Visual system evolution and the nature of the ancestral snake.

The dominant hypothesis for the evolutionary origin of snakes from 'lizards' (non-snake squamates) is that stem snakes acquired many snake features while passing through a profound burrowing (fossorial) phase. To investigate this, we examined the visual pigments and their encoding opsin genes in a range of squamate reptiles, focusing on fossorial lizards and snakes. We sequenced opsin transcripts isolated from retinal cDNA and used microspectrophotometry to measure directly the spectral absorbance of the photoreceptor visual pigments in a subset of samples. In snakes, but not lizards, dedicated fossoriality (as in Scolecophidia and the alethinophidian Anilius scytale) corresponds with loss of all visual opsins other than RH1 (λmax 490-497 nm); all other snakes (including less dedicated burrowers) also have functional sws1 and lws opsin genes. In contrast, the retinas of all lizards sampled, even highly fossorial amphisbaenians with reduced eyes, express functional lws, sws1, sws2 and rh1 genes, and most also express rh2 (i.e. they express all five of the visual opsin genes present in the ancestral vertebrate). Our evidence of visual pigment complements suggests that the visual system of stem snakes was partly reduced, with two (RH2 and SWS2) of the ancestral vertebrate visual pigments being eliminated, but that this did not extend to the extreme additional loss of SWS1 and LWS that subsequently occurred (probably independently) in highly fossorial extant scolecophidians and A. scytale. We therefore consider it unlikely that the ancestral snake was as fossorial as extant scolecophidians, whether or not the latter are para- or monophyletic.

Sexual dimorphism of the electrosensory system: a quantitative analysis of nerve axons in the dorsal anterior lateral line nerve of the blue-spotted Fantail Stingray (Taeniura lymma).

Quantitative studies of sensory axons provide invaluable insights into the functional significance and relative importance of a particular sensory modality. Despite the important role electroreception plays in the behaviour of elasmobranchs, to date, there have been no studies that have assessed the number of electrosensory axons that project from the peripheral ampullae to the central nervous system (CNS). The complex arrangement and morphology of the peripheral electrosensory system has a significant influence on its function. However, it is not sufficient to base conclusions about function on the peripheral system alone. To fully appreciate the function of the electrosensory system, it is essential to also assess the neural network that connects the peripheral system to the CNS. Using stereological techniques, unbiased estimates of the total number of axons were obtained for both the electrosensory bundles exiting individual ampullary organs and those entering the CNS (via the dorsal root of the anterior lateral line nerve, ALLN) in males and females of different sizes. The dorsal root of the ALLN consists solely of myelinated electrosensory axons and shows both ontogenetic and sexual dimorphism. In particular, females exhibit a greater abundance of electrosensory axons, which may result in improved sensitivity of the electrosensory system and may facilitate mate identification for reproduction. Also presented are detailed morphological data on the peripheral electrosensory system to allow a complete interpretation of the functional significance of the sexual dimorphism found in the ALLN.

Vision in elasmobranchs and their relatives: 21st century advances.

This review identifies a number of exciting new developments in the understanding of vision in cartilaginous fishes that have been made since the turn of the century. These include the results of studies on various aspects of the visual system including eye size, visual fields, eye design and the optical system, retinal topography and spatial resolving power, visual pigments, spectral sensitivity and the potential for colour vision. A number of these studies have covered a broad range of species, thereby providing valuable information on how the visual systems of these fishes are adapted to different environmental conditions. For example, oceanic and deep-sea sharks have the largest eyes amongst elasmobranchs and presumably rely more heavily on vision than coastal and benthic species, while interspecific variation in the ratio of rod and cone photoreceptors, the topographic distribution of the photoreceptors and retinal ganglion cells in the retina and the spatial resolving power of the eye all appear to be closely related to differences in habitat and lifestyle. Multiple, spectrally distinct cone photoreceptor visual pigments have been found in some batoid species, raising the possibility that at least some elasmobranchs are capable of seeing colour, and there is some evidence that multiple cone visual pigments may also be present in holocephalans. In contrast, sharks appear to have only one cone visual pigment. There is evidence that ontogenetic changes in the visual system, such as changes in the spectral transmission properties of the lens, lens shape, focal ratio, visual pigments and spatial resolving power, allow elasmobranchs to adapt to environmental changes imposed by habitat shifts and niche expansion. There are, however, many aspects of vision in these fishes that are not well understood, particularly in the holocephalans. Therefore, this review also serves to highlight and stimulate new research in areas that still require significant attention.

