Affect, attention, or anticipatory arousal? Human blink startle modulation in forward and backward affective conditioning

Affect modulates the blink startle reflex in the picture-viewing paradigm, however, the process responsible for reflex modulation during conditional stimuli (CSs) that have acquired valence through affective conditioning remains unclear. In Experiment 1, neutral shapes (CSs) and valenced or neutral pictures (USs) were paired in a forward (CS → US) manner. Pleasantness ratings supported affective learning of positive and negative valence. Post-acquisition, blink reflexes were larger during the pleasant and unpleasant CSs than during the neutral CS. Rather than affect, attention or anticipatory arousal were suggested as sources of startle modulation. Experiment 2 confirmed that affective learning in the picture–picture paradigm was not affected by whether the CS preceded the US. Pleasantness ratings and affective priming revealed similar extents of affective learning following forward, backward or simultaneous pairings of CSs and USs. Experiment 3 utilized a backward conditioning procedure (US → CS) to minimize effects of US anticipation. Again, blink reflexes were larger during CSs paired with valenced USs regardless of US valence implicating attention rather than anticipatory arousal or affect as the process modulating startle in this paradigm.

The effect of emotional and attentional load on attentional startle modulation

The interactive effects of emotion and attention on attentional startle modulation were investigated in two experiments. Participants performed a discrimination and counting task with two visual stimuli during which acoustic eyeblink startle-eliciting probes were presented at long lead intervals. In Experiment 1, this task was combined with aversive Pavlovian conditioning. In Group Attend CS+, the attended stimulus was followed by an aversive unconditional stimulus (US) and the ignored stimulus was presented alone whereas the ignored stimulus was paired with the US in Group Attend CS−. In Experiment 2, a non-aversive reaction time task US replaced the aversive US. Regardless of the conditioning manipulation and consistent with a modality non-specific account of attentional startle modulation, startle magnitude was larger during attended than ignored stimuli in both experiments. Blink latency shortening was differentially affected by the conditioning manipulations suggesting additive effects of conditioning and discrimination and counting task on blink startle.

Attentional blink modulation in a reaction time task: performance feedback, warning stimulus modality, and task difficulty

The present research investigated the effect of performance feedback on the modulation of the acoustic startle reflex in a Go/NoGo reaction time task. Experiment 1 (n = 120) crossed warning stimulus modality (acoustic, visual, and tactile) with the provision of feedback in a between subject design. Provision of performance feedback increased the number of errors committed and reduced reaction time, but did not affect blink modulation significantly. Attentional blink latency and magnitude modulation was larger during acoustic than during visual and larger during visual than during tactile warning stimuli. In comparison to control blinks, latency shortening was significant in all modality conditions whereas magnitude facilitation was not significant during tactile warning stimuli. Experiment 2 (n = 80) employed visual warning stimuli only and crossed the provision of feedback with task difficulty. Feedback and difficulty affected accuracy and reaction time. Whereas blink latency shortening was not affected, blink magnitude modulation was smallest in the Easy/No Feedback and the Difficult/Feedback conditions. (C) 2002 Elsevier Science B.V. All rights reserved.

The effects of affective picture stimuli on blink modulation in adults and children

Two experiments examined blink modulation during viewing of pleasant, neutral and unpleasant picture stimuli in non-selected adults (N = 21) and children (N = 60) and children with anxiety disorders (N = 12). Blink reflexes were elicited by a white noise probe of 105 dB at lead stimulus intervals of 60, 240, 3500, and 5000 ms and during intertrial intervals. Blink modulation during unpleasant pictures was significantly different from blink modulation during neutral pictures at the 60 ms lead interval in children whereas adults showed no significant differences. Picture content had no differential effect on the extent of blink modulation for adults or children at the 240 ms lead interval. At the long lead intervals, blink modulation during unpleasant and pleasant pictures was significantly larger than during neutral pictures in adults. Picture valence did not differentially affect the extent of blink modulation at long lead intervals in children. Comparing the extent of blink modulation in anxious and non-selected children, blinks were significantly modulated during unpleasant pictures at the 60 ms lead interval for both groups. However, the extent of blink modulation was larger overall at this very short lead interval in anxious children. Children did not differ at other lead intervals. (C) 2004 Elsevier B.V. All rights reserved.

