Vector calibration in Australian desert ants,Melophorus bagoti: Effects of a delay after the acquisition of vector information

Desert ants navigate by using two chief strategies: path integration, keeping track of the straight‐line distance and direction to the starting point as they travel, and landmark guidance, orientation based on the visual panorama. Both Cataglyphis ants in North Africa and Melophorus bagoti in Central Australia are known to adjust their vectors derived from path integration to compensate for mismatches between their outbound direction of travel and (the reverse of) the inbound direction of travel that takes them home, a process known as vector calibration. We created mismatches of 90° between the outbound vector and the homebound direction by displacing ants from a feeder before their homebound run. We examined temporal factors in vector calibration by varying the delay (0, 1 or 3 hr) between the outbound run to the feeder and the homebound run from the displacement site. According to the temporal weighting rule, such a delay should decrease the weight given to the vector information obtained from the outbound run. This in turn should favour reliance on the visual panorama and thus speed up calibration. Results did not support this prediction. At the displacement site, a delay had little effect on the extent of calibration or the speed of calibration (the number of trials to reach maximum calibration). Just before being displaced, ants were also tested in a test ring surrounded by high walls that obliterated the visual scenery. In the test ring, a delay made the ants less likely to rely on their vector: ants were often not oriented as a group. Otherwise, the ants in the test ring also did not calibrate any more or any faster.     

Path integration in desert ants, Cataglyphis: how to make a homing ant run away from home

Path integration is an ant's lifeline on any of its foraging journeys. It results in a homebound global vector that continually informs the animal about its position relative to its starting point. Here, we use a particular (repeated training and displacement) paradigm, in which homebound ants are made to follow a familiar landmark route repeatedly from the feeder to the nest, even after they have arrived at the nest. The results show that during the repeated landmark-guided home runs the ant's path integrator runs continually, so that the current state of the homebound vector increasingly exceeds the reference state. The dramatic result is that the homing ants run away from home. This finding implies that the ants do not rely on cartographic information about the locations of nest and feeder (e.g. that the nest is always south of the feeder), but just behave according to what the state of their egocentric path integrator tells them.     

Landmark cues can change the motivational state of desert ant foragers

Desert ants of the genus Cataglyphis rely on path integration vectors to return to the nest (inbound runs) and back to frequently visited feeding sites (outbound runs). If disturbed, e.g., experimentally displaced on their inbound runs, they continue to run off their home-bound vector, but if disturbed in the same way on their outbound runs, they do not continue their feeder-based vector, but immediately switch on the home-bound state of their path integration vector and return to the nest. Here we show that familiar landmarks encountered by the ants during their run towards the feeder can change the ants’ motivational state insofar that the ants even if disturbed continue to run in the nest-to-feeder direction rather than reverse their courses, as they do in landmark-free situations. Hence, landmark cues can cause the ants to change their motivational state from homing to foraging.     

Three-dimensional orientation in desert ants: context-independent memorisation and recall of sloped path segments

Desert ants (Cataglyphis fortis) navigate by a combination of path integration and landmark-based route memories. Their ability to correct sloped path segments to their ground distances enables them to orientate accurately even in undulating terrain. In this study, we tested whether or not ants incorporate vertical components of an itinerary into their route memory in similar ways as they do with visual landmarks and horizontal changes of direction. In two separate experiments, we trained desert ants to walk over artificial hills and later tested their acceptance of slopes within novel contexts. In the first paradigm, ants had to traverse a hill only on their outbound run, but not on their homebound trip. In a follow-up experiment, we confronted ramp-trained animals with descents in a completely new temporal and spatial context. The animals transferred their newly acquired acceptance of slopes from the outbound to the homebound run as well as to novel foraging trips. Cataglyphis obviously dissociates the experience of sloped path segments from the original context in which they appeared, thus reducing their significance as a navigational aid.     

