the right
hindleg (dCR) to stimulus dCS (rhythmic sound of a whistle) to avoid an
electric shock to skin. Each dog of either alimentary or defensive group was
trained separately. The training was conducted daily with 8-10 trials per
session.
This test situation was a large empty room with no furniture except for a
small table and a chair for the experimenter. The dog was allowed to move
around freely. Neither conditioned nor unconditioned stimuli had been used
here before.
In the alimentary group of 5 dogs, first a control neutral stimulus (nS), the
sound of a rattle (never used before with these dogs) was applied for 15
seconds. This stimulus evoked only a slight orienting reaction. A few minutes
later, the alimentary test stimulus (taCS), the sound of a flute (to which the
alimentary instrumental CR was established in the uE before) was applied for
15 seconds. In all dogs, the taCS evoked only a motor reaction such as
approaching the flute and sniffing it, or coming to the experimenter and
placing the forepaw on her knees. However, the instrumental reaction
previously trained to this stimulus in the uE was not performed.
In the defensive group of dogs, first a control nS, the sound of a rattle (never
used with these dogs before), was applied tor 15 seconds, evoking only a slight
orienting reaction. Then the test CS (sound of whistle in two dogs, and
flashing light in two other dogs) to which the defensive CR was established in
the usual situation before, was applied for another 15 seconds. It was found
that the test stimulus produced general anxious movements in-
46 Conditioning
eluding shaking head (in case when electric shock to the ear was used as
reinforcement in the uE). Nevertheless, the previously trained avoidance
reaction (dCR) was not performed.
In this series, the test situation was the same large empty room that was used
in the neutral test situation. However, the reinforcing stimuli (food for the
alimentary group, and electric shock for the defensive group), were given in
this situation previously.
In the alimentary group, the test was carried out on the five dogs previously
used in the neutral test situation. In the first two sessions each dog was
allowed to move freely and received a piece of meat every 20-30 seconds.
Food was either thrown on the floor (for 2 dogs) or was offered in a bowl
placed on a wooden beam lying along the wall of the room (for the other 3
dogs). Then during the third session, a new control stimulus, a certain type of
noise, was applied for 15 seconds, producing no distinct reaction. One minute
later, a conditioned stimulus, the sound of a flute (previously used in the
training in the usual situation), was applied for 15 seconds. It turned out that
the instrumental reaction previously trained in the uE was not performed by
either of the two dogs which obtained food on the floor; one of them came to
the experimenter and performed a “begging” movement. On the other hand,
the three other dogs that obtained food in the bowl placed on a wooden beam
did perform the aCR (placing their forepaw on the beam), previously trained
in the uE.
In four dogs of the defensive group, first the control stimulus (the sound of a
rattle) was applied for 15 seconds producing no distinct reaction. Then, one
and a half minutes later, the test conditioned stimulus (flashing light for two
dogs, and the sound of a whistle for two other dogs) was applied for 15
seconds. The avoid-
ance reaction, previously trained in the uE, was not performed by any of the
dogs, but shaking the head and anxious movements were observed. Then, an
electric shock was given and the procedure was repeated. As before, there was
not a clear reaction to the control stimulus (the rattle), but the application of
the test stimulus elicited touching the ear (to which the shock was previously
given in the uE) in one dog. only when, in addition, a band had been fastened
on his leg, did this dog perform the previously trained movement of lifting
this leg to the test stimulus. No previously trained reaction was performed by
another dog, even when the band was fastened to its leg. However, two other
dogs did perform the previously trained reaction when the test was preceded
by an electric shock.
These experiments showed that the previously used conditioned stimulus
elicited only a general defensive reaction but not the instrumental reaction
previously trained in theusual situation. Only when some facilitating
elements were present, such as a beam (for the alimentary group) or an
electric shock (for the defensive group) would the previously trained CR
appear.
In two dogs the alimentary reaction of lifting the right hindleg to the beating
of the metronome was trained in the usual situation (uE). On the same days,
another alimentary reaction of lifting the left foreleg in one of these dogs, and
begging movement in the other dog, to the tactile stimulus, were trained in
the test situation. When the metronome beating was applied in the test
situation, none of these dogs performed the reaction previously trained in the
usual situation, i.e., lifting the right hindleg. Instead, both dogs performed the
movement trained in the test situation (i.e., lifting the right foreleg in one dog
and begging movement in the other dog).
