Internal Forward Model Research Articles

The neural network of sensory attenuation: A neuroimaging meta-analysis.

Sensory attenuation refers to the reduction in sensory intensity resulting from self-initiated actions compared to stimuli initiated externally. A classic example is scratching oneself without feeling itchy. This phenomenon extends across various sensory modalities, including visual, auditory, somatosensory, and nociceptive stimuli. The internal forward model proposes that during voluntary actions, an efferent copy of the action command is sent out to predict sensory feedback. This predicted sensory feedback is then compared with the actual sensory feedback, leading to the suppression or reduction of sensory stimuli originating from self-initiated actions. To further elucidate the neural mechanisms underlying sensory attenuation effect, we conducted an extensive meta-analysis of functional magnetic resonance imaging (fMRI) and positron emission tomography (PET) studies. Utilizing activation likelihood estimation (ALE) analysis, our results revealed significant activations in a prominent cluster encompassing the right superior temporal gyrus (rSTG), right middle temporal gyrus (rMTG), and right insula when comparing external-generated with self-generated conditions. Additionally, significant activation was observed in the right anterior cerebellum when comparing self-generated to external-generated conditions. Further analysis using meta-analytic connectivity modeling (MACM) unveiled distinct brain networks co-activated with the rMTG and right cerebellum, respectively. Based on these findings, we propose that sensory attenuation arises from the suppression of reflexive inputs elicited by self-initiated actions through the internal forward modeling of a cerebellum-centered action prediction network, enabling the "sensory conflict detection" regions to effectively discriminate between inputs resulting from self-induced actions and those originating externally.

Facilitation of sensorimotor temporal recalibration mechanisms by cerebellar tDCS in patients with schizophrenia spectrum disorders and healthy individuals

Core symptoms in patients with schizophrenia spectrum disorders (SSD), like hallucinations or ego-disturbances, have been associated with a failure of internal forward models to predict the sensory outcomes of self-generated actions. Importantly, forward model predictions must also be able to flexibly recalibrate to changing environmental conditions, for example to account for additional delays between action and outcome. We investigated whether transcranial direct current stimulation (tDCS) can be used to improve these sensorimotor temporal recalibration mechanisms in patients and healthy individuals. While receiving tDCS on the cerebellum, temporo-parietal junction, supplementary motor area, or sham stimulation, patients with SSD and healthy control participants were repeatedly exposed to delays between actively or passively elicited button presses and auditory outcomes. Effects of this procedure on temporal perception were assessed with a delay detection task. Similar recalibration outcomes and faciliatory effects of cerebellar tDCS on recalibration were observed in SSD and healthy individuals. Our findings indicate that sensorimotor recalibration mechanisms may be preserved in SSD and highlight the importance of the cerebellum in both patients and healthy individuals for this process. They further suggest that cerebellar tDCS could be a promising tool for addressing deficits in action-outcome monitoring and related adaptive sensorimotor processes in SSD.

Hummingbirds use distinct control strategies for forward and hovering flight.

The detection of optic flow is important for generating optomotor responses to mediate retinal image stabilization, and it can also be used during ongoing locomotion for centring and velocity control. Previous work in hummingbirds has separately examined the roles of optic flow during hovering and when centring through a narrow passage during forward flight. To develop a hypothesis for the visual control of forward flight velocity, we examined the behaviour of hummingbirds in a flight tunnel where optic flow could be systematically manipulated. In all treatments, the animals exhibited periods of forward flight interspersed with bouts of spontaneous hovering. Hummingbirds flew fastest when they had a reliable signal of optic flow. All optic flow manipulations caused slower flight, suggesting that hummingbirds had an expected optic flow magnitude that was disrupted. In addition, upward and downward optic flow drove optomotor responses for maintaining altitude during forward flight. When hummingbirds made voluntary transitions to hovering, optomotor responses were observed to all directions. Collectively, these results are consistent with hummingbirds controlling flight speed via mechanisms that use an internal forward model to predict expected optic flow whereas flight altitude and hovering position are controlled more directly by sensory feedback from the environment.

Impairment of the internal forward model and feedback mechanisms for vocal sensorimotor control in post-stroke aphasia: evidence from directional responses to altered auditory feedback.

