Chapter 1: Introduction

Comparative vertebrate anatomy is the

study of the structure of vertebrates (descriptive morphology)
study of the functional significance of structure (functional morphology)
Because structure entails development of the individual (ontogenesis)
And individuals have ancestral history (phylogenesis)

Every area of biological inquiry provides relevant data to the discipline. Ecology, embryology, genetics, molecular biology, serology, biochemistry and
paleobiology are all sources of valuable date. Geology is an indispensable source.

To the extent that comparative anatomy is concerned with phylogenesis, it is a study of history and of animals that are extinct. It is interested in the
survival value of a structure (an adaptation), of the struggle for compatibility with an ever-changing environment, of the invasion of new territory by those
most effectively equipped for survival and of the extinction of species.

The history of vertebrates is a story from which is developing a genealogy based on the date just described. Comparative anatomy addresses curiosity
about the origin of species. Generalizations and conclusions arrived at add to the enlightenment of the mind.

Phylum Chordata: The Big Four

A dichotomy that categorizes animals based on the presence of vertebral columns doesnt recognize a group of small
marine animals that are transitional between invertebrates and vertebrates protochordates.
Protochordates have no vertebral column but they share with vertebrates and with no other animals a combination of 4
other morphological features
o Notochord
o Dorsal hollow central nervous system
o Post anal tail less critical to development
o Endostyle glandular groove in the floor of the pharynx that secretes mucus
These characteristics are fundamental in vertebrates that they are among the first to appear in vertebrate embryos.
Without most of them, no vertebrate could proceed beyond the earliest stages of embryonic development.
Protochordates and vertebrates have been merged into a single taxon because of the primacy of structures
o Kingdom Animalia
Phylum Chordate
Subphylum Urochordata
Subphylum Cephalochordata
Subphylum Craniata
o Hagfish (craniates without vertebrae)
o Vertebrate (craniates with vertebrae)
Chordates are animals that have a notochord in the embryo stage at least
Craniates are chordates with a neurocranium (brain case)
Vertebrates are chordates with vertebrae. (vertebrae appear during embryonic development after the notochord has
formed)

The Craniate Body: General Plan

All craniates conform to a generalized pattern of anatomic structure representing a collection of primitive (pleismorphic)
and unique (derived) anatomical features.
This is the result of expression of similar DNA inherited during the course of evolution.
Craniates also exhibit similar but not identical patterns of embryonic development
Morphology and developmental processes have been altered during the passage of time which as it lengthens provides
increasing opportunities for genetic changes that result in anatomic diversity.
Despite these changes, innumerable primitive structural and developmental similarities still exist

Regional Differentiation

Head
o Special sense organs for monitoring the environment
Brain large enough to receive and process information; provide apt stimuli to muscles
Cephalization developed to a greater degree in craniates
Jaws for some species for acquiring, retaining and macerating food
Gills for respiration in fishes
Trunk
o Coelom cavity that houses most of the viscera
o Body wall surrounds the coelom; consists of muscle, vertebral column and ribs
 o Paired pectoral and pelvic appendages (fins or limbs) in many but not all craniates
o Neck narrow extension of the trunk of amphibians, reptiles, mammals and lacks a coelom
Consists of vertebrae, muscles, spinal cord, nerves and elongated tubes (esophagus, blood vessels,
lymphatics, trachea) that connect structures of head and trunk
Post anal Tail
o Commences at the anus or vent hence post anal
o Consists almost exclusively of caudal continuation of body wall muscles, axial skeleton, nerves and blood
vessels
o Some adult craniates lack a post anal tail although its present in all embryos
Amphibians have tails but it is resorbed at metamorphosis
Birds have tails reduced to a nubbin
Humans have a vestigial post anal tail. Remnants remain as tailbone (coccyx)
o 2 pairs of appendages pectoral and pelvic are characteristic of most vertebrates but they are sometimes
vestigial or completely lost
The earliest known craniates lacked one or both pairs of these appendages
Lacking both pairs of appendages is seen in living jawless craniates (agnathans)
We do not see pelvic appendages until the origin of gnathostomes (craniates with jaws)

Bilateral Symmetry and Anatomic Planes

Craniates have 3 principal body axes

o Longitudinal (antero-posterior) axis
o Dorso-ventral axis
o Left right axis (shared among members of Bilateria)
With reference to the first 2, structures at one end of the axis are different from those at the other end
The left right axis terminates in identical structures at each end. Thus the head differs from the tail and the dorsum
differs from the venter but the right and left sides are mirror images of each other. An animal with this arrangement of
body parts exhibits bilateral symmetry.
It is sometimes convenient to discuss parts of the craniates body with reference to the 3 principal anatomic planes
o A cut through the left-right and the dorso-ventral axes (transverse plane) is a cross section
o A cut through the left-right and longitudinal axes (frontal plane) is a frontal section
o A cut through the longitudinal and dorso-ventral axes (sagittal plane) is a sagittal section
Sections parallel to the sagittal plane are parasagittal

