Topography of spatially directed conditioned responding: effects of context and trial duration.

Domesticated quail (Coturnix japonica) received a discrete conditioned stimulus (CS) at one end of the experimental chamber paired with the opportunity to copulate with a female quail (the unconditioned stimulus) in a goal box located 112 cm away. Approach to the CS (sign tracking) and approach to the goal area (goal tracking) were measured. The duration of exposure to the experimental context (C) was varied in Experiment 1, and the duration of the conditioning trials (T) was varied in Experiment 2 for independent groups, creating C/T ratios of 1.0, 1.5, 4.5, 45, and 180. Contrary to previous reports of a direct relation between the C/T ratio and conditioned responding, in the present experiments, a shift in the topography and stimulus control of conditioned behavior occurred. Low C/T ratios (1.0-4.5) produced goal tracking controlled by contextual cues, whereas high C/T ratios (45 and 180) produced sign tracking controlled by the discrete CS.

Pavlovian feed-forward mechanisms in the control of social behavior.

The conceptual and investigative tools for the analysis of social behavior can be expanded by integrating biological theory, control systems theory, and Pavlovian conditioning. Biological theory has focused on the costs and benefits of social behavior from ecological and evolutionary perspectives. In contrast, control systems theory is concerned with how machines achieve a particular goal or purpose. The accurate operation of a system often requires feed-forward mechanisms that adjust system performance in anticipation of future inputs. Pavlovian conditioning is ideally suited to subserve this function in behavioral systems. Pavlovian mechanisms have been demonstrated in various aspects of sexual behavior, maternal lactation, and infant suckling. Pavlovian conditioning of agonistic behavior has been also reported, and Pavlovian processes may likewise be involved in social play and social grooming. Several further lines of evidence indicate that Pavlovian conditioning can increase the efficiency and effectiveness of social interactions, thereby improving their cost/benefit ratio. We extend Pavlovian concepts beyond the traditional domain of discrete secretory and other physiological reflexes to complex real-world behavioral interactions and apply abstract laboratory analyses of the mechanisms of associative learning to the daily challenges animals face as they interact with one another in their natural environments.

Pavlovian conditioning of social-affiliative behavior in the Mongolian gerbil (Meriones unguiculatus).

Gerbils learned to approach a spatial-olfactory stimulus that signaled access to their pairmate. Experiments 1 and 3 used a discrimination procedure in which 1 conditioned stimulus (the CS+) was presented immediately before access to the pairmate and another (the CS-) was presented alone. Both male and female gerbils came to approach the CS+ sooner than the CS- and spent more time near the CS+ than the CS-. Discrimination learning was facilitated by making the CS+ and CS- spatially distinct (Experiment 3). Learning also was demonstrated in male gerbils, using a between-subjects design with a single CS. Pairing the CS with the opportunity for social interaction resulted in greater approach to the CS within 10 trials than presenting the CS and social opportunity in an unpaired fashion (Experiment 2). These findings demonstrate social-affiliative learning in the Mongolian gerbil. Similarities and differences between these findings and sexual conditioning effects in other species are discussed.

Differences in the sexual conditioned behavior of male and female Japanese quail (Coturnix japonica).

The conditioned responses of male and female Japanese quail (Coturnix japonica) were compared in a Pavlovian conditioning procedure in which presentation of a brief conditioned stimulus was immediately followed by the release of a copulation partner. Male quail vigorously approached the conditioned stimulus and were much more likely to enter the compartment housing their copulation partner than were female birds (Experiment 1). In females, sexual conditioning resulted in increased squatting (Experiment 2). This response was the reflection of sexual behavior rather than more general social behavior (Experiment 3). These findings provide the first definitive evidence of sexual learning in female quail and are consistent with the interpretation that sexual conditioning increases sexual arousal or receptivity in both sexes but the increase has different behavioral manifestations in male and female quail.

The behavior of the Japanese or domestic quail Coturnix japonica.

