Enhanced short-wavelength sensitivity in the blue-tongued skink Tiliqua rugosa.

Despite lizards using a wide range of colour signals, the limited variation in photoreceptor spectral sensitivities across lizards suggests only weak selection for species-specific, spectral tuning of photoreceptors. Some species, however, have enhanced short-wavelength sensitivity, which probably helps with the detection of signals rich in ultraviolet and short wavelengths. In this study, we examined the visual system of Tiliqua rugosa, which has an ultraviolet/blue tongue, to gain insight into this species' visual ecology. We used electroretinograms, opsin sequencing and immunohistochemical labelling to characterize whole-eye spectral sensitivity and the elements that shape it. Our findings reveal that T. rugosa expresses all five opsins typically found in lizards (SWS1, SWS2, RH1, RH2 and LWS) but possesses greatly enhanced short-wavelength sensitivity compared with other diurnal lizards. This enhanced short-wavelength sensitivity is characterized by a broadening of the spectral sensitivity curve of the eye towards shorter wavelengths while the peak sensitivity of the eye at longer wavelengths (560 nm) remains similar to that of other diurnal lizards. While an increased abundance of SWS1 photoreceptors is thought to mediate elevated ultraviolet sensitivity in a couple of other lizard species, SWS1 photoreceptor abundance remains low in this species. Instead, our findings suggest that short-wavelength sensitivity is driven by multiple factors which include a potentially red-shifted SWS1 photoreceptor and the absence of short-wavelength-absorbing oil droplets. Examining the coincidence of enhanced short-wavelength sensitivity with blue tongues among lizards of this genus will provide further insight into the co-evolution of conspecific signals and whole-eye spectral sensitivity.

Energy conservation characterizes sleep in sharks.

Sharks represent the earliest group of jawed vertebrates and as such, they may provide original insight for understanding the evolution of sleep in more derived animals. Unfortunately, beyond a single behavioural investigation, very little is known about sleep in these ancient predators. As such, recordings of physiological indicators of sleep in sharks have never been reported. Reduced energy expenditure arising from sustained restfulness and lowered metabolic rate during sleep have given rise to the hypothesis that sleep plays an important role for energy conservation. To determine whether this idea applies also to sharks, we compared metabolic rates of draughtsboard sharks (Cephaloscyllium isabellum) during periods ostensibly thought to be sleep, along with restful and actively swimming sharks across a 24 h period. We also investigated behaviours that often characterize sleep in other animals, including eye closure and postural recumbency, to establish relationships between physiology and behaviour. Overall, lower metabolic rate and a flat body posture reflect sleep in draughtsboard sharks, whereas eye closure is a poorer indication of sleep. Our results support the idea for the conservation of energy as a function of sleep in these basal vertebrates.

Visual Opsin Diversity in Sharks and Rays.

The diversity of color vision systems found in extant vertebrates suggests that different evolutionary selection pressures have driven specializations in photoreceptor complement and visual pigment spectral tuning appropriate for an animal's behavior, habitat, and life history. Aquatic vertebrates in particular show high variability in chromatic vision and have become important models for understanding the role of color vision in prey detection, predator avoidance, and social interactions. In this study, we examined the capacity for chromatic vision in elasmobranch fishes, a group that have received relatively little attention to date. We used microspectrophotometry to measure the spectral absorbance of the visual pigments in the outer segments of individual photoreceptors from several ray and shark species, and we sequenced the opsin mRNAs obtained from the retinas of the same species, as well as from additional elasmobranch species. We reveal the phylogenetically widespread occurrence of dichromatic color vision in rays based on two cone opsins, RH2 and LWS. We also confirm that all shark species studied to date appear to be cone monochromats but report that in different species the single cone opsin may be of either the LWS or the RH2 class. From this, we infer that cone monochromacy in sharks has evolved independently on multiple occasions. Together with earlier discoveries in secondarily aquatic marine mammals, this suggests that cone-based color vision may be of little use for large marine predators, such as sharks, pinnipeds, and cetaceans.

Functional morphology of the cornea of the Little Penguin Eudyptula minor (Aves).

