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 μm<sup>2</sup> 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 μm<sup>2</sup> 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.
| Identifer | oai:union.ndltd.org:ADTP/279143 |
| Creators | Lenore Litherland |
| Source Sets | Australiasian Digital Theses Program |
| Detected Language | English |
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