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Chapter 10 Urinary system

Key Points

The functional unit of the kidney is the nephron; each kidney contains many thousands of nephrons.
Each nephron is made of several parts, each of which has a role in modifying the glomerular filtrate.
Within the glomerular capsule of the nephrons, blood passes over a fine filter. Water and very small particles are removed – this is known as ultrafiltration and forms what is known as the glomerular filtrate.
Large particles such as red blood cells and plasma proteins remain in the blood.
The very dilute glomerular filtrate passes down the nephrons and undergoes a series of modifications which result in the formation of urine. The final volume of urine is significantly reduced and reflects the status of the extracellular fluid (ECF) of the body.
The function of the kidney is to regulate the volume and the osmotic concentration of the body fluids so that they remain constant – this is one of the homeostatic mechanisms of the body and is essential if the body is to function normally.
The kidney is also responsible for the excretion of nitrogenous waste in the urine.
Urine is stored in the bladder and removed from the body via the urethra.
Urine samples can provide useful diagnostic information.

If the metabolic processes of the body are to function effectively, the chemical composition and volume of the tissue fluid must be kept constant. The most important function of the urinary system – and principally that of the kidney – is to maintain this constant internal environment, described as homeostasis.

The urinary system lies in the abdominal and pelvic cavities. It is anatomically linked with the genital or reproductive system and may be referred to as the urogenital system. Both systems share the urethra which runs through the penis of the male and joins the vagina of the female.

The parts of the urinary system are:

A pair of kidneys
A pair of ureters
Bladder
Urethra.

The functions of the urinary system are:

To regulate the chemical composition and volume of the body fluids – osmoregulation
To remove nitrogenous waste products and excess water from the body – excretion
To act as an endocrine gland by the secretion of the hormone erythropoietin (see Ch. 6).

The kidney

There are two kidneys lying in the cranial abdominal cavity, one on each side of the midline ventral to the lumbar hypaxial muscles (Fig. 10.1). Each kidney is closely attached to the lumbar muscles by a covering of parietal peritoneum. There is no mesenteric attachment, as seen in other abdominal organs, and the kidney is described as being retroperitoneal. The right kidney lies slightly cranial to the left because the stomach has evolved to lie on the left side of the abdomen, pushing the left kidney out of position. Lying close to the cranial pole of each kidney are the ovaries of the female and the adrenal glands (Fig. 10.2).

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Fig. 10.1 The position of the urinary system in the bitch.

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Fig. 10.2 Ventrodorsal view of the urogenital system of the bitch.

Macroscopic structure

The kidneys of the cat and dog have a characteristic bean shape and the indented area is known as the hilus. This is the point at which blood vessels, nerves and the ureters enter and leave the kidney. The kidneys are normally a deep reddish-brown but the colour may be affected by any substance filtering through them. On a lateral radiograph of the abdomen, a normal kidney can be seen to be equivalent in size to approximately 2.5 vertebrae (Fig. 10.3). The outer surface may be surrounded by a layer of fat, which acts as an energy reserve and protects the kidney from external damage.

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Fig. 10.3 Radiograph of the kidney area. The bladder has been inflated with air for contrast and the kidney (arrows) can be seen to measure about 2.5 times the length of the lumbar vertebrae.

When examining the cut surface of a normal kidney cut longitudinally, it is possible to see four layers (Fig. 10.4). From the outside inwards these are:

1 Capsule – protective layer of irregular dense fibrous connective tissue closely attached to the cortex. This can be easily peeled away from a healthy kidney but any adhesions may indicate previous infection or damage
2 Cortex – dark red outermost layer of the kidney containing the renal corpuscles and convoluted tubules of the nephrons
3 Medulla – slightly paler than the cortex and it may be possible to see the triangular-shaped pyramids, which contain the collecting ducts and, between them, tissue containing the loops of Henle of the nephrons
4 Pelvis – this is basin-shaped and made of fibrous connective tissue which gives it a whitish appearance. Urine formed by the nephrons drains into the pelvis and out of the kidney via a single ureter.
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Fig. 10.4 Parasagittal section through the dog kidney.

