Fractionating dead reckoning: role of the compass,odometer, logbook,and home base establishment in spatial orientation

Rats use multiple sources of information to maintain spatial orientation. Although previous work has focused on rats’ use of environmental cues, a growing number of studies have demonstrated that rats also use self-movement cues to organize navigation. This review examines the extent that kinematic analysis of naturally occurring behavior has provided insight into processes that mediate dead-reckoning-based navigation. This work supports a role for separate systems in processing self-movement cues that converge on the hippocampus. The compass system is involved in deriving directional information from self-movement cues; whereas, the odometer system is involved in deriving distance information from self-movement cues. The hippocampus functions similar to a logbook in that outward path unique information from the compass and odometer is used to derive the direction and distance of a path to the point at which movement was initiated. Finally, home base establishment may function to reset this system after each excursion and anchor environmental cues to self-movement cues. The combination of natural behaviors and kinematic analysis has proven to be a robust paradigm to investigate the neural basis of spatial orientation.     

Motivation and vector navigation in honey bees

Animals that forage from a central place can keep track of their displacement relative to home through a process called "path integration." During a study of the stability of homing information over time, we noticed that honey bees held at a feeding place for several hours sometimes headed not in the homeward compass direction on their release, but in the reverse compass direction. This behavior suggested that the path integration system had been reset to a state corresponding to an outward flight to the food. Most models of insect navigation assume that it is the experience of reaching home that resets the path integration system, enabling the activation of vectors appropriate for subsequent outbound foraging trips. Here we provide evidence that this resetting can be influenced by motivational cues associated with food deprivation. The effect of food deprivation is independent of any positional cues provided by familiar landmarks or by experience in traveling toward a goal.     

Vision-independent odometry in the ant Cataglyphis cursor

In contrast to flying insects, in which distance estimation is visually mediated, self-induced image motion and use of familiar landmarks are known to play a minor role in ants. Here we show that strictly diurnal Cataglyphis cursor ants can gauge with accuracy the distance they have travelled even in complete darkness in the absence of any other cues, i.e. chemical or protocounting information. Thus, an ants odometer is a vision-independent system based on proprioceptive cues, implicating some form of step counting, which remain to be elucidated.     

Wind and sky as compass cues in desert ant navigation

While integrating their foraging and homing paths, desert ants, Cataglyphis fortis, depend on external compass cues. Whereas recent research in bees and ants has focused nearly exclusively on the polarization compass, two other compass systems—the sun compass and the wind (anemo) compass—as well as the mutual interactions of all these compass systems have received little attention. In this study, we show that of the two visual compass systems, it is only the polarization compass that invariably outcompetes the wind compass, while the sun compass does so only under certain conditions. If the ants are experimentally deprived of their polarization compass system, but have access simultaneously to both their sun compass and their wind compass, they steer intermediate courses. The intermediate courses shift the more towards the wind compass course, the higher the elevation of the sun is in the sky.     

Orientation and navigation during adult transport between nests in the ant Cataglypis iberica

  Cataglyphis iberica is a polydomous ant species in which adult transports between nests are frequently observed. When pairs of workers were captured and released at the same location, the transporters (Ts) fled directly towards their destination nest and reached it in most of the cases. The transportees (Te), on the other hand, fled in the opposite direction and only a third of them eventually reached their nest of departure. Additional experiments suggest that this result may be explained by the fact that the Ts ants have a memory of the compass direction of the nest they are heading to and that they adjust their course by using a sequence of memorised landmarks. As regards to the Te, the reversal of their direction of transport seems to be based essentially on celestial cues. Received: 20 October 1999 / Accepted in revised form: 10 May 2000     

