Morphogenesis of the dorsal pigmentary pattern in wild-type and mutant Rana pipiens

The transition from larval to adult pigmentary patterns during metamorphosis of wild-type. burnsi, and kandiyohi R. pipiens is described. Larval fusiform epidermal melanocytes form a pattern that exactly corresponds to the adult spotting pattern. It is concluded that the larval epidermal pattern expresses a “prepattern” in the larval tissue for the adult pattern. This “prepattern” is visible in kandiyohi, but not in burnsi, tadpoles. The role of the extracellular environment in pigmentary pattern determination is discussed. Gradual changes in all chromatophore densities accompany larval development, while abrupt changes accompany metamorphic climax. There is a net increase in all chromatophore densities by the completion of metamorphosis. Kandiyohi density changes differ quantitatively but not qualitatively from those of wild type. In both wild type and kandiyohi, differentiation of many new dermal melanophores in presumptive spot regions effects expression of the adult spotting pattern. The relative roles of chromatophore differentiation and mitosis in pattern expression, and the hormonal control thereof, are discussed.     

Cytophysiological basis of disruptive pigmentary patterns in the leopard frog Rana pipiens. I. Chromatophore densities and cytophysiology

Statistical analyses of chromatophore densities and morphological states in wild type, burnsi, and kandiyohi: adult Rana pipiens integument yielded the conclusion that disruptive pigmentary pattern is permanently defined by regional chromatophore densities. Spatial chromatophore patterning is enhanced and rendered more visible by the morphological and physiological differences among chromatophores within the various pattern regions. Specifically, (1) chromatophores were differentialy distributed among pattern regions, (2) greater complexity of gross disruptive patterning had greater underlying melanophores densities, (3) there were significant density differences among non-sib individual animals, among fertilization, and among shipment batches, but not among full-sibs, and (4)kandiyohi individuals had more while burnsi individuals had fewer total melanophores than wild type. A stellate morphology was significantly correlated with high melanophore density, although the relationship of melanophore density to melanophore morphology differed among pattern regions and among genotypes. The functional interrelationships of density and morphology, and their role in disruptive patterning and coloration, are discussed.     

The Distribution of Burnsi and Kandiyohi Frogs in Minnesota and Contiguous States

This paper contains new data and maps concerning distributionof the burnsi and kandiyohi pigment pattern variants of thenorthern leopard frog, Rana pipiens.     

The influence of long-term chromatic adaptation on pigment cells and striped pigment patterns in the skin of the zebrafish, Danio rerio

The striped pigment patterns in the flanks of zebrafish result from chromatophores deep within the dermis or hypodermis, while superficial melanophores associated with dermal scales add a dark tint to the dorsal coloration. The responses of these chromatophores were compared during the long-term adaptation of zebrafish to a white or a black background. In superficial skin, melanophores, xanthophores, and two types of iridophores are distributed in a gradient along the dorso-ventral axis independent of the hypodermal pigment patterns. Within one week the superficial melanophores and iridophores changed their density and/or areas of distribution, which adopted the dorsal skin color and the hue of the flank to the background, but did not affect the striped pattern. The increases or decreases in superficial melanophores are thought to be caused by apoptosis or by differentiation, respectively. When the adaptation period was prolonged for more than several months, the striped color pattern was also affected by changes in the width of the black stripes. Some black stripes disappeared and interstripe areas were emphasized with a yellow color within one year on a white background. Such long-term alteration in the pigment pattern was caused by a decrease in the distribution of melanophores and a concomitant increase in xanthophores in the hypodermis. These results indicate that morphological responses of superficial chromatophores contribute to the effective and rapid background adaptation of dorsal skin and while prolonged adaptation also affects hypodermal chromatophores in the flank to alter the striped pigment patterns.     

