Two brains in action: joint-action coding in the primate frontal cortex

Daily life often requires the coordination of our actions with those of another partner. After sixty years (1968-2018) of behavioral neurophysiology of motor control, the neural mechanisms which allow such coordination in primates are unknown. We studied this issue by recording cell activity simultaneously from dorsal premotor cortex (PMd) of two male interacting monkeys trained to coordinate their hand forces to achieve a common goal. We found a population of 'joint-action cells' that discharged preferentially when monkeys cooperated in the task. This modulation was predictive in nature, since in most cells neural activity led in time the changes of the "own" and of the "other" behavior. These neurons encoded the joint-performance more accurately than 'canonical action-related cells', activated by the action per se, regardless of the individual vs. interactive context. A decoding of joint-action was obtained by combining the two brains activities, using cells with directional properties distinguished from those associated to the 'solo' behaviors. Action observation-related activity studied when one monkey observed the consequences of the partner's behavior, i.e. the cursor's motion on the screen, did not sharpen the accuracy of 'joint-action cells' representation, suggesting that it plays no major role in encoding joint-action. When monkeys performed with a non-interactive partner, such as a computer, 'joint-action cells' representation of the "other" (non-cooperative) behavior was significantly degraded. These findings provide evidence of how premotor neurons integrate the time-varying representation of the self-action with that of a co-actor, thus offering a neural substrate for successful visuo-motor coordination between individuals.SIGNIFICANT STATEMENTThe neural bases of inter-subject motor coordination were studied by recording cell activity simultaneously from the frontal cortex of two interacting monkeys, trained to coordinate their hand forces to achieve a common goal. We found a new class of cells, preferentially active when the monkeys cooperated, rather than when the same action was performed individually. These 'joint-action neurons' offered a neural representation of joint-behaviors by far more accurate than that provided by the canonical action-related cells, modulated by the action per se regardless of the individual/interactive context. A neural representation of joint-performance was obtained by combining the activity recorded from the two brains. Our findings offer the first evidence concerning neural mechanisms subtending interactive visuo-motor coordination between co-acting agents

Temporal Evolution and Strength of Neural Activity in Parietal Cortex during Eye and Hand Movements

The role of area 7a in eye-hand movement was studied by recording from individual neurons while monkeys performed 7 different tasks, aimed at assessing the relative influence of retinal, eye, and hand information on neural activity. Parietal cell activity was modulated by visuospatial signals about target location, as well as by information concerning eye and/or hand movement, and position. The highest activity was elicited when the hand moved to the fixation point. The population activities across different memory tasks showed common temporal peaks when aligned to the visual instruction (visuospatial peak) or Go signal (motor peak) for eye, hand, and coordinated eye-hand movement. The motor peak was higher for coordinated eye-hand movement, and it was absent in a No-Go task. Two activation maxima were also observed during visual reaching. They had the same latency of the visuospatial and motor peaks seen in the memory tasks. Therefore, area 7a seems to operate through a common neural mechanism underlying eye, hand, or combined eye-hand movement. This mechanism is revealed by invariant temporal activity profiles and is independent from the effector selected and from the presence or absence of a visible target during movement. For comparative purposes, we have studied the temporal evolution of the population activity in the superior parietal lobule (SPL) during the same reaching tasks and during a saccade task. In SPL, the population activity was characterized by a single peak, time locked to the Go signal for eye, hand, or combined eye-hand movement. As in IPL, the time of occurrence of this peak was effector independent. The population activity remained unchanged when the position of the eye changed, suggesting that SPL is mostly devoted to the hand motor behavior

Decoding of path-guided apparent motion from neural ensembles in posterior parietal cortex

We compared quantitatively the psychometric capacity of human subjects to detect path-guided apparent motion (PAM) and the accuracy of cell ensembles in area 7a to code the same type of stimuli. Nine human subjects performed a detection task of PAM. They were instructed to indicate with a key-press whether they perceived a circularly moving object when five stimuli were flashed successively at the vertices of a regular pentagon. The stimuli were presented along a low contrast circular path with one of 33 speeds (150-600°/s). The average psychometric curve revealed that the threshold for PAM detection was 314°/s. The minimum and maximum thresholds for individual subjects were 277° and 378°/s, respectively. In addition, the activity of cells in area 7a that were modulated by the stimulus position in real or apparent motion was used in a multivariate linear regression analysis to recover the stimulus position over time. Real stimulus motion was decoded successfully from neural ensemble activity at all speeds. In contrast, the decoding of PAM was poor at low stimulus speeds but improved markedly above 300°/s: in fact, this was very close to the threshold above for human subjects to perceive continuous stimulus motion in this condition. These results suggest that the posterior parietal cortex is part of a high-level system that is directly involved in the dynamic representation of complex motion. © Springer-Verlag 2004

