Bird vision

1 Extraocular anatomy
The eye of a bird most closely resembles that of the rep-
tiles. Unlike the mammalian eye, it is not spherical, and
the atter shape enables more of its visual eld to be in fo-
cus. A circle of bony plates, the sclerotic ring, surrounds
the eye and holds it rigid, but an improvement over the
reptilian eye, also found in mammals, is that the lens is
pushed further forward, increasing the size of the image
on the retina.[2]

With forward-facing eyes, the bald eagle has a wide eld of

binocular vision.

Visual elds for a pigeon and an owl

Most birds cannot move their eyes, although there are ex-
Vision is the most important sense for birds, since good
ceptions, such as the great cormorant.[3] Birds with eyes
eyesight is essential for safe ight, and this group has a
on the sides of their heads have a wide visual eld, useful
number of adaptations which give visual acuity superior
for detecting predators, while those with eyes on the front
to that of other vertebrate groups; a pigeon has been de-
of their heads, such as owls, have binocular vision and can
scribed as two eyes with wings.[1] The avian eye resem-
estimate distances when hunting.[4] The American wood-
bles that of a reptile, with ciliary muscles that can change
cock probably has the largest visual eld of any bird, 360
the shape of the lens rapidly and to a greater extent than
in the horizontal plane, and 180 in the vertical plane.[5]
in the mammals. Birds have the largest eyes relative to
their size in the animal kingdom, and movement is con-
sequently limited within the eyes bony socket.[1] In ad-
dition to the two eyelids usually found in vertebrates, it
is protected by a third transparent movable membrane.
The eyes internal anatomy is similar to that of other ver-
tebrates, but has a structure, the pecten oculi, unique to
birds.
Some bird groups have specic modications to their vi- The nictitating membrane of a masked lapwing
sual system linked to their way of life. Birds of prey have
a very high density of receptors and other adaptations that The eyelids of a bird are not used in blinking. Instead
maximise visual acuity. The placement of their eyes gives the eye is lubricated by the nictitating membrane, a third
them good binocular vision enabling accurate judgement concealed eyelid that sweeps horizontally across the eye
of distances. Nocturnal species have tubular eyes, low like a windscreen wiper.[6] The nictitating membrane also
numbers of colour detectors, but a high density of rod covers the eye and acts as a contact lens in many aquatic
cells which function well in poor light. Terns, gulls and birds when they are under water.[7] When sleeping, the
albatrosses are amongst the seabirds which have red or lower eyelid rises to cover the eye in most birds, with the
yellow oil droplets in the colour receptors to improve dis- exception of the horned owls where the upper eyelid is
tance vision especially in hazy conditions. mobile.[8]

