Complete Chordata Notes

CHORDATA NOTES
I PAPER- I SEMESTER
M.SC. PREVIOUS
ZOOLOGY
DR.C.V.NARASIMHA MURTHY
M.Sc.,Ph.D.,M.Ed.,
P.G.DIP IN HIGHER EDUCATION,
P.G.DIPLOMA IN JOURNALISM
P.G.DIPLOMA IN PUBLICRELATIONS AND ADVERTISING
PROFICIENCY IN GENETIC ENGENEERING
ASSOCIATE PROFESSOR (CONTRACT)

DEPARTMENT OF ZOOLOGY
V.S.U.P.G.CENTER
KAVALI
2016
CHORDATA
UNIT III :
3.1 Evolutionary time scale, Eras, Periods & Epoch major events in evolutionary
Time Scale
3.2 Vertebrate integument and derivatives, Skin structure and function, Glands,
Scales, Horns, Claws, Nails, Hoofs, Feathers, Hair
3.3 Comparative anatomy of Heart, Aortic arches and portal system
3.4 Comparative account of excretory system.
UNIT IV:
4.1 Comparative Anatomy of Respiratory organs

4.2 Comparative anatomy of Brain and Spinal cord
4.3 Organs of vision: Structure and functional significance.
4.4 Organs of Hearing and tactile responses
SUGGESTED READING MATERIAL:

1. Barrington, EJ. W. Invertebrate Structure and Function. Thomas Nelson and Sons
Ltd., London.
2. Jagerstein, G. Evolution of Metazoan life cycle, Academic Press, New York &
London.
3. Hyman, L.H. The Invertebrates. Vol. 1-8. Mc Graw Hill Co., New York and London.
4. Barmes, R.D. Invertebrate Zoology, III edition. W.B. Saunders Co., Philadelphia.
7. Russel-Hwlter, W.D. Biology of Higher Invertebrates. The Mc Milan Co. Ltd.,
London.
8. Hyman, L.B. The Invertebrates Smaller Coelomate Groups, Vol. V. Mc.GrawHill,
Co., New York
9. Sedgwick, A. A Student Text Book of Zoology, Vol.II and III. Central Book Depot,
Allahabad
10. Parker, TJ., Haswell, W.A. Text Book of Zoology, Mc Milan Co.,
London.
11. Alexander, R.M. The Chordata. Cambridge University Press, London
12. Barrington, EJ. W. The Biology of Hemichordata and Protochordata. Oliver
and Boyd, Edinburgh.
13 Bourne, GH. The Structure and Functions of Nervous Tissue. Academic Press,
New York
14 Carter, GS. Structure and Habit Invertebrate Evolution Sedgwick and Jackson,
London.
2
Dr.C.V.Narasimha Murthy , M.Sc. Zoology- previous – chordata Notes 2016.
QESTION BANK
Short answers
1. Feathers
2. Scales
3. Spinal cord
4. Tactile Response
5. Nails
6. Malphegian body
7. Lungs
8. Fangs
Essay questions
1. Write about integument and its derivatives?
2. Write an essay on comparative anatomy of heart?
3. Write an essay on comparative anatomy of respiratory system.
4. Write an essay on organs of vision and hearing.
5. Give comparative account of aortic arches and Portal system?
6. Explain in detail about the evolutionary time scale?
8. Write an essay on organs of vision in detail
V.S.UNIVERSISTY, NELLORE
M.Sc. ZOOLOGY
FIRST YEAR - SEMISTER-I
PAPER –I – NOVEMBER 2015.
TIME: THREE HOURS MAX MARKS; 70
PART –A
ANSWER ANY FOUR QUESTIONS (EACH QUESTION CARRIES 5 MARKS) = 20
1.
2.
3.
4.
5. Feathers
6. Scales
7. Spinal cord
8. Tactile Response
PART -B
ANSWER ANY ONE QUESTION (EACH QUESTION CARRIES 12.5 MARKS) = 50
9.
A) .
3
B) .
Or
10. c).
d) .
11.
a).
b).
Or
12. c) .
d) .
UNIT III
13. Write about integument and its derivatives?
Or
14. Write an essay on comparative anatomy of heart?
UNIT IV
Or
16. Write an essay on organs of vision and hearing.
V.S.UNIVERSITY, NELLORE
M.Sc. ZOOLOGY
FIRST YEAR - SEMISTER-I
PAPER –I. NOVMBER 2014.
TIME: THREE HOURS MAX MARKS; 70
PART –A
ANSWER ANY FOUR QUESTIONS (EACH QUESTION CARRIES 5 MARKS) = 20
1.
2.
3.
4.
5. Nails
6. Malphegian body
7. Lungs
8. Fangs
PART -B
ANSWER ANY ONE QUESTION (EACH QUESTION CARRIES 12.5 MARKS) = 50
9.
A).
B).
Or
10. c).
d) .
11.
4
a).
b).
Or
12. c) .
d) .
UNIT III
13. Give comparative account of aortic arches and Portal system?
or
14, Explain in detail about the evolutionary time scale?
UNIT IV
Or
16. Write an essay on organs of vision in detail
GEOLOGICAL TIME SCALE
EVOLUTINARY TIME SCALE:
Define evolutionary time scale:
The Evolutionary Time scale or Geological time scale (GTS) is a system of

chronological measurement that relates stratigraphy to time, and is used by geologists,
paleontologists, and other Earth scientists to describe the timing and relationships
between events that have occurred throughout Earth’s history. The table of
Evolutionary scale is given below:
What does the Evolutionary time scale represent?
The geologic time scale divides up the history of the earth based on life-forms that
have existed during specific times since the creation of the planet. These divisions are
called geochronologic units (geo: rock, chronology: time). Most of these life-forms are
found as fossils, which are the remains or traces of an organism from the geologic past
that has been preserved in sediment or rock. Without fossils, scientists may not have
concluded that the earth has a history that long precedes mankind.
Divisions of Evolutionary time scale
The Geologic Time Scale is divided by the following divisions:
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1. Eons: Longest subdivision; based on the abundance of certain fossils
2. Eras: Next to longest subdivision; marked by major changes in the fossil record
3. Periods: Based on types of life existing at the time
Epochs: Shortest subdivision; marked by differences in life forms and can vary from
continent to continent.
Due to the fact that early geologists had no way of knowing how the discoveries of the
Earth were going to develop, geologist over time have put the time scale together piece
by piece. Units were named as they were discovered. Sometimes unit names were
borrowed from local geography, from a person, or from the type of rock that dominated
the unit.
Cambrian: From the Latin name for Wales. Named for exposures of strata found in a
type-section in Wales by British geologist Adam Sedgwick.
Devonian: Named after significant outcrops first discovered near Devonshire, England
Jurassic: Named for representative strata first seen in the Jura Mountains by German
geologist Humboldt in 1795)
Cretaceous: From the Latin “creta” meaning chalk by a Belgian geologist
While the units making up the time scale are called geochronologic units, the actual
rocks formed during those specific time intervals are called chronostratigraphic units.
The actual rock record of a period is called a system, so rocks from the Cambrian Period
are of the Cambrian system
The earliest time of the Earth is called the Hadean and refers to a period of time for
which we have no rock record, and the Archean followed, which corresponds to the ages
of the oldest known rocks on earth. These, with the Proterozoic Eon are called the
Precambrian Eon. The remainder of geologic time, including present day, belongs to the
Phanerozoic Eon.
While the units making up the time scale are called geochronologic units, the actual
rocks formed during those specific time intervals are called chronostratigraphic units.
The earliest time of the Earth is called the Hadean and refers to a period of time for
which we have no rock record, and the Archean followed, which corresponds to the ages
of the oldest known rocks on earth. These, with the Proterozoic Eon are called the
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Precambrian Eon. The remainder of geologic time, including present day, belongs to the
Phanerozoic Eon. The actual rock record of a period is called a system, so rocks from
the Cambrian Period are of the Cambrian system.
The oldest known rocks are about 3.8 billion (3800 million) years old. It
corresponds to 38, 000, 000 centuries. According to Geologists and Geo- physicists our
planet earth is 4.7 billion (4,700 million) years old.
In this ancient time scale evidences showed that the first life originated
billion (2,500 million) years ago. Hence from the formative stage for nearly 2,200
billion years there was no life on earth. Such a period in earth’s history is
known as the Azoic era (2,200 to 5,000 billion years ago). Evidences show that during
this era the earth was a hot sphere. Gradually the upper surface of the earth cooled
down. It resulted in solidification and formation of rocks and rocky terrain. Further
water molecules were formed resulting in accumulation of water and water reservoirs as
water masses and land surface were established. These transformations provided a
suitable condition for the origin of the first life.
Once the life originated and established, evolutionary changes took place.
Changes happened in the structure, organization and living methods of organisms,
depending on natural surroundings and changes in natural surroundings. Thus fauna and
flora started flourishing on earth. The water masses were fully exploited. Later land
surfaces were invaded by plants and animals.
The Geological time succeeding Azoic Era, was dramatic and rich in life. This
period is divided into three eras. These were significant periods in earth’s history. Of
these, the oldest era was the Paleozoic era. It ranged from 600 to 210 million years ago.
Thus its duration was nearly 390 million years. This era saw the origin and adaptive
radiation of sponges, starfishes, snails, insects, crabs, and terrestialised amphibians and
reptiles.
The Paleozoic era was followed by middle period named as Mesozoic era. This
era ranged from 65 to 210 million years ago. Its duration was 145 million years. During
this era, among animals the reptiles came to prominence. Hence this era is known as the
golden age of reptiles. Further this era saw the origin and development of birds and
reptiles.The period ranging from 65 million years till date is named as the Cenozoic era.
This era is characterized by rapid evolutionary changes in mammals. This era is known
as the Age of mammals.
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GEOLOGICAL TIME SCALE
A geological calendar has been formulated by assessing the age of rocks and
rock sediments. Based on age, and events, the ancient period from
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Era Duration Significance
Paleozoic 600 - 210 m. years ago “Cradle of ancient life”.
Mesozoic 210 - 65 m. years ago “Golden age of reptiles”.

f l”
earth’s history is formulated into eras-periods-epochs. Each division in the geological

calendar is clearly identified and demarcated. Incidents pertaining to earth surface, plant
and animals life are neatly recorded. The influence of geological and climatic changes
on the life and the evolution of the living organism had been well analyzed.
E P i d E h D ti
Quaternary Pleistocene 2-1
Tertiary Pliocene 7-2
Miocene 26 - 7
CENOZOIC Oligocene 38 - 26
Eocene 54 - 38
Paleocene 65 - 54
Cretaceous 130 - 65
Jurassic 160 - 130
Triassic 210 - 160
MESOZOIC
Permian 235 - 210
Pennsylvanian 255 - 235
Mississippian 275 - 255
Devonian 315 - 275
Silurian 350 - 315
Ordovician 440 - 350
Cambrian 600 - 440
Precambrian 440 and before
Periods Events
Quaternary - Pleistocene 2 - 1 m.years ago Human evolution
TERTIARY - Pliocene 7 - 2 m. years ago Rodents were successful
Mammals increased
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TERTIARY - Miocene 26 - 7 m. years ago Prairies were formed Horses
evolved. Carnivorous mammals
were dominant
TERTIARY - Oligocene 38 - 26 m. years ago Monkeys and apes originated
TERTIARY - Eocene 54 - 38 m. years ago Horses originated

TERTIARY - Paleocene 65 - 54 m. years ago First flowering plants Mammals
originated Dinosaurs disappeared.
CRETACEOUS 130 - 65 m.years ago Dinosaurs became extinct

JURASSIC 160 - 130 m. years ago Birds originated Modern bony
fishes.
TRIASSIC 210 - 160 m. years ago Dinosaurs evolved. Mammals
originated.
PERMIAN 235 - 210 m. years ago Origin of reptiles.
PENNSYLVANIAN 255 - 235 m. years ago Land living insects.
MISSISSIPPIAN 275 - 255 m. years ago Origin of Amphibia. Land living
insects, Forests.
DEVONIAN 315 - 275 m. years ago Age of fishes. Ferns and cycas.
SILURIAN 350 - 315 m. years ago Jawed fishes originated

ORDOVICIAN 440 - 350 m. years ago First vertebrates,
CAMBRIAN 600 - 440 m. years ago Thallophytes, Arthopods,

Molluscs, Echinoderms.
PRECAMBRIAN before 440 m. years ago Protozoans,
Poriferans and Annelids lived.
Paleozoic era :- This era produced revolutionary changes in the biosphere. Further this
era saw the origin and the radiation of several groups of animals and plants that
remained as the forefathers for the modern groups. Thus this era is known as the Cradle
of ancient life.
Cambrian period :- (600 to 440 million years ago) The period preceding Cambrian is
known as Pre-Cambrian period. During Precambrian time simple algae, protozoans,
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Poriferans, annelids, were well established. Thus the Cambrian started with the plants
and animals that were successful during the Precambrian period. During Cambrian
among plants Thallophytes were well established. They diversified into various groups.
(Chlorophyceae, Rhodophyceae etc.,). Among animals the aquatic arthropods and
echinoderms came to prominence. The fossils of such organism were obtained from
several places.
Ordovician period :- (440 to 350 million years ago)
This period was marked by formation of

coral rocks and mollusks and echinoderms.
Among plants the semi terrestrial bryophytes
were getting established. Interestingly this period
saw the origin of first vertebrates. These were the now extinct agnatha. (Jawless,
armoured fishes). The origin of early vertebrates was the major event that happened in
the evolution of animals. Among arthropods, the trilobites were more prominent during
this period.
Silurian period :- (350 to 315 million years ago)
The oldest and plant originated in this period. These plants possessed
conducting tissues. They colonized the land. Among invertebrates except for insects all
others flourished. The corals diversified. Several coral islands were formed. Jawed
fishes originated. The fishes developed scales and paired fins, for the first time jaws
originated in fishes. Origin of paired fins and jaws is considered as major events in
chordate evolution.
Devonian Period :- (315 to 275 million years ago)
It is a significant period in the Paleozoic era. During this period land living
plants were more successful. The forests were filled with varieties of ferns and cycas.
(non-flowering plants). Among aquatic animals fishes became dominant. They
diversified by adapting themselves to live in various aquatic ecosystems. The forefathers
of almost all modern fishes lived during this period. Due to these reasons this period is
called as the Age of fishes.
Mississippian Period :- (275 to 255 million years ago)
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The Auustralian lungg fish - Neocceratodus
Severaal changes haappened to the t land struccture. There were massivve upraisingg of land in
severaal places. Thiis resulted inn the formatiion of severaal mountain ranges. Hugge water
bodiess were brokeen into smaller lakes. Theese major ch hanges on eaarth’s surfacee were
known n as revolutio
ons(eg. Caleedonian revoolution). Suchh changes w were the causse for the
origin of lungs in fishes.
f Lunggs evolved foor the purposse of living ttemporarily on
o land. It
helpedd such fishes to find new w water bodiees. Such pracctices encouuraged the orrigin of the
amphibbians. The origin
o of landd living ampphibians were further inccreased by thhe
prolifeeration of sev
veral land livving insects..
Pennsylvanian :- (255 to 235 million yeaars ago)
The land liiving forms became

b morre successfull during this period. Therre were
huge forests
f of ferrns and cycass. Due to geootectonic chhanges severaal forests got buried
under the soil. Todday’s coal annd petroleum
m are obtaineed from suchh resources only.
o
Hence the Pennsyllvanian and thet earlier MMississippiann were colleectively know wn as
Carbonniferous (carrbon bearingg) period.
Permiian Period :- (235 to 210 million yeears ago)
It was the last periiod in the Paaleozoic era. This period was markedd by extinctioons of
severaal older groupps of animalls and plantss. Nearly 60%% of the organism that su urvived at
that tim
me became extinct.
e Somme of the ampphibians drammatically laiid land eggs (cleidoic
eggs). Specificallyy the group ofo organismss that laid such eggs are identified ass
Seymoouria. These are considerred as inter-cconnecting links
l betweeen amphibianns and
reptiles.
Mesozzoic Era :-
This middlle period in the

t history oof life was marked
m by thee prominence of land
living forms. Amoong animals thet reptiles bbecame morre dominant. They increaased in size
and in number. Heence this era is named ass the Golden age of reptiles.
Triasssic Period :-- (210 to 1600 million yeaars ago)
For the firsst time fossills of turtles, crocodiles, and

a dinosaurrs have beenn obtained
from thhis period. Fossil
F evidennces show thhat aquatic anda flying reptiles thrived during
this tim
me. The mam mmals originnated from reeptiles durin ng this periodd.
12
Dr.C.V.Narasimha Muurthy , M.Sc. Z
Zoology- previious – chordataa Notes 2016.
Jurassic Period :- (160 to 130 million years ago)
There was a marked adaptive radiation among dinosaurs. They diversified into
carnivores and herbivores forms. The first birds originated from the reptiles. The earliest
bird thus originated is known as the Archaeopteryx. The origin of birds was a major
physiological change among animals. From a more common poikilothermic condition
through feathers the birds became homoeothermic. The modern bony fishes were
diversified into several groups.
Cretaceous Period :- (130 to 65 million years ago)
The larger marine mollusks became extinct. The fossils of such organisms are available
in places like Ariyaloor, of Tamil Nadu, today.
The Dinosaurs of the Mesozoic era abruptly became extinct during this period.
Several reasons are given for the extinction of the dinosaurs. Fossils of dinosaurs were
not obtained from later periods.
Cenozoic Era :- (65 million years ago till date)
Triceratops - a horned dinosaur
Plenty of fossils of organisms belonging to this era had been obtained. All modern
animals and plants were represented in these fossils. This era is subdivided into Tertiary
and Quaternary periods. Further this era contains seven epochs. Through fossils we can
trace the origin and evolution of independent groups of animals, camels and man.
Paleocene epoch :-
Modern placental mammals originated during this time.
Eocene Epoch :-
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Ungulates originated. The ancestral form of modern horses lived during this epoch.
Oligocene epoch :-
Several animals with ancient characteristics became extinct. Modern mammalian

families were established. The apes originated during this epoch.
Miocene epoch :-
Several varieties of grasses evolved in Europe and N. America. Thus large Prairies were
formed. These changes encouraged the evolution of fast running herbivores mammals
and their predators. Thus the carnivorous mammals came to prominence.
Pliocene Epoch :-
The prairies enlarged still further in several regions. The rodents became more
successful. The mammals increased in number.
Pleistocene epoch :-
Several glaciations happened during this time. This epoch is popularly called the ‘Ice
age’. The evolution of horses and man reached the final stages during this period. The
melting of ice that happened 1,500 years ago is considered as the last stage of this
epoch. Today we are living in an inter-glacial Period.
Evolutionary significance of fossils
Fossils tell us the full story of evolution. Fossil studies reveal the course of evolution.
Through fossils the origin and evolution of specific groups of organisms can be
understood e.g. Horse evolution.
Fossils provide us clues regarding climatic conditions of various prehistoric periods.

