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.

     

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.     

Eye structure and foraging in King Penguins Aptenodytes patagonicus

Anterior eye structure and retinal visual fields were determined in King Penguins Aptenodytes patagonicus using keratometry and an ophthalmoscopic reflex technique. The cornea is relatively flat (radius 32.9 mm) and hence of low refractive power (10.2 dioptres in air) and this may be correlated with the amphibious nature of penguin vision. The large size of the eye and of the fully dilated pupil may be correlated with activity at low light levels. In air, the binocular field is long (vertical extent 180̀) and narrow (maximum width 29̀), with the bill placed approximately centrally—a topography found in a range of bird species which employ visual guidance of bill position when foraging. Upon immersion in water, the optical power of the cornea is abolished, with the effect that the monocular fields decrease and binocularity is lost. King Penguins have a pupil type which has not hitherto been recorded in birds. In daylight it contracts to a square-shaped pinhole but dilates to a large circular aperture in darkness. This change alters retinal illumination by 300-fold (2.5 log₁₀ units). When diving, this permits the retina to be pre-adapted to the low ambient light levels that the birds encounter upon reaching mesopelagic depths. These penguins also forage at depths where ambient light levels, even during the day, can fall below the equivalent of terrestrial starlight. Under these conditions, the birds must rely upon the detection of light from the photophores of their prey. In this they are aided by their absolutely large pupil size and broad cyclopean visual field.     

Visual fields in the Stone-curlew Burhinus oedicnemus

The Stone-curlew Burhinus oedicnemus is a short-billed terrestrial wading bird (Burhinidae; Charadriiformes) which forages primarily for surface living invertebrates in open, sparsely vegetated habitats during twilight and nighttime. Visual field topography in restrained alert birds was investigated using an ophthalmoscopic reflex technique. The visual fields have the following features: (1) Eye movements of significant amplitude appear to be absent. (2) The retinal binocular field is relatively small, with the bill placed near its centre. It extends vertically through 80° in the median sagittal plane with a maximum width of 18° occurring 5° above the bill. (3) With the head in a typical posture (eye-bill-tip angle approximately 15° below the horizontal), the binocular field stretches from 60° below to 20° above the horizontal. (4) The blind area behind the head is relatively narrow (15° at the horizontal), giving the bird near panoramic vision in the horizontal plane, but the widest blind area (32°) occurs directly above the head. (5) Monocular retinal fields in the horizontal plane are 182° wide and are asymmetric about the optic axis. (6) There is a blind sector of 7–12° at the margin of the optical fields, indicating that binocular field widths are not maximized. Interspecific comparisons of these visual field features suggest that the foraging of Stone-curlews is guided primarily by visual cues.     

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.     

Interspecific differences in the visual system and scanning behavior of three forest passerines that form heterospecific flocks

Little is known as to how visual systems and visual behaviors vary within guilds in which species share the same micro-habitat types but use different foraging tactics. We studied different dimensions of the visual system and scanning behavior of Carolina chickadees, tufted titmice, and white-breasted nuthatches, which are tree foragers that form heterospecific flocks during the winter. All species had centro-temporally located foveae that project into the frontal part of the lateral visual field. Visual acuity was the highest in nuthatches, intermediate in titmice, and the lowest in chickadees. Chickadees and titmice had relatively wide binocular fields with a high degree of eye movement right above their short bills probably to converge their eyes while searching for food. Nuthatches had narrower binocular fields with a high degree of eye movement below their bills probably to orient the fovea toward the trunk while searching for food. Chickadees and titmice had higher scanning (e.g., head movement) rates than nuthatches probably due to their wider blind areas that limit visual coverage. The visual systems of these three species seem tuned to the visual challenges posed by the different foraging and scanning strategies that facilitate the partitioning of resources within this guild.     

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.     

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.     

