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.     

Variation in migratory behavior influences regional genetic diversity and structure among American Kestrel populations (Falco sparverius) in North America

Birds employ numerous strategies to cope with seasonal fluctuations in high-quality habitat availability. Long distance migration is a common tactic; however, partial migration is especially common among broadly distributed species. Under partial migration systems, a portion of a species migrates, whereas the remainder inhabits breeding grounds year round. In this study, we identified effects of migratory behavior variation on genetic structure and diversity of American Kestrels (Falco sparverius), a widespread partial migrant in North America. American Kestrels generally migrate; however, a resident group inhabits the southeastern United States year round. The southeastern group is designated as a separate subspecies (F. s. paulus) from the migratory group (F. s. sparverius). Using mitochondrial DNA and microsatellites from 183 and 211 individuals, respectively, we illustrate that genetic structure is stronger among nonmigratory populations, with differentiation measures ranging from 0.060 to 0.189 depending on genetic marker and analysis approach. In contrast, measures from western North American populations ranged from 0 to 0.032. These findings suggest that seasonal migratory behavior is also associated with natal and breeding dispersal tendencies. We likewise detected significantly lower genetic diversity within nonmigratory populations, reflecting the greater influence of genetic drift in small populations. We identified the signal of population expansion among nonmigratory populations, consistent with the recent establishment of higher latitude breeding locations following Pleistocene glacial retreat. Differentiation of F. s. paulus and F. s. sparverius reflected subtle differences in allele frequencies. Because migratory behavior can evolve quickly, our analyses suggest recent origins of migratory American Kestrel populations in North America.     

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.     

To boldly go: individual differences in boldness influence migratory tendency

Partial migration, whereby only a fraction of the population migrates, is thought to be the most common type of migration in the animal kingdom, and can have important ecological and evolutionary consequences. Despite this, the factors that influence which individuals migrate and which remain resident are poorly understood. Recent work has shown that consistent individual differences in personality traits in animals can be ecologically important, but field studies integrating personality traits with migratory behaviour are extremely rare. In this study, we investigate the influence of individual boldness, an important personality trait, upon the migratory propensity of roach, a freshwater fish, over two consecutive migration seasons. We assay and individually tag 460 roach and show that boldness influences migratory propensity, with bold individuals being more likely to migrate than shy fish. Our data suggest that an extremely widespread personality trait in animals can have significant ecological consequences via influencing individual-level migratory behaviour.     

Regulation of migration in Mythimna separata (Walker) in China: a review integrating environmental, physiological, hormonal, genetic, and molecular factors

Each year the Mythimna separate (Walker), undertakes a seasonal, long-distance, multigeneration roundtrip migration between southern and northern China. Despite its regularity, the decision to migrate is facultative, and is controlled by environmental, physiological, hormonal, genetic, and molecular factors. Migrants take off on days 1 or 2 after eclosion, although the preoviposition period lasts ≈7 d. The trade-offs among the competing physiological demands of migration and reproduction are coordinated in M. separata by the "oogenesis-flight syndrome." Larvae that experience temperatures above or below certain thresholds accompanied by appropriate humidity, short photoperiod, poor nutrition, and moderate density tend to develop into migrants. However, there is a short window of sensitivity within 24 h after adult eclosion when migrants can be induced to switch to reproductive residents if they encounter extreme environmental factors including starvation, low temperature and long photoperiod. Juvenile hormone (JH) titer is low before migration but high titers are associated with termination of migratory behavior and the switch to reproduction. Early release of JH by the corpora allata in environmentally stressed 1-d old adults, otherwise destined by larval conditions to be migrants, switches them to residents. Offspring inherit parental additive genetic effects governing migratory behavior. However, they also retain flexibility in expression of both flight and reproductive life history traits. The insect neuropeptide, allatotropin, which activates corpora allata to synthesize JH, controls adult flight and reproduction. Future research directions to better understand regulation of migration in this species are discussed.     