Study of fossils simplifies phylogenetic discussions.
Some fossils like woolly mammoth can provide vital clues regarding genetical make up.
Important fossils
Ichthyostega - interconnecting link between fishes and amphibians. Seymouria -

Interconnecting link between Amphibians and Reptiles. Archaeopteryx - Ancestral form
of birds
Dinosaurs - Extinct group of reptiles.
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Hyracotherium - Early ancestor of horses.
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Extinct animals-mass extinction
Extinction may be defined as the termination of a lineage without is- sue or abrupt
disappearance of specific groups of organisms without leaving descendents. Extinctions are of two
types namely true extinctions and pseudo extinction. In true extinctions a particular lineage totally
disappears without any progeny or evolutionary descendents.
Pseudo extinction may also be called as phyletic extinction or phyletic transformation In this type a
group may disappear leaving descendents with evolutionary modifications. In horse evolution while the
earliest ancestor Eohippus became extinct its descendent survived to produce the modern ‘Equus’.
Extinctions can happen for the taxonomic groups such as a family or genus. Thus the extinction of
dinosaurs as a group is a case of true extinction. Similar extinctions happened to trilobites.
The process of extinction is not always predetermined. It happens due to abrupt changes in
environmental conditions or other biological factors. Through the use of computer models evolutionists
can now examine the probability of extinction for large as well as small groups of organisms. From the
fossil record it becomes evident that extinctions have occurred at regular intervals of time.
Pattern of extinction :-
Fossil record reveals a few patterns for extinction. Major groups of herbivorous vertebrates are more
susceptible than the carnivorous vertebrates for extinction. Larger organisms easily became extinct.
VanValen(1973) recorded constancy in the rate of extinction in a number of groups. He explained this
using ‘Mac Arthur’s law’. According to this law as every new adaptation encourages the survival of a
possessor it also decreases a fitness of other related species of that area.
Causes of extinction :-
Even though extinctions are regular events in the history of earth they are caused due to specific
reasons (1) A mass extinction may be due to drastic changes in the environmental conditions. (2) Any
adaptive advance in one species decreases the fitness of all other species. Thus according to Red
Queen’s hypothesis you have to keep running pretty fast, just in order to stay in the same place. (3)
Over specialization to a specific situation may cause extinction.(ex. Antlers.) (4) The spread of an
epidemic disease without any control can cause extinction. (5) An increase in the population strength of
herbivores animals can cause rapid food shortage and cause extinction for several inter-related groups.
(6) A sudden cosmic radiation can cause the death of large organisms. (7) A dust storm formed due to
falling of a meteorite is commonly mentioned as a cause for the disappearance of dinosaurs.
In the recorded history of earth, extinctions of major groups of organisms were due to natural
causes. By the end of Permian period of the Paleozoic Era, nearly 60% of the varieties then existed,
became extinct. Similar large scale extinctions have been observed by the end of Mesozoic era and
during Cenozoic time.
However the extinction of animals and plants during our time is mostly due to human
interference. Thus the cause for the modern extinctions is invariably human activities. The realization
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of a cause lead to starting of several international voluntary agencies to monitor and control
extinctions. “The red-data book brought out regularly by W. W. F (World Wide Fund for nature,
Formerly IUCN - International union for the conservation of nature and natural resources) provides a
list of animals and plants that are endangered or have become extinct.
VERTEBRATE INTEGUMENT AND DERIVATIVES, SKIN STRUCTURE AND FUNCTION,

GLANDS, SCALES, HORNS, CLAWS, NAILS, HOOFS, FEATHERS, HAIR
The word integument means covering. The integumentary system covers the outside of the
body. It protects internal structures, prevents the entry of infectious agents, reduces water loss, regulates
body temperature, produces vitamin D and detects stimuli such as touch, pain and temperature. Since
the integument performs several functions, it is commonly referred to as Jack of all trades.
SKIN STRUCTURE AND FUNCTION
Introduction
The integument or skin is the largest organ of the body, making up 16% of body weight, with a
surface area of 1.8 m2. It has several functions, the most important being to form a physical barrier
to the environment, allowing and limiting the inward and outward passage of water, electrolytes
and various substances while providing protection against micro-organisms, ultraviolet radiation,
toxic agents and mechanical insults. There are three structural layers to the skin: the epidermis, the
dermis and subcutis. Hair, nails, sebaceous, sweat and apocrine glands are regarded as derivatives of
skin (see Figure 1.1). Skin is a dynamic organ in a
constant state of change, as cells of the outer
layers are continuously shed and replaced by
inner cells moving up to the surface. Although
structurally
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Table 1.1 Layers of the skin.
Skin layer Description
Epidermis The external layer mainly composed of layers of keratinocytes but

also containing melanocytes, Langerhans cells and Merkel cells.
Basement membrane The multilayered structure

forming the dermoepidermal
junction.
Dermis The area of supportive connective tissue between the epidermis and the
underlying subcutis: contains sweat glands, hair roots, nervous cells and
fibres, blood and lymph vessels.
Subcutis The layer of loose connective tissue and fat beneath the dermis.
consistent throughout the body, skin varies in thickness according to anatomical site and age of the
individual.
Skin anatomy
The epidermis is the outer layer, serving as the physical and chemical barrier between the interior
body and exterior environment; the dermis is the deeper layer providing the structural support of
the skin, below which is a loose connective tissue layer, the subcutis or hypodermis which is
an important depot of fat (see Table 1.1).
Epidermis
The epidermis is stratified squamous epithelium. The main cells of the epidermis are the
keratinocytes, which synthesize the protein keratin. Protein bridges called desmosomes connect
the keratinocytes, which are in a constant state of transition from the deeper layers to the
superficial (see Figure 1.2). The four separate layers of the epidermis are formed by the differing
stages of keratin maturation. The epidermis varies in thickness from 0.05 mm on the eyelids to 0.8-
1.5 mm on the soles of the feet and palms of the hand. Moving from the lower layers upwards to
the surface, the four layers of the epidermis are:
1. stratum basale (basal or germinativum cell layer)
2. stratum spinosum (spinous or prickle cell layer)
3. stratum granulosum (granular cell layer)
4. stratum corneum (horny layer).
In addition, the stratum lucidum is a thin layer of translucent cells seen in thick epidermis. It
represents a transition from the stratum granulosum and stratum corneum and is not usually seen
in thin epidermis. Together, the stratum spinosum and stratum granulosum are sometimes referred
to as the Malphegian layer.
Stratum basale
The innermost layer of the epidermis which lies adjacent to the dermis comprises mainly dividing
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and non-dividing keratinocytes, which are attached to the basement membrane by

hemidesmosomes. As keratinocytes divide and differentiate, they move from this deeper layer to
the surface. Making up a small proportion of the basal cell population is the pigment
(melanin) producing melanocytes. These cells are characterized by dendritric processes, which stretch
between relatively large numbers of neighboring keratinocytes. Melanin accumulates in
melanosomes that are transferred to the adjacent keratinocytes where they remain as granules.
Melanin pigment provides protection against ultraviolet (UV) radiation; chronic exposure to light
increases the ratio of melanocytes to keratinocytes, so more are found in facial skin compared to the
lower back and a greater number on the outer arm compared to the inner arm. The number of
melanocytes is the same in equivalent body sites in white and black skin but the distribution and
rate of production of melanin is different. Intrinsic ageing diminishes the melanocyte population.
Merkel cells are also found in the basal layer with large numbers in touch- sensitive sites such
as the fingertips and lips. They are closely associated with cutaneous nerves and seem to be
involved in light touch sensation.
Stratum spinosum
As basal cells reproduce and mature, they move towards the outer layer of skin, initially forming the
stratum spinosum. Intercellular bridges, the desmosomes, which appear as 'prickles' at a
microscopic level, connect the cells. Langerhans cells are dendritric, immunologically active cells
derived from the bone marrow,
and are found on all epidermal surfaces but are mainly located in the middle of this layer. They play a
significant role in immune reactions of the skin, acting as antigen-presenting cells.
Stratum granulosum
Continuing their transition to the surface the cells continue to flatten, lose their nuclei and their
cytoplasm appears granular at this level.
Stratum corneum
The final outcome of keratinocytes maturation is found in the stratum corneum, which is made up
of layers of hexagonal-shaped, non-viable cornified cells known as corneocytes. In most areas of the
skin, there are 10-30 layers of stacked corneocytes with the palms and soles having the most. Each
corneocyte is surrounded by a protein envelope and is filled with water-retaining keratin
proteins. The cellular shape and orientation of the keratin proteins add strength to the stratum
corneum. Surrounding the cells in the extracellular space are stacked layers of lipid bilayer (see
Figure 1.3).
The resulting structure provides the natural physical and water-retaining barrier of the skin.
The corneocyte layer can absorb three times its weight in water but if its water content drops
below 10% it no longer remains pliable and cracks. The movement of epidermal cells to this layer
usually takes about 28 days and is known as the epidermal transit time.
Dermoepidermal junction/basement membrane
This is a complex structure composed of two layers. Abnormalities here result in the expression of rare
skin diseases such as bullous pemphigoid and epidermolysis bullosa. The structure is highly
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irregular, with dermal papillae from the papillary dermis projecting perpendicular to the skin
surface. It is via diffusion at this junction that the epidermis obtains nutrients and disposes of waste.
The dermoepidermal junction flattens during ageing which accounts in part for some of the visual
signs of ageing.
Dermis
The dermis varies in thickness, ranging from 0.6 mm on the eyelids to 3 mm on the back, palms and
soles. It is found below the epidermis and is
composed of a tough, supportive cell matrix. Two
layers comprise the dermis:
1. A thin papillary layer

2. A thicker reticular layer.
The papillary dermis lies below and connects with

the epidermis. It contains thin loosely arranged
collagen fibres. Thicker bundles of collagen run
parallel to the skin surface in the deeper reticular
layer, which extends from the base of the papillary
layer to the subcutis tissue. The dermis is made up of
fibroblasts, which produce collagen, elastin and
structural proteoglycans, together with Immuno-
competent mast cells and macrophages. Collagen
fibres make up 70% of the dermis, giving it strength and toughness. Elastin maintains normal
elasticity and flexibility while proteoglycans provide viscosity and hydration. Embedded
within the fibrous tissue of the dermis are the dermal vasculature, lymphatics, nervous cells and
fibres, sweat glands, hair roots and small quantities of striated muscle.
Subcutis
This is made up of loose connective tissue and fat, which can be up to 3 cm thick on the abdomen.
Blood and lymphatic vessels
The dermis receives a rich blood supply. A superficial artery plexus is formed at the papillary and
reticular dermal boundary by branches of the subcutis artery Branches from this plexus form
capillary loops in the papillae of the dermis, each with a single loop of capillary vessels, one arterial
and one venous. The veins drain into mid-dermal and subcutaneous venous networks. Dilatation or
constriction of these capillary loops plays a direct role in thermoregulation of the skin. Lymphatic
drainage of the skin occurs through abundant lymphatic meshes that originate in the papillae and feed
into larger lymphatic vessels that drain into regional lymph nodes.
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Nerve supply
The skin has a rich innervation with the hands, face and genitalia having the highest density of
nerves. All cutaneous nerves have their cell bodies in the dorsal root ganglia and both myelinated
and non-myelinated fibres are found. Free sensory nerve endings lie in the dermis where they detect
pain, itch and temperature. Specialized corpuscular receptors also lie in the dermis allowing
sensations of touch to be received by Meissner's corpuscles and pressure and vibration by Pacinian
corpuscles. The autonomic nervous system supplies the motor innervation of the skin: adrenergic
fibres innervate blood vessels, hair erector muscles and apocrine glands while cholinergic fibres
innervate exocrine sweat glands. The endocrine system regulates the sebaceous glands, which are not
innervated by autonomic fibres.
Derivative structures of the skin
Hair
Hair can be found in varying densities of growth over the entire surface of the body, exceptions
being on the palms, soles and glans penis. Follicles are most dense on the scalp and face and are
derived from the epidermis and the dermis. Each hair follicle is lined by germinative cells, which
produce keratin and melanocytes, which synthesize pigment. The hair shaft consists of an outer
cuticle, a cortex of keratinocytes and an inner medulla. The root sheath, which surrounds the hair
bulb, is composed of an outer and inner layer. An erector pili muscle is associated with the hair shaft
and contracts with cold, fear and emotion to pull the hair erect, giving the skin 'goose bumps'.
Nails
Nails consist of a dense plate of hardened keratin between 0.3 and 0.5 mm thick. Fingernails function
to protect the tip of the fingers and to aid grasping. The nail is made up of a nail bed, nail matrix and a
nail plate. The nail matrix is composed of dividing keratinocytes, which mature and keratinise into
the nail plate. Underneath the nail plate lies the nail bed. The nail plate appears pink due to adjacent
dermal capillaries and the white lunula at the base of the plate is the distal, visible part of the matrix.
The thickened epidermis which underlies the free margin of the nail at the proximal end is called
the hyponychium. Fingernails grow at 0.1 mm per day; the toenails more slowly (see Figure 1.4).
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Figure 1.4 Nail structure.
Sebaceous glands
These glands are derived from epidermal cells and are closely associated with hair follicles especially
those of the scalp, face, chest and back; they are not found in hairless areas. They are small in
children, enlarging and becoming active at puberty, being sensitive to androgens. They produce an
oily sebum by holocrine secretion in which the cells break down and release their lipid cytoplasm. The
full function of sebum is unknown at present but it does play a role in the following:1
1. Maintaining the epidermal permeability barrier, structure and differentiation

2. Skin-specific hormonal signaling
3. Transporting antioxidants to the skin surface
4. Protection from uv radiation.
Sweat glands
There are thought to be over 2.5 million on the skin surface and they are present over the
majority of the body. They are located within the dermis and are composed of coiled tubes, which
secrete a watery substance. They are classified into two different types: exocrine and apocrine.
• Eccrine glands are found all over the skin especially on the palms, soles, axillae and forehead. They
are under psychological and thermal control. Sympathetic (cholinergic) nerve fibres innervate
exocrine glands. The watery fluid they secrete contains chloride, lactic acid, fatty acids, urea,
glycoproteins and mucopolysaccharides.
• Apocrine glands are larger, the ducts of which empty out into the hair follicles.
They are present in the axillae, anogenital region and areolae and are under thermal control. They
become active at puberty, producing an odourless protein-rich secretion which when acted
upon by skin bacteria gives out a characteristic odour. These glands are under the control of
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sympathetic (adrenergic) nerve fibres.
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Skin functions
The skin is a complex metabolically active organ, which performs important physiological
functions that are summarized in Table 1.2.
Table 1.2 Functions of the skin.
• Provides a protective barrier against mechanical, thermal and physical injury and noxious
agents.
• Prevents loss of moisture.
• Reduces the harmful effects of UV radiation.
• Acts as a sensory organ.
• Helps regulate temperature control.
• Plays a role in immunological surveillance.
• Synthesizes vitamin D3 (cholecalciferol).
• Has cosmetic, social and sexual associations.
Barrier function and skin desquamation
As the viable cells move towards the stratum corneum they begin to clump proteins into
granules in the granular layer (see Figure 1.5). The granules are filled with the protein fillagrin,
which becomes complexed with keratin to prevent the breakdown of fillagrin by proteolytic
enzymes. As the degenerating cells move towards the outer layer, enzymes break down the
keratin-fillagrin complex. Fillagrin forms on the outside of the corneocytes while the water-
retaining keratin remains inside.
When the moisture content of the skin reduces, fillagrin is further broken down into free
amino acids by specific proteolytic enzymes in the stratum corneum. The breakdown of
fillagrin only occurs when the skin is dry in order to control the osmotic pressure and the amount of
water it holds: in healthy skin the water content of the stratum corneum is normally around 30%.
The free amino acids, along with other components such as lactic acid, urea and salts, are known as
'natural moisturizing factors' (NMF) and are responsible for keeping the skin moist and pliable due
to their ability to attract and hold water.
Lipids
The major factor in the maintenance of a moist, pliable skin barrier is the presence of
intercellular lipids. These form stacked bilayer that surround the corneocytes and incorporate water
into the stratum corneum. The lipids are derived from lamellar granules, which are released into
extracellular spaces of degrading cells in the granular layer; the membranes of these cells also release
lipids. Lipids include cholesterol, free fatty acids and sphingolipids.
Ceramide, a type of sphingolipid, is mainly responsible for generating the stacked lipid
structures that trap water molecules in their hydrophilic region. These stacked lipids surround the
corneocytes and provide an impermeable barrier by preventing the movement of water and NMF
out of the surface layers of the skin. After the age of 40 there is a sharp decline in skin lipids
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thus increasing our susceptibility to dry skin conditions.
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26
Figure 1.5 Barrier function of the epidermis.
Shedding (desquamation) of skin cells

Shedding the cells of the stratum corneum is an important factor in maintaining skin integrity and
smoothness. Desquamation involves the enzymatic process of dissolving the protein bridges, the
desmosomes, between the corneocytes, and the eventual shedding of these cells (see Figure 1.2).
The proteolytic enzymes responsible for desquamation are located intracellularly and function in
the presence of a well-hydrated stratum corneum. In the absence of water the cells do not
desquamate normally and the skin becomes roughened, dry, thickened and scaly. In normal healthy
skin there is a balance in the production and shedding of corneocytes. In diseases such as psoriasis,
which involve increased corneocyte production, and where decreases in shedding occur, the result
is dry, rough skin as the cells accumulate on the skin.
Remaining functions
UV protection
Melanocytes, located in the basal layer, and melanin have important roles in the skin's barrier function
by preventing damage by UV radiation. In the inner layers of the epidermis, melanin granules form a
protective shield over the nuclei of the keratinocytes; in the outer layers, they are more evenly
distributed. Melanin
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Table 1.3 Immune components of the skin.
Defense type Component Immune action
Structural Skin Impenetrable physical barrier to most external organisms.
Blood and Provision of transport network for

lymphatic cellular defense.
vessels
Cellular Langerhans cells Antigen presentation.
T lymphocytes Facilitate immune reactions. Self-regulating
through the action of T suppressor cells. Mast cells Facilitate

inflammatory skin reactions.
Keratinocytes Secrete inflammatory cytokines; have ability
to express surface immune reactive molecules.
Systemic Cytokines Cytokines: cell mediation

and chemicals produced by
eicosanoids components of the cellular
defense system.
Eicosanoids: non-specific
inflammatory mediators
produced by mast cells,
macrophages and keratinocytes.
Increase the number of cellular

Adhesi defense facilitators in an area by
on binding to T cells.
molecul
Activation of this initiates a host of
es
destructive mechanisms, including
Complem opsonisation, lysis, chemotaxis and
ent mast cell degranulation.
cascade
Enables immunological
recognition of antigens.
Immunogenetic Major
Histocompatib
ility complex
(MHC)
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absorbs UV radiation, thus protecting the cell's nuclei from DNA (deoxyribonucleic acid)
damage. UV radiation induces keratinocytes proliferation, leading to thickening of the
epidermis.
Thermoregulation
The skin plays an important role in maintaining a constant body temperature through
changes in blood flow in the cutaneous vascular system and evaporation of sweat from the
surface.
Immunological surveillance
As well acting as a physical barrier, skin also plays an important immunological role. It
normally contains all the elements of cellular immunity, with the exception of B cells.2
Immune components of the skin are given.
PROTOCHORDATE SKIN
Protochordata are marine animals and show aquatic adaptations in the skin. Amphioxus
skin is made of the outer epidermis and an inner dermis but epidermis is composed of single
layer of living cells which must be protected on the surface by a layer of cuticle and a basement
membrane that attaches epidermal cells with dermal tissue. Dermis is composed of loose
connective tissue and lacks melanocytes.
SKIN OF CYCLOSTOMATA
Cyclostomes are also marine animals which are parasitic or flesh-feeders. They lack
scales on the skin and are dark in colour but the larval skin is ciliated. Epidermis is made of all
living cells but is thick and multilayered and is covered with a layer of cuticle for protection.
There are unicellular mucous glands in epidermis.
Dermis of cyclostomes is composed of collagenous connective tissue, which is tough and
fibrous tissue that provides strength to the skin. Thus, the cyclostomes skin is simple, without
many modifications as seen in more advanced groups.
FISH SKIN
All fishes being aquatic possess more or less similarly adapted skin. They possess a
multilayered epidermis of living cells that is covered on the surface with a layer of cuticle. Fishes
lack a typical stratum corneum made of dead and cornified cells as found in tetra pods.
Multicellular glands also occur in some fishes, e.g. electric organs of eels and electric rays,
and luminescent glands in deep sea fishes. Mucous glands are all unicellular.
Important modifications of skin are dermal scales that cover the body
for protection.
Kinds of scales. The following types of scales are found in living and extinct
fishes:
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Cosmoid
scales. They
were found
in ancient
crossopteryg
ian fishes of
Devonian
period and
were many
layers thick
and strong
but not
flexible.
Apparently
ancient
crossopteryg
ian, with
thick and
hard scales covering the body, were not active and fast swimming animals.
Ganoid scales. They are thick rhomboidal scales having the surface coated with a hard enamel-
like material called ganoin and lacunae (pits) and canaliculae (fine canals) forming the surface
sculpture.
Cycloid and Ctenoid scales. These scales are found in modern teleosts and are thin, strong and
extremely flexible. They are large, oval in shape and made of isopodan and fibrous tissue.
Placoid scales. They are characteristics of cartilaginous fishes (Chondrichthys) and are hard and
microscopic in size.
Evolutionary modifications in scales of fishes

