Straight-line Orientation Research Articles

Correction: The effect of step size on straight-line orientation.

Correction: The effect of step size on straight-line orientation.

Drosophila require both green and UV wavelengths for sun orientation but lack a time-compensated sun compass.

Celestial orientation and navigation are performed by many organisms in contexts as diverse as migration, nest finding and straight-line orientation. The vinegar fly, Drosophila melanogaster, performs menotaxis in response to celestial cues during tethered flight and can disperse more than 10 km under field conditions. However, we still do not understand how spectral components of celestial cues and pauses in flight impact heading direction in flies. To assess individual heading, we began by testing flies in a rotating tether arena using a single green LED as a stimulus. We found that flies robustly perform menotaxis and fly straight for at least 20 min. Flies maintain their preferred heading directions after experiencing a period of darkness or stopping flight, even up to 2 h, but reset their heading when the LED changes position, suggesting that flies do not treat this stimulus as the sun. Next, we assessed the flies' responses to a UV spot alone or a paired UV-green stimulus - two dots situated 180deg apart to simulate the solar and antisolar hemispheres. We found that flies respond to UV much as they do to green light; however, when the stimuli are paired, flies adjust for sudden 90deg movements, performing sun orientation. Lastly, we found no evidence of a time-compensated sun compass when we moved the paired stimuli at 15degh-1 for 6 h. This study demonstrates that wavelength influences how flies respond to visual cues during flight, shaping the interpretation of visual information to execute an appropriate behavioral response.

A model of cue integration as vector summation in the insect brain.

Ball-rolling dung beetles are known to integrate multiple cues in order to facilitate their straight-line orientation behaviour. Recent work has suggested that orientation cues are integrated according to a vector sum, that is, compass cues are represented by vectors and summed to give a combined orientation estimate. Further, cue weight (vector magnitude) appears to be set according to cue reliability. This is consistent with the popular Bayesian view of cue integration: cues are integrated to reduce or minimize an agent's uncertainty about the external world. Integration of orientation cues is believed to occur at the input to the insect central complex. Here, we demonstrate that a model of the head direction circuit of the central complex, including plasticity in input synapses, can act as a substrate for cue integration as vector summation. Further, we show that cue influence is not necessarily driven by cue reliability. Finally, we present a dung beetle behavioural experiment which, in combination with simulation, strongly suggests that these beetles do not weight cues according to reliability. We suggest an alternative strategy whereby cues are weighted according to relative contrast, which can also explain previous results.

Weighted cue integration for straight-line orientation

SummaryAnimals commonly integrate multiple sources of information to guide their behavior. Among insects, previous studies have suggested that the relative reliability of cues affects their weighting in behavior, but have not systematically explored how well alternative integration strategies can account for the observed directional choices. Here, we characterize the directional reliability of an ersatz sun at different elevations and wind at different speeds as guiding cues for a species of ball-rolling dung beetle. The relative reliability is then shown to determine which cue dominates when the cues are put in conflict. We further show through modeling that the results are best explained by continuous integration of the cues as a vector-sum (rather than switching between them) but with non-optimal weighting and small individual biases. The neural circuitry in the insect central complex appears to provide an ideal substrate for this type of vector-sum-based integration mechanism.

Compass Cue Integration and Its Relation to the Visual Ecology of Three Tribes of Ball-Rolling Dung Beetles.

