Journal of Experimental Psychology: Copyright 199d by the American Psychological Association, Inc.

Animal Behavior Processes 0097-7403/96/S3.00

1996. Vol. 22, No. 1. 3-18

Dimensions of Stimulus Complexity

J. Gregor Fetterman
Indiana University-Purdue University Indianapolis

Animal learning research has increasingly used complex stimuli that approximate natural
objects, events, and locations, a trend that has accompanied a resurgence of interest in the role
of cognitive factors in learning. Accounts of complex stimulus control have focused mainly
on cognitive mechanisms and largely ignored the contribution of stimulus information to
perception and memory for complex events. It is argued here that research on animal learning
stands to benefit from a more detailed consideration of the stimulus and that James Gibson's
stimulus-centered theory of perception serves as a useful framework for analyses of complex
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stimuli. Several issues in the field of animal learning and cognition are considered from the
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Gibsonian perspective on stimuli, including the fundamental problem of defining the effective
stimulus.

In 1989, I participated in a conference at Dalhousie Uni-
versity organized by W. K. Honig. The participants were
experts in the field of learning and memory in nonhuman
animals, with interests in stimulus control, broadly defined,
involving complex stimulus events. The working title of the
symposium, the Dalhousie Conference on Complex/Ex-
tended Stimuli, was inspired by the recognition that stimu-
lus control research had increasingly incorporated complex,
naturalistic events in place of the simpler stimuli that had
been used in the past. But the working title gave way to one
that more accurately reflected dominant trends that empha-
size the role of cognitive processes in animal behavior
(Hulse, Fowler, & Honig, 1978). The conference was re-
named the Dalhousie Conference on Cognitive Aspects of
Stimulus Control, which became the title of the edited book
based on the proceedings (Honig & Fetterman, 1992).
As the preceding paragraph indicates, in the last 15-20
years psychologists have witnessed the rapid development
(some might say, a reemergence) of a subdiscipline of our
field devoted to the study of cognitive processes in nonhu-
man animals (see Hulse, 1993, for a recent commentary).
This growth is represented in many ways—through the
Editor's Note. This article is another in a series on various
topics to be invited by the Editor.
Preparation of this article was assisted by National Institute of
Mental Health Grant MH 48359 and National Science Foundation
Grant BNS 9407527. Some of these ideas appeared in earlier
publications including Stubbs, Dreyfus, and Fetterman (1984),
Fetterman, Stubbs, and MacEwen (1992), and Fetterman (1993).
I am most grateful to Alan Stubbs for many discussions over the
years on the issues raised in this article and for his helpful sug-
gestions on various drafts. I also extend thanks to Leon Dreyfus,
Peter Hanford, Vern Honig, Kathy Johnson, Peter Killeen, and
Geoff White for their comments on the article.
Correspondence concerning this article should be sent to J. Gre-
gor Fetterman, Department of Psychology, Indiana University-
Purdue University, 402 North Blackford Street, Indianapolis, In-
diana 46202. Electronic mail may be sent to gfetter@IndyVax.
IUPUI.EDU.
publication of edited books on animal cognition (Honig &
Fetterman, 1992; Hulse et al., 1978; Roitblat, Bever, &
Terrace, 1984; Zentall, 1993); from special issues of jour-
nals (e.g., Cognition in 1990, Journal of the Experimental
Analysis of Behavior in 1989, Learning & Motivation in
1987) and specialized conferences (e.g., Conference on
Comparative Cognition) and symposia and paper sessions
devoted to research on cognitive processes in animals;
through review papers (Roitblat & Von Fersen, 1992; Was-
serman, 1993); and, of course, from the empirical and
theoretical content of published research. These examples
are very familiar to the specialist, but the recent resurgence
of work on animal cognition has also contacted a broader
audience, and exemplars of this line of research are now
frequently included in introductory psychology textbooks
(e.g., Baron, 1995).
A detailed discussion of events that have contributed to
the resurgence of research on animal cognition is beyond
the scope of this article, however, to set the stage for what
follows, several contributing factors are noted. First, re-
search on animal learning and memory has been influenced
by theories, concepts, and methodologies from human cog-
nitive psychology. Numerous studies of animal cognition
have been predicated on prior research with humans (e.g.,
Blough, 1992; Cook, 1992b; Fountain & Rowan, 1995), and
these investigations have formed the basis of a modern
psychology of comparative cognition, resulting in both em-
pirical and theoretical advances. Second, a good deal of
the recent research on animal learning and memory has
used complex stimulus arrangements—arrays of elements
(Blough, 1979), textured patterns (Cook, 1992a), color pho-
tographs (Herrnstein, 1979), musical selections (Porter &
Neuringer, 1985), moving stimuli (Dittrich & Lea, 1993),
spatial locations (Olton & Samuelson, 1976), and the like—
stimulus events now easier to arrange and study because of
computers and related advances in technology such as touch
screen devices (Blough, 1986; Pisacreta & Rilling, 1987).
These stimuli seem more ecologically valid than the simple
unidimensional stimuli that traditionally have been used in
research on animal learning (e.g., lights of different wave-
 FETTERMAN
lengths, tones of different frequencies), and coping with
these complex events seems to place increased demands on
information-processing resources. The structure and func-
tion of these resources are, of course, at the core of current
research on animal cognition.
Whereas analyses of animal behavior in terms of under-
lying cognitive mechanisms (expectancies, attentional con-
straints, memory representations, common codes, etc.) have
gained widespread (but not universal) acceptance, I argue in
this article that our field stands to benefit by undertaking an
in-depth analysis of the complex stimuli that have become
commonplace in modern research in animal learning. There
is no shortage of cognitive theory that purports to explain
how animals encode, process, remember, and retrieve infor-
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mation, and how these internal processes are related to
behavior. However, there exists no corresponding theory of
the stimulus that concerns the information that is immedi-
ately available from stimuli. One might argue that the
veridicality of the coupling between distal (the stimulus
object) and proximal (the stimulus at the receptor) stimula-
tion dictates the contribution of cognitive mechanisms.
With Stevens (1951), I regard the definition of the effective
stimulus as a most important problem for psychology. How-
ever, conceptual analyses of cognitive processes in animals
have advanced much more rapidly than analyses of the
informational content of complex stimuli, as suggested by
the change in the title of the Dalhousie conference.
Research and theory on human perception has a long-
standing concern with the interplay between distal and
proximal stimuli (e.g., see Hochberg, 1988), but stimulus
analysis in the field of animal learning comes instead from
a different tradition. Research on stimulus control in ani-
mals is rooted in work on reflexology, in which stimuli were
treated as physical goads for action (e.g., see Staddon,
1983), in sensory psychology (Honig, 1993), and in the
influence of the stimulus-response (S-R) tradition in the
field of animal learning (Balsam, 1988; Hulse, 1993). Bal-
sam noted that the study of learning within the S-R frame-
work was viewed as a matter of finding out how stimuli that
previously did not evoke a response were later able to do so.
The framework was rarely questioned, only the conditions
necessary for establishing S-R bonds were investigated.
In the past, the study of stimulus control in animals (e.g.,
Terrace, 1966), as in most scientific study, relied on a
strategy of simplification and analysis of stimuli into ele-
mental units. Thus, simple, discrete, unidimensional stimuli
were used, which permitted precise control over stimulus
events and provided the psychological equivalent of the
atom, stimulus elements that could serve as the building
blocks of more complex objects that compose the natural
world. The modern prototype of this line of research was
described by Guttman and Kalish (1956), who provided an
elegant demonstration of stimulus control along the wave-
length dimension. In the ensuing 40 years, the experimental
analysis of stimulus control has proceeded mainly along the
lines of the Guttman and Kalish prototype (Honig & Ur-
cuioli, 1981), investigating many aspects of stimulus con-
trol, such as the role of attentional factors (Mackintosh,
1977) and inhibitory processes (Rilling, 1977).
