Melanosomes
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A melanosome is an
organelle In cell biology, an organelle is a specialized subunit, usually within a cell, that has a specific function. The name ''organelle'' comes from the idea that these structures are parts of cells, as organs are to the body, hence ''organelle,'' the ...
found in animal cells and is the site for synthesis, storage and transport of
melanin Melanin (; from el, μέλας, melas, black, dark) is a broad term for a group of natural pigments found in most organisms. Eumelanin is produced through a multistage chemical process known as melanogenesis, where the oxidation of the amino ...
, the most common light-absorbing pigment found in the animal kingdom. Melanosomes are responsible for color and
photoprotection Photoprotection is the biochemical process that helps organisms cope with molecular damage caused by sunlight. Plants and other oxygenic phototrophs have developed a suite of photoprotective mechanisms to prevent photoinhibition and oxidative stre ...
in animal cells and tissues. Melanosomes are synthesised in the skin in
melanocyte Melanocytes are melanin-producing neural crest-derived cells located in the bottom layer (the stratum basale) of the skin's epidermis, the middle layer of the eye (the uvea), the inner ear, vaginal epithelium, meninges, bones, and heart. ...
cells, as well as the eye in
choroidal The choroid, also known as the choroidea or choroid coat, is a part of the uvea, the vascular layer of the eye, and contains connective tissues, and lies between the retina and the sclera. The human choroid is thickest at the far extreme rear ...
melanocytes and retinal pigment epithelial (RPE) cells. In lower vertebrates, they are found in melanophores or
chromatophore Chromatophores are cells that produce color, of which many types are Biological pigment, pigment-containing cells, or groups of cells, found in a wide range of animals including amphibians, fish, reptiles, crustaceans and cephalopods. Mammals and ...
s.


Structure

Melanosomes are relatively large organelles, measuring up to 500 nm in diameter. They are bound by a bilipid membrane and are, in general, rounded, sausage-like, or cigar-like in shape. The shape is constant for a given species and cell type. They have a characteristic
ultrastructure Ultrastructure (or ultra-structure) is the architecture of cells and biomaterials that is visible at higher magnifications than found on a standard optical light microscope. This traditionally meant the resolution and magnification range of a co ...
on
electron microscopy An electron microscope is a microscope that uses a beam of accelerated electrons as a source of illumination. As the wavelength of an electron can be up to 100,000 times shorter than that of visible light photons, electron microscopes have a hi ...
, which varies according to the maturity of the melanosome, and for research purposes a numeric staging system is sometimes used.


Synthesis of melanin

Melanosomes are dependent for their pigment on certain enzymes, especially
tyrosinase Tyrosinase is an oxidase that is the rate-limiting enzyme for controlling the production of melanin. The enzyme is mainly involved in two distinct reactions of melanin synthesis otherwise known as the Raper Mason pathway. Firstly, the hydroxylat ...
, that synthesise the large polymers of
melanin Melanin (; from el, μέλας, melas, black, dark) is a broad term for a group of natural pigments found in most organisms. Eumelanin is produced through a multistage chemical process known as melanogenesis, where the oxidation of the amino ...
within the cell. Before it generates sufficient pigment to be seen on
light microscopy Microscopy is the technical field of using microscopes to view objects and areas of objects that cannot be seen with the naked eye (objects that are not within the resolution range of the normal eye). There are three well-known branches of micr ...
it is known as a pre-melanosome. Dysfunction or absence of the melanin-synthesising enzymes (in conditions such as
Chédiak–Higashi syndrome Chédiak–Higashi syndrome (CHS) is a rare autosomal recessive disorder that arises from a mutation of a lysosomal trafficking regulator protein, which leads to a decrease in phagocytosis. The decrease in phagocytosis results in recurrent pyogeni ...
) leads to various patterns of
albinism Albinism is the congenital absence of melanin in an animal or plant resulting in white hair, feathers, scales and skin and pink or blue eyes. Individuals with the condition are referred to as albino. Varied use and interpretation of the term ...
.


