A-LEVEL BIOLOGY (2014) NAKAPANKA JUDE MAYANJA 0774649208/0704716641

HOMEOSTASIS
Homeostasis is the relative constancy of the body’s internal environment regardless of the conditions in the external
environment.

Examples of components of the internal environment that are homeostatically controlled include;

a. Concentration of blood glucose at 90mg/100cm3

b. Average core body temperature at 37oC or 98.6oF
c. Blood pressure at the brachial artery averages near 120/80
d. Blood levels of ions such as Na+, Cl-, Ca2+e.t.c.
e. Concentration of carbon dioxide

Cybernetics is the study of control systems i.e. self-regulating systems which operate by means of feedback
mechanisms. A feedback mechanism is one in which an input stimulus causes an output response that ‘feeds back’
to the initial input.

The internal state of an animal’s body is a dynamic equilibrium i.e. many physical and chemical changes do occur
that the net result is that, the changes are maintained constant by feedback systems so as to enable cells function
efficiently.

Feedback may be positive or negative

a. Negative feedback
A mechanism in which the effect of deviation from the normal condition triggers a response that eliminates its
deviation in order to reduce further corrective action of the control system once the set point value has been
reached.
In negative feedback mechanism, a stimulus causes a sensory receptor to signal the regulatory centre in the
brain. The regulatory centre then signals an effector to respond and the response cancels/reverses the stimulus to
restore the condition to the norm.
b. Positive feedback
A mechanism in which the effect of deviation from the normal condition intensifies the original response such
that the change tends to proceed in the same direction as the initial stimulus.
Examples of positive feedback include;
i. A 10oC decrease in temperature doubles metabolic activity, releasing more heat that raises the activity
even more
ii. During child birth, stretching of the uterus by the foetus stimulates contraction and the contractions
stimulate further stretching. The cycle continues until the foetus is expelled.
iii. During blood clotting, to stop bleeding in order to keep blood volume constant. One clotting factor
activates another in a cascade that leads quickly to the formation of a clot. A cascade effect is the way
in which a small amount of, say a hormone, can cause a target organ to produce a large amount of the
product.

ESSENTIAL COMPONENTS OF A CONTROL SYSTEM

Each control system must have the following essential components;

a. Receptors/detectors
These are parts of the body hat constantly monitor and detect changes from the reference point/norm in the
internal environment and then signal the deviations.
Temperature receptors in the skin provide information on variation in temperature of external environment.
b. Control centre

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This is usually the brain that coordinates the information received from the various receptors and sends out
instructions which will correct this deviation.
Variations in temperature of external environment are conveyed to the hypothalamus of the brain.
c. Effector/responding organs
These are parts of the body that bring about the necessary changes needed to return the system to the reference
point/norm.
The hypothalamus initiates corrective responses in effectors like blood vessels and skin to restore the
temperature back to the normal. The average temperature is 37 oC.
d. Reference point/norm
This is the set level at which the system operates
e. Feedback loop
Hormones and/or nerve impulses that inform the receptor of any change in the system as a result of the action of
the effectors.

(Fig 19.1 & 19.2 R. Soper page 648)

TISSUE FLUID
This is the fluid that is derived from blood plasma by filtration through capillaries. It is found around tissue cells and
contains molecules that enter from or exit to the capillaries. The body’s internal environment consists of tissue fluid
and blood that bathes cells.

FORMATION OF TISSUE FLUID

Tissue fluid is formed by ultrafiltration i.e. hydrostatic pressure of blood forces small molecules to exist blood
capillaries via fine pores on the basement membrane but large molecules are held back.

Fluid movement in and out of capillaries depends on the balance between the blood pressure (hydrostatic pressure)
and osmotic pressure (solute potential). Osmotic pressure is created by the presence of salts and plasma proteins in
blood while blood pressure is created by the pumping action of the heart and the resistance to blood flow caused by
the small size of the lumen of capillaries.

At the arterial end of the capillary bed, blood pressure is higher than osmotic pressure of blood. This results into the
forced exit of small molecules like glucose, water, amino acids, ions, oxygen, and small plasma protein molecules
via fine pores on the basement membrane of arterioles BUT large plasma protein molecules and red blood cells are
retained.

