Brightness discrimination in the South African fur seal (Arctocephalus pusillus).

Underwater, the contrast between object and background is much larger reduced with increasing distance between object and observer than in air. For marine predators, such as pinnipeds, it would therefore be advantageous to possess a high sensitivity for brightness differences, since this would increase the distance at which prey can be detected visually. Few studies have examined the brightness discrimination thresholds of pinnipeds. Two studies with phocid seals have confirmed low brightness discrimination thresholds in pinnipeds whereas the threshold obtained for the South African fur seal seems to be twice as high as that of the phocids. However, the experiments with the South African fur seal have been conducted under inadequate conditions which likely resulted in an underestimation of the brightness discrimination ability of this species. The study at hand reinvestigated the brightness discrimination threshold of the South African fur seal under well controlled conditions. In a two alternative forced choice task, one fur seal was trained to indicate the position of the brighter of two gray discs presented on a black background on a monitor. The thresholds were determined for 11 standard intensities each tested against 8 lower comparison intensities. It was found that the fur seal was able to perceive brightness differences of 8-10%, which is better than the phocid species tested so far. For low standard intensities, however, the threshold increased which could to be due to a relative slow dark adaptation rate of the fur seal. The results are discussed in terms of the relevance of visual information for pinnipeds during foraging dives and are directly compared to the results obtained for the harbor seal which has been tested under the same conditions as the fur seal in a previous study.

Flow sensing by pinniped whiskers.

Beside their haptic function, vibrissae of harbour seals (Phocidae) and California sea lions (Otariidae) both represent highly sensitive hydrodynamic receptor systems, although their vibrissal hair shafts differ considerably in structure. To quantify the sensory performance of both hair types, isolated single whiskers were used to measure vortex shedding frequencies produced in the wake of a cylinder immersed in a rotational flow tank. These measurements revealed that both whisker types were able to detect the vortex shedding frequency but differed considerably with respect to the signal-to-noise ratio (SNR). While the signal detected by sea lion whiskers was substantially corrupted by noise, harbour seal whiskers showed a higher SNR with largely reduced noise. However, further analysis revealed that in sea lion whiskers, each noise signal contained a dominant frequency suggested to function as a characteristic carrier signal. While in harbour seal whiskers the unique surface structure explains its high sensitivity, this more or less steady fundamental frequency might represent the mechanism underlying hydrodynamic reception in the fast swimming sea lion by being modulated in response to hydrodynamic stimuli impinging on the hair.

The impact of glide phases on the trackability of hydrodynamic trails in harbour seals (Phoca vitulina).

The mystacial vibrissae of harbour seals (Phoca vitulina) constitute a highly sensitive hydrodynamic receptor system enabling the seals to detect and follow hydrodynamic trails. In the wild, hydrodynamic trails, as generated by swimming fish, consist of cyclic burst-and-glide phases, associated with various differences in the physical parameters of the trail. Here, we investigated the impact of glide phases on the trackability of differently aged hydrodynamic trails in a harbour seal. As fish are not easily trained to swim certain paths with predetermined burst-and-glide phases, the respective hydrodynamic trails were generated using a remote-controlled miniature submarine. Gliding phases in hydrodynamic trails had a negative impact on the trackability when trails were 15 s old. The seal lost the generated trails more often within the transition zones, when the submarine switched from a burst to a glide moving pattern. Hydrodynamic parameter analysis (particle image velocimetry) revealed that the smaller dimensions and faster decay of hydrodynamic trails generated by the gliding submarine are responsible for the impaired success of the seal tracking the gliding phase. Furthermore, the change of gross water flow generated by the submarine from a rearwards-directed stream in the burst phase to a water flow passively dragged behind the submarine during gliding might influence the ability of the seal to follow the trail as this might cause a weaker deflection of the vibrissae. The possible ecological implications of intermittent swimming behaviour in fish for piscivorous predators are discussed.

Hydrodynamic determination of the moving direction of an artificial fin by a harbour seal (Phoca vitulina).

Harbour seals can use their vibrissal system to detect and follow hydrodynamic trails left by moving objects. In this study we determined the maximum time after which a harbour seal could indicate the moving direction of an artificial fish tail and analysed the hydrodynamic parameters allowing the discrimination. Hydrodynamic trails were generated using a fin-like paddle moving from left to right or from right to left in the calm water of an experimental box. The blindfolded seal was able to recognise the direction of the paddle movement when the hydrodynamic trail was up to 35 s old. Particle Image Velocimetry (PIV) revealed that the seal might have perceived and used two different hydrodynamic parameters to determine the moving direction of the fin-like paddle. The structure and spatial arrangement of the vortices in the hydrodynamic trail and high water velocities between two counter-rotating vortices are characteristic of the movement direction and are within the sensory range of the seal.

