Jason Friedman

The Neural Processes Underlying Self-Agency

Self-agency (SA) is the individuals perception that an action is the consequence of his/her own intention. The neural networks underlying SA are not well understood. We carried out a novel, ecologically valid, virtual-reality experiment using blood oxygen level-dependent functional magnetic resonance imaging (fMRI) where SA could be modulated in real-time while subjects performed voluntary finger movements. Behavioral testing was also performed to assess the explicit judgment of SA. Twenty healthy volunteers completed the experiment. Results of the behavioral testing demonstrated paradigm validity along with the identification of a bias that led subjects to over- or underestimate the amount of control they had. The fMRI experiment identified 2 discrete networks. These leading and lagging networks likely represent a spatial and temporal flow of information, with the leading network serving the role of mismatch detection and the lagging network receiving this information and mediating its elevation to conscious awareness, giving rise to SA.

Task-dependent selection of grasp kinematics and stiffness in human object manipulation.

Object manipulation with the hand is a complex task. The task has redundancies at many levels, allowing many possibilities for the selection of grasp points, the orientation and posture of the hand, the forces to be applied at each fingertip and the impedance properties of the hand. Despite this inherent complexity, humans perform object manipulation nearly effortlessly. This article presents experimental findings of how humans grasp and manipulate objects, and examines the compatibility of grasps selected for specific tasks. This is accomplished by looking at the velocity transmission and force transmission ellipsoids, which represent the transmission ratios of the corresponding quantity from the joints to the object, as well as the stiffness ellipsoid which represents the directional stiffness of the grasp. These ellipsoids allow visualization of the grasp Jacobian and grasp stiffness matrices. The results show that the orientation of the ellipsoids can be related to salient task requirements.

Prehension synergies and control with referent hand configurations.

We used the framework of the equilibrium-point hypothesis (in its updated form based on the notion of referent configuration) to investigate the multi-digit synergies at two levels of a hypothetical hierarchy involved in prehensile actions. Synergies were analyzed at the thumb–virtual finger (VF) level (VF is an imaginary digit with the mechanical action equivalent to that of the four actual fingers) and at the individual finger level. The subjects performed very quick vertical movements of a handle into a target. A load could be attached off-center to provide a pronation or supination torque. In a few trials, the handle was unexpectedly fixed to the table and the digits slipped off the sensors. In such trials, the hand stopped at a higher vertical position and rotated into pronation or supination depending on the expected torque. The aperture showed non-monotonic changes with a large, fast decrease and further increase, ending up with a smaller distance between the thumb and the fingers as compared to unperturbed trials. Multi-digit synergies were quantified using indices of co-variation between digit forces and moments of force across unperturbed trials. Prior to the lifting action, high synergy indices were observed at the individual finger level while modest indices were observed at the thumb–VF level. During the lifting action, the synergies at the individual finger level disappeared while the synergy indices became higher at the thumb–VF level. The results support the basic premise that, within a given task, setting a referent configuration may be described with a few referent values of variables that influence the equilibrium state, to which the system is attracted. Moreover, the referent configuration hypothesis can help interpret the data related to the trade-off between synergies at different hierarchical levels.

The sources of two components of variance: an example of multifinger cyclic force production tasks at different frequencies

In a multifinger cyclic force production task, the finger force variance measured across trials can be decomposed into two components, one that affects the combined force output (“bad variance”) and one that does not (“good variance”). Previous studies have found similar time patterns of “bad variance” and force rate leading to an approximately linear relationship between them. Based on this finding and a recently developed model of multifinger force production, we expected the “bad variance” during cyclic force production to increase monotonically with the rate of force change, both within a cycle and across trials at different frequencies. Alternatively, “bad variance” could show a dependence on task frequency, not on actual force derivative values. Healthy subjects were required to produce cyclic force patterns to prescribed targets by pressing on unidimensional force sensors, at a frequency set by a metronome. The task was performed with only the index finger, and with all four fingers. In the task with all four fingers, the “good variance” increased approximately linearly with an increase in the force magnitude. The “bad variance” showed within-a-cycle modulation similar to that of the force rate. However, an increase in the frequency did not lead to an increase in the “bad variance” that could be expected based on the natural relationships between action frequency and the rate of force change modulation. The results have been interpreted in the framework of an earlier model of multifinger force production where “bad variance” is a result of variance of the timing parameter. The unexpected lack of modulation of the “bad variance” with frequency suggests a drop in variance of the timing parameter with increased frequency. This mechanism may serve to maintain a constant acceptable level of variance under different conditions.