Loft features reveal the functioning of the young pigeon’s navigational system

It is thought that young homing pigeons are able to use information acquired en route for their initial homeward orientation. However, the cues involved and mechanisms utilised are under discussion. Blocking light-dependent route-specific information during the first leg of an outward journey detour, together with analysis of pigeons that were raised under different loft conditions, allowed us to correctly evaluate the functioning of this mechanism and, more generally, the navigational map of birds. Pigeons from the same stock were raised and kept in two different lofts. The birds in the experimental groups were transported to the release sites via detours, and light-dependent information was denied during the first half of the outward journey (no compass information was available). Control birds were transported by the most direct route and had access to all available information. In general, the results showed that the low-loft birds preferred to use magnetic compass cues, whereas the high-loft birds preferred to use navigational map cues to collect information of the first part of the outward journey. The impairments observed in the homing performances of the experimental groups highlight the reliability of information collected inside the map area. Relevant to an understanding of the route-reversal mechanism was the evidence that this mechanism is able to function in the absence of compass information (birds raised in a wind-exposed loft show a detour effect). In systems where directional information could be provided by multiple sources, processing and extracting accurate course trajectories through a common mechanism may prove more efficient and reliable.     

Canopy compass in nocturnal homing of the subsocial shield bug, <Emphasis Type="Italic">Parastrachia japonensis</Emphasis> (Heteroptera: Parastrachiidae)

In contrast to an open environment where a specific celestial cue is predominantly used, visual contrast of canopies against the sky through the gap, known as canopy cues, is known to play a major role for visually guided insect navigators in woodland habitats. In this paper, we investigated whether a subsocial shield bug, Parastrachia japonensis, could gauge direction using canopy cues on a moonless night. The results show that they could perform the round trip foraging behaviour even in an experimental arena with only an artificial round gap opened in the ceiling of the arena and adjust their homing direction for a new azimuth when the gap was rotated. Thus, P. japonensis can use slightly brighter canopy cues as a compass reference but not complex landmarks during nocturnal homing behaviour.     

Spontaneous expression of magnetic compass orientation in an epigeic rodent: the bank vole,Clethrionomys glareolus

Magnetoreception has been convincingly demonstrated in only a few mammalian species. Among rodents, magnetic compass orientation has been documented in four species of subterranean mole rats and two epigeic (i.e. active above ground) species—the Siberian hamster and the C57BL/6J mouse. The mole rats use the magnetic field azimuth to determine compass heading; their directional preference is spontaneous and unimodal, and their magnetic compass is magnetite-mediated. By contrast, the primary component of orientation response is learned in the hamster and the mouse, but both species also exhibit a weak spontaneous bimodal preference in the natural magnetic field. To determine whether the magnetic compass of wild epigeic rodents features the same functional properties as that of laboratory rodents, we investigated magnetic compass orientation in the bank vole Clethrionomys glareolus (Cricetidae, Rodentia). The voles exhibited a robust spontaneous bimodal directional preference, i.e. built nests and slept preferentially along the north-south axis, and deflected their directional preference according to a shift in the direction of magnetic north, clearly indicating that they were deriving directional information from the magnetic field. Thus, bimodal, axially symmetrical directional choice seems to be a common feature shared by epigeic rodents. However, spontaneous directional preference in the bank vole appeared to be more pronounced than that reported in the hamster and the mouse. These findings suggest that bank voles are well suited for future studies investigating the adaptive significance and mechanisms of magnetic orientation in epigeic rodents.     

Landmark memories are more robust when acquired at the nest site than en route: experiments in desert ants

Foraging desert ants, Cataglyphis fortis, encounter different sequences of visual landmarks while navigating by path integration. This paper explores the question whether the storage of landmark information depends on the context in which the landmarks are learned during an ant's foraging journey. Two experimental set-ups were designed in which the ants experienced an artificial landmark panorama that was placed either around the nest entrance (nest marks) or along the vector route leading straight towards the feeder (route marks). The two training paradigms resulted in pronounced differences in the storage characteristics of the acquired landmark information: memory traces of nest marks were much more robust against extinction and/or suppression than those of route marks. In functional terms, this result is in accord with the observation that desert ants encounter new route marks during every foraging run but always pass the same landmarks when approaching the nest entrance.     

