UNIT 9 URINOGENITAL SYSTEM

Structure
9.1 Introduction
Objectives
9.2 The Urinary System in Chordates
Urinary System in Protochordates
Urinary System in Vertebrates
9.3 Embryonic Development of Urinary System
9.4 Uriniferous Tubules
9.5 The Kidney
Structure of Kidney
Blood Circulation in Kidney
9.6 Phylogeny and Succession of Kidneys
9.7 Functions of Urinary System
9.8 Variations in the Urinary System Plan
Habitat Related Structural Variations
Variations in Urinary Systems of Vertebrates belonging to different vertebrate Groups
9.9 The Genital System
Embryonic Origin of Gonads and Gametes
Functions of the Genital System
Genital System of Protochordates
9.10 Male Genital System'of Vertebrates
Testes
Male Genital Duct
Male Accessory Sex Glailds
Intromittent Organ5
9.11 Female Genital System of Vertebrates
Ovary
Female Genital Ducts
Female Accessory Glands
External Female Genitalia
Mammary Apparatus
9.12 Survey of Gonads in Vertebrates
9.13 Summary
9.14 Terminal Questions
9.15 Answers

9.1 INTRODUCTION
- - -

Excretion and osmoregulation as you are aware (Refer LSE-05 Unit 4) are two
homeostatic processes performed by the kidneys and urinary ducts, which form the
urinary system. This system is intimately associated both anatomically, and in
terms of embryonic origin with the genital system. The genital system includes the
gonads wlilcli generate gametes and the genital ducts that serve as passages for the
gametes - sperms In males and ova in females. Though functionally different the two
organ systems the urinary and the genital system are treated together as the urino-
genital system, since both develop from the same segmental blocks of trunk
mesoderm or ad-jacent tissues and share many of the ducts. Thus although the two
systems have nothing common functionally they are closely associated in their use of
common ducts and are studied under the broad heading of urinogenital system. In
this unit, you will learn about the embryonic development of the urinogenital system,
the basic plan in vertebrates and the variations in different chordate groups. You will
also study the adaptive variations of urinary system in vertebrate living in specialised
habitats.

Objectives
After going through this unit, you should be able to:
describe and illustrate the urinogenital system of the proto chordates -
cephalochordates and urochordates,
illustrate and describe the basic plan of urinogenital system in vertebrates,
give a comparative account of the urinogenital system of fish, amphibians, reptiles,
birds and mammals.
Functional Anatomy of
-
Chordates 11
describe the origin and embryonic development of kidneys, gonads and their
associated ducts,
discuss the morphological and physiological adaptations of the urinogenital system
of vertebrates.

9.2 THE URINARY SYSTEM IN CHORDATES

9.2.1 Urinary System in Protochordates
The urinogenital system of protochordates such as the Cephalochordate [e.g.
Branchiostoma (amphioxus)] ahd Urochordate, (e.g. Herdmania) is very different both in
structure and origin from that of the vertebrates (Refer unit 1).

Cephalochordata
Despite the similarities to vertebrates in other aspects of its anatomy, the specialised
organs of excretion in the cephalochordate, Branchiostoma (amphioxus) show no
relationship to any part otthe vertebrate kidney or other known fluid regulating structure.

The organs of excretion in Branchiostoma are the protonephridia - which are

ectodermal in origin (unlike vertebrate kidneys which originate from the mesoderm).
Protonephridia(Fig, 9.1) are segmentally arranged sac-like tubes which lie in coelomic
spaces or atrium abbve the pharynx. Each protonephridium opens into the peribranchial
or atrial space surrdunding the gills so that its excretory products released into the atrium,
can be flushed away by the outgoing stream of water. Internally, within the coelomic sac
the protonephridium terminates blindly (not opening) into the coelom. (see also Fig.
9.17a in this unit)
Dorsopharyngeal coelom
. / Coelomic fluid

Nucleus

II Fig. 9:l: The protonephridium of Broncl~iostoma.(Arrows show path of fluid flow)

i
The protonephridibl sac bears numerous, specialised tubular flame cells called the
solenocytes (Fig. 9.2.). Each solenocyte has a single flagellum projectrng downwards
into the protonephridial tube or canal. Constant beating of flagella probably forces fluid
into the protonephridia. The exact mechanism of functioning of solenocytes is still not
- Flagellum
clear. As the distql ends of solenocytes lie close to branchial (gills) blood vessels, the
solenocytes probably help to filter blood fluids from the branches of branchial blood
vessels. In the prdcess some fluid components such as ions are probably returned to the
body (Fig. 9.1).
Fig. 9.2: Enlarged view of a single
flame cell or solenocyte. Urochordata
In the urochordatg, Herdmania, neural gland is supposed to be the excretory organ (Fig.
9.3) and lies embedded in the mantle above the nerve ganglion or brain. The neural gland
consists of few central tubules from which arise peripheral tubules. The central tubules
open into a longitbdinal canal. The longitudinal canal opens by a ciliated funnel at the
base of dorsal tubercle. The excretory cells are called nephrocytes. They are present in
blood in order to kollect waste which are in the form of xanthene and urates. The waste
passes through th lumen of the neural gland and its duct and is finally discharged into
f
the pharynx. Neu a1 glands are endocrine in function as well, since they also control
oviposition, development and metamorphosis.
 Zlrinogenital System

Blood sinusci

hr,of ncunl

u
Fig. 9.3: Neural glar~dof Herdnranin.

9.2.2 Urinary System in Vertebrates

The basic plan of urinogenital system is common to all vertebrates. The urinary organ-
system lies in the abdominal region (Fig. 9.4 a and b) and consists of paired kidneys which
are the principal organs of excretion and osmoregulation. These kidneys lie dorsal to the
coelom, (retroperitoneal), one on each side of the dorsal aorta. The basic units of the
kidney are rhe minute, uriniferous tubules that end blindly and receive filterate from the
blood. The uriniferous tubule consists of two parts, the nephron and collecting tubule. The
'
primary function of the uriniferous tubules is removing excess water, salts, waste
metabolites and foreign substances from the blood. Numerous such tubules connect to
form an urinary or excretory duct system called ureter which carries urine from the
kidney. The ureter of each kidney opens into a urinary bladder which in most adult
tetrapods is a single median structure. Urinary bladder acts as a reservoir which stores
urine prior to its removal. The bladder develops as a ventral outgrowth from the cloaca.
The bladder opens to the body surface through a short tube, the urethra. The urethra is
thus a small tube which arises from the bladder and opens by a single,urinary opening
either into the cloaca or directly to the outside.
l-------'II~l~qt~Ir b n w r ard

Fig. 9.4: Ventral view ofthe human excretory systems in (a) male. (b) female.

9.3 EMBRYONIC DEVELOPMENT OF URINARY

SYSTEM
Urinogenital system is mesodermal in origin. The intermediate mesoderm which is
located in the dorsal and posterior body wall of the embryo forms the kidney. At the
onset of its differentiation this posterior region of the intermediate mesoderm expands to
form a nephric ridge that protrudes slightly from the dorsal wall of the body cavity (Fig.
9.5 a). Next the paired nephrotome which are the embryonic fore runner of the nephric
tubule (Fig. 9.5 b) are formed from the posterior part of intermediate mesodehn.

The nephroto~neis often segmented and soon differentiates into a series of segmentally
arranged tubules each of which contain a nephrocoel, which is a coelomic chamber that
Functional Anatomy of opens into the coelom by means of a ciliated peritoneal funnel. The medial end of the
Chordates - I1
nephrotome then widens to form a thin walled renal capsule into which enters a tuft of
arterial capillaries aalled glomerulus. From the lateral end of the nephrotome arise
outgrowths. 'These outgrowth fuse to form a common nephric dpct (Fig. 9.5 c). 'The
nephrotome is now properly called a nephric tubule and may retain its connection with
the coelom through the persistent peritoneal funnel. Thus the fundamental plan
underlying the excretory system consists of paired and segmented nephric tubules that
open on one end into the coelom and on the other end into the nephric duct with a
.glomerulus in behveen (Fig. 9.5 d). In most adult vertebrates, however, there is no
connection with the coelom through peritoneal funnel.

In the development of the urinary duct of ureter, longitudinal ducts appear first at the
anterior end of nephrogenous mesoderm as backwardly directed extensions of the
excretory or nephric tubules. The extensionsjoin to form the 'nephric duct' which opens
into the cloaca. Al~lthe nephric tubules ultimately open into this duct.
Nave luk n
\
Mewneph~
ric
duct

tubule

Fig. 9.5: Embryonia development of nephric tubules i n vertebrates. (a) Embryo (lower part) showing
location of developing kidney (nephric ridge); (b) section through embryo showing the
appearance of segmental nephrotome i n the posterior part of the intermediate mesoderm; (c)
diagrammlatic representationof developing kidney tubule and longitudinal nephric duct. note
the segmeatal arrangements; (d) the medial end of the nephrotomes differentiates into the first
part oftha nephric hrbule. the renal capsule into which the glomeruls grows. Arterial sprouts
from dorsbl aorta form the glomerules. The lateral ends of nephrotomes grow outward and fuse
with each ~ t h e to
r form the nephric duct. Sometimes the nephrotomes remain connected to the
coelom byi means of ciliated peritoneal funnel.

The number o f uriniferous tubules vary The functional units of the kidney as mentioned before are the minute uriniferous
from only a few hundred in the tubules (Fig. 9.6). Each tubule is tonstituted of (i) nephron (nephric tubule which is
kidneys of cyclostomes to over a
million per kidney i n mammals, in
the excretory unit) and the (ii) collecting tubule (Fig. 9.6) into which the nephron
whom the tubules o f both kidneys empties. Nephrbn (nephros: Gk Kidney); concentrates urine and passes it through the
combined constitute over 120 km of collecting tubule. The collecting tubule affects the concentration of urine and conveys it
tubing. to the minor calyx, (see Fig. 9.9 a) which is the beginning of the excretory duct.
 llrinogenital System

Fig. 9.6: llriniferous tubule - the basic excretory unit.

I) Nephron
Each nephron has: a) a dilated portion -renal (Malpighian) corpuscle and b) the
secretory tubule portion.

(a) Renal Corpuscle or Malpighian Corpuscle

The renal corpuscle is the proximal portion of each nephron and is made of (i) double
walled squamous epithelial capsule called Bowman's or renal capsule. The outer
layer of the capsule is called parietal layer and inner one is called the visceral layer.
Between the layers of Bowman's capsule is a urinary space receiving fluid filtered
through capillary wall and visceral layer. (ii) A tuft of inter-arterial capillaries called
glomerulus is surrounded by the Bowman's capsule. Each glomerulus is formed by
the subsequent branching of the renal artery (one of the major branches from the
dorsal aorta) into a capillarity bed. The visceral or inner layer of the Bowman's
capsule is made of stellate cells called podocytes. The visceral layer is in contact with
blood vessels of glomerulus. The point of eMry of the blood vessels forms the
vascular pole and the entry point of the nephric filtrate from the Bowman's capsule
into the proximal convoluted tubule is called the urinary pole. Fig. 9.7 shows a
complete uriniferous tubules, surrounded by its blood vascular system.

(b) Tubular portion

The tubule portion is subdivided into (a) proximal convoluted tubule (PCT) (b) loop
of Henle with thick and thin limbs and (c) distal convoluted tubule (DCT). The PCT
begins at the urinary pole of renal corpuscle (Fig. 9.8) and is Kited by simple
epithelial cells which have microvilli forming a brush border and lots of
mitochondria and the enzyme Na' K' ATpase . PCT has a wide lumen and is
surrounded by peritubular capillaries.

Henle's loop is U-shaped with thick descending and ascending limbs. Initially thick
(the thick part is the straight part or pars recta of the proximal tubule) the loop of
flenle thins near the turns then thickens again as it becomes part of the distal
convoluted tubule. Henle's loop has a large lumen lined by squamous epithelium
but no brush border.