Two dogs of the defensive group were trained to lift their right hindleg to the
flashing light in the usual situation, and to stand on hindlegs to the sound of
a buzzer in the test situation; both movements secured the animal against the
electric shock applied to the
48 Conditioning
left ear. Then each conditioned stimulus was tested in the other situation. It
was found that neither stimulus evoked the instrumental reaction which had
been trained to it before in the uE. Instead, it elicited the reaction which had
been trained to a different stimulus in the test situation, i.e., standing on
hindlegs to the sound of a buzzer in one dog, and lifting the hindleg to the
flashing light in the other.
In three dogs, the basic training was conducted in two different situations,
one with alimentary reinforcement (situation A) and the other with defensive
reinforcement (situation D). Situation A was a usual Pavlovian chamber.
Situation D was arranged on the floor in an empty part of a room located in
another building; the experimental part of situation D was separated from the
observation section with a wooden partition. In both situations, the recording
bands were fastened on all four legs of the dogs.
In situation A, the sound of a metronome for two dogs and a moving black
disc for the third dog were used as alimentary conditioned stimuli; the
instrumental reaction was placing the right foreleg on the feeder platform
(aCR). In situation D, the sound of a whistle was used as a defensive CS; the
instrumental reaction was lifting the right hindleg to avoid a shock to the
same leg (dCR). The training was conducted in alternate series each lasting
for several days in each situation. Then the test was performed in which the
defensive CS was applied in the alimentary situation, and the
TABLE 5.1
TABLE 5.2
When the defensive CS was applied in the alimentary situation for the first
time, one dog performed a slight but distinct lifting ol
50 Conditioning
the right foreleg (defensive CR). Another dog reacted to the defensive CS with
a slight lifting of the right foreleg (partial aCR), followed by shivering,
bristling of hair, and anxious behavior. No clear reaction to defensive CS was
seen in the third dog. When the defensive CS was repeated in the next trial,
all three dogs performed the lifting of the right hindleg (dCR). When a control
stimulus, the sound of bubbling water (never used with these dogs before)
was applied, a slight orienting reaction only was observed in one dog, while
two other dogs performed an alimentary CR. When the control stimulus was
repeated, all three dogs performed the alimentary CR.
The first application of the alimentary CS in the defensive situation did not
produce any motor reaction in two dogs, but the third dog performed 3 slight
movements of the right hindleg (i.e., the defensive movements). The
repetition of the alimentary stimulus in this situation again did not produce
any effect in the first two dogs. The third dog, however, first lifted its right
hindleg and then its right foreleg and held it in this position for several
seconds; this resembled the aCR trained in the alimentary situation.
Detour Behavior
51
52 Conditioning
The problem of detour behavior was also studied by Beritashvili and his
associates (Beritashvili 1941, Beritashvili and Akhmeteli 1941) who conducted
their observations on dogs, rabbits, hens, and pigeons. The results of their
studies led them to a conclusion that the animal performs the detour
behavior due to a psycho-neural process which reproduces a picture of the
external world in the animal’s mind thus guiding its movements towards the
food. Later, Roginski and Tikh (1956) studied detour behavior on chickens,
rats, and monkeys, and found that monkeys learned to make the detour
fastest, and chickens most slowly; they concluded, therefore, that the detour
reaction depends on the phylogenetic level of the animal. Pavlov (1949),
however, expressed the opinion that the detour behavior has an acquired,
conditioned reflex characteristic.
In the preliminary testing period of this study it was found that adult dogs
easily made a detour when food was placed behind a visible obstacle. A
question arose, therefore, as to whether this reaction was inborn or acquired
by experience. In order to answer this question, a series of experiments was
performed on puppies that had been kept in a controlled environment from
birth and never had any occasion to make a detour before.
The study was conducted on 28 puppies, male and female, 12-131 day old, of 8
different litters from 5 female and 8 male dogs. Mother and puppies of each
litter were housed in a large pen, 2x3.5 m at the bottom and about 1.5 m in
height. Their home compartment was empty and the pups had no occasion to
go around any object. They were first nursed by the mother, and later they
were fed solid food (mixture of cooked meat and cereal) with the mother.