A brief view of known landmarks reorientates path integration in hamsters

 In darkness, hamsters commute between their nest and a feeding site through path integration only, and therefore show cumulative errors in the return direction to the nest. We examined whether a brief presentation of familiar room cues could reset the path integrator. The hamsters could see the room cues either during, or at the end of, the outward journey to the food place, in a conflict situation where motion cues and visual information were set at variance. In both conditions, the animals used mainly visual information to return home. Thus, hamsters can determine their azimuth, and possibly their location, through a visual fix, and can reset their path integrator through the fix. This allows them to update their position during further locomotion in the dark and thus to compute a correct homing vector with respect to a visually induced reference frame. Taking episodic positional fixes may greatly enhance the functional value of path integration. Received: 17 May 2000 / Accepted in revised form: 11 September 2000     

Social prophylaxis through distant corpse removal in ants

Living in groups raises important issues concerning waste management and related sanitary risks. Social insects such as ants live at high densities with genetically related individuals within confined and humid nests, all these factors being highly favorable for the spread of pathogens. Therefore, in addition to individual immunity, a social prophylaxis takes place, namely, by the removal of risky items such as corpses and their rejection at a distance from the ant nest. In this study, we investigate how Myrmica rubra workers manage to reduce encounters between potentially hazardous corpses and nestmates. Using both field and laboratory experiments, we describe how the spatial distribution and the removal distance of waste items vary as a function of their associated sanitary risks (inert item vs. corpse). In the field, corpse-carrying ants walked in a rather linear way away from the nest entrance and had an equal probability of choosing any direction. Therefore, they did not aggregate corpses in dedicated areas but scattered them in the environment. In both field and laboratory experiments, ants carrying corpses dropped their load in more remote—and less frequented—areas than workers carrying inert items. However, for equidistant areas, ants did not avoid dropping corpses at a location where they perceived area marking as a cue of high occupancy level by nestmates. Our results suggest that ants use distance to the nest rather than other occupancy cues to limit sanitary risks associated with dead nestmates.     

Nestmate recognition in ants is possible without tactile interaction

Ants of the genus Camponotus are able to discriminate recognition cues of colony members (nestmates) from recognition cues of workers of a different colony (non-nestmates) from a distance of 1 cm. Free moving, individual Camponotus floridanus workers encountered differently treated dummies on a T-bar and their behavior was recorded. Aggressive behavior was scored as mandibular threat towards dummies. Dummies were treated with hexane extracts of postpharyngeal glands (PPGs) from nestmates or non-nestmates which contain long-chain hydrocarbons in ratios comparable to what is found on the cuticle. The cuticular hydrocarbon profile bears cues which are essential for nestmate recognition. Although workers were prevented from antennating the dummies, they showed significantly less aggressive behavior towards dummies treated with nestmate PPG extracts than towards dummies treated with non-nestmate PPG extracts. In an additional experiment, we show that cis-9-tricosene, an alkene naturally not found in C. floridanus' cuticular profile, is behaviorally active and can interfere with nestmate recognition when presented together with a nestmate PPG extract. Our study demonstrates for the first time that the complex multi-component recognition cues can be perceived and discriminated by ants at close range. We conclude that contact chemosensilla are not crucial for nestmate recognition since tactile interaction is not necessary.     

Homing in the wolf spider Lycosa tarantula (Araneae, Lycosidae): the role of active locomotion and visual landmarks

Previous studies on the homing of the wolf spider Lycosa tarantula have shown that it is carried out by path integration. Animals using this mechanism must measure the distance walked and the angles turned. This study aims to understand if wolf spider L. tarantula is able to estimate the walked distance in an outward path. As this information is more likely obtained by proprioceptive mechanisms, active or passive displacements have been performed. An active locomotion was found essential to estimate distances. During passive locomotion, spiders searched for their burrows near the release point while when displaced actively the inbound journey was longer than the outbound one. The possible use of visual landmarks near the burrow was also tested as a cue to complete the inbound journey. Our results did not show that L. tarantula used these visual landmarks to find the burrow. L. tarantula seems to use only proprioceptive information obtained during the outbound path to estimate the distance traveled.     