The distal convoluted tubule (DCT) is continued from the ascending limb of Henle's
loop and forms the last segment of the nephron. It is lined by cuboidal epithelium
which lacks brush border. At the vascular pole, DCT establishes contact with renal
corpuscle of the parent nephron (see Fig. 9.7 a, b and 9.8).
Functional Anatomy of
-
Chordates 11

\. tvvntl layer made orpodtxytca

Rovrmm s
apeprulc
I I

Fig. 9.7: The basic excdetory unit :a)uriniferous tubule showing its relationship to its surrounding blood
vascular systelm, b) an enlargement of the boxed area in a showing the juxta glomerular apparatus.

11) The Collecting Tubule and Collecting Duct

Urine passes fdom the distal convulated tubule to the collecting tubule. Collecting
tubules of nephrons join together to form collecting ducts or papillary ducts of
Bellini. These ducts widen near tips of renal or medullary pyramids of the kidney.

Distal tubule

Fig.9.8: Magnified view of the Borvman's capsule showing poles the jurta'glomerular cells and ~naculadensa.
 llrinogcnital System
Juxtaglomerular Apparatus
'The wall of afferent arteriole near the renal corpuscle is made up of modified smooth
muscle cells called juxtaglomerular cells which contain lots of Golgi bodies. The
secretion of these cells is Renin which helps to regulate blood volume. The wall of the
distal tubule at this region is modified and looks dark under the microscope, hence it is
called macula densa. The afferent arteriole juxtaglomerular sells and macula densa
form the juxtaglomerular apparatus (Fig. 9.7 b).

9.5 THE KIDNEY

- -

The kidneys are specialised for maintaining he appropriate levels of water and many
solutes in the body and for eliminating wastes of protein metabolism and in canying out
osmoregulation. The kidneys of vertebrates are especially important in eliminating
excess water and divalentions and conserving solutes. In this section we will just deal
with the kidney and the urinary system of vertebrates. Let us begin by examining the
structure of the kidney, using the mammalian kidney as an example. This will familiarise
you with the terminologies that are used to describe the anatomical complexity of the
vertebrate kidney. When the kidneys fail in a human being, the person can be put on an
artificial kidney temporarily. (See Box 9.1)

Box 9.1

COIand u r dilyrir lanpcmm b l h didpis

fluid fluid

In case of renal-failure in humans, the patient has to be put on an artificial kidney. Haemodialysis or CAPD (Continuous-
Ambulatory Peritoneal Dialysis) are employed as the techniques. Artificial kidney is a mechanical device which can
separate waste products from blood and works on the principle of dialysis. The process employed is called
haemodialysis (GK: haeme - blood; lysis - 'separate'). A semipermeable membrane made of porous cellophane
separates large non-diffusible particles like blood cells from small diffisible molecules of urea and other waste products.

9.5.1 Structure of Kid ey

P
e
The mammalian kidney (Fig. .9) has a smooth external surface. The typical
metanephric, mammalian kidn y (Refer subsection 9.2.2) is a bean:shaped organ
attached to the dorsal wall and is retroperitoneal in position. It has at its medial side a
cavity or depression called hilum or hilus through which the ureter leaves the kidney. At
'the exit point of the ureter, a renal vein also leaves the kidney and a renal artery and
-
- nerve enters it.
Functional Anatomy of -
-
Chordates I1

Fig. 9.9: Sccli)n of mammalian kidney showing cortex, medulla and departure of ureter.

A section of the kidney reveals that the kidney is surrounded by a capsule of connective
tissue, under which lies the cortex. The renal corpuscles and the convoluted portions of
the secretory tubules lie entirely in the cortex (Fig. 9.10). Immediately beneath the cortex
is the medulla of the kidney which is striated in appearance and contains the loops of
Henle and collecting tubules. The medulla is partially composed of large areas known as
medullary pyramid0 or renal pyramids (Fig. 9.9). The medullary pyramids are made up
of parallel tubules called medullary rays. Each medullary ray contains one or more
collecting tubules and nephrons. Parts of the cortex between the medullary rays form the
renal column of Bkrtini. The outer borders of the pyramids are subdivided into smaller
units called lobules. The collecting tubules lie in the pyramids but may extend well up
towards the cortex. The collecting tubules of nephrons join together to form collecting
ducts or papillary putts of Bellini. There ducts widen near tips of renal or medullary
pyramids. The inndr portion of each pyramid is in the form of a blunt papilla and projects
into an out pocketihg of the pelvis known as minor calyx (plural: calyces). several minor
calyces join together to enter the major calyx, which in turn opens into the renal pelvis.
Urine produced by the kidney enters the minor and then major calyx. The pelvis leads to
the ureter which empties into .the bladder (cloaca in monotremes). Urine which is stored
temporarily in the bladder passes to the outside through the urethera.

Mammalian kidneys have two types of nephrons and the ratio of each varies depending
on the species (Fig. 9.10). One type the cortical nephron, has its glomerulus in the outer
kidney cortex with p tubule that extends only in the outer region of the medulla. Cortical
nephrons which prddominate in the human kidqey (90%) have short loops of Henle. The
second type have their glomeruli in the deep cortex close to the edge of the kidney
medulla and therefore are called juxta medullary nephron. These which predominate
(100%) in mammalb of dry habits have long loops of Henle that dip deeply into the
rrtedulla. Kidney tubules show'tremendous variation in vertebrates.

Fig. 9.10: A diagrammbtic representation of section of mammalian kidney showing a cortical nephron
and juxta mddullary nephron.
Urinary bladder is present in all mammals. It is a muscular sac, derived from the ventral
cloaca1 wall. The bladder narrows down and connects to the outside by the urethra. The
lower ends of the ureters except in monotremes open directly into the bladder on its
dorsal surface. In monotremes they open into the urethra through small papillae near the
base of the bladder. At the junction of bladder and'brethra there is an abundance of
elastic tissue. Much of the bladder musculature in the form of bundles continues down to
the urethra. Urine flow ceases when the length of the urethra is increased and diameter of
lumen is reduced. In males the urethra passes through the penis to open at the tip of that
organ through the external urethral orifice or urethral meatus. In females the condition
varies. In some, as in rat and mouse the urethra opens independently to the outside,
passing through the clitoris; in others it enters a urinogenital sinus or vestibule, which is
the terminal part of the genital and urinary tract.

9.5.2 Blood Circulation in Kidney

Kidneys are highly vascular (Fig. 9.1 1). The renal artery delivers blood to each kidney.
Before entering the kidney it divides into two branches - one going to anterior part of
kidney and other to the posterior part. The branches give interlobular arteries which are
located in the renal medullery pyramids. At corticomedullary junction, they form the
arcuate artery. Fig. 9.1 1 shows the supply of blood to the nephron. Afferent arterioles
arise from the interlobular arteries and supply blood to the glomerular capillaries. Blood
passes from these capillaries into efferent arterioles. Efferent arterioles branch to form
peritubular capillary network which nourish PCT and DCT. They also carry away
absorbed ions and molecules.

The efferent arterioles associated with juxtamedullary nephrons form long thin capillary
vessels which run straight into the medulla and loop back towards the corticomedullary
boundary. These are called vasa recta or straight vessels. 'They provide nourishment
and oxygen to medulla since they contain filtered blood.

Renal veins take the same course as renal arteries. Blood from interlobular vein form
arcuate vessels. The interlobular vessels then converge to form renal vein through which
blood leaves the kidney (Fig. 9.1 1). .
Renal Interstitiurn: In both cortex and medulla, the spaces between the urinary tubules,
contain cells called interstitial cells, which secrete prostaglandin.
- .

- --

I;ig. 9.1 I: Scctiolb of kidney showing Mood circulation.

SAQ 1
Fill in the blanks with appropriate words.
i) The bench of capillaries surrounded by the Bowman's capsule is called
.......................................
ii) A notch present on the medial side of human kidney is called
......................................
iii) .........................artery delivers blood to each kidney.
iv) Urinogenital system is ..........................in origin.
Functional Anatomy of
-
Chordates 11 9.6 PHYLOGENY AND SUCCESSION OF KIDNEYS
- - - - - -

Comparative studieb of vertebrate development indicate that the ancestral kidney called
archinephros (ancieht kidney) of the earliest vertebrates extended along the length of the
co6lomic cavity and was made up of segmentally arranged tubules, each resembling an
invertebrate nephridium. Each tubule of the archinephros (also called holonephros)
opened at one end ihto the coelom by a nephrostome (ciliated funnel) and at the other end
into a common nephric duct called archinephric duct. The archinephric duct of each
kidney runs along the dorsal side of the coelom and opens into the cloaca. A segmental
kidney is found in the embryos of hagfishes and caecilians (Fig. 9.12 a).

Kidney of living vdrtebrates have developed from this primitive plan. During the
embryonic life of the amniote vertebrates, the three types of kidneys pronephros,
mesonephros and metanephros (Fig. 9.12) appear in succession. There is a succession of
three developmental stages of kidneys, so that this is termed 'succession of kidneys' or
'tripartite concept of kidney' (Fig. 9.12 b). Some but not all of these Stages are
observed also in other vertebrate groups.

F$. 9.12: Comparatb'e development of niale vet tebrate kidney. (a) archinephros (b) pronephros (c)
mesonephros (d) metanephros. (Light colour representsdegenerated or underdeveloped
structures and bright red the functional structures).

Pronephros
In all vertebrate embryos, the first kidney to appear is the pronephros. In it the pronephric
. tubules are anteriorly located and segmentally arranged. One end of these tubules open

1
into the coelom y a nephrostome while other end opens into the archinephric duct which
in this case is cal ed pronephric duct. The two pronephric duct, one on each side extend
posteriorly a ~ *en
d into the cloaca. Pronephros is located anteriorly in the body. Only in
the adult hagfishes (anamniote) does the pronephros become part of the'persistent kidney
also called "head kidneyn (Fig. 9. I2 b). In all other vertebrates it degenerates during
development an4 is replaced by a more centrally located mesonephros.
I
 llrinogenital System
The term 'opisthonephros" of the embryonic anamniotes is not quite comparable to'the
mesonephros of the embryonic amniotes, even though the two are structirally similar. By
convention the term mesonephros is used for the structure which appears during
embryonic development in reptiles, birds and mammals.

Mesonephros
In the vertebrate embryo, tubules arising from the middle of the nephric ridge constitute The term archinephric duct is applied to
the mesonephros (Fig. 9.12 c). Mesonephros is a sometimes known as the Wolffian the kidney duct in anamniotes. The
body. Mesonephric tubules are segmentally or metamerically arranged in the beginning name wolftian duct is given to the duct
in amniotes which forms in connection
but as more tubules rise this metamerisms is lost. The tubules open into an already with the pronephros and mesonephros.
existing pronephric duct and no new duct is formed. When the pronephros degenerates
the persistent pronephric duct is called mesonephric duct or the wolffian duct. The
mesonephric duct may enlarge at its posterior end to form the (i) urinary bladder for
storage of urine or (ii) the seminal vesicle for storage of sperms.

The mesonephros is the functional kidneys of embryonic amniotes (reptiles birds and
mammals) and contributes to the adult kidney (called an opisthonephros) in anamniotes
ofjawless vertebrates, fishes and amphibians (Fig. 9.12 and Fig. 9.13).

Fig. 9.13: Vertebrate excretory systems (a) comparison of the evoldtion (i-iv) and (b) embryonic
'
development (v-vii) of the vertebrate kidney and its duct.

Mesonephros may persist after birth in reptiles, the egg laying mammals (Prototheria) and
pouched mammals (the metatheria). In the adult amniotes, the functional kidney is the
metanephros. As soon as the metanephros becomes functional, Wolffian duct
(mesonephric duct) degenerates in females and persists in males as urinogenital duct.
Functional ~ n a t o n i yof Parts of the male genital ducts such as epididymis and ductus deferens as well as the
Chordates - I 1
seminal vesicles develop from mesonephric tubules.

Metanephros
During embryonic development of amniotes - (the reptiles, birds and mammals) the
pronephros, mesonephros and metanephros appear in succession. The adult kidney of
amniotes is the methnephros (Fig. 9.12 d). 'The metanephros develops as the posterior
part of the nephric ridge and during development, it gets displaced anteriorly and
laterally. The displaced metanephros develops its own new duct, the metanephric duct or
the ureter. The dilated tip of metanephric duct becomes the renal pelvis (Fig. 9.9).