1
2 2 ANATOMY OF THE EYE
The eye is also cleaned by tear secretions from the
lachrymal gland and protected by an oily substance from
the Harderian glands which coats the cornea and prevents
dryness. The eye of a bird is larger compared to the size
of the animal than for any other group of animals, al-
though much of it is concealed in its skull. The ostrich
has the largest eye of any land vertebrate, with an axial
length of 50 mm (2 in), twice that of the human eye.[1]
Bird eye size is broadly related to body mass. A study
of ve orders (parrots, pigeons, petrels, raptors and owls)
showed that eye mass is proportional to body mass, but
as expected from their habits and visual ecology, raptors
and owls have relatively large eyes for their body mass.[9]
Behavioural studies show that many avian species focus
on distant objects preferentially with their lateral and
monocular eld of vision, and birds will orientate them-
selves sideways to maximise visual resolution. For a pi-
geon, resolution is twice as good with sideways monocu-
lar vision than forward binocular vision, whereas for hu-
mans the converse is true.[1]
The European robin has relatively large eyes, and starts to sing
early in the morning.
The performance of the eye in low light levels depends
on the distance between the lens and the retina, and small
birds are eectively forced to be diurnal because their
eyes are not large enough to give adequate night vision.
Although many species migrate at night, they often col-
lide with even brightly lit objects like lighthouses or oil
platforms. Birds of prey are diurnal because, although
their eyes are large, they are optimised to give maximum
spatial resolution rather than light gathering, so they also
do not function well in poor light.[10] Many birds have an
asymmetry in the eyes structure which enables them to
keep the horizon and a signicant part of the ground in
focus simultaneously. The cost of this adaptation is that
they have myopia in the lower part of their visual eld.[1]
Birds with relatively large eyes compared to their body
mass, such as common redstarts and European robins sing
earlier at dawn than birds of the same size and smaller
body mass. However, if birds have the same eye size but
dierent body masses, the larger species sings later than
the smaller. This may be because the smaller bird has to
start the day earlier because of weight loss overnight.[11]
Overnight weight loss for small birds is typically 5-10%
and may be over 15% on cold winter nights.[11] In one
study, robins put on more mass in their dusk feeding when
nights were cold.[12]
Nocturnal birds have eyes optimised for visual sensitivity,
with large corneas relative to the eyes length, whereas di-
urnal birds have longer eyes relative to the corneal diame-
ter to give greater visual acuity. Information about the ac-
tivities of extinct species can be deduced from measure-
ments of the sclerotic ring and orbit depth. For the latter
measurement to be made, the fossil must have retained its
three-dimensional shape, so activity pattern cannot be de-
termined with condence from attened specimens like
Archaeopteryx, which has a complete sclerotic ring but
no orbit depth measurement.[13]
2 Anatomy of the eye
Anatomy of the avian eye
The main structures of the bird eye are similar to those
of other vertebrates. The outer layer of the eye consists
of the transparent cornea at the front, and two layers of
sclera a tough white collagen bre layer which sur-
rounds the rest of the eye and supports and protects the
eye as a whole. The eye is divided internally by the lens
into two main segments: the anterior segment and the
posterior segment. The anterior chamber is lled with a
watery uid called the aqueous humour, and the posterior
chamber contains the vitreous humour, a clear jelly-like
substance.
The lens is a transparent convex or 'lens shaped body with
a harder outer layer and a softer inner layer. It focuses the
light on the retina. The shape of the lens can be altered by
ciliary muscles which are directly attached to lens capsule
by means of the zonular bres. In addition to these mus-
cles, some birds also have a second set, Cramptons mus-
cles, that can change the shape of the cornea, thus giving
birds a greater range of accommodation than is possible

for mammals. This accommodation can be rapid in some
diving water birds such as in the mergansers. The iris is a
coloured muscularly operated diaphragm in front of the
lens which controls the amount of light entering the eye.
At the centre of the iris is the pupil, the variable circular
area through which the light passes into the eye.[2][14]
Hummingbirds are amongst the many birds with two foveae
The retina is a relatively smooth curved multi-layered
structure containing the photosensitive rod and cone cells
with the associated neurons and blood vessels. The den-
sity of the photoreceptors is critical in determining the
maximum attainable visual acuity. Humans have about
200,000 receptors per mm2 , but the house sparrow has
400,000 and the common buzzard 1,000,000. The pho-
toreceptors are not all individually connected to the optic
nerve, and the ratio of nerve ganglia to receptors is impor-
tant in determining resolution. This is very high for birds;
the white wagtail has 100,000 ganglion cells to 120,000
photoreceptors.[2]
Rods are more sensitive to light, but give no colour infor-
mation, whereas the less sensitive cones enable colour vi-
sion. In diurnal birds, 80% of the receptors may be cones
(90% in some swifts) whereas nocturnal owls have almost
all rods. As with other vertebrates except placental mam-
mals, some of the cones may be double cones. These can
amount to 50% of all cones in some species.[15]
Towards the centre of the retina is the fovea (or the less
specialised, area centralis) which has a greater density of
receptors and is the area of greatest forward visual acuity,
i.e. sharpest, clearest detection of objects. In 54% of
birds, including birds of prey, kingshers, hummingbirds
and swallows, there is second fovea for enhanced sideways
viewing. The optic nerve is a bundle of nerve bres which
carry messages from the eye to the relevant parts of the
brain and vice versa. Like mammals, birds have a small
blind spot without photoreceptors at the optic disc, under
which the optic nerve and blood vessels join the eye.[2]
The pecten is a poorly understood body consisting of
folded tissue which projects from the retina. It is well
supplied with blood vessels and appears to keep the retina
supplied with nutrients,[1] and may also shade the retina
from dazzling light or aid in detecting moving objects.[2] The four spectrally distinct cone pigments are derived
Pecten oculi is abundantly lled with melanin granules from the protein opsin, linked to a small molecule called
3
which have been proposed to absorb stray light entering
the bird eye to reduce background glare. Slight warm-
ing of pecten oculi due to absorption of light by melanin
granules has been proposed enhance metabolic rate of
pecten that is suggested to help increase secretion of nu-
trients into vitreous, eventually to be absorbed by avas-
cular retina of birds for improved nutrition.[16] Exra-high
enzymic activity of alkaline phosphatase in pecten oculi
has been proposed to support high secretory activity of
pecten to supplement nutrition of retina.[17]
The choroid is a layer situated behind the retina which
contains many small arteries and veins. These provide
arterial blood to the retina and drain venous blood. The
choroid contains melanin, a pigment which gives the inner
eye its dark colour, helping to prevent disruptive reec-
tions.
3 Light perception
370 nm 445 nm 508 nm 565 nm
1.0
Absorbance
0.5
0.0
330 nm 400 nm 500 nm 600 nm 700 nm
Ultraviolet
The four pigments in estrildid nches' cones extend the range of
colour vision into the ultraviolet.[18][19]
There are two sorts of light receptors in a birds eye,
rods and cones. Rods, which contain the visual pigment
rhodopsin are better for night vision because they are sen-
sitive to small quantities of light. Cones detect specic
colours (or wavelengths) of light, so they are more impor-
tant to colour-orientated animals such as birds.[20] Most
birds are tetrachromatic, possessing four types of cone
cells each with a distinctive maximal absorption peak.
In some birds, the maximal absorption peak of the cone
cell responsible for the shortest wavelength extends to the
ultraviolet (UV) range, making them UV-sensitive.[21] In
addition to that, the cones at birds retina are arranged
in a characteristic form of spatial distribution, known as
hyperuniform distribution, which maximizes its light and
color absorption. This form of spatial distributions are
only observed as a result of some optimization process,
which in this case can be described in terms of birds evo-
lutionary history.[22]
4 3 LIGHT PERCEPTION

