THE POSTCRANIAL SKELETAL
The primary function of the cranial skeleton was to protect and support
the brain and associated sensory organs. In contrast, the postcranial skeleton
is used less for protection but more for support of the body and for locomotion
The postcranial skeleton can be divided into two regions:
Trunk - includes the vertebral column, ribs and sternum (part
of axial skeleton)
Appendicular skeleton - limbs and girdles
Notochord vs. the Vertebral Column
In early vertebrates the notochord is a non-bony skeletal support for
swimming by lateral undulation
- consists of vacuolated cells surrounded by a thick fibrous
sheath that maintains rigidity, but is also flexible
- ventral and parallel to the spinal cord (= dorsal hollow nerve cord)
in vertebrates
- absent in hemichordates, present in the tail of most larval urochordates
but lost in the adults
- present in the body and tail of cephalochordates and vertebrates
- present in the embryo of all vertebrates but, as the vertebrae form
around it, it becomes constricted and in many groups, such as mammals,
there is almost nothing that remains.
The vertebral column is a basic support structure that is developed from
individual metameric units called vertebrae which replaces the notochord
The basic units of an individual vertebra are:
Vertebral/neural arch - surrounds and protects the spinal cord
Hemal arch - surrounds and protects the caudal artery and vein in
fishes
Neural spine - spinous process that projects dorsally from the neural
arch
Centrum - body of the vertebra which replaces the notochord; shape
depends on the vertebrate class (Fig. 8.4, p. 277):
-
Amphicoelous - centrum hollowed at anterior and posterior end, characteristic
of teleosts and early reptiles
-
Opisthocoelous - centrum convex on anterior surface, concave on
posterior surface, characteristic of some of the vertebrae of reptiles
and mammals
-
Procoelous - centrum concave on anterior surface, convex on posterior
surface, characteristic of some reptiles and amphibians
-
Acoelous - centrum flat on posterior and anterior surface, characteristic
of some reptiles and mammals
Transverse process - spinous process that projects laterally from
the centrum
Zygaphophysis - articular processes that extend forward and backward
from neural arches and help to strengthen the union between vertebrae
Intervertebral disks - pads derived from part of the notochord and
composed of fibrocartilage and connective tissue that lie between adjacent
centra that help to cushion the connection
In tetrapods, two general anatomical relationships occur between centra
and neural arches:
aspidospondyly - all arch elements (intercentrum, pleurocentrum,
and neural arch) remain as separate ossified elements
holospondyly - all vertebral elements are fused into a single piece
For most vertebrates (except for most fishes), regional variation in the
appearance of the vertebrae are recognized (Fig. 8.2, p. 277):
Cervical - vertebrae of neck (not found in fishes)
Atlas - the first cervical vertebra of terrestrial vertebrates which
articulates with the skull; nodding movements of the head occur between
the atlas and the skull
Axis - the second cervical vertebra of mammals; rotary movements
of the head occur between the atlas and axis
Thoracic - vertebrae of chest region, articulate with ribs
Lumbar - vertebrae of lower back
Sacral - lower vertebrae that are fused (three fused in dog and
cat, five fused in human) as the sacrum
Caudal - vertebrae of tail (pygostyle in birds forms attachment
for tail feathers)
Ribs and Sternum
The ribs and sternum serve several general purposes:
- help to strengthen the body wall and lend support to the thorax
- protect the organs of the thoracic region (circulatory and respiratory)
- used as the site of muscle attachment
- in amniotes they are used to assist in breathing
There are two general ways in which ribs can develop in different vertebrate
classes:
Intermuscular (dorsal) ribs - develop in the myosepta between
the myomeres and attach to centra of vertebrae, between the dorsal and
ventral muscle masses
Subperitoneal (ventral) ribs - form between the ventral muscles
and the lining of the coelom
With the exception of the agnathans that have no ribs at all, fishes generally
show both types of rib morphology
- may be found throughout the trunk, as is the case in salamanders
and reptiles, or only in the thoracic region, as is the case for mammals
and birds
Mammalian ribs articulate with the vertebrae at two points:
- the dorsal point of articulation tuberculum
- the ventral point of articulation capitulum
- also articulate differently with the sternum, and can be either true
ribs (directly attached to the sternum via costal cartilages - 7 pairs
in human), false ribs (attached indirectly to the sternum through
the costal cartilages of the other ribs - 5 pairs in human) and floating
ribs (do not attach to the sternum - 2 pairs in humans).
The sternum forms a complete enclosure of the chest region in conjunction
with the ribs and is connected closely to the shoulder girdle and ribs.
