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In the reddish-violet parts of the skin of the diadema pseudochromis Pseudochromis diadema, we found novel dichromatic chromatophores with a reddish pigment and reflecting platelets. We named these novel cells 'erythro-iridophores'. In standard physiological solution, erythro-iridophores displayed two hues, red and dark violet when viewed with an optical microscope under ordinary transmission light and epi-illumination optics, respectively. Under transmission electron microscopy, however, we observed no typical red chromatosomes, i.e., erythrosomes, in the cytoplasm. High-performance thin-layer chromatography (HPTLC) analysis of the pigment eluted from the erythro-iridophores indicated that carotenoid is the main pigment generating the reddish color. Furthermore, when the irrigating medium was a K(+)-rich saline solution, the color reflected from the erythro-iridophores changed from dark violet to sky blue, but the red coloration remained. The motile activities of the erythro-iridophores may participate in the changes in the reddish-violet shades of the pseudochromis fish., (© 2011 John Wiley & Sons A/S.)

An increase in the concentration of Ca(2+) ions in the external medium ([Ca(2+)](o)) induced pigment aggregation in melanophores of three species of freshwater teleosts examined. Denervated melanophores were refractory to elevations of [Ca(2+)](o). The pigment-aggregating action was inhibited by the sympathetic blocking agents, phentolamine, prazosin and yohimbine. Bretylium, an agent known to block the release of the neurotransmitter, interfered with the response effectively. Ca(2+) blockers, such as Mn(2+), verapamil and gallopamil, also inhibited the response, possibly by inhibiting Ca(2+) entry into the presynaptic elements of melanosome-aggregating fibers. The conclusion is that the increase in [Ca(2+)](o) may induce membrane depolarization of presynaptic nervous elements around the melanophores, which open the voltage-dependent Ca(2+) channels there. The liberation of adrenergic neurotransmitter follows, which induces the aggregation of pigment in melanophores.

Chromatophores, including melanophores, xanthophores, erythrophores, leucophores and iridophores, are responsible for the revelation of integumentary coloration in fish. Recently, blue chromatophores, also called cyanophores, were added to the list of chromatophores. Many of them are also known to possess cellular motility, by which fish are able to change their integumentary hues and patterns, thus enabling them to execute remarkable or subtle chromatic adaptation to environmental hues and patterns, and to cope with various ethological encounters. Such physiological color changes are indeed crucial for them to survive, either by protecting themselves from predators or by increasing their chances of feeding. Sometimes, they are also useful in courtship and mutual communications among individuals of the same species, leading to an increased rate of species survival. Such strategies are realized by complex mechanisms existing in the endocrine and/or nervous systems. Current studies further indicate that some paracrine factors such as endothelins (ETs) are involved in these processes. In this review, the elaborate mechanisms regulating chromatophores in these lovely aquatic animals are described.

Like melanophores of many teleosts, those of the dark chub, Zacco temmincki, and the common minnow, Z. platypus (Cyprinidae, Cypriniformes) responded to norepinephrine (NE) by the aggregation of pigment. It was further found that some melanophores were responsive to acetylcholine (ACh) in the same way. The response to NE was blocked by an alpha-adrenergic blocker, phentolamine, whereas the response to ACh was not. By contrast, two muscarinic cholinoceptor antagonists, namely, atropine and scopolamine, effectively blocked the action of ACh. The pigment aggregation due to the liberated sympathetic neurotransmitter was blocked by phentolamine but not by cholinergic blockers. These results suggest that, although the melanophores of these species are controlled in an orthodox manner by the sympathetic nervous system, some of them possess extra muscarinic cholinoceptors that also mediate the aggregation of pigment. The present report is the first to describe the presence of cholinoceptors on the chromatophores in species of fish other than those that belong to the order Siluriformes. The evolutionary implications of these findings are discussed.

The red abdominal skin of the neon tetra Paracheirodon innesi and the cardinal tetra P. axelrodi was found to blanch at night or in the dark. Melatonin added to the bathing medium caused blanching of the red skin. Microscopic observations of the erythrophores indicated that the erythrosomes aggregated in response to norepinephrine, melanin-concentrating hormone (MCH), and melatonin. Of these compounds, melatonin was the most effective. By contrast, many erythrophores were refractory to MCH. Alpha-melanophore-stimulating hormone, isoproterenol, adenosine, and ATP each caused dispersal of the pigment to some extent. Isoproterenol dispersed the pigment only when an alpha-adrenergic blocker, tolazoline, was present. It appears that the change in color of the abdominal skin is primarily due to increased secretion during the night of the pineal hormone melatonin, while other hormonal and nervous factors may modify the distribution of the pigment in the erythrophores.

Splenic white pulp of the Aleutian skate (Bathyraja aleutica), an elasmobranch, was investigated using light and transmission electron microscopy. The major cellular constituent was plasma cells, of the typical Marshalko type, characterized by well developed rough-endoplasmic reticulum and a Golgi complex. The morphology of the rough-endoplasmic reticulum was variable, being lamellar in some cells and spherical in others. Plasma cells with distended cisternae of rough-endoplasmic reticulum and cells with Russel bodies were often observed. Only a small number of lymphocytes were encountered. These findings indicate that the splenic white pulp is the major site for immunoglobulin production in this fish, thus confirming our previous immunocytochemical observation. Globules presumably containing immunoglobulin were found consistently associated with the Golgi complex. The secretion mechanism of immunoglobulin by plasma cells is discussed in connection with the globules.