The metanephrosis is distinguished in several ways from pronephros and mesonephros. It

is more caudally (posteriorly) located and is a much larger more compact structure
containing a very large number of nephric tubules. It is drained by a new duct, the ureter,
which develops when the old archinephric duct is relinquished to the reproductive system
of the male for sperm transport. Thus the three successive kidney types - prosnephros,
mesonephros, metahephros succeed each other embryologically and to some extent
phylogenetically in amniotes (Fig. 9.13).
>

9.7 FUNCTIONS OF URINARY SYSTEM

The urinary system has several functions (Fig. 9.14):
I. The kidneys produce urine which contains metabolic waste products such as
ammonia or urea or uric acid etc depending on the animal group. Urine passes
through ureters to the bladder. It is temporarily stored in the bladder and is then
released to the outside via urethra through urinary or urinogenital opening. Urine is
formed by filteration, reabsorption and secretion (Fig. 9.14 a).

WhmrWmnmEofblmdhlbaain~
h ~ g u ~hi& nk in blmd mamic p m s m rhu b Inmeifid
fluid oprorrocpron in Spodulmus simulued

wur 0' urs b k d N r l

I l q m h h of blmd v * . r mmd b k d p n r m r r .

b m o m Nm' radmrglm
Rnndnaal tubuk

4
Plnrmm volume ramred
C 4
Blmd prcuurc d ~ z e d

Fig. 9.14: Functions oqurinary system (a) removal of metabolic wastes by lilteratius, reabsorption and
secretion (uliine formation) (b) osnloregulation (c) regulation of blood volume.
 Zlrinogenital System
2. Kidneys regulate fluid and electroyte balance and hence serve in osmoregulation and
water bal-ance(Fig. 9.14 b).
3. Kidneys produce an enzyme renin which has a function in regulating blood pressure
(Fig. 9.14 c).
4. The cortical cells of kidneys secrete a hormone Erythropoietin which is associated
with production of blood cells. In case of loss of blood or hypoxia, erythropoietin
formation is stimulated. Erythropoietin acts on bone marrow to promote formation
of erythrocytes.
5. In certain male vertebrates, apart from carrying urine, the ducts from kidneys also
carry sperms and are thus called urinogenital ducts. In female vertebrates, ureters act
only as a passage for urine.

SAQ 2
i) Name the
a) Functional unit of the kidney.

b) Tuft of capillaries surrounded by renal capsule.

c) U-shaped loop between PCT and DCT.

.............................................................................................
d) Collecting duct.
.............................................................................................
ii) State the parts of the uriniferous tubule in its proper sequence.

iii) Tick the appropriate functions of the kidney.

a) Removal of metabolic waste.
b) Osmoregulation.
c) Transmission of gametes.
d) Secretion of renin and erythropoietin

iv) Match the following in column A with terms in column B.

Column A Column B
a) capillaries of efferent arterioles i. Prostaglandin
b) Renal interstitium ii. Juxtaglomerular apparatus
c) macula densa iii. Red blood cells
d) erythroporetin iv. vasarectae
v. Sperms

9.8 VARIATIONS IN THE URINARY SYSTEM PLAN

You have already learnt in section 9.7 of this unit that the two major functions of the
kidneys are (I) excretion of metabolic wastes and (ii) maintenance of water and
electrolyte balance (osmoregulation). It has been observed that apart from the small
variations in structure and function of kidneys that have evolved in the urinogenital
systems of vertebrates due to living in different habitats the pattern of the renal system is
fairly uniform and apparently simple both in the anamniotes and amniotes. However,
study does show that there are great differences from group to group in the construction .
of the urinary system in different vertebrate groups these namely Agnatha, Fish,
Amphibia, Reptiles and Mammals. This is due to difference in habitat.

9.8.1 Habitat Related Structural Variations

Vertebrates evolved in water. Some emerged on land. Life in freshwater and in
the sea offered different problems of osmotic concentration. Since kidneys were
the organs for osmoregulation, the kidney tubules underwent structural
modifications to meet the regulatory needs of each vertebrate group in its own
specific environment.

Kidneys of vertebrates are confronted with two kinds of problems:

Functional Anatomy of 1. Water eliminlation in the fresh water organisms and
Chordates - I1
2. Water consertvation in the marine and terrestrial organisms.

Fresh Water Organism

[Link] water organisms, kidneys have (i) large well developed glomeruli which produce
large volumes of glbmerular filterate. (ii) Prominent distal tubules absorb salts and amino
acids from the filtrate to retain them within the body and also absorb much less water.
Lots of water is eliminated in the urine. .

Marine Organism
Marine organisms face dehydration as the outside environment (sea water) has greater
osmotic concentration. So water tends to move out of the body of the marine organism.
The structural modifications in these organisms is adapted to retain water and are:
i) Loss of glomeruli (aglomerular kidney).
ii) Shortening or loss of distal segments of kidney tubules which are responsible for
reabsorption of salts. Both the above modifications cause water retention and
increased salt excretion so that homeostasis is maintained in these organisms.
iii) Marine fishes have extra renal structures for excretion of salts. Elasmorbranchs
(cartilaginous fishes) have chloride secreting glands on the surface of gills and also
rectal glands which secrete salts. Marine reptiles and birds have salt excreting -
paired large nasal glands located in a bony socket with duct opening into nostril.
Refer to unit 9 of LSE-05.

Terrestrial Organism
Terrestrial organisms also need to conserve water. Snakes have aglomerular kidneys for
this purpose. In mammals, the Henle's loop of the nephron concentrates the urine by
reabsorbing water firom it. The length of loop of Henle's varies with the environment of
the organism. For axample beaver which have lots of water in their immediate
environment have short, Henle's loops. Both long and short loops of Henle are present in
rabbit and humans which have an intermediate capacity for concentrating urine (see Fig.
9.6 again). The sand rat, on the other hand which is found in arid conditions has very
long loops of Henle's in order to reabsorb as much water as possible and so excretes a
very concentrated urine.

9.8.2 Variatians in Urinary Systems of Vertebrates belonging to

Differeat Groups
The vertebrate kiddeys themselves are varied in structure; the ducts vary as does the
urinary bladder. These variation are due to two major causes: (I) The kidney unlike
many organs must start functioning at an early stage of development in order to remove
the metabolic wastes of the rapidly growing embryo. The kidney is however, subject to
modification or replacement in the later developmental stages and adult existence (ii) as
the gonads (testis and ovary) lie adjacent to the kidneys. These organs specially the testis
tend to take over part of the tubes and tubules of the urinary structures as a conducting
system for their prdducts. Asa result of this the urinary organs have been markedly
modified in most vertebrate grcups. In this subsection we will survey the construction
and hence normal variations of urinogenital systems in various vertebrate groups (Fig.
9.15 a tog).

Agnatha (Cyclostome) o r Jawless Vertebrates

The adult kidney ofjawless vertebrases lamprey (Petryomyzon) is opisthonephros or
Mesonephrons and the tubules are derived from the posterior mesomere. The tubules
have lost all connection with the coelomic cavity but form a close association with the
glomerular blood vessels at the blind-ended kidney capsple. The absence of connection
between kidney tubules and coelom makes it evident that "coelomic fluid filter" was
replaced by "blood filter system" early in vertebrate evolution. This may be because
possibly coelomic fluid pressure is too low and variable to keep up with the fluid balance
demands of vertebrtates which have high metabolic rates.

In adult lamprey the kidney is elongated and flauened. The kidneys lie on either side of
the mid-dorsal line and each is suspended by a mesentery like membrane. The
 llrinogenital System
archinephric duct courses along the free edge of the kidney. Tlie pronepliros exists above
the ~nesonepllrosas the non functional "head kidney".

In the adult hagfish the opisthonephros consists simply of a series of tubule arranged in a
segmental manner along most of the length of the trunk, each tubule draining directly into
the archinephric duct or pronephros. In hagfish adults the archinephros becomes
modified to form a persistent head kidney. The remainder of the kidney, however is
separated from the head kidney and becomes the opisthonephros. The hagfish has only
forty glomeruli, each connected to the collecting duct (archinephric duct) by a s~nallneck
segment. (Fig. 9.15 a)

Fish
The kidneys'of fislies are opisthonephric or mesonephric and though tliey exhibit great
variation in shape, they are however funda~nentallysimilar in structure (Fig. 9.15 b and
c). In all species they are dorsal in position. In some species they extend almost the entire
length of coelom. In other fishes tliey may be more voluminous and the two sides may
sliow various degree of fiision. In still otliers they are sllort and confined to the posterior
part of the body cavity. Peritoneal funnels are only retained in a few forms, notably Anria,
Sturgeons and certain elasmobranchs. In some marine teleosts no external or internal
glomeruli are present, and so such kidneys are called as aglomerular kidneys.

Generally the kidneys of male fisli are longer than those of females, since the anterior
ends in male are appropriated by the reproductive system. In males of some groups small
modified kidney tubqles now called efferent ductules, connect the testes with tlie
archinepliric duct. This archinephric duct is termed as the ductus deferens and serves as
the passage for sperm transport. It may, in addition continue to carry wastes. However, in
such cases there is a marked tendency for the posterior portion of the opisthonephros to
take over the greater part of the excretory function, with one or more accessory ducts
developing which are responsible for carrying wastes directly to the cloaca or to the
outside. In Selacliians, Chondrosteans and some otliers the conncection of the testis and
archinephric duct usually occurs at the anterior end of tlie ~[Link] teleosts
however the testes and opisthonepliric kidneys are not connected. In them the duct from
the testes either join the archinephric ducts near the posterior ends or open independently
to the exterior.

In some fislies the dilation of the archinephric duct may form a bladder like enlargement
for temporary storage of urine (Fig. 9.15 c). In those fish where archinephric duct
functions as ductus deferens, enlargements called seminal vesicle and sperm sac may
develop to serve as temporary storage places for spermatozoa,

Amphibians
Tlie amphibian kidney is opisthonepllric or mesonephric and the anterior tubules ofitlie
opisthonepliros functions as tlie sperm duct. The primitive archinephric type of kidney is
found in tlle larval caecilians of the amphibians. The kidney is similar to the larval kidney
of the hagfish and consists of the distinct ~netamericarrangement of kidney tubules, renal
corpuscles and nephrostome. In the adult the kidney is opisthonephros. The
opisthonepliros is lobulated and extends the greater part of the length of the coelom. In
many larval amphibians a small head kidney with peritoneal connection may be present
but it does not persist in the adult.

The urodele amphibians have opisthonephric or mesonephric kidneys that are similar to
the elamobranchs (cartilaginous fisli). The kidneys have two regions (i) anterior narrow
region which in males is more genital than urinary in function and is referred to as
epididymis and (ii) posterior expanded portion which is the main part of the
opistho~iephrosand is called the 'kidney proper'. The archinepliric ducts run along the
lateral edge of tlie kidney a short distance from tlie kidney proper. Numerous collecting
ducts or tubules which are more developed in males than fe~iialesjoin at intervals to tlie
archinephric duct from the opisthonephros. The archinephric duct now called
mesonephric ducts serves in tlie male as a ductus deferens as well as transports wastes but
in females is concerned only with transporting wastes. The mesonephric ducts in both
sexes open into the cloaca on either side. through a small papillia (Fig. 9.15 d).
In the anurans the opisthonephric kidneys are dorsoventrally flattened have a more
posterior concentration of tubules. They are confined to the posterior part of the
abdominal cavity and so are retroperitoneal and dorsally located. Unlike in the urodeles
the anterior and posterior regions of the kidney are not clearly distinguished. An
yellowish, orange adrehal gland is closely attached to the ventral side of the kidney. In
females the kidneys anid reproductive system have no connection to each other. However,
in males they are intimlately connected. In males certain anterior kidney tubules become
modified into efferent ductules that connect the testis to the kidney while the mesonephric
duct serves to transport spermatozoa as well as urinary wastes. In these animals unlike the
urodeles the archinephtic duct is located within the kidney along its lateral margin. It
leaves the opisthonephfos near the posterior end and passes to the cloaca (Fig. 9.15 e).