Midway along the capillary bed where the blood pressure is lower, the two forces of blood pressure and osmotic
pressure essentially cancel other and the substances diffuse according to their concentration gradients i.e. glucose,
oxygen and other solutes diffuse out of the capillary while carbondioxide and other wastes diffuse into the capillary.
No net movement of water occurs.

At the venule end of the capillary bed, blood pressure is lower than osmotic pressure of blood, resulting into entry of
water, carbondioxide, wastes and solutes into the capillaries. However, the total amount of fluid exiting capillaries at
the arterial end exceeds that entering at the venule end. This is because the osmotic pressure causing the entry of
fluids at the venule end is lower than the blood pressure causing exit of fluid at the arterial end, resulting into failure
of some fluid flowing in capillaries, forming what is called tissue fluid.

(Roberts and Reiss, page 268)

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Note. Tissue fluid is drained by the lymphatic system, where it becomes lymph which eventually passes into the
veins at the same rate as it is formed, failure of which results in a condition known as oedema.

HOMEOSTASIS IN UNICELLULAR ORGANISMS AND CELLS OF MULTICELLULAR

ORGANISMS
At cellular level, the internal environment of a cell is its cytoplasm while the cell’s immediate surrounding
constitutes its external environment.

Tissue fluid in most animals and sap in plants, constitute the external environment of cells of multi cellular animals
and plants respectively, but form the internal environment of these organisms.

The constituents of a cell cytoplasm are modulated by the partial permeability of its cell membranes and the level of
activity of its enzymes.

The cell surface membrane selectively allows entry and exit of molecules at a strictly controlled rate by diffusion
gradient, osmotic gradients and active transport.

The nature and amounts of materials synthesised is controlled by the rates of protein synthesis and they catalyse
most anabolic and catabolic reactions within cells.

Therefore, relative constancy of the cell’s internal environment depends on supply of metabolites, utilisation of
cellular material or out put through activity of the modulators.

HOMEOSTATIC ROLE OF THE LIVER AND THE PANCREAS

A. Structure
The liver is the largest internal organ of the body, weighing about 1.5kg in humans, which is about 3.4% of the
total body mass. The liver’s external shape is of little importance, but the internal structures reveals precious
details
The liver is composed of structural and functional units called lobules, which are cylindrical in shape,
numbering over 1000 with each being approximately 1mm in diameter. Liver cells (hepatocytes) closely pack
in each lobule in various rows radiating outwards from the centre. Hepatocytes which are in contact with blood
vessels bear microvilli. Hepatocytes are characterised by similarity in structure and function, prominent nuclei,
Golgi complex, glycogen granules, peroxisomes, numerous mitochondria, fat granules and lysosomes.
Peroxisomes contain catalase and other oxidative enzymes responsible for detoxification.
Located between lobules are triads consisting of a branch of the hepatic artery which bring oxygenated blood
to the liver, a branch of the hepatic portal vein which brings nutrients from the gut and the bile duct that drains
bile from the liver. A central vein (branch of the hepatic vein) runs longitudinally mid-way each lobule and is
linked by sinusoids to the interlobular vessels (hepatic artery and hepatic portal vein). Sinusoids radiate from
the centre to the periphery of the lobule and their endothelial lining is perforated. Sinusoids alternate with the
bile canaliculi; small canals that carry bile. Attached to the walls of sinusoids are macrophagus cells called
kupffer cells

(Roberts and Reiss, page 277 fig 16.11 & Soper page 667 fig. 19.21)

B. How structure is related to function

i. Kupffer cells ingest worn-out red blood cells, bacteria and foreign particles from the blood flowing
through the liver
ii. Closeness of hepatocytes with sinusoids and canaliculi enables them to receive nutrients and expel
waste products
iii. The excellent blood supply provides nutrients to the cells and enables waste to be carried away
iv. Hepatocytes bear numerous mitochondria for ATP production required in providing energy that
facilitate some of the metabolic reactions