Ostracoderms and placoderms, which were ancestors of all modern fishes, had their
bodies covered with dermal bony armour for protection from predators. These fishes were
sluggish in movement and were bottom feeders of detritus. To attain flexibility, the bony armour
gave way to cosmoid scales which were still like thick bony plates covering the body. In ganoid
fishes, the spongy bone of cosmoid scales was lost. Cycloid and ctenoid scales of modern
teleosts are very thin and flexible and consist of only isopodan and fibrous tissue. Thus modern
teleosts are agile and quick swimmers.
AMPHIBIAN SKIN
Amphibian skin shows adaptation to terrestrial life but the animals have to return back to
water for reproduction and hence the skin also demonstrate certain adaptation to aquatic mode of
life. On land a major problem faced by the animal is of desiccation and strong solar radiation.
There are no scales on the skin of modern amphibians, although ancestral labyrinthodont
amphibians had dermal scales on their skin for protection. Also, the scaly protection gone,
amphibians evolved mulitcellular poison glands to deter potential predators.
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Amphibian skin is the simplest skin having no particular modifications as found in higher
groups of vertebrates but it has typical structure of terrestrial skin that is subsequently carried
through to reptiles, birds and mammals.
REPTILIAN SKIN
Reptiles are completely independent of water. They do not have to go back to water for
breeding as they are capable of laying a large cleioid, shelled egg that can develop and hatch on
land. Reptiles are adapted to life in arid climate and have very few glands. Then there are dermal
plates on the body of crocodiles and turtles.
Exoskeleton of Chelonia (tortoises and turtles) consists of a dorsal convex carapace and a
ventral flat plastron which are attached by ligaments on the lateral sides. Outer surface of the
carapace and plastron consists of epidermal plates called scutes while the inner side is made of
dermal plates.
Carapace is fused with the vertebral column and attached to the plastron on the sides.
BIRD SKIN
Birds possess thin skin that is loosely attached to body to allow free movement of wings
during flight. Basic structure is of typical tetrapod skin. Feathers are characteristic modifications
of bird skin, which not only cover the entire body but also help in flight. Glands are absent in
bird skin except the uropygial glands near the tail that secrete oil for preening feathers.
Modifications of skin include epidermal scales on the legs and feet, claws, beak, spur, comb etc.
Contour feathers. Also called quill feathers, they are hard feathers that cover the entire body
and form the contour or outline of the body by which we identify the bird. Ventral side of the
shaft has a longitudinal groove called umbilical groove. Barbs on either side of the shaft area
arranged parallel to each other and in such a way that they form a flat Vane that has aerodynamic
shape.
Down feathers. Also called plumules, they are soft feathers that serve as insulation between the
hard contour feathers and the delicate skin. Powder down is a term used to identify such feathers
in adults whereas nestling down appear in chicks.
Hair feathers. They are hair-like thin and long feathers, also called filoplumes or pin feathers
and are scattered all over the body. They are especially modified to guard nostrils, ear openings
and serve as eye lashes. They are made of long thin hair-like shaft to the top of which are
attached a bunch of thin barbs and barbules.
Development of feather
Feathers grown on certain certain tracts on the skin called feather tracts or pterylae and
feathers do not grow on the areas termed as apterylae or apteria but the growth is dense and
hence the entire body is covered with feathers. Precocial chicks as in fowl and other game birds
are covered with nestling down when they hatch from the eggs but altricial chicks such as
sparrow and pigeon hatch naked and development of feather can be observed during the growth
of chicks. First indication of the growth of feathers on skin is the appearance of pimple-like
elevations called feather primordium that changes into feather follicle in the middle of which the
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feather grows. The body of feather is produced by multiplication of epidermal cells assisted by
feather papilla.
MAMMALIAN SKIN
Mammalian skin is thick, tough, glandular and covered with hairs. Dermis is much
thicker than epidermis but general structure exhibits typical terrestrial skin characters. Epidermis
in some areas is very thick and can be differentiated into several layers. Dermis is thick and
composed of collagenous connective tissue which makes the entire skin tough and fibrous.
Mammalian skin has abundance of glands, particularly sudoriferous or sweat glands and
Sebaceous glands which are oil secreting glands attached to the hair follicle. Scent glands or
Hedonic glands found in musk deer and civets are also modifies epidermal glands.
Mammalian skin becomes sensory owing to the presence of large number of free
nerve endings and specialized receptors such as pacinian corpuscles, meissner’s corpuscles,
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tactile discs, Krause’s end bulbs etc. Also the base of hair follicle is supplied with nerve, making
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the hair sensory. Carnivores, rodents and many other mammals possess specialized hairs on the
upper lip called whiskers, which are sensitive to air currents, touch and vibrations.
Development of hair
Hairs grow on the skin in the embryonic stage and continue to grow throughout life.
When a hair has to grow, epidermal cells in that area multiply fast forming a mass of cells called
hair primordium that carries a dermal papilla at the base. Keratinization of epidermal cells inside
the hair follicle produce the hair root and shaft that grows out of skin. Length of the hair is
determined by the wear and tear to which the tip of the hair is exposed and the thickness of the
hair shaft.
Development of Mammary glands

Mammary glands are apocrine compound tubuloacinar glands that specialize in secreting
nourishing fluid called milk that is sucked from teats by the young ones of mammals.
Development of mammary glands is completed during embryonic stage in both male and female
sexes. The position and number of nipple formation along the milk line varies in different
species, e.g. one pair of axillary nipples are formed in flying lemurs, thoracic in primates, bats
and sirenians, one or two pairs of inguinal in ungulates.
When the mammary gland has to develop, epidermal cells of milk line in that particular
location start multiplying fast forming a mass of mammary tissue that grows and branches to
form a mass of considerable size. The development of mammary glands stops at this stage in
both sexes and further development takes place only in female at the time of puberty. Unlike in
hair or feather, development of mammary glands does not need the assistance of dermal papilla.
Monotremes possess primitive mammary glands which are perhaps modified sweat
glands that got attached to the hair follicles.
Horns in mammals
Mammals sport one pair of horns on the head which are organs and defense and offence
and sometimes serve as secondary sexual organs of male. The following five types of horns are
found in mammals:
1. True horns or Hollow horns, as found in family Bovidae that includes cattle.
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2. Pronghorns, found in pronghorn antelope.

3. Antlers, as found in deer family Cervidae.
4. Knob horns or Giraffe horns, found in giraffes.
5. Hair horns or Keratin fibre horns, found in rhinoceroses.
TRUE HORNS. These horns are found in cattle, sheep, goat and antelopes. They may be
straight, curved or spiralling as in antelopes and sheep and are made of bony core and a
keratinized epidermal sheath.
PRONGHORNS. They are found in pronghorn antelope, Antilocapra americana that inhabits
North American plains and also in Saiga antelope of Russia.
ANTLERS. They are found in the males of family Cervidae and are secondary sex organs meant
to attract females as well as to intimidate or fight the rival males. These horns are made of bony
core that grows from the frontal bones of the skull and is covered with living velvety skin.
KNOB HORNS. They are found in giraffes and okapi in both sexes and are permanent
structures. Structurally they are similar to the antlers with bony core covered with velvety skin.
HAIR HORNS. Found in rhinoceros, they are made of specialized hairs or keratinized fibres
that get agglutinated to form a pointed and hard horn.
OTHER MODIFICATIONS OF MAMMALIAN SKIN

1. Many mammals have epidermal scales as on the feet and tail of rats and mice.
2. There are dermal bony plates on the body of armadillo (order Edentata).
3. Ischial or Lilac callosities on monkey’s buttocks are cushion-like structures on
which the animal sits on hard branches and rocks.
4. Knee pads on the legs of camels are meant for absorbing shock when the animals
kneels and sits on the ground.
5. Tori are epidermal pads on fingers, palms and soles of feet of primates that help in
firm grasping branches of trees while climbing.
6. Baleens are horny epidermal plates numbering about 370 that hang from the roof
of the oral cavity of toothless whales. They are used to filter planktons such as
krill on which these giants of the sea feed.
7. Claws, nails and hooves are also cornified structures derived from the epidermis
of skin. They are made of two types of cornified tissue, the outer hard and shiny
tissue called unguis, under which lies a softer tissue called subunguis which lies in
contact with the living tissue.
COMPARATIVE ANATOMY OF HEART, AORTIC ARCHES AND PORTAL SYSTEM
Vertebrate Circulatory Systems:

1. Transport gases, nutrients, waste products, hormones, heat, & various other materials
2. Consist of heart, arteries, capillaries, & veins:
a. Arteries
i. carry blood away from the heart
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ii. have muscular, elastic walls

iii. terminate in capillary beds
b. Capillaries
i. have very thin walls (endothelium only)
ii. are the site of exchange between the blood and body cells
c. Veins
i. carry blood back to the heart
ii. have less muscle in their walls than arteries but the walls are very elastic
iii. begin at the end of capillary beds
d. Heart
i. a muscular pump (cardiac muscle)
ii. contains a pacemaker to regulate rate but rate can also be influenced by
the Autonomic Nervous System
SIMPLE CIRCULATORY SYSTEMS
The circulatory system varies from simple systems in invertebrates to more complex
systems in vertebrates. The simplest animals, such as the sponges (Porifera) and rotifers
(Rotifera), do not need a circulatory system because diffusion allows adequate exchange of
water, nutrients, and waste, as well as dissolved gases (figure a). Organisms that are more
complex, but still have only two layers of cells in their body plan, such as jellies (Cnidaria) and
comb jellies (Ctenophora), also use diffusion through their epidermis and internally through the
gastrovascular compartment. Both their internal and external tissues are bathed in an aqueous
environment and exchange fluids by diffusion on both sides (figure b). Exchange of fluids is
assisted by the pulsing of the jellyfish body.
Animals without circulatory systems
Simple animals consisting of a single cell layer, such as the (a) sponge, or only a few cell
layers, such as the (b) jellyfish, do not have a circulatory system. Instead, gases, nutrients, and
wastes are exchanged by diffusion.
For more complex organisms, diffusion is not efficient for cycling gases, nutrients, and
waste effectively through the body; therefore, more complex circulatory systems evolved. Closed
circulatory systems are a characteristic of vertebrates; however, there are significant differences
in the structure of the heart and the circulation of blood between the different vertebrate groups
due to adaptation during evolution and associated differences in anatomy.
Fish Circulatory Systems
Fish have a single circuit for blood flow and a two-chambered heart that has only a single
atrium and a single ventricle (figure a). The atrium collects blood that has returned from the
body, while the ventricle pumps the blood to the gills where gas exchange occurs and the blood
is re-oxygenated; this is called gill circulation. The blood then continues through the rest of the
body before arriving back at the atrium; this is called systemic circulation. This unidirectional
flow of blood produces a gradient of oxygenated to deoxygenated blood around the fish's
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systemic circuit. The result is a limit in the amount of oxygen that can reach some of the organs
and tissues of the body, reducing the overall metabolic capacity of fish.
(a) Fish have the simplest circulatory systems of the vertebrates: blood flows unidirectional from
the two-chambered heart through the gills and then to the rest of the body.
(b) Amphibians have two circulatory routes: one for oxygenation of the blood through the lungs
and skin, and the other to take oxygen to the rest of the body. The blood is pumped from a three-
chambered heart with two atria and a single ventricle.
(c) Reptiles also have two circulatory routes; however, blood is only oxygenated through the
lungs. The heart is three chambered, but the ventricles are partially separated so some mixing of
oxygenated and deoxygenated blood occurs, except in crocodilians and birds.
(d) Mammals and birds have the most efficient heart with four chambers that completely
separate the oxygenated and deoxygenated blood; it pumps only oxygenated blood through the
body and deoxygenated blood to the lungs.
Amphibian Circulatory Systems

In amphibians, reptiles, birds, and mammals, blood flow is directed in two circuits: one
through the lungs and back to the heart (pulmonary circulation) and the other throughout the rest
of the body and its organs, including the brain (systemic circulation).
Amphibians have a three-chambered heart that has two atria and one ventricle rather than
the two-chambered heart of fish (figure b). The two atria receive blood from the two different
circuits (the lungs and the systems). There is some mixing of the blood in the heart's ventricle,
which reduces the efficiency of oxygenation. The advantage to this arrangement is that high
pressure in the vessels pushes blood to the lungs and body. The mixing is mitigated by a ridge
within the ventricle that diverts oxygen-rich blood through the systemic circulatory system and
deoxygenated blood to the pulmocutaneous circuit where gas exchange occurs in the lungs and
through the skin. For this reason, amphibians are often described as having double circulation.
Reptile Circulatory Systems
Most reptiles also have a three-chambered heart similar to the amphibian heart that directs
blood to the pulmonary and systemic circuits (figure c). The ventricle is divided more effectively
by a partial septum, which results in less mixing of oxygenated and deoxygenated blood. Some
reptiles (alligators and crocodiles) are the most primitive animals to exhibit a four-chambered
heart. Crocodilians have a unique circulatory mechanism where the heart shunts blood from the
lungs toward the stomach and other organs during long periods of submergence; for instance,
while the animal waits for prey or stays underwater waiting for prey to rot. One adaptation
includes two main arteries that leave the same part of the heart: one takes blood to the lungs and
the other provides an alternate route to the stomach and other parts of the body. Two other
adaptations include a hole in the heart between the two ventricles, called the foramen of Panizza,
which allows blood to move from one side of the heart to the other, and specialized connective
tissue that slows the blood flow to the lungs. Together, these adaptations have made crocodiles
and alligators one of the most successfully-evolved animal groups on earth.
Mammal and Bird Circulatory Systems
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In mammals and birds, the heart is also divided into four chambers: two atria and two ventricles
(figure d). The oxygenated blood is separated from the deoxygenated blood, which improves the
efficiency of double circulation and is probably required for the warm-blooded lifestyle of
mammals and birds. The four-chambered heart of birds and mammals evolved independently
from a three-chambered heart.
COMPARATIVE ANATOMY OF HEART
All vertebrates possess a heart – a hollow muscular organ

composed of cardiac muscle fibres. The function of the heart is to
pump oxygen to all parts of the body. The evolution of the heart is
based on the separation of oxygenated blood from deoxygenated
blood for efficient oxygen transport. In fishes, the heart was like a
hollow tube. This evolved into the four-chambered heart in
mammals.
Vertebrate Hearts:
Piscean heart
Fish has only two chambers in its heart – one auricle and
one ventricle. Since both the auricle and the ventricle
remain undivided, only deoxygenated blood
passes through it. The deoxygenated blood
enters the gills for oxygenation from the
ventricle. It has additional chambers such as sinus
venous and conus arteriosus
Cartilaginous fishes
1. Single-circuit heart with 4 chambers: sinus
venous, atrium, ventricle, & conus
arteriosus
a. the sinus venous receives blood & is filled by suction when the ventricle contracts
& enlarges the pericardial cavity
b. the atrium is a thin-walled muscular sac; an A-V valve regulates flow between
atrium & ventricle
c. the ventricle has thick, muscular walls
d. the conus arteriosus leads into the ventral aorta (and a series of conal valves in the
conus arteriosus prevent the backflow of blood)
Teleosts - heart is similar to that of cartilaginous fishes, except a bulbous arteriosus (a muscular
extension of the ventral aorta) is present rather than a conus arteriosus (a muscular extension of
the ventricle)
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Amphibian heart
Amphibians, such as frogs, have three-chambered hearts, with two auricles and one ventricle.
The auricle is divided into a right and a left
chamber by an inter-auricular septum, while the
ventricle remains undivided.
Additional chambers such as sinus venous
and conus arteriosus are also present. The
oxygenated blood from the lungs enters the left
auricle and simultaneously, the deoxygenated
blood from the body enters the right auricle. Both
these auricles empty into the ventricle, wherein
the oxygenated and deoxygenated blood get
mixed to some extent.
Lungfish & amphibians - modifications are

correlated with the presence of lungs & enable oxygenated blood returning
from the lungs to be separated from deoxygenated blood returning from
elsewhere
1. Partial or complete partition within atrium (complete in anurans and
some urodeles)
2. Partial interventricular septum (lungfish) or ventricular trabeculae
(amphibians) to maintain separation of oxygenated & unoxygenated
blood
3. Formation of a spiral valve in the conus arteriosus of many dipnoans
and amphibians. The spiral valve alternately blocks & unblocks the
entrances to the left and right pulmonary arches (sending unoxygenated blood to the skin
& lungs).
4. Shortening of ventral aorta, which helps ensure that the oxygenated & unoxygenated
blood kept separate in the heart moves directly into the appropriate vessels
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5 = ventricle, 11 = right atrium, 12 = left atrium, 13 = conus arteriosus

Amniotes:
1 - Heart consists of 2 atria & 2 ventricles &, except in adult birds & mammals, a sinus venous
2 - Complete intertribal septum
3 - Complete interventricular septum only in crocodilians, birds, & mammals; partial septum
in other amniotes
Used by permission of John W. Kimball
Reptilian heart
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Reptiles have incomplete four-chambered hearts,

except for crocodiles, alligators, and gharials.
They have only one accessory chamber called
sinus venous. The reptilian heart also shows
mixed blood circulation.
Avian and mammalian hearts
They have two pairs of chambers for

separating oxygenated and deoxygenated bloods.
The heart is divided into four chambers. The
upper two chambers are called atria and

the lower two chambers are called ventricles. The
chambers are separated by a muscular wall that
prevents the mixing of the blood rich in oxygen
with the blood rich in carbon dioxide.
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Comparative anatomy of Aortic Arches
Aortic arches are paired blood vessels that emerge from the ventricle of the heart which are
basically similar in number and disposition in different vertebrates during the embryonic stages.
Embryonic aortic arches: During the embryonic stages—
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1. Six pairs of aortic arches develop in most gnathostomes and are named according to the
name of the visceral clefts.
2. These are designated by roman numerals.
3. The first aortic arch named Mandibular, proceeds upwards on either side of the pharynx.
4. Mandibular aortic arch turn backwards as lateral aorta which both join medially to form
the common dorsal aorta.
5. The second aortic arch becomes hyoid arch.
6. The third, fourth, fifth & sixth are called branchial arches.
Modification of aortic arches in different vertebrates:
1. The number of aortic arches is different in different adult vertebrates but they built on the
same fundamental plan in embryonic life.
2. The differences in number of aortic arches are due to the complexity of heart circulation
in the mode of living from aquatic to terrestrial respiration.
3. There is a progressive reduction of aortic arches in the vertebrate series during evolution.
1. Aortic arches in primitive vertebrates:
¾ Branchiostoma (amphioxus) has about 60 pairs of aortic arches, but Petromyzon has only
7 pairs and Myxine has 6 pairs of aortic arches.
2. Aortic arches in fishes:
In Elasmobranchs:
¾ The primitive elasmobranches, Heptanchus has only 7 pairs of aortic arches, where as
Selachins has only 6 pairs of aortic arches.
¾ In most sharks, Scoliodon, have 5 pairs of functional aortic arches; the first pair is
reduced or disappears or replaced by the non-functional gills.
In Teleosts:
¾ In most teleosts or bony fishes, the first and second aortic arches are tend to disappear &
thus only third, fourth, fifth & sixth pairs of aortic arches remain functional.
In Polypterus & Lungfishes (Dipnoi):
a) Due to the mode of living & respiration from aquatic to terrestrial, the first aortic arches
disappeared & thus there are third, fourth & sixth aortic arches which are functional.
b) A set of pulmonary artery arises from the sixth aortic arches, near the dorsal aorta.
Notes:
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1. In Elasmobranchs & dipnoans, each aortic arch has one afferent artery & two efferent
arteries in each gill.
2. In teleosts, each arch has one afferent & one efferent artery in each gill.
3. In tetrapods, the arteries do not break up; as a result, the gills are situated below the
artery.
3. Aortic arches in amphibians:
Due to presence of lung as the main respiratory organ, the importance of gills is diminished.
In urodeles:
1. Four pairs of aortic arches (third to sixth) are functional in general.

