Albinism in biology

This article is about the biological term. For the medical condition in humans, see Albinism.
"Claude", an albino alligator at the California Academy of Sciences
Individual with normal pigmentation
Examples of albino laboratory mammals
Albino mouse
Albino guinea-pig
Albino rat

Albinism in biology is the "Congenital absence of any pigmentation or coloration in a person, animal, or plant, resulting in white hair and pink eyes in mammals."[1] Varied use and interpretation of the terms means that written reports of albinistic animals can be difficult to verify. Albinism can reduce the survivability of an animal, for example, it has been suggested that albino alligators have an average survival span of only 24 hours due to the lack of protection from UV and their lack of camouflage to avoid predators.[2] Albino animals have characteristic pink or red eyes because the lack of pigment in the iris allows the blood vessels of the retina to be visible. Familiar albino animals include in-bred strains of laboratory animals (rats, mice, guinea-pigs and rabbits), but populations of naturally occurring albino animals exist in the wild, e.g. Mexican cave tetra. Albinism is a well-recognized phenomenon in mollusks, both in the shell and in the soft parts. It has been claimed by some, e.g.[3] that "albinism" can occur for a number of reasons aside from inheritance including genetic mutations, diet, living conditions, age, disease, or injury, however, this is contrary to definitions where the condition is inherited.

In plants, albinism is characterised by partial or complete loss of chlorophyll pigments and incomplete differentiation of chloroplast membranes. Albinism in plants interferes with photosynthesis which can reduce survivability.[4] Some plant variations may have white flowers or other parts. However, these plants are not totally devoid of chlorophyll. Terms associated with this phenomenon are "hypochromia" and "albiflora".[5]

Biological colouration

Biological pigments are substances produced by living organisms that have a colour resulting from selective colour absorption. What we perceive as "colour" are the wavelengths of light that are not absorbed by the pigment, but which instead are reflected. Biological pigments include plant pigments and flower pigments.

Animal colouration

Main article: Animal coloration

Animals can appear coloured due to two mechanisms, pigments and structural colours. Animals may have both biological pigments and structural colours, for example, some butterflies.

Pigments

Many animal body-parts, such as skin, eyes, feathers, fur, hair, scales and cuticles, contain pigments in specialized cells called chromatophores. These cells are found in a wide range of animals including amphibians, fish, reptiles, crustaceans and cephalopods. Mammals and birds, in contrast, have a class of cells called melanocytes for colouration. The term chromatophore can also refer to coloured, membrane-associated vesicles found in some forms of photosynthetic bacteria. Chromatophores are largely responsible for generating skin and eye colour in poikilothermic animals and are generated in the neural crest during embryonic development. Mature chromatophores are grouped into subclasses based on their colour under white light:

Structural colours

Animals can also appear coloured due to structural colour, the result of coherent scattering[6] perceived as iridescence. The structures themselves are colourless. Light typically passes through multiple layers and is reflected more than once. The multiple reflections compound one another and intensify the colours. Structural colour differs according to the observer's position whereas pigments appear the same regardless of the angle-of-view. Animals which show iridescence include mother of pearl seashells, fish and peacocks. These are just a few examples of animals with this quality, but it is most pronounced in the butterfly family.

Plant colouration

The primary function of pigments in plants is photosynthesis, which uses the green pigment chlorophyll along with several red and yellow pigments including porphyrins, carotenoids, anthocyanins and betalains.

Definition

Albino wallaby with her offspring

Definitions of albinism vary and are inconsistent. Whilst they are clear and precise for humans and other mammals, this is because the majority of mammals have only one pigment, melanin. Many animals have pigments other than melanin, and some also have structural colours. Some definitions of albinism, whilst taking most taxa into account, ignore others. So, "a person or animal with very pale skin, white hair or fur, and pink eyes caused by a medical condition that they were born with"[8] and "a person or animal with white skin and hair and pink eyes"[9] do not include feathers, scales or cuticles of birds, fish and invertebrates. Nor do they include plants. Some definitions are too broad to be of much use, e.g. "an animal or plant with a marked deficiency in pigmentation"[10]

Other definitions of albinism encompass most of the major animal taxa, but ignore the several other pigments that non-mammalian animals have and also structural colouration. For example, "Absence of the pigment melanin in the eyes, skin, hair, scales, or feathers."[11] refers only to the pigment melanin.

Because of the various uses of different terms applied to colouration, some authors have indicated that the colour of the eyes is the defining characteristic of albinism, e.g. "This leads to a good diagnostic feature with which to distinguish leucistic and albino individuals – the colour of the eye."[12]

The term "partial albino" is sometimes used in the literature, however, it has been stated that "A common misnomer is ‘partial albino’ – this is not possible since albinism affects the whole plumage of a bird, not just part"[12] and the definition of albinism precludes the possibility of "partial albinism" in which a mostly white bird shows some form of melanin pigmentation. "It is simply impossible, just like being ‘partially pregnant’ ".[13]

One definition states that "albinism, (from the Latin albus, meaning “white”), hereditary condition characterized by the absence of pigment in the eyes, skin, hair, scales, or feathers",[14] however, this does not encompass invertebrates, nor does it include plants. Furthermore, it could be interpreted that "...absence of pigment..." does not include an absence of structural colours.