Viewing and feeling touch modulates hand position for reaching

Action requires knowledge of our body location in space. Here we asked if interactions with the external world prior to a reaching action influence how visual location information is used. We investigated if the temporal synchrony between viewing and feeling touch modulates the integration of visual and proprioceptive body location information for action. We manipulated the synchrony between viewing and feeling touch in the Rubber Hand Illusion paradigm prior to participants performing a ballistic reaching task to a visually specified target. When synchronous touch was given, reaching trajectories were significantly shifted compared to asynchronous touch. The direction of this shift suggests that touch influences the encoding of hand position for action. On the basis of this data and previous findings, we propose that the brain uses correlated cues from passive touch and vision to update its own position for action and experience of self-location.

Intrinsic joint kinematic planning. I: Reassessing the Listing's law constraint in the control of three-dimensional arm movements

This study tested the validity of the assumption that intrinsic kinematic constraints, such as Listing’s law, can account for the geometric features of three-dimensional arm movements. In principle, if the arm joints follow a Listing’s constraint, the hand paths may be predicted. Four individuals performed ‘extended arm’, ‘radial’, ‘frontal plane’, and ‘random mixed’ movements to visual targets to test Listing’s law assumption. Three-dimensional rotation vectors of the upper arm and forearm were calculated from three-dimensional marker data. Data fitting techniques were used to test Donders’ and Listing’s laws. The coefficient values obtained from fitting rotation vectors to the surfaces described by a second-order equation were analyzed. The results showed that the coefficients that represent curvature and twist of the surfaces were often not significantly different from zero, particularly not during randomly mixed and extended arm movements. These coefficients for forearm rotations were larger compared to those for the upper arm segment rotations. The mean thickness of the rotation surfaces ranged between ≈1.7° and 4.7° for the rotation vectors of the upper arm segment and ≈2.6° and 7.5° for those of the forearm. During frontal plane movements, forearm rotations showed large twist scores while upper arm segment rotations showed large curvatures, although the thickness of the surfaces remained low. The curvatures, but not the thicknesses of the surfaces, were larger for large versus small amplitude radial movements. In conclusion, when examining the surfaces obtained for the different movement types, the rotation vectors may lie within manifolds that are anywhere between curved or twisted manifolds. However, a two-dimensional thick surface may roughly represent a global arm constraint. Our findings suggest that Listing’s law is implemented for some types of arm movement, such as pointing to targets with the extended arm and during radial reaching movements.

The Flexibility of Nonconsciously Deployed Cognitive Processes: Evidence from Masked Congruence Priming

Background It is well accepted in the subliminal priming literature that task-level properties modulate nonconscious processes. For example, in tasks with a limited number of targets, subliminal priming effects are limited to primes that are physically similar to the targets. In contrast, when a large number of targets are used, subliminal priming effects are observed for primes that share a semantic (but not necessarily physical) relationship with the target. Findings such as these have led researchers to conclude that task-level properties can direct nonconscious processes to be deployed exclusively over central (semantic) or peripheral (physically specified) representations. Principal Findings We find distinct patterns of masked priming for “novel” and “repeated” primes within a single task context. Novel primes never appear as targets and thus are not seen consciously in the experiment. Repeated primes do appear as targets, thereby lending themselves to the establishment of peripheral stimulus-response mappings. If the source of the masked priming effect were exclusively central or peripheral, then both novel and repeated primes should yield similar patterns of priming. In contrast, we find that both novel and repeated primes produce robust, yet distinct, patterns of priming. Conclusions Our findings indicate that nonconsciously elicited cognitive processes can be flexibly deployed over both central and peripheral representations within a single task context. While we agree that task-level properties can influence nonconscious processes, our findings sharply constrain the extent of this influence. Specifically, our findings are inconsistent with extant accounts which hold that the influence of task-level properties is strong enough to restrict the deployment of nonconsciously elicited cognitive processes to a single type of representation (i.e. central or peripheral).