The present study examined opposing and following vocal responses to altered auditory feedback (AAF) to determine how damage to left-hemisphere brain networks impairs the internal forward model and feedback mechanisms in post-stroke aphasia. Forty-nine subjects with aphasia and sixty age-matched controls performed speech vowel production tasks while their auditory feedback was altered using randomized ± 100 cents upward and downward pitch-shift stimuli. Data analysis revealed that when vocal responses were averaged across all trials (i.e., opposing and following), the overall magnitude of vocal compensation was significantly reduced in the aphasia group compared with controls. In addition, when vocal responses were analyzed separately for opposing and following trials, subjects in the aphasia group showed a significantly lower percentage of opposing and higher percentage of following vocal response trials compared with controls, particularly for the upward pitch-shift stimuli. However, there was no significant difference in the magnitude of opposing and following vocal responses between the two groups. These findings further support previous evidence on the impairment of vocal sensorimotor control in aphasia and provide new insights into the distinctive impact of left-hemisphere stroke on the internal forward model and feedback mechanisms. In this context, we propose that the lower percentage of opposing responses in aphasia may be accounted for by deficits in feedback-dependent mechanisms of audio-vocal integration and motor control. In addition, the higher percentage of following responses may reflect aberrantly increased reliance of the speech system on the internal forward model for generating sensory predictions during vocal error detection and motor control.

Morphological and Functional Principles Governing the Plasticity Reserve in the Cerebellum: The Cortico-Deep Cerebellar Nuclei Loop Model.

Cerebellar reserve compensates for and restores functions lost through cerebellar damage. This is a fundamental property of cerebellar circuitry. Clinical studies suggest (1) the involvement of synaptic plasticity in the cerebellar cortex for functional compensation and restoration, and (2) that the integrity of the cerebellar reserve requires the survival and functioning of cerebellar nuclei. On the other hand, recent physiological studies have shown that the internal forward model, embedded within the cerebellum, controls motor accuracy in a predictive fashion, and that maintaining predictive control to achieve accurate motion ultimately promotes learning and compensatory processes. Furthermore, within the proposed framework of the Kalman filter, the current status is transformed into a predictive state in the cerebellar cortex (prediction step), whereas the predictive state and sensory feedback from the periphery are integrated into a filtered state at the cerebellar nuclei (filtering step). Based on the abovementioned clinical and physiological studies, we propose that the cerebellar reserve consists of two elementary mechanisms which are critical for cerebellar functions: the first is involved in updating predictions in the residual or affected cerebellar cortex, while the second acts by adjusting its updated forecasts with the current status in the cerebellar nuclei. Cerebellar cortical lesions would impair predictive behavior, whereas cerebellar nuclear lesions would impact on adjustments of neuronal commands. We postulate that the multiple forms of distributed plasticity at the cerebellar cortex and cerebellar nuclei are the neuronal events which allow the cerebellar reserve to operate in vivo. This cortico-deep cerebellar nuclei loop model attributes two complementary functions as the underpinnings behind cerebellar reserve.

Predictive coordination of breathing during intra-personal speaking and listening

It has long been known that human breathing is altered during listening and speaking compared to rest: during speaking, inhalation depth is adjusted to the air volume required for the upcoming utterance. During listening, inhalation is temporally aligned to inhalation of the speaker. While evidence for the former is relatively strong, it is virtually absent for the latter. We address both phenomena using recordings of speech envelope and respiration in 30 participants during 14min of speaking and listening to one's own speech. First, we show that inhalation depth is positively correlated with the total power of the speech envelope in the following utterance. Second, we provide evidence that inhalation during listening to one's own speech is significantly more likely at time points of inhalation during speaking. These findings are compatible with models that postulate alignment of internal forward models of interlocutors with the aim to facilitate communication.

Exploring the internal forward model: action-effect prediction and attention in sensorimotor processing.

Action-effect predictions are believed to facilitate movement based on its association with sensory objectives and suppress the neurophysiological response to self- versus externally generated stimuli (i.e. sensory attenuation). However, research is needed to explore theorized differences in the use of action-effect prediction based on whether movement is uncued (i.e. volitional) or in response to external cues (i.e. stimulus-driven). While much of the sensory attenuation literature has examined effects involving the auditory N1, evidence is also conflicted regarding this component's sensitivity to action-effect prediction. In this study (n = 64), we explored the influence of action-effect contingency on event-related potentials associated with visually cued and uncued movement, as well as resultant stimuli. Our findings replicate recent evidence demonstrating reduced N1 amplitude for tones produced by stimulus-driven movement. Despite influencing motor preparation, action-effect contingency was not found to affect N1 amplitudes. Instead, we explore electrophysiological markers suggesting that attentional mechanisms may suppress the neurophysiological response to sound produced by stimulus-driven movement. Our findings demonstrate lateralized parieto-occipital activity that coincides with the auditory N1, corresponds to a reduction in its amplitude, and is topographically consistent with documented effects of attentional suppression. These results provide new insights into sensorimotor coordination and potential mechanisms underlying sensory attenuation.

