Table tennis players use superior saccadic eye movements to track moving visual targets.

Introduction: Table tennis players perform visually guided visuomotor responses countlessly. The exposure of the visual system to frequent and long-term motion stimulation has been known to improve perceptual motion detection and discrimination abilities as a learning effect specific to that stimulus, so may also improve visuo-oculomotor performance. We hypothesized and verified that table tennis players have good spatial accuracy of saccades to moving targets. Methods: University table tennis players (TT group) and control participants with no striking-sports experience (Control group) wore a virtual reality headset and performed two ball-tracking tasks to track moving and stationary targets in virtual reality. The ball moved from a predetermined position on the opponent's court toward the participant's court. A total of 54 conditions were examined for the moving targets in combinations of three ball trajectories (familiar parabolic, unfamiliar descent, and unfamiliar horizontal), three courses (left, right, and center), and six speeds. Results and discussion: All participants primarily used catch-up saccades to track the moving ball. The TT group had lower mean and inter-trial variability in saccade endpoint error compared to the Control group, showing higher spatial accuracy and precision, respectively. It suggests their improvement of the ability to analyze the direction and speed of the ball's movement and predict its trajectory and future destination. The superiority of the spatial accuracy in the TT group was seen in both the right and the left courses for all trajectories but that of precision was for familiar parabolic only. The trajectory dependence of improved saccade precision in the TT group implies the possibility that the motion vision system is trained by the visual stimuli frequently encountered in table tennis. There was no difference between the two groups in the onset time or spatial accuracy of saccades for stationary targets appearing at various positions on the ping-pong table. Conclusion: Table tennis players can obtain high performance (spatial accuracy and precision) of saccades to track moving targets as a result of motion vision ability improved through a vast amount of visual and visuo-ocular experience in their play.

Transcranial static magnetic field stimulation over hMT+ inhibits visual motion discriminability.

Visuomotor performance acting on a moving target is fundamentally based on visual motion discriminability, and its neural basis is presumed to be human MT (hMT+), a motion vision center of the dorsal visual pathway. In this study, we investigated whether and how the accuracy and speed of motion discrimination are affected by applying transcranial static magnetic field stimulation (tSMS) to hMT+, which reduces cortical excitability. Sixteen participants performed a motion direction discrimination (MDD) task using a random dot kinematogram before (Pre-test) and during (During-test) application of the tSMS over left hMT+. The correct rate of the MDD task was significantly lower in the During-test compared to the Pre-test, an effect not seen with the sham condition. The inhibition effects were observed only for the right visual field corresponding to hMT+ in the stimulated hemisphere. On the other hand, no modulatory effect of tSMS was observed in the reaction time. We, therefore, demonstrated the inhibitory effect of tSMS on the left hMT+ impairs the accuracy but not the speed of motion information processing in the contralateral visual field.

Acute exercise has specific effects on the formation process and pathway of visual perception in healthy young men.

Visual perception is formed over time through the formation process and visual pathway. Exercise improves visual perception, but it is unclear whether exercise modulates nonspecifically or specifically the formation process and pathway of visual perception. Healthy young men performed the visual detection task in a backward masking paradigm before and during cycling exercise at a mild intensity or rest (control). The task presented gratings of a circular patch (target) and annulus (mask) arranged concentrically as a visual stimulus and asked if the presence and striped pattern (feature) of the target were detected. The relationship between the orientations of the gratings of the target and the mask included iso-orientation and orthogonal orientation to investigate the orientation selectivity of the masking effect. The masking effect was evaluated by perceptual suppressive index (PSI). Exercise improved feature detection (∆PSI; Exercise: -20.6%, Control: 1.7%) but not presence detection (∆PSI; Exercise: 8.9%, Control: 29.6%) compared to the control condition, and the improving effect resulted from the attenuation of the non-orientation-selective (∆PSI; Exercise: -29.0%, Control: 16.8%) but not orientation-selective masking effect (∆PSI; Exercise: -3.1%, Control: 11.7%). These results suggest that exercise affects the formation process of the perceptual feature of the target stimulus by suppressively modulating the neural networks responsible for the non-orientation-selective surround interaction in the subcortical visual pathways, whose effects are inherited by the cortical visual pathways necessary for perceptual image formation. In conclusion, our findings suggest that acute exercise improves visual perception transiently through the modulation of a specific formation process of visual processing.

