Page 58 

Chapter 5 Urogenital system

Jane Williams

CHAPTER CONTENTS

The urinary system 58
The reproductive system 62

This chapter considers the anatomical features and the physiological control of the urinary system with particular reference to water balance and sodium retention. The anatomy of the male and female reproductive systems is discussed and the mutual components of the urogenital system are identified.

The urinary and reproductive systems are often collectively known as the urogenital system as there are a number of shared components between the two.

THE URINARY SYSTEM

The urinary system consists of:

Kidneys (a pair)
Ureters (a pair)
Bladder
Urethra.

Kidneys

The kidneys filter blood. They lie against the dorsal wall of the abdomen attached by a fibrous covering called the capsule. The right kidney is situated about half its own length in front of the left kidney. The hilus is an indentation through which the renal artery, vein, nerves and ureters enter and leave the kidney. Each kidney has a pad of fat located next to it for protection and as an energy store.

The internal structure of the kidney comprises three distinct areas (Figure 5.1):

1. Cortex (outer)
2. Medulla (middle)
3. Renal pelvis (inner).
image

Figure 5.1 Gross kidney anatomy.

The kidney is surrounded by a dense fibrous connective tissue covering the capsule. The cortex contains the renal corpuscles and convoluted tubules of the nephron. The medulla contains the collecting ducts and pyramids located between them and the loops of Henle of the nephrons. The renal pelvis is a whitish colour and made of dense connective tissue. It is effectively a funnel through which urine collects and drains from the kidney into the ureter.

Functions of the kidneys

Formation (excretion) of urine
Production of renin
Conversion of vitamin D to its active form
Production of erythropoietin
It plays a role in homeostasis (balancing the pH of the body).

The kidneys produce urine from water, salts and wastes that they filter from the blood. By adjusting the amount of water and salts they excrete, the kidneys perform a vital homeostatic service in maintaining the internal chemical balance of the body (Table 5.1). Urine is excreted in a continuous trickle, collecting in the renal pelvis.

Table 5.1 Urine analysis

pH Cat/dog 5–7
Slightly acidic due to carnivorous diet
Horse 7.42–7.45
Slightly alkaline due to diet
Specific gravity A measure of density or concentration of urine
Cat 1.020–1.040
Dog 1.016–1.060
Horse 1.020–1.060
Protein Should not appear in a healthy animal
Can be present if the urinary tract is damaged, or if there is increased permeability of glomerulus, chronic renal disease and cystitis
Blood Not normal
It can be due to physical damage, contaminant (prostate) or because an animal is in season (bitch)
Bile Can be due to liver disease or blockage of bile duct
Glucose Present if the renal threshold is exceeded, e.g. in diabetes mellitus
Ketones Produced when fats are oxidised (used for energy)
Can appear in diabetes mellitus and in malnutrition cases where fat is used by the body as a food source
Deposits Calculi, cells, casts of renal tubules may all be present
  Page 59 

Ureters

From the renal pelvis peristaltic contractions move the urine along the ureters. The ureters connect the kidneys to the urinary bladder; the length of ureters varies depending on the species. The ureters have thick walls of smooth muscle and are lined with mucous membrane and transitional epithelium to allow them to expand as the urine passes. They enter the bladder close to its neck on the dorsal aspect where there is a simple flap valve to prevent backflow of urine.

Bladder

The bladder is located in the lower, anterior portion of the pelvic cavity. It is made up of a double layer of smooth muscle and transitional epithelium (that allows stretch) that continues into the urethra. It is covered with peritoneum and its lining is invaginated (folded upon itself) which allows for great expansion. The trigone is the area between the ureteral openings and the neck of the bladder. As the volume of urine held increases, distension of the muscle wall stimulates nerve endings which in turn send impulses to the brain, producing a feeling of fullness. Impulses may then be sent back to stimulate urination. Urination is also known as micturition. Micturition is defined as the expulsion of urine from the bladder.

Urethra

The urethra leads from the bladder to outside the body in males and to the urethral orifice in females (which opens into the genital tract at the junction of the vagina and the vestibule).

