A Dickensian origin for Sherrington's enchanted loom?

Sherrington's image of the mind-brain as an 'enchanted loom' has intrigued scientists for decades. Its origin, however, has been unclear. Here, we make a new suggestion as to where it may have come from. The article sheds light on the connections between literary and scientific similes and metaphors.

Grip force regulates hand impedance to optimize object stability in high impact loads.

Anticipatory grip force adjustments are a prime example of the predictive nature of motor control. An object held in precision grip is stabilized by fine adjustments of the grip force against changes in tangential load force arising from inertia during acceleration and deceleration. When an object is subject to sudden impact loads, prediction becomes critical as the time available for sensory feedback is very short. Here, we investigated the control of grip force when participants performed a targeted tapping task with a hand-held object. During the initial transport phase of the movement, load force varied smoothly with acceleration. In contrast, in the collision, load forces sharply increased to very large values. In the transport phase, grip force and load force were coupled in phase, as expected. However, in the collision, grip force did not parallel load force. Rather, it exhibited a stereotyped profile with maximum ∼65 ms after peak load at contact. By using catch trials and a virtual environment, we demonstrate that this peak of grip force is pre-programmed. This observation is validated across experimental manipulations involving different target stiffness and directions of movement. We suggest that the central nervous system optimizes stability in object manipulation-as in catching-by regulating mechanical parameters including stiffness and damping through grip force. This study provides novel insights about how the brain coordinates grip force in manipulation involving an object interacting with the environment.

Selective antegrade cerebral perfusion attenuates brain metabolic deficit in aortic arch surgery: a prospective randomized trial.

BACKGROUND: Aortic arch surgery has a high incidence of brain injury. This may in part be caused by a cerebral metabolic deficit observed after hypothermic circulatory arrest (HCA). We hypothesized that selective antegrade cerebral perfusion (SACP) would attenuate this phenomenon. METHODS AND RESULTS: In a prospective randomized trial, 42 adult patients were allocated to either HCA (22) or SACP. HCA occurred at a nasopharyngeal temperature of 15 degrees C and SACP at a corporeal temperature of 25 degrees C with cerebral perfusion at 15 degrees C. Paired arterial and jugular venous samples were taken before and after arrest. Continuous transcranial Doppler monitoring of middle cerebral artery velocity (MCAV) was performed. Neuropsychometric testing was performed preoperatively and at 6 and 12 weeks postoperatively. There were 3 hospital deaths (7.1%), 2 strokes (4.8%), and 6 episodes of transient neurological deficit (14.3%). From before to after arrest, jugular bulb pO2 changed by -21.67 mm Hg (26.4) in the HCA group versus +2.27 mm Hg (18.8) in the SACP group (P=0.007). Oxygen extraction changed by +1.7 mL/dL (1.3) in the HCA group versus -1 mL/dL (2.4) in the SACP group (P<0.001). MCAV increased by 6.25 cm/s (9.1) in the HCA group and 19.2 cm/s (10.1) in the SACP group (P=0.001). Incidence of neuropsychometric deficit at 6 weeks was 6/12 (50%) in HCA patients and 8/10 (80%) in SACP patients (P=0.2), and at 12 weeks was 6/16 (38%) in HCA patients and 4/11 (36%) in SACP patients (P=1). CONCLUSIONS: SACP attenuates the metabolic changes seen after HCA. Further studies are required to assess optimal perfusion conditions and clinical outcome.

Absence of gaze direction effects on EEG measures of sensorimotor function.

OBJECTIVE: Gaze direction is known to modulate the activation patterns of sensorimotor areas as seen at the single cell level and in functional magnetic resonance imaging (fMRI). To determine whether such gaze direction effects can be observed in scalp-recorded electroencephalogram (EEG) measures of sensorimotor function we investigated somatosensory evoked potentials (SEPs) and steady state movement related cortical potentials (MRPs). METHODS: In two separate experiments, SEPs were elicited by electrical stimulation of the median nerve (experiment 1) and steady state MRPs were induced by 2 Hz tapping paced by an auditory cue (experiment 2), while subjects directed their gaze 15 degrees to the left or to the right. RESULTS: Gaze direction failed to produce any appreciable differences in the waveforms of the SEPs or MRPs. In particular, there was no effect on peak amplitude, peak latency and peak scalp topography measures of SEP and MRP components, or on spatial or temporal parameters of dipole models of the underlying cortical generators. Additional frequency domain analyses did not reveal reliable gaze-related changes in induced power at electrode sites overlying somatosensory and motor areas, or in coherence between pairs of parietal, central and frontal electrodes, across a broad range of frequencies. CONCLUSIONS: EEG measures of sensorimotor function, obtained in a non-visual motor task, are insensitive to modulatory effects of gaze direction in sensorimotor areas that are observable with fMRI.