The lack of clarity about the term is furthered when the name of an animal includes the term "albino" although the animals (clearly) do not have the condition. For example, the albino gaur has this name because it is ash-grey whereas other gaur are almost black, and the American albino horse is a chestnut with a dilution gene.

A clear definition appears to be – "Congenital absence of any pigmentation or coloration in a person, animal, or plant, resulting in white hair and pink eyes in mammals."[1] Whilst this does not state specifically that non-mammalian albino animals (or plants) are white, this can be inferred from "...absence of any pigmentation or coloration..." Due to the varied use and interpretation of the term "albino", written reports of albinistic organisms can often not be verified.

Mechanism and frequency

Melanin is an organic pigment that produces most of the colour seen in mammals. Depending on how it is created, melanin comes in two colour ranges, eumelanin (producing dark browns and blacks) and pheomelanin (producing light reddish tans and blondes). The dark and light melanins have their influence either alone or in conjunction, making either plain or multi-coloured coats. Sometimes, in a condition called agouti, they make multi-coloured individual hairs. The production of melanin occurs in melanocytes in a complex process involving the enzyme tyrosinase. Mammals have a gene that codes for the presence of tyrosinase in cells – called the TYR gene. If this gene is altered or damaged, melanin cannot be reliably produced and the mammal will become an albino. Besides the TYR gene, several other genes can cause albinism. This is because other hormones and proteins are involved in melanin production, the presence of which is genetically determined. In mice, a total of 100 genes are known to affect albinism.[15]

All the genetic traits for albinism are recessive traits. This means that their influence is hidden when paired with stronger traits. For the recessive albino trait to be expressed in a mammal, the offspring must inherit a recessive gene from both parents.[15]

Albinism occurs throughout the animal kingdom. At least 300 species in North America have had at least one reported instance of albinism. Albinism occurs once in every 10,000 mammal births, however, albinism is much more common in birds, occurring once in every 1,764 births.[16] Albinism in animals is most commonly seen in birds, reptiles and amphibians. It is more rarely seen in mammals and other species. It is often difficult to explain these occasional occurrences, especially when only one documented incidence has occurred such as only one albino gorilla and one albino koala bear.[17]

Some species, such as white peacocks, swans and geese, are not believed to be true albinos, as they do not have red eyes, rather, their colouration is suggested to be the expression of a white fur or feather gene, not a lack of melanin.[18]

Consequences

Melanin has several functions in most mammals and other animals; these are disrupted by albinism.

Abnormal eye development and appearance

Melanin functions in the normal development of various parts of the eye, including the iris, retina, eye muscles, and optic nerve. The absence of melanin results in abnormal development of eyes and leads to problems with focusing, and depth perception. The eyes of albino animals appear red because the colour of the red blood cells in the retina can be seen through the iris, which has no pigment to obscure this. Some albino animals may have pale-blue eyes due to other colour generating processes. Albino vertebrates exposed to intense light typically lose photoreceptors due to apoptosis.[19]

In all albino mammals studied, the centre of the retina is under-developed and there is a deficit of rod cells; the central ganglion cell density is approximately 25% below normal (except for the gray squirrel). In nearly all mammals, the overwhelming majority of photoreceptors are rods rather than cones. Albinism specifically affects the rod cells, but the number and distribution of the cones is unaffected. In contrast, the retina of birds is cone rich meaning that the vision of albino birds is affected less than albino mammals.[20]

Reduced protection from sunlight

Melanin protects the skin from ultra-violet radiation in sunlight. Melanosomes block harmful electromagnetic radiation from the sun while allowing beneficial frequencies to enter the body.

Survival disadvantages

Many animals with albinism lack their protective camouflage and are therefore less able to conceal themselves from their predators or prey: The survival rate of animals with albinism in the wild can be low,[21][22] however, it has been stated that in studies where animals had many places to hide, predators captured albino and normally coloured animals at the same rate.[15] Furthermore, albino animals may be excluded from families or other groups,[16] or rejected as mates.[15]

The novelty of albino animals has occasionally led to their protection by groups such as the Albino Squirrel Preservation Society. They have also been protected in studies on their ecology and behaviour.[23]

Reduced viability

Studies on medaka fish in the laboratory, i.e. with no predators, sufficient food supply, controlled temperatures, etc., found that albinos had considerably reduced viability; from 800 albino embryos, only 29 survived to full adulthood.[24] Early studies on fish led some researchers to describe albinism as a "semi-lethal mutation".[24]