Daily fluctuations in visual motion discriminability contribute to daily fluctuations in continuous visuomotor performance.

In ball sports such as table tennis, in which a ball moving at high speed is hit, an athlete's brain needs to process the motion information of the ball, predict the arrival point, and form a motor command to direct the racket there. Therefore, day-to-day fluctuations in visuomotor performance may be ascribed to fluctuations in visual motion discriminability, but it is not clear how the two are related. To examine this point, university table tennis players performed a motion direction discrimination (MDD) task and continuous visuomotor (CVM) task over 10 days as an estimation of visual motion discriminability and visuomotor performance, respectively. In the MDD task, using a joystick, participants distinguished the direction of a global coherent motion of target dots moving in the same direction on a PC monitor from innumerable dots moving in random directions. In the CVM task, participants hit sequential targets moving fast from right to left on the PC monitor by operating the cursor on the left side of the monitor up and down using the prehensile force of their thumb and index finger. The scores in the MDD and CVM tasks fluctuated day by day and showed a significant and moderate correlation between the MDD task score for the visual field in which the participants captured the target in the CVM task and the CVM task score. This correlation was confirmed even with the target moving from left to right. The fluctuations in the onset latency and the endpoint position of the cursor movement approaching the target were correlated with those of the visual motion discriminability, suggesting the contribution of motion vision to the speed and accuracy of the visuomotor performance. Moreover, these relationships were prominent in veteran players. For table tennis athletes, especially experienced players, fluctuations in the visual motion discrimination performance in a visual field specific for capturing a ball may be responsible for the fluctuations in continuous visuomotor (striking) performance.

The combination of acute exercise and eye closure has a synergistic effect on alpha activity.

Acute aerobic exercise increases the brain cortical activity in alpha frequency. Eye closure also increases alpha activity. However, whether the two have an additive or a synergistic effect on alpha activity has never been explored. This study observed electroencephalography (EEG) from fifteen participants seated on the cycle ergometer before, during, and after a cycling exercise with the eyes open and with them closed. Exercise intensity was set to a target heart rate (120-130 bpm), corresponding to light-to-moderate intensity exercise. Each epoch was 6 min and the last 4 min (eyes closed in the first 2 min and eyes open in the second 2 min) were analyzed. The EEG power spectrum densities were calculated for alpha frequency band activity (8-13 Hz). At rest, alpha activity was significantly greater with the eyes closed than open. Exercise significantly increased alpha activity in both eye conditions. More importantly, in the occipital site, the alpha-increasing effect of their combination was significantly greater than the sum of the effect of each, showing a synergistic effect. We concluded that acute light-to-moderate intensity exercise with the eyes closed has a synergistic effect on alpha activity.

Noradrenaline modulates neuronal and perceptual visual detectability via ß-adrenergic receptor.