Relative Wulst volume is correlated with orbit orientation and binocular visual field in birds

In mammals, species with more frontally oriented orbits have broader binocular visual fields and relatively larger visual regions in the brain. Here, we test whether a similar pattern of correlated evolution is present in birds. Using both conventional statistics and modern comparative methods, we tested whether the relative size of the Wulst and optic tectum (TeO) were significantly correlated with orbit orientation, binocular visual field width and eye size in birds using a large, multi-species data set. In addition, we tested whether relative Wulst and TeO volumes were correlated with axial length of the eye. The relative size of the Wulst was significantly correlated with orbit orientation and the width of the binocular field such that species with more frontal orbits and broader binocular fields have relatively large Wulst volumes. Relative TeO volume, however, was not significant correlated with either variable. In addition, both relative Wulst and TeO volume were weakly correlated with relative axial length of the eye, but these were not corroborated by independent contrasts. Overall, our results indicate that relative Wulst volume reflects orbit orientation and possibly binocular visual field, but not eye size.     

Development of the compound eyes of dragonflies (Odonata). III. Adult compound eyes

The distribution of ommatidial diameters and interommatidial angles, as determined by measuring the angles between the optic axes of adjacent ommatidia, are mapped across the surface of the compound eyes of a variety of species selected for different adult behaviors, developmental histories, and taxonomic positions. The size of the visual fields, prey capture foveas, foveas composed of large dorsal ommatidia, and other specializations in the numbers of ommatidia that view various directions in the visual field are discussed in relation to adult behavior. Advanced species have less resemblance between their larval and adult eyes than primitive species. In contrast to their larvae, adults increase the monocular resolution of each eye at the expense of binocular vision. Most species have foveas which view in approximately the anterior direction, instead of in a region of binocular overlap, and many species have foveal bands which view along the horizon. Some advanced perching species, which approach their prey and other odonates from below, have an additional vertical foveal band that views along a vertical plane from the anterior direction to a more dorsal direction. The most unusual foveal band is seen in active flying species. The large dorsal ommatidia of the migratory Anax junius, which cover approximately one third of the eye surface, view a narrow region of the visual field that extends along a plane from the most lateral direction of one eye to a dorsal direction, and continues without interruption to the most lateral direction of the other eye.     

The visual fields of two ground‐foraging birds,House Finches and House Sparrows,allow for simultaneous foraging and anti‐predator vigilance

In birds, differences in the extent and position of the binocular visual field reflect adaptations to varying foraging strategies, and the extent of the lateral portion of the field may reflect anti‐predator strategies. The goal of this study was to describe and compare the visual fields of two ground‐foraging passerines, House Finch Carpodacus mexicanus and House Sparrow Passer domesticus. We found that both species have a binocular field type that is associated with the accurate control of bill position when pecking. Both species have eye movements of relatively large amplitude, which can produce substantial variations in the configuration of the binocular fields. We propose that in these ground foragers, their relatively wide binocular fields could function to increase foraging efficiency by locating multiple rather than single food items prior to pecking events. The lateral fields of both species are wide enough to facilitate the detection of predators or conspecifics while head‐down foraging. This suggests that foraging and scanning are not mutually exclusive activities in these species, as previously assumed. Furthermore, we found some slight, but significant, differences between species: House Sparrow binocular fields are both wider and vertically taller, and the blind area is wider than in House Finches. These differences may be related to variations in the degree of eye movements and position of the orbits in the skull.     

Is the landing response of the housefly (Musca) driven by motion of a flow field?

The landing response of tethered flying housefliesMusca domestica elicited by motion of periodic gratings is analysed. The field of view of the compound eyes of a fly can be subdivided into a region of binocular overlap and a monocular region. In the monocular region the landing response is elicited by motion from front to back and suppressed by motion from back to front. The sensitivity to front to back motion in monocular flies (one eye covered with black paint) has a maximum at an angle 60°–80° laterally from the direction of flight in the equatorial plane. The maximum of the landing response to front to back motion as a function of the contrast frequencyw/ is observed at around 8 Hz. In the region of binocular overlap of monocular flies the landing response can be elicited by back to front motion around the equatorial plane if a laterally positioned pattern is simulataneously moved from front to back. 40° above the equatorial plane in the binocular region the landing response in binocular flies is elicited by upward motion, 40° below the equatorial plane in the binocular region it is elicited by downward motion. The results are interpreted as an adaptation of the visual system of the fly to the perception of a flow field having its pole in the direction of flight.     

