Behavior and Philosophy, 41, 33-59 (2013).

2015 Cambridge Center for Behavioral Studies

FILLING THE GAPS: SKINNER ON THE ROLE OF NEUROSCIENCE

IN THE EXPLANATION OF BEHAVIOR
Diego Zilio
UFES -Federal University of Esprito Santo

ABSTRACT: It is often said, especially in philosophy and the neuroscience literature, that
Skinner defended an anti-physiological position on the explanation of behavior. Aside from
this, behavior analysts who discuss the relation between behavior analysis and physiology
usually emphasize the independence of these two fields. Amid criticisms of Skinners
allegedly anti-physiological position and behavior analysts defense of their discipline as
an autonomous science, there is comparatively little discussion of Skinners positive views
on physiology. The goal of this paper is to present an analysis of these views, taking into
account Skinners writings from the 1930s to the 1990s. Among the topics to be
discussed are the definition of the object of study of physiology, its role in the explanation
of behavior, and the relation between behavior-analytic and physiological explanations. I
pay special attention to Skinners use of physiological hypotheses in developing his
theories of private events and perception, and I hope to counteract the ill-founded notion
that Skinner was always opposed to physiology in the study of behavior.
Key words: Skinner, behavior analysis, radical behaviorism, neuroscience, physiology,
explanation of behavior

Outside the behavior-analysis community, Skinner is known for his critical

positions regarding physiological explanations of behavior (e.g., Bradnan, 1982;
Garca-Hoz, 2004; Illard & Feldman, 2001; Kandel, 1976; Loucks, 1941;
Machamer, 2009; Panksepp, 1990; Razran, 1965; Staddon & Bueno, 1993). To this
day, the notion that Skinner defended an anti-physiological position continues to
be disseminated among neuroscientists and philosophers. For example, consider
this excerpt from a handbook on the philosophy of neuroscience:
Skinner () dismisses the use of physiological variables, because he believes
that they could add nothing of relevance to the science of behavior. All one
needed, for Skinner, were the environmental or stimulus conditions (), the
behavioral response, and the contingencies or conditions that led to
reinforcement. () Clearly in this radical Skinnerian sense, no neuroscientist
could be a behaviorist, for they, by virtue of discipline, must discuss variables
that are internal to the organism and irrelevant to a science of behavior.
(Machamer, 2009, p. 168)

Meanwhile, behavior-analytic writings often emphasize Skinners arguments

for the autonomy of behavior science from physiology (e.g., Baer, 1996; Carvalho
Neto, 1999; Carvalho Neto & Tourinho, 1999; Greenberg, 1983; Greenberg &

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Lambdin, 2007; Reese, 1996; Staddon, 2001; Starling, 2000; Tourinho, Teixeira &
Maciel, 2000).
Amid criticisms of Skinners supposedly anti-physiological stance and
behavior analysts own efforts at establishing behavior analysis as an autonomous
science, Skinners positive views on the role of physiology in the explanation of
behavior have been rather neglected. This situation perpetuates the misguided
ideaevident in Machamers (2009) quotethat behavior analysis and
neuroscience are actually incompatible with each other. To be sure, a significant
literature already deals with the relation between behavior analysis and
neuroscience by clarifying their possible relations and roles in the explanation of
behavior (e.g., Bullock, 1996; Carvalho Neto, 1999; Carvalho Neto & Tourinho,
1999; Catania, 2000; Donahoe, 1996, 2002; Donahoe & Palmer, 1994; Elcoro,
2008; Mechner, 2008; Moore, 2002; Mustaca, 2003; Schaal, 2003, 2005; Silva,
1987; Strumwasser, 1994; Thompson, 2007, 2008, Timberlake, Schaal &
Steinmetz, 2005) and the present article can be seen as an addition to these efforts.
What is lacking so far is a detailed, systematic analysis of Skinners views on the
topic that takes into account his writings from the 1930s to the 1990s. He wrote
in several occasions that behavior analysis had gaps and that those would be
filled by physiology, especially neuroscience. The goal of this paper is to discuss
these gaps.

Where is the Boundary?

To understand Skinners perspective on the role of neuroscience in the
explanation of behavior, we first need to examine how he defined the object of
physiological studies and how it differs from that of behavior analysis. Through
various definitions, Skinner actually emphasized different aspects of the research
topics of behavior analysis and physiology. I will classify these definitions
respectively as structural, causal, and explanatory.
The structural definition
The structural definition presents the object of study of physiology as the
physiological structures that make behavior possible (Skinner, 1947/1961a,
1957/1961d, 1963, 1974, 1975a, 1983, 1988, 1989d). For example, Skinner wrote
that as the science of physiology advances, it will presumably be possible to show
what is happening in various structures within the organism during particular
behavioral events (1947/1961a, p. 236); an organism behaves as it does because
of its current structure (1974, p. 17); it is a given organism at a given moment
that behaves, and it behaves because of its biological equipment at that moment.
Eventually neurology will tell us all we need to know about the equipment (1988,
p. 301); what they call mechanism seems to be the current state of the organism
described in the language of physics and biology. The organism behaves as it does
because of the present state (1988, pp. 304-305). According to Skinner (1974),
this structure is made of effectors and receptors, nerves, and a brain (p. 136).

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Thus, a biological system (Skinner, 1956/1961e, p. 215)the organismis

necessary for behavior to occur. Structure, equipment, state, and
mechanism are terms used by Skinner to generically indicate the physiological
configuration of the organism while it behaves in a given fashion. Importantly, (a)
this configuration is causally responsible for the organisms behavior, and (b) this
structure should not be view as static or permanent. It changes through time,
mostly as a consequence of the interactions between organism and environment.
Those changes in physiology are directly responsible for later behavioral changes.
Skinner made this point clear in numerous occasions (e.g., 1972, 1974, 1975b,
1985, 1988, 1988/1989c, 1993). For example:
A behavioral analysis acknowledges the importance of physiological research.
What an organism does will eventually be seen to be due to what it is, at the
moment it behaves, and the physiologist will someday give us all the details. He
will also tell us how it has arrived at that condition as a result of its previous
exposure to the environment as a member of the species and as an individual.
(Skinner, 1974, p. 249)