Cortico-spinal modularity in the parieto-frontal system: a new perspective on action control

: Classical neurophysiology suggests that the motor cortex (MI) has a unique role in action control. In contrast, this review presents evidence for multiple parieto-frontal spinal command modules that can bypass MI. Five observations support this modular perspective: (i) the statistics of cortical connectivity demonstrate functionally-related clusters of cortical areas, defining functional modules in the premotor, cingulate, and parietal cortices; (ii) different corticospinal pathways originate from the above areas, each with a distinct range of conduction velocities; (iii) the activation time of each module varies depending on task, and different modules can be activated simultaneously; (iv) a modular architecture with direct motor output is faster and less metabolically expensive than an architecture that relies on MI, given the slow connections between MI and other cortical areas; (v) lesions of the areas composing parieto-frontal modules have different effects from lesions of MI. Here we provide examples of six cortico-spinal modules and functions they subserve: module 1) arm reaching, tool use and object construction; module 2) spatial navigation and locomotion; module 3) grasping and observation of hand and mouth actions; module 4) action initiation, motor sequences, time encoding; module 5) conditional motor association and learning, action plan switching and action inhibition; module 6) planning defensive actions. These modules can serve as a library of tools to be recombined when faced with novel tasks, and MI might serve as a recombinatory hub. In conclusion, the availability of locally-stored information and multiple outflow paths supports the physiological plausibility of the proposed modular perspective

A view-based decision mechanism for rewards in the primate amygdala

Primates make decisions visually by shifting their view from one object to the next, comparing values between objects, and choosing the best reward, even before acting. Here, we show that when monkeys make value-guided choices, amygdala neurons encode their decisions in an abstract, purely internal representation defined by the monkey’s current view but not by specific object or reward properties. Across amygdala subdivisions, recorded activity patterns evolved gradually from an object-specific value code to a transient, object-independent code in which currently viewed and last-viewed objects competed to reflect the emerging view-based choice. Using neural-network modeling, we identified a sequence of computations by which amygdala neurons implemented view-based decision making and eventually recovered the chosen object’s identity when the monkeys acted on their choice. These findings reveal a neural mechanism in the amygdala that derives object choices from abstract, view-based computations, suggesting an efficient solution for decision problems with many objects

The orientation and kinematics of inner tidal tails around dwarf galaxies orbiting the Milky Way

Using high-resolution collisionless N-body simulations we study the properties of tidal tails formed in the immediate vicinity of a two-component dwarf galaxy evolving in a static potential of the Milky Way (MW). The stellar component of the dwarf is initially in the form of a disk and the galaxy is placed on an eccentric orbit motivated by CDM-based cosmological simulations. We measure the orientation, density and velocity distribution of the stars in the tails. Due to the geometry of the orbit, in the vicinity of the dwarf, where the tails are densest and therefore most likely to be detectable, they are typically oriented towards the MW and not along the orbit. We report on an interesting phenomenon of `tidal tail flipping': on the way from the pericentre to the apocentre the old tails following the orbit are dissolved and new ones pointing towards the MW are formed over a short timescale. We also find a tight linear relation between the velocity of stars in the tidal tails and their distance from the dwarf. Using mock data sets we demonstrate that if dwarf spheroidal (dSph) galaxies in the vicinity of the MW are tidally affected their kinematic samples are very likely contaminated by tidally stripped stars which tend to artificially inflate the measured velocity dispersion. The effect is stronger for dwarfs on their way from the peri- to the apocentre due to the formation of new tidal tails after pericentre. Realistic mass estimates of dSph galaxies thus require removal of these stars from kinematic samples.Comment: 8 pages, 7 figures, accepted for publication in MNRA

The complex hodological architecture of the macaque dorsal intraparietal areas as emerging from neural tracers and dw-mri tractography