2. The fifth pair is absent in Siren, Amphiuma.
3. The third pair forms the carotid artery & the fourth pair forms the systemic arches.
4. The radix or lateral aorta between third & fourth arches may persist as a vascular
connection called ductus caroticus.
5. The sixth pair forms the pulmonary arteries which supply blood to skin and lungs.
6. It retains connection with radix aorta, called ductus arteriosus.
In anurans:
1. At metamorphosis, with loss of gills, first, second & fifth aortic arches disappear
altogether.
2. Thus three pairs of aortic arches (third, fourth & sixth) are functional in general.
3. Carotid arch takes oxygenated blood to head region.
4. Systemic arch on each side continues to dorsal aorta to distribute blood elsewhere except
head & lung.
5. Pulmonary arch supplies venous blood exclusively to lungs for purification.
6. The ductus caroticus & ductus arteriosus are usually absent.
Notes:
• According to Kent & Miller (1997) the adult anurans have three pairs of aortic arches
(third, fourth & sixth) which are also retained by the amniotes or higher vertebrates.
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4. Aortic arches in reptiles:
Reptiles are fully terrestrial vertebrates in which gills disappear altogether and replaced by
lungs.
1. Only three functional arches (third, fourth & sixth) are present.
2. Right systemic arch (fourth) arises from left ventricle carrying oxygenated blood to the
carotid arch (third) to be sent into head.
3. Left systemic arch (fourth) leads from right ventricle carrying deoxygenated or mixed
blood to the body through dorsal aorta.
4. Pulmonary trunk (sixth) arises from right ventricle carrying deoxygenated blood to the
lungs for purification.
5. Generally ductus caroticus
& ductus arteriosus are
absent but in certain
snakes & lizards
(Uromasitx) the ductus
caroticus is present and in
some turtle & Sphenodon,
the ductus arteriosus is
present.
5. Aortic arches in birds &

mammals:
Birds & mammals are warm-

blooded because in both the
ventricle is completely divided so
that there is no mixing of
oxygenated & deoxygenated
bloods.
1. 6 arches develop in embryo, but only 3 arches (third, fourth & sixth) persist in the adult.
2. Single systemic aorta, right in birds & left in mammals, emerging from left ventricle and
carrying oxygenated blood.
3. Systemic aorta unites with the radix aorta to form dorsal aorta.
4. Subclavian artery present on the left side in birds & on the right side in mammals.
5. Third arch represents carotid arteries, which arise from systemic aorta.
6. Sixth arch arises from a single pulmonary trunk taking deoxygenated blood from right
ventricle to the lung.
7. Embryonic ductus caroticus & ductus arteriosus also disappear.
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Conclusion: Discussing the aortic arches in these vertebrate groups, it is clear that they are
originated from a common ancestral stock and their embryonic condition of aortic arches
supports the ‘recapitulation theory’ of Haeckel. (Kent & Miller, 1997)
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PORTAL VENOUS SYSTEM
In the circulatory system of animals, a portal venous system occurs when a capillary bed
pools into another capillary bed through veins, without first going through the heart. Both
capillary beds and the blood vessels that connect them are considered part of the portal venous
system.
They are relatively uncommon as the majority of capillary beds drain into veins which then drain
into the heart, not into another capillary bed. Portal venous systems are considered venous
because the blood vessels that join the two capillary beds are either veins or venules.
Portal circulatory systems differ from the typical circulatory route in that the blood passes
through two sets of smaller vessels before returning to the heart. Blood from the first set of
capillaries collects in portal vessels (sometimes called portal veins) which then begin to branch
again to supply a capillary network to a second location before entering a series of veins which
will lead to the heart.
HYPOPHYSCEAL PORTAL SYSTEM
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The hypophysial portal system is a system of blood vessels in the brain that connects the
hypothalamus with the anterior pituitary. Its main function is the transport and exchange of
hormones to allow a fast communication between both glands. The fenestrated structure of
capillaries in the hypophysial portal system facilitates a rapid exchange between the
hypothalamus and the pituitary, with only a small amount of hormones needed to stimulate an
accurate effect in the respective target organs in the body.
The pituitary gland is also supplied with a portal vessel. This small and short portal system
carries hormones directly from capillaries in the hypothalamic region of the brain to the target
tissues in the anterior pituitary.
RENAL PORTAL SYSTEM
A renal portal system which delivers portal blood to the kidneys is present in fish,
amphibians, and reptiles. Birds and crocodilians have a similar system, but the would-be portal
vessel forms a bypass around the second capillary bed, keeping the renal portal vein from acting
as a true portal system.
Origin:
The femoral and sciatic vein united together to form renal portal vein this vein open into kidney
and breaks into capillaries.
Destination: Kidney
Function: By the system filtration of metabolic

waste and excretion of these materials of the blood
by the kidney is done.
Significance: To separate the waste materials from

the blood before entering the heart while brings into
the kidney to filter and purify.
HEPATIC PORTAL SYSTEM
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Humans, and most other vertebrates, have a portal system which supplies the liver. The hepatic
portal system carries blood from the stomach, intestines, pancreas, and spleen through the
hepatic portal vein to the liver. In the liver the portal vessels begin to branch and supply blood to
the liver sinusoids, a form of liver capillary.
Origin: Different parts of the digestive system such as stomach , intestine, pancreas„ spleen etc
and wins come from hind limb are united together and enters into liver and breaks into capillaries
instead open to heart
Destination: Liver
Function: Absorbed food of alimentary canal and transport into liver. This food is differentiated
into the liver and sends them to the place of necessary and store the excess food.
Significance: The absorbed food is stored into liver not circulated meaning lastly into the body
and food is distributed when in need metabolized by the liver before reaching general circulation.
Blood flow to the liver is unique in that it receives both oxygenated and (partially)
deoxygenated blood. Blood passes from branches of the portal vein through cavities between
"plates" of hepatocytes called sinusoids. Blood also flows from branches of the hepatic artery
and mixes in the sinusoids to supply the hepatocytes with oxygen. This mixture percolates
through the sinusoids . A liver sinusoid is a type of sinusoidal blood vessel (with fenestrated,
discontinuous endothelium) that serves as a location for the oxygen-rich blood from the hepatic
artery and the nutrient-rich blood from the portal vein. Hepatocytes are separated from the
sinusoids by the space of Disse. Kupffer cells are located inside the sinusoids and can take up
and destroy foreign material such as bacteria. This blood then collects in a central vein which
drains into the hepatic vein. The hepatic vein subsequently drains into the inferior vena cava. The
hepatic artery provides 30 to 40% of the oxygen to the liver, while only accounting for 25% of
the total liver blood flow. The rest comes from the
partially deoxygenated blood from the portal vein.
Interestingly, the liver consumes about 20% of the
total body oxygen when at rest. That is why the total
liver blood flow is quite high, at about 1 liter a
minute and up to two liters a minute. That is on
average one-fourth of the average cardiac output at
rest.
COMPARATIVE ANATOMY OF
EXCRETORY SYSTEM
1. THE EXCRETORY SYSTEM
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All though the excretory system is a part of urinogeniatl system, the excretory
system has its own specialized structures and functions. Control of the salt and water
content and elimination of metabolic waste products are functions of the excretory
system. The end products of the metabolism include carbon dioxide, ammonia, urea, uric
acid, creatine, pigments and inorganic salts. The elimination of carbon dioxide is
accomplished by respiratory system. With certain exception, the other wastes are
eliminated by excretory or urinary organs. Usually these substances are in soluble in
water, in which case the final product, as eliminated by the body, is known as urine.
The Excretory System
Except for a few structure near the distal opening of some of the ducts, the entire
excretory system is divided from the embryonic mesomere.
Excretory Organs:
1. Organs which eliminate nitrogenous waste from blood are called excretory system. In
vertebrates urinary apparatus is used as excretory organ.
2. In vertebrates, skin, gills, lungs etc. eliminate CO2 from blood, but nitrogenous waste ate
eliminated by kidney.
3. Kidney is absent in Protochordata. For excretion different typed of organs are used in
Protochordata. Such some type of example are given bellow:
Balanoglossus have glomerulus.

Ascidia have pyloric glands, renal vesicle, partial vesicle, neural glands.
Amphioxus have solenocytes for excretion.
The main excretory system of vertebrates is kidney. It is different in structure in various

types in animals. It consists of three types of nephridia. A pair of kidneys is present in every
vertebrate, lying dorsal to coelom in trunk region, one or either side of dorsal aorta. They are all
built in accordance with a basic pattern.
Basic Structure of Vertebrate Kidney:
Nephron: Every vertebrate kidney has a basic structure. As for example, each kidney is
composed of a large number of units called viniferous tubbtes or neprons. Their number,
complexity and arrangement differ in different groups of vertebrates.
Origin of Uriniferous Tubules (Nephrons): Kidney tubules arise in the embryo in a linear
series from a special part of mesoderm called mesomere or nephrotome. It is the ribbon like
intermediate mesoderm running between segmental mesoderm (epimere) and lateral plate
(hypomere) or either side along the entire trunk from heart to cloacae.
Parts of Nephrons:
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A Uriniferous tubule is differentiated into three parts: (a) Peritoneal funnel, (b) Tubule
and (c) Malpighi an body.
(a) Peritoneal Funnel:

Near the free end of a uriniferous tubule is a funnel like ciliated structure called
peritoneal funnel. It opens into coelom (splanchnocoel) by a wide aperture, the coelomostome or
nephrostome. Nephrostome are usually confined to embryos and leave and considered vestiges
of a hypothetical primitive kidney.
(b) Malpighian Bodies: Bowman’s capsule and enclosed glomerulus together form . a renal
corpuscles or Malpighian bodies.
Bowman’s capsule: A tubule begins as a blind, cup like, hollow, doubled-walled Bowman’s
capsule.
Glomerulus: It encloses a tuft of blood capillaries called glomerulus.
It is supplied blood by a branch of renal artery, called afferent glomerular arteriole.

An efferent glomerular arteriole emerges out of glomerulus to join the capillary network
surrounding the tubule.
Encapsulated glomerulus are termed internal glomeruli which are common.
Those without a capsule and suspended freely coelomic cavity are called external
glomeruli embryos and larvae.
Capsule without glomeruli are termed aglomerular such as found in embryos, larvae and
some other fishes.
Nephric Tubules: At one end, the kidney tubule originates at a Bowman’s capsule and its walls
are continuous with the outer wall of the capsule. The tubule may be elongated and coiled. At the
other end it opens into a kidney duct called longitudinal duct.
Functions of Nephron:
1. Peritoneal funnel helps draining wastes from coelomic fluid

2. Malpighian bodies filter water, salts and other substances from blood.
3. During passage through Nephric tubules more substances are secreted into filtrate, while
some are reabsorbed.
4. Types of Kidneys:
Suring the evolution of the vertebrates, a series of kidney types replaced each other as the
functional excretory organs.
Archniphrose: It seems probable that the excretory organ of the primitive, ancestral vertebrate
was of a type that has been named archinephros.
.
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Structure of Archinephros: Kidney is thought to have considered of a series of archinepheric

tubules, one tubule on each side of each body segment. Each of these tubules opened into coelom
by a ciliated, funnel –shaped opening called a nephrostome. Glomeruli were suspended
segmentally in the coelomic cavity adjacent to the nephrostomes. Fluids eliminated by the
glomeruli were collected by the nephrostomes and passed down the archinephric tubules.
The tubules emptied into a pair of archinephros ducts, one on each side of the dorsal wall of the
body cavity. These ducts extended for the whole length of the body cavity.
Distribution of archinephros: No adults, living vertebrate possesses an archinephros, but it is

present in the larval form of the hagfish ^Myxine^ and the larvae of some of the caecilians
^Apoda^.
Pronephros: The most primitive type of kidney functional in an adult vertebrate is the
pronephros.
Structure of Pronephros:
The pronephros is believed to represent the most anterior part of the ancestral archinephros.
Tubules and Malpighian Bodies:

In the embryos of all vertebrates, the first kidney tubules arise from the anterior end of the
mesomere. In a few primitive forms of glomerulus which are delivered from segmental branches
of the dorsal aorta, project into the coelom near the nephrostomes. Such structures are known as
external glomerulus. Usually, however, the glomerulus is internal that is, it is almost completely
surrounded by the bowman’s capsule which projects from the tubule near its proximal end. The
nephrostomes persist and still retains its opening to the coelom. Whether the glomerulus are
external or internal seems to have no particular significance.
Because of the segmental nature of the pronephros and the relatively small number off
tubes and related structure, the diffuse pronephric kidney does not form a distinct organ
identifiable to naked eye. For example, there are thirteen tubules and in the lamprey, seven in the
3mm, human embryo and four in the shark embryo.
Ducts of Pronephrose:
The pronephric tubules fuse to form the pronephric ^originally archinephric^ ducts which
grows backward and opens posteriorly into the cloaca. While the pronephros may be important
to those organisms in which it is functional, its chief evolutionary significance seems to be the
part it plays in forming the pronepheric duct, which persist after the pronephros itself has
disapeard.
Distribution of Pronephrose: The pronephroic tubules continue to function in the adult hagfish
and in occasional teleosts. The pronephros is also a functional structure in many immature fishes
and in the larvae of some of the amphibians and appears transitorily in the embryos of all the
higher vertebrates.
Mesonephros and Opisthonephros:
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The mesonephros is the kidney tissue that develops posterior to the pronephros. Posterior
to the mesonephros, the meta nephros (that portion of the kidney tissue that gives rise to the adult
amniote kidney) develops. The adult kidney of the anamniotes involves not only the mesonephric
tissue but some of the metanephric tissue as well. Thus, while the kidney of the anamniotes is
frequently called a mesonephros, it is not completely homologous with the mesonephros of the
amniote embryo, and for this reason is preferably referred to as the opisthonephros.
Structure of Mesonephros and opisthonephrose:

These kidney structures form discrete organs that are readily apparent as kidneys. They
resemble each other closely in structure and function.
Tubules and Malpighian Bodies:

While the first few mesonephric tubules tend to be segmentally distributed, proliferation
of many such tubules in each of the body segments eventually obliterates all evidence of
metamerism. Thus, for the first time in evolution, the primary excretory organ becomes a
discrete, bulky structure, lying against the dorsal body wall and tending to bulge into the coelom.
The embryonic tubules are first formed as solid structures, but they soon develop lumina
and begin to elongate. One end off the tubule enlarges and is invaginated by the ball of
capillaries forming the glomerulus and thus becomes the double-walled Bowman’s capsule
described above. This capsule, with its included glomerulus, forms the renal or Malpighian
bodies. All glomerulus in the opisthonephros are surrounded by Bowman’s capsules and hence
are internal. In a few forms, peritoneal funnels (nephrostomes) may connect the tubules with the
coelom, but this seems to be the exception. Most species that have been studied have lost the
peritoneal funnels and the coelom is no longer continuous with the outside world through the
excretory system.
At first, as the mesonephric tubule develops and elongates it becomes ‘S’ shaped, and
then the proximal and distal loops of the S fold further to form typical proximal and distal
convolutions so characteristic of the mesonephric tubule.
Mesonephric duct:
The distal ends of the tubules establish connections with the old pronephros duct, which is
now known as the mesonephric duct ( also called Wolffian duct). The distal ends of several
tubules may unite before they join the mesonephric duct, and sometimes the tube formed from
the components of several tubules may establish an opening directly into the cloaca rather than
into the mesonephric duct. The mesonephric duct may also transport sperm to the outside.
Distribution of Mesonephros and Opisthonephros: Mesonephric part functions in the embryos

of reptiles, birds and mammals. The opisthonephros is the functional kidney of the adult
lamprey, the cartilaginous fishes, the bony fishes and the amphibians.
Metanephros: The metanephric kidney develops from the most posterior portion of the
mesomere and is the most compact of any of the vertebrate renal structures.
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Structure of metanephros: The body of the metanephros has a twofold origin. Part of it
develops form the posterior end of the opisthonephros, while another part forms as a new and
unique metanephric structure.
Malpighian Bodies and Nephric Tubules:
The metanephric kidney is basically made up of the same structure as the mesonephros-
the Malpighian bodies and they are associated tubules. No nephrostomes (peritoneal funnels) are
ever present in the metanephros. The metanephric tubules are somewhat more complex in the
mammals and (to a lasser extent) in the birds then are the mesonephric tubules. In these animals,
each tubule develops a long loop called the loop of Hanley with ascending and descending
portions between the proximal and distal convoluted segments. These loops functions primarily
in the re-absorption of water.
Collecting Tubules and Metanephric Duct:
In the amniote embryo, a diverticulum develops near the posterior end of the
mesonephric or Wolffian duct and grows outward and forward to make contact with the
developing metanephros. This diverticulum branches and rebranches within the metanephros to
form a large number of very fine collecting tubules. This collecting tubules establish connections
with the S- shaped renal tubules, and henceforth the waste products pass through them to the new
duct formed from the diverticulum. This duct is called a ureter. When this new pathway develops
and the embryonic mesonephros degenerates, the Wolffian duct no longer transports urine from
the kidney but remains as a reproductive duct transporting sperm from the testis.
Distribution of the Metanephros: The metanephros becomes functional in most reptile, bird
and mammal embryos and is the functional kidney of all adult amniotes.
The Excretory or Urinary System of Human
The human urinary system consists of the organs- Kidneys, Ureter, Urinary bladder and Urinary
duct (Urethra).
Kidney: The human kidneys are bean-shaped, reddish brown organs each about the size of a fist.
They are located on either side of the vertebral column where they are partially protected by the
lower rib-case.
Ureter: A ureter is a duct that carries urine from the kidneys to the urinary bladder. Each kidney
is connected to a ureter.
Urinary Bladder: Urinary Bladder, where urine is stored until it is voided from the body.
Urethra: Urethra is a single urinary duct by which excretory substances like urine is voided
from the body. In males, the urethra passes through the penis and in females, it opens ventral to
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the opening of the vagina.
The Urinary System of Human
Urine: Urine is a excretory substance made by kidneys is carried from the body by the other
organs in the urinary systems.
There is no connection between the Genital (Reproductive) and the Urinary systems in females
but there is a connection in males. In males, the urethra also carries sperm during ejaculation.
EXCRETORY SYSTEM OF LIZARD AND COMPARE WITH THAT PIGEON AND

RABBIT.
Lizard, bird and rabbit all these three animals come under the group amniota. The taking
away of nitrogenous un wanted waste products from the body is called excretion. If excretion
takes place not properly in the body they become poisonous. vertebrates main excretory organs
are called as kidneys. skin-gills-lungs-liver-intestine are also acts as accessory excretory organs.
The kidneys are made up with numerous Uriniferous tubules. Kidneys are located in dorsal side
of coelom.
A typical Uriniferous tubule having three parts-

1-ciliated peritoneal funnel
2-malpighian body
3-ciliated convoluted tube.
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EXCRETORY SYSTEM - EXCRETORY SYSTEM – EXCRETORY SYSTEM-

GARDEN LIZARD PIGEON RABBIT
1. Paired kidneys are dark red 1. Kidneys are dark red and
1. Kidneys are dark red and
and irregular in shape. These somewhat rectangular and flattened
bean shaped organs.
are flattened organs. organs.
2. Kidneys are located in the
posterior region of the 2. Kidneys are located in the
2. Kidneys are situated in the
abdominal cavity and attached posterior part of the
anterior part of the abdomen.
to the dorsal wall by a fold of abdominal cavity.
peritoneum.
3. The two kidneys are
3. Right and left kidneys are distinct. The right kidney
3. Same as in calottes
opposite to each other. lies much ahead than the left
kidney.
4. They are attached to the 4. They are fitted in the hollows of
4. Same as in calottes.
dorsal muscles. the pelvic girdle.
5. They are very near to the 5. They are a little away from the 5. They are well away from
median line kidneys are median line. Kidneys are Meta the median line. Kidneys are
Metanephros type. nephros type. meta nephros type.
6. Each kidney is a single
6. Each kidney has two lobes
5. Each kidney has three lobes They lobed structure. Inner side of
Anterior lobe is broad and
are anterior, median and posterior the kidney has a concave
posterior lobe is broad Hilus is
lobes. Hilus is absent. depression is known as the
absent.
'hilus'.
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7. The two kidneys are united