The benefits of simplification have a complement in the
problems engendered by the use of more complex stimuli,
stimuli that may be more representative of the natural en-
vironment but more difficult to analyze in terms of individ-
ual stimulus features that control responding (Cook, 1993).
Such stimuli are not readily manipulable in a way that lends
itself to rigorous experimental analysis. As noted, however,
the trend in recent years has been toward increasing use of
more complex stimulus events. In this article, I consider this
trend from the standpoint of stimulus analysis in the field of
human perceptual psychology and argue that current con-
cepts of the stimulus as an elemental unit of environmental
description are inadequate. Research on animal cognition
stands to benefit by considering the role of the stimulus
from the perspective of perceptual theory and, specifically,
from theories that adopt a less simplistic use of the stimulus
concept (cf. Schleidt, 1985).
Stimuli and Cognition
A fundamental question in the field of perception con-
cerns the relative contributions of the stimulus and the
organism to veridical perceptions. The prevailing view of
proximal stimulation, owing largely to the influence of
Helmholtz, is that the receptor surfaces do not afford suf-
ficient information to account for the perception of distal
properties of objects and events, thereby requiring cognitive
mechanisms (unconscious inferences) to piece the impov-
erished stimulus information together (see Hochberg, 1988,
for a review). By this account, elemental stimuli serve as the
basic units of perception, and the analysis of perception
involves figuring out how perceptual systems synthesize
simple stimuli into complex patterns. The flavor of the
elementarist point of view, in which the explanatory burden
is carried by cognitive mechanisms, is nicely captured in a
recent article by Graham (1992):
The visual system first breaks the information that is con-
tained in the visual stimulus into parts; then the visual system
puts the information back together again. But why take it apart
in the first place? Because the proximal stimulus—the light
falling on the retina—bears little direct resemblance to the
important aspects of the world that must be perceived, that is.
to the distal stimulus, (p. 55)
Graham's (1992) position is representative of views that
regard complex stimuli as composites of more elemental
features, but other views adopt a different perspective on the
nature of stimuli and on the role of cognitive mechanisms in
perception. The Gestalt psychologists, for example, viewed
an entire melodic sequence of notes as a single stimulus, and
such stimuli did not need to be elaborated by cognitive
structures. J. J. Gibson (e.g., 1966b) argued that the com-
plexities of distal events were directly revealed in proximal
stimuli, with no need for further elaboration or interpreta-
tion, or both, by cognitive processes (a view referred to as
direct perception). Juxtapose Graham's remarks with a
view offered by a distinguished proponent of J. J. Gibson's
position:
 COMPLEX STIMULI
I do not believe that it is essential to invent mechanisms for
assembling what we know we actually do perceive: things, a
spatial layout, events that take place in it. There is structure in
the array, relational information that does not have to be
pieced together because, like truth, it is already there. . . .
Living organisms search for information for invariant struc-
ture (in fact, an objective one in which not only I but other
animals too can locate things); of objects, with constant di-
mensions (not shrinking, expanding, or losing and gaining
substance as we move around) and reliable affordances that
meet our requirements for survival; and of events, in which
things happen with predictable relations (not randomly and
chaotically).... If I look for mechanisms therefore, they are
going to have to be ones that deal with relations and ones that
abstract information, rather than add on or integrate pieces.
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(E. J. Gibson, 1977, pp. 157, 159).
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The basic difference between these two approaches is
illustrated by the title of an article by Mace (1977), "James
Gibson's Strategy for Perceiving: Ask Not What's Inside
Your Head, but What Your Head's Inside of." Most readers
are familiar with findings on neural coding and feature
detectors, and they are familiar with cognitive theories of
various kinds, approaches that emphasize the role of internal
processes. Gibson and others (e.g., Cutting, 1986) do not
emphasize what goes on internally, but rather what happens
out in the world. Everyone is familiar with technological
advances that have afforded ever more realistic depictions
of the world. Renaissance painters created more realistic
works once they gained knowledge of perspective and prin-
ciples such as shading, haze, and relative size. The motion
picture industry has changed from black and white silent
films, to technicolor, to wide-screen color films with sur-
round sound, making the movie experience more realistic.
We are in the early stages of movie rides, where the screen
fills the visual field and the individual's chair moves in
relation to the visual changes. Audio recordings have
changed from Edison disks to hi-fi records to compact disks.
These advances have come about from increased knowledge
of the relevant stimuli, not from internally generated
changes in perception.
In the next section I consider in greater detail different
positions on the role of stimuli in perception and cognition.
The different views define a continuum that, at one extreme,
holds that many of the complexities of environmental events
are directly revealed in proximal stimulation (cf. E. J. Gib-
son, 1977); at the opposite end of this continuum processing
and elaboration of elemental stimuli are regarded as prereq-
uisites for perception (cf. Graham, 1992).
Parsing the Contributions of the
Stimulus and Organism
James Gibson's theory of perception (e.g., 1950, 1966a,
1966b, 1979) is an obvious point of departure for a detailed
consideration of the stimulus, because he outlined a frame-
work for perceiving in which the stimulus played a central
role. Traditional views held that the proximal stimulus for
visual perception consisted of a succession of mosaics of
light intensities on the retina from which the distal proper-
ties of the world were recovered through a reconstruction of
sensory elements into a unified perceptual whole (e.g.,
associations between successive snapshot views of the
world). By contrast, Gibson argued that the stimulus infor-
mation available as an organism moves about was much
richer than had been recognized previously and that this
information exactly specified objects and events in the
environment.
One of Gibson's key insights was the observation that,
whereas many local changes in proximal stimulation occur
at the receptor level, certain higher order patterns remain
invariant and specify both permanent properties of the en-
vironment and egocentric changes with respect to the world.
Gibson argued that perceptual systems have evolved to
extract this higher order information, which may be partic-
ular to one niche (e.g., perception of bird song) or may be
characteristic of any niche (e.g., space and time). According
to Gibson, these higher order stimuli—invariants—serve as
the stimuli for the perception of the environment.
The information about distance provided by a gradient of
deformation provides a good example of a stimulus invari-
ant (J. J. Gibson, 1950). Gibson pointed out that the rate of
change of a given point on the retinal image is inversely
proportional to the distance of its distal source. When we
move forward 1 m, for example, an object 2 m away
becomes 1 m away and is 50% closer (with greater relative
change), an object 10 m away becomes 9 m away (10%
closer), and an object 100 m away becomes 99 m away (1%
closer). Moving forward in the world produces continuous
change in visual stimulation, but at the same time there is an
invariant pattern of change over time involving the dis-
tances of objects, with objects 1 m, 10m, and 100 m away
undergoing different relative amounts of change. This pat-
tern is invariant over different contexts; as we walk forward
on a tiled floor, a rug, or on the beach, the relative gradient
changes at all distances as our pace speeds up or slows
down.
James Gibson and Animal Cognition
Several aspects of Gibson's views bear on the research
agenda for the field of animal learning and cognition and,
particularly, for research on complex stimulus control. The
first is a conceptual distinction between the energetic and
informational value of a stimulus (Garner, 1986). Research
on stimulus control in animals has typically used simple
stimuli whose energetic properties could be easily manipu-
lated (e.g., tone frequencies). Gibson pointed out, however,
that energetic variables do not convey the information nec-
essary for perception of complex events in the natural
world; such information is provided by invariants in the flux
of stimulation that he referred to as the optic array in the
case of vision.
