Action without awareness: reaching to an object you do not remember seeing

Background

Previous work by our group has shown that the scaling of reach trajectories to target size is independent of obligatory awareness of that target property and that “action without awareness” can persist for up to 2000 ms of visual delay. In the present investigation we sought to determine if the ability to scale reaching trajectories to target size following a delay is related to the pre-computing of movement parameters during initial stimulus presentation or the maintenance of a sensory (i.e., visual) representation for on-demand response parameterization.

Methodology/Principal Findings

Participants completed immediate or delayed (i.e., 2000 ms) perceptual reports and reaching responses to different sized targets under non-masked and masked target conditions. For the reaching task, the limb associated with a trial (i.e., left or right) was not specified until the time of response cuing: a manipulation that prevented participants from pre-computing the effector-related parameters of their response. In terms of the immediate and delayed perceptual tasks, target size was accurately reported during non-masked trials; however, for masked trials only a chance level of accuracy was observed. For the immediate and delayed reaching tasks, movement time as well as other temporal kinematic measures (e.g., times to peak acceleration, velocity and deceleration) increased in relation to decreasing target size across non-masked and masked trials.

Conclusions/Significance

Our results demonstrate that speed-accuracy relations were observed regardless of whether participants were aware (i.e., non-masked trials) or unaware (i.e., masked trials) of target size. Moreover, the equivalent scaling of immediate and delayed reaches during masked trials indicates that a persistent sensory-based representation supports the unconscious and metrical scaling of memory-guided reaching.     

Activity of descending contralateral movement detector neurons and collision avoidance behaviour in response to head-on visual stimuli in locusts

We recorded the activity of the right and left descending contralateral movement detectors responding to 10-cm (small) or 20-cm (large) computer-generated spheres approaching along different trajectories in the locust's frontal field of view. In separate experiments we examined the steering responses of tethered flying locusts to identical stimuli. The descending contralateral movement detectors were more sensitive to variations in target trajectory in the horizontal plane than in the vertical plane. Descending contralateral movement detector activity was related to target trajectory and to target size and was most sensitive to small objects converging on a direct collision course from above and to one side. Small objects failed to induce collision avoidance manoeuvres whereas large objects produced reliable collision avoidance responses. Large targets approaching along a converging trajectory produced steering responses that were either away from or toward the side of approach of the object, whereas targets approaching along trajectories that were offset from the locust's mid-longitudinal body axis primarily evoked responses away from the target. We detected no differences in the discharge properties of the descending contralateral movement detector pair that could account for the different collision avoidance behaviours evoked by varying the target size and trajectories. We suggest that descending contralateral movement detector properties are better suited to predator evasion than collision avoidance.     

A Real-Time Brain-Machine Interface Combining Motor Target and Trajectory Intent Using an Optimal Feedback Control Design

Real-time brain-machine interfaces (BMI) have focused on either estimating the continuous movement trajectory or target intent. However, natural movement often incorporates both. Additionally, BMIs can be modeled as a feedback control system in which the subject modulates the neural activity to move the prosthetic device towards a desired target while receiving real-time sensory feedback of the state of the movement. We develop a novel real-time BMI using an optimal feedback control design that jointly estimates the movement target and trajectory of monkeys in two stages. First, the target is decoded from neural spiking activity before movement initiation. Second, the trajectory is decoded by combining the decoded target with the peri-movement spiking activity using an optimal feedback control design. This design exploits a recursive Bayesian decoder that uses an optimal feedback control model of the sensorimotor system to take into account the intended target location and the sensory feedback in its trajectory estimation from spiking activity. The real-time BMI processes the spiking activity directly using point process modeling. We implement the BMI in experiments consisting of an instructed-delay center-out task in which monkeys are presented with a target location on the screen during a delay period and then have to move a cursor to it without touching the incorrect targets. We show that the two-stage BMI performs more accurately than either stage alone. Correct target prediction can compensate for inaccurate trajectory estimation and vice versa. The optimal feedback control design also results in trajectories that are smoother and have lower estimation error. The two-stage decoder also performs better than linear regression approaches in offline cross-validation analyses. Our results demonstrate the advantage of a BMI design that jointly estimates the target and trajectory of movement and more closely mimics the sensorimotor control system.     

