Intentional and unintentional contributions to nonspecific preparation: electrophysiological evidence

The authors hypothesized that there are distinct intentional and unintentional influences on nonspecific preparation for a future event. In 2 experiments, participants responded to an imperative stimulus (S-sub-2) that was presented equiprobably either 400 ms or 1,200 ms after the offset of a warning stimulus (S-sub-1). During the S-sub-1-S-sub-2 interval, the authors measured the contingent negative variation (CNV), an event-related brain potential reflecting nonspecific preparation. S-sub-1 provided either no information or reliable information about the duration of the impending S-sub-1-S-sub-2 interval, thereby allowing an intentional influence on the state of preparation. The effect of S-sub-1 information on the CNV was approximately additive to the effect of the S-sub-1-S-sub-2 interval that was used on the preceding trial. This supports the view that the preceding S-sub-1-S-sub-2 interval contributes unintentionally to the state of nonspecific preparation guided by a process of trace conditioning.     

Sequential effects within a short foreperiod context: evidence for the conditioning account of temporal preparation

Responses to an imperative stimulus (IS) are especially fast when they are preceded by a warning signal (WS). When the interval between WS and IS (the foreperiod, FP) is variable, reaction time (RT) is not only influenced by the current FP but also by the FP of the preceding trial. These sequential effects have recently been proposed to originate from a trace conditioning process, in which the individuals learn the temporal WS-IS relationship in a trial-by-trial manner. Research has shown that trace conditioning is maximal when the temporal interval between the conditioned and unconditioned stimulus is between 0.25 and 0.60 s. Consequently, one would predict that sequential effects occur especially within short FP contexts. However, this prediction is contradicted by Karlin [Karlin, L. (1959). Reaction time as a function of foreperiod duration and variability. Journal of Experimental Psychology, 58, 185-191] who did not observe the typical sequential effects with short FPs. To investigate temporal preparation for short FPs, three experiments were conducted, examining the sequential FP effect comparably for short and long FP-sets (Experiment 1), assessing the influence of catch trials (Experiment 2) and the case of a very dense FP-range (Experiment 3) on sequential FP effects. The results provide strong evidence for sequential effects within a short FP context and thus support the trace conditioning account of temporal preparation.     

The foreperiod effect revisited: conditioning as a basis for nonspecific preparation

The foreperiod (FP) is the interval between a warning stimulus and the imperative stimulus. It is a classical finding that both the duration and the intertrial variability of FP considerably affects response time. These effects are invariably attributed to the participant's state of nonspecific preparation at the moment the imperative stimulus is presented. In this article, we examined a proposal by Los, S. A. (1996) [On the origin of mixing costs: exploring information processing in pure and mixed blocks of trials. Acta Psychologica, 94, 145-188] that the real-time development of nonspecific preparation during FP relies on the same principle as trace conditioning. To this end, we adjusted the formal conditioning model developed by Machado, A. (1997) [Learning the temporal dynamics of behavior. Psychological Review, 104 (2), 241-265], and fitted this model to a representative data set we obtained from nine participants. Although the model accounted for only a moderate proportion of the variance, it accurately reproduced several key features of the data. We therefore concluded that the model is a promising first step toward a theory of nonspecific preparation.     

The effect of a cross-trial shift of auditory warning signals on the sequential foreperiod effect

When a warning signal (WS) precedes an imperative signal (IS) by a certain amount of time (the foreperiod, FP), responses are speeded. Moreover, this effect is modulated by the FP length in the previous trial. This sequential FP effect has lately been attributed to a trace-conditioning mechanism according to which individuals learn (and re-learn) temporal relationships between the WS and the IS. Recent evidence suggests that sensory WS attributes are critical to trigger time-related response activation. Specifically, when WS modality is shifted in subsequent trials (e.g., from auditory to visual modality), the sequential FP effect becomes attenuated. This study examined whether the sequential FP effect is reduced only by between-modality shifts or whether this attenuation generalizes to cross-trial shifts of WS attributes within modalities. We compared dimensional (low vs. high tone frequency) and qualitative shifts (pure tone vs. noise) of equal-intense auditory WS events. The results of four experiments revealed that shifts of tone frequency did not, whereas shifts of qualitative tone characteristics did attenuate the sequential FP effect. These results support the view that the WS acts as a trigger cue that unintentionally activates responses at previously reinforced critical moments.     

