Competitive Dynamics in MSTd: A Mechanism for Robust Heading Perception Based on Optic Flow

Human heading perception based on optic flow is not only accurate, it is also remarkably robust and stable. These qualities are especially apparent when observers move through environments containing other moving objects, which introduce optic flow that is inconsistent with observer self-motion and therefore uninformative about heading direction. Moving objects may also occupy large portions of the visual field and occlude regions of the background optic flow that are most informative about heading perception. The fact that heading perception is biased by no more than a few degrees under such conditions attests to the robustness of the visual system and warrants further investigation. The aim of the present study was to investigate whether recurrent, competitive dynamics among MSTd neurons that serve to reduce uncertainty about heading over time offer a plausible mechanism for capturing the robustness of human heading perception. Simulations of existing heading models that do not contain competitive dynamics yield heading estimates that are far more erratic and unstable than human judgments. We present a dynamical model of primate visual areas V1, MT, and MSTd based on that of Layton, Mingolla, and Browning that is similar to the other models, except that the model includes recurrent interactions among model MSTd neurons. Competitive dynamics stabilize the model’s heading estimate over time, even when a moving object crosses the future path. Soft winner-take-all dynamics enhance units that code a heading direction consistent with the time history and suppress responses to transient changes to the optic flow field. Our findings support recurrent competitive temporal dynamics as a crucial mechanism underlying the robustness and stability of perception of heading.     

Systematic Biases in Human Heading Estimation

Heading estimation is vital to everyday navigation and locomotion. Despite extensive behavioral and physiological research on both visual and vestibular heading estimation over more than two decades, the accuracy of heading estimation has not yet been systematically evaluated. Therefore human visual and vestibular heading estimation was assessed in the horizontal plane using a motion platform and stereo visual display. Heading angle was overestimated during forward movements and underestimated during backward movements in response to both visual and vestibular stimuli, indicating an overall multimodal bias toward lateral directions. Lateral biases are consistent with the overrepresentation of lateral preferred directions observed in neural populations that carry visual and vestibular heading information, including MSTd and otolith afferent populations. Due to this overrepresentation, population vector decoding yields patterns of bias remarkably similar to those observed behaviorally. Lateral biases are inconsistent with standard Bayesian accounts which predict that estimates should be biased toward the most common straight forward heading direction. Nevertheless, lateral biases may be functionally relevant. They effectively constitute a perceptual scale expansion around straight ahead which could allow for more precise estimation and provide a high gain feedback signal to facilitate maintenance of straight-forward heading during everyday navigation and locomotion.     

Haptic perception of object curvature in Parkinson's disease

Background

The haptic perception of the curvature of an object is essential for adequate object manipulation and critical for our guidance of actions. This study investigated how the ability to perceive the curvature of an object is altered by Parkinson''s disease (PD).

Methodology/Principal Findings

Eight healthy subjects and 11 patients with mild to moderate PD had to judge, without vision, the curvature of a virtual “box” created by a robotic manipulandum. Their hands were either moved passively along a defined curved path or they actively explored the curved curvature of a virtual wall. The curvature was either concave or convex (bulging to the left or right) and was judged in two locations of the hand workspace–a left workspace location, where the curved hand path was associated with curved shoulder and elbow joint paths, and a right workspace location in which these joint paths were nearly linear. After exploring the curvature of the virtual object, subjects had to judge whether the curvature was concave or convex. Based on these data, thresholds for curvature sensitivity were established. The main findings of the study are: First, 9 out 11 PD patients (82%) showed elevated thresholds for detecting convex curvatures in at least one test condition. The respective median threshold for the PD group was increased by 343% when compared to the control group. Second, when distal hand paths became less associated with proximal joint paths (right workspace), haptic acuity was reduced substantially in both groups. Third, sensitivity to hand trajectory curvature was not improved during active exploration in either group.

Conclusion/Significance

Our data demonstrate that PD is associated with a decreased acuity of the haptic sense, which may occur already at an early stage of the disease.     