The distribution of bumblebee colour patterns worldwide: possible significance for thermoregulation, crypsis, and warning mimicry

Bumblebee colour patterns can be highly variable within species, but are often closely similar among species. The present study takes a quantitative approach to survey bumblebee colour patterns in order to address some of the most basic questions concerning resemblances: (1) do colour‐pattern groups exist; (2) are species within colour‐pattern groups geographically clumped; and (3) are some colour‐pattern groups associated with particular kinds of habitat? The results using data for 632 worker patterns from all of the world’s bumblebee species show that: (1) there are many repeating colour patterns, forming relatively few groups of species with similar patterns; (2) colour‐pattern groups can be recognized using simple rules; and (3) species within the 24 largest colour‐pattern groups are significantly aggregated in particular areas of the world. Three principal divisions of colour‐pattern groups are associated with three likely functions: (1) the darkest bumblebees are associated primarily with the tropics, where a thermoregulatory function is suggested; (2) the palest bumblebees are associated with intermediate northern latitudes, where a cryptic function in drying grasslands is suggested; and (3) the intermediate, strongly banded bumblebees are widespread, although these patterns predominate where banding may have advantages as collective warning signals to predators (Müllerian mimicry). Further studies are needed to test these explanations. © The Natural History Museum, London. Journal compilation © 2007 The Linnean Society of London, Biological Journal of the Linnean Society, 2007, 92 , 97–118.     

Melanosome and erythrosome positioning regulates cAMP-induced movement in chromatophores from spotted triplefin, Grahamina capito

This study investigated regulation of uniform positioning of melanosomes and erythrosomes in chromatophores from spotted triplefin Grahamina capito from New Zealand, by modulating levels of intracellular cAMP. Elevated cAMP levels, caused by forskolin treatment, inhibited aggregation and induced rapid dispersion of melanosomes and erythrosomes. The dispersing organelles moved to and accumulated at the cell periphery, leading to an abnormal hyperdispersed state with a melanosome- or erythrosome-depleted cell center. Minutes after hyperdispersion, these organelles reversed direction and moved towards the center again to finally distribute throughout the cells. When chromatophores with initially dispersed melanosomes or erythrosomes were treated with forskolin, no hyperdispersion was seen, but the erythrosomes aggregated slowly. Disassembly of actin by latrunculin resulted in a similar but constant hyperdispersed melanosome and erythrosome distribution. The results show that cAMP not only disperses but also aggregates melanosomes and erythrosomes, and that it is the intracellular position of these organelles that determine the directionality of the cAMP-induced movement. To ascertain the even distribution in the dispersed state, regulatory components associated with the actin cytoskeleton in the cell periphery might modify activity of cytoplasmic dynein or kinesin upon contact with dispersing melanosomes or erythrosomes.     

Spatial patterns in a two-tiered semi-arid shrubland in southeastern Spain

Abstract. Single species and bivariate distribution patterns in a semi-arid shrubland in southeastern Spain, dominated by the tall leguminous shrub Retama sphaerocarpa, were investigated by second-order spatial analysis based on Ripley's K-function. Shrubs were significantly clumped because of a strong association of dwarf shrubs, mostly Artemisia barrelieri, under the canopy of Retama. Retama shrubs were randomly distributed, but when different size-classes were analysed separately, the pattern changed from significantly clumped to random and then to regular with increasing canopy diameter, suggesting increasing intraspecific competition with shrub size. Artemisia was significantly clumped at all scales because of aggregation under the canopy of large Retama shrubs. The association between the species became stronger with increasing canopy diameter of Retama shrubs, suggesting that facilitation prevailed over interspecific competition because of niche separation in different tiers, both above and below ground. Retama shrub size thus determined both the type of pattern for its own size class and tier, and the scale and intensity of the association with its understorey shrubs.     

Genomic conflict drives patterns of X‐linked population structure in Drosophila neotestacea

Intragenomic conflict has the potential to cause widespread changes in patterns of genetic diversity and genome evolution. In this study, we investigate the consequences of sex‐ratio (SR) drive on the population genetic patterns of the X‐chromosome in Drosophila neotestacea. An SR X‐chromosome prevents the maturation of Y‐bearing sperm during male spermatogenesis and thus is transmitted to ~100% of the offspring, nearly all of which are daughters. Selection on the rest of the genome to suppress SR can be strong, and the resulting conflict over the offspring sex ratio can result in the accumulation of multiple loci on the X‐chromosome that are necessary for the expression of drive. We surveyed variation at 12 random X‐linked microsatellites across 16 populations of D. neotestacea that range in SR frequency from 0% to 30%. First, every locus was differentiated between SR and wild‐type chromosomes, and this drives genetic structure at the X‐chromosome. Once the association with SR is accounted for, the patterns of differentiation among populations are similar to the autosomes. Second, within wild‐type chromosomes, the relative heterozygosity is reduced in populations with an increased prevalence of drive, and the heterozygosity of SR chromosomes is higher than expected based on its prevalence. The combination of the relatively high prevalence of SR drive and the structuring of polymorphism between the SR and wild‐type chromosomes suggests that genetic conflict because of SR drive has had significant consequences on the patterns of X‐linked polymorphism and thus also probably affects the tempo of X‐chromosome evolution in D. neotestacea.     