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v. The liver is large, providing a large surface area for metabolic reactions to occur
vi. Hepatocytes bear numerous peroxisomes containing catalase and other oxidative enzymes responsible
for detoxification of poisonous substances in the liver
vii. Its tissue is elastic, enabling expansion to store a large volume of blood
viii. Hepatocytes are similar in structure (undifferentiated) enabling them to perform various metabolic
functions
C. Functions of the liver
Some of the functions of the liver, out of the estimated 500 that it performs, are digestive, regulatory and
excretory.
I. It maintains a steady blood glucose concentration by conversion of glucose into glycogen (if above
the norm) and vice versa (if below the norm), under the influence of hormones. The liver’s
carbohydrate metabolism involves the following;
i. Glycogenesis, promoted by insulin
ii. Glucogenolysis, promoted by glucagon
iii. Lactic acid metabolism, initiated by the enzyme lactate dehydrogenase
iv. Gluconeogenesis, promoted by cortisone and adrenaline hormone

(Fig 19.4 R.Soper page 650 & Fig 13.6 Chilton page 107)

II. The liver regulates amino acids and proteins (protein metabolism) in the body
i. Excess amino acids are not stored in the body, any surplus is gotten rid of by the liver through
deamination. Deamination involves removal of the amino group (-NH2) from the amino acid
to form ammonia, which in mammals is converted through a series of reactions (ornithine
cycle) to urea, CO(NH2)2, which is shed from the liver cells into the blood stream and
transported to the kidney for excretion. The amino acid residue is fed into the carbohydrate
metabolism pathway and oxidised to release energy.
ii. The liver also carries out transamination i.e. it transfers amino groups from amino acids to
other organic compounds to form amino acids that are deficient in the diet.

(Fig 4.7.3 Ann Fullick page 331 & Fig 16.7 Roberts and Reiss page 274)

III. The liver regulates lipids (lipid metabolism) in the body

i. Excess carbohydrate is converted to fat
ii. Stored fat are de-saturated prior to oxidation
iii. It synthesizes and degrades phospholipids and cholesterol
iv. It synthesizes lipid transporting globulins

Excess cholesterol in blood is excreted into bile by the liver to avoid accumulation, which can
result in arteriosclerosis (narrowing of the lumen of arteries) which can cause thrombosis (blood
clotting in blood vessels). If cholesterol is greatly in excess in bile it forms gall stones, which can
block the bile duct.

IV. The liver forms red blood cells in the foetus and breaks down worn out red blood cells in adults. The
liver’s kupffer cells break down worn out red blood cells to form the bile pigment, bilirubin, which
is excreted in bile. The haemoglobin is broken down into globin, a protein and haem, from which
iron is removed and stored
V. The liver detoxifies poisonous substances i.e. naturally occurring compounds absorbed by the body
which can be toxic if allowed to accumulate are rendered harmless by the liver cells (hepatocytes)
e.g. ethanol is oxidised to ethanal. Products of detoxification are usually excreted, but sometimes
they are stored

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VI. It synthesizes plasma proteins from amino acids. They include albumin (a transport molecule),
globulin (a transport molecule of hormones), prothrombin and fibrinogen (clotting factors)
VII. The liver produces bile, which is a mixture of salts and cholesterol. Bile emulsifies fats during
digestion in the duodenum
VIII. The liver stores fat soluble vitamins A, D, E, K and water soluble vitamins B 12 and C.
IX. The liver stores minerals like iron, potassium, copper, zinc and trace elements
X. The liver stores up to 1500cm3 of blood in its vast network of blood vessels, hence acting as a blood
reservoir during emergency cases
XI. The liver destroys all hormones after exerting their effects in the body

REGULATION OF BLOOD GLUCOSE

The concentration of glucose is 90-100mg of glucose per 100cm3 approximately 0.1% or 0.1g per 100cm3 of blood.
The glucose level may lower to 70mg per 100cm3 of blood, this is known as hypoglycaemia or rise to 150-200mg
of blood and this is known as hyperglycaemia.