Neuronal representation of optic flow experienced by unilaterally blinded flies on their mean walking trajectories

Asymmetries in the optic flow on both eyes may indicate an unintended turn of an animal and evoke compensatory optomotor responses. On a straight path in an evenly structured environment, the optic flow on both eyes is balanced corresponding to a state of optomotor equilibrium. When one eye is occluded an optomotor equilibrium is expected to be reached on a curved path provided that the translatory optic flow component is cancelled by a superimposed rotation. This hypothesis is tested by analysing how the HSE cell, a constituent element of the fly's optomotor system, represents optic flow in behavioural situations. The optic flow as seen on the average trajectory of freely walking monocular flies is reconstructed. This optic flow is used as stimulus of the HSE cell in electrophysiological experiments and as input of a model of the fly's optomotor system. The responses of the HSE cell and of the model fluctuate around the resting potential. On average, they are much smaller than the responses evoked by optic flow experienced on a straight path. These results corroborate the hypothesis that the mean trajectory of monocular flies corresponds to a path of optomotor equilibrium. Accepted: 29 February 2000     

Mantled Howler (<Emphasis Type="Italic">Alouatta palliata</Emphasis>) Arboreal Pathway Networks: Relative Impacts of Resource Availability and Forest Structure

Numerous studies have concluded that primates move about their environments in a nonrandom manner, frequently traveling between consecutive foraging sites along relatively straight-line paths. However, primates do not always take the most direct path between resources, and a number of species have been observed to travel repeatedly along a network of the same arboreal pathways. In this study, I used spatially explicit techniques to examine quantitatively what mantled howler monkey groups on Barro Colorado Island, Panama, accomplish by selecting nonlinear paths between resources and by repeatedly using the same paths within an arboreal network. Results show that chosen arboreal paths between sites where foraging occurred have higher levels of resource availability and canopy connectivity than comparable straight-line paths between the same sites. When comparing the relative importance of these factors, autologistic models of pathway choice indicate that though canopy connectivity is related to the location of repeatedly used arboreal pathway networks, the most statistically significant predictor is resource availability (both on a path and within a visual detection distance of a path). These results provide support for the hypothesis that repeated use of arboreal pathway networks aids in resource monitoring and acquisition. In addition, statistical models developed from 1 primary focal group’s travel patterns had high predictive value when employed to generate likely locations for arboreal pathways in the home ranges of 3 neighboring groups. This finding has important implications for studies of primate habitat use and seed dispersal, as it suggests that different groups consistently use similar characteristics when deciding on travel paths.     

Three-dimensional Particle Tracking Velocimetry for Turbulence Applications: Case of a Jet Flow

3D-PTV is a quantitative flow measurement technique that aims to track the Lagrangian paths of a set of particles in three dimensions using stereoscopic recording of image sequences. The basic components, features, constraints and optimization tips of a 3D-PTV topology consisting of a high-speed camera with a four-view splitter are described and discussed in this article. The technique is applied to the intermediate flow field (5 <x/d <25) of a circular jet at Re ≈ 7,000. Lagrangian flow features and turbulence quantities in an Eulerian frame are estimated around ten diameters downstream of the jet origin and at various radial distances from the jet core. Lagrangian properties include trajectory, velocity and acceleration of selected particles as well as curvature of the flow path, which are obtained from the Frenet-Serret equation. Estimation of the 3D velocity and turbulence fields around the jet core axis at a cross-plane located at ten diameters downstream of the jet is compared with literature, and the power spectrum of the large-scale streamwise velocity motions is obtained at various radial distances from the jet core.     

Parietal area VIP neuronal responses to heading stimuli are encoded in head-centered coordinates

The ventral intraparietal area (VIP) is a multimodal parietal area, where visual responses are brisk, directional, and typically selective for complex optic flow patterns. VIP thus could provide signals useful for visual estimation of heading (self-motion direction). A central problem in heading estimation is how observers compensate for eye velocity, which distorts the retinal motion cues upon which perception depends. To find out if VIP could be useful for heading, we measured its responses to simulated trajectories, both with and without eye movements. Our results showed that most VIP neurons very strongly signal heading direction. Furthermore, the tuning of most VIP neurons was remarkably stable in the presence of eye movements. This stability was such that the population of VIP neurons represented heading very nearly in head-centered coordinates. This makes VIP the most robust source of such signals yet described, with properties ideal for supporting perception.     

