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Journal: The Journal of experimental biology


Many functions have been postulated for the aerodynamic role of the avian tail during steady-state flight. By analogy with conventional aircraft, the tail might provide passive pitch stability if it produced very low or negative lift. Alternatively, aeronautical principles might suggest strategies that allow the tail to reduce inviscid, induced drag: if the wings and tail act in different horizontal planes, they might benefit from biplane-like aerodynamics; if they act in the same plane, lift from the tail might compensate for lift lost over the fuselage (body), reducing induced drag with a more even downwash profile. However, textbook aeronautical principles should be applied with caution because birds have highly capable sensing and active control, presumably reducing the demand for passive aerodynamic stability, and, because of their small size and low flight speeds, operate at Reynolds numbers two orders of magnitude below those of light aircraft. Here, by tracking up to 20,000, 0.3 mm neutrally buoyant soap bubbles behind a gliding barn owl, tawny owl and goshawk, we found that downwash velocity due to the body/tail consistently exceeds that due to the wings. The downwash measured behind the centreline is quantitatively consistent with an alternative hypothesis: that of constant lift production per planform area, a requirement for minimizing viscous, profile drag. Gliding raptors use lift distributions that compromise both inviscid induced drag minimization and static pitch stability, instead adopting a strategy that reduces the viscous drag, which is of proportionately greater importance to lower Reynolds number fliers.


Crocodilians are among the most vocal non-avian reptiles. Adults of both sexes produce loud vocalizations known as ‘bellows’ year round, with the highest rate during the mating season. Although the specific function of these vocalizations remains unclear, they may advertise the caller’s body size, because relative size differences strongly affect courtship and territorial behaviour in crocodilians. In mammals and birds, a common mechanism for producing honest acoustic signals of body size is via formant frequencies (vocal tract resonances). To our knowledge, formants have to date never been documented in any non-avian reptile, and formants do not seem to play a role in the vocalizations of anurans. We tested for formants in crocodilian vocalizations by using playbacks to induce a female Chinese alligator (Alligator sinensis) to bellow in an airtight chamber. During vocalizations, the animal inhaled either normal air or a helium/oxygen mixture (heliox) in which the velocity of sound is increased. Although heliox allows normal respiration, it alters the formant distribution of the sound spectrum. An acoustic analysis of the calls showed that the source signal components remained constant under both conditions, but an upward shift of high-energy frequency bands was observed in heliox. We conclude that these frequency bands represent formants. We suggest that crocodilian vocalizations could thus provide an acoustic indication of body size via formants. Because birds and crocodilians share a common ancestor with all dinosaurs, a better understanding of their vocal production systems may also provide insight into the communication of extinct Archosaurians.

Concepts: Reptile, Acoustics, Sound, Resonance, Crocodilia, Alligatoridae, Formant, Alligator


While evidence suggests that warming may impact cognition of ectotherms, the underlying mechanisms remain poorly understood. A possible, but rarely considered mechanism is that metabolic response of ectotherms to warming associate with changes in brain morphology and functioning. Here we compared aerobic metabolism, volume of brain, boldness, and accuracy of maze solving of common minnows (Phoxinus phoxinus) acclimated for eight months to either their current optimal natural (14 °C) or warm (20 °C) water temperature. Metabolic rates indicated increased energy expenditure in warm acclimated fish, but also at least partial thermal compensation as warm acclimate fish maintained high aerobic scope. Warm acclimated fish had larger brain than cool acclimated fish. Volume of dorsal medulla relative to the overall brain size was larger in warm than cool acclimated fish, but proportion of other brain regions did not differ between the temperature treatments. Warm acclimated fish did not differ in boldness but did more errors than cool acclimated fish in exploring the maze across four trials. Inter-individual differences in numbers of exploration errors were repeatable across the four trials of the maze test. Our findings suggest that in warm environments, maintaining a high aerobic scope which is important for the performance of physically demanding tasks, can come at the cost of changes in brain morphology and impairment of the capacity to explore novel environments. This trade-off could have strong fitness implications for wild ectotherms.