The urethra has two sphincters close to where it leaves the bladder:

1. The internal sphincter, which is under involuntary control
2. The external sphincter, which is under voluntary control.

The urethra is shorter in females than in males. In females it only transports urine but in males it transports semen (sperm plus seminal fluid) and urine. The length of urethra in the male discourages bacterial infection due to its length whereas the female urethra is shorter, leading to a higher incidence of infection. Conversely the length and smaller diameter of the male urethra encourage blockages along its length and particularly at areas where the urethra curves, e.g. the ischial arch. Bladder control depends upon an animal’s learned ability to facilitate or inhibit the reflex action that causes micturition. In dogs and cats this ability is not inherent and requires time to learn.

  Page 60 

Nephron

Each kidney contains millions of nephrons that regulate the composition of the blood and excrete waste. A nephron comprises a renal corpuscle and a renal tubule.

Renal corpuscle

The renal corpuscle is made up of a Bowman’s capsule and a glomerulus
The Bowman’s capsule is a double-walled, hollow cup of cells
The glomerulus is a spherical tuft of capillaries fed by an afferent arteriole and drained by an efferent arteriole
The glomerulus projects into the kidney capsule.

Renal tubule

This consists of three main regions:

1. The proximal convoluted tubule
2. The loop of Henle (which goes from the cortex into the medulla and loops back into the cortex)
3. The distal convoluted tubule.

The proximal convoluted tubule is lined with simple cuboidal or columnar epithelium which has microvilli to increase the surface area for absorption. The loop of Henle is lined with simple squamous epithelium and the distal convoluted tubule is lined with cuboidal epithelium but contains no microvilli. Each distal convoluting tubule empties into a collecting duct, lined with columnar epithelium, with lots of distal tubules which empty into each collecting duct. The collecting ducts drain into the renal pelvis at the apices of the pyramids of the medulla. The calyces are defined as the indentations between the pyramids of the medulla.

Renal filtration and absorption

Figure 5.2 shows absorption in the renal nephron.

image

Figure 5.2 Absorption in the renal nephron. A, active; P, passive; 1, proximal convoluted tubule; 2, loop of Henle; 3, distal convoluted tubule.

An afferent arteriole delivers blood to the glomerulus. It then drains from the glomerulus into a much narrower efferent arteriole, producing a high hydrostatic pressure within the glomerulus. The hormone renin can also be employed by the body to constrict the glomerulus further and increase the pressure
Blood is under high pressure in the glomerulus and under low pressure in the Bowman’s capsule
The result is the passage of fluid or ultrafiltration; the fluid produced is known as the glomerular filtrate or primitive urine and drains into the Bowman’s capsule
The glomerular filtrate consists mainly of plasma minus the larger plasma proteins and blood cells. It contains 99% water and 1% chemicals in solution
The amount of glomerular filtrate produced vastly exceeds how much urine is produced. Therefore the reabsorption system must be highly effective to prevent excessive water loss from the body.

Note: the renal blood supply is via the renal artery and vein which travel past the glomerulus to the kidney.

Reabsorption

Reabsorption is the passage of substances from the lumen of renal tubules into the renal capillaries to return them to the systemic circulation.