Tracking of biogenic hydrodynamic trails in harbour seals (Phoca vitulina).

For seals hunting in dark and murky waters one source of sensory information for locating prey consists of fish-generated water movements, which they can detect using their highly sensitive mystacial vibrissae. As water movements in the wake of fishes can persist for several minutes, hydrodynamic trails of considerable length are generated. It has been demonstrated that seals can use their vibrissae to detect and track hydrodynamic trails generated artificially by miniature submarines. In the present study, we trained a harbour seal to swim predefined courses, thus generating biogenic hydrodynamic trails. The structure of these trails was measured using Particle Image Velocimetry. A second seal was trained to search for and track the trail after the trail-generating seal had left the water. Our trail-following seal was able to detect and accurately track the hydrodynamic trail, showing search patterns either mostly congruent with the trail or crossing the trail repeatedly in an undulatory way. The undulatory trail-following search pattern might allow a seal to relocate a lost trail or successfully track a fleeing, zigzagging prey fish.

Mental rotation of perspective stimuli in a California sea lion (Zalophus californianus).

The time it takes humans to discriminate rotated objects from their mirror images increases linearly with the rotation angle. This phenomenon is probably due to an analogue mode of visual information processing during which an object's mental representation is rotated in a time-consuming process called mental rotation. As the speed of mental rotation in humans depends on rotation axis, we tested the ability of a California sea lion to mentally rotate perspective line drawings of three-dimensional objects about four axes. In a matching-to-sample experiment the animal was presented with the image and a mirror image of a block sample that had previously been shown upright. Both image and mirror image were rotated by a multiple of 60 degrees about the object's x-, y-, z-axis, or a skew axis (an axis oblique to these standard orthogonal axes). The animal's choice and reaction times were recorded using a computer-controlled touch-screen device. Mean reaction times and errors generally increased with angular disparity supporting the model of mental rotation for three-dimensional objects. Linear regression analysis of mean reaction times yielded high correlation coefficients only for three axes. The slope of reaction time functions indicated the highest mental rotation speed for the skew axis. This contrasts with the priority of mental rotation axes in humans suggesting that due to special ecological demands a different mode of orientation invariance evolved in marine mammals.

Hydrodynamic trail-following in harbor seals (Phoca vitulina).

Marine mammals often forage in dark or turbid waters. Whereas dolphins use echolocation under such conditions, pinnipeds apparently lack this sensory ability. For seals hunting in the dark, one source of sensory information may consist of fish-generated water movements, which seals can detect with their highly sensitive whiskers. Water movements in the wake of fishes persist for several minutes. Here we show that blindfolded seals can use their whiskers to detect and accurately follow hydrodynamic trails generated by a miniature submarine. This shows that hydrodynamic information can be used for long-distance prey location.

Haptic discrimination of size and texture in squirrel monkeys (Saimiri sciureus).

The present study analyzed haptic abilities of four squirrel monkeys. Using a two-alternative forced-choice procedure, stimuli were presented in a visually opaque box, allowing unrestrained test subjects to grab through a small opening and touch the discriminanda. Difference thresholds were determined by a modified method of limits. In the first experiment we determined size difference thresholds for the discrimination of circular cylinders using standard stimuli differing in diameter from 10 mm to 35 mm. In the second experiment a texture difference threshold was obtained for the discrimination of grooved surfaces (groove width 2-7 mm). The squirrel monkeys achieved a mean size difference threshold of 8% stimulus difference. The linear increase of absolute thresholds as a function of the starting stimulus size showed that haptic size discriminations in squirrel monkeys correspond to Weber's law. Three of the animals achieved a texture difference of 10% stimulus difference, while one monkey showed a distinctively lower haptic acuity. An analysis of the exploratory behavior points to a subject-related difference in the significance of cutaneous and kinesthetic information during size discriminations. Whereas differences in the animals' exploratory behavior did not correlate with the size difference threshold a subject achieved, different thresholds for texture discrimination can be explained by the different exploratory procedures the monkeys used to touch grooved surfaces. The low difference thresholds determined for the squirrel monkeys in the present study point to the significance of unrestrained test conditions for the assessment of the haptic capacity of a species.

Selective heating of vibrissal follicles in seals (Phoca vitulina) and dolphins (Sotalia fluviatilis guianensis).

