Experimental studies of the development of migratory orientation mechanisms

Summary Recent experimental studies (since ca 1985) on the ontogeny of orientation mechanisms in migratory birds are reviewed. The processes and interactions are synthesized into a framework that may help identify critical research questions. Birds that grow up in the earth's magnetic field develop the ability to perform appropriate migratory orientation even in the absence of any experience with relevant visual cues. In two species, large changes in direction during the course of migration seem to be controlled by an endogenous time program. In one of these, the pied flycatcher (Ficedula hypoleuca), the correct magnetic orientation seems to occur only when the magnetic fields appropriate to the latitudes encountered en route were experienced at the proper seasonal time. The magnetic compass may be modified by visual experience with either the daytime or night sky. Celestial rotation may be the calibrating reference in this case, as it is in the development of the star compass. Young Savannah sparrows (Passerculus sandwichensis) learn to perform compass orientation at sunset based on polarized skylight. This compass capability seems to be calibrated by magnetic directions. Some problems of experimental design and the interpretation of results from experiments on development are discussed.     

Magnetic inclination compass: A basis for the migratory orientation of birds in the Northern and Southern Hemisphere

We conducted orientation experiments with Silvereyes,Zosterops lateralis, Australian passerine migrants, to see whether birds living in the Southern Hemisphere in a magnetic field with an upward inclination orient in the same way as birds in the Northern Hemisphere that experience a downward inclination of the magnetic field. Tested indoors in the local geomagnetic field, the birds preferred southerly directions corresponding to their migratory direction in spring. In a magnetic field with a reversed vertical component, they reversed their directional tendencies. This shows that the magnetic compass of Silvereyes also functions as an inclination compass based on the inclination of the field lines instead of the polarity.     

Overcast tests with clock-shifted pigeons

In order to investigate the pigeon's compass mechanism, a series of overcast tests with clock-shifted birds were run at two familiar release sites. While controls were able to assume a correct homeward direction, the experimental birds' initial orientation cannot be explained either on the basis of a time-compensated sun compass or of a time-independent magnetic compass. Speculative explanations of our paradoxical results are attempted.     

Sensory basis of bird orientation

Summary Sensory information which may be essential for the complex process of orientation of birds is described in this article. The use of vibrational, visual, chemical, olfactory, magnetic cues and their receptive mechanisms, as far as they are known, are explained. Special reference is given to the behavioral and physiological aspects of magnetic sensitivity.     

Sensory basis of bird orientation

Sensory information which may be essential for the complex process of orientation of birds is described in this article. The use of vibrational, visual, chemical, olfactory, magnetic cues and their receptive mechanisms, as far as they are known, are explained. Special reference is given to the behavioral and physiological aspects of magnetic sensitivity.     

The sun compass

Summary The sun compass was discovered by G. Kramer in caged birds showing migratory restlessness. Subsequent experiments with caged brids employing directional training and clock shifts, carried out by Hoffmann and by Schmidt-Koenig, showed that sun azimuth is used and sun altitude ignored. McDonald found the accuracy to be ±3^o–±5^o. According to Hoffmann and to Schmidt-Koenig, caged birds trained at medium northern latitudes were able to allow for the sun's apparent movement north of the arctic circle but not in equatorial and trans-equatorial latitudes.In homing experiments, and employing clock shifts, Schmidt-Koenig demonstrated that the sun compass is used by homing pigeons during initial orientation. This finding supports the existence of a map and compass navigational system. Pigeons living in equatorial latitudes utilize the sun compass even under the extreme solar conditions of equinox (Ranvaud, Schmidt-Koenig, Ganzhorn et al.). The use of the sun compass during zenith passage of the sun is being investigated.     

