Effects of differential cue availability in an active avoidance CS for young and adult rats

In Experiment I, groups of 22- and 140+ -day-old rats were trained in acquisition and extinction of 1-way avoidance with a CS that consisted of the opening of a guillotine door 5 sec before US onset or a combination of door-opening plus a tonal signal that remained on until the occurrence of the motor response. Under both CS conditions, avoidance acquisition was similar at each age level. The extinction data indicated comparable performance for the young subjects but differential performance and greater resistance to extinction for the adult subjects. Adults trained with the door-opening CS persisted in responding for an entire series of 100 extinction trials, whereas the adults trained with the compound CS extinguished well within the 100-trial limit. A 2nd experiment included 10 pre-exposures of the simple or compound CS's prior to avoidance training. Although the pups were insensitive to pre-exposure effects, the adults that were pre-exposed and trained with identical CS'5 showed evidence of pre-exposure effects. Results of both experiments were interpreted as indicative of differential cue saliency between ages.     

Age-related differences in avoidance behavior in rats following CS preexposure

Weanling and adult rats were exposed to either 0 or 10 conditioned stimulus (CS) presentations prior to 1-way active-avoidance (AA) training. Although CS exposure retarded avoidance acquisition in the adults, it produced no effect in the pups during avoidance learning. Neither adult nor young rats demonstrated preexposure effects in extinction. A general extinction analysis showed that pups had less resistance to extinction than the adults, despite a comparable avoidance learning criterion between age groups. The lack of preexposure influences on performance by the pups was compared to previous findings of response inhibitory deficits in immature rats. The results were considered in light of selective attention interpretations of latent inhibition.     

Effects of posttraining damage to the pedunculopontine tegmental nucleus on conditioned stimulus transfer in two-way active avoidance in rats

The effects of posttraining excitotoxic lesions of the pedunculopontine tegmental nucleus (PPTg) on two-way active avoidance after changing the conditioned stimulus (CS) used during prelesion training were examined. Prelesion training was carried out with either a tone or a light as the CS, and this CS was changed during postlesion training. Replacing the tone with a light reduced the performance of control and lesioned rats, but the degree of reduction was higher in the latter. Replacing the light with a tone had slight detrimental effects in lesioned rats but not in controls. Thus, posttraining PPTg lesions slowed down the reacquisition of shuttle-box avoidance under conditions of CS transfer, an effect that may be attributable to disruption of attention and/or gating of sensory stimuli.     

Avoidance learning under delayed shock termination in prenatally x-irradiated rats

Groups of MP1 male albino rats were exposed to x-irradiation of 200 rad on Day 17 of gestation. Irradiated and control rats were trained in a shuttlebox with either immediate or .5-sec delayed termination of both the electric shock (US) and the conditioned stimulus (CS) on escape trials. An avoidance response always produced prompt CS termination. Irradiated rats showed a higher degree of avoidance behavior than controls and, under the delay condition, tended to run rapidly in the CS-US interval.     

Generalization of conditioned suppression during salicylate-induced phantom auditory perception in rats.

Tonal frequency generalization was examined in a total of 114 pigmented male rats, 60 of which were tested under the influence of salicylate-induced phantom auditory perception, introduced before or after lick suppression training. Thirty control subjects received saline injections, and the remaining 24 subjects served as noninjected controls of tonal background effects on generalization. Rats were continuously exposed to background noise alone or with a superimposed tone. Offset of background noise alone (Experiment I), or combined with onset or continuation of the tone (Experiments II and III) served as the conditioned stimulus (CS). In Experiment I, tone presentations were introduced only after suppression training. Depending on the time of salicylate introduction, a strong and differential influence on generalization gradients was observed, which is consistent with subjects' detection of salicylate-induced, high-pitched sound. Moreover, when either 12- or 3 kHz tones were introduced before or after Pavlovian training to mimic salicylate effects in 24 rats, the distortions in generalization gradients resembled trends obtained from respective salicylate injected groups. Experiments II and III were aimed at evaluating the masking effect of salicylate-induced phantom auditory perception on external sounds, with a 5- or a 10-kHz tone imposed continuously on the noise or presented only during the CS. Tests of tonal generalization to frequencies ranging from 4- to 11- kHz showed that in this experimental context salicylate-induced perception did not interfere with the dominant influence of external tones, a result that further strengthens the conclusion of Experiment I.     

