REFLECTIONS ON BEHAVIOR ANALYSIS AND EVOLUTIONARY BIOLOGY: A SELECTIVE REVIEW OF EVOLUTION SINCE DARWIN—THE FIRST 150 YEARS. EDITED BY M. A. BELL,D. J. FUTUYAMA,W. F. EANES, & J. S. LEVINTON

This review focuses on parallels between the selectionist sciences of evolutionary biology and behavior analysis. In selectionism, complex phenomena are interpreted as the cumulative products of relatively simple processes acting over time—natural selection in evolutionary biology and reinforcement in behavior analysis. Because evolutionary biology is the more mature science, an examination of the factors that led to the triumph of natural selection provides clues whereby reinforcement may achieve a similar fate in the science of behavior.     

Socio-cultural selection and the sculpting of the human genome: Cultures’ directional forces on evolution and development

This paper argues for culture as a significant determinant of the modern human genome. As progress in the fields of genetics and evolutionary biology have gained greater insights into evolutionary process, aspects of classical proposals explaining how phenotypic responses to environmental experience could make their way into the genotype have returned in new guises. Recent proposals recognize environmental context as a key source of genetic variation by directly altering selection pressures to mask and unmask adaptive value of expressed traits, by reorganizing the combination and expression of genes during ontogeny to provide novel variants for selection, and by influencing developmental trajectories through epigenetic systems acutely sensitive to ontogenetic contexts. The emergence of a robust human socio-cultural niche, shielding humans from more proximate evolutionary pressures that marked our ancestral evolution, has arguably provided the strongest directive force on modern human evolution. Language is discussed as an exemplar of a cultural niche with a powerful self-organizing dynamic and the potential to dramatically alter the human genome.     

A Test of the Habit-Goal Framework of Depressive Rumination and Its Relevance to Cognitive Reactivity

The aim of the present study was to test predictions derived from the habit-goal framework of depressive rumination and investigate its relevance to cognitive reactivity—another well-known vulnerability factor to depression. Formerly depressed (FD; n = 20) and never depressed (ND; n = 22) participants completed self-report measures of rumination, cognitive reactivity, and habitual characteristics of rumination (e.g., lack of awareness, control, intent). A standard mood-induction task was also used to measure cognitive reactivity and an outcome-devaluation task to measure general habit vs. goal-directed behavior control. Habitual characteristics of ruminative thoughts were greater in the FD group and were related to depressive brooding and cognitive reactivity, but not reflective pondering. Reliance on habit on the outcome-devaluation task was strongly correlated with number of depression episodes, although group differences were not observed in general habit vs. goal-directed control. Habitual characteristics of rumination (e.g., greater automaticity) may explain reactivity and persistence of negative thoughts in depression. Habitual behavior control may contribute to inflexible responding and vulnerability for depression episodes.     

When do adaptive developmental mechanisms yield maladaptive outcomes?

This article discusses 3 ways in which adaptive developmental mechanisms may produce maladaptive outcomes. First, natural selection may favor risky strategies that enhance fitness on average but which have detrimental consequences for a subset of individuals. Second, mismatch may result when organisms experience environmental change during ontogeny, for instance, because they move from one environment to another. Third, organisms may learn about their environment in order to develop an appropriate phenotype; when cues indicate the environmental state probabilistically, as opposed to deterministically, sampling processes may produce mismatch. For each source of maladaptation, we present a selection of the relevant empirical research and illustrate how models from evolutionary biology can be used to make predictions about maladaptation. We also discuss what data can be collected to test these models in humans. Our goal is to show that evolutionary approaches not only yield insights into adaptive outcomes but can also illuminate the conditions leading to maladaptation. This perspective provides additional nuance to the dialectic between the developmental psychopathology model and evolutionary developmental psychology.     