Second, the assumption that perception normally involves
learning to abstract higher order invariants (cf. E. J. Gibson,
1977) appears to be relevant to a substantial literature con-
cerning the capacity of nonhuman animals for abstract
learning; it is common to find that animals have difficulty
 FETTERMAN

acquiring discriminations based on abstract properties of reconstruct the positions of pieces on a board when the
stimuli such as the same-different concept (e.g., Carter & arrangement is a result of moves in an actual game; amateur
Werner, 1978; Premack, 1978). Such observations do not chess players are not nearly as accurate in reconstructing the
square with a view that everyday perception involves a array. When, however, the arrangement of the pieces is
process of abstraction unless, of course, perceptual systems random, chess masters fare no better than the amateur.
are attuned to the higher order regularities of natural envi-
ronments and, as a consequence, organisms readily learn to
extract only these regularities (Shepard, 1984). Alternative Positions
Third, Gibson (e.g., 1960, 1966a) proposed that a stimu-
Whereas Gibson argued that directly apprehended invari-
lus may endure over time and space, in contrast to tradi-
ant stimulation serves as the basis of perception, most
tional views in which temporally extended stimuli have
psychologists ascribe a much larger role to cognitive mech-
been construed as composites of localized elements (Honig,
anisms. For example, Julian Hochberg (1982, 1988) has
1993). When discrete stimuli are spread out over time,
strongly emphasized the role of mental structure in percep-
explanatory accounts seemingly must include memory
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tion in the classic Helmholtzian sense. Hochberg has also

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mechanisms (the problem of action across temporal distanc-

taken issue with Gibson's position on a number of grounds.
es). Gibson, however, did not subscribe to the sharp demar-
For instance, he pointed out that whereas there are numer-
cation between perception and memory (the razor's edge
ous invariances in distal stimulation, there is not a great deal
that separates the present from the past and the future) that
of evidence about whether organisms respond directly to
is characteristic of most of psychology. This issue is rele-
this higher order information (see Hochberg, 1982). If this
vant to recent research on animal cognition in which the
information, which is admittedly present in the distal stim-
stimulus consists of a temporal sequence of events. Many
ulus, has no psychological import, then Gibson's stimulus-
experiments on animal cognition have used stimuli that take
centered theory would be of little consequence for the
place over time, as with a sequence of tonal stimuli (e.g.,
psychology of perception; but this is an unresolved empir-
Hulse, Cynx, & Humpal, 1984) or a piece of music (e.g.,
ical issue.
Porter & Neuringer, 1985) or a small number of brief light
If Gibson and Hochberg represent the extremes, the views
flashes that must be discriminated from a larger number
of Wendall Garner (1974, 1986) and Roger Shepard (1981,
(e.g., W. A. Roberts & Mitchell, 1994). If such events are
1984, 1987) fall somewhere in between. Both acknowledge
perceived as holistic stimuli, traditional views of perception
that there are circumstances under which perception is
and memory may need to be reconsidered. Later in this
largely determined by proximal stimulation because the
article I consider the radical notion of direct memory.
invariant information is sufficient to specify the significant
Finally, Gibson's approach to perception is decidedly
environmental events. In other cases, however, perception
ecological, because he argued that perception should be
occurs under conditions in which proximal stimulation may
studied as it occurs in natural environments, where organ-
not resolve alternative interpretations of distal events (e.g.,
isms enjoy free mobility. Such environments, according to
a fleeting glance of an object at night). Perception in such
Gibson, afford the higher order stimuli that serve as the
cases, according to Garner and Shepard, depends a good
basis of everyday perception. Gibson regarded laboratory
deal on internal inferential processes that reflect the con-
experiments, in which an observer might be allowed a brief
straints of the external world (Shepard, 1984).
monocular view of a stimulus, as ecologically invalid.
In the remainder of this article, I consider various topics
These arguments resonate with recent trends in the field of
in the field of animal cognition, with special attention to the
animal learning emphasizing the role of ecological factors
issues raised by Gibson in regard to the concept of the
in learning and cognition (e.g., Johnston, 1981; Shettle-
stimulus in psychology. In particular, the overview consid-
worth, 1993).
ers whether analyses of complex stimulus control based on
It should be clear that Gibson was mainly concerned with
cognitive mechanisms give proper consideration to the role
analyzing the coupling between distal and proximal stimu-
of the stimulus.
lation. Accordingly, Gibson downplayed (some say, did not
address) the role of perceptual and cognitive mechanisms,
and he has been severely criticized for this position (Hoch- Stimulus Complexity
berg, 1982; Ullman, 1980). However, careful reading of
Gibson shows that organismic variables play an important if I begin with the distinction between simple and complex
underappreciated role in his approach. For example, Gibson stimuli, a distinction fundamental to recent research on
(1966b) regarded animals as information seekers and dis- animal cognition and to perceptual theory. In spite of the
tinguished between information obtained by a passive ob- differences among perceptual theorists, there is general
server versus that received by an active observer, noting that agreement that the stimulus information for perception is
stimulus information available to a locomoting animal is complex (see Garner, 1986), but complexity can be a diffi-
much richer than that available to its immobile counterpart. cult term to operationally define. In the case of recent work
Most important, Gibson emphasized that what is extracted on animal cognition, the term has been used in a somewhat
from the environment depends on learning (E. J. Gibson, informal way.
1991). Consider a well-known example from cognitive psy- Although stimulus complexity may be partly in the eye of
chology (Chase & Simon, 1973). Chess masters can readily the beholder, many psychologists would agree that complex
 COMPLEX STIMULI
stimuli are not easily described in terms of one single simple
dimension; often such stimuli can only be specified with a
great deal of description (e.g., color photographs) or at a
fairly high level of abstraction (e.g., relational concepts such
as same-different). A photograph is said to be worth a
thousand words, and the effective stimuli in discriminations
involving sets of photographs are harder to describe than
those involving red and green hues. Although this distinc-
tion between simple and complex stimuli has a good deal of
face validity, some evidence indicates that such intuitive
distinctions do not always accord with experimental out-
comes. For example, studies of picture perception involving
naive human observers (people without prior experience
viewing pictures) have shown that participants looking at
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simple line drawings may be confused by the depictions,
whereas these same people are able to report accurately
about the content of color photographs (Hagen, 1980). Such
stimuli may be more complex in their specification but more
easily discriminated because of their similarity to the natural
scenes they represent.
Research with nonhuman animals also demonstrates that
stimuli that may appear complex from the experimenter's
point of view may not be perceived as such by the subject.
Research on concept attainment in pigeons, involving many
different color photographs, demonstrates that animals can
acquire the discrimination quite rapidly (after several ex-
perimental sessions and sometimes within the very first
training session; e.g., Herrnstein, 1985; Mallot & Siddall,
1972), at least as quickly as seemingly simpler discrimina-
tions such as red versus green. Discriminations based on
nominally simpler projective line drawings do, however,
appear difficult for animals to learn (Cerella, 1977). Simi-
larly, spatial memory tasks involving complex arrays of
distal cues are also learned very quickly (Olton, 1979).
