A spitting image: specializations in archerfish eyes for vision at the interface between air and water

Archerfish are famous for spitting jets of water to capture terrestrial insects, a task that not only requires oral dexterity, but also the ability to detect small camouflaged prey against a visually complex background of overhanging foliage. Because detection of olfactory, auditory and tactile cues is diminished at air–water interfaces, archerfish must depend almost entirely on visual cues to mediate their sensory interactions with the aerial world. During spitting, their eyes remain below the water''s surface and must adapt to the optical demands of both aquatic and aerial fields of view. These challenges suggest that archerfish eyes may be specially adapted to life at the interface between air and water. Using microspectrophotometry to characterize the spectral absorbance of photoreceptors, we find that archerfish have differentially tuned their rods and cones across their retina, correlated with spectral differences in aquatic and aerial fields of view. Spatial resolving power also differs for aquatic and aerial fields of view with maximum visual resolution (6.9 cycles per degree) aligned with their preferred spitting angle. These measurements provide insight into the functional significance of intraretinal variability in archerfish and infer intraretinal variability may be expected among surface fishes or vertebrates where different fields of view vary markedly.     

Characterization of the Aerial Escape Response of the African Butterfly Fish, Pantodon buchholzi Peters

We studied the startle response of the African butterfly fish, Pantodon buchholzi (Osteoglossomorpha, Osteoglossoidea). It is an upward movement, mediated by abduction of the pectoral fins, and is elicited by mechanical and visual stimuli. Because this fish inhabits the first few centimeters beneath the water surface, its startle response results in an aerial excursion that may be described as ballistic-like, following a motion as defined by linear acceleration. We show that the aerial excursion is well-modeled by a parabola. On average, a fish jumps no more than twice its height and travels horizontally about five times its standard length. The fish may exhibit variable in-flight trunk and fin movements, but neither increases the travel distance in air following the initial in-water propulsive event. Similar vertical jumps also occur entirely within the water column suggesting that this motor behavior of Pantodon is a general escape behavior analogous to a Mauthner neuron-induced escape response. The variability in its posture in air and its direction of motion after reentering the water enhances this act of vertical flight as a step in this fish's escape behavior. The aerial aspect of its escape behavior is only a consequence of its position in the water column.     

The Evolution of Lung-Gill Bimodal Breathing and the Homology of Vertebrate Respiratory Pumps

SYNOPSIS. A comparative analysis of actinopterygian and sarcopterygianaerial buccal pumps indicates that the primitive pattern ofair transfer differs fundamentally between these two clades.Actinopterygian fishes ventilate their lungs with a four-strokebuccal pump: the buccal cavity expands and fills with expiredair, compresses to expel expired air, expands again to takein fresh air, and then compresses again to pump fresh air intothe lungs. Lungfishes, caudates, and anurans expand and compressthe buccal cavity only once per expiratory-inspiratory cycle,and thus use a two-stroke pump. Both of these bidirectional,aerial buccal pumps evolved from unidirectional, aquatic buccalpumps. The two-stroke aerial pump and the primitive aquaticpump used for gill irrigation share slow movements and may bothbe triggered by the same central rhythm generator. These similaritiessuggest that the two-stroke buccal pump evolved from the gillirrigation pump. Similarly, the four-stroke pump shares rapidmovement and afferent triggering with aquatic suction feedingand coughing, suggesting that the four-stroke pump may haveevolved from a combination of two suction feeding or coughingmovements. Thus the differences between the actinopterygianand sarcopterygian aerial buccal pumps may be due to their independentevolution from different aquatic buccal pumps, rather than dueto divergence from a single aerial buccal pump.     

On the structure of the aerial visual field of aquatic animals distorted by refraction

The aerial visual field of aquatic animals living near the water surface is distorted by refraction. The imaging of aerial objects by one or two submerged eyes is studied. The aerial binocular image field is determined for pairs of submerged eyes in horizonal and vertical planes. These two image spaces have significantly different structures. Aquatic animals have to correct for refraction, adapting themselves to the former aerial image field in order to recognize aerial predators or to capture such prey. The other aerial image space is only of theoretical interest.     

