The eye of the humboldt penguin, Spheniscus humboldti: visual fields and schematic optics

Construction of a schematic eye indicates that the eye of Spheniscus humboldti is aquatic in design. The lens has a power of 100 dioptres (D) while (in air) the cornea has a power of 29 D. In air, the eye is myopic (approximately 28 D) but in water it is emmetropic. Minimum pupil size would seem insufficient to allow the pupil to function as a stenopaic aperture and increase depth of focus sufficiently to overcome the eye's aerial myopia. Entry into water reduces maximum image brightness by approximately three times. In air, the maximum width of the retinal binocular field is 45 degrees and this occurs approximately 10 degrees above the line of the bill. The bill intrudes into the retinal field and binocular field width in the plane containing the bill and the optic axes is 28 degrees. The vertical extent of the binocular field is 125 degrees. In the plane containing the optic axes the cyclopean field equals 282 degrees and the optic axes diverge by 116 degrees. In this plane the mean uniocular field is 155 degrees with the temporal hemifield approximately 11 degrees larger than the nasal hemifield. Entry into water reduces the widths of the visual fields such that maximum binocular field width is only 17 degrees and the vertical extent is reduced to about 80 degrees. Binocular vision is lost in the plane of the bill, and the uniocular retinal field is reduced by 32 degrees and the cyclopean field by 36 degrees.     

Ecomorphology of eye shape and retinal topography in waterfowl (Aves: Anseriformes: Anatidae) with different foraging modes

Despite the large body of literature on ecomorphological adaptations to foraging in waterfowl, little attention has been paid to their sensory systems, especially vision. Here, we compare eye shape and retinal topography across 12 species representing 4 different foraging modes. Eye shape was significantly different among foraging modes, with diving and pursuit-diving species having relatively smaller corneal diameters compared to non-diving species. This may be associated with differences in ambient light intensity while foraging or an ability to tightly constrict the pupil in divers in order to facilitate underwater vision. Retinal topography was similar across all species, consisting of an oblique visual streak, a central area of peak cell density, and no discernible fovea. Because the bill faces downwards when the head is held in the normal posture in waterfowl, the visual streak will be held horizontally, allowing the horizon to be sampled with higher visual acuity. Estimates of spatial resolving power were similar among species with only the Canada goose having a higher spatial resolution. Overall, we found no evidence of ecomorphological adaptations to different foraging modes in the retinal ganglion cell layer in waterfowl. Rather, retinal topography in these birds seems to reflect the ‘openness’ of their habitats.     

Eye size in birds and the timing of song at dawn

Why do different species of birds start their dawn choruses at different times? We test the hypothesis that the times at which different species start singing at dawn are related to their visual capability at low light intensities. Birds with large eyes can achieve greater pupil diameters and hence, all other things being equal, greater visual sensitivity and resolution than birds with small eyes. We estimated the maximum pupil diameter of passerine birds by measuring the diameter of the exposed eye surface, and measured the times of the first songs at dawn of songbirds present in different bird communities, and the light intensities at these times. Using phylogenetic comparative analyses, we found that songbirds with large eyes started to sing at lower light intensities (and therefore earlier) than species with smaller eyes. These relationships were stronger when differences in body size were controlled for statistically, and were consistent between two phylogenies and when species were treated as independent data points. Our results therefore provide robust support for the hypothesis that visual capability at low light levels influences the times at which birds start to sing at dawn.     

Vision and the foraging technique of Great Cormorants Phalacrocorax carbo: pursuit or close‐quarter foraging?

Predatory diving birds, such as cormorants (Phalacrocoracidae), have been generally regarded as visually guided pursuit foragers. However, due to their poor visual resolution underwater, it has recently been hypothesized that Great Cormorants do not in fact employ a pursuit-dive foraging technique. They appear capable of detecting typical prey only at short distances, and primarily use a foraging technique in which prey may be detected only at close quarters or flushed from a substratum or hiding place. In birds, visual field parameters, such as the position and extent of the region of binocular vision, and how these are altered by eye movements, appear to be determined primarily by feeding ecology. Therefore, to understand further the feeding technique of Great Cormorants we have determined retinal visual fields and eye movement amplitudes using an ophthalmoscopic reflex technique. We show that visual fields and eye movements in cormorants exhibit close similarity with those of other birds, such as herons (Ardeidae) and hornbills (Bucerotidae), which forage terrestrially typically using a close-quarter prey detection or flushing technique and/or which need to examine items held in the bill before ingestion. We argue that this visual field topography and associated eye movements is a general characteristic of birds whose foraging requires the detection of nearby mobile prey items from within a wide arc around the head, accurate capture of that prey using the bill, and visual examination of the caught prey held in the bill. This supports the idea that cormorants, although visually guided predators, are not primarily pursuit predators, and that their visual fields exhibit convergence towards a set of characteristics that meet the perceptual challenges of close-quarter prey detection or flush foraging in both aquatic and terrestrial environments.     