The behavior of an organism at time t can be causally associated with what

the organism is at t. Here no mention is made of phylogenetic or ontogenetic
history. Explaining behavior at moment t is done solely in terms of the organisms
physiological state or configuration at t. Skinner (1988) goes even further
along these lines:
Of course differences in structure are important, but they are themselves the
results of contingencies of selection. Structures are what is selected. Organisms
have evolved in such a way that when they are hungry, food strongly reinforces
their behavior. Other reinforcers are merely conditioned. But both reinforce
because of the structure of the organism. The structure is not an initiating cause
of anything, however. (p. 434, italics added)

It is the organisms physiological configuration, or what Skinner calls

structure, that is the object of selection by phylogenetic and ontogenetic
contingencies. This object is not behavior, at least not directly. Contingencies
select behavior only to the extent that they select physiological structures or
configurations responsible for behavior (cf. Glenn & Madden, 1995). The point
here is that in Skinners view the selection of behavior is indirect; it depends on the
intermediary link, the physiological configuration of the organism. This does not
imply, however, that all of the causal factors related to behavior are to be found in
physiology. We cannot say of the physiological configuration at time t that it is the
initial and only cause of behavior at t, because this configuration is itself a product
of phylogenetic and ontogenetic contingencies.
The causal definition
According to Skinner (1986), in a given episode the environment acts upon
the organism, something happens inside, the organism then acts upon the

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environment, and certain consequences follow (p. 716). This excerpt suggests a
causal chain made of environmental events, physiological processes and behavior.
Environmental events change the physiological configuration of an organism;
these changes cause some behavior; the behavior, by its turn, changes the
environment. Figure 1 illustrates this causal chain.

Figure 1: Skinners (1986) causal chain.

According to Skinner (1986), the first, third, and fourth of these events is the
field of a science of behavior, which undertakes to discover how they are related to
each other. What happens inside is another part of the story (p. 716). In terms of
Figure 1, behavior analysis deals with the events labeled (1), (3), and (4); events
which, by definition, constitute the three-term contingency, the basic unit of
analysis of operant behavior (cf. Skinner, 1969a). The remaining link in the causal
chain, event (2), is the proper domain of physiology, especially neuroscience.
When Skinner (1988) says that the nervous system mediates the relations studied
in an experimental analysis of behavior (p. 257), he is fully consistent with Figure
1.
Whereas Figure 1 presents events rather simplistically as a single chain-like
sequence, Skinner (1956/1961e, p. 206) formulates a richer and more heuristic
description of the same basic idea. Here it is, using Skinners own illustration this
time (Figure 2).

Figure 2: Skinners heuristic causal chain. Adapted from Skinner (1956/1961e, p. 206).

Instead of a point-to-point description of the events that constitute the causal

chain, here we find a distinction between the behavior of organism (the dependent

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variable) and the environmental events related to this organisms phylogenetic and
ontogenetic histories (the independent variables). According to Figure 2, the
behavioral repertoire of an organism at time t is a product of its species history
and of its own history of interactions with the environment. But what about the
organism, the mediational variable of this causal chain? In Skinners
(1956/1961e) words:
This does not mean, of course, that the organism is conceived of as actually
empty, or that continuity between input and output will not eventually be
established. The genetic development of the organism and the complex
interchanges between organism and environment are the subject matters of
appropriate disciplines. Some day we shall know, for example, what happens
when a stimulus impinges upon the surface of an organism, and what happens
inside the organism after that, in a series of stages the last of which is the point
at which the organism acts upon the environment and possibly changes it. At
that point we lose interest in this causal chain. Some day, too, we shall know
how the ingestion of food sets up a series of events, the last of which to engage
our attention is a reduction in the probability of all behavior previously
reinforced with similar food. Some day we may even know how to bridge the
gap between the behavioral characteristics common to parents and offspring.
But all these inner events will be accounted for with techniques of observation
and measurement appropriate to the physiology of the various parts of the
organism, and the account will be expressed in terms appropriate to that subject
matter. () Its task [physiologys] is to account for the causal relations between
input and output which are the special concern of a science of behavior.
Physiology should be left free to do this in its own way. (pp. 214-215)

One day it will be possible to know how the environment affects the organism
by causing physiological changes, which are in turn responsible for the occurrence
of behavior. Skinner also brings into discussion the causal/mediational role of
physiological events: They are paramount to the very existence of causal relations
between the independent and dependent variables (what Skinner calls inputs and
outputs in this excerpt) studied in behavior analysis.
The explanatory definition
Skinner often discussed the possibility of direct manipulation of physiological
variables with the aim of predicting and controlling behavior. His discussions
exemplify the third aspect of what would be the object of study of physiology from
Skinners perspective. Let us start with the following excerpt:
Eventually a science of the nervous system based upon direct observation rather
than inference will describe the neural states and events, which immediately
precede instances of behavior. We shall know the precise neurological
conditions which immediately precede, say, the response, No, thank you. ()
However, we may note here that we do not have and may never have this sort of
neurological information at the moment it is needed in order to predict a specific
instance of behavior. It is even more unlikely that we shall be able to alter the

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nervous system directly in order to set up the antecedent conditions of a

particular instance. The causes to be sought in the nervous system are, therefore,
of limited usefulness in the prediction and control of specific behavior. (Skinner,
1953/1965, pp. 28-29, italics added)

Here we find a more detailed description of the idea already present in

Skinner (1956/1961e) about the possibility of knowing one day the physiological
events causally responsible for a particular behaviorthe events labeled (2) in the
causal chain of Figure 1. Skinner talks specifically about neuroscience and, most
importantly, he says that neurophysiological events must be directly observed. It is
impossible to predict and control behavior by manipulating neurophysiological
events without any direct knowledge of neurophysiology. Skinner (1953/1965)
goes further:
To what extent is it helpful to be told, He drinks because he is thirsty? If to be
thirsty means nothing more than to have a tendency to drink, this is mere
redundancy. If it means that he drinks because of a state of thirst, an inner causal
event is invoked. If this state is purely inferential - if no dimensions are assigned
to it which would make direct observation possible - it cannot serve as an
explanation. But if it has physiological () properties, what role can it play in a
science of behavior? The physiologist may point out that several ways of raising
the probability of drinking have a common effect: they increase the
concentration of solutions in the body. Through some mechanism not yet well
understood, this may bring about a corresponding change in the nervous system
which in turn makes drinking more probable. () we have a causal chain
consisting of three links: (1) an operation performed upon the organism from
without for example, water deprivation; (2) an inner conditionfor example,
physiological () thirst; and (3) a kind of behaviorfor example, drinking.
Independent information about the second link would obviously permit us to
predict the third without recourse to the first. () The second link is useless in
the control of behavior unless we can manipulate it. (pp. 33-34, italics added)