Dependence on aerial respiration and its influence on microdistribution in the invasive freshwater snail <Emphasis Type="Italic">Pomacea canaliculata</Emphasis> (Caenogastropoda,Ampullariidae)

The invasive Neotropical snail Pomacea canaliculata is usually regarded as amphibious, although the relative significance of aerial and aquatic respiration is unknown. To investigate the degree of dependence on aerial respiration and its influences on microdistribution, experiments were performed in the laboratory and under seminatural and natural conditions. Restriction of aerial respiration negatively affected survivorship, activity and feeding, its effects worsening with temperature and water fouling; females were more seriously affected than males although the effect depended on reproductive effort. Snails were unevenly distributed relative to the access to air, both in a stream and in an outdoor tank, being concentrated less than 2–4 m from the nearest emergent substratum. Accessibility to air would be an important trait of waterbodies prone to invasions of P. canaliculata, especially in tropical areas. The effectiveness of some control measures could be increased by focusing on areas where snails are concentrated due to their dependence on air.     

Effect of cattail (<Emphasis Type="Italic">Typha</Emphasis> spp<Emphasis Type="Italic">.</Emphasis>) mowing on water beetle assemblages: changes of environmental factors and the aerial colonization of aquatic habitats

Some studies found no, or weak evidence that dense monotypic cattail (Typha spp.) stands exclude water beetle species from aquatic habitats, or modify aquatic beetle assemblages. Other studies suggest that cattail may reduce the chance of aerial water beetle colonization, and decreases water temperature; negatively affecting these insects. We examined the response of aquatic beetle assemblages to the mowing of cattail (Typha angustifolia L., T. latifolia L.) in a freshwater marsh. Following removal of cattail at the water level in experimental plots, aquatic beetles were sampled both in mowed and intact (control) plots weekly, through a month in the spring of 2008. Aquatic beetles were more abundant in mowed plots. Species richness was the same, but it showed different patterns in mowed and intact plots. Shannon’s diversity was similar between treatments, while evenness was lower in mowed plots. 29% of the aquatic beetles showed a strong preference for mowed plots, and 15% preferred the control plots. Water temperature was an important factor, with mowed plots having higher water temperatures because of increased solar radiation. Polarization visibility of the water surface was also a factor, since aerially colonizing (flying) aquatic beetles use horizontally polarized light reflected from the water surface to seek potential locations. Using imaging polarimetry, we showed that mowing strongly enhanced the water-reflected polarized light signal, because it reduced the screening effect of cattail leaves, which made the visual detection of water easier. Our results suggest that cattail mowing is a useful method in aquatic beetle conservation: it increases the chance of aerial colonization due to the enhanced polarization visibility of the water surface, and creates a habitat for more abundant assemblages otherwise excluded by the monodominant dense cattail stands. Thus, sustaining hemi-marsh conditions with vegetated and mowed areas is advisable to maximize overall aquatic beetle diversity.     

The eye of the humboldt penguin, Spheniscus humboldti: visual fields and schematic optics

Construction of a schematic eye indicates that the eye of Spheniscus humboldti is aquatic in design. The lens has a power of 100 dioptres (D) while (in air) the cornea has a power of 29 D. In air, the eye is myopic (approximately 28 D) but in water it is emmetropic. Minimum pupil size would seem insufficient to allow the pupil to function as a stenopaic aperture and increase depth of focus sufficiently to overcome the eye's aerial myopia. Entry into water reduces maximum image brightness by approximately three times. In air, the maximum width of the retinal binocular field is 45 degrees and this occurs approximately 10 degrees above the line of the bill. The bill intrudes into the retinal field and binocular field width in the plane containing the bill and the optic axes is 28 degrees. The vertical extent of the binocular field is 125 degrees. In the plane containing the optic axes the cyclopean field equals 282 degrees and the optic axes diverge by 116 degrees. In this plane the mean uniocular field is 155 degrees with the temporal hemifield approximately 11 degrees larger than the nasal hemifield. Entry into water reduces the widths of the visual fields such that maximum binocular field width is only 17 degrees and the vertical extent is reduced to about 80 degrees. Binocular vision is lost in the plane of the bill, and the uniocular retinal field is reduced by 32 degrees and the cyclopean field by 36 degrees.     