Motor intention activity in the macaque's lateral intraparietal area. I. Dissociation of motor plan from sensory memory.

1. The lateral intraparietal area (area LIP) of the monkey's posterior parietal cortex (PPC) contains neurons that are active during saccadic eye movements. These neurons' activity includes visual and saccade-related components. These responses are spatially tuned and the location of a neuron's visual receptive field (RF) relative to the fovea generally overlaps its preferred saccade amplitude and direction (i.e., its motor field, MF). When a delay is imposed between the presentation of a visual stimulus and a saccade made to its location (memory saccade task), many LIP neurons maintain elevated activity during the delay (memory activity, M), which appears to encode the metrics of the next intended saccadic eye movements. Recent studies have alternatively suggested that LIP neurons encode the locations of visual stimuli regardless of where the animal intends to look. We examined whether the M activity of LIP neurons specifically encodes movement intention or the locations of recent visual stimuli, or a combination of both. In the accompanying study, we investigated whether the intended-movement activity reflects changes in motor plan. 2. We trained monkeys (Macaca mulatta) to memorize the locations of two visual stimuli and plan a sequence of two saccades, one to each remembered target, as we recorded the activity of single LIP neurons. Two targets were flashed briefly while the monkey maintained fixation; after a delay the fixation point was extinguished, and the monkey made two saccades in sequence to each target's remembered location, in the order in which the targets were presented. This "delayed double saccade" (DDS) paradigm allowed us to dissociate the location of visual stimulation from the direction of the planned saccade and thus distinguish neuronal activity related to the target's location from activity related to the saccade plan. By imposing a delay, we eliminated the confounding effect of any phasic responses coincident with the appearance of the stimulus and with the saccade. 3. We arranged the two visual stimuli so that in one set of conditions at least the first one was in the neuron's visual RF, and thus the first saccade was in the neuron's motor field (MF). M activity should be high in these conditions according to both the sensory memory and motor plan hypotheses. In another set of conditions, the second stimulus appeared in the RF but the first one was presented outside the RF, instructing the monkey to plan the first saccade away from the neuron's MF. If the M activity encodes the motor plan, it should be low in these conditions, reflecting the plan for the first saccade (away from the MF). If it is a sensory trace of the stimulus' location, it should be high, reflecting stimulation of the RF by the second target. 4. We tested 49 LIP neurons (in 3 hemispheres of 2 monkeys) with M activity on the DDS task. Of these, 38 (77%) had M activity related to the next intended saccade. They were active in the delay period, as expected, if the first saccade was in their preferred direction. They were less active or silent if the next saccade was not in their preferred direction, even when the second stimulus appeared in their RF. 5. The M activity of 8 (16%) of the remaining neurons specifically encoded the location of the most recent visual stimulus. Their firing rate during the delay reflected stimulation of the RF independently of the saccade being planned. The remaining 3 neurons had M activity that did not consistently encode either the next saccade or the stimulus' location. 6. We also recorded the activity of a subset of neurons (n = 38) in a condition in which no stimulus appeared in a neuron's RF, but the second saccade was in the neuron's MF. In this case the majority of neurons tested (23/38, 60%) became active in the period between the first and second saccade, even if neither stimulus had appeared in their RF. Moreover, this activity appeared only after the first saccade had started in all but two of

Motor intention activity in the macaque's lateral intraparietal area. II. Changes of motor plan.

1. In the companion paper we reported that the predominant signal of the population of neurons in the lateral intraparietal area (area LIP) of the monkey's posterior parietal cortex (PPC) encode the next intended saccadic eye movement during the delay period of a memory-saccade task. This result predicts that, should be monkey change his intention of what the next saccade will be, LIP activity should change accordingly to reflect the new plan. We tested this prediction by training monkeys to change their saccadic plan on command and recording the activity of LIP neurons across plan changes. 2. We trained rhesus monkeys (Macaca mulatta) to maintain fixation on a light spot as long as this spot remained on. During this period we briefly presented one, two, or three peripheral visual stimuli in sequence, each followed by a delay (memory period, M). After the final delay the fixation spot was extinguished, and the monkey had to quickly make a saccade to the location of the last target to have appeared. The monkey could not predict which stimuli, nor how many, would appear on each trial. He thus had to plan a saccade to each stimulus as it appeared and change his saccade plan whenever a stimulus appeared at a different location. 3. We recorded the M period activity of 81 area LIP neurons (from 3 hemispheres of 2 monkeys) in this task. We predicted that, if a neuron's activity reflected the monkey's planned saccade, its activity should be high while the monkey planned a saccade in the neuron's motor field (MF), and low while the planned saccade was in the opposite direction. The activity of most of the neurons in our sample changed in accordance with our hypothesis as the monkey's planned saccade changed. 4. In one condition the monkey was instructed by visual stimuli to change his plan from a saccade in the neuron's preferred direction to a saccade planned in the opposite direction. In this condition activity decreased significantly (P < 0.05) in 65 (80%) of 81 neurons tested. These neurons' activity changed to reflect the new saccade plan even though the cue for this change was not presented in their RF. 5. As a control we randomly interleaved, among trials requiring a plan change, trials in which the monkey had to formulate two consecutive plans to make a saccade in the neuron's preferred direction. The activity remained unchanged (P < 0.05) in 22 of 31 neurons tested (79%), indicating that the neurons continued to encode the same saccade plan. 6. In a variant of the task, the cue to the location of the required saccade was either a light spot or a noise burst from a loudspeaker. Of 22 neurons tested in this task, 16 (73%) showed activity changes consistent with plan changes cued by visual or auditory stimuli. 7. Alterations in the monkey's intentions, even in the absence of overt behavior, are manifested in altered LIP activity. These activity changes could be induced whether visual or auditory cues were used to indicate the required plan changes. Most LIP neurons thus do not encode only the locations of visual stimuli, but also the intention to direct gaze to specific locations, independently of whether a gaze shift actually occurs.