Hearing disorders

Pigmentation disorders such as albinism are occasionally associated with hearing impairments in mice, rats, guinea pigs and cats.[25]

In mammals

Artificially selected

Examples of pet mammals artificially selected for albinism
Albino ferret
Albino rabbit

Intentionally bred albinistic strains of some animal species are commonly used as model organisms in biomedical research and also as pets. Examples include the BALB/c mouse and Wistar and Sprague Dawley rat strains, laboratory rabbits, guinea-pigs and ferrets. Albino axolotl (an amphibian) are also used widely in the laboratory as their transparent skin allows observation of the underlying tissues during limb regeneration.[26] Some researchers have argued that albino animals are not always the best choice for scientific studies due to the consequences of albinism (e.g. hearing and visual impairments).[27]

Many individual albino mammals are in captivity and were caught as youngsters. It is doubtful whether these individuals would have survived to become adults without the protection and care they receive in captivity.

Naturally occurring

"Snowflake", a blue-eyed albino Western Lowland gorilla caught in the wild

It has been claimed that "Squirrels are the only known albino mammal to survive successfully in the wild".[20] In all albino mammals studied, the eye color is green, blue or red due to absence of eumelanin in the iris, and the centre of the retina is under-developed with a deficit of rod cells; the central ganglion cell density is approximately 25% below normal. In nearly all mammals, the overwhelming majority of photoreceptors are rods rather than cones. Albinism specifically affects the rod cells, but the number and distribution of the cones is unaffected. The retina of the squirrel (Sciurus carolinensis) is unusual for mammals as it is rich in cones. Central cell densities are less than 5% lower in albino squirrels than in pigmented individuals. This relatively minor disruption to vision is thought to assist in the survivability of albino squirrels in the wild. This is supported by observations that the behaviour of albinos in the wild, e.g. leaping from branch to branch, is similar to pigmented squirrels.[20]

A 2012 survey of the literature reported that in India, there were several records of albino mammals including the tiger, lesser rat-tailed bat, chital, common palm civet, Northern palm squirrel five-striped palm squirrel and wild boar.[28]

Albino macaques have been reported in several occasions including a toque macaque (M. sinica), rhesus macaque (M. mulatta), and bonnet macaque (M. radiata).[28]

A study on albinistic prairie voles (Microtus ochrogaster) found that albinism in this species conferred an advantage for the males compared to the wild-type; albino males had higher mount frequencies than wild-type males. In addition, the albinos had greater differential fertilizing capacity.[29]

Marine mammals

The costs of albinism for marine mammals may include reduced heat absorption in colder waters, poor camouflage from predators, increased sensitivity to sunlight, and impaired visual communication. Despite the costs, some individuals do reach adult age and breeding status.[30]

Albino dolphins were first sighted in the Gulf of Mexico in 1962. Since 1994, three further individuals have been seen. These tend to be pink in colour due to blood vessels showing through the blubber and unpigmented skin.[31]

A report published in 2008 stated that in marine mammals, "anomalously white" individuals have been reported for 21 cetacean species and 7 pinniped species but there were no known reports of anomalously white sea otters (Enhydra lutris) or sirenians. [30]

Whales and dolphins also may appear white if extensively scarred, or covered with a fungus, such as Lobo's disease (also known as "lobomycosis").[30]

Famous albino mammals

Famous albino mammals include "Migaloo", a humpback whale living off the coast of Australia; Pinky, a bottlenose dolphin living in and around in Calcasieu Lake, Louisiana; "Carolina Snowball", a popular albino bottlenose dolphin displayed at the Miami Seaquarium in the early 1960s; "Snowflake", a Barcelona Zoo gorilla, and "Mahpiya Ska", (Sioux for "White Cloud"), a buffalo in Jamestown, North Dakota.[32][33] The inspiration for Herman Melville's novel Moby-Dick was a sperm whale known as Mocha Dick.

In birds

Kookaburras
Albino kookaburra
Blue winged kookaburra with normal pigmentation

The most important pigments that determine plumage coloration in birds are melanines and carotenoids. The latter are ingested in food and transformed into colour pigments by enzymes. Aberrations in this pigmentation are mostly caused by food deficiencies and usually do not have a genetic basis. Well-known examples are flamingos which owe their distinct pink colour to the presence of red carotenoids in their natural food. When these carotenoids are in short supply, these birds appear white after the next moult. Mutations causing changes in carotenoid-based colour pigments are rare; melanine mutations occur much more frequently. Two types of melanin, eumelanin and phaeomelanin, are present in birds. In the skin and eyes, only eumelanin is present. In some bird species, the colour is completely caused by eumelanin, however, both types of melanin are found in most species. In birds, albinism has been defined as "a total lack of both melanins in feathers, eyes and skin as a result of an inherited absence of tyrosinase",[13] however, this ignores the effects of other pigments and structural colours.