Calibration processes in desert ant navigation: vector courses and systematic search

This study investigates the ability of desert ants to adapt their path integration system to an "open-jaw" training paradigm, in which the point of arrival (from the nest) does not coincide with the point of departure (to the nest). Upon departure the ants first run off their home vector and then start a systematic search for the nest. Even if they are subjected to this training-around-a-circuit procedure for more than 50 times in succession, they never adopt straight homeward courses towards the nest. Their path integration vector gets slightly recalibrated (pointing a bit closer to the nest), and their search pattern gets asymmetric (with its search density peak shifted towards the nest), but the bipartite structure of the inbound trajectory invariably remains. These results suggest (1). that the ants cannot learn separate inbound and outbound vectors (i.e. vectors that are not 180 degrees reversals of each other), (2). that the recalibrated vector is dominated by the ant's outbound course, (3). that the recalibration of the vector and the modification of the search geometry are fast and flexible processes occurring whenever the ant experiences a mismatch between the stored and actual states of its path integrator.     

How do insects represent familiar terrain?

We argue here that ants and bees have a piecemeal representation of familiar terrain. These insects remember no more than what is needed to sustain the separate and parallel strategies that they employ when travelling between their nest and foraging sites. One major strategy is path integration. The insect keeps a running tally of its distance and direction from the nest and so can always return home. This global path integration is enhanced by long-term memories of significant sites that insects store in terms of the coordinates (direction and distance) of these sites relative to the nest. With these memories insects can plan routes that are steered by path integration to such sites. Quite distinct from global path integration are memories associated with familiar routes. Route memories include stored views of landmarks along the route with, in some cases, local vectors linked to them. Local vectors by encoding the direction and/or distance from one landmark to the next, or from one landmark to a goal, help an insect keep to a defined route. We review experiments showing that although local vectors can be recalled by recognising landmarks, the global path integration system is independent of landmark information and that landmarks do not have positional coordinates associated with them. The major function of route landmarks is thus procedural, telling an insect what action to perform next, rather than its location relative to the nest.     

Optimal cue integration in ants

In situations with redundant or competing sensory information, humans have been shown to perform cue integration, weighting different cues according to their certainty in a quantifiably optimal manner. Ants have been shown to merge the directional information available from their path integration (PI) and visual memory, but as yet it is not clear that they do so in a way that reflects the relative certainty of the cues. In this study, we manipulate the variance of the PI home vector by allowing ants (Cataglyphis velox) to run different distances and testing their directional choice when the PI vector direction is put in competition with visual memory. Ants show progressively stronger weighting of their PI direction as PI length increases. The weighting is quantitatively predicted by modelling the expected directional variance of home vectors of different lengths and assuming optimal cue integration. However, a subsequent experiment suggests ants may not actually compute an internal estimate of the PI certainty, but are using the PI home vector length as a proxy.     

Use of Long-Term Stored Vector Information in the Neotropical Ant <Emphasis Type="BoldItalic">Gigantiops destructor</Emphasis>

We investigated how the formicine ant Gigantiops destructor can use vector information to navigate within the cluttered environment of the rain forest. Displaced foragers use skylight information to move in the theoretical feeder-to-nest direction, whether they are prevented from updating their path-integrator during foraging or captured at the departure from their nest, i.e. with a current accumulator state very close to zero. Only ants that have collected food are able to download a long-term stored reference vector pointing in the nest direction, irrespective of the current accumulator state of their path-integrator stored in a working memory and independent of familiar landmarks. Depending on the release sites, ants that became lost at a maximum distance of 50 cm could still hit and recognize their familiar route, or they engaged in a systematic search for it centered on the release sites. In contrast to Cataglyphis desert ants, Gigantiops ants do not rely primarily on the current accumulator state of their egocentric path integrator. Such a long-term vector-based navigation primed by food capture is well adapted for a tropical ant foraging during periods spanning several hours. This could prevent the numerous cumulative errors in the evaluation of the angles steered that might result from a continuously running path-integrator operating during complex foraging patterns performed at ground or arboreal levels and during passive displacement in response to heavy rain.     