Measuring effects of linear obstacles on wildlife movements: accounting for the relationship between step length and crossing probability

Animal movements in the landscape are influenced by linear features such as rivers, roads and power lines. Prior studies have investigated how linear features, particularly roads, affect movement rates by comparing animal's movement rate measured as step lengths (i.e., the distance between consecutive observations such as GPS locations) before, during and after crossing of a linear feature. The null hypothesis has been that the length of crossing steps should not differ from other steps, and a deviation from this, mainly that steps are longer during crossing, has been taken as support for a disturbance effect of the linear feature. However, based on the simple relationship between the length of a step and its probability to cross a linear feature, we claim that this assumption is inappropriate to test for behavioural responses to linear features. The probability is related to the proportion of the total length of the trajectory (i.e., the path of movement) a step constitutes. Consequently, care should be taken when formulating hypotheses about how animal moves in relation to linear features in the landscape. Statistical tests should be set up with respect to the expected length based on the distribution of step lengths in the trajectory. We propose two methods that accounts for the bias in crossing frequency that is caused by step lengths, and illustrates their applications by using simulated animal trajectories as well as empirical data on reindeer in an area with a power line.     

Drifting towards a diffuse control model of exploratory motor learning: A comparison of global and within-trial performance measures

 Accurate measurement is crucial for understanding the processes that underlie exploratory patterns in motor learning. Accordingly, measures of learning should be sensitive to the changes that take place during skill acquisition. Most studies, however, use trial-based global measures that assess performance but do not actually measure gradual changes taking place within trials. The present study attempted to remedy this shortcoming by analysing a visual adaptation task, and comparing traditional global measures of learning with new, within-trial measures. Movement time was the only global measure sensitive to changes in the movement trajectory during learning. Three new measures were expected to reveal changes to the movement trajectory that are associated with learning: (i) the length of runs, (ii) change of trajectory angle in relation to the target, and (iii) drift (change in distance from the target). All three measures were sensitive to learning and indicated a gradual straightening of the movement trajectories over trials. Furthermore, three different methods to partition trajectories into segments were examined. The new within-trial measures, irrespective of partitioning method, prove promising for the development of a diffuse control model of exploratory learning. Received: 5 February 2001 / Accepted in revised form: 16 January 2002     

Analysis of kinematic invariances of multijoint reaching movement

 There is a no unique relationship between the trajectory of the hand, represented in cartesian or extrinsic space, and its trajectory in joint angle or intrinsic space in the general condition of joint redundancy. The goal of this work is to analyze the relation between planning the trajectory of a multijoint movement in these two coordinate systems. We show that the cartesian trajectory can be planned based on the task parameters (target coordinates, etc.) prior to and independently of angular trajectories. Angular time profiles are calculated from the cartesian trajectory to serve as a basis for muscle control commands. A unified differential equation that allows planning trajectories in cartesian and angular spaces simultaneously is proposed. Due to joint redundancy, each cartesian trajectory corresponds to a family of angular trajectories which can account for the substantial variability of the latter. A set of strategies for multijoint motor control following from this model is considered; one of them coincides with the frog wiping reflex model and resolves the kinematic inverse problem without inversion. The model trajectories exhibit certain properties observed in human multijoint reaching movements such as movement equifinality, straight end-point paths, bell-shaped tangential velocity profiles, speed-sensitive and speed-insensitive movement strategies, peculiarities of the response to double-step targets, and variations of angular trajectory without variations of the limb end-point trajectory in cartesian space. In humans, those properties are almost independent of limb configuration, target location, movement duration, and load. In the model, these properties are invariant to an affine transform of cartesian space. This implies that these properties are not a special goal of the motor control system but emerge from movement kinematics that reflect limb geometry, dynamics, and elementary principles of motor control used in planning. All the results are given analytically and, in order to compare the model with experimental results, by computer simulations. Received: 6 April 1994/Accepted in revised form: 25 April 1995     

