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Species-specific defense reactions and avoidance learning: An evaluative review

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 Species-Specific Defense Reactions a n d
Avoidance Learning
An Evaluative Review
MARY CRAWFORD, PH.D., AND F R E D A . M A S T E R S O N , P H . D .
West Chester State College
West Chester, Pennsylvania
and University of Delaware
Newark, Delaware
Abstract--Bolles (1970) proposed a theory of avoidance learning, the species-specific
defense reaction (SSDR) hypothesis, which emphasized innate constraints on the re-
sponse repertoire of rats in aversive situations and minimized the role of reinforcement in
avoidance learning. The present paper describes Bolles' (1970, 1971, 1972, 1975, 1978)
development of SSDR theory and reviews the empirical tests of its assumptions and
predictions. It is concluded that the SSDRs described by Bolles, along with some others,
are highly probable in aversive situations but that the response repertoire is not limited to
them. Further, there is strong evidence for reinforcement effects in the establishment and
maintenance of at least some avoidance responses.

BOLLES' (1970) species-specific defense reac-
tion (SSDR) theory was a major departure from
p r e v i o u s a p p r o a c h e s to a v o i d a n c e learning.
F r o m its first v e r s i o n on, the t h e o r y has
minimized or even denied the role of reinforce-
ment in the acquisition of avoidance responses,
relying, instead, on the selective elicitation of
innate defense reactions. In its emphasis on in-
nate reactions, SSDR theory is one of the first
learning theories to incorporate the notion of
biological constraints on learning. In its deem-
phasis o f reinforcement, the theory presents a
highly parsimonious account of avoidance learn-
ing. A critic might call it too parsimonious, but
the critic would then be required to demonstrate
the necessity for a greater emphasis on rein-
forcement. Thus, SSDR theory has taken its
place beside two-factor theory (Mowrer 1960,
Rescorla and Solomon 1967) as another perspec-
tive which must be contrasted with any newer
approach.
The SSDR theory of avoidance behavior was
the first to focus on the animal itself. In his origi-
nal statement of the SSDR hypothesis, Bolles
(1970) reminded learning theorists that laboratory
avoidance [earning tasks have often been used to
"'demonstrate" how animals could survive in the
wild. Instead, he proposed, theorists should look
at the rat as a small mammal that must defend
itself in natural situations. It seemed logical that
the rat would have evolved a repertoire of defen-
sive behaviors which would strongly influence its
behavior in the laboratory. Thus, Bolles pro-
posed that, rather than attempt to explain the
rat's survival in its natural habitat through exam-
ples taken from its behavior in the laboratory, we
might better attempt to explain the puzzling in-
consistencies in its laboratory behavior through
an examination of its natural defensive reactions.
The purpose of this paper is to review the em-
pirical precedents for SSDR theory, the de-
v e l o p m e n t o f the t h e o r y , and the result o f
subsequent direct empirical tests of the theory.

The writing of this paper was supported in part by a
Faculty Equivalent Time Award from West Chester
State College to M. Crawford.
The authors are grateful to R. C. Bolles for comments
on an earlier version and to D. Herrmann for providing
resources and support for research while the first au-
thor was in residence at Hamilton College.
Address reprint requests to: Mary Crawford, De-
partment of Psychology, West Chester State College,
West Chester, Pennsylvania 19380.
Empirical Basis for the Theory
Years before Bolles introduced the SSDR
hypothesis, each of the most influential theories
then c u r r e n t had left i m p o r t a n t p r o b l e m s
uresolved (for reviews, see Bolles 1972b, Selig-
man and Johnston 1973, Mineka 1979). Out of
these, one problem emerged as the most severe:
the typical rate of avoidance learning of the labo-

0093-2213/82/1000/0204/$01.35 O J. B. Lippincott Company

204
Volume 17
Number 4
SSDRs AND AVOIDANCE
ratory rat varies greatly depending on what be-
havior the experimenter selects as avoidance
r e s p o n s e - - w h a t Bolles (1970) called the re-
sponse problem.
The r e s p o n s e p r o b l e m c o m p l i c a t e d the
analysis of avoidance behavior in terms of con-
trolling contingencies, an approach initiated by
Kamin (1956), who studied the relative contribu-
tion of warning signal (WS) offset and shock
avoidance to learning of a shuttle avoidance task.
The response problem repeatedly surfaced in
contingency analysis studies. For example, Bolles
and Seelbach (1964) studied the effect of escape
and punishment contingencies on simple, high-
operant-level responses and found that some re-
sponses, but not others, could be brought under
aversive control. Mogenson, Mullin, and Clark
(1965) found that delaying WS offset (which
Kamin had shown to affect shuttle avoidance)
had little effect on wheel-running or wheel-
turning avoidance responses. Bolles, Stokes, and
Younger (1966) examined the importance of WS
offset, shock escape, and shock avoidance to
avoidance learning in both the shuttlebox and the
running wheel, thus replicating and extending
Kamin's (1956) study, and found that the relative
contributions of the three contingencies vary
with the required response. BoUes (1969) found
differential learnability of turning, rearing, and
running responses in a wheel and differential im-
portance for the shock escape contingency in
their acquisition. BoUes and Tuttle (1967) demon-
strated a failure of WS offset to reinforce the
learning of a new instrumental response in the
absence of shock. Several experimenters (e.g.,
Masterson 1970, Meyer, Cho, and Wesemann
1960) f o u n d l i t t l e l e a r n i n g of a b a r - p r e s s
avoidance response with the WS offset con-
tingency as nominal reinforcer.