Openin8 of umer into c l o r r

Bird (Aves)

Fig. 9.15: Urinary systemsof vertebrates (a) cyclostome (b) cartilaginous flsh (c) bony fish (d) salamander
(amphibian), (e) frog (amphibian), (f) lizard (g) bird.
A thin walled, bilobed bladder which is thought to be endodermal in origin and
homologous with the urinary bladder is present. It is derived from the cloaca1 wall and
opens into the amphibian cloaca a short distance beyond the openings af the aichinephric
ducts. The archinephric duct and bladder are not connected directly, and so the thin,
watery urine tirst passes directly into the cloaca.

Reptiles
Reptiles are the tirst co~npletelyterrestrial vertebrates adapted to fresh water,
estuarine and even marine habitats. Kidney in reptiles are metanephric. In reptiles
the kidneys are usually small and compact with lobulated surface arid confined to the
posterior half of the abdominal cavity, generally to the pelvic region (Fig. 9.15 f).
The posterior part of the kidney narrows down on each side and in some lizards the
hind parts may even fuse. The degree of symmetry varies in reptiles especially in
case of snakes and limbless lizards who due to their elongated body shape possess
excessively lobulated, long, narrow kidneys; often one kidney may be entirely
posterior to the other.

In snakes and crocodilians a urinary bladder is absent. Most turtles and lizards
however have well-developed and usually bilobed bladders which open into the
cloaca. Except in turtles the ureters open separately in the cloaca. In turtles the ureters
are connected to the bladder. In some turtles a pair of accessory urinary bladders is
also connected with the cloaca. These functions as accessory organs of respiration. In
females they may be filled with water. which is used to soften the ground when they
prepare their nest.

Living reptiles have only about a few thousand nephrons in which loops of Henle are
absent and renal corpuscle are poorly developed. The glomeruli are small in order to
conserve water.

Birds
The kidneys in all birds are situated in the pelvic region ofthe body cavity and their
posterior ends are frequently united. They are complex, lobed structures with short
ureters, which open independently into the cloaca (Fig. 9.15 g). The kidneys contain some
mammalian-like kidney tubules with loops of Henle (some short and some long loops)
that parallel collecting ducts. Birds mostly, however have reptilian type tubules that do
not have loops. Urinary bladder is absent in all birds except for ostrich. Urinary wastes
are ~nainlyin the form of uric acid and are eliminated via the cloaca along with the faeces.
The ability of l<idney to concentrate urine and thereby conserve water is not as good as in
mammals. but is better than in reptiles. The cloaca and even the posterior large intestine
may further modify and concentrats urine by resorption of water and ions.

The mammalian urinary system has already been dealt is the previous sections. Refer
subsection 9.2.2, sections 9.4, 9.5 and fig. 9.4 for details.

SAQ 3
i) Tick the correct alternative:
a) Marine organisms possess
I) sI101-t distal tubule
2) long distal tubule
3) very long distal tubule
4) short proximal tubule.

b) Snakes have aglomerular kidney in order:

I) to conserve water
2) to remove water
3) to conserve protein
4) to remove protein.

c) The desert kangaroo rat in order to combat dehydration :

I) has long Hen le's loop for reabsorption
 2) utilises metabolic water
3) eats waterladen vegetation
4) all of the above.
-

9.9 THE GENITAL SYSTEM

The genital system is tlie reproductive systetn and is made up of tlie gonads,
gonoducts and genital openings. The gonads as you have read are mesodermal in
origin. They are called testes (sing testis) in males and ovaries in female. Both the
testes and tlie ovaries produce gametes and secrete hormones and are termed primary
sex organs.

The gonads are usdally paired structure, though unpaired gonads occurs in some forms
such as cyclostomds aawless fish), certain fish as well as female birds of most species.
This is the result oteither fusion of paired gonads or the unilateral degeneration of one
gonad. Evidence of metamerism of gonads in chordates is only found is the primitive
ampliioxus - cephdochordate.

You will recall that ann~nniolcsinclude In vertebrates the dvaries and testes come to be attached to the dorsal body wall by
cartilagillous and bony tishes and mesentery like bantls of tissue, the mesorchium in the male and mesovarium in the
amphibians while amniotes include the fernale. In most ve~ebratesthe gametes (ova in females and spermatozoa in males)
reptiles, birds and mammals.
produced by the gdnads are transported outside the body by means of the deferens
ducts (vas deferensb in males and by oviducts in females. In a few forms like
cyclostonie ducts ake absent in both sexes. Eggs and sperm escape from the body
cavity through genital pores. The deferent ducts as you will recall are usually the
mesonephric or tlie Wolffian ducts which also serve to transport urinary wastes from
the opisthonephric ior mesonephric kidneys in those animals in which these kidneys
function either during embryonic or adult life (in male anaminotes). In arnniotes in
which the metanepliros is tlie functional kidney and in which the mesonephros
degenerates, the Wolffian duct of the male on each side persists to become the
ductus deferens (male genital duct).

In most vertebrates in both sexes when the reproductive ducts first develop they open
posteriorly into the cloaca. This relationship persists throughout life in many vertebrates,
but in others tlie cloaca1 region becomes modified so that the reproductive ducts either
open separately to the outside or in the males atleast, join the excretory ducts to emerge
by a common opening or urinogenital opening.

In many aquatic vertebrates fertilisation is external while in all terrestrial vertebrates

except for anurari amphibians and even in many aquatic species it is internal. In some
animals transport of spermetazoa from male to female is brought about by apposition of
the cloaca (c1oaca:sing) of the two sexes in most animals, however, males have
copulatory organs which are used in an intromittent manner to deposit the spermatazoa
into the reproductive tract of the female. Various types of copulatory organs are found in
the vertebrate groups.

In both sexes all the structures or organs which help to bring the germ cells or products of
the primary sex organs together are termed as accessory sex organs. These include the *
reproductive ducts, associated glands and intromittent organs. Secondary sex characters
are indirectly concarned with sex but play a part in the reproductive scheme. Sexual
differences in such secondary sex characters like plumage, body size and strength, as well
as vocal apparatus are only indirectly related to reproduction.

Cloaca (Roman term for sewer) is present in a variety of vertebrates. It occurs as a

ventral pocket, at tlie back end of the trunk of the animal, and opens to the exterior.
The orifices of tlie digestive, genital and urinary systems open into it. The cloaca
appears to be a primitive vertebrate feature. Primitively, tlie gut. urinary duct and
genital ducts all terminate in a common, short chamber, the cloaca that empties to the
outside.

Most tetrapods (amphibians, birds reptiles, egg laying mammals) have retained a distinct
cloaca, although tnost mammals have not.
SAQ 4
Fill in the blanks witli appropriate words.
i) The primary sex organs are the ........................... and ................................
ii) Mammary glands are .................................... sex organs.
iii) ...................................... duct becomes the ductus deferens.
iv) The .................................are attached to the dorsal body wall by the
mesovariu~n.
v) When reproductive ducts first develop, they open into ........................... .........
vi) The tissue that s ~ ~ s p e n the
d s testis from tlie dorsal body wall is called
....................................
vii) Copulatory organ is an .................................... organ.

9.9.1 Embryonic Origin of Gonads and Gametes

In ;impliibian males. 1l1e cloaca1 glands
The sex of an individual depends basically upon the nature of its chromosomal are prescnl \\,liicli secrele scell1 in order
inheritance. The early development of tlie embryo is mainly because of the organisation lo allracl the female for mating. In them
already present in tlie unfertilised egg, the influence of the sperm and the hereditary clocal glands also secrete ;I jelly like
substance \vhicli holds logether a
features it introduces, all which are not obvious until a relatively late stage. We thus package of sperlns which is called
observe that for some time the sex organs of tlie embryo remain in an indifferent stage spcrn~[Link] l'emale picks up the
during which the dcvelopmcnt of the gonads and their ducts proceed considerably far spermrilopliore in her cloaca for
without any indication of whether tlie embryo would develop into a male or female. fcr~ilisatio~lof her eggs.
Finally the embryo develops its specific sex features, presumably due to hormonal
activity.

Embryonic origin of Gonads

Maleness in chordates develop basically on a female plan. Both the gonads and kidneys
develop from tlie intermediate mesoderm (mesomere) of the embryo (Fig. 9.16 a).

In vertebrates there is a great difference in the rapidity with which different organ
systems develop. The nervous system for instance, grows very rapidly in early stages
while the genital organs are one of the slowest to develop. The gonads make their
appearance only at a stage when most of the other organ systems have been blocked out
and the coelolilic cavities are well developed. Paired longitudinal swellings the genital
ridges develop along the roof of tlie coelom, medial to the embryonic kidney and on
either side of the dorsal mesentery (Fig. 9.16 b). These ridges gives rise to the gonads.

The gonads developing from such ridges are elongate to begin with but often in later
stages become short and compact with a L I S L Itendency
~~ for anterior concentration of
tissue. The germinal epithelium of the ridges are continuous witli the mesodermal lining
of the rest of the coelom and form the more important structural parts of the gonad.

Mesenchyma lying beneath the epithelium forms the connective tissue and in higher
vertebrates at least gives rise lo tlie special interstitial tissues that are thought to be a
source of gonad hormone.

Before the end of the indifferent stage, the gonad generally develops in many cases into a
swollen structure which extends out into the coelomic cavity from its dorsal wall in the
neigbourhood of the developing kidney and is often supported by a special mesentry. From
the germinal epithelium that cover the gonad surface, finger like structures called the
primary sex cords grows inward into the underlying mesencliyma of the gonad (Fig. 9.16 b).

The gonads of both sexes initially contain germ cells. These germ cells do not arise in the
genital ridge nor even in the ad-jacent mesoderm. In fact they do not arise in the embryo at
all. They f?rst rise in remote sites outside the embryo in the extra-embryonic endoderm.
From here they migrate to tlie indifferent gonad where they get located permanently. In
females the germ cells establish residence in the cortex while in males they get located in
the medulla.

As gonads mature they enlarge and are pushed downwards where they lie suspended by a
dorsal mesentry termed mesorchium in males and mesovarium in females.
Functional Anatomy of
Chordates - 11

~ p l n s LO^
l --& Glomerular capsule -Lx

Embryo cross section

Glomerular capsul

c Porm~tionof female
reproductive system

Van defdrens
Urinary bladder i

Proslalc

U
Formution of malc
re()rsductive syrtem

Fig. 9.16: Development ofgonads in vertebrates showing modification of the indifferent gonad into ovary or
testes. Thegonadal structure, whether primary cortex or medulla is derived from embryonic
mesoderm (a) the primodial germ cells which give rise to spermatozoa or egg are initially located
i n the beginning, within the embryonic endoderm and then migrate through the mesenteries
during development into the indifferent gonad (b) the germ cells arrange themselves between the
medullary and cortical cells of the ind~fferentgonad (c) the cortical tissues predominate in the
formation of the ovary while (d) the medullary tissues predominates i n the for motion of testes.

9.9.2 Functions of the Genital System

The primary function of the genital system is production of gametes, the sperms and
the eggs respectively by the male and female gonads under the influence of
hypothalamol - pituitary hormones.
Gonoducts of the genital system transmit gametes to the place most suitable for
fertilisation. It may be water in which eggs are laid by the female and sperms
dropped on them by the male (external fertilisation) or sperms may be discharged
just as far into the female tract where egg and sperm can meet (internal fertilisation).
Transport of sperms into female body is specially performed by accessory
reproductive organs or intromittent organs, to ensure fertilisation.
The female gonoduct is different from that of the male. It is specialised for retention
of: (i) egg (in case of egg laying or oviparous organisms), e.g. fish, amphibia,
reptiles and binds (ii) the fertilised egg with the developing embryo (in case of
ovovivipardus organisms which lay the egg after embryo has developed to a cemin
stage) e.g. certain snakes or (iii) the developing embryo (in case of viviparous
organisms which deliver full fledged young ones) e.g. mammals.
 llrinogenital System
Gonads secrete sex hormones which are steriod in chemical nature. The male hormone
secreted by the testis is testosterone. The female hormones are malnly estrogen and
progesterone. The sex hormones control the production of secondary sex characters,
difference in structure, physiology and behaviour by which male and female can be
distinguished. The secondary sex characters are especially prominent during breeding season.