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Blood supply

Arterial blood is carried from the aorta in a single renal artery to each kidney (Fig. 10.4). This carries 20% of cardiac output. Within the tissue of the kidney, the renal artery divides into several interlobar arteries, which pass between the renal pyramids and into the cortex. Here capillaries supply the renal tubules and also give off numerous capillary networks known as glomeruli (sing. glomerulus) (Fig. 10.5). Each glomerulus supplies an individual nephron. The capillaries then recombine to form interlobar veins, which enter the single renal vein. This carries venous blood to the caudal vena cava.

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Fig. 10.5 A renal nephron.

Blood entering the kidney carries oxygen, nutrients and waste products from the tissues of the body; blood leaving the kidney carries carbon dioxide produced by the kidney tissues, but the nitrogenous waste products have been removed via the glomeruli.

Microscopic structure

The functional unit of the kidney is the nephron (Fig. 10.5). Each kidney contains about a million nephrons, which are closely packed together. They are responsible for the filtration of blood and the production of urine. Each nephron is a long tubule divided into several parts:

Glomerular capsule – a cup-shaped structure enclosing a network of blood capillaries called the glomerulus (Fig. 10.6). The capsule may also be known as Bowman’s capsule. The capsule and the glomerulus form the renal corpuscle. The basement membrane of the inner surface of the capsule, which is in close contact with the endothelium of the glomerular capillaries, is lined by podocyte cells between which are tiny pores (Fig 10.6). These pores are of such a size that they will allow the passage of fluid and small molecules, but restrict the passage of larger molecules. The outer surface of Bowman’s capsule is continuous with the epithelium of the proximal convoluted tubule. Fluid filtered by the capsule drains into the space between the two layers and continues into the next part of the nephron.
Proximal convoluted tubule – a long, twisted tube leading from the neck of the capsule and lying in the renal cortex (Fig. 10.5). The tubules are lined in simple cuboidal or columnar epithelium. The side of the epithelium directed towards the lumen of the tubule is lined by fine microvilli forming a ‘brush border’. This increases the surface area for the reabsorption of water and electrolytes.
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Loop of Henle – a U-shaped part of the tube leading from the proximal convoluted tubule and dipping down into the renal medulla. The tubule is lined in simple squamous epithelium which is thicker in the ascending loop than in the descending loop (Fig. 10.5).
Distal convoluted tubule – a short but less twisted part of the tube than the proximal convoluted tubule. It lies in the renal cortex and is lined in cuboidal epithelium without a brush border.
Collecting duct – each duct receives urine from several nephrons and conducts it through the pyramids into the renal pelvis. The ducts are lined in columnar epithelium.
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Fig. 10.6 Structure of the basement membrane of the glomerular capsule.

Renal function – the formation of urine

Blood is filtered by the kidneys and the resulting filtrate undergoes a series of modifications within the renal tubules to produce urine. This urine is very different in composition and volume from the original filtrate. For every 100 L of fluid filtered from the blood only 1 L is produced as urine – 99% of the original filtrate is reabsorbed back into the blood. The changes made to the filtrate reflect the status of the extracellular fluid (ECF) and in particular that of the blood plasma.

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The physiological processes occurring in the renal nephrons are:

Osmosis – passage of water from a weaker to a stronger solution across a semi-permeable membrane
Diffusion – passage of a substance from a high concentration to a low concentration
Reabsorption – passage of a substance from the lumen of the renal tubules into the renal capillaries and so back into the circulation; this is an active process and requires energy
Secretion – passage of chemical substances from the renal capillaries into the lumen of the tubules and out of the body in the urine; this is an active process and requires energy.

Blood enters the kidney and is carried to the capillaries forming the glomeruli.

Glomerulus

Blood pressure within each glomerulus is high because:

The blood has come straight from the renal artery and the aorta, both of which carry blood under high pressure
The smooth muscle in the walls of the efferent arteriole leaving the glomerulus is able to constrict, under the control of the hormone renin, and thus regulate the pressure of blood in the glomerulus.

High pressure in the glomerulus forces fluid and small molecules out of the blood through the pores of the basement membrane into the capsule lumen. Larger-sized particles such as red blood cells, plasma proteins and any substance bound to protein molecules, e.g. hormones are retained in the blood. This process is known as ultrafiltration and the filtrate is referred to as the glomerular filtrate or primitive urine. The glomerular filtrate is very dilute and contains 99% water and 1% chemical solutes.