Ontogenesis of learning. II. Variation in the rat's reflexive and learned responses to acoustic stimulation

The onotogenesis of the rat's reflexive and simple learned reactions to acoustic stimulation was investigated in five experiments. The pattern of restults suggests that (a)reflexive reactions to tonal stimulation emerge earleir in ontogenesis that the capacity to learn about thses stimuli, and (b)the age at which simple Pavlovian conditioning is first obsrved depedens on the nature of the acoustic conditioned stimulus (CS). Specifically, these data suggest that pups are not capable of simple Pavlovian conditioning to a 2000hyphen;Hz tone (CS) paired with an oral infusion of sucrose (US) until they are 14–15 days old. The same acoustic stimulus will, however, evoke reflexive reactions (startle inspiration) in pups as young as 10–12 days old. Additionally when a different auditory stimulus (an intermittent trains of clicks) serves as the CS, there is evidence of conditioning in pups only 12–13 days old. It is suggested that these age-related dissociations in the emergence of reflexive and simple learned responses to sound reflect a caudal-to-rostral maturational sequence of the components of the ascending auditory system.     

Early Pavlovian conditioning impairs later Pavlovian conditioning

Four experiments tested the effects in the rat of very early experience with stimuli to be used later for Pavlovian conditioning. Beginning on postnatal Day 12, prior to the development of substantial detection and effective perception of visual and auditory stimuli, rats were given five daily experiences with either lights or tones and a footshock known to be an effective unconditioned stimulus at these ages. Twenty-four hours after the last of these experiences, pairings of either the light or tone and the unconditioned stimulus were given with parameters established to yield a moderate degree of conditioning in untreated preweanlings (Experiment 1). Experiment 2 determined that early experience with paired or unpaired presentations of either the light or tone and the unconditioned stimulus resulted in a failure to condition to these same lights or tones on postnatal Day 17, although nontreated pups from the same litters conditioned quite effectively. Experiment 3 determined that this early conditioning experience with either paired or unpaired presentations of the lights or tones and the unconditioned stimulus yielded impaired conditioning on postnatal Day 17 in the alternative sensory modality as well, although again nontreated siblings conditioned quite effectively. Experiment 4 replicated the results of each of Experiments 2 and 3 and determined in addition that despite the impairment in conditioning that resulted from early paired or unpaired experience with the stimuli of conditioning, early experience with the individual stimuli of conditioning—with only the CS, the US, or the context—did not result in a similar impairment in conditioning. Although the results were unexpected, they may be understood in part in terms of intersensory competition during development, and there is precedent in the literature for similar interfering effects of early learning on later learning in a variety of species. © 1996 John Wiley & Sons, Inc.     

Conditioned hippocampal cellular response to a behaviorally silent conditioned stimulus.

Multiunit activity (MUA) was chronically recorded in the hippocampal CA3 field of rats using a blocking paradigm with conditioned suppression of lever pressing for food as the measure of conditioning. In Experiment 1, a classical blocking paradigm demonstrated the good conditionability of 2 stimuli (a light and a tone) and their respective ability to block each other. In Experiment 2, MUA was recorded in CA3 cells in rats submitted to a similar paradigm. Four groups received either tone (groups B and B1) or click (groups BC and B1C) conditioned stimulus (CS) presentations that were followed immediately by an electrical footshock (unconditioned stimulus, US). The rats were then given either 40 (groups B and BC) or 1 (groups B1 and B1C) tone + light-footshock presentations. During test sessions, the animals showed MUA responses to the added CS (light), with no conditioned suppression of lever presses occurring during CS presentations. The results of Experiment 3 strongly suggest that hippocampal increase in cellular activity to the light appeared at the first compound trial presentation. Conditioning to the light obtained by increasing the US intensity after the single compound trial suggests that the hippocampal response reflects a redundant CS-US association (light-shock). Long-term retention tests given 45 days after the end of conditioning revealed that behavioral and hippocampal responses could still be detected but only in response to the stimulus that had elicited a behavioral response during learning.     