SAQ 5
Choose the correct alternatives from the parenthesis.
i) Reproductive organs develop from the embryonic (mesodennleaoderm) germ layer.
ii) The finger like process developing from germinal epithelium are called
(primarylsecondary) sex cords.
iii) Organism that lay eggs in which the embryo develops are called (viviparous1oviparous).
iv) Mammals are (ovoviviparous1viviparous).
v) Estrogen is a (malelfemale) hormone.

9.9.3 Genital System of Protochordates

Branchiostorna is a dioecious chordate. The gonads of both male and female are similar
in arrangement, with distinctly segmented gonads (i.e. is many gonads) that are arranged
below the muscle segmenrs or myomeres (Fig. 9.17 a). Approximately 26 pairs of gonads
on either side of the atrium also called peribranchial space project into it from the inner
surface of the body wall. The most anterior gonads are located around the middle of the
pharyngeal region. Each gonad (or segment) is a hollow sac which is lined with an (I)
outer coelomic epithelium and an (2) inner germinal epithelium. Gametes develop within
the cavity of the gonad and are expelled through the wall into the artrial cavity by means
of temporary pores. From this cavity the gametes pass directly to the outside through the
atriopore. In these animals fertilisation is external. Overall, the reproductive organs of the
Branchiostonta are not similar to those of vertebrates, The ovarian follicle also appears to
differ from the vertebrates because a follicular epithelium is absent.

Urochordates (Tunicates) are he~maphrodites,that is, they have both sex organs in the
same individual. In other words they are hermaphrodite. Ducts from gonads enter the
atrium, a space surrounding the viscera (organs of the body) and then to the outside by the
atriopore (Fig. 9.17 b).
I * w wn11
~ I ~ Uhwn
I of smvr

Ntwlurrd

Ihwnl nonu

IXnsul ruhmie wnal

t;cladrmral
rfilhcrllum

Pnnr?nr

tnmrrerrr
mu.**

I t~bnn
n AlfNn
Fig. 0.17: Genital systclns ol' protochordates (a) cross section ol'Brmrrltic~stomo, showing segmental
gonads (b) saggital section of a tunicste showing its reproductive systems.

I. Paired gonads -testes (singular - testis)

2. Paired urinogenital ducts
I
Functional Anatomy of
-
Chordates 11
3. Single urinogenital opening
Let us begin our study of the male genital system with the anatomy of the testes.

9.10.1 Testes
Testes of all vertebrates have similar construction. The typical testis is a compact organ
whose shape however varies in members of different vertebrate classes. In some forms,
testis is composed of: (i) elongated tubules called the seminiferous tubules in which the
primordial g e m ce~llsdevelop into mature sperm and which connect by means of ducts to
the outside e.g. anulran amphibians and amniotes. (ii) in others, the testes consists of
rounded cavities called seminiferous ampullae or spermatic ampullae or spermatic cysts -
e.g. cyclostomes, fish and urodeles. Both the rounded ampullae and elongated tubules, at
first consist of solid masses of cells which later develop the cavities or lumina.

Seminiferous Tubules
The testes in anurans, amniotes (reptiles, birds and mammals) and even some teleosts, are
composed largely ofiseminiferous tubules which are coiled tubes and whose walls contain
cells that produce sperm (Fig. 9.18 a).
I

Fig. 9.18: nlwmmnlian testis (a) section through testis show~ngspcrm passage (b) nn enlargement of the
boxed nrer in (a) showing the details of seminiferous tubules and rete testis (c) section of kidne)
showing seminiferolds tubules and a group of Leydig cells between the tubules (d) enlargement
of the boxed area in kc) showing dcvc\oping sperms and sertoli cell.
 Ilrinogenital System
The testes are surrounded by a capsule, the tunica albuginae. Seminiferoustubules may
constitute upto 90 per cent of the testis. The tubule walls consist of a multilayered
germinal epithelium containing spermatogonic cells and Sertoli cells which are nutritive
in function and in which the heads of the maturing sperms are embedded. Sertoli cells
probably also produce, most of the fluid in which the sperms are suspended, while
leaving the testis by means of active filteration from blood plasma (Fig. 9.17 b).
Seminiferous tubules may end blind at the tunic or outermost tissue layer, and pass
toward the centre, becoming tortuous, that is, full of twists and bends, before emptying
into a system of collecting tubule, the rete testis. Such an arrangement is characteristic of
frogs. In certain amniotes like the rat for example the tubules may be open ended, running
a zig-zag course from the rete to the periphery and back again. The average length of such
tubules is 30 cms, and they seldom communicate with each other. In many mammals, the
tubules are grouped into lobules separated by connective-tissue septa, or walls. This
arrangement allows the packing in of an extensive quantity of germinal epithelium into a
small space (Fig. 9.18 a). The tubules are inconspicuous and epithelium is inactive in
immature males and between breeding season in the adult males of those species which
have specific breeding seasons.

In some species however, spermatogenesis or production of sperms in adult males

proceed at a variable pace throughout the year. An active epithelium may exhibit all
stages of developing sperms. The lumen or tubule cavity contains the tails of many
sperms whose heads are embedded in thc Sertoli cells, free sperms and fluid that is
resorbed. In mammals in any single zone along a tubule, all sperm are at the same stage
of maturation; adjacent zones contain different generation of sperms and a period of
sperm formation and discharge is followed by interval of inactivity.

Seminiferous Ampullae
Cyclostomes, most fishes and tailed amphibian have seminiferous cysts also called
spermatogonial cysts, spermatocysts or sperm follicles or ampullae or crypts or sacs
acini or capsules in which sperms develop but in which the epithelium is not germinal.
Sperms mature within the ampullae, among cells called the Sertoli cells. The Sertoli cells
appear to be partially nutritive in function. Once the sperms mature, the walls of the
ampullae break down and release the sperm into the coelomic cavity. The arrangement of
the germinal epithelium is different from that of seminiferous tubules. Sperlnatogenic
cells migrate into the cysts from a permanent germinal layer, which dependins on the
species may be among cysts at the periphery of the testes or in a ridge along one margin
of the testis. The spermatogenic cells after invading the thin, non- germinal epithelium of
the cells, multiply producing a large number of sperms. The cysts become swollen and
whitish in colour and the entire testis swells up as well becoming granular in appearance.
When sperms mature they separate from the epithelium and move freely in the cystic
fluid. Finally the cysts burst and the sperms are shed into the duct. In the case of
cyclostomes and a few teleosts the sperms are released into the coelom. The cysts
collapse on becoming totally empty. They are then either replaced by new ones or
become repopulated by additional spermatogenic cells.

The spaces between the seminiferous tubules or spermatogenic cysts in the testes are
filled with testicular stroma which consist chiefly of connective tissue, blood, lymphatic
vessels and nerves. Stroma is more abundant in some vertebrates than in other. Glandular
interstitial cells called Leydig cells are also present in most if not all vertebrates. The
Leydig cells are thought to be a primary source of androgens or male hormones. These
cells are not always easily distinguishable. The capillar%wystemof the rat testis and
probably that of many other vertebrates is such that blood which bathes the Leydig cells
flows to the tubules. In most vertebrates however the adult gonads retain a position in the
upper part of the coelomic cavity. In vertebrates except mammal the testis lie within the
body. This is also the case in many and sometimes in all members of the mammalian
orders Monotremata, Insectivora, Hyracoidea, Edentata, Sirenia, Cetaceae and
Proboscidiea.

In some male mammals like in most marsupials, ungulates, carnivores and primates after
infancy the testes descend into special pouch called scrotum where they are lodged
permanently. The scrotum or scrotal sac is a temperature regulating device. The scrotal
sacs are paired structures which project externally from the floor of the abdominal cavity,
each connected to the abdominal cavity by a inguinal canal lined with peritoneal
membrane. During development the testis move backward and downward from their
Functional Anatomy of original position into these sacs each accompanied by its duct, blood vessels, lymphatic
Chordates - 11
vessels and by a fold of its proper mesentery called gubernaculum. A few mammals
have a pouch into which the testes descend and from which they can be retracted by the
action of the musclie (cremaster) in the scrotal sac. 'Ihese include a few rodents such as
ground squirrels, most bats and some primitive primates (loris, potto). In some cases the
sacs remain in open connection with the abdominal cavity, and the testes may be
withdrawn into the body between breeding season through the iguinal canal which acts as
path of descent and retraction of the testes to the sac. In descending, the testes carry along
a spermatic duct, blood and lymphatic vessels and a nerve supply wrapped in peritoneum
which collectively constitute the spermatic cords. In rabbits, most rodents and some
insectivores the scrotal sacs are absent instead they have a wide inguinal canal into which
the testes may be withdrawn and from which they are retracted when in danger of injury.
In these mammals, the descended testes cause a temporary bulge in the perineal region
(i.e. between the anus and urinogenital opening). In a small number of mammals the
testes permanently occupy the perineal location. In some mammals including man (Fig.
9.4 a) the sacs may be permanently closed off, but sometimes a weak spot in the
abdominal wall of this region may rupture leading to a condition known as iguinal hernia.

9.10.2 Male Genital Duct

The male gonoducts or genital ducts which in most vertebrates serve to transport
spermatozoa to the outside of the body are the archinephric ducts or the Wolffian ducts,
that are formed in connection with the developing kidneys. You will recall that the name
archinephric duct is used for the kidney duct in anamniotes. Although the male genital
ducts are similar to the female genital ducts, the male genital ducts have a more complex
history and organisation.

The original function of these ducts as you will recall, is elimination of urinary wastes. In
a number of fishes ~ n amphibians
d certain modified kidney tubules are employed in
carrying spermatozoa from the testis to the archinephric duct. 'They are known as efferent
ductules and the archinephric duct is then termed as the ductus deferens. Even in the
amniotes in which the mesonephros degenerates, its duct persists to become the ductus
deferens and epididymis (ductuh epididymidis). This ductus epididymidis establishes
connections with the testis via effh-ent ducts or ductules (Refer Fig. 9.18 a) which in
this case are the modified and persistent mesonephric tubules. Reproductive ducts are
lacking in amphioxlus and modem jawless fish, the cyclostomes.

In modern jawless fish the testicular ampullae rupture to release sperm into the coelom
from where they pass out of the body by means of the genital pores (Fig. 9.19). In most
vertebrates however the sperms never enter the coelom but go directly into genital ducts.
In all jawed vertebrates the testes are joined to one or more ducts called the sperm ducts
through which the sperm leave the body All sperm ducts include parts of the kidney
ducts and so they are called urinogenital ducts. The exception are the teleost fish in which
the ducts are not derived from the kidney but from the testis itself. In most vertebrates the
sperms mature as they pass through the genital ducts but in teleosts they seem to be fully
mature at the time they leave the testis. The genital ducts terminate at the urinogenital
sinus which often empties into the cloaca. Uninogenital ducts may also carry urine from
the adjoining kidney though that part of the kidney stops functioning in some species.
From the testis is formed a tubular network called the rete tubule or rete testis. The rete
testis within the testis forms a network of thin walled ductules or minute ducts that collect
sperms from the seminiferous tubules (Fig. 9.18 a).

The rete is drained by a number of small ducts called the efferent ducts or vasa
efferentia. The vasa efferentia are usually modified kidney tubules and are usually less
than 10 in number. The kidney tubules or vasa efferentia drain into the archinephric or
mesonephric duct aalled the deferent duct in male which empties into the urinogenital
sinus. The efferent and deferent ducts are collectively termed as epididymis and may be
used primarily for sperm transport (Fig. 9.18 a).

