Definition and research guide
This page combines a MeSH definition, publication statistics and selected research abstracts. It is a starting point for reading the literature, not a medical textbook or a systematic evidence review.
Content available: 4 complete abstracts (838 words), 10 references. Full article text is not stored or reproduced here; follow the PMID, DOI or PMC links to read the original sources and their access terms.
Definition
The most anterior portion of the uveal layer, separating the anterior chamber from the posterior. It consists of two layers - the stroma and the pigmented epithelium. Color of the iris depends on the amount of melanin in the stroma on reflection from the pigmented epithelium.
Definition: NLM Medical Subject Headings (MeSH) scope note (Public domain (U.S. National Library of Medicine); attribution requested). View the MeSH record.
Where it fits in the MeSH classification
- Broader topics
- Anterior Eye Segment; Uvea
- Narrower topics
- Pupil
Research landscape
PubMed indexes 24,830 records tagged with the MeSH heading “Iris” (counts retrieved 2026-10-31). 3,237 of them (13.0%) were published from 2020 onward, and the busiest period so far is through 1999 with 14,337 records.
Reviews account for 876 records (3.5%) and meta-analyses for 36. Randomized controlled trials number 744 (3.0%) as a share of all records on the topic.
Show the data
| Period | Records |
|---|---|
| through 1999 | 14,337 |
| 2000-2009 | 3,207 |
| 2010-2019 | 4,049 |
| 2020 onward | 3,237 |
Publication-type chart data
| Type | Records | Share |
|---|---|---|
| Reviews | 876 | 3.5% |
| Meta-analyses | 36 | 0.1% |
| Randomized controlled trials | 744 | 3.0% |
Method: PubMed query "Iris"[MeSH Terms] with publication-type and date filters, retrieved 2026-10-31.
Reading the evidence in anatomy and physiology
Anatomy describes how body structures are built and physiology describes how they normally work, so most findings here are descriptive or mechanistic rather than evidence about treatment. Much of what is known comes from dissection, imaging, tissue studies and experiments in animals, and measurements in healthy volunteers can differ from those in people with disease or at different ages. Normal ranges vary between individuals and populations. Check whether a result was measured in people or in another species, the age and health of the participants, and the method used.
Evidence from open-access studies
These open-access studies come from PubMed’s best-match results for this heading (papers whose main topic it is), ranked here by citation count, with one recent addition. Each abstract is reproduced in full under its recorded open licence so that you can read the authors’ own words. They illustrate the literature; they are not a systematic summary of it. Citation counts measure attention, not reliability or clinical benefit.
| Study / authors | Year | Source label / design | Europe PMC citations | Licence / source |
|---|---|---|---|---|
| Pupil Dilation and Microsaccades Provide Complementary Insights into the Dynamics of Arousal and Instantaneous Attention during Effortful Listening | 2023 | Research article Design not recorded in structured metadata | 38 | cc by PMID 37127361 · DOI · PMC10312051 |
| From pupil to the brain: New insights for studying cortical plasticity through pupillometry | 2023 | Review article Design not recorded in structured metadata | 35 | cc by PMID 37063384 · DOI · PMC10102476 |
| Self-regulating arousal via pupil-based biofeedback | 2024 | Research article Design not recorded in structured metadata | 34 | cc by PMID 37904022 · DOI · PMC10810759 |
| Increased Baseline Pupil Size Linked to Uncertainty Avoidance in Decision Making | 2026 | Research article Design not recorded in structured metadata | 1 | cc by PMID 41603188 · DOI · PMC12848648 |
Pupil Dilation and Microsaccades Provide Complementary Insights into the Dynamics of Arousal and Instantaneous Attention during Effortful Listening [1]
Abstract reading summary — source excerpts
- Opening excerpt
- Listening in noisy environments requires effort- the active engagement of attention and other cognitive abilities- as well as increased arousal.
- Closing excerpt
- These results reveal the push-pull process through which auditory attention interfaces with the (visual) attention network that controls microsaccades, establishing microsaccades as a powerful tool for measuring auditory attention and its deficits.
- Interpretation limits
- Verbatim excerpts from the authors’ abstract; the closing excerpt may describe a finding, limitation or recommendation. Design, population, effect size and uncertainty have not been independently extracted or appraised. Read the complete abstract and linked paper before interpreting the findings.
