Melanosomes: Biogenesis, Properties, and Evolution of an Ancient Organelle.

Melanosomes are organelles that produce and store melanin, a widespread biological pigment with a unique suite of properties including high refractive index, semiconducting capabilities, material stiffness, and high fossilization potential. They are involved in numerous critical biological functions in organisms across the tree of life. Individual components such as melanin chemistry and melanosome development have recently been addressed, but a broad synthesis is needed. Here, we review the hierarchical structure, development, functions, and evolution of melanosomes. We highlight variation in melanin chemistry and melanosome morphology and how these may relate to function. For example, we review what is known of the chemical differences between different melanin types (eumelanin, pheomelanin, allomelanin) and whether/how melanosome morphology relates to chemistry and color. We integrate the distribution of melanin across living organisms with what is known from the fossil record and produce hypotheses on its evolution. We suggest that melanin was present in life forms early in evolutionary history and that melanosomes evolved at the origin of organelles. Throughout, we discuss the (sometimes gaping) holes in our knowledge and suggest areas that need particular attention as we move forward in our understanding of these still-mysterious organelles and the materials that they contain.