Skinner again describes the heuristic causal chain discussed earlier, but now
he focuses on the manipulation of links in the chain. An aspect of the organisms
history (in this case, the operation of deprivation, 1) modifies in some way the
neurophysiology of the organism (2), resulting in what we may call state of
thirst, which in turn increases the probability of drinking behavior (3). The second
link in the explanation of behavior can be dealt with in different ways, however.
We cannot infer the state of thirst from the frequency of drinking behavior. This
would be circular reasoning: inferring that the organism is thirsty because it is
drinking water, and explaining the drinking by saying that the organism is thirsty.
Inferring the second link from the first (for example, to say that the organism is
thirsty because of the long time since it last drank water) would be useless as
well, because this inference would depend on our previous knowledge of the

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history of organism. As far as Skinner is concerned, an inferential second link

cannot serve in the explanation of behavior: To be useful, explanations in terms of
the second link should deal with events that are observable and susceptible of
manipulation. Otherwise, we face the risks of explanations based on hypothetical
constructs (Moore, 2008; Skinner, 1974, 1950, 1953/1965, 1988; Zuriff, 1985):
The most objectionable practice is to follow the causal sequence back only as far
as a hypothetical second link (Skinner, 1953/1965, p. 34).
What if the second link is actually physiological? What if the state of thirst
is described accurately as a certain physiological process? In that case, according
to Skinner, we could associate different ways of raising the probability of drinking
behavior (deprivation of water, ingestion of salt, etc.) with their physiological
effects and associate drinking behavior with the physiological events that precede
it. When Skinner wrote his comments, the relevant knowledge was unavailable, but
he was aware that this was only a technological limitation of the physiology of the
times (Skinner, 1969b, 1972, 1974, 1975b, 1987, 1988). Eventually, directly
knowledge of the second link would be possible and then (only then) it could serve
to the explanation of behavior. This point is clear in Skinners (1972, 1974) talk of
the omniscient physiologist:
There are gaps in time and space between behavior and the environmental
events to which it is attributed, and it is natural to try to fill them with an
account of the intervening state of the organism. We do this when we summarize
a long evolutionary history by speaking of genetic endowment. Should we not
do the same for a personal history? An omniscient physiologist should be able to
tell us, for example, how a person is changed when a bit of his behavior is
reinforced, and what he thus becomes should explain why he subsequently
behaves in a different way. () An omniscient physiologist should be able to do
the same for comparable states in the field of behavior. He should also be able to
change behavior by changing the organism directly rather then by changing the
environment. (1972, p. 18)

The omniscient physiologist is Skinners argumentative device to defend the

possibility, at least in principle, of explaining behavior by studying the second link.
The omniscient physiologist is able to establish causal relations between events
related to the history of organism and changes in his physiological configuration
(first and second links) as well as between those physiological events and the
organisms subsequent behavior (second and third links). Therefore, the
omniscient physiologist can predict and control behavior through the observation
and manipulation of physiological events.

Explanations and Gaps

Skinners most common attribution of function to neuroscience is that of
filling in the gaps of a behavioral analysis (1953/1965, p. 54; 1956/1961e, p. 214;
1969b, p. 24; 1972, p. 18; 1974, pp. 214-215; 1975b, p. 43; 1985, p. 297; 1987, p.

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782; 1988, p. 122, 128, 257, 470; 1989a, p. 56; 1989d, p. 18; 2009, p. 69). For
example:
The promise of physiology is of a different sort. New instruments and methods
will continue to be devised, and we shall eventually know much more about the
kinds of physiological processes, chemical or electrical, which take place when
a person behaves. The physiologist of the future will tell us all that can be known
about what is happening inside the behaving organism. His account will be an
important advance over a behavioral analysis, because the latter is necessarily
historicalthat is to say, it is confined to functional relations showing
temporal gaps. Something is done today which affects the behavior of an
organism tomorrow. No matter how clearly that fact can be established, a step is
missing, and we must wait for the physiologist to supply it. He will be able to
show how an organism is changed when exposed to contingencies of
reinforcement and why the changed organism then behaves in a different way,
possibly at a much later date. What he discovers cannot invalidate the laws of a
science of behavior, but it will make the picture of human action more nearly
complete. (Skinner, 1974, pp. 214-215, italics added)

Gaps arise because behavior analysis leaves out the second link of the causal
chain in Figure 2. Basically, behavior analysts manipulate contingencies (the first
link in the chain) in order to the see the effects on behavior (third link). No matter
how detailed, a functional analysis of the relations between the first and third links
will always be incomplete, because it does not include information about the
second, physiological, link. Notice that the gap in analysis is tolerable only if we
assume transitivity between the causal links in the chain. In this context,
transitivity means that if A causes B and B causes C, then A causes C. If
contingencies of selection (A) are responsible for changes in an organisms
physiology (B) and if the latter changes result in behavior (C), then, assuming
these events are orderly related to each other, contingencies of selection are
ultimately responsible for the behavior of organisms. Unless there is a weak spot
in our causal chain so that the second link is not lawfully determined by the first,
or the third by the second, then the first and third links must be lawfully related
(Skinner, 1953/1965, p. 35).
Behavior analysis qualifies as an incomplete science to the extent that it lacks
information about the physiological variables responsible for behavior. But
behavior analysis is still capable of providing explanations by studying the
relations between contingencies and behavior change. The explanatory gap will be
filled by physiology. So in this sense, physiology can also provide explanations
of behavior. Here, however, it becomes paramount to discuss how behavioranalytic and physiological explanations relate to each other. In particular, are we
talking about the same notion of explanation?
From a radical behaviorist perspective, scientific explanation basically
consists in describing functional relations between events (Skinner, 1931/1961c,
1938/1966a, 1947/1961a, 1953/1965, 1966b; cf. Donahoe, 2004; Hayes &

