A framework for the study of genetic variation in migratory behaviour

Evolutionary change results from selection acting on genetic variation. For migration to be successful, many different aspects of an animal's physiology and behaviour need to function in a co-coordinated way. Changes in one migratory trait are therefore likely to be accompanied by changes in other migratory and life-history traits. At present, we have some knowledge of the pressures that operate at the various stages of migration, but we know very little about the extent of genetic variation in various aspects of the migratory syndrome. As a consequence, our ability to predict which species is capable of what kind of evolutionary change, and at which rate, is limited. Here, we review how our evolutionary understanding of migration may benefit from taking a quantitative-genetic approach and present a framework for studying the causes of phenotypic variation. We review past research, that has mainly studied single migratory traits in captive birds, and discuss how this work could be extended to study genetic variation in the wild and to account for genetic correlations and correlated selection. In the future, reaction-norm approaches may become very important, as they allow the study of genetic and environmental effects on phenotypic expression within a single framework, as well as of their interactions. We advocate making more use of repeated measurements on single individuals to study the causes of among-individual variation in the wild, as they are easier to obtain than data on relatives and can provide valuable information for identifying and selecting traits. This approach will be particularly informative if it involves systematic testing of individuals under different environmental conditions. We propose extending this research agenda by using optimality models to predict levels of variation and covariation among traits and constraints. This may help us to select traits in which we might expect genetic variation, and to identify the most informative environmental axes. We also recommend an expansion of the passerine model, as this model does not apply to birds, like geese, where cultural transmission of spatio-temporal information is an important determinant of migration patterns and their variation.

Animal migration: is there a common migratory syndrome?

Ornithologists, and especially northern hemisphere ornithologists, have traditionally thought of migration as an annual return movement of populations between regular breeding and non-breeding grounds. Problems arise because selection does not ordinarily act on populations and because organisms of many taxa (including birds) are clearly migrants, but fail to undertake movements of the kind described. There are also extensive return movements that are not migratory. I propose that it is more useful to think of migration as a syndrome of behavioral and other traits that function together within individuals, and that such a syndrome provides a common ground across taxa from aphids to albatrosses. Large-scale return movements of populations are one outcome of the syndrome. Similar behavioral and physiological traits serve both to define migration and to provide a test for it. I use two insect (Hemipteran) examples to illustrate migratory syndromes and to demonstrate that, in many migrants, behavior and physiology correlate with life history and morphological traits to form syndromes at two levels. I then compare the two Hemipterans with migration in birds, butterflies, and fish to assess the question of whether there are migratory syndromes in common between these diverse migrants. Syndromes are more similar at the level of behavior than when morphology and life history traits are included. Recognizing syndromes leads to important evolutionary questions concerning migration strategies, trade-offs, the maintenance of genetic variance and the responses of migratory syndromes to both similar and different selective regimes.

Ecological factors associated with the breeding and migratory phenology of high-latitude breeding western sandpipers

Environmental conditions influence the breeding and migratory patterns of many avian species and may have particularly dramatic effects on long-distance migrants that breed at northern latitudes. Environment, however, is only one of the ecological variables affecting avian phenology, and recent work shows that migration tactics may be strongly affected by changes in predator populations. We used long-term data from 1978 to 2000 to examine the interactions between snowmelt in western Alaska in relation to the breeding or migration phenologies of small shorebirds and their raptor predators. Although the sandpipers' time of arrival at Alaskan breeding sites corresponded with mean snowmelt, late snowmelts did delay breeding. These delays, however, did not persist to southward migration through British Columbia, likely due to the birds' ability to compensate for variance in the length of the breeding season. Raptor phenology at an early stopover site in British Columbia was strongly related to snowmelt, so that in years of early snowmelt falcons appeared earlier during the sandpipers' southbound migration. These differential effects indicate that earlier snowmelt due to climate change may alter the ecological dynamics of the predator-prey system.

Is re-feeding efficiency compromised by prolonged starvation during aestivation in the green striped burrowing frog, Cyclorana alboguttata?

We examined several morphological parameters of the gastrointestinal tract, digesta passage rates, and nutrient assimilation efficiencies of Green-striped Burrowing frogs (Cyclorana alboguttata) following prolonged fasting during three months of aestivation and compared these with frogs that had been continuously fed. Whole animal digesta passage rates were significantly reduced following three months aestivation as a result of a decreased digesta evacuation rate from the stomach. Furthermore, food was selectively retained in the small intestine for an increased time following three months of aestivation. Overall digestibility of food and nitrogen, carbon, and energy extraction efficiencies were not significantly different from control values following three months of aestivation. These findings suggest that C. alboguttata employs reduced digesta passage rates so as to maximize nutrient assimilation efficiency following prolonged food deprivation during aestivation. (C) 2003 Wiley-Liss, Inc.