Effects of conditioned stimulus pre-exposure on human electrodermal conditioning to fear-relevant and fear-irrelevant stimuli

The effects of conditioned stimulus (CS) pre-exposure and fear-relevance of the CS on human Pavlovian electrodermal conditioning were investigated. A differential delayed conditioning paradigm was used with a CS-unconditioned stimulus (US; shock) interval of 8 s. In Experiment 1, 64 subjects were randomized into four groups, two of which received fear-relevant stimuli and the other two fear-irrelevant stimuli. Half of the subjects were pre-exposed to the to-be-CSs and the other half to two not-to-be-CSs, with 15 exposure of each stimulus. During acquisition, subjects received 8 reinforced and 8 nonreinforced CS+ and CS- trials, and during the extinction phase 15 nonreinforced trials of each CS. Pre-exposure to the to-be-CSs retarded conditioning for the first and second interval anticipatory responses (FIRs and SIRs); that is, a latent inhibition effect was demonstrated, although the results for the FIR were inconclusive. The expected effects of fear-relevance were not revealed. Experiment 2 addressed the question whether the long pre-exposure period interfered with the frequently observed "preparedness effect" of higher resistance to extinction to fear-relevant stimuli. The design was similar to that of Experiment 1, but for half of the subjects the acquisition phase was initiated immediately after a short rest period, and for the other half acquisition started after an extended rest period, equal to the duration of the pre-exposure phase in Experiment 1. Twenty extinction trials of each CS were presented. A reliable difference in arousal in terms of spontaneous fluctuations was produced by the rest periods, but although differential conditioning was observed, no effect of fear-relevance was seen during extinction.     

Age-related disruption of trace but not delay classical conditioning of the rabbit's nictitating membrane response

Young (6 months of age) and old (36-60 months) New Zealand albino rabbits underwent classical conditioning of the nictitating membrane response in either a delay conditioning (Experiment 1) or a trace conditioning (Experiment 2) paradigm. There was no difference between old and young animals in acquisition of the conditioned response in the delay paradigm, nor were there any age-related differences in generalization to the tone conditioned stimulus (CS) or in sensitivity to the tone CS or eye shock unconditioned stimulus. In the trace conditioning paradigm, however, old animals acquired the conditioned response significantly more slowly than young rabbits. Because the same stimulus parameters and the same response were used in both experiments, it is unlikely that age-related differences in trace conditioning were due to stimulus sensitivity, motivation, or fatigue. The results are discussed in terms of how brain changes that accompany aging could differentially affect these two types of classical conditioning.     

Time and stimulus specificity in extinction of the conditioned nictitating membrane response in the rabbit (Oryctolagus cuniculus)

The present experiments demonstrated that, in the rabbit nictitating membrane preparation, a conditioned response (CR) can be selectively eliminated in one portion of a conditioned stimulus (CS) while it is still paired with the unconditioned stimulus (US). Rabbits were initially trained with two stimuli (tone, light). Each was paired with the US by using a mixture of two CS-US interstimulus intervals (ISIs): 200 ms and 1,200 ms in Experiment 1; 150 ms and 500 ms in Experiment 2. The CRs showed double peaks, one for each ISI. Subsequently, one CS (A) was trained with only the longer ISI, whereas the other CS (B) continued to be trained with both ISIs. Consequently, the CR peak based on the shorter ISI disappeared for CSA but not for CSB. The later CR peaks during both CSA and CSB were maintained. These results support time-based models of conditioning. Implications for proposed mechanisms of extinction are discussed.     

Ontogeny of eyeblink conditioned response timing in rats

Eyeblink conditioned response (CR) timing was assessed in adult and infant rats. In Experiment 1, adult rats were trained with a 150-ms tone conditioned stimulus (CS) paired with a periorbital shock unconditioned stimulus (US; presented at 200- or 500-ms interstimulus intervals [ISIs]). The rats acquired CRs with 2 distinct peaks that occurred just before the US onset times. Experiments 2 and 3 examined developmental changes in CR timing in pups trained on Postnatal Days 24-26 or 32-34. Experiment 3 used a delay conditioning procedure in which the tone CS continued throughout the ISIs. Pups of both ages exhibited robust conditioning. However, there were age-related increases in the percentage of double-peaked CRs and in CR timing precision. Ontogenetic changes in eyeblink CR timing may be related to developmental changes in cerebellar cortical or hippocampal function.     