Genomic legacy of migration in endangered caribou

Wide-ranging animals, including migratory species, are significantly threatened by the effects of habitat fragmentation and habitat loss. In the case of terrestrial mammals, this results in nearly a quarter of species being at risk of extinction. Caribou are one such example of a wide-ranging, migratory, terrestrial, and endangered mammal. In populations of caribou, the proportion of individuals considered as “migrants” can vary dramatically. There is therefore a possibility that, under the condition that migratory behavior is genetically determined, those individuals or populations that are migratory will be further impacted by humans, and this impact could result in the permanent loss of the migratory trait in some populations. However, genetic determination of migration has not previously been studied in an endangered terrestrial mammal. We examined migratory behavior of 139 GPS-collared endangered caribou in western North America and carried out genomic scans for the same individuals. Here we determine a genetic subdivision of caribou into a Northern and a Southern genetic cluster. We also detect >50 SNPs associated with migratory behavior, which are in genes with hypothesized roles in determining migration in other organisms. Furthermore, we determine that propensity to migrate depends upon the proportion of ancestry in individual caribou, and thus on the evolutionary history of its migratory and sedentary subspecies. If, as we report, migratory behavior is influenced by genes, caribou could be further impacted by the loss of the migratory trait in some isolated populations already at low numbers. Our results indicating an ancestral genetic component also suggest that the migratory trait and their associated genetic mutations could not be easily re-established when lost in a population.     

Detecting genetic structure in migrating bowhead whales off the coast of Barrow, Alaska

We develop a general framework for analysing and testing genetic structure within a migratory assemblage that is based on measures of genetic differences between individuals. We demonstrate this method using microsatellite DNA data from the Bering-Chukchi-Beaufort stock of bowhead whales (Balaena mysticetus), sampled via Inuit hunting during the spring and autumn migration off Barrow, Alaska. This study includes a number of covariates such as whale ages and the time separation between captures. Applying the method to a sample of 117 bowhead whales, we use permutation methods to test for temporal trends in genetic differences that can be ascribed to age-related effects or to timing of catches during the seasons. The results reveal a pattern with elevated genetic differences among whales caught about a week apart, and are statistically significant for the autumn migration. In contrast, we find no effects of time of birth or age-difference on genetic differences. We discuss possible explanations for the results, including population substructuring, demographic consequences of historical overexploitation, and social structuring during migration.     

Multi-cellular logistics of collective cell migration

During development, the formation of biological networks (such as organs and neuronal networks) is controlled by multicellular transportation phenomena based on cell migration. In multi-cellular systems, cellular locomotion is restricted by physical interactions with other cells in a crowded space, similar to passengers pushing others out of their way on a packed train. The motion of individual cells is intrinsically stochastic and may be viewed as a type of random walk. However, this walk takes place in a noisy environment because the cell interacts with its randomly moving neighbors. Despite this randomness and complexity, development is highly orchestrated and precisely regulated, following genetic (and even epigenetic) blueprints. Although individual cell migration has long been studied, the manner in which stochasticity affects multi-cellular transportation within the precisely controlled process of development remains largely unknown. To explore the general principles underlying multicellular migration, we focus on the migration of neural crest cells, which migrate collectively and form streams. We introduce a mechanical model of multi-cellular migration. Simulations based on the model show that the migration mode depends on the relative strengths of the noise from migratory and non-migratory cells. Strong noise from migratory cells and weak noise from surrounding cells causes "collective migration," whereas strong noise from non-migratory cells causes "dispersive migration." Moreover, our theoretical analyses reveal that migratory cells attract each other over long distances, even without direct mechanical contacts. This effective interaction depends on the stochasticity of the migratory and non-migratory cells. On the basis of these findings, we propose that stochastic behavior at the single-cell level works effectively and precisely to achieve collective migration in multi-cellular systems.     

The genetic basis of smoltification-related traits in Oncorhynchus mykiss

The timing and propensity for migration between fresh- and seawater is a key theme in the diversity of life histories within the salmonid fishes. Across salmonid species, life-history strategies range from wholly freshwater-resident populations, to migratory and nonmigratory variation within populations, to populations and species that are primarily migratory. Despite the central theme of migration to the evolution of these fishes, the genetic architecture of migration-related processes is poorly understood. Using a genetic cross of clonal lines derived from migratory and nonmigratory life-history types of Onchorhynchus mykiss (steelhead and rainbow trout, respectively), we have dissected the genetic architecture of the complex physiological and morphological transformation that occurs immediately prior to seaward migration (termed smoltification). Quantitative trait loci (QTL) analyses were used to identify the number, effects, and genomic location of loci associated with smoltification-related traits, including growth and condition factor, body coloration, morphology, and osmoregulatory enzymes during the smoltification period. Genetic analyses revealed numerous QTL, but one locus in particular is associated with multiple traits in single and joint analyses. Dissecting the genetic architecture of this highly complex trait has profound implications for understanding the genetic and evolutionary basis of life-history diversity within and among migratory fishes.     