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Brownstein, 1986; Lacey, 1984; Laurenti & Lopes, 2008; Moore, 2008), and an
explanation is valued mainly to the extent that it promotes prediction and control
of the phenomena being explained (Skinner, 1938/1966a, 1953/1965; cf. Hayes &
Brownstein, 1986; Lacey, 1984; Moore, 2010; Palmer, 2009; Smith, 1992). But
caution is needed when dealing with the to-explain-is-to-describe motto often
found in the behavior-analysis literature (e.g., Baum, 1994; Catania, 2007; Reese,
1996), because the mere description of relations between events does not
invariably count as an explanation of these events (cf. Marr, 2003; Tonneau, 2008).
Let us say that a scientist observes the occurrence of an event A and,
following it, the occurrence of another event B. After a long period of observation,
the scientist notes that A always occurs before B, that B always follows A, and that
A and B never occur alone. These observations eventually start to control the
scientists explanatory verbal behavior. She starts saying, every time A occurs,
B occurs and, B is always preceded by A. Explaining takes more than mere
description, however; explaining demands an active role of the scientist in
manipulating the events under study (Woodward, 2003). We can assume a
functional relation between A and B only if we see effects in B when manipulating
A. Even prediction is not enough for explanation. Through a series of observations,
a scientist can predict the occurrence of B after seeing A occur. It is thus
possible to predict events from observation alone, without any kind of
manipulation. A consistent but fortuitous correlation between events can provide
useful information for strictly predictive purposes. Explanation, however, goes
beyond description and prediction to include manipulation and control. If the cooccurrence of A and B is merely fortuitous, then the manipulation of A will have
no impact on the occurrence of B; but if it turns out that we can control B by
manipulating A, then we have solid grounds to assume that A and B are
functionally related.
In sum, explanation is not merely a narrative activity. Explanation requires the
discovery of conditions (or variables) necessary for the production of a given
phenomenon. For Skinner (1957), to explain is to specify what conditions are
relevant to the occurrence of the behaviorwhat are the variables of which it is a
function? (p. 10). The process of discovering these conditions in turn requires
manipulating events and analyzing the effects of those manipulations. To explain a
phenomenon B, as a form of verbal behavior, thus consists in describing the
variables responsible for the production of B. The scientist in our example would
say, every time I produce A, B also occurs or, every time I prevent A, B also
fails to occur. Achieving the stage of functional description (that is, explanation)
requires the active role of the scientist, who is not merely a passive observer.
Assuming this view of explanation, to say that the behavior-analytic
explanation of behavior has gaps means that it does not include some of the
variables that are responsible for the production of behavior. These variables (and
among them, those studied by physiology or neuroscience) should be seen as
important as the ones studied in behavior analysis, for the good reason that in their
absence there would be no behavior to explain. Alongside behavior analysis,
therefore, physiology can provide explanations of behavior by discovering some of

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the variables responsible for its occurrence. Physiology discovers functional
relations between contingencies of selection and physiological processes (first and
second links) as well as between physiological processes and behavior (second and
third links).
Are physiological explanations reductive? Skinner (1963) wrote, The
physiologist studies structures and processes without which behavior could not
occur. He is in a position to supply a reductionist explanation beyond the reach
of an analysis which confines itself to terminal variable (p. 957, italics added).
Skinner use of the term reduction may be misguided, however. Analyzing the
concept of reduction is no easy task (e.g., Bickle, 1998; Horst, 2007; Schaffner,
1967), but it is hard to see how an explanation of behavior that focuses on
contingencies (or history) could reduce to an explanation based on neural
processes. We cannot reduce the occurrence of a stimulus (a historical event) to
any given brain activity, for example (Marr & Zilio, 2013). Admittedly, in some
cases physiological explanations might be used instead of behavior-analytical
explanations, particularly when we are ignorant of historical variables. Skinner
(1953/1965) even said that physiological explanations would be preferable in this
context:
Independent information about the second link would obviously permit us to
predict the third without recourse to the first. It would be a preferred type of
variable because it would be non-historic; the first link may lie in the past
history of the organism, but the second is a current condition. (p. 34, italics
added)

In the absence of knowledge about the real nervous system, however, an

explanation based on historical variables is better than those appealing to an
inferential, conceptual nervous system.

Demarcation Criteria
When Skinner (1986) delineated the causal chain between environmental,
physiological and behavioral events, he also asserted behavior and physiology are
not two ways of approaching the same subject (p. 716, italics added). The same
idea can be found in a later excerpt: brain processes are not another 'aspect of
behavior; they are another part of what an organism does (Skinner, 1989a, p. 56).
Here Skinner suggests a clear-cut separation between the events studied by
behavior analysis (i.e., environmental and behavioral events) and those studied by
physiology; these two sciences do not deal with the same object and therefore
cannot be distinct ways of approaching the same phenomenon.
At other places, however, Skinner suggests that physiological events are part
of behavior and not of a different domain: I have never questioned the importance
of physiology or in particular brain science or its relevance to behavior. What is
happening inside the skin of an organism is part of its behavior (Skinner, 1989a,
pp. 129-130, italics added). This quote suggests the opposite of the earlier one
(1989a, p. 56). Are we facing a contradiction in behavior-analytic thinking? I will