Harbor Seals (Phoca vitulina) Can Perceive Optic Flow under Water

Optic flow, the pattern of apparent motion elicited on the retina during movement, has been demonstrated to be widely used by animals living in the aerial habitat, whereas underwater optic flow has not been intensively studied so far. However optic flow would also provide aquatic animals with valuable information about their own movement relative to the environment; even under conditions in which vision is generally thought to be drastically impaired, e. g. in turbid waters. Here, we tested underwater optic flow perception for the first time in a semi-aquatic mammal, the harbor seal, by simulating a forward movement on a straight path through a cloud of dots on an underwater projection. The translatory motion pattern expanded radially out of a singular point along the direction of heading, the focus of expansion. We assessed the seal''s accuracy in determining the simulated heading in a task, in which the seal had to judge whether a cross superimposed on the flow field was deviating from or congruent with the actual focus of expansion. The seal perceived optic flow and determined deviations from the simulated heading with a threshold of 0.6 deg of visual angle. Optic flow is thus a source of information seals, fish and most likely aquatic species in general may rely on for e. g. controlling locomotion and orientation under water. This leads to the notion that optic flow seems to be a tool universally used by any moving organism possessing eyes.     

Non-invasive evaluation of face and motion perception in humans

The neural mechanisms for the perception of face and motion were studied using psychophysical threshold measurements, event-related potentials (ERPs), and functional magnetic resonance imaging (fMRI). A face-specific ERP component, N170, was recorded over the posterior temporal cortex. Removal of the high-spatial-frequency components of the face altered the perception of familiar faces significantly, and familiarity can facilitate the cortico-cortical processing of facial perceptions. Similarly, the high-spatial-frequency components of the face seemed to be crucial for the recognition of facial expressions. Aging and visuospatial impairments affected motion perception significantly. Two distinct components of motion ERPs, N170 and P200, were recorded over the parietal region. The former was related to horizontal motion perception while the latter reflected the perception of radial optic flow motion. The results of fMRI showed that horizontal movements of objects and radial optic flow motion were perceived differently in the V5/MT and superior parietal lobe. We conclude that an integrated approach can provide useful information on spatial and temporal processing of face and motion non-invasively.     

Flow measurements in a highly curved atherosclerotic coronary artery cast of man.

Flow visualization and wall pressure measurements were made in a polyurethane cast of a cadaver coronary artery with a significant "s" shaped reverse curvature. A sucrose solution was used to simulate the kinematic viscosity of blood, with flow rates in the physiologic range. Flow visualization demonstrated significant secondary flow patterns in the wall vicinity, which increased with increasing Reynolds number. Random dye dispersion was observed at a Reynolds number of about 400, but not at 200. Dye filament patterns in the transition between the first and second curved region were predominantly influenced by the second curved region at lower Reynolds numbers, and by the first curved region at higher Re. Local wall pressure measurements demonstrated a significant centrifugal effect with large radial pressure differences across the casting. Flow resistances for the casting were considerably greater than reference Poiseuille flow values, and increased further with pulsatile flow.     

Cardinal directions for visual optic flow.

As we move through our environment, the flow of deforming images on the retinae provides a rich source of information about the three-dimensional structure of the external world and how to navigate through it. Recent evidence from psychophysical [1] [2] [3] [4], electrophysiological [5] [6] [7] [8] [9] and imaging [10] [11] studies suggests that there are neurons in the primate visual system - in the medial superior temporal cortex - that are specialised to respond to this type of complex 'optic flow' motion. In principle, optic flow could be encoded by a small number of neural mechanisms tuned to 'cardinal directions', including radial and circular motion [12] [13]. There is little support for this idea at present, however, from either physiological [6] [7] or psychophysical [14] research. We have measured the sensitivity of human subjects for detection of motion and for discrimination of motion direction over a wide and densely sampled range of complex motions. Average sensitivity was higher for inward and outward radial movement and for both directions of rotation, consistent with the existence of detectors tuned to these four types of motion. Principle component analysis revealed two clear components, one for radial stimuli (outward and inward) and the other for circular stimuli (clockwise and counter-clock-wise). The results imply that the mechanisms that analyse optic flow in humans tend to be tuned to the cardinal axes of radial and rotational motion.     