Mechanisms underlying the generation of averaged modified trajectories

In this work we have studied what mechanisms might possibly underlie arm trajectory modification when reaching toward visual targets. The double-step target displacement paradigm was used with inter-stimulus intervals (ISIs) in the range of 10-300 ms. For short ISIs, a high percentage of the movements were found to be initially directed in between the first and second target locations (averaged trajectories). The initial direction of motion was found to depend on the target configuration, and on : the time difference between the presentation of the second stimulus and movement onset. To account for the kinematic features of the averaged trajectories two modification schemes were compared: the superposition scheme and the abort-replan scheme. According to the superposition scheme, the modified trajectories result from the vectorial addition of two elemental motions: one for moving between the initial hand position and an intermediate location, and a second one for moving between that intermediate location and the final target. According to the abort-replan scheme, the initial plan for moving toward the intermediate location is aborted and smoothly replaced by a new plan for moving from the hand position at the time the trajectory is modified to the final target location. In both tested schemes we hypothesized that due to the quick displacement of the stimulus, the internally specified intermediate goal might be influenced by both stimuli and may be different from the location of the first stimulus. It was found that the statistically most successful model in accounting for the measured data is based on the superposition scheme. It is suggested that superposition of simple independent elemental motions might be a general principle for the generation of modified motions, which allows for efficient, parallel planning. For increasing values of the inferred locations of the intermediate targets were found to gradually shift from the first toward the second target locations along a path that curved toward the initial hand position. These inferred locations show a strong resemblance to the intermediate locations of saccades generated in a similar double-step paradigm. These similarities in the specification of target locations used in the generation of eye and hand movements may serve to simplify visuomotor integration. Received: 22 June 1994 / Accepted in revised form: 15 September 1994     

Statistical decision theory and trade-offs in the control of motor response

We present a novel approach to the modeling of motor responses based on statistical decision theory. We begin with the hypothesis that subjects are ideal motion planners who choose movement trajectories to minimize expected loss. We derive predictions of the hypothesis for movement in environments where contact with specified regions carries rewards or penalties. The model predicts shifts in a subject's aiming point in response to changes in the reward and penalty structure of the environment and with changes in the subject's uncertainty in carrying out planned movements. We tested some of these predictions in an experiment where subjects were rewarded if they succeeded in touching a target region on a computer screen within a specified time limit. Near the target was a penalty region which, if touched, resulted in a penalty. We varied distance between the penalty region and the target and the cost of hitting the penalty region. Subjects shift their mean points of contact with the computer screen in response to changes in penalties and location of the penalty region relative to the target region in qualitative agreement with the predictions of the hypothesis. Thus, movement planning takes into account extrinsic costs and the subject's own motor uncertainty.     

Formation and control of optimal trajectory in human multijoint arm movement

In this paper, we study trajectory planning and control in voluntary, human arm movements. When a hand is moved to a target, the central nervous system must select one specific trajectory among an infinite number of possible trajectories that lead to the target position. First, we discuss what criterion is adopted for trajectory determination. Several researchers measured the hand trajectories of skilled movements and found common invariant features. For example, when moving the hand between a pair of targets, subjects tended to generate roughly straight hand paths with bell-shaped speed profiles. On the basis of these observations and dynamic optimization theory, we propose a mathematical model which accounts for formation of hand trajectories. This model is formulated by defining an objective function, a measure of performance for any possible movement: square of the rate of change of torque integrated over the entire movement. That is, the objective function CT is defined as follows: (formula; see text) We overcome this difficult by developing an iterative scheme, with which the optimal trajectory and the associated motor command are simultaneously computed. To evaluate our model, human hand trajectories were experimentally measured under various behavioral situations. These results supported the idea that the human hand trajectory is planned and controlled in accordance with the minimum torque-change criterion.     

Dynamic optimization of the sit-to-stand movement

The purpose of this study was to clarify criteria that can predict trajectories during the sit-to-stand movement. In particular, the minimum jerk and minimum torque-change models were examined. Three patterns of sit-to-stand movement from a chair, i.e., upright, natural, and leaning forward, were measured in five young participants using a 3-D motion analysis device (200 Hz). The trajectory of the center of mass and its smoothness were examined, and the optimal trajectories predicted by both models were evaluated. Trajectories of the center of mass predicted by the minimum torque-change model, rather than the minimum jerk model, resembled the measured movements in all rising movement patterns. The upright pattern required greater extension torque of the knee and ankle joints at the instant of seat-off. The leaning-forward pattern required greater extension hip torque and higher movement cost than the natural and upright patterns. These results indicate that the natural sit-to-stand movement might be a result of dynamic optimization.     