M e a n w h i l e , it was b e c o m i n g a p p a r e n t to
avoidance learning experimenters that a clear
continuum of task difficulty existed for the labo-
ratory rat. At one extreme is the rapid, certain
learning of one-way running or jumping responses
( M a a t s c h 1959, T h e i o s and D u n a w a y 1964,
Theios, Lynch, and Lowe 1966). At the other
extreme is the erratic, very slow and incomplete
learning of bar press or wheel-turn-
m a n i p u l a n d u m a v o i d a n c e r e s p o n d i n g with
s t a n d a r d c o n t i n g e n c i e s ( M e y e r , Cho, a n d
Wesemann 1960, D ' A m a t o and Schiff 1964, Mas-
terson 1970, Coons, Anderson, and Myers 1960).
Responses such as shuttlebox running (Theios
and Dunaway 1964, Theios et al. 1966) and wheel
running (BoUes et al. 1966) are intermediate in
difficulty. This continuum of difficulty persists
over a wide variety of task parameters. Its impact
has been nicely summed up by Bolles (1978,
p. 90):
205
The differences between the learning curves obtained
in different situations are really quite remarkable.
These differences dwarf the effects of the experimen-
tal variables usually manipulated in avoidance-
learning experiments. Thus, white parameters such
as the properties of the shock, the properties of the
warning signal, the kind of rat, or various experimen-
tal contingencies can be shown to have reliable ef-
fects on performance in this or that situation, all such
effects are rather puny compared with the enormous
differences that are typically found between different
situations. I suspect that if all the avoidance-learning
data ever reported could be combined and subjected
to a gigantic analysis of variance, we would find that
about 80% of the total variance would be attributable
to situational factors, and only about 10% could be
attributed to all the other parameters that are usually
considered to be of theoretical interest. (The remain-
ing 10% would be error variance.) The existence of
these great situational differences is, I repeat, the
most salient fact about avoidance learning.
The growing recognition of a response variable
unaccounted for by any theory, plus the interac-
tion of this response variable with contingencies
such as WS offset and shock escape, increasingly
demanded a new approach. Much of the analysis
of the response problem and its interaction with
programed contingencies had been done by Bol-
les and his students. For example, it was BoUes
(1969) who reanalyzed the classic experiment of
M o w r e r and Lamoreaux (1946), long cited in
support of a contribution of the escape con-
tingency to avoidance learning, and provided
evidence that the importance of this contingency
depends on the required avoidance response. It
was Bolles (1970) who first summarized the evi-
dence and concluded that the WS offset and es-
c a p e c o n t i n g e n c i e s , long c o n s i d e r e d crucial
sources of reinforcement, could not alone explain
the data. And it was BoUes (1970) who formulated
the first theory of avoidance learning that recog-
nized and attempted to solve those problems. The
SSDR hypothesis was a much-needed alternative
to earlier theories, a fresh approach to avoidance
learning. We turn now to the development of the
SSDR theory.
Development of SSDR Theory
Original 1970 Statement
The hypothesis that Bolles (1970) originated
was that fear absolutely limits the rat's behavioral
repertoire to a small set of SSDRs--freezing,
fleeing, and fighting. Avoidance learning could
occur rapidly only if the required response were
one of the rat's SSDRs. In this case, the mecha-
nism underlying learning was seen as the sup-
pression of inappropriate SSDRs through
punishment. In other words, an SSDR that suc-
cessfully avoided shock would become more
206 CRAWFORD AND
probable, not because it was reinforced, but be-
cause less successful SSDRs would drop out. In
the original version of SSDR theory, rapid acquis-
ition of avoidance responses is attributed solely
to punishment, with positive reinforcement play-
ing no role.
In addition to the above described rapid learn-
ing of SSDRs, BoUes proposed a mechanism
whereby non-SSDRs could be learned slowly.
The idea was that certain events become con-
ditioned as safety signals which tend to alleviate
fear and thus partially remove the restriction that
all behaviors be SSDRs. WS offset, the onset of
an avoidance-response-contingent feedback sig-
nal (FS), and proprioceptive stimuli generated by
an avoidance response all predict absence of
shock and hence should become Pavlovian in-
hibitors of fear. As fear is thus inhibited, the
restriction of behaviors to SSDRs becomes more
and more relaxed, until non-SSDRs such as
lever-pressing can occur. In addition, Bolles
raises the possibility that these safety signals
might reinforce slowly learned avoidance re-
sponses. In effect, Bolles has preserved a mod-
ified form of two factor theory to explain the slow
learning of non-SSDRs.