The visual ecology of avian photoreceptors.

The spectral sensitivities of avian retinal photoreceptors are examined with respect to microspectrophotometric measurements of single cells, spectrophotometric measurements of extracted or in vitro regenerated visual pigments, and molecular genetic analyses of visual pigment opsin protein sequences. Bird species from diverse orders are compared in relation to their evolution, their habitats and the multiplicity of visual tasks they must perform. Birds have five different types of visual pigment and seven different types of photoreceptor-rods, double (uneven twin) cones and four types of single cone. The spectral locations of the wavelengths of maximum absorbance (lambda(max)) of the different visual pigments, and the spectral transmittance characteristics of the intraocular spectral filters (cone oil droplets) that also determine photoreceptor spectral sensitivity, vary according to both habitat and phylogenetic relatedness. The primary influence on avian retinal design appears to be the range of wavelengths available for vision, regardless of whether that range is determined by the spectral distribution of the natural illumination or the spectral transmittance of the ocular media (cornea, aqueous humour, lens, vitreous humour). Nevertheless, other variations in spectral sensitivity exist that reflect the variability and complexity of avian visual ecology.

Variations in cone photoreceptor abundance and the visual ecology of birds.

The relative abundance and topographical distribution of retinal cone photoreceptors was measured in 19 bird species to identify possible correlations between photoreceptor complement and visual ecology. In contrast to previous studies, all five types of cone photoreceptor were distinguished, using bright field and epifluorescent light microscopy, in four retinal quadrants. Land birds tended to show either posterior dorsal to anterior ventral or anterior dorsal to posterior ventral gradients in cone photoreceptor distribution, fundus coloration and oil droplet pigmentation across the retina. Marine birds tended to show dorsal to ventral gradients instead. Statistical analyses showed that the proportions of the different cone types varied significantly across the retinae of all species investigated. Cluster analysis was performed on the data to identify groups or clusters of species on the basis of their oil droplet complement. Using the absolute percentages of each oil droplet type in each quadrant for the analysis produced clusters that tended to reflect phylogenetic relatedness between species rather than similarities in their visual ecology. Repeating the analysis after subtracting the mean percentage of a given oil droplet type across the whole retina (the 'eye mean') from the percentage of that oil droplet type in each quadrant, i.e. to give a measure of the variation about the mean, resulted in clusters that reflected diet, feeding behaviour and habitat to a greater extent than phylogeny.

Visual pigments, cone oil droplets and ocular media in four species of estrildid finch.

A microspectrophotometric study was conducted on the retinal photoreceptors of four species of bird: cut-throat finches (Amadina fasciata), gouldian finches (Erythrura gouldiae), white-headed munias (Lonchura maja) and plum-headed finches (Neochmia modesta). Spectral characteristics of the photoreceptors in all four species were very similar. Rods contained a medium-wavelength-sensitive visual pigment with a wavelength of maximum absorbance at 502-504 nm. Four spectrally distinct types of single cone contained a visual pigment with wavelength of maximum absorbance at either 370-373 nm (ultraviolet-sensitive), 440-447 nm (short-wavelength-sensitive); 500 nm (medium-wavelength-sensitive) or 562-565 nm (long-wavelength-sensitive). Oil droplets in the ultraviolet-sensitive single cones showed no detectable absorption between 330 nm and 800 nm. Oil droplets in the short-, medium-, and long-wavelength-sensitive single cones had cut-off wavelengths at 415-423 nm, 510-520 nm and 567-575 nm, respectively. Double cones contained the visual pigment with wavelength of maximum absorbance at 562-565 nm observed in long-wavelength-sensitive single cones. Only the principal member of the double cone pair contained an oil droplet (P-type, cut-off wavelength at 414-489 nm depending on species and retinal location). Spectral transmittance of the intact ocular media of each species was measured along the optic axis. Wavelengths of 0.5 transmittance for all species were very similar (316-318 nm).

Visual pigments, oil droplets, ocular media and cone photoreceptor distribution in two species of passerine bird: the blue tit (Parus caeruleus L.) and the blackbird (Turdus merula L.).