Spicule form in calcareous sponges (Porifera: Calcarea). The principle of uniplanar curvature

A study of the shapes of secondary sagittal tri- and quadri-radiate spicules of the calcareous sponge Leuconia fistulosa has revealed that the paired rays appear kinked, with straight sections in between the kinks, when the spicules are viewed along the optic axis of the constituent calcite. The secondary sagittal spicules thus conform with monaxons and with per-regular and primary sagittal tri- and quadri-radiates, in general, in that whenever a ray is curved, the curvature is always in a single plane that includes the optic axis. The kinks are not necessarily distributed in symmetrical fashion along the paired rays. A mechanism is proposed whereby uniplanar curvature could be achieved.     

Path curvature discrimination: dependence on gaze direction and optical flow speed

Many experimental approaches to the control of steering rely on the tangent point (TP) as major source of information. The TP is a good candidate to control self-motion. It corresponds to a singular and salient point in the subject's visual field, and its location depends on the road geometry, the direction of self-motion relative to the road and the position of the driver on the road. However, the particular status of the TP in the optical flow, as a local minimum of flow speed, has often been left aside. We therefore assume that the TP is actually an optimal location in the dynamic optical array to perceive a change in the trajectory curvature. In this study, we evaluated the ability of human observers to detect variations in their path curvature from optical flow patterns, as a function of their gaze direction in a virtual environment. We simulated curvilinear self-motion parallel to a ground plane. Using random-dot optic flow stimuli of brief duration and a two-alternative forced-choice adaptive procedure, we determined path curvature discrimination thresholds, as a function of gaze direction. The discrimination thresholds are minimal for a gaze directed toward a local minimum of optical flow speed. A model based on Weber fraction of the foveal velocities (ΔV/V) correctly predicts the relationship between experimental thresholds and local flow velocities. This model was also tested for an optical flow computation integrating larger circular areas in central vision. Averaging the flow over five degrees leads to an even better fit of the model to experimental thresholds. We also found that the minimal optical flow speed direction corresponds to a maximal sensitivity of the visual system, as predicted by our model. The spontaneous gazing strategies observed during driving might thus correspond to an optimal selection of relevant information in the optical flow field.     

Moving muscle points provide accurate curved muscle paths in a model of the cervical spine

Muscle paths in musculoskeletal models have been modeled using several different methods; however, deformation of soft tissue with changes in posture is rarely accounted for, and often only the neutral posture is used to define a muscle path. The objective of this study was to model curved muscle paths in the cervical spine that take into consideration soft tissue deformation with changes in neck posture. Two subject-specific models were created from magnetic resonance images (MRI) in 5 different sagittal plane neck postures. Curved paths of flexor and extensor muscles were modeled using piecewise linear lines-of-action in two ways; (1) using fixed via points determined from muscle paths in the neutral posture and (2) using moving muscle points that moved relative to the bones determined from muscle paths in all 5 postures. Accuracy of each curved modeled muscle path was evaluated by an error metric, the distance from the anatomic (centroid) muscle path determined from the MRI. Error metric was compared among three modeled muscle path types (straight, fixed via and moving muscle point) using a repeated measures one-way ANOVA (α=0.05). Moving muscle point paths had 21% lower error metric than fixed via point paths over all 15 pairs of neck muscles examined over 5 postures (3.86 mm vs. 4.88 mm). This study highlights the importance of defining muscle paths in multiple postures in order to properly define the changing curvature of a muscle path due to soft tissue deformation with posture.     

Desert ants Cataglyphis fortis use self-induced optic flow to measure distances travelled

While foraging, desert ants of the genus Cataglyphis use a vector navigation (route integration) system for homing. Any vector navigation system requires that the animal is able to evaluate the angles steered and the distances travelled. Here we investigate whether the ants acquire the latter information by monitoring self-induced optic flow. To answer this question, the animals were trained and tested within perspex channels in which patterns were presented underneath a transparent walking platform. The patterns could be moved at different velocities (up to > 0.5 the ant's walking speed) in the same or in the opposite direction relative to the direction in which the animal walked. Experimental manipulations of the optic flow influenced the ant's homing distances (Figs. 2 and 4). Distance estimation depends on the speed of self-induced image motion rather than on the contrast frequency, indicating that the motion sensitive mechanism involved is different from mechanisms mediating the optomotor response. Experiments in which the ants walked on a featureless floor, or in which they wore eye covers (Fig. 6), show that they are able also to use additional (probably kinesthetic) cues for assessing their travel distance. Hence, even though optic flow cues are not the only ones used by the ants, the experiments show that ants are obviously able to exploit such cues for estimation of travel distance.     

