Introduction to the Skeleton and the Head Skeleton

KK pp. 177-183, H&G 114-115

The importance of the skeleton to zoologists

Durable part of the fossil record. Conservative enough to show broad outlines of vertebrate phylogeny. Plastic enough to respond to specific adaptations good for identifying animals. Informs us about the soft parts of animals through scars of muscle attachments, traces of pathways of nerves, blood vessels, etc.

Vertebrate Connective Tissues

Kardong uses a very broad definition of connective tissue that includes cartilage, bone and blood. (H&G does not.) Their common property is that they are composed of cells in an extracellular matrix, which can be liquid, gel-like or hard.

KK 5.19

Fibrous Connective Tissue

KK 177-178 A protein matrix containing fibroblasts that produce collagen and elastic fibers. Derived from mesenchyme. Periosteum connective tissue that surrounds bone Perichondrium - connective tissue that surrounds cartilage Perimyseum - connective tissue that surrounds muscle

Tendons - connect muscles to bones Ligaments connect bones to bones Aponeuroses and fasciae connect muscles broadly to other tissues

Cartilage KK p. 178, H&G 114

A glycoprotein matrix produced by chrondrocytes. May contain collagen and elastic fibers. Usually from mesoderm. Hyaline cartilage translucent cartilage that caps bones, has few fibers. Fibrocartilage contains lots of collagen fibers, e.g., intervertebral discs of mammals. Elastic cartilage contains lots of elastic fibers, e.g., external ear or pinna. Calcified cartilage calcium salts make this cartilage hard e.g., shark skeleton.
Hyaline cartilage

Bone KK 187, H&G 14-115

Bone is a matrix of hydroxyapatite (calcium phosphate) containing osteocytes (bone cells) and collagen fibers. May be compact or spongy, and contains nerves and blood vessels. It is a living tissue, modifying itself according to the stresses put on it. Dermal Bone Develops from the dermatome and mesenchyme and is essentially external and protective. It is sometimes called membrane bone, although this term may also be applied to the external layers of endochondral bones. Endochondral Bone Formed by the replacement of embryonic cartilage by ossification from within, and from the outside by the periosteum. Derived from mesenchyme, schleratome, and neural crest cells.

KK 5.23, H&G 6.3

Structure of Bone
Outer layer of lamellar bone laid down by periosteum.

Inner network of canals constantly being reworked by bone osteocytes.

Interior of spongy or cancellous bone.

Formation of an endochondral bone from embryonic cartilage.

Ossification (in black) begins on the diaphysis (shaft) from the periosteum and then internally from the ends (epiphyses) of the bone. Cartilage remains on the articular surface of the epiphyses.
Why are embryonic bones made of cartilage?

KK 5.24, H&G 9.24

The Head Skeleton

Chondrocranium cartilage or endochrondal bone enclosing and protecting the brain and special sense organs. Splanchnocranium - cartilage or endochondral bone supporting the gills and their derivatives. Dermatocranium shield-like dermal bones which form a protective covering. The dermatocranium is not endochondral bone and does not begin as cartilage.

The Chondrocranium in Chondrichthyes

Among adult vertebrates, the chondrocranium is most complete in Chondrichthyes, which lack the dermal components of the skull.

KK 7.18b, H&G 8.9

In these diagrams, the chondrocranium is in blue and the splanchnocranium is in yellow.

Chondrocranium of Embryos
The chondrocranium is also relatively complete in embryos, where it appears early in development. It includes: -capsules around the three special sense organs (including the eye) -trabecular and parachordal cartilages that form the basal plate -the occipital arch (neural arches of cranial vertebrae)

KK 7.3, H&G 8.3

The Splanchocranium
The splanchnocranium supports the jaws and gill openings. It is composed of cartilage that may ossify in the adult. KK 7.5, H&G 8.5

Development of the Splanchocranium

Splanchnocranium is derived from embryonic neural crest. It is best seen in sharks, or in embryos of other vertebrates before dermal bones replace or cover it. Note that some dermal bones (e.g., maxilla, squamosal) have contributions from neural crest.

KK 7.4

Origins of Jaws
The jaws of vertebrates were originally cartilaginous supports for the anterior gill openings. Modern vertebrates have 7 visceral arches, the first two being the mandibular and hyoid arches. Some believe there were arches ahead of the mandibular arch in early vertebrates (i.e., the pink arch in this diagram).

Note the spiracle.

KK 7.7, H&G 8.5

Fate of the anterior visceral skeleton 1

KK 7.8, H&G 8.7 The mandibular arch is mostly replaced by dermal bones in bony fishes and tetrapods. The hyomandibula links the jaws of fish to the rest of the skull of most fish. The homologues of the palatoquadrate and mandibular (Meckels) cartilage are the quadrate and articular bones at the jaw hinge of bony fish and early tetrapods. The tetrapod homologue of the hyomandibula is the stapes.

articular

Sharks can extend upper and lower jaws to bite using the hyomandibula as a lever.
KK 7.19

Fate of the anterior visceral skeleton 2:

A new role for the hyomadibula and the inner ear in tetrapods
KK 17.39

In the first tetrapods, the inner ear was likely used for a new function; to detect vibrations from the ground through the jaw. (The inner ear was not originally used for hearing, but several fish do so, as do tetrapods.) The hyomandibula (now called the stapes) may have originally provided a connection from jaw to ear.

In modern Amphibia and reptiles, the tympanum (eardrum) covers the spiracle opening, and the stapes lies in the spiracle (now called the eustachian tube) to connect the tympanum to the inner ear, conducting vibrations from the air.

Fate of the anterior visceral Skeleton 3

Further changes in mammals

The essential role of the visceral skeleton did not change through the rest of tetrapod evolution until mammals. The quadrate and articular remained jaw hinges. In mammals, the quadrate and articular join the stapes as ossicles of the middle ear.

A dermal bone of the lower jaw, the angular, also leaves the jaw to contribute to the ear as the tympanic bulla

KK7.55, H&G 8.14

The middle ear of mammals

Paleontologists distinguish reptiles from mammals based on whether they have an articular or a malleus.

angular Homologues hyomandibula = stapes Meckels = articular = malleus palatoquadrate = quadrate = incus angular = tympanic KK 17.36c, H&G 19.11

We can verify these homologies by following their ontogeny, as they migrate and change shape during development.

You can also see in this embryo (Armadillo, an edentate mammal) where the rest of the visceral skeleton ends up in tetrapods; it contributes to the hyoid apparatus, larynx and trachea.

The rest of the visceral skeleton in tetrapods

Parts of visceral arches 2 through 7 contribute to: -the hyoid apparatus (support for the attachment of the tongue) -the larynx -the rings of cartilage supporting the trachea or windpipe. -the alisphenoid bone

KK 7.54, H&G 8.13

Summary of the three components of the tetrapod head skeleton, by development

Chondrocranium = blue, splanchnocranium = yellow, dermatocranium = pink

KK 7.11, H&G 8.4

Summary of the three components of the adult mammal head skeleton

KK 7.54, H&G 8.4, 8.6

Chondrocranium = blue, splanchnocranium = yellow, dermatocranium = pink