Individual differences in migratory behavior shape population genetic structure and microhabitat choice in sympatric blackcaps (Sylvia atricapilla)

In migratory birds, traits such as orientation and distance are known to have a strong genetic background, and they often exhibit considerable within‐population variation. How this variation relates to evolutionary responses to ongoing selection is unknown because the underlying mechanisms that translate environmental changes into population genetic changes are unclear. We show that within‐population genetic structure in southern German blackcaps (Sylvia atricapilla) is related to individual differences in migratory behavior. Our 3‐year study revealed a positive correlation between individual migratory origins, denoted via isotope (δ²H) values, and genetic distances. Genetic diversity and admixture differed not only across a recently established migratory polymorphism with NW‐ and SW‐migrating birds but also across δ²H clusters within the same migratory route. Our results suggest assortment based on individual migratory origins which would facilitate evolutionary responses. We scrutinized arrival times and microhabitat choice as potential mechanisms mediating between individual variation in migratory behavior and assortment. We found significant support that microhabitat choice, rather than timing of arrival, is associated with individual variation in migratory origins. Moreover, examining genetic diversity across the migratory divide, we found migrants following the NW route to be genetically more distinct from each other compared with migrants following the traditional SW route. Our study suggests that migratory behavior shapes population genetic structure in blackcaps not only across the migratory divide but also on an individual level independent of the divide. Thus, within‐population variation in migratory behavior might play an important role in translating environmental change into genetic change.     

Adoption of alternative migratory tactics: a view from the ultimate mechanism and threshold trait changes in a salmonid fish

Partial migration, in which a portion of the population migrates while the rest of the population remains as residents, is a common form of migration. Alternative migratory tactics (AMTs) of partial migration are often determined by polygenic threshold traits. However, the ultimate mechanisms that drive inter‐population variations in threshold traits are not well understood. We present a simple schematic model to explain how the threshold trait changes with fitness consequences under opposing natural and artificial selection forces. We conducted a field test to evaluate the effects of migration difficulty (as a natural selective force) and selective captive breeding (as an artificial selective force) on threshold traits of a partially migratory fish. Male masu salmon Oncorhynchus masou in the Shari River system have AMTs divided into three population categories of hatchery, wild/above the waterfall, and wild/below the waterfall (control). The wild/above the waterfall salmon live in a high‐migration‐cost situation, and the threshold trait changed in a direction that promoted residency. In hatchery salmon, which are produced by migrant‐selective captive breeding, the threshold trait changed in a direction that promoted migration. In contrast, Dolly Varden charr Salvelinus malma displayed only resident tactics, and the threshold trait did not differ between the populations above and below the waterfall, indicating that environment did not explain the variation in the threshold trait. Our results support the model and suggest that opposing natural and artificial selection forces drive variations in the threshold traits and migratory patterns in the studied species. Our conceptual framework for the ultimate mechanism may help to better understand adoption of AMTs and production of diverse intraspecific traits in migratory animals.     

Consequences of the genetic threshold model for observing partial migration under climate change scenarios