Pseudopodia and tanning

In some melanocytes, the melanosomes remain static within the cell. In others the cell can extend its surface lengthwise as temporary projections known as
pseudopodia A pseudopod or pseudopodium (plural: pseudopods or pseudopodia) is a temporary arm-like projection of a eukaryotic cell membrane that is emerged in the direction of movement. Filled with cytoplasm, pseudopodia primarily consist of actin filament ...
, which carry melanosomes away from the center of the cell, thereby increasing the cell's effectiveness in absorbing light. The pseudopodial process (aka the ''tanning'' process) happens slowly in dermal melanocytes in response to
ultraviolet light Ultraviolet (UV) is a form of electromagnetic radiation with wavelength from 10 nm (with a corresponding frequency around 30  PHz) to 400 nm (750  THz), shorter than that of visible light, but longer than X-rays. UV radiation i ...
and to production of new melanosomes and increased ''donation'' of melanosomes to adjacent
keratinocyte Keratinocytes are the primary type of Cell (biology), cell found in the epidermis (skin), epidermis, the outermost layer of the skin. In humans, they constitute 90% of epidermal skin cells. Basal cells in the stratum basale, basal layer (''str ...
s, which are typical skin surface cells. Donation occurs when some keratinocytes engulf the end of the melanocyte pseudopodia, which contain many melanosomes. Cytoplasmic
dynein Dyneins are a family of cytoskeletal motor proteins that move along microtubules in cells. They convert the chemical energy stored in ATP to mechanical work. Dynein transports various cellular cargos, provides forces and displacements importa ...
will carry the vesicles containing the melanin to the center of the cell, which causes melanosomes to sequester the keratinocyte's nucleus, providing optimal protection from UV rays. These changes are responsible for tanning of human skin after exposure to UV light or sunlight.