Simple SummaryTo escape competition for food at the dung pat, ball-rolling dung beetles shape a piece of dung into a sphere and roll it away. To maintain their bearing, these beetles integrate directional information from a range of celestial cues. For the majority of diurnal dung beetles living in open habitats, the most dominant of these cues is the sun. It has recently been demonstrated that beetles living in closed habitats, with closely spaced trees and tall grass, rely predominantly on directional information provided by polarised skylight rather than the sun. Taken together, these findings suggests that the orientation strategy of the beetle is influenced by the animal’s visual ecology. To further investigate the relative weighting of cues in the orientation system of beetles, and its relation to their visual ecology, we investigated the orientation strategy of ball-rollers from three different dung beetle tribes, all present within the same savanna biome. We find that species within a tribe share the same orientation strategy, but that this strategy differs across tribes. We conclude that, despite dramatic intertribal differences in body size and external eye design, the dynamic heading direction network of the South African ball-rolling dung beetles is well adapted to guide the foraging insect in the habitat that it normally traverses.To guide their characteristic straight-line orientation away from the dung pile, ball-rolling dung beetles steer according to directional information provided by celestial cues, which, among the most relevant are the sun and polarised skylight. Most studies regarding the use of celestial cues and their influence on the orientation system of the diurnal ball-rolling beetle have been performed on beetles of the tribe Scarabaeini living in open habitats. These beetles steer primarily according to the directional information provided by the sun. In contrast, Sisyphus fasciculatus, a species from a different dung-beetle tribe (the Sisyphini) that lives in habitats with closely spaced trees and tall grass, relies predominantly on directional information from the celestial pattern of polarised light. To investigate the influence of visual ecology on the relative weight of these cues, we studied the orientation strategy of three different tribes of dung beetles (Scarabaeini, Sisyphini and Gymnopleurini) living within the same biome, but in different habitat types. We found that species within a tribe share the same orientation strategy, but that this strategy differs across the tribes; Scarabaeini, living in open habitats, attribute the greatest relative weight to the directional information from the sun; Sisyphini, living in closed habitats, mainly relies on directional information from polarised skylight; and Gymnopleurini, also living in open habitats, appear to weight both cues equally. We conclude that, despite exhibiting different body size, eye size and morphology, dung beetles nevertheless manage to solve the challenge of straight-line orientation by weighting visual cues that are particular to the habitat in which they are found. This system is however dynamic, allowing them to operate equally well even in the absence of the cue given the greatest relative weight by the particular species.

The effect of step size on straight-line orientation.

Moving along a straight path is a surprisingly difficult task. This is because, with each ensuing step, noise is generated in the motor and sensory systems, causing the animal to deviate from its intended route. When relying solely on internal sensory information to correct for this noise, the directional error generated with each stride accumulates, ultimately leading to a curved path. In contrast, external compass cues effectively allow the animal to correct for errors in its bearing. Here, we studied straight-line orientation in two different sized dung beetles. This allowed us to characterize and model the size of the directional error generated with each step, in the absence of external visual compass cues (motor error) as well as in the presence of these cues (compass and motor errors). In addition, we model how dung beetles balance the influence of internal and external orientation cues as they orient along straight paths under the open sky. We conclude that the directional error that unavoidably accumulates as the beetle travels is inversely proportional to the step size of the insect, and that both beetle species weigh the two sources of directional information in a similar fashion.

Straight-line orientation in the woodland-living beetle Sisyphus fasciculatus

To transport their balls of dung along a constant bearing, diurnal savannah-living dung beetles rely primarily on the sun for compass information. However, in more cluttered environments, such as woodlands, this solitary compass cue is frequently hidden from view by surrounding vegetation. In these types of habitats, insects can, instead, rely on surrounding landmarks, the canopy pattern, or wide-field celestial cues, such as polarised skylight, for directional information. Here, we investigate the compass orientation strategy behind straight-line orientation in the diurnal woodland-living beetle Sisyphus fasciculatus. We found that, when manipulating the direction of polarised skylight, Si. fasciculatus responded to this change with a similar change in bearing. However, when the apparent position of the sun was moved, the woodland-living beetle did not change its direction of travel. In contrast, the savannah-living beetle Scarabaeus lamarcki responded to the manipulation of the solar position with a corresponding change in bearing. These results suggest that the dominant compass cue used for straight-line orientation in dung beetles may be determined by the celestial cue that is most prominent in their preferred habitat.

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.