retinal, which is closely related to vitamin A. When the pigments in birds give increased discrimination.[20]
pigment absorbs light the retinal changes shape and alters Each cone of a bird or reptile contains a coloured oil
the membrane potential of the cone cell aecting neu- droplet; these no longer exist in mammals. The droplets,
rons in the ganglia layer of the retina. Each neuron in which contain high concentrations of carotenoids, are
the ganglion layer may process information from a num- placed so that light passes through them before reaching
ber of photoreceptor cells, and may in turn trigger a nerve the visual pigment. They act as lters, removing some
impulse to relay information along the optic nerve for fur- wavelengths and narrowing the absorption spectra of the
ther processing in specialised visual centres in the brain. pigments. This reduces the response overlap between pig-
The more intense a light, the more photons are absorbed
ments and increases the number of colours that a bird can
by the visual pigments; the greater the excitation of each discern.[20] Six types of cone oil droplets have been iden-
cone, and the brighter the light appears.[20]
tied; ve of these have carotenoid mixtures that absorb
at dierent wavelengths and intensities, and the sixth type
has no pigments.[25] The cone pigments with the lowest
maximal absorption peak, including those that are UV-
sensitive, possess the 'clear' or 'transparent' type of oil
droplets with little spectral tuning eect.[26]
The colours and distributions of retinal oil droplets vary
considerably among species, and is more dependent on
the ecological niche utilised (hunter, sher, herbivore)
than genetic relationships. As examples, diurnal hunters
like the barn swallow and birds of prey have few coloured
droplets, whereas the surface shing common tern has
a large number of red and yellow droplets in the dorsal
retina. The evidence suggests that oil droplets respond to
natural selection faster than the cones visual pigments.[23]
Even within the range of wavelengths that are visible to
humans, passerine birds can detect colour dierences that
humans do not register. This ner discrimination, to-
gether with the ability to see ultraviolet light, means that
many species show sexual dichromatism that is visible to
birds but not humans.[27]
Migratory songbirds use the Earths magnetic eld, stars,
the Sun, and other unknown cues to determine their mi-
gratory direction. An American study suggested that mi-
gratory Savannah sparrows used polarised light from an
area of sky near the horizon to recalibrate their mag-
netic navigation system at both sunrise and sunset. This
Diagram of a bird cone cell suggested that skylight polarisation patterns are the pri-
mary calibration reference for all migratory songbirds.[28]
By far the most abundant cone pigment in every However, it appears that birds may be responding to sec-
bird species examined is the long-wavelength form of ondary indicators of the angle of polarisation, and may
iodopsin, which absorbs at wavelengths near 570 nm. not be actually capable of directly detecting polarisation
This is roughly the spectral region occupied by the direction in the absence of these cues.[29]
red- and green-sensitive pigments in the primate retina,
and this visual pigment dominates the colour sensitivity
of birds.[23] In penguins, this pigment appears to have 3.1 Ultraviolet sensitivity
shifted its absorption peak to 543 nm, presumably an
adaptation to a blue aquatic environment.[24] There are two types of colour vision in birds: violet
The information conveyed by a single cone is limited: by sensitive (VS) and ultraviolet sensitive (UVS).[30] UVS
itself, the cell cannot tell the brain which wavelength of birds have a visual pigment in the cones of their retinas
light caused its excitation. A visual pigment may absorb that absorbs UV light, allowing them to see the ultravi-
two wavelengths equally, but even though their photons olet portion of the spectrum. The major clades of birds
are of dierent energies, the cone cannot tell them apart, that have UVS vision are Palaeognathae (ratites and tina-
because they both cause the retinal to change shape and mous), Charadriiformes (shorebirds, gulls, and alcids),
thus trigger the same impulse. For the brain to see colour, Trogoniformes (trogons), Psittaciformes (parrots), and
it must compare the responses of two or more classes of Passeriformes (perching birds, representing more than
cones containing dierent visual pigments, so the four half of all avian species).[30]
 5