The cornea is a specialized component of the vertebrate eye that provides protection, refractive power, transparency for optical imaging and mechanical support. However, the corneas of birds have received little attention with no comprehensive study of their functional morphology. Using light microscopy and both scanning and transmission electron microscopy, the first description of the ultrastructure of all of the main components of the cornea in two different-sized individuals of the Little Penguin Eudyptula minor is presented. Two types of microprojections protrude from the surface of the cornea with a predominance of microridges and microvilli found in central (flattened) and peripheral regions, respectively. Epithelial cell density is higher in peripheral cornea, especially in the larger (older) individual, while there is a reduction of epithelial cell density with age. The cornea comprises a thick epithelium uniquely attached to the basement membrane with numerous incursions rather than anchoring fibres and anchoring plaques as is found in other vertebrate corneas. Posterior to Bowman's layer, the orthogonally-arranged collagen fibril lamellae in the stroma form extensive branches and anastomoses. Desçemet's membrane is well-developed with an anterior or foetal portion with long banding. However, the thickness of Desçemet's membrane is larger in the older individual with the inclusion of an additional irregular pale-staining posterior portion. Polygonal endothelial cells extend across the cornea as a monolayer with often tortuous cell junctions. Endothelial cell density increases towards the periphery, but decreases with age. Primary cilia are observed protruding through the central region of some endothelial cells into the anterior segment but subsurface structures resembling cilia suggest that these features may be more common. The ultrastructure of the corneal components reveals a range of functional adaptations that reflect the amphibious lifestyle of this seabird.

A comparison of the ultrastructure of the cornea of the pre- and post-metamorphic axolotl (Ambystoma mexicanum, Amphibia).

The corneal ultrastructure of the pre- and post-metamorphic stages of the neotenic axolotl Ambystoma mexicanum is examined using light microscopy and both scanning and transmission electron microscopy to reveal whether there are any morphological changes associated with a switch in lifestyle. Although the complement of corneal layers remains the same, there are significant quantitative changes in corneal, epithelial and stromal thickness, epithelial and endothelial cell size and density, and the thickness of Bowman's layer and Desçemet's membrane. Microholes in the epithelium and vertical sutures within the stroma are predominant features in the pre-metamorphic stage but are rarely seen in the post-metamorphic stage. There are also significant quantitative centro-peripheral differences in the thickness of the whole cornea, primarily due to differences in the thickness of the stroma in both metamorphic stages. These changes may reflect the physiological demands on the cornea as it switches from a purely aquatic to an amphibious lifestyle, which includes venturing onto land.

Behavioural sleep in two species of buccal pumping sharks (Heterodontus portusjacksoni and Cephaloscyllium isabellum).

Sleep is known to occur in most, if not all, animals studied thus far. Recent studies demonstrate the presence of sleep in flatworms and jellyfish, suggesting that this behaviour evolved early in the evolution of animals. Sharks are the earliest known extant, jawed vertebrates and may play an important role in understanding the evolutionary history of sleep in vertebrates, and yet, it is unknown whether they sleep. The Port Jackson (Heterodontus portusjacksoni) and draughtsboard (Cephaloscyllium isabellum) sharks are both benthic, buccal pumping species and remain motionless for extended periods of time. Whether these periods of prolonged inactivity represent sleep or quiet wakefulness is unknown. A key criterion for separating sleep from other quiescent states is an increased arousal threshold. We show here that inactive sharks of both species require significantly higher levels of electric stimulation before they show a visible response. Sharks deprived of rest, however, show no significant compensatory increase in restfulness during their normal active period following enforced swimming. Nonetheless, increased arousal thresholds in inactive animals suggest that these two species of shark sleep - the first such demonstration for members of this group of vertebrates. Further research, including electrophysiological studies, on these and other sharks, is required for a comprehensive understanding of sleep in cartilaginous fishes.

The effect of ecological factors on eye morphology in the western rainbowfish, Melanotaenia australis.