RATIONALE: Noradrenaline (NA) is a neuromodulator secreted from noradrenergic neurons in the locus coeruleus to the whole brain depending on the physiological state and behavioral context. It regulates various brain functions including vision via three major adrenergic receptor (AR) subtypes. Previous studies investigating the noradrenergic modulations on vision reported different effects, including improvement and impairment of perceptual visual sensitivity in rodents via ß-AR, an AR subtype. Therefore, it remains unknown how NA affects perceptual visual sensitivity via ß-AR and what neuronal mechanisms underlie it. OBJECTIVES: The current study investigated the noradrenergic modulation of perceptual and neuronal visual sensitivity via ß-AR in the primary visual cortex (V1). METHODS: We performed extracellular multi-point recordings from V1 of rats performing a go/no-go visual detection task under the head-fixed condition. A ß-AR blocker, propranolol (10 mM), was topically administered onto the V1 surface, and the drug effect on behavioral and neuronal activities was quantified by comparing pre-and post-drug administration. RESULTS: The topical administration of propranolol onto the V1 surface significantly improved the task performance. An analysis of the multi-unit activity in V1 showed that propranolol significantly suppressed spontaneous activity and facilitated the visual response of the recording sites in V1. We further calculated the signal-to-noise ratio (SNR), finding that the SNR was significantly improved after propranolol administration. CONCLUSIONS: Pharmacological blockade of ß-AR in V1 improves perceptual visual detectability by modifying the SNR of neuronal activity.

Acetylcholine from the nucleus basalis magnocellularis facilitates the retrieval of well-established memory.

Retrieval deficit of long-term memory is a cardinal symptom of dementia and has been proposed to associate with abnormalities in the central cholinergic system. Difficulty in the retrieval of memory is experienced by healthy individuals and not limited to patients with neurological disorders that result in forgetfulness. The difficulty of retrieving memories is associated with various factors, such as how often the event was experienced or remembered, but it is unclear how the cholinergic system plays a role in the retrieval of memory formed by a daily routine (accumulated experience). To investigate this point, we trained rats moderately (for a week) or extensively (for a month) to detect a visual cue in a two-alternative forced-choice task. First, we confirmed the well-established memory in the extensively trained group was more resistant to the retrieval problem than recently acquired memory in the moderately trained group. Next, we tested the effect of a cholinesterase inhibitor, donepezil, on the retrieval of memory after a long no-task period in extensively trained rats. Pre-administration of donepezil improved performance and reduced the latency of task initiation compared to the saline-treated group. Finally, we lesioned cholinergic neurons of the nucleus basalis magnocellularis (NBM), which project to the entire neocortex, by injecting the cholinergic toxin 192 IgG-saporin. NBM-lesioned rats showed severely impaired task initiation and performance. These abilities recovered as the trials progressed, though they never reached the level observed in rats with intact NBM. These results suggest that acetylcholine released from the NBM contributes to the retrieval of well-established memory developed by a daily routine.

Spatial Accuracy of Predictive Saccades Determines the Performance of Continuous Visuomotor Action.

In a table tennis rally, players perform interceptive actions on a moving ball continuously in a short time, such that the acquisition process of visual information is an important determinant of the performance of the action. However, because it is technically hard to measure gaze movement in a real game, little is known about how gaze behavior is conducted during the continuous visuomotor actions and contributes to the performance. To examine these points, we constructed a novel psychophysical experiment model enabling a continuous visuomotor task without spatial movement of any body parts, including the arm and head, and recorded the movement of the gaze and effector simultaneously at high spatiotemporal resolution. In the task, Gabor patches (target) moved one after another at a constant speed from right to left at random vertical positions on an LC display. Participants hit the target with a cursor moving vertically on the left side of the display by controlling their prehensile force on a force sensor. Participants hit the target with the cursor using a rapid-approaching movement (rapid cursor approach, RCA). Their gaze also showed rapid saccadic approaching movement (saccadic eye approach, SEA), reaching the predicted arrival point of the target earlier than the cursor. The RCA reached in or near the Hit zone in the successful (Hit) trial, but ended up away from it in the unsuccessful (Miss) trial, suggesting the spatial accuracy of the RCA determines the task's success. The SEA in the Hit trial ended nearer the target than the Miss trial. The spatial accuracy of the RCA diminished when the target disappeared 100 ms just after the end of the SEA, suggesting that visual information acquired after the saccade acted as feedback information to correct the cursor movement online for the cursor to reach the target. There was a target speed condition that the target disappearance did not compromise RCA's spatial accuracy, implying the possible RCA correction based on the post-saccadic gaze location information. These experiments clarified that gaze behavior conducted during fast continuous visuomotor actions enables online correction of the ongoing interceptive movement of an effector, improving visuomotor performance.

