The periosteum: a simple tissue with many faces, with special reference to the antler-lineage periostea.

Periosteum is a thin membrane covering bone surfaces and consists of two layers: outer fibrous layer and inner cambium layer. Simple appearance of periosteum has belied its own complexity as a composite structure for physical bone protection, mechano-sensor for sensing mechanical loading, reservoir of biochemical molecules for initiating cascade signaling, niche of osteogenic cells for bone formation and repair, and "umbilical cord" for nourishing bone tissue. Periosteum-derived cells (PDCs) have stem cell attributes: self-renewal (no signs of senescence until 80 population doublings) and multipotency (differentiate into fibroblasts, osteoblasts, chondrocytes, adipocytes and skeletal myocytes).