Ecological factors such as spatial habitat complexity and diet can explain variation in visual morphology, but few studies have sought to determine whether visual specialisation can occur among populations of the same species. We used a small Australian freshwater fish (the western rainbowfish, Melanotaenia australis) to determine whether populations showed variation in eye size and eye position, and whether this variation could be explained by environmental (light availability, turbidity) and ecological (predation risk, habitat complexity, invertebrate abundance) variables. We investigated three aspects of eye morphology - (1) eye size relative to body size, (2) pupil size relative to eye size and (3) eye position in the head - for fish collected from 14 sites in a major river catchment in northwest Western Australia. We found significant variation among populations in all three measures of eye morphology, but no effect of sex on eye size or eye position. Variation in eye diameter and eye position was best explained by the level of habitat complexity. Specifically, fish occurring in habitats with low complexity (i.e. open water) tended to have smaller, more dorsally located eyes than those occurring in more complex habitats (i.e. vegetation present). The size of the pupil relative to the size of the eye was most influenced by the presence of surrounding rock formations; fish living in gorge habitats had significantly smaller pupils (relative to eye size) than those occupying semi-gorge sites or open habitats. Our findings reveal that different ecological and environmental factors contribute to habitat-specific visual specialisations within a species.

Convergence of Olfactory Inputs within the Central Nervous System of a Cartilaginous and a Bony Fish: An Anatomical Indicator of Olfactory Sensitivity.

The volume of the olfactory bulbs (OBs) relative to the brain has been used previously as a proxy for olfactory capabilities in many vertebrate taxa, including fishes. Although this gross approach has predictive power, a more accurate assessment of the number of afferent olfactory inputs and the convergence of this information at the level of the telencephalon is critical to our understanding of the role of olfaction in the behaviour of fishes. In this study, we used transmission electron microscopy to assess the number of first-order axons within the olfactory nerve (ON) and the number of second-order axons in the olfactory peduncle (OP) in established model species within cartilaginous (brownbanded bamboo shark, Chiloscyllium punctatum [CP]) and bony (common goldfish, Carassius auratus [CA]) fishes. The total number of axons varied from a mean of 18.12 ± 7.50 million in the ON to a mean of 0.38 ± 0.21 million in the OP of CP, versus 0.48 ± 0.16 million in the ON and 0.09 ± 0.02 million in the OP of CA. This resulted in a convergence ratio of approximately 50:1 and 5:1, respectively, for these two species. Based on astroglial ensheathing, axon type (unmyelinated [UM] and myelinated [M]) and axon size, we found no differentiated tracts in the OP of CP, whereas a lateral and a medial tract (both of which could be subdivided into two bundles or areas) were identified for CA, as previously described. Linear regression analyses revealed significant differences not only in axon density between species and locations (nerves and peduncles), but also in axon type and axon diameter (p < 0.05). However, UM axon diameter was larger in the OPs than in the nerve in both species (p = 0.005), with no significant differences in UM axon diameter in the ON (p = 0.06) between species. This study provides an in-depth analysis of the neuroanatomical organisation of the ascending olfactory pathway in two fish taxa and a quantitative anatomical comparison of the summation of olfactory information. Our results support the assertion that relative OB volume is a good indicator of the level of olfactory input and thereby a proxy for olfactory capabilities.

Sexual selection and the evolution of sperm morphology in sharks.

Post-copulatory sexual selection, and sperm competition in particular, is a powerful selective force shaping the evolution of sperm morphology. Although mounting evidence suggests that post-copulatory sexual selection influences the evolution of sperm morphology among species, recent evidence also suggests that sperm competition influences variation in sperm morphology at the intraspecific level. However, contradictory empirical results and limited taxonomic scope have led to difficulty in assessing the generality of sperm morphological responses to variation in the strength of sperm competition. Here, we use phylogenetically controlled analyses to explore the effects of sperm competition on sperm morphology and variance in sharks, a basal vertebrate group characterized by wide variation in rates of multiple mating by females, and consequently sperm competition risk. Our analyses reveal that shark species experiencing greater levels of sperm competition produce sperm with longer flagella and that sperm flagellum length is less variable in species under higher sperm competition risk. In contrast, neither the length of the sperm head and midpiece nor variation in sperm head and midpiece length was associated with sperm competition risk. Our findings demonstrate that selection influences both the inter- and intraspecific variation in sperm morphology and suggest that the flagellum is an important target of sexual selection in sharks. These findings provide important insight into patterns of selection on the ejaculate in a basal vertebrate lineage.