The tests were carried out separately for each pup, in a large, empty room
(different from the pen). The experimental space was partially divided by a
wire-net partition as schematically shown in fig. 6.1.
FIGURE 6.1
Samples of detour behavior of puppies (1-4 months old) that were controlled
from birth and never had occasion to make the detour before. (Photocopies of
sketches of detour behavior of single puppies, made by the experimenter
during the test session).
Straight line, wood partition; wavy line, wire-net partition; square at the
bottom of each sketch, starting place; empty circle at the front of wire-net,
site of the bowl and milk during preparatory sessions; circle with a cross, site
of the bowl behind the wire-net partition during the test session; the broken
line with small arrows traces of detour behavior.
54 Conditioning
net partition (similar to that used in the room) where they were allowed to
run both in front of and behind the wire-net.
It turned out that most of the 12 pups of the group that had been allowed to
walk in both parts of the experimental compartment before the test easily
found the way to the food in seconds; only a few pups in this group needed
more time to make a detour. Similar results were obtained on 5 pups that
were allowed to run both in front of and behind the partition in their
preliminary training in the garden.
On the other hand, none of the 11 pups that had been allowed to remain only
in front of the partition before the test could find the way to the food.
Independent of age, they only repetitively scratched the partition at the site
where the food was visible behind the wire-netting. This behavior was
observed for several consecutive sessions until finally, earlier or later, each
pup started to walk back and forth alongside the net, and accidentally found
the passage to the bowl (see a schematic pathway of a pup in fig. 6.1). On an
average, 4-6 sessions were needed for each pup to learn the detour. It is
interesting that age did not influence the response. When the test was
conducted on 4-month-old pups, their learning of the detour took even longer
than learning in the 2-month-old pups.
Once the pup learned to make a detour in a room, it was able to make it also
in a different surrounding, namely in the garden with the use of the same (or
similar) partition and the same procedure. This means that the situational
stimuli were recognized by the pup as the same as those used in the room
before, and thus, the detour reaction was performed.
In this case, both the stimulus amid the reaction are complex. The stimulus is
composed of permanent situational elements such as visual stimuli of the
wire-net partition, the bowl, and the passage. The motor reaction involves
both excitatory and inhibitory processes. First, the initial motor reaction to
the bowl (scratching the partition in front of the animal in an attempt to
reach the food behind the partition) had to be fully extinguished, i.e.,
inhibited by not obtaining the reinforcement (food). Then only the correct
motor reaction, a U-turn, rewarded by food, can gradually develop
(fig6 1 )- . u u ,
Of course, there are more instances of behavior where both stimuli and
reactions are even more complex. The manipulatory reactions of animals of
various species described by Thorndike (1911) and chain reflexes analyzed by
Skinner (Skinner 1938, Ferster and Skinner 1957) may serve as examples of
complex conditioned behavior. The directional instrumental reactions studied
by Lawicka (1959) and Konorski and Lawicka (1959) also belong
to this category.
56 Conditioning
FIGURE 6.2
The second part of the feeding behavior is the act of eating itself. A closer look
at this act shows that it is quite complex and dependent on the kind of food to
be consumed. In mammals, the first feeding behavior is sucking. The fact that
it appears early after birth indicates that sucking behavior is inborn (although
it is not excluded that it can occur even before birth in the womb by sucking
the surrounding fluids). Later in life, the young start to follow the mother or
other adults of their colony in seeking food and eating that food (Galef 1977,
1978; Galef and Clark 1972). This suggests that the motor ability to eat
various kinds of food (other than mother’s milk) is acquired by experience
later in life. This ability is based on a number of inborn reflexes, such as
mastication, i.e., the movements of opening (digastric reflex) and closing
(masseteric reflex) the mouth (Clemente et al. 1966; Chase and McGinty 1970
a,b; Chase and Babb 1973) as well as the movements of the tongue and other
parts of the mouth cavity. The motor performance may vary depending on the
kind of food to be eaten. The motor (instrumental) reaction of eating a piece
of meat is different than the motor reaction of eating creamy pudding or
drinking milk. Due to the repetitive experience with various kinds of food, the
act of eating gradually becomes more and more com-
58 Conditioning
plex. Even later in life some further adjustments in the act of eating may take
place when the unknown kinds of food are being offered. Therefore, the food,
which is commonly called an “unconditioned stimulus,” turns out to be a
specific conditioned stimulus , and the act of eating becomes a conditioned
instrumental reaction. In other words, the alimentary instrumental
conditioned reaction is reinforced not by food as an unconditioned stimulus
but by food as a conditioned stimulus when it is seen or sniffed by the
subject. In this case, only the final effect of eating, the sensory satisfaction
resulting from food consumption, can be considered the “reward” or the
“reinforcement” (Wyrwicka 1975).