Visual coverage and scanning behavior in two corvid species: American crow and Western scrub jay

Inter-specific differences in the configuration of avian visual fields and degree of eye/head movements have been associated with foraging and anti-predator behaviors. Our goal was to study visual fields, eye movements, and head movements in two species of corvids: American crow (Corvus brachyrhynchos) and Western scrub jay (Aphelocoma californica). American crows had wider binocular overlap, longer vertical binocular fields, narrower blind areas, and higher amplitude of eye movement than Western scrub jays. American crows can converge their eyes and see their own bill tip, which may facilitate using different foraging techniques (e.g., pecking, probing) and manufacturing and handing rudimentary tools. Western scrub jays had a higher head movement rate than American crows while on the ground, and the opposite between-species difference was found when individuals were perching. Faster head movements may enhance the ability to scan the environment, which may be related to a higher perceived risk of predation of Western scrub jays when on the ground, and American crows when perching. The visual field configuration of these species appears influenced mostly by foraging techniques while their scaning behavior, by predation risk.     

The visual field and visually guided behavior in the zebra finch (Taeniopygia guttata)

Summary Measurements were made of the physical properties of the visual system of the zebra finch, a bird with laterally placed eyes. The use of the visual system in pecking and courtship behavior was examined. It was demonstrated that the optical axis and the fovea of the eye point in a direction about 62° from the sagittal axis of the head. The visual field of each eye covers about 170° in the horizontal plane. In the frontal region there is an overlap of about 30°–40° where the birds can see binocularly; caudally there is a gap in the visual field of 60°. The point of best binocular viewing is in the sagittal plane at 16.5° below the beak.Concerning movement detection, the upper threshold is 540°/s for the binocular (frontal) part of the visual field and about 1100°/s for the monocular (lateral) part. Most fixations before pecking occur monocularly. A preference for one eye during pecking was not detected. During the courtship song, a male bird directs its head towards the female. The results are discussed in comparison with findings in pigeons and chickens.     

Visual fields and foraging ecology of Blacksmith Lapwings Vanellus armatus

The visual fields of Blacksmith Lapwings Vanellus armatus show the characteristics of visual guided foragers that use precision pecking for prey capture – a binocular field of narrow width and limited vertical extent, with the projection of the bill close to its centre and a large blind area above and behind the head. The topography of the total field, particularly the binocular field, is similar to that of European Golden Plovers Pluvialis apricaria. We suggest that the ‘foot‐trembling’ behaviour associated with foraging in Plovers is not under visual guidance but forces the escape of hidden prey, which is detected when the prey item moves into the binocular field to enable its capture in the bill. Foot‐trembling thus functions to extend the effective foraging area of a bird beyond the limits of its visual field.     

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.     

Reverse lateralization of visual discriminative abilities in the European starling

Previous experiments on visual feature discrimination abilities have consistently shown a right-eye system lateralization in pigeons, Columba livia, and young domestic chickens, Gallus gallus domesticus, both nonpasserine species. Recently, however, it has been shown that photoreceptor distribution in the left and right retinas are asymmetrical in the European starling, Sturnus vulgaris, a passerine species. Single cone receptors are significantly more abundant in the left retina, which suggests that starlings should perform visual discrimination tasks more proficiently with the left eye, in contrast to previous findings with nonpasserines. We tested this hypothesis using the technique of monocular occlusion. In the first experiment, starlings were tested on a simultaneous visual discrimination task in three conditions: binocular (both eyes), left monocular (left eye only) and right monocular (right eye only). Subjects in the binocular and left-monocular conditions achieved significantly higher performance scores on the discrimination task than birds in the right-monocular condition. A second experiment found similar results, with birds in the left-monocular condition learning the discrimination task more than twice as quickly as those in the right-monocular condition. Subsequent tests with the alternative eye for both groups indicated no interocular transfer. These findings suggest that visual discriminative abilities in starlings are asymmetrical, and that they are lateralized in the opposite eye system than has been reported for all other species tested to date.     