Operant and nonoperant vocal responding in the mynah: Complex schedule control and deprivation-induced responding

Several recent studies have been concerned with operant responses that are also affected by nonoperant factors, (e.g., biological constraints, innate behavior patterns, respondent processes). The major reason for studying mynah vocal responding concerned the special relation of avian vocalizations to nonoperant emotional and reflexive systems. The research strategy was to evaluate operant and nonoperant control by comparing the schedule control obtained with the vocal response to that characteristic of the motor responses of other animals. We selected single, multiple, and chain schedules that ordinarily produce disparate response rates at predictable times. In multiple schedules with one component where vocal responding (“Awk”) was reinforced with food (fixed-ratio or fixed-interval schedule) and one where the absence of vocal responding was reinforced (differential reinforcement of other behavior), response rates never exceeded 15 responses per minute, but clear schedule differences developed in response rate and pause time. Nonoperant vocal responding was evident when responding endured across 50 extinction sessions at 25% to 40% of the rate during reinforcement. The “enduring extinction responding” was largely deprivation induced, because the operant-level of naive mynahs under food deprivation was comparable in magnitude, but without deprivation the operant level was much lower. Food deprivation can induce vocal responding, but the relatively precise schedule control indicated that operant contingencies predominate when they are introduced.     

Functions of the environment in behavioral evolution

This paper explores some of the ways in which the environment functions with respect to behavior within an explanatory framework analogous to that of evolutionary biology. In both the behavioral and organic domains, the environment functions differently with respect to individual occurrences and evolutionary units. Within the behavioral domain, the problem of accounting for an occurrence of an operant instance differs from that of accounting for the existence of the operant unit of which the instance is a part. Maintaining these distinctions in levels of analysis within the behavioral domain, we focus first on operant units and operant instances as products of evolutionary processes occurring in the behavioral domain and second upon the causal role of the environment with respect to the existence of operant units and the occurrence of operant instances. The environment's function is selective with respect to origin, maintenance, suppression, and extinction of behavioral populations. At the level of operant instances, the environment has instantiating functions-evocative or alterative. Evocative functions are exemplified by discriminative relations, and alterative functions include both conditional and motivative relations. Implications are considered regarding extension of the analogy to more complex behavior-environment relations.     

Variation and selection: The evolutionary analogy and the convergence of cognitive and behavioral psychology

The empirical and theoretical work of both operant and cognitive researchers has increasingly appealed to evolutionary concepts. In particular, both traditional operant studies of extinction-induced behavior and cognitive investigations of creativity and problem solving converge on the fundamental evolutionary principles of variation and selection. These contemporary developments and their implications for the alleged preparadigmatic status of psychology are discussed.     

Countercontrol in behavior analysis

Countercontrol is a functional class of behavior that is part of Skinner's analysis of social behavior. Countercontrol refers to behavioral episodes comprised of socially mediated aversive controlling conditions and escape or avoidance responses that do not reinforce, and perhaps even punish, controllers' responses. This paper suggests that neglect of countercontrol in modern behavior analysis is unfortunate because the concept applies to interpersonal and social relations the fundamental operant principle that human behavior is both controlled and controlling-humans are not passive and inflexible. Countercontrol is addressed here in terms of conceptual status, contemporary developments in behavior analysis, its importance in a behavior-analytic approach to freedom and cultural design, applications, and research. The main conclusion is that Skinner's formulation of counter-control is scientifically supported and worthy of increased prominence in behavior analysis.     

Evolution and ontogeny of stress response to social challenges in the human child

The stress response systems of the human child are highly sensitive to social challenges. Because stress hormones can have negative developmental and health consequences, this presents an evolutionary paradox: Why would natural selection have favored mechanisms that elevate stress hormone levels in response to psychosocial stimuli? Two complementary hypotheses are considered: (a) maladaptation to the novelty of chronic stress in social environments, and (b) adaptive neural reorganization that facilitates the ontogeny of social competencies. Data on salivary cortisol, morbidity, and social environment from an 18 year study of child health in a rural community on the island of Dominica are examined from the perspective of these alternative hypotheses. Results indicate that difficult family environments and traumatic events are associated with elevated cortisol levels and higher morbidity. The long-term effects of traumatic early experiences on cortisol profiles are complex and indicate domain-specific effects, with normal recovery from physical stressors, but heightened response to negative-affect social challenges.     

Four routes of cognitive evolution

Four routes of cognitive evolution are distinguished: phylogenetic construction, in which natural selection produces qualitative change to the way a cognitive mechanism operates (language); phylogenetic inflection, in which natural selection biases the input to a cognitive mechanism (imprinting and spatial memory); ontogenetic construction, in which developmental selection alters the way a cognitive mechanism operates (face recognition and theory of mind); and ontogenetic inflection, in which developmental selection changes the input to a cognitive mechanism (imitation). This framework integrates findings from evolutionary psychology (i.e., all research on the evolution of mentality and behavior). In contrast with human nativist evolutionary psychology, it recognizes the adaptive significance of developmental processes, conserves the distinction between cognitive and noncognitive mechanisms, and encompasses research on human and nonhuman animals.     

