Celestial Compass Research Articles

Route retracing: way pointing and multiple vector memories in trail-following ants.

Maintaining positional estimates of goal locations is a fundamental task for navigating animals. Diverse animal groups, including both vertebrates and invertebrates, can accomplish this through path integration. During path integration, navigators integrate movement changes, tracking both distance and direction, to generate a spatial estimate of their start location, or global vector, allowing efficient direct return travel without retracing the outbound route. In ants, path integration is accomplished through the coupling of pedometer and celestial compass estimates. Within path integration, it has been theorized navigators may use multiple vector memories for way pointing. However, in many instances, these navigators may instead be homing via view alignment. Here, we present evidence that trail-following ants can attend to segments of their global vector to retrace their non-straight pheromone trails, without the confound of familiar views. Veromessor pergandei foragers navigate to directionally distinct intermediate sites via path integration by orienting along separate legs of their inbound route at unfamiliar locations, indicating these changes are not triggered by familiar external cues, but by vector state. These findings contrast with path integration as a singular memory estimate in ants and underscore the system's ability to way point to intermediate goals along the inbound route via multiple vector memories, akin to trapline foraging in bees visiting multiple flower patches. We discuss how reliance on non-straight pheromone-marked trails may support attending to separate vectors to remain on the pheromone rather than attempting straight-line shortcuts back to the nest.

Celestial compass sensor mimics the insect eye for navigation under cloudy and occluded skies

Insects use the sun’s position (even when concealed) as a compass for navigation by filtering celestial light intensity and polarisation through their compound eyes. To replicate this functionality, we present a sensor that imitates essential aspects of insect eyes, particularly the fan-like arrangement of polarised light receptors in their dorsal rim area. Our sensor comprises a ring of eight pairs of photodiodes (evaluating two orthogonal orientations of polarised light) to analyse the skylight coming from different directions. Because the layout of our sensor aligns with the polarised light pattern in the sky, a circular-mean model that integrates information spatially across the analysers can estimate the solar azimuth. When using the same sensor design, our model achieves lower compass errors than alternative (and computationally more complex) algorithms, especially under cloudy and occluded skies. Thus, the morphology and processing of the insect celestial compass provide an efficient and robust directional input for navigation.

Physiological properties of the visual system in the Green Weaver ant, Oecophylla smaragdina

The Green Weaver ants, Oecophylla smaragdina are iconic animals known for their extreme cooperative behaviour where they bridge gaps by linking to each other to build living chains. They are visually oriented animals, build chains towards closer targets, use celestial compass cues for navigation and are visual predators. Here, we describe their visual sensory capacity. The major workers of O. smaragdina have more ommatidia (804) in each eye compared to minor workers (508), but the facet diameters are comparable between both castes. We measured the impulse responses of the compound eye and found their response duration (42 ms) was similar to that seen in other slow-moving ants. We determined the flicker fusion frequency of the compound eye at the brightest light intensity to be 132 Hz, which is relatively fast for a walking insect suggesting the visual system is well suited for a diurnal lifestyle. Using pattern-electroretinography we identified the compound eye has a spatial resolving power of 0.5 cycles deg−1 and reached peak contrast sensitivity of 2.9 (35% Michelson contrast threshold) at 0.05 cycles deg−1. We discuss the relationship of spatial resolution and contrast sensitivity, with number of ommatidia and size of the lens.

Intricacies of running a route without success in night-active bull ants (Myrmecia midas).