Polarization-Sensitive Interneurons in the Optic Lobe of the Desert Ant Cataglyphis bicolor

 Desert ants, Cataglyphis bicolor (Hymenoptera), navigate by using compass information provided by skylight polarization. In this study, electrophysiological recordings were made from polarization-sensitive interneurons (POL-neurons) in the optic lobe of Cataglyphis. The POL-neurons exhibit a characteristic polarization opponency. They receive monochromatic input from the UV receptors of the specialized dorsal rim area of the compound eye. Both polarization opponency and monochromacy are features also found in the POL-neurons of crickets (Orthoptera). Received: 29 September 1999 / Accepted in revised form: 13 December 1999     

The sea-finding behavior of hatchling olive ridley sea turtles, Lepidochelys olivacea, at the beach of San Miguel (Costa Rica)

Newly hatched olive ridley sea turtles (Lepidochelys olivacea) were tested for their directional preferences in a sand-filled circular arena in total darkness. Hatchlings that had crawled about 5 m on the beach, toward the sea preferred the southwesterly direction that would have brought them to the water line, whereas hatchlings that had been denied this experience headed eastward, a direction of unclear origin. These data suggest that a short crawl across the natural beach can set the direction in which the young turtles subsequently move. The crawling experience was sufficient to acquire the compass course that they later follow, probably with the help of a magnetic compass, not only in the water, but already while still on land.     

Ant-lepidopteran associations along African forest edges

Working along forest edges, we aimed to determine how some caterpillars can co-exist with territorially dominant arboreal ants (TDAAs) in tropical Africa. We recorded caterpillars from 22 lepidopteran species living in the presence of five TDAA species. Among the defoliator and/or nectarivorous caterpillars that live on tree foliage, the Pyralidae and Nymphalidae use their silk to protect themselves from ant attacks. The Notodontidae and lycaenid Polyommatinae and Theclinae live in direct contact with ants; the Theclinae even reward ants with abundant secretions from their Newcomer gland. Lichen feeders (lycaenid; Poritiinae), protected by long bristles, also live among ants. Some lycaenid Miletinae caterpillars feed on ant-attended membracids, including in the shelters where the ants attend them; Lachnocnema caterpillars use their forelegs to obtain trophallaxis from their host ants. Caterpillars from other species live inside weaver ant nests. Those of the genus Euliphyra (Miletinae) feed on ant prey and brood and can obtain trophallaxis, while those from an Eberidae species only prey on host ant eggs. Eublemma albifascia (Erebidae) caterpillars use their thoracic legs to obtain trophallaxis and trophic eggs from ants. Through transfer bioassays of last instars, we noted that herbivorous caterpillars living in contact with ants were always accepted by alien conspecific ants; this is likely due to an intrinsic appeasing odor. Yet, caterpillars living in ant shelters or ant nests probably acquire cues from their host colonies because they were considered aliens and killed. We conclude that co-evolution with ants occurred similarly in the Heterocera and Rhopalocera.     

In search of the sky compass in the insect brain

Like many vertebrate species, insects rely on a sun compass for spatial orientation and long- range navigation. In addition to the sun, however, insects can also use the polarization pattern of the sky as a reference for estimating navigational directions. Recent analysis of polarization vision pathways in the brain of orthopteroid insects sheds some light onto brain areas that might act as internal navigation centers. Here I review the significance, peripheral mechanisms, and central processing stages for polarization vision in insects with special reference to the locust Schistocerca gregaria. As in other insect species, polarization vision in locusts relies on specialized photoreceptor cells in a small dorsal rim area of the compound eye. Stages in the brain involved in polarized light signaling include specific areas in the lamina, medulla and lobula of the optic lobe and, in the midbrain, the anterior optic tubercle, the lateral accessory lobe, and the central complex. Integration of polarized-light signals with information on solar position appears to start in the optic lobe. In the central complex, polarization-opponent interneurons form a network of interconnected neurons. The organization of the central complex, its connections to thoracic motor centers, and its involvement in the spatial control of locomotion strongly suggest that it serves as a spatial organizer within the insect brain, including the functions of compass orientation and path integration. Time compensation in compass orientation is possibly achieved through a neural pathway from the internal circadian clock in the accessory medulla to the protocerebral bridge of the central complex.     

