Detecção de predadores por dicromatas e tricromatas humanos e a sua implicação na evolução da visão de cores em primatas

Among placental mammals, primates are the only ones to present trichromatic color vision. However, the distribution of trichromacy among primates is not homogeneous: Old World primates shows an uniform trichromacy (with all individuals being trichromats) and New World primates exhibit a color vision polymorphism (with dichromatic males and dichromatic or trichromatic females). Visual ecology studies have investigated which selective pressures may have been responsible for the evolution of trichromacy in primates, diverging from the dichromat standard found in other mammals. Cues associated with foraging and the socio-reproductive status were analyzed, indicating a trichromatic advantage for the rapid detection of visually conspicuous objects against a green background. However, dichromats are characterized by an efficient capture of cryptic and camouflaged stimuli. These advantages regarding phenotype may be responsible for the maintenance of the visual polymorphism in New World primates and for the high incidence of color blindness in humans (standing around 8% in Caucasian men). An important factor that has not yet been experimentally taken into account is the predation risk and its effect on the evolution of trichromacy in primates. To answer this question, we prepared and edited pictures of animals with different coats: oncillas (Leopardus spp.), puma (Puma concolor) and ferret (Galictis cuja). The specimens were taxidermized and the photographs were taken in three different vegetation scenarios (dense forest, cerrado and grassland). The images of the predators were manipulated so that they fit into two categories of stimulus size (small or large). After color calibration and photo editing, these were presented to 40 humans (20 dichromats and 20 trichromats) by a computer program, which presented a set of four photos at a time (one picture containing the taxidermized animal amid the background vegetation and three depicting only the background vegetation) and recorded the response latency and success rate of the subjects. The results show a trichromatic advantage in detecting potential predators. The predator detection was influenced by the background, the predator species, the dimension of the stimulus and the observer s visual phenotype. As humans have a high rate of dyschromatopsias, when compared to wild Catarrhini or human tribal populations, it is possible that the increased rate of dichromats is a result of reduced pressure for rapid predator detection. Since our species came to live in more cohesive groups and resistant to attack by predators, with the advent of agriculture and the formation of villages, it is possible that the lower risk of predation has reduced the selection in favor of trichromats

Seletividade espacial de cor do potencial cortical provocado visual pseudoaleatório

A seletividade espacial para cor tem sido investigada usando métodos eletrofisiológicos invasivos e não invasivos, e métodos psicofísicos. Em eletrofisiologia cortical visual não invasiva este tópico foi investigado usando métodos convencionais de estimulação periódica e extração de respostas por promediação simples. Novos métodos de estimulação (apresentação pseudo-aleatória) e extração de respostas corticais não invasivas (correlação cruzada) foram desenvolvidos e ainda não foram usados para investigar a seletividade espacial de cor de respostas corticais. Este trabalho objetivou introduzir esse novo método de eletrofisiologia pseudoaleatória para estudar a seletividade espacial de cor. Foram avaliados 14 tricromatas e 16 discromatópsicos com acuidade visual normal ou corrigida. Os voluntários foram avaliados pelo anomaloscópio HMC e teste de figuras de Ishihara para caracterizar a visão de cores quanto à presença de tricromacia. Foram usadas redes senoidais, 8º de ângulo visual, vermelho-verde para 8 frequências espaciais entre 0,2 a 10 cpg. O estímulo foi temporalmente modulado por uma sequência-m binária em um modo de apresentação de padrão reverso. O sistema VERIS foi usado para extrair o primeiro e o segundo slice do kernel de segunda ordem (K2.1 e K2.2, respectivamente). Após a modelagem da resposta às frequências espaciais com função de diferença de gaussianas, extraiu-se a frequência espacial ótima e banda de frequências com amplitudes acima de ¾ da amplitude máxima da função para servirem como indicadores da seletividade espacial da função. Também foi estimada a acuidade visual cromática pelo ajuste de uma função linear aos dados de amplitude a partir da frequência espacial do pico de amplitude até a mais alta frequência espacial testada. Em tricromatas, foi encontrada respostas cromáticas no K2.1 e no K2.2 que apresentaram seletividade espacial diferentes. Os componentes negativos do K2.1 e do K2.2 apresentaram sintonia passa-banda e o componente positivo do K2.1 apresentou sintonia passa-baixa. A acuidade visual estimada de todos os componentes estudados foi próxima àquelas encontradas por Mullen (1985) e Kelly (1983). Diferentes componentes celulares podem estar contribuindo para a geração do VECP pseudoaleatório. Este novo método se candidata a ser uma importante ferramenta para a avaliação não invasiva da visão de cores em humanos.