Magnetic orientation and celestial cues in migratory orientation

Summary Young birds on their first migration possess innate information on the direction of their migration route. It is present in two forms, using celestial rotation and the geomagnetic field as references. These two systems, together with information provided by factors associated with sunset, interact in a complex way to establish the migratory direction. During ontogeny, celestial rotation appears to be dominant; during migration, however, celestial cues appear to be controlled by the magnetic field. The factors associated with sunset — the view of the setting sun, the characteristic pattern of polarized light — are important secondary cues which seem to derive their directional significance from the magnetic field. Their role appears to be more variable, with possible species-specific differences. During spring migration and later autumn migrations, flying in the migratory direction is complemented by navigational processes which enable the birds to return to a specific home site known from previous stays.     

A magnetic pulse leads to a temporary deflection in the orientation of migratory birds

Migratory Australian Silvereyes were treated with a strong magnetic pulse designed to alter the magnetization of the small magnetite particles that are found in birds' heads. Prior to the treatment, the birds preferred the northeasterly migratory direction. The pulse initially resulted in a 90° clockwise shift of orientation; however, within about a week, the birds seemed to return to their original headings. These findings, which seem to suggest an involvement of magnetite in migratory orientation, are in contrast with previous findings which indicated that it is a light-dependent process. They are discussed in view of the current concepts on magnetoreception and on the role of magnetic information in avian orientation.     

Wind and orientation of migrating birds: A review

Summary Migratory flights are strongly affected by wind, and birds have developed many adaptations to cope with wind effects. By day, overland migrants at high altitudes may often allow crosswinds to drift their tracks laterally from the preferred heading. In contrast, many birds at low altitude adjust their headings to compensate for drift, and may overcompensate to allow for previous drift. The relative motion of landscape features is probably used to sense drift, at least by day. By night, some overland migrants compensate fully for drift but others do not; no pattern is obvious. Over the sea, compensation is rarely if ever total; wave patterns may allow partial compensation. Other adaptations can include reduction of drift by flying at times and/or altitudes without strong crosswinds. Some birds recognize the need to change course to allow for previous wind displacement, and reorient at least roughly toward the original route or destination. Some juveniles en route to previously-unvisited wintering grounds seem to have this ability, but corroboration is needed. Such reorientation may not require a true navigation ability. However, some birds have unexplained abilities to sense the wind while aloft.     

Mechanism of neurogenesis in adult avian brain

Summary Adult neurogenesis in birds offers unique opportunities to study basic questions addressing the birth, migration and differentiation of neurons. Neurons in adult canaries originate from discrete proliferative regions on the walls of the lateral ventricles. They migrate away from their site of birth, initially at high rates, along the processes of radial cells. The rates of dispersal diminish as the young neurons invade regions devoid of radial fibers, probably under the guidance of other cues. The discrete sites of birth in the ventricular zone generate neurons that end up differentiating throughout the telencephelon. New neurons may become interneurons or projection neurons; the latter connect two song control nuclei between neostriatum and archistriatum. Radial cells, that in mammals disappear as neurogenesis comes to an end, persist in the adult avian brain. The presence of radial cells may be key to adult neurogenesis. Not only do they serve as guides for initial dispersal, they also divide and may be the progenitors of new neurons.     

Mechanism of neurogenesis in adult avian brain

Adult neurogenesis in birds offers unique opportunities to study basic questions addressing the birth, migration and differentiation of neurons. Neurons in adult canaries originate from discrete proliferative regions on the walls of the lateral ventricles. They migrate away from their site of birth, initially at high rates, along the processes of radical cells. The rates of dispersal diminish as the young neurons invade regions devoid of radial fibers, probably under the guidance of other cues. The discrete sites of birth in the ventricular zone generate neurons that end up differentiating throughout the telencephalon. New neurons may become interneurons or projection neurons; the latter connect two song control nuclei between neostriatum and archistriatum. Radial cells, that in mammals disappear as neurogenesis comes to an end, persist in the adult avian brain. The presence of radial cells may be key to adult neurogenesis. Not only do they serve as guides for initial dispersal, they also divide and may be the progenitors of new neurons.     

Evolutionary aspects of orientation and migration in birds

Summary Evolutionary aspects of avian migratory orientation and navigation are reviewed. A theoretical approach to the evolution of complex (endogenously programmed) migratory behavior is presented using the comparative method of ordering a progression in existing behavioral characteristics from dispersal, to facultative migration, to obligate migration.     

