Understanding the migratory orientation program of birds: extending laboratory studies to study free flying migrants in a natural setting

For many years, orientation in migratory birds has primarily been studied in the laboratory. Although a laboratory-based setting enables greater control over environmental cues, the laboratory-based findings must be confirmed in the wild in free-flying birds to be able to fully understand how birds orient during migration. Despite the difficulties associated with following free-flying birds over long distances, a number of possibilities currently exist for tracking the long distance, sometimes even globe-spanning, journeys undertaken by migrating birds. Birds fitted with radio transmitters can either be located from the ground or from aircraft (conventional tracking), or from space. Alternatively, positional information obtained by onboard equipment (e.g., GPS units) can be transmitted to receivers in space. Use of these tracking methods has provided a wealth of information on migratory behaviors that are otherwise very difficult to study. Here, we focus on the progress in understanding certain components of the migration-orientation system. Comparably exciting results can be expected in the future from tracking free-flying migrants in the wild. Use of orientation cues has been studied in migrating raptors (satellite telemetry) and thrushes (conventional telemetry), highlighting that findings in the natural setting may not always be as expected on the basis of cage-experiments. Furthermore, field tracking methods combined with experimental approaches have finally allowed for an extension of the paradigmatic displacement experiments performed by Perdeck in 1958 on the short-distance, social migrant, the starling, to long-distance migrating storks and long-distance, non-socially migrating passerines. Results from these studies provide fundamental insights into the nature of the migratory orientation system that enables experienced birds to navigate and guide inexperienced, young birds to their species-specific winter grounds.

Differential effects of magnetic pulses on the orientation of naturally migrating birds

In migratory passerine birds, strong magnetic pulses are thought to be diagnostic of the remagnetization of iron minerals in a putative sensory system contained in the beak. Previous evidence suggests that while such a magnetic pulse affects the orientation of migratory birds in orientation cages, no effect was present when pulse-treated birds were tested in natural migration. Here we show that two migrating passerine birds treated with a strong magnetic pulse, designed to alter the magnetic sense, migrated in a direction that differed significantly from that of controls when tested in natural conditions. The orientation of treated birds was different depending on the alignment of the pulse with respect to the magnetic field. These results can aid in advancing understanding of how the putative iron-mineral-based receptors found in birds' beaks may be used to detect and signal the intensity and/or direction of the Earth's magnetic field.

Testing the role of sensory systems in the migratory heading of a songbird

The identification of the sensory cues and mechanisms by which migratory birds are able to reach the same breeding and wintering grounds year after year has eluded biologists despite more than 50 years of intensive study. While a number of environmental cues have been proposed to play a role in the navigation of birds, arguments still persist about which cues are essential for the experience based navigation shown by adult migrants. To date, few studies have tested the sensory basis of navigational cues used during actual migration in the wild: mainly laboratory based studies or homing during the non-migratory season have been used to investigate this behaviour. Here we tested the role of olfactory and magnetic cues in the migration of the catbird (Dumetella carolinensis) by radio tracking the migration of birds with sensory manipulations during their actual migratory flights. Our data suggest that adult birds treated with zinc sulphate to produce anosmia were unable to show the same orientation as control adults, and instead reverted to a direction similar to that shown by juveniles making their first migration. The magnetic manipulation had no effect on the orientation of either adults or juveniles. These results allow us to propose that the olfactory sense may play a role in experience based migration in adult catbirds. While the olfactory sense has been shown to play a role in the homing of pigeons and other birds, this is the first time it has been implicated in migratory orientation.

True navigation in migrating gulls requires intact olfactory nerves

During migratory journeys, birds may become displaced from their normal migratory route. Experimental evidence has shown that adult birds can correct for such displacements and return to their goal. However, the nature of the cues used by migratory birds to perform long distance navigation is still debated. In this experiment we subjected adult lesser black-backed gulls migrating from their Finnish/Russian breeding grounds (from >60°N) to Africa (to < 5°N) to sensory manipulation, to determine the sensory systems required for navigation. We translocated birds westward (1080 km) or eastward (885 km) to simulate natural navigational challenges. When translocated westwards and outside their migratory corridor birds with olfactory nerve section kept a clear directional preference (southerly) but were unable to compensate for the displacement, while intact birds and gulls with the ophthalmic branch of the trigeminal nerve sectioned oriented towards their population-specific migratory corridor. Thus, air-borne olfactory information seems to be important for migrating gulls to navigate successfully in some circumstances.