Reabsorption occurs in the renal tubule
Substances are reabsorbed by active transport or by passive diffusion
Water is reabsorbed in the proximal convoluted tubule: some reabsorption occurs in the distal convoluted tubule by active transport under the control of antidiuretic hormone (ADH)
Sodium ions are reabsorbed in the proximal convoluted tubule by active transport and the ascending limb of the loop of Henle. A small amount is reabsorbed in the distal convoluted tubule. Potassium ions are secreted into the filtrate in the distal convoluted tubule to replace the reabsorbed sodium ions
Chloride ions are reabsorbed in the proximal convoluted tubules by diffusion as a secondary result of sodium absorption due to ionic attraction
Glucose is reabsorbed in the proximal convoluted tubule when it exceeds the renal threshold. The renal threshold is the ‘normal’ concentration of a certain chemical or substance that should be in the body; once this is exceeded it is excreted. In diabetes mellitus, due to the failure in insulin production, excessive glucose is circulating in the blood and is not converted to glycogen but will be filtered within the glomerular filtrate, thus exceeding the renal threshold, and it is excreted in the urine
Amino acids are reabsorbed in both proximal and distal convoluted tubules and should be completely absorbed back into the body
Any protein that is filtered, which is a small amount, is reabsorbed in the proximal convoluted tubule
A small amount of urea is reabsorbed in the proximal convoluted tubule then secreted into the loop of Henle
Acids and bases are selectively absorbed and secreted, e.g. hydrogen, bicarbonate and ammonia ions to aid the body acid–base balance
  Page 61 
Drugs and toxins are actively secreted into the filtrate to remove them from the body.

Remember – substances that are reabsorbed in the nephron are taken back into the body, usually via the blood, to be utilised; they are normally substances which are useful.

Water reabsorption

The loop of Henle regulates the volume and concentration of urine that will be produced in response to the body’s status
Water is reabsorbed by passive and active transport: passive (osmosis) in the proximal convoluted tubule and active in the distal convoluted tubule
Water is reabsorbed by osmosis in the descending loop of Henle, resulting in very concentrated filtrate in the loop; in the ascending limb the walls are impermeable to water molecules but actively reabsorb sodium ions. As a result of this process water molecules are drawn back into the filtrate. On balance the concentration of the filtrate will be constant at the beginning and end of the loop of Henle but the volume of urine will be reduced. This aids certain species, e.g. gerbils, which have long loops of Henle which help to preserve body water
Water reabsorption in the distal convoluted tubule is under the control of ADH
ADH is secreted from the posterior pituitary gland in response to water balance in the body
The tubules of the distal convoluted tubule are lined with simple epithelial cells and microvilli to increase their surface area
There are also lots of mitochondria to provide the energy required for active transport
  Page 62 
When the body needs to conserve water, ADH is secreted and it acts upon the distal convoluted tubule, increasing its permeability to water and resulting in very concentrated urine
If the body needs to get rid of excess water, ADH production will be reduced, therefore less water will be reabsorbed by the distal convoluted tubule
For example, in diabetes insipidus, the body cannot make sufficient ADH and is therefore unable to concentrate urine regardless of the intake volume.

The distal convoluted tubule can be thought of like a tube containing holes that can change their diameter. Under normal circumstances the holes are small and water molecules cannot fit through them back into the blood. However when times are hard and water is scarce, ADH increases the diameter of the holes and they are now big enough for the water molecules to fit through, increasing the amount of water in the blood stream.

Sodium reabsorption

This is under the control of aldosterone and has a secondary effect on water reabsorption
Aldosterone is produced in the adrenal cortex
When plasma sodium levels fall the kidney produces a substance called renin (note the spelling difference between this renin and rennin, which is produced in the digestive system)
Renin acts on substances in the plasma (angiotensin) that stimulate the renal cortex to increase the secretion of aldosterone
Aldosterone acts on the distal convoluted tubule and increases the rate of absorption of sodium and chloride ions
Angiotensin also increases blood pressure, thereby increasing the glomerular filtration rate and pressure.

Osmoregulation is an important aspect of homeostasis and the control of water and sodium reabsorption in the body plays an important role in the regulation and balance of body water.

Species variation – urinary tract

The urinary system of the bird has a pair of kidneys, a pair of ureters but no bladder; they excrete nitrogenous waste in the form of uric acid and urates suspended in urinary water. The semisolid fluid leaves the ureters and enters the urodeum of the cloaca, then returns to the colon for more water reabsorption to occur before excretion. Reptile kidneys do not possess loops of Henle and therefore produce very dilute urine; snakes also do not have a bladder.

THE REPRODUCTIVE SYSTEM

Many generalisations can be made for sexually reproducing animals even though the details of the reproductive process vary with species.

