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Skeletal structure  117




                                                 forces                morphogens


                                                     mesenchymal stem cells


                                 osteogenesis    chondrogenesis    myogenesis    ligamentagenesis




                                  osteoblast      chondroctye       myoblast





                                  osteocyte       hypertrophic      myotube          ligament
                                                  chondrocyte                       fibroblast
                                    bone           cartilage        muscle          ligament

                                 Figure 7.3. All skeletal tissues arise from a single cell type, the mesenchymal stem
                                 cell. Differentiation into bone, cartilage, muscle, or ligament occurs in response to
                                 the mechanical and biochemical stimuli of the stem cell’s environment.


                                 osteoblasts. Upon activation, osteoblasts secrete osteoid, the organic extra-
                                 cellular matrix into which mineral crystals are deposited. The organic
                                 matrix contains 90 per cent collagen and a ground substance consisting of
                                 large protein polysaccharides and a variety of matrix proteins. Gaps in the
                                 collagen fibrils serve as mineral nucleation sites for calcium phosphate,
                                 which forms the inorganic phase. The result is a composite of a ductile
                                 polymeric (collagen) phase and a strong ceramic phase. This combination
                                 gives bone its high mechanical strength and toughness.
                                    Approximately 15 per cent of osteoblasts become entrapped in their
                                 own matrix to become osteocytes. Osteocytes have a vast three-dimen-
                                 sional network of cell processes (canaliculi), providing nourishment and
                                 cell–cell interactions. Because they are located throughout bone tissue and
                                 have an extensive canalicular network, osteocytes are assumed to be a vital
                                 component of sensing mechanical signals. Nutrients are essential for the
                                 vitality of bone tissue and are obtained from the blood supply, limiting
                                 most osteocytes to lie within 150 m of a blood vessel, resulting in a high
                                 cellular density: 25000 osteocytes within a square millimetre of bone
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