7. The two kidneys are separate and 7. The two kidneys are
posteriorly forming a V -
do not fuse with each other. distinct.
shaped structure.
8. The two ureters are narrow,
8. The ureters open into the
thin-walled ducts extending
urinary bladder. Ureters arise
behind from the kidneys to the 8. Same as in Calotes.
from the hilus of each
cloaca, where these open into
kidney.
the urodaeum.
9. Each ureter is expanded in
9. There is no pelvis. 9. There is no pelvis. its kidney into a funnel like
pelvis.
10. In males the ureters join at
its posterior end with its 10 The ureters do not join with the
10. Ureters open separately
corresponding vas deferens vas deferens and both open
into the urinary bladder.
and both open by a common separately into the cloaca.
urino-genital aperture.
11. A thin walled urinary 11. Urinary bladder is a
bladder opens on the ventral 11. Urinary bladder is absent large, median, pear, shaped,
side of cloaca thin walled transparent sac.
12. Urinary bladder 12.Urinary bladder opens
communicates with urodaeum 12. _ into the urethra or
thrumph its ventral wall. unnogenital canal.
13. Calotes is uricotelic animal
13. Urine consists mainly of uric 13. Urine consists mainly of
Urine consists n.ainly of uric
acid cotelic animal ureaureotelic animal
acid.
14. Urine is excreted in a semi 14. Urine is excreted ir a semisolid 14. Urine is passed out in a
solid state. state (Bird droppings). fluid state.
Schematic presentation of three stages of kidney development: pronephros, mesonephros,

and metanephros.
Pronephros is the earliest nephric stage in humans, and constitutes the mature kidney in most
primitive vertebrates. It extends from the 4th to the 14th somites and consists of 6-10 pairs of
tubules. These spill into a pair of primary ducts that are formed at the same level, extend
caudally, and eventually reach and spill into the cloaca. The pronephros is a vestigial structure
that disappears completely by the 4th week of human embryonic life.
Mesonephros develops by the formation of mesonephric tubules from the intermediate

mesoderm, it is the principal excretory organ during early embryonic life (4—8 weeks). It
gradually degenerates, although parts of its duct system become associated with the male
reproductive organs.
Metanephros arises caudal to the mesonephros at five weeks of development; it is the

permanent and functional kidney in higher vertebrates. It is derived from the intermediate
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mesodermm. The ureteeric bud arisees as a diverrticulum from

m the Wollfiian duct, close to the
entrance to the cloacaa and grows towards andd inside the metanephric
m c mesenchym me.
REESPIRATO ORY SYSTE EM

Respiraation is the process
p of obbtaining oxyggen from thee external
environm
ment & elimiinating Carbondiaoxide.
1. Externnal respiratioon - oxygen and carbon dioxide
exchaanged betweeen the externnal environm ment &
the bo
ody cells
2. Intern
nal respiratioon - cells usee oxygen for ATP
produuction (& prooduce carbonn dioxide in the
processs)
Adaptations for external respirration:

1 - Primaryy organs in adult
a vertebrrates are exteernal &
innternal gills, swim bladdders or lungs, skin, & thee buccopharyyngeal mucoosa
2 - Less coommon respiiratory devicces include filamentous
fi o
outgrowths o the posterrior
of
trrunk & thigh h (African haairy frog), linning of the cloaca,
c & linning of esophhagus
Respirattory organs:
Cutaneoous respiratiion
a. respiration
n through thee skin can taake place in air,
a water, orr both
b. most impo ortant amongg amphibianss (especiallyy the family Plethodontid
P dae)
Gills
62
Dr.C..V.Narasimha Murthy
M , M.Scc. Zoology- preevious – chordaata Notes 20166.
63
Cartilaginous fishes:
1. 5 ‘naked’ gill slits
2. Anterior & posterior walls of the 1st 4 gill chambers have a gill surface (demibranch).
Posterior wall of last (5th) chamber has no demibranch.
3. Interbranchial septum lies between 2 demibranchs of a gill arch
4. Gill rakers protrude from gill cartilage & ‘guard’ entrance into gill chamber
5. 2 demibranchs + septum & associated cartilage, blood vessels, muscles, & nerves =
holobranch
Bony fishes (teleosts):

1. usually have 5 gill slits
2. operculum projects backward over gill chambers
3. interbranchial septa are very short or absent
Agnathans:
1. 6 - 15 pairs of gill pouches
2. pouches connected to pharynx by afferent branchial (or gill)
ducts & to exterior by efferent branchial (or gill) ducts
Larval gills:
1. External gills
a. outgrowths from the external surface of 1 or more gill
arches
b. found in lungfish & amphibians
2. Filamentous extensions of internal gills
a. project through gill slits
b. occur in early stages of development of elasmobranchs
3. Internal gills - hidden behind larval operculum of late anuran tadpoles
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Swim bladder & origin of lungs - most vertebrates develop

an outpocketing of pharynx or esophagus that becomes one
or a pair of sacs (swim bladders or lungs) filled with gases
derived directly or indirectly from the atmosphere.
Similarities between swim bladders & lungs indicate they are
the same organs.
Vertebrates without swim bladders or lungs include
cyclostomes, cartilaginous fish, and a few teleosts (e.g.,
flounders and other bottom-dwellers).
Swim bladders:
1. May be paired or unpaired (see diagram above)
2. have, during development, a pneumatic duct that usually connects to the esophagus. The
duct remains open (physostomous) in bowfins and lungfish, but closes off (physoclistous)
in most teleosts.
3. Serve primarily as a hydrostatic organ (regulating a fish's specific gravity)
4. Gain gas by way of a 'red body' (or red gland); gas is resorbed via the oval body on
posterior part of bladder may also play important roles in:
a. Hearing - some freshwater teleosts (e.g., catfish, goldfish, & carp) 'hear' by way
of pressure waves transmitted via the swim bladder and small bones called
weberian ossicles (see diagram below)
b. sound production - muscles attached to
the swim bladder contract to move air
between 'sub-chambers' of the bladder.
The resulting vibration creates sound in
fish such as croakers, grunters, &
midshipman fish.
c. Respiration - the swim bladder of
lungfish has number subdivisions or septa (to increase surface area) & oxygen and
carbon dioxide is exchanged between the bladder & the blood
Lungs & associated structures
1. Larynx
a. Tetrapods besides mammals - 2 pair of cartilages: artytenoid & cricoid
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65
b. Mammals
- paired
arytenoids
+ cricoid
+ thyroid
+ several
other
small
cartilages
including
the
epiglottis (closes glottis when swallowing)
c. Amphibians, some lizards, & most mammals - also have vocal cords stretched
across the larynx.
2. Trachea & syrinx
a. Trachea
i. usually about as long as a vertebrates neck (except
in a few birds such as cranes)
ii. reinforced by cartilaginous rings (or c-rings)
iii. splits into 2 primary bronchi &, in birds only,
forms the syrinx at that point
3. Lungs
a. Amphibian lungs
i. 2 simple sacs
ii. internal lining may be smooth or have simple sacculations or pockets
iii. air exchanged via positive-pressure ventilation
b. Reptilian lungs
i. simple sacs in Sphenodon & snakes
ii. Lizards, crocodilians, & turtles lining is septets, with lots of chambers &
sub chambers
iii. air exchanged via positive-pressure ventilation
4.
a. Avian lungs - modified from those of reptiles:
i. air sacs (diverticula of lungs) extensively distributed throughout most of
the body
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ii. arrangement of air ducts in lungs ----> no passageway is a dead-end

iii. air flow through lungs (parabronchi) is unidirectional
5.
a. Mammalian lungs:
i. multi chambered & usually divided into lobes
ii. air flow is bidirectional:
6. Trachea <---> primary bronchi <---> secondary bronchi <---> tertiary bronchi <--->
bronchioles <---> alveoli
7 . Air exchanged via negative pressure ventilation, with pressures changing due to
contraction & relaxation of diaphragm & intercostals muscles
GILLS
Gills in Protochordates
A large and sieve-like pharynx in majority of

these animals performs dual function of respiration
and trapping food particles which are brought in
through the current of water. The primitive
pterobranch hemichordates (Cephalodiscus and
Rhabdopleura) have either no gill slits or have very
few and sport tentaculated arms, which other than
food gathering, also function as efficient respiratory
organs. Balanoglossus possesses a large pharynx
having as many as 700 pairs of gill slits, which
appears to be a necessity in the burrowing habitat of the animal.
The free-living urochordates, such as Salpa and Doliolum do not possess many stigmata
or gill slits as their entire body is permeable to oxygen but in the sedentary ascidians pharynx is
prominently enlarged and perforated with no less than 200,000 stigmata for filter-feeding.
Cephalochordates use pharynx for both filter-feeding and respiration and hence carry
150-200 pairs of gill slits.
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Respiratory organs of Cyclostomes
Agnathans have 6-15 pairs of gill pouches, which are lateral extensions of pharynx and
contain gill lamellae within. Cyclostomes are called marsipobranchs, which means “pouched
gills”, since the gill lamellae are housed in gill pouches. The hagfish, Myxine has only 6 pairs of
gill pouches whose ducts join together and open to the exterior by a single pair of openings,
while Bdellostoma carries 6-15 pairs of gill pouches that vary in different species and open to the
outside independently. In Myxine behind the gill pouches there is a single pharyngocutaneous
duct on the left side, which is a modified gill pouch which drains excess water that fails to enter
the gill pouches. The lamprey, Petromyzon, has 8 embryonic and 7 adult paired gill pouches that
open to the exterior by independent openings.
Respiratory organs in Elasmobranchs
Most Elasmobranchs possess 5 pairs of gill slits and a pair of spiracles. There is no
operculum covering the gill slits in cartilaginous fishes. A demibranch is a bunch of gill
lamellae attached on one side of the interbranchial septum. Hence, there are altogether 9 pairs of
demibranchs in Elasmobranchs. Between the two demibranchs lies the interbranchial septum,
which is supported by gill cartilages. Anterior to the first gill slit is a spiracle or pseudobranch. In
free swimming sharks and dogfishes water generally enters through the mouth.
Blood to the gills is supplied by five pairs of afferent branchial arteries coming from
ventral aorta and hence they bring deoxygenated blood from heart. Blood is then oxygenated in
gills and is collected by the loops of four pairs of efferent branchial arteries and carried to the
paired dorsal aorta, the two sides of which meet posteriorly to form single median dorsal aorta
that supplies oxygen-rich blood to the whole body.
Gills of bony fishes
In bony fishes gills are covered with an operculum that is made of flattened skeletal
plates and there is no spiracle as in Elasmobranchs. There are 4 pairs of gill pouches, each
containing two demibranchs, making the total number of demibranchs in bony fishes as 8 pairs
or four pairs of complete gills or holobranchs. Teleosts always breathe with their mouth open and
eject expiratory water by opening operculum. Gills in Chondrostei, Holostei and the lungfish
Neoceratodus exhibit partial reduction in their interbranchial septa, which happens to be
somewhat intermediate condition between Elasmobranchs and teleosts.
EXTERNAL GILLS
External gills develop from the outer wall of pharynx or from the exposed portion of
branchial arch. They occur in larval lampreys, few larval fishes, Polypterus, lungfishes, some
larval teleosts and all larvae and some adults of amphibians. There is a single pair of larval gill in
the chondrosteian bony fish, Polypterus, which has a long axis carrying gill lamellae. The
African and South American lung fishes possess 4 pairs of feathery external gills. The larval
forms of some amphibians and some adult urodeles possess external gills which arise simply as
folds of skin on the surface of the III, IV and V branchial arches but weakly supported by the
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skeletal system. Perennibranch amphibians as Necturus and Proteus retain external gills
throughout life along with 2 or 3 pairs of gill slits, which are functionless as the water does not
pass through pharynx. Instead, gills are waved in water by means of muscles attached at the base
of gill axis for respiration. The larvae of limbless amphibian, Caecilia, have a pair of
exceptionally large leaf-like gills with profuse blood supply. Salamanders that inhabit hill
streams, e.g. Eurycea and Salamandrina, which belong to family Plethodontidae have neither
gills nor lungs for respiration and survive only on cutaneous respiration.
AIR BLADDER
Barring agnathans, cartilaginous fishes and few bottom dwelling teleosts, all fishes carry
a gas-filled air bladder on the dorsal side of the gut, which serves as hydrostatic organ. On the
ventral side of the bladder there occurs a highly vascularised area called red gland that is
supplied by intestinal artery and portal vein and which has unique capability of extracting free
oxygen from the blood and release it into the air bladder in order to make it inflate. Small pouch-
like diverticula called oval that can be closed or opened by sphincter muscles is the site of
reabsorption of gases. Secretion and absorption of gases in swim bladder occurs under the
control of autonomic nervous system, based on the depth at which a fish is swimming.
In Cypriniformes (Teleosti), a series of four small bones (tripus, intercalarium,
scaphium and claustrum), derived from the first three vertebrae and called Weberian Ossicles,
connect the anterior end of air bladder with the sinus impar of membranous labyrinth. Sound
vibrations received by air bladder from the surrounding water are conveyed to the internal ear
through this unique apparatus to bestow some hearing ability to these fishes.
In some fishes as for example ganoids, carps and catfishes, a pneumatic duct connects
the air bladder with oesophagus. Such condition is called physostomous (Gr. physo=bag;
stoma=opening). Fishes which do not have such a pneumatic duct connecting the air bladder are
called physoclistous (Gr. physo=bag; clista=closed).
The comparative study of air bladders in different groups of fishes and striking similarity
between the swim bladder and lung suggest a phylogenetic relationship between the two. The
conventional belief is that lungs evolved from the air bladder of fishes. However, recent
evidences point to the contrary that lungs evolved first in fishes for supplementing oxygen from
air and then they got transformed into swim bladder as the oxygen concentration in water
increased.
Accessory Respiratory Organs in Fishes
Many species of fishes developed breathing organs other than gills for supplementing
deficiency of oxygen in water. These are as follows:
Dendritic Organs
They are also called arborescent organs as they are highly vascularised tree-like,
branched structures produced by the second and fourth gill arches and located in the
suprabranchial chamber, posterior to the gills. Paired gill fans at the opening of branchial
chamber force air over the dendritic organs as the fishes gulp air. Dendritic organs are found in
catfishes such as Clarias.
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Labyrinthine Organs
These are rosette-like concentric plates of tissue present in the suprabranchial chamber
of climbing perch (Anabas), Trichogaster, Osphromanus and Polycanthus. Respiration takes
place when these fishes gulp air. Perches can migrate from one pond to another by breathing air
through labyrinthine organs and using pectoral fin spines to walk on land.
Pneumatic Sac
It is a tube like extrabranchial diverticulum that extends up to tail in some cat fishes such
as Heteropneustes, which can survive out of water for considerable time using these organs for
air breathing.
Air Chamber
Air chamber is a small, highly vascularised sac located behind the gills of some fishes,
e.g. Ophiocephalus, Macropodus and cuchia eel (Amphipnous). These fishes can gulp air and use
it as air breathing organ.
Buccopharyngeal epithelium
Mud skippers (Periophthalmus, Balaeophthalmus) possess vascularised buccopharyngeal

epithelium and also a respiratory tail. They skip around in swamy areas, breathing air by
buccopharyngeal epithelium or keep their tail in water for aquatic respiration.
Integument
Eels (Anguilla) breathe through skin while migrating from the American and European
rivers to Sargasso Sea in Bermuda. As much as 60% exchange of gases takes place through the
highly vascularised skin.
Gut epithelium
Fishes such as Callichthys, Hypostomus, Doras, Misgurnus, Cobitis can suck and release
water through anus and exchange of gases can take place in the rectal lining. In giant loach
(Cobitis) and Misgurus lining of stomach and intestine is used as respiratory organ.
LUNGS
Lungs of Polypterus and the ganoid fish Calamoichthys are asymmetrical and connected
by pneumatic duct on the ventral side of pharynx. The blood is supplied to lung by pulmonary
artery that emerges off the 6th aortic arch, but unlike in lungfishes venous blood returns to
hepatic vein.
Lungs of Dipnoi (Choanichthys) are bilobed or paired as in Protopterus (African lung
fish) and Lepidosiren (South American Lung fish) and are connected to oesophagus via a
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pneumatic duct. But the Australian lung fish (Neoceraodus) has a single lung that is used as
hydrostatic organ.
In tetrapods, embryonic lungs arise from pharyngeal wall as a hollow mid-ventral
evagination that subsequently bifurcates to form two lungs that carry an envelope of peritoneum.
AMPHIBIA
Lungs of amphibians are two simple sacs, narrow and elongated in urodeles and bulbous
in anurans enclosed in a single peritoneal membrane and supplied by pulmonary arteries and
drained by pulmonary veins. Left lung in limbless amphibians is rudimentary. Lungs are
vestigial in salamanders inhabiting hill streams where in fast flowing water buoyancy would not
be a desirable trait.
Amphibians lack ribs and hence use floor of the buccal cavity to force air in and out of
the lungs. Frogs and toads modify 2nd, 3rd and 4th visceral arches to produce a plate-like
hyobranchial apparatus that lies in the floor of oral cavity and is connected to squamosal bone of
skull by petrohyal muscle and to sternum by sternohyal muscle. One breathing cycle is
completed in four steps in anurans that is affected by contraction of these two muscles.
Breathing in frog requires much faster movement of hyoid plate as compared to lungs and
considerable amount of gas exchange takes place in bucco-pharyngeal region too. Cutaneous
respiration also contributes to major part of oxygen supply to the body of amphibians.
REPTILES
Lungs are narrow and elongated in snakes and lizards extending up to two-third of the
body cavity but are more bulbous in turtles and crocodiles. The left lung is rudimentary in
limbless lizards and snakes. There are well formed alveoli in lungs which are housed securely in
a pair of pleuro-peritoneal cavities. Breathing in snakes and lizards is carried out by a
combination of hyoid plate, nostril valves and ribs or only by the movement of rib cage, while
tortoises and turtles make use of muscles surrounding peritoneal membranes. Crocodiles are the
only living reptiles that possess a muscular diaphragm for breathing as do the mammals.
BIRDS
Lungs are secured into pleural cavities and extend into membranous air sacs that occupy
all available space in the body cavity and also penetrate into bone marrow cavities. This makes
the bones pneumatic in birds and help to reduce body weight which is so necessary in flight.
Majority of birds have 5 pairs of air sacs, namely, cervical at the base of neck; interclavicular
often united across midline; anterior thoracic placed lateral to the heart; posterior thoracic
within the oblique septum and abdominal within the abdominal cavity. Sometimes there are also
axillary air sacs near the pectoral muscles. The flight muscles inflate and deflate the air sacs like
bellows with each stroke of wings.
Air duct system is unique in birds as there are no alveoli as found in reptiles and
mammals. Trachea divides into two bronchi which enter the lungs and branch into mesobronchi
that again divide to form parabronchi. From each parabronchus, bunches of air capillaries arise
which loop back into their own lumen to form anastomosis that eventually leads into the air sacs.
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Air capillaries are minute and only one cellular layer thick and contain respiratory epithelium
and rich network of blood capillaries.
During inspiration and expiration, air passses through the air capillary anastomosis into
the air sacs and back twice making it a double respiration.
MAMMALS
Mammalian lungs increase efficiency by increasing the surface area of respiratory

epithelium of alveoli whose number goes up to millions and lungs become almost like semisolid
sponge with little empty lumen inside. Lungs are enclosed in double peritoneal membranes, the
outer parietal pleura and inner visceral pleura that enclose the fluid-filled pleuro-peritoneal
cavity in between.
Trachea which is commonly known as windpipe opens in pharynx by a slit-like glottis
and posteriorly divides into two primary bronchi that enter lungs and branch off to secondary and
tertiary bronchi which ultimately lead to fine capillaries called bronchioles. Each bronchiole is
connected to several alveoli by alveolar ducts. The number of alveoli in human lungs is
estimated to be about 750 million which collectively carry an enormous surface area of about
100 m2 and if stretched.
A muscular diaphragm located between the thoracic and abdominal cavities moves in the
antero-posterior direction and forces air in and out of the lungs. External costal muscles and
internal costal muscles are attached between the ribs and sternum, the former increases the
thoracic space while the latter decreases it to carry out what is known as thoracic respiration or
common panting after strenuous physical exercise.
Whales possess enormous nasal chambers in the head that can store large quantity of air
when diving deep in the sea. Passage of air from nasal chambers to lungs is controlled by a pair
of valves.
SOUND PRODUCING ORGANS