Serotonin improves behavioral contrast sensitivity of freely moving rats.

Serotonin (5-HT) is a neuromodulator secreted from serotonergic neurons located in the pons and upper brain stem in a behavioral context-dependent manner. The serotonergic axon terminals innervate almost the whole brain, causing modulatory actions on various brain functions including vision. Our previous study demonstrated the visual responses of neurons in the primary visual cortex (V1) of anesthetized monkeys were modulated by the activation of 5-HT receptors depending on the response magnitude, in which 5-HT2A receptor-selective agonists enhanced weak visual responses but not strong responses. This observation suggests that the activation of serotonergic receptors modulates neuronal visual information processing to improve the behavioral detectability of a stimulus. However, it remains unknown if 5-HT improves visual detectability at the behavioral level. To investigate this point, visual detectability was measured as contrast sensitivity (CS) in freely moving rats using a two-alternative forced-choice visual detection task (2AFC-VDT) combined with the staircase method. The grating contrast was decreased or increased step by step after a correct choice (hit) or incorrect choice (miss), respectively. CS was evaluated as an inverse value of the visual contrast threshold. The effect of the intraperitoneal administration of fluoxetine (FLX, 5 mg/kg), a selective serotonin reuptake inhibitor, on CS was tested. The CS of rats was significantly higher in FLX than control conditions, and the drug effect showed specificity for the spatial frequency (SF) of a grating stimulus, in which CS improvement was observed at optimal SF but not non-optimal high SF. Thus, we conclude that endogenously-secreted serotonin in the brain improves visual detectability, which may be mediated by vision-related neurons acquiring SF information of the visual stimulus.

Caffeine improves contrast sensitivity of freely moving rats.

Caffeine (1, 3, 7-trimethylxanthine) is a well-known central nervous system stimulant that affects various brain functions such as attention, memory and sensation. However, it remains unclear whether and how caffeine modulates visual ability such as contrast sensitivity (CS) and the CS-spatial frequency (SF) relationship. To investigate these points, we tested the effects of caffeine on the perceptual CS of rats under three SF conditions. CS was measured using a two-alternative forced choice (2AFC) grating detection task combined with a staircase method. Intraperitoneal administration of caffeine 30 min prior to the task improved CS in an SF-dependent manner, in which the improving effect was observed at 0.1 cycles/degree (cpd) of the optimal SF for rats but not at 0.5 or 1 cpd. We concluded that caffeine, a representative ingredient contained in foods or drinks consumed daily, leads to an improvement of perceptual visual sensitivity.

Discretion for behavioral selection affects development of habit formation after extended training in rats.

As training progresses, animals show a transition from goal-dependent behavior to goal-independent behavior (habitual responses). Habit formation is influenced by several factors, including the amount of training and action-outcome contingency. However, it remains unknown whether and how discretion for behavioral selection influences habit formation. To this end, we trained male rats in two types of two-alternative forced-choice task: visual association and nonvisual association tasks. In the first type of task, rats learned the association between reward and a visual cue, the position of which was randomly changed per trial so that rats had to make a judgmental decision about which choice delivered the reward in each trial (discreet judgment group); in the second type of task, the rats learned that a reward was delivered after either choice following task initiation (uncontrolled judgment group). To test the sensitivity to contingency manipulation, the extinction tests were conducted in short- and long-term trained groups, with the result that the overtrained rats in the uncontrolled judgment group, but not the other three groups, showed less sensitivity. To further investigate the reward sensitivity in the long-term trained groups from another perspective, we continuously and periodically altered the reward size for each trial. The rats of the discreet judgment group changed intertrial intervals depending on reward size, while this tendency was weaker in the uncontrolled judgment group. These results suggest that discreet judgment maintained goal-directed rat behavior, whereas uncontrolled judgment led to the development of habit-like behavior.