Characterization of Opsin Gene Alleles Affecting Color Vision in a Wild Population of Titi Monkeys (Callicebus brunneus)

The color vision of most platyrrhine primates is determined by alleles at the polymorphic X-linked locus coding for the opsin responsible for the middle- to long-wavelength (M/L) cone photopigment. Females who are heterozygous at the locus have trichromatic vision, whereas homozygous females and all males are dichromatic. This study characterized the opsin alleles in a wild population of the socially monogamous platyrrhine monkey Callicebus brunneus (the brown titi monkey), a primate that an earlier study suggests may possess an unusual number of alleles at this locus and thus may be a subject of special interest in the study of primate color vision. Direct sequencing of regions of the M/L opsin gene using feces-, blood-, and saliva-derived DNA obtained from 14 individuals yielded evidence for the presence of three functionally distinct alleles, corresponding to the most common M/L photopigment variants inferred from a physiological study of cone spectral sensitivity in captive Callicebus. Am. J. Primatol. 73:189-196, 2011. (C) 2010 Wiley-Liss, Inc.

Ecology and evolution of primate colour vision

More than one hundred years ago, Grant Allen suggested that colour vision in primates, birds and insects evolved as an adaptation for foraging on colourful advertisements of plants-fruits and flowers. Recent studies have shown that well developed colour vision appeared long before fruits and flowers evolved. Thus, colour vision is generally beneficial for many animals, not only for those eating colourful food. Primates are the only placental mammals that have trichromatic colour vision. This may indicate either that trichromacy is particularly useful for primates or that primates are unique among placental mammals in their ability to utilise the signals of three spectrally distinct types of cones or both. Because fruits are an important component of the primate diet, primate trichromacy could have evolved as a specific adaptation for foraging on fruits. Alternatively, primate trichromacy could have evolved as an adaptation for many visual tasks. Comparative studies of mammalian eyes indicate that primates are the only placental mammals that have in their retina a pre-existing neural machinery capable of utilising the signals of an additional spectral type of cone. Thus, the failure of non-primate placental mammals to evolve trichromacy can be explained by constraints imposed on the wiring of retinal neurones.

Cone topography and spectral sensitivity in two potentially trichromatic marsupials, the quokka (Setonix brachyurus) and quenda (Isoodon obesulus)

The potential for trichromacy in mammals, thought to be unique to primates, was recently discovered in two Australian marsupials. Whether the presence of three cone types, sensitive to short- (SWS), medium-(MWS) and long-(LWS) wavelengths, occurs across all marsupials remains unknown. Here, we have investigated the presence, distribution and spectral sensitivity of cone types in two further species, the quokka (Setonix brachyurus) and quenda (Isoodon obesulus). Immunohistochemistry revealed that SWS cones in the quokka are concentrated in dorso-temporal retina, while in the quenda, two peaks were identified in naso-ventral and dorso-temporal retina. In both species, MWS/LWS cone spatial distributions matched those of retinal ganglion cells. Microspectrophotometry (MSP) confirmed that MWS and LWS cones are spectrally distinct, with mean wavelengths of maximum absorbance at 502 and 538 nm in the quokka, and at 509 and 551 nm, in the quenda. Although small SWS cone outer segments precluded MSP measurements, molecular analysis identified substitutions at key sites, accounting for a spectral shift from ultraviolet in the quenda to violet in the quokka. The presence of three cone types, along with previous findings in the fat-tailed dunnart and honey possum, suggests that three spectrally distinct cone types are a feature spanning the marsupials.