An albino bird has a white beak, white plumage, non-coloured skin, white talons and pink or red eyes. Albinism is only seen in about 1 of every 1,800 birds. The two most common species of albino birds are the common house swallow and the American robin.[17] Famous albino birds include "Snowdrop", a Bristol Zoo penguin.[34]

In one study, albinism in birds has been categorised according to the extent of pigment absence.[35] –

However, it has been argued that the definition of albinism precludes the possibility of "partial albinism" in which a mostly white bird shows some form of melanin pigmentation. "It is simply impossible, just like being ‘partially pregnant’.[13]

In fish

As with other animals, it has been stated that for fish to be properly described as "albino", they must have a white body and pink or red eyes.[24]

Artificially selected

Examples of fish artificially selected for albinism
Albino catfish
Albino Peacock Cichild

Several types of aquarium fish are called albino, e.g. albino driftwood catfish, albino festivium, albino red tailed shark and albino tiger shark, however, photographs of these four fish indicate that the latter three have dark eyes and/or pigmented fins.[36]

Zebrafish have three types of chromatophores – iridophores, melanophores and xanthophores which produce silver, black and yellow pigmentation respectively. Zebrafish that lack iridophores are known as "roy mutants", those which lack melanophores as "albino mutants" and those which lack both melanophores and iridophores as "ruby mutants". The gross eye morphology, feeding and swimming behaviours between wild-type and albino zebrafish were indistinguishable, except under dim or bright light or low contrast.[37] In mammals, albinism is occasionally associated with hearing impairments. However, when tested, there was no differences in responses between wild-type and albinistic European wels (Silurus glanis) and South American bronze catfish (Corydoras aeneus). Similarly, Mexican blind cave fish (Astyanax mexicanus) do not differ in hearing sensitivity from the normally pigmented and eyed surface-dwelling populations. Fish lack melanin in the inner ear, meaning that hearing in fishes is less likely to be affected by albinism than in mammals.[25]

Naturally occurring

There are several reports of total albinism in both freshwater and marine fish, however, frequently captured albino fish are only reported in aquarium magazines and local newspapers.[38]

The incidence of albinism can be artificially increased in fish by exposing the eggs to heavy metals (e.g. arsenic, cadmium, copper, mercury, selenium, zinc).[39]

In the wild, albinism is reasonably common in the teleosts, especially the Pleuronectiformes (flatfish), however, it is rarely reported in the elasmobranchs.[40] Albinism has been reported in several catfishes, hagfish, lampreys, sharks, rays and numerous teleost fishes, e.g. grunts or cyprinids.[25]

In Actinopterygii

Albino and normally pigmented channel catfish (Ictalurus punctatus) differ in their characteristics. Normal individuals of this species are superior to albinos in body weight and total length. Albinos crossed with other albinos require 11 days longer to spawn and produce smaller egg masses. These masses contain eggs of lighter weight with poorer hatchability than crosses of normal fish. The albino fish have lower survival rates than normal fish but dress-out percentages are nearly equal.[41]

Some wild cave fish have populations that are albinistic. The Mexican cave tetra is a species that has evolved specialized characteristics in a series of independent caves. One of these is albinism linked to the Oca2 gene, a known pigmentation gene, This trait has evolved independently in at least two caves.[42]

In Chondrichthyes

In the class chondrichthyes, several species of naturally occurring albino rays and sharks have been recorded. Furthermore, an albino individual spotted ratfish (Hydrolagus colliei) from the order Chimaeriformes has been reported.[43]

Albinistic individuals of the following shark species have been reported:[44][45]

A study published in 2006 reported albinistic individuals of the following ray species:[45]

In Hyperoartia

There are several reports of albino lampreys and it has been estimated that this occurs at a frequency of one in 100,000 normal individuals.[24]

In Sarcopterygii

Albinism in the African lungfish has been reported on at least two occasions.[46]

In reptiles

"Albino" California kingsnake

Many reptiles labeled as albino are, in fact, not completely lacking in all colour pigments. They are actually amelanistic, not albino. Reptiles often possess at least two pigments. Among the most common are xanthin (yellow) and etherin (red). An amelanistic reptile therefore, may still have pale yellow, orange, or red pigmentation.

