Human temporal perception and the circadian system : effects of age, sleep pressure and light

Our intention with this work was to examine cognitive aspects of temporal regulation in humans and delineate possible connections to the circadian and sleep homeostatic systems. In the first experiment reported here, we assessed cognitive temporal orientation on the twenty-four hours scale, operationalized here as the conscious awareness about time of day in absence of external time cues. We studied two age groups under conditions of both high and low sleep pressure. Both age groups tended to overestimate actual time of day under both condtions, but this overestimation was more pronounced in older participants and significantly so under conditions of sustained wakefulness. Under both sleep pressure conditions, both age groups displayed a circadian oscillation in estimation errors which ran parallel to the endogenously generated oscillation in core body temperature. In the younger participants, under conditions of sustained wakefulness, this pattern was combined with an overall increase in the magnitude of estimation errors. Our results gave evidence that interval timing may provide a basis for cognitive temporal orientation via a mechanism of temporal integration. In two further studies, we assessed interval timing in young, healthy male participants for several stimulus magnitudes in the seconds range using temporal production and temporal reproduction under sustained wakefulness and controlled multiple 75 min sleep episodes. During temporal production conducted under controlled napping conditions all durations were over-produced and a slight linear increase in this behavior was observed across the protocol. During the reproduction task under napping conditions, smaller durations were over-reproduced but larger durations were under-reproduced, particularly during the biological night. During temporal production conducted under sustained wakefulness, all durations were under-produced. During reproduction, smaller durations were over-reproduced, especially during the second half of the protocol. Larger durations were under-reproduced under conditions of sustained wakefulness. Our findings suggest a rather weak effect of circadian phase on interval timing and a slightly stronger effect of elevated sleep pressure. We also observed a large degree of inter-individual variability in interval timing behavior which probably reflects the influence of further variables. Finally, we present an investigation into the effects of differential computer screen illumination on temporal perception. We found that interval timing responses are typically lowered under LED-screen illumination when compared to exposure to a CCFL-illuminated computer screen. In summary, we have demonstrated a possible relationship between interval timing and cognitive temporal orientation and were able to show that interval timing needs to be tested under strictly controlled conditions using several methods and stimulus durations in parallel. Our results regarding the effects of differential screen illumination technologies on interval timing should be of great interest to the development of biologically and ergonomically optimized user interfaces for information technology

Modeling circadian and sleep-homeostatic effects on short-term interval timing

Short-term interval timing i.e., perception and action relating to durations in the seconds range, has been suggested to display time-of-day as well as wake dependent fluctuations due to circadian and sleep-homeostatic changes to the rate at which an underlying pacemaker emits pulses; pertinent human data being relatively sparse and lacking in consistency however, the phenomenon remains elusive and its mechanism poorly understood. To better characterize the putative circadian and sleep-homeostatic effects on interval timing and to assess the ability of a pacemaker-based mechanism to account for the data, we measured timing performance in eighteen young healthy male subjects across two epochs of sustained wakefulness of 38.67 h each, conducted prior to (under entrained conditions) and following (under free-running conditions) a 28 h sleep-wake schedule, using the methods of duration estimation and duration production on target intervals of 10 and 40 s. Our findings of opposing oscillatory time courses across both epochs of sustained wakefulness that combine with increasing and, respectively, decreasing, saturating exponential change for the tasks of estimation and production are consistent with the hypothesis that a pacemaker emitting pulses at a rate controlled by the circadian oscillator and increasing with time awake determines human short-term interval timing; the duration-specificity of this pattern is interpreted as reflecting challenges to maintaining stable attention to the task that progressively increase with stimulus magnitude and thereby moderate the effects of pacemaker-rate changes on overt behavior