The in vitro effects of synthetic salmon melanin-concentrating hormone (MCH) on chromatophores of four teleost species were studied. In the erythrophores of the platyfish (Xiphophorus maculatus) and the swordtail (Xiphophorus helleri), and in the xanthophores and amelanotic melanophores of the medaka (Oryzias latipes), pigment aggregation took place in response to MCH even in the absence of Ca2+. In contrast to this, the leucophores of the medaka responded to MCH by the pigment dispersion but only when Ca2+ was present. The motile iridophores of the blue damselfish (Chrysiptera cyanea), which play a predominant role in coloration and its changes, were not affected by the hormone. Pharmacological studies employing various blocking agents suggest that the pigment-aggregating action of MCH is probably mediated through specific receptors possessed by the erythrophores, xanthophores, or amelanotic melanophores, while the pigment-dispersing action on the leucophores might be revealed through the receptors for melanophore-stimulating hormone (MSH).

Elevation of [K+]0 elicited pigment aggregation within melanophores, and, at the same time, release of both norepinephrine (NE) and purines from split fin preparations of the tilapia, Sarotherodon niloticus, doubly labeled with [14C]NE and [3H]adenosine. This concomitant release increased with the increase in [K+]0 up to 50 mM, although a further increase in [K+]0 caused a progressive decrease in the outputs. The release of either NE or purines was significantly reduced by bretylium. It was concluded that, upon stimulation of the nerve, both the true transmitter, NE, and the co-transmitter, adenine nucleotides, are liberated for an exquisite control of chromatophore movements.

The mechanism regulating the movements of the unique motile iridophores of the blue damselfish, Chyrsiptera cyanea, was studied. The reaction in which the cells become reflective to light rays of longer wavelength, i.e. from the near u.v. region to the green region, was designated as the "coloring response", while the reverse process was labeled the "clearing response". Both nervous stimuli and adrenergic agonists gave rise to the coloring response, which could be antagonized by alpha adrenolytic agents. The clearing response was accelerated by adenosine and inhibited by theophylline. None of the hormonal substances tested had any effect on the motile response of the cells. It was concluded that the motile iridophores are solely under the control of the sympathetic adrenergic system, and that the co-transmitter, adenosine, may function to antagonize quickly the true transmitter-induced colored state of the cells.

Both acetylcholine and catecholamines showed melanin-aggregating action within melanophores on an isolated bony plate of the mailed catfish Corydoras paleatus. Chromatic nervous stimulation either by an electrical field or by an elevation of [K+]0 brought about melanosome aggregation. Alpha adrenolytic agents antagonized the melanin-aggregating effects either of catecholamines or of nervous stimuli. Muscarinic cholinolytics interfered with the action of acetylcholine, but did not have any effect on the responses to nervous stimuli. In addition to the alpha adrenoceptors which participate in sympathetic-melanophore transmission, muscarinic cholinoceptors of unknown functional significance, which also mediate melanosome aggregation in the cell, exist in Corydoras melanophores.

1. Synthetic alpha-MSH was tested for its action on dermal and epidermal melanophores of the siluroid catfish. Parasilurus asotus. 2. MSH did not produce melanosome aggregation in these melanophores. 3. It was, however, very effective in bringing about pigment dispersion in dermal melanophores. 4. Epidermal melanophores were also responsive to MSH, although the rate and the degree of the response was smaller than those of the dermal melanophores. 5. A possible role of MSH in skin color changes in vivo is discussed.

Sympathetic innervation to the melanophores of a siluroid Parasilurus has been the sole instance of such innervation among lower vertebrates, in which the peripheral transmission to the effector cells is peculiarly cholinergic (Fujii and Miyashita, 1976). In an effort to find a similar case, we studied the nature of transmission of melanophores of a glass catfish Kryptopterus. Electrical nervous stimulation brought about melanosome aggregation in the melanophores. While catecholamines were found ineffective, acetylcholine and its analogues were potently active in aggregating pigment. Atropine or scopolamine interferred with the action of both nervous stimulation and acetylcholine. Physostigmine, on the other hand, augmented the cholinergic effects. The conclusion was that the transmission was cholinergic, being mediated by cholinoceptors of muscarinic type, as in the case of Parasilurus.

Using the guppy, Lebistes reticulatus, and the siluroid catfish, Parasilurus asotus , the effects of purine and pyrimidine derivatives on the movement of melanophores were studied. All the substances tested did not aggregate pigment within melanophores. Adenosine and adenine nucleotides were very effective in dispersing melanosomes within the cell, although adenine itself lacked such action. Derivatives of other purines than adenine and of pyrimidines did not disperse melanosomes. The pigment dispersion induced by adenine derivatives was specifically antagonized by methylxanthines. It was concluded that adenosine receptors are present on the melanophore membrane, which take part in the darkening reaction of fishes.

Effects of cyclic nucleotides on motile iridophores were examined in the blue damselfish, Chrysiptera cyanea. All of the cyclic nucleotides tested, i.e. cAMP, 2',3'-cAMP and cGMP, accelerated the clearing response of the cells even at concentrations of 10(-5) or 10(-4) M. The action of these nucleotides was effectively antagonized by methylxanthines. These results suggest that the effect of cyclic nucleotides on damselfish iridophores is mediated by adenosine receptors in a similar fashion to the action of adenosine.

Using split fin preparations of a tilapia, Sarotherodon niloticus, release of ATP from the adrenergic melanosome-aggregating nerve was studied. Associated with the melanosome aggregation, an apparent release of ATP was detected following electrical nervous stimulation. ATP-release increased with increases in stimulus intensity. Spontaneous release of a small amount of ATP was also detected. It was concluded that ATP may be liberated as a co-transmitter along with the true one, norepinephrine, and it may function to help melanophores recover from the effect of the latter, thus enabling fish to change their hue very rapidly.