Migration is a widespread phenomenon across the animal kingdom as a response to seasonality in environmental conditions. Partially migratory populations are populations that consist of both migratory and residential individuals. Such populations are very common, yet their stability has long been debated. The inheritance of migratory activity is currently best described by the threshold model of quantitative genetics. The inclusion of such a genetic threshold model for migratory behavior leads to a stable zone in time and space of partially migratory populations under a wide range of demographic parameter values, when assuming stable environmental conditions and unlimited genetic diversity. Migratory species are expected to be particularly sensitive to global warming, as arrival at the breeding grounds might be increasingly mistimed as a result of the uncoupling of long‐used cues and actual environmental conditions, with decreasing reproduction as a consequence. Here, we investigate the consequences for migratory behavior and the stability of partially migratory populations under five climate change scenarios and the assumption of a genetic threshold value for migratory behavior in an individual‐based model. The results show a spatially and temporally stable zone of partially migratory populations after different lengths of time in all scenarios. In the scenarios in which the species expands its range from a particular set of starting populations, the genetic diversity and location at initialization determine the species’ colonization speed across the zone of partial migration and therefore across the entire landscape. Abruptly changing environmental conditions after model initialization never caused a qualitative change in phenotype distributions, or complete extinction. This suggests that climate change‐induced shifts in species’ ranges as well as changes in survival probabilities and reproductive success can be met with flexibility in migratory behavior at the species level, which will reduce the risk of extinction.     

Migratory costs and the evolution of egg size and number in introduced and indigenous salmon populations

The trade-off between reproductive investment and migration should be an important factor shaping the evolution of life-history traits among populations following their radiation into habitats with different migratory costs and benefits. An experimentally induced difference in migratory rigor for families of chinook salmon (Oncorhynchus tshawytscha), of approximately 86 km and 413 m elevation, exacted a cost to somatic energy reserves (approximately 17% reduction in metabolizable mass) and ovarian investment (13.7% reduction in ovarian mass). This cost was associated with a reduction in egg size and paralleled the phenotypic pattern of divergence between two introduced New Zealand populations of common origin, presently breeding at sites with different migration distances. The genetic pattern of divergence of these same populations, detected under common rearing, was consistent with compensation for migratory costs (the population that migrates farther invested more in ovarian mass), but egg number more than egg size was associated with this evolution. These evolutionary patterns are consistent with what is known of the inheritance of these traits and with trade-offs and constraints favoring initial evolution in offspring number over offspring size. Analysis of egg number-size patterns of other Pacific salmon populations in their native range supported the hypothesis that migration strongly influences patterns of reproductive allocation, favoring a higher ratio of egg number to egg size with greater migration distance.     

Evolutionary genetics of partial migration – the threshold model of migration revis(it)ed

Partial migration is a common and widespread phenomenon in animal populations. Even though the ecological causes for the evolution and maintenance of partial migration have been widely discussed, the consequences of the genetics underlying differences in migration patterns have been little acknowledged. Here, I revise current ideas on the genetics of partial migration and identify open questions, focussing on migration in birds. The threshold model of migration describing the inheritance and phenotypic expression of migratory behaviour is strongly supported by experimental results. As a consequence of migration being a threshold trait, high levels of genetic variation can be preserved, even under strong directional selection. This is partly due to strong environmental canalization. This cryptic genetic variation may explain rapid de novo evolution of migratory behaviour in resident populations and the high prevalence of partial migration in animal populations. To date the threshold model of migration has been tested only under laboratory conditions. For obtaining a more realistic representation of migratory behaviour in the wild, the simple threshold model needs to be extended by considering that the threshold of migration or the liability may be modified by environmental effects. This environmental threshold model is valid for both facultative and obligate migration movements, and identifies genetic accommodation as an important process underlying evolutionary change in migration status. Future research should aim at identifying the major environmental variables modifying migration propensity and at determining reaction norms of the threshold and liability across variation in these variables.     

The influence of seasonal migration on range size in temperate North American passerines

Seasonal migration has been alternately proposed to promote geographic range size in some contexts and to constrain it in others, but it remains unclear if migratory behavior has a general effect on range size. Because migration involves movement, most hypotheses about the relationship between migration and range size invoke an influence of migration on the process of dispersal-mediated range expansion. Intuitively, a positive relationship between migratory behavior and dispersal ability could bolster range expansion among migratory species, yet some biogeographic patterns suggest that long-distance migration may instead impede range expansion, especially in the temperate zone. We conducted a comparative analysis of the relationship between migratory behavior and range size by testing the effect of migratory status, migration distance and morphological dispersal ability on breeding range size among all temperate North American passerines. Further, we assessed whether these traits affect range expansion into suitable habitat by analyzing their relationship with range filling (the proportion of climatically-suitable area occupied, or ‘filled’ by a species). Contrary to previous studies, we found migration and dispersal ability to be poor predictors of range size and range filling in North America. Rather, most variation in range size is explained by latitude. Our results suggest that migratory behavior does not affect range size within the scale of a continent, and furthermore, that temperate North American passerines’ breeding ranges are not influenced by their dispersal abilities. To better understand why migratory behavior appears to promote range size in some contexts and constrain it in others, future studies should investigate how migratory behavior affects dispersal at the individual level, as well as the relationship between the evolution of migratory behavior and the breadth of species’ climatic niches.     