The 1970 model also entertained the notion
that an SSDR must be functionally effective:
The critical feature of jumping as a flight response
appears to be whether it is functionally effective in
the sense that it actually makes flight possible. The
possibility of flight appears to be much more impor-
tant in establishing a flight response than either its
topographical features or even whether it is effective
in avoiding shock [Bolles 1970, p. 35].
.... Running will not be acquired as an R
(avoidance response), at least not very readily, un-
less the running response is effective for flight, that
is, effective in the functional sense that it takes the rat
out of the situation [Bolles 1970, p. 35].
Bolles' 1970 paper did not explicitly link func-
tional e f f e c t i v e n e s s with r e i n f o r c e m e n t ;
however, the two seem closely related.
While not detailed in the 1970 theory, the moti-
vational basis for avoidance learning is assumed
to be classically conditioned fear. The effect of
this fear, at least initially, is to shrink the animal's
response repertoire:
I am suggesting that the immediate and inevitable
effect of severe aversive stimulation on a domesti-
cated animal is to convert it, at least temporarily, into
a wild animal by restricting its response repertoire to
a narrow class o f . . . SSDRs [BoUes 1970, p. 33,
italics added].
[Fear] simply limits S's response repertoire to a
very restricted set of species-specific defense reac-
tions [BoUes 1971, p. 220].
[SSDRs axe] in a sense the only responses available
to the frightened rat, and we cannot profitably require
MASTERSON Pay. J. Biol. Sci.
October-December 1982
it to learn any more than to do one rather than the
other in any aversive situation [BoUes 1971, p. 189,
italics added].
At this initial stage, Bolles argues for an all-or-
none character for fear motivation. While point-
ing out that avoidance behavior is motivated in
the sense that the probability or "vigor" of an
avoidance response sometimes can be made to
vary with the aversiveness of the situation, he
argues that amount of fear motivation may be
largely irrelevant:
Many animals" defense reactions have an all-or-none
character so that what has to be accounted for is the
associative control of the response rather than its
motivation. A slight fear may be as hard to find as a
slight pregnancy. The [animal] either freezes or
leaves or it does not, and the question is what stimuli
control these behaviors, not how motivated are they
[BoUes 1971, p. 224].
Later in learning, the effect of fear has a more
continuous character, so that the response reper-
toire gradually expands beyond SSDRs as fear
becomes progressively inhibited by safety sig-
nals. E v e n t u a l l y , n o n - S S D R s , such as bar-
pressing, may occur.
The 1972 Version
While conceptually similar to the original ver-
sion of the theory, the 1972 version explicates its
relation to the motivation-reinforcement view
exemplified by Mowrer (1947). Bolles retains the
role of fear as motivation but explicitly denies the
role of r e i n f o r c e m e n t for r a p i d l y l e a r n e d
avoidance responses:
[SSDRs] have no direct associative linkage with the
CS, although they can be brought under CS stimulus
control with the use of differential punishment.
Avoidance behavior is motivated, as Mowrer con-
tended (that remains), but is not reinforced [BoUes
1972b, p. 131].
Thus, as in the original version, Bolles views fear
as the motivator and punishment of inappropriate
SSDRs as the learning mechanism for rapidly
acquired avoidance behavior.
In rejecting a reward mechanism, Bolles was
influenced in part by Bindra's attempts to explain
the acquisition of instrumental behaviors in Pav-
lovian terms (Bindra 1972, I974, and 1978) and by
the emergence of phenomena such as autoshap-
ing (Brown and Jenkins 1968) that lent support to
Bindra's approach (Bolles 1981, personal com-
munication). Bindra's is the most recent in a
series of attempts to derive instrumental learning
from Pavlovian learning mechanisms. Earlier at-
tempts at such a reduction were proposed by
Mowrer (1956) and Sheffield (1966). Bindra's ap-
proach is more successful due to its use of innate
Volume 17 SSDRs AND AVOIDANCE
Number 4
sensorimotor patterns guided by conditional in-
centive stimuli. Yet there remains room for
doubt as to the desirability of totally abandoning
operant reinforcement principles. For a discus-
sion of this and related issues, see Bindra (1978).
Consistent with his denial of any role for rein-
forcement in rapid avoidance learning, Bolles re-
pudiated his earlier inclusion of a functional ef-
fectiveness factor. This time he explicitly iden-
tified the notion of functional effectiveness with
reinforcement but denied its importance in the
light of Bolles and Riley's (1973) experiments on
freezing as an SSDR. BoUes and Riley found that
rats rapidly restrict their behavior to freezing
when freezing avoids shock. Since freezing pro-
duces minimal stimulus change,
It therefore becomes very difficult to defend any
hypothesis which attributes the reinforcement of
avoidance behavior to the consequences of the re-
sponse. Indeed, it becomes difficult to defend any
hypothesis which attributes the acquisition of
avoidance behavior to reinforcement [BoUes 1972b,
p. 127].