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Proximal convoluted tubule

Approximately 65% of all the resorptive processes take place here (Fig. 10.7). These are:

Reabsorption of water and sodium – sodium (Na+) and chloride (Cl) ions are actively reabsorbed from the filtrate into the blood. Water is reabsorbed by osmosis in response to the movement of Na+ ions. 80% of all the Na+ and Cl ions in the filtrate are reabsorbed at this point.
Reabsorption of glucose – in the normal animal the glomerular filtrate contains glucose which is all reabsorbed back into the blood, so normal urine does not contain glucose. This reabsorption occurs up to a certain level referred to as the renal threshold.
Concentration of nitrogenous waste – the main waste product is urea, produced as a result of protein metabolism by the liver. Water reabsorption concentrates the levels of urea in the tubules.
Secretion of toxins and certain drugs – these are actively secreted into the filtrate. Penicillin and its more modern derivatives, given therapeutically, are secreted from the blood and carried to the bladder in the urine. It is therefore a useful antibiotic for the treatment of bladder infections.
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Fig. 10.7 Diagrammatic representation of the processes occurring in the different parts of the nephron.

In cases of diabetes mellitus, there is too much glucose in the blood (hyperglycaemia). This overflows into the glomerular filtrate and the tubules reabsorb it up to the limit of the renal threshold. The excess passes on down the tubules and out in the urine – glucosuria is a diagnostic sign of diabetes mellitus.

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Loop of Henle

The function of the loop of Henle is to regulate the concentration and volume of the urine according to the status of the ECF. The glomerular filtrate flows first into the descending loop and then into the ascending loop, both of which lie in the medulla (Fig. 10.7).

Descending loop of Henle – the walls of the tubule are permeable to water but do not contain the mechanism to reabsorb Na+. Water is drawn out of the tubule by osmosis – pulled by the concentration of Na+ ions in the surrounding medullary tissue. The filtrate becomes more concentrated as it passes down the loop and reaches maximum concentration at the tip.
Ascending loop of Henle – the walls are impermeable to water but contain sodium pumps that actively reabsorb Na+ into the medullary tissue and capillaries. This draws water from the descending loop. Water cannot be drawn from the ascending loop as it is impermeable. The filtrate becomes less concentrated because the Na+ ions have been removed.

The net result of this mechanism is that the filtrate is the same concentration when it enters the loop of Henle as it is when it leaves and passes into the distal convoluted tubule, but it is reduced in volume as water has been removed. This water is reabsorbed into the blood – it has been conserved for use by the body. If the animal is dehydrated, more water is reabsorbed; if it is overhydrated, more water will be lost in the filtrate.

Distal convoluted tubule

It is here that the final adjustments are made to the chemical constituents of the urine in response to the status of the ECF. This is achieved by:

Reabsorption of sodium (Na+) and secretion of potassium (K+) – the amount of reabsorption of Na+ is much smaller than occurs in the proximal convoluted tubules. Reabsorption of water by osmosis as a result of reabsorption of Na+, varies and is controlled by the hormone aldosterone produced by the adrenal glands. K+ is excreted into the urine to replace the Na+ ions (Fig. 10.7).
Regulation of acid/base balance by the excretion of hydrogen (H+) ions – the normal pH of blood is 7.4.
If pH falls, i.e. the blood becomes more acid due to excess H+ ions, the kidney excretes H+ ions into the urine via the distal convoluted tubule. The pH of the blood returns to normal.
If pH rises, i.e. the blood becomes more alkaline due to reduced amounts of H+ ions, the kidney stops excreting H+ ions, retaining them in the blood. The pH of the blood falls to normal.

Collecting duct

Here the final adjustments are made to the volume of water in the urine in response to the status of the ECF (Fig. 10.7). The hormone antidiuretic hormone (ADH), produced by the posterior pituitary gland, is able to change the permeability of the duct walls to water.

Diuretics are drugs used to increase the volume of urine produced and so reduce the volume of fluid in the body. One of the most commonly used – furosemide – is a ‘loop diuretic’, which acts on the loops of Henle. It prevents reabsorption of Na+ so water is not resorbed by osmosis and remains in the urine as it flows through the tubules.

