CHAPTER 8: VERTEBRAE, RIBS, STERNA

Axial Skeleton
Skull
Form in lateral, ventral body
Ribs
Sterna
walls
Vertebral column forms around, sometimes invade
notochord during ontogenesis
VERTEBRAL COLUMN
Keystone of vertebrate skeleton
Segmented, arched rod flanked by axial musculature (where
head is attached):

In fishes - from w/c the rest of the body is suspended

In most tetrapods from w/c trunk is suspended

between anterior and posterior limbs
Provides protective bony tunnel for spinal cord
Essential architectural component of axial locomotor apparatus
in vertebrates (except agnathans)
In fishes:

Locomotion resistance by pushing laterally against

water with movements of trunk and tail; segmental
muscles attached to vertebrae, ribs and myosepta

Articulation in vertebrae side-to-side movement only

In land:

Articulation changed dorsoventral flexibility; also

providing strong skeletal bowlike arch for suspending the
trunk

Functional separation of muscles used in locomotion from

those used in ventilation
Centra, Arches and Processes
Modern vertebrae typically consist of centrum, neural arch,
and apophyses

Centra occupy position occupied by notochord

Neural arches - perched on the centra;

interconnecting ligaments and successive arches
encloses the vertebral canal (neural canal)

Haemal arches (chevron bones) inverted beneath

centra of tail; house caudal artery and vein

Transverse process (diapophyses) most

common, articulate w/ ribs projecting laterad into
skeletogenous septum separating epaxial and
hypaxial muscles of body wall; serve as attachment
for some muscles that extend/flex vertebral column

Zygapophyses paired processes at the cephalic

end of trunk vertebrae (prezygapophyses) and at
caudal end (postzygapophyses); chiefly in tetrapods
interlocking arrangement limits dorsoventral
flexion of column in region of trunk; absence of
zygapophyses in tetrapod tail makes it flexible

Parapophyses lateral projection from the centra;

serve as articulation site for capitulum

Hypapophyses prominent midventral projections

from centra; site of attachment for certain
muscles/tendons
Morphogenesis of Vertebrae
Typical vertebra:

Arises from mesenchyme cells that stream out of

sclerotomes of mesodermal somites, surround
notochord and neural tube and produce blastema for
future vertebra
Chondroblasts that differentiate w/in blastema deposit
cartilaginous centrum, neural arch and hemal arch resulting
to cartilaginous vertebra w/ notochord in centrum
Except in cartilaginous fishes cartilage is removed, bone is
deposited
Notochord remain depending on species:

Chondrichthyes cartilaginous vertebra never

replaced by bone

Teleosts, urodeles and apodans membrane bone is

deposited in perichordal blastema; but arches arise
from cartilage

Many fishes, amphibians except anurans not only

cartilage and membrane bone deposited around
notochord (perichordal cartilage/bone),
chondroblasts invade notochord sheath and deposit
cartilage in sheath and sometimes in notochord itself;
centra contain chordal cartilage
In forming centra sclerobasts from caudal half of one somite
and cephalic half of next somite stream to intersegmental
location around notochord to establish perichordal blastema;
centra typically intersegmental, myomeres are segmental
Diplospondyly two centra per segment; provide tail w/
increased flexibility for locomotion
Fishes intervertebral ligaments; Tetrapods intervertebral
joints

Vertebral Column of Fishes

Vertebral column exhibits only two major morphologic regions
of specialization: trunk (dorsals) and tail (caudal)
Living agnathans strange vertebral column; lateral neural
cartilages (only skeletal elements) fuse to form a single
longitudinal dorsolateral cartilaginous plate perforated by
foramina for spinal nerves
Hagfishes lns limited to tail
LNS may be vestigial or primitive, or may bear no phylogenetic
relationship to vertebrae
Sharks

Notochord is present throughout adult vertebral

column, constricted within each centrum; centra
(chordial and perichordial cartilage) concave at each
end amphicelous

Vertebral canal consists of paired dorsal plates

constituting neural arch, paired dorsal intercalary
plates and in some species, wedge-shaped
supradorsal cartilages

Squalus acanthias lack supradorsals; neural arch

and intercalary plates perforated by foramina for
dorsal and ventral roots of spiral nerves for blood
vessels

Scyllium foramina emerge between plates; hemal

arches consist of paired ventral plates, ventral
intercalary plates common in between hemal arches
Fibroelastic ligament overlies and connect neural spines along
entire length of vertebral column
Few fishes w/ exceptional columns:

