Is there any role of membrane bilayer-skeleton interaction in maintaining the transmembrane phospholipid asymmetry in erythrocytes?

Erythrocyte membrane phospholipids are asymmetrically distributed in two surfaces of the membrane bilayer. This asymmetry in these cells, on one hand, has been considered to arise from the membrane skeleton-bilayer interactions, while on the other, it has been thought to originate from an ATP-dependent aminophospholipid pump. A critical analysis of these two proposals, in the light of the existing literature, reveals that neither the membrane skeleton nor the aminophospholipid pump is adequate per se to maintain the phospholipid asymmetry. Instead, evidence is presented to show that the phospholipid pump together with the membrane skeleton is required for generation and maintenance of the transmembrane phospholipid asymmetry in native erythrocytes