Just follow your nose: homing by olfactory cues in ants

How is an ant-equipped with a brain that barely exceeds the size of a pinhead-capable of achieving navigational marvels? Even though evidences suggest that navigation is a multimodal process, ants heavily depend on olfactory cues-of pheromonal and non-pheromonal nature-for foraging and orientation. Recent studies have directed their attention to the efficiency of pheromone trail networks. Advances in neurophysiological techniques make it possible to investigate trail pheromone processing in the ant's brain. In addition to relying on pheromone odours, ants also make use of volatiles emanating from the nest surroundings. Deposited in the vicinity of the nest, these home-range markings help the ants to home after a foraging run. Furthermore, olfactory landmarks associated with the nest enhance ants' homing abilities.     

Interactions of the polarization and the sun compass in path integration of desert ants

Desert ants, Cataglyphis fortis, perform large-scale foraging trips in their featureless habitat using path integration as their main navigation tool. To determine their walking direction they use primarily celestial cues, the sky’s polarization pattern and the sun position. To examine the relative importance of these two celestial cues, we performed cue conflict experiments. We manipulated the polarization pattern experienced by the ants during their outbound foraging excursions, reducing it to a single electric field (e-)vector direction with a linear polarization filter. The simultaneous view of the sun created situations in which the directional information of the sun and the polarization compass disagreed. The heading directions of the homebound runs recorded on a test field with full view of the natural sky demonstrate that none of both compasses completely dominated over the other. Rather the ants seemed to compute an intermediate homing direction to which both compass systems contributed roughly equally. Direct sunlight and polarized light are detected in different regions of the ant’s compound eye, suggesting two separate pathways for obtaining directional information. In the experimental paradigm applied here, these two pathways seem to feed into the path integrator with similar weights.     

Desert ants (Melophorus bagoti) navigating with robustness to distortions of the natural panorama

Many insects are known to use the terrestrial visual panorama for navigation. Research suggests that large-scale panoramic properties are often used for orientation rather than individual objects, usually called landmarks. We degraded the natural panorama encountered by Australian red honey ants, Melophorus bagoti, to test how robust their orientation based on the terrestrial panorama is. Foraging ants were lured to a feeder at a constant location. Trained ants were allowed to run home individually with food, but were captured just before they entered their nest. The tested ant was brought back to the location of the feeder, now covered, and allowed to run home again under different distortions of the natural panorama. In one experiment, a large tract of the view on one side of the feeder was obstructed by a tall plastic sheet. In a second experiment, the visual heights of terrestrial objects were altered by raising or lowering the ant by 80 cm. Under both kinds of distortions, the ants continued to be well oriented in the homeward direction. Navigation based on the natural terrestrial panorama proved robust to large distortions.     

Views,landmarks, and routes: how do desert ants negotiate an obstacle course?

The Australian desert ant Melophorus bagoti often follows stereotypical routes through a cluttered landscape containing both distant panoramic views and obstacles (plants) to navigate around. We created an artificial obstacle course for the ants between a feeder and their nest. Landmarks comprised natural objects in the landscape such as logs, branches, and tussocks. Many ants travelled stereotypical routes home through the obstacle course in training, threading repeatedly the same gaps in the landmarks. Manipulations altering the relations between the landmarks and the surrounding panorama, however, affected the routes in two major ways. Both interchanging the positions of landmarks (transpositions) and displacing the entire landmark set along with the starting position of the ants (translations) (1) reduced the stereotypicality of the route, and (2) increased turns and meanders during travel. The ants might have used the entire panorama in view-based travel, or the distal panorama might prime the identification and use of landmarks en route. Despite the large data set, both options (not mutually exclusive) remain viable.     

Can the antCataglyphis cursor (Hymenoptera: Formicidae) encode global landmark-landmark relationships in addition to isolated landmark-goal relationships?