ultraviolet light (insects, reptiles, and crustaceans have

UVS vision as well), some predators of UVS birds can-
not see ultraviolet light. This raises the possibility that
ultraviolet vision gives birds a channel in which they
can privately signal, thereby remaining inconspicuous to
predators.[35] However, recent evidence does not appear
to support this hypothesis.[36]

4 Perception

4.1 Contrast sensitivity

Contrast is dened as the dierence in brightness be-

tween two stimuli, divided by the sum of brightness of
both stimuli. Contrast sensitivity is the inverse of the
smallest contrast that can be detected, a contrast sensi-
tivity of 100 means that the smallest contrast that can
be detected is 1%. Birds have comparably low con-
trast sensitivity to mammals. Humans have been shown
to detect contrasts as low as 0.5-1%,[37] whereas most
birds tested have require ca 10% contrast to show a be-
havioural response.[38][39][40] A contrast sensitivity func-
tion describes an animals ability to resolve detect the con-
The common kestrel, like other raptorial birds, have a very low
trast of patterns of dierent spatial frequency (detail).
sensitivity to UV light. For stationary viewing experiments the contrast sensitiv-
ity is highest for a medium spatial frequency and lower
for higher and lower spatial frequencies.[41]
UVS vision can be useful for courtship. Many birds show
plumage patterns in ultraviolet that are invisible to the
human eye; some birds whose sexes appear similar to the 4.2 Movement
naked eye are distinguished by the presence of ultravi-
olet reective patches on their feathers. Male blue tits
have an ultraviolet reective crown patch which is dis-
played in courtship by posturing and raising of their nape
feathers.[31] Male blue grosbeaks with the brightest and
most UV-shifted blue in their plumage are larger, hold the
most extensive territories with abundant prey, and feed
their ospring more frequently than other males do.[20]
The bills appearance is important in the interactions of
the blackbird. Although the UV component seems unim-
portant in interactions between territory-holding males, A red kite ying at a bird feeding station in Scotland
where the degree of orange is the main factor, the female
responds more strongly to males with bills with good UV- Birds can resolve rapid movements better than humans,
reectiveness.[32] for whom ickering at a rate greater than 50 light pulse
UVS vision may also give birds an advantage in foraging cycles per second appears as continuous movement. Hu-
for food. The waxy surfaces of many fruits and berries mans cannot therefore distinguish individual ashes of a
reect UV light that might advertise their presence.[20] uorescent light bulb oscillating at 60 light pulse cycles
Common kestrels may be able to locate the trails of voles per second, but budgerigars and chickens have icker or
visually. These small rodents lay scent trails of urine and light pulse cycles per second thresholds of more than 100
faeces that could reect UV light, making them visible light pulse cycles per second which is safe to strictly say
to the kestrels, particularly in the spring before the scent not Hertz A Coopers hawk can pursue agile prey through
marks are covered by vegetation.[33] However, this view woodland and avoid branches and other objects at high
has been challenged by the nding of low UV sensitivity speed; to humans such a chase would appear as a blur.[5]
in raptors and weak UV reection of mammal urine.[34] Birds can also detect slow moving objects. The move-
While birds are not unique in their ability to perceive ment of the sun and the constellations across the sky is
6 5 VARIATIONS ACROSS BIRD GROUPS

imperceptible to humans, but detected by birds. The abil-

ity to detect these movements allows migrating birds to
properly orient themselves.[5]
To obtain steady images while ying or when perched on
a swaying branch, birds hold the head as steady as possible
with compensating reexes. Maintaining a steady image
is especially relevant for birds of prey.[5]