Changes in visual fields associated with web reduction in the spider family uloboridae

When visual fields of the primitive orb-weaver, Waitkera waitkerensis, are reconstructed using measurements taken from intact lenses and cross and longitudinal sections of the prosoma, they show that this species has complete visual surveillance, but that none of the visual fields of its eight eyes overlap. The more advanced orb-weaver, Uloborus glomosus, also has eight eyes, but each eye has a greater visual angle, giving this species a complex pattern of overlapping visual fields. Uloborids that spin reduced webs are characterized by reduction or loss of the four anterior eyes and other carapace modifications necessary for them to effectively monitor and manipulate their reduced webs. The eyes of these uloborids have greater visual angles than those of orb-weavers, resulting primarily from perimetric expansion of their retinal hemispheres. Additionally, the axes of their visual fields are more ventrally directed due to greater dorsal than ventral retinal expansion and to ventral redirection of the entire eye. Consequently, even though the anterior lateral eyes of the triangle-weaver Hyptiotes cavatus lack retinae, the species' six functional eyes permit complete visual surveillance and exhibit visual overlap. The single-line-weaver, Miagrammopes animotus, has lost its four anterior eyes, and with them much of the anterior vision and all of the visual overlap found in the other species. However, changes similar to those of H. cavatus permit this species to retain most if its dorsal and ventral visual surveillance. Thus, ocular changes act in consort to maintain relatively complete visual surveillance in the face of eye loss and other major carapace modifications necessary for the operation of reduced webs.     

Visual fields, eye movements, and scanning behavior of a sit-and-wait predator, the black phoebe (Sayornis nigricans)

Foraging mode influences the dominant sensory modality used by a forager and likely the strategies of information gathering used in foraging and anti-predator contexts. We assessed three components of visual information gathering in a sit-and-wait avian predator, the black phoebe (Sayornis nigricans): configuration of the visual field, degree of eye movement, and scanning behavior through head-movement rates. We found that black phoebes have larger lateral visual fields than similarly sized ground-foraging passerines, as well as relatively narrower binocular and blind areas. Black phoebes moved their eyes, but eye movement amplitude was relatively smaller than in other passerines. Black phoebes may compensate for eye movement constraints with head movements. The rate of head movements increased before attacking prey in comparison to non-foraging contexts and before movements between perches. These findings suggest that black phoebes use their lateral visual fields, likely subtended by areas of high acuity in the retina, to track prey items in a three-dimensional space through active head movements. These head movements may increase depth perception, motion detection and tracking. Studying information gathering through head movement changes, rather than body posture changes (head-up, head-down) as generally presented in the literature, may allow us to better understand the mechanisms of information gathering from a comparative perspective.     

Hawk Eyes I: Diurnal Raptors Differ in Visual Fields and Degree of Eye Movement

Background

Different strategies to search and detect prey may place specific demands on sensory modalities. We studied visual field configuration, degree of eye movement, and orbit orientation in three diurnal raptors belonging to the Accipitridae and Falconidae families.

Methodology/Principal Findings

We used an ophthalmoscopic reflex technique and an integrated 3D digitizer system. We found inter-specific variation in visual field configuration and degree of eye movement, but not in orbit orientation. Red-tailed Hawks have relatively small binocular areas (∼33°) and wide blind areas (∼82°), but intermediate degree of eye movement (∼5°), which underscores the importance of lateral vision rather than binocular vision to scan for distant prey in open areas. Cooper''s Hawks'' have relatively wide binocular fields (∼36°), small blind areas (∼60°), and high degree of eye movement (∼8°), which may increase visual coverage and enhance prey detection in closed habitats. Additionally, we found that Cooper''s Hawks can visually inspect the items held in the tip of the bill, which may facilitate food handling. American Kestrels have intermediate-sized binocular and lateral areas that may be used in prey detection at different distances through stereopsis and motion parallax; whereas the low degree eye movement (∼1°) may help stabilize the image when hovering above prey before an attack.

Conclusions

We conclude that: (a) there are between-species differences in visual field configuration in these diurnal raptors; (b) these differences are consistent with prey searching strategies and degree of visual obstruction in the environment (e.g., open and closed habitats); (c) variations in the degree of eye movement between species appear associated with foraging strategies; and (d) the size of the binocular and blind areas in hawks can vary substantially due to eye movements. Inter-specific variation in visual fields and eye movements can influence behavioral strategies to visually search for and track prey while perching.