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try to answer this question by discussing two kinds of empirical work, one that
addresses physiological events as parts of behavioral contingencies, the other
dealing with physiology as a mechanism underlying behavior.
With respect to empirical work of the first kind, Skinner (1953/1965) defined
the environment as any event in the universe capable of affecting the organism
(p. 257). The effective environment thus comprises conditional and discriminative
stimuli, and this sense of environment, operant contingencies can incorporate
physiological events as antecedent stimuli, consequences, or responses (BarnesHoles, 2003; Branch, 2006; Silva, Gonalves & Garcia-Mijares, 2007). The effects
of substance administration, for example, can function as discriminative stimuli for
operant behavior, an important issue not only with respect to drug taking but also
with respect to private events (Branch, 2006). Lubinski and Thompsons (1987,
1993) experimental model for the interpersonal communication of private states
thus involves (among other things) the discriminative control of behavior by the
physiological effects of pentobarbital, cocaine, and saline administration. The
important point here is that physiological effects can function as discriminative
stimuli; they can be part of the behavioral relations studied in behavior analysis.
Notice that Lubinski and Thompsons goal was purely behavioral: [to] determine
whether nonhuman animals could learn to interact communicatively (i.e., exchange
discriminative stimuli with each other), based on events in their internal milieu
(1987, p. 1). The background knowledge that informed Lubinski and Thompsons
empirical demonstration was also behavioral: from the definition of private events,
tact and mand, to research on symbolic communication in non-human animals and
to the discriminative effects of drug administration (Lubinski & Thompson, 1987,
1993).
As an example of empirical work of the second kind, consider an experiment
by Campeau, Miserendino and Davis (1992) that also deals with substance
administration, but differs from Lubinski and Thompsons (1987) study in the way
physiological variables are addressed. The difference is already evident in the
authors stated purpose: to test the effects of intra-amygdala infusion of AP5 on
the acquisition and expression of aversive conditioning to an auditory stimulus as
measured with fear-potentiated startle (Campeau et a1. 1992, p. 569). In this case,
the administration of a chemical substance (AP5) is justified by its effects on a
particular neurophysiological mechanism and not because of its function as part of
behavioral contingencies. Indeed, it is known that the amygdalaa group of nuclei
located within the medial temporal lobeis part of the mechanism for establishing
the behavioral relations associated with fear (LeDoux, 2000). The procedure
(called fear conditioning) used to document this role consists of pairing an
aversive stimulus, usually shock, with a neutral stimulus (LeDoux, 2000) and
recording the resulting changes in heart rate, arterial pressure, skin conductance,
hormonal secretion, and motor activity (Davis & Whalen, 2001; LeDoux, 2000).
A detailed description of the physiological mechanisms of fear conditioning
would be beyond the scope of this article. Suffice it to say that in order to increase
the synaptic efficacy relevant to fear conditioning, NMDA receptors in the lateral
amygdala must be unblocked, since they are usually blocked by Mg2+ molecules

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(Sah, Westbrook, & Lthi, 2008). Conversely, any manipulation that maintains the
NMDA receptors in a blocked state should make fear conditioning impossible.
That is exactly the hypothesis tested by Campeau et al. (1992). They maintained
the NMDA receptors in a state of blocking by the intra-amygdala infusion of AP5
((2R)-amino-5-phosphonovaleric acid), a NDMA receptor antagonist. The results
showed that the infusion of AP5 before the learning period (when the subjects
were submitted to the aforementioned contingency) blocked the establishment of
fear conditioning.
Based on these two examples (many more could be found in the technical
literature), we can see that (a) studying the operant discrimination of effects
produced by the administration of certain substances and (b) studying the effects of
the administration of certain substances on a neurophysiological mechanism are
not quite the same thing. We may allude to this distinction as involving a double
aspect of physiological events, provided we remember that the two aspects are
not intrinsic to the events themselves, but only concerns how two scientific
communities interact with a shared object of study (Zilio, 2013). Thus, the
distinction ultimately involves the different contingencies that control the scientific
behavior of behavior analysts and neuroscientists. Physiological events can be
involved in behavioral contingencies, as stimuli (antecedent and consequent) or
responses. If studied as such, they are part of the domain of behavior analysis; they
are part of behavior, as Skinner put it. If, however, physiological events are
studied as mechanisms (or components thereof) associated with the occurrence of
behavior, then they qualify as physiology, in particular neurophysiology. Now they
fit Skinners definition of physiological events as intermediary links between
historical variables and the behavior of an organism. In this context, Skinners
causal chain is best viewed not as a representation of distinct events linearly
related to each other, but as an abstraction that distinguishes the objects of study of
behavior analysis and physiology according to how two scientific communities
interact with the same objects.
The distinction is clearer when we shift from a discussion of the object of
analysis to a discussion of the role of physiology in the explanation of behavior.
According to Skinner, the overall goal of physiology as a scientific discipline is to
explain the physiological mechanisms associated with the occurrence of behavioral
relations: How contingencies of operant reinforcement affect physiological
processes is no doubt an important question (Skinner, 1988/1989c, p. 82). Again:
The behaving organism will eventually be described and explained by the
anatomist and physiologist. As far as behavior is concerned, they will give us an
account of the genetic endowment of the species and tell how that endowment
changes during the lifetime of the individual and why, as a result, the individual
then responds in a given way on a given occasion. (Skinner, 1975b, p. 42)

This excerpt is important because it describes the how and why of

physiology according to Skinner. The how points to the study of physiological
mechanisms that allow an organism to respond to the environment and how these
mechanisms change during the organisms interactions with the environment. The

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why, on the other hand, stands for explaining why an organism behaves in such a
way at a given time. Why questions are typically associated with the behavioranalytic study of phylogenetic and ontogenetic contingencies. Skinner, however,
implies that physiology can also offer its own answers to why questions by
clarifying the functional relations between physiological processes and behavior.
The same idea showed up elsewhere in the Skinnerian corpus: A great deal
goes on inside the skin, and physiology will eventually tell us more about it. It will
explain why behavior is indeed related to the antecedent events of which it can be
shown to be a function (Skinner, 1971b, p. 195). Put differently, physiology will
explain why there are ordered functional relations between the first and third
links (phylogenetic and ontogenetic histories, and behavior, respectively) in
Skinners causal chain.

Skinners Reliance on Physiology: Some Examples

Beyond his general comments on the objects and goals of physiology and
behavior analysis, Skinner also raised specific issues about behavior that may have
required a physiological answer. Some of his interpretations and conceptual
analyses about behavior also involved physiological hypotheses. In this section I
will discuss a few cases of Skinners interpretations that rely on physiology.
Motivation, discrimination and reinforcement
One of the issues that Skinner thought would be answered by physiology
concerned the role of motivational variables in the explanation of behavior:
One of my first papers [Skinner, 1932] was on the state of hunger (or drive'),
and I have been interested in states off and on ever since. The organism we
observe and possibly study can certainly be said to be in a given state at a given
time. The state will eventually be directly observed by those who have the
proper instruments and methods - namely, anatomists and physiologists. They
will complete the account offered by an experimental analysis of behavior,
which necessarily has temporal and spatial gaps. (Skinner, 1988, p. 122)

Hunger or drive are commonly viewed as motivational states. What

Skinner is saying here is that these states should be studied directly (through direct
observation and manipulation instead of inference) and therefore physiologically. I
have already discussed this example in relation to a history of deprivation and the
resulting behavior. Elsewhere, Skinner (1966b) explicitly relates the issue to
motivation and emotion: other independent variables are found in the classical
fields of motivation and emotion. The experimental analyst does not manipulate
inner states as such (p. 215).
A similar idea can be found in Skinners (1988) remarks on a study of
discriminative control:
The experiment throws additional light on the nature of the control exerted by
stimuli. Eventually a physiological explanation will be available. Meanwhile, I

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do not see that anything is gained by referring to a hidden mechanism. On the

contrary, representing a datum with a hidden mechanism makes it harder to
integrate with other data from other experiments, which could also be done
without postulating a hidden mechanism (p. 133).

