Field vole (Microtus agrestis) seasonal spacing behavior: the effect of predation risk by mustelids

There are numerous studies showing that predation risk may change different aspects of the behavior of prey, such as habitat use, activity pattern, and foraging. Prey should exhibit the strongest antipredatory response against their most deadly predator. Small mustelids are considered the most important mammalian predators of voles. Nevertheless, there is no general agreement as to whether strong antipredatory reactions exist in natural free-living populations of voles. Here, we studied the field vole Microtus agrestis spatial reaction to high predation risk from small mustelids in the breeding (August) and nonbreeding (October) seasons under natural conditions. Voles were exposed to a caged weasel (Mustela nivalis) and a stoat (Mustela erminea), as well as to the odors of these predators. The reactions of 30 field voles were monitored with radiotelemetry. The field voles were found to display antipredator reactions that varied with season. In the breeding period, in response to predation risk, voles reduced locomotory activity and daily-range size, whereas in the nonbreeding period they did not. Changes in home range position were similar for control and treatment voles, in both the breeding and nonbreeding periods. The results indicate that mustelid predators modify the spatial behavior of small rodents in natural conditions depending on season. This might be a reflection of differences in state-dependent responses to predation from sexually active or inactive individuals. This suggests that the basic antipredatory reaction of voles under high predation risk from small mustelids limits their locomotory activity.     

Avian magnetic compass: fast adjustment to intensities outside the normal functional window

To determine how fast birds can adapt to magnetic intensities outside the normal functional window of their magnetic compass, we tested migratory birds in a magnetic field of 92,000 nT, twice the intensity of the local geomagnetic field at the test site in Frankfurt a.M., Germany. In the local field, robins showed a significant preference of their southerly migratory direction, whereas in the 92,000-nT field, they were initially disoriented. However, when the birds were preexposed to 92,000 nT for 1 h before being tested, they were able to orient under this intensity, and their behavior did not differ from that in the geomagnetic field. These data show that birds require only a short time to adjust to magnetic intensities, which they cannot spontaneously use for orientation. Interpreting these findings in view of the radical pair model (Ritz et al. 2000), this means that they can learn rather quickly to interpret novel activation patterns on their retina.     

Light-dependent magnetic compass in Iberian green frog tadpoles

Here, we provide evidence for a wavelength-dependent effect of light on magnetic compass orientation in Pelophylax perezi (order Anura), similar to that observed in Rana catesbeiana (order Anura) and Notophthalmus viridescens (order Urodela), and confirm for the first time in an anuran amphibian that a 90° shift in the direction of magnetic compass orientation under long-wavelength light (≥500 nm) is due to a direct effect of light on the underlying magnetoreception mechanism. Although magnetic compass orientation in other animals (e.g., birds and some insects) has been shown to be influenced by the wavelength and/or intensity of light, these two amphibian orders are the only taxa for which there is direct evidence that the magnetic compass is light-dependent. The remarkable similarities in the light-dependent magnetic compasses of anurans and urodeles, which have evolved as separate clades for at least 250 million years, suggest that the light-dependent magnetoreception mechanism is likely to have evolved in the common ancestor of the Lissamphibia (Early Permian, ~294 million years) and, possibly, much earlier. Also, we discuss a number of similarities between the functional properties of the light-dependent magnetic compass in amphibians and blue light-dependent responses to magnetic stimuli in Drosophila melanogaster, which suggest that the wavelength-dependent 90° shift in amphibians may be due to light activation of different redox forms of a cryptochrome photopigment. Finally, we relate these findings to earlier studies showing that the pineal organ of newts is the site of the light-dependent magnetic compass and recent neurophysiological evidence showing magnetic field sensitivity in the frog frontal organ (an outgrowth of the pineal).     

Abundance thresholds and the underlying ecological processes: Field voles Microtus agrestis in a fragmented landscape

Field voles (Microtus agrestis) were trapped in 14 field margins and their behavioural and demographic parameters measured. Strong support was found for thresholds in margin width below which vole abundance was extremely low. Narrow margins were male biased with individuals moving greater distances and a large proportion of males behaved as transient individuals. However, no effect was observed on the age structure or survival of the population. Individuals were able to compensate for the lack of habitat through alterations in their behaviour sufficiently to maintain their survival. Within intensive agro-ecosystems, narrow strips between crops are important links for voles between wider margins and, if available, other more suitable habitats. Maintenance of narrow margins, along with larger areas of suitable habitat, is therefore effective in farmed landscapes for sustaining populations of specialist species where they show sufficient flexibility.     

