How does the ant know the way home without guiding clues on the desert plain?

Diana Tofan1 Group 30413 E-mail: diana_tofan@ymail.com

Abstract. Desert ants of the genus Cataglyphis have fascinating navigation abilities. On their foraging excursions, they cover distances up to hundred thousand body lengths. Having found food, they return to their nests with high precision, using path integration as their major navigation aid. This essay focuses on how these ants measure their travelling distances and how they use the location of the sun in the sky for guidance. It also presents the pedometer hypothesis which states that ants count their steps, along with some experiments done in this area. Although path integration is error prone due to its susceptibility to accumulate errors, ants use landmarks additionally.

1. Introducing the Cataglpyhis genus Most animals actively search for food and this requires an ability to navigate efficiently in their own habitat. The most important navigational task is to return to certain places. The desert ants of the genus Cataglyphis live in flat and rather hostile habitats, with high ambient temperatures, where they search for food, mostly insects. In contrast to other ant species, they do not rely on pheromones to mark their trail. The lack of scent trails is due to constraints of their habitat. Since the position of the bait is unpredictable, each ant performs large excursions on its own until it finds a food item. From this point, the ant returns to its nest on a rather straight way. Not only is the navigational feat quite remarkable, but also the physical endurance is fascinating. Foraging excursions can extend as far as some hundred meters, corresponding to ten-thousands of body lengths, and the ants move at the astonishing average speed of 50 cm/s and even more. 2. Path integration There are three principal strategies to solve the problem of returning to a specific point in space. The first, route following, consists of applying some marks to the outward trail and to follow these marks back to the starting point. Another method is to build up an internal representation of the surroundings, in which the geometrical relations between the locations are stored, and to use this cognitive map as a reference system to navigate in known areas, by comparing external cues with their internal counterparts. The North-African ant Cataglyphis fortis follows the third strategy, which is called path integration. Path integration requires the ability to start at a fixed point, visit several locations, and return directly to the original point. To accomplish this, the ant simply computes its return distance and direction vector from the vectors joining the locations on its route. This saves retracing steps. Path integration involves a complexity of neuronal mechanisms that lies between the route following and the map strategies mentioned above.

To whom any correspondence should be addressed.

Figure 1. The three orientation strategies to return to a point in space (left: route following, middle: path integration, right: cognitive map)

Example: Figure 2 represents a foraging path made up of several segments, starting from the nest at point (0, 0). Each segment is represented by a vector, for example the first segment is given by the vector v1 = (1, 2). The sum of the first i vectors gives us a vector which points from the origin (0, 0) to the current location. The negative of the vector points from the current location straight back to the nest. So to know our way back home, we don't need to remember all the vectors we travelled along we simply add the current one to the last total and take the negative.

Figure 2. Adding the first two vectors v1 and v2 gives a new vector v that points from the origin to the position after two steps. The same works when we add the first i vectors: the resulting vector points from the origin to our position after i steps.

Path integration is error-prone. ven small s!stematic errors tend to accumulate in the process. "ndeed, ants that were trained to cover large distances in a linear channel showed increasing errors in their distance estimates. #ow can ants avoid a $ potentiall! disastrous $ long term accumulation of errors% &ne solution is to reset the path integrator to a 'ero state at certain places. Ants reset their path integrator to 'ero when reentering the nest after a foraging trip, so the! avoid error accumulation to carr! over across repeated foraging e(cursions. )emarka*l!, coming ver! close to the nest entrance, while not entering the nest, is not sufficient to start the reset process.

+onetheless, it is still not well understood how the ants o*tain the necessar! accurac! of the path integrator in e(cursions over ten thousands of *od! lengths. "ndeed, ants do emplo! two kinds of safet! programs that help to compensate for errors of the navigational tool kit, as well as in case of accidental displacements of an animal *! wind gusts. The first is the s!stematic search strateg! that ants appl! if the! do miss the nest after having run off the home vector. Ants can even ad,ust their search program to the uncertaint! that increases with distance.