Metamerism

Craniates show the primitive feature of metamerism serial repetition of structures in the longitudinal axis of the body
Its clearly manifested in craniates embryos and retained in many adult systems
No external evidence is seen in most adult reptiles and mammals because the skin is not metameric
However, if the skin is stripped from the trunk and tail of fishes, most amphibians and some reptiles, one sees a series
of muscle segments that are reflections of the embryonic metamerism

Craniate Characteristics

In addition to retaining primitive characters (bilateral symmetry, deuterostome development and chordate big four), craniates
exhibit a unique combination of morphological features
1. Cranium
2. 3 part brain
3. Neural crest and its derivatives
4. Paired external sense organs (olfactory, optic, otic with a single semicircular canal and lateral line system with
unicellular sense organs
5. Cartilage

Vertebrate Characteristics

1. Vertebral column (primitively seen as isolated elements associated with an unrestricted notochord)
2. 2 semicircular canals
3. Electroreception
4. Lateral line system with multi cellular neuromasts
5. A number of additional soft tissue specializations

Structures Common to Craniates

Notochord and Vertebral Column
Notochord first skeletal structure to appear in craniates embryos
o At its peak of embryonic development, it is a rod of living cells located ventral to the CNS and dorsal to the
alimentary canal extending from the midbrain to the tip of the tail.
o The part of the notochord in the head becomes incorporated in the floor of the skull and except in hagfish, the
part in the trunk and tail becomes surrounded by cartilaginous or bony vertebrae (rigid support)
o Typical vertebra
Centrum deposited around the notochord
Neural arch forms over spinal chord
Processes
o In the tail, a hemal arch may surround the caudal artery and vein
o Fate of notochord in adult craniates is variable
Fishes, urodeles, some lizards persists the length of the trunk and tail, constricted within each
centrum
Modern reptiles, birds and mammals notochord almost obliterated during development
Vestige remains in mammals within the invertebral disks that separate successive centra
Vestige consists of pulpy nucleus (soft spherical mass of CT)
Lampreys notochord grows along with the animal and paired lateral neural cartilages become
perched on the notochord lateral to the spinal cord. (reminiscent of neural arches)
o When a notochord persists as an important part of the adult axial skeleton, it develops a strong outer elastic
and inner fibrous notochord sheath
Pharynx
A pharynx that is perforated by openings to either the exterior or an atrium is common to hemichordates and chordates
Pharynx is a vital part of the craniates embryo
o Produces fish gills, tetrapod lungs, skeleton and musculature of the jaws, endocrine glands
o Gives rise to the middle ear cavity of tetrapods
o Provides in humans, during fetal life the initial cells of the immune system