How do ants resolve conflicts between different sets of navigational cues during navigation? When two cue sets point to diametrically opposite directions, theories predict that animals should pick one set of cues or the other. Here we tested how nocturnal bull ants Myrmecia midas adjust their paths along established routes if route following does not lead to their entry into their nest. During testing, foragers were repeatedly placed back along their homeward route up to nine times, a procedure called rewinding. This procedure produced an accumulating path integrator, or vector, in diametric opposition to the learned landmark views of the route. Repeated rewinding made some individuals head initially in the nest-to-feeder vector direction, but all ants ended up using the visual scene for homing, demonstrating the importance of view-based homing in this species. Repeated rewinding, however, led to path deteriorations; with increased path meander and scanning, results also found in desert ants. After nine rewinding trips, ants were displaced off their route in further manipulations, to a site near the nest, an unfamiliar site, or with the terrestrial surround entirely covered. The results show that a change in visual conditions diminished the weight accorded to path integration: the off-route ants no longer headed off in the vector direction as they did on the immediately preceding trial. They relied on celestial compass cues in other ways for homing. Experiment 2 showed the effects of rewinding in the unaltered natural habitat were not view-specific in these bull ants. (PsycInfo Database Record (c) 2023 APA, all rights reserved).

Anisotropic charge trapping in phototransistors unlocks ultrasensitive polarimetry for bionic navigation

Being able to probe the polarization states of light is crucial for applications from medical diagnostics and intelligent recognition to information encryption and bio-inspired navigation. Current state-of-the-art polarimeters based on anisotropic semiconductors enable direct linear dichroism photodetection without the need for bulky and complex external optics. However, their polarization sensitivity is restricted by the inherent optical anisotropy, leading to low dichroic ratios of typically smaller than ten. Here, we unveil an effective and general strategy to achieve more than 2,000-fold enhanced polarization sensitivity by exploiting an anisotropic charge trapping effect in organic phototransistors. The polarization-dependent trapping of photogenerated charge carriers provides an anisotropic photo-induced gate bias for current amplification, which has resulted in a record-high dichroic ratio of >104, reaching over the extinction ratios of commercial polarizers. These findings further enable the demonstration of an on-chip polarizer-free bionic celestial compass for skylight-based polarization navigation. Our results offer a fundamental design principle and an effective route for the development of next-generation highly polarization-sensitive optoelectronics.

Predicting performance of naïve migratory animals, from many wrongs to self-correction

Migratory orientation of many animals is inheritable, enabling inexperienced (naïve) individuals to migrate independently using a geomagnetic or celestial compass. It remains unresolved how naïve migrants reliably reach remote destinations, sometimes correcting for orientation error or displacement. To assess naïve migratory performance (successful arrival), we simulate and assess proposed compass courses for diverse airborne migratory populations, accounting for spherical-geometry effects, compass precision, cue transfers (e.g., sun to star compass), and geomagnetic variability. We formulate how time-compensated sun-compass headings partially self-correct, according to how inner-clocks are updated. For the longest-distance migrations simulated, time-compensated sun-compass courses are most robust to error, and most closely resemble known routes. For shorter-distance nocturnal migrations, geomagnetic or star-compass courses are most robust, due to not requiring nightly cue-transfers. Our predictive study provides a basis for assessment of compass-based naïve migration and mechanisms of self-correction, and supports twilight sun-compass orientation being key to many long-distance inaugural migrations.

Receptive field structures for two celestial compass cues at the input stage of the central complex in the locust brain.

Successful navigation depends on an animal's ability to perceive its spatial orientation relative to visual surroundings. Heading direction in insects is represented in the central complex (CX), a navigation center in the brain, to generate steering commands. In insects that navigate relative to sky compass signals, CX neurons are tuned to celestial cues indicating the location of the sun. The desert locust CX contains a compass-like representation of two related celestial cues: the direction of unpolarized direct sunlight and the pattern of polarized light, which depends on the sun position. Whether congruent tuning to these two compass cues emerges within the CX network or is inherited from CX input neurons is unclear. To address this question, we intracellularly recorded from GABA-immunoreactive TL neurons, which are input elements to the locust CX (corresponding to R neurons in Drosophila), while applying visual stimuli simulating unpolarized sunlight and polarized light across the hemisphere above the animal. We show that TL neurons have large receptive fields for both types of stimuli. However, faithful integration of polarization angles across the dorsal hemisphere, or matched-filter ability to encode particular sun positions, was found in only two out of 22 recordings. Those two neurons also showed a good match in sun position coding through polarized and unpolarized light signaling, whereas 20 neurons showed substantial mismatch in signaling of the two compass cues. The data, therefore, suggest that considerable refinement of azimuth coding based on sky compass signals occurs at the synapses from TL neurons to postsynaptic CX compass neurons.