A question can be asked whether or not the act of eating can depend on the
visual or olfactory stimuli related to a particular portion of food,
independently of the situation where this food has been offered. It seems that
environmental factors may still influence the reaction of eating. One of these
factors is the state of satiation. In this case, the act of eating may not occur. In
another case, the unsatisfactory environment such as a dirty tablecloth or the
presence of a constant loud noise, etc., may inhibit the consumption. There
are many such factors restraining eating.
On the other hand, there are also factors that facilitate eating. An example is
the already quoted study by Wyrwicka and Chase (1972) in which cats were
offered food in two containers; eating from one of these containers was
rewarded by desirable hypothalamic stimulation, while eating from the other
container was not. The cats chose to eat only from the container where they
obtained the pleasant stimulation.
Experiments with completely satiated goats which refused to eat any more
can serve as another example. These animals, however, started to eat again
when they obtained a pleasant light electrical stimulation in the “feeding
center” in the lateral hypothalamus each time they ate, even when this was
leading to overeating (Wyrwicka et al. 1959, 1960, 1982, 1988).
Presleep Behavior
Internal Organs
Intensive studies were undertaken by N.E. Miller and his associates on the
possibility of producing instrumental conditioning related to internal organs
innervated by the autonomic nervous system. Miller and DiCara (1967)
demonstrated that rats immobilized by an injection of curara and artificially
respirated learned either to increase or decrease their heart rate (dependent
of the experimental design) to obtain the rewarding electrical stimulation in
the basal forebrain bundle (cf. Olds and Milner 1954; Olds 1962). In other
experiments, the curarized rats learned to increase their heart rate (or
decrease it, dependent on the chosen experimental procedure) to avoid an
electric shock to the skin (DiCara and Miller 1968a). Similar results were
reported by Throwhill (1967).
Using the same method, it was possible to change the vasomotor activity, i.e.,
to obtain either vasoconstriction or vasodilation, when the required activity
was reinforced by desired electrical stimulation in the basal forebrain bundle
(DiCara and Miller 1968b). Another study showed that thirsty, noncurarized
dogs learned to increase salivation when rewarded with access to water; it was
observed that salivation in these dogs increased with the training (Miller and
Carmona 1967). It was also possible to increase the rate of urine formation in
one group and decrease it in another group of curarized rats when electrical
stimulation in the basal forebrain bundle was offered as a reward (Miller and
DiCara (1968).
61
62 Conditioning
vous system. In order to answer this question, a high dose of curara (4.8.
mg/kg over 3 hours) was used, resulting in full stoppage of muscular activity;
in spite of complete immobilization of the animal, rats still learned to change
their heart rate (DiCara and Miller 1968c).
Several studies have shown that electrical activity of the brain (EEG) can be
conditioned, i.e., it becomes an instrumental conditioned reaction. Attempts
of conditioning of the electroencephalic function of the brain were made by
various investigators. One such study was that of Olds and Olds (1961), who
recorded unitary discharges from paleocortical structures of the rat’s brain.
Each time the number of discharges from the unit increased, the rewarding
electrical stimulation of the medial forebrain bundle was applied. As a result,
the number of discharges became considerably greater.
EEG arousal. Each time the EEG reaction occurred to the tone, the shock was
withheld. After a number of repetitions of this procedure, EEG arousal began
to appear in each trial while the cat remained lying down and apparently
asleep as shown by electromyographic recording (EMG) from the animal’s
neck muscles. In this experiment desynchronization of the EEG pattern
became an instrumental avoidance reaction.
64 Conditioning
SMR burst appeared more often and at regular intervals (Wyrwicka and
Sterman 1968). When a milk reward was withheld during the extinction
procedure, the frequency of the SMR bursts temporarily increased at the
beginning of extinction. The finding of a temporary increase of the
instrumental reaction, which is usually observed at the beginning of the
extinction process (see Konorski 1948,p. 217; 1967, p. 361), supports the view
that the sensorimotor rhythm became conditioned as an instrumental type
reaction.