Impact of age, sleep pressure and circadian phase on time-of-day estimates

Orientation and self-location within the temporal fabric of the environment involves multiple organismic systems. While temporal self-location on the physiological level has been known for some time to be based on a 'biological clock' located within the hypothalamus, the mechanisms that participate in temporal position finding on the cognitive level are not yet fully understood. In order to probe the mechanisms that underlie this faculty, verbal estimates on time-of-day were collected at 3.75-h intervals from 16 young (7 males, 8 females; 20-31 years) and 16 older (8 males, 8 females; 57-74 years) subjects in a balanced crossover design during 40-h epochs of prolonged wakefulness and 40-h epochs of sleep satiation spent under constant routine conditions. An overestimation of clock time during prolonged wakefulness was found in both age-groups, with significantly larger errors for the older group (young: 0.5+/-0.2h; older: 1.5+/-0.2h, p>0.05). In both age-groups, estimation errors ran roughly parallel to the time course of core body temperature. However a significant interaction between time-of-day and age-group was observed (rANOVA, p>0.05): younger subjects exhibited similar estimation errors as the older subjects after 16 h of prior wakefulness, whereas the latter did not manifest decrements under high sleep pressure. Data collected under conditions of sleep satiation also displayed a diurnal oscillation in estimation errors and a general overestimation (young: 0.8+/-0.2h; older: 1.3+/-0.3h, p>0.05). Here however, the age-groups did not differ significantly nor was there an interactive effect between time-of-day and age-group. The effects of age, duration of wake time and circadian phase on temporal position finding are in line with predictions based on the idea that awareness about current position in time is derived from interval timing processes

Valence and agency influence striatal response to feedback in patients with major depressive disorder

BACKGROUND: Reduced sensitivity to positive feedback is common in patients with major depressive disorder (MDD). However, findings regarding negative feedback are ambiguous, with both exaggerated and blunted responses being reported. The ventral striatum (VS) plays a major role in processing valenced feedback, and previous imaging studies have shown that the locus of controls (self agency v. external agency) over the outcome influences VS response to feedback. We investigated whether attributing the outcome to one's own action or to an external agent influences feedback processing in patients with MDD. We hypothesized that depressed participants would be less sensitive to the feedback attribution reflected by an altered VS response to self-attributed gains and losses. METHODS: Using functional MRI and a motion prediction task, we investigated the neural responses to self-attributed (SA) and externally attributed (EA) monetary gains and losses in unmedicated patients with MDD and healthy controls. RESULTS: We included 21 patients and 25 controls in our study. Consistent with our prediction, healthy controls showed a VS response influenced by feedback valence and attribution, whereas in depressed patients striatal activity was modulated by valence but was insensitive to attribution. This attribution insensitivity led to an altered ventral putamen response for SA - EA losses in patients with MDD compared with healthy controls. LIMITATIONS: Depressed patients with comorbid anxiety disorder were included. CONCLUSION: These results suggest an altered assignment of motivational salience to SA losses in patients with MDD. Altered striatal response to SA negative events may reinforce the belief of not being in control of negative outcomes contributing to a cycle of learned helplessness

Functional lateralization of the anterior insula during feedback processing

Effective adaptive behavior rests on an appropriate understanding of how much responsibility we have over outcomes in the environment. This attribution of agency to ourselves or to an external event influences our behavioral and affective response to the outcomes. Despite its special importance to understanding human motivation and affect, the neural mechanisms involved in self-attributed rewards and punishments remain unclear. Previous evidence implicates the anterior insula (AI) in evaluating the consequences of our own actions. However, it is unclear if the AI has a general role in feedback evaluation (positive and negative) or plays a specific role during error processing. Using functional magnetic resonance imaging and a motion prediction task, we investigate neural responses to self- and externally attributed monetary gains and losses. We found that attribution effects vary according to the valence of feedback: significant valence × attribution interactions in the right AI, the anterior cingulate cortex (ACC), the midbrain, and the right ventral putamen. Self-attributed losses were associated with increased activity in the midbrain, the ACC and the right AI, and negative BOLD response in the ventral putamen. However, higher BOLD activity to self-attributed feedback (losses and gains) was observed in the left AI, the thalamus, and the cerebellar vermis. These results suggest a functional lateralization of the AI. The right AI, together with the midbrain and the ACC, is mainly involved in processing the salience of the outcome, whereas the left is part of a cerebello-thalamic-cortical pathway involved in cognitive control processes important for subsequent behavioral adaptations