H2O masers and protoplanetary disk dynamics in IC 1396 N

We report H$_2$O maser line observations of the bright-rimmed globule IC 1396 N using a ground-space interferometer with the 10-m RadioAstron radio telescope as the space-based element. The source was not detected on projected baselines >2.3 Earth diameters, which indicates a lower limit on the maser size of L >0.03 AU and an upper limit on the brightness temperature of 6.25 x 10$^{12}$ K. Positions and flux densities of maser spots were determined by fringe rate mapping. Multiple low-velocity features from -4.5 km/s to +0.7 km/s are seen, and two high-velocity features of V$_{LSR}$=-9.4 km/s and +4.4 km/s are found at projected distances of 157 AU and 70 AU, respectively, from the strongest low-velocity feature at V$_{LSR}$=$\sim$0.3 km/s. Maser components from the central part of the spectrum fall into four velocity groups but into three spatial groups. Three spatial groups of low-velocity features detected in the 2014 observations are arranged in a linear structure about 200 AU in length. Two of these groups were not detected in 1996 and possibly are jets which formed between 1996 and 2014. The putative jet seems to have changed direction in 18 years, which we explain by the precession of the jet under the influence of the gravity of material surrounding the globule. The jet collimation can be provided by a circumstellar protoplanetary disk. There is a straight line orientation in the V$_{LSR}$-Right Ascension diagram between the jet and the maser group at V$_{LSR}$=$\sim$0.3 km/s. However, the central group with the same position but at the velocity V$_{LSR}$$\sim$-3.4 km/s falls on a straight line between two high-velocity components detected in 2014. Comparison of the low-velocity positions from 2014 and 1996, based on the same diagram, shows that the majority of the masers maintain their positions near the central velocity V$_{LSR}$=$\sim$0.3 km/s during the 18 year period.

A Snapshot-Based Mechanism for Celestial Orientation

In order to protect their food from competitors, ball-rolling dung beetles detach a piece of dung from a pile, shape it into a ball, and roll it away along a straight path [1]. They appear to rely exclusively on celestial compass cues to maintain their bearing [2-8], but the mechanism that enables them to use these cues for orientation remains unknown. Here, we describe the orientation strategy that allows dung beetles to use celestial cues in a dynamic fashion. We tested the underlying orientation mechanism by presenting beetles with a combination of simulated celestial cues (sun, polarized light, and spectral cues). We show that these animals do not rely on an innate prediction of the natural geographical relationship between celestial cues, as other navigating insects seem to [9, 10]. Instead, they appear to form an internal representation of the prevailing celestial scene, a "celestial snapshot," even if that scene represents a physical impossibility for the real sky. We also find that the beetles are able to maintain their bearing with respect to the presented cues only if the cues are visible when the snapshotis taken. This happens during the "dance," a behavior in which the beetle climbs on top of its ball and rotates about its vertical axis [11]. This strategy for reading celestial signals is a simple but efficient mechanism for straight-line orientation.

The role of the sun in the celestial compass of dung beetles

Recent research has focused on the different types of compass cues available to ball-rolling beetles for orientation, but little is known about the relative precision of each of these cues and how they interact. In this study, we find that the absolute orientation error of the celestial compass of the day-active dung beetle Scarabaeus lamarcki doubles from 16° at solar elevations below 60° to an error of 29° at solar elevations above 75°. As ball-rolling dung beetles rely solely on celestial compass cues for their orientation, these insects experience a large decrease in orientation precision towards the middle of the day. We also find that in the compass system of dung beetles, the solar cues and the skylight cues are used together and share the control of orientation behaviour. Finally, we demonstrate that the relative influence of the azimuthal position of the sun for straight-line orientation decreases as the sun draws closer to the horizon. In conclusion, ball-rolling dung beetles possess a dynamic celestial compass system in which the orientation precision and the relative influence of the solar compass cues change over the course of the day.

Dung Beetles Use the Milky Way for Orientation

When the moon is absent from the night sky, stars remain as celestial visual cues. Nonetheless, only birds, seals, and humans are known to use stars for orientation. African ball-rolling dung beetles exploit the sun, the moon, and the celestial polarization pattern to move along straight paths, away from the intense competition at the dung pile. Even on clear moonless nights, many beetles still manage to orientate along straight paths. This led us to hypothesize that dung beetles exploit the starry sky for orientation, a feat that has, to our knowledge, never been demonstrated in an insect. Here, we show that dung beetles transport their dung balls along straight paths under a starlit sky but lose this ability under overcast conditions. In a planetarium, the beetles orientate equally well when rolling under a full starlit sky as when only the Milky Way is present. The use of this bidirectional celestial cue for orientation has been proposed for vertebrates, spiders, and insects, but never proven. This finding represents the first convincing demonstration for the use of the starry sky for orientation in insects and provides the first documented use of the Milky Way for orientation in the animal kingdom.