The effective time course of preparation

In reaction time (RT) research on nonspecific preparation, the preparation period is often identified with the foreperiod (FP), the interval between the offset of a neutral warning stimulus (S1) and the onset of the reaction stimulus (S2). However, the “effective preparation period” may be longer than FP: nonspecific preparation may start prior to FP (e.g., at the onset of S1) and/or continue after it (i.e., in parallel with the reaction process). In four experiments, we factorially varied FP and an additional factor (S1-duration; S2-luminance; stimulus–response compatibility) that probed the effective preparation period outside the bounds of FP. By examining how equivalent RT–FP functions obtain at the different levels of the additional factor, we showed that nonspecific preparation (1) starts at the onset of S1 for brief FPs but at its offset for longer FPs and (2) continues in parallel with S2-encoding but stops prior to response selection.     

Dynamic adjustment of temporal preparation: Shifting warning signal modality attenuates the sequential foreperiod effect

We examined sequential effects in the variable foreperiod (FP) paradigm, which refer to the finding that responses to an imperative signal (IS) are fast when a short FP trial is repeated but slow when it is preceded by a long FP trial. The effect has been attributed to a trace-conditioning mechanism in which individuals learn the temporal relationship between a warning signal (WS) and the IS in a trial-by-trial manner. An important assumption is that the WS in a current trial (i.e., trial FP_n) acts as a conditioned stimulus, such that it automatically triggers the conditioned response at the exact critical moment that was imperative in the previous trial (i.e., trial FP_n−1). According to this assumption, a shift from one WS modality in trial FP_n−1 to another modality in trial FP_n is expected to eliminate or at least reduce the sequential FP effect. This prediction was tested in three experiments that included a random variation of WS modality and FP length within blocks of trials. In agreement with the prediction, a shift in WS modality attenuated the asymmetry of the sequential FP effect.     

Unlikely allies: embodied social cognition and the intentional stance

I argue that proponents of embodied social cognition (ESC) can usefully supplement their views if they enlist the help of an unlikely ally: Daniel Dennett. On Dennett??s view, human social cognition involves adopting the intentional stance (IS), i.e., assuming that an interpretive target??s behavior is an optimally rational attempt to fulfill some desire relative to her beliefs. Characterized this way, proponents of ESC would reject any alliance with Dennett. However, for Dennett, to attribute mental states from the intentional stance is not to attribute concrete, unobservable mental causes of behavior. Once this is appreciated, the kinship between IS??understood as a model of our quotidian interpretive practices??and ESC is apparent: both assume that quotidian interpretation involves tracking abstract, observable, behavioral patterns, not attributing unobservable, concrete, mental causes, i.e., both assume social cognition is possible without metapsychology. I argue that this affinity constitutes an opportunity: proponents of ESC can use IS as a characterization of the subpersonal basis for social cognition. In the process, I make my interpretation of IS more precise and relate it to current empirical literature in developmental psychology.     

Biasing the development of response topography with nonspecific positive and negative evocative verbal stimuli

Conditioned response (CR) rate and development of CR latency, rise time, and airpuff attenuation were examined for V- and C-form responders using two nonspecific command words, do and don't, as conditioned stimuli (CSs) in single-cue, double-cue, and differential eyelid conditioning. In both single-cue and differential conditioning, regardless of the command word used to signal the unconditioned stimulus (UCS), the Vs produced a higher response rate and learned a better UCS-attenuating response topography than the Cs. However, in a double-cue conditioning paradigm in which both command words were presented alone on different trials and reinforced, response latency was longer and puff attenuation poorer among Vs than when the UCS was signaled by a unique cue. In contrast, adding a second reinforced cue actually enhanced the development of puff-avoidant CR topographies among Cs compared to single-cue conditioning. These results and others indicate that response topography development is to some extent a labile process that can be biased toward either good or poor puff-avoidant properties and that the factors responsible for influencing CR topography differ for Vs and Cs.     

