What Is the Function of Melanin?
Warwick L. Morison, MD
An informal survey of my colleagues revealed that
most dermatologists believe the primary func-
tion of melanin in humans is to act as a sunscreen to
protect us from ultraviolet (UV) radiation in sun-
light; the leading textbooks of dermatology support
this notion. Recently, while reading materials out-
side of our specialty, I was surprised to find that
dermatologists might be alone in the scientific com-
munity in their acceptance of this view. Further-
more, the evidence for melanin having developed to
serve other functions appeared to be as strong or
stronger than the evidence that it is primarily a
sunscreen. Interestingly, a decade or so ago, there
was considerable discussion about this subject
among dermatologists who had an interest in photo-
biology,1,2 but, recently, we appear to have accepted
as dogma that melanin is a sunscreen. Dogma, in any
discipline, should be examined periodically. With
many recent advances in the biology of melanin, it is
probably an appropriate time to again discuss the
primary function of melanin.
Perhaps, before proceeding, I should clearly define
the question: What is the primary function of mela¬
nin that determines its presence in human skin?
Melanin does absorb UV radiation and therefore
does function as a sunscreen, but this is not the issue.
Keratin absorbs UV radiation, but we do not regard
it as a sunscreen in the usual sense. Ears function to
support earrings, but we acquired ears for quite a
different reason. We are concerned with whether
melanin was developed as an adaptation to protect us
from the harmful effects of UV radiation.
WHY IS MELANIN CONSIDERED A SUNSCREEN?
In 1820, Sir Everard Home3 was the first to
propose the concept that melanin is a sunscreen; the
evidence supporting it has not changed very much in
the intervening 165 years. The logic behind the
concept is simple. Prehumans were thought to have
first appeared in a grassland environment in tropical
and subtropical Africa. In this environment, a sun¬
screen would have been necessary to protect them
from the damaging effects of UV radiation. Because
melanin in the skin provided such protection, there
was an evolutionary advantage in being black in this
sun-drenched environment. There are some prob¬
lems with the assumptions on which this argument
rests. Fossil finds are consistent with an origin of
Accepted for publication March 19, 1985.
From the Department of Dermatology, The Johns Hopkins
Medical Institutions, Baltimore.
Reprint requests to the Department of Dermatology, The Johns
Hopkins Medical Institutions, 600 N Wolfe St, Baltimore, MD
21205 (Dr Morison).
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prehumans in Africa, but recent finds46 indicate that
hominids developed in a forest environment, wherein
there would be no need for a sunscreen. Further¬
more, there is no evidence to support or refute the
assumption that prehumans were black.
Two principal observations are used to support the
concept that the main function of melanin is to act as
a sunscreen. First, albinos living in the tropics
develop solar kératoses during childhood and die of
skin cancer in early adult life. Second, transplanted
Europeans have a high incidence of skin cancer in
tropical and subtropical climates. These are facts
but, for several reasons, are irrelevant to the role
of melanin as a sunscreen. First, as Hamil¬
ton7"""91224,says in his thoughtful treatise on pigmen¬
tation, function should not be interpreted on
the basis of genetic anomalies that delete a normal
characteristic or on sudden transplantation of a
population to an entirely new environment. Evolu¬
tion is a slow process involving many interconnected
adaptations and cannot anticipate evolutionary acci¬
dents. Second, for melanin to be an evolutionary
adaptation to prevent the harmful effects of sun¬
light, those effects must interfere with reproduction.
This elementary point was made by noted photobiol-
ogist Harold Blum8 many years ago, when he consid¬
ered and rejected the possibility that sunburn and
skin cancer interfered with the reproductive capacity
of hominids. The study by Okoro' of 1,000 albinos in
Nigeria supports this view. In this extreme situation,
skin cancer does not cause death in persons under 25
years of age. Presumably, because of their short life
spans, hominids would have reproduced long before
this age.
Therefore, we can conclude that hominids proba¬
bly lived in forests and did not need a sunscreen and
that even if they did need a sunscreen, this require¬
ment would not have provided pressure for an evolu¬
tionary adaptation. However, before discarding the
notion that melanin is a sunscreen, we should consid¬
er a few other questions.
IS MELANIN A GOOD SUNSCREEN?
This can be answered by comparing the properties
of melanin with those of the ideal sunscreen.
1. A sunscreen should be substantive and should
not wash off; melanin satisfies this requirement.
2. A sunscreen should efficiently and maximally
absorb in the UV region and should not absorb
visible or infrared radiation, because absorption of
these longer wavelengths will only heat the body.