Dipnoans, chondrostei and coelacanth Latimeria

develop no centra; notochord present and
unconstructed, its fibrous sheath contains little
cartilage/bone

Associated notochord in each body segment are

paired basidorsal, basiventral, interdorsal and
interventral cartilages that are products of separate
chondrification centers in embryonic perichordal
blastema

Extinct holocephalans vertebral column sonsisted

of thick rings of calcified castilage that had been
deposited in notochord sheath; rings more numerous
than body segments
Teleosts:

Well-ossified amphicelous vertebra; core of each

centrum is a dumbbell-shaped vacuole where
notochord previously existed. Space occupied by
porous cartilagelike material that probably includes
notochordal remnants

Centra and neural arches of successive vertebrae

are interconnected by complex system of
collagenous and elastic ligaments that facilitate
lateral undulation of body for locomotion

In posterior trunk, ligaments are frequently ossified to

form rods

Neural spines are often very tall, sometimes

surrounded by supraneural bones

Variety of processes (mosly unique to fishes)

protrude from arches and centra
Elasmobranchs and bony fishes two centra and two sets of
neural and hemal arches in each metamere of tail; number of

centra is double the number of myomeres and spinal nerves

(diplospondylous condition)
Amia two centra per body segment in postanal portion of
vertebral column, but only one bear arches; other is termed
intercentrum (not homologous w/ hypocentra/intercentra)
Embryonic basidorsal and basiventral cartilages are
incorporated in centra, interdorsal and interventrals are
incorporated in the intercentra
No movement porssible in first vertebra and skull in fishes;
united by cartilage/nonelastic connective tissue

Evolution of Tetrapod Vertebrae

Rhipidistian fishes and early labyrinthodons vertebrae consist
of several bones per segment

Hypocentrum (intercentrum) large, median, Ushaped anterior bone that cradled notochord

Pleurocentra small wedges of bone overlying the

notochord dorsolaterally

Independent left and right laminae of bone lateral to

spinal cord that provided a neural arch; each lamina
rested in notch between pleuro and hypo

Notochord continuous throughout but constricted at

level of hypocentrum

Rachitomous (consists of several pieces)

Later tetrapod vertebrae except urodeles and apodans
modification of rachitomous condition; increased prominence of
pleurocentra and concomitant reduction of hypocentra
Modern adult tetrapod vertebra composite structure
Urodeles and apodans lack evidence of
pleurocentra/hypocentra
Vertbrae of urodeles, apodans, Sphenodon and basal lizards
amphicelous
In most other tetrapods concavity has disappeared anteriorly,
posteriorly or both ends of centra opisthocelous, procelous
or acelous, respectively.
Anurans and modern nonavian reptiles procelous

Condition evolved from a buildup of a hypocentrum of

chordal cartilage between two centra and eventual
coalescence of intercentrum w/ centrum ahead of it.

Coalescence of hypocentrum w/ centrum immediately

behind it resulted to ophithocelous
(salamanders)

Mammals acelous (fibrocartilaginous

intervertebral disc bet. Successive vertebra and
pulpy nucleus in its core
Modified intervertebral joints permit more flecibility to column
Regional Specialization in Tetrapod Columns
Adaptation to life in land:

Push of posterior appendage transmitted via the

pelvic girdle to vertebrae modified into sacral
vertebrae

Shortening of ribs on some of anterior vertebrae,

increasing mobility of intervertebral joints which
became the neck; cervical vertebrae

Long ribs restricted to vertebrae in anterior portion of

trunk, preserving a bony cage: thoracic vertebrae

Remaining vertebrae anterior to sacral: lumbar

vertebrae
Snakes have longest columns, 400 or more w/ no
specialization
Apodans limbless, as many as 250 or more vertebrae
Urodeles 100
Anurans shortest columns and limited flexibility
Turtle and birds only cervical and caudal segments of column
are flexible; trunk vertebrae fused to synsacrum in birds and
carapace in turtles
The Craniovertebral Junction and Neck Vertebrae
Amphibians cervical vertebrae lacks processes; smooth
concave facets that articulate w/ 2 occipital condyle