HOW THE LIVER AND PANCREAS INTERACT TO MAINTAIN GLUCOSE LEVELS

CONSTANT
Hyperglycaemia stimulates the beta cells of the islets of Langerhans in the pancreas to secrete the hormone insulin
into blood. Insulin binds to body cells with insulin receptors and leads to processes which reduce glucose
concentration, for example;

i. Increased cellular respiration in muscle and liver cells to form carbondioxide and water
ii. Increased glycogenesis (formation of glycogen from glucose) in muscle and liver cells
iii. Increased concentration of glucose to fat and protein in adipose tissue
iv. Increased uptake of glucose in muscle cells

Hypoglycaemia inhibits insulin secretion but stimulates alpha cells of the islets of Langerhans in the pancreas to
secrete the hormone glucagon into blood. Glucagon binds to liver cells since they are the only ones with glucagon
receptors, causing them to increase blood glucose level through;

i. Increased glycogenolysis (hydrolysis of glycogen to glucose)

ii. Increased formation of glucose from amino acids and glycerol. The formation of glucose from non-
carbohydrate sources is called gluconeogenesis.

(Fig 19.22 R.Soper page 668)

Note:

1. Insulin and glucagon are not the only hormones that control the blood glucose concentration. For example;
i. Adrenaline (from adrenal medulla) causes hydrolysis of glycogen during acute stress or
excitement and the usage of glucose and thus increases and reduces its concentration in blood
respectively.
ii. Cortisol (from the adrenal cortex) causes formation of glucose from amino acids and glycerol
when glycogen exhausts, hence increasing glucose concentration in blood.
iii. Growth hormone (from the anterior pituitary)increases the glucose concentration in blood
through fat breakdown.
iv. Thyroxine (from the thyroid gland) stimulates the metabolic rate e.g. increased glucose
breakdown.
2. In some people there may be insufficient secretion of insulin or the cells may be insensitive to insulin,
resulting into a condition known as diabetes mellitus. Insulin dependent diabetes is caused by insufficient

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secretion of insulin while insulin independent diabetes results from insensitivity of cells to insulin.
Symptoms of diabetes include;
 Hyperglycemia
 Glycosuria (glucose in urine)
 Frequent copious urine
 Visual disturbances
 Itching of genitals
 Fatigue
 Rapid weight loss
 Drowsiness
 Skin disorders e.g. boils
 General weakness

Liver disorders

a. Hepatitis, which is an inflammation, caused by hepatitis viruses A, B, C, D and E.

b. Jaundice, which is characterised by a yellowish tint to the white of the eyes and a lightly pigmented skin
c. Cirrhosis, the liver becomes fatty, then fibrous and it is common in alcoholics
d. Liver cancer which is caused by exposure to chemicals like cigarette smoke, radiations e.g. X-rays, or
genetic pathways.

THERMAL REGULATION (TEMPERATURE CONTROL)

The temperature of animals must be regulated because;

I. Most body enzymes at effectively within a narrow temperature range of 35-38oC. Temperatures above
450C denature enzymes and other proteins. Temperatures below the narrow range inactivate enzymes.
Inactivation and denaturation of enzymes are both fatal.
II. Excessively high or low temperatures disorganise the structure and functioning of cell surface membranes,
and consequently affects entry and exit of substances resulting into death of the organism.

Terms associated with thermoregulation

a. Endotherm, an organism capable of maintaining a stable body temperature independent of the environmental
temperature by generating heat metabolically when environmental temperature is low e.g. mammals and birds
b. Endothermy, the ability of animals to maintain a constant internal body temperature
Advantages
 Animals are able to exploit various environments regardless of the existing temperatures
 Enzyme controlled reactions proceed without much interruption most of the time
 Since high metabolic reactions are maintained all the time, plenty of energy is availed to support body
processes
Disadvantages
 There’s high food intake during low environmental temperatures to support the metabolic reactions
that liberate heat
 Enzyme controlled reactions are slowed during low temperatures because enzymes become inactive
 Enzyme controlled reactions are slowed during low temperatures to avoid overheating of the body, and
efficient insulation when the external temperature is too low.
c. Homeotherm, an organism capable of maintaining a stable body temperature independent of the environmental
temperature e.g. mammals and birds
d. Ectotherm, an organism whose body temperature is regulated by behaviour or by the surrounding e.g. reptiles,
fish, amphibians, insects e.t.c (all other animals except mammals and birds)
Advantage

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 There is low food intake since regulation of temperature is by behavioural means and from the
environment
Disadvantages
 Animals have limited environments to exploit depending on the existing external temperature
 Their activities are limited during instances of extreme temperature
e. Poikilotherm, an animal with a body temperature that fluctuates with that of the external environment e.g.
retiles, fish, amphibians, insects e.t.c. (all other animals except mammals and birds)

(Fig 19.6 R. Soper page 653)

HEAT GAIN AND LOSS IN ORGANISMS

Heat gain, heat is gained as a by-product of metabolism from exothermic reactions

Heat loss, heatis lost by evaporation of water from the body during sweating and from body surfaces like the mouth
and respiratory surfaces of land dwelling animals.