Magnetosensation during re-learning walks in desert ants (Cataglyphis nodus)

At the beginning of their foraging careers, Cataglyphis desert ants calibrate their compass systems and learn the visual panorama surrounding the nest entrance. For that, they perform well-structured initial learning walks. During rotational body movements (pirouettes), naïve ants (novices) gaze back to the nest entrance to memorize their way back to the nest. To align their gaze directions, they rely on the geomagnetic field as a compass cue. In contrast, experienced ants (foragers) use celestial compass cues for path integration during food search. If the panorama at the nest entrance is changed, foragers perform re-learning walks prior to heading out on new foraging excursions. Here, we show that initial learning walks and re-learning walks are structurally different. During re-learning walks, foragers circle around the nest entrance before leaving the nest area to search for food. During pirouettes, they do not gaze back to the nest entrance. In addition, foragers do not use the magnetic field as a compass cue to align their gaze directions during re-learning walk pirouettes. Nevertheless, magnetic alterations during re-learning walks under manipulated panoramic conditions induce changes in nest-directed views indicating that foragers are still magnetosensitive in a cue conflict situation.

Light pollution forces a change in dung beetle orientation behavior

Increasing global light pollution1,2 threatens the night-time darkness to which most animals are adapted. Light pollution can have detrimental effects on behavior,3-5 including by disrupting the journeys of migratory birds,5,6 sand hoppers,7-9 and moths.10 This is particularly concerning, since many night-active species rely on compass information in the sky, including the moon,11,12 the skylight polarization pattern,13,14 and the stars,15 to hold their course. Even animals not directly exposed to streetlights and illuminated buildings may still experience indirect light pollution in the form of skyglow,3,4 which can extend far beyond urban areas.1,2 While some recent research used simulated light pollution to estimate how skyglow may affect orientation behavior,7-9 the consequences of authentic light pollution for celestial orientation have so far been neglected. Here, we present the results of behavioral experiments at light-polluted and dark-sky sites paired with photographic measurements of each environment. We find that light pollution obscures natural celestial cues and induces dramatic changes in dung beetle orientation behavior, forcing them to rely on bright earthbound beacons in place of their celestial compass. This change in behavior results in attraction toward artificial lights, thereby increasing inter-individual competition and reducing dispersal efficiency. For the many other species of insect, bird, and mammal that rely on the night sky for orientation and migration, these effects could dramatically hinder their vital night-time journeys.

Role of the pheromone for navigation in the group foraging ant, Veromessor pergandei.

Navigation is comprised of a variety of strategies which rely on multiple external cues to shape a navigator's behavioral output. Here, we explored in the ant Veromessor pergandei, the interactions between the information provided by the pheromone trail and the home vector guided by the celestial compass. We found that a cross sensory interaction between the pheromone cue and the path integrator underlies correct orientation during the inbound journey. The celestial compass provides directional information, while the presence of the trail pheromone acts as a critical context cue, triggering distinct behaviors (vector orientation, search, and backtracking). While exposed to the pheromone, foragers orient to the vector direction regardless of vector state, while in the pheromone's absence, the current remaining vector determines the forager's navigational behavior. This interaction also occurs in foragers with no remaining path integrator, relying on the activation of a celestial compass-based memory of the previous trip. Such cue interactions maximize the foragers' return to the nest and inhibit movement off the pheromone trail. Finally, our manipulations continuously rotated foragers away from their desired heading, yet foragers were proficient at counteracting these changes, steering to maintain a correct compass heading even at rotational speeds of ~ 40°/s.