Noradrenaline Improves Behavioral Contrast Sensitivity via the ß-Adrenergic Receptor.

Noradrenaline (NA) is released from the locus coeruleus in the brainstem to almost the whole brain depending on the physiological state or behavioral context. NA modulates various brain functions including vision, but many questions about the functional role of its effects and mechanisms remain unclear. To explore these matters, we focused on three questions, 1) whether NA improves detectability of a behavior-relevant visual stimulus, 2) which receptor subtypes contribute to the NA effects, and 3) whether the NA effects are specific for visual features such as spatial frequency (SF). We measured contrast sensitivity in rats by a two-alternative forced choice visual detection task and tested the effects of NA receptor blockers in three SF conditions. Propranolol, a ß-adrenergic receptor inhibitor, significantly decreased contrast sensitivity, but neither prazosin nor idazoxan, α1- and α2-adrenergic receptor inhibitors, respectively, had an effect. This ß blocker effect was observed only at optimal SF. These results indicate that endogenous NA enhances visual detectability depending on stimulus spatial properties via mainly ß-adrenergic receptors.

Cholinergic and serotonergic modulation of visual information processing in monkey V1.

The brain dynamically changes its input-output relationship depending on the behavioral state and context in order to optimize information processing. At the molecular level, cholinergic/monoaminergic transmitters have been extensively studied as key players for the state/context-dependent modulation of brain function. In this paper, we review how cortical visual information processing in the primary visual cortex (V1) of macaque monkey, which has a highly differentiated laminar structure, is optimized by serotonergic and cholinergic systems by examining anatomical and in vivo electrophysiological aspects to highlight their similarities and distinctions. We show that these two systems have a similar layer bias for axonal fiber innervation and receptor distribution. The common target sites are the geniculorecipient layers and geniculocortical fibers, where the appropriate gain control is established through a geniculocortical signal transformation. Both systems exert activity-dependent response gain control across layers, but in a manner consistent with the receptor subtype. The serotonergic receptors 5-HT1B and 5HT2A modulate the contrast-response curve in a manner consistent with bi-directional response gain control, where the sign (facilitation/suppression) is switched according to the firing rate and is complementary to the other. On the other hand, cholinergic nicotinic/muscarinic receptors exert mono-directional response gain control without a sign reversal. Nicotinic receptors increase the response magnitude in a multiplicative manner, while muscarinic receptors exert both suppressive and facilitative effects. We discuss the implications of the two neuromodulator systems in hierarchical visual signal processing in V1 on the basis of the developed laminar structure.

Spatiotemporal characteristics of surround suppression in primary visual cortex and lateral geniculate nucleus of the cat.

In the primary visual cortex (V1), a neuronal response to stimulation of the classical receptive field (CRF) is predominantly suppressed by a stimulus presented outside the CRF (extraclassical receptive field, ECRF), a phenomenon referred to as ECRF suppression. To elucidate the neuronal mechanisms and origin of ECRF suppression in V1 of anesthetized cats, we examined the temporal properties of the spatial extent and orientation specificity of ECRF suppression in V1 and the lateral geniculate nucleus (LGN), using stationary-flashed sinusoidal grating. In V1, we found three components of ECRF suppression: (1) local and fast, (2) global and fast, and (3) global and late. The local and fast component, which resulted from within 2° of the boundary of the CRF, started no more than 10 ms after the onset of the CRF response and exhibited low specificity for the orientation of the ECRF stimulus. These spatiotemporal properties corresponded to those of geniculate ECRF suppression, suggesting that the local and fast component of V1 is inherited from the LGN. In contrast, the two global components showed rather large spatial extents ∼5° from the CRF boundary and high specificity for orientation, suggesting that their possible origin is the cortex, not the LGN. Correspondingly, the local component was observed in all neurons of the thalamocortical recipient layer, while the global component was biased toward other layers. Therefore, we conclude that both subcortical and cortical mechanisms with different spatiotemporal properties are involved in ECRF suppression.

