The laminar pattern of proprioceptive activation in human primary motor cortex.

The primary motor cortex (M1) is increasingly being recognized for its vital role in proprioceptive somatosensation. However, our current understanding of proprioceptive processing at the laminar scale is limited. Empirical findings in primates and rodents suggest a pronounced role of superficial cortical layers, but the involvement of deep layers has yet to be examined in humans. Submillimeter resolution functional magnetic resonance imaging (fMRI) has emerged in recent years, paving the way for studying layer-dependent activity in humans (laminar fMRI). In the present study, laminar fMRI was employed to investigate the influence of proprioceptive somatosensation on M1 deep layer activation using passive finger movements. Significant M1 deep layer activation was observed in response to proprioceptive stimulation across 10 healthy subjects using a vascular space occupancy (VASO)-sequence at 7 T. For further validation, two additional datasets were included which were obtained using a balanced steady-state free precession sequence with ultrahigh (0.3 mm) in-plane resolution, yielding converging results. These results were interpreted in the light of previous laminar fMRI studies and the active inference account of motor control. We propose that a considerable proportion of M1 deep layer activation is due to proprioceptive influence and that deep layers of M1 constitute a key component in proprioceptive circuits.