The Australo-Papuan bird migration system: Another consequence of Wallace's Line

With respect to its avian migrants, Australo-Papua is a largely self-contained region. Only some 30 species of shorebird and 10 species of land bird migrate from Asia to Australo-Papua to winter. Possible factors precluding migration of Asian birds include long over-water distances and lack of suitable habitats in Australia or New Guinea. Whether evolved within the region or descended from Asian relatives, Australasian species all confine their migratory movements east of Wallace's Line. The most likely factors restricting migration to the region are climate and habitat. The open forests and arid habitats of Australasia are absent from south-east Asia, so there is little, if any, selection promoting extra-regional migration. The arid or semi-arid but otherwise mild climates of much of Australia favour partial migration and widely varying movement patterns and pathways, although with an underlying north to south component. Movements also occur between mainland Australia and New Guinea and Tasmania. The restriction of migration within Australo-Papua and the high variability of migratory pathways have important implications for the ecology and evolution of the Australasian avifauna.

Potential interactions between humans and non-breeding shorebirds on a subtropical intertidal flat

The primary aim of this study was to investigate whether bait harvesting, with all its inherent effects, occurring in the intertidal zone of a subtropical estuary, had an impact on a migratory shorebird, the eastern curlew Numenius madagascariensis. In a large-scale manipulative study (units of experiment were 1 ha plots), callianassid shrimp Trypaea australiensis populations were harvested simulating the technique (manual pumping) and the levels of harvesting intensity per unit area (347 shrimp per hectare per harvesting event) exhibited by bait-collectors in SE Australia and South Africa. It was found that at present levels of harvesting intensity per unit area (approximately 1% of standing stock removed per harvesting event) there is no threat to the stocks of Trypaea exploited by the curlews in Moreton Bay, Australia. However, the results show that the curlews themselves apply a considerable predation pressure on Trypaea. Based on the birds' foraging rates and densities, it was estimated that they would consume up to 100% of the initial Trypaea stock over the course of a non-breeding season (October to March). However, the stable seasonal trend in the density of the size-cohort of Trypaea preyed upon by the curlews indicates that the existing rates of predation are easily counterbalanced, e.g. through continuous density-dependent recruitment of these crustaceans. We suggest that this mechanism will provide for a stable foraging environment for both the shorebirds and bait collectors.

Why do eastern curlews Numenius madagascariensis feed on prey that lowers intake rate before migration?

Eastern curlews Numenius madagascariensis spending the nonbreeding season in eastern Australia foraged on three intertidal decapods: soldier crab Mictyris longicarpus, sentinel crab Macrophthalmus crassipes and ghost-shrimp Trypaea australiensis. Due to their ecology, these crustaceans were spatially segregated (=distributed in 'patches') and the curlews intermittently consumed more than one prey type. It was predicted that if the curlews behaved as intake rate maximizers, the time spent foraging on a particular prey (patch) would reflect relative availabilities of the prey types and thus prey-specific intake rates would be equal. During the mid-nonbreeding period (November-December), Mictyris and Macrophthalmus were primarily consumed and prey-specific intake rates were statistically indistinguishable (8.8 versus 10.1 kJ x min(-1)). Prior to migration (February), Mictyris and Trypaea were hunted and the respective intake rates were significantly different (8.9 versus 2.3 kJ x min(-1)). Time allocation to Trypaea-hunting was independent of the availability of Mictyris. Thus, consumption of Trypaea depressed the overall intake rate. Six hypotheses for consuming Trypaea before migration were examined. Five hypotheses: the possible error by the predator, prey specialization, observer overestimation of time spent hunting Trypaea, supplementary prey and the choice of higher quality prey due to a digestive bottleneck, were deemed unsatisfactory. The explanation for consumption of a low intake-rate but high quality prey (Trypaea) deemed plausible was diet optimisation by the Curlews in response to the pre-migratory modulation (decrease in size/processing capacity) of their digestive system. With a seasonal decrease in the average intake rate, the estimated intake per low tide increased from 1233 to 1508 kJ between the mid-nonbreeding and pre-migratory periods by increasing the overall time spent on the sandflats and the proportion of time spent foraging.

