Extinction of conditioned sexual responses in male Japanese quail (Coturnix japonica): role of species-typical cues

The authors examined how a conditioned stimulus (CS) that included species-typical cues affected the acquisition and extinction of conditioned sexual responses in male quail (Coturnix japonica). Some subjects were conditioned with a CS that supported sexual responses and included a taxidermic head of a female quail. Others were conditioned with a similar CS that lacked species-typical cues. Pairing the CSs with access to live females increased CS-directed behavior, with the head CS eliciting significantly more responding than the no-head CS. Responding to the head CS persisted during the 42-day, 126-trial extinction phase; responses to the no-head CS extinguished. Responding declined when the cues were removed or the subjects were sexually satiated. Possible functions and mechanisms of these effects are discussed.     

Sexual fetishism in a quail (Coturnix japonica) model system: test of reproductive success

In the present study, the authors explored the reproductive consequences of fetishistic behavior in a previously developed animal model of sexual fetishism (F. K?ksal et al., 2004). Male domesticated quail (Coturnix japonica) received sexual conditioning trials in which a terrycloth object (the conditioned stimulus [CS]) was paired with the opportunity to copulate with a female quail (the unconditioned stimulus). Approximately half of the male quail came to copulate with the CS object and were considered to have developed fetishistic behavior. Each of the male quail was then tested with a female quail, whose eggs were incubated to determine rates of fertilization. The CS object was present for 30 s before and during the copulation test. Fetishistic male quail were slower to achieve cloacal contact with the female quail and showed less efficient copulatory behavior. However, they fertilized a greater proportion of eggs than nonfetishistic male quail. These results are unexpected from previous studies of the relationship between reproductive success and copulatory behavior and are discussed in terms of how fetishistic behavior directed toward an inanimate object may modify male-female interactions.     

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.     

An animal model of fetishism

An animal model of sexual fetishism was developed with male Japanese quail based on persistence of conditioned sexual responding during extinction to an inanimate object made of terrycloth (Experiments 1 and 3). This persistent responding occurred only in subjects that came to copulate with the terrycloth object, suggesting that the copulatory behavior served to maintain the fetishistic behavior. Sexual conditioning was carried out by pairing a conditioned stimulus (CS) with the opportunity to copulate with a female (the unconditioned stimulus or US). Copulation with the CS object and persistent responding did not develop if the CS was a light (Experiment 1) or if conditioning was carried out with a food US (Experiment 2). In addition, subjects that showed persistence in responding to the terrycloth CS did not persist in their responding to a light CS (Experiment 3). The results are consistent with the hypothesis that conditioned copulatory behavior creates a form of self-maintenance that leads to persistent responding to an inanimate object. The development of an animal model of such fetishistic behavior should facilitate experimental analysis of the phenomenon.     

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.     

Effects on 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; in the deprivation condition, the males were not allowed to copulate for a day 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.     

Olfactory fear conditioning paradigm in rats: Effects of midazolam,propranolol or scopolamine

In rodents, fear conditioned responses are more pronounced toward olfactory stimulus, since olfaction is a dominant sense in these subjects. The present study was outlined to investigate if the association between coffee odor (CS1) and electrical footshock (US) would be an effective model for the study of fear-induced behavior and whether compounds used in humans for emotional-related disorders such as midazolam, propranolol, or scopolamine, applied during the different stages of fear conditioning (acquisition, consolidation and expression), affect the defensive responses to both, the olfactory CS1, and the context (CS2) where the CS1 had been presented (second order conditioning). The results revealed that five pairings between coffee odor (CS1) and electrical footshock (US) were able to elicit consistent defensive responses and a second order conditioning to the context (CS2). Midazolam (0.375–0.5 mg/kg; i.p.) treatment was able to interfere with the CS1–US association and with the consolidation of the aversive information. The propranolol (5–10 mg/kg; i.p.) treatment interfered with the CS1–US association, with the retention of fear memory and with the CS1–CS2 association. Propranolol also attenuated the expression of conditioned fear responses when applied before the CS1 test session. Scopolamine (0.6–1.2 mg/kg; i.p.) treatment impaired the acquisition of CS1–US and CS1–CS2 associations, and also disrupted the expression of conditioned fear responses when injected prior to the CS1 test session. These findings have pointed out the usefulness for the olfactory fear conditioning paradigm to investigate drug effects on the acquisition, consolidation and expression of fear conditioned responses.     