Aerial and aquatic visual acuity of the grey bichir Polypterus senegalus,as estimated by optokinetic response

The present study assessed the aerial and aquatic visual abilities of juvenile grey bichir Polypterus senegalus, fish capable of terrestrial locomotion, by measuring the optokinetic response to stimuli of varying speed and spatial frequency. In water, fish tracked slow-moving (2° s⁻¹) stimuli moderately well and fast-moving stimuli very poorly. Spatial acuity was very low compared with many other species, with maximum response observed at 0.05–0.075 stimulus cycles per degree of visual arc; however, it should be noted that adult fish, with their larger eyes, are likely to have somewhat improved spatial acuity. Low spatial acuity and limited stimulus tracking ability might be expected in a nocturnal ambush predator such as P. senegalus, where gaze stabilization may be less crucial and other sensory inputs may have greater importance in perception of the environment. In air, spatial and temporal acuity were both poorer by every measure, but some visual ability persisted. As the eye shows no anatomical specialization for aerial vision, poor vision was expected; however, the large decrease in saccade velocity observed in air trials was unexpected. Stimulus parameters typically have little effect on the characteristics of the saccade, so this finding may suggest that the function of the reflex system itself could be compromised in the aerial vision of some fishes capable of terrestrial locomotion.     

Visual fields in Blue Ducks Hymenolaimus malacorhynchos and Pink-eared Ducks Malacorhynchus membranaceus: visual and tactile foraging

Blue Ducks Hymenolaimus malacorhynchos (Anatidae), an IUCN Red Listed Endangered species, reside in headwaters of New Zealand rivers and feed primarily on aquatic invertebrates. However, whether such food items are detected by tactile or visual cues is unknown. That Blue Ducks may use tactile cues when foraging is suggested by the presence of specialized flaps of thickened, keratinized epidermis containing Herbst's corpuscles along the ventral margins of the upper mandibles near the bill tip. Similar bill flaps are found only in one other duck species, Pink-eared Ducks Malacorhynchus membranaceus , that surface filter-feed on a range of planktonic organisms. Using an ophthalmoscopic reflex technique we determined the visual fields of both species. In Blue Ducks the eyes are frontally placed resulting in a relatively wide binocular field into which the narrow tapering bill intrudes. There is a large blind area to the rear of the head. This visual field topography is similar to that of other visually guided foragers including those that take mobile prey from the water column, e.g. penguins (Spheniscidae). By contrast, Pink-eared Duck visual fields show features found in other tactile feeding ducks: a narrow frontal binocular field with the bill falling at the periphery, and comprehensive visual coverage of the celestial hemisphere. We conclude that although Blue Ducks may take prey from rock surfaces they are primarily visual feeders of the water column and we suggest therefore that their foraging may be significantly disrupted by changes in water clarity. This introduces a previously unconsidered factor into the selection of sites for population enhancement or re-introductions, a current conservation focus.     