Consider another surprising example involving temporal
discrimination. We (Fetterman, Dreyfus, & Stubbs, 1989)
trained pigeons on two versions of a task in which subjects
discriminated the relation between a red light of one dura-
tion and a green light of a different duration. In one condi-
tion, the discrimination involved a judgment of which color
was longer; one choice was reinforced when red was longer,
and the alternate choice was reinforced when green was
longer. In other conditions, choices were reinforced accord-
ing to whether the ratio of the red to the green duration was
less or greater than a criterion ratio, such as 2:1; that is, one
choice was reinforced if the ratio was less than 2:1 and the
other was reinforced when that ratio was greater than 2:1.
Although the ratio-based discrimination seems more com-
plex (and it may be for humans; Fetterman, Dreyfus, &
Stubbs, 1993), the pigeons were equally accurate on the two
variants of the temporal pair comparison; recent research,
comparing acquisition in different groups of pigeons, dem-
onstrated that the two discriminations were acquired at the
same rate (Fetterman & Dreyfus, 1994).
Precise comparisons of simple and complex tasks is not
without complication, because both the stimulus dimensions
and the task requirements typically vary. Learning any
discrimination involves a mixture of instructional and per-
ceptual learning. But many examples suggest that ideas
7
regarding stimulus complexity are rooted in anthropocentric
biases; these examples further suggest that analyses of stim-
uli along the lines suggested by Gibson may yield a better
understanding of the information that mediates such com-
plex discriminations. For instance, most radial maze tasks
provide complex arrays of extramaze cues that clearly fa-
cilitate spatial navigation and orientation (Olton, 1978), but
surprisingly little research has been directed at the role of
extramaze stimuli (cf. Cook, 1993). Such research may
suggest an important role for invariants that afford ready
navigation about the maze environment.
Many tasks involving seemingly complex stimuli may be
readily acquired because there is higher order information—
in variances—directly available for perception, not requir-
ing further elaboration (Pick, 1983). This point of view
suggests a taxonomy of stimuli based not on stimulus en-
ergetics, as in the past, but on the relation between stimuli
and distal events. Such a taxomony provides an ecological
foundation for analyses of stimulus control along the lines
suggested by both J. J. Gibson (e.g., 1966b) and Brunswik
(1956), and this foundation is consistent with recent trends
in the field of animal learning and memory toward an
increasing concern with ecological factors.
Stimulus Invariances
J. J. Gibson referred to unchanging aspects of the stimu-
lus array as invariants, and he held that invariants served as
the basic stimulus unit. He described many examples of
invariances in stimulus patterns (e.g., texture gradients, mo-
tion gradients), but an exact determination of the number of
stimulus invariants is impossible. It is fair to say, however,
that the notion of stimulus invariance applies to a variety of
situations. For example, Bassili (1978) attached tiny lights
to people's faces and instructed them to portray different
emotions. The portrayals occurred in a darkened room so
that only the lights were visible, and the pattern of light
movements was filmed. Participants watching the films
easily recognized the emotions depicted even though the
actor's face was not visible (see Cutting, Profitt, & Koz-
lowski, 1977, for similar results). In this section, I consider
the applicability of the invariance concept to research on
animal cognition.
Invariances in Pictures
Color photographs arguably constitute the prototype of
complex stimulus arrangements in the field of animal cog-
nition. Beginning with the seminal work of Herrnstein and
Loveland (1964), the use of photographic images as dis-
criminative stimuli has increased dramatically over the last
30 years. Photographs of natural locations and objects lend
an aura of ecological validity to stimulus control research
under the presumption that such stimuli are representative
of natural environments, possessing "configural or emer-
gent attributes of the more complicated object stimuli and
categories that compose the natural world" (Cook, 1993, p.
175). The widespread popularity of pictorial stimuli is
 8 FETTERMAN
largely due to the rapid increase in research on concept
attainment in animals, but photographs have also been used
as stimuli in experiments on serial list learning (e.g., Ter-
race, 1993), in serial memory tasks (e.g., Wright, Santiago,
Sands, Kendrick, & Cook, 1985), and as discriminative
stimuli under concurrent schedule procedures (Vaughan &
Herrnstein, 1987).
J. J. Gibson (e.g., 1971) suggested that pictures provide
some of the invariant cues present in the natural environ-
ment (e.g., pictorial cues to depth, shading, relative size); it
follows from this position that the ability to perceive infor-
mation in the natural environment is the only prerequisite
for recognizing object-picture correspondences. Gibson
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also noted that, for humans at least, pictures have a dual
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reality. We know a picture is not the real thing (the per-
spective attitude), but people would hate to lose photos of
loved ones in a fire because photographs are often said to
capture the essence of someone (the naive attitude). Clearly
we are concerned with pictorial perception in animals in the
naive sense.
Although animals readily learn to discriminate between
pictorial stimuli based on various natural categories, little is
known about the basis of such discriminations. Some ques-
tion whether nonhuman animals acquire anything like a
human concept (e.g., Lea, 1984; Lea & Ryan, 1990), argu-
ing rather that the discriminations are based on collections
of meaningless (to the subject) features (e.g., Von Fersen &
Lea, 1990) such as decomposed line patterns and blobs of
colors. But the more common if tacitly held assumption is
that animals perceive some correspondence between photo-
graphs of objects and the objects themselves. This assump-
tion is supported by the finding that conceptual discrimina-
tions show good transfer between familiar and novel
instances of the concept. For example, a pigeon trained to
discriminate between 40 pictures containing trees and 40
not containing trees will show nearly perfect transfer of the
discrimination to new photographs, similarly instantiating
the tree versus no-tree concept (e.g., Herrnstein, 1979).
Further support for the correspondence notion is based on
comparisons between concept discrimination training and
pseudoconcept training, in which the same photographs are
arbitrarily assigned to positive and negative categories.
Thus one pigeon might be trained with 80 photographs, one
half with trees and one half without trees, receiving rein-
forcement for pecking at slides containing the tree feature
but not for pecking at slides without trees; another pigeon
would receive training with the same 80 photographs but
with tree and non-tree slides equally divided between pos-
itive and negative categories. The pseudoconcept discrimi-
nation is much more difficult for animals to learn (Edwards
& Honig, 1987; Herrnstein & De Villiers, 1980; Wasser-
man, Kiedinger, & Bhatt, 1988).
Other evidence, however, suggests that animals may not
extract the same information from photographs as do hu-
mans. Pigeons are relatively insensitive to distortions of
pictorial stimuli (e.g., inversions, mirror reversals) that dis-
rupt discriminations in monkeys and humans (Phelps &
Roberts, 1994). Similarly, pigeons learn serial lists com-
prised of either color photographs or kaleidoscopic images
with equal ease; humans, on the other hand, have greater
difficulty learning lists comprised of meaningless kaleido-
scopic images (Wright, Cook, et al., 1990; Wright et al.,
1985). Familiarity with real objects and locations likewise
does not produce robust effects on pictorial discriminations.
Studies of pigeons' conceptual discriminations based on
different locations have shown that familiarity with the
pictured locations may facilitate acquisition of the discrim-
ination compared with subjects without the familiarization
experience (Kendrick, 1992; Wilkie, Wilson, & Kardal,
1989); but the benefits of such real-world experiences are
rather small.