Pharyngeal pouches and slits

o A series of pharyngeal pouches arises as diverticula of the endoderm of the foregut and the pouches grow
toward the surface of the animal. These pouches establish the limits of the pharynx
o Simultaneously, an ectodermal groove grows out of each pouch
o Soon, only a think membrane, the branchial plate separates the groove from the pouch
o If and when the branchial plate ruptures, a passageway is formed between the pharyngeal lumen and the
exterior of the animal. This passageway is a pharyngeal slit
o The slits may be permanent (fishes for water for gills) or temporary (tetrapods)
o 8 is the largest number of pouches that develop in any jawed craniates. (sharks)
o Living agnathans have as many as 15 slits
o Pharyngeal slits are temporary if the animal is going to live on land.
4/6 pharyngeal pouches in frog embryos give rise to gill slits in tadpoles
In reptiles (including birds) and mammals, the slits are transitory
3/5 pouches in chicks rupture to the exterior and close again
Only one or two of the more anterior pouches of mammals may rupture
Cervical fistulas occasionally seen in human beings are usually the result of failure of the cervical
sinus, housing the 4rd and 4th slits to close
o Although the pharyngeal pouches of tetrapods rarely give rise to permanent slits, the first one becomes the
auditory tube and middle ear cavity of tetrapods and the second persists as the pouch of the palatine tonsil of
mammals. The walls of several pouches give rise to endocrine tissue in all craniates
Pharyngeal Arches
o Each embryonic pouch or slit is separated from the next by a column of tissue pharyngeal arch
o Each arch contains 4 basic components or blastemas from which these components develop
Supportive skeletal elements
Striated muscles that operate the arch
Branches of 5th, 7th, 9th and 10th cranial nerves which innervate the muscles and provide sensory input
to the brain
Aortic arch connects the ventral and dorsal aortas
o The boundaries of a pharyngeal arch can be identified from the exterior when there are ectodermal grooves or
pharyngeal slits to serve as landmarks.
o If the grooves disappear or the slits close (in tetrapods), the boundaries of the arches are lost
o Pharyngeal (or visceral) skeleton skeleton of the pharyngeal arches; collective
 o Branchiomeric muscles of the arches
o First or mandibular arch constituted by upper and lower jaws and associates muscles, nerves and vessels
Develops anterior to the first pouch
o Second or hyoid arch
o Remaining arches referred to by number, starting with the 3 rd (3rd-7th visceral arches; alternatively these latter
arches may be referred to as the 1st-5th branchial arches representing unmodified branchial arches in fish)
Dorsal Hollow Central Nervous System
CNS consists of brain and spinal cord and contains a central lumen the neurocoel
The dorsal location and the lumen result from the observation that the CNS typically arises as a longitudinal neural
groove in what will become the dorsal surface of the embryo
The groove closes over or rolls up and sinks beneath the surface to become a hollow neural tube located dorsal to the
notochord. The tube is wider anteriorly and this part becomes the brain with its ventricles.
Neurulation process of forming the neural tube
In living agnathans and neopterygian fishes (bowfins, gars and teleosts) the basic pattern of Neurulation has become
slightly modified. Instead of forming a groove, the surface ectoderm dorsal to the notochord proliferates a wedge
shaped neural keel. Eventually the keel separates from the surface and a cavity forms within it by rearrangement of the
cells in its interior. The result is a typical dorsal hollow CNS
Cranial and spinal nerves connect the CNS with the various organs of the body
The nerves along with the associated ganglia and plexuses, constitute the PNS
The spinal nerves of most craniates are metameric, arising at the level of each embryonic body segment and passing to
the skin and muscles derived from that segment and to the viscera
Up to 16 cranial nerves arise from the brain of fishes and amphibians and 13-18 in reptiles and mammals (changes are
the result of gains, losses or fusion of cranial nerves or their components.
2 gains or reorganizations in the tetrapods are spinal nerves that became trapped within the skull

Other Craniate Characteristics

Integument

Skin of craniates consists of epidermis (multilayered) and dermis

Many varieties of defensive, lubricatory, nutritive, pheromonal and homeostasis-maintaining glands develop from the
epidermis and empty on the surface in one taxon or another.
The epidermis of terrestrial craniates forms a variety of cornified (horny) appendages such as spines, reptilian scales,
feathers, hair, claws and hooves
Bone in the dermis provides heavy coat of armor for catfishes and other relicts of the Mesozoic era and contributes to
the dense sharp, pointed scales of sharks and to the think, flexible scales of modern bony fishes
Dense fibroid (collagen-bearing) tissue is common in the dermis
In bovines, the dermis is made into leather through a training process
The epidermis of terrestrial craniates has a surface layer of dead (cornified) cells that protect from dehydration

Respiratory Mechanisms

Most craniates carry on external respiration by means of highly vascularized membranes (gills) located on the
pharyngeal arches or derived from the pharyngeal floor (lungs)
In some species, respiration takes place through the skin or the lining of the oral and pharyngeal cavities
In developing embryos in a porous eggshell or within the body of a parent, gaseous exchange occurs via special extra
embryonic membranes
Coelom

Craniate trunk is built like a tube within a tube, the outer tube (body wall) and inner tube (gut) being separated by a
cavity, the coelom
In fishes, amphibians and some non avian reptiles, the coelom is partitioned into a pericardial cavity housing the heart,
and a pleuroperitoneal cavity housing most of the other viscera including lungs in tetrapods
Transverse septum separates the pericardial and pleuroperitoneal cavities
In other reptiles and mammals, the heart occupies a pericardial cavity, each lung occupies a separate pleural cavity and
the digestive tract caudal to the esophagus occupies an abdominal (peritoneal) cavity
In most male mammals, a pair of scrotal cavities houses the testes; scrotal cavities arise as evaginations from
abdominal cavity
The coelom is totally enclosed by a peritoneal membrane that lines the body wall and invests the coelomic viscera
(parietal and visceral peritoneum) these membranes are continuous via dorsal mesenteries and when present, via
ventral mesenteries
 The few visceral organs that do not develop mesenteries chiefly the kidneys lie against the dorsal body wall just
external to the peritoneum therefore they are retroperitoneal (in some species, the kidneys may be suspended in the
coelom Necturus)