Tolerance of forced air emergence by a fish with a broad vertical distribution, the rockpool blenny, Hypsoblennius gilberti (Blenniidae)

Many intertidal fishes, particularly among the Blenniidae and Cottidae, possess amphibious adaptations, including the ability to breathe in air and to avoid desiccation in terrestrial conditions. These traits are absent in subtidal species of blennies and cottids. Hypsoblennius gilberti, the rockpool blenny, is found in shallow rockpools in the mid to high intertidal areas of Southern California, and deeper to 18 m in the subtidal zone. This broad vertical distribution could indicate that this blenny is adapted for tidal air emergence, although H. gilberti has not been observed out of water in its natural habitat. H. gilberti does not emerge voluntarily from hypoxic sea water in the laboratory, but it easily withstands 3 h out of water. The aerial respiratory exchange ratio (CO2 released compared to O2 consumed) is 0.70, similar to that of amphibious intertidal fishes in air, indicating sufficient release of metabolically produced CO2 while emerged. There is no increase in aquatic respiration following emergence. However, unlike other amphibious fishes that maintain aerial oxygen consumption at a level similar to aquatic oxygen consumption, H. gilberti has an aerial oxygen consumption rate one-third that in water. H. gilberti can recover rapidly from terrestrial water loss, and shows no change in evaporative water loss rates at 93% and 77% relative humidities. The amphibious capabilities in H. gilberti, even if rarely used, permit survival in air during tidal emergence. These findings suggest that H. gilberti may demonstrate an intermediate condition between the amphibious species of intertidal fishes that regularly emerge from water, and the subtidal fishes that do not survive air emergence and are completely restricted to an aquatic habitat.     

Aquatic feeding by moose Alces alces in a Canadian lake

The aquatic feeding behaviour of moose, and the abundance, species, and chemical composition of aquatic plants, were studied in a small Canadian lake which attracted many animals. Feeding was much more common in June than later in the summer, and somewhat more common during the morning and evening than in (he afternoon. Individual adult moose appeared to use the lake intensively during 1- to 4-day visits. Compared to deciduous browse, the aquatic plants had high levels of sodium and iron, less fat, and similar levels of crude protein, crude fibre, sulphate and other minerals. In the preferred feeding areas, compared to other parts of the lake, plants were more abundant, had a different species composition, and were richer in iron and calcium. Recent flooding, a flow of water through the feeding areas, and a bicarbonate-rich tributary may all have contributed to the lake's attraction for moose.     

Anti-Predator Behavioral Responses of Mosquito Pupae to Aerial Predation Risk

Aquatic insects have two potential sources of predation risk: aquatic predators and aerial predators. Our goal was to assess anti-predator responses of Culex pipiens to aerial predation. By simulating predator attacks, we assessed (a) the distance fled in relation to depth and group size, (b) the distribution of individuals at different depths, and (c) the duration of surfacing events to obtain air in scenarios with varying predation risk. Pupae located closer to the surface fled deeper into the water, and the number of conspecifics decreased the distance fled. When the risk of predation increased, more individuals were found deeper in the water column, and the interval between two consecutive surfacing events increased. Culex pipiens shows a trade-off between avoiding aerial predation and maintaining oxygen acquisition, which may be regulated by the need to conserve energy reserves.     

The visual system of the guitar fish (Rhinobatos productus)

Summary The retinal projections in adult and juvenile guitar fish (Rhinobatos productus) were determined with the aid of the Nauta-Fink-Heimer techniques. The visual system was found to be more extensive and more differentiated than in any other elasmobranch studied to date. Massive projections exist to the dorsal and ventral thalamus, tectum and pretectum, in addition to the usual weak contributions to the hypothalamus and the ventral mesencephalic tegmentum. The projection to the lateral tectum is significantly less distinct than that to the medial part of this structure, suggesting that the ventral visual field has a smaller input, perhaps due to the fact that this visual field is normally aimed at the body of this flat fish.     

