Living elasmobranchs occupy every major aquatic ecosystem throughout the world (Compagno 2003; Compagno et al. 2005). Sensory ecology can be a good determinant in comprehending the processes occurring between an organism and its natural environment (Weissburg and Browman 2005). By utilizing ecophysiological tools, insight into the adaptive responses of the sensory systems to their ever-changing ecological niche can help explain behavioral and life history characteristics (Hueter 1991; Litherland 2009). Aquatic animals show structural and physiological adaptations in their visual sense specific to the ecological requirements of their habitat (Hart et al. 2004), implying that vision is an important modality. The visual system of the smooth dogfish (Mustelus canis, family Triakidae) was examined using corneal electrophysiological methods to determine the visual spectral range, irradiance sensitivity, and speed of vision (flicker fusion frequency, FFF). The smooth dogfish, a shallow water bottom feeder inhabiting inshore waters along the eastern United States, was found to be extremely sensitive to dim light (-3.1- 0.1 log light intensity), and have a slow FFF (13 Hz), thus being well adapted to the scotopic conditions of the turbid coastal inshore waters. This prompted a second set of experiments focusing on the chromatic adaptations of the photoreceptor cells and retina function following light adaptation. Light adaptation increased the photopic threshold by 2.0 log light units of intensity (LLI). However, the temporal resolution was not dramatically increased (to 17 Hz), indicating that the retinal integration time is very slow for this species under all circumstances. The spectral sensitivity peak for M. canis (470 nm) was found to be significantly blue-shifted in comparison to other members of the Triakidae family (Crescitelli et al. 1995; Sillman et al. 1996). Smooth dogfish appear to forgo high spatial and temporal resolution for the enhancement of photon capture. The sandbar shark inhabits the same inshore estuaries during the summer months but has a visual system with a higher temporal resolution (FFF, 54 Hz) and a brighter photopic threshold (1.2 LLI-50% max) (Litherland 2009). Furthermore, other elasmobranch or telelost species inhabiting similar photic environments also exhibit faster temporal resolution; little skate (FFF, 30 Hz), weakfish (FFF, 40 Hz), red drum (FFF, 50 Hz), spotted sea trout (FFF, 60 Hz), and Atlantic croaker (FFF, 58 Hz) (Horodysky et al. 2008; McComb et al. 2010). Coastal seas tend to contain more dissolved organics and particulates than the clear oceanic waters of the epipelagic and pelagic zones (McFarland 1986), therefore the retina of smooth dogfish has adapted to be extremely sensitive to dim light, has a long integration time, a low flicker fusion frequency and temporal resolution, and retinal cells that are able to adjust to changing light conditions. All of these factors contribute to the visual system to provide optimal visual ability to enable smooth dogfish to accurately exploit its surroundings.

Made available in DSpace on 2014-12-17T15:37:13Z (GMT). No. of bitstreams: 1 PedroZPMRM_DISSERT.pdf: 485516 bytes, checksum: 5e7cc13b6cdb53047d1dea3153509081 (MD5) Previous issue date: 2012-05-29 / Conselho Nacional de Desenvolvimento Cient?fico e Tecnol?gico / 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 / Dentre os mam?feros placent?rios, os primatas s?o os ?nicos a apresentarem uma vis?o de cores tricromata. Contudo, a distribui??o da tricromacia dentre os primatas n?o ? homog?nea: primatas do Velho Mundo apresentam uma tricromacia uniforme (com todos os indiv?duos sendo tricromatas) e primatas do Novo Mundo apresentam um polimorfismo de vis?o de cores (com machos dicromatas e f?meas dicromatas ou tricromatas). Estudos em ecologia visual t?m investigado que press?es seletivas podem ter sido respons?veis pela evolu??o da tricromacia em primatas, divergindo do padr?o dicromata encontrado nos demais mam?feros. Pistas associadas ao forrageio e ao contexto s?cio-reprodutivo foram analisadas, indicando uma vantagem tricromata na detec??o r?pida de objetos visualmente consp?cuos no ambiente. Entretanto, dicromatas s?o caracterizados pela captura eficiente de est?mulos cr?pticos e camuflados. Estas vantagens relativas aos fen?tipos podem ser respons?veis pela manuten??o do polimorfismo visual em primatas do Novo Mundo e pelo alto ?ndice de daltonismo em humanos (situando-se em torno de 8% em homens caucasianos). Um importante fator que ainda n?o foi levado experimentalmente em conta ? o risco de preda??o e o seu efeito na evolu??o da tricromacia em primatas. Para responder esta pergunta, n?s preparamos e editamos fotografias de animais com pelagens distintas: gatos-do-mato (Leopardus spp.), puma (Puma concolor) e fur?o (Galictis cuja). Os exemplares estavam taxidermizados e as fotografias foram capturadas em tr?s diferentes cen?rios de vegeta??o (mata fechada, cerrado e campo aberto). As imagens dos predadores foram manipuladas para que eles se encaixassem em duas categorias de tamanho de est?mulo (pequenos ou grandes). Ap?s a calibra??o das cores e edi??o das fotos, estas foram apresentadas a 40 humanos (20 dicromatas e 20 tricromatas) por um programa de computador, o qual apresentava um conjunto de quatro fotos por vez (uma foto contendo o animal taxidermizado em meio ? vegeta??o de fundo e outras tr?s contendo apenas a vegeta??o de fundo) e registrava a lat?ncia de resposta e a taxa de acerto dos sujeitos. Os resultados apontam uma vantagem tricromata na detec??o de potenciais predadores. A detec??o dos predadores foi influenciada pelo cen?rio de fundo, pelo tipo de predador, pela sua dimens?o e pelo fen?tipo visual do observador. Como os humanos apresentam uma elevada taxa de discromatopsias, quando comparados com popula??es selvagens de outros Catarrhini ou mesmo popula??es humanas tribais, ? poss?vel que o aumento no ?ndice de dicromatas seja resultado de uma press?o reduzida de detec??o r?pida de predadores. Uma vez que nossa esp?cie passou a viver em grupos mais coesos e resistentes aos ataques de predadores, com o advento da agropecu?ria e a forma??o de vilas, ? poss?vel que o menor risco de preda??o tenha relaxado a sele??o a favor de tricromatas