Effects of conditioned stimulus preexposure on human electrodermal conditioning

Two experiments investigated the effects of conditioned stimulus (CS) preexposure on Pavlovian differential conditioning and extinction of the skin conductance response. In both experiments, half the subjects were exposed to 20 presentations each of the CS+ and CS-, and the other half were exposed to control stimuli. CS duration was 8 sec. The unconditioned stimulus in Experiment 1 (N = 48) was a 1000 Hz tone of 80 dB which signalled a reaction time requirement, and in Experiment 2 (N = 48), it was a 1 sec burst of white noise at 105 dB. The results of Experiment 1 indicated that no-preexposure groups displayed more CS+/CS- differentiation than preexposure groups during acquisition and more resistance to extinction, at least for the first interval anticipatory response. In addition, the results of Experiment 2 indicated that no-preexposure groups displayed more differentiation than preexposure groups in terms of the second interval anticipatory response. These data constitute a demonstration of the latent inhibition effect with human subjects, and imply that there is an intrinsic relationship between the orienting response and the conditioning process.     

Appetitive conditioning in neonatal rats: conditioned ingestive responding to stimuli paired with oral infusions of milk

Components of rat pups' ingestive responses to orally infused milk came to be elicited by a novel odor conditioned stimulus (cedar) that had been repeatedly paired with milk infusions (Experiment I). Pups responded specifically to one odor, and they did not generalize their conditioned responding to either another odor or an unscented airstream (Experiment II). Ingestive responses could also be conditioned to a vibrotactile CS paired with milk, although levels of conditioned responding were lower than were obtained with an odor CS (Experiment III). Pups' internal state determined the effectiveness of training, in that pups that were removed from their dam for 24 hr showed reliable conditioned responding, while nondeprived pups and dehydrated pups did not (Experiment IV). Finally, pups showed retention of conditioned responding for at least several days after training (Experiment V).     

Ontogeny of two-way avoidance in male and female rats

In Experiment I, 17-, 21-, 36-, 51-, 90-, and 200-day-old male and female rats were given a single session of 100 two-way avoidance (TWA) trials. In the 2nd experiment, males and females of these ages and 15 and 28 days of age that were obtained from a different source, weaned at a later age, and housed differently received TWA training. Results of both studies showed that avoidance of 15-, 17-, and 21-day-old rats is low, but avoidance increases from 21 to 51 days of age. Avoidance of 95- and 200-day-old animals was generally lower than 51-day-old rats. No significant gender differences appeared until 90 days of age; at this age avoidance of males was lower than females. In Experiment III, CS intensity was varied and the US intensity was lower than that used in Experiment I and II. Avoidance of 21- and 90-day-old rats was higher with a more intense CS, but 21-day-olds were still lower than adults. Avoidance of 17-day-old rats was not affected by CS intensity.     

Disrupted conditioned inhibition of the rabbit nictitating membrane response following mesencephalic lesions

Following acquisition of conditioned inhibition (CI) of the nictitating membrane, response, rabbits received radio-frequency lesions of the mesencephalon. The reinforced conditioned stimulus (CS) was light onset and the nonreinforced CS was the same light compounded with a tone. The unconditioned stimulus was electric shock to the paraorbital region of the eye. In accord by an earlier study by Mis [14], lesions disrupted conditioned inhibition by reducing the capacity to suppress conditioned responses on nonreinforced trials; conditioned responding on reinforced trials was unaffected. Lesions of the pretectum in the region of the posterior commissure and of the periaqueductal grey produced the greatest disruption CI, and fiber degeneration from these regions, as assessed by the Fink-Heimer method, projecting in a ventro-anterior direction to the posterior hypothalamus was implicated in CI disruption. This observation, together with indirect evidence that the dorsal noradrenergic bundle is involved in CI, suggested that conditioned inhibitors may act via a brainstem-hypothalamus reward system to antagonize the motivational support of the conditioned response.     