Attention vs. emotion: Affective learning and the startle reflex

The eye-blink startle reflex can be modulated by attentional and emotional processes. The reflex is facilitated during stimuli that engage attention. A linear pattern of emotional modulation has also been consistently demonstrated: the reflex is facilitated during unpleasant stimuli and attenuated during pleasant stimuli. However, during anticipation of pleasant or unpleasant stimuli it is unclear whether emotion or attention drives startle reflex modulation. This study used a differential learning procedure to investigate whether startle modulation during anticipation of a salient stimulus reflected emotional or attentional processes. In acquisition, a CS+ was paired with a pleasant or unpleasant US and a CS- was presented alone. In extinction, blink startle magnitude was measured during CS+ and CS-. Post-acquisition valence ratings and affective priming showed that CS+ had acquired the same affective value as the pleasant or unpleasant US with which it was paired. No differences in modulation of blink startle reflexes during pleasant CS+ and unpleasant CS+ were found throughout extinction. Blink startle facilitation occurred during CS+ but not CS- across the first third of extinction. Thus, attentional rather than emotional processes appeared to facilitate blink startle during anticipation of salient stimuli.

Verbal instruction abolishes fear conditioned to racial out-group faces

Previous research has shown resistance to extinction of fear conditioned to racial out-group faces, suggesting that these stimuli may be subject to prepared fear learning. The current study replicated and extended previous research by using a different racial out-group, and testing the prediction that prepared fear learning is unaffected by verbal instructions. Four groups of Caucasian participants were trained with male in-group (Caucasian) or out-group (Chinese) faces as conditional stimuli; one paired with an electro-tactile shock (CS+) and one presented alone (CS). Before extinction, half the participants were instructed that no more shocks would be presented. Fear conditioning, indexed by larger electrodermal responses to, and blink startle modulation during the CS+, occurred during acquisition in all groups. Resistance to extinction of fear learning was found only in the racial out-group, no instruction condition. Fear conditioned to a racial out-group face was reduced following verbal instructions, contrary to predictions for the nature of prepared fear learning.

The effects of unconditional stimulus valence and conditioning paradigm on verbal, skeleto-motor, and autonomic indices of human Pavlovian conditioning

The effects of unconditional stimulus (US) valence (aversive electro-tactile stimulus vs. nonaversive imperative stimulus of a RT task) and conditioning paradigm (delay vs. trace) on affective learning as indexed by verbal ratings of conditional stimulus (CS) pleasantness and blink startle modulation and on relational learning as indexed by electrodermal responses were investigated. Affective learning was not affected by the conditioning paradigm; however, electrodermal responses and blink latency shortening indicated delayed learning in the trace procedure. Changes in rated CS pleasantness were found with the aversive US, but not with the non-aversive US. Differential conditioning as indexed by electrodermal responses and startle modulation was found regardless of US valence. The finding of significant differential blink modulation and electrodermal responding in the absence of a change in rated CS pleasantness as a result of conditioning with a non-aversive US was replicated in a second experiment. These results seem to indicate that startle modulation during conditioning is mediated by the arousal level of the anticipated US, rather than by the valence of the CS. (C) 2002 Elsevier Science (USA). All rights reserved.