Comparative fuel metabolism in Gentoo and King Penguins: adaptation to brief versus prolonged fasting

To compare fuel utilization in large birds adapted to brief or prolonged fasting, protein and lipid utilization were quantified in the Gentoo Penguin (Pygoscelis papua) and the King Penguin (Aptenodytes patagonica). The inshore feeder Gentoo Penguin fasts for only a few days in its colony, while King Penguin chicks starve for several months in the subantarctic winter and male King Penguins starve for 5–6 weeks at the beginning of their breeding cycle. After an initial decrease in both daily body mass loss and nitrogen excretion during the first days of food deprivation, these two parameters thereafter stabilized at low values. At that time, protein utilization accounted for 15% of total energy expenditure in Gentoo Penguins and only 6% in King Penguin chicks during winter, the remainder (85% and 94%, respectively) being provided by fat oxidation. Similar percentages in fuel metabolism as seen in chicks during winter were reached in fasting adult King Penguins and spring chicks. However, a seasonal adaptation occurs in fasting chicks because energy expenditure is lower during winter. As previously described in starved mammals, the effectiveness in protein sparing could be related to the initial adiposity of the birds: the larger the fat stores (about 9% and 30% in Gentoo Penguins and winter chicks of King Penguins, respectively), the longer the fast duration and the better the level of protein conservation.     

Visual fields in hornbills: precision-grasping and sunshades

Retinal visual fields were determined in Southern Ground Hornbills Bucorvus leadbeateri and Southern Yellow-billed Hornbills Tockus leucomelas (Coraciiformes, Bucerotidae) using an ophthalmoscopic reflex technique. In both species the binocular field is relatively long and narrow with a maximum width of 30° occurring 40° above the bill. The bill tip projects into the lower half of the binocular field. This frontal visual field topography exhibits a number of key features that are also found in other terrestrial birds. This supports the hypothesis that avian visual fields are of three principal types that are correlated with the degree to which vision is employed when taking food items, rather than with phylogeny. However, unlike other species studied to date, in both hornbill species the bill intrudes into the binocular field. This intrusion of the bill restricts the width of the binocular field but allows the birds to view their own bill tips. It is suggested that this is associated with the precision-grasping feeding technique of hornbills. This involves forceps-like grasping and manipulation of items in the tips of the large decurved bill. The two hornbill species differ in the extent of the blind area perpendicularly above the head. Interspecific comparison shows that eye size and the width of the blind area above the head are significantly correlated. The limit of the upper visual field in hornbills is viewed through the long lash-like feathers of the upper lids and these appear to be used as a sunshade mechanism. In Ground Hornbills eye movements are non-conjugate and have sufficient amplitude (30–40°) to abolish the frontal binocular field and to produce markedly asymmetric visual field configurations.     

The visual fields of Common Guillemots Uria aalge and Atlantic Puffins Fratercula arctica: foraging,vigilance and collision vulnerability

Significant differences in avian visual fields are found between closely related species that differ in their foraging technique. We report marked differences in the visual fields of two auk species. In air, Common Guillemots Uria aalge have relatively narrow binocular fields typical of those found in non‐passerine predatory birds. Atlantic Puffins Fratercula arctica have much broader binocular fields similar to those that have hitherto been recorded in passerines and in a penguin. In water, visual fields narrow considerably and binocularity in the direction of the bill is probably abolished in both auk species. Although perceptual challenges associated with foraging are similar in both species during the breeding season, when they are piscivorous, Puffins (but not Guillemots) face more exacting perceptual challenges when foraging at other times, when they take a high proportion of small invertebrate prey. Capturing this prey probably requires more accurate, visually guided bill placement and we argue that this is met by the Puffin's broader binocular field, which is retained upon immersion; its upward orientation may enable prey to be seen in silhouette. These visual field configurations have potentially important consequences that render these birds vulnerable to collision with human artefacts underwater, but not in air. They also have consequences for vigilance behaviour.     