This paper reviews the literature pertaining to the behavior of the Japanese or domestic quail Coturnix japonica. Details are given of the classification, characteristics, domestication and the economic and research potential of the species. Further sections deal with sensation and perception (including taste and smell, vision and hearing), maintenance behavior (including feeding and drinking, dust bathing and thermoregulation), development and aging (including vocalization, filial imprinting, sexual imprinting, fear and avoidance responses, sexual maturation and aging), adult learning (including habituation, instrumental conditioning, Pavlovian conditioning and observational learning), photoperiodism, reproductive behavior (including courtship and mating, hormonal control and ontogeny of sexual differentiation, and male and female sexual behavior), parental behavior (including nest-site selection and nest building, incubation behavior and its hormonal control, and hen-chick relationships), and aggressive behavior and dominance (including agonistic behavior and the hormonal control of aggressive behavior).

One-trial appetitive conditioning in the sexual behavior system.

One-trial Pavlovian conditioning has been demonstrated with aversive reinforcers such as electric shock and lithium chloride. The present experiments using male Japanese quail were conducted to determine if an appetitive unconditioned stimulus (US), the opportunity to copulate with a female quail, can also generate one-trial learning. In Experiment 1, subjects were exposed to a contextual conditioned stimulus (CS) for 35 sec and then given copulatory opportunity. In Experiment 2, male birds were given 0, 2, or 4 min of exposure to the contextual CS before presentation of a female. In both cases, control groups received comparable exposure to both the contextual CS and the sexual US, but in an unpaired fashion. Subjects were tested for sexual learning by reintroducing them to the experimental context in the presence of a test object that incorporated the taxidermically prepared head of a female quail. In both Experiments 1 and 2, subjects for whom the contextual CS and sexual US had been paired displayed significantly more approach to the test object than did unpaired subjects.

The topography of sexually conditioned behaviour: effects of a trace interval.

In a trace conditioning procedure, subjects were presented with a 30-sec conditioned stimulus (CS) followed by a 30-sec trace interval. Delayed conditioning consisted of a 60-sec CS presentation followed by an unconditioned stimulus (US). Although conditioning developed with both procedures, the topography of the conditioned response differed. Sexual conditioned approach was evident in all of the subjects during the presentation of the CS. Trace-conditioned subjects moved away from the area where the CS had been presented during the trace interval but remained closer to the CS location than did an unpaired control. This reduction in the spatial specificity of the conditioned response was interpreted from a behaviour systems perspective. The trace interval presumably increased the perceived separation between the CS and the US and therefore elicited conditioned behaviour less specifically directed towards the CS.

Sign tracking versus goal tracking in the sexual conditioning of male Japanese quail (Coturnix japonica).

Pairings of a conditioned (CS) and unconditioned (US) stimulus can result in approach to either the CS (sign tracking) or the US (goal tracking). Increasing the spatial or temporal separation between the CS and a US food has been reported to results in a decline in sign tracking and an increase in goal tracking. In the present experiments, sign tracking was the predominant response of male quail (Coturnix japonica) to a CS that was presented up to 91 cm from the door through which a female was presented as the US (Experiment 1). Contrary to reported results with food reinforcement, goal tracking was not facilitated by increasing the spatial separation between the CS and US (Experiment 2) or by introducing a trace interval between them (Experiment 3). The preponderance of sign tracking may reflect an adaptive specialization of sexual conditioning or may be related to the absence of magazine training and the use of a moving US in the sexual conditioning procedures.

Learning and male-male sexual competition in Japanese quail (Coturnix japonica).

Male Japanese quail (Coturnix japonica) were trained individually to discriminate between 2 sounds presented at opposite ends of an outdoor aviary. One of the sounds (the positive conditioned stimulus [CS+]) was associated with the release of a female, and the other (the negative CS [CS-]) was presented alone. Which of the 2 sounds served as the CS+ (and which served as the CS-) was counterbalanced across subjects. The subjects came to approach their CS+ but did not move away from their CS-. After having been conditioned individually, the subjects were tested in pairs, with a single female released after the presentation of a stimulus that was the CS+ for one of the males and the CS- for the other male. During most of these tests, the male for whom the prefemale stimulus was the CS+ copulated with the female before the male for whom the prefemale stimulus was the CS-. These results indicate that learning can have an important role in competition for access to a reproductive partner.

