Biology subjects

Ruuskanen, V.

Publications and source records attributed to Ruuskanen, V..

8 recordsLinked to original sources

bioRxiv2026

Larger Pupils are Associated with Improved Visual, but not Auditory, Near-Threshold Detection

Prestimulus pupil size is associated with near-threshold detection performance in both the visual and auditory domain, a relationship that is commonly attributed to arousal. However, given that larger pupils also let more light into the eye, in the visual domain this relationship is likely also driven by optical effects. To better understand this, we investigated how pupil size, skin conductance, and electroencephalographic (EEG) measures relate to detection performance in both a visual and an auditory task. We found that larger pupils were associated with higher sensitivity in the visual condition but lower sensitivity in the auditory condition. Skin conductance was negatively related to visual sensitivity, but unrelated to auditory sensitivity. EEG power measures were not related to sensitivity in either condition, though pupil size was positively correlated with alpha and beta power. Together, these results suggest that in visual detection the relationship between pupil size and performance is driven by both arousal and optics, whereas in auditory detection the relationship is driven solely by arousal. More broadly, our findings highlight the pupil as an active and functional component of the visual system.

neuroscience↗

bioRxiv2026

Subtle Pupil-Size Changes Associated With Exploration Do Not Affect Visual Sensitivity

When we feel restless and easily distracted, continuously switching tasks (exploration), our pupils tend to be large. In contrast, when we are calmly focused on a single task (exploitation), our pupils tend to be small. According to the Adaptive Gain Theory (AGT), a switch from exploitation to exploration is associated with an increase in norepinephrine in the locus coeruleus, which in turn triggers pupil dilation. However, the AGT does not provide a functional explanation of why exploration triggers pupil dilation. One possibility is that visual sensitivity, which increases with pupil size, is especially important during exploration. We set out to provide evidence consistent with this functional explanation, as well as to replicate two key previous results. Participants performed a four-armed bandit task, which induces both exploration and exploitation behavior. During the task, participants also needed to detect an occasional and unpredictable near-threshold peripheral flash. We replicated two key results: pupils were larger during exploration than during exploitation; and increased pupil size (overall, independent of exploration status) was associated with increased visual sensitivity. However, most importantly, we did not find that visual sensitivity was higher during exploration than during exploitation; probably, the reliable-yet-tiny increase in pupil size during exploration was too small to affect visual sensitivity. We conclude that key previous results are replicable; however, common experimental paradigms, such as the four-armed bandit task, induce only small changes in exploration behavior. Therefore, more powerful paradigms are required in order to test functional explanations of pupil-size changes during exploration and exploitation.

animal behavior and cognition↗

bioRxiv2026

Pupil Constriction Causes Activity in the Human Retina and Visual System

Pupil responses shape the earliest stages of visual perception by regulating the amount of light that enters the eye. How this affects visual processing is still poorly understood. Here we present evidence that pupil constriction causes activity in the human retina and visual system, independently of visual stimulation. Healthy human participants (N=119) viewed brief visual stimuli (light increments or decrements) while pupil size, retinal activity (electroretinogram), and brain activity (electroencephalogram) were recorded. As expected, visual stimulation triggered an initial burst of retinal activity followed by pupil constriction (the pupil light response). Importantly, we used trial-to-trial variability in constriction latency to reveal a previously unknown retinal response that is locked to pupil constriction, rather than to visual stimulation. Presumably, and in line with similar findings in mice, this constriction-locked retinal activity is a response to the sudden decrease in retinal light exposure that accompanies pupil constriction (although contribution of iris muscle activity is not conclusively ruled out). A similar constriction-locked response emerged later over visual cortex. Given these findings, an important open question is how the visual system maintains brightness constancy despite pupil-induced retinal and cortical activity.

neuroscience↗

bioRxiv2025

Pupil size correlates with near-threshold detection performance irrespective of stimulus colour, eccentricity, or retinal adaptation-state

In visual near-threshold detection tasks, larger pre-stimulus pupil size is associated with improved accuracy. However, previous studies have used black and white peripherally presented stimuli, leaving open the question of whether the relationship persists when targets differ in their colour or eccentricity. Here we addressed this question with three experiments that systematically varied the lighting conditions and target properties in a visual near-threshold detection task (data collected in 2022-2024). Light conditions ranged from dark to dim to bright, with the dark condition including a period of dark-adaptation. Possible target colours were blue and red on a black background (dark condition), blue and red on a grey background (dim condition), or yellow and cyan on a white background (bright condition). Possible target eccentricities ranged from parafoveal to peripheral, in a continuous manner (experiment 3) or as two predefined near and far eccentricities (experiments 1 & 2). Across all experiments we show that larger pre-stimulus pupil size is associated with improved performance. This large-pupil advantage is not systematically modulated by the colour or eccentricity of the targets, the illumination of the testing room, or retinal adaptation-state. We conclude that the phenomenon is robust, indicating that pupil size affects vision in a behaviourally relevant manner, regardless of the exact conditions. Significance statementBy showing that pupil size is robustly associated with performance in visual near-threshold detection tasks, we provide evidence for the functional role of endogenous pupil size fluctuations in visual perception.

neuroscience↗

bioRxiv2024

The interplay of spontaneous pupil-size fluctuations and EEG power in near-threshold detection