Neurochemistry of the afferents to the rat cochlear root nucleus: Possible synaptic modulation of the acoustic startle

Afferents to the primary startle circuit are essential for the elicitation and modulation of the acoustic startle reflex (ASR). In the rat, cochlear root neurons (CRNs) comprise the first component of the acoustic startle circuit and play a crucial role in mediating the ASR. Nevertheless, the neurochemical pattern of their afferents remains unclear. To determine the distribution of excitatory and inhibitory inputs, we used confocal microscopy to analyze the immunostaining for vesicular glutamate and GABA transporter proteins (VGLUT1 and VGAT) on retrogradely labeled CRNs. We also used reverse transcription-polymerase chain reaction (RT-PCR) and immunohistochemistry to detect and localize specific neurotransmitter receptor subunits in the cochlear root. Our results show differential distributions of VGLUT1- and VGAT-immunoreactive endings around cell bodies and dendrites. The RT-PCR data showed a positive band for several ionotropic glutamate receptor subunits, M1-M5 muscarinic receptor subtypes, the glycine receptor alpha 1 subunit (GlyR alpha 1), GABA(A), GABA(B), and subunits of alpha 2 and beta-noradrenergic receptors. By immunohistochemistry, we confirmed that CRN cell bodies exhibit positive immunoreaction for the glutamate receptor (GluR) 3 and NR1 GluR subunits. Cell bodies and dendrites were also positive for M2 and M4, and GlyR alpha 1. Other subunits, such as GluR1 and GluR4 of the AMPA GluRs, were observed in glial cells neighboring unlabeled CRN cell bodies. We further confirmed the existence of nor-adrenergic afferents onto CRNs from the locus coeruleus by combining tyrosine hydroxylase immunohistochemistry and tract-tracing experiments. Our results provide valuable information toward understanding how CRNs might integrate excitatory and inhibitory inputs, and hence how they could elicit and modulate the ASR. (C) 2008 IBRO. Published by Elsevier Ltd. All rights reserved.

A fast cholinergic modulation of the primary acoustic startle circuit in rats

Cochlear root neurons (CRNs) are the first brainstem neurons which initiate and participate in the full expression of the acoustic startle reflex. Although it has been suggested that a cholinergic pathway from the ventral nucleus of the trapezoid body (VNTB) conveys auditory prepulses to the CRNs, the neuronal origin of the VNTB-CRNs projection and the role it may play in the cochlear root nucleus remain uncertain. To determine the VNTB neuronal type which projects to CRNs, we performed tract-tracing experiments combined with mechanical lesions, and morphometric analyses. Our results indicate that a subpopulation of non-olivocochlear neurons projects directly and bilaterally to CRNs via the trapezoid body. We also performed a gene expression analysis of muscarinic and nicotinic receptors which indicates that CRNs contain a cholinergic receptor profile sufficient to mediate the modulation of CRN responses. Consequently, we investigated the effects of auditory prepulses on the neuronal activity of CRNs using extracellular recordings in vivo. Our results show that CRN responses are strongly inhibited by auditory prepulses. Unlike other neurons of the cochlear nucleus, the CRNs exhibited inhibition that depended on parameters of the auditory prepulse such as intensity and interstimulus interval, showing their strongest inhibition at short interstimulus intervals. In sum, our study supports the idea that CRNs are involved in the auditory prepulse inhibition of the acoustic startle reflex, and confirms the existence of multiple cholinergic pathways that modulate the primary acoustic startle circuit. © 2013 Springer-Verlag Berlin Heidelberg.

The effect of stimulus modality and task difficulty on attentional modulation of blink startle

The effects of the sensory modality of the lead Stimulus and of task difficulty on attentional modulation of the electrical and acoustic blink reflex were examined. Participants performed a discrimination and counting task with either two acoustic, two visual, or two tactile lead stimuli. In Experiment 1, facilitation of the electrically elicited blink was greater during task-relevant than during task-irrelevant lead stimuli. Increasing task difficulty enhanced magnitude facilitation for acoustic lead stimuli. In Experiment 2, acoustic blink facilitation was greater during task-relevant lead stimuli, but was unaffected by task difficulty. Experiment 3 showed that a further increase in task difficulty did not affect acoustic blink facilitation during visual lead stimuli. The observation that blink reflexes are facilitated by attention in the present task domain is consistent across a range of stimulus modality and task difficulty conditions.