Cataglyphis desert ants are impressive navigators. When the foragers roam the desert, they employ path integration. For these ants, distance estimation is one key challenge. Distance information was ...thought to be provided by optic flow (OF)—that is, image motion experienced during travel—but this idea was abandoned when stride integration was discovered as an odometer mechanism in ants. We show that ants transported by nest mates are capable of measuring travel distance exclusively by the use of OF cues. Furthermore, we demonstrate that the information gained from the optic flowmeter cannot be transferred to the stride integrator. Our results suggest a dual information channel that allows the ants to measure distances by strides and OF cues, although both systems operate independently and in a redundant manner.
The diurnal thermophilic Saharan silver ant,
, is the fastest of the North African
desert ant species. These highly mobile ants endure the extreme temperatures of their sand dune environment with ...outstanding behavioural, physiological and morphological adaptations. Surprisingly,
has comparatively shorter legs than its well-studied sister species
from salt pan habitats. This holds despite the somewhat hotter surface temperatures and the more yielding sand substrate. Here, we report that
employs a different strategy in reaching high running speeds, outperforming the fastest known runs of the longer-legged
ants. Video analysis across a broad range of locomotor speeds revealed several differences to
Shorter leg lengths are compensated for by high stride frequencies, ranging beyond 40 Hz. This is mainly achieved by a combination of short stance phases (down to 7 ms) and fast leg swing movements (up to 1400 mm s
). The legs of one tripod group exhibit almost perfect synchrony in the timings of their lift-offs and touch-downs, and good tripod coordination is present over the entire walking speed range (tripod coordination strength values around 0.8). This near synchrony in leg movement may facilitate locomotion across the yielding sand dune substrate.
Air-cushioned spheres are widely used as treadmills to study behavioural and neurophysiological questions in numerous species. We describe an improved spherical treadmill design that reliably ...registers the path and walking behaviour of an animal walking on top of the sphere. The simple and robust set-up consists of a very light hollowed styrofoam ball supported by an air stream in a hollow half sphere and can be used indoors and outdoors. Two optical mouse sensors provided with lenses of 4.6 mm focal length detect the motion of the sphere with a temporal resolution of more than 200 frames s
and a spatial resolution of less than 0.2 mm. The treadmill can be used in an open- or closed-loop configuration with respect to yaw of the animal. The tethering allows animals to freely adjust their body posture and in the closed-loop configuration to quickly rotate around their yaw axis with their own moment of inertia. In this account, we present the first evidence of naturalistic homing navigation on a spherical treadmill for two species of
desert ants. We were able to evaluate with good precision the walking speed and angular orientation at any time. During homing the ants showed a significant difference in walking speed between the approach and search phases; moreover, they slowed down significantly as soon as they reached zero vector state, the fictive nest position.
Ant Odometer: Stepping on Stilts and Stumps Wittlinger, Matthias; Wehner, Rüdiger; Wolf, Harald
Science (American Association for the Advancement of Science),
06/2006, Letnik:
312, Številka:
5782
Journal Article
Recenzirano
Desert ants, Cataglyphis, navigate in their vast desert habitat by path integration. They continuously integrate directions steered (as determined by their celestial compass) and distances traveled, ...gauged by as-yet-unknown mechanisms. Here we test the hypothesis that navigating ants measure distances traveled by using some kind of step integrator, or "step counter." We manipulated the lengths of the legs and, hence, the stride lengths, in freely walking ants. Animals with elongated ("stilts") or shortened legs ("stumps") take larger or shorter strides, respectively, and concomitantly misgauge travel distance. Travel distance is overestimated by experimental animals walking on stilts and underestimated by animals walking on stumps.
Navigation plays an essential role for many animals leading a mobile mode of life, and for central place foragers in particular. One important prerequisite for navigation is the ability to estimate ...distances covered during locomotion. It has been shown that Cataglyphis desert ants, well-established model organisms in insect navigation, use two odometer mechanisms, namely, stride and optic flow integration. Although both mechanisms are well established, their mode of interaction to build one odometer output remains enigmatic. We tackle this problem by selectively covering the ventral eye parts in Cataglyphis fortis foragers, the eye regions responsible for optic flow input in odometry. Exclusion of optic flow cues was implemented during different sections of outbound and inbound travel. This demonstrated that the two odometers have separate distance memories that interact in determining homing distance. Possible interpretations posit that the two odometer memories (i) take on different relative weights according to context or (ii) compete in a winner-take-all mode. Explanatory values and implications of such interpretations are discussed. We are able to provide a rough quantitative assessment of odometer cue interaction. An understanding of the interaction of different odometer mechanisms appears valuable not only for animal navigation research but may inform discussions on sensor fusion in both behavioural contexts and potential technical applications.
