Bones and Cartilage

Developmental and Evolutionary Skeletal Biology

Brian K. Hall

• Publisher: Academic Press
• Year: 2014
• eBook ISBN: 9780124166851
• Edition: 2
• Topic: Biological Sciences
• Subtopic: Orthopedics
• Index: Biological Sciences

Bones and Cartilage provides the most in-depth review and synthesis assembled on the topic, across all vertebrates. It examines the function, development and evolution of bone and cartilage as tissues, organs and skeletal systems. It describes how bone and cartilage develop in embryos and are maintained in adults, how bone is repaired when we break a leg, or regenerates when a newt grows a new limb, or a lizard a new tail. The second edition of Bones and Cartilage includes the most recent knowledge of molecular, cellular, developmental and evolutionary processes, which are integrated to outline a unified discipline of developmental and evolutionary skeletal biology. Additionally, coverage includes how the molecular and cellular aspects of bones and cartilage differ in different skeletal systems and across species, along with the latest studies and hypotheses of relationships between skeletal cells and the most recent information on coupling between osteocytes and osteoclasts All chapters have been revised and updated to include the latest research. - Offers complete coverage of every aspect of bone and cartilage, with updated references and extensive illustrations - Integrates development and evolution of the skeleton, as well a synthesis of differentiation, growth and patterning - Treats all levels from molecular to clinical, embryos to evolution, and covers all vertebrates as well as invertebrate cartilages - Includes new chapters on evolutionary skeletal biology that highlight normal variation and variability, and variation outside the norm (neomorphs, atavisms) - Updates hypotheses on the origination of cartilage using new phylogenetic, cellular and genetic data - Covers stem cells in embryos and adults, including mesenchymal stem cells and their use in genetic engineering of cartilage, and the concept of the stem cell niche

Part I

Vertebrate Skeletal Tissues

Outline

Part I. Vertebrate Skeletal Tissues

Chapter 1 Vertebrate Skeletal Tissues

Chapter 2 Bone

Chapter 3 Vertebrate Cartilages

Part I. Vertebrate Skeletal Tissues

When Tess made me too weepy, I turned to the timeless serenity of the frontal bones.

A. d’A. Bellairs, 1989. The Isle of Sea Lizards. p. 93. Durrell Institute of Conservation and Ecology, University of Kent, Canterbury.

The importance for the classification of life of the fundamental differences between skeletal tissues was recognised by Aristotle (384–322 BC), who separated the cartilaginous fishes (Chondrichthyes) from the bony fishes (Osteichthyes) on the basis of the presence of a cartilage- or a bone-based endoskeleton, respectively. The four classes recognised by Linnaeus (Mammalia, Aves, Amphibia, Pisces) were recognised as representing a single group of animals with bone by J. G. C. Barsch, who, in 1788, named them Knochenthiere and Ossea. Renamed by Jean-Baptist Lamarck in 1801 as animaux à vertebres, we now know them as Vertebrates, the name given to them by Ernst Haeckel in 1866 (Nielsen, 2012).

Bone and cartilage are two of four vertebral skeletal tissues. The other two are dentine and enamel. This may seem an unlikely list. Normally, we think of cartilage and bone as skeletal tissues and of enamel and dentine as dental tissues associated with teeth. Enamel and dentine, however, arose evolutionarily as skeletal tissues in the exoskeletons of early jawless and toothless vertebrates. These four tissues are distinguished by the nature of their cells and by the nature of the extracellular matrices produced by those cells. Osteoblasts and osteocytes are bone-forming cells. Chondroblasts, chondrocytes and hypertrophic chondrocytes are cartilage-forming cells. Odontoblasts synthesise and deposit dentine matrix. Ameloblasts synthesise and deposit enamel matrix.

Chapter 1 introduces the four skeletal tissues and the four cell types, traces their origins to the outset of vertebrate evolution 450 million years ago, examines the concept of vertebrate endo- and exoskeletons and introduces four tissues – cementum, enameloid, chondroid, chondroid bone – that are intermediate between bone and cartilage, bone and dentine and dentine and enamel.