3. Another methods to approximate distances Ants can approximate distances by counting their steps and use the location of the sun in the sky as a tool in representing the direction back to the nest. Through evolution, ants have developed neural circuits in their brain which can take information about distance and direction and produce an output which is a continuously updated home vector. The ants use this information when they have found food or if they are attacked by a predator and need to get back home quick. 3.1. The location of the sun In a simple experiment, scientists used a set of mirrors to alter the perceived location of the sun in the sky. And almost immediately, as soon as the mirrors were in place, the ants altered their paths, using the new location of the sun as their guide. This raised a new problem though: the sun moves across the sky. In another experiment, the researchers waited for an ant leave the nest, find the bait, and then trapped it under a box for a few hours. In that time, the sun would have moved across the sky, and wouldnt have been a reliable tool to use to calculate the return trajectory. The key to this experiment was that while trapped under the box, the ants would not have been able to continuously update their representation of the nests direction based on the suns motion. If they rely on the sun as an external cue, then they would make systematic errors when attempting to navigate back to the nest after being released from their temporary prisons. On the other hand, if they simply use the suns location as a way to calibrate their own internal clocks, then they would have no trouble getting back home. In another impressive win for the ants, they indeed accurately found their way home. This means that these desert ants maintain a mental representation of the passage of time, even when the movement of the sun is hidden from them. Taken together, this tells us something about how ants determine the direction home: they use the suns motion across the sky in order to calibrate an internal mental clock. 3.2. Counting steps Researchers proposed the pedometer hypothesis: perhaps ants simply count the number of steps that they take. Pig bristles were glued to the ants legs, creating ants on stilts. By varying the length of the stride (longer for stilts, shorter for stumps), and measuring the distance the ant walks to get back home, we can see if it was counting the steps that it takes. If ants count their steps, than those on stilts should overestimate the distance home, and those with stumps should underestimate the distance home. It turned out that they did indeed count their steps. The stumpy ants underestimated the distance home, and the stilt-walkers overestimated the distance home. More evidence comes from some ants who were given stilts or stumps prior to leaving the nest: these ants had no problem getting back home, since their legs were of equivalent length for both the outbound and return parts of their journey.

Figure 3. Ant legs 4. Measuring the inclinations

Figure 4. Ant on stilts

Figure 5. Testing ants- orientation in ./

(Training channel in the foreground. The nest site on the left was surrounded *! an enclosure in order to lead the ants into the hill channel. The feeder was situated at the end of the 0th hill. Also visi*le are the two longer test channels, laid out in parallel to the training course, to which the ants were transferred from the feeder and released at the far end.)

The conclusion from these results was that ants must have a means of measuring the inclination of their path when walking over hill! terrains, and must *e a*le to feed this information into their path integration module to compute the *ase line distance from the respective slopes and walking distances. These results underline the notion that the ant1s pedometer must *e still more comple( than the stride integrator. /espite the intensive efforts, it is still unclear how the ants actuall! measure ascending or descending path segments. An investigation of the potential contri*ution of several groups of hair sensors, *! shaving or immo*ili'ing hair fields, has not revealed an! clue of the mechanism used *! these ants so far.

5. Conclusion
Although an ant-s *rain weighs onl! 2.1 milligrams, it produces a clever *ehavior when it comes to finding its wa! *ack to the nest. " think ants are smarter than the average insect, *ecause the! can-t distinguish o*,ects ver! clearl! and are still a*le to reach their goals *! using comple( mechanisms which impl! local geometr!. 6. References 314 Andrew Philippides and Paul 5raham 6211 Finding Your Way Home Without Knowing Where You Are (7ondon: Plus 8aga'ine) 364 9ason 5 5oldman 6216 Desert Ants Are Better Than Most High School Students at Trigonometry (+ew :ork: ;cientific American) [3] Bernhard Ronacher 2008 Path Integration as the Basic Navigation Mechanism of the Desert Ant Cataglyphis Fortis (Berlin: Myrmecological News) [4] David Andel and Rdiger Wehner Path Integration in Desert Ants, Cataglyphis: How to Make a Homing Ant Run Away From Home (London: The Royal Society)