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Chapter 1, The Reptilian Spectacle - UWSpace - University of ...

Chapter 1, The Reptilian Spectacle - UWSpace - University of ...

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Vision has been credited with sparking the incredible phyletic diversification <strong>of</strong> animals during the<br />

Cambrian explosion 545 million years ago (Nilsson 1996; Land and Nilsson 2002). Given the<br />

remarkable value <strong>of</strong> light perception and image formation, it is perhaps no wonder that most species<br />

have evolved structures, behaviours and biochemical mechanisms to protect the integrity <strong>of</strong> their eyes<br />

during the courses <strong>of</strong> their life cycles. While many invertebrates have eyes supported by hard chitinous<br />

material (eg. arthropods) and others are able to regenerate damaged or excised eyes (eg. gastropods,<br />

Flores Scarsso and Pellegrino de Iraldi, 1973), vertebrates have comparatively fragile eyes in that the<br />

optically transmissive window to the outside world, the cornea, is rather delicate compared to their<br />

integument and has limited regenerative abilities beyond the renewal <strong>of</strong> the epithelium and scarring <strong>of</strong><br />

the stroma. Vertebrates have thus evolved protective extra-ocular structures, with eyelids and<br />

nictitating membranes (i.e. “third” eyelids) being the most familiar examples among terrestrial species.<br />

Another protective structure that evolved among some vertebrates, terrestrial and aquatic alike,<br />

consists <strong>of</strong> a layer <strong>of</strong> transparent integument overlaying the eyes, which acts as a permanent,<br />

immovable shield against the external environment. <strong>The</strong>se integumentary “spectacles” are found in<br />

some fishes (which typically lack eyelids altogether) and a few amphibians, but find their greatest<br />

terrestrial presence among reptiles, with some lizards having them, and snakes in particular being<br />

ubiquitously equipped with them.<br />

Given that there is no analogue to the spectacle in mammals (or birds), and that it thus<br />

precludes spectacled animals as models for most human ocular conditions, it is perhaps no source <strong>of</strong><br />

wonder that gaps exist in our knowledge <strong>of</strong> it in such diverse areas as its anatomy, physiology, optics,<br />

evolution, and its implications to ecology and ethology. This review aims to bring together the current<br />

state <strong>of</strong> knowledge <strong>of</strong> the reptilian spectacle -- its anatomy and physiology, its adaptive significance,<br />

and its evolution -- and in doing so to highlight areas <strong>of</strong> limited knowledge, some <strong>of</strong> which will be<br />

addressed by the experiments described in the 3 chapters that follow. This review will begin with a<br />

description <strong>of</strong> the anatomy <strong>of</strong> spectacles, from a somewhat historical perspective, to provide a context<br />

for further discussions <strong>of</strong> their other biological characteristics.<br />

3

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