Contextual conditioning and the control of copulatory behavior by species-specific sign stimuli in male Japanese quail

Three experiments were conducted to identify species-specific sign stimuli sufficient to elicit copulatory behavior in male Japanese quail and to determine how learning is involved in the control of behavior by these sign stimuli. In Experiment 1, sexually experienced subjects were tested for copulatory behavior with a live female quail and with a model consisting of a female quail's head and neck mounted in front of a foam pad. Comparable levels of copulatory behavior were observed in the two tests, indicating that static visual cues provided by a female quail's head and neck are sufficient to elicit copulatory behavior in this species. Experiment 2 showed that male birds that previously received numerous opportunities to copulate with a live female quail in the test situation were significantly more likely to copulate with the head + neck model than were sexually inexperienced subjects. Experiment 3 showed that prior sexual experience with live quail facilitated responding to the head + neck model only if the sexual experience was provided in the same place where subjects were later tested with the model. This last finding suggests that sexual experience facilitates control of copulatory behavior by species-specific sign stimuli through contextual conditioning. Contextual conditioning may lower the threshold for sexual behavior with the result that a stimulus as impoverished as an immobile model containing only the head and neck of a female quail becomes sufficient to elicit normal levels of copulatory behavior. The results are also discussed as an example of conditioned stimulus facilitation of responding to an unconditioned stimulus.     

THE “WHERE IS IT?” REFLEX: AUTOSHAPING THE ORIENTING RESPONSE

The goal of this review is to compare two divergent lines of research on signal-centered behavior: the orienting reflex (OR) and autoshaping. A review of conditioning experiments in animals and humans suggests that the novelty hypothesis of the OR is no longer tenable. Only stimuli that represent biological “relevance” elicit ORs. A stimulus may be relevant a priori (i.e., unconditioned) or as a result of conditioning. Exposure to a conditioned stimulus (CS) that predicts a positive reinforcer causes the animal to orient to it throughout conditioning. Within the CS-US interval, the initial CS-directed orienting response is followed by US-directed tendencies. Experimental evidence is shown that the development and maintenance of the conditioned OR occur in a similar fashion both in response-independent (classical) and response-dependent (instrumental) paradigms. It is proposed that the conditioned OR and the signal-directed autoshaped response are identical. Signals predicting aversive events repel the subject from the source of the CS. It is suggested that the function of the CS is not only to signal the probability of US occurrence, but also to serve as a spatial cue to guide the animal in the environment.     

Footshock intensity and generalization in contextual and auditory-cued fear conditioning in the rat

The relationship between US (footshock) intensity and the two conditioned freezing responses (to acoustic CS and to "context") was investigated in fear conditioning. Administered footshock intensity was 0.00, 0.15, 0.30, 0.60, 0.90, and 1.20 mA to six different groups of 70-day-old male Albino Wistar rats. To measure contextual freezing, the animals were again placed inside the conditioning apparatus without acoustic CS and US presentation. To measure acoustic CS freezing, the animals were placed in a totally different apparatus and only the acoustic CS was presented. The 0.15 mA footshock intensity was not sufficient to condition the animals, in fact no freezing was exhibited as in the non-shocked control group. The 0.30 mA footshock intensity was sufficient only to condition the animals to the acoustic CS, whereas the 0.60 mA was sufficient to condition the animals both to acoustic CS and to context. Footshock intensities (0.90 and 1.20 mA) did not elicit any significant increase in conditioned freezing for either acoustic CS or context but at the highest one the generalization phenomenon appeared (freezing in the different context before presentation of acoustic CS). Acoustic CS freezing to all over-threshold intensities was longer than that to context. In conclusion, freezing responses to acoustic CS and context after increasing footshock intensities follow distinct patterns, and intermediate footshock intensities (0.60 and 0.90 mA) appear to be the most useful for eliciting conditioned freezing responses to acoustic CS and to context without inducing a generalized fear status contamination.     