4.3 Edges and shapes

When an object is partially blocked by another, hu-
mans unconsciously tend to make up for it and com-
plete the shapes (See Amodal perception). It has how-
ever been demonstrated that pigeons do not complete oc-
cluded shapes.[42] A study based on altering the grey level
of a perch that was coloured dierently from the back-
ground showed that budgerigars do not detect edges based
on colours.[43] Hawk-eyed is a byword for visual acuity

4.4 Magnetic elds
The perception of magnetic elds by migratory birds
has been suggested to be light dependent.[44] Birds move
their head to detect the orientation of the magnetic
eld,[45] and studies on the neural pathways have sug-
gested that birds may be able to see the magnetic
elds.[46] The right eye of a migratory bird contains pho-
toreceptive proteins called cryptochromes. Light excites
these molecules to produce unpaired electrons that inter-
act with the Earths magnetic eld, thus providing direc-
tional information.[47][48]
the deep central fovea of raptors can create a telephoto
optical system,[49] increasing the size of the retinal im-
age in the fovea and thereby increasing the spatial resolu-
tion. Behavioural studies show that some large eyed rap-
tors (Wedge-tailed eagle, Old world vultures) and have ca
2 times higher spatial resolution than humans, but many
medium and small sized raptors have comparable or lower
spatial resolution.[50][51][52][53][54][55]
fovea

5 Variations across bird groups

5.1 Diurnal birds of prey
The visual ability of birds of prey is legendary, and the
keenness of their eyesight is due to a variety of factors.
Raptors have large eyes for their size, 1.4 times greater
than the average for birds of the same weight,[9] and
the eye is tube-shaped to produce a larger retinal im-
age. The resolving power of an eye depends both on
the optics, large eyes with large appertures suers less
from diraction and can have larger retinal images due
to a long focal length, and on the density of receptor
spacing. The retina has a large number of receptors
per square millimeter, which determines the degree of
visual acuity. The more receptors an animal has, the
higher its ability to distinguish individual objects at a dis-
tance, especially when, as in raptors, each receptor is
typically attached to a single ganglion.[1] Many raptors
have foveas with far more rods and cones than the hu-
man fovea (65,000/mm2 in American kestrel, 38,000 in
humans) and this provides these birds with spectacular
long distance vision. It is proposed that the shape of
pecten
Each retina of the black-chested buzzard-eagle has two
foveae[56]
The forward-facing eyes of a bird of prey give binocular
vision, which is assisted by a double fovea.[2] The raptors
adaptations for optimum visual resolution (an American
kestrel can see a 2mm insect from the top of an 18m
tree) has a disadvantage in that its vision is poor in low
light level, and it must roost at night.[1] Raptors may have
to pursue mobile prey in the lower part of their visual
eld, and therefore do not have the lower eld myopia
adaptation demonstrated by many other birds.[1] Scaveng-
ing birds like vultures do not need such sharp vision, so
a condor has only a single fovea with about 35,000 re-
ceptors mm2 . Vultures, however have high physiological
5.3 Water birds 7