Eye structure and amphibious foraging in albatrosses

Anterior eye structure and retinal visual fields were determined in grey-headed and black-browed albatrosses, Diomedea melanophris and D. chrysostoma (Procellariiformes, Diomedeidae), using keratometry and an ophthalmoscopic reflex technique. Results for the two species were very similar and indicate that the eyes are of an amphibious optical design suggesting that albatross vision is well suited to the visual pursuit of active prey both on and below the ocean surface. The corneas are relatively flat (radius ca. 14.5 mm) and hence of low absolute refractive power (ca. 23 dioptres). In air the binocular fields are relatively long (vertical extent ca. 70 degrees) and narrow (maximum width in the plane of the optic axes 26–32 degrees), a topography found in a range of bird species that employ visual guidance of bill position when foraging. The cyclopean fields measure approximately 270 degrees in the horizontal plane, but there is a 60 degrees blind sector above the head owing to the positioning of the eyes below the protruding supraorbital ridges. Upon immersion the monocular fields decrease in width such that the binocular fields are abolished. Anterior eye structure, and visual field topography in both air and water, show marked similarity with those of the Humboldt penguin.     

Visual fields in woodcocks Scolopax rusticola (Scolopacidae; Charadriiformes)

Woodcocks, Scolopax rusticola, are long-billed terrestrial wading birds (Scolopacidae; Charadriiformes) which forage primarily by probing in soft substrates for invertebrates. Visual field topography in restrained alert birds was investigated using an ophthalmoscopic reflex technique.

1. Eye movements of significant amplitude are absent.
2. The retinal binocular field is long and narrow. It extends through 190° in the median sagittal plane. When the head adopts a normal posture (bill at an angle of 40° below the horizontal) the binocular field stretches from 25° above the bill to 5° above the horizontal behind the head. Thus, woodcocks have comprehensive visual coverage of the hemisphere above them but the bill falls outside the visual field. Maximum binocular field width equals 12° and occurs perpendicular to the line of the bill. To the rear of the head binocular field width is less than 5° except in an area 40° above the horizontal where it increases to 7°.
3. Monocular retinal fields in the horizontal plane are 182° wide. There is no blind sector at the margin of the optical fields.
4. The general structure of woodcock skulls facilitates panoramic vision in a horizontal plane.
5. Interspecific comparisons are consistent with the hypothesis that visual field topography among birds is closely associated with the role of vision in foraging. Comprehensive visual coverage of the celestial hemisphere probably occurs only in species, such as woodcocks, which rely primarily upon senses other than vision to guide foraging.

     

Sex determination of African Penguins Spheniscus demersus using bill measurements: method comparisons and implications for use

African Penguins Spheniscus demersus are sexually dimorphic; on average, males are larger than females but measurements overlap making sex determination difficult through observations alone. We developed a discriminant function, using bill length and depth from a sample of birds sexed from gonad visualisation during post-mortem, which correctly classified 93% of the individuals. Cross-validation correctly assigned 90% of DNA-sexed birds and 91% of birds sexed by partner measurement comparisons. The use of discriminant function score cutpoints, while leaving 16% and 29% of birds unclassified, improved accuracy of birds sexed by DNA to 97% and of those sexed by partner comparison to 99%. Bill depth was found to be a discriminating variable. However, two techniques for measuring bill depth are currently in use for African Penguins. While these measurements are correlated (r = 0.85), they differ on average by 1.4?mm hindering accuracy of sex determination when using a discriminant function developed from the other bill depth measurement. Exploration of adult bill morphology of birds sexed from DNA at different colonies suggests the discriminant functions can be applied throughout the African Penguins’ South African range.     

Vision and the foraging technique of skimmers (Rynchopidae)