The basic components of the male reproductive system are:

Male gonads or testes which produce spermatozoa
Transport of sperm to exterior of body via the sperm duct
A specialised area of the sperm duct which produces nourishment – the seminal vessels
A copulatory organ into which the sperm duct empties – the penis.

The basic components of the female reproductive system are:

The female gonads or ovaries which produce ova
A tube for transporting ova, called the oviduct
An area for storage of the ova and development of the fertilised egg – the uterus
The terminal portion of the oviduct which opens to the exterior of body – the vagina.

Not all male and female animals possess all the modifications. Some simple animals have complex reproductive systems whereas some complex animals have simple reproductive systems. The reproductive systems of the male and female will be considered individually, taking the dog as an example of a mammal.

Male reproductive system

Figure 5.3 shows the male reproductive tract.

image

Figure 5.3 Male reproductive tract.

(Reproduced from Aspinall V 2006 The Complete Textbook of Veterinary Nursing. Butterworth Heinemann, London, with permission.)

Functions of the male reproductive system include:

Production, storage and nourishment of the spermatozoa (male gamete)
Transport of spermatozoa to the female genital tract for fertilisation to occur.

It comprises the:

Testes
Epididymides
Vasa deferentia/deferent ducts
Prostate gland
(In some species, e.g. the cat, there is an additional gland – the bulbourethral gland)
Urethra
Penis
Prepuce.

Note: the urethra and penis are common to both the urinary and reproductive tracts.

  Page 63 

Testes

These descend from the abdomen through the inguinal ring which then closes as the animal develops. They are developed from undifferentiated gonad tissue in the fetus. When the testes move through the inguinal ring the perineum lining that is pulled through with them develops into the scrotum.

The scrotum has two layers, an inner and an outer layer:

1. Internal tunica vaginalis – the inner layer
2. External tunica vaginalis – the external layer.

At the same time the blood vessels, nerves and ducts of the glands also descend through the inguinal ring and collectively form the spermatic cord. The testes may remain in the abdomen for various reasons:

A cryptorchid animal has no testes descended
A monorchid animal has only one testis descended
  Page 64 
A rig is an animal (usually a horse) that has some testicular tissue still present in the abdomen and exhibits sexual characteristics postcastration.

Testes normally begin their descent around 8–10 weeks of age and should be palpable within the groin if not actually present in the scrotum. The testes remain relatively small until puberty (but this is dependent on breed and species). Running concurrent with the spermatic cord is a thin strip of muscle originating from the external oblique, called the cremaster muscle. The cremaster muscle retains the ability to move the testes closer to or away from the body to help control the temperature at which the spermatozoa are stored.

In small mammals the inguinal ring remains open and the cremaster muscle can withdraw the testes into the abdomen.

Temperature control of the testes is important as spermatozoa production and storage can be affected by fluctuations. Ideally the temperature of the testes should be lower than body temperature. The blood supply to the testes originates from the abdominal aorta along the spermatic cord to the associated epididymus where it becomes involuted and forms the pampiniform plexus. The pampiniform plexus allows the blood to cool and pulsations to smooth out before entering the gonad, preserving the lower temperature required for optimum sperm production and storage.

Surgical removal of testes is commonly known as castration or orchidectomy. Early castration can prevent the development of the secondary sexual characteristics:

Thickening of muscle mass
Development of erectile tissue in the penis (dog)
Distinctive urine smell and voice of tomcat.

The scrotum comprises the testis and epididymus within the tunica vaginalis.

Internal structure of the testis

The testis is made up of hundreds of convoluted tubules called seminiferous tubules. These are supported by connective tissue and cells. There are two types of cell that line the seminiferous tubules:

1. Spermatogenic cells: these divide by meiosis to produce the male gametes or spermatozoa
2. Sertoli cells: these produce oestrogens and part of the spermatic fluid (nourish spermatozoa).

The cells located within the connective tissue matrix are called the cells of Leydig. Cells of Leydig are interstitial cells which produce testosterone. Testosterone is thought of as male hormone and is responsible for regulating spermatogenesis and for the development and maintenance of the secondary sexual characteristics. Testosterone has an inhibitory effect on the hypothalamus and subsequently the anterior pituitary gland from producing interstitial cell-stimulating hormone (ICSH) and luteinising hormone (LH) until puberty.