Disruption of contextual freezing, but not contextual blocking of fear-potentiated startle, after lesions of the dorsal hippocampus

The role of the dorsal hippocampus in contextual fear conditioning was investigated with a contextual blocking paradigm. In Experiment 1, rats were given pairings of a light conditioned stimulus (CS) and footshock after preexposure either to footshock or to the context alone. The group preexposed to footshock showed poorer fear conditioning to the light CS, as measured by the fear-potentiated startle reflex. In Experiment 2, a group preexposed to footshock in the same context showed poorer fear conditioning to the light CS than did a group preexposed to footshock in a different context, indicating contextual blocking of fear-potentiated startle. In Experiment 3, lesions of the dorsal hippocampus had no effect on contextual blocking, even though contextual freezing was disrupted. The sparing of contextual blocking indicated that contextual memory was intact following hippocampal lesions, despite the disruption of contextual freezing.     

Intertrial responses of rats in two-way avoidance learning to visual and auditory stimuli.

Learning and performance of two-way avoidance were investigated in a total of 68 rats trained with either a visual (change in illumination to darkness or to light) or an auditory (white noise of 70 or 60 dB intensity) conditioned stimulus (CS). Experiment I showed that the darkness CS produce lower avoidance performance and a much higher rate of intertrial responses than either a auditory or a compound (visual plus auditory) CS. A monotonic within-session increase of avoidance performance and a similar, but less regular increase of intertrial responses were found at the beginning of training in each group. In later sessions such trends were observed only in rats trained with a visual CS. Experiments II and III showed also rapid transfer of avoidance response and a corresponding change of intertrial response rate with the change of CS modality. When the compound CS was used, the effects of the visual element were completely overcome by the auditory one. Rats trained with a visual CS in Experiment IV showed a positive correlation between avoidance performance and the number of intertrial responses, which was more pronounced in earlier than in later training sessions. We consider the rise of intertrial behaviour as an adaptive response to the increase of task difficulty. CSi of different modalities differ not only in relative saliency, but also in the discriminability between their onset and offset. The modality of the CS influences not only avoidance performance but also the course of learning.     

The influence of natural preference for tactile stimuli on appetitive learning in rat pups

Stimuli of the same modality tend to be organized along a “natural preference scale.” This study examined the ability of six- and nine-day-old rat pups to acquire appetitive learning, when the CS was one of two differently “naturally preferred” tactile stimuli (floor textures: rug and plywood). In Experiment 1, all pups showed a relative natural preference for the rug texture over the plywood texture. Pups conditioned on the plywood texture (exposed to a sibling pup as the US) showed a robust increase in preference for the conditioned texture whereas pups conditioned on the rug texture showed a different and more moderate pattern of acquisition. Developmental differences were found only in extinction of the conditioned response: six-day-old (but not nine-day-old) pups displayed extinction of the response over four trials. Experiment 2 indicated that preexposure to the rug CS prior to conditioning (latent inhibition) did not interfere with the learning process on the rug texture. Two major alternative explanations for the differential learning patterns are discussed: motivational and catecholaminergic influences on learning. The results suggest that natural preferences may modulate early learning and memory processes. © 1997 John Wiley & Sons, Inc. Dev Psychobiol 30: 29–30, 1997     

Potentiation of conditioned freezing following dorsomedial prefrontal cortex lesions does not interfere with fear reduction in mice

In Experiment 1, an auditory conditioned stimulus (CS) was paired with footshock, except when it was preceded by another stimulus (a visual conditioned inhibitor [CI]). After conditioning, all mice displayed less CS-evoked freezing when the CI-CS compound was presented than when the CS was presented alone. However, lesions of the dorsomedial prefrontal cortex (dmPFC) potentiated CS-evoked freezing on each of the 2 sessions (i.e., CI CS and CS alone). In Experiment 2, mice were submitted to fear extinction (CS-alone presentation for 3 days). Lesioned mice exhibited a higher level of freezing behavior than controls on each of the 3 sessions. However, lesioned mice and controls displayed the same rate of reduction of freezing over the 3 days of extinction. These data in mice support previous studies in rats, which suggests that the dmPFC is not critical for either conditioned inhibition or extinction of acquired freezing behavior.