Navigation mit Duftkarte und Sonnenkompaß: Das Heimfindevermögen der Brieftauben

Pigeon homing, investigated as a paradigmatic example of bird navigation, appears to be based on two mechanisms of orientation whose functions correspond to those of map and compass. Tasks of the latter are usually accomplished by a sun compass, taking into account the sun's movement and time of day. Under overcast skies, the magnetic field of the earth may be used for compass orientation. The "map" part of the system, responsible for site localization, makes use of olfactory perception of atmospheric trace compounds, which must be concluded to contain positional information in unfamiliar areas up to several hundreds of kilometers from home.     

Magnetic compass orientation of European robins under 565 nm green light

European robins tested under monochromatic green light with a peak wavelength of 565 nm at an intensity of 2.1 mW m^-2 in the local geomagnetic field preferred their migratory direction, heading southward in autumn and northward in spring. Inverting of the vertical component of the magnetic field caused the robins to reverse their headings, indicating that the birds used a magnetic inclination compass to locate their migratory direction. The behavior recorded under green light at an intensity of 2.1 mW m^-2 is thus not different from that previously recorded under "white" light; it represents normal migratory orientation.     

Attempts to train goldfish to respond to magnetic field stimuli

Two different conditioning procedures, one appetitive and the other aversive, were used in separate attempts to demonstrate response to magnetic fields in the goldfish, Carassius auratus. Our results lead us to question those of an orientation experiment by Becker, although we recognize the possibility that goldfish may be sensitive primarily to magnetic field direction rather than intensity and that their directional sensitivity may be evidenced most readily by orientation in the field.     

Habitat quality of wildflower strips for common voles (Microtus arvalis) and its relevance for agriculture

Ecological compensation areas have been widely promoted in agriculture in the last decade. Besides their positive effects on biological diversity they also bear a risk of sheltering potential pest species such as common voles (Microtus arvalis). To assess the influence of wildflower strips on the spatio-temporal behavior of voles and their impact on adjacent crop fields, a common vole population living in a wildflower strip was monitored from May to October 2000 and from March to September 2001. A new system for automatic radio tracking was used which allowed tracking at 1 min intervals and up to 1000 bearings per vole per day. Voles showed small home ranges with a median size of 125 m² (minimum convex polygon) and 30 m² (Kernel), respectively. Home ranges were stable with a median overlap of 90% for consecutive days, were almost exclusively within the wildflower strip and contained several core areas per range. A polyphasic activity pattern with a phase length of 1.7 h was found during summer with a trend towards diurnal activity. Overall wildflower strips were high-quality habitats for voles and sustained high population densities without increased risk of voles dispersing into adjacent fields.     

Magnetoreception in birds: no intensity window in “fixed direction” responses

Under 502 nm turquoise light combined with 590 nm yellow light and in total darkness, European robins, Erithacus rubecula, no longer prefer their migratory direction, but exhibit so-called fixed direction responses that do not show the seasonal change between spring and autumn. We tested robins under these light conditions in the local geomagnetic field of 46 μT, a field of twice this intensity, 92 μT, and a field of three times this intensity, 138 μT. Under all three magnetic conditions, the birds preferred the same easterly direction under turquoise-and-yellow light and the same northwesterly direction under dark, while they were oriented in their seasonally appropriate direction under control conditions. “Fixed direction” responses are thus not limited to a narrow intensity window as has been found for normal compass orientation. This can be attributed to their origin in the magnetite-based receptor in the upper beak, which operates according to fundamentally different principles than the radical pair mechanism in the retina mediating compass orientation. “Fixed direction” responses are possibly a relict of a receptor mechanism that changed its function, now mainly providing information on magnetic intensity.     

<Emphasis Type="Italic">Tenebrio</Emphasis> beetles use magnetic inclination compass

Animals that guide directions of their locomotion or their migration routes by the lines of the geomagnetic field use either polarity or inclination compasses to determine the field polarity (the north or south direction). Distinguishing the two compass types is a guideline for estimation of the molecular principle of reception and has been achieved for a number of animal groups, with the exception of insects. A standard diagnostic method to distinguish a compass type is based on reversing the vertical component of the geomagnetic field, which leads to the opposite reactions of animals with two different compass types. In the present study, adults of the mealworm beetle Tenebrio molitor were tested by means of a two-step laboratory test of magnetoreception. Beetles that were initially trained to memorize the magnetic position of the light source preferred, during the subsequent test, this same direction, pursuant geomagnetic cues only. In the following step, the vertical component was reversed between the training and the test. The beetles significantly turned their preferred direction by 180 degrees. Our results brought until then unknown original findings that insects, represented here by the T. molitor species, use-in contrast to another previously researched Arthropod, spiny lobster-the inclination compass.     