Read complete abstract (360 words)
Listening in noisy environments requires effort- the active engagement of attention and other cognitive abilities- as well as increased arousal. The ability to separately quantify the contribution of these components is key to understanding the dynamics of effort and how it may change across listening situations and in certain populations. We concurrently measured two types of ocular data in young participants (both sexes): pupil dilation (PD; thought to index arousal aspects of effort) and microsaccades (MS; hypothesized to reflect automatic visual exploratory sampling), while they performed a speech-in-noise task under high- (HL) and low- (LL) listening load conditions. Sentences were manipulated so that the behaviorally relevant information (keywords) appeared at the end (Experiment 1) or beginning (Experiment 2) of the sentence, resulting in different temporal demands on focused attention. In line with previous reports, PD effects were associated with increased dilation under load. We observed a sustained difference between HL and LL conditions, consistent with increased phasic and tonic arousal. Importantly we show that MS rate was also modulated by listening load. This was manifested as a reduced MS rate in HL relative to LL. Critically, in contrast to the sustained difference seen for PD, MS effects were localized in time, specifically during periods when demands on auditory attention were greatest. These results demonstrate that auditory selective attention interfaces with the mechanisms controlling MS generation, establishing MS as an informative measure, complementary to PD, with which to quantify the temporal dynamics of auditory attentional processing under effortful listening conditions.SIGNIFICANCE STATEMENT Listening effort, reflecting the "cognitive bandwidth" deployed to effectively process sound in adverse environments, contributes critically to listening success. Understanding listening effort and the processes involved in its allocation is a major challenge in auditory neuroscience. Here, we demonstrate that microsaccade rate can be used to index a specific subcomponent of listening effort, the allocation of instantaneous auditory attention, that is distinct from the modulation of arousal indexed by pupil dilation (currently the dominant measure of listening effort). These results reveal the push-pull process through which auditory attention interfaces with the (visual) attention network that controls microsaccades, establishing microsaccades as a powerful tool for measuring auditory attention and its deficits.
Abstract reproduced from the open-access article under the recorded cc by licence; © the authors. PMID 37127361 · DOI · PMC10312051
From pupil to the brain: New insights for studying cortical plasticity through pupillometry [2]
Abstract reading summary — source excerpts
- Opening excerpt
- Pupil size variations have been associated with changes in brain activity patterns related with specific cognitive factors, such as arousal, attention, and mental effort.
- Closing excerpt
- Finally, we will review data suggesting that pupillometry could provide a quantitative and complementary measure of cortical plasticity also in pre-clinical studies.
- Interpretation limits
- Verbatim excerpts from the authors’ abstract; the closing excerpt may describe a finding, limitation or recommendation. Design, population, effect size and uncertainty have not been independently extracted or appraised. Read the complete abstract and linked paper before interpreting the findings.
Read complete abstract (194 words)
Pupil size variations have been associated with changes in brain activity patterns related with specific cognitive factors, such as arousal, attention, and mental effort. The locus coeruleus (LC), a key hub in the noradrenergic system of the brain, is considered to be a key regulator of cognitive control on pupil size, with changes in pupil diameter corresponding to the release of norepinephrine (NE). Advances in eye-tracking technology and open-source software have facilitated accurate pupil size measurement in various experimental settings, leading to increased interest in using pupillometry to track the nervous system activation state and as a potential biomarker for brain disorders. This review explores pupillometry as a non-invasive and fully translational tool for studying cortical plasticity starting from recent literature suggesting that pupillometry could be a promising technique for estimating the degree of residual plasticity in human subjects. Given that NE is known to be a critical mediator of cortical plasticity and arousal, the review includes data revealing the importance of the LC-NE system in modulating brain plasticity and pupil size. Finally, we will review data suggesting that pupillometry could provide a quantitative and complementary measure of cortical plasticity also in pre-clinical studies.
Abstract reproduced from the open-access article under the recorded cc by licence; © the authors. PMID 37063384 · DOI · PMC10102476
Self-regulating arousal via pupil-based biofeedback [3]
Abstract reading summary — source excerpts
- Opening excerpt
- The brain's arousal state is controlled by several neuromodulatory nuclei known to substantially influence cognition and mental well-being.
- Closing excerpt
- We provide evidence that pupil-based biofeedback makes the brain's arousal system accessible to volitional control, a finding that has tremendous potential for translation to behavioural and clinical applications across various domains, including stress-related and anxiety disorders.
- Interpretation limits
- Verbatim excerpts from the authors’ abstract; the closing excerpt may describe a finding, limitation or recommendation. Design, population, effect size and uncertainty have not been independently extracted or appraised. Read the complete abstract and linked paper before interpreting the findings.
Read complete abstract (150 words)
The brain's arousal state is controlled by several neuromodulatory nuclei known to substantially influence cognition and mental well-being. Here we investigate whether human participants can gain volitional control of their arousal state using a pupil-based biofeedback approach. Our approach inverts a mechanism suggested by previous literature that links activity of the locus coeruleus, one of the key regulators of central arousal and pupil dynamics. We show that pupil-based biofeedback enables participants to acquire volitional control of pupil size. Applying pupil self-regulation systematically modulates activity of the locus coeruleus and other brainstem structures involved in arousal control. Furthermore, it modulates cardiovascular measures such as heart rate, and behavioural and psychophysiological responses during an oddball task. We provide evidence that pupil-based biofeedback makes the brain's arousal system accessible to volitional control, a finding that has tremendous potential for translation to behavioural and clinical applications across various domains, including stress-related and anxiety disorders.
Abstract reproduced from the open-access article under the recorded cc by licence; © the authors. PMID 37904022 · DOI · PMC10810759
Increased Baseline Pupil Size Linked to Uncertainty Avoidance in Decision Making [4]
Abstract reading summary — source excerpts
- Opening excerpt
- Uncertainty is a key contributor to decision making, and humans show inconsistent attitudes towards it.