To account for slower learning, where the re-
quired avoidance response is not an SSDR, the
1970 views of the importance of safety signals (SSs)
is retained and expanded. It is argued that SSs and
FSs have been assumed to differ because they are
established differently (noncontingently and con-
tingently, respectively) and because most exper-
iments have tested FSs for reinforcing properties
and SSs for motivational effects. However, SSs
and FSs are postulated to be functionally equiva-
lent. Both predict safety, and both serve to rein-
force contingent behavior and "deactivate" or
inhibit ongoing fear-motivated behavior.
The 1975 Version
An earlier paper (Bolles 1972a) had concep-
tualized fear motivation in terms of an S-S* ex-
pectancy (where S is an initially neutral stimulus
such as WS onset and S* is a biologically signifi-
cant event such as shock). But it was not until
1975 that BoUes united this view of motivation
with his view of the mechanisms underlying the
rapid and slow learning of avoidance behavior.
The 1975 version of SSDR theory completed
the trend away from operant reinforcement ex-
planations. The 1970 version denied the role of
reward in rapid avoidance learning but retained
the reward-like concept of functional effective-
ness. The 1972 version of the theory eliminated
functional effectiveness. At this stage of de-
velopment, all that remained of operant rein-
forcement mechanisms was the suppression of
ineffective SSDRs by punishment and, possibly,
the reinforcement of avoidance responses by
207
safety signals. The 1975 version of SSDR theory
eliminated all forms of operant reinforcement.
The earlier assumption that the mechanism un-
derlying rapid learning is punishment-produced
suppression of competing SSDRs was replaced
by the view that SSDRs are not subject to any
form of learning. Freezing, fleeing, and fighting
are seen as respondents elicited by shock or by
shock cues; no response learning at all occurs in
avoidance situations. The rat simply learns that
certain cues (danger signals) predict shock, and
other cues (safety signals) predict safety: motiva-
tion is a matter of S-S* expectancies. Environ-
mental stimuli usually support one or another
SSDR and lead to its emergence, e.g., if an ap-
paratus "looks like a place to freeze," the animal
will freeze in the presence of danger signals.
Safety signals (SSs) are still viewed as important
in situations where the required response is not
an unambiguous SSDR. In these more slowly
learned situations, SSs can inhibit fear and thus
allow non-SSDRs to emerge. More importantly,
SSs can support and direct a particular defensive
behavior. Unlike earlier versions, the 1975
hypothesis views SSs as primarily motivational
rather than reinforcing. The animal approaches
SSs because it is motivated by its expectancy of
shock, but it is not reinforced by SS onset.
The 1978 Version
Reinforcement returned to the theory in its
most recent formulation (Bolles 1978), albeit in a
minimal role. As in the 1975 model, motivation is
conceptualized in terms of the elicitation of rela-
tively fixed units of behavior, the SSDRs, by
S-S* expectancies. Also as in the 1975 model,
neither reinforcement nor punishment is involved
in the acquisition of an avoidance response which
approximates an SSDR. For learning of non-
SSDRs, however, BoUes returned to the view
developed earlier (Bolles 1972b) of SSs as rein-
forcers for response learning.
Summary of SSDR Theory Development
In summary, the SSDR theory presented by
Bolles (1970) and later modified (Bolles 1971,
1972a, 1972b, 1975 and 1978) has remained con-
stant in its assumption that fear dramatically
limits the rat's response repertoire. It has increas-
ingly specified the motivational processes pre-
sumed to be operating, culminating in a concep-
tualization of motivation in terms of expectancies
of danger and safety. It originally postulated two
distinct learningprocesses, a punishment mech-
anism for rapid suppression of inappropriate
SSDRs and a safety-signal reinforcement process
208 CRAWFORD AND MASTERSON
governing the gradual acquisition of non-SSDRs.
Both were discarded (1975) in favor of the view
that SSDRs are respondent behaviors differen-
tially elicited by supporting stimuli in the envi-
ronment and that the gradual learning of non-
SSDRs is governed primarily by motivational,
rather than reinforcing, properties of SSs. The
most recent statement of the theory (Bolles 1978)
does, however, retain a minor role for the rein-
forcing properties of SSs.
Evaluation of the Theory
One potential problem with the development of
rigorous empirical tests of SSDR theory is that its
predictions lack specificity. For example, Selig-
man and Hager (1972, p. 212) have pointed out
that it is difficult to determine preexperimentally
which responses are SSDRs and which SSDR will
emerge in a particular situation. Hineline and
Harrison (1978) compared lever-biting, assumed
to be an SSDR, and the "arbitrary" response of
lever-pressing, as avoidance responses. Al-
though lever-biting occurred frequently with
some rats, the response did not come under the
control of the avoidance contingency. In con-
trast, lever-pressing was established and was
controlled by the avoidance contingency.
Hineline and Harrison characterize the SSDR
approach as insufficiently precise for other than
post hoc explanation of these data and caution
against its uncritical acceptance. It is not clear,
however, whether the difficulty lies entirely with
the theory or, at least in part, with the failure of
psychologists to systematically identify and clas-
sify SSDRs.