Underwater hearing in sea snakes (Hydrophiinae): first evidence of auditory evoked potential thresholds.

The viviparous sea snakes (Hydrophiinae) are a secondarily aquatic radiation of more than 60 species that possess many phenotypic adaptations to marine life. However, virtually nothing is known of the role and sensitivity of hearing in sea snakes. This study investigated the hearing sensitivity of the fully marine sea snake Hydrophis stokesii by measuring auditory evoked potential (AEP) audiograms for two individuals. AEPs were recorded from 40 Hz (the lowest frequency tested) up to 600 Hz, with a peak in sensitivity identified at 60 Hz (163.5 dB re. 1 µPa or 123 dB re. 1 µm s-2). Our data suggest that sea snakes are sensitive to low-frequency sounds but have relatively low sensitivity compared with bony fishes and marine turtles. Additional studies are required to understand the role of sound in sea snake life history and further assess these species' vulnerability to anthropogenic noise.

Evidence for Sleep in Sharks and Rays: Behavioural, Physiological, and Evolutionary Considerations.

Sleep is widespread across the animal kingdom. However, most comparative sleep data exist for terrestrial vertebrates, with much less known about sleep in amphibians, bony fishes, and invertebrates. There is an absence of knowledge on sleep in cartilaginous fishes. Sharks and rays are amongst the earliest vertebrates, and may hold clues to the evolutionary history of sleep and sleep states found in more derived animals, such as mammals and birds. Here, we review the literature concerning activity patterns, sleep behaviour, and electrophysiological evidence for sleep in cartilaginous (and bony) fishes following an exhaustive literature search that found more than 80 relevant studies in laboratory and field environments. Evidence for sleep in sharks and rays that respire without swimming is preliminary; evidence for sleep in continuously swimming fishes is currently absent. We discuss ways in which the latter group might sleep concurrent with sustained movement, and conclude with suggestions for future studies in order to provide more comprehensive data on when, how, and why sharks and rays sleep.

Retinal Topography in Two Species of Baleen Whale (Cetacea: Mysticeti).

Little is known about the visual systems of large baleen whales (Mysticeti: Cetacea). In this study, we investigate eye morphology and the topographic distribution of retinal ganglion cells (RGCs) in two species of mysticete, Bryde's whale (Balaenoptera edeni) and the humpback whale (Megaptera novaeanglia). Both species have large eyes characterised by a thickened cornea, a heavily thickened sclera, a highly vascularised fibro-adipose bundle surrounding the optic nerve at the back of the eye, and a reflective blue-green tapetum fibrosum. Using stereology and retinal whole mounts, we estimate a total of 274,268 and 161,371 RGCs in the Bryde's whale and humpback whale retinas, respectively. Both species have a similar retinal topography, consisting of nasal and temporal areas of high RGC density, suggesting that having higher visual acuity in the anterior and latero-caudal visual fields is particularly important in these animals. The temporal area is larger in both species and contains the peak RGC densities (160 cells mm-2 in the humpback whale and 200 cells mm-2 in Bryde's whale). In the Bryde's whale retina, the two high-density areas are connected by a weak centro-ventral visual streak, but such a specialisation is not evident in the humpback whale. Measurements of RGC soma area reveal that although the RGCs in both species vary substantially in size, RGC soma area is inversely proportional to RGC density, with cells in the nasal and temporal high-density areas being relatively more homogeneous in size compared to the RGCs in the central retina and the dorsal and ventral retinal periphery. Some of the RGCs were very large, with soma areas of over 2,000 µm2. Using peak RGC density and eye axial diameter (Bryde's whale: 63.5 mm; humpback whale: 48.5 mm), we estimated the peak anatomical spatial resolving power in water to be 4.8 cycles/degree and 3.3 cycles/degree in the Bryde's whale and the humpback whale, respectively. Overall, our findings for these two species are very similar to those reported for other species of cetaceans. This indicates that, irrespective of the significant differences in body size and shape, behavioural ecology and feeding strategy between mysticetes and odontocetes (toothed whales), cetacean eyes are adapted to vision in dim light and adhere to a common "bauplan" that evolved prior to the divergence of the two cetacean parvorders (Odontoceti and Mysticeti) over 30 million years ago.