In animals

In many species of
fish Fish are aquatic, craniate, gill-bearing animals that lack limbs with digits. Included in this definition are the living hagfish, lampreys, and cartilaginous and bony fish as well as various extinct related groups. Approximately 95% of li ...
,
amphibian Amphibians are tetrapod, four-limbed and ectothermic vertebrates of the Class (biology), class Amphibia. All living amphibians belong to the group Lissamphibia. They inhabit a wide variety of habitats, with most species living within terres ...
s,
crustacean Crustaceans (Crustacea, ) form a large, diverse arthropod taxon which includes such animals as decapods, seed shrimp, branchiopods, fish lice, krill, remipedes, isopods, barnacles, copepods, amphipods and mantis shrimp. The crustacean group ...
s, and
reptile Reptiles, as most commonly defined are the animals in the class Reptilia ( ), a paraphyletic grouping comprising all sauropsids except birds. Living reptiles comprise turtles, crocodilians, squamates (lizards and snakes) and rhynchocephalians ( ...
s, melanosomes can be highly mobile within the cell in response to hormonal (or sometimes neural) control, which leads to visible changes in colour that are used for behavioural signaling or
photoprotection Photoprotection is the biochemical process that helps organisms cope with molecular damage caused by sunlight. Plants and other oxygenic phototrophs have developed a suite of photoprotective mechanisms to prevent photoinhibition and oxidative stre ...
. Melanosomes found in certain fish species contain
pigments A pigment is a colored material that is completely or nearly insoluble in water. In contrast, dyes are typically soluble, at least at some stage in their use. Generally dyes are often organic compounds whereas pigments are often inorganic compou ...
that control the color of the fish's
scales Scale or scales may refer to: Mathematics * Scale (descriptive set theory), an object defined on a set of points * Scale (ratio), the ratio of a linear dimension of a model to the corresponding dimension of the original * Scale factor, a number w ...
.
Molecular motors Molecular motors are natural (biological) or artificial molecular machines that are the essential agents of movement in living organisms. In general terms, a motor is a device that consumes energy in one form and converts it into motion or mecha ...
, when signaled, will either carry melanosomes containing pigments out to the periphery of the cell, or concentrate them at the center. The motor protein
dynein Dyneins are a family of cytoskeletal motor proteins that move along microtubules in cells. They convert the chemical energy stored in ATP to mechanical work. Dynein transports various cellular cargos, provides forces and displacements importa ...
is responsible for concentrating the melanosomes toward the center of the cell, or the "minus end" of
microtubule Microtubules are polymers of tubulin that form part of the cytoskeleton and provide structure and shape to eukaryotic cells. Microtubules can be as long as 50 micrometres, as wide as 23 to 27  nm and have an inner diameter between 11 an ...
s. Conversely, the protein
kinesin A kinesin is a protein belonging to a class of motor proteins found in eukaryotic cells. Kinesins move along microtubule (MT) filaments and are powered by the hydrolysis of adenosine triphosphate (ATP) (thus kinesins are ATPases, a type of enzy ...
is responsible for dispersing the melanosomes to the periphery of the cell, and are ''plus end'' directed motors. Because the plus ends of microtubules are oriented towards the periphery, kinesin will carry melanosomes to the periphery. Dispersing melanosomes to the periphery causes the cell to appear darker; concentrating melanosomes towards the center will cause the cell to appear lighter color. This is how a
photoprotective Photoprotection is the biochemical process that helps organisms cope with molecular damage caused by sunlight. Plants and other oxygenic phototrophs have developed a suite of photoprotective mechanisms to prevent photoinhibition and oxidative st ...
system works for the fish on a molecular level. Recently, melanosomes were found in spiders as well. The beautiful and rapid colour changes seen in many
cephalopod A cephalopod is any member of the molluscan class Cephalopoda (Greek plural , ; "head-feet") such as a squid, octopus, cuttlefish, or nautilus. These exclusively marine animals are characterized by bilateral body symmetry, a prominent head ...
s such as
octopus An octopus ( : octopuses or octopodes, see below for variants) is a soft-bodied, eight- limbed mollusc of the order Octopoda (, ). The order consists of some 300 species and is grouped within the class Cephalopoda with squids, cuttle ...
es and
squid True squid are molluscs with an elongated soft body, large eyes, eight arms, and two tentacles in the superorder Decapodiformes, though many other molluscs within the broader Neocoleoidea are also called squid despite not strictly fitting t ...
, are based on a different system, the
chromatophore Chromatophores are cells that produce color, of which many types are Biological pigment, pigment-containing cells, or groups of cells, found in a wide range of animals including amphibians, fish, reptiles, crustaceans and cephalopods. Mammals and ...
organ.