Efficient training protocol for rapid learning of the two-alternative forced-choice visual stimulus detection task.

The potential of genetically engineered rodent models has accelerated demand for training procedures of behavioral tasks. Such training is generally time consuming and often shows large variability in learning speed between animals. To overcome these problems, we developed an efficient and stable training system for the two-alternative forced-choice (2AFC) visual stimulus detection task for freely behaving rodents. To facilitate the task learning, we introduced a spout-lever as the operandum and a three-step training program with four ingenuities: (1) a salient stimulus to draw passive attention, (2) a reward-guaranteed trial to keep motivation, (3) a behavior-corrective trial, and (4) switching from a reward-guaranteed trial to a nonguaranteed one to correct behavioral patterns. Our new training system realizes 1-week completion of the whole learning process, during which all rats were able to learn effortlessly the association between (1) lever-manipulation and reward and (2) visual stimulus and reward in a step-by-step manner. Thus, our new system provides an effective and stable training method for the 2AFC visual stimulus detection task. This method should help accelerate the move toward research bridging the visual functions measured in behavioral tasks and the contributing specific neurons/networks that are genetically manipulated or optically controlled.

Blockade of muscarinic receptors impairs the retrieval of well-trained memory.

Acetylcholine (ACh) is known to play an important role in memory functions, and its deficit has been proposed to cause the cognitive decline associated with advanced age and Alzheimer's disease (the cholinergic hypothesis). Although many studies have tested the cholinergic hypothesis for recently acquired memory, only a few have investigated the role of ACh in the retrieval process of well-trained cognitive memory, which describes the memory established from repetition and daily routine. To examine this point, we trained rats to perform a two-alternative forced-choice visual detection task. Each trial was started by having the rats pull upward a central-lever, which triggered the presentation of a visual stimulus to the right or left side of the display monitor, and then pulling upward a stimulus-relevant choice-lever located on both sides. Rats learned the task within 10 days, and the task training was continued for a month. Task performance was measured with or without systemic administration of a muscarinic ACh receptor (mAChR) antagonist, scopolamine (SCOP), prior to the test. After 30 min of SCOP administration, rats stopped manipulating any lever even though they explored the lever and surrounding environment, suggesting a loss of the task-related associative memory. Three hours later, rats were recovered to complete the trial, but the rats selected the levers irrespective of the visual stimulus, suggesting they remembered a series of lever-manipulations in association with a reward, but not association between the reward and visual stimulation. Furthermore, an m1-AChR, but not nicotinic AChR antagonist caused a similar deficit in the task execution. SCOP neither interfered with locomotor activity nor drinking behavior, while it influenced anxiety. These results suggest that the activation of mAChRs at basal ACh levels is essential for the recall of well-trained cognitive memory.

Effects of stimulus spatial frequency, size, and luminance contrast on orientation tuning of neurons in the dorsal lateral geniculate nucleus of cat.

It is generally thought that orientation selectivity first appears in the primary visual cortex (V1), whereas neurons in the lateral geniculate nucleus (LGN), an input source for V1, are thought to be insensitive to stimulus orientation. Here we show that increasing both the spatial frequency and size of the grating stimuli beyond their respective optimal values strongly enhance the orientation tuning of LGN neurons. The resulting orientation tuning was clearly contrast-invariant. Furthermore, blocking intrathalamic inhibition by iontophoretically administering γ-aminobutyric acid (GABA)A receptor antagonists, such as bicuculline and GABAzine, slightly but significantly weakened the contrast invariance. Our results suggest that orientation tuning in the LGN is caused by an elliptical classical receptive field and orientation-tuned surround suppression, and that its contrast invariance is ensured by local GABAA inhibition. This contrast-invariant orientation tuning in LGN neurons may contribute to the contrast-invariant orientation tuning seen in V1 neurons.

Cholinesterase inhibitor, donepezil, improves visual contrast detectability in freely behaving rats.