Transfer tests from pictures to real objects (or vice versa)
provide the most direct evidence about picture-object rela-
tions, but the results of such tests are somewhat ambiguous,
contrary to predictions based on Gibson's analysis. Cole and
Honig (1994) trained pigeons to discriminate between color
slides of different locations within a large room; subjects
were then trained to find food in the actual room. Different
subjects were given congruent transfer training (location of
food in the actual room was the same as the positive slide
locations) or incongruent training (positive slide locations
and the location of food in the room differed). The congru-
ent transfer subjects learned to find food more quickly than
the incongruent group, supporting the picture-object corre-
spondence thesis. Unfortunately, this positive transfer effect
turned out to be asymmetrical. When subjects were first
trained in the actual room and then transferred to the pic-
torial discrimination, the congruent versus incongruent reg-
imens did not result in differential transfer to the pictured
locations. Similar asymmetries have been observed in other
studies of object-picture transfer (see reviews of this liter-
ature by Cabe, 1980, and Wilkie, Wilson, & MacDonald,
1992).
Although it is common to describe complex discrimina-
tions involving photographs in terms of the conceptual
language game of the human species, such descriptions are
probably premature, and potentially misleading. There is
not a great deal of evidence that nonhuman animals, espe-
cially pigeons, perceive pictorial stimuli in the same way
humans do. It seems more likely that subjects base concep-
tual discriminations on polymorphous rules involving col-
lections of individual features (Lea & Ryan, 1990).
There may be good reasons for using photographs as
discriminative stimuli (see Wright, 1992) in experiments on
cognitive processes in animals, but the notion that photo-
graphs are somehow more naturalistic because they repre-
sent real objects and locations is, I believe, misguided. A
more productive comparative psychology of concept attain-
ment would stand to benefit by using artificial stimuli that
are equally devoid of meaning for humans and other ani-
mals (Homa, Sterling, & Trepel, 1981; Pearce, 1989). Al-
though such stimuli do not possess the apparent ecological
validity of naturalistic color photographs, they may afford
the possibility of more meaningful cross-species compari-
sons.
 COMPLEX STIMULI
Invariances in Relational Patterns
Many stimulus invariances involve relationships between
features of complex stimuli. It is well-known, for instance,
that the relationship between the illumination of an object
and that of its surround (the luminance ratio) remains
roughly constant under changing conditions of illumination
(Hochberg, 1982), as does the perception of relative bright-
ness (brightness constancy). The debate over the mecha-
nisms of brightness constancy echoes one of the major
themes of this article—the relative contributions of invari-
ant stimulus information and cognition to perception. In one
view the invariance of the luminance ratio serves to explain
brightness constancy, whereas in another the constancy
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results from internal computational mechanisms (e.g., Dem-
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ber & Warm, 1979).
Relational learning tasks provide an excellent illustration
of the potential role of stimulus invariance in learning,
because, by definition, such tasks require subjects to ab-
stract a higher order rule from varying instances of individ-
ual stimuli. Same-different tasks involving matching-to-
sample and related procedures constitute the modern
prototype of relational learning tasks, but many other pro-
cedures have been used. For instance, Fetterman and Drey-
fus (1986) trained pigeons on a relational timing task in
which two colors were presented in succession, with the
colors lasting different durations. At the end of the se-
quence, different choices were reinforced depending on
which color lasted longer. The pigeons readily learned to
discriminate the temporal relation between the two colors.
Another line of research on relational learning has in-
volved the perception of auditory patterns. Hulse and col-
leagues (e.g., Hulse et al., 1984), for example, trained star-
lings to discriminate between sequences of tones according
to whether the sequences rose or fell in pitch. The birds
acquired this discrimination nominally based on relative
pitch information. On the face of it, these examples suggest
that nonhuman animals learn to discriminate invariances in
relational stimulus patterns, consistent with Gibson's thesis.
In contrast with the results from the learning of relational
tasks, the results of transfer tests with novel stimulus values
suggest that nonhuman animals are generally incapable of
extracting invariances in relational patterns. If an animal has
learned the invariant pattern structure, changes in the pattern
constituents should have little or no effect on performance,
but animals often show little evidence of transfer. For ex-
ample, Hulse et al. (1984) found that the discrimination
between ascending and descending pitch sequences was lost
when the tone frequencies were changed from those used in
the original training (for similar findings see D'Amato,
1988; D'Amato & Colombo, 1988; D'Amato & Salmon,
1984). Similarly, Fetterman and Dreyfus (1986) reported
that the discrimination of relative time did not transfer to
novel pairs of intervals outside the range of values used
during training. And, a substantial literature on the same-
different rule suggests that subjects normally learn the par-
ticulars of individual stimuli, not a more general rule in-
volving stimulus relations (e.g., Premack, 1978).
These findings suggest that nonhuman animals are gen-
erally incapable of extracting invariances from stimulus
patterns, at least under relatively artificial laboratory con-
ditions. However, recent research indicates that pigeons and
other species can learn to respond to invariances in stimulus
patterns. Such learning appears to demand that subjects
receive exposure to a large number of stimuli before transfer
tests are given and that hierarchical and lower order features
of the stimuli not be completely confounded. In the work of
Fetterman and Dreyfus (1986), for instance, pigeons were
trained with 12 pairs of durations, receiving a substantial
number of trials that contained duration pairs in which a
single interval by itself was predictive of the higher order
relation. Dreyfus, Fetterman, Stubbs, and Montello (1992)
used 74 unique pairs of stimuli constructed in order to
minimize the confounding between absolute and invariant
stimulus properties. Under these circumstances pigeons
showed good transfer to novel pairs of stimuli (see also
Stubbs et al., 1994).
Page, Hulse, and Cynx (1989) used a similar strategy to
train starlings to discriminate between ascending and de-
scending pitch sequences. One stimulus set contained 64
patterns in which relative and absolute pitch information
were completely uncorrelated; in a second set, which con-
tained 8 patterns, both relative and absolute pitch were
predictive. Starlings were initially unable to learn the dis-
crimination involving 64 patterns; however, birds that re-
ceived training on the smaller set of confounded patterns
subsequently showed good transfer to the larger stimulus
set. Similar results have been obtained in investigations
involving the same-different concept (Wright, Cook,
Rivera, Sands, & Delius, 1988; Wright, Shyan, & Jitsumori,
1990).
Many discussions of relational learning in nonhuman
animals concern capacities for abstract learning among dif-
ferent species, with suggestions that some species prefer to
process stimuli in an absolute fashion (e.g., D'Amato,
Salmon, & Colombo, 1985; Wright et al., 1988). However,
the recent findings indicate an important role for stimulus
information in the sense described by Gainer (1974, 1986),
wherein the information conveyed by an individual pattern
can only be understood by reference to the possible alter-
natives. A pair of stimuli (e.g., one ascending and one
descending pitch sequence) might be perceived one way
when the stimuli are presented in isolation and another way
when the stimuli are part of a larger pool of items.
It is common to describe the results of relational learning
experiments in an either-or fashion; subjects may appear
bound to the specifics of the training stimuli, or perhaps as
a result of elaborate training regimens a subject may behave
in ways that lead us to conclude that a higher order rule has
been learned. But this dichotomy may be overly simplistic
on at least two counts. First, although certain conditions
may afford the perception of higher order relationships,
perception of invariant structure does not preclude an ability
to respond to concrete levels of stimulation (e.g., J. J. Gib-
son, 1966a). For example, faces may be perceived as holis-
tic patterns, but differences in facial expression may also be
noticed and reacted to. In the case of relational learning
tasks, subjects may learn to perceive both the invariant
 10 FETTERMAN
structure and the details of the events that compose rela-
tional stimulus patterns. Page et al. (1989), for instance,
found that starlings used both relative and absolute pitch
information in discriminating between ascending and de-
scending pitch sequences. Likewise, Dreyfus et al. (1992)
found pigeons' discriminations of the relative durations of
temporal intervals to be based on both the temporal rela-
tionship between the intervals and the absolute durations of
individual stimuli (see also Dreyfus, 1992; Wright, Cook, &
Kendrick, 1989, for similar findings).