Target tissue influences the peripheral trajectory of mouse primary sensory olfactory axons

Primary olfactory neurons situated in the nasal septum project axons within fascicles along a highly stereotypical trajectory en route to the olfactory bulb. The ventral fascicles make a distinct dorsovental turn at the rear of the septum so as to reach the olfactory bulb. In the present study we have used a brain and nasal septum coculture system to examine the role of target tissue on the peripheral trajectory of olfactory sensory axons. In cultures of isolated embryonic nasal septa, olfactory axons form numerous parallel fascicles that project caudally in the submucosa, as they do in vivo. The ventral axon fascicles in the septum, however, often fail to turn, and do not project dorsally towards the roof of the nasal cavity. The presence of olfactory bulb, cortical, or tectal tissue apposed to the caudal end of the septum rescued this phenotype, causing the ventral fascicles to follow a normal in vivo-like trajectory. Ectopic placements of the explants revealed that brain tissue is not tropic for olfactory axons but appears to maintain the peripheral trajectory of growing axons in the nasal septum. Although primary olfactory axons are able to penetrate into olfactory bulb in vitro, they only superficially enter cortical tissue, whereas they do not grow into tectal explants. The ability of axons to differentially grow into different brain regions was shown to be unrelated to the migratory behavior of olfactory ensheathing cells, indicating that olfactory axons are directly responsive to guidance cues in the brain. (C) 2004 Wiley Periodicals, Inc.

Angiostrongrylus cantonensis as a cause of cerebrospinal disease in a yellow-tailed black cockatoo (Calyptorhynchus funereus) and two tawny frogmouths (Podargus strigoides)

A captive yellow-tailed black cockatoo (Calyptorhynchus funereus) and 2 free-living tawny frogmouths (Podargus strigoides), both native Australian species, were presented with neurologic signs including depression and pelvic limb paresis and paralysis. Despite supportive treatment, all 3 birds died or were euthanatized. On histologic examination, sections of metastrongyloid nematode larvae were found in the central nervous system of all 3 birds, whereas intact larvae, identified as Angiostrongylus cantonensis, were recovered from the brain and spinal cord of 2 birds. Angiostrongylus cantonensis, the rat lungworm. has an obligatory migratory phase through the host's central nervous system, which can cause severe pathologic lesions. Natural infections in accidental hosts have been documented only in mammals, and to our knowledge, angiostrongyliasis in avian hosts has not been previously reported.

Expression analysis of the Epha1 receptor tyrosine kinase and its high-affinity ligands Efna1 and Efna3 during early mouse development

Interaction of Eph receptor tyrosine kinases with their membrane bound ephrin ligands initiates bidirectional signaling events that regulate cell migratory and adhesive behavior. Whole-mount in situ hybridization revealed overlapping expression of the Epha1 receptor and its high-affinity ligands ephrin A1 (Efna1) and ephrin A3 (Efna3) in the primitive streak and the posterior paraxial mesoderm during early mouse development. These results show complex and dynamic expression for all three genes with expression domains that are successively complementary, overlapping, and divergent. (c) 2006 Elsevier B.V. All rights reserved.

The transcriptional coactivator Querkopf controls adult neurogenesis

The adult mammalian brain maintains populations of neural stem cells within discrete proliferative zones. Understanding of the molecular mechanisms regulating adult neural stem cell function is limited. Here, we show that MYST family histone acetyltransferase Querkopf (Qkf, Myst4, Morf)-deficient mice have cumulative defects in adult neurogenesis in vivo, resulting in declining numbers of olfactory bulb interneurons, a population of neurons produced in large numbers during adulthood. Qkf-deficient mice have fewer neural stem cells and fewer migrating neuroblasts in the rostral migratory stream. Qkf gene expression is strong in the neurogenic subventricular zone. A population enriched in multipotent cells can be isolated from this region on the basis of Qkf gene expression. Neural stem cells/progenitor cells isolated from Qkf mutant mice exhibited a reduced self-renewal capacity and a reduced ability to produce differentiated neurons. Together, our data show that Qkf is essential for normal adult neurogenesis.

Global flows of foreign talent: Identity anxieties in Singapore's ethnoscape

This paper draws on Appadurai's (1996) concept of ethnoscapes — the global flow of people or what has become increasingly popularized as the global flow of talent. Singapore has initiated a foreign talent policy to compete for a global pool of talent to make up for its shortfall of indigenous work-force. The rationale for recruiting foreign talent is informed by a nationalist competitive ideology to sustain Singapore in the new knowledge-based economy. This paper examines the competing and dissenting discourses surrounding the foreign talent policy. It argues that the mobility of migratory flow has transformative and disruptive effects at the level of culture and the identity landscape of Singapore, where its discursive cultural boundaries are drawn according to a nationalist framework. Drawing on theories and concepts of ‘diaspora’, ‘hybridity’, and ‘third space’, these are the political and cultural issues that this paper attempts to tease out. [ABSTRACT FROM AUTHOR]