The spatial pattern and canopy-understory association of trees in a cool temperate,mixed forest in western Japan

The spatial distribution pattern of trees and the association between canopy and understory individuals were examined with reference to the distribution of tree crowns in a cool temperate, mixed forest in Ohdaigahara, western Japan. Line transect and contact sampling methods were used to examine the pattern over various spatial scales. These methods are useful to detect patterns over a large study area. The dominance ofChamaecyparis obtusa on steep slopes forming large patches suggested that the distribution of this species is a consequence of landslides. UnderstoryFagus crenata showed a clumped distribution, and the relative coverage of this species was larger in canopy gaps than under a closed canopy. Understory individuals ofAbies homolepis showed a positive association with canopy trees ofF. crenata but a negative association with conspecific canopy individuals. These patterns suggested thatF. crenata regenerates in canopy gaps and is replaced byA. homolepis. The dynamics of these two species are consistent with the process of gap dynamics. The effects of both small- and large-scale disturbance must be evaluated to understand the mechanisms of patch formation and the coexistence of forest tree species.     

Spatial patterns on the sagebrush steppe/Western juniper ecotone

Analysis of the spatial patterns of woody plants is important to better understand the ecological processes that govern the worldwide expansion of woody plants across semi-arid ecosystems. Second-order characteristics of a marked spatial point pattern of western juniper (Juniperus occidentalis subsp. occidentalis) were analyzed using Ripley’s K-functions and the pair-correlation function g. The marked point process of crown diameters was produced via two-dimensional wavelet analysis of a fine scale aerial photograph at the woodland-steppe ecotone in the Reynolds Creek watershed in the Owyhee Mountains, southwestern Idaho. Colonization of J. occidentalis stems from mature juniper trees growing in rocky, fire resistant areas. Although these areas introduce components of natural heterogeneity within the landscape, the selected study area is situated within a single soil type, and we modeled the expansion of juniper plants into previously juniper-free sagebrush steppe as a homogeneous point process with constant intensity. Through this research we have identified two statistically significant spatial scales characteristic of J. occidentalis on the woodland/steppe ecotone: (1) We observed inhibition between J. occidentalis plants at distances <15 m, resulting in a regular pattern, rather than clumped or random. This short-distance inhibition can be attributed to competition for water and other resources. Recruitment of young J. occidentalis occurs significantly more often in a direction away from older plants, maximizing the utilization of water and light resources, and perpetuating the spread of the species into previously juniper-free shrub-steppe. (2) J. occidentalis on the ecotone exhibits significant clustering within a 30–60 m radius. Bivariate point pattern analyses provide evidence that, within a distance of 50–70 m, there is a spatial dependence in tree size such that medium trees are more likely than small trees to be close to large trees. We attribute these phenomena to the fact that juniper seeds are commonly dispersed by berry-eating birds with small territories (0.3–1 ha). Beyond a distance of 50–70 m, juniper plants are randomly distributed, suggesting that additional long-range seed dispersal processes are at work. We further acknowledge the importance of including a reference to spatial scale when formulating hypotheses in statistical analysis of spatio-temporal point patterns.     

Range use of the forest chimpanzees of Kibale: Implications for the understanding of chimpanzee social organization

We contrast the range use patterns of male and female chimpanzees (Pan troglodytes) based on repeated sightings over three years of 19 individuals in the Kanyawara community of Kibale Forest Reserve, Uganda. Depending on how home range size was calculated, male chimpanzees used an area that was 1.5 to 2 times greater than that of females. There was no difference between the sexes in whether their home ranges were used in a clumped or uniform fashion. However, males were more likely to be seen in boundary areas than females. These results are discussed in light of previously proposed models of chimpanzee social organization. It is concluded that the scenario in which females have smaller core areas within the defended home range of the males is most strongly supported by the range use patterns observed in Kibale chimpanzees. © 1993 Wiley-Liss, Inc.     