Many animals live in groups because of the potential benefits associated with defense and foraging. Group living may also induce a ‘calming effect’ on individuals, reducing overall metabolic demand. This effect could occur by minimising the need for individual vigilance and reducing stress through social buffering. However, this effect has proved difficult to quantify. We examined the effect of shoaling on metabolism and body condition in the gregarious damselfish Chromis viridis Using a novel respirometry methodology for social species, we found that the presence of shoal-mate visual and olfactory cues led to a reduction in the minimum metabolic rate of individuals. Fish held in isolation for 1 week also exhibited a reduction in body condition when compared with those held in shoals. These results indicate that social isolation as a result of environmental disturbance could have physiological consequences for gregarious species.

Concepts: Photosynthesis, Carbon dioxide, Metabolism, Energy, Cellular respiration, Nicotinamide adenine dinucleotide, Coral, Coral reef


A prominent feature of gliding flight in snakes of the genus Chrysopelea is the unique cross-sectional shape of the body, which acts as the lifting surface in the absence of wings. When gliding, the flying snake Chrysopelea paradisi morphs its circular cross-section into a triangular shape by splaying its ribs and flattening its body in the dorsoventral axis, forming a geometry with fore-aft symmetry and a thick profile. Here, we aimed to understand the aerodynamic properties of the snake’s cross-sectional shape to determine its contribution to gliding at low Reynolds numbers. We used a straight physical model in a water tunnel to isolate the effects of 2D shape, analogously to studying the profile of an airfoil of a more typical flyer. Force measurements and time-resolved (TR) digital particle image velocimetry (DPIV) were used to determine lift and drag coefficients, wake dynamics and vortex-shedding characteristics of the shape across a behaviorally relevant range of Reynolds numbers and angles of attack. The snake’s cross-sectional shape produced a maximum lift coefficient of 1.9 and maximum lift-to-drag ratio of 2.7, maintained increases in lift up to 35 deg, and exhibited two distinctly different vortex-shedding modes. Within the measured Reynolds number regime (Re=3000-15,000), this geometry generated significantly larger maximum lift coefficients than many other shapes including bluff bodies, thick airfoils, symmetric airfoils and circular arc airfoils. In addition, the snake’s shape exhibited a gentle stall region that maintained relatively high lift production even up to the highest angle of attack tested (60 deg). Overall, the cross-sectional geometry of the flying snake demonstrated robust aerodynamic behavior by maintaining significant lift production and near-maximum lift-to-drag ratios over a wide range of parameters. These aerodynamic characteristics help to explain how the snake can glide at steep angles and over a wide range of angles of attack, but more complex models that account for 3D effects and the dynamic movements of aerial undulation are required to fully understand the gliding performance of flying snakes.

Concepts: Fluid dynamics, Aerodynamics, Snake, Lift, Airfoil, Angle of attack, Chrysopelea, Lift coefficient


The diurnal thermophilic Saharan silver ant, Cataglyphis bombycina, is the fastest of the North African Cataglyphis desert ant species. These highly mobile ants endure the extreme temperatures of their sand dune environment with outstanding behavioural, physiological and morphological adaptations. Surprisingly, C. bombycina has comparatively shorter legs than its well-studied sister species Cataglyphis fortis from salt pan habitats. This holds despite the somewhat hotter surface temperatures and the more yielding sand substrate. Here, we report that C. bombycina employs a different strategy in reaching high running speeds, outperforming the fastest known runs of the longer-legged C. fortis ants. Video analysis across a broad range of locomotor speeds revealed several differences to C. fortis Shorter leg lengths are compensated for by high stride frequencies, ranging beyond 40 Hz. This is mainly achieved by a combination of short stance phases (down to 7 ms) and fast leg swing movements (up to 1400 mm s-1). The legs of one tripod group exhibit almost perfect synchrony in the timings of their lift-offs and touch-downs, and good tripod coordination is present over the entire walking speed range (tripod coordination strength values around 0.8). This near synchrony in leg movement may facilitate locomotion across the yielding sand dune substrate.


Nociception allows for immediate reflex withdrawal whereas pain allows for longer-term protection via rapid learning. We examine here whether shore crabs placed within a brightly lit chamber learn to avoid one of two dark shelters when that shelter consistently results in shock. Crabs were randomly selected to receive shock or not prior to making their first choice and were tested again over 10 trials. Those that received shock in trial 2, irrespective of shock in trial 1, were more likely to switch shelter choice in the next trial and thus showed rapid discrimination. During trial 1, many crabs emerged from the shock shelter and an increasing proportion emerged in later trials, thus avoiding shock by entering a normally avoided light area. In a final test we switched distinctive visual stimuli positioned above each shelter and/or changed the orientation of the crab when placed in the chamber for the test. The visual stimuli had no effect on choice, but crabs with altered orientation now selected the shock shelter, indicating that they had discriminated between the two shelters on the basis of movement direction. These data, and those of other recent experiments, are consistent with key criteria for pain experience and are broadly similar to those from vertebrate studies.