The spectral absorption characteristics of the retinal photoreceptors of the blue tit (Parus caeruleus) and blackbird (Turdus merula) were investigated using microspectrophotometry. The retinae of both species contained rods, double cones and four spectrally distinct types of single cone. Whilst the visual pigments and cone oil droplets in the other receptor types are very similar in both species, the wavelength of maximum sensitivity (lambda(max)) of long-wavelength-sensitive single and double cone visual pigment occurs at a shorter wavelength (557 nm) in the blackbird than in the blue tit (563 nm). Oil droplets located in the long-wavelength-sensitive-single cones of both species cut off wavelengths below 570-573 nm, theoretically shifting cone peak spectral sensitivity some 40 nm towards the long-wavelength end of the spectrum. This raises the possibility that the precise lambda(max) of the long-wavelength-sensitive visual pigment is optimised for the visual function of the double cones. The distribution of cone photoreceptors across the retina, determined using conventional light and fluorescence microscopy, also varies between the two species and may reflect differences in their visual ecology.

Retinal asymmetry in birds.

Vertebrate sensory systems are generally based on bilaterally symmetrical sense organs. It is evident, nevertheless, that birds preferentially use either their left or right eye for viewing novel or familiar stimuli [1], and perform visual discrimination tasks under monocular viewing conditions better with one eye than with the other [2] [3]. Because of the nearly complete contralateral decussation of the optic nerves in birds [4], it has been assumed that this division of labour is due solely to cerebral hemispheric specialisation, generated as a result of uneven photostimulation of the eyes of the developing embryo during the last three or four days before hatching [5] [6]. Here, however, we present evidence that in the European starling, Sturnus vulgaris, even the retinae are morphologically asymmetrical in terms of photoreceptor distribution. This is the first evidence for such asymmetry in any bird and suggests that retinal photoreceptor composition should be assessed during studies involving the lateralisation of visually mediated behaviours.

Visual pigments, cone oil droplets, ocular media and predicted spectral sensitivity in the domestic turkey (Meleagris gallopavo).

A microspectrophotometric survey conducted on the retinal photoreceptors of the domestic turkey (Meleagris gallopavo) revealed the presence of five different types of vitamin A1-based visual pigment (rhodopsin) in seven different types of photoreceptor. A single class of rod contained a medium wavelength-sensitive visual pigment (wavelength of maximum absorbance, lambda max, 504 nm). Four different types of single cone contained visual pigment maximally sensitive to wavelengths in either the long (LWS, lambda max 564 nm), medium (MWS, lambda max 505 nm), short (SWS, lambda max 460 nm) or violet (VS, lambda max 420 nm) spectral ranges. The LWS, MWS and SWS single cones contained pigmented oil droplets with cut-off wavelengths (lambda cut) at 514, 490 and 437 nm, respectively. The VS single cone contained a transparent oil droplet which displayed no significant absorbance above 330 nm. A single class of double cone was also identified, both the principal and accessory members of which contained the LWS cone visual pigment. The principal member contained an oil droplet with a lambda cut at 436 nm. No oil droplet was observed in the accessory member. The use of a glycerol-based cell mountant, which reduced wavelength dependent measurement artefacts in the microspectrophotometric measurements, is described. Predictions of cone effective spectral sensitivity, incorporating measurements of the spectral transmission of the ocular media, suggest that turkeys have considerable sensitivity to wavelengths in the ultraviolet-A (UV-A, 315-400 nm) spectral range. This has implications for both the visual ecology of wild birds and the welfare of intensively farmed individuals.

Negative pressure induced airway and pulmonary injury.

PURPOSE: To describe negative pressure injury occurring during the use of a laryngeal mask airway (LMA) in which airway bleeding rather than pulmonary oedema was the major complication. CLINICAL FEATURES: A patient presented to the day surgery unit for resection of a ganglion cyst on her right wrist. She underwent general anaesthesia using an LMA, and experienced severe laryngospasm and transient hypoxaemia (oxygen saturation to 66%) seven minutes after incision. This resolved within 90 sec of succinylcholine administration. Nonetheless, the LMA was removed, a tracheal tube was inserted atraumatically and positive pressure ventilation was maintained until the time of emergence, when fresh blood appeared in the tracheal tube. The blood ultimately became frothy, resembling pulmonary oedema fluid. Haemoptysis, continued postoperatively and led to the hospitalization of this ambulatory patient. CONCLUSION: Rapid development of large subatmospheric pressures, as can occur during severe laryngospasm, may disrupt the tracheobronchial vasculature causing airway bleeding. This bleeding should be distinguished from negative pressure pulmonary oedema.