Conditioned changes in pain reactivity I: A discrete CS elicits hypoalgesia, not hyperalgesia, on the tail-flick test

Prior work suggests that a stimulus which has been paired with an aversive event may elicit either an increase or decrease in pain reactivity. In four experiments we attempted to isolate the variables which determine the direction of the conditioned response. In all experiments, one stimulus was paired with shock (the CS+) while another was nonreinforced (the CS−). Pain reactivity was assessed by measuring tail withdrawal from radiant heat (the tail-flick test). Experiment 1 tested whether the amount of training plays a critical role. We found that the CS+ elicits longer tail-flick latencies irrespective of whether subjects received 6 or 24 CS-US pairings. Although there was some evidence that subjects may exhibit a general hyperalgesia at 2 min after the presentation of the conditioned stimulus, the results of Experiment 2 revealed that this effect was an artifact caused by differences in the amount of time between test trials. The results of Experiment 2 also showed that the difference observed between the CS+ and CS− was not due to a CS− induced hyperalgesia. Rather, it reflects a CS+ induced hypoalgesia. Experiment 3 evaluated whether the time of day at which testing occurs is important. We found that subjects exhibit conditioned hypoalgesia when tested in either the light or dark portion of the light-dark cycle. In Experiment 4 we examined whether it matters if subjects are tested in the training context or a neutral context. We found that subjects exhibit conditioned hypoalgesia during the CS+ irrespective of where they are tested. The results suggest that a CS+ elicits hypoalgesia, not hyperalgesia, on the tail-flick test over a wide range of conditions.     

A new model for sexual conditioning: the ring dove (Streptopelia risoria)

Sexual conditioning investigations have primarily used rats and domestic quail as subjects. Although much has been learned from such experiments, the relative simplicity of rat and quail reproductive behavior prohibits investigation of certain experimental questions about sexual conditioning. In contrast, the reproductive behavior of male and female ring doves (Streptopelia risoria) is complex, involving courtship, bonding, and parental care. In the present experiment, male ring doves that were presented with a visual conditioned stimulus paired with access to their pair-bonded mate (the unconditioned stimulus) showed sexual conditioned responding in the presence of the conditioned stimulus. These results represent the first evidence of sexual conditioning in ring doves and illustrate the potential of ring doves as a useful model for future sexual conditioning investigations.     

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.     

Learning with arbitrary versus ecological conditioned stimuli: Evidence from sexual conditioning

Laboratory investigations of Pavlovian conditioning typically involve the association of an arbitrary conditioned stimulus (CS) with an unconditioned stimulus (US) that has no inherent relation to the CS. However, arbitrary CSs are unlikely to become conditioned outside the laboratory, because they do not occur often enough with the US to result in an association. Learning under natural circumstances is likely only if the CS has a preexisting relation to the US. Recent studies of sexual conditioning have shown that in contrast to an arbitrary CS, an ecologically relevant CS is resistant to blocking, extinction, and increases in the CS-US interval and results in sensitized responding and stronger second-order conditioning. Although the mechanisms of these effects are not fully understood, these findings have shown that signature learning phenomena are significantly altered when the kinds of stimuli that are likely to become conditioned under natural circumstances are used. The implications of these findings for an ecological approach to the study of learning are discussed.     

Extinction of goal tracking also eliminates the conditioned reinforcing effects of an appetitive conditioned stimulus

Previous studies have suggested that the effects of extinction are response-specific. The present study investigated whether an extinction treatment that eliminated goal tracking elicited by an appetitive conditioned stimulus (CS) would also eliminate the conditioned reinforcing effects of that CS. Rats were first trained on a goal-tracking procedure in which an auditory CS was paired with a food unconditioned stimulus. Animals learned to approach the location where the food was delivered. In a subsequent phase, rats in one group received extinction training that eliminated the goal-tracking elicited by the CS. Rats in the other group did not experience extinction of the food-paired CS. Then, both groups received a test for conditioned reinforcement in which leverpresses resulted in the brief presentation of the stimulus previously paired with food. This stimulus did not act as a conditioned reinforcer in the group that had been subjected to extinction training, but did serve as a conditioned reinforcer in the group that did not experience extinction. These results indicate that the effects of extinction generalize from the approach-eliciting to the conditioned reinforcing effects of an appetitive CS.     

