The nature and role of learning in the orientation and migratory behavior of fishes

Synopsis Fish migration may be viewed as the product of two processes; the selection and tracking of optimal environmental conditions through time and space, and the use of predictive information about environmental structure to bias movements towards a goal. The establishment and maintenance of directional bias is based on the interaction of experience and instinct. The preoccupation of much fish orientation research with innate fixed patterns of behavior on one hand and hydrodynamics on the other has led us to underestimate the possibility that orientation is a flexible process relying on developmental sequences, calibration of the motor-sensory interaction based on experience and the learning of environmental pattern. Evidence illustrating how experience and learning may influence the direction of movement and how the goal is recognized is presented according to two general categories: (a) imprinting and early experience and (b), spatial learning, including the social transmission of migratory routes and directions. In the first category, the olfactory hypothesis of salmon homing is briefly reviewed and new data presented describing olfactory imprinting in Atlantic salmon,Salmo salar. In the second category, evidence is presented demonstrating the modifiability of sun-compass orientation and the ability of some fish species to learn the spatial distribution of landmarks. The role of social transmission in the migration of coral reef fishes is reviewed. The possible role of these learning phenomena in the formation of familiar area maps, route-based and location-based navigation and the critical distance factor is considered. The relationship between life history and the nature of learning in migratory orientation is discussed     

Animal navigation: general properties of directed walks

The ability to locomote is a defining characteristic of all animals. Yet, all but the most trivial forms of navigation are poorly understood. Here we report and discuss the analytical results of an in-depth study of a simple navigation problem. In principle, there are two strategies for navigating a straight course. One is to use an external directional reference and to continually reorient with reference to it. The other is to monitor body rotations from internal sensory information only. We showed previously that, at least for simple representations of locomotion, the first strategy will enable an animal or mobile agent to move arbitrarily far away from its starting point, but the second strategy will not do so, even after an infinite number of steps. This paper extends and generalizes the earlier results by demonstrating that these findings are true even when a very general model of locomotion is used. In this general model, error components within individual steps are not independent, and directional errors may be biased. In the absence of a compass, the expected path of a directed walk in general approximates a logarithmic spiral. Some examples are given to illustrate potential applications of the quantitative results derived here. Motivated by the analytical results developed in this work, a nomenclature for directed walks is proposed and discussed. Issues related to path integration in mammals and robots, and measuring the curvature of a noisy path are also addressed using directed walk theory.     

Behavioral analysis of polarization vision in tethered flying locusts

For spatial navigation many insects rely on compass information derived from the polarization pattern of the sky. We demonstrate that tethered flying desert locusts (Schistocerca gregaria) show e-vector-dependent yaw-torque responses to polarized light presented from above. A slowly rotating polarizer (5.3° s^–1) induced periodic changes in yaw torque corresponding to the 180° periodicity of the stimulus. Control experiments with a rotating diffuser, a weak intensity pattern, and a stationary polarizer showed that the response is not induced by intensity gradients in the stimulus. Polarotaxis was abolished after painting the dorsal rim areas of the compound eyes black, but remained unchanged after painting the eyes except the dorsal rim areas. During rotation of the polarizer, two e-vectors (preferred and avoided e-vector) induced no turning responses: they were broadly distributed from 0 to 180° but, for a given animal, were perpendicular to each other. The data demonstrate polarization vision in the desert locust, as shown previously for bees, flies, crickets, and ants. Polarized light is perceived through the dorsal rim area of the compound eye, suggesting that polarization vision plays a role in compass navigation of the locust.     

Quantitative evaluation of the major determinants of human gait

Accurate knowledge of the isolated contributions of joint movements to the three-dimensional displacement of the center of mass (COM) is fundamental for understanding the kinematics of normal walking and for improving the treatment of gait disabilities. Saunders et al. (1953) identified six kinematic mechanisms to explain the efficient progression of the whole-body COM in the sagittal, transverse, and coronal planes. These mechanisms, referred to as the major determinants of gait, were pelvic rotation, pelvic list, stance knee flexion, foot and knee mechanisms, and hip adduction. The aim of the present study was to quantitatively assess the contribution of each major gait determinant to the anteroposterior, vertical, and mediolateral displacements of the COM over one gait cycle. The contribution of each gait determinant was found by applying the concept of an ‘influence coefficient’, wherein the partial derivative of the COM displacement with respect to a prescribed determinant was calculated. The analysis was based on three-dimensional measurements of joint angular displacements obtained from 23 healthy young adults walking at slow, normal and fast speeds. We found that hip flexion, stance knee flexion, and ankle-foot interaction (comprised of ankle plantarflexion, toe flexion and the displacement of the center of pressure) are the major determinants of the displacements of the COM in the sagittal plane, while hip adduction and pelvic list contribute most significantly to the mediolateral displacement of the COM in the coronal plane. Pelvic rotation and pelvic list contribute little to the vertical displacement of the COM at all walking speeds. Pelvic tilt, hip rotation, subtalar inversion, and back extension, abduction and rotation make negligible contributions to the displacements of the COM in all three anatomical planes.     