Further research focused on precise localization of the brain region where the
sensorimotor rhythm is initiated. Research of Sterman and Clemente (1962),
Clemente and Sterman (1963, 1967), Clemente et al. (1963), Sterman and
Wyrwicka (1967), Sterman et al. 1969) revealed that this region is situated in
the basal forebrain area (BFA). According to the stereotaxic atlas of the cat’s
brain of Jasper and Ajmone- Marsan (1960), the coordinates for this area are
A 13.0-15.0, L 2.5-3.0, H—4.0, although some differences from cat to cat may
exist. Bilateral lesions in the basal forebrain area in cats resulted in
suppression of sleep (McGinty and Sterman 1968).
In experiments by Babb and Chase (1974), cats were trained to produce the
sensorimotor rhythm (SMR) while the masseteric and
66 Conditioning
In a special study of Chase and Harper (1971), cats were trained to produce a
synchronized EEG pattern of 12-14 cps of sensorimotor activity to get a milk
reward. During the presence of SMR, somatic motor activity was inhibited; at
the same time, heart rate decreased, and the respiratory pattern became more
regular than during control periods. In a later study, neurons in the thalamus
were found to discharge in a “burst- pause” manner during SMR, similar to
that found in quiet sleep (Harper 1973, Harper and Sterman 1972). These data
suggest that the sensorimotor rhythm is related to motor inhibition.
67
68 Conditioning
in the previous chapter demonstrate that with the use of a specific reward, it
is possible to obtain the required EEG pattern both in animal and human
subjects (Olds and Olds 1961, Miller 1969 / experiments of Carmona/, Kamiya
1962, Wyrwicka 1964, and others).
The finding of the inhibitory character of the sensorimotor rhythm led to the
question of whether it would be possible to use the SMR as a natural
treatment in conditions involving disregulation of the motor system in
epilepsy in humans.
dents were limited only to some small movements such as wrinkling of the
brow associated with the left lateral deviation of the eyes, crossing of the right
arm to the left knee, and falling to the left. The majority of seizure incidents
were currently nocturnal, occurring in the later portion of a night s sleep.
The frequency of incidents varied from 2 to 4 per month at the onset to 1 per
3 months one year later. During the 12-month period immediately preceding
the treatment, seizures were present irregularly at an average rate of about 2
per month. The subject had variable success in using a combination of drugs
such as Dilantin, Mysoline, Peganone, Dianox, and Ritalin. In the preceding
12 months she also daily used Dilantin and Mebarol.
The training techniques included the EEG recording from the scalp
sensorimotor and occipital areas, using needle electrodes. Placement of
electrodes was standardized by reference to the international (10-20) system.
The electrocardiogram (EKG) and electromyogram (EMG) were also
recorded. During the sessions the patient was sitting comfortably on a
reclining chair. The instructions were delivered through standard tape
recording. The device recording the SMR activity (feedback) consisted of a
rectangular unit suspended from the ceiling several feet above and in front of
the patient. The unit contained two rows of ten small lamps covered with
transparent colored Plexiglas. When the EEG activity of appropriate
amplitude and duration appeared, the lamps in the top row were successively
lighted; each advance was accompanied by the sounding of a single chime
which was used as a reward. With the eleventh successful EEG response the
lights of the top row were extinguished and the lamps of the bottom row were
activated; each time this was accompanied by a double chime reward. Usually,
200 to 300 rewards were obtained during a single session lasting from 30
minutes to one hour. The sessions were conducted once a week during the
first month and then twice during the following three months.
The results of this treatment were dramatic. Initially, the patient showed a
moderately low level of SMR feedback. Later, however, after adjustment of
the reward from 12-14 cycles/sec (c/sec) to 11-13 c/sec, a gradual increase in
the production of SMR feed-
70 Conditioning
back was observed. After three sessions there were no seizures for the
following period of 3 months, except one mild nocturnal seizure. At the same
time, some changes occurred in the personality of this patient. Being
previously a quiet and rather timid, non-assertive individual, the patient
progressively became more outgoing, with personal, confidence and even an
enhanced interest in her appearance. She also reported a shorted latency to
sleep onset, and a more rapid awakening in the morning. Moreover, this
patient voluntarily ceased her previous practice of supplementing
anticonvulsant medication while experiencing an aura, without seizure, and
reported a reduction in such occurrences. In a later phase of the study she
obtained an additional small monetary reward for producing SMR; this
resulted in the highest rate of performance she had ever showed.