Finger interaction in a three-dimensional pressing task

Accurate control of forces produced by the fingers is essential for performing object manipulation. This study examines the indices of finger interaction when accurate time profiles of force are produced in different directions, while using one of the fingers or all four fingers of the hand. We hypothesized that patterns of unintended force production among shear force components may involve features not observed in the earlier studies of vertical force production. In particular, we expected to see unintended forces generated by non-task fingers not in the direction of the instructed force but in the opposite direction as well as substantial force production in directions orthogonal to the instructed direction. We also tested a hypothesis that multi-finger synergies, quantified using the framework of the uncontrolled manifold hypothesis, will help reduce across-trials variance of both total force magnitude and direction. Young, healthy subjects were required to produce accurate ramps of force in five different directions by pressing on force sensors with the fingers of the right (dominant) hand. The index finger induced the smallest unintended forces in non-task fingers. The little finger showed the smallest unintended forces when it was a non-task finger. Task fingers showed substantial force production in directions orthogonal to the intended force direction. During four-finger tasks, individual force vectors typically pointed off the task direction, with these deviations nearly perfectly matched to produce a resultant force in the task direction. Multi-finger synergy indices reflected strong co-variation in the space of finger modes (commands to fingers) that reduced variability of the total force magnitude and direction across trials. The synergy indices increased in magnitude over the first 30% of the trial time and then stayed at a nearly constant level. The synergy index for stabilization of total force magnitude was higher for shear force components when compared to the downward pressing force component. The results suggest complex interactions between enslaving and synergic force adjustments, possibly reflecting the experience with everyday prehensile tasks. For the first time, the data document multi-finger synergies stabilizing both shear force magnitude and force vector direction. These synergies may play a major role in stabilizing the hand action during object manipulation.

Processing of low spatial frequency faces at periphery in choice reaching tasks.

Various aspects of face processing have been associated with distinct ranges of spatial frequencies. Configural processing of faces depends chiefly on low spatial frequency (LSF) information whereas high spatial frequency (HSF) supports feature based processing. However, it has also been argued that face processing has a foveal-bias (HSF channels dominate the fovea). Here we used reach trajectories as a continuous behavioral measure to study perceptual processing of faces. Experimental stimuli were LSF-HSF hybrids of male and female faces superimposed and were presented peripherally and centrally. Subject reached out to touch a specified sex and their movements were recorded. The reaching trajectories reveal that there is less effect of (interference by) LSF faces at fovea as compared to periphery while reaching to HSF targets. These results demonstrate that peripherally presented LSF information, carried chiefly by magnocellular channels, enables efficient processing of faces, possibly via a retinotectal (subcortical) pathway.

Faster, stronger, lateralized: low spatial frequency information supports face processing.

Distinct visual pathways are selectively tuned for processing specific spatial frequencies. Recently, Awasthi, Friedman, and Williams (2011) reported fast categorisation of faces at periphery, arguing for primacy of low spatial frequency (LSF) information in face processing. However, previous studies have also documented rapid categorization of places and natural scenes. Here, we tested if the LSF advantage is face specific or also involved in place perception. We used visually guided reaching as a continuous behavioral measure to examine the processing of LSF and high spatial frequency (HSF) hybrids, presented at the periphery. Subjects reached out and touched targets and their movements were recorded. The trajectories revealed that LSF interference was both 95 ms earlier and stronger for faces than places and was lateralized to the left visual field. The early processing of LSF information supports the assumption that faces are prioritised and provides a (neural) framework for such specialised processing.