In birds, the position and extent of the region of binocular vision appears to be determined primarily by feeding ecology. Of prime importance is the degree to which vision is used for the precise control of bill position when foraging. Skimmers (Rynchops, Rynchopidae, Charadriiformes) exhibit a unique foraging behaviour and associated structural adaptations. When foraging they fly low and straight over water with the mouth open and the mandible partially submerged. Items that are hit by the lower mandible are grasped by a rapid reflex bill closure. It is believed that this unique ‘skimming’ foraging technique is guided by tactile rather than visual cues. It is predicted therefore that the visual fields of skimmers will have similar topography to those of other tactile feeding birds. We determined retinal visual fields in Black Skimmers Rynchops niger using an ophthalmoscopic reflex technique. Contrary to expectation the visual fields of Black Skimmers are not like those of other tactile feeders. They show high similarity with those of birds that feed by precision‐pecking. The projection of the bill tip when the mouth is closed and when open (as in skimming) falls within the frontal binocular field and there is an extensive blind area above and behind the head. We argue that this visual field topography functions to achieve accurate bill positioning with respect to the water surface when skimming and, because foraging skimmers cannot determine the identity of what they are seizing as they skim, to permit the visual identification of prey items held between the mandibles after they have been taken from the water surface. When skimming, only a small portion of the binocular field, approximately 5° wide and extending 5° above the horizontal, looks in the direction of travel. The small size of this forward‐facing region of binocularity in skimmers suggests that control of locomotion in birds does not necessarily require extensive binocularity in the direction of travel.     

Finding your mate in a seabird colony: contrasting strategies of the Guillemot Uria aalge and King Penguin Aptenodytes patagonicus

Capsule King Penguins recognize their mates by voice, but Guillemots do not need acoustic cues even though their calls show individual variation.

Aims To determine whether the structure of Guillemot calls could allow individual recognition, as with King Penguin, and whether acoustic cues are used to locate mates among a dense mass of conspecifics at a colony.

Methods Observations were made on breeding Guillemots and King Penguins. Calls made by birds returning to their mates were recorded, the signals digitized and the calls analysed. Calls were later played back to the mates of the birds concerned and the effects noted on both them and their neighbours.

Results Both Guillemots and King Penguins emitted calls on return to the breeding site which contained individual signatures and were therefore potentially usable for mate recognition. In King Penguins, auditory recognition was essential for finding a mate, whereas in Guillemots most of the arriving birds located their mate in a dense crowd of conspecifics without the help of acoustic signals. Guillemots could differentiate neighbours from strangers without auditory cues.

Conclusion Calls are essential for the successful identification of mates by King Penguins but not by Guillemots.     

Visual fields in the Black-crowned Night Heron Nycticorax nycticorax: nocturnality does not result in owl-like features

Compared with diurnally active species, the eyes of nocturnally active herons (Ardeidae) are relatively larger and more widely separated. These features are found in comparisons between the nocturnally foraging Black-crowned Night Heron Nycticorax nycticorax and the diurnally active Cattle Egret Bubulcus ibis. Casual viewing of the Black-crowned Night Heron gives the impression of a somewhat owl-like appearance, with an apparently wide degree of binocular overlap. Visual fields and eye movements were determined in two alert, restrained Black-crowned Night Herons with the use of an ophthalmoscopic technique. A wide degree of binocular overlap was not confirmed, and the Black-crowned Night Heron's visual field closely resembles those of diurnally foraging herons (Western Reef Heron Egretta gularis schistacea , Squacco Heron Ardeola ralloides , Cattle Egret). The Black-crowned Night Heron's binocular field is vertically long and narrow (maximum width 22̀), with the bill placed approximately at the centre. Monocular and cyclopean field widths in the horizontal plane equal 171̀ and 320̀, respectively. Retinal binocular overlap can be abolished by eye movements. There is a blind sector (10–13̀ wide) at the margin of each eye's optical field, and this results in the functional retinal binocular field being much narrower than the optical binocular fields. It is these blind sectors which give rise to the appearance of a much wider binocular field. The visual field characteristics of this heron species may be best understood in relation to the foraging technique of capturing agile, evasive prey directly in the bill. The comparatively large size of the Black-crowned Night Herons' eyes may be associated with activity over a wide range of natural light levels but does not give rise to binocular fields larger than those of diurnal heron species.     

Examining the microclimate hypothesis in Amazonian birds: indirect tests of the ‘visual constraints’ mechanism