In order to obtain more data, the following study was undertaken on 8 poorly
controlled seizure patients by Sterman and MacDonald (1978). Four of these
patients were rewarded alternately for facilitation and suppression of 12-15
c/sec SMR activity. All these patients showed a seizure reduction only during
positive reward tor the 12—15 c/sec pattern. Four other patients were
rewarded for the facilitation and then suppression of 18-23 c/sec rhythm.
Three patients of this group showed significant seizure reduction.
age from 18 to 35 years, and their history included diagnosed epilepsy from 8
to 24 years. The techniques used for this group included not only the training
with the use of various frequencies of waves but also sleep EEG spectra,
clinical EEG and anticonvulsant blood levels. The training was conducted
mainly in the patient’s home and lasted 3 months.
72 Conditioning
Theoretical Comments
73
74 Conditioning
The experimental data show that a single conditioned stimulus produces the
definite instrumental CR only in the situation in which it was trained. Once
the situation has been changed, the conditioned stimulus is not able to
produce the correct instrumental conditioned reaction. Sometimes, a
diminished and modified reaction may be present, hardly resembling the
correct CR.
Theoretical Comments 75
The above described data support a view that the process of conditioning
involves both the intermittent conditioned stimulus and the situation in
which this stimulus appears. However, due to the experimental procedure
which consists of giving the reinforcement only to the intermittent CS and
never to the permanent complex of situational stimuli, the reaction to the
situation alone ceases to appear. This is a partial inhibition of the conditioned
reflex to the situation. In other words, a differentiation takes place between
the intermittent CS+situation and the situation alone.
This does not mean that the CR to the situation alone completely disappears.
It may still reappear in various circumstances, such as a sudden disturbance
of the usual experimental procedure, etc. Moreover, the previously described
experiments showed that the conditioned stimulus retains its acquired
property of producing the CR only in the situation in which the training took
place. When the CS was applied in a different situation, the CR was absent.
This was especially evident in the case of instrumental conditioned reflexes.
76 Conditioning
However, the problem of the situation persisted and in many cases seemed to
be the decisive factor in behavior. One such instance is the study of Denisov
and Kupalov (1933, Kupalov 1961), described in chapter 4. Briefly, in
experiments on dogs, two kinds of illumination of the experimental chamber
were used, full illumination in some sessions, and dim illumination in other
sessions. When the conditioned reflex was trained in full light and then tested
in dim light, its value remained at a high level during several successive
sessions and only then decreased. Vice versa, after training in dim light with
reflex testing in full light, its value remained low for the next several sessions.
Kupalov (1961) suggested that this was an effect of conditioning to the
experimental situation. According to Kupalov, however, the situation is not
able to evoke the conditionwed reaction but can change the tonus of some
parts of the brain. Therefore, he called the reflex to situation a shortened or
truncated conditioned reflex.
Theoretical Comments 77
duces another instrumental CR, that which had been trained in this new
situation. This phenomenon leads to the conclusion that, in fact, the
situational complex of stimuli dominates over the intermittent stimulus.
78 Conditioning
Theoretical Comments 79
Let us add a few words of explanation of the term internal situation. It can be
understood as a complex of memories of previous experiences of the
organism. These memories may serve as an internal background for the final
reaction. However, which of these memories are used as the main factors for
the reaction has to be
'
Chapter 2 deals with the motor conditioned behavior, which was not included
in Pavlov’s theory. According to Pavlov, the performance of a movement
produces sensory impulses analogous to those produced by visual or acoustic
stimuli. When these impulses are followed by an unconditioned stimulus
(e.g., food in the mouth) they become a conditioned stimulus (CS). Pavlov,
however, did
81
82 Conditioning
The observations of Konorski and Miller include the fact that the trained CRII
appears not only to the intermittent conditioned stimulus (CS), but also to
the situation in which it is applied, in the absence of the CS.
