The role of medial parieto occipital cortex in visuospatial attention and reach planning: electrophysiological studies in human and non-human primates

We usually perform actions in a dynamic environment and changes in the location of a target for an upcoming action require both covert shifts of attention and motor planning update. In this study we tested whether, similarly to oculomotor areas that provide signals for overt and covert attention shifts, covert attention shifts modulate activity in cortical area V6A, which provides a bridge between visual signals and arm-motor control. We performed single cell recordings in monkeys trained to fixate straight-ahead while shifting attention outward to a peripheral cue and inward again to the fixation point. We found that neurons in V6A are influenced by spatial attention demonstrating that visual, motor, and attentional responses can occur in combination in single neurons of V6A. This modulation in an area primarily involved in visuo-motor transformation for reaching suggests that also reach-related regions could directly contribute in the shifts of spatial attention necessary to plan and control goal-directed arm movements. Moreover, to test whether V6A is causally involved in these processes, we have performed a human study using on-line repetitive transcranial magnetic stimulation over the putative human V6A (pV6A) during an attention and a reaching task requiring covert shifts of attention and reaching movements towards cued targets in space. We demonstrate that the pV6A is causally involved in attention reorienting to target detection and that this process interferes with the execution of reaching movements towards unattended targets. The current findings suggest the direct involvement of the action-related dorso-medial visual stream in attentional processes, and a more specific role of V6A in attention reorienting. Therefore, we propose that attention signals are used by the V6A to rapidly update the current motor plan or the ongoing action when a behaviorally relevant object unexpectedly appears at an unattended location.

The posterior parietal cortex: a bridge between vision and action

The present work takes into account three posterior parietal areas, V6, V6A, and PEc, all operating on different subsets of signals (visual, somatic, motor). The work focuses on the study of their functional properties, to better understand their respective contribution in the neuronal circuits that make possible the interactions between subject and external environment. In the caudalmost pole of parietal lobe there is area V6. Functional data suggest that this area is related to the encoding of both objects motion and ego-motion. However, the sensitivity of V6 neurons to optic flow stimulations has been tested only in human fMRI experiments. Here we addressed this issue by applying on monkey the same experimental protocol used in human studies. The visual stimulation obtained with the Flow Fields stimulus was the most effective and powerful to activate area V6 in monkey, further strengthening this homology between the two primates. The neighboring areas, V6A and PEc, show different cytoarchitecture and connectivity profiles, but are both involved in the control of reaches. We studied the sensory responses present in these areas, and directly compared these.. We also studied the motor related discharges of PEc neurons during reaching movements in 3D space comparing also the direction and depth tuning of PEc cells with those of V6A. The results show that area PEc and V6A share several functional properties. Area PEc, unlike V6A, contains a richer and more complex somatosensory input, and a poorer, although complex visual one. Differences emerged also comparing the motor-related properties for reaches in depth: the incidence of depth modulations in PEc and the temporal pattern of modulation for depth and direction allow to delineate a trend among the two parietal visuomotor areas.

Essays in Economic Geography and Long-Term Development

This dissertation has two main themes: first, the economic impact of tourism on cities and, secondly, the determinants of European long-run development, with a focus on the pre-Industrial era. The common thread is the attempt to develop economic geography models that incorporate spatial frictions and are liable to be given empirical content. Chapter 1, written in conjunction with G. Alfredo Minerva, provides an empirical analysis of the relationship between tourism and economic activity across Italian municipalities, and lays down the basic elements of an urban theory of tourism in an a-spatial setting. Chapter 2 extends these ideas to a quantitative urban framework to study the economic impact and the welfare consequences of tourism into the city of Venice. The model is given empirical content thanks to a large collection of data at the Census tract level for the Municipality of Venice, and then used to perform counterfactual policty analysis. In chapter 3, with Matteo Santacesaria, we consider a setting where agents are continuously distributed over a two-dimensional heterogeneous geography, and are allowed to do business at a finite set of markets. We study the equilibrium partition of the economic space into a collection of mutually-exclusive market areas, and provide condition for this equilibrium partition to exist and to be unique. Finally, chapter 4 "The rise of (urban) Europe: a Quantitative-Spatial analysis", co-authored with Matteo Cervellati and Alex Lehner, sets up a quantitative economic geography model to understand the roots of the Industrial Revolution, in an attempt to match the evolution of the European urban network, and the corresponding city-size distribution, over the period A.D. 1000-1850. It highlights the importance of agricultural trade across cities for the emergence of large manufacturing hubs.