Later on Skinner (1988) argues: as a behavioral problem, discrimination

raises no question if one stops talking about input-processing mechanisms. The
behavioral facts are well known. They are often astonishing, and a neurological
explanation is badly needed (p. 471). According to Skinner, physiology could
also explain (at least in part) the difference between elicitation and discrimination:
a discriminative stimulus which brings about the emission of a response (which
'sets the occasion' for the response) differs quantitatively in its action from the
eliciting stimulus and must be 'explained' by a different neural mechanism
(Skinner, 1938/1966a, p. 430). In this context, Skinner (1988) again stresses that
only the physiological variables that are directly observable and susceptible of
manipulation (as opposed to hypothetical, inferential links) are worth pursuing. No
less than in the case of motivational and emotional states, a non-hypothetical
physiology should be able to shed light on the mechanisms that mediate behavioral
processes of elicitation and discrimination.
The study of the physiological mechanisms of reinforcement is another clear
example of how Skinner relied on neuroscience to elucidate important behavioral
issues. This seems to especially important in the light of the alleged circularity in
the concept of reinforcement (e.g., Skinner, 1953/1965, 1957/1961d, 1971a, 1972,
1973, 1988; see also: Paniagua, 1985; Prado Jr., 1982; Schick, 1971; Tonneau,
2008). In Skinners (1953/1965) words:
The only defining characteristic of a reinforcing stimulus is that it reinforces.
The only way to tell whether or not a given event is reinforcing to a given
organism under given conditions is to make a direct test. We observe the
frequency of a selected response, then make an event contingent upon it and
observe any change in frequency. If there is a change, we classify the event as
reinforcing to the organism under the existing conditions. There is nothing
circular about classifying events in terms of their effects; the criterion is both
empirical and objective. It would be circular, however, if we then went on to
assert that a given event strengthens an operant because it is reinforcing. (pp.
72-73)

Behavior analysis provides a classificatory description of presently

reinforcing events. If we want to go further and know why reinforcers reinforce,
then we should turn to other scientific fields, including physiology: a biological
explanation of reinforcing power is perhaps as far as we can go in saying why an
event is reinforcing (Skinner, 1953/1965, p. 84); in the field of behavior, it is
the reinforcing consequences which alter the probability that an organism will
subsequently behave in a given way. The physiological mechanisms which make
this possible (themselves a product of natural selection) are not yet known
(Skinner, 1973, p. 264); finally:

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Another possibility is that neurological conditions may be arranged which will
have a positive effect. A step in this direction has been taken by James Olds with
his discovery that weak electrical stimulation of certain parts of the brain,
through permanently implanted electrodes, has an effect similar to that of
positive reinforcement. In one of Olds' experiments, a rat presses a lever to give
itself mild electrical stimulation in the anterior hypothalamus. When every
response is so "reinforced," behavior is sustained in strength for long periods of
time. () Although there remain some puzzling differences between behavior
so reinforced and behavior reinforced with food, Olds' discovery in an important
step toward our understanding of the physiological mechanisms involved in the
operation of the environmental variable. (Skinner, 1957/1961d, pp. 122-124)

Skinner rarely mentioned specific examples of research in neuroscience, so

his reference to Olds work should not be taken lightly. Skinner may have given
credit to Olds research because the latter was dealing with the nervous system in
actual physiological terms and not as a conceptual/inferential system.
Private events
In Skinners remarks on private events we find him not only raising issues to
be addressed by physiology but also advancing physiological hypotheses that
would support his own conceptions. In a generic sense, private events involve
behavioral relations (i.e., relations between stimuli and responses) that are
accessible only to the behaving person (cf. Moore, 1980, 2009; Palmer, 2009;
Tourinho, 2006a, 2006b). I have argued elsewhere (Zilio, 2010; Zilio & Dittrich,
2014) that the core of Skinners theory of private events is not so much their lack
of accessibility than the special kind of contact that makes private relations
possible. Here is Skinners (1963) take on the subject:
The fact of privacy cannot, of course, be questioned. Each person is in special
contact with a small part of the universe enclosed within his own skin. To take a
noncontroversial example, he is uniquely subject to certain kinds of
proprioceptive and interoceptive stimulation. Though two people may in some
sense be said to see the same light or hear the same sound, they cannot feel the
same distension of a bile duct or the same bruised muscle. (When privacy is
invaded with scientific instruments, the form of stimulation is changed; the
scales read by the scientist are not the private events themselves). (p. 952, italics
added)

Behavioral relations do not float around but require anatomical and

physiological support, and for something in the world to become a stimulus,
some kind of sensory apparatus is needed. According to Skinner (1953/1965, 1963,
1969b, 1972, 1974), there are three ways of contacting the world: (a) via the
exteroceptive nervous system, responsible for contacting the environment that is
also accessible to others (through their own exteroceptive nervous system); (b) via
the interoceptive nervous system, responsible for contacting the digestive,
circulatory and respiratory systems; and (c) via the proprioceptive nervous system,
responsible for contacting posture and movement. It is proprioceptive and