Retinal Morphology and Visual Specializations in Three Species of Chimaeras, the Deep-Sea R. pacifica and C. lignaria, and the Vertical Migrator C. milii (Holocephali).

The majority of holocephalans live in the mesopelagic zone of the deep ocean, where there is little or no sunlight, but some species migrate to brightly lit shallow waters to reproduce. This study compares the retinal morphology of two species of deep-sea chimaeras, the Pacific spookfish (Rhinochimaera pacifica) and the Carpenter's chimaera (Chimaera lignaria), with the elephant shark (Callorhinchus milii), a vertical migrator that lives in the mesopelagic zone but migrates to shallow water to reproduce. The two deep-sea chimaera species possess pure rod retinae with long photoreceptor outer segments that might serve to increase visual sensitivity. In contrast, the retina of the elephant shark possesses rods, with an outer-segment length significantly shorter (a mean of 34 µm) than in the deep-sea species, and cones, and therefore the potential for color vision. The retinal ganglion cell distribution closely follows that of the photoreceptor populations in all three species, but there is a lower peak density of these cells in both deep-sea species (215-275 cells/mm2 vs. 769 cells/mm2 in the elephant shark), which represents a significant increase in the convergence of visual information (summation ratio) from photoreceptors to ganglion cells. It is evident that the eyes of deep-sea chimaeras have increased sensitivity to detect objects under low levels of light, but at the expense of both resolution and the capacity for color vision. In contrast, the elephant shark has a lower sensitivity, but the potential for color discrimination and a higher visual acuity.

Evolution of Vertebrate Phototransduction: Cascade Activation.

We applied high-throughput sequencing to eye tissue from several species of basal vertebrates (a hagfish, two species of lamprey, and five species of gnathostome fish), and we analyzed the mRNA sequences for the proteins underlying activation of the phototransduction cascade. The molecular phylogenies that we constructed from these sequences are consistent with the 2R WGD model of two rounds of whole genome duplication. Our analysis suggests that agnathans retain an additional representative (that has been lost in gnathostomes) in each of the gene families we studied; the evidence is strong for the G-protein α subunit (GNAT) and the cGMP phosphodiesterase (PDE6), and indicative for the cyclic nucleotide-gated channels (CNGA and CNGB). Two of the species (the hagfish Eptatretus cirrhatus and the lamprey Mordacia mordax) possess only a single class of photoreceptor, simplifying deductions about the composition of cascade protein isoforms utilized in their photoreceptors. For the other lamprey, Geotria australis, analysis of the ratios of transcript levels in downstream and upstream migrant animals permits tentative conclusions to be drawn about the isoforms used in four of the five spectral classes of photoreceptor. Overall, our results suggest that agnathan rod-like photoreceptors utilize the same GNAT1 as gnathostomes, together with a homodimeric PDE6 that may be agnathan-specific, whereas agnathan cone-like photoreceptors utilize a GNAT that may be agnathan-specific, together with the same PDE6C as gnathostomes. These findings help elucidate the evolution of the vertebrate phototransduction cascade from an ancestral chordate phototransduction cascade that existed prior to the vertebrate radiation.

Electrophysiological measures of temporal resolution, contrast sensitivity and spatial resolving power in sharks.

In most animals, vision plays an important role in detecting prey, predators and conspecifics. The effectiveness of vision in assessing cues such as motion and shape is influenced by the ability of the visual system to detect changes in contrast in both space and time. Understanding the role vision plays in shark behaviour has been limited by a lack of knowledge about their temporal resolution, contrast sensitivity and spatial resolution. In this study, an electrophysiological approach was used to compare these measures across five species of sharks: Chiloscyllium punctatum, Heterodontus portusjacksoni, Hemiscyllium ocellatum, Mustelus mustelus and Haploblepharus edwardsii. All shark species were highly sensitive to brightness contrast and were able to detect contrast differences as low as 1.6%. Temporal resolution of flickering stimuli ranged from 28 to 44 Hz. Species that inhabit brighter environments were found to have higher temporal resolution. Spatial resolving power was estimated in C. punctatum, H. portusjacksoni and H. ocellatum and ranged from 0.10 to 0.35 cycles per degree, which is relatively low compared to other vertebrates. These results suggest that sharks have retinal adaptations that enhance contrast sensitivity at the expense of temporal and spatial resolution, which is beneficial for vision in dimly lit and/or low contrast aquatic environments.