Dung beetles ignore landmarks for straight-line orientation

Upon locating a suitable dung pile, ball-rolling dung beetles shape a piece of dung into a ball and roll it away in a straight line. This guarantees that they will not return to the dung pile, where they risk having their ball stolen by other beetles. Dung beetles are known to use celestial compass cues such as the sun, the moon and the pattern of polarised light formed around these light sources to roll their balls of dung along straight paths. Here, we investigate whether terrestrial landmarks have any influence on straight-line orientation in dung beetles. We find that the removal or re-arrangement of landmarks has no effect on the beetle's orientation precision. Celestial compass cues dominate straight-line orientation in dung beetles so strongly that, under heavily overcast conditions or when prevented from seeing the sky, the beetles can no longer orient along straight paths. To our knowledge, this is the only animal with a visual compass system that ignores the extra orientation precision that landmarks can offer.

The Dung Beetle Dance: An Orientation Behaviour?

An interesting feature of dung beetle behaviour is that once they have formed a piece of dung into a ball, they roll it along a straight path away from the dung pile. This straight-line orientation ensures that the beetles depart along the most direct route, guaranteeing that they will not return to the intense competition (from other beetles) that occurs near the dung pile. Before rolling a new ball away from the dung pile, dung beetles perform a characteristic “dance,” in which they climb on top of the ball and rotate about their vertical axis. This dance behaviour can also be observed during the beetles' straight-line departure from the dung pile. The aim of the present study is to investigate the purpose of the dung beetle dance. To do this, we explored the circumstances that elicit dance behaviour in the diurnal ball-rolling dung beetle, Scarabaeus (Kheper) nigroaeneus. Our results reveal that dances are elicited when the beetles lose control of their ball or lose contact with it altogether. We also find that dances can be elicited by both active and passive deviations of course and by changes in visual cues alone. In light of these results, we hypothesise that the dung beetle dance is a visually mediated mechanism that facilitates straight-line orientation in ball-rolling dung beetles by allowing them to 1) establish a roll bearing and 2) return to this chosen bearing after experiencing a disturbance to the roll path.

Line detection in images through regularized hough transform

The problem of determining the location and orientation of straight lines in images is of great importance in the fields of computer vision and image processing. Traditionally the Hough transform, (a special case of the Radon transform) has been widely used to solve this problem for binary images. In this paper, we pose the problem of detecting straight lines in gray-scale images as an inverse problem. Our formulation is based on use of the inverse Radon operator, which relates the parameters determining the location and orientation of the lines in the image to the noisy input image. The advantage of this formulation is that we can then approach the problem of line detection within a regularization framework and enhance the performance of the Hough-based line detector through the incorporation of prior information in the form of regularization. We discuss the type of regularizers that are useful for this problem and derive efficient computational schemes to solve the resulting optimization problems enabling their use in large applications. Finally, we show how our new approach can be alternatively viewed as one of finding an optimal representation of the noisy image in terms of elements chosen from a dictionary of lines. This interpretation relates the problem of Hough-based line finding to the body of work on adaptive signal representation.

Visual cues used by ball-rolling dung beetles for orientation.

Ball rolling by dung beetles is considered to be a derived behaviour that evolved under pressure for space, and from competitors at the dung pat. Straight-line orientation away from the pat using a celestial cue should be the most successful rolling strategy to move dung to an unknown burial site. We tested this hypothesis in the field and the laboratory by presenting five species of ball-rolling beetles with different orientation tasks, involving reaction to obstacles as well as to reflected sunlight and artificial light sources. Beetles were found to consistently orientate along a chosen route, usually in the direction of the sun. Beetles rolling dung balls successfully negotiated barriers and returned to the original path as did beetles falling from ramps, or rotated about a fixed point while rolling a ball. The sun was found to be the main orientation cue, which could be substituted by reflected or artificial light. However, beetles reoriented themselves less accurately in response to lights in the laboratory, than they did to the reflected sun in the field. It is probable that phototactic orientation using the sun, which is widespread amongst arthropods, has been incorporated in the straight-line foraging behaviour that has evolved in ball-rolling dung beetles.