ICSH and LH are responsible for the development of the seminiferous tubules, where the sperm are produced, i.e. no sperm are produced until after puberty.

When puberty is achieved, testosterone no longer inhibits the hypothalamus and the production of ICSH and LH results in the development of the seminiferous tubules and increased testosterone production by the cells of Leydig, i.e. sperm are produced and male characteristics are formed.

Testosterone

Testosterone is a male hormone
It is responsible for spermatogenesis
It produces the secondary sexual characteristics
Testosterone controls the descent of the testes
It inhibits the hypothalamus from stimulating the anterior pituitary to produce ICSH and LH until puberty.

Epididymus

One epididymus is located upon each of the testis. It may be palpable during examination of the testis. It consists of many coiled tubules where the spermatozoa are stored. Spermatozoa are transported to the epididymus via the vas efferentia (efferent ducts).

Vas deferens (deferent duct)

The vas deferens lie within the spermatic cord. They transport the sperm from the epididymus through the inguinal ring to the urethra. They empty into the urethra through the tissue of the prostate gland.

Structure of spermatozoa

Sperm comprise a head (acrosome and nucleus), a mid-piece(mitochondria) and a tail (flagella for movement).

Spermatogenesis is the process of sperm production. It is a complicated process with many stages. The age of the animal affects the number of spermatozoa produced, with fertility levels declining as animals age. The size of an animal may also play a role: large dogs produce more sperm than small dogs. High or low temperatures, radiation, drugs, toxins, vitamin A deficiency and limited diets can affect spermatozoa production. Abstinence reduces fertility due to blockage of tubules by old spermatozoa and it is normal practice for two coverings to occur to negate this issue.

  Page 65 

Prostate gland

The prostate gland produces a large proportion of the seminal fluid – the fluid that nourishes the sperm and provides it with a transport medium. The ejaculate comprises spermatozoa and seminal fluid. The prostate is a bilobed structure that surrounds the urethra at the level of the pelvis brim. It can often be palpated and may cause problems in older uncastrated animals as it may develop neoplasia which results in an increase in size and excretory problems, as enlargement can cause obstruction to faeces.

Bulbourethral glands

These are only found in certain species, e.g. cat. Their function is to contribute a proportion of the seminal fluid and they are located further down the urethra than the prostate, near to the perineum.

Penis

The penis runs from the ischial arch through the perineum to its conclusion between the hind legs. It differs greatly in structure between species. The distal portion is protected by the prepuce. The prepuce is a sheath or covering constructed from abdominal skin tissue suspended from the ventral abdominal wall. It is lined with mucous epithelium for lubrication and its function is to protect the distal penis or glans penis. It is pushed back to expose the glans penis during coitus. Problems can occur if the prepuce is too small for the erect penis and surgical intervention may be necessary.

Structure of the penis

The penis comprises the following structures:

Urethra
Os penis
Corpus cavernosum penis
Corpus spongiosum penis
Retractor penile muscle.

Two strips of erectile tissue or crura run along the penis from the root to almost the tip. This tissue is known as the corpus cavernosum and houses the urethra in the middle of the two layers. The corpus spongiosum runs beneath the corpus cavernosum and urethra and expands proximally to form the bulb (important to achieve the tie) and the tip or glans penis. At the glans penis the corpus spongiosum completely engulfs the urethra. The os penis is a small bone found within penile tissue in some species, e.g. the dog. It is v-shaped with the urethra running along a groove upon it.

Retractor penis muscle

The retractor penis muscle originates from the first coccygeal vertebrae and its role is to pull the penis back into the protective prepuce.

Female reproductive system

The female reproductive system (Figure 5.4) comprises the following:

Ovaries (female gonads)
Oviducts or fallopian tubes
Uterus (horns and body and cervix)
Vagina
Vestibule
Clitoris
Vulva.
image

Figure 5.4 Overview of female reproductive tract.

