The Spectrum of Darkness
An in-depth examination of hyperpigmentation, its genetic underpinnings, adaptive advantages, and diverse manifestations across species.
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What is Melanism?
Congenital Excess of Melanin
Melanism is defined as the congenital excess of melanin in an organism, leading to a distinctly dark pigmentation. This genetic phenomenon results in individuals appearing significantly darker than their typical counterparts within a species.
Pseudomelanism (Abundism)
A related but distinct variant of pigmentation is pseudomelanism, also known as abundism. This condition is characterized by dark spots or enlarged stripes that cover a substantial portion of an animal's body, creating an appearance that mimics true melanism without being a complete darkening of the entire organism.
Differentiating from Melanosis
It is crucial to distinguish melanism from melanosis. Melanosis refers to the pathological deposition of black matter, often associated with malignant characteristics, leading to pigmented tumors. Unlike melanism, which is a congenital trait, melanosis is a morbid condition.
Cultural Significance
Beyond its biological implications, melanism can hold cultural significance. For instance, melanistic guinea pigs (Cavia porcellus) are exceptionally rare and are utilized in rituals by Andean curanderos, highlighting a unique intersection of biology and traditional practices.
Adaptive Melanism
Evolutionary Advantage
Adaptive melanism describes instances where dark pigmentation confers a survival and reproductive advantage within a specific environment. This often occurs because darker individuals are better camouflaged, making them less conspicuous to predators. Conversely, some predators, such as leopards, leverage their dark coloration as a foraging advantage during nocturnal hunts.
Genetic Inheritance
Typically, adaptive melanism is a heritable trait. In many cases, a dominant allele is responsible for the excessive melanin production, leading to its pronounced expression in the phenotype. However, in certain species, such as the Zelandoperla fenestrata stoneflies, adaptive melanism associated with Batesian mimicry is controlled by a recessive allele at the ebony locus.
Diverse Animal Examples
Adaptive melanism is observed across a wide range of animal taxa. Notable examples include various mammals like squirrels, numerous felid (cat) species, canids, and even reptiles such as coral snakes. This widespread occurrence underscores the significant evolutionary benefits that dark coloration can provide.
Industrial Melanism
Pollution-Driven Evolution
Industrial melanism is a specific evolutionary phenomenon observed primarily in insects, famously exemplified by the peppered moth (Biston betularia). It occurs in regions subjected to industrial pollution, where environmental changes drive selective pressures favoring darker pigmented individuals.
Camouflage Advantage
The primary hypothesis for industrial melanism is that darker individuals gain a significant advantage through natural selection because they are better camouflaged against backgrounds darkened by industrial soot and pollutants. This reduces their visibility to predators, increasing their survival rates.
Reversal with Pollution Reduction
A compelling aspect of industrial melanism is its reversibility. As industrial pollution levels decreased in many areas, the selective advantage shifted back to lighter forms. Consequently, the frequency of melanism in affected populations declined, demonstrating the dynamic nature of natural selection in response to environmental changes.
Alternative Explanations
While camouflage is the leading explanation, other theories have been proposed to account for the prevalence of industrial melanism. These include the possibility that melanin pigment enhances the function of immune defenses in darker individuals, or that the darker coloration provides a thermal advantage by absorbing more heat, particularly in cooler environments.
Feline Melanism
A Common Polymorphism
Melanistic coat coloration is a common polymorphism found in 11 out of 37 known felid species. While it can reach high frequencies within certain populations, it rarely achieves complete fixation, meaning both melanistic and non-melanistic forms typically coexist.
Geographic Distribution
The "black panther" is a well-known example of a melanistic leopard, frequently observed in the equatorial rainforests of Malaya and the tropical rainforests on the slopes of African mountains like Mount Kenya. Melanic forms of jaguars are also common in specific regions of South America. Even bobcats, typically spotted, have been recorded with melanistic forms in subtropical Florida.
Genetic Underpinnings
Research has elucidated the genetic basis of melanism in various felids. In jaguars, a dominant mode of inheritance was confirmed, linked to a 15-base pair inframe deletion in the MC1R gene. For jaguarundis, a 24-base pair deletion causes an incompletely dominant allele for melanism. In domestic cats, a 2-base pair deletion in the agouti signaling peptide coding region of the agouti gene leads to black coloration. These distinct genetic alterations suggest that melanism has arisen independently at least four times within the cat family.
Hypothesized Advantages
Several hypotheses attempt to explain the persistence and prevalence of melanism in felines. One theory suggests a selective advantage for ambush predators, as darker coats provide better camouflage in low-light environments. Other theories propose that genes associated with melanism may confer resistance to viral infections or offer a thermal advantage, as black fur absorbs more solar radiation, aiding warmth in cooler, high-altitude habitats.
Avian Melanism
Rare Sightings
While less common than in some other animal groups, melanism can occur in birds, sometimes leading to striking observations. For instance, an exceptionally rare black flamingo was sighted on the Mediterranean island of Cyprus in April 2015, captivating ornithologists and the public alike.
Domesticated Breeds
In domesticated birds, melanism is intentionally bred for in certain varieties. The Silkie and Ayam Cemani chicken breeds are prime examples. Ayam Cemani, an uncommon breed from Indonesia, possesses a dominant gene causing fibromelanosis, resulting in the chicken being entirely black, including its feathers, beak, and even internal organs.
Game Bird Release
Melanistic common pheasants are specifically bred and released as game birds. Their distinct dark plumage makes them a unique target for hunters, demonstrating a human-influenced aspect of melanism in avian populations.
Urban Adaptations
Melanism is quite prevalent in feral rock doves, particularly in areas where the species is abundant. The degree of pigmentation can vary significantly, ranging from slightly darker wings to an almost entirely black plumage. This suggests an adaptive response to urban environments, possibly related to camouflage against sooty buildings or other environmental factors.