Larval size constraints determine directional ontogenetic shifts in the visual system of teleosts1

We summarize characteristic sequences of morphological change in the teleost visual system from larvae to large adults at the level of the retina, the optic tract and the optic tectum. These shifts include sizes and ratios of cone and rod receptor cells, sizes and types of retinal ganglion cells and optic tract fibers as well as features of the optic tectum. Teleost larvae are the smallest vertebrates known. We suggest that the utilization of color contrasts as an adaptive benefit dictates the starting point of morophological development, which is a pure cone retina in most fish larvae. The direction of morphological and functional shifts in the teleost visual system during growth is determined by continuous retinal stretch, which allows for improving visual abilities. The larval visual system probably provides just adequate photopic (cone-)acuity for plankton feeding, but limited space in the retina hampers optimization of both, photopic resolving power and sensitivity Limited space also Irevents the simultaneous development of the scotopic (rod-)system. Over a wide range of body sizes, morphological parameters change, photopic and scotopic resolving power, acuity and sensitivity improve. Size constraints in the teleost visual system and lifefong shifts in sensory capacities are discussed with respect to ecology and the niche concept.     

The breathing pattern and the ventilatory response to aquatic and aerial hypoxia and hypercarbia in the frog Pipa carvalhoi

Anuran amphibians are known to exhibit an intermittent pattern of pulmonary ventilation and to exhibit an increased ventilatory response to hypoxia and hypercarbia. However, only a few species have been studied to date. The aquatic frog Pipa carvalhoi inhabits lakes, ponds and marshes that are rich in nutrients but low in O(2). There are no studies of the respiratory pattern of this species and its ventilation during hypoxia or hypercarbia. Accordingly, the aim of the present study was to characterize the breathing pattern and the ventilatory response to aquatic and aerial hypoxia and hypercarbia in this species. With this purpose, pulmonary ventilation (V(I)) was directly measured by the pneumotachograph method during normocapnic normoxia to determine the basal respiratory pattern and during aerial and aquatic hypercarbia (5% CO(2)) and hypoxia (5% O(2)). Our data demonstrate that P. carvalhoi exhibits a periodic breathing pattern composed of single events (single breaths) of pulmonary ventilation separated by periods of apnea. The animals had an enhanced V(I) during aerial hypoxia, but not during aquatic hypoxia. This increase was strictly the result of an increase in the breathing frequency. A pronounced increase in V(I) was observed if the animals were simultaneously exposed to aerial and aquatic hypercarbia, whereas small or no ventilatory responses were observed during separately administered aerial or aquatic hypercarbia. P. carvalhoi primarily inhabits an aquatic environment. Nevertheless, it does not respond to low O(2) levels in water, although it does so in air. The observed ventilatory responses to hypercarbia may indicate that this species is similar to other anurans in possessing central chemoreceptors.     

High temperature acclimation alters the emersion behavior in the crab Neohelice granulata

An increase in environmental temperature can deleteriously affect organisms. This study investigated whether the semiterrestrial estuarine crab Neohelice granulata uses emersion behavior as a resource to avoid thermal stress and survive higher aquatic temperatures. We also examined whether this behavior is modulated by exposure to high temperature; whether, during the period of emersion, the animal loses heat from the carapace to the medium; and whether this behavior is altered by the temperature at which the animal has been acclimated. The lethal temperature for 50% of the population (LT₅₀) was determined through 96-h mortality curves in animals acclimated at 20 °C and 30 °C. The behavioral profile of N. granulata during thermal stress was based on monitoring crab movement in aerial, intermediary, and aquatic zones. Acclimation at a higher temperature and the possibility of emersion increased the thermotolerance of the crabs and the synergistic effect of acclimation temperature. The possibility of leaving the hot water further increased the resistance of these animals to thermal stress. We observed that when the crab was subjected to thermal stress conditions, it spent more time in the aerial environment, unlike under control conditions. Under the experimental conditions, it made small incursions into the aquatic environment and stayed in the aerial environment for a longer time in order to cool its body temperature. The animals acclimated at 20 °C and placed into water at 35 °C remained in the aerial zone. The animals acclimated and maintained at 30 °C (control) that were placed in water at 35 °C with the possibility of emerging into hot air transited more frequently between the aquatic and aerial zones than did the animals that were put in water at 35 °C with the possibility of emerging into a cooler air environment. We conclude that emergence behavior allows N. granulata to survive high temperatures and that this behavior is influenced by acclimation temperature.     