Light-dependent magnetoreception in birds: interaction of at least two different receptors

Passerine migrants require light from the blue-green part of the spectrum for magnetic compass orientation; under yellow light, they are disoriented. European robins tested under a combination of yellow light and blue or green light showed a change in behavior, no longer preferring their seasonally appropriate migratory direction: in spring as well as in autumn, they preferred southerly headings under blue-and-yellow and northerly headings under green-and-yellow light. This clearly shows that yellow light is not neutral and suggests the involvement of at least two types of receptors in obtaining magnetic compass information, with the specific interaction of these receptors being rather complex.     

Magnetic compass orientation of migratory birds in the presence of a 1.315 MHz oscillating field

The radical pair model of magnetoreception predicts that magnetic compass orientation can be disrupted by high frequency magnetic fields in the Megahertz range. European robins, Erithacus rubecula, were tested under monochromatic 565 nm green light in 1.315 MHz fields of 0.48 T during spring and autumn migration, with 1.315 MHz being the frequency that matches the energetic splitting induced by the local geomagnetic field. The birds responses depended on the alignment of the oscillating field with respect to the static geomagnetic field: when the 1.315 MHz field was aligned parallel with the field lines, birds significantly preferred northerly directions in spring and southerly directions in autumn. These preferences reflect normal migratory orientation, with the variance slightly increased compared to control tests in the geomagnetic field alone or to tests in a 7.0 MHz field. However, in the 1.315 MHz field aligned at a 24° angle to the field lines, the birds were disoriented in both seasons, indicating that the high frequency field interfered with magnetoreception. These finding are in agreement with theoretical predictions and support the assumption of a radical-pair mechanism underlying the processes mediating magnetic compass information in birds.     

Loft features reveal the functioning of the young pigeon’s navigational system

It is thought that young homing pigeons are able to use information acquired en route for their initial homeward orientation. However, the cues involved and mechanisms utilised are under discussion. Blocking light-dependent route-specific information during the first leg of an outward journey detour, together with analysis of pigeons that were raised under different loft conditions, allowed us to correctly evaluate the functioning of this mechanism and, more generally, the navigational map of birds. Pigeons from the same stock were raised and kept in two different lofts. The birds in the experimental groups were transported to the release sites via detours, and light-dependent information was denied during the first half of the outward journey (no compass information was available). Control birds were transported by the most direct route and had access to all available information. In general, the results showed that the low-loft birds preferred to use magnetic compass cues, whereas the high-loft birds preferred to use navigational map cues to collect information of the first part of the outward journey. The impairments observed in the homing performances of the experimental groups highlight the reliability of information collected inside the map area. Relevant to an understanding of the route-reversal mechanism was the evidence that this mechanism is able to function in the absence of compass information (birds raised in a wind-exposed loft show a detour effect). In systems where directional information could be provided by multiple sources, processing and extracting accurate course trajectories through a common mechanism may prove more efficient and reliable.     

The sea-finding behavior of hatchling olive ridley sea turtles, Lepidochelys olivacea, at the beach of San Miguel (Costa Rica)

Newly hatched olive ridley sea turtles (Lepidochelys olivacea) were tested for their directional preferences in a sand-filled circular arena in total darkness. Hatchlings that had crawled about 5 m on the beach, toward the sea preferred the southwesterly direction that would have brought them to the water line, whereas hatchlings that had been denied this experience headed eastward, a direction of unclear origin. These data suggest that a short crawl across the natural beach can set the direction in which the young turtles subsequently move. The crawling experience was sufficient to acquire the compass course that they later follow, probably with the help of a magnetic compass, not only in the water, but already while still on land.     

Retinal cryptochrome in a migratory passerine bird: a possible transducer for the avian magnetic compass

The currently discussed model of magnetoreception in birds proposes that the direction of the magnetic field is perceived by radical-pair processes in specialized photoreceptors, with cryptochromes suggested as potential candidate molecules mediating magnetic compass information. Behavioral studies have shown that magnetic compass orientation takes place in the eye and requires light from the blue-green part of the spectrum. Cryptochromes are known to absorb in the same spectral range. Because of this we searched for cryptochrome (CRY) in the retina of European robins, Erithacus rubecula, passerine birds that migrate at night. Here, we report three individually expressed cryptochromes, eCRY1a, eCRY1b, and eCRY2. While eCRY1a and eCRY2 are similar to the cryptochromes found in the retina of the domestic chicken, eCRY1b has a unique carboxy (C)-terminal. In light of the radical-pair model, our findings support a potential role of cryptochromes as transducers for the perception of magnetic compass information in birds.     