Ontogeny of the star compass in birds: pied flycatchers (Ficedula hypoleuca) can establish the star compass in spring

The star compass of birds, as well as the sun compass, is not innate. To possess either of them, the birds have to observe the rotating sky and determine its' center of rotation (in the star compass case) or the sun's movement (in the sun compass). Young birds are believed to learn how to use the star compass before their first migration, even though the evidence of this is lacking. Here, we test whether hand-raised Pied flycatchers (Ficedula hypoleuca) which had not established the star compass prior to their first autumn migration can gain it later in their ontogeny, in spring. We also attempted to examine if the observation of diurnal celestial cues (the sun and the polarized light) prior to autumn migration would affect the process of the star compass learning in spring. When tested in the vertical magnetic field under the natural starry sky, the group of birds which observed the stars in spring as the first celestial cues could choose the migratory direction. In contrast, the birds, which had never seen the stars, were not able to use the nightly celestial cues in the vertical magnetic field. However, the group of birds which had seen the daytime celestial cues till autumn and the stars at spring was disoriented, although this might be due to the small sample size. Our data suggest the possibility that the star compass may be learned in spring and emphasize the necessity for the further research in the interaction of celestial compasses.

A red knot as a black swan: how a single bird shows navigational abilities during repeat crossings of the Greenland Icecap

Despite the wealth of studies on seasonal movements of birds between southern nonbreeding locations and High Arctic breeding locations, the key mechanisms of navigation during these migrations remain elusive. A flight along the shortest possible route between pairs of points on a sphere (‘orthodrome’) requires a bird to be able to assess its current location in relation to its migration goal and to make continuous adjustment of heading to reach that goal. Alternatively, birds may navigate along a vector with a fixed orientation (‘loxodrome’) based on magnetic and/or celestial compass mechanisms. Compass navigation is considered especially challenging for summer migrations in Polar regions, as continuous daylight and complexity in the geomagnetic field may complicate the use of both celestial and magnetic compasses here. We examine the possible use of orientation mechanisms during migratory flights across the Greenland Icecap. Using a novel 2 g solar‐powered satellite transmitter, we documented the flight paths travelled by a female red knotCalidris canutus islandicaduring two northward and two southward migrations. The geometry of the paths suggests that red knots can migrate across the Greenland Icecap along the shortest‐, orthodrome‐like, path instead of the previously suggested loxodrome path. This particular bird's ability to return to locations visited in a previous year, together with its sudden course changes (which would be appropriate responses to ambient wind fields), suggest a map sense that enables red knots to determine location, so that they can tailor their route depending on local conditions.

Path integration: how details of the honeybee waggle dance and the foraging strategies of desert ants might help in understanding its mechanisms.

Path integration is a navigational strategy that gives an animal an estimate of its position relative to some starting point. For many decades, ingenious and probing behavioural experiments have been the only window onto the operation of path integration in arthropods. New methods have now made it possible to visualise the activity of neural circuits in Drosophila while they fly or walk in virtual reality. Studies of this kind, as well as electrophysiological recordings from single neurons in the brains of other insects, are revealing details of the neural mechanisms that control an insect's direction of travel and other aspects of path integration. The aim here is first to review the major features of path integration in foraging desert ants and honeybees, the current champion path integrators of the insect world, and second consider how the elaborate behaviour of these insects might be accommodated within the framework of the newly understood neural circuits. The discussion focuses particularly on the ability of ants and honeybees to use a celestial compass to give direction in Earth-based coordinates, and of honeybees to use a landscape panorama to provide directional guidance for path integration. The possibility is raised that well-ordered behaviour might in some cases substitute for complex circuitry.