Proposed mechanisms for the decline of terrestrial and understory insectivorous birds in the tropics include a related subset that together has been termed the ‘microclimate hypothesis’. One prediction from this hypothesis is that sensitivity to bright light environments discourages birds of the dimly lit rainforest interior from using edges, gaps, or disturbed forest. Using a hierarchical Bayesian framework and capture data across time and space, we tested this by first determining vulnerability based on differences in within‐species capture rates between disturbed and undisturbed forest for 64 bird species at the Biological Dynamics of Forest Fragments Project in central Amazonian Brazil. We found that 35 species (55%) were vulnerable to anthropogenic habitat degradation, whereas only four (6%) were more commonly captured in degraded forest. To infer visual sensitivity, we then examined two different characters: eye size (maximum pupil diameter) relative to body mass and the initiation time of dawn song, which presumably reflects a species’ visual capacity under low light intensities. We predicted that species with large relative eye sizes and birds with earlier dawn songs would exhibit increased vulnerability in degraded habitats with bright light. Contrary to our predictions, however, vulnerability was positively correlated with the mean start time of dawn song. This indicates that species that wait to initiate dawn song are also more vulnerable to habitat degradation. After correcting for body size, there was no effect of eye size on vulnerability. Together, our results do not provide quantitative support for the light sensitivity mechanism of the microclimate hypothesis. More sensitive experimental tests, such as behavioral assays with controlled light environments, especially in a comparative framework, are needed to rigorously evaluate the role of light sensitivity as an aspect of the microclimate hypothesis among Neotropical birds.     

Eye morphology in cathemeral lemurids and other mammals

The visual systems of cathemeral mammals are subject to selection pressures that are not encountered by strictly diurnal or nocturnal species. In particular, the cathemeral eye and retina must be able to function effectively across a broad range of ambient light intensities. This paper provides a review of the current state of knowledge regarding the visual anatomy of cathemeral primates, and presents an analysis of the influence of cathemerality on eye morphology in the genus Eulemur. Due to the mutual antagonism between most adaptations for increased visual acuity and sensitivity, cathemeral lemurs are expected to resemble other cathemeral mammals in having eye morphologies that are intermediate between those of diurnal and nocturnal close relatives. However, if lemurs only recently adopted cathemeral activity patterns, then cathemeral lemurids would be expected to demonstrate eye morphologies more comparable to those of nocturnal strepsirrhines. Both predictions were tested through a comparative study of relative cornea size in mammals. Intact eyes were collected from 147 specimens of 55 primate species, and relative corneal dimensions were compared to measurements taken from a large sample of non-primate mammals. These data reveal that the five extant species of the cathemeral genus Eulemur have relative cornea sizes intermediate between those of diurnal and nocturnal strepsirrhines. Moreover, all Eulemur species have relative cornea sizes that are comparable to those of cathemeral non-primate mammals and significantly smaller than those of nocturnal mammals. These results suggest that Eulemur species resemble other cathemeral mammals in having eyes that are adapted to function under variable environmental light levels. These results also suggest that cathemerality is a relatively ancient adaptation in Eulemur that was present in the last common ancestor of the genus (ca. 8-12 MYA).     

Differences in foraging ecology determine variation in visual fields in ibises and spoonbills (Threskiornithidae)

Variations in visual field topography among birds have been interpreted as adaptations to the specific perceptual challenges posed by the species’ foraging ecology. To test this hypothesis we determined visual field topography in four bird species which have different foraging ecologies but are from the same family: Puna Ibis Plegadis ridgwayi (probes for prey in the soft substrates of marsh habitats), Northern Bald Ibis Geronticus eremita (surface pecks for prey in dry terrestrial habitats), African Spoonbill Platalea alba and Eurasian Spoonbill Platalea leucorodia (bill‐sweeps for prey in shallow turbid waters). All four species employ tactile cues provided by bill‐tip organs for prey detection. We predicted that the visual fields of these species would show general features similar to those found in other birds whose foraging is guided by tactile cues from the bill (i.e. bill falling outside the frontal binocular field and comprehensive visual coverage of the celestial hemisphere). However, the visual fields of all four species showed general features characteristic of birds that take food directly in the bill under visual guidance (i.e. a narrow and vertically long binocular field in which the projection of the bill tip is approximately central and with a blind area above and behind the head). Visual fields of the two spoonbills were very similar but differed from those of the ibises, which also differed between themselves. In the spoonbills, there was a blind area below the bill produced by the enlarged spatulate bill tip. We discuss how these differences in visual fields are related to the perceptual challenges of these birds’ different foraging ecologies, including the detection, identification and ingestion of prey. In particular we suggest that all species need to see binocularly around the bill and between the opened mandibles for the identification of caught prey items and its transport to the back of the mouth. Our findings support the hypothesis that sensory challenges associated with differences in foraging ecology, rather than shared ancestry or the control of locomotion, are the main determinants of variation in visual field topography in birds.     

