Björn Mauck

Seal whiskers detect water movements

How do pinnipeds orientate themselves under water? As most pinniped species feed in conditions under which visibility is drastically reduced, for example at night, at great depths or in murky waters, it has been particularly unclear how they succeed in finding food. Here we show that harbour seals (Phoca vitulina) can use their whiskers to detect minute water movements. The high sensitivity of this sensory system should allow a seal to gain hydrodynamic information resulting from movements of other aquatic animals, such as prey, predators or conspecifics.

High olfactory sensitivity for dimethyl sulphide in harbour seals.

Productive areas are patchily distributed at sea and represent important feeding grounds for many marine organisms. Although pinnipeds are known to travel on direct routes and return regularly to particular feeding sites, the environmental information seals use to perform this navigation is as yet unknown. As atmospheric dimethyl sulphide (DMS) has been demonstrated to be a reliable indicator for profitable foraging areas, we tested seals for their ability to smell DMS at concentrations typical for the marine environment. Using a go/no-go response paradigm we determined the DMS detection threshold in two harbour seals (Phoca vitulina vitulina). DMS stimuli from 8.05×108 to 8 pmol (DMS) m−3(air) were tested against a control stimulus using a custom-made olfactometer. DMS-thresholds determined for both seals (20 and 13 pmol m−3) indicate that seals can detect ambient concentrations associated with high primary productivity, e.g. in the North Atlantic. Thus, seals possess an extraordinarily high olfactory sensitivity for DMS, which could provide a sensory basis for identifying or orienting to profitable foraging grounds.

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

SUMMARY 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.

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

SUMMARY 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.

Hydrodynamic discrimination of wakes caused by objects of different size or shape in a harbour seal (Phoca vitulina)

SUMMARY Harbour seals can use their mystacial vibrissae to detect and track hydrodynamic wakes. We investigated the ability of a harbour seal to discriminate objects of different size or shape by their hydrodynamic signature and used particle image velocimetry to identify the hydrodynamic parameters that a seal may be using to do so. Hydrodynamic trails were generated by different sized or shaped paddles that were moved in the calm water of an experimental box to produce a characteristic signal. In a two-alternative forced-choice procedure the blindfolded subject was able to discriminate size differences of down to 3.6 cm (Weber fraction 0.6) when paddles were moved at the same speed. Furthermore the subject distinguished hydrodynamic signals generated by flat, cylindrical, triangular or undulated paddles of the same width. Particle image velocimetry measurements demonstrated that the seal could have used the highest velocities and the steepness of the gradients within the wake to discriminate object size, beside the size of counter-rotating vortices and the spatial extension of a wake. For shape discrimination the subject could have used the spatial extension of the whole wake, in addition to the arrangement of the vortices. We tested whether the seal used highest velocities, the steepness of the gradients and the spatial extension of the wake in a second set of experiments by varying moving speed and paddle size, respectively. The subject was still able to discriminate between the respective object sizes, but the minimum detectable size difference increased to 4.4 cm (Weber fraction 3.6). For the shape discrimination task, the seal was only able to distinguish flat from triangular paddles. Our results indicate that the seals discrimination abilities depend on more than one hydrodynamic parameter.

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° 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.

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

SUMMARY 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.

Identity concept formation during visual multiplechoice matching in a harbor seal (Phoca vitulina)

Identity concept formation was tested in a harbor seal using a visual multiple-choice matching-tosample task. The seal was first trained on a two-alternative matching task. After criterion (≥80% correct choices in two successive sessions) was reached with two sets of two stimuli (Figure 3, Blocks A and B), stimulus sets were enlarged to six objects (Blocks C-G). After the seal reached criterion immediately with two successive sets (Blocks F and G), multiple-choice matching was introduced, first using stimulus sets of four familiar objects (Blocks H-M). After the seal reached the criterion immediately with two successive sets (Blocks L and M), completely new objects were used in two further stimulus sets (Blocks N and O). The seal immediately applied the matching rule in all four sessions (≥80% correct choices). In two further sessions with problems composed of all 38 familiar stimuli, the seal again reached the criterion (Block P). In the final, transfer session, 20 new problems were composed of 80 unknown stimuli (Block Q). The seal immediately applied the matching rule in these one-trial tests, showing that harbor seals can conceptualize complex visual information.

Visual Information Processing in the Lion-Tailed Macaque (Macaca silenus): Mental Rotation or Rotational Invariance?

Mental rotation is a widely accepted concept indicating an image-like mental representation of visual information and an analogue mode of information processing in certain visuospatial tasks. In the task of discriminating between image and mirror-image of rotated figures, human reaction times increase with the angular disparity between the figures. In animals, tests of this kind yield inconsistent results. Pigeons were found to use a time-independent rotational invariance, possibly indicating a non-analogue information processing system that evolved in response to the horizontal plane of reference birds perceive during flight. Despite similar ecological demands concerning the visual reference plane, a sea lion was found to use mental rotation in similar tasks, but its processing speed while rotating three-dimensional stimuli seemed to depend on the axis of rotation in a different way than found for humans in similar tasks. If ecological demands influence the way information processing systems evolve, hominids might have secondarily lost the ability of rotational invariance while retreating from arboreal living and evolving an upright gait in which the vertical reference plane is more important. We therefore conducted mental rotation experiments with an arboreal living primate species, the lion-tailed macaque. Performing a two-alternative matching-to-sample procedure, the animal had to decide between rotated figures representing image and mirror-image of a previously shown upright sample. Although non-rotated stimuli were recognized faster than rotated ones, the animal’s mean reaction times did not clearly increase with the angle of rotation. These results are inconsistent with the mental rotation concept but also cannot be explained assuming a mere rotational invariance. Our study thus seems to support the idea of information processing systems evolving gradually in response to specific ecological demands.

An analysis of visual oddity concept learning in a California sea lion (Zalophus californianus)

We tested a California sea lion for visual oddity learning by presenting problems composed of three two-dimensional black-and-white stimuli, two identical (S−) and one different (S+). In the first experimental stage, a single problem per session was presented until learning criterion was reached. In the second experimental stage, all problems were presented only five times in succession; then a new problem was introduced (six problems/session). In the third experimental stage, each problem was presented only once. The sea lion mastered all stages of oddity learning. A final transfer test with oddity problems composed of completely new stimuli yielded performance significantly above chance. Data analyses suggested learning of specific stimulus properties in the first stage, learning set formation in the second stage, but oddity conceptualization in the third stage.