- possessed only by tetrapods, with the exceptions of snakes and turtles
(completely lacking)
- acts to protect the thoracic region
- serves as a site for attachment of the pectoral limbs
- aids in rib movements
In birds, the sternum is completely ossified:
• ratite - lacking a distinctive keel or sternum for attachment
of breast muscles
• carinate - having a massively enlarged sternum to support
flight muscles
In mammals, the sternum is divided into three regions (anterior to posterior):
• manubrium
• sternebrae (ossified bony elements)
• xiphisternum and xiphoid cartilage
Several vertebrates, including lizards and crocodiles also posess gastralia:
- ribs of dermal origin restricted to the sides of the ventral body
wall
- do not articulate with the vertebrae but act as an accessory skeletal
system for muscle attachment and support for the abdomen
Definitions
Acoelous - centrum flat on posterior and anterior surface, characteristic
of some reptiles and mammals
Amphicoelous - centrum hollowed at anterior and posterior end, characteristic
of teleosts and early reptiles
Amphistylic - jaw is supported both by the hyomandibular and by a direct
connection between the jaw and the chondrocranium; found in primitive cartilaginous
fishes
Atlas - the first cervical vertebra of terrestrial vertebrates, which
articulates with the skull; nodding movements of the head occur between
the atlas and the skull
Autostylic - upper jaw (pterygoquadrate cartilage) articulates or is
fused with the chondrocranium, lower jaw forms from the mandibular cartilage,
and the jaw remains unsupported by the hyomandibular found in lungfishes,
and in the tetrapod ancestors
Axis - the second cervical vertebra of mammals; rotary movements of
the head occur between the atlas and axis
Calcified cartilage - cartilage containing deposited calcium salts.
Found in the vertebrae of cartilaginous fish
Carinate - having a massively enlarged sternum to support flight muscles
Cartilage - a firm but elastic skeletal tissue whose matrix contains
proteoglycan molecules that bind with water.
Choanae - internal nostrils that pass from the external nares to the
lungs
Chondrocranium - anterior part of the axial skeleton that encases the
special sense organs and contributes to the skeletal elements encasing
the brain
Chondrocyte - a mature cartilage cell
Composite materials - heterogeneous materials composed of many different
elements, that is generally much stronger than homogenous materials.
Cricoid cartilage - cartilage of the larynx that develops from the
hyobranchial apparatus
Dermatocranium - consists of dermal bones that encase the chondrocranium
and splanchnocranium and contribute to the braincase, jaws, and skeletal
elements of the mouth (teeth)
Elastic cartilage - cartilage containing elastin fibers that appears
yellowish. Found primarily on external ear and epiglottis.
Fibrocartilage - cartilage containing collagen fibers. Found in the
intervertebral disks and pubic symphysis
Fontanelle - temporary gaps between bones, such as those that occur
during the formation and fusion of the cranial bones
Heterodont - teeth modified for different functions
Hyaline cartilage - cartilage with a clear translucent matrix. Found
primarily on the ends of ribs and on the trachea
Hyomandibular cartilage - dorsal part of the hyoid arch that articulates
with the chondrocranium
Hyostylic - upper jaw loses any major direct connection with the chondrocranium
and the upper and lower jaws are supported solely by the hyomandibular;
found in elasmobranchs and most bony fishes
Lacunae - small cavities, such as those in bone or cartilage, that
contain osteocytes or chondrocytes
Membrane bone - superficial bones that lie in or just beneath the skin
and develop from the direct deposition of bone in connective tissue
Opisthocoelous - centrum convex on anterior surface, concave on posterior
surface, characteristic of some of the vertebrae of reptiles and mammals
Perichondrium - the connective tissue that lies on the outside of cartilaginous
tissue
Procoelous - centrum concave on anterior surface, convex on posterior
surface, characteristic of some reptiles and amphibians
Pygostyle - the fused caudal vertebrae of a bird that support the tail
feathers
Ratite - lacking a distinctive keel or sternum for attachment of breast
muscles
Sacrum - the union of two or more vertebrae and their ribs, by which
the pelvis articulates with the vertebral column
Sclerotic bones - bones that surround the orbit but do not articulate
with it, and provide reinforcement for the eye
Splanchnocranium - or visceral arches that support and move the gills
and contribute to production of the jaws in gnathostomes
Thecodont - also called homodont, or undifferentiated tooth structure
Turbinate bones - bones of the nasal cavity that increase the surface
area available for olfaction
Wormian bones - intermediary bones, or small islands of bone that occur
between sutures in the skull
Zygaphophysis - articular processes that extend forward and backward
of neural arches and help to strengthen union between vertebrae