Precardiac cell migration: fibronectin localization at mesoderm-endoderm interface during directional movement

The pathway of directional movement of chick precardiac mesoderm cells was studied by indirect immunofluorescence and by scanning electron microscopy. Directional movement of the precardiac cells begins at stage 6 from the lateral sides of the embryo at the level of Hensen's node. The cells move anteriorly in an arc to the embryo's midline. By stage 8 the cells arrive at the lateral sides of the anterior intestinal portal and movement ceases. The interval of this directional movement is approximately 10 hr. During migration the precardiac cells are in close association with the underlying endoderm. As migration proceeds, the cells encounter increasing amounts of fibrils in the substratum at the mesoderm-endoderm interface. Concomitant with increasing fibril formation there is an increase in fibronectin (FN) in the heart-forming region. During stage 5 FN first appears in the lateral heart-forming regions and increases in amount during the period of cell migration. By stage 7 a concentration difference of FN is apparent in the lateral regions with more FN cephalad and decreasing amounts caudad. At stages 7 and 8 large amounts of extracellular FN-associated fibrils are observed at the lateral sides of the anterior intestinal portal where the cells stop moving. The precardiac cells moving into this region are oriented perpendicular to the anterior intestinal portal and in close association with these fibrils. There is no evidence that the fibrillar meshwork forming the substratum of the precardiac mesoderm cells is physically oriented as a guide for directional movement. The correlations between FN distribution at the mesoderm-endoderm interface and directional cell movement suggest that the precardiac cells may migrate by haptotaxis, i.e., by moving along the substratum toward areas of greater adhesiveness.     

Plasticity and ontogenetic drift of biomass allocation in response to above- and below-ground resource availabilities in perennial herbs: a case study of <Emphasis Type="Italic">Alternanthera philoxeroides</Emphasis>

Changes in plant biomass allocation in response to varying resource availabilities may result from ontogenetic drift caused by allometric growth (i.e., apparent plasticity), a true adjustment of ontogenetic trajectories (true plasticity) or both (complex plasticity). Given that the root allocation of annual species usually decreases during the growth, the developmentally explicit model predicts that annual herbs will exhibit true plasticity in root allocation under above-ground resource limitation and apparent plasticity for moderate stress of below-ground resource. For perennial species, the root allocation of which increases during growth, the reverse patterns would be expected. In this study, we tested the developmentally explicit model with a perennial weed, Alternanthera philoxeroides (Mart.) Griseb. We report its adaptive changes and ontogenetic drift of root allocation in response to different resource levels (i.e., light, water and nutrient availability) by comparing root allocation on both an age and a size basis. The root allocation of A. philoxeroides increased with the size (i.e., ontogenetic drift) during the growth, and exhibited significant changes in response to different resource availabilities. Furthermore, the root allocation in response to water or nutrient availability exhibited typical complex plasticity, while the light stress only slowed down the growth, with the ontogenetic trajectory unchanged (apparent plasticity). The contrasting responses to above-ground and below-ground stresses were consistent with the prediction of the developmentally explicit model.     

Action evaluation is modulated dominantly by internal sensorimotor information and partly by noncausal external cue

Previous studies demonstrated that human motor actions are not always monitored by perceptual awareness and that implicit motor control plays a key role in performing actions. In addition, appropriate evaluation of our own motor behavior is vital for human life. Here we combined a reaching task with a visual backward masking paradigm to induce an implicit motor response that is congruent or incongruent with the visual perception. We used this to investigate (i) how we evaluate such implicit motor response that could be inconsistent with perceptual awareness and (ii) the possible contributions of reaching error, external visual cues, and internal sensorimotor information to this evaluation. Participants were instructed, after each trial, to rate their own reaching performance on a 5-point scale (i.e., smooth-clumsy). They also needed to identify a color presented at a fixation point that could be changed just after the reaching start. The color was linked to the prime-mask congruency (i.e., congruent-green, incongruent-blue) in the practice phase, and then inconsistent pairs (congruent-blue or incongruent-green) were introduced in the test phase. We found early trajectory deviations induced by the invisible prime stimulus, and such implicit motor responses are significantly correlated with the action evaluation score. The results suggest the "conscious" action evaluation is properly monitoring online sensory outcomes derived by implicit motor control. Furthermore, statistical path analyses showed that internal sensorimotor information from the motor behavior modulated by the invisible prime was the predominant cue for the action evaluation, while the color-cue association learned in the practice phase in some cases biases the action evaluation in the test phase.     