Interpopulation differences in activity patterns ofMacaca sylvanus in the Moroccan Middle Atlas

We studiedMacaca sylvanus populations from a) a protected, relatively undamaged climax Cedar forest and b) a degraded, mixed Cedar-Oak forest. Animals exhibit considerable differences in demographic, social and time budget characteristics. In the degraded forest there were fewer social groups of smaller average group size and lower population density. Differences in habitat quality, patchiness and intensity of human activities between the two localities are reflected in macaque activity patterns. In the degraded forest the animals allocate less time to feeding, foraging (clumped food resources, smaller group sizes) and Vigilance (more habituated to human presence). They allocate more time to resting and they are involved more frequently in grooming (probably an indication of social tension within groups). In contrary social structures and wild stock, population is declining.     

The phylogenetic significance of colour patterns in marine teleost larvae

Ichthyologists, natural‐history artists, and tropical‐fish aquarists have described, illustrated, or photographed colour patterns in adult marine fishes for centuries, but colour patterns in marine fish larvae have largely been neglected. Yet the pelagic larval stages of many marine fishes exhibit subtle to striking, ephemeral patterns of chromatophores that warrant investigation into their potential taxonomic and phylogenetic significance. Colour patterns in larvae of over 200 species of marine teleosts, primarily from the western Caribbean, were examined from digital colour photographs, and their potential utility in elucidating evolutionary relationships at various taxonomic levels was assessed. Larvae of relatively few basal marine teleosts exhibit erythrophores, xanthophores, or iridophores (i.e. nonmelanistic chromatophores), but one or more of those types of chromatophores are visible in larvae of many basal marine neoteleosts and nearly all marine percomorphs. Whether or not the presence of nonmelanistic chromatophores in pelagic marine larvae diagnoses any major teleost taxonomic group cannot be determined based on the preliminary survey conducted, but there is a trend toward increased colour from elopomorphs to percomorphs. Within percomorphs, patterns of nonmelanistic chromatophores may help resolve or contribute evidence to existing hypotheses of relationships at multiple levels of classification. Mugilid and some beloniform larvae share a unique ontogenetic transformation of colour pattern that lends support to the hypothesis of a close relationship between them. Larvae of some tetraodontiforms and lophiiforms are strikingly similar in having the trunk enclosed in an inflated sac covered with xanthophores, a character that may help resolve the relationships of these enigmatic taxa. Colour patterns in percomorph larvae also appear to diagnose certain groups at the interfamilial, familial, intergeneric, and generic levels. Slight differences in generic colour patterns, including whether the pattern comprises xanthophores or erythrophores, often distinguish species. The homology, ontogeny, and possible functional significance of colour patterns in larvae are discussed. Considerably more investigation of larval colour patterns in marine teleosts is needed to assess fully their value in phylogenetic reconstruction. © 2013 The Authors. Zoological Journal of the Linnean Society published by John Wiley & Sons Ltd on behalf of The Linnean Society of London     

Interpreting biogeographical boundaries among Afrotropical birds: spatial patterns in richness gradients and species replacement

Aim We analyse the geographical distribution of 1911 Afrotropical bird species using indices of three simple biogeographic patterns. The first index, the frequency of species with range edges (T_e), is formulated to map directly the density of species distribution limits, for comparison with the results of traditional biogeographical classification and ordination procedures, in order to show variations in the strength and breadth of transition zones. The other two indices are formulated to seek to distinguish as directly as possible between two components within these transition-zone patterns: contributions from gradients in species richness (T_g); and contributions from replacements among species (T_r). We test the ability of these indices to discover the same boundaries among Afrotropical bird faunas as one popular procedure for classifying areas (TWINSPAN) and then use them to look for geographical trends in the different kinds of transition zones. Location The analysis is restricted to the sub-Saharan or Afrotropical region, excluding the Arabian Peninsula, Madagascar and all offshore islands. Methods We record the presence of each species in 1961 1°×1° grid cells of the map. To apply the three indices, each (core) grid cell in turn is compared with its neighbouring eight cells in the grid. The range edges index (T_e) counts the number of species with range edges between the core cell and the surrounding cells. The richness gradients index (T_g) counts the largest difference in species richness measured diametrically across the core cell in any direction when there is a consistent trend in richness along this line of three cells. The species replacements index (T_r) counts the number of species pairs recorded within a nine-cell neighbourhood that are not corecorded within any of the cells. Values for each of the 1961 grid cells are calculated and used to produce colour-scale maps of transition zones. Results Large-scale spatial patterns of variation in density of range edges (T_e) are consistent with classifications of the same data and with most previous biogeographical classifications proposed for the region. Variation in richness gradients (T_g) and species replacements (T_r) explain different parts of this pattern, with transition zones around humid forests in the equatorial region being dominated by species replacement, and transition zones around deserts (most extensive in the north and south) being dominated by richness gradients. Main conclusions The three indices distinguish the spatial arrangement and intensity of different kinds of transition zones, thereby providing a first step towards a more rigorous mechanistic understanding of the different processes by which they may have arisen and are maintained. As an example of one such pattern shown by our analyses of Afrotropical birds, there is evidence for a broad latitudinal trend in the nature of transition zones in faunal composition (following the latitudinal distribution of the different kinds of habitat transitions), from being dominated by species replacements near the equator to being dominated by richness gradients further from the equator.     