Response Constraints
One of the most salient features of SSDR
theory is its assumption that, initially, fear com-
pletely restricts the rat's response repertoire to
SSDRs. It follows that, in order for any learning
to take place, at least early in avoidance training,
the required response must be an SSDR. It also
follows that the learning of a non-SSDR as
avoidance response must necessarily be slow and
erratic, involving as it does reinforcement by SSs
which are only gradually established.
The extreme response restriction proposed by
the SSDR hypothesis for the initial stages of
learning can be evaluated both conceptually and
empirically. First, it is conceptually difficult to
reconcile with a continuum along which there are
gradations of fear. If we imagine fear as poten-
tially ranging from zero to maximum along this
continuum, where does the rat's response reper-
toire shrink to SSDRs? It seems likely that the
Pay. J. Biol, Sei.
October-December 1982
repertoire narrows gradually as fear increases,
rather than suddenly at a particular point. The
absolute response restriction rule also lacks his-
torical precedent. Traditionally, ethologists have
viewed motivation as continuously modulating
the thesholds for instinctive behaviors. Seen
from this perspective, fear would be expected to
increase the probabilities of SSDRs but not to
absolutely exclude non-SSDRs. Alternatively, it
is possible that an orienting response would pre-
dominate at low levels of fear, with SSDRs ap-
pearing only after a certain threshold had been
reached. The orientation response may vary con-
tinously, while after some threshold value the
SSDR may be all-or-none. At any rate, the strong
response restriction rule of the SSDR hypothesis
presents an exception to the flexibility of typical
mammalian behavior, as Bolles (1972) has
pointed out.
Empirical evidence on the restriction of the
frightened rat's behavior to SSDRs and the slow,
erratic learning of non-SSDRs comes mainly
from studies of bar-press avoidance learning.
Bar-pressing would seem to be the perfect pro-
totype of a non-SSDR. The fact that rats learn it
as an avoidance response only with great diffi-
culty thus is consistent with the theory. The fact
that they do sometimes achieve moderately good
performance is explained (Bolles 1970 and 1971)
as the result of adapting the freezing SSDR to the
situation. The rat learns to freeze on the bar, so
that the bar is pressed by the animal's reflexive
lurch in response to shock onset. Since this
strategy minimizes shock, fear gradually dissi-
pates, and the response repertoire gradually ex-
pands. With reinforcement provided as SSs are
established, the response that began as a variant
of the freezing SSDR may eventually become an
operant bar-press response.
There is evidence that rats typically do remain
immobile and in contact with the bar during in-
tertrial intervals (ITIs) and that shock onset does
typically elicit a brief lurch which is recorded as a
short-latency escape response (Davis and Burton
1974 and 1976, Peterson and Lyon 1975). Also
consistent with Bolles' interpretation is evidence
that bar-holding, which can be seen as a minor
variant of freezing, is readily learned as an
avoidance response in a Sidman procedure
(Davis and Burton 1976) and that when rats are
required to lift a bar to escape shock, they solve
the problem by freezing under the bar and lurch-
ing upward at shock onset (Davis, Hirschorn, and
Hurwitz 1973).
On the other hand, there is evidence that,
under certain special conditions, bar-press
avoidance can be readily acquired. Masterson
(1970) and Crawford and Masterson (1978) both
Volurar
Number 4
17 SSDRs AND AVOIDANCE
trained rats in a bar-press avoidance task. For
some groups, a bar-press led to standard
avoidance contingenies--WS offset, FS onset,
and shock escape or avoidance. Learning in these
groups followed the extremely slow and uncer-
tain course typical in bar-press situations. For
other groups, a bar-press led to the same con-
tingencies plus the opening of a door allowing the
rat to run to a safe place. Learning was much
more rapid in these groups; in one experiment
(Masterson 1970), the animals readily achieved
asymptotic avoidance performance of 100%. The
bar-press response was clearly an operant, non-
reflexive, and learned without passing through a
freezing-on-the-bar stage. Thus, such a stage,
while it may be typical, is not a universal or
necessary condition for acquisition of bar-press
avoidance.
It is also useful to examine SSDR theory's view
of exactly what responses constitute the rat's
SSDR repertoire. (Evidence for reinforcement
effects on these behaviors will be examined in the
next section). As described earlier, Bolles (1970)
postulated freezing, fleeing, and fighting as
SSDRs. There is abundant evidence that freezing
and fleeing are innate and highly probable re-
sponses for the rat in fear-eliciting situations
(Blanchard and Blanchard 1971 and 1972,
Rakover 1975). Fighting, or attack behavior, is
less probable and appears to be closely linked to
primary aversive stimulation plus the existence
of an appropriate object to attack. For example,
under some circumstances shock elicits attacks
directed toward other rats (Ulrich and Azrin
1962) or toward mice (Myer and Baenninger
1966). Knutson (1971) reported that rats paired
with a n o n s h o c k e d rat developed freezing
postures that reduced or avoided shock in
preference to attacking the other rat. However,
some aggression did occur.
