The operant and the classical in conditioned orientation of Drosophila melanogaster at the flight simulator

Ever since learning and memory have been studied experimentally, the relationship between operant and classical conditioning has been controversial. Operant conditioning is any form of conditioning that essentially depends on the animal's behavior. It relies on operant behavior. A motor output is called operant if it controls a sensory variable. The Drosophila flight simulator, in which the relevant behavior is a single motor variable (yaw torque), fully separates the operant and classical components of a complex conditioning task. In this paradigm a tethered fly learns, operantly or classically, to prefer and avoid certain flight orientations in relation to the surrounding panorama. Yaw torque is recorded and, in the operant mode, controls the panorama. Using a yoked control, we show that classical pattern learning necessitates more extensive training than operant pattern learning. We compare in detail the microstructure of yaw torque after classical and operant training but find no evidence for acquired behavioral traits after operant conditioning that might explain this difference. We therefore conclude that the operant behavior has a facilitating effect on the classical training. In addition, we show that an operantly learned stimulus is successfully transferred from the behavior of the training to a different behavior. This result unequivocally demonstrates that during operant conditioning classical associations can be formed.     

Instrumental cardiovascular conditioning: A review

This paper reviews experiments, hypotheses, and current controversies about instrumental cardiovascular conditioning. Demonstrations of such conditioning in curarized animals challenged a differentiation between instrumental and classical learning on the basis of their respective effector systems but did not prove direct operant learning by the autonomic nervous system. In humans, ethical prohibition of curarization and lack of adequate controls for respiration and muscle tension have resulted in incomplete understanding of the roles of voluntary, somatic mediators. Despite a variety of potential clinical applications of biofeedback, the available literature lacks studies of its efficacy compared to more standard modes of therapy. The physiological mechanisms and central neural pathways involved ininstrumental cardiovascular conditioning remainalmost totally unknown.     

Feeding behavior of Aplysia: a model system for comparing cellular mechanisms of classical and operant conditioning

Feeding behavior of Aplysia provides an excellent model system for analyzing and comparing mechanisms underlying appetitive classical conditioning and reward operant conditioning. Behavioral protocols have been developed for both forms of associative learning, both of which increase the occurrence of biting following training. Because the neural circuitry that mediates the behavior is well characterized and amenable to detailed cellular analyses, substantial progress has been made toward a comparative analysis of the cellular mechanisms underlying these two forms of associative learning. Both forms of associative learning use the same reinforcement pathway (the esophageal nerve, En) and the same reinforcement transmitter (dopamine, DA). In addition, at least one cellular locus of plasticity (cell B51) is modified by both forms of associative learning. However, the two forms of associative learning have opposite effects on B51. Classical conditioning decreases the excitability of B51, whereas operant conditioning increases the excitability of B51. Thus, the approach of using two forms of associative learning to modify a single behavior, which is mediated by an analytically tractable neural circuit, is revealing similarities and differences in the mechanisms that underlie classical and operant conditioning.     

Operant Reward Learning in Aplysia

Anticipating the future has a decided evolutionary advantage, and researchers have found many evolutionarily conserved mechanisms by which humans and animals learn to predict future events. Researchers often study such learned behavior using conditioning experiments. The marine snail Aplysia has been at the forefront of research into the cellular and molecular mechanisms of classical conditioning. Recently, Aplysia has also gained a reputation as a valuable model system for operant reward learning. Its feeding behavior can be operantly conditioned in the intact animal as well as in reduced preparations of the nervous system. The reward signal relies on dopamine transmission and acts in conjunction with activity in an identified neuron (B51) to bring about operant memory.     

