An experimental displacement and over 50 years of tag-recoveries show that monarch butterflies are not true navigators.

Monarch butterflies (Danaus plexippus) breeding in eastern North America are famous for their annual fall migration to their overwintering grounds in Mexico. However, the mechanisms they use to successfully reach these sites remain poorly understood. Here, we test whether monarchs are true navigators who can determine their location relative to their final destination using both a "compass" and a "map". Using flight simulators, we recorded the orientation of wild-caught monarchs in southwestern Ontario and found that individuals generally flew in a southwest direction toward the wintering grounds. When displaced 2,500 km to the west, the same individuals continued to fly in a general southwest direction, suggesting that monarchs use a simple vector-navigation strategy (i.e., use a specific compass bearing without compensating for displacement). Using over 5 decades of field data, we also show that the directional concentration and the angular SD of recoveries from tagged monarchs largely conformed to two mathematical models describing the directional distribution of migrants expected under a vector-navigation strategy. A third analysis of tagged recoveries shows that the increasing directionality of migration from north to south is largely because of the presence of geographic barriers that guide individuals toward overwintering sites. Our work suggests that monarchs breeding in eastern North America likely combine simple orientation mechanisms with geographic features that funnel them toward Mexican overwintering sites, a remarkable achievement considering that these butterflies weigh less than a gram and travel thousands of kilometers to a site they have never seen.

Physiological characterization of the compound eye in monarch butterflies with focus on the dorsal rim area.

The spectral, angular and polarization sensitivities of photoreceptors in the compound eye of the monarch butterfly (Danaus plexippus) are examined using electrophysiological methods. Intracellular recordings reveal a spectrally homogenous population of UV receptors with optical axes directed upwards and >or=10 degrees to the contralateral side. Based on optical considerations and on the opsin expression pattern (Sauman et al. 2005), we conclude that these UV receptors belong to the anatomically specialized dorsal rim area (DRA) of the eye. Photoreceptors in the main retina with optical axes <10 degrees contralateral or ipsilateral have maximal sensitivities in the UV (lambda(max)

Do monarch butterflies use polarized skylight for migratory orientation?

To test if migratory monarch butterflies use polarized light patterns as part of their time-compensated sun compass, we recorded their virtual flight paths in a flight simulator while the butterflies were exposed to patches of naturally polarized blue sky, artificial polarizers or a sunny sky. In addition, we tested butterflies with and without the polarized light detectors of their compound eye being occluded. The monarchs' orientation responses suggested that the butterflies did not use the polarized light patterns as a compass cue, nor did they exhibit a specific alignment response towards the axis of polarized light. When given direct view of the sun, migratory monarchs with their polarized light detectors painted out were still able to use their time-compensated compass: non-clockshifted butterflies, with their dorsal rim area occluded, oriented in their typical south-southwesterly migratory direction. Furthermore, they shifted their flight course clockwise by the predicted approximately 90 degrees after being advance clockshifted 6 h. We conclude that in migratory monarch butterflies, polarized light cues are not necessary for a time-compensated celestial compass to work and that the azimuthal position of the sun disc and/or the associated light-intensity and spectral gradients seem to be the migrants' major compass cue.

Connecting the navigational clock to sun compass input in monarch butterfly brain.

Migratory monarch butterflies (Danaus plexippus) use a time-compensated sun compass to navigate to their overwintering grounds in Mexico. Although polarized light is one of the celestial cues used for orientation, the spectral content (color) of that light has not been fully explored. We cloned the cDNAs of three visual pigment-encoding opsins (ultraviolet [UV], blue, and long wavelength) and found that all three are expressed uniformly in main retina. The photoreceptors of the polarization-specialized dorsal rim area, on the other hand, are monochromatic for the UV opsin. Behavioral studies support the importance of polarized UV light for flight orientation. Next, we used clock protein expression patterns to identify the location of a circadian clock in the dorsolateral protocerebrum of butterfly brain. To provide a link between the clock and the sun compass, we identified a CRYPTOCHROME-staining neural pathway that likely connects the circadian clock to polarized light input entering brain.

Functional morphology of the ear in fossorial rodents, Microtus arvalis and Arvicola terrestris.

Functionally relevant features and parameters of the outer, middle, and inner ear were studied morphologically and morphometrically in two species of voles, smaller Microtus arvalis and larger Arvicola terrestris. The findings in these fossorial (i.e., burrowing) rodents with components of surface activity were compared with respective findings reported for taxonomically related muroid rodents representing the same size classes but different eco-morphotypes: obligate subterranean rodents (Ellobius talpinus and Spalax ehrenbergi superspecies) and generalized rodents (Mus domesticus and Rattus norvegicus). The ear in voles was characterized by traits reported for subterranean rodents. The eardrum was round, without a distinct pars flaccida, and had an area of 5.4 mm2 in M. arvalis and 9 mm2 in A. terrestris. The middle ear exhibited reduced goniale, enlarged incus nearly parallel to the manubrium of the malleus. The malleus-incus lever ratio amounted to 2.1 (M. arvalis) and 2.0 (A. terrestris). The malleus-incus complex weighed about 0.8 mg in both vole species. The stapedial footplate had an area of 0.3 mm2 in M. arvalis and 0.4 mm2 in A. terrestris. The cochlea had 2.3 coils in both vole species; the basilar membrane was 8.5 mm and 10.5 mm long in M. arvalis and A. terrestris, respectively. There were on average 1,030 (M. arvalis) and 1,220 (A. terrestris) inner hair cells, and 3,760 (M. arvalis) and 4,250 (A. terrestris) outer hair cells in the organ of Corti. In quantitative terms, all these (as well as some further) traits and parameters were intermediate (related to body size) between those reported for generalized rodents on the one hand and subterranean ones on the other. The sound transmission system of the ear seems to be best tuned to frequencies of about 8-16 kHz with a high-frequency cut-off at about 50-60 kHz. The ear of A. terrestris seems to be tuned to somewhat lower frequencies than that in M. arvalis. In this aspect as well as regarding hearing sensitivity (as judged from the mechanical transmission parameters), voles can be considered intermediate not only in their lifestyle but also in their hearing abilities between the subterranean rodents (mole-vole and blind mole-rat) and the surface dwellers (house mouse and Norway rat).

Cryptochromes and neuronal-activity markers colocalize in the retina of migratory birds during magnetic orientation.

Migratory birds can use a magnetic compass for orientation during their migratory journeys covering thousands of kilometers. But how do they sense the reference direction provided by the Earth's magnetic field? Behavioral evidence and theoretical considerations have suggested that radical-pair processes in differently oriented, light-sensitive molecules of the retina could enable migratory birds to perceive the magnetic field as visual patterns. The cryptochromes (CRYs) have been suggested as the most likely candidate class of molecules, but do CRYs exist in the retina of migratory birds? Here, we show that at least one CRY1 and one CRY2 exist in the retina of migratory garden warblers and that garden-warbler CRY1 (gwCRY1) is cytosolic. We also show that gwCRY1 is concentrated in specific cells, particularly in ganglion cells and in large displaced ganglion cells, which also showed high levels of neuronal activity at night, when our garden warblers performed magnetic orientation. In addition, there seem to be striking differences in CRY1 expression between migratory and nonmigratory songbirds at night. The difference in CRY1 expression between migrants and nonmigrants is particularly pronounced in the large displaced ganglion cells known to project exclusively to a brain area where magnetically sensitive neurons have been reported. Consequently, cytosolic gwCRY1 is well placed to possibly be the primary magnetic-sensory molecule required for light-mediated magnetoreception.