Neuroethology and ecomorphology seek to understand ecology and behaviour from the perspective of specialised adaptations of sensory systems, such as vision. Sharks display a large variety of visual specialisations reflecting the diversity of different ecological niches they occupy. Many shark species are long-lived and wide ranging and often select different habitats for reproduction, growth, and feeding. Habitat complexity, ambient lighting conditions and feeding strategies can therefore change throughout a shark’s lifetime or between populations. Few comprehensive investigations of visual function exist for sharks as studies typically focus on a narrow aspect of visual function or a particular life history stage. Consequently, there is limited data on within-species plasticity of visual function in response to acclimation to different visual environments or ontogenetic development. The aim of this thesis is to undertake a functional analysis of the shark visual system. An integrated approach is employed to investigate optical, anatomical and physiological specialisations, linking such specialisations to known habitat and/or behavioural traits, with particular emphasis on ontogenetic, inter-population and inter-specific variability. Fundamental capabilities of the visual system are examined, including optical quality, eye morphology, spectral range, irradiance sensitivity, spatial and temporal resolution, contrast discrimination, and temporal and spatial summation. The main study species is the sandbar shark (<i>Carcharhinis plumbeus</i>; Carcharhinidae), a cosmopolitan species of ecological and economic importance. <i>C. plumbeus</i> occupies a wide range of natural habitats from highly turbid coastal estuaries, to relatively clear waters off the outer continental shelves and near pristine clear waters over the slopes of oceanic islands. This provides an opportunity to explore the relationship between habitat variability and the adaptation of visual specialisations and subsequent behaviour. For inter-specific comparison, the visual systems of two other species of shark with contrasting ecological niches are also assessed: the shortspine spurdog (<i>Squalus mitsukurii</i>; Squalidae) and the tiger shark (<i>Galeocerdo cuvier</i>; Carcharhinidae). The study finds marked differences in visual specialisations of the three species studied. The eyes of <i>S. mitsukurii</i> are adapted to enhance retinal illumination within a dim light environment with a large eye, immobile pupil, reflective tapetum and a relatively high optical sensitivity (2.72 μm² steradians). Visual features include a short wavelength lenticular filter, a high spatial resolving power (7.2 cycles/degree) and a large binocular overlap in the dorsal visual field, suggesting adaptations may facilitate the visualisation of bioluminescent prey. In contrast, the eyes of <i>C. plumbeus</i> are optimised for vision under variable light conditions with a mobile pupil and an occlusible tapetum. The sandbar shark shows an optical sensitivity of 1.11 μm² steradians. Visual resolution is highest in the lateral visual field, reaching a peak spatial resolution of 8.9 cycles/degree. An ERG derived spectral response curve for this species indicates maximal response to blue light between 460-490 nm. Interestingly, the tiger shark is maximally sensitive to a brighter range of light intensities compared to sandbar sharks, implying that tiger sharks occupy a more photopic light environment. However, sandbar sharks have a visual system with higher temporal resolution, as evaluated by the ERG response, (54 Hz) than tiger sharks (38 Hz). These results may reflect a difference in the importance of motion perception between <i>C. plumbeus</i> and <i>G. cuvier</i>. Phenotypic variability in visual function is shown between different populations of <i>C. plumbeus</i> occupying habitats with different ambient light conditions. This study provides new evidence of plasticity of visual function in response to acclimation to different visual environments within the same species. Sandbar sharks show an adaptive plasticity in visual sensitivity and temporal resolution, which appears to enable both temporal and population-specific adaptations to local light environments. In addition, the eyes of <i>C. plumbeus</i> and <i>S. mitsukurii</i> continue to grow even in adulthood. Visual performance, with respect to spatial resolving power and optical sensitivity, improve with eye growth. For example, peak spatial resolution increases with eye growth from 4.3 to 8.9 cycles/degree in <i>C. plumbeus</i> and from 5.7 to 7.2 cycles/degree in <i>S. mitsukurii</i>. These studies suggest that the light environment strongly influences visual function in this ancient class of vertebrates. Anthropogenically induced changes in water clarity may, therefore, impact on visually-mediated behaviours such as prey detection, agonistic signals or vertical migration. Anatomical and physiological parameters obtained from these studies provide a platform from which to model visual behaviours such as 1). Prey detection capabilities, 2). The impacts of water clarity on the limits of visually-mediated behaviour, and 3). The visual strategies that would allow sharks to maximise visual function, such as spatial and temporal summation under low light conditions. In conclusion, neuroethological studies can be a useful means to enrich information obtained from life-history and tagging studies and, together, can inform us of the functional role of sharks in marine ecosystems.