The epididymis of amniotes which is a highly coiled duct that drains the vasa efferentia
usually serves as a temporary storage place for sperm. In mammals the first part of the
epididymis consists of a head, body and tail that wrap around the testis and then gradually
straightens out to become the spermatic duct. The epididymis secretes substances that
prolong the life of the stored sperm and increase their capacity for motility.
 Urinogcnitwl Systcn~
9.10.3 Male Accessory Sex Glands
Vertebrates males, especially of terrestrial species may have several accessory sex glands.
Some amphibians (salamanders) have cloacal and pelvic glands which secrete a jelly
package to enclose sperm thus forming the spermatophore. They may also have other
functions as well. In mammals a number of relatively large and complex accessory glands
occur. The testis and epididymis secrete fluids that form part of the semen. Other glands
are the prost ate gland. vesicular glands, bulbourethral glands, urethral glands (or Littre's
gland) and coagulating glands. All of these open into the urethral part of the urinogenital
sinus. Not all the glands are present in every species. However, the major mammalian sex
glands are the prostrate, the bulbourethral and the ampullary glands and the seminal
vesicles all of which are the outgrowths of the spermatic duct or of the urethra and all
four occur in elephants and horses and in most moles, bats, rodents, rabbits, cattle and
primates.

The prost ate is the ~iiostwidely distributed mammalian accessory sex gland. It empties
into the urethra by multiple ducts. Many rodents, insectivores and lagomorphs have three
separate prostatic lobes. In a few mammals which include some carnivores and primates,
the prostrate is a single mass with lobules and encircles the urethra at the base of tlie
bladder. In many rodents as well as in some other mammals, the semen coagulates
quickly after ejaculation due to secretion from a male coagulating gland which is usually
considered part of the prostatic mass. Coagulated semen forms a vaginal plug that
temporarily prevents copulation.

Balbourethral or Cowper's glands arise from the urethra near the penis and are
surrounded by the muscle of tlie urethra or penis. Usually there is one pair of Cowper's
gland in the mammals, except for in some marsupials where as many as 3 pairs have been -
found.

An ampullary swelling on the spermatic duct near the urethra is present in many
mammals. However in a small number of mammals a separate ampullary gland is formed
as an outgrowth of the duct.

Seminal vesicles are paired, typically elongated and coiled fibromuscular sacs that empty
into either the spermatic duct or the urethra. These vesicles contribute the sugar fructose
and citric acid to the semen but do not serve as sperm reservoir.

9.10.4 Intromittent Organs

In most aquatic forms external fertilisation takes place and water provides the medium by
which the spermatozoa reach the eggs. In terrestrial forms, however, a liquid environment
is needed to transport the spermatozoans so internal fertilisation is the rule. The necessary
fluids are produced by both male and female. In a number of terrestrial vertebrates
spermatozoa are transferred from male to female by cloacal apposition but in most
terrestrial forms, and even in many aquatic species where fertilisation is internal the
males develop. organs called intromittent or copulatory organs which are used by them
during internal fertiliation to introduce sperms suspended in seminal fluid into the female
tract.

A number of types of copulatory organs exists among vertebrates all of which are not
homologous.

lntromittent organ in Anamniotes .

Fish - Among fishes copulation with internal fertilisation occurs only in elasmobranchs,
holocephalians and some teleosts. In elasmobranchs copulation is accomplished by means
of clasping organs called claspers (Fig. 9.19 a) which are modifications of the medial
portions of the pelvic fins of males. During copulation one clasper is inserted into the
female cloaca. Sperms leave the male cloaca, enter a groove on the clasper and are
flushed by water squirted from siphon sacs within the body wall of the male into the
female cloaca. In the teleosts which have internal fertilisation the anterior border of the
anal fin of the male may be elongated posteriorly to form an intromittent organ called the
gonopodium (Fig. 9.19 b).
Functional Anatomy of
-
Chordates II

Fig. 9.19: Intromittent organs of Ashes (a) clasper, u modified pelvic fin of male elasmobranch Squalus
acanthias (dbgfish) and (b) gonopodium, which is a modification of the anterior part o f the anal
fin in males, of those teleosts. who exhibit internal fertilisation.

Amphibians - Copulatory organs are absent in urodeles and anurans though internal
fertilisation does oacur in urodeles. In these animals the male deposits spermatophores,
which are actually small packets of spermatozoa held together by secretions of cloacal
glands. The female picks up the spermatophore by the muscular movements of the cloacal
lips. A dorsal diverticulum of the cloaca, the spermatheca serves for storage of the
spermatozoa which are thus available for fertilising the ova as they pass down the
oviducts to the cloaca.

lntromittent organs of Amniotes

lntromittent organs are very well developed in reptiles and mammals and few birds like
the ostrich, drakes and ganders. In the amniotes intromittent organs are of two kinds:
(1) Hemipenes are paired, saclike organs devoid of erectile tissues lying in pockets
under the skin near the cloaca. Hemipenes can be everted or retracted eg. Snakes and
lizards (Fig. 9.20).
(2) Penis is an unpaired erectile organ, eg. male turtles, crocodiles, few birds and
mammals (Fig. 9.21 and 9.22).

Reptiles: The only reptile lacking copulatory organs is Sphenodon. In other reptiles as
you have learnt, two types of structures occur. In snakes lizards the hemipenes is present.
They are everted during copulation by propulsor and retractor muscles and filling of
blood sinuses in the hemipenes (Fig. 9.20).

In turtles and crocodiles, the single penis is derived from paired thickenings or ridges in
the anterior and venltral walls of the cloaca and is composed of connective and erectile
tissues. The paired masses of erectile tissue are called corpora cavernosa. A groove
along the dorsal surface serves as a passage for the spermatozoa. During the mating act,
the corpora cavernoha are filled and distended with blood, making the penis firm and
enlarged and erect. The penis can be extruded and retracted.

Fig. 9.20: lleniipenes of nlrlc reptile (lizard).

Birds: A penis occurs only in the males of ducks, geese, swans and ostriches. It is a
single structure, buillt on the same plan as'that of turtles and crocodilians. In the remaining
birds sperms are transmitted through cloaca (Refer unit 16 of LSE-10).

Mammals: A single penis is typical of mammals. In monotremes, under normal

conditions, the penis lies in the cloacal floor. It is similar to the organs of turtles,
crocodilians and bird except that the groove on the dorsal side becomes a closed tube.
 tlrinogenital System
In addition the tube is surrounded by a single mass of erectile tissue the corpus
spongiosum which is separate from another pair of erectile tissue mass called the corpora
cavernosa. The canal in monotremes is supposed to carry only spermatozoa since the
urethra has a separate opening into the cloaca (Fig. 9.2 1).

Fig. 9.21: Diagrammatic longitudinal section through cloacal region of male monotreme showing a
retracted penis.

In the rest of the male mammals the openings are not separate (Fig. 9.22). The urethra in
these animals and subsequent groups serves as a passage for both urine and seminal fluid.
The two corpora cavernosa are separated by a septum and the corpus spongiosum
surrounds the urethra. The end or tip of the penis is enlarged to form a sensitive swollen
glass, containing erectile tissue and numerous nerve endings which make it extremely
sensitive to certain stimuli. It is is continuous with the corpus spongiosum. The glans is
covered by a thin and delicate skin called foreskin or prepuce.

Fig. 9.22: Sag$:.-.! oicw of pelvis of human male showing the urinogenital system and interior of testis and
its duct.

. .
SAQ 6
Correctthe following sentences if necessary.
i) In cyclostomes, sperms develops in seminiferous tubules within the testis.
ii) Leydig cells secrete the hormone estrogen.
iii) The Inale genital ducts are called fallopian ducts.
iv) The accessory sex glands namely prostrate, Cowper's gland, Bulboutheral gland are
found in female vertebrates.
v) The intromittent organ in lizard is called gonopodium.
Functional A ~ ~ a t o ~ofn y
Chordates - I1

The genital system in vertebrate female consist of:

I. Paired gonads - ovaries (sometimes one due to degeneration)
2. Paired oviducts
3. Single female genital opening
(Refer to Fig. 9.23 a and b)

9.11.1 Ovary
,411ovarian Ibllicle consisls o f an oocyt Ovaries lie within the body cavity and are suspended by the dorsal mesentry, the
or immature egg enclosed in a mesovarium, through which pass blood and lytnph vessels and nerves. Primitive
epithelium. The cells ol~epitheliu~n &\re , vertebrate ovaries are found in hagfish, in which a mesentry-like fold of gonadal tissue
referred to variously as follicular cell or
nurse ccll or granulose cells. In
stretches across nearly the entire length of the body cavity. In the hagfish, the unique
cyclostomes. teleosts and amphibians feature is that the fuhctional ovarian tissue occupies only the forward half of the-gonadal
the epithelium is one layer thick. In the mass while the rear part contains the rudimentary testicular tissue. In most fishes except
hagfish and in those vertehrates like in very primitive forms, the ovaries are similarly elongated. In tetrapods except for
elasmohranches. reptiles. birds and
monotremes in which the oocytcs have
mammals the ovaries are usually confined to the middle third or h6lf of the body cavity.
heavy yolk deposit. the epithelium particularly during the non breeding seasons. The ovaries of mammals undergl.
appears to be two cells thick. In moderately caudal displacement in order to be located between the kidney and the pelvis
mammals above thc level n f (Fig. 9.23 a).
oionotremes. the tollicular epithelium
appears to be many ccll thick.
The shape of an ovary depends on many factors like whether one egg or thousands are
discharged (ovulated), whether the eggs are immature or ripe; whether mature eggs are
small or large, or whether pigments occur in the egg cytoplasm, such as those responsible
for yellow yolk. The appearance of the ovary is also affected by other factors such as the
season of the year in seasonal breeders. (as ovary enlarges during the breeding seasons
and diminishes tn size between seasons) the age of the animal (whether juvenile or
reproductively active or senile especially in birds and niammals) and the fate of the
ovulated egg follicles or sacs.

Fig. 9.23: Mammalian female reproductive s)'stcni. (a) saggital view of pelvis of human fcmales 'showing
female reproduktive system. (b) cut away of ovary showing internal structures.

Ovaries are characterised as saccular or hollow or lacunate (i.e. compartmented) or

cotlipact structures. However, most ovaries are similar in construction. They are covered
with a germinal epithelium that is'continuous with the peritoneum lining the body
cavity. Beneath the epithelium the ovary has a layer of connective tissue called tunica
albuginea which is considerably thinner than that surrounding the testis. Below this the
ovary has an external cortex and internal medulla. The cortex which is the thick outer
layer lies immediately internal to tunica albuginea and contains future eggs and at one
time or another, eggs in ovarian follicles (developing eggs) (Fig. 9.23 b). The cortex
also contains remnants o f ovulated follicles and in mammals. and clusters of interstitial
cells that in some species are glandular. The cortical components are embedded in a
supportive frame work o f connective, vascular and neural tissue that form the stroma.
Internal to the cortex is the vascular, meso-dermal medulla which consists o f blood and
lymph vessels, nerves and connective tissue. 'The medulla lacks germinal elements and
exhibits no significant cyclical activity. It is usually inconspicuous and is continuous
with the dorsal mesentry. In the cyclostomes the medulla and dorsal mesentry are
indistinguishable from each other. On the contrary in mammals the medulla i s almost
completely surrounded by the cortex and converges on the mesovarium at a narrow
hilus, at which nerves and vessels enter the ovary. In the medulla o f the mammalian
ovary near the hilus arc small masses o f blind tubules or solid cords called the rete
ovari. The 'rete ovari' are homologous (of the same embryonic origin) with rete testis
in the male. The rudimentary right ovary o f the birds usually consists o f only medullary
tissue.

9.11.2 Female Genital Ducts

Eggs are released into the coelo~niccavity. Once in the coelom, the mature eggs enter the
female genital duct called oviduct which parallels the archinephric duct in its embryonic
course. The oviduct usually joins the urinogenital sinus near the cloaca. Oviducts except
in teleosts and some fishes are modifications o f the mullerian ducts and develop in every
male or female embryo (except in cyclostomes) as a pair o f longitudinal ducts. In males,
the Mullerian duct disappears or becomes rudimentary. In females, it grows larger to
become the reproductive tract or gonoduct. Its smooth muscles and cilia o f the ciliated
cells, in its lining propel eggs along the tract.

