How the clear-sky angle of polarization pattern continues underneath clouds: full-sky measurements and implications for animal orientation.
One of the biologically most important parameters of the cloudy sky is the proportion P of the celestial polarization pattern available for use in animal navigation. We evaluated this parameter by measuring the polarization patterns of clear and cloudy skies using 180 degrees (full-sky) imaging polarimetry in the red (650 nm), green (550 nm) and blue (450 nm) ranges of the spectrum under clear and partly cloudy conditions. The resulting data were compared with the corresponding celestial polarization patterns calculated using the single-scattering Rayleigh model. We show convincingly that the pattern of the angle of polarization (e-vectors) in a clear sky continues underneath clouds if regions of the clouds and parts of the airspace between the clouds and the earth surface (being shady at the position of the observer) are directly lit by the sun. The scattering and polarization of direct sunlight on the cloud particles and in the air columns underneath the clouds result in the same e-vector pattern as that present in clear sky. This phenomenon can be exploited for animal navigation if the degree of polarization is higher than the perceptual threshold of the visual system, because the angle rather than the degree of polarization is the most important optical cue used in the polarization compass. Hence, the clouds reduce the extent of sky polarization pattern that is useful for animal orientation much less than has hitherto been assumed. We further demonstrate quantitatively that the shorter the wavelength, the greater the proportion of celestial polarization that can be used by animals under cloudy-sky conditions. As has already been suggested by others, this phenomenon may solve the ultraviolet paradox of polarization vision in insects such as hymenopterans and dipterans. The present study extends previous findings by using the technique of 180 degrees imaging polarimetry to measure and analyse celestial polarization patterns. (+info)
Speeds and wingbeat frequencies of migrating birds compared with calculated benchmarks.
Sixteen species of birds passing Falsterbo in southwest Sweden during the autumn migration season were observed using short-range optical methods. Air speeds and wingbeat frequencies were measured, reduced to sea level, and compared with benchmark values computed by Flight.bas, a published flight performance program based on flight mechanics. The benchmark for air speed was the calculated sea-level value of the minimum power speed (V(mp)). The mean speeds of three raptor species that flew by flap-gliding were below V(mp), apparently because the flap-glide cycle involved slowing down below V(mp) when gliding and accelerating back up to V(mp) when flapping. The mean speeds of 11 species that flew by continuous flapping were between 0.82V(mp) and 1.27V(mp). Two passerine species that flew by bounding had mean speeds of 1.70V(mp) and 1.96V(mp), but these high mean speeds reflected their ability to fly faster against head winds. These results do not support predictions from optimal migration theory, which suggest that migrating birds 'should' fly faster, relative to V(mp). However, observations were restricted for technical reasons to birds flying below 200 m and may not represent birds that were seriously committed to long-distance migration. The benchmark wingbeat frequency (f(ref)) was derived from dimensional reasoning, not from statistical analysis of observations. Observed wingbeat frequencies ranged from 0.81f(ref) to 1.05f(ref), except in the two bounding species, whose wingbeat frequencies appeared anomalously high. However, the mechanics of bounding with a power fraction q imply that gravity during the flapping phase is increased by a factor 1/q, and when the value of gravity was so adjusted in the expression for f(ref), the wingbeat frequencies of the two bounding species were predicted correctly as a function of the power fraction. In small birds with more muscle power than is required to fly at speeds near V(mp), bounding is an effective method of adjusting the specific work in the muscle fibres, allowing conversion efficiency to be maximised over a wide range of speeds. (+info)
Light-dependent magnetoreception in birds: the behaviour of European robins, Erithacus rubecula, under monochromatic light of various wavelengths and intensities.
To investigate how magnetoreception is affected by the wavelength and intensity of light, we tested European robins, Erithacus rubecula, under monochromatic lights of various wavelengths at two intensities using oriented behaviour as an indicator of whether the birds could derive directional information from the geomagnetic field. At a quantal flux of 7 x 10(15) quanta s(-1) m(-2), the birds were well oriented in their migratory direction east of North under 424 nm blue, 510 nm turquoise and 565 nm green light, whereas they were disoriented under 590 nm yellow light. Increasing the intensity of light at the same wavelengths more than sixfold to 43 x 10(15) quanta s(-1) m(-2) resulted in a change in behaviour: under bright blue and green light, the birds now showed a preference for the East-West axis, with the majority of headings at the western end; under bright turquoise light, they oriented unimodally towards a direction slightly west of North. Under bright yellow light, the birds continued to be disoriented. These findings suggest a rather complex relationship between the receptors involved in magnetoreception. Magnetoreception appears to follow rules that are different from those of vision, suggesting that light-dependent magnetoreception may involve receptors and neuronal pathways of its own. (+info)
Effects of duration and time of food availability on photoperiodic responses in the migratory male blackheaded bunting (Emberiza melanocephala).