The effect of on‐ or off‐line extinction of a first‐order conditioned stimulus on a second‐order conditioned response in rats

In a water‐licking experiment with rats, the effects of the extinction of a first‐order conditioned stimulus (CS1) on the suppression of licking established for a second‐order conditioned stimulus (CS2) were explored. Extinction of the first‐order conditioned response (CR1) attenuated the conditioned suppression induced by CS2 when the rats had been allowed to lick water during the CR1 extinction phase. However, if water had been unavailable during the CR1 extinction phase, suppression by CS2 was not affected. The latter result is consistent with other studies of rats' conditioned suppression and suggests that the underlying mechanism of second‐order conditioning in this experiment is a connection between CS2 and the response elicited by CS1 rather than a CS2‐CS1 connection. The former result was interpreted as the CR1 extinction phase encouraging the rats to lick water despite the fear elicited by CS1, and thus, in testing, they licked despite the fear elicited by CS2.     

对体液免疫反应的条件反射性调节

以饮糖精水作为条件刺激(conditioned stimulus,CS),腹腔注射免疫抑制剂环磷酰胺作为非条件刺激(unconditioned stimulus,UCS)训练Wistar大鼠,3天后腹腔注射卵清蛋白(ovalbunfin,OVA)抗原,观察再次单独条件刺激对原发性体液免疫反应的作用。结果发现．一次CU-UCS结合训练导致CS组大鼠对再现糖精水产生厌恶反应,外周血中抗OVA-IgG抗体水平显著低于UCS组。两次CS-UCS结合训练并多次给予条件刺激后,CS组大鼠抗OVA-IgG的条件性免疫抑制效应与一次CS-UCS结合训练及再次给予一次条件刺激的反应类同。这些结果证明条件刺激增强了环磷酰胺对动物原发性体液免疫反应的抑制作用．这种条件性体液免疫抑制作用是相对稳定和有限度的,不易受条件反射建立参数的影响。     

Inhibition and superconditioning

Superconditioning is said to occur when learning an association between a conditioned stimulus (CS) and unconditioned stimulus (US) is facilitated by pairing the CS with the US in the presence of a previously established conditioned inhibitor. Previous demonstrations of superconditioning have been criticized because their control conditions have allowed alternative interpretations. Using a within–subjects autoshaping procedure, the present study unambiguously demonstrated superconditioning. The results support the view that superconditioning is the symmetric opposite of blocking.     

Social transmission of Pavlovian fear: fear-conditioning by-proxy in related female rats

Pairing a previously neutral conditioned stimulus (CS; e.g., a tone) to an aversive unconditioned stimulus (US; e.g., a foot-shock) leads to associative learning such that the tone alone will elicit a conditioned response (e.g., freezing). Individuals can also acquire fear from a social context, such as through observing the fear expression of a conspecific. In the current study, we examined the influence of kinship/familiarity on social transmission of fear in female rats. Rats were housed in triads with either sisters or non-related females. One rat from each cage was fear conditioned to a tone CS+ shock US. On day two, the conditioned rat was returned to the chamber accompanied by one of her cage mates. Both rats were allowed to behave freely, while the tone was played in the absence of the foot-shock. The previously untrained rat is referred to as the fear-conditioned by-proxy (FCbP) animal, as she would freeze based on observations of her cage-mate’s response rather than due to direct personal experience with the foot-shock. The third rat served as a cage-mate control. The third day, long-term memory tests to the CS were performed. Consistent with our previous application of this paradigm in male rats (Bruchey et al. in Behav Brain Res 214(1):80–84, 2010), our results revealed that social interactions between the fear conditioned and FCbP rats on day two contribute to freezing displayed by the FCbP rats on day three. In this experiment, prosocial behavior occurring at the termination of the cue on day two was significantly greater between sisters than their non-sister counterparts, and this behavior resulted in increased freezing on day three. Our results suggest that familiarity and/or kinship influences the social transmission of fear in female rats.     

Conditioned taste aversion is enhanced when the unconditioned stimulus is presented in a different context

Using a conditioned taste aversion procedure with rats as the subjects, two experiments examined the effect of presenting a conditioned stimulus (CS saccharin solution) in one context followed by an unconditioned stimulus (US LiCl) in a different context. Experiment 1 showed that animals which received the above-mentioned procedure (Group D) showed a more marked conditioned aversion to the CS than animals which were given both the CS and the US in the same context (Group S). Experiment 2 found that in both Group D and Group S, aversion to the CS increased when the subjects were exposed to the conditioned context after the conditioning. These findings supported the argument that the strength of the CS-US association acquired during conditioning is compared with that of the context-US to determine the magnitude of aversion revealed to the CS.