Amphibian Melanism
The Alpine Salamander
A notable example of melanism in amphibians is the alpine salamander (Salamandra atra). One of its subspecies, S. atra atra, is entirely black, presenting a striking contrast to other salamander species and even other subspecies within its own genus.
Pigmentary System
The dark pigmentation in these amphibians originates from specialized cells called melanophores, which are responsible for producing the compound melanin. In contrast, the yellow spots observed in some other subspecies are attributed to pigment-producing cells known as xanthophores.
Phenotype Variation
The alpine salamander exhibits varying levels of melanin pigmentation across its four subspecies. Interestingly, the fully black phenotypes of S. atra atra do not develop xanthophores, indicating a complete genetic suppression of yellow coloration in these individuals.
Evolutionary Trajectory
DNA history studies suggest that the ancestral alpine salamander phenotype likely featured a black base with some yellow spots. This implies that the fully black coloration observed in S. atra atra evolved over time, indicating a strong selective pressure for this melanistic trait across many generations. Both fully melanistic and spotted alpine salamanders are known to produce a toxin from their skin, suggesting this defense mechanism is independent of their specific color pattern.
Human Pigmentation
Not True Melanism
It is important to clarify that true melanism, defined as a mutation resulting in completely dark skin across the entire body, does not exist in humans. Human skin pigmentation is a complex polygenic trait, not a singular melanistic mutation.
Melanin's Protective Role
In humans, the amount of melanin is determined by multiple dominant alleles (AABBCC), with different ethnicities exhibiting varying quantities. Melanin is the primary determinant of skin pigmentation and plays a crucial role in protecting the body from harmful ultraviolet (UV) radiation from the sun.
Latitudinal Adaptation
The variation in human skin color is an evolutionary adaptation linked to geographic location. Lighter skin, characterized by less melanin, is an adaptation related to the prehistoric migration of humans away from equatorial regions. At higher latitudes, reduced sunlight exposure necessitates less melanin to allow for sufficient synthesis of vitamin D in the skin.
Ethnic Diversity
While individuals from regions such as Sub-Saharan Africa, South Asia, Southeast Asia, Australia, and Melanesia may possess very dark skin, this is a natural and normal variation in human pigmentation, not melanism in the biological sense of a congenital excess due to a specific mutation.
Related Conditions
Peutz-Jeghers Syndrome
Peutz-Jeghers syndrome is a rare genetic disorder characterized by the development of macules with hyperpigmentation on the lips and oral mucosa, a condition often referred to as melanosis. In addition to these distinctive pigmented lesions, individuals with this syndrome also develop benign polyps throughout their gastrointestinal tract.
Sociopolitical Interpretations
Digital Discourse
The term "melanism" has found usage outside of its scientific context, appearing in discussions on platforms such as Usenet, internet forums, and blogs. In these digital spaces, its meaning has sometimes been reinterpreted or applied metaphorically.
Afrocentrist Perspectives
Within certain Afrocentrist social movements, the term "melanism" has been employed to articulate the belief that dark-skinned humans represent the original population from which all other human skin colors originated. This perspective was notably promoted by figures such as Frances Cress Welsing, reflecting a specific sociopolitical interpretation of human pigmentation.
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References
References
- King, R.C., Stansfield, W.D., Mulligan, P.K. (2006). A Dictionary of Genetics, 7th ed., Oxford University Press
- Majerus, M. E. (2009). Industrial melanism in the peppered moth, Biston betularia: an excellent teaching example of Darwinian evolution in action. Evolution: Education and Outreach, 2(1), 63â74.
- McIntyre, N. E. (2000). Ecology of urban arthropods: a review and a call to action. Annals of the Entomological Society of America, 93(4), 825â835.
- Cook, L. M., Saccheri, I. J., 2013. The peppered moth and industrial melanism: evolution of a natural selection case study. Journal of Heredity 110:207â12
- Grant, B. S., Wiseman L. L., 2002. Recent history of melanism in American peppered moths. Journal of Heredity 93:86-90.
- Brakefield, P. M., Liebert, T. G., 2000. Evolutionary dynamics of declining melanism in the peppered moth in the Netherlands. Proceedings of the Royal Society of London Biology 267:1953â1957.
- Grant, B. S., Cook, A. D., Clarke, C. A., & Owen, D. F. (1998). Geographic and temporal variation in the incidence of melanism in peppered moth populations in America and Britain. Journal of Heredity, 89(5), 465â471.
- Mikkola, K., & Rantala, M. J. (2010). Immune defence, a possible nonvisual selective factor behind the industrial melanism of moths (Lepidoptera). Biological Journal of the Linnean Society, 99(4), 831â838.
- Mikkola, K., Albrecht, A., 1988. The melanism of Adalia-bipunctata around the Gulf of Finland as an industrial phenomenon (Coleoptera, Coccinellidae). Annales Zoologici Fennici 25:177â85.
- Muggleton, J., Lonsdale, D., Benham, B. R., 1975. Melanism in Adalia-bipunctata L (ColCoccinellidae) and its relationship to atmospheric pollution. Journal of Applied Ecology 2:451â464.
- De Jong, P. W., Verhoog, M. D., Brakefield, P. M., 1992. Sperm competition and melanic polymorphism in the 2-spot ladybird, Adalla bipunctata (Coleoptera, Coccinellidae). Journal of Heredity 70:172â178.
- Ulmer, F. A. (1941) Melanism in the Felidae, with special reference to the Genus Lynx. Journal of Mammalogy 22 (3): 285â288.
- Seidensticker, J., Lumpkin, S. (2006). Smithsonian Q & A: the ultimate question and answer book. Cats. Collins, New York
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