The antCataglyphis cursor was tested for its landmark-based homing in a laboratory setting. Workers were induced to go down a tube at the center of an arena to forage. On the periphery of the arena were four different black shapes serving as the only distinguishing visual landmarks, i.e., a cross, a circle, a triangle, and a square. The purpose was to show that the spatial memory of ants represents something of the overall arrangement of landmarks. When first released into the arena, the ants were not oriented toward home in their navigation. After 2 days of free access in the usual landmark setup, the ants learned to orient rapidly significantly goalward. When landmarks were all removed, they did not orient in any direction significantly. When the landmarks were rotated by 90°, their compass positions were changed but their relative positions maintained, and the ants rotated their heading by a similar amount. This rotated homing direction implies that the array of landmarks was used as the only source of directional determination. When the landmark nearest their home was absent, but the other three were in their usual places, the ants were slightly homeward oriented at one-quarter of the way, but not at one-half of the way when the other landmarks were behind them. When the landmarks were randomly permuted, both their compass positions and their overall spatial relationships were altered, and the ants were not significantly oriented in any direction. These results indicate that spatial memory in the antC. cursor encodes global landmark-landmark relations. Thus, ants can abstract certain topological properties of their environment.     

How do insects use path integration for their navigation?

 We combine experimental findings on ants and bees, and build on earlier models, to give an account of how these insects navigate using path integration, and how path integration interacts with other modes of navigation. At the core of path integration is an accumulator. This is set to an initial state at the nest and is updated as the insect moves so that it always reports the insect's current position relative to the nest. Navigation that uses path integration requires, in addition, a way of storing states of the accumulator at significant places for subsequent recall as goals, and a means of computing the direction to such goals. We discuss three models of how path integration might be used for this process, which we call vector navigation. Vector navigation is the principal means of navigating over unfamiliar terrain, or when landmarks are unavailable. Under other conditions, insects often navigate by landmarks, and ignore the output of the vector navigation system. Landmark navigation does not interfere with the updating of the accumulator. There is an interesting symmetry in the use of landmarks and path integration. In the short term, vector navigation can be independent of landmarks, and landmark navigation needs no assistance from path integration. In the longer term, visual landmarks help keep path vector navigation calibrated, and the learning of visual landmarks is guided by path integration. Received: 6 June 1999 / Accepted in revised form: 20 March 2000     

Some psychophysics of the pigeon's use of landmarks

1. Three pigeons (Columba livia) were trained to find hidden food in a sunken well (3.3 cm in diameter) at a constant place within an (160 cm x 160 cm) experimental box (Fig. 1). After learning the location, the animals were tested occasionally with the well and food absent. Landmarks in the experimental box might be transformed on such tests. 2. Changing the height or width of a nearby landmark had no systematic influence on the position of peak search. Translating a nearby landmark, however, led to a shift in peak search position. All three birds then searched most somewhere between the original goal location, as defined by the unmoved landmarks, and the goal location as defined by the shifted landmark. Within a limited range of landmark shift, the peak shift as a function of landmark shift is linear (Fig. 3). 3. To explain the data (Fig. 7), the pigeon records at the location of the goal the algebraic vectors from a number of landmarks to the goal. These vectors have both a direction and a distance component. When searching for the goal again in the experimental box, it computes independently for each landmark a navigation vector. This is arrived at by vector-adding the algebraic vector from the bird's current position to the landmark in question, supplied by perception, to the corresponding landmark-goal vector in its record. The pigeon moves in the direction and distance specified by a weighted average of the independently calculated navigation vectors. For positive vector weights, vector geometry guarantees that the bird would search somewhere between the original goal and the goal according to the shifted landmark. The extent to which it shifts toward the shifted goal reflects the vector weight given to the shifted landmark.     