Pavlovian heart rate and jaw movement conditioning in the rabbit: effects of medial prefrontal lesions

An experiment was conducted in which jaw movements (JM) and heart rate (HR) were concomitantly assessed in rabbits during simple Pavlovian conditioning. A 2-s 1200-Hz tone was the conditioned stimulus (CS) and an intraoral 1-cc pulse of 0.5 M sucrose-water solution was the unconditioned stimulus (US). Sham and medial prefrontal (mPFC)-lesioned animals received paired CS/US training with a 70- to 75-dB CS and were compared with sham- and mPFC-lesioned animals that received explicitly unpaired CS/US presentations. The percentages of JM CRs were significantly greater in the paired than the unpaired groups, but mPFC lesions had no effect on this measure. Conditioned HR decelerations occurred only in the paired groups and then only during the first session of training. Moreover, these CS-evoked cardiac decelerations were somewhat attenuated by the mPFC lesion. CS-evoked HR accelerations, which were significantly greater in unpaired than in paired animals, occurred during the four subsequent sessions. These results suggest that a CS-evoked cardioinhibitory process, mediated by the mPFC, is engendered by Pavlovian appetitive conditioning, as has been previously demonstrated for aversive conditioning. However, during JM conditioning these inhibitory changes are quickly replaced by tachycardia, possibly related to increased nonspecific somatomotor activity, since the tachycardia was somewhat greater in the unpaired animals.     

Attentional and intentional cueing in a Simon task: An EEG-based approach

Advance information about the location of a stimulus (attentional cueing) does not affect the Simon effect (a shortening of manual response times whenever the position of a stimulus that is irrelevant for the task corresponds to the side of the response). However, advance information about the side of a response (intentional cueing) enhances the Simon effect. At first sight, these well-established results contradict two important assumptions about the origin of the Simon effect: (a) the effect originates at least in part in a covert shift of visual attention that forces the preparation of a response towards the location of the attentional shift and (b) interference between stimulus location and response side takes place within a response selection stage. We replicated the behavioral finding in a study that measured event-related potentials (ERPs) of the EEG. ERPs indicated that the mechanisms causing the Simon effect remain widely unaffected by advance information. Clear evidence for both response preparation and attentional shifts in the cue-target interval was found. Additionally, ERPs suggested that the increment of the Simon effect by intentional cueing might be due to perceptual factors rather than to an alteration in the mechanisms involved in the generation of a regular Simon effect. The implications of these data for the role of attention and of response selection in Simon tasks are discussed.     

Reminder-induced recovery of associations to an overshadowed stimulus

The possibility of reversing the deficit produced by overshadowing through the use of memory reactivation was investigated. Using lick suppression as a measure of associative strength, water-deprived rats were conditioned in a Pavlovian paradigm which produced reliable overshadowing of a flashing light by a tone. It was found, however, that exposure to the overshadowed stimulus outside of the conditioning context during the retention interval (reminder treatment) caused an increase in lick suppression during testing in animals that had undergone overshadowing, relative to nonreminded overshadowed animals. Subjects that received the reminder treatment but were conditioned without overshadowing showed no increase in lick suppression. Additional control groups ensured that the increase in suppression observed in the overshadowed subjects following reminder treatment was not due to nonspecific fear. The results suggest that the performance deficit produced by overshadowing is due at least in part to a reversible failure to efficiently retrieve associations to the overshadowed stimulus at the time of testing, rather than a failure to form those associations during conditioning.     

Overshadowing of subsequent events and recovery thereafter

Four experiments using a conditioned lick suppression preparation with rats were conducted to examine whether overshadowing of subsequent events could be obtained in Pavlovian backward conditioning (i.e. unconditioned stimulus [US] before conditioned stimulus [CS]), and to determine whether such overshadowing could be reversed without further training with the overshadowed CS, as has been reported in overshadowing of antecedent events. In Experiment 1, a backward-conditioned CS overshadowed a second backward-conditioned CS. Two posttraining manipulations, extinction of the overshadowing CS (Experiment 2) and shifting of the temporal relationship of the overshadowing CS to the US (Experiment 3), increased responding to the overshadowed CS. These results constitute the first unambiguous demonstration of stimulus competition between subsequent events using first-order conditioning, and they show that, like overshadowing with forward conditioning, such overshadowing is due, at least in part if not completely, to a failure to express information that had been acquired.     