Celotno besedilo
Dostopno za:
DOBA, IZUM, KILJ, NUK, PILJ, PNG, SAZU, SIK, UILJ, UKNU, UL, UM, UPUK
Cataglyphis ants are renowned for their impressive navigation skills, which have been studied in numerous experiments during forward locomotion. However, the ants' navigational performance during ...backward homing when dragging large food loads has not been investigated until now. During backward locomotion, the odometer has to deal with unsteady motion and irregularities in inter-leg coordination. The legs' sensory feedback during backward walking is not just a simple reversal of the forward stepping movements: compared with forward homing, ants are facing towards the opposite direction during backward dragging. Hence, the compass system has to cope with a flipped celestial view (in terms of the polarization pattern and the position of the sun) and an inverted retinotopic image of the visual panorama and landmark environment. The same is true for wind and olfactory cues. In this study we analyze for the first time backward-homing ants and evaluate their navigational performance in channel and open field experiments. Backward-homing Cataglyphis fortis desert ants show remarkable similarities in the performance of homing compared with forward-walking ants. Despite the numerous challenges emerging for the navigational system during backward walking, we show that ants perform quite well in our experiments. Direction and distance gauging was comparable to that of the forward-walking control groups. Interestingly, we found that backward-homing ants often put down the food item and performed foodless search loops around the left food item. These search loops were mainly centred around the drop-off position (and not around the nest position), and increased in length the closer the ants came to their fictive nest site.
North African desert ants, Cataglyphis fortis, are established model organisms in animal navigation research. Cataglyphis re-visit plentiful feeding sites, but their decision to return to a feeder ...and the organization of food searches has been little studied. Here we provide a review of recent advances regarding this topic. At least two parameters determine the ants' assessment of site quality, namely, amount of food available and reliability of food encounter on subsequent visits. The amount of food appears to be judged by the concentration of items at the food uptake site. Initially the amount of food in a feeder dominates the foragers' decision to return, whereas learning about reliability takes precedence in the course of a few visits. The location of a worthwhile site is determined by the animals' path integration system. In particular, the distance of the feeding site is memorized as the arithmetic average of the distances covered during the previous outbound and homebound journeys. Feeding sites that are small and inconspicuous cannot be approached directly with sufficient certainty, due to inevitable inaccuracies of the path integrator. Instead, desert ants steer downwind of the goal to encounter the odor plume emanating from the food and they follow this plume to the feeder. The angle steered downwind reflects the animals' maximal navigation error and is adjusted according to experience. In summary, food searches of desert ants provide an unexpected wealth of features that may advance our understanding of search, navigation, and decision strategies. There are several aspects that warrant further scrutiny.
Path integration, although inherently error-prone, is a common navigation strategy in animals, particularly where environmental orientation cues are rare. The desert ant Cataglyphis fortis is a ...prominent example, covering large distances on foraging excursions. The stride integrator is probably the major source of path integration errors. A detailed analysis of walking behaviour in Cataglyphis is thus of importance for assessing possible sources of errors and potential compensation strategies. Zollikofer (J Exp Biol 192:95–106, 1994a) demonstrated consistent use of the tripod gait in Cataglyphis, and suggested an unexpectedly constant stride length as a possible means of reducing navigation errors. Here, we extend these studies by more detailed analyses of walking behaviour across a large range of walking speeds. Stride length increases linearly and stride amplitude of the middle legs increases slightly linearly with walking speed. An initial decrease of swing phase duration is observed at lower velocities with increasing walking speed. Then it stays constant across the behaviourally relevant range of walking speeds. Walking speed is increased by shortening of the stance phase and of the stance phase overlap. At speeds larger than 370 mms⁻¹, the stride frequency levels off, the duty factor falls below 0.5, and Cataglyphis transitions to running with aerial phases.
► We tried to impair stride integration by amputating two of the six walking legs. ► Homing accuracy and homing certainty remain unaffected by walking disturbances. ► This was true even if ventral ...optic flow was eliminated, a 2nd odometry mechanism. ► Ventral optic flow input does not enhance search certainty in homing ants.
Desert ants gauge walking distance by means of a stride integrator and, to a minor extent, by optic flow integration. With the present experiments we attempt to interfere with both, stride integration and optic flow input in order to reveal possible interactions of the two modes of odometry and further functional details of the stride integrator.
We tried to impair stride integration by amputating two of the six walking legs. Amputation of left middle and right hind legs had especially severe effects since it left only the right front leg in one of the support tripods that are alternately used in walking. We tried to impair optic flow input – which is used for distance estimation to a minor extent – by covering both ventral eye halves. These two sets of manipulations were carried out in combination to study possible compensatory effects, for instance, of optic flow input in the case of an impaired stride integrator.
Unexpectedly, none of the manipulations we carried out had significant effects on homing performance. This was true with regard to homing distance estimation (as determined by the centres of the ants’ nest searches) and homing certainty (as determined by the search spreads). These results corroborate the surprising robustness of odometry by stride integration, and they indicate that leg proprioceptive feedback is used for stride integration. The question of a possible interaction of optic flow input and stride integration remains open.
Like many other animals, insects are capable of returning to previously visited locations using path integration, which is a memory of travelled direction and distance. Recent studies suggest that ...Drosophila can also use path integration to return to a food reward. However, the existing experimental evidence for path integration in Drosophila has a potential confound: pheromones deposited at the site of reward might enable flies to find previously rewarding locations even without memory. Here, we show that pheromones can indeed cause naïve flies to accumulate where previous flies had been rewarded in a navigation task. Therefore, we designed an experiment to determine if flies can use path integration memory despite potential pheromonal cues by displacing the flies shortly after an optogenetic reward. We found that rewarded flies returned to the location predicted by a memory-based model. Several analyses are consistent with path integration as the mechanism by which flies returned to the reward. We conclude that although pheromones are often important in fly navigation and must be carefully controlled for in future experiments, Drosophila may indeed be capable of performing path integration.
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