Collective cell migration has distinct directionality and speed dynamics

When a constraint is removed, confluent cells migrate directionally into the available space. How the migration directionality and speed increase are initiated at the leading edge and propagate into neighboring cells are not well understood. Using a quantitative visualization technique—Particle Image Velocimetry (PIV)—we revealed that migration directionality and speed had strikingly different dynamics. Migration directionality increases as a wave propagating from the leading edge into the cell sheet, while the increase in cell migration speed is maintained only at the leading edge. The overall directionality steadily increases with time as cells migrate into the cell-free space, but migration speed remains largely the same. A particle-based compass (PBC) model suggests cellular interplay (which depends on cell–cell distance) and migration speed are sufficient to capture the dynamics of migration directionality revealed experimentally. Extracellular Ca²⁺ regulated both migration speed and directionality, but in a significantly different way, suggested by the correlation between directionality and speed only in some dynamic ranges. Our experimental and modeling results reveal distinct directionality and speed dynamics in collective migration, and these factors can be regulated by extracellular Ca²⁺ through cellular interplay. Quantitative visualization using PIV and our PBC model thus provide a powerful approach to dissect the mechanisms of collective cell migration.     

Olfactory navigation in birds

Summary Many bird species rely on an osmotactic mechanism to find food sources even at a considerable distance. Pigeons also rely on local odours for homeward orientation, and they integrate those perceived during passive transportation with those at the release site. It is possible to design experiments in which birds are given false olfactory information, and predictions about the effects of this can be made and tested. Pigeons build up their olfactory map by associating wind-borne odours with the directions from which they come; this was shown by experiments which aimed at preventing, limiting or altering this association. Some objections have been made to this conclusion; namely that even anosmic pigeons are sometimes homeward oriented, that they may be demotivated in flying or disturbed in their general behaviour, and that olfactory cues may be only one component of pigeo's navigational repertoire. The most recent experiments, however, confirm that pigeons derive directional information from atmospheric odouts. The lack of any knowledge about the chemical nature and distribution of the odorants which allow pigeons to navigate hinders progress in this area of research.     

Spatiotemporal programs and genetics of orientation

Summary A number of migratory bird species have endogenous annual rhythms that regulate the entire annual cycle including the migratory portion. Moreover, captive migrants display inherited migratory activity; this could theoretically also be used by free-living migrants as a time-program for migration. Finally, this heritable migratory activity is oriented in a seasonally appropriate direction even in naive birds. These, characteristics should enable inexperienced migrants isolated from contact with experienced conspecifics to utilize a heritable vector-navigation program to migrate from the breeding grounds to the winter quarters. That is, migrants should reach goal areas they have never experienced by migrating in programmed directions, for as long a period as the genetically fixed time-program for migratory activity induces them to do so. The time-course of migration as established by trapping stations, theoretical influences of environmental variables on migratory programs, and also compensatory behavior and migratory backup measures, are discussed. The present evidence supports the view that a large number of migrants are essentially brought to their wintering areas by vector-navigation systems.     

Magnetische Kernresonanz und Festkörperphysik

Summary Nuclear magnetic resonance (NMR) deals with isotopes possessing a nuclear magnetic dipole moment. The nuclei are used as probes to explore the internal magnetic fields in solids. Thus NMR has become a new tool in solid-state physics. Some typical examples of the application of NMR to solid-state problems are discussed.The location and orientation of water molecules in crystals are determined by NMR with relative ease. The second moment of an NMR line shape yields information on the structure as well as on the mobility of the atoms. The various types of resonance frequency shifts may be used for the understanding of the interactions and bonds in the solid and the dynamics of the crystal lattice.Very important are nuclei possessing both a magnetic dipole and an electric quadrupole moment. They serve as very sensitive detectors of the internal electric fields prevailing in solids. Apart from further structural information, NMR spectra influenced by electric quadrupole interaction may be used for studying such topics as aluminium-silicon ordering in feldspars or phase transitions in ferroelectrics.The utility of NMR in treating problems of the bonding is exemplified by showing that the covalent effects in the alkali halides are very small.Some investigations of ferromagnetism, antiferromagnetism and superconductivity are mentioned.     

