a midsole body with a proximal surface and a distal surface and having a forefoot region and a heel region; wherein the midsole body has a vertical axis extending perpendicular to the proximal surface and the distal surface, a medial sidewall portion extending away from the proximal surface, and a lateral sidewall portion extending away from the proximal surface and spaced apart from the medial sidewall portion across the vertical axis; wherein the midsole body has an interior portion having a first density and an outer skin covering the interior portion, separated from the interior portion by a boundary, and having a second density that is greater than the first density; wherein the midsole body has perforated holes established by perforations through the midsole body; wherein the outer skin extends along each of the perforated holes from the proximal surface to the distal surface and forms a cylinder around each of the perforated holes; and

a plurality of proprioceptive elements, wherein each proprioceptive element is disposed in a different one of the perforated holes as an integral portion of the midsole body surrounded by the perforations and is movable along a central axis of the respective proprioceptive element in a direction toward the proximal surface under a force along the respective central axis at the distal surface;

wherein the outer skin is most resistant to compression along a length of the cylinder such that the midsole body has a greater compressive stiffness under loading along each central axis than under loading transverse to each central axis, or than under loading at an oblique angle to each central axis;

wherein a first set of the plurality of proprioceptive elements are disposed in the forefoot region, and have a third density that is different than the first density and the second density;

wherein a second set of the plurality of proprioceptive elements are disposed in the heel region, and have a fourth density that is different than the first density, the second density, and the third density; and wherein a third set of the plurality of proprioceptive elements are disposed in the medial sidewall portion and in the lateral sidewall portion; and

wherein each central axis of the plurality of proprioceptive elements disposed in the medial sidewall portion and the lateral sidewall portion is angled relative to the vertical axis and is configured to align with a direction of a lateral loading force on the midsole body at a location of the respective proprioceptive element such that as each proprioceptive element moves along the respective central axis toward the proximal surface under the lateral loading force at the distal surface, the respective proprioceptive element protrudes further at the proximal surface than when not under the lateral loading force and is thereby configured to provide tactile feedback to a wearer of the sole structure at the medial sidewall portion and the lateral sidewall portion.