Is behaviorism false because there is no such thing as conditioning? Popper and skinner on learning

Sir Karl Popper has claimed that behaviorism is misguided because it holds that conditioning occurs through repetition. According to Popper, there is no such thing as learning through repetition. To the limited extent that philosophers of science have concerned themselves with behaviorism, this attack is one of the most direct and unique in that the battleground is not over the value of mentalism/cognitivism but a bold claim that conditioning—the heart and soul of behaviorism—is fictitious. This paper examines the soundness of Popper's argument against behaviorism by examining whether operant and classical conditioning rely on learning through repetition and suffer from other problems Popper attributes to the notion of conditioning. Although Popper correctly attributes certain properties to classical conditioning, he fails to undermine the empirical evidence that such conditioning occurs. Second, we claim that although B. F. Skinner is never entirely clear, operant conditioning does not rely on repetition nor does it suffer from the other problems Popper attributes to conditioning. Thus, Popper's argument also fails because of his assumption that all conditioning is classical conditioning and therefore his misattributing properties of classical conditioning to operant conditioning. We conjecture that the earlier polemics of John Watson and Watson's sole reliance on classical conditioning probably contributed to Popper's confusion on this point.     

Learning during stressful times

Stressful life events can have profound effects on our cognitive and motor abilities, from those that could be construed as adaptive to those not so. In this review, I discuss the general notion that acute stressful experience necessarily impairs our abilities to learn and remember. The effects of stress on operant conditioning, that is, learned helplessness, as well as those on classical conditioning procedures are discussed in the context of performance and adaptation. Studies indicating sex differences in learning during stressful times are discussed, as are those attributing different responses to the existence of multiple memory systems and nonlinear relationships. The intent of this review is to highlight the apparent plasticity of the stress response, how it might have evolved to affect both performance and learning processes, and the potential problems with interpreting stress effects on learning as either good or bad. An appreciation for its plasticity may provide new avenues for investigating its underlying neuronal mechanisms.     

Redundant amodal properties facilitate operant learning in 3-month-old infants

The current study examined the role redundant amodal properties play in an operant learning task in 3-month-old human infants. Prior studies have suggested that the presence of redundant amodal information facilitates detection and discrimination of amodal properties and potentially functions to influence general learning processes such as associative conditioning. The current study examined how human infants use redundant amodal information (visual and haptic) about the shape of an object to influence learning of an operant response. Infants learned to kick to move a mobile of cylinders while either holding a cylinder, a rectangular cube, or no object. Kick rate served as the dependent measure. The results showed that infants given matching redundant amodal properties (e.g., viewed cylinders while holding a cylinder) showed facilitated operant learning whereas infants given mismatching redundant amodal properties showed inhibited operant learning. These results support and extend the Intersensory Redundancy Hypothesis by demonstrating that amodal redundancy influences complex learning processes such as operant conditioning.     

Identification of a reinforcement pathway necessary for operant conditioning of head waving in Aplysia californica

The marine mollusc Aplysia californica exhibits a wide range of nonassociative and associative forms of learning. Recently, we found that the learning repertoire of Aplysia includes operant conditioning (Cook & Carew, 1986, 1989b). The behavior we examined is a naturally occurring, side-to-side head-waving response used by Aplysia in seeking food, obtaining a foothold, and egg laying. Aplysia can be operantly conditioned to reduce head-waving to one side of their body if such a response results in exposure to bright uniform-field illumination, which the animals find aversive. An essential step toward achieving a mechanistic understanding of operant conditioning is to identify and characterize the reinforcement pathway used during the learning. Toward this end, we wished to determine which of the peripheral visual pathways in Aplysia are critical for performance of the operant task. Previous experiments indicated that photic input from the optic and rhinophore nerves functionally inhibited motor neurons that participate in the operant response (head-waving), while photic input from the oral veil nerves excited these same motor neurons (Cook & Carew, 1989c). These findings suggested the hypothesis that one or both of these pathways could play an important role in mediating reinforcement during training. To explore this possibility we operantly trained animals that had received chronic bilateral transections of either the optic and rhinophore nerves or the oral veil nerves C1-C3 (in conjunction with transection of the optic and rhinophore nerves). We found that operant conditioning was not disrupted by ablation of input from the eyes and rhinophores. By contrast, ablation of input from the oral veil (together with that from the eyes and rhinophores) abolished operant conditioning. Thus, the oral veil nerves play a critical modulatory role in operant conditioning of head-waving. This observation further suggested that photic input from the oral veil is conveyed to the CNS via the oral veil nerves. In a final experiment we confirmed that stimulation of the oral veil with light evokes increased afferent activity in the oral veil nerves C1-C2. These results support the idea that the oral veil nerves contain processes that are critical components of the reinforcement pathway for operant conditioning of head-waving.     