The California Academy of Science, in the Steinhart Aquarium, as of 2015, has on display an albino American alligator named "Claude". The alligator is blind because of lack of pigment in its eyes.[47] The albino alligator hatched from the egg in 1995 in Florida, and was brought to the Academy in 2008. This alligator would not have survived in the wild because its whiteness would have made it too easy a prey object. The only known albino alligators are in captivity.[48]

In snakes, partial absence of pigment is more common than absolute albinism. For snakes that are usually patterned in colours, they appear as a faint blue, peach or yellowish. In these cases, there has been a genetic mutation in the melanin and pigment delivery. The appearance comes from the inability for full colours to be present, such as black, red, brown and others. The eyes of an albino snake are typically red or pink. Albino snakes can remain in the sunlight for several hours with minimal harm. Corn snakes and snakes of larger types, such as a boa or diamondback snakes, are the most commonly affected by albinism often appearing to be a pinkish or yellowish colour.[17]

Albino tortoises and turtles are uncommon; Sulcata tortoises are the most likely type of turtle to express albinism. The shells have an almost yellow colouration and they have pink eyes. For turtles, a pure white colour is nearly impossible, even with albinism. Albino turtles can have a longer lifespan than many other albino animals; their hard shells help to prevent predation and other environmental challenges. Vision and sensory organs are slightly affected.[17]

An albino anole has been reported and photographed.[49] In 2007, it was reported that an albino stumpy-tail lizard (possibly a shingleback lizard), approximately 12 cm long and about one-year-old, had been found in Victoria, Australia.[50] (Also see )

There are three known "albino" strains of leopard gecko, however, breeders state that albinos are generally recognized by their lack of black pigment and having red eyes is not a requirement to be considered an albino. These three strains are called "tremper albios", "rainwater albinos" and "bell albinos".[51]

In amphibians

As with reptiles, many amphibians labeled as albino are, in fact, not completely lacking in all colour pigments. They are actually amelanistic, not albino. Amphibians have six types of chromatophore in their skin, i.e. melanophores, xantophores, erythrophores, leucophores, cyanophores and iridophores.[52] An amelanistic amphibian therefore, may still have various pigmentation.

The incidence of albinism in frogs salamanders and newts is relatively higher than other taxa. It has been estimated that one on four hundred of these animals is albino. When albino tadpoles hatch, they are almost transparent. This may help camouflage them initially, however, after two weeks, when their hind legs begin to emerge, they become milky white. A survey in 2001 found hundreds of tiny albino plains leopard frogs, but when the researcher returned a few months later, not a single albino adult could be found.[26]

In European Salamandridae, albinism has been recorded in the fire salamander (Salamandra salamandra), gold-striped salamander (Chioglossa lusitanica), Italian crested newt (Triturus carnifex), marbled newt (Triturus marmoratus), Iberian ribbed newt (Pleurodeles waltl), alpine newt (Ichthyosaura alpestris) and two sub-species of the smooth newt (Lissotriton vulgaris vulgaris and Lissotriton vulgaris meridionalis).[52]

In invertebrates

Albinism in mollusks has been recognized to be a hereditary phenomenon at least since 1900.[53] Albinism in mollusks can exist to a variable degree: sometimes an individual snail will have a normally pigmented body, but the shell will be completely without the normal pigmentation because of a defect in the cells of the mantle. Shells of certain mollusk species can be translucent when they lack the normal pigmentation.[54]

In insects

The neurohormone [His7]-corazonin induces darkening of the cuticle of Locusta migratoria. The Okinawa strain of this species is deficient in [His7]-corazonin and is albino. One of the typical features of Locusta migratoria ia that they are gregarious locusts. However, the albino strain shows more solitarious behaviour.[55]

The yellow mutation in fruit flies is a mutation causing a congenital lack of normal pigment; it is a similar phenomenon to albinism in other organisms.[56]

"Albino" individuals of normally red citrus red mites (Panonychus citri) occasionally appear in laboratory colonies, however, these still contain green and yellow pigments. This albinism does not affect mortality.[57]

In echinoderms

The Japanese sea cucumber (Apostichopus japonicus) is an echinoderm which is caught in the wild or cultivated for food. Normal Japanese sea cucumbers start to develop pigmentation when they are about 1 cm long. The upper side becomes a dull, yellowish -brown to maroon and the underside a light brown. The body walls of adult, albino Japanese sea cucumbers contain only 0.24% melanin compared to 3.12% in normal adults. The difference in melanin content becomes visually apparent at 60 days of age. The epidermis is thinner in the albinos and contains fewer melanocytes. Albino individuals are similar to normal individuals in growth rate, digestion rate and fertility.[58]

Astacin is the main carotenoid in marine crustaceans (and fish). It has been shown that adding astacin to the feed can improve the skin and muscle colour of marine organisms and thereby increase their commercial and ornamental value.[58]

In arachnids

"Depigmented" arthropods have been found, usually in cave populations.[59]

Related pigment disorders

In some animals, albinism-like conditions may affect other pigments or pigment-production mechanisms:

In plants

In plants, albinism is characterised by partial or complete loss of chlorophyll pigments and incomplete differentiation of chloroplast membranes. Albinism in plants interferes with photosynthesis which can reduce survivability.[4] Some plant variations may have white flowers or other parts. However, these plants are not totally devoid of chlorophyll. Terms associated with this phenomenon are "hypochromia" and "albiflora".[5]

Plants that are pale simply from being in the dark are termed etiolated.