Geographic and clonal variation in the milkweed-oleander aphid,Aphis nerii (Homoptera: Aphididae), for winged morph production,life history,and morphology in relation to host plant permanence

Summary Populations of the milkweed-oleander aphid,Aphis nerii, were sampled in California, Iowa and Puerto Rico. Among these localities the aphid's host plants differ greatly in permanence. I compared populations for migratory potential, measured as the proportion of winged offspring produced in response to being crowded, and for life history and morphometric traits of the subsequent adult winged aphids. I predicted a negative correlation between degree of host plant permanence and migratory potential. As predicted, aphids from Iowa, where migration on to temporary hosts must occur each year, produce a greater proportion of winged offspring (37.7%) than those from California (25.7%) or Puerto Rico (31.6%) where hosts are more permanent. However, hosts in Puerto Rico appear to be more permanent than those in California, yet the difference between populations for migratory potential was opposite to that predicted. Within California the prediction again held: aphids collected from the most impermanent sites produce the greatest proportion of winged offspring. There were no population differences for any life history or morphometric traits of winged aphids that are important contributors to fitness or migratory ability such as time to reproductive maturity, fecundity or wing length. Nor did any traits covary with migratory potential. Thus, there does not appear to be an association of life history and morphology with migratory potential that could enhance the colonizing ability of migrant aphids. I was unable to detect population differentiation for life history and morphology even though there is ample genetic variation within populations on which selection could act and an absence of constraints arising from genetic correlations that could prevent appropriate evolution of traits within populations. The exploitation of temporary host plants therefore occurs by an increase in the number of colonists produced and not by change in life history or morphology of those colonists.     

Are color or high rearing density related to migratory polyphenism in the band-winged grasshopper, Oedaleus asiaticus?

Locusts represent an impressive example of migratory polyphenism, with high densities triggering a switch from a solitarious, shorter dispersal range, and sometimes greenish phenotype to a gregarious and sometimes darker form exhibiting behavioral, morphological and physiological traits associated with long-distance migratory swarms. While such polyphenism has been well documented in Locusta migratoria and Schistocerca gregaria, the extent to which other grasshoppers exhibit this type of migratory polyphenism is unclear. Anecdotally, the Chinese grasshopper, Oedaleus asiaticus, forms migratory swarms comprised mostly of a darker, brown-colored morph, but also exhibits a non-migratory green-colored morph that predominates at low densities. In a population in Inner Mongolia not currently exhibiting migratory swarms, we found that while green and brown O. asiaticus are found concurrently across our sampled range, only brown grasshoppers were found in high densities. Differences between field-collected brown and green forms matched some but not key predictions associated with the hypothesis that the brown form is morphologically and physiologically specialized for gregarious migration. Controlling for body mass, brown forms had more massive thoraxes, abdomens and legs, and higher metabolic rates, but not more flight muscle or lipid stores. Further, the brown and green grasshoppers did not differ in gregarious behavior, and neither would fly in multiple lab and field trials. Lab or field-rearing at high densities for one-to-multiple juvenile instars caused grasshoppers to exhibit some morphological traits predicted to benefit migration (larger wings and a shift in relative mass from abdomen to thorax), but did not change color or induce flight behavior. One hypothesis to explain these data is that a migratory form of O. asiaticus is partially triggered by high field densities, but that existing ecological conditions blocked full expression of such traits (and outbreak swarms). Alternatively, color variation in this species may more tightly linked to other functions in this species such as crypsis or disease resistance, and mechanisms other than late-juvenile rearing density (e.g. genetic variation, maternal effects) may be more critical for promoting variation in color and/or migratory polyphenism.     