Melanin absorbs UV radiation, but it also absorbs
visible light efficiently, and its absorption extends
into the near-infrared region. The reason pigmented
 people do not sunbathe is that they are too uncom¬
fortably hot when in direct sunlight, not because
they are already tanned.
3. A sunscreen should be limited mainly to the
stratum corneum, so absorbed radiation can be radi¬
ated back into the environment and not transfer¬
red within the body as heat. Melanin is produced
in the basal layer and is found throughout the epi¬
dermis.
4. A sunscreen should not be involved in photo¬
chemical processes that could be harmful to the
organism. Melanin is a photosensitizer that is capa¬
ble of killing cells10; pheomelanin is mutagenic"; and
cystinyldopa, a precursor of pheomelanin, photore-
acts with DNA (M. Chedekel, PhD, oral communica¬
tion, 1984). Pheomelanin has not been demonstrated
directly in skin, but much circumstantial evidence
indicates that this potentially harmful substance is
present in some individuals. In addition, one study
reported that after exposure to UV radiation, fewer
sunburn cells were present in vitiliginous skin com¬
pared with normal pigmented skin.12 This finding
suggests that melanin photosensitizes, killing epi¬
dermal cells. These observations suggest that we
cannot regard melanin as an innocuous sunscreen.
Therefore, of the four properties that constitute an
ideal sunscreen, melanin exhibits only one.
DOES MELANIN BEHAVE AS A SUNSCREEN?
If melanin is a sunscreen, we would expect (1) a
uniform distribution of melanin across the surface of
the skin, so as to provide protection for all cells in an
even manner and (2) that melanin should function
most efficiently in black skin.
Melanin satisfies neither of these two criteria.
Melanosomes in the keratinocytes of black skin are
single, large, and only partially degraded by the time
they reach the stratum corneum; thus, this layer
contains some melanosomes and some more evenly
distributed melanin dust.13 In contrast, the melano¬
somes in the keratinocytes of fair skin are complete¬
ly degraded, so that only melanin dust is present in
the stratum corneum.13 Although measurements to
quantitate these differences have not been reported,
it seems likely that melanin is a more efficient
sunscreen per unit of melanin in fair skin compared
with black skin. This finding is perhaps the reverse
of what one would expect. We are forced to conclude
that melanin is not only far from an ideal sunscreen
but also that it does not behave as if its prime
function is to be a sunscreen.
WHY DO WE TAN WHEN
EXPOSED TO SUNLIGHT?
Almost everybody believes that tanning is a natu¬
ral defense mechanism to protect the skin from
subsequent exposure to sunlight. However, when we
examine the specificity of the response, analyze the
epidermal response to UV radiation, and remember
our first classes in pathology, it is easy to see that
this concept is probably incorrect. Melanogenesis
occurs in response to various traumas, such as
ionizing radiation, contact dermatitis, and skin abra-
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sion. Do we seriously consider the pigmentation that
follows an abrasion to be a protective device to
prevent another abrasion? Certainly not. Pigmenta¬
tion occurs because melanocytes have been killed and
damaged. In response, the remaining cells multiply
and produce melanin, which is a nonspecific response
to trauma. Similarly, if we examine the epidermal
response to UV radiation, we find that it has all the
hallmarks of a nonspecific hyperplasia in response to
trauma. After exposure to UV radiation, melano¬
cytes and keratinocytes increase in number, leading
to increased production of melanin14 and keratin,15
respectively; both melanocytes and keratinocytes
increase by a factor of two to six times. The epider¬
mis undergoes nonspecific hyperplasia in response to
the damage and death of cells in the same manner
that many other tissues respond to damage. If a
portion of our liver is excised, the remainder hyper¬
trophies. No one can seriously believe this response
by the liver to be a device to protect itself from being
cut again.
It might be argued that the shorter wavelengths of
UV radiation in sunlight are more efficient and
reliable means of activating melanocytes than are
other traumas. This concept is true, but the illusion
of specificity merely reflects the gradation of the
trauma and the optical properties of skin, so that a
certain percentage of the photons of UV radiation
are absorbed by each layer of the skin. In contrast,
an abrasion is not graded but reaches a certain level,
so that it might just remove the first few cell layers
and not affect the basal melanocytes, involve the
basal melanocytes, or remove the whole basal layer
of the epidermis as well as part of the dermis. The
result can be no alteration of pigmentation,
increased pigmentation, or depigmentation. The par¬
allels for UV radiation are UV-C (254-nm) radiation,
which produces insignificant pigmentation, because
it is absorbed by the superficial epidermis; UV-B (280
to 320-nm) radiation, which is efficient in stimulat¬
ing melanocytes; and a blistering dose of sunlight,
which often results in hypopigmentation.