Absence of processes and nature of amphibian

craniovertebral joint permits limited dorsoventral
rocking of skull

Change from one occipital condyle of early

amphibians

Amniotes larger number of cervical vertebrae than

amphibians; long flexible neck

First two vertebrae:

o
Atlas (first vertebrata) - ringlike because
centrum is missing; 2 superior articular
facets on its cephalic ends for articulation
of occipital condyles
o
Axis (second vertebrata) odontoid
process as centrum of atlas; process held
by tranverse atlantal ligament
o
Proatlas resembles a neural arch and
derived from bilateral cartilaginous
blastemas; interposed between skull and
atlas in crocodilians, Sphenodon and
hedgehogs first appeared in basal
amniotes

Skull and atlas rocks as a unit in odontoid process

(axis of rotation); rocking facilitated by absence of
zygapophyses in atlas

Sphenodon odontoid process chondrifies as

independent element, unites w/ axis only after
hatching
Flexible neck asset in gathering food and avoiding enemies
in land
Flexibility of neck exceptional in birds because of nature of
articulations of vertebrae

Caudal ends of centra saddle-shaped:

heterocelous enable both lateral and dorsoventral
flexion of neck

Heterocelous centra, atlas-axis complex and largest

number of cervical vertebrae in any tetrapos
enable turning of heads 180 degrees to the rear:
o
12 common
o
25 swans
Turtle uniquely flexible neck

Ball-and-socket joints bet. successive procelous

centra enable head and neck to be completely
retracted into shell by flexing neck into several folds
dorsoventrally

8 cervical vertebrae
Remainder column of birds and turtles (except caudal
vertebrae) rigid
Mammals always 7 cervical vertebrae

Exception: 3 edentates (2 sloths and great anteater)

w/ 6, 8 or 9; manatees w/ 6

Length of centra, not number, that determines neck

length in mammals

Stabilizing the Hind Limbs:

Sacrum and Synsacrum
Sacral vertebrae has stout transverse processes strong
enough to bear thrust of pelvic girdle as hind limbs push
against earth in locomotion
Stout ribs ankylosed to distal ends of transverse processes

Amphibians 1 sacral vertebra

Reptiles including birds and opposums 2

Mammals 3-5 except perisodactyles 8, edentates

13; ankylose to form sacrum
Do not differentiate in tetrapods that lack hind limbs
Modern birds last thoracic vertebra, all lumbar, sacrals and
first few caudals and their ribs ankylose to form synsacrum

Synsacrum fuses with pelvic girdle compact pelvis

is formed, providing a rigid brace for tetter-totterlike
stance of birds

Thoracic vertebra anterior to synsacrum unite

completely little flexibility in avian backbone caudal
to neck; minimizes number of muscles needed to
keep body streamlined during flight, reducing cost in
energy expenditure

Archeopteryx rudimentary synsacrum; trunk

vertebrae not fused, pelvic girdle proportionally
smaller than todays birds

Weight of body when standing counterbalanced by

long tail
Mammals synsacrum: Armadillos

Consists of up to 13 fused sacral and caudal

vertebrae

Tail Vertebrae: Urostyle, Pygostyle and Coccyx

Caudal vertebrae 50 or more in early tetrapods
Reduced number in modern ones; toward end of tail arches
and processes become shorter and rudimentary until consist
solely of small cylindrical centra
Anurans unique urostyle; develops from continuous
elongated perichordal cartilage at base of larval tail, grows and
ossifies after tail is lost after metamorphosis

In some species, postsacral vertebra composition

evident from centra, vestigial arches, transverse
process and nerve foramina (part of cranial end of
urostyle)
Reptiles: Lizards

When captured by tail, break off at end distal to site

of capture

Tail regenerates

Autotomy implemented by zone of soft tissue

dividing each tail vertebra into cephalic and caudal
sections, location at level of myosepta

Sudden opposing reflex muscle contractions on sides

of septum cause of break
Archaeopteryx long tail
Modern birds still have tail but not prominent

Pigeons 15 caudal vertebrae: 5 ankylosed w/

synsacrum, 6 independent, last 4 fused to form
pygostyle (skeleton of visible part of tail)

Pygostyle develops as 4 separate cartilaginous

centra
Mammals few as 3 caudal vertebra and as many as 50

Tail of sperm whales 24

Apes, humans 4/5 vestigial caudal vertebrae

comparable to pygostyle of birds; lack arches, most
have rudimentary transverse processes, last 3/4
diminish in size and fuse to form coccyx (in human
at 25 yrs. of age)