If the animal is dehydrated, the volume of the plasma and ECF will be reduced. ADH will be produced and the permeability of the collecting ducts to water will be increased. Water will be drawn through the walls by the high concentration of Na+ ions in the surrounding medullary tissue (secreted by the ascending loop of Henle), and into the plasma and ECF (Fig. 10.7).

As a result of the above processes, the ultrafiltrate that passed through the glomerular capsule has now become concentrated urine by the repeated effects of reabsorption and secretion.

Osmoregulation – control of renal function

Osmoregulation ensures that the volume of the plasma and the concentration of dissolved chemicals in the plasma and other tissue fluids remains constant. Homeostasis is maintained and the body can function normally. Several factors are involved in the control of osmoregulation (Table 10.1) and it is achieved in two ways:

Control of the amount of water lost from the body
Control of the amount of salt (NaCl) lost from the body.

Table 10.1 Factors involved in osmoregulation

Controlling factor Function
Renin (hormone) Produced by the glomeruli of the kidney in response to low arterial pressure
Angiotensinogen (plasma protein) Converted to angiotensin by the action of renin
Angiotensin (protein) Causes vasoconstriction; stimulates the release of aldosterone from the adrenal cortex
Aldosterone (hormone – mineralocorticoid) Secreted by the cortex of the adrenal gland; acts mainly on the distal convoluted tubules but has a lesser effect on the collecting ducts; regulates the reabsorption of Na+ ions
Antidiuretic hormone (ADH or vasopressin) Secreted by the posterior pituitary gland; mainly affects the collecting ducts by changing their permeability to water; also has an effect on the distal convoluted tubules
Baroreceptors Found in the walls of the blood vessels; monitor arterial blood pressure
Osmoreceptors Found in the hypothalamus; monitor the osmotic pressure of the plasma; affect the thirst centre of the brain and influence the secretion of ADH

Control of water loss

Water is lost by the healthy animal in urine, faeces, sweat and in respiration. Very small amounts may also be lost in vaginal secretions and tears. If this water loss is not replaced by food and drink, or if the water loss is excessive, e.g. in vomiting, diarrhoea or haemorrhage, the total volume of ECF falls and the animal is described as being dehydrated.

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A dehydrated patient will show:

Lowered blood pressure – a fall in the volume of ECF results in a fall in blood plasma volume and consequently a fall in blood pressure
A rise in osmotic pressure – a fall in plasma volume results in a rise in the concentration of Na+ ions in the blood and as a result a rise in the osmotic pressure of the blood.

Note: Osmotic pressure is the pressure needed to prevent osmosis from occurring. It is dependent on the number of particles of undissolved molecules and ions in a solution. Blood pressure is the pressure exerted on the inside of the walls of the arterial blood vessels by the blood. It is detected by baroreceptors in the walls of the vessels.

In dehydration, osmoregulatory mechanisms will start to work (Fig. 10.8). The fall in blood pressure and rise in osmotic pressure ultimately result in an increase in the reabsorption of water from the collecting ducts, under the control of the hormone ADH (see Ch. 6). The osmoreceptors also stimulate the thirst centre in the brain and the animal feels the need to drink. Both of these pathways result in a rise in plasma volume and an increase in arterial blood pressure. Dehydrated animals should be given fluid parenterally (intravenously) or, in less severe cases, provided with access to drinking water. They will excrete reduced quantities of concentrated urine.

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Fig. 10.8 Mechanism controlling the amount of water lost from the kidney.

Raised blood pressure will result in the opposite effect, and the animal will produce increased quantities of more dilute urine.

Control of sodium levels

Sodium is taken into the body in the form of salt (NaCl) in food. It is lost in sweat, faeces and urine. Sodium is found in the ionised form Na+ in all the body fluid compartments and plays a fundamental part in determining arterial blood pressure. High Na+ levels in the diet draw water into the plasma by osmosis. This raises blood volume and blood pressure also rises. Conversely, if an animal has a low level of Na+, less water is drawn into the plasma and blood volume falls – this lowers the arterial blood pressure. The osmoregulatory mechanisms take steps to correct this, resulting in a compensatory rise in blood pressure (Fig. 10.9).

