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Iris

Anatomy and Physiology · NLM MeSH · updated 2026-10-31 · version 1

Citations verified against PubMedNot yet clinician-reviewed10 referencesApproximately 4 min read

The most anterior portion of the uveal layer, separating the anterior chamber from the posterior.

Table of Contents

1. Definition2. Research landscape3. Reading the evidence4. Open-access studies5. Key reviews6. References7. About this article

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.

PubMed records by publication periodthrough 1999: 14,337 records; 2000-2009: 3,207 records; 2010-2019: 4,049 records; 2020 onward: 3,237 records14,337through 19993,2072000-20094,0492010-20193,2372020 onward
Records in PubMed indexed with this heading, by publication period. Real counts from NCBI E-utilities; the final period covers only part of the current year. Volume reflects how much has been published, not how strong the evidence is.
Show the data
PeriodRecords
through 199914,337
2000-20093,207
2010-20194,049
2020 onward3,237
Share of records by publication typeReviews: 3.5 percent; Meta-analyses: 0.1 percent; Randomized controlled trials: 3.0 percentReviews3.5% (876)Meta-analyses0.1% (36)Randomized controlled trials3.0% (744)
Share of all PubMed records on this heading that carry each publication type. Types overlap with other records, so the shares do not sum to 100%.
Publication-type chart data
Publication types among 24,830 PubMed records
TypeRecordsShare
Reviews8763.5%
Meta-analyses360.1%
Randomized controlled trials7443.0%

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 comparison — selected abstracts, not a systematic review
Study / authorsYearSource label / designEurope PMC citationsLicence / source
Pupil Dilation and Microsaccades Provide Complementary Insights into the Dynamics of Arousal and Instantaneous Attention during Effortful Listening
Contadini-Wright C, Magami K, Mehta N, Chait M
2023Research article
Design not recorded in structured metadata
38cc by
PMID 37127361 · DOI · PMC10312051
From pupil to the brain: New insights for studying cortical plasticity through pupillometry
Viglione A, Mazziotti R, Pizzorusso T
2023Review article
Design not recorded in structured metadata
35cc by
PMID 37063384 · DOI · PMC10102476
Self-regulating arousal via pupil-based biofeedback
Meissner SN, Bächinger M, Kikkert S, Imhof J, Missura S, Carro Dominguez M, et al.
2024Research article
Design not recorded in structured metadata
34cc by
PMID 37904022 · DOI · PMC10810759
Increased Baseline Pupil Size Linked to Uncertainty Avoidance in Decision Making
Kakaei E, Schlecht A, Hauser TU
2026Research article
Design not recorded in structured metadata
1cc by
PMID 41603188 · DOI · PMC12848648

Pupil Dilation and Microsaccades Provide Complementary Insights into the Dynamics of Arousal and Instantaneous Attention during Effortful Listening [1]

Research article Contadini-Wright C, Magami K, Mehta N, Chait M · · 2023 · cited 38 times (Europe PMC)

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]

Review article Viglione A, Mazziotti R, Pizzorusso T · · 2023 · cited 35 times (Europe PMC)

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]

Research article Meissner SN, Bächinger M, Kikkert S, Imhof J, Missura S, Carro Dominguez M, et al. · · 2024 · cited 34 times (Europe PMC)

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]

Research article Kakaei E, Schlecht A, Hauser TU · · 2026 · cited 1 times (Europe PMC)

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]
    Joshi S, Gold JI · Trends Cogn Sci · 2020 · cited 558 times (Europe PMC)
  • Research article Pupil dilation as an index of effort in cognitive control tasks: A review [6]
    van der Wel P, van Steenbergen H · Psychon Bull Rev · 2018 · cited 483 times (Europe PMC)
  • Research article A circuit for pupil orienting responses: implications for cognitive modulation of pupil size [7]
    Wang CA, Munoz DP · Curr Opin Neurobiol · 2015 · cited 271 times (Europe PMC)
  • Research article Best Practices and Advice for Using Pupillometry to Measure Listening Effort: An Introduction for Those Who Want to Get Started [8]
    Winn MB, Wendt D, Koelewijn T, Kuchinsky SE · Trends Hear · 2018 · cited 232 times (Europe PMC)
  • Research article The Pupil Dilation Response to Auditory Stimuli: Current State of Knowledge [9]
    Zekveld AA, Koelewijn T, Kramer SE · Trends Hear · 2018 · cited 218 times (Europe PMC)
  • Research article Neuromodulatory Correlates of Pupil Dilation [10]
    Larsen RS, Waters J · Front Neural Circuits · 2018 · cited 213 times (Europe PMC)

References

  1. 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
  2. Viglione A, Mazziotti R, Pizzorusso T. From pupil to the brain: New insights for studying cortical plasticity through pupillometry. . 2023. PMID 37063384 · DOI · PMC10102476
  3. 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
  4. Kakaei E, Schlecht A, Hauser TU. Increased Baseline Pupil Size Linked to Uncertainty Avoidance in Decision Making. . 2026. PMID 41603188 · DOI · PMC12848648
  5. Joshi S, Gold JI. Pupil Size as a Window on Neural Substrates of Cognition. Trends Cogn Sci. 2020. PMID 32331857 · DOI · PMC7271902
  6. 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
  7. Wang CA, Munoz DP. A circuit for pupil orienting responses: implications for cognitive modulation of pupil size. Curr Opin Neurobiol. 2015. PMID 25863645 · DOI
  8. 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
  9. Zekveld AA, Koelewijn T, Kramer SE. The Pupil Dilation Response to Auditory Stimuli: Current State of Knowledge. Trends Hear. 2018. PMID 30249172 · DOI · PMC6156203
  10. 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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At a glance

24,830
PubMed records on this heading
13%
published since 2020
876
review articles
744
randomized trials
10
verified references below

Counts come from PubMed on 2026-10-31; they show how much has been written, not how strong the evidence is.

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  • ChoroidAnatomy and Physiology
  • ConjunctivaAnatomy and Physiology

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Sources

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