Migratory navigation in birds: new opportunities in an era of fast-developing tracking technology

Birds have remained the dominant model for studying the mechanisms of animal navigation for decades, with much of what has been discovered coming from laboratory studies or model systems. The miniaturisation of tracking technology in recent years now promises opportunities for studying navigation during migration itself (migratory navigation) on an unprecedented scale. Even if migration tracking studies are principally being designed for other purposes, we argue that attention to salient environmental variables during the design or analysis of a study may enable a host of navigational questions to be addressed, greatly enriching the field. We explore candidate variables in the form of a series of contrasts (e. g. land vs ocean or night vs day migration), which may vary naturally between migratory species, populations or even within the life span of a migrating individual. We discuss how these contrasts might help address questions of sensory mechanisms, spatiotemporal representational strategies and adaptive variation in navigational ability. We suggest that this comparative approach may help enrich our knowledge about the natural history of migratory navigation in birds.

Assortative mating as a mechanism for rapid evolution of a migratory divide.

There have been numerous recent observations of changes in the behavior and dynamics of migratory bird populations, but the plasticity of the migratory trait and our inability to track small animals over large distances have hindered investigation of the mechanisms behind migratory change. We used habitat-specific stable isotope signatures to show that recently evolved allopatric wintering populations of European blackcaps Sylvia atricapilla pair assortatively on their sympatric breeding grounds. Birds wintering further north also produce larger clutches and fledge more young. These findings describe an important process in the evolution of migratory divides, new migration routes, and wintering quarters. Temporal segregation of breeding is a way in which subpopulations of vertebrates may become isolated in sympatry.

Indicators of the impact of climate change on migratory species

The Bonn Convention on the Conservation of Migratory Species of Wild Animals adopted a Resolution in 2005 recognising the impacts of climate change on migratory species. It called on Contracting Parties to undertake more research to improve our understanding of these impacts and to implement adaptation measures to reduce foreseeable adverse effects. Given the large diversity of taxa and species affected by climate change, it is impossible to monitor all species and effects thereof. However, it is likely that many of the key ecological and physical processes through which climate change may impact wildlife could be monitored using a suite of indicators, each comprising parameters of species/populations or groups of species as proxies for wider assemblages, habitats and ecosystems. Herein, we identify a suite of 17 indicators whose attributes could reveal negative impacts of climate change on the global status of migratory species: 4 for birds, 4 for marine mammals, 2 for sea turtles, 1 for fish, 3 for land mammals and 3 for bats. A few of these indicators would be relatively straightforward to develop, but most would require additional data collation, and in many cases methodological development. Choosing and developing indicators of the impacts of climate change on migratory species is a challenge, particularly with endangered species, which are subject to many other pressures. To identify and implement conservation measures for these species, indicators must account for the full ensemble of pressures, and link to a system of alerts and triggers for action.

True navigation in birds: from quantum physics to global migration

Birds are capable of true navigation, the ability to return to a known goal from a place they have never visited before. This is demonstrated most spectacularly during the vast migratory journeys made by these animals year after year, often between continents and occasionally global in nature. However, it remains one of the great unanswered questions in science, despite more than 50 years of research in this field. Nevertheless, the study of true navigation in birds has made significant advances in the previous 20 years, in part thanks to the integration of many disciplines outside its root in behavioural biology, to address questions of neurobiology, molecular aspects, and the physics of sensory systems and environmental cues involved in bird navigation, often involving quantum physics. However, true navigation remains a controversial field, with many conflicting and confusing results making interpretation difficult, particularly for those outside or new to the field. Unlike many general texts on migration, which avoid discussion of these issues, this review will present these conflicting findings and assess the state of the field of true navigation during bird migration.