Concepts: Shock, Blue crab, Portunidae, Carcinus maenas, Portunoidea, Carcinus, Carcinus aestuarii, Crustaceans of the Atlantic Ocean


During winter at temperate and high latitudes, the low ambient temperatures, limited food supplies and short foraging periods mean small passerines show behavioural, morphological and physiological adaptations to reduce the risk of facing energy shortages. Peripheral tissues vasoconstrict in low ambient temperatures to reduce heat loss and cold injury. Peripheral vasoconstriction has been observed with food restriction in captivity but has yet to be explored in free-ranging animals. We experimentally food restricted both wild and captive great tits (Parus major) during winter months and measured surface temperatures of the bill and eye region using thermal imaging, to investigate whether birds show rapid local heterothermic responses, which may reduce their thermoregulatory costs when facing a perceived imminent food shortage. Our results of a continuously filmed wild population showed that bill temperature was immediately reduced in response to food restriction compared with when food was available ad libitum, an apparent autonomic response. Such immediacy implies a ‘pre-emptive’ response before the bird experiences any shortfalls in energy reserves. We also demonstrate temporal variation in vasoconstriction of the bill, with bill temperature gradually rising throughout the food restriction after the initial drop. Eye-region temperature in the wild birds remained at similar levels throughout food restriction compared with unrestricted birds, possibly reflecting the need to maintain steady circulation to the central nervous and visual systems. Our findings provide evidence that birds selectively allow the bill to cool when a predictable food supply is suddenly disrupted, probably as a means of minimising depletion of body reserves for a perceived future shortage in energy.


Mammalian hair cells possess only a limited ability to repair damage after trauma. In contrast, sea anemones show a marked capability to repair damaged hair bundles by means of secreted repair proteins (RPs). Previously, it was found that recovery of traumatized hair cells in blind cavefish was enhanced by anemone-derived RPs; therefore, the ability of anemone RPs to assist recovery of damaged hair cells in mammals was tested here. After a 1 h incubation in RP-enriched culture media, uptake of FM1-43 by experimentally traumatized murine cochlear hair cells was restored to levels comparable to those exhibited by healthy controls. In addition, RP-treated explants had significantly more normally structured hair bundles than time-matched traumatized control explants. Collectively, these results indicate that anemone-derived RPs assist in restoring normal function and structure of experimentally traumatized hair cells of the mouse cochlea.

Concepts: Animal, Cochlea, Mammal, Murinae, Rodent, Mouse, Hair cell, Sea anemone


Although the functional properties of shark skin have been of considerable interest to both biologists and engineers because of the complex hydrodynamic effects of surface roughness, no study to date has successfully fabricated a flexible biomimetic shark skin that allows detailed study of hydrodynamic function. We present the first study of the design, fabrication and hydrodynamic testing of a synthetic, flexible, shark skin membrane. A three-dimensional (3D) model of shark skin denticles was constructed using micro-CT imaging of the skin of the shortfin mako (Isurus oxyrinchus). Using 3D printing, thousands of rigid synthetic shark denticles were placed on flexible membranes in a controlled, linear-arrayed pattern. This flexible 3D printed shark skin model was then tested in water using a robotic flapping device that allowed us to either hold the models in a stationary position or move them dynamically at their self-propelled swimming speed. Compared with a smooth control model without denticles, the 3D printed shark skin showed increased swimming speed with reduced energy consumption under certain motion programs. For example, at a heave frequency of 1.5 Hz and an amplitude of ±1 cm, swimming speed increased by 6.6% and the energy cost-of-transport was reduced by 5.9%. In addition, a leading-edge vortex with greater vorticity than the smooth control was generated by the 3D printed shark skin, which may explain the increased swimming speeds. The ability to fabricate synthetic biomimetic shark skin opens up a wide array of possible manipulations of surface roughness parameters, and the ability to examine the hydrodynamic consequences of diverse skin denticle shapes present in different shark species.

Concepts: Roughness, Shark, Surface metrology, Lamniformes, Lamnidae, Shortfin mako shark, Isurus, Longfin mako shark