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Connective and supportive tissues (textus connectivus)  87



                  matrix. The collagen fibres are arranged in spirals along the  tion as part of lifelong bone remodelling. Loss of bone
       VetBooks.ir  long axis of the central canal. In alternate lamellae, the spiral  mass with advancing age is associated with a reduction in
                  collagen fibres are oriented in different directions, forming a  osteocytes and delayed mineralisation.
                  criss-crossing network. Cross-linkages are formed between
                                                                    At the internal and external surfaces of the bone, the
                  adjacent lamellae. These features impart stability to the  lamellae are arranged in circular sheets, forming the inter-
                  bone under forces of pressure and tension.     nal circumferential lamellae (adjacent to the endosteum)
                     In additional to its structural role, lamellar bone – like  and external circumferential lamellae (lying against the
                  all tissues derived from connective tissue – also plays a sig-  internal surface of the periosteum). Collagen fibres that
                  nificant part in metabolism. This is enabled in large part  bind the periosteum to the bone (fibrae perforantes,
                  by functional interactions between the bone cells, blood  Sharpey’s fibres) are incorporated into the external cir-
                  vessels and connective tissue within the osteon.  cumferential lamellae. The endosteum, composed of
                     Osteocytes  within lacunae are regularly arranged  osteoprogenitor cells and connective tissue, lines the inter-
                  between the concentric lamellae around the central canal  nal circumferential lamellae and the trabeculae (Figure
                  (Figures 3.34 to 3.37). Their long cytoplasmic processes  3.34).
                  radiate into interconnecting canaliculi (canaliculi ossei)   Trabecular (spongy) bone also contains lamellae, but
                  extending from the lacunae. The cellular extensions estab-  these are not organised into osteonal systems. Remodelling
                  lish contact with the processes of other osteocytes via  processes occur at a particularly high rate in spongy
                  gap junctions. This arrangement permits the transport  bone.
                  of substances between the blood vessel in the Haversian
                  canal and the bone matrix (both to and from the vessel).  New bone formation (osteogenesis)
                  Channels running transversely through the bone (perfo-  The formation of new bone occurs in two ways.
                  rating canals, Volkmann’s canals) connect central canals  Development of bone directly from mesenchymal con-
                  with each other, and with the endosteum and periosteum.  nective tissue without a cartilaginous precursor phase is
                  Through this interconnecting network of blood vessels,  referred to as intramembranous ossification (primary or
                  bone is a well-vascularised tissue (Figure 3.34).  direct ossification). This type of bone formation gives rise
                     Lamellar bone serves as a metabolically active calcium  to certain flat bones of the skull and the bony collar of
                  depot. Under the influence of parathyroid hormone, acti-  developing long bones. It is also observed in repairing bone
                  vation of osteolytic osteocytes or, in larger areas of bone  fractures. Formation of bone based on a cartilaginous tem-
                  resorption, multinuclear osteoclasts, within bone lacunae  plate is termed endochondral ossification (secondary
                                                      2+
                  and canaliculi results in rapid liberation of Ca  and PO   or indirect ossification). This results first in (immature)
                                                              3−
                                                             4
                  ions from the mineralised matrix. These ions are swiftly   woven bone that is gradually replaced by (mature) lamellar
                  conveyed to the circulation by the intraosseous transport   bone (sometimes referred to as replacement bone) (Figures
                  system.                                        3.40 to 3.42).
                     Any change in the mechanical load on a bone results
                  in dynamic adaptation of its internal structure. Non-  INTRAMEMBRANOUS OSSIFICATION
                  functional osteons are broken down, leaving remnants  In intramembranous ossification, osteoprogenitor cells
                  referred to as interstitial lamellae (Figures 3.34 and 3.35).  differentiate into osteoblasts (Figures 3.38 and 3.39) that
                  The structure of compact bone undergoes gradual altera-  produce collagen fibres and osteoid. Through continued













                                                                            3.38  Intramembranous ossification:
                                                                            bone of cranium (juvenile dog). Near
                                                                            capillaries, osteoprogenitor cells within
                                                                            loose connective tissue differentiate into
                                                                            osteoblasts, which become arranged in
                                                                            sheets and liberate osteoid. As the bone
                                                                            matrix becomes mineralised, osteoblasts
                                                                            transform into osteocytes. Goldner’s
                                                                            Masson trichrome stain (x480).









       Vet Histology.indb   87                                                                                   16/07/2019   14:56
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