Functional connectivity between parietal and temporal lobes mediates internal forward models during speech production

Internal forward models hypothesize functional links between motor and sensory systems for predicting the consequences of actions. Recently, the cascaded theory proposes that somatosensory estimation in the inferior parietal lobe (IPL) can be a relay computational structure, converting motor signals into predictions of auditory consequences in a serial processing manner during speech production. The study used fMRI with functional connectivity (FC) analyses to investigate the proposed cascaded processes using three speech tasks: overt articulation (OA), silent articulation (SA) and imagined articulation (IA). The FC results showed that connectivity between aIPL and STG was increased in OA compared with SA, suggesting that the relationship between somatosensory and auditory estimations can be modulated by speech tasks. Moreover, stronger connectivity between IFGoper and pIPL, and between pIPL and STG were observed in SA and IA compared with OA. These results are consistent with a cascaded process in the internal forward models.

Botulinum Toxin-A Treatment of Post-Stroke Spasticity Is Associated with Changes in Interhemispheric Parietal Cortex Connectivity (P12-8.001)

<h3>Objective:</h3> To assess resting-state functional connectivity (rsFC) of the ipsilesional superior parietal lobule (SPL) and intraparietal sulcus (IPS) using functional magnetic resonance imaging (fMRI) in chronic stroke patients with hemiparesis both with and without post-stroke spasticity (PSS) and to explore the relationship between SPL/IPS resting-state functional connectivity (rsFC) and PSS severity. <h3>Background:</h3> Posts-stroke spasticity hampers effective rehabilitation of the paretic arm and lowers the quality of life. Effective PSS treatment with botulinum neurotoxin (BoNT) is associated with transient decrease in task-related fMRI activation of the ipsilesional SPL and IPS. We hypothesized that this would be reflected in changes in rsFC of the SPL/IPS. <h3>Design/Methods:</h3> Fourteen chronic stroke patients with upper limb weakness and PSS (the PSS group) and 8 patients with comparable weakness but no PSS (the control group) underwent clinical and spasticity evaluation and 3 fMRI examinations, at baseline (W0) and 4 and 11 weeks after BoNT (W4 and W11, respectively). Seed-based rsFC of the atlas-based SPL and IPS was evaluated using a group×time interaction analysis and the change in rsFC of SPL/IPS was correlated with change in PSS (modified Ashworth scale), integrity of the ipsilesional somatosensory afferent pathway (evoked potential N20 latency), and age. <h3>Results:</h3> In the PSS group, transient improvement in PSS was associated with increase in rsFC between the ipsilesional IPS and the contralesional SPL at W4. The interhemispheric connectivity was negatively correlated with PSS severity at baseline and with PSS improvement at W4. These changes in the posterior parietal interhemispheric rsFC suggest that previously observed task-related changes in SPL/IPS were independent of actual task performance. <h3>Conclusions:</h3> We propose adaptation of the internal forward model as the possible underlying mechanism and discuss its potential association with increased limb use or improved motor performance, as well as its potential contribution to the clinical effects of BoNT. <b>Disclosure:</b> The institution of Prof. Hlustik has received research support from Czech Health Research Council (AZV CR). Dr. Hok has nothing to disclose. Tomas Veverka has nothing to disclose. Markéta Trnecková has nothing to disclose. Pavel Otruba has nothing to disclose. Jana Zapletalova has nothing to disclose. Zbynek Tudos has nothing to disclose. Prof. Lotze has received publishing royalties from a publication relating to health care. Dr. Kanovsky has nothing to disclose.

Multiple step saccades are generated by internal real-time saccadic error correction.