Second, complex events may consist of different levels of
higher order relations, from the concrete to the abstract,
constituting a hierarchy of stimulus structure; that is, some
events have embedded superordinate structures (cf. E. J.
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Gibson & Levin, 1975). Animals may learn to respond to
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some or all of these levels, depending on the predictive
relations between the various levels and significant out-
comes (cf. Herrnstein, 1990). For instance, living organisms
may be classified at different taxonomic levels (e.g., animal,
mammal, carnivore, dog, cocker spaniel), and the observed
level(s) of classification may be determined both by the
physical variation in individual stimuli and by the conse-
quences associated with varying levels of categorization
(e.g., W. A. Roberts & Mazmanian, 1988).
How Big Is a Stimulus?
"The fact is that the general shape and emphases of any
theory of perception and of information processing depends
strongly on this matter of the size of the effective stimulus"
(Hochberg, 1982, p. 192).
"I argue that there are limits on the intervals of space and
time over which we can integrate information available in
the sensory arrays and these limits are themselves lawful in
ways that cry out for explanation" (Shepard, 1984, p. 419).
"When does a sequence constitute a single and when a
number of separate stimuli; also, can a single enduring
stimulus exist throughout a changing sequence?" (J. J. Gib-
son, 1960, p. 696).
These passages speak to a controversial aspect of James
Gibson's views—that a temporal sequence of events may be
perceived as a single stimulus without the aid of memory.
Gibson (e.g., 1966a) placed successive and simultaneous
stimulation on equal footing and set no boundaries on the
temporal extent of an effective stimulus. 1 In this section I
take up the issue of whether extended temporal sequences
may function as unitary stimulus events, an issue of some
significance for perceptual and cognitive theory.
Many experiments on animal cognition involve simulta-
neous arrays of stimulus elements, as in the relative numer-
osity discriminations reported by Honig (e.g., Honig &
Stewart, 1989), the texture gradients used by Cook (e.g.,
Cook, 1992b), or the simultaneous list learning procedures
studied by Terrace (e.g., Terrace, 1993). In other experi-
ments the stimulus elements are presented in a temporal
sequence, as in the work of Hulse and colleagues (e.g.,
Hulse et al., 1984), in serial list learning tasks (e.g., Wright
et al., 1985), or in studies of sequential numerosity discrim-
inations (e.g., Alsop & Honig, 1991; Fernandes & Church.
1982; Fetterman, 1993). It is customary to distinguish be-
tween the apprehension of simultaneous and successive
stimulus patterns in terms of perceiving versus remember-
ing, but this dichotomy may be based on preconceptions
about stimuli as discrete, momentary events, as noted by
Gibson.
Consider an example involving numerosity discrimina-
tion. Honig and Stewart (1989) presented pigeons with
simultaneous arrays composed of red and blue dots. The
pigeons were initially trained to discriminate between two
homogeneous arrays (i.e., all red vs. all blue) and then were
given generalization tests with arrays containing different
proportions of the red and blue elements. Honig and Stewart
obtained orderly generalization gradients based on the pro-
portion of each element in the arrays, and they found that
the discrimination transferred to new arrays that contained
different total numbers of elements. One might say that the
effective stimulus consisted of the emergent dimension of
relative numerosity based on the entire pattern of elements
in the array (Honig, 1992).
Alsop and Honig (1991) used a similar task, but one in
which the elements (red and blue light flashes) were pre-
sented in succession; under their procedure, different
choices were reinforced depending on whether the sequence
was comprised of more red or more blue flashes. As under
the simultaneous procedure, discriminative performance
was a function of the proportion of the different elements in
the sequence. However, there were clear effects of the
location of an element in the sequence, with light flashes
appearing later in the sequence exerting more control over
choice. In addition, increases in the temporal gap between
successive flashes (from 0.33 s to 0.66 s to 1.0 s) resulted in
decreases in accuracy. The differential weighting of pattern
elements as a function of serial position is consistent with an
explanation of sequential numerosity discrimination based
on memory processes, but not with Gibson's suggestion that
such extended patterns may be directly apprehended with-
out the aid of memory.
Some results appear to favor the idea that animals can
integrate a series of events over substantial periods of time,
however. Stubbs, Dreyfus, and Fetterman (1984) developed
a temporal integration task in which pigeons viewed a
sequence of four colored lights that lasted for different
1
Consider an example involving a primitive life form, the
bacterium. This organism swims about in a medium in two distinct
patterns, runs and tumbles; runs are movements in a straight line,
and tumbles are movements involving random changes in direc-
tion. The occurrence of the different patterns is related to gradients
of chemical attractants in which the frequency of tumbling de-
creases as the bacterium moves up the gradient and increases as it
moves down the gradient. Note that the chemical gradient is a
complex stimulus and that control of behavior by the stimulation is
distributed over time and space. One could assume that this uni-
cellular organism stores discrete representations of its environment
for comparison with current inputs, but it seems more parsimoni-
ous to assume that the chemical gradient provides a complex
stimulus that directly controls the observed pattern of behavior (cf.
Staddon, 1983; Turvey & Shaw, 1979).
 COMPLEX STIMULI
durations. The series consisted of red (R) and green (G)
lights presented in alternation, R,-G1-R2-G2; many differ-
ent individual durations were used, with each color lasting
5 s on average and a mean sequence duration of 20 s. At the
end of the color sequence, different choices were reinforced
depending on which color pair (R, + R2 or G! + G2) had
lasted longer. The birds' choices were related to the relative
durations of the two color pairs in a very orderly way, and
mean accuracy for three pigeons was about 75% correct.
One pigeon was exposed to a second condition in which the
average durations of the individual components of the se-
quence were increased fourfold, from 5 s to 20 s, providing
a mean sequence length of 80 s. Performance under this
extended sequence condition was identical to that observed
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with the shorter sequence. These results are somewhat sur-
prising but consistent with Gibson's thesis that a single
stimulus may endure over substantial periods of time.
Discrimination of complex stimuli often involves inte-
grating a series of discrete events, such as individual notes
in bird song (Weisman & Ratcliffe, 1987), elements of a
numerical series (Alsop & Honig, 1991), sequences of
choices in a radial maze (Olton & Samuelson, 1976), or
individual components of complex patterns of reward
(Hulse & O'Leary, 1982). It is important to know the extent
to which animals can perceive a temporally extended series
of events as a unitary stimulus because the limits of per-
ceptual integration set the boundary conditions for theories
of information processing (Hochberg, 1982; Shepard,
1984). Experimental analyses of perceptual time horizons
may serve to constrain admissable theories of the percep-
tion, processing, and retention of stimulus structure.
Role of Memory
It is conventional to treat perception of extended patterns
in terms of associative and memory mechanisms that link
together and store individual pattern elements, as in the case
of Alsop and Honig (1991). By this convention, identifying
a span of integration depends on establishing a sharp de-
marcation between perceiving and remembering so that
events inside and outside the perceptual window engage
distinctly different mechanisms. However, J. J. Gibson
(1966a) pointed out, as did others before him, that it is
difficult to say exactly where perception leaves off and
memory begins.
The stream of experience does not consist of an instantaneous
present and a linear past receding into the distance; it is not a
"traveling razor's edge" dividing the past from the future.
Perhaps the present has a certain duration. If so, it should be
possible to find out when perceiving stops and remembering
begins. But it has not been possible. (J. J. Gibson, 1979, p.
253).