Functional lateralization of the anterior insula during feedback processing

Effective adaptive behavior rests on an appropriate understanding of how much responsibility we have over outcomes in the environment. This attribution of agency to ourselves or to an external event influences our behavioral and affective response to the outcomes. Despite its special importance to understanding human motivation and affect, the neural mechanisms involved in self-attributed rewards and punishments remain unclear. Previous evidence implicates the anterior insula (AI) in evaluating the consequences of our own actions. However, it is unclear if the AI has a general role in feedback evaluation (positive and negative) or plays a specific role during error processing. Using functional magnetic resonance imaging and a motion prediction task, we investigate neural responses to self- and externally attributed monetary gains and losses. We found that attribution effects vary according to the valence of feedback: significant valence × attribution interactions in the right AI, the anterior cingulate cortex (ACC), the midbrain, and the right ventral putamen. Self-attributed losses were associated with increased activity in the midbrain, the ACC and the right AI, and negative BOLD response in the ventral putamen. However, higher BOLD activity to self-attributed feedback (losses and gains) was observed in the left AI, the thalamus, and the cerebellar vermis. These results suggest a functional lateralization of the AI. The right AI, together with the midbrain and the ACC, is mainly involved in processing the salience of the outcome, whereas the left is part of a cerebello-thalamic-cortical pathway involved in cognitive control processes important for subsequent behavioral adaptations

Evening exposure to a light-emitting diodes (LED)-backlit computer screen affects circadian physiology and cognitive performance

Many people spend an increasing amount of time in front of computer screens equipped with light-emitting diodes (LED) with a short wavelength (blue range). Thus we investigated the repercussions on melatonin (a marker of the circadian clock), alertness, and cognitive performance levels in 13 young male volunteers under controlled laboratory conditions in a balanced crossover design. A 5-h evening exposure to a white LED-backlit screen with more than twice as much 464 nm light emission {irradiance of 0,241 Watt/(steradian × m(2)) [W/(sr × m(2))], 2.1 × 10(13) photons/(cm(2) × s), in the wavelength range of 454 and 474 nm} than a white non-LED-backlit screen [irradiance of 0,099 W/(sr × m(2)), 0.7 × 10(13) photons/(cm(2) × s), in the wavelength range of 454 and 474 nm] elicited a significant suppression of the evening rise in endogenous melatonin and subjective as well as objective sleepiness, as indexed by a reduced incidence of slow eye movements and EEG low-frequency activity (1-7 Hz) in frontal brain regions. Concomitantly, sustained attention, as determined by the GO/NOGO task; working memory/attention, as assessed by "explicit timing"; and declarative memory performance in a word-learning paradigm were significantly enhanced in the LED-backlit screen compared with the non-LED condition. Screen quality and visual comfort were rated the same in both screen conditions, whereas the non-LED screen tended to be considered brighter. Our data indicate that the spectral profile of light emitted by computer screens impacts on circadian physiology, alertness, and cognitive performance levels. The challenge will be to design a computer screen with a spectral profile that can be individually programmed to add timed, essential light information to the circadian system in humans

Amygdala response to self-critical stimuli and symptom improvement in psychotherapy for depression

Background: Cognitive–behavioural therapy is efficacious in the treatment of major depressive disorder but response rates are still far from satisfactory. Aims: To better understand brain responses to individualised emotional stimuli and their association with outcome, to enhance treatment. Method: Functional magnetic resonance imaging data were collected prior to individual psychotherapy. Differences in brain activity during passive viewing of individualised self-critical material in 23 unmedicated out-patients with depression and 28 healthy controls were assessed. The associations between brain activity, cognitive and emotional change, and outcome were analysed in 21 patients. Results: Patients showed enhanced activity in the amygdala and ventral striatum compared with the control group. Non-response to therapy was associated with enhanced activity in the right amygdala compared with those who responded, and activity in this region was negatively associated with outcome. Emotional but not cognitive changes mediated this association. Conclusions: Amygdala hyperactivity may lessen symptom improvement in psychotherapy for depression through attenuating emotional skill acquisition