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Fig. 10.9 Mechanism controlling sodium (and therefore blood pressure) in the kidney.

Regulation of Na+ in the plasma occurs mainly in the distal convoluted tubule and is controlled by the hormone aldosterone, secreted by the cortex of the adrenal gland (see Ch. 6).

Excretion

Excretion is the removal of waste products from the body. These are formed within the tissues as a result of metabolic processes and are useless, surplus to the requirements of the body or potentially harmful to the body tissues. Excretion by the kidneys removes:

Water – in varying amounts dependent on the volume of the ECF and controlled by osmoregulatory processes
Inorganic ions – the amount depends on the osmotic pressure of the blood and other body fluids and is controlled by osmoregulatory processes
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Nitrogenous waste products – these result from the metabolism of protein taken into the body in food. Protein is broken down by the process of digestion into amino acids which are carried to the liver by the blood and converted into body protein. Surplus amino acids cannot be stored by the body so undergo a process of deamination in which they are broken down. Ammonia, which is extremely toxic to the cells, especially those of the nervous system, is formed as a by-product. Within the liver ammonia combines with carbon dioxide in a series of reactions known as the ornithine cycle and urea is formed:

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Urea passes into the circulation and is carried to the kidney where it is excreted in the urine.

Kidney disease can present in many forms and the symptoms are related to the fact that all kidney functions are affected and the homeostatic mechanisms cease to work. Chronic renal failure commonly occurs in older cats and dogs and develops when the nephrons are gradually replaced by fibrous connective tissue as a normal ‘old age’ change. Symptoms, many of which are related to rising levels of urea in the blood, develop when 75% of the nephrons have ceased to function.

Products of detoxification – hormones, certain drugs and poisons are inactivated within the liver and excreted by the kidney.

Other parts of the urinary system

The ureters

Urine formed by the nephrons leaves each kidney by a single ureter at a point known as the hilus (Fig. 10.4). Each ureter is a narrow muscular tube running caudally towards the bladder, one on each side of the midline in the dorsal abdomen (Figs 10.2, 10.3). Each ureter is suspended from the dorsal body wall by a fold of visceral peritoneum, the mesoureter.

Urine is pushed along the tube towards the bladder by peristaltic waves brought about by contraction of the smooth muscle fibres forming the ureter walls. Transitional epithelium lines the ureters and allows for expansion as urine passes along.

The bladder

Each ureter enters the single bladder close to the neck. The ureters underrun the bladder mucosa for a short distance before opening into the lumen at an oblique angle. This area, known as the trigone, acts as a valve preventing the backflow of urine along the ureter (Fig. 10.2).

The bladder is a pear-shaped hollow organ. The rounded end points cranially, while the narrow end or neck points caudally and usually lies within the pelvic cavity. Its function is to collect and store urine. When full, the bladder extends into the abdomen, pulling the neck ventrally over the edge of the pelvic brim with the ventral surface of the bladder touching the abdominal floor; when empty most of the bladder lies in the pelvic cavity.

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In cross-section, the bladder consists of an inner lining of transitional epithelium, which enables the walls to expand when filling with urine. A submucosal layer of elastic tissue and smooth muscle is arranged in folds to allow expansion. The smooth muscle fibres are continuous with those of the internal bladder sphincter. The bladder is surrounded by a layer of peritoneum, which covers only the cranial end lying in the abdomen. All organs in the pelvic cavity are surrounded by connective tissue and muscle and are not covered in peritoneum.

Urolithiasis is the formation of mineral ‘stones’ known as uroliths or calculi in the bladder. These may cause a blockage in the urinary tract. The most common site for blockage in the cat is at the tip of the penis; the most common site in the male dog is as the urethra bends over the ischial arch and as it passes through the os penis. Blockage is rare in the female animal because the urethra is shorter and more distensible.

The neck of the bladder ends in the bladder sphincter, whose function is to control the flow of urine out of the bladder and down the urethra. It consists of two concentric parts:

Internal sphincter – made of smooth muscle; under involuntary control
External sphincter – ring of striated muscle; under voluntary control.