Gibson's position might seem naive in light of a volumi-
nous literature on short-term remembering in both human
and nonhuman animals, a literature that seemingly has es-
tablished the bounds of working memory (Honig & Thomp-
son, 1982; W. A. Roberts & Grant, 1976; Roitblat, 1984).
However, efforts to quantify the time course of forgetting
11
(e.g., Anderson & Schooler, 1991; Harnett, McCarthy, &
Davison, 1984; Loftus, 1985; White, 1985, 1991; Wixted,
1989; Wixted & Ebbesen, 1991) have produced results
consistent with Gibson's position in the sense that some of
the candidate functions posit a constant rate of forgetting
(e.g., the exponential), making it impossible to identify a
discrete point in time at which perceiving leaves off and
remembering takes over.
The development of quantitative models of memory has
also highlighted possible parallels between perceptual and
memorial processes. For example, power functions of the
form
M = AT (1)
provide an excellent account of forgetting curves across a
range of conditions involving different species, stimulus
materials, and memory paradigms (Anderson & Schooler,
1991; Wixted & Ebbesen, 1991). In Equation 1, M stands
for the measure of memory performance, T for the retention
interval, and A and b are the parameters of the model. The
exponent, b, can be taken as an estimate of the rate of
forgetting. The perceptual counterpart to the power function
model is, of course, Stevens' Power Law,
S = klb, (2)
which describes the relation between physical (I) and sub-
jective (S) magnitudes for judgments of many perceptual
dimensions such as loudness, brightness, time, and weight,
with each of these dimensions having its own characteristic
exponent (Stevens, 1975).
The similarity of the mathematical form of psychophys-
ical and forgetting functions could be purely coincidental,
but there may be underlying correspondences between per-
ceiving and remembering, as White (1991), Staddon (1983,
1984), and Wixted and Ebbesen (1991) have suggested.
White (1991), for example, proposed a theory of direct
remembering based on the idea that remembering involves
a discrimination between events across temporal distances
and that such discriminations are analogous to making dis-
tinctions between objects at different spatial distances (cf.
Staddon, 1984). In this view, increasing the temporal dis-
tance between the stimuli to be discriminated attenuates
their discriminability just as increasing spatial distances
make objects more difficult to tell apart. Thus, the salience
of memories may decline over time according to the same
function that describes changes in the perceived intensity of
a stimulus (e.g., a light or sound) as its distance varies
(Staddon, 1984; Wixted & Ebbesen, 1991).
This approach places the analysis of memory in the con-
text of psychophysics and draws attention to the role of
stimuli in remembering (see Fetterman, 1995, for a recent
example of a psychophysical analysis of remembering).
Many psychophysical experiments have shown that the
exponent of the psychophysical function depends on the
underlying physical continuum; for example, the dimen-
sions of brightness, length, loudness, and shock intensity
yield representative exponent values of 0.33, 1.0, 0.67, and
3.5, respectively (Stevens, 1975). Similarly, the exponent of
 12 FETTERMAN
the forgetting curve embodied in Equation 1 may be a
function of the to-be-remembered events. Not much is
known about the role of stimuli in remembering, however,
because primary emphasis has been given to intervening
cognitive variables (rehearsal; common coding; response-
outcome expectancies, etc.). The limited evidence suggests
that stimulus characteristics are relevant variables. For ex-
ample, pigeons forget discriminations based on hue samples
more rapidly than those based on line orientations (Farthing,
Wagner, Gilmour, & Waxman, 1977; Urcuioli & Zentall,
1986). Pigeons also forget visual discriminations more rap-
idly than auditory discriminations (Kraemer & Roberts,
1984), whereas monkeys forget auditory discriminations
more rapidly (Colombo & D'Amato, 1986). At the very
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least, psychophysical analyses of delayed stimulus control
(i.e., memory) along the lines of Fetterman (1995) can
provide a comprehensive portrait of delayed dimensional
control and may illuminate parallels between perceiving and
remembering.
Multidimensional Stimuli
Complex stimuli normally involve multiple and related
sources of stimulus information, and questions naturally
arise about the relative contributions of different stimulus
features to the perception of the whole. It has become
conventional in analyses of conditioning and learning to
treat complex stimuli as compounds of elemental features
(e.g., Kamin, 1969) and to assume an associative competi-
tion between individual elements that predict a common
outcome (e.g., presentation of an unconditioned stimulus).
Many experiments have demonstrated that increments in the
associative value of a target cue depend on the predictive
values of other cues that compose the entire associative
context (see Bouton, 1994, for a recent review).
The assumption of stimulus competition is further abetted
by studies of the role of attentional factors in learning that
suggest that paying attention to one dimension reduces
control exerted by other, correlated dimensions. In a widely
cited illustration of this result, Reynolds (1961) trained
pigeons to peck at a stimulus compound comprised of a
white triangle superimposed on a red background; once the
pecking response was established stimulus control was as-
sessed through generalization tests with the individual ele-
ments. The results of the generalization test indicated that
the instrumental response was conditioned to a single aspect
of the compound, in spite of the fact that both elements were
equally predictive of reinforcement. Reynolds interpreted
these results in terms of a process of selective attention.
Similar results and conclusions have emerged from research
with compound matching-to-sample procedures (e.g., Riley
& Brown, 1991; Riley & Leith, 1976; Santi, Grossi, &
Gibson, 1982).
Whereas manipulations of the predictive validities of
elemental cues support the idea that learning involves the
detection of correlations between stimuli or between stimuli
and responses, simple correlational models may not fully
represent the causal texture of complex environments.
Brunswik (1956; Tolman & Brunswik, 1935) long ago
pointed out the advantages of studying behavior in situa-
tions in which the natural covariation between variables was
preserved because such situations were more representative
of naturalistic conditions. In particular, the cue competition
framework may not be a useful model of situations in which
multiple sources of stimulus information are not completely
redundant predictors of some target event; under such cir-
cumstances there is much to be gained by using more than
one predictor (cf. Fetterman, 1993; Fetterman, Stubbs, &
Dreyfus, 1986; Fetterman, Stubbs, & MacEwen, 1992).
Consider an example involving foraging behavior. Opti-
mal foraging models assume that animals search and obtain
food so as to maximize energy intake for the time spent
foraging (e.g., Stephens & Krebs, 1986), an assumption that
seemingly requires sophisticated computational mecha-
nisms. Many believe, however, that the mechanisms of
foraging involve heuristic decision rules that greatly sim-
plify the maximization problem. For example, a forager
might use a rule based on the time since the last prey
encounter as the basis for deciding when to leave one patch
for another (e.g., Brunner, Kacelnik, & Gibbon, 1992;
Kacelnik, Brunner, & Gibbon, 1990), and such simple tem-
poral decision rules may afford a reasonable approximation
to strategies based on complex algorithms embodied in
optimal foraging models. But other sources of information
about prey densities might also be predictive (e.g., total
number of prey captured), and this information could be
combined with temporal cues to more accurately anticipate
the optimal patch departure time.
Several findings are consistent with this idea. Kamil,
Yoerg, and Clements (1988) trained blue jays on a simu-
lated foraging task involving two food patches that provided
reinforcements according to different scheduled probabili-
ties. The higher probability schedule provided a fixed num-
ber of prey items so that food was no longer available once
the allotted reinforcers were obtained (the "sudden death"
patch). The alternative ("nondepleting") patch provided re-
inforcement at a lower, constant probability that was inde-
pendent of the number of reinforcements earned. Under
these conditions an optimal forager should first obtain all of
the food available under the richer schedule and then switch
immediately to the leaner, constant probability alternative.