LEARNING THEORY APPROACHES TO PERSUASION

This review synthesizes relevant research dealing with the processes of learning and suggests its applications to compliance gaining. The two major issues addressed are: (1) to what degree can learning theories explain the acquisition of new attitudes and behaviors, and (2) to what degree are attitudinal and behavioral changes governed by learning theory principles? The learning theories discussed are grouped into three categories: stimulus-response or connectionist approaches; cognitive approaches; and stochastic, mathematical, and cybernetic approaches. The stimulus-response models, which encompass most of the research examined in this paper, are further broken down into four types: (1) classical conditioning, (2) contiguity models, (3) instrumental (or operant) conditioning and (4) models including drive and drive reduction. Principles and major research evidence from numerous learning theories are reviewed and analyzed, and suggestions are made as to how this evidence may aid in the construction of more complete theories of persuasion and attitude change.     

The infant vocal-conditioning literature: a theoretical and methodological critique

Most studies of operant conditioning of infant vocalization do not fully use the experimental-analysis-of-behavior methodology that behavioral researchers have developed to study operant phenomena. This could be a contributing factor in their failure to correctly operationalize the definition of reinforcement, severely limiting the amount of information they can provide about operant learning in infants. Furthermore, single-subject-design studies may be added to supplement or replace group experimental designs in the study of infant vocal conditioning if we are to recognize the different learning processes that affect infant learning. Finally, single-subject experimental designs may be crucial to the development of an effective technology of early language intervention.     

Elicited versus emitted behavior: Time to abandon the distinction

The concept of emitted behavior was formulated as a part of the original argument for the validity of a new kind of learning called operant conditioning. The rationale for operant conditioning contrasted it with Pavlovian or classical conditioning, which was (and remains) fundamentally based on responses to conditioned and unconditioned stimuli. Classical conditioned responses were said to be elicited. In contrast, operant behavior was viewed as emitted and controlled primarily by response consequences rather than antecedents. I argue that the distinction between emitted and elicited behavior is no longer warranted for three major reasons. First, the distinction was based on a view of Pavlovian conditioning that is no longer viable. Second, the distinction is incompatible with both empirical data and contemporary conceptualizations of operant behavior. Third, the only way to overcome these problems is to define emitted and elicited in terms of the type of conditioning (operant and classical) that produces these behaviors, but that approach makes the definitions circular and does not avoid implications of the terms that are misleading and counterproductive in light of contemporary research and thinking.     

Genetic dissociation of acquisition and memory strength in the heat-box spatial learning paradigm in Drosophila

Memories can have different strengths, largely dependent on the intensity of reinforcers encountered. The relationship between reinforcement and memory strength is evident in asymptotic memory curves, with the level of the asymptote related to the intensity of the reinforcer. Although this is likely a fundamental property of memory formation, relatively little is known of how memory strength is determined. Memory performance at different levels in Drosophila can be measured in an operant heat-box conditioning paradigm. In this spatial learning paradigm, flies learn and remember to avoid one-half of a dark chamber associated with a temperature outside of the preferred range. The reinforcement temperature has a strong effect on the level of learning in wild-type flies, with higher temperatures inducing stronger memories. Additionally, two mutations alter memory-acquisition curves, either changing acquisition rate or asymptotic memory level. The rutabaga mutation, affecting a type-1 adenylyl cyclase, decreases the acquisition rate. In contrast, the white mutation, modifying an ABC transporter, limits asymptotic memory. The white mutation does not negatively affect classical olfactory conditioning but actually improves performance at low reinforcement levels. Thus, memory acquisition/memory strength and classical olfactory/operant spatial memories can be genetically dissociated. A conceptual model of operant conditioning and the levels at which rutabaga and white influence conditioning is proposed.     