Visual accommodation and active pursuit of prey underwater in a plunge-diving bird: the Australasian gannet

Australasian gannets (Morus serrator), like many other seabird species, locate pelagic prey from the air and perform rapid plunge dives for their capture. Prey are captured underwater either in the momentum (M) phase of the dive while descending through the water column, or the wing flapping (WF) phase while moving, using the wings for propulsion. Detection of prey from the air is clearly visually guided, but it remains unknown whether plunge diving birds also use vision in the underwater phase of the dive. Here we address the question of whether gannets are capable of visually accommodating in the transition from aerial to aquatic vision, and analyse underwater video footage for evidence that gannets use vision in the aquatic phases of hunting. Photokeratometry and infrared video photorefraction revealed that, immediately upon submergence of the head, gannet eyes accommodate and overcome the loss of greater than 45 D (dioptres) of corneal refractive power which occurs in the transition between air and water. Analyses of underwater video showed the highest prey capture rates during WF phase when gannets actively pursue individual fish, a behaviour that very likely involves visual guidance, following the transition after the plunge dive's M phase. This is to our knowledge the first demonstration of the capacity for visual accommodation underwater in a plunge diving bird while capturing submerged prey detected from the air.     

Influence of temperature and size on the respiration of a tropical intertidal gastropod,Morula granulata (Duclos), in air and water

Summary The effect of temperature on the size related metabolism of a tropical intertidal gastropod,Morula granulata (Duclos), was studied both in air and water. An increase was observed in the aerial and aquatic respiration with increasing size at all temperatures studied. Significant differences (P<0.05) were found between the regression values (b) and also the intercept values (a) of aerial and aquatic respiration at different temperatures. The aerial regression values at any given temperature were significantly higher (P<0.05) than the aquatic values. Aerial and aquatic respiration ofM. granulata at different temperatures are compared and discussed in relation to the habitat of the animal.     

Ventilation of the branchial chambers in the amphibious West African freshwater crab,Sudanonautes (Convexonautes) aubryi monodi (Balss, 1929) (Brachyura,Potamonautidae)

The anatomy of the respiratory system of the savanna-zone African freshwater crab, Sudanonautes (Convexonautes) aubryi monodi [Balss, 1929], has been examined and has been found to be adapted for both aerial and aquatic gas exchange. The activities of the scaphognathites and the directions of flow of the ventilatory stream have been recorded in stressed, active and resting specimens during their exposure to a wide range of conditions from deep water to dry land.Ventilation of the branchial chambers during aquatic gas exchange in Sudanonautes kept in deep water is shown to consist of a rapid, predominantly forward water flow similar to that of fully-aquatic species. Ventilation of the branchial chambers during aerial gas exchange in Sudanonautes on land is shown to consist of a relatively slow forward air flow. This flow is continuous in post-operative crabs, pulsatile in active crabs and completely immobile in resting crabs.A second method of ventilation of the branchial chambers during aerial gas exchange is shown to consist of a pulsatile reversed air flow. This occurs (1) when Sudanonautes is kept in very shallow water and active or stressed; (2) when it has recently moved on to land; and (3) when it is completely immersed and exhibiting aerial gas exchange under water. The unusual phenomenon of aerial gas exchange under water is reported here for the first time in any species of crab.Bimodal ventilation of the branchial chambers occurs in stressed or active crabs partly immersed in shallow water. This consists of an alternation between forward water flow and reversed air flow.The morphology of the branchial chambers in Sudanonautes, and observational data on the patterns of ventilation of the branchial chambers, are discussed in relation to those described for other air-breathing decapod crustaceans.     

Ein septum papillaris im auge von Pantodon buchholzi pet. (Teleostei,Osteoglossiformes)

A horizontal septum subdivides the eye of the surface feeding fish Pantodon buchholzi into an upper and lower part. The numbers and ratio of the receptor cells and succeeding neurons are different in both parts. The functional importance of these facts is discussed.