PROBLEM BEHAVIOR DURING PREFERENCE ASSESSMENTS: AN EMPIRICAL ANALYSIS AND PRACTICAL RECOMMENDATIONS

Preferences of 2 children with developmental disabilities, whose functional analyses indicated that their problem behavior was maintained by access to tangible items, were assessed using three formats (i.e., paired stimulus [PS], multiple‐stimulus without replacement [MSWO], and free operant [FO]). The experimenter administered each format five times and compared levels of problem behavior across formats in a multielement design. Both participants exhibited problem behavior in PS and MSWO formats but not in the FO format. Results are discussed in terms of recommendations for practitioners.     

EVALUATION AND TRAINING OF YES‐NO RESPONDING ACROSS VERBAL OPERANTS

Topographically similar verbal responses may be functionally independent forms of operant behavior. For example, saying yes or no may have different functions based on the environmental conditions in effect. The present study extends previous research on both the assessment and acquisition of yes and no responses across contexts in children with language deficits and further examined the functional independence of topographically similar responses. All participants in the present study acquired yes and no responses within verbal operants (e.g., mands). However, generalization of the responses across novel verbal operants (e.g., tacts to intraverbals) did not occur without additional training, thus supporting Skinner's (1957) assertion of functional independence of verbal operants.     

The effect of type of behavior on behavior change caused by type of upcoming food substance

Previous research suggests that rats will decrease their consumption of a low-valued substance if a high-valued one will soon be available (anticipatory contrast), but will increase their rate of operant responding for a low-valued substance if a high-valued one will soon be available (positive induction). The present experiments tested whether rats would increase their operant rate of licking or lever pressing for 1% liquid-sucrose reinforcement when 32% sucrose reinforcement was upcoming in the same session. Results indicated that upcoming 32% sucrose increased rates of lever pressing for 1% sucrose, but did not produce similar increases in rates of licking. In fact, upcoming 32% sucrose significantly reduced lick rates in Experiment 2. The present results suggest that the different changes in behavior may be linked to the specific response that the subjects must engage in to obtain the reward (i.e., licking vs. lever pressing), and not to the function of the behavior (i.e., consummatory vs. operant) or to how frequently the substances are available (i.e., continuously vs. intermittently).     

Procedure for operant conditioning of the dog

Most reports arising from operant conditioning procedures have little or no emphasis on the actual behavior shaping or acquisition phase of the responses which are the cumulative frequency of the well-practiced act. There is a need for more detail for beginners or those desiring a clearer understanding of procedures leading to the finally reported data. The procedure reported here is typically used to compare animal subjects or conditions, such as drugs, both in the acquisition and final phases of behavior. Beginning with an unconditional response (e.g. feeding) a bridging stimulus is paired in classical conditioning fashion. From that point on the bridging stimulus and UCS (feeding—or shock) are used immediately as reward or reinforcement only for responses “in the direction of the final desired behavior. The number of timed standardized behavior shaping sessions to criterion is the best index of acquisition phase performance and the total number bar presses or the rate of bar pressing (slope) is the usual index of operant responding. With timid animals it is often necessary to administer tranquilizers.     

A general account of selection: biology, immunology, and behavior.

Authors frequently refer to gene-based selection in biological evolution, the reaction of the immune system to antigens, and operant learning as exemplifying selection processes in the same sense of this term. However, as obvious as this claim may seem on the surface, setting out an account of "selection" that is general enough to incorporate all three of these processes without becoming so general as to be vacuous is far from easy. In this target article, we set out such a general account of selection to see how well it accommodates these very different sorts of selection. The three fundamental elements of this account are replication, variation, and environmental interaction. For selection to occur, these three processes must be related in a very specific way. In particular, replication must alternate with environmental interaction so that any changes that occur in replication are passed on differentially because of environmental interaction. One of the main differences among the three sorts of selection that we investigate concerns the role of organisms. In traditional biological evolution, organisms play a central role with respect to environmental interaction. Although environmental interaction can occur at other levels of the organizational hierarchy, organisms are the primary focus of environmental interaction. In the functioning of the immune system, organisms function as containers. The interactions that result in selection of antibodies during a lifetime are between entities (antibodies and antigens) contained within the organism. Resulting changes in the immune system of one organism are not passed on to later organisms. Nor are changes in operant behavior resulting from behavioral selection passed on to later organisms. But operant behavior is not contained in the organism because most of the interactions that lead to differential replication include parts of the world outside the organism. Changes in the organism's nervous system are the effects of those interactions. The role of genes also varies in these three systems. Biological evolution is gene-based (i.e., genes are the primary replicators). Genes play very different roles in operant behavior and the immune system. However, in all three systems, iteration is central. All three selection processes are also incredibly wasteful and inefficient. They can generate complexity and novelty primarily because they are so wasteful and inefficient.     