Thus the three responses proposed as SSDRs
by Bolles have remained viable. It is possible that
other responses may be SSDRs as well. Search-
ing behavior, for example, is characteristic of the
rat's initial exposures to shock and persists until
several seconds after shock, when freezing be-
comes dominant. At other times, a more cautious
searching behavior may occur, punctuated by pe-
riods of freezing. This latter kind of searching
probably diminishes over trials of avoidance
learning as the rat becomes familiar with the ap-
paratus. As a result of searching, the rat may
discover stimulus configurations that support
either freezing or flight.
Digging and clawing may be seen as searching
SSDRs, since they can be interpreted as attempts
to obtain an escape route. That is, searching in-
volves not only an examination of the dangerous
209
place, aimed at finding an opening, but can be an
active effort to create an opening. Furthermore,
recent research on defensive burial of objects
shows that, under some circumstances, the rat
approaches and buries sources ofaversive stimu-
lation (Pinel and Treit 1980).
Still another SSDR is thigmotaxis. Rats in
novel situations, as well as in the presence of
cues previously paired with shock, display
strong tendencies to avoid open areas (Crawford
et al. 1981, Grossen and Kelly 1972). Defensive
digging, thigmotaxis, and searching may serve
varying functions and merit integration into
SSDR theory.
Learning Mechanisms and Their
Motivational Basis
One of the most striking features of the SSDR
approach is its rejection of a reinforcement prin-
ciple for rapid learning. As described earlier,
Bolles (1970) entertained the idea that an SSDR
must be "functionally effective" to increase in
probability but later (1972, 1975) argued against the
importance of the stimulus consequences of re-
sponding. The notion of functional effectiveness
was perhaps too hastily discarded. There is evi-
dence that flight is rapidly learned as an
avoidance response only when it is effective in
the sense of gaining access to a different location.
For example, rats rapidly learn to jump to a ledge
to avoid shock (Maatsch 1959) but learn very
slowly, and to a low asymptote, when jumping
does not lead to "getting away" (Masterson,
Whipple, and Benner 1972). Bolles (1975) cites an
unpublished study by Duncan in which running
was not learned as an avoidance response when
the requirement was simply that the rat run a
fixed distance in any direction. Furthermore,
Modaresi (1975) found that rats provided with a
safe platform (inserted on the appropriate side of
the apparatus at the start of each trial) showed
unusually rapid learning of a shuttle avoidance
task. Finally, Masterson, Crawford, and Bartter
(1978) developed a "brief escape" variant of
one-way avoidance in which rats are returned to
the shock box immediately after running into the
safe box, and spend the ITI in the shock box. Rats
learn the brief escape task fairly rapidly, suggest-
ing that even brief exposures to a safe place are
rewarding.
Thus, available evidence suggests that func-
tional effectiveness--running away vs. simply
running--is important. And fuctional effective-
ness is easily conceptualized in terms of rein-
forcement: a response is effective only if it pro-
duces certain rewarding stimulus changes.
210 CRAWFORD AND MASTERSON
Perhaps the most clearcut demonstration of the
rewarding effect of flight-associated stimulus
changes is provided by the data of Crawford and
Masterson (1978). As described earlier, rats
rapidly learned a bar-press avoidance response
when it provided access to a safe place but not
when it provided only standard bar-press
avoidance contingencies. Furthermore, rats
learned equally well when a bar-press resulted in
their being carried to the safe place. Finally,
Crawford and Masterson also showed (experi-
ment 2) that stable asymptotic avoidance per-
formance deteriorated rapidly and completely
when safe-place access was discontinued, even
though WS offset, FS onset, and shock escape/
avoidance contingencies remained in effect.
These results show that actual performance of an
SSDR is not necessary for fairly rapid avoidance
learning. They also show that fearful rats are not
restricted to performing SSDRs. But most impor-
tantly, they are best explained in terms of the
reinforcing properties of safe-place access. The
stimulus changes provided by WS offset, FS on-
set, and shock escape/avoidance were insuffi-
cient to establish bar-press avoidance respond-
ing. But the stimulus changes associated with a
safe place were crucial to both the establishment
and maintenance of avoidance responding.
As described earlier, BoUes partially based his
1972b rejection of a reinforcement principle for
rapid avoidance learning on evidence that freez-
ing is respondent behavior (Bolles and Riley
1973). The BoUes and Riley experiments were
conducted in a standard laboratory apparatus.
Conceivably, use of a richer, naturalistic situa-
tion may reveal a functional effectiveness com-
ponent for freezing. Consider the case when a rat
spies a potential predator at a considerable dis-
tance. While flight is presumably an option, it
may not be necessary. Assuming the rat is near a
burrow to which it can, if the need arises, quickly
flee, it would be adaptive for the rat to try freezing
first. If freezing is functionally effective in the
sense that the predator fails to notice the rat,
flight will not be required, and the predator will
eventualy go elsewhere and the rat can go about its
business. Indeed, it might be adaptive for rats to
save running down a burrow as a last resort (as
when the predator comes too close), because the
activity might attract the predator's attention to
the burrow. It seems possible, then, that the func-
tional effectiveness of freezing may be observ-
able only in a naturalistic situation.
Similar arguments can be made for fighting.