Albino redwoods are rare examples of an albino tree with white needles; despite its lack of chlorophyll it may grow to substantial size as a parasite, usually on the base of the Redwood tree from which it first grew.[63][64][65] Only about sixty examples of albino redwoods are known.[64]

Albinism has frequently occurred in progeny of Black Tartarian, Bing and Hedelfingen varieties of sweet cherry.[66]

In human culture

The Albino Squirrel Preservation Society was founded at the University of Texas at Austin in 2001. Members of the society at the University of North Texas petitioned for an election to name their albino squirrel as the university's secondary mascot. The University of Louisville in Kentucky also has a documented population of albino squirrels.[67]

Albino animals are often kept as pets, for example, African clawed frog, guinea pigs and peacocks.[67]

See also

Wikimedia Commons has media related to Albinism.

References

  1. ↑ 1.0 1.1 "Albinism". The Free Dictionary. Retrieved January 31, 2015.
  2. ↑ "NEW Albino Alligators". The Georgia Aquarium. Retrieved January 20, 2015.
  3. ↑ McCardle, H. "Albinism in wild vertebrates". Texas State University. Retrieved January 24, 2015.
  4. ↑ 4.0 4.1 Kumari, M.; Clarke, H.J.; Small, I.; Siddique, K.H.M. (2009). "Albinism in plants: A major bottleneck in wide hybridization, androgenesis and doubled haploid culture". Critical Reviews in Plant Sciences. 28 (6): 393–409. doi:10.1080/07352680903133252.
  5. ↑ 5.0 5.1 "Albiflora". Retrieved January 28, 2015.
  6. ↑ Mäthger, L.M.; Denton, E.J.; Marshall, N.J.; Hanlon, R.T. (2009). "Mechanisms and behavioural functions of structural coloration in cephalopods". J. R. Soc. Interface 6: 149–163. doi:10.1098/rsif.2008.0366 (inactive 2015-03-29).
  7. ↑ Taylor, J.D. (1969). "The effects of intermedin on the ultrastructure of amphibian iridophores". Gen Comp Endocrinol 12 (3): 405–16. doi:10.1016/0016-6480(69)90157-9. PMID 5769930.
  8. ↑ "Albino". Retrieved January 31, 2015.
  9. ↑ "Albino". Cambridge Dictionaries On-line. Retrieved January 31, 2015.
  10. ↑ "Albino". Albino.com. Retrieved January 31, 2015.
  11. ↑ "Albinism". Mirriam-Webster. Retrieved January 31, 2015.
  12. ↑ 12.0 12.1 "Leucisam and albinism". British Trust for Ornithology. Retrieved January 31, 2015.
  13. ↑ 13.0 13.1 13.2 de Grouw, H. (2006). "Not every white bird is an albino: sense and nonsense about colour aberrations in birds." (PDF). Dutch Birding 28: 79–89.
  14. ↑ "Albinism". Encyclopædia Britannica. Retrieved January 27, 2015.
  15. ↑ 15.0 15.1 15.2 15.3 Binkley, S.K. "Color on, color off" (PDF). Retrieved January 22, 2015.
  16. ↑ 16.0 16.1 Nasr, S.R. "How albinism works". Howstuffworks. Retrieved January 23, 2015.
  17. ↑ 17.0 17.1 17.2 17.3 "Albino animals". Wonders of the Nature. Retrieved January 24, 2015.
  18. ↑ "Albinism in animals". Science illustrated. 2011. Retrieved January 24, 2015.
  19. ↑ Allison, W.T.; Hallows, T.E.; Johnson, T.; Hawryshyn; C.W.; Allen, D.M. (2006). "Photic history modifies susceptibility to retinal damage in albino trout". Visual Neuroscience 23 (1): 25–34. doi:10.1017/s0952523806231031. PMID 16597348.
  20. ↑ 20.0 20.1 20.2 Esteve, J.V.; Jeffery, G. (1998). "Reduced retinal deficits in an albino mammal with a cone rich retina: A study of the ganglion cell layer at the area centralis of pigmented and albino grey squirrels". Vision Research 38 (6): 937–940. doi:10.1016/s0042-6989(97)00229-0. PMID 9624442.
  21. ↑ Ilo Hiler, Albinos. Young Naturalist. The Louise Lindsey Merrick Texas Environment Series, No. 6, pp. 28–31. Texas A&M University Press, College Station (1983)
  22. ↑ Dobosz, ByS; Kohlmann, K.; Goryczko, K.; Kuzminski, H. (July 2008). "Growth and vitality in yellow forms of rainbow trout". Journal of Applied Ichthyology 16 (3): 117–20. doi:10.1046/j.1439-0426.2000.00147.x.