In most teleost fishes with tubular ovaries, eggs do not enter the coelom at large but
go directly into a special genital duct called gonaduct which is named so, because it is
apparently derived directly from the gonad. The gonaduct encloses a small part o f the
main coelom (Fig. 9.24). In higher mammals the oviduct differentiates into three
regions (i) Fallopian tube (ii) uterus and (iii) vagina. The oviduct o f most other
vertebrates is open at both ends, at the urinogenital sinus which may open into the
cloaca and near the ovary by means o f infundibulum which is also called pre
ampulla. The end next to the ovary is ringed with mobile, finger like ciliated
projections called fimbriae that actively envelops the ovary near the time o f ovulation
(see Fig. 9.24).

Oviducts may be long or short and may have glandular portions that are modified to
be secretory in function. As a result along the course, the oviducts may differentiate
into (i) a region that provide protective and nutrient ~naterialon the eggs, (ii) uterus
(pl: uteri) to lodge the developing embryo in case o f viviparous animals. (iii) the
terminal segment o f the female genital duct which i s modified to receive the
intromittent organ.

External cervical os

Fig. 9.24: Thc rcproductivc system of human female showing oviduct. ovary, uterus and vagina.
Ful~ctiona!\ I : I ! I J , : I ~ .,f'

Chordates 11 -
Uterus is the muscular middle part of the oviduct (Refer to Fig. 9.23 a and Fig. 9.24
again). Its muscles form the myometrium and its inner lining is called endometrium.
The endometrium becomes highly vascularised before the blastocyst stage (or developing
embryo) implants.

In all mammals the uterus narrows to form the posterior cervix (Fig. 9.23 and Fig. 9.24).
Lips of the cervix enclose the uteri opening or '0s uteri' through which sperms rise upto
the upper part of oviduct for fertilisation. The cervix dilates during delivery of the baby.

Uterus shows a num~berof variations in mammals. In primitive mammals like

monotremes (egg laying mammals) and marsupials there are two uteri (duplex uterus).
In most mammals hbwever, the distal parts of the two uteri are fused together to give a
bipartite or bicornuate uterus. In higher primates there is a complete fusion of the two
uteri to form a single simplex uterus (Fig. 9.25).

Duplex Uterus
-
Bipartite uterus

Bicomuate utcrnus Simplex uterus

Fig. 9.25: Diagram showing the fusion of posterior ends of paired uteri in females of placental n~ammals.
(The uterus And part of the vagina have been cut open.)

Vagina
'The uterus leads through the narrow cervix into the vagina. Vagina is the fused terminal
part of the Mullarian duct. It opens into urinogenital sinus (Fig. 9.22 ) or is extended to
open directly to theoutside. It is a muscular distensible tube which receives the penis
during mating. It has convolutions on it which are called vaginal rugae. Vagina opens to
the exterior by the female genital opening.

Vagina is absent in egg laying mammals. Marsupials, have paired vagina that opens into
urinogenital sinus. To match the two vaginae, male penis is forked at the tip and one tip
enters one lateral vaginal canal to discharge semen.

9.1 1.3 Female Accessory Glands

You have already read in subsection 9.10.3 about the various accessory glands associated
with the male genital system. You will similarly find that the accessory glands are also'
associated with the female genital system of many vertebrates. Oviducts of many female
vertebrates havcglands in the oviduct called the albumin gland that coat the egg with
albumin. Other glands associatea with the oviduct are the shell gland or nidamental gland
that cover the egg with shell material; mucous cell or oviducal tubular glands that secrete
a jelly like material. Amniotes that lay large eggs may have in the oviduct mucous
secreting glands cal~ledvaginal mucous gland that coat the egg prior to expulsion,
possibly to lubricate it. Some fishes have adhesive glands that coat the eggs with a sticky
secretion so that the egg scan adhere together or to appropriate objects. In some
vertebrates that retain the developing embryos within the body, special glands of the
oviduct evolve in order to nourish the young. These glands secrete into the oviduct so that
young present there absorb or ingest the secretion. This secretion is called uterine milk or
en1bryotrophe.

9.1 1.4 External Female Genitalia

In females the external genitalia in comparison to males is feebly developed. Maximum
developme~ito f female genitalia is in tlie Order Primates o f mammals. In liu~nanfemales
it is well developed and is described below:

External Genitalia o f Human Female

The external genital organs o f the female are termed vulva (Refer to Fig. 9.24). The
internal genitalia o f the female consists o f an outermost structure called labia majora
which are a pair o f skin folds and contain adipose tissue. Within the cleft formed by these
folds are the labia minora, which are a smaller pair o f skin folds that are highly
vascularised but have no fatty tissue. At the anterior end o f the vulva, these two interior
skin folds partly enclose tlie clitoris, a small organ for sexual stimulation. The opening o f
the urethra is about midway between the clitoris and vaginal opening. The vaginal
opening is located behind the urinary meatues and is much larger than the urinary
opening. I t is covered by a thin mucous membrane the hymen.

Bartholin's glands (or bulbovestibular glands) or greater vestibular glands are two
bean shaped glands, one on either side o f the vaginal opening. These secrete a lubricating
fluid. The two glands open by a single duct between hymen and labia minora. Bartholin
glands are homologous to male bulbourethral glands. A group o f tiny mucous glands,
the lesser vestibular glands also called Skenes glands open into the vestibule..

9.1 1.5 Mammary Apparatus

Mammals are named so because o f the characteristic presence o f the mammary apparatus,
which includes (i) mammary glands, (ii) elevated nipples which are the outlets for
secretion o f these glands and (iii) breasts o r mammae which are the integumentary
.swellings due to localised presence o f these glands (Fig. 9.26).

Mammary glands are modified sweat glands. They are present in both male and female
mammals but they become well developed only in females as their development is
controlled during puberty by the ovarian hormones estrogen and progesterone.

Each mammary gland is divided into a number o f lobes and each lobe has several
lobules. Lobules are formed o f connective tissue in which secretory cells called alveoli
are embedded. Alveoli are arranged in grape like clusters around minute ducts.
Progesterone stimulates the growth o f alveoli and estrogen stimulates growth o f ducts.

, Nipple

Fig. 9.26: Human female hreast showing mammary apparatus.

Ducts from lobules units to fonn a single lactiferous or milk carrying duct per lobe. Each
duct opens by a pore at the nipple. The nipple is bordered by a pigmented area called
areola containing several sebaceous glands wh,ich appear as small nodules underthe
skin. The nipples vary in number and location'in different mammals, The number
Functional Anatonly of depends on the nunbber of offsprings born in a litter. The position of mammae depends on
Cl~ordates- I 1
their availability to the suckling offspring. In monkeys and other primates which are
arboreal, the mamlnae are pectoral. Humans, who have descended from arboreal
ancestors also have pectoral nipples. In ruminants which suckle their young, while
standing, the mammae are elongated and project downwards. In pigs they are arranged on
the sides and so the mother lies down to suckle the young. The number of nipples vary
Horse, bats, whales and humans have a pair of nipple.

9.12 SURVEY OF GONADS IN VERTEBRATES

Jawless vertebrates: In the jawless fishes the gonads begin as paired structures but fuse
later in development into a medial gonad. Sperms forni in the ampullae rather than in
seminiferous tubulbs. In both male and female forms, special genital ducts are absent. The
gametes that are reileased from the gonads pass through the coelomic cavity and out of the
body via genital pores. The lack of genital ducts appears to be a primitive vertebrate
feature. The complete separation of the anus from the [Link] appears to be a
specialisation.

Jawed vertebrates
Cartilaginous fish -Gonad structure, origin, as well as pattern of genital ducts of ..
cartilaginous fishes are typical for jawed vertebrates in general (Fig. 9.27 a and b) and
indicate a common origin of vertebrate genital ducts from archinephric tubules. The
Mullerian duct originates as a new structure in jawed vertebrates. The testes are of
primitive arnpullaty type. Modifications for internal fertilisation include pelvic fins called
claspers (Refer subsection 9.10.4). The many variations on internal development among
cartilaginous fishes indicate no common pattern or clearly defined relationship to other
fishes or to tetrapods.

Fig. 9.27: Reproductive system of cartilaginons fishes (a) male urinogeoital system of a shark (b) female
reproduttive system of shark Squabrs. The left ovary has been removed.

Bony fishes: Mlost extant teleosts have specialised hollow ovaries and testes with genital
ducts called gonoducts that are derived from the gonads (Fig. 9.28 a and b). However
some bony fishes similar to jawless fish have coelomic transport. In them. extensions of
the gonads in the adult produce hollow testes or ovaries that continue as tubes to transport
the egg or sperm respectively to exit from pores through the posterior body wall. The
testes may be (i) ampullary (or acinar) as in jawless fish and sharks or (ii) tubular as in
most tetrapods. Tubular testes and the genital ducts associated with the kidneys seem to
be primitive features of bony fishes.
 IJrinogenitaI System
Kidney

Testis

Sperm duct

Archinephric duct

Urinary bladder

Urogenital papilla

Fig. 9.28: Reproductive s y s t e l ~o~f bony fial~cs(a) 111aleu r i ~ ~ o g c l ~ isystenls

tal o f tnale sea Il~)rsc,
Hippocnntp~rs(b) firmale reproductive system o f holly fist1 Anrict.

Amphibians: In amphibians the gonads and urinogenital ducts are basically like those of
primitive bony fishes and tetrapods (Fig. 9.29). Some oviparous and all viviparous forms
have developed internal fertilisation which may have originally been an adaptation for a
terrestrial mode of life. Caecilians appear to be the most of terrestrial of the amphibians in
terms of reproductive adaptation. They all have internal fertilisation and appear to have
developed mechanisms to protect the egg, by brooding, egg retention or more colnplete
viviparity. Living viviparous amphibians include several species of salamander,
approximately five species of frogs and about 20 species of caecilians. The oviducts of
females of some viviparous species of frogs, salamanders and caecilian produce nutrient
secretions called uterine milk for the young. No placenta is however known to develop
within the oviduct.

Bidder's d
\ Fat bodv

''u Cloaca

Fig. 9.29: ReyroQuctive s y s t e ~o ~

f a~~ o p l ~ i b i a n(a)
s male urisogeaital orgalls ol' load Blrfo anrericctnus,
showiag bidder's calla1 (b) female uril~ogenitalsystem of toad &fi, with right ovary
removed.

Reptiles: In reptiles the ~nesonephrictubules are taken over entirely by the testes for
sperm transport (Fig. 9.30 a and b). Urine is then carried across by a new duct, the ureter.
The oviducts in the females, similar to that of amphibia and most fishes are also not
involved with kidney or more appropriately excretory function. The intromittent organs,
the hemipenis and penis of the reptiles indicate a clear adaptation towards terrestrial
mode of life and hence internal fertilisation that has been phylogenetically continued into
mammalian descendants of reptiles. In some species the oviducts of female may retain
living sperm for a long period of time after copulation, upto four years in the case of some
turtles.
Functional Allatom) of
Chordates - I1

Fig. 9.30: Reproduclive systems o f reptiles (a) urinogenital system of the n ~ a l elizard Culores (b) female
urinoge~~ital
system of lizard, Calotrs.

Birds: Birds typically have reptilian reproductive organs except that in females only the left
gonad develops in to an ovary (Fig. 9.27 c). The right gonad develops upto a certain stage,
after which it regresses and remains undifferentiated. A penis occurs only in primitive birds.

Fig. 9.31: Reproduttive system ofAves (a) male urinogenital system of pigeon, Colrmrba (b) female
uri~~ogenlital
system of Colrmtba.

'The oviduct at its posterior end has pockets called sperm nests that store sperm. As the
egg leaves the ovary, it enters the infundibulum where it is fertilised. The fertilised egg is
forced along the oviduct in which a spiral band of material called chalazae is added at
each end, of the egg, as are also added thick and thin layers of albumen, shell membranes
and the shell, around the egg.