Centra of coccyx still identifiable but last one is mere

nodule of bone

People tailbone reccurently sore fractured coccyx

in a fall that caused them to land on the end of spine

Adult rhesus monkeys have prehensile tail the length

averages nearly 50% of monkeys sitting height
RIBS
-

Fishes
-

Articulate w/ vertebrae and extend into body wall

Formed intersegmentally by scleroblasts from 2 successive
mesodermal somites in same manner as centra
Sclerotomal origin for entire rib, necessary interaction bet.
sclerotome and myotome for successful dvlt of rib; most bony
ribs are of endochondral origin
Gastralia (abdominal ribs) in some reptiles not ribs, but
riblike membrane bones that maybe remnants of ancestral
body dermal exoskeleton
Basal actinopterygian Polypterus and some teleosts 2 sets of
ribs, dorsal and ventral, associated w/ each trunk vertebra

Dorsal ribs pass laterad into horizontal

skeletogenous septum separating into exapial and
hypaxial muscles

Ventral ribs develop in myosepta and arch ventrad

in lateral body wall external to parietal peritoneum as
in mammalian thorax; stop short of midventral line in
fishes

Commencing at vent, ventral ribs approach one

another beneath centrum and unite to form hemal
arches of tail
Most fishes only have ventral ribs (except perch)
Sharks and others only have dorsal ribs
Skates, chimaeras and few unusual teleosts (sea horses) no
ribs
Living agnathans no ribs; correlated with absence of centra
Fish ribs strengthen myosepta to w/c longitudinally disposed
locomotion muscles of myomeres are attached

Tetrapods
Early tetrapods ribs associated w/ vertebrae from atlas to
nearly base of tail; later became restricted in length and
number (short ones ankylosing with transverse processes)
Long ribs of anterior thoracic region sternum
Tetrapod rib occupies same location of ventral ribs of fishes
encircle coelomic cavities
Tetrapod rib and ventral fish rib not homologous (tetrapod rib
homologous to dorsal fish rib)
Most are bicipital

Tuberculum dorsal head; in early tetrapods,

articulated w/ extremity of transverse process

Capitulum ventral head; early tetrapods,

articulated w/ hypocentrum
Reduction of size of hypocentrum in amniotes through time,
articulation site of capitulum was altered (site changes based
on length of trunk). Articulated in several fashions:

2 adjacent demifacets; one on caudal end of

centrum, other on cephalic end of next more posterior
centrum

W/ parapophysis, when latter was present

In facet on a single centrum

Some mammals, tuberculum reduced to a vestige in one
region of the column or another
Some tetrapods, trend of fushion of 2 heads until only
tuberculum remained (crocodilians)
Typical thoracic ribs of amniotes consist of 2 parts:

Costal rib adjacent to vertebrae

Sternal rib distal, more ventral

Some ventral ribs generally articulate w/ sternum
Sternal ribs may remain cartilaginous, as in humans, w/c they
are also referred to as costal cartilages
Vertebral column, costal rib, sternal rib and sternum
provide skeletal enclosure for thoracic viscera of amniotes
No skeletal cage in amphibians
Amphibians
Anurans and urodeles all ribs have become short and
ankylosed to transverse processes (anurans)
Apodans long ribs

Present in association w/ all except the first vertebra

and a few more posterior ones; play vital role in
locomotion
Only anuran ribs not bicipital
Non-avian reptiles
Lizards and crocodilians long ribs on many of trunk
vertebrae, short ribs in most of neck

Sphenodon long ribs in most of trunk and ribs

continue into tail

Geckos ribs associated w/ every cervical vertebra

More specialized lizards lack ribs on atlas and axis

Draco half dozen of posterior ribs of trunk are

greatly elongated; can be rotated to elevate a broad
fold of body wall skin (patagium) w/c then becomes
winglike membrane on w/c lizards soars, folded when
not in use
Turtle no cervical ribs, ribs of trunk are fused w/ costal plates
of carapace

2 sacral ribs are not fused w/ carapace; short and

expanded distal ends ankylosed to ilia of pelvic girdle
(brace girdle against vertebral column)

Slender, riblike processes may be associated w/

some of the caudal vertebrae
Snakes - long, curved ribs beginning at second vertebra and
continuing far into tail

No sternum for ribs to attach to; ventral ends of ribs

have ligamentous connections w/ integumentary
scutes

Ribs participate in locomotion

Cobras long, curved ribs in neck can be rotated

outward like Dracos causing a fold of loose skin in
the neck to spread fold inflated w/ air from the
lungs