Magnetic compass orientation in birds and its physiological basis

A current model suggests that magnetoreception of compass information starts with light-dependent primary processes. Light-dependency of magnetoreception is supported by behavioral experiments with homing pigeons and caged migratory birds. Three passerine species showed normal orientation under dim monochromatic light from the blue-green range of the spectrum, while they were disoriented under yellow and red light. A sevenfold increase in intensity and pre-exposure to specific wavelengths caused changes in behavior. The behavioral data indicate a complex relationship between the wavelength of light and magnetoreception, suggesting the involvement of more than one type of receptors. Extracellular recordings from the nucleus of the basal optic root and the tectum opticum identified units that responded to changes in magnetic North. Each unit showed a peak in a distinct spatial direction, so that the input of these units, processed collectively and integrated, would indicate compass directions.     

Vector navigation in desert ants, Cataglyphis fortis: celestial compass cues are essential for the proper use of distance information

Foraging desert ants navigate primarily by path integration. They continually update homing direction and distance by employing a celestial compass and an odometer. Here we address the question of whether information about travel distance is correctly used in the absence of directional information. By using linear channels that were partly covered to exclude celestial compass cues, we were able to test the distance component of the path-integration process while suppressing the directional information. Our results suggest that the path integrator cannot process the distance information accumulated by the odometer while ants are deprived of celestial compass information. Hence, during path integration directional cues are a prerequisite for the proper use of travel-distance information by ants.     

Fractionating dead reckoning: role of the compass,odometer, logbook,and home base establishment in spatial orientation

Rats use multiple sources of information to maintain spatial orientation. Although previous work has focused on rats’ use of environmental cues, a growing number of studies have demonstrated that rats also use self-movement cues to organize navigation. This review examines the extent that kinematic analysis of naturally occurring behavior has provided insight into processes that mediate dead-reckoning-based navigation. This work supports a role for separate systems in processing self-movement cues that converge on the hippocampus. The compass system is involved in deriving directional information from self-movement cues; whereas, the odometer system is involved in deriving distance information from self-movement cues. The hippocampus functions similar to a logbook in that outward path unique information from the compass and odometer is used to derive the direction and distance of a path to the point at which movement was initiated. Finally, home base establishment may function to reset this system after each excursion and anchor environmental cues to self-movement cues. The combination of natural behaviors and kinematic analysis has proven to be a robust paradigm to investigate the neural basis of spatial orientation.     

Neurobiology of the homing pigeon—a review

Homing pigeons are well known as good homers, and the knowledge of principal parameters determining their homing behaviour and the neurological basis for this have been elucidated in the last decades. Several orientation mechanisms and parameters—sun compass, earth’s magnetic field, olfactory cues, visual cues—are known to be involved in homing behaviour, whereas there are still controversial discussions about their detailed function and their importance. This paper attempts to review and summarise the present knowledge about pigeon homing by describing the known orientation mechanisms and factors, including their pros and cons. Additionally, behavioural features like motivation, experience, and track preferences are discussed. All behaviour has its origin in the brain and the neuronal basis of homing and the neuroanatomical particularities of homing pigeons are a main topic of this review. Homing pigeons have larger brains in comparison to other non-homing pigeon breeds and particularly show increased size of the hippocampus. This underlines our hypothesis that there is a relationship between hippocampus size and spatial ability. The role of the hippocampus in homing and its plasticity in response to navigational experience are discussed in support of this hypothesis.     

The effect of clock-shift on the initial orientation of wild rock doves (<Emphasis Type="Italic">Columba l. livia</Emphasis>)

Previous experiments on wild rock doves ( Columba l. livia Gmelin) released within their familiar area revealed an evident effect of fast-shifting, although shifted doves, but not controls, tended to orient homeward. Such an outcome suggested a possible influence of the release time per se on the directional choices of the tested doves. In the present study, this hypothesis was investigated by comparing the orientation of slow-shifted birds to that of two control groups released at different times of the day. As would be expected if doves make use of a time-compensated sun compass, the bearings of shifted birds were deflected clockwise with respect to controls. The time of release itself seemed to influence only the scatter of the vanishing bearings of controls. These findings testify to the substantial similarity of clock-shift effects on the initial orientation of rock doves to those on homing pigeons released from familiar sites.