Mammals
Primitive mammals like the rnonotrernes(egg laying mammals) generally have a reptilian
reproductive apparatus except for the fact that the penis is tubular instead of being grooved
and the lnalnmqry glands are present. Monotreme have a distinct cloaca. The ureter in
lnonotremes similar to reptiles and birds lies between the Wolffian or Mullerian ducts
instead of lying lateral to the reproductive duct as in eutherian mammals. The oviducts of
monotremes are relatively unspecialised as compared to most mammals. They are unfused
and open individually into the urinogenital sinus. The oviducts are specialised to the extent
that they produfe a shell similar to that of reptilian eggs and also secrete uterine milk
called embryotirophe which is absorbed by the embryo and used for nutrition.
The details of the reproductive system of the rest (Fig. 9.32 a and b) of the mammal
groups have been given in various sections of this unit, however a figure of the
reproductive systern of a typical mam~iialis given below.

Kidney

Ureter ' Ampullary gland

Sem~nalvesicle
Coagulat~nggland

Prostate gland
Co\l/per's gland

Urinary bladder Preputial gland

Vas dererens -
Glans 4 \
Testis

Anus -

Fig. 9.32: Reproductive 3ystem of'rnatnnlal (a) tnale ~~l-itlogential

systern ot't~talerat. Rt~llrrs(b) l ' c ~ ~ ~ a l c
urinogrnitsl system of female rat Rollus.

SAQ 7
Fill in the blanks and colnpare your answers with those given at the end of this unit.
i) The four types of ~na~nmalian uteri are ......................., ............................,
.............................and ..........................
ii) The muscle layer of the uterus is called ...............................
iii) The sequence of organs of mammalian female genital system are:
two ovaries 3 - ....................... >- .................... +
.................. >- genital opening.
iv) Caecilians among amphibians have ........................fertilisation.
v) The oviducts of females of some viviparous species of amphibian produce nutrient
secretions called ...............................for the young.
vi) In female birds only the ........................gonad develops into the ovary.
vii) The oviducts of lnonotreme secrete uterine milk called ................................

9.13 SUMMARY
Both the urinary and reproductive organs arise e~nbryologicallyfrom the same or
adjacent tissue and maintain close anatomical and sometimes functional association,
throughout the organise life.
Several types of kidneys are found within the chordate groups.
The organs of excretion of the protochordates Branchisfonza(Cephalochordata) and
Herdnlania (Urochordata) show no relationship to any part of the vertebrate kidney
or other know11fluid regulatory structure.
The excretory organs of the vertebrates consist of paired kidneys and their associated
ducts.
The various types of kidneys that are found in the vertebrates have been derived from
a primitive structure termed as archinephros or holonephros.
The archinephros (holonephros) consisted of paired archinephric ducts which extended
the length of the coelom and were joined by segmentally in arranged tubules, one pair
to each segment. The free end of each tubule opened into the coelom by means of a
ciliated, funnel shaped nephrostome. Each tubule was intimately associated with a
small knot of inter arterial capillaries known as glomerulus. The larval stages of the
hagfish and calcilians exhibit an archinephric condition.
Kidneys of living vertebrates have developed from this primitive plan of the
archinephros type of kidney. The various types of vertebrate kidneys may be
regarded as successive stages that have evolved in a craniocaudal direction from the
original archinephros. They may be clearly observed during the embryonic
development of the amniote vertebrate where there is a succession of three
developmental stages of kidney - pronephros, mesonephros and metanephros. Some
but not all these stages are also seen in other vertebrate groups.
In adult ana~nniotes- cyclostomes, fishes and amphibians, the anterior part of the
primitive archinephros usually becomes modified or degenerates. In the embryo it
appears as a transitory structure called the pronephros. In a few lower vertebrates the
pronephros persists in the adult stage and is called the head kidney. A head kidney is
found in the adult hagfish and in certain teleosts. The part of the anamniote kidney
that remains and forms the adult kidney and is called the opisthonephros. ~his'retains
the archinephric duct but differs from the pronephros in that several kidney tubules
may be present In each segment and the tubules lose their peritoneal connections. In
the opisthonephric kidney there is a general tendency for concentration of kidney
tubules towardsthe posterior end. The anterior end loses its importance as an
excretory organ and in males is the appropriate by the reproductive system, with the
result that the archinephric duct becomes known as ductus deferens.
In arnniotes. of the three types of kidneys - pronephros, mesonephros and
metanephros which appear in craniocaudal direction during embryonic develo ment,
only the metanephros persists to form the adult kidney. The archinephric duct n
amniotes is termed as Wolffian duct.
P
In male amniotes, the Wollfian duct gives rise to the epididymis, ductus deferens and
certain other parts of the reproductive system.
The urinary system of most vertebrates includes two kidneys and two ureters. A
urinary bladder and urethra occur in all mammals and some vertebrates.
Ducts from the kidney lead to cloaca in most forms. In teleost fishes they open
directly to the outside.
In marnmals ducts of the kidney enter the urinary bladder (except in monotremes).
Urinary bladder, may be present as ventral out pocketings of the wall of the cloaca.
In mammals the bladder opens to the outside through a urethra which in males of all
forms except far monotermes is also used by the reproductive system.
Each mammalian kidney as well as that of most vertebrates is composed of a renal
capsule, enclosing a cortex and an internal medulla. Numerous individual tubules
called uriniferous tubules are the basic excretory unit of the kidney which produce
urine. Each uriniferous tubules consists of i) nephron, an excretory unit ii) collecting
tubule.
Each nephron consists of a cup shaped Bowman's or renal capsule, proximal
convulated tubwle (PCT), loop of Henle and distal convoluted tubule (DCT). Blood
vessels associated with the nephron are an afferent arteriole, glomerular capill,aries,
an efferent arteriole, and peritubular capillaries.
Kidneys are the principal organs of excretion and osmoregulation in vertebrates.
The various functions of the urinary system are (i) excretion of metabolic waste
products (ii) regulation of fluid, electrolyte balance and blood pressure (iii) sicretion
of hormone erythropoetin which is associated with production of blood cells in case
of loss of blood or hypoxia.
The kidneys of vertebrates are confronted with two kinds of problems since they
occupy diverse habitats like those in which water (i) may not be available for
example in terrestrial environment or marine environment or (ii) may be in
abundance for instance in fresh water environment. Thus the kidneys and tubules of
vertebrates are adapted according to the environment they inhabit.
The urinogenital systems of various groups also differ to some extent.

:NITAL SYSTEM

The reproductive system of chordates consist of primary sex organs, the gonads
which produce gametes.
The primary sex organs or gonads are testes in male and ovaries in female. The
testes produce the spermatozoa and the ovaries the ova.
 llrinogenitnl System
The vertebrate reproductive system consist o f primary and accessory sex organs.
The accessory sex organs are ducts and glands which provide passage for transport o f
gametes (eggs or spermatozoa to the outside o f the body). The archinephric duct in
anamniotes or the Wolffian duct in amniotes becomes the reproductive duct or ductus
deferens in the male. In females the Mullerian duct forms the oviduct. Ducts are
absent in amphioxus and cyclostomes.
In males, spermatozoa are formed within seminiferous ampullae or tubules in the testes.
Each ductus deferens establishes connections with the testes usually by means o f
epididy~uisand persistent kidney tubules called efferent ductules. In anamniotes the
epididymis and ductus deferens consists o f parts o f the archinephric duct which may
serve in some cases as an excretory duct as well as for passage o f the spermatozoa. In
amniotes l~oweverthe epididynlis and ductus deferens are formed from the persistent
Wolffian duct. which is entirely dissociated from the ureter o f tlle adult metanephric
kidney.
The testes in vertebrates are located in the abdominal cavity, except in a number o f
mammals. I n most mammals they are either temporarily or permanently located in a
scrotum outside the body proper.
I n male vertebrates especially mammals accessory glands are associated with
primary sex organs. These secrete seminal fluid in which spermatozoa are suspended
and which are essential for thc viability of the spermatozoa and for transport through
the reproductive tract both male and female.
Fertilisatio~iin vertebrates nlay be external or internal. Most vertebrates with
internal fertilisation have copulatory organs which are used in transferring
spermatozoa from male to female. In fishes these usually consist o f modified fins.
In snakes and lizards paired hemipenis are present; but in turtles, crocodilians,
certain birds and all mammals. a single penis is used for this purpose. The urethra *
corning from the urinary bladder passes through the penis in all mammals except
in monotremes. It thus serves for passage o f both urinary and seminal fluids.
Eggs develop within ovaries and when fully developed they break out ofthe ovaries
into the coelo~[Link] oviduct generally opens into tile coelom by a funnel-shaped
ostium. The oviducts o f teleosts are derived in a different manner and may not be
homologous with those o f other vertebrates.
In forms below rnammals the paired oviducts are separate and usually open
independently into a cloaca. In most higher mammals a cloaca is absent in the adult
and each oviduct differentiates into three regions (i) fallopian tube (ii) uterus (iii) .
vagina. Various degrees of fusion o f the paired uterus occur in mammals, resulting
in different types o f uteri.
The external genitalia o f females called vulva is feebly developed in co~nparisonto
males. Maximum development o f female genitalia is in the primates.
Accessory glands are associated with the female reproductive and secrete mucous.
The parallel development of male and female reproductive system is very striking.
The various structures in one sex have obvious homologous with those o f the
opposite sex.

9.14 TERMINAL QUESTIONS

I. Why are the urinary and reproductive systems usually studied under a common
urinogenital system?
...................................................................................................
Functional Anatomy of
Chordates - I l
2. Draw a well labelled diagram o f the uriniferous tubule o f mammal.

3. Label the given diagram o f human kidney.

4. Write short notes on (a) kidney blood circulation (b) types o f mammalian uteri.
(a). .................................................. :.............................................

...................................................................................................
(b). ...............................................................................................
...................................................................................................

5. What are intromittent organs? Describe the reptilian intromittent organ.

...................................................................................................
 llrinogc~~ital
System

6. List the functions of the excretory systems.

...................................................................................................

7. Describe the a~nniotetestis.

...................................................................................................
...................................................................................................
...................................................................................................
...................................................................................................
...................................................................................................
...................................................................................................
Functional Xaatoniy ol'
Chordates i l l

9.15 ANSWERS
Self-assessment Questions
1. i) glomerulus; ii) hilum or hilus; iii) renal; iv) ~nesoderrnal

2. i) a) nephron; b) glomerulus; c) loop of Henle d) duct of Belleni

ii) renal corpuscle/glomerulus and Bowman's capsule, proxi~nalconvoluted
tubule, descending limb, Henle's loop, ascending limb, distal convoluted
tubule, collecting tubule.
iii) a, b, c
i v) A B
a) capillaries of efferent arterioles vas a rectae
b) renal interstitium prostaglandin
c) macula densa juxta glomerular apparatus
d) erythropoietin red blood cells

3. a-i; b-i;c-1

4. i) testis ovary; ii) secondary; iii) Wolffian/Mesonepliric; iv) ovaries;

v) cloaca; vi) mesorchium; vii) intromitten.

5. i) mesoderm; ii) primary; iii) oviparous; iv) viviparous; v) female

6. i) In cyclostomes sperms develop in seminiferous ampullae of testis.

ii) Leydig cells secrete the hormone testosterone.
iii) The male genital ducts are called ductus deferens or vas defens.
iv) The accessory sex glands, prostrate glands and bulboutheral glands are found
in male vertebrates.
v) The intromittent organ in lizard is called hemipenes.

7. i) Duplex, bipartite, bicornuate, simplex:

ii) Myometrium;
iii) Two oviducts (fallopian tubes) >- uterus>- vagina;
iv) internal;
v) uterine milk;
vi) left;
vii) embryotrophe

Terminal Questions
1. Through the urinrary and reproductive system have nothing in common functionally;
they are usually studied under a common urinogenital system because both develop
from the some segmental blocks of trunk mesoderm or adjacent tissues and share
many of the ducts.
2. Draw figure on basis of Fig. 9.6 of this unit.
3. Label figure on basis of Fig. 9.9.
4. Refer a) subsection 9.5.2; b) subsection 9.11.2 - uterus.
5. Refer subsection 9.10.4. Intromittent organs of amniotes.
6. Refer Fig. 9.14 (b).
7. Refer section 9.10 testis and seminiferous tubules.