Multiple step saccades (MSSs) are an atypical form of saccade that consists of a series of small-amplitude saccades. It has been argued that the mechanism for generating MSS is due to the automatic saccadic plan. This argument was based on the observation that trials with MSS had shorter saccadic latency than trials without MSS in the reactive saccades. However, the validity of this argument has never been verified by other saccadic tasks. Alternatively, we and other researchers have speculated that the function of MSS is the same as that of the corrective saccade (CS), i.e., to correct saccadic errors. Thus, we propose that the function of the MSS is also to rectify saccadic errors and generated by forward internal models. The objective of the present study is to examine whether the automatic theory is universally applicable for the generation of MSSs in various saccadic tasks and to seek other possible mechanisms, such as error correction by forward internal models. Fifty young healthy subjects (YHSs) and fifty elderly healthy subjects (EHSs) were recruited in the present study. The task paradigms were prosaccade (PS), anti-saccade (AS) and memory-guided saccade (MGS) tasks. Saccadic latency in trials with MSS was shorter than without MSS in the PS task but similar in the AS and MGS tasks. The intersaccadic intervals (ISI) were similar among the three tasks in both YHSs and EHSs. Our results indicate that the automatic theory is not a universal mechanism. Instead, the forward internal model for saccadic error correction might be an important mechanism.

Interhemispheric parietal cortex connectivity reflects improvement in post-stroke spasticity due to treatment with botulinum toxin-A

In post-stroke spasticity (PSS), effective treatment with botulinum neurotoxin (BoNT) is associated with transient decrease in activation of the ipsilesional superior parietal lobule (SPL) and intraparietal sulcus (IPS). We hypothesized that this would be reflected in changes in resting-state functional connectivity (rsFC) of the SPL/IPS. Our aim was therefore to assess rsFC of the ipsilesional SPL/IPS in chronic stroke patients with hemiparesis both with and without PSS and to explore the relationship between SPL/IPS rsFC and PSS severity. To this end, fourteen chronic stroke patients with upper limb weakness and PSS (the PSS group) and 8 patients with comparable weakness but no PSS (the control group) underwent clinical evaluation and 3 fMRI examinations, at baseline (W0) and 4 and 11 weeks after BoNT (W4 and W11, respectively). Seed-based rsFC of the atlas-based SPL and IPS was evaluated using a group×time interaction analysis and a correlation analysis with PSS severity (modified Ashworth scale), integrity of the ipsilesional somatosensory afferent pathway (evoked potential N20 latency), and age. In the PSS group, transient improvement in PSS was associated with increase in rsFC between the ipsilesional IPS and the contralesional SPL at W4. The interhemispheric connectivity was negatively correlated with PSS severity at baseline and with PSS improvement at W4. We propose adaptation of the internal forward model as the putative underlying mechanism and discuss its possible association with increased limb use, diminished spastic dystonia, or improved motor performance, as well as its potential contribution to the clinical effects of BoNT.

The otolith vermis: A systems neuroscience theory of the Nodulus and Uvula.

The Nodulus and Uvula (NU) (lobules X and IX of the cerebellar vermis) form a prominent center of vestibular information processing. Over decades, fundamental and clinical research on the NU has uncovered many aspects of its function. Those include the resolution of a sensory ambiguity inherent to inertial sensors in the inner ear, the otolith organs; the use of gravity signals to sense head rotations; and the differential processing of self-generated and externally imposed head motion. Here, I review these works in the context of a theoretical framework of information processing called the internal model hypothesis. I propose that the NU implements a forward internal model to predict the activation of the otoliths, and outputs sensory predictions errors to correct internal estimates of self-motion or to drive learning. I show that a Kalman filter based on this framework accounts for various functions of the NU, neurophysiological findings, as well as the clinical consequences of NU lesions. This highlights the role of the NU in processing information from the otoliths and supports its denomination as the “otolith” vermis.

The effect of self-generated versus externally generated actions on timing, duration, and amplitude of blood oxygen level dependent response for visual feedback processing.

It has been widely assumed that internal forward models use efference copies to create predictions about the sensory consequences of our own actions. While these predictions have frequently been associated with a reduced blood oxygen level dependent (BOLD) response in sensory cortices, the timing and duration of the hemodynamic response for the processing of video feedback of self‐generated (active) versus externally generated (passive) movements is poorly understood. In the present study, we tested the hypothesis that predictive mechanisms for self‐generated actions lead to early and shorter neural processing compared with externally generated movements. We investigated active and passive movements using a custom‐made fMRI‐compatible movement device. Visual video feedback of the active and passive movements was presented in real time or with variable delays. Participants had to judge whether the feedback was delayed. Timing and duration of BOLD impulse response was calculated using a first (temporal derivative [TD]) and second‐order (dispersion derivative [DD]) Taylor approximation. Our reanalysis confirmed our previous finding of reduced BOLD response for active compared to passive movements. Moreover, we found positive effects of the TD and DD in the supplementary motor area, cerebellum, visual cortices, and subcortical structures, indicating earlier and shorter hemodynamic responses for active compared to passive movements. Furthermore, earlier activation in the putamen for active compared to passive conditions was associated with reduced delay detection performance. These findings indicate that efference copy‐based predictive mechanisms enable earlier processing of action feedback, which might have reduced the ability to detect short delays between action and feedback.

