Rapid color changes of amphibians are mediated by three types of dermal chromatophores, xanthophores, iridophores, and melanophores, which comprise a morphologically and physiologically distinct structure, the dermal chromatophore unit. Xanthophores, the outermost element, are located immediately below the basal lamella. Iridophores, containing light-reflecting organelles, are found just beneath the xanthophores. Under each iridophore is found a melanophore from which processes extend upward around the iridophore. Finger-like structures project from these processes and occupy fixed spaces between the xanthophores and iridophores. When a frog darkens, melanosomes move upward from the body of the melanophore to fill the fingers which then obscure the overlying iridophore. Rapid blanching is accomplished by the evacuation of melanosomes from these fingers. Pale coloration ranging from tan to green is provided by the overlying xanthophores and iridophores. Details of chromatophore structure are presented, and the nature of the intimate contact between the chromatophore types is discussed.
The fundamentally diverse vertebrate pigment cells, melanophores, xanthophores, and iridophores, contain pigmentary organelles known, respectively, as melanosomes, pterinosomes, and reflecting platelets. Their pigments are mealanins pteridines, and purines. Mosaic pigment cells containing more than one type of organelle have been observed and mosaic organelles containing more than one type of pigment have been discovered. It is proposed that the various pigment cells are derived from a stem cell that contains a primordial organelle of endoplasmic reticular origin. This primordial organelle can differentiate into any of the known pigmentary organelles.
Summary Although the various vertebrate classes, from fishes to mammals are each distinctive, they possess many common features making it important to understand their comparative biology. One general feature that has long commanded interest is the integumental pigmentary system. Thus, much is known about particular pigment cells; however, the basis for some specific colors, such as blue, has escaped the scrutiny of the comparative approach. Regardless of Class, blue is almost always a structural color based upon incoherent or coherent scatter of blue wavelengths from the animal surface. The source of scatter may be intracellular or extra‐cellular. A main intracellular scatterer is the surface of reflecting platelets of iridophores of lower vertebrates. Extra‐cellular scatter is widespread and thought to occur from ordered dermal collagen arrays in primitive fishes, birds and mammals including humans. Among birds, feather structures provide major means for extra‐cellular light scatter. There is only one known example of blue color deriving from a blue pigment found within a pigment cell. For amphibians, reptiles and birds, the scatter of blue wavelengths, together with the presence of yellow pigmentation, is fundamental for the expression of green coloration.
Primary spermatogonia have highly lobate nuclei and can be distinguished as pale and dark types on the basis of nuclear and cytoplasmic features. Nuclei of secondary spermatogonia are also lobate. Primary spermatocytes have spherical nuclei and display synaptinemal complexes in late zygotene-pachytene. Spermatocytes are connected by intercellular bridges, which persist through spermiogenesis. During spermiogenesis no acrosomal granule is formed. The acrosomal vesicle is large and forms in the apical part of the cell. A helical system of perinuclear microtubules accompanies the phase of nuclear elongation. Microtubules disappear in late spermatids and there forms a compact bundle of filaments which extends into the subacrosomal area. These filaments probably derive from the breakdown of the microtubules. A mitochondrial sleeve is formed around the proximal portion of the tail and much of it is cast off in the mature spermatid. The tail is composed of a spirally coiled contractile element and a stiff fibrous axial rod connected together by an undulating membrane. The axial rod and the axoneme-associated rodlet derive from a dense, juxtacentriolar fibrous mass. Sertoli cells surrounding the spermatogonial and spermatocyte cysts are slender and have oblong nuclei. In contrast, those associated with spermatids are columnar and have deeply indented nuclei. They possess many Golgi complexes, elongated mitochondria, cisternae of smooth endoplasmic reticulum, lysosome-like bodies, masses of glycogen particles, few lipid droplets, and an array of microtubules running longitudinally around the elongating spermatid nuclei. Desmosomes are formed between adjacent Sertoli cells.
Homeotherms are generally considered to lack classical active dermal pigment cells (chromatophores) in their integument, attributable to the development of an outer covering coat of hair or feathers. However, bright colored dermal pigment cells, comparable to chromatophores of lower vertebrates, are found in the irides of many birds. We propose that, because of its exposed location, the iris is an area in which color from pigment cells has sustained a selective advantage and appears to have evolved independently of the general integument. In birds, the iris appears to have retained the potential for the complete expression of all dermal chromatophore types. Differences in cell morphology and the presence of unusual pigments in birds are suggested to be the result of evolutionary changes that followed the divergence of birds from reptiles. By comparison, mammals appear to have lost the potential for producing iridophores, xanthophores, or erythrophores comparable to those of lower vertebrates, even though some species possess brightly colored irides. It is proposed that at least one species of mammal (the domestic cat) has recruited a novel iridial reflecting pigment organelle originally developed in the choroidal tapetum lucidum. The potential presence of classical chromatophores in mammals remains open, as few species with bright irides have been examined.
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