activity of many important enzymes to suit their distant

clarity of vision.[57]
Like other birds investigated raptors do also have
coloured oil droplets in their cones.[52][53][58] The gener- fovea
ally brown, grey and white plumage of this group, and
the absence of colour displays in courtship suggests that
colour is relatively unimportant to these birds.[2]
In most raptors a prominent eye ridge and its feathers ex- pecten
tends above and in front of the eye. This eyebrow gives
birds of prey their distinctive stare. The ridge physically
protects the eye from wind, dust, and debris and shields
it from excessive glare. The osprey lacks this ridge, al-
though the arrangement of the feathers above its eyes
[56]
serves a similar function; it also possesses dark feathers in Each owls retina has a single fovea
front of the eye which probably serve to reduce the glare
from the water surface when the bird is hunting for its
vironment. There are few coloured oil droplets, which
staple diet of sh.[5]
would reduce the light intensity, but the retina contains a
reective layer, the tapetum lucidum. This increases the
5.2 Nocturnal birds amount of light each photosensitive cell receives, allowing
the bird to see better in low light conditions.[2] Owls nor-
mally have only one fovea, and that is poorly developed
except in diurnal hunters like the short-eared owl.[59]
Besides owls, bat hawks, frogmouths and nightjars also
display good night vision. Some bird species nest deep
in cave systems which are too dark for vision, and nd
their way to the nest with a simple form of echolocation.
The oilbird is the only nocturnal bird to echolocate,[61] but
several Aerodramus swiftlets also utilise this technique,
with one species, Atiu swiftlet, also using echolocation
outside its caves.[62][63]

5.3 Water birds

A powerful owl photographed at night showing reective tapeta

lucida

Owls have very large eyes for their size, 2.2 times greater
than the average for birds of the same weight,[9] and posi-
tioned at the front of the head. The eyes have a eld over- Terns have coloured oil droplets in the cones of the eye to improve
lap of 5070%, giving better binocular vision than for di- distance vision
urnal birds of prey (overlap 3050%).[59] The tawny owls
retina has about 56,000 light-sensitive rods per square Seabirds such as terns and gulls that feed at the surface
millimetre (36 million per square inch); although earlier or plunge for food have red oil droplets in the cones of
claims that it could see in the infrared part of the spectrum
their retinas. This improves contrast and sharpens dis-
have been dismissed.[60] tance vision, especially in hazy conditions.[2] Birds that
Adaptations to night vision include the large size of the have to look through an air/water interface have more
eye, its tubular shape, large numbers of closely packed deeply coloured carotenoid
[23]
pigments in the oil droplets
retinal rods, and an absence of cones, since cone cells than other species.
are not sensitive enough for a low-photon nighttime en- This helps them to locate shoals of sh, although it is un-
8 7 NOTES

certain whether they are sighting the phytoplankton on by aerial predators. Two aspects of its optical structure
which the sh feed, or other feeding birds.[64] suggest that the eye of this species is adapted to vision at
Birds that sh by stealth from above the water have to night. In the shearwaters eyes the lens does most of the
correct for refraction particularly when the sh are ob- bending of light necessary to produce a focused image
served at an angle. Reef herons and little egrets appear to on the retina. The cornea, the outer covering of the eye,
be able to make the corrections needed when capturing is relative at and so of low refractive power. In a diur-
sh and are more successful in catching sh when strikes nal bird like the pigeon, the reverse is true; the cornea is
are made at an acute angle and this higher success may be highly curved and is the principal refractive component.
The ratio of refraction by the lens to that by the cornea is
due to the inability of the sh to detect their predators.[65]
Other studies indicate that egrets work within a preferred 1.6 for the shearwater and 0.4 for the pigeon; the gure
for the shearwater is consistent with that for a range of
angle of strike and that the probability of misses increase [68]
when the angle becomes too far from the vertical leading nocturnal birds and mammals.
to an increased dierence between the apparent and real The shorter focal length of shearwater eyes give them a
depth of prey.[66] smaller, but brighter, image than is the case for pigeons,
Birds that pursue sh under water like auks and divers so the latter has sharper daytime vision. Although the
have far fewer red oil droplets,[2] but they have special Manx shearwater has adaptations for night vision, the ef-
exible lenses and use the nictitating membrane as an ad- fect is small, and it is likely that these
[68]
birds also use smell
ditional lens. This allows greater optical accommodation and hearing to locate their nests.
for good vision in air and water.[7] Cormorants have a It used to be thought that penguins were far-sighted on
greater range of visual accommodation, at 50 dioptres, land. Although the cornea is at and adapted to swim-
than any other bird, but the kingshers are considered to ming underwater, the lens is very strong and can com-
have the best all-round (air and water) vision.[2] pensate for the reduced corneal focusing when out of
water.[59] Almost the opposite solution is used by the
hooded merganser which can bulge part of the lens
through the iris when submerged.[59]