An ant-inspired celestial compass applied to autonomous outdoor robot navigation

Desert ants use the polarization of skylight and a combination of stride and ventral optic flow integration processes to track the nest and food positions when traveling, achieving outstanding performances. Navigation sensors such as global positioning systems and inertial measurement units still have disadvantages such as their low resolution and drift. Taking our inspiration from ants, we developed a 2-pixel celestial compass which computes the heading angle of a mobile robot in the ultraviolet range. The output signals obtained with this optical compass were investigated under various weather and ultraviolet conditions and compared with those obtained on a magnetometer in the vicinity of our laboratory. After being embedded on-board the robot, the sensor was first used to compensate for random yaw disturbances. We then used the compass to keep the Hexabot robot’s heading angle constant in a straight forward walking task over a flat terrain while its walking movements were imposing yaw disturbances. Experiments performed under various meteorological conditions showed the occurrence of steady state heading angle errors ranging from 0.3∘ (with a clear sky) to 2.9∘ (under changeable sky conditions). The compass was also tested under canopies and showed a strong ability to determine the robot’s heading while most of the sky was hidden by the foliage. Lastly, a waterproof, mono-pixel version of the sensor was designed and successfully tested in a preliminary underwater benchmark test. These results suggest this new optical compass shows great precision and reliability in a wide range of outdoor conditions, which makes it highly suitable for autonomous robotic outdoor navigation tasks. A celestial compass and a minimalistic optic flow sensor called M2APix (based on Michaelis–Menten Auto-adaptive Pixels) were therefore embedded on-board our latest insectoid robot called AntBot, to complete the previously mentioned ant-like homing navigation processes. First the robot was displaced manually and made to return to its starting-point on the basis of its absolute knowledge of the coordinates of this point. Lastly, AntBot was tested in fully autonomous navigation experiments, in which it explored its environment and then returned to base using the same sensory modes as those on which desert ants rely. AntBot produced robust, precise localization performances with a homing error as small as 0.7% of the entire trajectory.

Two Compasses in the Central Complex of the Locust Brain.

Many migratory insects rely on a celestial compass for spatial orientation. Several features of the daytime sky, all generated by the sun, can be exploited for navigation. Two of these are the position of the sun and the pattern of polarized skylight. Neurons of the central complex (CX), a group of neuropils in the central brain of insects, have been shown to encode sky compass cues. In desert locusts, the CX holds a topographic, compass-like representation of the plane of polarized light (E-vector) presented from dorsal direction. In addition, these neurons also encode the azimuth of an unpolarized light spot, likely representing the sun. Here, we investigate whether, in addition to E-vector orientation, the solar azimuth is represented topographically in the CX. We recorded intracellularly from eight types of CX neuron while stimulating animals of either sex with polarized blue light from zenithal direction and an unpolarized green light spot rotating around the animal's head at different elevations. CX neurons did not code for elevation of the unpolarized light spot. However, two types of columnar neuron showed a linear correlation between innervated slice in the CX and azimuth tuning to the unpolarized green light spot, consistent with an internal compass representation of solar azimuth. Columnar outputs of the CX also showed a topographic representation of zenithal E-vector orientation, but the two compasses were not linked to each other. Combined stimulation with unpolarized green and polarized blue light suggested that the two compasses interact in a nonlinear way.SIGNIFICANCE STATEMENT In the brain of the desert locust, neurons sensitive to the plane of celestial polarization are arranged like a compass in the slices of the central complex (CX). These neurons, in addition, code for the horizontal direction of an unpolarized light cue possibly representing the sun. We show here that horizontal directions are, in addition to E-vector orientations from the dorsal direction, represented in a compass-like manner across the slices of the CX. However, the two compasses are not linked to each other, but rather seem to interact in a cell-specific, nonlinear way. Our study confirms the role of the CX in signaling heading directions and shows that different cues are used for this task.

The brain behind straight-line orientation in dung beetles.

For many insects, celestial compass cues play an important role in keeping track of their directional headings. One well-investigated group of celestial orientating insects are the African ball-rolling dung beetles. After finding a dung pile, these insects detach a piece, form it into a ball and roll it away along a straight path while facing backwards. A brain region, termed the central complex, acts as an internal compass that constantly updates the ball-rolling dung beetle about its heading. In this review, we give insights into the compass network behind straight-line orientation in dung beetles and place it in the context of the orientation mechanisms and neural networks of other insects. We find that the neuronal network behind straight-line orientation in dung beetles has strong similarities to the ones described in path-integrating and migrating insects, with the central complex being the key control point for this behavior. We conclude that, despite substantial differences in behavior and navigational challenges, dung beetles encode compass information in a similar way to other insects.