- Closing excerpt
- The specificity of our results suggests that uncertainty processing is dynamic and depends on (potentially noradrenergic) endogenous pupil fluctuations.
- Interpretation limits
- Verbatim excerpts from the authors’ abstract; the closing excerpt may describe a finding, limitation or recommendation. Design, population, effect size and uncertainty have not been independently extracted or appraised. Read the complete abstract and linked paper before interpreting the findings.
Read complete abstract (134 words)
Uncertainty is a key contributor to decision making, and humans show inconsistent attitudes towards it. Although excessive uncertainty-avoidance or uncertainty-seeking are hallmark symptoms of several mental conditions, the neural mechanism underlying uncertainty seeking and avoidance remains unclear. Here, we probed whether changes in pupil-linked arousal are indicative of uncertainty avoidance in humans. Investigating baseline pupil size to capture endogenous fluctuations across two experiments (N1 = 24, N2 = 21), we found that pretrial pupillary responses (as early as 700 ms prior to the onset of a trial) were closely related to uncertainty attitudes during multiarmed bandit tasks. Although increased baseline pupil size signalled avoidance in uncertainty-related decisions, it did not foreshadow value processing per se. The specificity of our results suggests that uncertainty processing is dynamic and depends on (potentially noradrenergic) endogenous pupil fluctuations.
Abstract reproduced from the open-access article under the recorded cc by licence; © the authors. PMID 41603188 · DOI · PMC12848648
Key reviews and meta-analyses
The most cited review-type papers among PubMed’s best-match results for this heading, listed with bibliographic details only. Follow the links for the full text and access terms.
- Research article Pupil Size as a Window on Neural Substrates of Cognition [5]
- Research article Pupil dilation as an index of effort in cognitive control tasks: A review [6]
- Research article A circuit for pupil orienting responses: implications for cognitive modulation of pupil size [7]
- Research article Best Practices and Advice for Using Pupillometry to Measure Listening Effort: An Introduction for Those Who Want to Get Started [8]
- Research article The Pupil Dilation Response to Auditory Stimuli: Current State of Knowledge [9]
- Research article Neuromodulatory Correlates of Pupil Dilation [10]
References
- Contadini-Wright C, Magami K, Mehta N, Chait M. Pupil Dilation and Microsaccades Provide Complementary Insights into the Dynamics of Arousal and Instantaneous Attention during Effortful Listening. . 2023. PMID 37127361 · DOI · PMC10312051
- Viglione A, Mazziotti R, Pizzorusso T. From pupil to the brain: New insights for studying cortical plasticity through pupillometry. . 2023. PMID 37063384 · DOI · PMC10102476
- Meissner SN, Bächinger M, Kikkert S, Imhof J, Missura S, Carro Dominguez M, et al.. Self-regulating arousal via pupil-based biofeedback. . 2024. PMID 37904022 · DOI · PMC10810759
- Kakaei E, Schlecht A, Hauser TU. Increased Baseline Pupil Size Linked to Uncertainty Avoidance in Decision Making. . 2026. PMID 41603188 · DOI · PMC12848648
- Joshi S, Gold JI. Pupil Size as a Window on Neural Substrates of Cognition. Trends Cogn Sci. 2020. PMID 32331857 · DOI · PMC7271902
- van der Wel P, van Steenbergen H. Pupil dilation as an index of effort in cognitive control tasks: A review. Psychon Bull Rev. 2018. PMID 29435963 · DOI · PMC6267528
- Wang CA, Munoz DP. A circuit for pupil orienting responses: implications for cognitive modulation of pupil size. Curr Opin Neurobiol. 2015. PMID 25863645 · DOI
- Winn MB, Wendt D, Koelewijn T, Kuchinsky SE. Best Practices and Advice for Using Pupillometry to Measure Listening Effort: An Introduction for Those Who Want to Get Started. Trends Hear. 2018. PMID 30261825 · DOI · PMC6166306
- Zekveld AA, Koelewijn T, Kramer SE. The Pupil Dilation Response to Auditory Stimuli: Current State of Knowledge. Trends Hear. 2018. PMID 30249172 · DOI · PMC6156203
- Larsen RS, Waters J. Neuromodulatory Correlates of Pupil Dilation. Front Neural Circuits. 2018. PMID 29593504 · DOI · PMC5854659
About this article
- Sources
- NLM Medical Subject Headings (definition, synonyms); NCBI PubMed (counts, identifiers); Europe PMC (open-access abstracts and citation counts).
- Verification
- Every reference was resolved against PubMed on 2026-10-31; title, year and DOI matched.
- Review state
- automated source verified; no human expert review has been recorded yet.
- Version
- 1, last updated 2026-10-31.
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Evidence strength
EvidenceHelix's computed view of the study designs behind this topic: how common meta-analyses, randomized trials and reviews are, how recent the literature is, and a plain-language reading. It describes the design mix of the literature. It is not a quality appraisal of any study, and it is not medical advice.