Reproduction of tracking movements and target positioning of the forearm in humans in the absence of visual control

In 17 healthy volunteers, we studied movements of the forearm, which included episodes of positioning on the target level. The trajectory of the non-ballistic (relatively slow) movement looked like a double trapezium (flexion of the elbow joint from the state of full extension, 0 deg, positioning on the 50 deg level, further flexion to the limit angle of 100 deg, and a similar reverse sequence). The command trajectory and the trajectory of the realized movement were visualized with movements of cursors on a monitor in time/joint angle coordinates. We compared parameters of the tracking movements (in the presence of visual feedback) and their blindfold reproduction (with the complete absence of visual control). It was found that blindfold reproduction movements differ from sample tracking movements and their reproduction with partial limitation of visual control [16] in higher peak velocities and shorter durations, i.e., a trend toward conversion of such movements into ballistic ones was observed. Under conditions of elimination of visual control, movements that led to positioning were mostly hypermetric, i.e., positioning was usually accompanied by positive systematic errors (whose sign coincided with the direction of the preceding movement phase). The mean intragroup value of the systematic error of the first positioning (after flexion to the target level) was +6.73 ± 1.15 deg, while the respective mean for the second positioning (after extension to the same level) was +4.00 ± 1.31 deg. The nonlinear properties of stretch receptors of muscles whose activity provides the formation of a proprioceptive estimate of the joint angle are considered the crucial reason for systematic errors of blindfold positioning.Neirofiziologiya/Neurophysiology, Vol. 36, Nos. 5/6, pp. 393–404, September–December, 2004.This revised version was published online in April 2005 with a corrected cover date and copyright year.     

Digit ratio predicts sense of direction in women

The relative length of the second-to-fourth digits (2D:4D) has been linked with prenatal androgen in humans. The 2D:4D is sexually dimorphic, with lower values in males than females, and appears to correlate with diverse measures of behavior. However, the relationship between digit ratio and cognition, and spatial cognition in particular, has produced mixed results. In the present study, we hypothesized that spatial tasks separating cue conditions that either favored female or male strategies would examine this structure-function correlation with greater precision. Previous work suggests that males are better in the use of directional cues than females. In the present study, participants learned a target location in a virtual landscape environment, in conditions that contained either all directional (i.e., distant or compass bearing) cues, or all positional (i.e., local, small objects) cues. After a short delay, participants navigated back to the target location from a novel starting location. Males had higher accuracy in initial search direction than females in environments with all directional cues. Lower digit ratio was correlated with higher accuracy of initial search direction in females in environments with all directional cues. Mental rotation scores did not correlate with digit ratio in either males or females. These results demonstrate for the first time that a sex difference in the use of directional cues, i.e., the sense of direction, is associated with more male-like digit ratio.     

Light-triggered action potentials in the liverwort Conocephalum conicum

The response to light of a liverwort, Conocephalum conicum L., measured as a change in the resting potential, consists of two stages. The first stage is a slight depolarization dependent on light intensity. This plays the role of a generator potential (GP) which induces the second stage - an action potential of the all-or-none character. Action potentials induced by light and by electrical stimuli have the same properties, i.e. identical time course, propagation velocity, and refractory periods. A summation occurs of sub threshold light stimuli and of light and electrical stimuli. The presence of 5⋅10-⁻⁶ M DCMU cancelled the light response and blocked - by inhibition of the electron transport chain - the mechanism leading to GP generation. However, this effect did not produce any change in the response to electrical stimuli.     

A procedure to quantify the feed intake response of growing pigs to perturbations

Improving robustness of farm animals is one of the goals in breeding programmes. However, robustness is a complex trait and not measurable directly. The objective of this study was to quantify and characterize (elements of) robustness in growing pigs. Robustness can be analysed by examining the animal’s response to perturbations. Although the origin of perturbations may not be known, their effect on animal performance can be observed, for example, through changes in voluntary feed intake. A generic model and data analysis procedure was developed (1) to estimate the target trajectory of feed intake, which is the amount of feed that a pig desires to eat when it is not facing any perturbations; (2) to detect potential perturbations, which are deviations of feed intake from the estimated target trajectory; and (3) to characterize and quantify the response of the growing pigs to the perturbations using voluntary feed intake as response criterion. The response of a pig to a perturbation is characterized by four parameters. The start and end times of the perturbation are ‘imposed’ by the perturbing factor, while two other parameters describe the resistance and resilience potential of the pig. One of these describes the immediate reduction in daily feed intake at the start of the perturbation (i.e., a ‘resistance’ trait) while another parameter describes the capacity of the pig to adapt to the perturbation through compensatory feed intake to rejoin the target trajectory of feed intake (i.e., a ‘resilience’ trait). The procedure has been employed successfully to identify the target trajectory of feed intake in growing pigs and to quantify the pig’s response to a perturbation.     