Chapter 4 describes various studies on rats in which the situation turned out
to be a potent factor producing CRs to some situational elements, such as the
neuroleptic effects in the front of the room where injections of
chlorpromazine were previously given (but not in the front of other rooms).
Similarly, a specific conditioned circling was observed in the environment
where apomor-phine injections were previously given. Chapter 4 also
includes a description of the situational effects on the behavior of human
84 Conditioning
This includes the case of heterogeneous CRs. When the CS, originally trained
to produce a defensive CR, was tested in the alimentary situation (in which
alimentary CRs were trained to different CSs) it evoked some general
defensive behavior (such as escape) but not the previously trained
instrumental CR. The same was observed when an alimentary CS was tested
in a defensive situation: it evoked a general alimentary posture such as
begging, but not the previously acquired CR. However, an introduction of
elements from the original situation (such as an object resembling the
feeder) led to the performance of the correct CR. These observations show
that the intermittent stimulus may retain its ability to evoke the acquired
instrumental CR only in the situation in which it was originally trained (see
Table 5.2).
Conclusions
Pavlov’s theory, however, has left some questions unanswered, one of which
is the role of the environmental background or situation in which the
conditioned reflex (CR) occurs.
Each change in the situation can alter the performance of the CR.
The CR can be established not only to an intermittent stimulus but also to the
complex of permanent situational stimuli. An example of CR to the situation
is the appearance of the intertrial CRs, observed often in instrumental CR
studies. These CRs to the situation are usually eliminated by non-reintorcing
them.
86 Conditioning
The “truncated” conditioned reflex is nothing else than a conditioned reflex to
the situation, partly inhibited by non-reinforcing it.
The examples described in this book are only a few selected cases showing the
crucial role of the situation in behavior. More studies on this topic are
needed.
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References 95
Babb, M., 57, 65 Beritashvili, J.S., 41, 52, 78 Black, A.H., 62, 70 Bogen, K.M., 9
Bond, I.K., 65 Brobeck, J.R., 25 Brown, B.B., 63 Buffy, A., 37 Bykov, K.M., 9, 56
Chase, M.H., 57, 58, 65, 66. 75, 76 Clark, M.M., 57 Clemente, C.D., 57, 59, 64
Corson, S.A., 36 Cott, A., 70
Davis, E.G., 65 Descartes, R., 5 Di Cara, L.V., 61, 67 Donhoffer, H., 65 Doty,
R.W., 32 Eikelboom, 36 Ellertsen, B., 71 Evans, D.R., 65
Ferster, C.B., 18, 19, 55 Finch, C.A., 70 Finley, W.W., 70, 71 Fisher, W., 51
Fonberg, E., 24 Fox, S.S., 63 Friar, L., 68
Izquierdo, I., 62
John, E.R., 65
Kamiya, J., 63, 68 Kaplan, B.J., 70, 71 Kimble, G.A., 14 Kimmel, H.D. 28
Khananashvili, M.M., 35 Klove, H., 71 Knauss, T.K., 57
Konorski, J., 2, 6. 10, 11, 12, 13, 14, 15, 17, 18, 21, 22, 31, 32, 33, 36, 43,55,64,78
Kozak, W., 43-44 Kohler, W., 51 Krasnogorski, N.I., 11, 15 Krulikowski, D., 91
(first author L.F.Lubar)
98 Conditioning
Murrin, M.R., 28
Pavlov, I.P., 2, 5, 6, 7, 8, 9, 10, 11, 12, 13, 14,41,52 Poser, E.G., 65 Poulos, C.X.,
35 Pressman, cit. by Asratyan, 28
Sachs, 65
Sterman, M.B., 13, 57, 65, 66, 68, 70 Stewart, J., 36 Stroganov, E.G., 41
Struchkov, M.I., cit. by Asratyan Szwejkowska, G., 33, 35
Thorndike, E.L., 17, 18, 51, 55 Throwhill, J.A., 61 Teyrowski, V., 51 Tikh, N.A.,
52 Thorpe, W.H., 51-52 Tolman, E.C., 51,
Ungerstedt, U., 37
Zamyatina, 29
Subject Index
Hypothalamus, 56
100 Conditioning
Releasing stimuli, 24
one situation, 41-42; two similar situations, 42; two different situations, 42-
43
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