Spatially tuned auditory responses in area LIP of macaques performing delayed memory saccades to acoustic targets.

1. The lateral intraparietal area (area LIP) of the macaque's posterior parietal cortex (PPC) lies in the dorsal stream of extrastriate visual areas. It receives extensive visual inputs and sends outputs to several eye movement centers. It contains neurons with visual and saccade-related responses suggesting a role of area LIP in programming saccadic eye movements to visual targets. Because primates can also orient to nonvisual stimuli, we investigated whether LIP neurons process stimuli of other modalities besides the visual one by comparing their activity in auditory and visual saccade tasks. 2. We recorded the activity of single neurons of Macaca mulatta monkeys while they performed memory saccades to acoustic and visual targets. We analyzed the activity during stimulus presentation (stimulus period, S) and during the delay (memory period, M) between stimulus presentation and the saccade to its remembered location. 3. Among 80 area LIP neurons tested, we found 44 that had S period and/or M period responses following presentation of the auditory stimulus. Most of these responses were spatially tuned, i.e., selective for the left or right stimulus location (27 of 29 S responses; 25 of 29 M responses). 4. The majority of neurons with responses in the auditory memory saccade task also responded in the visual version of the task. Eighty-nine percent (24/27) were clearly bimodal in the S period, and 88% (23/26) were bimodal in the M period. 5. Almost all the neurons with spatially tuned auditory responses that were bimodal were also spatially tuned in their visual responses (20/22 for S responses; 18/19 for M responses). The spatial tuning for the two modalities was the same in 85% (17/20) of the tested neurons for the S responses, and in 83% (15/18) of the tested neurons for the M responses. 6. Area LIP contains a population of neurons that respond to both visual and auditory stimuli. This result is consistent with our finding that the memory activity of many LIP cells encodes the next planned saccade. If cells are coding planned movements, they should be active independently of the sensory modality of the target for the movement, as was the case for most of the neurons described in the present study.

Saccade-related activity in the lateral intraparietal area. I. Temporal properties; comparison with area 7a.

1. The cortex of the inferior parietal lobule (IPL) contains neurons whose activity is related to saccadic eye movements. The exact role of the IPL in relation to saccades remains, however, unclear. In this and the companion paper, we approach this problem by quantifying many of the spatial and temporal parameters of the saccade-related (S) activity. These parameters have hitherto been largely unstudied. 2. The activity of single neurons was recorded from Macaca mulatta monkeys while they were performing a delayed-saccade task. The analysis presented here is based on 161 neurons recorded from the lateral intraparietal area (LIP), a recently defined subdivision of the IPL; and 54 neurons recorded from the neighboring part of the IPL, area 7a. Overall, 409 IPL neurons were isolated in this study. 3. The typical activity of IPL neurons during the delayed-saccade task has three basic phases: light sensitive (LS), memory (M), and S. These basic phases are common to neurons of both areas LIP and 7a. In each phase (LS, M, and S), individual neurons may or may not be active. Most LIP neurons, however, are active in more than one phase. 4. To compare the activity levels of different neurons, the actual firing rate was weighted by each neuron's background level, yielding an "activity index" for each neuron, in each phase of the task. We calculated the activity index for the LS and M phases and for three phases related to the saccade: a presaccadic (Pre-S), a saccade-coincident (S-Co), and a postsaccadic (Post-S) phase. For area LIP neurons the median values of the activity index were high for the LS, M, Pre-S, and S-Co activities, and slightly lower in the Post-S period. In area 7a the median values were low for the LS phase and, in particular, for the M and Pre-S phases, somewhat higher coincident with the saccade, and high post-saccadically. 5. In area LIP, in each phase, 49-63% of the neurons had excitatory activity, and 10-17% had inhibitory responses. 6. In contrast, in area 7a excitatory responses were most frequent in the Post-S phase (56%). Excitation was particularly infrequent during M (28%) and Pre-S (22%). The incidence of inhibitory responses varied too (4-18%). The time course of inhibition was roughly opposite that of excitation; the highest frequency of inhibitory responses occurred during the saccade.(ABSTRACT TRUNCATED AT 400 WORDS)

Saccade-related activity in the lateral intraparietal area. II. Spatial properties.