<Emphasis Type="Italic">Tenebrio</Emphasis> beetles use magnetic inclination compass

Animals that guide directions of their locomotion or their migration routes by the lines of the geomagnetic field use either polarity or inclination compasses to determine the field polarity (the north or south direction). Distinguishing the two compass types is a guideline for estimation of the molecular principle of reception and has been achieved for a number of animal groups, with the exception of insects. A standard diagnostic method to distinguish a compass type is based on reversing the vertical component of the geomagnetic field, which leads to the opposite reactions of animals with two different compass types. In the present study, adults of the mealworm beetle Tenebrio molitor were tested by means of a two-step laboratory test of magnetoreception. Beetles that were initially trained to memorize the magnetic position of the light source preferred, during the subsequent test, this same direction, pursuant geomagnetic cues only. In the following step, the vertical component was reversed between the training and the test. The beetles significantly turned their preferred direction by 180 degrees. Our results brought until then unknown original findings that insects, represented here by the T. molitor species, use-in contrast to another previously researched Arthropod, spiny lobster-the inclination compass.     

Intercontinental chemical variation in the invasive ant Wasmannia auropunctata (Roger) (Hymenoptera Formicidae): a key to the invasive success of a tramp species

Unicoloniality emerges as a feature that characterizes successful invasive species. Its underlying mechanism is reduced intraspecific aggression while keeping interspecific competitiveness. To that effect, we present here a comparative behavioural and chemical study of the invasive ant Wasmannia auropunctata in parts of its native and introduced ranges. We tested the hypothesis that introduced populations (New Caledonia archipelago) have reduced intraspecific aggression relative to native populations (e.g., Ilhéus area, Brazil) and that this correlates with reduced variability in cuticular hydrocarbons (CHCs). As predicted, there was high intraspecific aggression in the Brazilian populations, but no intraspecific aggression among the New Caledonian populations. However, New Caledonian worker W. auropunctata remained highly aggressive towards ants of other invasive species. The chemical data corresponded with the behaviour. While CHCs of ants from the regions of Brazil diverged, the profiles of ants from various localities in New Caledonia showed high uniformity. We suggest that in New Caledonia W. auropunctata appears to behave as a single supercolony, whereas in its native range it acts as a multicolonial species. The uniformity of recognition cues in the New Caledonia ants may reflect a process whereby recognition alleles became fixed in the population, but may also be the consequence of a single introduction event and subsequent aggressive invasion of the ecosystem. Chemical uniformity coupled with low intraspecific but high interspecific aggression, lend credence to the latter hypothesis.     

Selective olfactory attention of a specialised predator to intraspecific chemical signals of its prey

Prey-specialised predators have evolved specific cognitive adaptations that increase their prey searching efficiency. In particular, when the prey is social, selection probably favours the use of prey intraspecific chemical signals by predatory arthropods. Using a specialised ant-eating zodariid spider, Zodarion rubidum, which is known to prey on several ant species and possesses capture and venom adaptations more effective on Formicinae ants, we tested its ability to recognise chemical cues produced by several ant species. Using an olfactometer, we tested the response of Z. rubidum towards air with chemical cues from six different ant species: Camponotus ligniperda, Lasius platythorax and Formica rufibarbis (all Formicinae); and Messor structor, Myrmica scabrinodis and Tetramorium caespitum (all Myrmicinae). Z. rubidum was attracted to air carrying chemical cues only from F. rufibarbis and L. platythorax. Then, we identified that the spiders were attracted to airborne cues coming from the F. rufibarbis gaster and Dufour's gland, in particular. Finally, we found that among several synthetic blends, the decyl acetate and undecane mixture produced significant attraction of spiders. These chemicals are produced only by three Formicine genera. Furthermore, we investigated the role of these chemical cues in the communication of F. rufibarbis and found that this blend reduces their movement. This study demonstrates the chemical cognitive capacity of Z. rubidum to locate its ant prey using chemical signals produced by the ants. The innate capacity of Z. rubidum to olfactory detect different ant species is narrow, as it includes only two ant genera, confirming trophic specialisation at lower than subfamily level. The olfactory cue detected by Zodarion spiders is probably a component of the recruitment or trail pheromone.