Beyond task-space exploration: On the role of variance for motor control and learning.

This conceptual analysis on the role of variance for motor control and learning should be taken as a call to: (a) overcome the classic motor-action controversy by identifying converging lines and mutual synergies in the explanation of motor behavior phenomena, and (b) design more empirical research on low-level operational aspects of motor behavior rather than on high-level theoretical terms. Throughout the paper, claim (a) is exemplified by deploying the well-accepted task-space landscape metaphor. This approach provides an illustration not only of a dynamical sensorimotor system but also of a structure of internal forward models, as they are used in more cognitively rooted frameworks such as the theory of optimal feedback control. Claim (b) is put into practice by, mainly theoretically, substantiating a number of predictions for the role of variance in motor control and learning that can be derived from a convergent perspective. From this standpoint, it becomes obvious that variance is neither generally “good” nor generally “bad” for sensorimotor learning. Rather, the predictions derived suggest that specific forms of variance cause specific changes on permanent performance. In this endeavor, Newell’s concept of task-space exploration is identified as a fundamental learning mechanism. Beyond, we highlight further predictions regarding the optimal use of variance for learning from a converging view. These predictions regard, on the one hand, additional learning mechanisms based on the task-space landscape metaphor—namely task-space formation, task-space differentiation and task-space (de-)composition—and, on the other hand, mechanisms of meta-learning that refer to handling noise as well as learning-to-learn and learning-to-adapt. Due to the character of a conceptual-analysis paper, we grant ourselves the right to be highly speculative on some issues. Thus, we would like readers to see our call mainly as an effort to stimulate both a meta-theoretical discussion on chances for convergence between classically separated lines of thought and, on an empirical level, future research on the role of variance in motor control and learning.

The role of grasping demands on tactile suppression.

Tactile perception is impaired in a limb that is moving compared to when it is static. A possible mechanism that explains this phenomenon is an internal forward model that estimates future sensory states of the moving limb and suppresses associated feedback signals arising from that limb. Because sensorimotor estimations are based on an interplay of efferent and afferent feedback signals, the strength of tactile suppression may also depend on the relative utilization of sensory feedback from the moving limb. To test how the need to process somatosensory feedback influences movement-induced tactile suppression, we asked participants to perform reach-to-grasp movement of different demands: the target object was covered with materials of different frictional properties and the task was performed both with and without vision. As expected, participants performed the grasping movement more carefully when interacting with objects of low than high friction surfaces and when performing the task without vision. This denotes a greater need for somatosensory guidance of the digits to appropriately position the digits on the object. Accordingly, tactile suppression was weaker when grasping low than high friction objects, but only when grasping without vision. This suggests that movement-induced tactile suppression is modulated by grasping demands. Tactile suppression is downregulated when the need to process somatosensory feedback signals from that moving limb increases, like in situations when somatosensory input about the digit's state is the sole source of sensory information and when this information is particularly important for the task at hand.

Mental practice modulates functional connectivity between the cerebellum and the primary motor cortex

SummaryOur brain has the extraordinary capacity to improve motor skills through mental practice. Conceptually, this ability is attributed to internal forward models, which are cerebellar neural networks that can predict the sensory consequences of motor commands. In our study, we employed single and dual-coil transcranial magnetic stimulations to probe the level of corticospinal excitability and cerebellar-brain inhibition, respectively, before and after a mental practice session or a control session. Motor skill (i.e., accuracy and speed) was measured using a sequential finger tapping-task. We found that mental practice enhanced both speed and accuracy. In parallel, the functional connectivity between the cerebellum and the primary motor cortex changed, with less inhibition from the first to the second. These findings reveal the existence of neuroplastic changes within the cerebellum, supporting the involvement of internal models after mental practice.

Cerebellar Representations of Errors and Internal Models.