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interoceptive stimulations that make private behavioral relations possible, and

these relations are private to the extent that they are supported by a specific
physiological apparatus. Proprioceptive and interoceptive stimuli may have a
certain intimacy. They are likely to be especially familiar. () They may well be
of a special kind (Skinner, 1963, p. 953). In Skinners (1963) example, two
persons may be said to see the same light because both are contacting the stimulus
via their exteroceptive nervous system. One cannot feel another persons pain,
however, because the contact with the source of stimulation differs among people.
Dentists access the inflamed teeth of their patients via the dentists exteroceptive
nervous systems. In contrast, patients access their own inflamed teeth via their own
interoceptive and proprioceptive nervous systems (unless the patients also happen
to look at their teeth in a mirror).
One important consequence of the special contact definition of private
events is that they are not accessible through scientific instruments such as
microscopes, brain scans, and so on. The latter only enhance scientists
exteroceptive contact with the source of stimulation. They do not turn private
relations into public ones. Regardless of which apparatuses she relies on, the
dentist does not contact the source of stimulationan inflamed tooththe way her
patient does. As Skinner (1957/1961b) puts it: As physical states in the
individual, [private events] are a part of the physical world, but the individual
himself has a special connection with them. My aching tooth is mine in a very real
sense because none of you can possibly get nerves into it (p. 201, italics added).
Note that private does not means internal or inside the skin. These
connotations are metaphorical (and imprecise) descriptions at best. One stimulus is
private, not because it is part of an organisms physiology as opposed to the
external environment, but because the organism has as special contact with this
stimulus. This is why it is impossible to invade privacy. As Skinner (1963)
argued, as soon as others access the source of stimulation, the form of stimulation
is changedit is not private anymore. None of this implies, however, that
exteroceptive studies of a source of private stimulation are pointless; on the
contrary, these studies can supply important information about private events
(Skinner, 1957). The dentist can diminish the patients pain by delivering an
anesthetic to the inflamed tooth. Merely seeing the inflamed tooth is additional
evidence (along with vocal and facial expressions) that the patient is telling the
truth, that is, that he is really in pain. So, it usually matters to know the
characteristics of a source S of private stimulation because S is an additional
variable that can control our evaluating others verbal descriptions of private
events. Knowing more of S is also important for teaching members of our social
community to discriminate private stimuli more precisely: if events hitherto
classified as private can now be directly observed by the verbal community, the
community can arrange better contingencies in teaching its members to talk about
them (Skinner, 1969a, p. 262).
A similar analysis applies to covert behavior, with some differences worth
discussing. According to Skinner (1953/1965, 1963, 1969a, 1972, 1974), we
contact physiological stimulation privately through our interoceptive and

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proprioceptive nervous systems. Could there also be pathways responsible for

private relations and located wholly in the central nervous system (Moore, 2001,
2002, 2009; Natsoulas, 1983, 1985) Schnaitter (1984) answers affirmatively:
Skinner seems to assume that a sensory receptor in the peripheral nervous
system must be stimulated in order for a feeling to occur. This is simply not so.
For example, receptors exist within the central nervous system itself.
Osmoreceptors located in the hypothalamus are functionally related to drinking
and presumably to the experience of thirst. But it is not necessary for any
receptors to be stimulated at all for experiences to occur? The phenomenon of
dreaming is probably the clearest illustration of this. The current
neurophysiology of dreaming suggests that an endogenously cycling brainstem
structure regulates midline thalamic pacemaker projections to the cerebral
cortex: cortical desynchrony is correlated with the reported dream state. (p. 7)

The assumption that private relations involve only interoceptive and

proprioceptive stimulation is hardly essential to Skinners views on privacy,
however. Consider the following:
I agree that the kind of thinking which seems to be merely covert behavior
(truncated, unemitted, reduced, impotent behavioral acts) may be so reduced
that there is no muscular involvement to be sensed proprioceptively. Must we
appeal to some minute behavior which never reaches a muscle? If so, it is a
problem for the physiologist. There is a possibility that the effect is sensory. We
may see ourselves behaving rather than actually behave. I believe that my
analysis of seeing makes this a possible alternative. (Skinner, 1988, p. 331)

Thus, private behavioral relations can occur without any involvement of

muscular activity or even interoceptive and proprioceptive stimulation. Arguably,
this is the core of what we usually define as covert behavior (loosely speaking:
thinking, problem solving, imagining, internal speech, and so on). The
nervous system is massively interconnected. Brain areas responsible for a given
functions are connected to other areas responsible for other functions (cf. Baars &
Gage, 2010; Gazzaniga, Ivry, & Mangun, 2002). Brain areas related to verbal
behavior, for example, are connected to other areas associated with mental
images (cf. Bartolomeo, 2008; Farah, 2000; Gulys, 2009; Kosslyn, Thompson &
Ganis, 2006). So when a person covertly says to himself (internal speech) that he
is seeing Picassos Guernica in the absence of the actual painting (seeing in the
absence of the thing seen), he may very well be responding discriminatively (and
privately) to the neurophysiological effects once elicited by the actual painting but
that are now under control of other variables.
To dispel possible misunderstanding, I have not been arguing that differences
between private and public events are only a matter of anatomy or physiology.
What I have been saying is that anatomo-physiological considerations are a key
factor in Skinners distinction between private and public events. Physiology plays
an important part in the study of private relations by providing knowledge about
variables that we can use in teaching and assessing descriptions of private events

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(and not because it can be used to make private relations public; see Zilio, 2010;
Zilio & Dittrich, 2014).
Perception
Skinners theory of perception is an especially important example of his
appealing to physiology in solving theoretical issues in behavior analysis. His
theory essentially is a behavioral analysis of perception (cf. Lopes & Abib, 2002;
Zilio, 2010) that incorporates respondent and operant perceptual relations. In both
cases, the same logic applies: an antecedent condition (1) sets the occasion for a
perceptual response (2) that in turn may act as a discriminative stimulus (3) for a
persons verbal description of perceptual experience (4).
Among the antecedent conditions of perception are unconditional stimuli. In
this case, stimuli are responsible for eliciting unconditional perceptual responses
(Skinner, 1953/1965). The visitor of the Reina Sofa Museum in Madrid sees
Picassos Guernica in front of him. As a complex unconditional stimulus, the
painting elicits unconditional visual responsesthat is, neurophysiological
changes produced by stimulation (Zilio, 2010). According to Skinner (1963),
when a man sees red, he may be seeing the physiological effect [the visual
response] of a red stimulus; when he merely imagines red, he may be seeing the
same effect [the visual response] re-aroused (p. 957).
Now assume that the visitor of the Reina Sofa Museum also listens to
Beethovens Appassionata when in front of Guernica. This produces the Pavlovian
pairing of an auditory stimulus (Beethovens Appassionata) and a visual stimulus
(Picassos Guernica). As a result, every time the visitor listens to this particular
sonata, visual responses of Guernica will be elicitedan example of what Skinner
(1953/1965) called conditional visual responses. Assume further that the visitor
comes back to his home country and tries to remember the details of Guernica a
few months later: that is, he manipulates his environment so as to increase the
probability of responses of the class, seeing Guernica. He plays Beethovens
compositions on his stereo, read books about Picasso, and so on. He may also emit
operant visual responses (Skinner, 1953). After manipulating the environment so
as to see Guernica in the absence of the painting, the visitor now says: I am
seeing Guernica. In this case, the visual response is acting as discriminative
stimulus for the verbal description Im seeing Guernica (Skinner, 1963). In this
context, Skinner is well known for distinguishing seeing something from seeing
that we are seeing it:
Someone shows you a picture of a group of scientists, among them Einstein. He
asks, "Is Einstein there?" and you say, Yes, as you have been taught to do in
thousands of comparable cases. But suppose he asks, Do you see Einstein?
and you say, Yes. What have you reported? Did you, in response to his
question, simply look at Einstein a second time? If so, how do you distinguish
between seeing Einstein and seeing that you are seeing Einstein? A
possibility, which needs to be considered, is that in reporting that you see
Einstein, you are reporting a response rather than a stimulus. No matter how