The effects of the duration and time of food availability on stimulation of the photoperiodic responses (fattening and gain in body mass, and growth and development of testes) were investigated in the migratory blackheaded bunting (Emberiza melanocephala). Two experiments were performed. Experiment I examined the effects of a reduction in the duration of food supply in buntings that were subjected to long day lengths (16h:8h L:D) and received food ad libitum (group I) or for restricted durations, coinciding with the end of the lights-on period, of 8h (group II) and 4h (group III). Buntings of group I gained in body mass, whereas there was a mixed response in group II (half the birds gained and half lost body mass), and all birds of group III lost body mass. There was no effect on testis growth in groups I and II, but testes grew more slowly in group III. Experiment 2 investigated the effects of both the duration and the time of food availability. Of five groups of birds, group I was exposed to an 8h:16h L:D photoperiod, and groups II-V were exposed to 16h:8h L:D. Whereas birds of groups I and II received food ad libitum, those of groups III-V were fed only for 5 h, at zt 0-5 (group III), zt 5.5-10.5 (group IV) or zt 11-16 (group V), where zt = zeitgeber time and zt 0 refers to the beginning of the lights-on period. Apart from duration, the timing of food availability also had an effect on photoperiodic stimulation under the 16h:8h L:D photoperiod. Birds that were fed ad libitum fattened and gained in body mass, whereas among restricted feeding groups, only birds in the group fed during the first 5 h (zt 0-5, group III) showed a significant increase in body mass (albeit considerably lower than in the ad libitum group). Birds fed during the middle 5h (zt 5.5-10.5, group IV) showed an intermediate response, and those fed during the last 5h (zt 11-16, group V) lost body mass. Testicular growth was suppressed in birds that were fed for 5 h in the evening, but not in those fed for the same period in the morning or in the middle of the long day. Taken together, these results show that the duration of food supply and/or the time of day at which food is available affect photoperiodic stimulation of fattening and gain in body mass as well as the growth and development of gonads in the blackheaded bunting. (+info)
Complex bird clocks.
The circadian pacemaking system of birds comprises three major components: (i) the pineal gland, which rhythmically synthesizes and secretes melatonin; (ii) a hypothalamic region, possibly equivalent to the mammalian suprachiasmatic nuclei; and (iii) the retinae of the eyes. These components jointly interact, stabilize and amplify each other to produce a highly self-sustained circadian output. Their relative contribution to overt rhythmicity appears to differ between species and the system may change its properties even within an individual depending, for example, on its state in the annual cycle or its photic environment. Changes in pacemaker properties are partly mediated by changes in certain features of the pineal melatonin rhythm. It is proposed that this variability is functionally important, for instance, for enabling high-Arctic birds to retain synchronized circadian rhythms during the low-amplitude zeitgeber conditions in midsummer or for allowing birds to adjust quickly their circadian system to changing environmental conditions during migratory seasons. The pineal melatonin rhythm, apart from being involved in generating the avian pacemaking oscillation, is also capable of retaining day length information after isolation from the animal. Hence, it appears to participate in photoperiodic after-effects. Our results suggest that complex circadian clocks have evolved to help birds cope with complex environments. (+info)
Juvenile hormone regulation of longevity in the migratory monarch butterfly.
Monarch butterflies (Danaus plexippus) of eastern North America are well known for their long-range migration to overwintering roosts in south-central Mexico. An essential feature of this migration involves the exceptional longevity of the migrant adults; individuals persist from August/September to March while their summer counterparts are likely to live less than two months as adults. Migrant adults persist during a state of reproductive diapause in which both male and female reproductive development is arrested as a consequence of suppressed synthesis of juvenile hormone. Here, we describe survival in monarch butterflies as a function of the migrant syndrome. We show that migrant adults are longer lived than summer adults when each are maintained under standard laboratory conditions, that the longevity of migrant adults is curtailed by treatment with juvenile hormone and that the longevity of summer adults is increased by 100% when juvenile hormone synthesis is prevented by surgical removal of its source, the corpora allatum. Thus, monarch butterfly persistence through a long winter season is ensured in part by reduced ageing that is under endocrine regulation, as well as by the unique environmental properties of their winter roost sites. Phenotypic plasticity for ageing is an integral component of the monarch butterflies' migration-diapause syndrome. (+info)
Deriving dispersal distances from genetic data.
Dispersal is one of the most important factors determining the genetic structure of a population, but good data on dispersal distances are rare because it is difficult to observe a large sample of dispersal events. However, genetic data contain unbiased information about the average dispersal distances in species with a strong sex bias in their dispersal rates. By plotting the genetic similarity between members of the philopatric sex against some measure of the distance between them, the resulting regression line can be used for estimating how far dispersing individuals of the opposite sex have moved before settling. Dispersers showing low genetic similarity to members of the opposite sex will on average have originated from further away. Applying this method to a microsatellite dataset from lions (Panthera leo) shows that their average dispersal distance is 1.3 home ranges with a 95% confidence interval of 0.4-3.0 home ranges. These results are consistent with direct observations of dispersal from our study population and others. In this case, direct observations of dispersal distance were not detectably biased by a failure to detect long-range dispersal, which is thought to be a common problem in the estimation of dispersal distance. (+info)
Unexpected coherence and conservation.
The effects of migration in a network of patch populations, or metapopulation, are extremely important for predicting the possibility of extinctions both at a local and a global scale. Migration between patches synchronizes local populations and bestows upon them identical dynamics (coherent or synchronous oscillations), a feature that is understood to enhance the risk of global extinctions. This is one of the central theoretical arguments in the literature associated with conservation ecology. Here, rather than restricting ourselves to the study of coherent oscillations, we examine other types of synchronization phenomena that we consider to be equally important. Intermittent and out-of-phase synchronization are but two examples that force us to reinterpret some classical results of the metapopulation theory. In addition, we discuss how asynchronous processes (for example, random timing of dispersal) can paradoxically generate metapopulation synchronization, another non-intuitive result that cannot easily be explained by the standard theory. (+info)