Jochen Zeil

Catchment areas of panoramic snapshots in outdoor scenes

We took panoramic snapshots in outdoor scenes at regular intervals in two- or three-dimensional grids covering 1 m2 or 1 m3 and determined how the root mean square pixel differences between each of the images and a reference image acquired at one of the locations in the grid develop over distance from the reference position. We then asked whether the reference position can be pinpointed from a random starting position by moving the panoramic imaging device in such a way that the image differences relative to the reference image are minimized. We find that on time scales of minutes to hours, outdoor locations are accurately defined by a clear, sharp minimum in a smooth three-dimensional (3D) volume of image differences (the 3D difference function). 3D difference functions depend on the spatial-frequency content of natural scenes and on the spatial layout of objects therein. They become steeper in the vicinity of dominant objects. Their shape and smoothness, however, are affected by changes in illumination and shadows. The difference functions generated by rotation are similar in shape to those generated by translation, but their plateau values are higher. Rotational difference functions change little with distance from the reference location. Simple gradient descent methods are surprisingly successful in recovering a goal location, even if faced with transient changes in illumination. Our results show that view-based homing with panoramic images is in principle feasible in natural environments and does not require the identification of individual landmarks. We discuss the relevance of our findings to the study of robot and insect homing.

Eyes, eye stalks and the visual world of semi-terrestrial crabs

Summary1.We used in vivo optical techniques to study compound eye organisation and the extent of eye stalk development in 17 species of semi-terrestrial crabs (Brachyura).2.‘Narrow-fronted’ species (Ocypodidae, Mictyridae) which have their eyes close together on elongated, vertically oriented eye stalks have a narrow acute zone for vertical resolving power along the horizon (Figs. 2, 4, 6), while ‘broad-fronted’ species (Grapsidae, Xanthidae, Portunidae) which have their eyes far apart on short eye stalks lack this specialisation (Figs. 3, 4, 6).3.There is no pronounced acute zone for horizontal resolving power in either ‘type’ of crab (Fig. 5).4.A quantitative comparison of different species shows that acute zones for vertical resolving power are closely associated with long eye stalks (Fig. 6).5.In contrast to ‘broad-fronted’ species, ‘narrow-fronted’ species are known to inhabit relatively flat terrains. We show that elongated eye stalks and acute zones for vertical resolving power are specific adaptations to the problems of spatial vision in a flat environment and could enable animals to gain depth and size information monocularily from retinal position and retinal size alone (Figs. 8, 9).6.We discuss the preconditions of spatial vision in differently structured environments and how they might have shaped the evolution of different designs of visual systems in semi-terrestrial crabs.

Visual homing: an insect perspective

The ability to learn the location of places in the world and to revisit them repeatedly is crucial for all aspects of animal life on earth. It underpins animal foraging, predator avoidance, territoriality, mating, nest construction and parental care. Much theoretical and experimental progress has recently been made in identifying the sensory cues and the computational mechanisms that allow insects (and robots) to find their way back to places, while the neurobiological mechanisms underlying navigational abilities are beginning to be unravelled in vertebrate and invertebrate models. Studying visual homing in insects is interesting, because they allow experimentation and view-reconstruction under natural conditions, because they are likely to have evolved parsimonious, yet robust solutions to the homing problem and because they force us to consider the viewpoint of navigating animals, including their sensory and computational capacities.

Butterfly wing colours: scale beads make white pierid wings brighter

The wing–scale morphologies of the pierid butterflies Pieris rapae (small white) and Delias nigrina (common jezabel), and the heliconine Heliconius melpomene are compared and related to the wing–reflectance spectra. Light scattering at the wing scales determines the wing reflectance, but when the scales contain an absorbing pigment, reflectance is suppressed in the absorption wavelength range of the pigment. The reflectance of the white wing areas of P. rapae, where the scales are studded with beads, is considerably higher than that of the white wing areas of H. melpomene, which has scales lacking beads. The beads presumably cause the distinct matt–white colour of the wings of pierids and function to increase the reflectance amplitude. This will improve the visual discrimination between conspecific males and females.

The information content of panoramic images I: The rotational errors and the similarity of views in rectangular experimental arenas

Animals relocating a target corner in a rectangular space often make rotational errors searching not only at the target corner but also at the diagonally opposite corner. The authors tested whether view-based navigation can explain rotational errors by recording panoramic snapshots at regularly spaced locations in a rectangular box. The authors calculated the global image difference between the image at each location and the image recorded at a target location in 1 of the corners, thus creating a 2-dimensional map of image differences. The authors found the most pronounced minima of image differences at the target corner and the diagonally opposite corner--conditions favoring rotational errors. The authors confirmed these results in virtual reality simulations and showed that the relative salience of different visual cues determines whether image differences are dominated by geometry or by features. The geometry of space is thus implicitly contained in panoramic images and does not require explicit computation by a dedicated module. A testable prediction is that animals making rotational errors in rectangular spaces are guided by remembered views.