In macaque monkeys, dorsal intraparietal areas are involved in several daily visuomotor actions. However, their border and sources of cortical afferents remain loosely defined. Combining retrograde histologic tracing and MRI diffusion-based tractography, we found a complex hodology of the dorsal bank of the intraparietal sulcus (db-IPS), which can be subdivided into a rostral intraparietal area PEip, projecting to the spinal cord, and a caudal medial intraparietal area MIP lacking such projections. Both include an anterior and a posterior sector, emerging from their ipsilateral, gradient-like connectivity profiles. As tractography estimations, we used the cross-sectional area of the white matter bundles connecting each area with other parietal and frontal regions, after selecting regions of interest (ROIs) corresponding to the injection sites of neural tracers. For most connections, we found a significant correlation between the proportions of cells projecting to all sectors of PEip and MIP along the continuum of the db-IPS and tractography. The latter also revealed “false positive” but plausible connections awaiting histologic validation

The complex hodological architecture of the macaque dorsal intraparietal areas as emerging from neural tracers and DW-MRI tractography

In macaque monkeys, dorsal intraparietal areas are involved in several daily visuo-motor actions. However, their border and sources of cortical afferents remain loosely defined. Combining retrograde histological tracing and MRI diffusion-based tractography we found a complex hodology of the dorsal bank of the IPS, which can be subdivided into a rostral area PEip, projecting to the spinal cord, and a caudal area MIP lacking such projections. Both include a rostral and a caudal sector, emerging from their ipsilateral, gradient-like connectivity profiles. As tractography estimations, we used the cross-sectional volume of the white matter bundles connecting each area with other parietal and frontal regions, after selecting ROIs corresponding to the injection sites of neural tracers. For most connections, we found a significant correlation between the proportions of cells projecting to all sectors of PEip and MIP along the continuum of the dorsal bank of the IPS and tractography. The latter also revealed “false positive” but plausible streamlines awaiting histological validation

A wide-area view of the Phoenix dwarf galaxy from VLT/FORS imaging

We present results from a wide-area photometric survey of the Phoenix dwarf galaxy, one of the rare dwarf irregular/ dwarf spheroidal transition type galaxies (dTs) of the Local Group (LG). These objects offer the opportunity to study the existence of possible evolutionary links between the late- and early- type LG dwarf galaxies, since the properties of dTs suggest that they may be dwarf irregulars in the process of transforming into dwarf spheroidals. Using FORS at the VLT we have acquired VI photometry of Phoenix. The data reach a S/N~10 just below the horizontal branch of the system and consist of a mosaic of images that covers an area of 26' x 26' centered on the coordinates of the optical center of the galaxy. Examination of the colour-magnitude diagram and luminosity function revealed the presence of a bump above the red clump, consistent with being a red giant branch bump. The deep photometry combined with the large area covered allows us to put on a secure ground the determination of the overall structural properties of the galaxy and to derive the spatial distribution of stars in different evolutionary phases and age ranges, from 0.1 Gyr to the oldest stars. The best-fitting profile to the overall stellar population is a Sersic profile of Sersic radius R_S = 1.82'+-0.06' and m=0.83+-0.03. We confirm that the spatial distribution of stars is found to become more and more centrally concentrated the younger the stellar population, as reported in previous studies. This is similar to the stellar population gradients found for close-by Milky Way dwarf spheroidal galaxies. We quantify such spatial variations by analyzing the surface number density profiles of stellar populations in different age ranges; [Abridged]Comment: 21 pages; 11 figures. Accepted for publication in MNRA

The ACS LCID Project: On the origin of dwarf galaxy types: a manifestation of the halo assembly bias?

We discuss how knowledge of the whole evolutionary history of dwarf galaxies, including details on the early star formation events, can provide insight on the origin of the different dwarf galaxy types. We suggest that these types may be imprinted by the early conditions of formation rather than being only the result of a recent morphological transformation driven by environmental effects. We present precise star formation histories of a sample of Local Group dwarf galaxies, derived from colour-magnitude diagrams reaching the oldest main-sequence turnoffs. We argue that these galaxies can be assigned to two basic types: fast dwarfs that started their evolution with a dominant and short star formation event, and slow dwarfs that formed a small fraction of their stars early and have continued forming stars until the present time (or almost). These two different evolutionary paths do not map directly onto the present-day morphology (dwarf spheroidal vs dwarf irregular). Slow and fast dwarfs also differ in their inferred past location relative to the Milky Way and/or M31, which hints that slow dwarfs were generally assembled in lower density environments than fast dwarfs. We propose that the distinction between a fast and slow dwarf galaxy reflects primarily the characteristic density of the environment where they form. At a later stage, interaction with a large host galaxy may play a role in the final gas removal and ultimate termination of star formation.Comment: 7 pages, 3 figures, ApJ Letters, submitted. Comments welcom