Conditioned inhibition of social approach in male Japanese quail (Coturnix japonica) using visual exposure to a female.

Male Japanese quail (Coturnix japonica) learned to approach lights that predicted visual exposure to a female quail and showed significantly less approach to lights that predicted the absence of a female quail. Following discrimination training, subjects were given transfer summation tests in which a stimulus positively correlated with female exposure (CS +) was presented alone, simultaneously with a novel stimulus, and simultaneously with a stimulus negatively correlated with female exposure (CS -). Approach to the CS + was lower when the CS + and CS - were presented together than when the CS + was presented alone or when the CS + was presented with a novel stimulus. These findings demonstrate conditioned inhibition of sexual conditioned approach in Japanese quail.

Animal research in psychology. More than meets the eye of the general psychology student.

The general psychology course provides a unique opportunity to present the science of psychology to a wide audience. Informing the general public about the importance of animal research in psychology is especially important given contemporary concerns about animal rights and animal welfare. A study of 8 leading introductory psychology textbooks indicated that with the exception of principles of conditioning and learning, the contributions of animal research to psychology were often not explicitly acknowledged. In addition, major findings from animal research were frequently presented as if they had been obtained with humans. In obscuring the contributions of animal research, introductory psychology textbooks miss the opportunity to ensure that public policy be based on accurate information about the significance of this research to many areas of psychological science.

Effects of sexual conditioning of devaluing the US through satiation.

Male Japanese quail were conditioned to approach a visual CS that predicted an opportunity to copulate with a female quail. Subsequently the subjects were tested in both non-reinforced and reinforced test trials while sexually satiated and sexually deprived. In the satiation condition the sexual US was devalued by allowing the male subjects to copulate repeatedly with female birds just prior to testing. The subjects displayed significantly less approach to the CS when sexually satiated than when sexually deprived. These results are compatible with a mechanism in which performance of Pavlovian sexual approach conditioning is mediated by activation of a memorial representation of the US.

Stimulus control of copulatory behavior in sexually naive male Japanese quail (Coturnix japonica): effects of test context and stimulus movement.

Sexually experienced male quail (Coturnix japonica) are more likely to engage in copulatory behavior than sexually naive ones. These experiments suggest that sexual experience in a particular place may facilitate later copulatory responding because of increased familiarity with the contextual cues of the environment. Male quail in Experiment 1 did not copulate reliably with taxidermic models of females in a novel context, even though some of the subjects were allowed to copulate with female quail in their home cages. In contrast, sexually naive males in Experiments 2 and 3 copulated vigorously with taxidermic models of females in a familiar context. In Experiment 4, sexually naive males tested in an unfamiliar context were more likely to copulate with a moving than with a static model. The stimulus control of copulatory behavior in sexually naive male quail was similar to that in sexually experienced ones but only in familiar contexts.

Topography of sexually conditioned behavior in male Japanese quail (Coturnix japonica) depends on the CS-US interval.

The interval between exposure to a conditioned stimulus (CS) to male quail and access to a female (the unconditioned stimulus [US]) was varied from 0.5 to 20 min using a Pavlovian delayed conditioning procedure. Increasing the CS-US interval altered the spatial distribution of sexual conditioned behavior. With a short CS-US interval (1 min), conditioning resulted in the Ss remaining close to the CS and increasing their locomotor behavior near the CS. With a long CS-US interval (20 min), the Ss approached the CS to some degree, but their locomotor behavior was increased in areas further removed from the CS. Results are interpreted within the context of a behavior systems approach to the study of learning and indicate that the typical finding of an inverse relation between conditioned responding and the CS-US interval may be an artifact of the use of a limited range of behavioral measures.

Formulation of a behavior system for sexual conditioning.