  23. ↑ Menzel, R.W. (1958). "Further notes on the albino catfish". The Journal of Heridity 49 (6): 284–293.
  24. ↑ 24.0 24.1 24.2 24.3 Purdom, C.F., ed. (1993). Genetics and Fish Breeding. Chapman and Hall, London. ISBN 9780412330407. Retrieved February 2, 2015.
  25. ↑ 25.0 25.1 25.2 Lechner, W.; Ladich, F. (1987). "How do albino fish hear?". Journal of Zoology 283 (3): 186–192. doi:10.1111/j.1469-7998.2010.00762.x. PMID 21552308.
  26. ↑ 26.0 26.1 Halls, K.M. (2004). Albino Animals. Lerner Publishing Group, Inc, Minneapolis. ISBN 9781581960198.
  27. ↑ Creel, D. (June 1980). "Inappropriate use of albino animals as models in research". Pharmacology, Biochemistry, and Behavior 12 (6): 969–7. doi:10.1016/0091-3057(80)90461-X. PMID 7403210.
  28. ↑ 28.0 28.1 Anil Mahabal, P.D. Rane and S.K. Pati (2012). "A case of total albinism in the Bonnet Macaque Macaca radiata (Geoffroy) from Goa" (PDF). ZOO's PRINT 27 (12): 22.
  29. ↑ Dewsbury, D.A.; Ward, S.E. (1985). "Effects of albinism on copulatory behavior and sperm competition in prairie voles (Microtus ochrogaster)". Bulletin of the Psychonomic Society 23 (1): 68–70. doi:10.3758/bf03329782.
  30. ↑ 30.0 30.1 30.2 Fertl, D. and Rosel, P.E. (2008). "Albinism". In William F. Perrin, Bernd Würsig, and J.G.M. Thewissen. Encyclopaedia of Marine Mammals (2nd edition). Elsevier, Amsterdam. pp. 22–24. ISBN 978-0-12-373553-9.
  31. ↑ "Great White! 7 Albino Wonders of the Animal World". Web Ecoist. Retrieved January 20, 2015.
  32. ↑ "Rare Pink Dolphin Seen in Louisiana Lake – Science News | Science & Technology | Technology News". FOXNews.com. 2007-07-03. Archived from the original on 24 February 2010. Retrieved 27 February 2010.
  33. ↑ "Albino dolphin in northern Gulf of Mexico" (PDF). NOAA. Retrieved January 30, 2015.
  34. ↑ "Albino Animals from Snowflake the white gorilla to White Diamond the alligator". The Telegraph
  35. ↑ Alaja, P. and Mikkola, H. "Albinism in the Great Gray Owl (Strix nebulosa) and Other Owls." (PDF). Retrieved January 24, 2015.
  36. ↑ "List of aquarium fish species by name". Retrieved February 2, 2015.
  37. ↑ Ren, J.Q.; McCarthy, W.R.; Zhang, H.; Adolphc, A.R.; Lia, L. (2002). "Behavioral visual responses of wild-type and hypopigmented zebrafish". Vision Research 42 (3): 293–299. doi:10.1016/S0042-6989(01)00284-X. PMID 11809482.
  38. ↑ Leal, M.E. et al. (2013). "First record of partial albinism in two catfish species of Genidens (Siluriformes: Ariidae) in an estuary of Southern Brazil". Brazilian Archives of Biology and Technology 56 (2). Retrieved January 24, 2015.
  39. ↑ Brito, Marcelo F. G. de; Caramaschi, Érica P. (2005). "An albino armored catfish Schizolecis guntheri (Siluriformes: Loricariidae) from an Atlantic Forest coastal basin". Neotropical Ichthyology 3. doi:10.1590/S1679-62252005000100009.
  40. ↑ Nakaya K. (1973). "Albino zebra shark from the Indian ocean with comments on albinism in elasmobranchs" (PDF). Japanese Journal of Icththyology 20 (2): 120–122.
  41. ↑ Bondari, K. (1984). "Comparative performance of albino and normally pigmented channel catfish in tanks, cages, and ponds.". Aquaculture 37 (4): 293–301. doi:10.1016/0044-8486(84)90295-3.
  42. ↑ Protas, Meredith E; Hersey, Candace; Kochanek, Dawn; Zhou, Yi; Wilkens, Horst; Jeffery, William R; Zon, Leonard I; Borowsky, Richard; Tabin, Clifford J (2006). "Genetic analysis of cavefish reveals molecular convergence in the evolution of albinism". Nature Genetics 38 (1): 107–111. doi:10.1038/ng1700. PMID 16341223.
  43. ↑ Reum, J.C.P., Paulsen, C.E., Petsch, T.W. and Parker-Setter, S.L. (2008). "First record of an albino Chimeaeriforme fish." (PDF). Northewestern Naturalist 9: 60–62.