Conceptual approaches to avian navigation systems

Summary The general basis of migratory orientation in birds is most probably an endogenous time-and-direction program. Directions are selected with respect to celestial and geomagnetic clues. Using these clues, a bird may reach a large population-specific area; however, it will hardly be able to find a particular location, for instance its previous breeding site. Homing to a familiar site over several hundred kilometres of unfamiliar terrain appears to be based on the smelling of atmospheric trace compounds. Conceptual approaches to the mechanism of olfactory navigation have as yet only reached an early state of speculation.     

The early history of ship magnetism: The Airy-Scoresby controversy

With the advent of iron-built ships in the early nineteenth century the problem of managing a magnetic compass on board presented considerable difficulty. Prominent among the early scientists who tackled the problem were George Biddell Airy and William Scoresby. Airy had provided a mechanical system, employing correctors in the form of steel magnets and wrought iron masses, by which the ship's magnetic field at the compass position is neutralized. He based his system on the concept that the magnetism acquired by an iron ship during her construction remained with the ship throughout her life. Scoresby, on the other hand, thought differently. After having conducted a thorough and systematic series of experiments on iron plates and bars, he concluded that the magnetic character of an iron ship is liable to sudden and unexpected change. Scoresby argued, therefore, that Airy's system was defective and, indeed, dangerous. The aim of this paper is to discuss the two views which brought Airy and Scoresby into a conflict which is documented in a series of letters to the editor of The Athenaeum.     

高纯铝再结晶晶界面的择尤取向

相比于传统的晶体择尤取向（晶体织构,简称织构）,晶界面择尤取向是一个新的科学问题.系统地研究晶界面择尤取向的形成规律和演化机制对合理调控显微组织结构以显著改善材料的各种使用性能具有非常重要的科学意义和应用价值.本文利用基于体视学原理和电子背散射衍射技术的晶界特征分布五参数表征分析方法,研究了冷轧变形高纯铝（99.99%）再结晶晶界面的择尤取向问题.结果表明,在再结晶及晶粒长大过程中,晶界面择尤取向由最初的{2 2 3}晶面转变为{1 1 1}晶面,最后取向在{1 0 0}晶面上.分析指出,冷轧变形高纯铝再结晶组织中,小角度晶界（取向差介于2°~15°）是其晶界构成的主体部分;在晶粒长大过程中,晶体织构由{1 1 0}〈1 1 2〉黄铜织构为主转变为以{0 1 1}〈1 0 0〉高斯织构和{0 2 3}〈1 0 0〉织构共存为主,受晶面能和晶体织构的影响,小角度晶界一般择尤取向于可形成能量较低的倾侧型小角度晶界所对应的晶面上.这是导致高纯铝再结晶晶界面择尤取向随晶粒长大而发生上述规律性变化的主要原因.     

Differential roles of multiple adhesion molecules in cell migration: granule cell migration in cerebellum

The migration of cerebellar granule cells from the external granular layer to the internal granular layer is mediated by the radical Bergmann glial fiber. Recent works have shown that cell adhesion molecules, extra-cellular matrix proteins and proteolytic enzymes or their activators are involved in this process. Immuno-localization studies showed differential temporal and spatial expression patterns of different adhesion molecules, their isoforms, and post-translational modification during different stages of granule cell migration. Functional perturbation experiments using cerebellar explant cultures demonstrated that several adhesion molecules as well as plasminogen activator are involved in granule cell migration and are required in different stages. Other systems used to study granule cell migration including dissociated microwell cultures and granule cell deficient mouse mutants are discussed in the context of adhesion molecules. The results accumulated so far suggest that the migration of granule cells is a complex process in which the cooperation of a group of molecules with different functions, some for adhesion some for de-adhesion, are required to fulfill the different needs during the migratory course.