Significance of microhabitat heterogeneity in the spatial pattern and size-class structure of Anastatica hierochuntica L.

Anastatica hierochuntica L. (Brassicaceae) is a desert monocarpic annual species characterized by a topochory/ombrohydrochory type of seed dispersal. The hygrochastic nature of the dry skeletons (dead individuals) permits controlling seed dispersal by rain events. The amount of dispersed seeds is proportional to the intensity of rainfall. When light showers occur, seeds are released and remain in the site. Seeds dispersed in the vicinity of the mother or source plant (primary type of seed dispersal) resulted in clumped pattern and complicated interrelationships among size-classes of the population. Following heavy rainfall, most seeds are released and transported into small patches and shallow depressions which collect runoff water. The dead A. hierochuntica skeletons demonstrate site-dependent size-class structure, spatial pattern and spatial interrelationships in different microhabitats. Four microhabitat types have been sampled: runnels, patches and simple and compound depressions in two sites (gravel and sand). Ripley's K-function was used to analyze the spatial pattern in populations of A. hierochuntica skeletons in the study microhabitats. Clumped patterns were observed in nearly all of the study microhabitats. Populations of A. hierochuntica in the sand site were more productive than in the gravel site and usually had more individuals in the larger size-classes. In the compound-depression microhabitat, the degree of clumping decreased from the core zone to the intermediate zone then shifted into overdispersed pattern in the outer zone. At the within size-class level, the clumped pattern dominated in small size classes but shifted into random and overdispersed patterns in the larger size classes. Aggregation between small and large size-classes was not well-defined but large individuals were found closer to the smaller individuals than to those of their own class. In relation to the phytomass and the size-class structure, the outer zone of the simple depression and the outer and intermediate zones of the compound depression microhabitats were the most productive sites.     

Spatial patterns in seed bank and vegetation of semi-natural mountain meadows

Soil seed bank composition and vegetative spatial patterns were studied in four mountain meadow communities in the Broto Valley (N. Spain), in order to analyse the differences that might exists between the two life forms. Soil and vegetation samples were taken at 1 m intervals from 10 m × 10 m quadrats in each meadow in one-year study. The spatial distribution of species was analysed along with the calculation of an autocorrelation coefficient which takes account of the relative position of samples: (Moran’s I). The results indicate that the abundance of the majority of the species in the seed bank and in the vegetation are randomly distributed, the percentage of species with a clumped distribution only exceeds 35% in the vegetation of one meadow and none of the taxa identified showed a uniform spatial organisation. The species that were distributed in the seed bank in a clumped pattern in more than one meadow were those of the pioneer species (Anagallis arvensis, Centaurium erythraea, Lamium purpureum and Stellaria media). All of these formed long-term persistent seed banks but were absent in the established vegetation in these meadows. According to the results, there exist not only differences between the spatial distribution of the species present in the same community, but also, that some species change their pattern of distribution according to the life form and to the grassland type in which they are found.     

Do spatial effects play a role in the spatial distribution of desert‐dwelling Acacia raddiana?