Chapter 2 is devoted to bone as a skeletal tissue, including discovery of the basic structure of bone, distinctions between cellular and acellular bone, the features and gene products of osteocytes and the processes of intramembranous and endochondral ossification. Growth, modelling, remodelling and ageing of bone are discussed, as is the significance of the presence of bone in sharks and rays.

Chapter 3 is devoted to the three types of vertebrate cartilages (hyaline, elastic, fibrous), chondrones as fundamental structural units of cartilage, how cartilage grows, cartilage in sharks and rays and the specialised cartilage and proteins found in lampreys and hagfish.

Chapter 1

Vertebrate Skeletal Tissues

In this chapter I introduce, define and provide the major characteristics of the four vertebrate mineralised tissues – cartilage, bone, enamel and dentine – and the four cell types that produce them – chondroblasts, osteoblasts, ameloblasts and odontoblasts. All four began as skeletal tissues/cells at the outset of vertebrate evolution. I define and discuss the endoskeleton and the exoskeleton. I also introduce four tissues (cementum, enameloid, chondroid, chondroid bone) that are intermediate between the four skeletal tissues.

Keywords

bone; cartilage; chondroblasts; chondrocytes; chondroid; chondroid bone; cilia; collagens; dentine; direct ossification; enamel; enameloid; endochondral ossification; endoskeleton; evolution; exoskeleton; Haversian canals; indirect ossification; intermediate tissues; intramembranous ossification; mineralised tissues; modes of ossification; osteoblasts; osteocalcin; osteoclasts; osteocytes; osteopontin; skeletal tissues; teeth

In the pioneering stages of Natural Science we recognize the work of collecting, describing and classifying the typical units as a fundamental necessity. The study of their morphology and their history belongs to a more advanced period.

J. M. Petrie, 1914, Handbook for New South Wales, British Association for the Advancement of Science, Sydney, Australia, cited from L. Gilbert, 1997, p. 283.

Historically, the distinction between cartilage and bone has been recognised since at least the time of Aristotle (384–322 BC), who recognised and separated the cartilaginous fishes (Chondrichthyes) from the bony fishes (Osteichthyes) by the presence of either a cartilaginous or an osseous skeleton. Two thousand fifty years later, the oldest articulated chondrichthyan, the 50- to 75-cm-long Doliodus problematicus, was described from the Early Devonian (409 million years ago [mya]) of New Brunswick, Canada. Previously known only from isolated dermal scales and teeth, this specimen includes the braincase, prismatic mineralised cartilage and paired spines associated with the pectoral fins, previously unknown as a feature of cartilaginous fishes (R. F. Miller et al., 2003; Turner and Miller, 2005; and see Chapter 3).

Skeletal tissues are even more ancient, with origins reaching back perhaps three-quarters of a billion years. Nevertheless, the number of skeletal tissues or organs is limited. Thomas et al. (2000) evaluated 182 characters of skeletal design as possible design options in morphospace, that is, as a three-dimensional (3D) representation of the distribution of all known morphologies. Of these 182 characters, 146 were already in use in animals of the Burgess Shale fauna 530 mya. Indeed, within 15 million years of the appearance of the crown groups of the major phyla in the fossil record, 80% of the design element...

Table of contents

• Cover image
• Title page
• Table of Contents
• Copyright
• Preface
• Epigraphs
• Part I: Vertebrate Skeletal Tissues
• Part II: Origins and Types of Skeletal Tissues
• Part III: Unusual Modes of Skeletogenesis
• Part IV: Stem and Progenitor Cells
• Part V: Skeletogenic Cells
• Part VI: Embryonic Origins
• Part VII: Getting Started
• Part VIII: Similarity and Diversity
• Part IX: Maintaining Cartilage in Good Times and in Bad
• Part X: Growing Together and Growing Apart
• Part XI: Staying Apart
• Part XII: Limb Buds
• Part XIII: Limbs and Limb Skeletons
• Part XIV: Backbones and Tails
• Part XV: Evolutionary Skeletal Biology
• References
• Index