The thermal characteristics of the mystacial vibrissae of harbour seals (Phoca vitulina) and of the follicle crypts on the rostrum of the dolphin Sotalia fluviatilis guianensis were measured using an infrared imaging system. Thermograms demonstrate that, in both species, single vibrissal follicles are clearly defined units of high thermal radiation, indicating a separate blood supply to these cutaneous structures. It is suggested that the high surface temperatures measured in the area of the mouth of the follicles is a function of the sinus system. In seals and dolphins, surface temperature gradually decreased with increasing distance from the centre of a follicle, indicating heat conduction from the sinus system via the follicle capsule to adjacent tissues. It is suggested that the follicular sinus system is a thermoregulatory structure responsible for the maintenance of high tactile sensitivity at the extremely low ambient temperatures demonstrated for the vibrissal system of seals. The vibrissal follicles of odontocetes have been described as vestigial structures, but the thermograms obtained in the present study provide the first evidence that, in Sotalia fluviatilis, the follicles possess a well-developed sinus system, suggesting that they are part of a functional mechanosensory system.

Salinity discrimination in harbour seals: a sensory basis for spatial orientation in the marine environment?

Salinity variations can be considered as a potential source of information for orientation in the marine environment. To use this kind of environmental information marine animals must be able to detect these salinity differences. Therefore we determined salinity-difference thresholds of two harbour seals for the discrimination of seawater solutions as a function of the salinity level (15-35@1000) and compared them with the thresholds of human subjects. Whereas in humans thresholds increased with increasing salinity level, thresholds of seals decreased with increasing salinity level. Both seals achieved best sensitivity at 30/1000 salinity, where they detected a salinity difference < or = 4%. These data indicate that the ability of seals to detect salinity differences of seawater is well tuned to the natural occurrence of this environmental information. Their high gustatory resolving power for differences in seawater salinity is suggested to meet the basic requirements for chemosensory orientation of seals in the marine habitat.

Structure and innervation of the vibrissal follicle-sinus complex in the Australian water rat, Hydromys chrysogaster.

Light and electron microscopic techniques were used to examine the structure and innervation of the mystacial vibrissal follicle-sinus complex (F-SC) in the Australian water rat. The F-SCs of this semiaquatic rodent show the same morphologic elements described in terrestrial rats but differ in size, structure, and innervation. Most striking is the size of the water rat's caudal F-SCs, measuring 6.3 mm in length and 2.4 mm in diameter. The sinus system is divisible into a ring sinus and a cavernous sinus and shows a distinct asymmetry. At the highest level of the cavernous sinus, the outer root sheath forms a ridge in the direction of the trabeculae, which bind the ridge to the capsule. A ringwulst is present only in small and medium-sized F-SCs. The mean number of myelinated axons counted in the deep vibrissal nerve (DVN) of most caudal F-SCs was 537, indicating an innervation density of the water rat's vibrissal system at least 2.5 times as high as that of terrestrial rats. The total number of nerve fibers of the small superficial nerves was less than 10% of that of the DVN. These fibers innervate almost exclusively the area of the inner conical body. Structural specializations of the water rat F-SC are discussed as an analogous development in mammals adapted to the aquatic environment, primarily in terms of thermoregulation, whereas its high degree of innervation is assessed to lend support to the hypothesis that the vibrissal system is of special significance in aquatic mammals.

Ambient temperature does not affect the tactile sensitivity of mystacial vibrissae in harbour seals.

Vibrissae provide pinnipeds with tactile information primarily in the aquatic environment, which is characterized by its high thermal conductivity and large potential cooling power. Since studies of thermal effects on human tactile sensitivity have revealed that cooling below normal skin temperature impairs sensitivity, the present study investigates the tactile sensitivity of the vibrissal system of harbour seals at varying ambient temperatures. Using plates bearing gratings of alternating grooves and ridges, the texture difference thresholds of two adult seals were determined under water. We took advantage of the natural difference in ambient temperature between summer and winter. Mean water temperature was 1. 2 degreesC during the winter and 22 degreesC during the summer. During the cold season, the thermal status of both seals was examined using an infrared-sensitive camera system. The texture difference threshold of both seals remained the same (0.18 mm groove width difference) under both test conditions. The thermographic examination revealed that the skin areas of the head where the mystacial and supraorbital vibrissae are located show a substantially higher degree of thermal emission than do adjacent skin areas. This suggests that, in the vibrissal follicles of harbour seals, no vasoconstriction occurs during cold acclimation, so that the appropriate operating temperature for the mechanoreceptors is maintained.

Underwater audiogram of a tucuxi (Sotalia fluviatilis guianensis).