RELATIONSHIPS BETWEEN EXPERIMENTAL AND CLINICAL PSYCHOLOGY: THE CASE OF BEHAVIOR THERAPY

Behavior therapy is the outstanding example of the beneficial influence of experimental research on clinical practice. Initial applications of laboratory research to clinical disorders consisted primarily of the principles and procedures of operant and classical conditioning. The development and evaluation of the token economy and biofeedback methods are used to illustrate the contribution of operant conditioning, whereas the derivation and efficacy of exposure-based treatments for phobic and obsessive-compulsive disorders exemplify the utility of classical and avoidance conditioning concepts. As behavior therapy has matured and grown more clinically sophisticated, its theoretical and research bases have broadened to include areas such as vicarious learning and self-regulatory processes. Most recently, concepts from modern cognitive and social psychology (e.g., information processing, attribution theory) have begun to guide clinical applications. In turn, different therapeutic approaches have influenced the nature of laboratory research in some areas. Nonetheless, a gap still exists between experimental research and clinical practice. The solution to this perennial problem is not to make scientists of practitioners, but to recognize the unique contributions of basic researchers, clinical investigators, and practitioners, within a consistent framework that specifies the complex interrelationships among the different levels of analysis along the continuum of basic research to clinical practice. Such a framework is sketched-out together with some comments on the reasons for relative paucity of controlled clinical research in the U.S.A. and suggestions for bringing the influence of research findings to bear on practice.     

The history of imitation in learning theory: the language acquisition process

The concept of imitation has undergone different analyses in the hands of different learning theorists throughout the history of psychology. From Thorndike's connectionism to Pavlov's classical conditioning, Hull's monistic theory, Mowrer's two-factor theory, and Skinner's operant theory, there have been several divergent accounts of the conditions that produce imitation and the conditions under which imitation itself may facilitate language acquisition. In tracing the roots of the concept of imitation in the history of learning theory, the authors conclude that generalized imitation, as defined and analyzed by operant learning theorists, is a sufficiently robust formulation of learned imitation to facilitate a behavior-analytic account of first-language acquisition.     

Reinforcement in an in vitro analog of appetitive classical conditioning of feeding behavior in Aplysia: blockade by a dopamine antagonist

In a recently developed in vitro analog of appetitive classical conditioning of feeding in Aplysia, the unconditioned stimulus (US) was electrical stimulation of the esophageal nerve (En). This nerve is rich in dopamine (DA)-containing processes, which suggests that DA mediates reinforcement during appetitive conditioning. To test this possibility, methylergonovine was used to antagonize DA receptors. Methylergonovine (1 nM) blocked the pairing-specific increase in fictive feeding that is usually induced by in vitro classical conditioning. The present results and previous observation that methylergonovine also blocks the effects of contingent reinforcement in an in vitro analog of appetitive operant conditioning suggest that DA mediates reinforcement for appetitive associative conditioning of feeding in Aplysia.     

The Vienna comparative cognition technology (VCCT): An innovative operant conditioning system for various species and experimental procedures

This article describes a laboratory system for running learning experiments in operant chambers with various species. It is based on a modern version of a classical learning chamber for operant conditioning, the so-called ??Skinner box??. Rather than constituting a stand-alone unit, as is usually the case, it is an integrated part of a comprehensive technical solution, thereby eliminating a number of practical problems that are frequently encountered in research on animal learning and behavior. The Vienna comparative cognition technology combines modern computer, stimulus presentation, and reinforcement technology with flexibility and user-friendliness, which allows for efficient, widely automatized across-species experimentation, and thus makes the system appropriate for use in a broad range of learning tasks.     