Differential fear conditioning induces reciprocal changes in the sensory responses of lateral amygdala neurons to the CS(+) and CS(-)

In classical fear conditioning, a neutral sensory stimulus (CS) acquires the ability to elicit fear responses after pairing to a noxious unconditioned stimulus (US). As amygdala lesions prevent the acquisition of fear responses and the lateral amygdaloid (LA) nucleus is the main input station of the amygdala for auditory afferents, the effect of auditory fear conditioning on the sensory responsiveness of LA neurons has been examined. Although conditioning was shown to increase CS-evoked LA responses, the specificity of the changes in responsiveness was not tested. Because conditioning might induce nonspecific increases in LA responses to auditory afferents, we re-examined this issue in conscious, head-restrained cats using a differential conditioning paradigm where only one of two tones (CS(+) but not CS(-)) was paired to the US. Differential conditioning increased unit and field responses to the CS(+), whereas responses to the CS(-) decreased. Such changes have never been observed in the amygdala except in cases where the CS(-) had been paired to the US before and fear responses not extinguished. This suggests that fear conditioning is not only accompanied by potentiation of amygdalopetal pathways conveying the CS(+) but also by the depression of sensory inputs unpaired to noxious stimuli.     

Expectancy-learning and evaluative learning in human classical conditioning: affective priming as an indirect and unobtrusive measure of conditioned stimulus valence

It has been argued that in classical conditioning two processes might be operative. First, one may learn that the conditioned stimulus (CS+) is a valid predictor for the occurrence of the biologically negative or positive event (US; expectancy-learning). Second, one may learn to perceive the conditioned stimulus itself as a negative or positive stimulus, depending on the valence of the event it has been associated with (evaluative learning). Until the present, however, both forms of learning have been investigated using rather different conditioning procedures. Using a differential aversive conditioning preparation with pictures of human faces as CSs and an electrocutaneous stimulus as US, we were able to demonstrate that both forms of learning can co-occur. Moreover, the extent of evaluative learning in this aversive conditioning procedure did not significantly differ from the amount of evaluative learning in an evaluative conditioning procedure with positive and negative adjectives as USs, which was administered to the same participants. In the present study evaluative learning was not only indexed by direct evaluative ratings, but we introduced affective priming as an indirect and unobtrusive, reaction time based measure of stimulus valence. Finally, imagery instructions during acquisition did not facilitate expectancy-learning nor evaluative learning.     

Psychophysiological and behavioral differences as a function of age and Parkinson’s disease

Differences related to age and to specific neurological (Parkinsonian) damage were studied by contrasting, respectively, old (mean age 64 years) and young (27 years) subjects (N=15), and old-normal (N=15) and old-Parkinsonian (N=15) subjects. Both behavioral as well as psychophysiological dependent variables were employed. The behavioral measure was performance on a discrimination learning task, while the psychophysiological measures were based on habituation to a repeated tone stimulus and on a Pavlovian differential conditioning preparation. The behavioral task showed predominantly age-related differences, with the young learning faster. Another age-related difference was interpreted as showing a more insight-like learning process in the young. Response-bias (beta) values did not differ between groups. Age-related differences also emerged more clearly than specific-neurological-damage differences in the psychophysiological data. The older subjects manifested markedly less autonomic conditioning, which was probably due to a lowering of reactivity, as well as the emergence of habituation to the (loud-noise) unconditional stimulus. A correlational analysis of discrimination conditioning also yielded age-related differences. Most notably, reactivity played a greater role in conditioning in the old than in the young subjects. The results illustrated how psychophysiological measures can provide information that complements those provided by behavioral measures.     