Finally, if we do interpret the epidermal response
to UV radiation to be a protective device, then we
should change our notion that melanin is the sun¬
screen, because keratin appears to be superior to
melanin in this respect. Repeated exposure of vitilig¬
inous skin can give a sun-protective factor (SPF) of
1516 in the absence of melanocytes, but selective
activation of melanocytes by UV-A radiation only
gives an SPF of 2 or 3." If these two observations are
combined, we could expect chronic exposure to UV
radiation to result in an SPF of about 18 due to the
combined action of melanin and keratin as sun¬
screens. An SPF of 18 was indeed reported after four
weeks of irradiation of normal skin.16
THE FUNCTION OF MELANIN
We have reached the point of suggesting that
melanin is not an evolutionary adaptation to protect
humans from sunlight; however, we still have not
determined its function. Several theories have been
advanced, and each is based on one of the properties
 of melanin. Of these theories, two have some logic
and factual support: the "camouflage" hypothesis,
which is based on melanin's property of color, and
the "heat absorption" hypothesis, which is based on
the capacity of melanin to absorb radiation.
Melanin as a Camouflage
If indeed hominids developed in a forest environ¬
ment, some sort of camouflage would be essential.
Melanin could be very effective for this purpose.
Black skin reflects only 16% of visible light, whereas
fair skin reflects 45%.18 Since we see objects by light
reflected from them, an albino would be very easily
seen by predators and prey in the shadows of a
forest. However, before we conclude that melanin
developed as camouflage, it is necessary to establish
that hominids developed in forests. Evidence sup¬
porting the theory that forests were the homes of
prehumans has recently been summarized by Corth."
Perhaps the most important evidence is that recent
fossil finds have placed early hominids in forests and
that the fossil record is now consistent with a
movement of bipedal hominids to open savanna in
relatively recent times.4"6
The physiology and anatomy of vision provides
additional strong evidence that hominids lived in
forests. First, the spectral sensitivity for photopic
vision in humans peaks at about 550 nm, which
closely corresponds with the spectral composition of
light on a forest floor.20 Sunlight entering the top of a
forest canopy is absorbed, reflected, and scattered by
vegetation. Therefore, light on a forest floor is
yellow-green, with a fairly narrow peak at around
550 nm. In contrast, the sunlight spectrum under an
open sky has a flat, broad peak between 400 and 700
nm. Second, color discrimination in humans corre¬
lates well with the spectral composition of light in a
forest. Third, humans have a fovea in the retina,
which is an adaptation to a forest environment. A
fovea provides equal visual sensitivity in vertical and
horizontal planes, which is necessary when predators
and prey are above and around and when there is no
visible horizon.21 The retinal topography of animals
that inhabit grasslands is quite different. These
animals have a visual streak across the retina that
provides maximum visual sensitivity in a horizontal
plane; thus, they have panoramic vision.21 An excel¬
lent example of this difference in retinal topography
is provided by a comparison of the closely related
tree and red kangaroos. The tree kangaroo, whose
habitat is obvious, has a fovea, whereas the red
kangaroo, which lives in open grassland, has a visual
streak.22 Finally, and perhaps most important, the
human eye cannot tolerate exposure to bright sun¬
light. Exposure to bright sunlight for more than a
day results in a 50% to 90% reduction in retinal
sensitivity to the visible component of sunlight.23
This defect can persist for more than a week. Obvi¬
ously, the eyes have it—we are not adapted to living
in sunlight under an open sky.
Several points have been raised against the theory
that melanin was acquired for camouflage in a forest
environment. The first is that in a dense tropical
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forest, there would be little reason for the evolution
of beings with an upright posture, because locomo¬
tion would be restricted by undergrowth and vines.
However, this argument is based on rain forests; the
forests inhabited by hominids could have been more
open, with little undergrowth. Furthermore, an
upright posture is also a disadvantage in grasslands
because of the loss in speed of locomotion. In both
rain forests and grasslands, the advantages must
outweigh the disadvantages. Obviously, the pygmies
of Africa apparently have coped quite well with a
bipedal upright posture in a forest. The second point
is that a uniform black skin is not ideal camouflage
in a forest and that some variability and a brown
color would provide better concealment. These inter¬
esting points presuppose that we know the color of
prehumans; however, we do not. Prehumans might
have been streaked and brown, and present-day
Africans might be uniformly black as a later adapta¬
tion to some other aspect of their environment.