Polarized skylight-based heading measurements: a bio-inspired approach.

Many insects such as desert ants, crickets, locusts, dung beetles, bees and monarch butterflies have been found to extract their navigation cues from the regular pattern of the linearly polarized skylight. These species are equipped with ommatidia in the dorsal rim area of their compound eyes, which are sensitive to the angle of polarization of the skylight. In the polarization-based robotic vision, most of the sensors used so far comprise high-definition CCD or CMOS cameras topped with linear polarizers. Here, we present a 2-pixel polarization-sensitive visual sensor, which was strongly inspired by the dorsal rim area of desert ants' compound eyes, designed to determine the direction of polarization of the skylight. The spectral sensitivity of this minimalistic sensor, which requires no lenses, is in the ultraviolet range. Five different methods of computing the direction of polarization were implemented and tested here. Our own methods, the extended and AntBot method, outperformed the other three, giving a mean angular error of only 0.62° ± 0.40° (median: 0.24°) and 0.69° ± 0.52° (median: 0.39°), respectively (mean ± standard deviation). The results obtained in outdoor field studies show that our celestial compass gives excellent results at a very low computational cost, which makes it highly suitable for autonomous outdoor navigation purposes.

Neuroarchitecture of the dung beetle central complex.

Despite their tiny brains, insects show impressive abilities when navigating over short distances during path integration or during migration over thousands of kilometers across entire continents. Celestial compass cues often play an important role as references during navigation. In contrast to many other insects, South African dung beetles rely exclusively on celestial cues for visual reference during orientation. After finding a dung pile, these animals cut off a piece of dung from the pat, shape it into a ball and roll it away along a straight path until a suitable place for underground consumption is found. To maintain a constant bearing, a brain region in the beetle's brain, called the central complex, is crucially involved in the processing of skylight cues, similar to what has already been shown for path-integrating and migrating insects. In this study, we characterized the neuroanatomy of the sky-compass network and the central complex in the dung beetle brain in detail. Using tracer injections, combined with imaging and 3D modeling, we describe the anatomy of the possible sky-compass network in the central brain. We used a quantitative approach to study the central-complex network and found that several types of neuron exhibit a highly organized connectivity pattern. The architecture of the sky-compass network and central complex is similar to that described in insects that perform path integration or are migratory. This suggests that, despite their different orientation behaviors, this neural circuitry for compass orientation is highly conserved among the insects.

The Dung Beetle Compass

What do a burly rower, a backstroke swimmer and a hard-working South African dung beetle all have in common? The answer is: they all benefit from moving along a straight path, and do so moving backwards. This, however, is where the similarity ends. While the rower has solved this navigational challenge by handing the task of steering to the coxswain, who faces the direction of travel, and the swimmer is guided down her lane by colourful ropes, the beetle puts its faith in the sky. From here, it utilises a larger repertoire of celestial compass cues than is known to be used by any other animal studied to date. A robust internal compass, designed to interpret directional information, has evolved under the selective pressure of shifting today's lunch efficiently out of reach of competitors, also drawn to the common buffet. While this is a goal that beetles might share with the hungry athletes, they reach it with drastically different brain powers; the brain of the beetle is several times smaller than a match head, containing fewer than a million neurons.

Does size affect orientation using celestial cues?

Insects are well known to orient using celestial cues. The pattern of polarised skylight is the dominant celestial compass information that insects use, which they detect using a specialised set of ommatidia. The number of ommatidia decreases with body size, and it is unknown how this reduction in the number of ommatidia affects the precision of orienting using celestial cues. We investigated this in eight different ant species that had varying numbers of ommatidia. We captured ants returning home, displaced them to an unfamiliar location and measured their precision in determining heading direction using celestial cues. The heading direction of the ants measured at a fixed distance from the release and also at a distance scaled to their body size was not correlated with the number of ommatidia. However, both the path straightness and walking speed were lower in smaller ants indicating the ability to orient at a finer scale was affected by miniaturisation.