Computing the direction of heading from affine image flow

Observers moving through a three-dimensional environment can use optic flow to determine their direction of heading. Existing heading algorithms use cartesian flow fields in which image flow is the displacement of image features over time. I explore a heading algorithm that uses affine flow instead. The affine flow at an image feature is its displacement modulo an affine transformation defined by its neighborhood. Modeling the observer's instantaneous motion by a translation and a rotation about an axis through its eye, affine flow is tangent to the translational field lines on the observer's viewing sphere. These field lines form a radial flow field whose center is the direction of heading. The affine flow heading algorithm has characteristics that can be used to determine whether the human visual system relies on it. The algorithm is immune to observer rotation and arbitrary affine transformations of its input images; its accuracy improves with increasing variation in environmental depth; and it cannot recover heading in an environment consisting of a single plane because affine flow vanishes in this case. Translational field lines can also be approximated through differential cartesian motion. I compare the performance of heading algorithms based on affine flow, differential cartesian flow, and least-squares search.     

Optic flow drives human visuo-locomotor adaptation

Two strategies can guide walking to a stationary goal: (1) the optic-flow strategy, in which one aligns the direction of locomotion or "heading" specified by optic flow with the visual goal; and (2) the egocentric-direction strategy, in which one aligns the locomotor axis with the perceived egocentric direction of the goal and in which error results in optical target drift. Optic flow appears to dominate steering control in richly structured visual environments, whereas the egocentric- direction strategy prevails in visually sparse environments. Here we determine whether optic flow also drives visuo-locomotor adaptation in visually structured environments. Participants adapted to walking with the virtual-heading direction displaced 10 degrees to the right of the actual walking direction and were then tested with a normally aligned heading. Two environments, one visually structured and one visually sparse, were crossed in adaptation and test phases. Adaptation of the walking path was more rapid and complete in the structured environment; the negative aftereffect on path deviation was twice that in the sparse environment, indicating that optic flow contributes over and above target drift alone. Optic flow thus plays a central role in both online control of walking and adaptation of the visuo-locomotor mapping.     

生态系统中流动路径多样性的测度

路径作为生态网络组成的一个基本单元,是生态系统中物质、能量流动的通道。生态系统的结构和功能是通过生态系统中不同路径上能量和物质的流动来实现的。路径多样性是反映生态系统营养结构多样性的一个重要侧面。本文在定义基路径和等价基路径的基础上,给出了生态系统中流动路径多样性的测度     

On the barn owl’s visual pre-attack behavior: I. Structure of head movements and motion patterns

Barn owls exhibit a rich repertoire of head movements before taking off for prey capture. These movements occur mainly at light levels that allow for the visual detection of prey. To investigate these movements and their functional relevance, we filmed the pre-attack behavior of barn owls. Off-line image analysis enabled reconstruction of all six degrees of freedom of head movements. Three categories of head movements were observed: fixations, head translations and head rotations. The observed rotations contained a translational component. Head rotations did not follow Listing’s law, but could be well described by a second-order surface, which indicated that they are in close agreement with Donder’s law. Head translations did not contain any significant rotational components. Translations were further segmented into straight-line and curved paths. Translations along an axis perpendicular to the line of sight were similar to peering movements observed in other animals. We suggest that these basic motion elements (fixations, head rotations, translations along a straight line, and translation along a curved trajectory) may be combined to form longer and more complex behavior. We speculate that these head movements mainly underlie estimation of distance during prey capture.     

Visual Acceleration Perception for Simple and Complex Motion Patterns

Humans are able to judge whether a target is accelerating in many viewing contexts, but it is an open question how the motion pattern per se affects visual acceleration perception. We measured acceleration and deceleration detection using patterns of random dots with horizontal (simpler) or radial motion (more visually complex). The results suggest that we detect acceleration better when viewing radial optic flow than horizontal translation. However, the direction within each type of pattern has no effect on performance and observers detect acceleration and deceleration similarly within each condition. We conclude that sensitivity to the presence of acceleration is generally higher for more complex patterns, regardless of the direction within each type of pattern or the sign of acceleration.