fovea

6 See also
Visual system
pecten

7 Notes
Each retina of the Manx shearwater has one fovea and an elon-
gated strip of high photoreceptor density[56]
Tubenosed seabirds, which come ashore only to breed and
spend most of their life wandering close to the surface of
the oceans, have a long narrow area of visual sensitivity
on the retina[1] This region, the area giganto cellularis,
has been found in the Manx shearwater, Kerguelen petrel,
great shearwater, broad-billed prion and common diving-
petrel. It is characterised by the presence of ganglion cells
which are regularly arrayed and larger than those found in
the rest of the retina, and morphologically appear similar
to the cells of the retina in cats. The location and cellu-
lar morphology of this novel area suggests a function in
the detection of items in a small binocular eld projecting
below and around the bill. It is not concerned primarily
with high spatial resolution, but may assist in the detec-
tion of prey near the sea surface as a bird ies low over
it.[67]
The Manx shearwater, like many other seabirds, visits its
breeding colonies at night to reduce the chances of attack
[1] Gntrkn, Onur, Structure and functions of the eye in
Sturkie (1998) 118
[2] Sinclair (1985) 88100
[3] White, Craig R.; Day, N; Butler, PJ; Martin, GR;
Bennett, Peter (July 2007). Bennett, Peter, ed.
Vision and Foraging in Cormorants: More like
Herons than Hawks?" (PDF). PLoS ONE. 2 (7): e639.
doi:10.1371/journal.pone.0000639. PMC 1919429 .
PMID 17653266.
[4] Martin, Graham R.; Katzir, G (1999). Visual elds in
short-toed eagles, Circaetus gallicus (Accipitridae), and
the function of binocularity in birds. Brain, Behaviour
and Evolution. 53 (2): 5566. doi:10.1159/000006582.
PMID 9933782.
[5] Jones, Michael P; Pierce Jr, Kenneth E.; Ward, Daniel
(April 2007). Avian vision: a review of form and
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Robert G. Cook, ed. (2001). Avian Visual Cognition
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[68] Martin, Graham R.; Brooke, M. de L. (1991). The

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8 References
Burton, Robert (1985). Bird Behaviour. London:
Granada Publishing. ISBN 0-246-12440-7.

Sinclair, Sandra (1985). How Animals See: Other

Visions of Our World. Beckenham, Kent: Croom
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12 10 TEXT AND IMAGE SOURCES, CONTRIBUTORS, AND LICENSES

10 Text and image sources, contributors, and licenses

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Bird vision Source: https://en.wikipedia.org/wiki/Bird_vision?oldid=776793984 Contributors: Shyamal, Jimfbleak, Andrewman327,
Raul654, Gilgamesh~enwiki, Bender235, Desultor, 99of9, Sabines Sunbird, Wtmitchell, Woohookitty, BD2412, Rjwilmsi, Nihiltres,
RussBot, Mithridates, Gaius Cornelius, Dugosz, SCZenz, Epipelagic, Asams10, BorgQueen, Sardanaphalus, SmackBot, Chris the speller,
Colonies Chris, Snowmanradio, Richard001, Dicklyon, Ravewolf, JMK, Danlev, ShelfSkewed, MaxEnt, Mojo Hand, Headbomb, Atavi,
Rie, Hassocks5489, R'n'B, Adamd1008, VolkovBot, The Sky May Be, Cloudswrest, Viridiavus~enwiki, Wingedsubmariner, Eddy-
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DOI bot, SuperlativeHors, Luckas-bot, Yobot, Fraggle81, Julia W, Vincnet, AnomieBOT, Jim1138, Royote, JackieBot, Citation bot,
GrouchoBot, Sfoleor, Prari, LucienBOT, Citation bot 1, Pinethicket, Jonesey95, Tom.Reding, Trappist the monk, Vrenator, Zephyric,
Woodlot, FoxRaweln, DARTH SIDIOUS 2, RjwilmsiBot, EmausBot, John of Reading, Dcirovic, K6ka, RaymondSutanto, ClueBot NG,
Onano, Slartibartfastibast, Braincricket, Helpful Pixie Bot, BG19bot, Ccevo2011, BattyBot, Cyberbot II, ChrisGualtieri, Bw mutley, The-
JJJunk, SD5bot, Dexbot, Sigvit, Jochen Burghardt, Bernatel, Anrnusna, JaconaFrere, Monkbot, SantiLak, Parrispower, Lythronaxargestes,
Ccevol2014, Nowpedia, Ollind, CAPTAIN RAJU, The Garages Dragon, Bismvth, OlleLindiLund, Peter.olsson84, GreenC bot, Henryc-
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