Detection of near-threshold stimuli depends on the properties of the stimulus and the state of the observer. In visual detection tasks, improved sensitivity is associated with larger pre-stimulus pupil size. However, it is still unclear whether this association is due to optical effects (more light entering the eye), correlations with arousal, correlations with cortical excitability (as reflected in alpha power), or a mix of these. To better understand this, we investigated the relative contributions of pupil size and power in the alpha, beta, and theta frequency bands on near-threshold detection. We found that larger pre-stimulus pupil size is associated with improved sensitivity and a more liberal criterion (the likelihood of reporting that a stimulus was detected). Importantly, the relationship between pupil size and sensitivity was not mediated by any of the measured neural variables; the relationship with criterion however was mediated by power in the beta band (12-30 Hz). Pupil size was positively correlated with power in the alpha and beta bands. Additionally, we show that pre-stimulus theta suppression (4 - 8 Hz) is associated with improved sensitivity and a more liberal criterion. Taken together, our results show an independent effect of pupil size on detection sensitivity that is not driven by cortical excitability, but may be driven by optical effects, arousal, or a mix of both.

neuroscience↗

bioRxiv2023

Behaviorally irrelevant feature matching increases neural and behavioral working memory readout: Support for activity-silent working memory

There is an ongoing debate about whether working memory (WM) maintenance relies on persistent activity and/or short-term synaptic plasticity. This is a challenging question, because neuroimaging techniques in cognitive neuroscience measure activity only. Recently, neural perturbation techniques have been developed to tackle this issue, such as visual impulse perturbation or "pinging", which reveals (un)attended WM content during maintenance. There are contrasting explanations of how pinging reveals WM content, which is central to the debate. Pinging could reveal mnemonic representations by perturbing content-specific networks or by increasing the neural signal-to-noise ratio of active neural states. Here we tested the extent to which the neural impulse response is patterned by the WM network, by presenting two different impulse stimuli. If the impulse interacts with WM networks, the WM-specific impulse response should be enhanced by physical overlap between the initial memory item and the subsequent external perturbation stimulus. This prediction was tested in a working memory task by matching or mismatching task-irrelevant spatial frequencies between memory items and impulse stimuli, as well as probes. Matching probe spatial frequency with memory items resulted in faster behavioral response times and matching impulse spatial frequency with memory items increased the specificity of the neural impulse response as measured from EEG. Matching spatial frequencies did neither result in globally stronger neural responses nor in a larger decrease in trial-to-trial variability compared to mismatching spatial frequencies. The improved neural and behavioural readout of irrelevant feature matching provide evidence that impulse perturbation interacts directly with the memory representations.

neuroscience↗

bioRxiv2023

Baseline Pupil Size Seems Unrelated to Fluid Intelligence, Working Memory Capacity, and Attentional Control

Over the past few years, several studies have explored the relationship between resting-state baseline pupil size and cognitive abilities, including fluid intelligence, working memory capacity, and attentional control. However, the results have been inconsistent. Here we present the findings from two experiments designed to replicate and expand previous research, with the aim of clarifying previous mixed findings. In both experiments, we measured baseline pupil size while participants were not engaged in any tasks, and assessed fluid intelligence using a matrix task. In one experiment we also measured working memory capacity (letter-number-sequencing task) and attentional control (attentional-capture task). We controlled for several personal and demographic variables known to influence pupil size, such as age and nicotine consumption. Our analyses revealed no relationship between resting-state pupil size (average or variability) and any of the measured constructs, neither before nor after controlling for confounding variables. Taken together, our results suggest that any relationship between resting-state pupil size and cognitive abilities is likely to be weak or non-existent.

neuroscience↗

bioRxiv2023

Causal Effects of Pupil Size on Visual Processing

The size of the eyes pupils determines how much light enters the eye and also how well this light is focused. Through this route, pupil size shapes the earliest stages of visual processing. Yet causal effects of pupil size on vision are poorly understood and rarely studied. Here we report the effects of both experimentally induced and spontaneous changes in pupil size on visual processing as measured through EEG. We compare these to the effects of stimulus intensity and covert visual attention, because previous studies have shown that these factors all have comparable effects on some common measures of early visual processing, such as detection performance and steady-state visual evoked potentials; yet it is still unclear whether these are superficial similarities, or rather whether they reflect similar underlying processes. Using a mix of neural-network decoding, ERP analyses, and time-frequency analyses, we find that induced pupil size, spontaneous pupil size, stimulus intensity, and covert visual attention all affect EEG responses, mainly over occipital and parietal electrodes, but--crucially--that they do so in qualitatively different ways. Induced and spontaneous pupil-size changes mainly modulate activity patterns (but not overall power or intertrial coherence) in the high-frequency beta range; this may reflect a causal effect of pupil size on oculomotor activity and/ or visual processing. In addition, spontaneous (but not induced) pupil size tends to correlate positively with intertrial coherence in the alpha band; this may reflect a non-causal relationship, mediated by arousal. Taken together, our findings suggest that pupil size has qualitatively different effects on visual processing from stimulus intensity and covert visual attention. This shows that pupil size causally affects visual processing, and provides concrete starting points for further study of this important yet understudied earliest stage of visual processing.

animal behavior and cognition↗