Sound producing organs, larynx and syrinx, are associated
with trachea through which air can be forced from lungs into the sound
box to produce sound. Larynx in urodeles is so simple that it has only a
pair of lateral cartilages, called Guardian cartilages that surround the
glottis and this apparatus is incapable of producing any sound in these
animals.
Larynx of frog is made of a cricoid cartilage which is a
modification of the first tracheal ring and a pair of arytenoid cartilages,
which support a pair of vocal cord that vibrates to produce sound.
Males of frogs and toads in addition possess a pair of vocal sacs which
are evagination of oral cavity and serve as resonance chambers to
amplify sound.
Reptiles are silent animals but possess larynx, albeit without a vocal cord, in the absence
of which they can at best produce a hissing sound.
Birds inherited a rudimentary larynx from their reptilian ancestors and hence evolved a
secondary sound producing organ called syrinx located at the junction of trachea and bronchi
and hence called bronchotracheal type. The tympanic chamber has a bony ridge at base called
pessulus that supports the unpaired membrana semilunaris, which vibrates to produce sound.
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In mammals larynx consists of paired arytenoid cartilages that support vocal cord. Base
of the sound box is made of a ring-like cricoid cartilage and the unpaired thyroid cartilage forms
the surrounding walls of the tympanic chamber. A pair of fleshy vocal cords is stretched across
the cephalic part of the vocal chamber supported by the paired cartilages of Santorini. The cord
vibrates to produce sound that is modulated in the oral cavity.
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COMPARATIVE ANATOMY OF BRAIN AND SPINAL CORD
Brain
The brain is safely kept inside the cranial vault. Inside the skull the brain is surrounded by three
protective coverings. They may be grouped under two divisions.
1. Pachymenix -It includes the duramater.
2. Leptomeninges - In includes the arachnoid mater and pia mater.
The duramater is the outermost membrane. It is thick and inelastic in nature. The arachnoid
mater is the middle covering over the brain. In between arachnoid and piamater there is a space
called the subarachnoid space. It contains cerebro-spinal fluid and blood vessels. The piamater is
a delicate membrane closely applied
to the brain. This membrane
contains blood capillaries supplying
blood to the brain cells.
The human brain weighs about
1.3 Kg. It contains more than a
billion neurons. Based on
embryological development the
brain can be divided as follows.
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Prosencephalon (Fore brain) –

It consists of the cerebrum and the diencephalon. The cerebrum is the largest part of the
brain. It is divided into right and left hemispheres by a longitudinal fissure. However, at the base
the two hemispheres are connected by a sheet of nerve fibres called the corpus callossum.the
outer surface of the cerebrum is called the cortex or grey mater. It is 2 to 4 mm thick. The inner
content of the cerebrum is the white mater. The surface of the cerebrum has several folds called
the gyri. They greatly increase the surface area of the cortex. The shallow grooves in between the
gyri are called the sulci. A central sulcus runs in the lateral surface of the cerebrum from superior
to inferior region.
Each cerebral hemisphere is divided into four lobes. They are the frontal at the front, the
parietal towards the top of the head, the temporal on the side and the occipital at the rear. The
diencephalon contains the thalamus and hypothalamus. This region is found between the
cerebrum and the brain stem.
The thalamus has a cluster of nuclei which act as the relays for particular sensory
pathways. Just beneath the thalamus, the hypothalamus is present. It contains reflex centres
linked to the autonomic system. A funnel shaped stalk called the infundibulum extends from its
floor. It is connected to the neurohypophysis of the pituitary gland.
Mesencephalon (mid brain) –
It is the smallest region of the brainstem. On its dorsal surface there are four rounded bodies
called the carpora quadrigemina.
Rhombencephalon (hind brain) –
The three main regions of the rhombencephalon are the medulla oblongata, the pons varoli and
the cerebellum.
1. The cerebellum consists of two hemispheres. Its surface has many ridges called folia. The
cerebellum consists of three parts. They are the small anterior flocconodular lobe, a
narrow central vermis and two large lateral hemispheres.
2. The pons is just superior to the medulla oblongata. It contains ascending and descending
nerve tracts.
3. The medulla oblongata is about 3 cm long. It is continuous with the spinal cord. It
remains as a bridge between the brain and the spinal cord.
4. Brain stem - The medulla oblongata, pons and mid brain form the brain stem. It connects
the spinal cord to the brain. Ten of the twelve cranial nerves enter or exit the brain
through the brain stem.
Spinal cord - The spinal cord extends from the foramen magnum to the level of the second
lumbar vertebra. It is considerably shorter than the vertebral column. There are two enlargements
in the spinal cord. They are the cervical and lumbar enlargements. Below the lumbar
enlargement the spinal cord tapers to form a cone like region called the conus medullaris. A
connective tissue filament called the filum terminale extends inferiorly from conus medullaris to
the coccyx. The conus medullaris and the nerves extending below resemble a horse’s tail. Hence
it is called cauda equina.
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A cross section of the spinal cord reveals a central grey portion and a peripheral white
portion. The white matter consists of nerve tracts and the grey matter consists of neuron cell
bodies and dendrites. The dorsal and ventral sides have long fissures. There are 31 pairs of spinal
nerves arising from the spinal cord. Each nerve has a dorsal root and a ventral root from the
spinal cord. The dorsal roots have dorsal root ganglia.
Ventricles
The entire CNS remains as a hollow tube. The tube inside the adult brain forms
ventricles. Each cerebral hemisphere contains a large cavity called the lateral ventricle. It
corresponds to the hypothetical first and second ventricles. The two lateral ventricles
communicate with the third ventricle located in the centre of the diencephalon. This connection
is made through two interventricular foramina (foramen of Monro). The third ventricle inturn
opens into the fourth ventricle found inside the medulla oblongata. This communication happens
through a narrow canal called the cerebral aqueduct (aqueduct of sylvius). The fourth ventricle is
continuous with the central canal of the spinal cord. The central canal extends nearly to the full
length of the cord.
Cerebro-spinal fluid (CSF)
This fluid fills the ventricles of the brain and the central canal of the spinal cord. About
80-90 % of CSF is produced by specialized cells called ependymal cells within the lateral
ventricles. Remaining 10-12 % is produced by similar cells in the 3rd and 4th ventricles. These
ependymal cells, their supportive tissue and the associated blood vessels together are called
choroid plexuses. The plexuses are formed by invagination of the vascular piamater into the
ventricles.
Brain is the anterior end of spinal cord that has enlarged to take care of the sense organs
which are located on the anterior end of the body in a bilaterally symmetrical animal that moves
ahead in an anterior-posterior axis. This is called cephalon, which as it evolves further, divides
into three parts, namely, prosencephalon, mesencephalon and rhombencephalon. As the brain
develops further by increasing the number of neurons, it further divides into different parts, each
one having assigned its own specific function.
Prosencephalon divides into Telencephalon and Diencephalon, the former includes
olfactory lobes (Rhinencephalon) and Cerebral hemispheres that coordinate the activities of the
entire brain. The roof of cerebrum is called Pallium and the floor that generally contains nerve
fibres is known as corpus striatum.
Diencephalon is a small part of brain, generally covered by enormously enlarged
cerebral hemispheres. This is an extremely important part of brain which functions as switch
board to cerebrum. Dorsal part of diencephalon is called epithalamus and the ventral part
hypothalamus while the lateral parts are called thalami that contain relay centres to connect
dorsal and ventral parts of thalamus.
Anterior part of epithalamus contains a glandular area called anterior choroid plexus
(Tela Choroidea) which secretes cerebro-spinal fluid. Two dorsal processes of epithalamus, the
anterior paraphysis supports parietal body and the posterior epiphysis bears pineal body. These
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two bodies function as photoreceptors in lower vertebrates and gradually transform into
endocrine organs and biological clock in higher vertebrates.
The ventral hypothalamus has the optic chiasma (crossing of optic nerves) on the
anterior side and a ventral median evagination called infundibulum which supports pituitary
gland or hypophysis. There is an olfactory area, mammillary body on the posterior side of
hypothalamus.
Mesencephalon is concerned with sight and hearing. Its dorsal side is called Tectum and
the ventral fibre bundles are called Crura cerebri or cerebral peduncle. The tectum has a pair of
bulging optic lobes on the anterior side and a pair of auditory lobes on the posterior side. In
lower animals auditory lobes are insignificant and optic lobes are prominent. This is called
Corpora bigemina. Higher vertebrates such as mammals and snakes have corpora
quadrigemina, which means they have optic and auditory lobes of equal size.
Metencephalon is called cerebellum which is quite enlarged in active animals. In
mammals cerebellum contains bundles of branching fibres of white matter called Arbor Vitae.
The bulging ventral side of cerebellum is called pons varolli and it contains criss-crossing fibres
of neurons.
Myelencephalon or medulla oblongata is the posterior part of the brain which does not
undergo much modification in vertebrates since it controls the autonomic functions of body. The
ventral side contains RAS (Reticular Activating and Inhibiting System) which keep the brain
attentive and awake. Dorsal side exhibits the posterior choroid plexus, which secretes
cerebrospinal fluid that flows into the brain ventricles and to meninges through a median
Foramen of Megendie and the paired Foramina of Luschka. Medulla is attached with cranial
nerves which bring sensory impulses from the body.
Brain is hollow inside; the cavities are called ventricles which are lined by ciliated
epithelium, ependyma. Ventricles of the two cerebral hemispheres are called lateral ventricles,
or Telocoel or I and II ventricles which are connected together with a foramen of Monro. The
third ventricle extends from diencephalon to mesencephalon and the IV ventricle is larger inside
metencephalon and myelencephalon. The third and fourth ventricles are connected together by a
tube-like connection called Iter or aqueduct of Sylvius.
MENINGES
Meninges are protective layers around the brain. The outermost layer is fibrous dura
mater (meaning tough mother) which, though tightly attached to the periostial layer of skull, still
encloses a narrow epidural space that is filled with perimeningeal fluid. The second layer under
dura mater is Arachoid, so named because of spider web like appearance due to presence of villi
for the absorption of cerebrospinal fluid. Between the dura mater and arachnoid exists the
subdural space and between arachnoid and the lower pia mater is the subarachnoid space. The
innermost layer of meninges is the delicate pia mater which is intimately attached with the brain
tissue and extends deep into the sulci and fissures. It carries blood vessels and nerves.
The three separate meningeal layers are found in mammals only, while in amphibia,
reptiles and birds, arachnoid and pia mater fuse to form a single pia-arachnoid layer below the
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subdural space. Fishes have a single meninx primitiva that is separated from the skull bone by
perimeningeal tissue.
CYCLOSTOME BRAIN
Cyclostome brain is very primitive owing to their parasitic and detritus feeding habits.
Cerebral hemispheres are small and smooth. Olfactory lobes are well developed as these animals
detect suitability of their hosts by acute sense of smell. For the same reason thalamus, which
contains olfactory relay centres, is enlarged with a prominent median olfactory area called
habenula. Optic lobes are small because of primitive or rudimentary eyes. Cerebellus which is
related with balance and posture is reduced due to parasitic mode of life. Medulla oblongata is
quite well developed and receives six pairs of cranial nerves but there is no pons varolli on the
ventral side. Pineal and parietal bodies are present in lampreys but absent in hag fishes.
FISH BRAIN
Active bony fishes and sharks have well developed brain but bottom dwelling fishes have
reduced brain organs. Olfactory lobes are large in sharks and they can detect their injured prey
by the smell of blood from a distance of about a kilometer. But in majority of bony fishes optic
lobes are reduced. Cerebral hemispheres are quite large but smooth and white. Pineal and
parietal bodies are generally reduced in fishes. On the ventral side of diencephalon, there is
saccus vaculosus posterior to the pituitary that serves as sense organ. Optic lobes are well
developed as most fishes are gifted with large eyes but in deep sea fishes they are reduced.
Cerebellum is highly enlarged in sharks as well as in active bony fishes and also has lateral
extensions called restiform bodies or auricular lobes which connect medulla with cerebellum.
They help in maintenance of balance. Cerebellum is smaller in rays, lung fishes, ganoid fishes
and deep sea fishes. Medulla oblongata has no particular variation except in deep sea fishes in
which there are large vagal lobes on the lateral side which receive impulses from taste buds that
are scattered all over the body as pit organs.
URODELE BRAIN
Urodele brain is primitive and reflects their sluggish nature and under-developed sense
organs. Olfactory lobes, optic lobes and cerebellum are reduced and cerebral hemispheres are
also small and smooth. Pineal and parietal bodies are present but reduced. There is no saccus
vaculosus and corpus striatum is weak.
ANURAN BRAIN
Frogs and toads possess a better developed brain as compared with urodeles. Olfactory
lobes are large and fused at base that gives better sense of smell to frog. Cerebral hemispheres
are larger with a centralized gray area called Archipallium that controls olfactory sense, while
the lateral areas are white and are called paleopallium. Parietal body is reduced but pineal is
well developed and probably a photoreceptor. Ancient amphibians possessed a third eye over the
pineal-parietal complex. Optic lobes are well developed but there are no auditory lobes and
hence the mesencephalon is called corpora bigemina. Cerebellum is reduced but medulla is
enlarged to make the brain a reflex brain or spinal brain.
REPTILIAN BRAIN
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Brain becomes large by the enlargement of corpus striatum. Cerebral hemispheres are
large and oval but the surface is white and smooth except in crocodiles which develop gray
matter called neopallium similar to mammals. Olfactory lobes are well developed in snakes and
lizards which have olfactory sense organs on the tongue, but reduced in turtles and crocodiles.
Parietal body is well developed in lizards and in Sphenodon it lies under a lens-like transparent
area called the third eye. Pineal body is absent in crocodiles. Optic lobes are well developed in
all reptiles and corpora quadrigemina is found only in snakes. Cerebellum is reduced in all
reptiles owing to their creeping habit.
BIRD BRAIN
Bird brain is characterized by enormous enlargement of cerebral hemispheres, optic lobes
and cerebellum. Cerebral hemispheres become enlarged owing to enlargement of corpus
striatum which is called hyperstriatum but pallium is thin and surface has only white matter.
Olfactory lobes are highly reduced, attached to the anterior end of the cerebral hemispheres.
Optic lobes are enormous as birds are gifted with the best eye sight in animal kingdom. Parietal
body is absent and pineal small in most of the birds. Being bipedal and flying animals, birds need
good control over muscles and tendons, which comes from a trilobed highly enlarged
cerebellum. The middle lobe of cerebellum is called vermis as it has transverse folds and the
lateral lobes are called flocculi. Birds’ brain is instinctive brain that can carry out complex but
stereotype functions such as nest building. Spinal cord still has equal control over the body.
MAMMALIAN BRAIN
Mammalian brain is highly developed and keeps complete control over the body
functions. Cerebral hemispheres are enormously enlarged and the surface is folded into
depressions (sulci) and raised portions (gyri) so that large surface area can be accommodated in
the small space of the skull. Gray matter has spread to the surface which is called neopallium.
Two cerebral hemispheres are connected by thick bundles of nerve fibres called corpus
callosum, which is not found in monotremes and marsupials. Olfactory lobes are highly
enlarged, so that mammalian brain is sometimes called nose-brain. Parietal body is absent and
pineal is usually present except in animals like Armadillo, Sirenia and Edentates. Mammals
being active animals, cerebellum is highly enlarged and trilobed and all lobes possess gyri and
sulci. Nerve cells form bundles of branching fibres called Arbor Vitae. Medulla is short as
compared to the large brain but pons varolli is enlarged.
COMPARATIVE ANATOMY OF BRAIN AND SPINAL CORD
Brain
The brain is safely kept inside the cranial vault. Inside the skull the brain is surrounded by three
protective coverings. They may be grouped under two divisions.
3. Pachymenix -It includes the duramater.
4. Leptomeninges - In includes the arachnoid mater and pia mater.
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The duramater is the outermost membrane. It is thick and inelastic in nature. The arachnoid
mater is the middle covering over the brain. In between arachnoid and piamater there is a space
called the subarachnoid space. It contains cerebro-spinal fluid and blood vessels. The piamater is
a delicate membrane closely applied to the brain. This membrane contains blood capillaries
supplying blood to the brain cells.
The human brain weighs about 1.3 Kg. It contains more than a billion neurons. Based on
embryological development the brain can be divided as follows.
Prosencephalon (Fore brain) –
It consists of the cerebrum and the diencephalon. The cerebrum is the largest part of the
brain. It is divided into right and left hemispheres by a longitudinal fissure. However, at the base
the two hemispheres are connected by a sheet of nerve fibres called the corpus callossum. The
outer surface of the cerebrum is called the cortex or grey mater. It is 2 to 4 mm thick. The inner
content of the cerebrum is the white mater. The surface of the cerebrum has several folds called
the gyri. They greatly increase the surface area of the cortex. The shallow grooves in between the
gyri are called the sulci. A central
sulcus runs in the lateral surface of
the cerebrum from superior to
inferior region.
Each cerebral hemisphere is
divided into four lobes. They are the
frontal at the front, the parietal
towards the top of the head, the
temporal on the side and the
occipital at the rear. The
diencephalon contains the thalamus
and hypothalamus. This region is
found between the cerebrum and the brain stem.
The thalamus has a cluster of nuclei which act as the relays for particular sensory
pathways. Just beneath the thalamus, the hypothalamus is present. It contains reflex centres
linked to the autonomic system. A funnel shaped stalk called the infundibulum extends from its
floor. It is connected to the neurohypophysis of the pituitary gland.
Mesencephalon (mid brain) –
It is the smallest region of the brainstem. On its dorsal surface there are four rounded bodies
called the carpora quadrigemina.
Rhombencephalon (hind brain) –
The three main regions of the rhombencephalon are the medulla oblongata, the pons varoli and
the cerebellum.
5. The cerebellum consists of two hemispheres. Its surface has many ridges called folia. The
cerebellum consists of three parts. They are the small anterior flocconodular lobe, a
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narrow central vermis and two large lateral hemispheres.