Visually mediated species and neighbour recognition in fiddler crabs (Uca mjoebergi and Uca capricornis)

Mating signals are often directed at numerous senses and provide information about species identity, gender, receptiveness, individual identity and mate quality. Given the diversity of colourful body patterns in invertebrates, surprisingly few studies have examined the role of these visual signals in mate recognition. Here, we demonstrate the use of claw coloration as a species recognition signal in a fiddler crab (Uca mjoebergi). Furthermore, we show that distinct carapace colour patterns in Uca capricornis enable males to discriminate between their female neighbours and unfamiliar females. This is the first empirical evidence of the social importance of colour markings in fiddler crabs and the first example of visually mediated species and neighbour recognition in invertebrates other than insects.

The Information Content of Panoramic Images II: View-Based Navigation in Nonrectangular Experimental Arenas

Two recent studies testing navigation of rats in swimming pools have posed problems for any account of the use of purely geometric properties of space in navigation (M. Graham, M. A. Good, A. McGregor, & J. M. Pearce, 2006; J. M. Pearce, M. A. Good, P. M. Jones, & A. McGregor, 2004). The authors simulated 1 experiment from each study in a virtual reality environment to test whether experimental results could be explained by view-based navigation. The authors recorded a reference image at the target location and then determined global panoramic image differences between this image and images taken at regularly spaced locations throughout the arena. A formal model, in which an agent attempts to minimize image differences between the reference image and current views, generated trajectories that could be compared with the search performance of rats. For both experiments, this model mimics many aspects of rat behavior. View-based navigation provides a sufficient and parsimonious explanation for a range of navigational behaviors of rats under these experimental conditions.

Fundamental differences in the optical structure of the eyes of nocturnal and diurnal mosquitoes

Abstract We have studied the anatomy and optics of the eyes of a range of mosquito species from the wholly dark-active blood-feeding Anopheles gambiae to the diurnal plant-feeder Toxorhynchites brevipalpis. Consistent with studies by Satô in the 1950s, we find that dark-active and crepuscular species have short fused rhabdoms with a conical construction. This maximises the amount of light the rhabdoms receive from the almost hemispherical wide-aperture lenses. Toxorhynchites, on the other hand, has long narrow rhabdomeres that are separated from each other over their entire length, and so resemble the open rhabdoms of advanced flies (Brachycera and Cyclorrhapha). These findings are confirmed by studies of the pseudopupil, whose form indicates the layout of the rhabdomere tips in the focal plane of each ommatidial lens. In anopheline species the pseudopupil is a single undivided ellipse, indicating a fused rhabdom structure, whereas in Toxorhynchites there is a ring of six outer elements surrounding a central one. This means that each rhabdomere views a separate direction in space, and our measurements indicate that, as in higher Diptera, adjacent rhabdomeres share their fields of view with one of the rhabdomeres in the immediately adjacent ommatidia. This in turn means that in the diurnal type of mosquito eye there is a basis for neural superposition, but the fused construction of anopheline rhabdoms precludes this. The Aedes species studied were similar to Anopheles but with lenses of less extreme aperture, and Sabethes cyaneus, a diurnal blood-feeder, was intermediate in structure, with fused conical rhabdoms in the centre of the eye and unfused rhabdomeres around the periphery.

Homing in fiddler crabs (Uca lactea annulipes and Uca vomeris : Ocypodidae)

Abstract Fiddler crabs emerge from burrows on intertidal sand- and mudflats to feed during low tide. In the species studied here (Uca lactea annulipes, Uca vomeris) a crab normally wanders no more than about 1 m away from its burrow and, when frightened, dashes back along a straight line to take cover. Feeding crabs tend to move sideways, without changing orientation, along paths radiating from the burrow. When they move along circumferential paths they adjust their orientation so that one side continues to point towards the burrow. The crabs do not need to see the burrow in order to stay aligned with the home vector, and they are not misled by a dummy hole close to their own burrow unless they have come to within about 10 cm of it. The home runs of crabs end within a few centimeters of a burrow that is covered with a sheet of sandpaper and then give way to search runs, centred upon a position slightly short of the burrow location. Feeding crabs can be displaced on sandpapers and their subsequent home runs end at a position where the burrow would be, had there been no displacement. Landmarks close to the burrow do not influence the home runs of displaced crabs. Crabs that are rotated on a sheet of sandpaper, counter-turn to keep their original orientation constant. Fiddler crabs thus employ path integration with external compass information and close range visual guidance for homing.

Considerations on the use of video playbacks as visual stimuli: The Lisbon workshop consensus

Abstract This paper is the consensus of a workshop that critically evaluated the utility and problems of video playbacks as stimuli in studies of visual behavior. We suggest that video playback is probably suitable for studying motion, shape, texture, size, and brightness. Studying color is problematic because video systems are specifically designed for humans. Any difference in color perception must lead to a different color sensation in most animals. Another potentially problematic limitation of video images is that they lack depth cues derived from stereopsis, accommodation, and motion parallax. Nonetheless, when used appropriately, video playback allows an unprecedented range of questions in visual communication to be addressed. It is important to note that most of the potential limitations of video playback are not unique to this technique but are relevant to all studies of visual signaling in animals.