Optic-nerve-transmitted eyeshine, a new type of light emission from fish eyes.

BACKGROUND: Most animal eyes feature an opaque pigmented eyecup to assure that light can enter from one direction only. We challenge this dogma by describing a previously unknown form of eyeshine resulting from light that enters the eye through the top of the head and optic nerve, eventually emanating through the pupil as a narrow beam: the Optic-Nerve-Transmitted (ONT) eyeshine. We characterize ONT eyeshine in the triplefin blenny Tripterygion delaisi (Tripterygiidae) in comparison to three other teleost species, using behavioural and anatomical observations, spectrophotometry, histology, and magnetic resonance imaging. The study's aim is to identify the factors that determine ONT eyeshine occurrence and intensity, and whether these are specifically adapted for that purpose. RESULTS: ONT eyeshine intensity benefits from locally reduced head pigmentation, a thin skull, the gap between eyes and forebrain, the potential light-guiding properties of the optic nerve, and, most importantly, a short distance between the head surface and the optic nerves. CONCLUSIONS: The generality of these factors and the lack of specifically adapted features implies that ONT eyeshine is widespread among small fish species. Nevertheless, its intensity varies considerably, depending on the specific combination and varying expression of common anatomical features. We discuss whether ONT eyeshine might affect visual performance, and speculate about possible functions such as predator detection, camouflage, and intraspecific communication.

Retinal temporal resolution and contrast sensitivity in the parasitic lamprey Mordacia mordax and its non-parasitic derivative Mordacia praecox.

Lampreys and hagfishes are the sole extant representatives of the early agnathan (jawless) vertebrates. We compared retinal function of fully metamorphosed, immature Mordacia mordax (which are about to commence parasitic feeding) with those of sexually mature individuals of its non-parasitic derivative Mpraecox We focused on elucidating the retinal adaptations to dim-light environments in these nocturnally active lampreys, using electroretinography to determine the temporal resolution (flicker fusion frequency, FFF) and temporal contrast sensitivity of enucleated eyecups at different temperatures and light intensities. FFF was significantly affected by temperature and light intensity. Critical flicker fusion frequency (cFFF, the highest FFF recorded) of M. praecox and M. mordax increased from 15.1 and 21.8 Hz at 9°C to 31.1 and 36.9 Hz at 24°C, respectively. Contrast sensitivity of both species increased by an order of magnitude between 9 and 24°C, but remained comparatively constant across all light intensities. Although FFF values for Mordacia spp. are relatively low, retinal responses showed a particularly high contrast sensitivity of 625 in M. praecox and 710 in M. mordax at 24°C. This suggests selective pressures favour low temporal resolution and high contrast sensitivity in both species, thereby enhancing the capture of photons and increasing sensitivity in their light-limited environments. FFF indicated all retinal photoreceptors exhibit the same temporal response. Although the slow response kinetics (i.e. low FFF) and saturation of the response at bright light intensities characterise the photoreceptors of both species as rod-like, it is unusual for such a photoreceptor to be functional under scotopic and photopic conditions.

Functional diversity of the lateral line system among populations of a native Australian freshwater fish.

Fishes use their mechanoreceptive lateral line system to sense nearby objects by detecting slight fluctuations in hydrodynamic motion within their immediate environment. Species of fish from different habitats often display specialisations of the lateral line system, in particular the distribution and abundance of neuromasts, but the lateral line can also exhibit considerable diversity within a species. Here, we provide the first investigation of the lateral line system of the Australian western rainbowfish (Melanotaenia australis), a species that occupies a diversity of freshwater habitats across semi-arid northwest Australia. We collected 155 individuals from eight populations and surveyed each habitat for environmental factors that may contribute to lateral line specialisation, including water flow, predation risk, habitat structure and prey availability. Scanning electron microscopy and fluorescent dye labelling were used to describe the lateral line system in M. australis, and to examine whether the abundance and arrangement of superficial neuromasts (SNs) varied within and among populations. We found that the SNs of M. australis were present in distinct body regions rather than lines. The abundance of SNs within each body region was highly variable, and also differed among populations and individuals. Variation in SN abundance among populations was best explained by habitat structure and the availability of invertebrate prey. Our finding that specific environmental factors explain among-population variation in a key sensory system suggests that the ability to acquire sensory information is specialised for the particular behavioural needs of the animal.