After decades of study, a comprehensive understanding of cerebellar function remains elusive. Several hypotheses have been put forward over the years, including that the cerebellum functions as a forward internal model. Integrated into the forward model framework is the long-standing view that Purkinje cell complex spike discharge encodes error information. In this brief review, we address both of these concepts based on our recordings of cerebellar Purkinje cells over the last decade as well as newer findings from the literature. During a high-dimensionality tracking task requiring continuous error processing, we find that complex spike discharge provides a rich source of non-error signals to Purkinje cells, indicating that the classical error encoding role ascribed to climbing fiber input needs revision. Instead, the simple spike discharge of Purkinje cells carries robust predictive and feedback signals of performance errors, as well as kinematics. These simple spike signals are consistent with a forward internal model. We also show that the information encoded in the simple spike is dynamically adjusted by the complex spike firing. Synthesis of these observations leads to the hypothesis that complex spikes convey behavioral state changes, possibly acting to select and maintain forward models.

Human motor learning is robust to control-dependent noise.

Noises are ubiquitous in sensorimotor interactions and contaminate the information provided to the central nervous system (CNS) for motor learning. An interesting question is how the CNS manages motor learning with imprecise information. Integrating ideas from reinforcement learning and adaptive optimal control, this paper develops a novel computational mechanism to explain the robustness of human motor learning to the imprecise information, caused by control-dependent noise that exists inherently in the sensorimotor systems. Starting from an initial admissible control policy, in each learning trial the mechanism collects and uses the noisy sensory data (caused by the control-dependent noise) to form an imprecise evaluation of the performance of the current policy and then constructs an updated policy based on the imprecise evaluation. As the number of learning trials increases, the generated policies mathematically provably converge to a (potentially small) neighborhood of the optimal policy under mild conditions, despite the imprecise information in the learning process. The mechanism directly synthesizes the policies from the sensory data, without identifying an internal forward model. Our preliminary computational results on two classic arm reaching tasks are in line with experimental observations reported in the literature. The model-free control principle proposed in the paper sheds more lights into the inherent robustness of human sensorimotor systems to the imprecise information, especially control-dependent noise, in the CNS.

Movement Planning Determines Sensory Suppression: An Event-related Potential Study

Sensory suppression refers to the phenomenon that sensory input generated by our own actions, such as moving a finger to press a button to hear a tone, elicits smaller neural responses than sensory input generated by external agents. This observation is usually explained via the internal forward model in which an efference copy of the motor command is used to compute a corollary discharge, which acts to suppress sensory input. However, because moving a finger to press a button is accompanied by neural processes involved in preparing and performing the action, it is unclear whether sensory suppression is the result of movement planning, movement execution, or both. To investigate this, in two experiments, we compared ERPs to self-generated tones that were produced by voluntary, semivoluntary, or involuntary button-presses, with externally generated tones that were produced by a computer. In Experiment 1, the semivoluntary and involuntary button-presses were initiated by the participant or experimenter, respectively, by electrically stimulating the median nerve in the participant's forearm, and in Experiment 2, by applying manual force to the participant's finger. We found that tones produced by voluntary button-presses elicited a smaller N1 component of the ERP than externally generated tones. This is known as N1-suppression. However, tones produced by semivoluntary and involuntary button-presses did not yield significant N1-suppression. We also found that the magnitude of N1-suppression linearly decreased across the voluntary, semivoluntary, and involuntary conditions. These results suggest that movement planning is a necessary condition for producing sensory suppression. We conclude that the most parsimonious account of sensory suppression is the internal forward model.

Time for Space and the Stability of Prospective Control: Reaching-to-Grasp Gibson.

Gibson formulated an approach to goal-directed behavior using prospective information in the context of visually guided locomotion and manual behavior. The former was Gibson's paradigm case, but it is the rapidity of targeted reaching that has provided the special challenge for stable control. Recent treatments of visually guided reaching assume that internal forward models are required to generate stable behavior given delays caused by neural transmission times. Internal models are representations of the sort eschewed by Gibson in favor of prospective information. Reaching is usually described as guided using relative distances of hand and target, but prospective information is usually temporal rather than spatial. We describe proportional rate control models that incorporate time dimensioned prospective information and show they remain stable in the face of delays. The use of time-dimensioned prospective information removes the need for internal models for stable behavior despite neural transmission delays and allows Gibson's approach to prevail.