6. The pons is just superior to the medulla oblongata. It contains ascending and descending
nerve tracts.
7. The medulla oblongata is about 3 cm long. It is continuous with the spinal cord. It
remains as a bridge between the brain and the spinal cord.
8. Brain stem - The medulla oblongata, pons and mid brain form the brain stem. It connects
the spinal cord to the brain. Ten of the twelve cranial nerves enter or exit the brain
through the brain stem.
Spinal cord - The spinal cord extends from the foramen magnum to the level of the second
lumbar vertebra. It is considerably shorter than the vertebral column. There are two enlargements
in the spinal cord. They are the cervical and lumbar enlargements. Below the lumbar
enlargement the spinal cord tapers to form a cone like region called the conus medullaris. A
connective tissue filament called the filum terminale extends inferiorly from conus medullaris to
the coccyx. The conus medullaris and the nerves extending below resemble a horse’s tail. Hence
it is called cauda equina.
A cross section of the spinal cord reveals a central grey portion and a peripheral white
portion. The white matter consists of nerve tracts and the grey matter consists of neuron cell
bodies and dendrites. The dorsal and ventral sides have long fissures. There are 31 pairs of spinal
nerves arising from the spinal cord. Each nerve has a dorsal root and a ventral root from the
spinal cord. The dorsal roots have dorsal root ganglia.
Ventricles
The entire CNS remains as a hollow tube. The tube inside the adult brain forms
ventricles. Each cerebral hemisphere contains a large cavity called the lateral ventricle. It
corresponds to the hypothetical first and second ventricles. The two lateral ventricles
communicate with the third ventricle located in the centre of the diencephalon. This connection
is made through two interventricular foramina (foramen of Monro). The third ventricle inturn
opens into the fourth ventricle found inside the medulla oblongata. This communication happens
through a narrow canal called the cerebral aqueduct (aqueduct of sylvius). The fourth ventricle is
continuous with the central canal of the spinal cord. The central canal extends nearly to the full
length of the cord.
Cerebro-spinal fluid (CSF)
This fluid fills the ventricles of the brain and the central canal of the spinal cord. About
80-90 % of CSF is produced by specialized cells called ependymal cells within the lateral
ventricles. Remaining 10-12 % is produced by similar cells in the 3rd and 4th ventricles. These
ependymal cells, their supportive tissue and the associated blood vessels together are called
choroid plexuses. The plexuses are formed by invagination of the vascular piamater into the
ventricles.
Brain is the anterior end of spinal cord that has enlarged to take care of the sense organs
which are located on the anterior end of the body in a bilaterally symmetrical animal that moves
ahead in an anterior-posterior axis. This is called cephalon, which as it evolves further, divides
into three parts, namely, prosencephalon, mesencephalon and rhombencephalon. As the brain
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develops further by increasing the number of neurons, it further divides into different parts, each
one having assigned its own specific function.
Prosencephalon divides into Telencephalon and Diencephalon, the former includes
olfactory lobes (Rhinencephalon) and Cerebral hemispheres that coordinate the activities of the
entire brain. The roof of cerebrum is called Pallium and the floor that generally contains nerve
fibres is known as corpus striatum.
Diencephalon is a small part of brain, generally covered by enormously enlarged
cerebral hemispheres. This is an extremely important part of brain which functions as switch
board to cerebrum. Dorsal part of diencephalon is called epithalamus and the ventral part
hypothalamus while the lateral parts are called thalami that contain relay centres to connect
dorsal and ventral parts of thalamus.
Anterior part of epithalamus contains a glandular area called anterior choroid plexus
(Tela Choroidea) which secretes cerebro-spinal fluid. Two dorsal processes of epithalamus, the
anterior paraphysis supports parietal body and the posterior epiphysis bears pineal body. These
two bodies function as photoreceptors in lower vertebrates and gradually transform into
endocrine organs and biological clock in higher vertebrates.
The ventral hypothalamus has the optic chiasma (crossing of optic nerves) on the
anterior side and a ventral median evagination called infundibulum which supports pituitary
gland or hypophysis. There is an olfactory area, mammillary body on the posterior side of
hypothalamus.
Mesencephalon is concerned with sight and hearing. Its dorsal side is called Tectum and
the ventral fibre bundles are called Crura cerebri or cerebral peduncle. The tectum has a pair of
bulging optic lobes on the anterior side and a pair of auditory lobes on the posterior side. In
lower animals auditory lobes are insignificant and optic lobes are prominent. This is called
Corpora bigemina. Higher vertebrates such as mammals and snakes have corpora
quadrigemina, which means they have optic and auditory lobes of equal size.
Metencephalon is called cerebellum which is quite enlarged in active animals. In
mammals cerebellum contains bundles of branching fibres of white matter called Arbor Vitae.
The bulging ventral side of cerebellum is called pons varolli and it contains criss-crossing fibres
of neurons.
Myelencephalon or medulla oblongata is the posterior part of the brain which does not
undergo much modification in vertebrates since it controls the autonomic functions of body. The
ventral side contains RAS (Reticular Activating and Inhibiting System) which keep the brain
attentive and awake. Dorsal side exhibits the posterior choroid plexus, which secretes
cerebrospinal fluid that flows into the brain ventricles and to meninges through a median
Foramen of Megendie and the paired Foramina of Luschka. Medulla is attached with cranial
nerves which bring sensory impulses from the body.
Brain is hollow inside; the cavities are called ventricles which are lined by ciliated
epithelium, ependyma. Ventricles of the two cerebral hemispheres are called lateral ventricles,
or Telocoel or I and II ventricles which are connected together with a foramen of Monro. The
third ventricle extends from diencephalon to mesencephalon and the IV ventricle is larger inside
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metencephalon and myelencephalon. The third and fourth ventricles are connected together by a
tube-like connection called Iter or aqueduct of Sylvius.
MENINGES
Meninges are protective layers around the brain. The outermost layer is fibrous dura
mater (meaning tough mother) which, though tightly attached to the periostial layer of skull, still
encloses a narrow epidural space that is filled with perimeningeal fluid. The second layer under
dura mater is Arachoid, so named because of spider web like appearance due to presence of villi
for the absorption of cerebrospinal fluid. Between the dura mater and arachnoid exists the
subdural space and between arachnoid and the lower pia mater is the subarachnoid space. The
innermost layer of meninges is the delicate pia mater which is intimately attached with the brain
tissue and extends deep into the sulci and fissures. It carries blood vessels and nerves.
The three separate meningeal layers are found in mammals only, while in amphibia,
reptiles and birds, arachnoid and pia mater fuse to form a single pia-arachnoid layer below the
subdural space. Fishes have a single meninx primitiva that is separated from the skull bone by
perimeningeal tissue.
AMYGDALA:
Lying deep in the center of the limbic emotional brain, this powerful structure, the size
and shape of an almond, is constantly alert to the needs of basic survival including sex,
emotional reactions such as anger and fear. Consequently it inspires aversive cues, such as
sweaty palms, and has recently been associated with a range of mental conditions including
depression to even autism. It is larger in male brains, often enlarged in the brains of sociopaths
and it shrinks in the elderly.
BRAIN STEM:
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The part of the brain that connects to the spinal cord. The brain stem controls functions
basic to the survival of all animals, such as heart rate, breathing, digesting foods, and sleeping. It
is the lowest, most primitive area of the human brain.
CEREBELLUM:
Two peach-size mounds of folded tissue located at the top of the brain stem, the
cerebellum is the guru of skilled, coordinated movement (e.g., returning a tennis serve or
throwing a slider down and in) and is involved in some learning pathways.
CEREBRUM:
This is the largest brain structure in humans and accounts for about two-thirds of the
brain’s mass. It is divided into two sides — the left and right hemispheres—that are separated by
a deep groove down the center from the back of the brain to the forehead. These two halves are
connected by long neuron branches called the corpus callosum which is relatively larger in
women’s brains than in men’s. The cerebrum is positioned over and around most other brain
structures, and its four lobes are specialized by function but are richly connected. The outer 3
millimeters of “gray matter” is the cerebral cortex which consists of closely packed neurons that
control most of our body functions, including the mysterious state of consciousness, the senses,
the body’s motor skills, reasoning and language.
The Frontal Lobe is the most recently-evolved part of the brain and the last to develop in
young adulthood. It’s dorso-lateral prefrontal circuit is the brain’s top executive. It organizes
responses to complex problems, plans steps to an objective, searches memory for relevant
experience, adapts strategies to accommodate new data, guides behavior with verbal skills and
houses working memory. Its orbitofrontal circuit manages emotional impulses in socially
appropriate ways for productive behaviors including empathy, altruism, interpretation of facial
expressions. Stroke in this area typically releases foul language and fatuous behavior patterns.
The Temporal Lobe controls memory storage area, emotion, hearing, and, on the left side,
language.
The Parietal Lobe receives and processes sensory information from the body including
calculating location and speed of objects.
The Occipital Lobe processes visual data and routes it to other parts of the brain for
identification and storage.
HIPPOCAMPUS:
Located deep within the brain, it processes new memories for long-term storage. If you
didn't have it, you couldn't live in the present, you'd be stuck in the past of old memories. It is
among the first functions to falter in Alzheimer's.
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HYPOTHALAMUS:
Located at the base of the brain where

signals from the brain and the body’s
hormonal system interact, the hypothalamus
maintains the body’s status quo. It monitors
numerous bodily functions such as blood
pressure and body temperature, as well as
controlling body weight and appetite.
THALAMUS:
Located at the top of the brain stem, the thalamus acts as a two-way relay station, sorting,
processing, and directing signals from the spinal cord and mid-brain structures up to the
cerebrum, and, conversely, from the cerebrum down the spinal cord to the nervous system.
The evolution of Brain
The structure and function of the vertebrate brain have long been the subject of scientific
inquiry. Despite ongoing research, scientists are still not sure how the brain performs many of its
functions. For instance, scientists continue to look for the mechanism the brain employs to store
memories, and they do not understand how some memories can be “locked away,” only to
surface in times of stress. The brain is the most complex vertebrate organ ever to evolve, and it
can perform a bewildering variety of complex functions. His baby coyote is greeting the
approaching evening. His howling is not as impressive as his dad's—a good performance takes
practice. His brain is learning by repetition how to control the vocal cords.
Casts of the interior braincases of fossil agnathans, fishes that swam 500 million years
ago, have revealed much about the early evolutionary stages of the vertebrate brain. Although
small, these brains already had the three divisions, shown in figure 28.9, that characterize the
brains of all contemporary vertebrates: (1) the hindbrain, or rhombencephalon (colored yellow);
(2) the midbrain, or mesencephalon (colored green); and (3) the forebrain, or prosencephalon
(colored blue and purple).
The brain of a primitive fish.
The basic organization of the vertebrate brain can be seen in the brains of primitive
fishes. The brain is divided into three regions that are found in differing proportions in all
vertebrates: the hindbrain, which is the largest portion of the brain in fishes; the midbrain, which
in fishes is devoted primarily to processing visual information; and the forebrain, which is
concerned mainly with olfaction (the sense of smell) in fishes. In terrestrial vertebrates, thefore
brain plays a far more dominant role in neural processing than it does in fishes.
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The hindbrain was the major component of these early brains, as it still is in fishes today.
Composed of the cerebellum and medulla oblongata, the fish hindbrain may be considered an
extension of the spinal cord devoted primarily to coordinating motor reflexes. Tracts containing
large numbers of axons run like cables up and down the spinal cord to the hindbrain. The
hindbrain integrates the many sensory signals coming from the muscles and coordinates the
pattern of motor responses.
Much of this coordination is carried on within a small extension of the hindbrain called
the cerebellum (“little cerebrum”). In more advanced vertebrates, the cerebellum plays an
increasingly important role as a coordinating center and is correspondingly larger than it is in the
fishes. In all vertebrates, the cerebellum processes data on the current position and movement of
each limb, the state of relaxation or contraction of the muscles involved, and the general position
of the body and its relation to the outside world. These data are gathered in the cerebellum and
synthesized, and the resulting commands are issued to efferent pathways.
In fishes, the remainder of the brain is devoted to the reception and processing of sensory
information. The midbrain is composed primarily of the optic lobes (also called the optic
tectum), which receive and process visual information, while the forebrain is devoted to the
processing of olfactory (smell) information. The brains of fishes continue growing throughout
their lives. This continued growth is in marked contrast to the brains of other classes of
vertebrates, which generally complete their development by infancy. The human brain continues
to develop through early childhood, but no new neurons are produced once development has
ceased, except in the hippocampus, involved in long-term memory.
The Dominant Forebrain
Starting with the amphibians and continuing more prominently in the reptiles, sensory
information is increasingly centered in the forebrain. This pattern was the dominant evolutionary
trend in the further development of the vertebrate brain. The areas of the brain are color-coded in
figure 28.10 so that you can follow this trend in brain development, with the cerebrum of the
forebrain becoming larger in mammals.
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In sharks and other fishes, the hindbrain is predominant, and the rest of the brain serves
primarily to process sensory information. In amphibians and reptiles, the forebrain is far larger,
and it contains a larger cerebrum devoted to associative activity. In birds, which evolved from
reptiles, the cerebrum is even more pronounced. In mammals, the cerebrum covers the optic
tectum and is the largest portion of the brain. The dominance of the cerebrum is greatest in
humans, where it envelops much of the rest of the brain.
The forebrain in reptiles, amphibians, birds, and mammals is composed of two elements:
the diencephalon and the Telencephalon. The diencephalon consists of the thalamus and
hypothalamus (colored blue in figure 28.9). The thalamus is an integrating and relay center
between incoming sensory information and the cerebrum. The hypothalamus participates in basic
drives and emotions and controls the secretions of the pituitary gland, which in turn regulates
many of the other endocrine glands of the body (see chapter 30). Through its connections with
the nervous system and endocrine system, the hypothalamus helps coordinate the neural and
hormonal responses to many internal stimuli and emotions. The Telencephalon, or “end brain,” is
located at the front of the forebrain and is devoted largely to associative activity (colored purple
in figures 28.9 and 28.10). In mammals, the Telencephalon is called the cerebrum.
The Expansion of the Cerebrum
There is a remarkable difference in weight of brain between fishes, amphibians, and reptiles (to
the left of the branch point in the figure), and birds and mammals (to the right of the branch point
in the figure). Mammals in particular have brains that are particularly large relative to their body
size. This is especially true of porpoises and humans. The increase in brain size in mammals
largely reflects the great enlargement of the cerebrum, the dominant part of the mammalian
brain. The cerebrum is the center for correlation, association, and learning in the mammalian
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brain. It receives sensory data from throughout the body and issues motor commands to the
body.
CYCLOSTOME BRAIN
Cyclostome brain is very primitive owing to their parasitic and detritus feeding habits.
Cerebral hemispheres are small and smooth. Olfactory lobes are well developed as these animals
detect suitability of their hosts by acute sense of smell. For the same reason thalamus, which
contains olfactory relay centres, is enlarged with a prominent median olfactory area called
habenula. Optic lobes are small because of primitive or rudimentary eyes. Cerebellus which is
related with balance and posture is reduced due to parasitic mode of life. Medulla oblongata is
quite well developed and receives six pairs of cranial nerves but there is no pons varolli on the
ventral side. Pineal and parietal bodies are present in lampreys but absent in hag fishes.
FISH BRAIN
Active bony fishes and sharks have well developed brain but bottom dwelling fishes have
reduced brain organs. Olfactory lobes are large in sharks and they can detect their injured prey
by the smell of blood from a distance of about a kilometer. But in majority of bony fishes optic
lobes are reduced. Cerebral hemispheres are quite large but smooth and white. Pineal and
parietal bodies are generally reduced in fishes. On the ventral side of diencephalon, there is
saccus vaculosus posterior to the pituitary that serves as sense organ. Optic lobes are well
developed as most fishes are gifted with large eyes but in deep sea fishes they are reduced.
Cerebellum is highly enlarged in sharks as well as in active bony fishes and also has lateral
extensions called restiform bodies or auricular lobes which connect medulla with cerebellum.
They help in maintenance of balance. Cerebellum is smaller in rays, lung fishes, ganoid fishes
and deep sea fishes. Medulla oblongata has no particular variation except in deep sea fishes in
which there are large vagal lobes on the lateral side which receive impulses from taste buds that
are scattered all over the body as pit organs.
URODELE BRAIN
Urodele brain is primitive and reflects their sluggish nature and under-developed sense
organs. Olfactory lobes, optic lobes and cerebellum are reduced and cerebral hemispheres are
also small and smooth. Pineal and parietal bodies are present but reduced. There is no saccus
vaculosus and corpus striatum is weak.
ANURAN BRAIN
Frogs and toads possess a better developed brain as compared with urodeles. Olfactory
lobes are large and fused at base that gives better sense of smell to frog. Cerebral hemispheres
are larger with a centralized gray area called Archipallium that controls olfactory sense, while
the lateral areas are white and are called paleopallium. Parietal body is reduced but pineal is
well developed and probably a photoreceptor. Ancient amphibians possessed a third eye over the
pineal-parietal complex. Optic lobes are well developed but there are no auditory lobes and
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hence the mesencephalon is called corpora bigemina. Cerebellum is reduced but medulla is
enlarged to make the brain a reflex brain or spinal brain.
REPTILIAN BRAIN
Brain becomes large by the enlargement of corpus striatum. Cerebral hemispheres are
large and oval but the surface is white and smooth except in crocodiles which develop gray
matter called neopallium similar to mammals. Olfactory lobes are well developed in snakes and
lizards which have olfactory sense organs on the tongue, but reduced in turtles and crocodiles.
Parietal body is well developed in lizards and in Sphenodon it lies under a lens-like transparent
area called the third eye. Pineal body is absent in crocodiles. Optic lobes are well developed in
all reptiles and corpora quadrigemina is found only in snakes. Cerebellum is reduced in all
reptiles owing to their creeping habit.
BIRD BRAIN
Bird brain is characterized by enormous enlargement of cerebral hemispheres, optic lobes
and cerebellum. Cerebral hemispheres become enlarged owing to enlargement of corpus
striatum which is called hyperstriatum but pallium is thin and surface has only white matter.
Olfactory lobes are highly reduced, attached to the anterior end of the cerebral hemispheres.
Optic lobes are enormous as birds are gifted with the best eye sight in animal kingdom. Parietal
body is absent and pineal small in most of the birds. Being bipedal and flying animals, birds need
good control over muscles and tendons, which comes from a trilobed highly enlarged
cerebellum. The middle lobe of cerebellum is called vermis as it has transverse folds and the
lateral lobes are called flocculi. Birds’ brain is instinctive brain that can carry out complex but
stereotype functions such as nest building. Spinal cord still has equal control over the body.
MAMMALIAN BRAIN
Mammalian brain is highly developed and keeps complete control over the body
functions. Cerebral hemispheres are enormously enlarged and the surface is folded into
depressions (sulci) and raised portions (gyri) so that large surface area can be accommodated in
the small space of the skull. Gray matter has spread to the surface which is called neopallium.
Two cerebral hemispheres are connected by thick bundles of nerve fibres called corpus
callosum, which is not found in monotremes and marsupials. Olfactory lobes are highly
enlarged, so that mammalian brain is sometimes called nose-brain. Parietal body is absent and
pineal is usually present except in animals like Armadillo, Sirenia and Edentates. Mammals
being active animals, cerebellum is highly enlarged and trilobed and all lobes possess gyri and
sulci. Nerve cells form bundles of branching fibres called Arbor Vitae. Medulla is short as
compared to the large brain but pons varolli is enlarged.
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THE SPINAL CORD
The spinal cord is a cable of neurons extending from the brain down through the
backbone, which shows a darker gray area in the center that consists of neuron cell bodies, which
form a column down the length of the cord. This column is
surrounded by a sheath of axons and dendrites, which make
the outer edges of the cord white because they are coated
with myelin. The spinal cord is surrounded and protected by
a series of bones called the vertebrae. Spinal nerves pass out
to the body from between the vertebrae. Messages between
the body and the brain run up and down the spinal cord, like
an information highway.
Pairs of spinal nerves can be seen extending out from the spinal cord. Along these nerves, the
brain and spinal cord communicate with the body.
In each segment of the spine, motor nerves extend out of the spinal cord to the muscles. Motor
nerves from the spine control most of the muscles below the head. This is why injuries to the
spinal cord often paralyze the lower part of the body. A muscle is paralyzed and cannot move if
its motor neurons are damaged.
1. Transmission of nerve impulses. Neurons in the white matter of the spinal cord transmit
sensory signals from peripheral regions to the brain and transmit motor signals from the
brain to peripheral regions.
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2. Spinal reflexes. Neurons in the gray matter of the spinal cord integrate incoming sensory
information and respond with motor impulses that control muscles (skeletal, smooth, or
cardiac) or glands.
The spinal cord is an extension of the brainstem that begins at the foramen magnum and
continues down through the vertebral canal to the first lumbar vertebra (L 1). Here, the spinal
cord comes to a tapering point, the conus medullaris. The spinal cord is held in position at its
inferior end by the filum terminale, an extension of the pia mater that attaches to the coccyx.
Along its length, the spinal cord is held within the vertebral canal by denticulate ligaments,
lateral extensions of the pia mater that attach to the dural sheath.
The following are external features of the spinal cord (see Figure 1):
1. Spinal nerves emerge in pairs, one from each side of the spinal cord along its length.
2. The cervical nerves form a plexus (a complex interwoven network of nerves—nerves
converge and branch).
3. The cervical enlargement is a
widening in the upper part of
the spinal cord (C 4–T 1).
Nerves that extend into the
upper limbs originate or
terminate here.
4. The lumbar enlargement is a
widening in the lower part of
the spinal cord (T 9–T 12).
Nerves that extend into the
lower limbs originate or
terminate here.
5. The anterior median fissure
and the posterior median
sulcus are two grooves that
run the length of the spinal
cord on its anterior and posterior surfaces, respectively.
6. The cauda equina are nerves that attach to the end of the spinal cord and continue to run
downward before turning laterally to other parts of the body.
7. There are four plexus groups: cervical, brachial, lumbar, and sacral.The thoracic nerves
do not form a plexus.
8. Roots are branches of the spinal nerve that connect to the spinal cord. Two major roots
form the following:
9. A ventral root (anterior or motor root) is the branch of the nerve that enters the ventral
side of the spinal cord. Ventral roots contain motor nerve axons, transmitting nerve
impulses from the spinal cord to skeletal muscles.
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1. A dorsal root (posterior or sensory root) is the branch of a nerve that enters the dorsal
side of the spinal cord. Dorsal roots contain sensory nerve fibers, transmitting nerve
impulses from peripheral regions to the spinal cord.
2. A dorsal root ganglion is a cluster of cell bodies of a sensory nerve. It is located on the
dorsal root.
3. Gray matter appears in the center of the spinal cord in the form of the letter H (or a pair
of butterfly wings) when viewed in cross section:
4. The gray commissure is the crossbar of the H.
5. The anterior (ventral) horns are gray matter areas at the front of each side of the H. Cell
bodies of motor neurons that stimulate skeletal muscles are located here.
6. The posterior (dorsal) horns are gray matter areas at the rear of each side of the H. These
horns contain mostly interneurons that synapse with sensory neurons.
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7. The lateral horns are small projections of gray matter at the sides of H. These horns are
present only in the thoracic and lumbar regions of the spinal cord. They contain cell
bodies of motor neurons in the sympathetic branch of the autonomic nervous system.
8. The central canal is a small hole in the center of the H crossbar. It contains CSF and runs
the length of the spinal cord and connects with the fourth ventricle of the brain.
9. White columns (funiculi) refer to six areas of the white matter, three on each side of the
H. They are the anterior (ventral) columns, the posterior (dorsal) columns, and the lateral
columns.
10. Fasciculi are bundles of nerve tracts within white columns containing neurons with
common functions or destinations:
11. Ascending (sensory) tracts transmit sensory information from various parts of the body to
the brain.
12. Descending (motor) tracts transmit nerve impulses from the brain to muscles and glands.
Spinal Cord Regeneration
In the past, scientists have tried to repair severed spinal cords by installing nerves from another
part of the body to bridge the gap and act as guides for the spinal cord to regenerate. But most of
these experiments have failed because the nerve bridges did not go from white matter to gray
matter. Also, there is a factor that inhibits nerve growth in the spinal cord. After discovering that
fibroblast growth factor stimulates nerve growth, neurobiologists tried gluing on the nerves, from
white to gray matter, with fibrin that had been mixed with the fibroblast growth factor.
Three months later, rats with the nerve bridges began to show movement in their lower bodies. In
further analyses of the experimental animals, dye tests indicated that the spinal cord nerves had
regrown from both sides of the gap. Many scientists are encouraged by the potential to use a
similar treatment in human medicine. However, most spinal cord injuries in humans do not
involve a completely severed spinal cord; often, nerves are crushed. Also, although the rats with
nerve bridges did regain some locomotory ability, tests indicated that they were barely able to
walk or stand.
ORGANS OF VISION: STRUCTURE AND FUNCTIONAL SIGNIFICANCE.