Abstract. We investigated the spatial pattern of A. raddiana in the Negev desert of Israel in order to gain insights into the factors and processes driving the dynamics of this species. Using a scale‐dependent measure, the ring statistic, we analysed both patterns observed in the field and time series of spatial tree distributions produced by a simulation model. In the field, random spacing was the predominant pattern observed. However seedlings were clumped on small scales. We ran the model under two contrasting scenarios representing hypotheses that explain the clumping of seedlings and the random distribution of trees. One hypothesis is that there is spatial heterogeneity in seed distribution, germination and seedling mortality, but that these heterogeneities are not correlated with each other in space. The second hypothesis assumes a correlation between these heterogeneities leading to areas suitable for establishment. However, the suitability of the sites is temporally variable. Furthermore, the second hypothesis assumes density‐dependent tree mortality due to competition. Both hypotheses lead to spatial distributions that are in qualitative agreement with the patterns observed in the field. Therefore, the classical view that a clumped seedling distribution and a random pattern of older trees is due to clumped regeneration and density‐dependent mortality may not hold for Acacia trees in the Negev.     

Molecular and developmental contributions to divergent pigment patterns in marine and freshwater sticklebacks

Pigment pattern variation across species or populations offers a tractable framework in which to investigate the evolution of development. Juvenile threespine sticklebacks (Gasterosteus aculeatus) from marine and freshwater environments exhibit divergent pigment patterns that are associated with ecological differences. Juvenile marine sticklebacks have a silvery appearance, whereas sticklebacks from freshwater environments exhibit a pattern of vertical bars. We investigated both the developmental and molecular basis of this population‐level variation in pigment pattern. Time course imaging during the transition from larval to juvenile stages revealed differences between marine and freshwater fish in spatial patterns of chromatophore differentiation as well as in pigment amount and dispersal. In freshwater fish, melanophores appear primarily within dark bars whereas iridophores appear within light bars. By contrast, in marine fish, these chromatophores are interspersed across the flank. In addition to spatially segregated chromatophore differentiation, pigment amount and dispersal within melanophores varies spatially across the flank of freshwater, but not marine fish. To gain insight into the molecular pathways that underlie the differences in pigment pattern development, we evaluated differential gene expression in the flanks of developing fish using high‐throughput cDNA sequencing (RNA‐seq) and quantitative PCR. We identified several genes that were differentially expressed across dark and light bars of freshwater fish, and between freshwater and marine fish. Together, these experiments begin to shed light on the process of pigment pattern evolution in sticklebacks.     

Yellow Marking and Pteridine Contents in the Integument of Albino Armadillidium vulgare

The albino mutant strain in the woodlice, Armadillidium vulgare, was investigated with respect to the yellow patterns on the dorsal integument. Pigment cells were observed with electron microscope in order to determine the cell types of yellow markings. Quantitative analyses of pteridines in the albino were carried out by HPLC. The result indicated that the albino integument contain sepiapterin, biopterin, pterin, isoxanthopterin as in the wild type and the red mutant strain. The total amount of the four pteridines in the albino was about half as much as that in the red phenotype for both males and females, respectively. Males and females showed almost the same totals and ratios of the four pteridines in the albino and red phenotypes. Therefore, pteridine contents in both phenotypes of A. vulgare may not be related to the activity of androgenic gland hormone. Yellow chromatophores of the albino and red phenotypes were morphologically identical, emitting a yellow fluorescence. These cells contained numerous electron-lucent pigment organelles which were similar to pteridine granules of the wild type.     

Chromatophore activity during natural pattern expression by the squid Sepioteuthis lessoniana: contributions of miniature oscillation

Squid can rapidly change the chromatic patterns on their body. The patterns are created by the expansion and retraction of chromatophores. The chromatophore consists of a central pigment-containing cell surrounded by radial muscles that are controlled by motor neurons located in the central nervous system (CNS). In this study we used semi-intact squid (Sepioteuthis lessoniana) displaying centrally controlled natural patterns to analyze spatial and temporal activities of chromatophores located on the dorsal mantle skin. We found that chromatophores oscillated with miniature expansions/retractions at various frequencies, even when the chromatic patterns appear macroscopically stable. The frequencies of this miniature oscillation differed between "feature" and "background" areas of chromatic patterns. Higher frequencies occurred in feature areas, whereas lower frequencies were detected in background areas. We also observed synchronization of the oscillation during chromatic pattern expression. The expansion size of chromatophores oscillating at high frequency correlated with the number of synchronized chromatophores but not the oscillation frequency. Miniature oscillations were not observed in denervated chromatophores. These results suggest that miniature oscillations of chromatophores are driven by motor neuronal activities in the CNS and that frequency and synchrony of this oscillation determine the chromatic pattern and the expansion size, respectively.