1. Single-neuron activity was recorded from the inferior parietal lobule (IPL) of Macaca mulatta monkeys while they were performing delayed saccades and related tasks. Temporal characteristics of this activity were presented in the companion paper. Here we focus on the spatial characteristics of the activity. The analysis was based on recordings from 145 neurons. All these neurons were from the lateral intraparietal area (LIP), a recently defined subdivision of the IPL. 2. Delayed saccades were made in eight directions. Direction-tuning curves were calculated for each neuron, during each of the following activity phases that were described in the companion paper: light sensitive (LS), delay-period memory (M), and saccade related (S); the latter further partitioned into presaccadic (Pre-S), saccade coincident (S-Co), and postsaccadic (Post-S). 3. Width and preferred direction were calculated for each direction-tuning curve. We studied the distributions of widths and preferred directions in LIP's neuronal population. In each case we included only neurons that showed clear excitatory activity in the phases in question. 4. Width was defined as the angle over which the response was higher than 50% of its maximal net value. Width distributions were similar for all phases studied. Widths varied widely from neuron to neuron, from very narrow (less than 45 degrees) to very wide (close to 360 degrees). Median widths were approximately 90 degrees in all phases. 5. Preferred-direction distributions were also similar for various phases. All directions were represented in each distribution, but contralateral directions were more frequent (e.g., 69% for S-Co). 6. For each neuron the alignment of the preferred directions of its various phases was determined. Distributions of alignments were calculated (again, phases that were not clearly excitatory were disregarded). On the level of the neuronal population LS, M, and Pre-S were well aligned with each other. S-Co was also aligned with these phases, but less precisely. 7. A set of "narrowly tuned" neurons was selected by imposing a constraint of narrow (width, less than 90 degrees) LS and S-Co direction tuning. In this set of neurons, the LS and S-Co preferred directions were very well aligned (median, 12 degrees). The fraction of narrowly tuned neurons in the population was 40% (25/63). Thus, in a large subpopulation of area LIP, a fairly precise alignment exists between sensory and motor fields. 8. An additional set of 82 area LIP neurons were recorded while the monkey performed delayed saccades to 32 targets located on small, medium, and large imaginary circles.(ABSTRACT TRUNCATED AT 400 WORDS)

Sensorimotor transformation during eye movements to remembered visual targets.

For eye movements made to visual targets, the brain must transform the retinotopic coordinate frame of the visual system to that of the oculomotor plant. Ideally, responses should exactly match target demands. However, during eye movements to remembered targets, responses are spatially distorted. The transformation does not retain accurate retinotopic registration, having both constant and variable components of error. Generally, the constant pattern of distortion appears as a hypermetria for upward saccades and a hypometria for downward movements. Most of the error accumulates during the first 800 msec of memory-contingent delay. The results are interpreted with respect to theories of how spatial information may be coded and transformed.

Eye position effects on visual, memory, and saccade-related activity in areas LIP and 7a of macaque.

We studied the effect of eye position on the light-sensitive, memory, and saccade-related activities of neurons of the lateral intraparietal area and area 7a in the posterior parietal cortex of rhesus monkeys. A majority of the cells showed significant effects of eye position, for each of the 3 types of response. The direction tuning of the light-sensitive, memory and saccade responses did not change with eye position but the magnitude of the response did. Since previous work showed a similar effect for the light-sensitive response of area 7a neurons (Andersen and Mountcastle, 1983; Andersen et al., 1985b), the present results indicate that this modulating effect of eye position may be a general one, as it is found in 3 types of responses in 2 cortical areas. Gain fields were mapped by measuring the effect of eye position on the magnitude of the response at 9 different eye positions for each neuron. The gain fields were usually planar or largely planar for all 3 types of response in both areas, indicating that the magnitude of the response usually varies linearly with both horizontal and vertical eye position. A similar observation was made previously for the gain fields of the light-sensitive response of area 7a neurons (Andersen et al., 1985b). Although gain fields sloped in all directions for the population of cells, the gain field slopes of the light-sensitive, memory and saccade responses for individual cells were usually similar. It is proposed that these eye position effects play an important role in making coordinate transformations for visually guided movement.