Empirical and conceptual developments that led to the formulation of a behavior system for the sexual conditioning of male Japanese quail are described. Initial efforts concentrated on conditioning with localized conditioned stimuli and on identifying behavioral indices of conditioning. Later, learning about species-typical cues and about contextual cues was also explored, and it became evident that different types of cues control different aspects of sexual behavior. The results were used to formulate a behavior system containing both response and stimulus dimensions. In this system, contextual cues and local cues are assumed to elicit only general search behavior unconditionally. In contrast, unconditioned responses to species-typical cues of a female quail include general search, focal search, and copulatory behavior. General search, focal search, and copulatory behavior can become conditioned to local cues. Conditioning can also modify focal search behavior elicited by species-typical cues and can result in various modulatory influences between different types of stimuli. The behavior system approach provides a framework for organizing the diverse sexual conditioning effects and suggests future directions for investigation.

The nature of sexual reinforcement.

Sexual reinforcers are not part of a regulatory system involved in the maintenance of critical metabolic processes, they differ for males and females, they differ as a function of species and mating system, and they show ontogenetic and seasonal changes related to endocrine conditions. Exposure to a member of the opposite sex without copulation can be sufficient for sexual reinforcement. However, copulatory access is a stronger reinforcer, and copulatory opportunity can serve to enhance the reinforcing efficacy of stimulus features of a sexual partner. Conversely, under certain conditions, noncopulatory exposure serves to decrease reinforcer efficacy. Many common learning phenomena such as acquisition, extinction, discrimination learning, second-order conditioning, and latent inhibition have been demonstrated in sexual conditioning. These observations extend the generality of findings obtained with more conventional reinforcers, but the mechanisms of these effects and their gender and species specificity remain to be explored.

Sexual approach conditioning: unconditioned stimulus factors.

Unconditioned stimulus (US) factors were investigated in a Pavlovian sexual conditioning paradigm with male Japanese quail (Coturnix coturnix japonica). Copulation with a female quail was more effective than exposure to a female without copulation, but the latter also produced conditioned responding (Experiment 1). The greater effectiveness of copulatory opportunity as a US was probably not due to nonassociative effects of copulation (Experiment 2). Visual cues of the female, as presented on a taxidermic model, were not effective unless the subjects had prior sexual experience that increased their response to the model (Experiments 3a and 3b). Successful conditioning with noncopulatory female exposure is significant because it allows for sexual learning to occur in a broader range of circumstances than does conditioning with copulation and because it permits conducting multiple trials per day because males do not become satiated as rapidly.

Sexual approach conditioning: tests of unconditioned stimulus devaluation using hormone manipulations.

Contents of learning that result from conditioned-unconditioned stimulus pairings in sexual approach conditioning were explored with male Japanese quail (Coturnix coturnix japonica). Sexual motivation of subjects conditioned to approach an arbitrary stimulus in a Pavlovian sexual conditioning paradigm was reduced by exposing them to a short photoperiod. Decreased sexual motivation resulted in a decline in sexually conditioned approach behavior (Experiments 1 and 2). Responding was restored when subjects were returned to a long photoperiod (Experiment 1) and when exogenous testosterone was administered (Experiment 2). Decreased sexual motivation did not affect food-conditioned approach behavior (Experiment 3). These results suggest that sexually conditioned approach behavior is mediated by a representation of the unconditioned stimulus, which is activated by the conditioned stimulus.

Increased responding to female stimuli as a result of sexual experience: tests of mechanisms of learning.

Sexual experience increases the response of males to stimuli provided by female conspecifics in a variety of species. The mechanisms of learning involved in this type of phenomenon were explored in two experiments with Japanese quail. The results indicated that instrumental conditioning with copulatory opportunity is not necessary for the acquisition of responding to female cues, and responding is not facilitated by learning about the location of the female. However, the response of males to female stimuli (as well as to arbitrary stimuli associated with access to a female) was enhanced by the presence of sexually conditioned contextual cues. Substantial levels of responding also occurred to female stimuli in a context where the subjects never encountered a female quail before. This latter outcome is consistent with the possibility that stimuli from a female become directly associated with sexual reinforcement during the course of sexual experience. Similar forms of learning may be involved in the effects of sexual experience on the response of mammalian species to female odours.