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obscure its dimensions, the behavior called seeing must be involved, and you
must be reporting it, rather than the presence of the thing seen, when you report
that you see something. You may be reporting the same thing when you report
that you see something which isn't really there - when you are merely
imagining how Einstein looked. Seeing something in memory is not
necessarily seeing a copy. () When I recall how something looked, I may
simply be recalling how I once looked at something. There was no copy inside
me when I first looked at it, and there is none now. I am simply doing again
what I once did when I looked at something, and I can tell you that I am doing
so. (Skinner, 1967, pp. 329-330)

Skinners theory of seeing contrasts with the notion that perceptual responses
in the absence of the thing seen are cognitive and/or neural representations of the
original stimulus in the minds eye (cf. Lopes & Abib, 2002; Tourinho, 1994;
Zilio, 2010). In Skinners theory of perception there is no need for such copies or
representations: what is seen is the presentation, not a representation (Skinner,
1985, p. 292).
Figure 3 summarizes Skinners model of perception. Perceptual activities are
viewed as responses (and not simply as effects caused by stimulation) because they
are sensitive to the history of organism: They can occur as a function of operant or
respondent contingencies. In this sense, they are no less behavioral than motor
responses. Also, just as motor responses of different intensities and topographies
are classified as bar pressing as long as they fulfill a common criterion (e.g., the
deflection of the lever), perceptual responses qualify as seeing Guernica (for
example) because of their shared discriminative effects (saying, I see Guernica).
As in the motor case, perceptual stimulation may not cause identical neural
changes each time it occurs. The responses can and do vary, but are lumped in the
same class when we describe what we are seeing as something that we saw
before. As in the motor case, there is a limit to within-class variation too.
Evolving perceptual responses can differ from the original one to such an extent
that the visitor stops saying he is seeing Guernicahe now sees something else.

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Figure 3: Skinners model of perception. 1, 2, and 3 indicate an unconditional visual

relation, a conditional visual relation, and an operant visual relation, respectively.
Continuous arrows with filled and unfilled arrowheads represent relations of elicitation
and discriminative control, respectively. Dotted arrows with unfilled arrowheads represent
stimulus pairings.

Neurophysiological hypothesizing plays a complex role in Skinners theory of

perception. Skinner assumes that different kinds of stimuliunconditional,
conditional, and discriminativecan produce perceptual responses of the same
class, that is, similar neurophysiological effects. He also assumes that perceptual
responses occur even when we see something in the absence of the thing seen
and that we can in turn respond to these responses, which are now acting as
discriminative stimuli. The plausibility and validation of Skinners theory of
perception ultimately depends on what neuroscience will tell us about the workings
of the brain. As it stands, recent neurophysiological findings seem to indicate that
Skinner was on the right track. Although his take on neuroscience may appear
primitive by current standards, more recent research (e.g., Borst & Kosslyn, 2008;
Cichy, Heinzle, & Haynes, 2012; Kosslyn, Thompson & Ganis, 2006; Kreiman,
Koch & Fried, 2000) has uncovered important similarities between the
neurophysiological processes related to visual perception (unconditional vision)
and seeing in the absence of the thing seen (also known as mental imagery).
Kosslyn, Thompson and Ganis (2006), for example, conclude that visual imagery
evokes many of the same processing mechanisms used in visual perception (p.
135). As Skinner anticipated, the perceptual responses produced by unconditional
relations and those under other kinds of stimulus control (such as conditional and
discriminative) are clearly similar to one another.

Final Thoughts
The main goal of this paper was to discuss some of Skinners positive (noncritical) views about physiology. A first step in the argument was to clarify the

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differences, according to Skinner, between the domains of behavior analysis and
physiology as well as the role of both sciences in the explanation of behavior. I
also discussed specific examples from Skinners works of how physiology could
inform behavior analysis by providing explanations of motivational phenomena,
discrimination, elicitation, and the neural mechanisms of reinforcement. Close
attention was given to Skinners use of physiological notions in developing his
theories of private events and perception. The Skinnerian concept of privacy is
based, at least in part, on the special contact afforded by different physiological
systems, and his criticisms of representational theories of imagery embody a
specific behavioral/neurophysiological theory of perception.
These considerations converge toward one conclusion: Skinner never
defended an anti-physiological position on behavior. It is one thing to focus on
historical variables and to leave the study of other factors to physiology, quite
another to deny the importance of the latter. Skinner can hardly be accused of that.
In his own words: A behavioral account is incomplete, in part because it leaves a
great deal to neurology (1985, p. 297).

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Authors Note
Writing this article was supported by FAPESP - So Paulo Research Foundation
(2009/18324-1/2013/17950-1). The article itself is based on a talk presented at the
1st Amazonian Simposium on Behavior Research and Theory (Universidade
Federal do Par, Belm, Brasil, November 2013). I would like to thank Maria
Helena Leite Hunziker and Franois Tonneau for their helpful comments on earlier
drafts. Correspondence concerning this article should be addressed to Diego Zilio,
Federal University of Esprito Santo (UFES)- Department of Social and
Developmental Psychology, Fernado Ferrari Ave, 514. Goiabeiras. Vitoria- ES.
Brazil. 29075-910

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