Search strategies of ants in landmark-rich habitats

Search is an important tool in an ant’s navigational toolbox to relocate food sources and find the inconspicuous nest entrance. In habitats where landmark information is sparse, homing ants travel their entire home vector before searching systematically with ever increasing loops. Search strategies have not been previously investigated in ants that inhabit landmark-rich habitats where they typically establish stereotypical routes. Here we examine the search strategy in one such ant, Melophorus bagoti, by confining their foraging in one-dimensional channels to determine if their search pattern changes with experience, location of distant cues and altered distance on the homebound journey. Irrespective of conditions, we found ants exhibit a progressive search that drifted towards the fictive nest and beyond. Segments moving away from the start of the homeward journey were longer than segments heading back towards the start. The right tail distribution of segment lengths was well fitted by a power function, but slopes less than −3 on a log-log plot indicate that the process cannot be characterized as Lévy searches that have optimal slopes near –2. A double exponential function fits the distribution of segment lengths better, supporting another theoretically optimal search pattern, the composite Brownian walk.     

Mapping the navigational knowledge of individually foraging ants,Myrmecia croslandi

Ants are efficient navigators, guided by path integration and visual landmarks. Path integration is the primary strategy in landmark-poor habitats, but landmarks are readily used when available. The landmark panorama provides reliable information about heading direction, routes and specific location. Visual memories for guidance are often acquired along routes or near to significant places. Over what area can such locally acquired memories provide information for reaching a place? This question is unusually approachable in the solitary foraging Australian jack jumper ant, since individual foragers typically travel to one or two nest-specific foraging trees. We find that within 10 m from the nest, ants both with and without home vector information available from path integration return directly to the nest from all compass directions, after briefly scanning the panorama. By reconstructing panoramic views within the successful homing range, we show that in the open woodland habitat of these ants, snapshot memories acquired close to the nest provide sufficient navigational information to determine nest-directed heading direction over a surprisingly large area, including areas that animals may have not visited previously.     

Navigating desert ants (Cataglyphis fortis) learn to alter their search patterns on their homebound journey

In navigating home, desert ants first run off a global vector estimated on their outbound journey, and then engage in systematic search consisting of ever‐increasing loops interrupted by returns to the starting point of search. Desert ants (Cataglyphis fortis; Wehner, 1983 ) were trained to travel 6 m down a channel to a food source. Different groups of ants were trained to return home in another channel, from distances of 6 m (control), 9 m or 12 m. Ants at the feeder were then tested in a long test channel. The measure of where the ants first turned back on a test gave an estimate of the length of the global vector calculated on their outbound trip. The median distance of search on a 5‐min test gave an estimate of the centre of the search pattern. Relative to controls, the experimental ants did not increase their estimated length of global vector, but changed their search patterns, searching on average further from the start than the controls. Tests of the outbound journey, however, revealed no differences between groups. Desert ants can learn to modify their search pattern based on experience.     

Snapshot memories and landmark guidance in wood ants

Insects are thought to pinpoint a place by using memorized "snapshots," i.e., two-dimensional retinotopic views of the surrounding landmarks recorded when at the place (reviewed in ). Insects then reach the place by moving until their current view matches their snapshot. To determine when snapshots are recalled, and how differences between view and snapshot are translated into appropriate movements, we analyzed the approaches of wood ants to a feeding site that was located in the center of an array of two or three cylinders. In ants, contrary to flying hymenopterans, body orientation and direction of travel are collinear, so that an ant approaching an object always looks at it with frontal visual field. On their way to a food site, ants fixated and approached a cylinder predominantly when its angular size was smaller than when viewed from the food site. This finding implies that ants store snapshots at this place while fixating landmarks with frontal retina, so simplifying the later alignment of snapshots with their current view. It also means that ants recall snapshots well in advance of reaching the place. Although snapshots are centered on a landmark, we show that they extend at least 120 degrees into the periphery.     

Visual homing in analog hardware

Insects of several species rely on visual landmarks for returning to important locations in their environment. The "average landmark vector model" is a parsimonious model which reproduces some aspects of the visual homing behavior of bees and ants. To gain insights in the structure and complexity of the neural apparatus that might underly the navigational capabilities of these animals, the average landmark vector model was implemented in analog hardware and used to control a mobile robot. The experiments demonstrate that the apparently complex task of visual homing might be realized by simple and mostly peripheral neural circuits in insect brains.