The blue jays' behavior approximated the optimal strategy.
More important, analyses indicated that patch departure
decisions were based on more than one source of informa-
tion, including the number of prey obtained in the richer
patch, the number of successive trials without a prey item
(the run of bad luck), and the birds' history of receiving a
certain number of reinforcers in the richer patch. No single
simple cue was involved.
Fetterman (1993) trained pigeons to discriminate between
different fixed-ratio (FR) samples by reinforcing one choice
after the smaller sample (e.g., FR 10) and another after the
larger one (e.g., FR 20). This number-based discrimination
was readily learned and multiple regression analyses using
both the number of responses emitted and the time taken to
emit the responses as predictors of choice demonstrated that
 COMPLEX STIMULI
the birds' numerosity discriminations were based on both
dimensions (see Fetterman et al., 1986, for related findings).
Gallistel (1990) presented evidence that animals acquire
representations of the duration and number of significant
environmental events and suggested that these representa-
tions provide the computational ingredients for more com-
plex representations of the rate of events in time (e.g.,
number of reinforcers per hour). His account presupposes
that animals monitor both the number and duration of events
in the foraging setting, a supposition consistent with the
theme of control by multiple aspects of complex environ-
ments.
Whereas modern views of learning emphasize the impor-
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tance of simple correlations between events, the present
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discussion favors an extension of this approach. It recog-
nizes that complex events may involve multiple sources of
information that, taken as a whole more accurately predict
significant outcomes than does a single source of stimulus
information. Multiple regression may be the appropriate
statistical model in situations involving complex stimuli.
But, of course, multiple predictors give informational lever-
age only when the predictors are less than perfectly corre-
lated (the problem of multicolinearity). Perfectly correlated
predictors (e.g., a tone and light that both predict a shock)
often result in blocking or overshadowing of one predictor
by the other, a result consistent with a framework empha-
sizing cue competition. However, because complex envi-
ronments often do not involve complete redundancy among
predictive cues there is some advantage to be gained by
relying on multiple sources of stimulus information.
Role of the Stimulus in Timing
Although much of the recent research in animal cognition
has used complex multidimensional events, research on
time perception in nonhuman animals historically has had
very little concern with the role of stimuli. Most experi-
ments have involved a single unchanging stimulus (e.g., a
light or tone) lasting for some duration (e.g., Fetterman &
Killeen, 1992; S. Roberts, 1981; Stubbs, 1968). Such ex-
periments are equivalent to classic studies of depth percep-
tion in which stimulus information about distance was re-
duced to a minimum (e.g., Holway & Boring, 1941). In the
absence of stimulus-based information about the duration of
events, explanations of timing in animals have quite natu-
rally focused on cognitive mechanisms such as an internal
clock (Church, 1984) that may mediate temporal discrimi-
nations. Pulses from the internal clock provide the under-
lying temporal structure for the homogeneous stimuli typi-
cally used in animal timing research.
By contrast, the role of stimulus events has been a major
topic of empirical and theoretical analyses of human timing.
Research on time perception in humans provides abundant
evidence that stimulus factors play an important role in
temporal experience (e.g., Poynter, 1989). For example,
intervals that contain more stimuli seem longer than inter-
vals containing fewer stimuli (e.g., a flickering light vs. a
steady light; Thomas & Brown, 1974). This filled-duration
13
illusion has been observed under a variety of conditions
involving manipulations of the number of events that com-
pose the interval to be estimated (see Poynter & Homa,
1983); similar results have been obtained with stimulus
complexity as the independent variable (i.e., intervals con-
taining more complex stimuli are judged longer than inter-
vals containing less complex stimuli; Block, 1978). Many
other task and stimulus factors have been shown to affect
humans' temporal judgments, including the amount of work
done (Burnside, 1971), task demands (Sawyer, Meyers, &
Huser, 1994), signal modality (Goldstone & Lhamon,
1974), and stimulus movement (Brown, 1995).
The effects of stimulus structure have led to the develop-
ment of models of human temporal perception based on
cognitive activity (Block, 1990) in much the same way that
research involving complex stimuli with nonhuman animals
has drawn attention to the role of cognitive factors in
learning. Memory storage models (e.g., Ornstein, 1969), for
instance, propose that temporal estimates depend on the
amount of memory occupied by representations of stimulus
events; events that take up more space in memory appear to
last longer than those that take up less space. An alternative
cognitive model places primary emphasis on information-
processing effort, hypothesizing that perceived duration is
directly related to the amount of cognitive resources allo-
cated to temporal events (Burnside, 1971; Hicks, Miller, &
Kinsbourne, 1976). Once again, in the temporal domain as
in other areas of perception, Gibson (e.g., 1975) argued for
the primacy of stimulus information in perceiving. Gibson's
account of temporal perception focused on the role of stim-
ulus change in perceiving time, a view that regards stimulus
change as the index of the passage of time. From Gibson's
standpoint, studies of timing in animals, in which the stimuli
typically are unchanging lights or tones, are profoundly
lacking in ecological validity because the stimuli lack the
temporal structure that is characteristic of natural environ-
ments.
Although there have been no systematic efforts to assess
the role of stimuli in animal timing, bits and pieces of data
suggest that stimuli can play an important role in animals'
temporal judgments. For example, Duncan and Fantino
(1972) found that pigeons' estimates of a forthcoming delay
to reinforcement depended on the stimuli that signaled the
delay; the psychological distance to a reward was judged to
be longer when the delay signal was a segmented stimulus
(e.g., a red light followed by a yellow light) than when the
signal was a single stimulus (e.g., a blue light). Other
stimulus manipulations affect the accuracy of temporal dis-
criminations; filled durations are discriminated more accu-
rately than unfilled durations (Mantanus, 1981), durations
of access to food are discriminated more accurately than
light durations (Spetch & Wilkie, 1981), and light durations
are discriminated more accurately than tone durations
(W. A. Roberts, Cheng, & Cohen, 1989).
Gibson's approach places temporal perception in the
same context as other complex perceptual phenomena such
as space and motion perception, again suggesting an impor-
tant role for stimulus information. According to his ap-
proach, there are multiple sources of information about the
 14 FETTERMAN
duration of naturally occurring events, just as there are for
the distances of things. Standard experimental procedures
for studying timing in animals might obscure the potential
information if the stimuli to be discriminated are simple
unchanging lights or sounds that differ in duration only.
Natural environments are complex, however, and preserva-
tion of the inherent temporal structure (Brunswik, 1956) of
such environments will surely shed additional light on the
functions and mechanisms of timing behavior.
Conclusion
The concept of the stimulus needs to be liberalized.
This article is intended solely for the personal use of the individual user and is not to be disseminated broadly.
Animals have evolved to pick up predictive invariants in
This document is copyrighted by the American Psychological Association or one of its allied publishers.
patterns of stimuli (J. J. Gibson, 1979; Shepard, 1984); what
may appear to be a highly complex pattern within a tradition
that holds that a stimulus is something discrete and momen-
tary may be simple from the animal's point of view. A
stimulus may consist of patterns of change, relations be-
tween features, and so on. Current reductionist approaches
to stimulus control should be supplemented by psychophys-
ical analyses that relate behavior to higher order stimulus
variables. Many ecologically valid situations involve mul-
tiple sources of stimulus information that often are not
completely redundant predictors of outcomes. Approaches
that emphasize cue competition should be complemented by
those that consider the ways in which nonredundant stimuli
that compose complex events influence behavior.
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Received April 11, 1995
Revision received May 15, 1995
Accepted May 15, 1995 •