Bolles (1975, p. 363) describes his unsuccessful
attempts to condition attack behavior as an
avoidance response. We have a similar history of
unsuccessful attempts in our own laboratory.
l'av. J. Biol. Sci.
October-December 1982
Furthermore, Hineline and Harrison (1978)
report only limited success in conditioning lever-
biting as an avoidance response. However,
aggressive behaviors are not totally immune to as-
sociative control: Maier, Anderson, and Leiber-
man (1972) showed that exposure to inescapable
(but not to escapable) shock reduced the fre-
quency of later shock-induced fighting. And Az-
rin, Hutchinson, and McLaughlin (1965) demon-
strated that rats can learn an operant response for
the opportunity to attack another rat during
shock. These bits of evidence are insufficient to
draw firm conclusions about the functional effec-
tiveness of fighting in rats. Perhaps fighting, like
freezing, can be best examined for functional ef-
fectiveness in naturalistic situations; both may
prove to be susceptible to reinforcement by only
a very limited class of reinforcers.
The case for SSDRs as respondents rests
mainly on the evidence for freezing, as did the
case for response restriction. Freezing may well
be respondent behavior; however, it does not
logically follow that flight is a respondent. There
are respondent components connected with other
motivational systems--for example, salivation
with hunger motivation--yet it has seldom been
argued that the existence of salivation forces us to
regard all food-directed behavior as respondent.
Defense motivation provides a similar example:
the common view that urination, defecation, and
piloerection are respondents has not been
generalized to include flight in the respondent
category. The discovery of one more respondent
(freezing) need not force this generalization,
especially given the direct evidence cited above
for the importance of consequences in flight
avoidance responding. Experimental studies of
other possible SSDRs, such as searching and de-
fensive digging, could give a more general picture
of the role of reinforcement in rapid avoidance
learning.
A role for reinforcement in the learning of
non-SSDRs is suggested by Bolles (1970) and de-
veloped more fully later (Bolles 1972b): environ-
mental stimuli and feedback from the animal's
own response become safety signals through re-
peated pairing with the absence of shock. Bolles
(1972b) argues for both a motivating ("de-
activating") and a reinforcing role for SSs. Non-
contingently established SSs (e.g., Rescorla and
LoLordo 1965) and contingently established FSs
(e.g., Bolles and Grossen 1969) are postulated to
be functionally equivalent in that both predict
safety. Bolles (1972, p. 137) acknowledges that
the evidence for such functional equivalence is
"'sketchy and incomplete, and not yet very com-
pelling," and proposes some methods for further
empirical verification.
Volume 17
Number 4
SSDRs AND AVOIDANCE
One obvious way to establish the functional
equivalence of FSs and SSs would be to demon-
strate that prior Pavlovian conditioning of a signal
as an SS enhances its value as an FS when it is
subsequently made contingent on an avoidance
response. Though the issue has been studied for
the past decade, there is as yet no unequivocal
demonstration of such an effect in the literature.
Morris (1975) found that rats learned shuttle
avoidance faster when the FS had been pre-
trained as an SS, thus appearing to provide evi-
dence for functional equivalence. However, these
results are rendered somewhat ambiguous by the
fact that his non-pretrained FS group performed
no better than a no-FS group. In other words,
Morris failed to replicate the basic FS effect. The
evidence for equivalent reinforcing properties of
FSs and SSs remains as uncompelling as Bolles
characterized it to be in 1972.
Bolles (1975) shifted from emphasizing the
reinforcing properties of SSs to emphasizing their
motivational properties. The learning of non-
SSDRs is thus explained in terms of the animal's
tendency to approach SSs and to withdraw from
danger signals. Certainly there is evidence for
such sign-tracking in appetitive situations: pi-
geons approach keys whose illumination has been
paired with food and withdraw from those signal-
ing time out from food availability (Wasserman,
Franklin, and Hearst 1974). But the evidence is
less compelling for aversive situations. In a study
by Bartter and Masterson (1980), rats were given
inescapable shocks in a tilt-floor cage in which a
light stimulus could be presented on a key at
either end. For the paired group of rats, the light
preceeded the shocks. For the unpaired group,
the light predicted shock-free periods. For the
random group, light and shock presentations
were unrelated. Though the unpaired group
showed a small preference for the lit side of the
cage. the differences between the percent pref-
erence scores of the groups were relatively
small, certainly insufficient to account for the
acquisition of shuttlebox avoidance responding.
A larger aversive sign-tracking effect in rats has
been demonstrated by Leclerc and Reberg (1980)
using presentations of a platform as the condi-
tional stimulus. Thus, their SS was a supporting
stimulus for flight, so that the SS approach mech-
anism postulated by Bolles is confounded with
his SSDR elicitation mechanism. Since Bolles'
theory maintains a separation between these two
mechanisms, these results are not directly
applicable as a test of the theory.