Acetylcholine (ACh) modulates neuronal activities in extensive brain regions to play an essential role in various brain functions including attention, learning and memory, and cognition. Although ACh is known to modulate information processing in the primary visual cortex (V1) in many species including rodent, its functional role in visual ability has remained unknown. We examined whether and how ACh influences behavioral contrast detectability in rat. The detectability was assessed as the contrast sensitivity (CS) to a grating stimulus. Measurements were performed in a two-alternative forced-choice task combined with a staircase method in freely behaving rats. The contrast sensitivity function of rats under the no drug condition showed a low-pass spatial frequency (SF) tuning peaking at 0.1 cycles/degree (cpd) of SF (SF(peak)) that bottomed at 0.5 cpd (SF(bottom)), which was sensitive to the stimulus size, but to neither the temporal frequency nor orientation of the stimulus. The stimulus size was correlated with the CS only at the low SF range. The effect of donepezil on the size- and SF-dependency of the CS was examined using three stimulus conditions: an easy detectability condition with large grating at SF(peak), a difficult detectability condition with small grating at SF(peak), and an upper limit SF condition with large grating at SF(bottom). Donepezil improved the CS at SF(peak), especially in the difficult detectability condition. Therefore, we conclude that ACh plays an important role in enhancing behavioral CS at sensitive SF ranges, but not in improving the upper limit of SF.

Modulation-specific and laminar-dependent effects of acetylcholine on visual responses in the rat primary visual cortex.

Acetylcholine (ACh) is secreted from cholinergic neurons in the basal forebrain to regions throughout the cerebral cortex, including the primary visual cortex (V1), and influences neuronal activities across all six layers via a form of diffuse extrasynaptic modulation termed volume transmission. To understand this effect in V1, we performed extracellular multi-point recordings of neuronal responses to drifting sinusoidal grating stimuli from the cortical layers of V1 in anesthetized rats and examined the modulatory effects of topically administered ACh. ACh facilitated or suppressed the visual responses of individual cells with a laminar bias: response suppression prevailed in layers 2/3, whereas response facilitation prevailed in layer 5. ACh effects on the stimulus contrast-response function showed that ACh changes the response gain upward or downward in facilitated or suppressed cells, respectively. Next, ACh effects on the signal-to-noise (S/N) ratio and the grating-phase information were tested. The grating-phase information was calculated as the F1/F0 ratio, which represents the amount of temporal response modulation at the fundamental frequency (F1) of a drifting grating relative to the mean evoked response (F0). In facilitated cells, ACh improved the S/N ratio, while in suppressed cells it enhanced the F1/F0 ratio without any concurrent reduction in the S/N ratio. These effects were predominantly observed in regular-spiking cells, but not in fast-spiking cells. Electrophysiological and histological findings suggest that ACh promotes the signaling of grating-phase information to higher-order areas by a suppressive effect on supragranular layers and enhances feedback signals with a high S/N ratio to subcortical areas by a facilitatory effect on infragranular layers. Thus, ACh distinctly and finely controls visual information processing in a manner that is specific for the modulation and cell type and is also laminar dependent.

Cholinergic modulation of response gain in the rat primary visual cortex.

Acetylcholine (ACh) is known to modulate neuronal activity in the rodent primary visual cortex (V1). Although cholinergic modulation has been extensively examined in vitro, far less is understood regarding how ACh modulates visual information processing in vivo. We therefore extracellularly recorded visual responses to drifting sinusoidal grating stimuli from V1 of anesthetized rats and tested the effects of ACh administered locally by microiontophoresis. ACh exerted response facilitation or suppression in individual neurons across all cortical layers without any laminar bias. We assessed ACh effects on the stimulus contrast-response function, finding that ACh increased or decreased the response to varying stimulus contrasts in proportion to the magnitude of the control response without changing the shape of the original contrast-response function, which describes response gain control but not contrast gain control. Our results indicate that ACh serves as a gain controller in the visual cortex of rodents.