Ecophysiological consequences of non-random leaf orientation in the prairie compass plant,Silphium laciniatum

Summary The prairie compass plant (Silphium laciniatum L.) has vertical leaves that are characteristically oriented in a north-south plane (i.e., the flat surfaces of the lamina face east and west). We explored the consequences of this orientation by determining basic photosynthetic and water use characteristics in response to environmental factors and by determining total daily photosynthesis and water use of leaves held in different orientations. Average maximum CO₂ exchange rate (CER) of leaves near Ames, IA was constant at 22 micromol m^–2 s^–1 from May through August and then declined. CER did not exhibit a distinct lightsaturation point. CER at photon flux densities near full sunlight was constant from 22 to 35°C leaf temperature but declined at higher temperatures. However, leaf temperatures rarely exceed 35°C during the growing season. There was no change in the pattern of response of CER to temperature over the growing season. We constrained leaves to face east-west (EW,=natural), to face north-south (NS), or to be horizontal (HOR) on eight days in 1986–1988. EW leaves had the highest light interception, leaf temperatures, CER, and transpiration early and late in the day, whereas HOR leaves had the highest values in the middle of the day. Integrations of CER and transpiration over the eight daytime periods showed EW and HOR leaves to have equivalent carbon gain, higher than that of NS leaves. HOR leaves had the highest daily transpiration. Daily water use efficiency (WUE, carbon gained/water lost) was always highest in EW leaves, with the HOR leaves having 16% lower WUE and NS leaves having 33% lower WUE. The natural orientation of compass plant leaves results in equivalent or higher carbon gain and in increased WUE when compared to leaves with other possible orientations; this is likely to have a selective advantage in a prairie environment.     

Surgical lesion of the anterior optic tract abolishes polarotaxis in tethered flying locusts, <Emphasis Type="Italic">Schistocerca gregaria</Emphasis>

Many insects can detect the polarization pattern of the blue sky and rely on polarization vision for sky compass orientation. In laboratory experiments, tethered flying locusts perform periodic changes in flight behavior under a slowly rotating polarizer even if one eye is painted black. Anatomical tracing studies and intracellular recordings have suggested that the polarization vision pathway in the locust brain involves the anterior optic tract and tubercle, the lateral accessory lobe, and the central complex of the brain. To investigate whether visual pathways through the anterior optic tract mediate polarotaxis in the desert locust, we transected the tract on one side and tested polarotaxis (1) with both eyes unoccluded and (2) with the eye of the intact hemisphere painted black. In the second group of animals, but not in the first group, polarotaxis was abolished. Sham operations did not impair polarotaxis. The experiments show that the anterior optic tract is an indispensable part of visual pathways mediating polarotaxis in the desert locust.     

Moon orientation in adult and young sandhoppers

In the last 30 years, lunar orientation has received little attention from students of animal orientation. Even in Talitrus saltator, the first animals in which the lunar compass was demonstrated, research did not continue. Our studies have demonstrated that: 1) chronometrically compensated lunar orientation is independent of the earth's magnetic field (an ever present non-chronometric orientation reference); 2) lunar orientation is independent of the lunar shape; and 3) the lunar compass is also used by young animals born in the laboratory (without experience in nature). Accepted: 30 September 1998     

Contradictory results on the role of polarized light in compass calibration in migratory songbirds

Experiments with migrating birds on the interaction between magnetic and celestial cues have produced heterogeneous results. A recent study claimed that the magnetic compass in passerine migrants is calibrated by the pattern of polarized light at sunset and sunrise and that the area just above the horizon is crucial for this calibration. To test the latter hypothesis, we performed a similar experiment with Australian Silvereyes. It produced contrary results, however, the birds, in spite of observing the natural polarization pattern at sunrise and sunset down to the horizon in an altered magnetic field, continued in their normal southerly magnetic direction when subsequently tested in the local geomagnetic field—the conflict between magnetic and polarized light cues had not caused them to recalibrate their magnetic compass. This contradicts the assumption that skylight polarization patterns generally serve as a primary calibration reference for migratory songbirds.