Polarized skylight does not calibrate the compass system of a migratory bat

In a recent study, Greif et al. (2014) demonstrated a functional role of polarized light for a bat species confronted with a homing task. These non-migratory bats appeared to calibrate their magnetic compass by using polarized skylight at dusk, yet it is unknown if migratory bats also use these cues for calibration. During autumn migration, we equipped Nathusius' bats, Pipistrellus nathusii, with radio transmitters and tested if experimental animals exposed to a 90° rotated band of polarized light during dusk, would head in a different direction compared with control animals. After release, bats of both groups continued their journey in the same direction. This observation argues against the use of a polarization-calibrated magnetic compass by this migratory bat and questions that the ability of using polarized light for navigation is a consistent feature in bats. This finding matches with observations in some passerine birds that used polarized light for calibration of their magnetic compass before but not during migration.

Merely Birds of Passage: Lady Hariot Dufferin's Travel Writings and Medical Work in India, 1884-88

This article examines the travel writings and medical work in India of Lady Hariot Dufferin, Vicereine of India between 1884 and 1888. Lady Dufferin accompanied her husband, the Viceroy Lord Dufferin, through various social and political engagements in India, and carved her own niche in colonial and postcolonial history as a pioneer in the medical training of women in India. The article examines her travel writings on India and explores the nature of her complicity in the Raj, as well as the gendered nature of the separate public role she created for herself in relation to her 'zenana work' in providing medical care for the women of India. The author suggests that, through her work, Lady Dufferin challenges and extends the theoretical paradigms of postcolonialist and feminist critiques of empire.

Caged birds and cloning machines: how student imagery 'speaks' to us about cultures of schooling and student participation

With the advent of the United Nations Convention on the Rights of the Child (CRC), there is an increasing requirement that schools ensure children and young people's views are voiced, listened to and taken seriously on matters of significance. Encouraging these shifts by law is one thing; changing the culture in schools is another. For a significant proportion of schools, actively engaging students' voices on how they experience education poses a significant challenge and crucial gaps may exist between the rhetoric espoused and a school's readiness for genuine student involvement. This ethnographic study illuminates tensions that persist between headteachers' espoused views of how students are valued and students' creative images of their actual post-primary schooling experience. If cultures of schooling are to nurture the true spirit of democratic pupil participation implied by changes in the law, there is a need to develop genuine processes of student engagement in which students and staff can collaborate towards greater shared understandings of a school's priorities.

Two Birds, One Stone: Combining Student Assessment and Socio-Legal Research

We describe here a method of assessment for students. A number of short-comings of traditional assessment methods, especially essays and examinations, are discussed and an alternative assessment method, the student project, is suggested. The method aims not just to overcome the short-comings of more traditional methods, but also to provide over-worked and under-resourced academics with viable primary data for socio-legal research work. Limitations to the method are discussed, with proposals for minimising the impact of these limitations. The whole �student project� approach is also discussed with reference to the Quality Assurance Agency benchmark standards for law degrees, standards which are expected of all institutions in the UK.

Early birds are sexy: male age, dawn song and extrapair paternity in blue tits, Cyanistes (formerly Parus) caeruleus

Sexual selection theory predicts that signals reflecting the relative quality of individuals should be used in mate choice. Females could base their choice of copulation partners on male secondary sexual traits that honestly signal male age, as predicted by the age-based indicator mechanism. Studies have shown that female blue tits prefer older males and that aspects of dawn song reflect male quality, but it remains unknown whether dawn song characteristics correlate with male age. We compared dawn song characteristics of second-year (SY) and older (ASY) male blue tits (cross-sectional analysis), and tested for age-related changes within individuals (longitudinal analysis) and differential overwinter survival of SY males. We further investigated the relation between dawn song and paternity gain and loss. We found that ASY male blue tits began to sing earlier relative to sunrise than did SY males. This difference in the onset of dawn singing was due to age-related changes in individual performance rather than differential survival of individuals with varying expression of the trait. Males that began to sing earlier at dawn had more mating partners, and were more likely to gain extrapair paternity. Our findings suggest that the onset of dawn song can provide a simple mechanism for females to assess the relative quality of their mate and of neighbouring males. We propose that females use the onset of singing as a cue for their choice of extrapair partners. (c) 2006 The Association for the Study of Animal Behaviour. Published by Elsevier Ltd. All rights reserved.