Functions of the female reproductive system

Production of ova (female gametes)
To provide a location for fertilisation (fallopian tubes)
To prepare the genital tract for implantation
To provide a location for implantation and subsequently pregnancy to develop.

Ovaries

The ovaries are located in the abdomen, one behind each kidney. They are held in place by the ovarian ligament which attaches the ovary to the kidney capsule. Their blood supply is provided by the ovarian artery developed from the aorta; it is very convoluted and runs alongside the ovarian ligament to the ovary itself. The ovaries lie within a fold of the mesentery called the mesovarium, which has an opening within it to allow the mature ova to depart. This opening is known as the ovarian bursa. When an animal is born it has a definitive number of undeveloped ova within each ovary. Depending upon species ova may be released individually or in a multitude.

A primary ovarian follicle develops into a mature or ripe ovarian follicle or graafian follicle under hormonal influence. The almost ripe follicle bulges out of the surface of the ovary until it is ready to pop and release the ovum (female gamete). The follicle consists of the ovum and a large quantity of fluid. The rupture of the follicle and release of the ovum occur at ovulation. The ovum is ejected and usually caught by the hair-like projections or fimbriae of the infundibulum at the head of the oviduct. The ruptured follicle then develops into the yellow body or corpus luteum (which has a role in pregnancy by producing progesterone).

  Page 66 

Oviduct or fallopian tube

The oviduct is a narrow tube that transports the ovum from the ovary to the horn of the uterus. It consists of the infundibulum that encloses the ovary and the tube portion that runs to the uterus.

Uterus (womb)

Most animals are multiparous, i.e. they give birth to many young, including the cat and the dog. These animals have a uterus comprising two uterine horns. The uterine horns lead to the body of the uterus which in turn leads to the cervix or entrance of the uterus. The uterus is located within the broad ligament or mesometrium of the uterus and is dorsal to the abdomen. The exception to this location is when the uterus is enlarged during pregnancy, when the weight of the uterus brings it into a more ventral position.

The uterus consists of layers of smooth muscle, the myometrium and a mucous epithelial lining or endometrium. The cervix is located at the base of the uterus and is a muscular sphincter that closes the uterus except during mating or parturition. The round ligament runs from the ovary through the inguinal ring where it terminates in a pad of fat known as the vaginal process. The round ligament is the female equivalent of the cremaster muscle.

Vagina

The vagina extends from the cervix to the external urethral orifice. It is completely contained within the pelvis. At the end of the vagina the genital tract deviates sharply downwards and passes into the vestibule. The vestibule is utilised by both the urinary and reproductive systems. The muscles in the wall of the vestibule are very strong and help maintain the tie during mating in the dog.

Vulva

The vulva forms the boundary between the genital tract and the outside world. It takes the form of protective folds of skin or vulval lips. Contained within the vulval lips is the female equivalent to the penis, the clitoris. The clitoris is a piece of erectile tissue and lies in the clitoral fossa, a small cavity just inside the base of the vulva. The vulval lips are normally soft but during oestrus become enlarged and turgid.

Reproductive variations

Birds

Sexing is difficult and a number of methods are employed:
image Visual inspection of cloacal anatomy
image Examination of chromosomal karyotype and/or comparison of hormone levels from blood sampling
image Surgical sexing under anaesthetic by laparoscopy
image DNA techniques (feathers) are used in most parrot species
Stress and aggression are often seen when mixing birds:
image Birds may be incompatible
image One may dominate the other, which may be species-dependent
image Stress can lead to infertility
Birds can form permanent pair bonds which take time to establish (1–2 years, e.g. swans)
Reproductive cycles depend on daylight, quality, quantity, temperature and an increase in food supply
Generally larger birds lay one egg
Another generalisation is the higher the latitude, i.e. the longer days, then the more eggs are produced
Some birds will lay more if eggs are removed from their nest, e.g. chickens
Most lay one clutch of eggs a year and some on alternate years, e.g. penguins
The interval between laying eggs is approximately 24 hours but all will hatch at the same time (chicks can communicate with each other while still in their shells and synchronise their watches!)
Size, colour and shape of eggs vary
Male birds can either have or not have a phallus
In the intromittent form (those with protruding phallus), e.g. ducks, geese, swans:
image The phallus lies on the floor of the cloaca and is erected by engorgement of lymphatic fluid
In the non-intromittent form, e.g. domestic fowl and Passeriformes:
image Two lateral folds become momentarily engorged at ejaculation and protrude to meet the protruding oviduct of the female and internal fertilisation occurs.