Ideomotor silence: the case of complete paralysis and brain–computer interfaces (BCI)

The paper presents some speculations on the loss of voluntary responses and operant learning in long-term paralysis in human patients and curarized rats. Based on a reformulation of the ideomotor thinking hypothesis already described in the 19th century, we present evidence that instrumentally learned responses and intentional cognitive processes extinguish as a consequence of long-term complete paralysis in patients with amyotrophic lateral sclerosis (ALS). Preliminary data collected with ALS patients during extended and complete paralysis suggest semantic classical conditioning of brain activity as the only remaining communication possibility in those states.     

Cerebellar Substrates for Error Correction in Motor Conditioning

The authors evaluate a mapping of Rescorla and Wagner's (1972) behavioral model of classical conditioning onto the cerebellar substrates for motor reflex learning and illustrate how the limitations of the Rescorla-Wagner model are just as useful as its successes for guiding the development of new psychobiological theories of learning. They postulate that the inhibitory pathway that returns conditioned response information from the cerebellar interpositus nucleus back to the inferior olive is the neural basis for the error correction learning proposed by Rescorla and Wagner (Gluck, Myers, & Thompson, 1994; Thompson, 1986). The authors' cerebellar model expects that behavioral processes described by the Rescorla-Wagner model will be localized within the cerebellum and related brain stem structures, whereas behavioral processes beyond the scope of the Rescorla-Wagner model will depend on extracerebellar structures such as the hippocampus and related cortical regions. Simulations presented here support both implications. Several novel implications of the authors' cerebellar error-correcting model are described including a recent empirical study by Kim, Krupa, and Thompson (1998), who verified that suppressing the putative error correction pathway should interfere with the Kamin (1969) blocking effect, a behavioral manifestation of error correction learning. The authors also discuss the model's implications for understanding the limits of cerebellar contributions to associative learning and how this informs our understanding of hippocampal function in conditioning. This leads to a more integrative view of the neural substrates of conditioning in which the authors' real-time circuit-level model of the cerebellum can be viewed as a generalization of the long-term memory module of Gluck and Myers' (1993) trial-level theory of cerebellar-hippocampal interaction in motor conditioning.     

An Analysis of Feedback from a Behavior Analytic Perspective

The present paper presents a systematic analysis from a behavior analytic perspective of procedures termed feedback. Although feedback procedures are widely reported in the discipline of psychology, including in the field of behavior analysis, feedback is neither consistently defined nor analyzed. Feedback is frequently treated as a principle of behavior; however, its effects are rarely analyzed in terms of well-established principles of learning and behavior analysis. On the assumption that effectiveness of feedback procedures would be enhanced when their use is informed by these principles, we sought to provide a conceptually systematic account of feedback effects in terms of operant conditioning principles. In the first comprehensive review of this type, we compare feedback procedures with those of well-defined operant procedures. We also compare the functional relations that have been observed between parameters of consequence delivery and behavior under both feedback and operant procedures. The similarities observed in the preceding analyses suggest that processes revealed in operant conditioning procedures are sufficient to explain the phenomena observed in studies on feedback.     

The dynamics of operant conditioning

Existing models of operant learning are relatively insensitive to historical properties of behavior and applicable to only limited data sets. This article proposes a minimal set of principles based on short-term and long-term memory mechanisms that can explain the major static and dynamic properties of operant behavior in both single-choice and multiresponse situations. The critical features of the theory are as follows: (a) The key property of conditioning is assessment of the degree of association between responses and reinforcement and between stimuli and reinforcement; (b) the contingent reinforcement is represented by learning expectancy, which is the combined prediction of response-reinforcement and stimulus-reinforcement associations; (c) the operant response is controlled by the interplay between facilitatory and suppressive variables that integrate differences between expected (long-term) and experienced (short-term) events; and (d) very-long-term effects are encoded by a consolidated memory that is sensitive to the entire reinforcement history. The model predicts the major qualitative features of operant phenomena and then suggests an experimental test of theoretical predictions about the joint effects of reinforcement probability and amount of training on operant choice. We hypothesize that the set of elementary principles that we propose may help resolve the long-standing debate about the fundamental variables controlling operant conditioning.