Characterization of the cortisol response to incremental exercise in physically active young men

This study examined the cortisol response to incremental exercise; specifically to see if there was an increase in blood cortisol levels at low intensity exercise (i.e., < 60% VO2 intensity threshold) and determine whether a linear relationship existed between the blood cortisol responses and exercise of increasing workloads (i.e., intensity). Healthy, physically active young men (n = 11) completed exercise tests involving progressive workload stages (3 min) to determine peak oxygen uptake responses (VO2). Blood specimens were collected at rest and at the end of each stage and analyzed for cortisol. Results showed cortisol was significantly increased from resting levels at the end of the first exercise stage (80 W; 41.9 +/- 5.4% peak VO2) and remained significantly elevated from rest until the exercise ended. Interestingly, however, the cortisol concentrations observed at 80 W through 200 W did not significantly differ from one another. Thereafter, during the final two stages of exercise the cortisol concentrations increased further (p < 0.01). The subjects exceeded their individual lactate thresholds over these last two stages of exercise. Regression modeling to characterize the cortisol response resulted in significant regression coefficients (r = 0.415 [linear] and r = 0.655 [3rd order polynominal], respectively; p < 0.05). Comparative testing (Hotelling test) between the two regression coefficents revealed the polynominal model (sigmoidal curve) was the significantly stronger of the two (p = 0.05). In conclusion, the present findings refute the concept that low intensity exercise will not provoke a significant change in blood cortisol levels and suggest the response to incremental exercise involving increasing exercise workloads (i.e., intensities) are not entirely linear in nature. Specifically, a sigmoid curve more highly accurately characterizes the cortisol response to such exercise.     

Classes and narrowing selectivity of olfactory receptor neurons of Xenopus laevis tadpoles

In olfactory receptor neurons (ORNs) of aquatic animals amino acids have been shown to be potent stimuli. Here we report on calcium imaging experiments in slices of the olfactory mucosa of Xenopus laevis tadpoles. We were able to determine the response profiles of 283 ORNs to 19 amino acids, where one profile comprises the responses of one ORN to 19 amino acids. 204 out of the 283 response profiles differed from each other. 36 response spectra occurred more than once, i.e., there were 36 classes of ORNs identically responding to the 19 amino acids. The number of ORNs that formed a class ranged from 2 to 13. Shape and duration of amino acid-elicited [Ca2+]i transients showed a high degree of similarity upon repeated stimulation with the same amino acid. Different amino acids, however, in some cases led to clearly distinguishable calcium responses in individual ORNs. Furthermore, ORNs clearly appeared to gain selectivity over time, i.e., ORNs of later developmental stages responded to less amino acids than ORNs of earlier stages. We discuss the narrowing of ORN selectivity over stages in the context of expression of olfactory receptors.     

Inter-Individual Variability of Stone Marten Behavioral Responses to a Highway

Efforts to reduce the negative impacts of roads on wildlife may be hindered if individuals within the population vary widely in their responses to roads and mitigation strategies ignore this variability. This knowledge is particularly important for medium-sized carnivores as they are vulnerable to road mortality, while also known to use available road passages (e.g., drainage culverts) for safely crossing highways. Our goal in this study was to assess whether this apparently contradictory pattern of high road-kill numbers associated with a regular use of road passages is attributable to the variation in behavioral responses toward the highway between individuals. We investigated the responses of seven radio-tracked stone martens (Martes foina) to a highway by measuring their utilization distribution, response turning angles and highway crossing patterns. We compared the observed responses to simulated movement parameterized by the observed space use and movement characteristics of each individual, but naïve to the presence of the highway. Our results suggested that martens demonstrate a diversity of responses to the highway, including attraction, indifference, or avoidance. Martens also varied in their highway crossing patterns, with some crossing repeatedly at the same location (often coincident with highway passages). We suspect that the response variability derives from the individual''s familiarity of the landscape, including their awareness of highway passage locations. Because of these variable yet potentially attributable responses, we support the use of exclusionary fencing to guide transient (e.g., dispersers) individuals to existing passages to reduce the road-kill risk.