Reptiles

Copulatory organs of male snakes and lizards are paired and found at the sides of the base of the tail just behind the tail vent
At rest they are retracted and it is possible to palpate a pocket-like sac which can be used to determine sexing (females won’t have one if you probe, except for pythons where it is half-depth in females compared with the males)
In chelonians and crocodiles the hemipenes are situated in the ventral wall of the cloaca; these can be extruded into the female’s cloaca to facilitate passage of sperm
Internal fertilisation occurs
The nesting place is important to all reptiles – they usually dig holes
They often have problems whereby they become egg-bound because captive animals keep hold of eggs looking for an ideal nesting place, or due to stress, infections of the oviduct or mineral imbalances (calcium)
Ova are fertilised in the salpinx (located anterior to the cloaca)
Reproductive cycles are closely related to environmental conditions:
image Mature, good condition and healthy
image Space and habitat
image Daylight length and quality
image Temperature
image Moisture
image Food
Eggs can be laid or develop in a female who subsequently will give birth to live young.

Fish

External/internal fertilisation occurs
Hit-and-miss process
Gonads produce normal hormones which can also play a role in colour changes
Most species have a male and female but hermaphroditism, bisexuality, parthenogenesis and gynogenesis occur
Sex reversal in adults is not uncommon
The number of eggs produced is usually determined by the amount of parental care given, e.g. cod release millions of eggs into open water but sticklebacks that build nests and protect them only produce a few tens of eggs
Some fish will brood their young in their mouths
Fertilisation methods vary:
image Eggs and sperm are released into water
image Copulation: fertilised eggs are released
image Copulation: internal incubation – live young are released, e.g. guppy
Testes are found adjacent to the swim bladder and can weigh up to 12% of total body weight
Sperm are carried from the testes by the vas deferens to the external environment by a genital opening at the urinary papilla
In some primitive species these ducts are absent and sperm are released into the body cavity before leaving through the genital opening
Ovaries vary in complexity but are usually paired and can account for up to 70% of adult weight
In advanced species a small oviduct conducts eggs to the genital opening whilst in others ova are released into the peritoneal cavity, then find their way through to the genital opening.
  Page 68 

Bibliography

Aspinall V. The complete textbook of veterinary nursing. London: Butterworth Heinemann; 2006.

Aspinall V, O’Reilly M. Introduction to veterinary anatomy and physiology. Oxford: Butterworth Heinemann; 2004.

Campbell N, Reece J, Mitchell L. Biology, 5th edn. San Francisco: Benjamin Cummings; 1999.

Colville T, Bassett J M. Clinical anatomy and physiology for veterinary technicians. St Louis: Mosby; 2002.

Dyke K M, Sack W O, Wensing C J G. Text book of veterinary anatomy, 2nd edn. Philadelphia: W B Saunders; 1996.

Green N P O, Stout G W, Taylor D J. Biological science. Cambridge: Cambridge University Press; 1991.

Lane D R, Cooper B, Turner L. BSAVA Textbook of Veterinary Nursing. Oxford: BSAVA; 2007.

Moore M. Manual of veterinary nursing. Oxford: BSAVA; 2000.

Pratt P W. Principles and practice of veterinary technology. Philadelphia: Mosby; 1998.

Roberts M B V. Biology: a functional approach. UK: Nelson, Surrey; 1986.

Simpson G. Practical veterinary nursing, 3rd edn. Oxford: BSAVA; 1994.