The urethra

The urethra is a tube which conveys urine caudally from the bladder through the pelvic cavity to the outside. Its structure varies according to the sex of the animal, and in the case of the male animal, between cats and dogs.

Female – the urethra is a short tube, opening into the floor of the reproductive tract at the junction of the vagina and vestibule. The opening is known as the external urethral orifice and is located in the centre of a small ridge, the urethral tubercle. This is a useful landmark when catheterising a bitch and can be seen when using a speculum.
Male – the urethra is divided into two parts: the pelvic urethra and the penile urethra. There is a difference between the dog and the tomcat:
Dog (see Ch. 11, Fig. 11.1). Close to the neck of the bladder, there are three openings into the urethra – one of these is from the prostate gland and two are from the deferent ducts or vas deferens (one from each testis). The urethra runs caudally through the pelvic cavity, over the edge of the ischial arch where it is joined by erectile tissue to form the penis. The penile urethra opens to the outside at the tip of the penis.
Tomcat (see Ch. 11, Fig. 11.2). There is a short length of urethra, cranial to the openings from the prostate gland and the deferent ducts, known as the preprostatic urethra, which is not found in the dog. The urethra continues caudally and opens to the outside in the perineal area, ventral to the anus. There is no penile urethra lying outside the pelvic cavity. Close to the end of the urethra are the openings from the paired bulbourethral glands.

From the point at which the deferent ducts join it, the urethra conveys both urine and sperm to the outside of the body.

Micturition

Micturition (often incorrectly referred to as urination) is the act of expelling urine from the bladder. It is normally a reflex activity but can be overridden by voluntary control from the brain. The steps involved are:

Bladder distends with urine formed by the kidneys
Stretch receptors in the smooth muscle of the bladder wall are stimulated and send nerve impulses to centres in the appropriate segment of the spinal cord
Nerve impulses are transmitted via parasympathetic nerves back to the smooth muscle, and initiate contraction
Nerve impulses also stimulate relaxation in the internal bladder sphincter and urine is expelled.

If it is inappropriate for the animal to micturate, the brain overrides the reflex pathway and prevents the bladder sphincter from relaxing. At a more appropriate time, the brain stimulates both the external and internal sphincters: they relax and urine is released. Voluntary control develops as the young animal matures – in puppies and kittens it is not fully developed until the animal is about 10 weeks old.

Urinalysis

Urine is derived from the ultrafiltrate of plasma so it reflects the health status of the whole animal. The analysis of urine, or urinalysis, is a useful diagnostic tool. Normal urine contains only water, salts and urea. The clinical parameters used to evaluate a sample of urine are shown in Table 10.2.

Table 10.2 Normal values shown by the urine of the dog and cat

Clinical parameter Normal value Comments
Daily volume Dog: 20–100 ml/kg body weight Polyuria – increased volume of urine
Oliguria – reduced volume of urine
  Cat: 10–12 ml/kg body weight Anuria – absence of urine
Appearance Clear, yellow, characteristic smell Tomcat urine has an unpleasant strong smell; old samples smell ammoniacal
pH 5–7 Carnivorous diet produces acid urine; herbivorous diet produces alkaline urine
Specific gravity (s.g.) Dog: 1.016–1.060 Reflects the concentration of urine; exercise, high environmental temperatures and dehydration will cause a rise in specific gravity
  Cat: 1.020–1.040
Protein None Proteinuria – presence of protein.
May indicate damage to nephrons, chronic renal failure, inflammation of the urinary tract
Blood None Haematuria – presence of blood
Haemoglobinuria – presence of haemoglobin, due to rupture of red cells
May indicate damage or infection to the urinary tract.
Glucose None Glucosuria – presence of glucose
May indicate diabetes mellitus; levels of glucose in the filtrate exceed the renal threshold and excess is excreted in the urine
Ketones None Ketonuria – presence of ketones
May be accompanied by acid pH and smell of ‘peardrops’ in urine and on the breath
Bile None Bilirubinuria – presence of bile
Indicator of some form of liver disease
Crystals and casts In small quantities, these may be considered to be normal Crystalline or colloidal material coalesce to form a cast of the renal tubules and are flushed out by the urine; in large quantities crystals may form calculi or uroliths and block the tract