Melanin as a Heat Absorber
Radiant energy from sunlight absorbed by mela¬
nin is converted to heat, which is partially dissipated
into the body. As a consequence, the rise in body
temperature produced by exposure to sunlight is
greater in a pigmented person than in a nonpig-
mented person. The heat load produced by the pres¬
ence of melanin in the skin can be regarded as a
disadvantage, because it decreases performance and
causes discomfort, if a pigmented person is living in
tropical sunlight.24 However, if exposure to sunlight
is taken in moderation, pigmentation can be an
advantage, because it contributes to the maintenance
of normal body temperature and thus conserves
metabolic heat, which comes from food. This concept
forms the basis of the heat absorption theory of the
function of melanin."1*""195)
Prehumans slept without the benefit of fire or
clothing. Although they were presumably living
mainly in the tropical and subtropical regions of
Africa, nighttime temperatures in those areas are
often below 20 °C. The critical environmental tem¬
perature for humans is 27 to 29 °C; at lower temper¬
atures, deep body temperature progressively falls
during the night, and this deficit must be replenished
before the next night. Measurements of the extent of
the decline in body temperature in Bushmen of the
Kalahari desert and Aborigines of Australia25 indi¬
cate that a 2-°C drop in body temperature would not
be unusual. As Hamilton7(ppl"195) proposed, this drop
could represent a deficit of 1,300 calories in a 50-kg
man that would have to be restored by metabolic or
environmental heat. Having melanin to absorb sun¬
light, therefore, could be an advantage, because it
would permit rapid restoration of body temperature
and obviate the need for additional food. The heat
absorption theory might also explain the immediate
darkening of pigment that occurs in pigmented peo¬
ple upon exposure to sunlight, since this phenomenon
would accelerate absorption of heat.
The main argument against the heat absorption
theory is that overheating could be a problem during
 the hottest portion of the day. However, this argu¬
ment implies that prehumans did not have the good
sense, as the inhabitants of the tropics and subtrop-
ics today, to have a siesta in the shade.
Other Functions of Melanin
Melanin pigmentation was once considered a
mechanism to prevent excessive production of vita¬
min D by the skin, but this idea has now been
discounted. It has also been proposed that melanin is
a hormone; however, this concept appears to have
died for lack of support. One intriguing, if not com¬
pelling, hypothesis is that melanin is a threat display
organ.26 According to this hypothesis, certain parts of
the body are developed or modified to act as threat
displays in social relationships—a prominent nose or
chin, a bullneck, and a hairy chest are a few ex¬
amples. Dark colors in certain species are considered
to be threat displays; hence, melanin pigmentation
has been proposed as a threat display organ. A point
raised in favor of this hypothesis is that in almost all
mammals where there is sexual dimorphism in body
color, the male mammal is darker. This concept
might explain why women wore white gloves and
carried parasols but now, in the era of feminism,
vigorously indulge in sunbathing. As I said, it is an
intriguing, if not compelling, hypothesis.
CONCLUSION
The evidence that melanin is an adaptation to
living in a sun-drenched environment does not with¬
stand scientific scrutiny. Melanin certainly absorbs
UV radiation; otherwise, it possesses few properties
to suggest that its prime function is as a sunscreen.
Furthermore, melanin does not appear to behave as a
sunscreen. The formation of melanin is not a specific
response to sunlight exposure but instead forms part
of a nonspecific response of the epidermis to damage
and death of cells. Besides, hominids living in forests
had no need for a sunscreen. What they did need was
camouflage and a device to keep their bodies warm.
Logic and the available evidence support the theory
that melanin pigmentation is an evolutionary adap¬
tation for concealment and heat absorption.
Consideration of the function of melanin is not
just idle philosophical musing; it has important
ramifications for our health and environment that
should be of concern to dermatologists. By regarding
melanin as a sunscreen, we unwittingly agree that
sunbathing and use of suntan parlors is providing
natural protection. In the "natural" society we live
in, we can expect a natural sunscreen to be perceived
as the best. Tanning is merely evidence that the skin
has been unnaturally damaged and will, in the near
future, become unnaturally aged. Furthermore, by
accepting melanin as an adaptation acquired by our
ancestors to protect them from sunlight, we imply
that sunlight under an open sky is natural light for
humans. Such a belief leads to prophecies of the
grave biologic dangers we may encounter if environ¬
mental lighting does not mimic sunlight.27 Of course,
we quickly reject the idea of sunlight under an open
sky being natural as we walk outside without sun-
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glasses on a bright sunny day and start squinting
and screwing up our faces in discomfort.
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