Digestive System

The digestive tracts have specialized regions for the acquisition, processing, temporary storage, digestion and
absorption of food and for elimination of unabsorbed residue
These include the oral cavity, pharynx, esophagus, stomach and intestine
Intestine is often coiled or has within it a spiral valve either of which increases the absorptive area without increasing the
body length. The tract may also exhibit one or more diverticula or ceca that play various roles in the digestion
The digestive tract in most craniates terminates in a cloaca, a common chamber that also receives the urinary and
reproductive ducts. It opens to the exterior via vent
In modern fishes and few tetrapods, the cloaca becomes very shallow or nonexistent
In mammals other than monotremes, the embryonic cloaca becomes subdivided into 2-3 passageways each with its
own exit to the exterior. The exit from the intestine is called the anus

Urogenital Organs

Kidneys and gonads arise close together in the roof of the coelom and the 2 systems share certain passageways
Kidneys (nephroi) are the chief organs for elimination of water in those species in which this is necessary (the ease of
diffusion in an aquatic environment is seen as a reduction of water elimination in many marine teleosts. In contrast, the
reduced elimination of water in desert animals represents a highly derived system for resorption)
The kidneys also assist in maintaining an appropriate electrolyte balance in the blood
In most basal fishes, fluid wastes accumulate in the coelom and are removed by simple kidney tubules reminiscent of
the nephridia of earthworms. In most craniates however, more complex kidney tubules collect fluids directly from blood
capillaries. In both instances, the tubules lead to a pair of longitudinal ducts that empty into the cloaca or a urinary
bladder or less commonly lead to the exterior
Reproductive organs include gonads, ducts, accessory glands, storage chambers and copulatory mechanisms
Early in development, all craniates are bisexual having gonadal and duct primordial for both sexes
Jawless craniates lack reproductive ducts. Their sperm and eggs are shed into the coelom and exit via a urogenital
papilla located immediately behind the anus

Circulatory System

Whole blood, consisting of plasma and formed elements chiefly RBC and WBC and platelets is confined to arteries,
veins, capillaries and sinusoids (sinusoids are broad channels rather than tubes they are a prominent feature of
invertebrate circulatory systems and are more common in fishes than tetrapods)
Craniates also have lymph system that collect some of the interstitial tissue fluids and conduct them to large veins
The heart which forms immediately ventral to the embryonic pharynx, remains in that location, close to the gills in fishes.
In tetrapods, it is somewhat displaced caudad during subsequent development.
The heart pumps blood forward into a ventral aorta which is foreshortened in tetrapods through aortic arches and into a
dorsal aorta in which blood courses caudad.
Branches from the anterior most aortic arches carry blood to the head.
Fishes have a single circuit heart. Blood passes from the heart to the gills where oxygen is acquired then to the body
tissues where oxygen is given up and then back to the heart to be recirculated. The loss of gills and reliance on lungs
eventually resulted in 2-circuit hearts.

Skeleton

Cartilage, bones and ligaments make up a jointed framework of rigid components that give the body its shape, protects
the vital organs and provides the site of attachment of locomotors and other muscles
The framework consists of a longitudinal axial skeleton, principally skull and vertebral column; a pharyngeal skeleton,
best developed in fishes in which it supports the gills and an appendicular skeleton.
As in invertebrates, mineralization of ordinary CT gives skeletal components their rigidity

Muscles

The axial skeleton is moved principally by trunk and tail muscles that are metameric. The appendicular skeleton is
operated primitively by budlike extensions of the body wall muscles onto the fins or limbs.
Muscles of the pharyngeal arches (branchiomeric muscles) operate the pharyngeal skeleton
All these muscles appear striated when viewed by light microscopy
 Cardiac muscle is a special variety of striated muscle
Non-striated (smooth) muscles are found principally in the walls of tubes and vessels

Sense Organs
Craniates have a wider variety of sense organs than any other animals which in craniates are concentrated anteriorly on
the head. These monitor the constantly changing external and internal environments.
Exteroreceptors: mechanoreceptors, chemoreceptors, electroreceptors and thermoreceptors and receptors for radiation
Mechanoreceptors are stimulated chiefly by vibrations in the water or air, by barometric pressure or direct contact
With the exception of receptors for radiation, exteroreceptors are distributed widely on the head trunk and tail of fishes.
In tetrapods, exteroreceptors are confined to the head except those for touch and temperature
Proprioreceptors monitor the activity of muscles, joints and tendons
Visceral receptors monitor the rest of the internal environment