Using a go/no go response paradigm, a tucuxi (Sotalia fluviatilis guianensis) was trained to respond to pure-tone signals for an underwater hearing test. Auditory thresholds were obtained from 4 to 135 kHz. The audiogram curve shows that this Sotalia had an upper limit of hearing at 135 kHz; from 125 to 135 kHz sensitivity decreased by 475 dB/oct. This coincides with results from electrophysiological threshold measurements. The range of best hearing (defined as 10 dB from maximum sensitivity) was between 64 and 105 kHz. This range appears to be narrower and more restricted to higher frequencies in Sotalia fluviatilis guianensis than in other odontocete species that had been tested before. Peak frequencies of echolocation pulses reported from free-ranging Sotalia correspond with the range of most sensitive hearing of this test subject.

Mental rotation in a California sea lion (Zalophus californianus).

Mental rotation is a widely accepted concept that suggests an analogue mode of visual information-processing in certain visuospatial tasks. Typically, these tasks demand the discrimination between the image and mirror-image of rotated figures, for which human subjects need an increasing reaction time depending on the angular disparity between the rotated figures. In pigeons, tests of this kind yielded a time-independent rotational invariance, suggested as being the result of a non-analogue information-processing that has evolved in response to the horizontal plane that birds perceive from above while flying. Given that marine mammals use the water surface as the horizontal plane for orientation while diving, the ability of a California sea lion to mentally rotate two-dimensional shapes was tested. Using a successive two-alternative matching-to-sample procedure, the animal had to decide between the image and mirror-image of a previously shown sample. Both stimuli were rotated by a multiple of 30 degrees with respect to the sample. The animal's reaction time was measured by a computer-controlled touch-screen device, rewarding the animal for pressing its snout against the stimulus matching the sample. A linear regression analysis of the animal's mean reaction time against the angular rotation of the stimulus yielded a significant correlation coefficient. Thus, the present data can be explained by the mental rotation model, predicting an image-like representation of visual stimuli in this species. The present results therefore correspond well with those found for human subjects, but are inconsistent with the data reported for pigeons.

Sensitivity of the mystacial vibrissae of harbour seals (Phoca vitulina) for size differences of actively touched objects.

We studied the capability of one male and one female harbour seal Phoca vitulina to discriminate diameter differences of circular discs by means of active touch with their mystacial vibrissae. To prevent the animals from perceiving visual information, they were blindfolded during trials. In a two-alternative forced-choice procedure, the seals were required to choose the larger of two Perspex discs. Weber fractions c (the ratio of the lowest diameter difference detected by the seals on 75% of occasions to the starting disc diameter D, delta D/D = c) were determined for three standard discs (diameters 1.12cm, 5.04cm and 8.74cm) by the psychophysical method of limits. While the seals achieved Weber fractions of 0.29 (male) and 0.26 (female) at the smallest standard disc, their performance improved with increasing disc size, resulting in an approximately constant Weber fraction of 0.13 (male) and 0.08 (female) for the two larger standard discs. The difference in performance between the two seals did not reflect a real difference in sensitivity, but may best be explained by a difference in choice behaviour. As a measure of tactile acuity, the Weber fractions obtained for the larger standard discs indicate that harbour seals can use their mystacial vibrissae as efficiently for active touch as monkeys use their hands.

Tactile size discrimination by a California sea lion (Zalophus californianus) using its mystacial vibrissae.

The capability of a blindfolded California sea lion (Zalophus californianus) to discriminate diameter differences of circular discs by means of active touch with its mystacial vibrissae was studied. Using a forced choice paradigm the sea lion was required to choose the larger of two simultaneously presented perspex discs. Absolute difference thresholds (delta D) were determined for 3 standard discs (1.12 cm phi, 2.52 cm phi, 8.74 cm phi) by the psychophysical method of constants. Increasing disc size resulted in an increase in the absolute difference threshold from 0.33 cm for the smallest disc size to 1.55 cm for the largest disc size. The relative difference threshold (Weber fraction) remained approximately constant at a mean value of 0.26. According to a video analysis the sea lion did not move its vibrissae when touching the discs. Instead, it performed precisely controlled lateral head movements, with the touched disc located centrally between the vibrissae of both sides of the muzzle. Since the extent of these head movements was identical at discs to be compared, discs of different size must have led to different degrees of deflection of vibrissae involved in the tactile process, resulting in quantitatively different mechanical stimulations of mechanoreceptors in the follicles. This suggests that the accuracy of the sea lion's size discrimination was determined by the efficiency of two sensory systems: the mechanosensitivity of follicle receptors as well as kinaesthesis.