A comparison of neuronal reactions during classical and instrumental conditioning under similar conditions

During the elaboration of an instrumental reflex, it is not obligatory to use a conditioned stimulus, which signals the necessity to generate an instrumental reaction in order to receive reinforcement. However, the presence of a conditioned stimulus simplifies analysis of instrumental reaction, which in this case is the response to the conditioned stimulus. On the other hand, it is necessary to distinguish between instrumental and classical conditioning, since in both cases the response to a conditioned stimulus increases. We studied neuronal analogs of classical and instrumental conditioning in the identified neurons responsible for the defensive closure of the pneumostome in the Helix mollusk under the same conditions. During classical conditioning, a mollusk received punishment after a tactile stimulus. During instrumental conditioning, a mollusk received punishment when an identified neuron did not generate an action potential in response to a tactile stimulus. The appearance of a painful stimulus did not depend on the generation or failure of a spike in the related control neuron. Another tactile stimulus, which was never paired with an unconditioned stimulus, was used as a discriminated stimulus. We also compared the behavior of such identified neurons during pseudoconditioning. The experiments were carried out in a semi-intact preparation. We examined how responses to the tactile and painful stimuli changed during different forms of training. It was shown that the dynamics of neuronal responses to a conditioned tactile stimulus were much more complex during instrumental conditioning and consisted of several phases. Throughout a learning session, neural system consecutively acquired information as to which kind of learning was presented, whether a reaction of the neural system must be generated or inhibited and which instrumental reaction is correct. We have demonstrated that response to a painful stimulus during classical conditioning decreases after short-term initial increase. However, during instrumental learning, the neurons controlling instrumental action remained highly sensitive to the unconditioned stimulus. Meanwhile, foreign neurons decreased their responses to the unconditioned stimulus. We may tentatively conclude that classical and instrumental paradigms are fundamentally different at the cellular level.     

Timing at the Start of Associative Learning

Some theories of associative learning imply that time plays a fundamental role in the acquisition process. Consistent with these theories, this paper presents evidence that the time from the onset of a conditioned stimulus (CS) until presentation of the unconditioned stimulus (US) is learned very rapidly at the start of training. We report two autoshaping studies and a study on aversive conditioning in goldfish in which we examine timing at the start of conditioning. We also review data from a number of other conditioning preparations, including fear-potentiated startle, appetitive conditioning in rats, and eyeblink conditioning in rabbits, that report conditioned response (CR) timing early in training. Acquisition speed and the very first expressions of conditioned responding often show sensitivity to the time of US presentation. In instances where temporal control is slowly expressed, it is likely due to performance factors, not to slow learning about time. In fact, the learning about time may be a necessary condition for associative learning.     

Task switches under Go/NoGo conditions and the decomposition of switch costs

Alternating switches between two simple S-R tasks are combined with Go/NoGo tasks. Non-switches after Go trials are assumed to selectively profit from stimulus driven repetition benefits, whereas switches after NoGo trials are assumed to be selectively delayed by stimulus driven negative priming. Intentionally driven reconfiguration costs are assessed by RT differences between switches after Go trials (no negative priming) and non-switches after NoGo trials (no repetition benefits). Experiment 1 indicates that with short preparation time repetition benefits, negative priming costs, and intentional components contribute approximately additively to switch costs. Experiment 2 confirms that the delay of switches after NoGo trials is indeed due to negative priming. Experiments 3 and 4 show that repetition benefits and intentional components of switch costs are properly assessed only if the settings assure that participants reconfigure the required task set in NoGo as well as Go trials.     

Blocking and unconditioned response diminution in human classical autonomic conditioning

This paper summarizes the results of three experiments on blocking and unconditioned response diminution in human SCR conditioning. Recent theories of conditioning propose that reduced processing of the unconditioned stimulus due to conditioning results in reduced unconditioned response magnitude and in blocking of conditioning of a second conditioned stimulus when it is in compound with the previously conditioned stimulus. In these experiments, clear evidence of diminution of the unconditioned response was obtained in all three studies, but blocking was found only in the third study, which employed a within-subject design and in which the blocking was observed only after at least one extinction trial on both the blocked and nonblocked stimuli. Order of testing was not a factor in this delayed blocking effect. These results do not support the view that blocking occurs because of reduced processing of the unconditioned stimulus in the unconditioned response diminution sense.