VERTEBRATE EYE
All vertebrates have eyes that are similar in basic construction plan. This eye must have
evolved very early in chordate evolution and then got modified subsequently to increase its
efficiency in higher vertebrates.
The eye is formed of 3 coats or tunics.
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Coats or tunic Regions

1. Outer or fibrous - sclera & cornea
2. Middle or vascular - choroid, ciliary body & iris
3. Inner or nervous - retina
The sclera is the white outer layer of the eye. It covers posterior five- sixths of the
eye. This firm layer provides shape and protects the internal structures. A small region of
the sclera can be seen as the “white of the eye”.
In the front, the outer layer forms a transparent region called the cornea. It permits
entry of light. The cornea is
made up of a connective
tissue having collagen,
elastic fibres and
proteoglycans.
The middle tunic of
the eyeball is the vascular
tunic. It contains most of the
blood vessels. The vascular
tunic contains melanin
containing pigment cells. It
appears black in colour. A
major part of the vascular
tunic is found in association with the sclera and called the choroid. Anteriorly this layer
forms the ciliary body and iris.
The ciliary body consists of

smooth muscles called the ciliary
muscles. Contraction of the ciliary
muscles can change the shape of the lens.
The anterior compartment is
divided into two chambers. There is an
anterior chamber found between the
cornea and iris. A smaller posterior
chamber lies between the iris and lens.
These two chambers are filled with a
substance called the aqueous humor. It
helps to maintain intraocular pressure.
The posterior compartment of the eye is much larger and it contains a
transparent jellylike substance called vitreous humor.
The eye lens is a unique biological structure. It is transparent and biconvex. It is
made up of long columnar epithelial cells called lens fibres. These fibres have an
accumulation of proteins called crystalline. The lens is placed between the two eye
compartments by suspensory ligaments.
The functioning of the eye is aided by accessory structures. They include the
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eyebrows, eyelids, conjunctiva and lacrimal apparatus. The eyebrows prevent the sweat
during perspiration from running down into the eye. They help to shade the eyes from
direct sunlight.
The eyelids and associated lashes protect the eyes from foreign objects. The medial
region where the eyelids join has a small reddish-pink mound called the caruncle. It contains
modified sebaceous and sweat glands. There are two or three rows of hairs attached to the
free edges of eyelids. Modified sweat glands called the ciliary glands open into the
follicles of the eyelashes. It keeps them lubricated. The inner margin of the eyelids
contain Melbomian glands. These glands produce sebum for lubricating the eyelids.
The inner surface of the eyelids and the anterior surface of the eye are covered by a
thin, transparent mucous membrane called the conjunctiva.
The lacrymal glands or tear glands are situated in the superolateral corner of the
eye orbit. They produce tear at the rate of about 1 ml / day. It helps to moisten the eye
surface and wash away foreign substances. At the corners of the eye there are small
openings called the puncta. Each punctum inturn opens into a lacrymal canaliculus. The
lacrimal canaliculi open into a lacrimal sac. This sac enters into a nasolacrimal duct which
opens into the inferior nasal concha. These ducts help to drain the excess tear. The entire
organization related to ‘tear’ is called the lacrymal apparatus.
COMPARATIVE ANATOMY OF EYE
Protochordata do not have true eyes; Amphioxus has pigmented eye spots on the ventro-
lateral sides of the spinal cord for detecting light and shades.
CYCLOSTOME EYE
Hagfishes have rudimentary eyes that are hardly one mm in diameter but Petromyzon has
well developed eyes. The eye ball is flat and eye lids are absent. Lens is permanently spherical
and there are no ciliary bodies. Sclera and choroid are fused and pupil has fixed diameter.
FISH EYE
Eyelids are immovable and permanently open and nictitating membrane covers the eye in
front. In elasmobranchs, cornea is flat in front, lens permanently spherical which is moved
forward and backward by protector lentis muscle. There is no fovea in area centralis but there is
a layer of shiny guanine crystals called tapetum behind retina for night vision. Sclera is
cartilaginous in elasmobranchs.
In bony fishes also the eye ball is elliptical and lens permanently spherical that almost
touches the cornea in front. Lens is moved forward and backward by a muscle called campanula
Halleri that is attached to the processus falciformis at the back of the eye ball. There is no fovea
on retina and the layer of reflecting cells is called argentea.
AMPHIBIAN EYE
Limbless amphibians have rudimentary eyes owing to their burrowing habits. Urodeles
also have small eyes which are essentially fish like in structure. However, anurans have evolved
an eye that is suitable for vision in terrestrial conditions.
Frog eye is oval in shape with three distinct layers, fibrous sclera, vascular choroid and
sensory retina. Eye lids are movable and lower eye lid is larger and also carries a nictitating
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membrane. Lachrymal glands are also located on the lower side of the eye ball and a harderian
gland is present on the back of the eye ball. Lens is flat on the front side but convex on the
posterior side. Ciliary body is well developed but lens is moved forward and backward for
focusing by a protractor lentis muscle. The whole eye ball can be pulled back into the eye orbit
by a retractor bulbi muscle and can be brought forward by levator bulbi muscle. Frog
possesses limited colour vision.
REPTILIAN EYE
Reptilian eye is made on amphibian plan and is adapted for terrestrial environs. Sclera is
cartilaginous or bony and retina possesses a fovea centralis. Lizards also have a conus
papillaris protruding on the lower side of the fovea. It probably has nourishing function. Lens is
soft and flexible and can alter focal length by squeezing with the help of ciliary muscles.
Lachrymal glands are well developed except in Sphenodon and snakes. Lower eye lid is still
larger and nictitating membrane is present. In snakes eyelids are fused and become transparent to
form a spectacle in front of the eye. Crocodiles have a pecten as in birds. Colour vision seems to
be present in some lizards and turtles.
BIRD EYE
Bird eye is best developed among all vertebrates which is a necessity for a flying animal.
Bird eye ball is large and depressed in the area of powerful ciliary body. Sclerotic coat is tough
with cartilages and bones. Retina is highly sensitive having abundance of cones and a very deep
fovea centralis. A pecten is present in birds which increases contrast and reduces glare in bright
sunshine. Lens is highly flexible and can be squeezed by two powerful muscles, namely, muscle
of Crampton and muscle of Bruke. Iris is supplied with striated muscles. Nictitating membrane
is used to cover the front of eyeball when the bird is in flight. Except for the nocturnal birds, all
birds possess colour vision.
MAMMALIAN EYE
In mammals sclera is made of fibrous tissue and continues in front as cornea. Retina has
a fovea but there is no pecten. Iris can change the diameter of pupil by radial and circular
muscles. Lens is soft and capable of changing focal length by altering its thickness with the help
of circular and meridional muscles present in the ciliary body. Upper eyelid is large in
mammals and lachrymal gland is located on the upper side of the eyeball. Nictitating membrane
is present in some mammals such as horse, anteaters, panda and caribou. A modification of iris
called umbraculum is found in cats, camels etc. which cuts excessive glare in daytime.
ORGANS OF HEARING AND TACTILE RESPONSES
VERTEBRATE EAR
The ears are the organs of hearing and balance. They have three parts, namely external,
middle and inner ears.
External ear - The fleshy part outside the head is called the pinna. It is made up of elastic
cartilage and skin. It is followed by the external auditory meatus. This passage is lined with hairs
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and ceruminous glands. These glands produce cerumen or earwax. The hair and wax prevent
foreign objects from reaching the ear drum. The ear drum or tympanic membrane is an oval,
three layered structure. It separates outer and middle ears.
Middle ear - It is an air filled cavity. It contains three auditory ossicles called the malleus, incus
and stapes. The handle of malleus is in contact with the inner surface of the ear drum. The head
of the malleus is attached to the incus. While the stapes on one side is attached to the incus, its
other side fits into the oval window. The oval window leads to the inner ear.
Inner ear - This region has tunnels and chambers inside the temporal bone called the bony
labyrinth. The bony labyrinth contains three regions called the cochlea, vestibule and
semicircular canals. The oval window found in between the middle and inner ears
communicates with the vestibule of the inner ear. The organs of the inner ear perceive the
sound.
Actual ear is a delicate organ located in the temporal region of the skull. It is called
membranous labyrinth, which evolved in primitive vertebrates for maintenance of balance and
posture. Hearing function of this organ evolved for the first time in anurans when they started
living in terrestrial environment, where sound waves travel faster and to longer distances. A
typical membranous labyrinth
consists of three canals and three
sacs. The three canals are attached
at right angle to each other and to
the sacs in anterior, posterior and
horizontal positions. The three
sacs which are also collectively
called vestibule are: Utriculus,
sacculus and lagena, each one of
which possesses a depression
inside with a sensory organ called
macula that detects the position of
the body in relation to gravity, or static sense. Each canal ends in an ampulla that also carries a
sense organ called crista, which is responsible for detection of the movement of head in any
direction, or kinetic sense. Both these senses together maintain the balance and posture of the
animal during its activity. Endolymph fills the membranous labyrinth and is secreted and drained
in meninges. Perlilymph surrounds the membranous labyrinth for protection.
The structure of crista is like any neuromast organ of fishes. It consists of sensory cells
with hairs on the top and nerve fibres attached below and many supporting cells that nourish and
protect the sensory cells. Hairs of sensory cells are embedded in a gelatinous mass called cupula
terminalis in which some crystals are embedded just above the hair tips. As the animal moves
these crystals also move and touch the hairs of sensory cells, which produce nerve impulses that
take the kinetic sense to the brain. This is called dynamic equilibrium maintained by the
semicircular canals.
Macula is the sensory organ of sacs. Its sensory cells also have hairs on the top and nerve
at the base but the cells are of two types, namely, cylindrical cells and flask shaped cells which
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bear stereocilia and kinocilia embedded in a gelatinous mass. A paper thin otolith made of
calcium carbonate and proteins floats over the cilia and its position can be disturbed by the
animal bending or tilting in any direction in relation to gravity, disturbing the cilia in the process
and generating a nerve impulse that informs the brain of static sense.
CYCLOSTOMES
Cyclostomes have degenerated membranous labyrinth due to their parasitic mode of life.
Petromyzon has only two semicircular canals, namely the anterior and posterior and only two
sacs, utriculus and sacculus, lagena being absent. As they have no paired fins, lampreys and
hagfishes do not seem to have three dimensional sense of things. In hagfishes, the anterior and
posterior canals also fuse together to make a ring like structure. Sacculus and utriculus also fuse
together to form a single sac.
FISHES
Fishes use membranous labyrinth for balance and posture only, since the middle and
external ears are absent but some fishes can perceive water borne sounds to some extent. For
example, in fishes of the family Ostariophysi four pieces of Weberian ossicles (scaphium,
claustrum, intercalarium and tripus) connect the swim bladder with membranous labyrinth
and transport sound vibrations for hearing.
Fish membranous labyrinth is well developed with three canals and three sacs, and crista
and macula well formed. In elasmobranchs a recessus utriculus is attached on the lower side of
utriculus. Macula of utriculus is sometime called pars neglecta in fishes.
AMPHIBIA
Urodela and apoda do not have external and middle ear and consequently inner ear is
meant for maintenance of balance and posture only. Hearing power evolved in anurans by the
modification of lagena which produces an organ of reception of sound waves, called basilar
papilla. On the surface, on either side of head, a tympanic membrane is stretched over a ring of
cartilage called annulus tympanicus, which is a modification of pterygoquadrate cartilage of the
first visceral arch. Sound vibrations received by this membrane are transmitted to basilar papilla
through a columella bone, which is a modification of hyomandibular cartilage of the hyoid
arch. Thus, in frog a mechanism has evolved to receive sound vibrations from outside and
transmit them to the sense organ of lagena, so that the nerve impulse can convey information
about the sound to brain.
REPTILES & BIRDS

In reptiles and birds lagena elongates and basilar papilla elaborated to form the organ of
corti, which is much more sensitive to perceive sound vibrations. Columella bone is called
stapes, which is further elongated by the addition of an extrastapaedial cartilage. For better
protection, tympanic membrane sinks into the temporal bone, producing an auditory canal,
through which sound vibrations travel from outside to the tympanic membrane. In crocodiles
lagena is coiled like cochlea of mammals. In reptiles both Eustachian tubes join together to
open into pharynx by a single opening.
In snakes there are no external and middle ear cavities and hence sound vibrations travel
through ground to the lower jaw and from there to quadrate bone to columella, which carries
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them to lagena of inner ear for perception of hearing. Therefore, snakes cannot hear sounds
travelling through air.
Birds have essentially similar mechanism of hearing as do the reptiles, except that ear is
much more sensitive in birds due to elongation of lagena.
MAMMALS
Mammals possess the best developed hearing power among all vertebrates. Lagena
elongates to form a spirally coiled cochlea. In middle ear cavity, instead of one bone, mammals
have three ear ossicles, namely, incus ( a derivative of pterygoquadrate), malleus (a derivative of
articular or meckel’s cartilage) and stapes (derived from hyomandibular), which transmit sound
vibrations from tympanic membrane to the fenestra ovalis that leads to scala vestibuli of
cochlea. Mammals also have various sizes of ear pinna to collect
sound waves and direct them to the auditory canal.
Cochlea is a specialized and highly sensitive sense organ of
hearing. Its cross section reveals three long chambers, namely,
scala vestibuli, scala media and scala tympani, the middle
chamber is filled with endolymph while the other two are filled
with perilymph. The organ of corti is attached to the basilar
membrane and carries sensory hair cells and supporting cells and
the cochlear nerve at the base. A tectorial membrane floats in the
middle of scala media and touches the hairs of sensory cells when it
vibrates by the sound vibrations, resulting in the generation of a
nerve impulse that travels via the cochlear nerve to the brain. Base of cochlea is more sensitive
to high frequency vibrations while the apical portion can detect very faint vibrations. Sound
waves from stapes enter scala vestibuli through fenestra ovalis or oval window, then travel to
the entire length of cochlea and return back via scala tympani and escape to the middle ear cavity
through fenestra rotunda or round window.
Human ear is capable of hearing sound waves ranging from 20 to 20,000 hertz but
different mammals possess capacity to hear sounds of different frequencies. For instance,
whales, dolphins and porpoises can hear ultrasonic sounds of up to 150,000 hertz and elephants
can also hear infrasonic sounds of 10-14 hertz.
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Complete Chordata Notes

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CHORDATA NOTES

ASSOCIATE PROFESSOR (CONTRACT)

4.1 Comparative Anatomy of Respiratory organs

SUGGESTED READING MATERIAL:

15. Write an essay on comparative anatomy of respiratory system.

GEOLOGICAL TIME SCALE

EVOLUTINARY TIME SCALE:

Define evolutionary time scale:

The Evolutionary Time scale or Geological time scale (GTS) is a system of

What does the Evolutionary time scale represent?

Divisions of Evolutionary time scale

The Geologic Time Scale is divided by the following divisions:

Cretaceous: From the Latin “creta” meaning chalk by a Belgian geologist

Mesozoic 210 - 65 m. years ago “Golden age of reptiles”.

earth’s history is formulated into eras-periods-epochs. Each division in the geological

TERTIARY - Oligocene 38 - 26 m. years ago Monkeys and apes originated

TERTIARY - Eocene 54 - 38 m. years ago Horses originated

CRETACEOUS 130 - 65 m.years ago Dinosaurs became extinct

SILURIAN 350 - 315 m. years ago Jawed fishes originated

CAMBRIAN 600 - 440 m. years ago Thallophytes, Arthopods,

PRECAMBRIAN before 440 m. years ago Protozoans,

Poriferans and Annelids lived.

Ordovician period :- (440 to 350 million years ago)

This period was marked by formation of

Silurian period :- (350 to 315 million years ago)

Devonian Period :- (315 to 275 million years ago)

Mississippian Period :- (275 to 255 million years ago)

Pennsylvanian :- (255 to 235 million yeaars ago)

The land liiving forms became

Permiian Period :- (235 to 210 million yeears ago)

This middlle period in the

Triasssic Period :-- (210 to 1600 million yeaars ago)

For the firsst time fossills of turtles, crocodiles, and

Cretaceous Period :- (130 to 65 million years ago)

Cenozoic Era :- (65 million years ago till date)

Triceratops - a horned dinosaur

Modern placental mammals originated during this time.

Several animals with ancient characteristics became extinct. Modern mammalian

Evolutionary significance of fossils

Fossils provide us clues regarding climatic conditions of various prehistoric periods.

Ichthyostega - interconnecting link between fishes and amphibians. Seymouria -

Extinct animals-mass extinction

VERTEBRATE INTEGUMENT AND DERIVATIVES, SKIN STRUCTURE AND FUNCTION,

SKIN STRUCTURE AND FUNCTION

Table 1.1 Layers of the skin.

Skin layer Description

Epidermis The external layer mainly composed of layers of keratinocytes but

Basement membrane The multilayered structure

and non-dividing keratinocytes, which are attached to the basement membrane by

Dermoepidermal junction/basement membrane

1. A thin papillary layer

The papillary dermis lies below and connects with

Blood and lymphatic vessels

Derivative structures of the skin

Figure 1.4 Nail structure.

1. Maintaining the epidermal permeability barrier, structure and differentiation

sympathetic (adrenergic) nerve fibres.

Table 1.2 Functions of the skin.

Barrier function and skin desquamation

thus increasing our susceptibility to dry skin conditions.

Figure 1.5 Barrier function of the epidermis.

Shedding (desquamation) of skin cells

Table 1.3 Immune components of the skin.