Ontogenetic Shifts in Brain Organization in the Bluespotted Stingray Neotrygon kuhlii (Chondrichthyes: Dasyatidae).

Fishes exhibit lifelong neurogenesis and continual brain growth. One consequence of this continual growth is that the nervous system has the potential to respond with enhanced plasticity to changes in ecological conditions that occur during ontogeny. The life histories of many teleost fishes are composed of a series of distinct stages that are characterized by shifts in diet, habitat, and behavior. In many cases, these shifts correlate with changes in overall brain growth and brain organization, possibly reflecting the relative importance of different senses and locomotor performance imposed by the new ecological niches they encounter throughout life. Chondrichthyan (cartilaginous) fishes also undergo ontogenetic shifts in habitat, movement patterns, diet, and behavior, but very little is known about any corresponding shifts in the size and organization of their brains. Here, we investigated postparturition ontogenetic changes in brain-body size scaling, the allometric scaling of seven major brain areas (olfactory bulbs, telencephalon, diencephalon, optic tectum, tegmentum, cerebellum, and medulla oblongata) relative to the rest of the brain, and cerebellar foliation in a chondrichthyan, i.e., the bluespotted stingray Neotrygon kuhlii. We also investigated the unusual morphological asymmetry of the cerebellum in this and other batoids. As in teleosts, the brain continues to grow throughout life, with a period of rapid initial growth relative to body size, before slowing considerably at the onset of sexual maturity. The olfactory bulbs and the cerebellum scale with positive allometry relative to the rest of the brain, whereas the other five brain areas scale with varying degrees of negative allometry. None of the major brain areas showed the stage-specific differences in rates of growth often found in teleosts. Cerebellar foliation also increases at a faster rate than overall brain growth. We speculate that changes in the olfactory bulbs and cerebellum could reflect increased olfactory and locomotor capabilities, which may be associated with ontogenetic shifts in diet, habitat use, and activity patterns, as well as shifts in behavior that occur with the onset of sexual maturity. The frequency distributions of the three cerebellar morphologies exhibited in this species best fit a 2:1:1 (right-sided:left-sided:intermediate) distribution, mirroring previous findings for another stingray species.

The Central Nervous System of Jawless Vertebrates: Encephalization in Lampreys and Hagfishes.

Lampreys and hagfishes are the sole surviving representatives of the early agnathan (jawless) stage in vertebrate evolution, which has previously been regarded as the least encephalized group of all vertebrates. Very little is known, however, about the extent of interspecific variation in relative brain size in these fishes, as previous studies have focused on only a few species, even though lampreys exhibit a variety of life history traits. While some species are parasitic as adults, with varying feeding behaviors, others (nonparasitic species) do not feed after completing their macrophagous freshwater larval phase. In addition, some parasitic species remain in freshwater, while others undergo an anadromous migration. On the basis of data for postmetamorphic individuals representing approximately 40% of all lamprey species, with representatives from each of the three families, the aforementioned differences in life history traits are reflected in variations in relative brain size. Across all lampreys, brain mass increases with body mass with a scaling factor or slope (α) of 0.35, which is less than those calculated for different groups of gnathostomatous (jawed) vertebrates (α = 0.43-0.62). When parasitic and nonparasitic species are analyzed separately, with phylogeny taken into account, the scaling factors of both groups (parasitic α = 0.43, nonparasitic α = 0.45) approach those of gnathostomes. The relative brain size in fully grown adults of parasitic species is, however, less than that of the adults of nonparasitic species, paralleling differences between fully grown adults and recently metamorphosed individuals of anadromous species. The average degree of encephalization is found in anadromous parasitic lampreys and might thus represent the ancestral condition for extant lampreys. These results suggest that the degree of encephalization in lampreys varies according to both life history traits and phylogenetic relationships.