Recent in vitro studies have shown that acetylcholine (ACh) selectively reduces the efficacy of lateral cortical connections via a muscarinic mechanism, while boosting the efficacy of thalamocortical/feed-forward connections via a nicotinic mechanism. This suggests that high levels of ACh should reduce center-surround interactions of neurons in primary visual cortex, making cells more reliant on feed-forward information. In line with this hypothesis, we show that local iontophoretic application of ACh in primate primary visual cortex reduced the extent of spatial integration, assessed by recording a neurons' length tuning. When ACh was externally applied, neurons' preferred length shifted toward shorter bars, showing reduced impact of the extra-classical receptive field. We fitted a difference and a ratio of Gaussian model to these data to determine the underlying mechanisms of this dynamic change of spatial integration. These models assume overlapping summation and suppression areas with different widths and gains to be responsible for spatial integration and size tuning. ACh significantly reduced the extent of the summation area, but had no significant effect on the extent of the suppression area. In line with previous studies, we also show that applying ACh enhanced the response in the majority of cells, especially in the later (sustained) part of the response. These findings are similar to effects of attention on neuronal activity. The natural release of ACh is strongly linked with states of arousal and attention. Our results may therefore be relevant to the neurobiological mechanism of attention.
Cortical processing is strongly influenced by the actions of neuromodulators such as acetylcholine (ACh). Early studies in anaesthetized cats argued that acetylcholine can cause a sharpening of orientation tuning functions and an improvement of the signal-to-noise ratio (SNR) of neuronal responses in primary visual cortex (V1). Recent in vitro studies have demonstrated that acetylcholine reduces the efficacy of feedback and intracortical connections via the activation of muscarinic receptors, and increases the efficacy of feed-forward connections via the activation of nicotinic receptors. If orientation tuning is mediated or enhanced by intracortical connections, high levels of acetylcholine should diminish orientation tuning. Here we investigate the effects of acetylcholine on orientation tuning and neuronal responsiveness in anaesthetized marmoset monkeys. We found that acetylcholine caused a broadening of the orientation tuning in the majority of cells, while tuning functions became sharper in only a minority of cells. Moreover, acetylcholine generally facilitated neuronal responses, but neither improved signal-to-noise ratio, nor reduced trial-to-trial firing rate variance systematically. Acetylcholine did however, reduce variability of spike occurrences within spike trains. We discuss these findings in the context of dynamic control of feed-forward and lateral/feedback connectivity by acetylcholine.
Face perception plays a crucial role in primate social communication. We have investigated the pattern of eye movements produced by rhesus monkeys (Macaca mulatta) as they viewed images of faces. Eye positions were recorded accurately using implanted eye coils, while neutral upright, inverted and scrambled images of monkey and human faces were presented on a computer screen. The monkeys exhibited a similar eye scan pattern while viewing familiar and unfamiliar monkey face images, or while viewing monkey and human face images. No differences were observed in the distribution of viewing times, number of fixations, time into the trial of first saccade to local facial features, and the temporal and spatial characteristics of viewing patterns across the facial images. However, there was a greater probability of re-fixation of the eye region of unfamiliar faces during the first few seconds of the trial suggesting that the eyes are important for the initial encoding of identity. Indeed, the highest fixation density was found in the eye region of all the face images. The viewing duration and the number of fixations per image decreased when inverted or scrambled faces were presented. The eye region in these modified images remained the primary area of fixation. However, the number of fixations directed to the eyes decreased monotonically from the upright images through the inverted versions to the scrambled face images. Nonetheless, the eyes remain the most salient facial substructure regardless of the arrangement of other features, although the extent of salience which they attain may depend both on the low level properties of the eyes and on the global arrangement of facial features.
This paper mainly concerns the essential normality of graded submodules. Essentially all of the basic Hilbert modules that have received attention over the years are p-essentially normal-including the d-shift Hilbert module, the Hardy and Bergman modules of the unit ball. Arveson conjectured graded submodules over the unit ball inherit this property and provided motivations to seek an affirmative answer. Some positive results have been obtained by Arveson and Douglas. However, the problem has been resistant. In dimensions d = 2, 3, this paper shows that the Arveson's conjecture is true. In any dimension, the paper also gives an affirmative answer in the case of the graded principal submodule. Finally, the paper is associated with K -homology invariants arising from graded quotient modules, by which geometry of the quotient modules and geometry of algebraic varieties are connected. In dimensions d = 2, 3, it is shown that K -homology invariants determined by graded quotients are nontrivial. The paper also establishes results on p-smoothness of K -homology elements, and gives an explicit expression for K -homology invariant in dimension d = 2.
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