In fossils

Recent (2008) discoveries by Xu Xing, a Chinese
paleontologist Paleontology (), also spelled palaeontology or palæontology, is the scientific study of life that existed prior to, and sometimes including, the start of the Holocene epoch (roughly 11,700 years before present). It includes the study of fossi ...
, include
fossil A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved ...
ized feathers in rock formations dating from the
Jurassic period The Jurassic ( ) is a geologic period and stratigraphic system that spanned from the end of the Triassic Period million years ago (Mya) to the beginning of the Cretaceous Period, approximately Mya. The Jurassic constitutes the middle period of ...
(200 to 150 million years ago) to the late
Paleogene The Paleogene ( ; British English, also spelled Palaeogene or Palæogene; informally Lower Tertiary or Early Tertiary) is a geologic period, geologic period and system that spans 43 million years from the end of the Cretaceous Period million yea ...
and
Neogene period The Neogene ( ), informally Upper Tertiary or Late Tertiary, is a geologic period and system that spans 20.45 million years from the end of the Paleogene Period million years ago ( Mya) to the beginning of the present Quaternary Period Mya. ...
s (66 to 2 million years ago). The feathers contain preserved residues of carbon that were previously thought to be traces of bacteria that decomposed feather tissues; however these (residues) are in fact microscopic organic imprints of fossilized melanosomes. Some of these structures still maintain an
iridescent Iridescence (also known as goniochromism) is the phenomenon of certain surfaces that appear to gradually change color as the angle of view or the angle of illumination changes. Examples of iridescence include soap bubbles, feathers, butterfl ...
color typical of feather and fur tissues. It is conjectured that these microscopic structures could be further studied to reveal the original colors and textures of softer tissues in fossils. "The discovery of ultra-structural detail in feather fossils opens up remarkable possibilities for the investigation of other features in soft-bodied fossils, like fur and even internal organs," said
Derek Briggs Derek Ernest Gilmor Briggs (born 10 January 1950) is an Irish palaeontologist and taphonomist based at Yale University. Briggs is one of three palaeontologists, along with Harry Blackmore Whittington and Simon Conway Morris, who were key in th ...
of the
Yale University Yale University is a private research university in New Haven, Connecticut. Established in 1701 as the Collegiate School, it is the third-oldest institution of higher education in the United States and among the most prestigious in the wo ...
study team. Melanosomes were used to discover the true colors of fossil ''
Anchiornis huxleyi ''Anchiornis'' is a genus of small, four-winged paravian dinosaurs, with only one known species, the type species ''Anchiornis huxleyi'', named for its similarity to modern birds. The Latin name ''Anchiornis'' derives from a Greek word meaning " ...
'' by a collaborative team including members from the
Beijing Museum of Natural History The Beijing Museum of Natural History (BMNH; ) is located at 126 Tian Qiao Nan Street, Dongcheng District, Beijing, 100050, and is the most popular natural history museum in China.Peking University Peking University (PKU; ) is a public research university in Beijing, China. The university is funded by the Ministry of Education. Peking University was established as the Imperial University of Peking in 1898 when it received its royal charter ...
,
Yale University Yale University is a private research university in New Haven, Connecticut. Established in 1701 as the Collegiate School, it is the third-oldest institution of higher education in the United States and among the most prestigious in the wo ...
, the
Peabody Museum of Natural History The Peabody Museum of Natural History at Yale University is among the oldest, largest, and most prolific university natural history museums in the world. It was founded by the philanthropist George Peabody in 1866 at the behest of his nephew Othn ...
, the
University of Akron The University of Akron is a public research university in Akron, Ohio. It is part of the University System of Ohio. As a STEM-focused institution, it focuses on industries such as polymers, advanced materials, and engineering. It is classified ...
, and the
University of Texas at Austin The University of Texas at Austin (UT Austin, UT, or Texas) is a public research university in Austin, Texas. It was founded in 1883 and is the oldest institution in the University of Texas System. With 40,916 undergraduate students, 11,075 ...
. Melanosomes have also been found in fossils from ''Tupandactylus'' cf. ''imperator'' pteurosaurs in the Lower Cretaceous Crato Formation, in the
Araripe Basin The Araripe Basin () is a rift basin covering about ,Neto et al., 2013, p.1 in Ceará, Piauí and Pernambuco states of northeastern Brazil. It is bounded by the Patos and Pernambuco lineaments, and is situated east of the Parnaíba Basin, southwes ...
, in
Brazil Brazil ( pt, Brasil; ), officially the Federative Republic of Brazil (Portuguese: ), is the largest country in both South America and Latin America. At and with over 217 million people, Brazil is the world's fifth-largest country by area ...
.


Templating

Melanosomes are believed to template melanin polymerization by way of
amyloid Amyloids are aggregates of proteins characterised by a Fibril, fibrillar morphology of 7–13 Nanometer, nm in diameter, a beta sheet (β-sheet) Secondary structure of proteins, secondary structure (known as cross-β) and ability to be Staining, ...
ogenesis of the protein Pmel17, which is present in abundant quantities in melanosomes.


References


External links

* - "Integument: pigmented skin" {{organelles Articles containing video clips Organelles Skin anatomy