Migration costs drive convergence of threshold traits for migratory tactics

Partial migration of some, but not all, members of a population is a common form of migration. We evaluated how migration costs influence which members migrate in 10 populations of two salmonid species. The migratory patterns of both species were evaluated based on the size at maturity for resident males, which is the threshold trait that determines the migratory tactics used within a population. In both species, this size was smaller in males located further from the sea, where migration costs are presumably higher. Moreover, the threshold sizes at maturity in males were correlated between both species. Our results suggest that migration costs are a significant convergent selective force on migratory tactics and life-history traits in nature.     

Adcyap1 polymorphism covaries with breeding latitude in a Nearctic migratory songbird,the Wilson's warbler (Cardellina pusilla)

Understanding the genetic background of complex behavioral traits, showing multigenic control and extensive environmental effects, is a challenging task. Among such traits, migration is known to show a large additive genetic component. Yet, the identification of specific genes or gene regions explaining phenotypic variance in migratory behavior has received less attention. Migration ultimately depends on seasonal cycles, and polymorphism at phenological candidate genes may underlie variation in timing of migration or other aspects of migratory behavior. In this study of a Nearctic–Neotropical migratory songbird, the Wilson's warbler (Cardellina pusilla), we investigated the association between polymorphism at two phenological candidate genes, Clock and Adcyap1, and two aspects of the migratory phenotype, timing of spring migration through a stopover site and inferred latitude of the breeding destination. The breeding destination of migrating individuals was identified using feather deuterium ratio (δ²H), which reliably reflects breeding latitude throughout the species' western breeding range. Ninety‐eight percent of the individuals were homozygous at Clock, and the rare heterozygotes did not deviate from homozygous migration phenology. Adcyap1 was highly polymorphic, and allele size was not significantly associated with migration date. However, Adcyap1 allele size significantly positively predicted the inferred breeding latitude of males but not of females. Moreover, we found a strong positive association between inferred breeding latitude and Adcyap1 allele size in long‐distance migrating birds from the northern sector of the breeding range (western Canada), while this was not the case in short‐distance migrating birds from the southern sector of the breeding range (coastal California). Our findings support previous evidence for a role of Adcyap1 in shaping the avian migratory phenotype, while highlighting that patterns of phenological candidate gene–phenotype associations may be complex, significantly varying between geographically distinct populations and even between the sexes.     

鸟类迁徙:在全球变暖趋势下的演化、调控与发展(英文)

最近几十年的研究证实 ,鸟类迁徙在很大程度上受到遗传因素的直接控制。有证据表明 ,存在某种先天的迁徙动因并涉及以下几方面的遗传调控 :(1)迁徙过程的起始、持续以及结束 ;(2 )迁徙活动量 ,即决定鸟类飞行距离的遗传参数 ;(3)迁徙方向 ;(4)生理参数 ,特别是迁徙期间的脂肪贮存 ,以及对于那些部分个体迁徙的鸟种而言 ,决定个体迁徙与否的生理参数。双因素选择实验表明 ,部分迁徙群经由几个世代的选择即可转变成完全的迁徙群或非迁徙群。新迁徙方向以及由此导致的新越冬区的改变 ,也能在野生鸟类中迅速实现。至少在以往研究得最为透彻的鸟种 (黑顶林莺Sylviaatricapilla)中 ,“迁徙”或“非迁徙”是先天性的 ,与特异性迁徙活动量相关 (尤如一时间程序 ) ,前者 (迁徙的 )已证实是由一种阈机制所控制的。一项新的鸟类迁徙理论假设 ,即使好些完全迁徙的类群 ,较低水平的迁徙活动量选择也会导致阈的异位 ,低于这一阈值就会出现非迁徙个体。因此 ,通过选择作用 ,一个迁徙型种群可以通过部分迁徙型转变为非迁徙型。这种中间阶段在现存鸟类中十分普遍。它始见于生物演化早期 ,就鸟类而言 ,可能在原始鸟类就已具备。模型运算表明 ,在施以强定向选择情况下 ,迁徙鸟类经过约 4 0年可转变为留鸟 ,反之亦然。这就解