The responses of the pupil of Gekko gekko to external light stimulus

1. The responses of the pupil of a nocturnal gecko (Gekko gekko) to external light stimulus were studied. 2. The responses of the pupil are determined by light entering the pupil and not by light acting directly on the iris. 3. The responses of the pupil are very uniform in sensitivity including spectral sensitivity for light coming in different directions to the eye. 4. The possible change in area of the pupil is more than 300-fold and probably represents an effort to shield the pure rod retina from saturating light intensities. 5. The pupil continues to contract sharply for changes in external light intensity which give retinal illuminations corresponding to 10⁶ quanta/sec. striking a retinal rod. 6. There is a large degree of spatial summation of the response; circular external light fields subtending 5 and 140° giving the same illumination at the pupil give approximately the same pupil response. 7. The spectral sensitivity curve agrees with the absorption curve of an extracted pigment from a closely related gecko described by Crescitelli in the followig paper. It is similar to the human scotopic curve but its maximum is displaced about 20 to 30 mµ towards the red end of the spectrum. The fall in sensitivity towards the red end of the spectrum is described by the equation See PDF for Equation     

Pupil dilation and visual field in the piked dogfish, <Emphasis Type="Italic">Squalus acanthias</Emphasis>

The relatively large eye and pupil of the Piked Dogfish, Squalus acanthias, is visually arresting. However, knowledge of its basic visual characteristics lags far behind other areas in this generally well studied species. This study quantifies pupil dilation in a species that is naturally exposed to a broad range of light intensities and finds that the pupil area in the dark adapted state is 35.3% of the total eye area, an increase of 12.4% from the light adapted state. The anterior and posterior extents of the horizontal visual field are assessed and compared with both morphological and electrophysiological techniques and the results are integrated with the measured head yaw to derive the anterior convergence distance and blind area. The position of the eyes and the triangular, pointed snout of S. acanthias provides excellent anterior vision, which likely facilitates foraging upon its mobile prey.     

The subtlety of simple eyes: the tuning of visual fields to perceptual challenges in birds

Birds show interspecific variation both in the size of the fields of individual eyes and in the ways that these fields are brought together to produce the total visual field. Variation is found in the dimensions of all main parameters: binocular region, cyclopean field and blind areas. There is a phylogenetic signal with respect to maximum width of the binocular field in that passerine species have significantly broader field widths than non-passerines; broadest fields are found among crows (Corvidae). Among non-passerines, visual fields show considerable variation within families and even within some genera. It is argued that (i) the main drivers of differences in visual fields are associated with perceptual challenges that arise through different modes of foraging, and (ii) the primary function of binocularity in birds lies in the control of bill position rather than in the control of locomotion. The informational function of binocular vision does not lie in binocularity per se (two eyes receiving slightly different information simultaneously about the same objects from which higher-order depth information is extracted), but in the contralateral projection of the visual field of each eye. Contralateral projection ensures that each eye receives information from a symmetrically expanding optic flow-field from which direction of travel and time to contact targets can be extracted, particularly with respect to the control of bill position.     

The circadian response of intrinsically photosensitive retinal ganglion cells

Intrinsically photosensitive retinal ganglion cells (ipRGC) signal environmental light level to the central circadian clock and contribute to the pupil light reflex. It is unknown if ipRGC activity is subject to extrinsic (central) or intrinsic (retinal) network-mediated circadian modulation during light entrainment and phase shifting. Eleven younger persons (18-30 years) with no ophthalmological, medical or sleep disorders participated. The activity of the inner (ipRGC) and outer retina (cone photoreceptors) was assessed hourly using the pupil light reflex during a 24 h period of constant environmental illumination (10 lux). Exogenous circadian cues of activity, sleep, posture, caffeine, ambient temperature, caloric intake and ambient illumination were controlled. Dim-light melatonin onset (DLMO) was determined from salivary melatonin assay at hourly intervals, and participant melatonin onset values were set to 14 h to adjust clock time to circadian time. Here we demonstrate in humans that the ipRGC controlled post-illumination pupil response has a circadian rhythm independent of external light cues. This circadian variation precedes melatonin onset and the minimum ipRGC driven pupil response occurs post melatonin onset. Outer retinal photoreceptor contributions to the inner retinal ipRGC driven post-illumination pupil response also show circadian variation whereas direct outer retinal cone inputs to the pupil light reflex do not, indicating that intrinsically photosensitive (melanopsin) retinal ganglion cells mediate this circadian variation.