Birds
-

First 2 pairs of ribs in many carinates articulate w/ last 2

cervical vertebrae at base of neck; movable and lack sternal
segment
Next 5 pairs thoracic ribs w/ bony sternal segments;
constitute major skeleton of thoracic basket (chest cavity)
thin, flat and broad and most bear broad uncinate processes
Thin broad ribs and uncinate processes provide birds w/
lightweight but sturdy thoracic body wall skeleton for
attachment of powerful muscles required for flight
Ribs caudal to thoracic basket ankylosed to underside of
synsacrum (as are all trunk ribs in turtles)
Developing embryos exhibit transitory rudimentary ribs deep
into tail
Uncinate processes found also in some lizards and in some
early labyrinthodonts

Mammals
Recognizable ribs in mammals confined to thorax
Number ranges from 9 pairs in some whales to 24 in some
sloths; 12 pairs are common
Larger then 10, rest are floating; costal cartilages fail to
reach the sternum
Humans and apes other than orangutans 12 pairs of thoracic
ribs; apes addl 2 pairs of lumbars, humans frequently have
an addl rib , cervical or lumbar, as an anomaly
Mammalian embryos 2 heads of a vestigial bicipital rib
develop in association w/ each cervical vertebra. One of the
heads attached to a transverse process, the other to a centrum

Two heads create in bet. transverse foramen on

each cervical vertebra

Consecutive foramena provide a bony tunnel,

arteriovertebral canal, w/c transmits a vertebral
artery from its source at the base of neck to brain

Transverse foramina develop in birds, but number of

vertebrae involved depends on the species

Vertebral artery exists in other reptiles but not

enclosed in canal
TETRAPOD STERNUM
Sternum tetrapod structure and predominantly amniote
Endochondral origin
Presence, size and anatomical relationships are correlated w.
extent to w/c forelimbs are employed in locomotion
Serves chiefly as base against w/c pectoral girdles and in ribs
are braced (in amniotes) and as the anatomical origin of
ventral muscles of forelimbs
Largest in pterosaurs, carinate birds and bats (w/c require
strong muscles for flight)
Reduced or absent in tetrapods lacking forelimbs
Acquired additional roles in some tetrapods (assisting in
breathing movements in birds)
Sternum of amphibians only well differentiated in anurans

Necturus consist of more/less isolated nodules of

cartilage associated w/ the linea alba immediately
caudal to coracoid cartilages of pectoral girdle and
extending into proximal ends of adjacent myosepta

Small but discrete sternal element present in other

salamanders in same location as Necturus

Apodans no trace of sternum

Reptiles

Lizards agile climbers and runners; strong pectoral

girdles firmly braced against substantial shieldshaped sternum of cartilage/replacement bone

Crocodiles sternum is a simple cartilaginous plate

attached to the precoracoid of pectoral girdle;
caudally continuous w/ expanse of mineralized
fibrous membrane incorporating gastralia

Turtles no sternum; 2 halves of pectoral girdle are

united ventrally by fibrous ligaments/ by mere wedge
of cartilage
Birds

Sterna of modern birds developed keel/carina to

w/c massive pectoral flight muscles are attached

Dvlt of carina in pterosaurs and birds convergent

evolution

Archaeopteryx small and lacked a keel, indication

that first birds are not strong fliers
Mammals

Terrestrial mammals composed of series of bony

segments (sternebrae); comparatively narrow to that
of reptiles

Xiphisternum last sternebra, bears a cartilaginous/

bony xiphoid process

Marine mammals relatively short sternum, only

small number of ribs reach sternum and individual
sternebrae tend to be fused; in cetacean, may be
correlated w/ galloping mode of swimming
necessitated by lack of functionally competent
flippers
Nature of earliest sterna little evidence of a sternum in
labyrinthodont amphibians

Either it was not preserved or had not yet come to

being

Tendency of collagenous tissues (w/c includes

myosepta and linea alba) to undergo ossification
when subjected to continued mechanical stress

May be ossification in cephalic portion of linea alba

that ultimately produced sterna response to
sustained mechanical stresses resulting from
locomotion on land

Sternum may have arisen several times in

evolution of tetrapods perhaps independently in
urodeles, in anurans, and in an amphibian line
leading to amniotes
Amniote sternum arises as paired mesenchymal bars that later
unite and undergo chondrogenesis. In many, mamms:

Presternal and suprasternal blastemas also

develop
o
Presternal contributes to manubrium
o
Suprasternal sometimes contribute to
manubrium; may give rise to independent
suprasternal ossicles