Evolution and the nonlegal equivalent of aggressive criminal behavior

Within the framework of modern evolutionary theory, arguments are reviewed that the nonlegal equivalent of aggressive criminal behavior may have evolved by natural selection among mammals, particularly primates, as part of their overall approach to reproduction. If so, the commission of aggressive crimes (or their nonlegal equivalent) by humans, and even efforts to prevent fellow social group members from being victimized by aggressive crimes, may also be partially explainable in natural selection terms. The plausibility of this deduction was explored, first, by specifying the three elements that a human act must have to be regarded as an aggressive crime. Summarily, these were that (1) injury to a victim must be a likely result of the act, (2) the act must be intended, and (3) the act must elicit negative responses from those witnessing it. The primate behavior literature was examined for evidence that some behavior of nonhumans met all three conditions. Affirmative results were obtained. Therefore, while further research is in order, human aggressive criminal behavior, as well as human efforts to control it, seem to have close parallels in other primates. This would be consistent with the notion that aggressive criminal behavior (along with its condemnation by fellow group members) is part of a social system produced and sustained by natural selection.     

DELAYED REINFORCEMENT OF OPERANT BEHAVIOR

The experimental analysis of delay of reinforcement is considered from the perspective of three questions that seem basic not only to understanding delay of reinforcement, but, also, by implication, the contributions of temporal relations between events to operant behavior. The first question is whether effects of the temporal relation between responses and reinforcers can be isolated from other features of the environment that often accompany delays, such as stimuli or changes in the temporal distribution or rate of reinforcement. The second question is that of the effects of delays on operant behavior. Beyond the common denominator of a temporal separation between reinforcers and the responses that produce them, delay of reinforcement procedures differ from one another along several dimensions, making delay effects circumstance dependent. The final question is one of interpreting delay of reinforcement effects. It centers on the role of the response—reinforcer temporal relation in the context of other, concurrently operating behavioral processes.     

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.     

Arbitrarily applicable comparative relations: experimental evidence for a relational operant

Arbitrarily applicable derived relational responding has been argued by relational frame theorists to be a form of operant behavior. The present study examined this idea with 4 female participants, ages 4 to 5 years old, who could not perform a series of problem-solving tasks involving arbitrary more than and less than relations. In a combined multiple baseline (across responses and participants) and multiple probe design (with trained and untrained stimuli), it was shown that reinforced multiple-exemplar training facilitated the development of arbitrary comparative relations, and that these skills generalized not just across stimuli but also across trial types. The sequence of training identified potential prerequisites in the development of comparative relations (e.g., nonarbitrary comparative relations). Taken as a whole, the present data, along with previous work by others in this area, suggest that relating arbitrary events comparatively is an operant. The implications of this conclusion for the analysis of complex behavior are discussed.     

Behavior analysis and neuroscience: Complementary disciplines

Behavior analysis and neuroscience are disciplines in their own right but are united in that both are subfields of a common overarching field—biology. What most fundamentally unites these disciplines is a shared commitment to selectionism, the Darwinian mode of explanation. In selectionism, the order and complexity observed in nature are seen as the cumulative products of selection processes acting over time on a population of variants—favoring some and disfavoring others—with the affected variants contributing to the population on which future selections operate. In the case of behavior analysis, the central selection process is selection by reinforcement; in neuroscience it is natural selection. The two selection processes are inter‐related in that selection by reinforcement is itself the product of natural selection. The present paper illustrates the complementary nature of behavior analysis and neuroscience through considering their joint contributions to three central problem areas: reinforcement—including conditioned reinforcement, stimulus control—including equivalence classes, and memory—including reminding and remembering.