  44. ↑ Martin, R.A. "Albinism in sharks". ReefQuest Centre for Shark Research. Retrieved February 2, 2015.
  45. ↑ 45.0 45.1 Sandoval-Celtillo, J., Mariano-Melendez, E. and Villavicencio-Garayzar, C. (2006). "New records of albinism in two elasmobranchs: the tiger shark Galeocerdo cuvier and the giant electric ray Narcine entemedor". Cybium 30 (2): 191–192. Retrieved February 2, 2015.
  46. ↑ Hubbard, G.B.; Fletcher, K.C. (1985). "A seminoma and a leiomyosarcoma in an albino African lungfish (Protopterus dolloi)". Journal of Wildlife Diseases 21 (1): 72–74. doi:10.7589/0090-3558-21.1.72. PMID 3981753.
  47. ↑ The Swamp. California Academy of Sciences
  48. ↑ Guthrie, Julian (September 14, 2010) "Albino alligator Claude to mark 15th Birthday". sfgate.com
  49. ↑ "Albino anole". 2012. Retrieved March 8, 2012.
  50. ↑ "Rare albino lizard found in backyard". ABC. 2007. Retrieved March 8, 2015.
  51. ↑ Sage, P. (2006). "Guide to Leopard Gecko Morphs and Genetics". Retrieved March 8, 2015.
  52. ↑ 52.0 52.1 Modesti, A., Aguzzi, S. and Manenti, R. (2011). "A case of complete albinism in Lissotriton vulgaris meridionalis." (PDF). Herpetology Notes 4: 395–396.
  53. ↑ Taylor, John William (1900). Monograph of the Land & Freshwater Mollusca of the British Isles 1. Taylor brothers. p. 92.
  54. ↑ Taylor, John William (1900). Monograph of the Land & Freshwater Mollusca of the British Isles 1. Taylor brothers. p. 90.
  55. ↑ Hoste, B.; Simpson, S.J.; Tanaka, S.; De Loof, A.; Breuer M. (2002). "A comparison of phase-related shifts in behavior and morphometrics of an albino strain, deficient in [His7]-corazonin, and a normally colored Locusta migratoria strain". Journal of Insect Physiology 48 (8): 791–801. doi:10.1016/s0022-1910(02)00106-3. PMID 12770057.
  56. ↑ Service, Elizabeth. Fruit Fly Phenotypes. unc.edu.
  57. ↑ Wilkes, J.T. (1963). "A recessive albinism in the citrus red mite". Annals of the Entomological Society of America 56 (6): 792–795. doi:10.1093/aesa/56.6.792.
  58. ↑ 58.0 58.1 Zhao, H., Chen, M. and Yang, H. (2015). "Albinism". In Hongsheng Yang, Jean-Francois Hamel and Annie Mercier. The Sea Cucumber Apostichopus japonicus: History, Biology and Aquaculture. Academic Press. pp. 211–226. ISBN 978-0-12-799953-1.
  59. ↑ Locket, N.A. (1986). "Albinism and eye structure in an Australian scorpion, Urodacus yaschenkoi (Scorpiones, Scorpionidae)". Journal of Arachnology 14 (1): 101–115. JSTOR 3705557.
  60. ↑ Hesford, Clive (January 1998). "The Parblue Puzzle: Part 4—Common Parblue Varieties: The Cockatiel [Nymphicus hollandicus]" in The Genetics of Colour in the Budgerigar and Other Parrots
  61. ↑ "Amphibian Biology & Physiology: Caudata" at Amphibian Information Resource: An Educational Web Project About Amphibian Species; sourced December 2006, actual authoring/publication date unspecified.
  62. ↑ "Feather Colors: What We See" by Dr. Julie Feinstein of the American Museum of Natural History (NY), in Birder's World Magazine online archive; sourced December 2006, actual authoring/publication date unspecified.
  63. ↑ Stienstra, T. (11 October 2007). "It's no snow job – handful of redwoods are rare albinos". San Francisco Chronicle. Retrieved December 6, 2010.
  64. ↑ 64.0 64.1 Krieger, L. M. (2010-11-28). "Albino redwoods hold scientific mystery". MercuryNews.com. San Jose Mercury News. Retrieved 2012-11-23.
  65. ↑ "A Creepy Monster of the Forest: The Albino, Vampiric Redwood Tree". Discoblog. Discover Magazine. 2010-12-08. Retrieved 2012-11-23.
  66. ↑ Kerr, E.A. (1963). "Inheritance of crinkle, variegation, and albinism in sweet cherry". Canadian Journal of Botany 41 (10): 1395–1404. doi:10.1139/b63-122.
  67. ↑ 67.0 67.1 "10 Most Incredible Albino Animals on Earth". Scribol. Retrieved January 20, 2015.