Green (1978) and Green and Rachlin (1977)
have shown that a light danger signal displayed on
a pecking key reduces the number of pecks pi-
geons will make to produce food. In their proce-
211
dure, two pecking keys supported equal concur-
rent variable interval (VI) schedules for food re-
ward. Presentation of a light danger signal on one
key decreased the relative rate of responding on
that key, a specific directional effect, and also
reduced the absolute amount of responding on
both keys combined, a nonspecific or diffuse ef-
fect. The extent of each effect was determined by
factors such as shock intensity and CS localizabil-
ity. While the directional effect is consistent with
the withdrawal from danger signals expected in
aversive sign-tracking, it may reflect little more
than a reluctance of pigeons to peck a danger
signal. That is, pigeons and rats may be tolerant
of relatively close proximities to a light danger
signal so long as they need not establish contact.
The sign-tracking explanation is unsatisfyingly
vague in its specification of exactly how certain
SSs are established. For example, consider
BoUes' (1975) explanation of shuttlebox avoidance
learning. The rat initially freezes because the
shuttlebox is perceived as a place to freeze. After
a few trials, however, cues such as WS onset and
apparatus cues become established as danger
signals. Safety signals become established, t o o - -
response-produced feedback, WS offset, and the
"other end" of the shuttlebox. The latter cue is
crucial because it is the only SS to which the rat
can direct its approach behavior. But how does
the "other end" of the shuttlebox become an SS?
Both ends are also danger signals. And "'the other
end" is not a place but is constantly redefined by
the changing relationship of the rat to the ap-
paratus. Pity the poor rat trying to establish an
expectancy of safety--as soon as it reaches the
"'other end," the other end becomes the other
end !
The 1975 SSDR theory shares a problem that
has traditionally plagued expectancy theories,
the problem of specifying how expectancies are
to be translated into action. What expectancy
theories tack is the specification of processes
that control the interaction of incoming sen-
sory data with memory representations of past
events to produce behavior. How are these mem-
ory representations stored, retrieved, and com-
bined? Expectancy theories have not delineated
the mechanisms that bridge the gap between per-
ceiving and responding. Perhaps it was this
problem that influenced Bolles (1978) to return
a SS-reinforcement mechanism to the theory.
Summary and Conclusions
The SS DR hypothesis proposed that fear abso-
lutely limits the rat's response repertoire to a
small class of relatively fixed behaviors, the
SSDRs. The evidence is consistent with a similar
212 CRAWFORD AND
but less stringent rule: the fearful rat is highly
likely to emit SSDRs but is not totally restricted
to doing so. The SSDR hypothesis also proposed
three SSDRs, freezing, fleeing, and fighting. Evi-
dence indicates that these responses are indeed
crucial to the defense repertoire and suggests that
others may also be added to the list. The model's
arguments that SSDRs are respondents (Bolles
1975) and that rapid avoidance acquisition in-
volves no reinforcement appear to be most defen-
sible in the case of freezing. Little evidence is
available for fighting. Flight responses, on the
other hand, appear to be established and main-
tained by their consequences. Investigation of
fighting and other possible SSDRs (e.g., search-
ing. defensive digging) may help give a more gen-
eral picture o f the role of reinforcing conse-
quences in defensive behavior. Finally, the 1975
SSDR model's reliance on danger and safety sig-
nals as elicitors of approach and withdrawal be-
havior awaits further confirmation of a reliable
and sizeable sign-tracking effect in aversive
situations.
The SSDR model was one o f the first learning
theories to question the ecologic validity of labo-
ratory conditioning tasks. Further, it was one of
the first to incorporate biological constraints on
learning. Its publication encouraged theorists to
confront the response problem and, more gener-
ally, to consider the adaptive significance of
avoidance learning phenonema.
By denying a major role for instrumental re-
sponse learning, SSDR theory joins a tradition
begun by Sheffield (1966) and Mowrer (1956) and
elaborated by Bindra (1972, 1974, and 1978). In
this tradition, the structure of behavior previ-
ously attributed solely to instrumental response
learning is now attributed solely to innate re-
sponses (which can be guided by Pavlovian con-
ditional stimuli). A major contribution of Bolles'
theory has been to develop this argument for the
case o f animal defense behavior. It is now neces-
sary to integrate other phenomena, such as the
rapid aquisition of flight as an avoidance re-
sponse, which clearly indicate an instrumental
response learning process. Toward this goal, we
have elsewhere proposed a defense motivation
s y s t e m t h e o r y of a v o i d a n c e learning which
e q u a t e s r e i n f o r c e m e n t with c o n s u m m a t o r y -
stimulus feedback from innate defense responses
(Masterson and Crawford 1982).
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Nominations for Membership

in the Pavlovian Society
The Society is always interested in considering nominees
for membership who have special interests and expertise in
the advancement of the biological sciences. The Society
deemphasizes bureaucratic procedures in accordance with
the desire of its founder. W. Horsley Gantt. so that maximal
effort can be expended for scientific accomplishment. The
Society is interdisciplinary and international.
Members who wish to nominate candidates should return a
brief biographical sketch or curriculum vita to David Randall,
Ph.D., Chairman, Membership Committee, Department of
Physiology, U n i v e r s i t y of K e n t u c k y , Medical School,
Lexington, Kentucky 40506.
Especially interested individuals who do not know a mem-
ber of the Society who wish to apply for membership may
nominate themselves.

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