Heat may be gained or lost to the environment by radiation. This is the transfer of energy in the form of
electromagnetic waves.

Heat may be gained or lost to the environment by convection. This is the transfer of heat by currents of air or water

Heat may be gained or lost to the environment by conduction. This is the transfer of heat by the collisions of
molecules. Conduction is particularly important between organisms and the ground or water, since air is a poor
conductor of heat

THE HUMAN SKIN

Functions of the skin

i. It is the major organ involved in temperature regulation in the body

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ii. It provides protection against mechanical damage, ultra violet radiation from the sun, microorganism
invasion and water loss by underlying tissues
iii. It is a sense organ, containing sensory nerve endings for detecting temperature, touch, pressure and pain
iv. It is an excretory organ of urea, salt and excess water
v. It manufactures vitamin D when exposed to sunlight. The dermis contains lipids called steroids which are
converted into vitamin D by ultraviolet light.

Response of endotherms to variation in the external temperature

Response to hot conditions

a. Physical and physiological means

- Metabolic rate reduces to minimise on the heat generated in the body
- Erector pili muscles relax to lower hairs/fur, so that no insulating layer of air is trapped near the skin
surface enabling much heat to be lost.
- Panting occurs in dogs, birds and cats to increase evaporation of heat from the lungs, pharynx and other
moist surfaces to cool the body
- Sweat production by sweat glands increases to enable evaporation of water from the skin surface
- Vasodilation occurs i.e. superficial capillaries dilate to increase blood flow so that much heat can be lost by
conduction and radiation
b. Behavioural means (in man)
- Taking cold drinks
- Putting on light clothing
- Moving to shady places
- Switching on a fun
- Taking a bath
- Being active mainly at night (nocturnability

(Fig 2 Kent page 154)

Response to cold conditions

a. Physical and physiological means

- Shivering, which is due to the involuntary contractions of the skeletal muscles, occurs so as to generate heat
metabolically
- Metabolic rate increases to generate extra heat in the body. This occurs particularly in muscles and liver
cells. Special brown fat may also be metabolized.
- Erector pili muscles contract to cause hairs/fur to ‘stand on end’ to trap an insulating layer of air near the
skin to reduce heat loss by convection
- Sweat production by sweat glands reduces/stops to reduce evaporation of heat from the skin surface
- Vasoconstriction occurs i.e. superficial capillaries narrow to reduce blood flow so that heat loss by
conduction and radiation can be minimised
b. Behavioural means (in man)
- Taking hot drinks
- Putting on thick clothes
- Turning on heat in houses
- Moving near a heat source e.g. fire

(Fig 15.6 Roberts page 236)

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EFFECT OF CHANGING THE ENVIRONMENTAL TEMPERATURE

Two separate groups of naked people, one is exposed to gradually cooling air while another is exposed to gradually
increasing air temperature. Their metabolic rates, physical changes and internal core body temperature are observed.
Naked people are used to avoid physical interference of clothes or covering such that observations made are based
on physiological responses only.

(Fig 15.6 Roberts page 238)

Low critical temperature, this is a low environmental temperature at which physical mechanisms like
vasoconstrictions and erection of hair fail to maintain body temperature constant, triggering a rise to generate heat to
maintain body temperature constant

Lower lethal temperature, this is extremely low temperature at which increased metabolic rate fails to generate
enough heat to maintain body temperature constant, resulting into death of the organism

Hypothermia, is a condition that results when heat loss greatly exceeds heat gain from metabolism due to prolonged
exposure to cold, resulting into great reduction in core body temperature of the organism

High critical temperature, this is a high environmental temperature at which physical mechanisms like sweating
and vasodilation fail to maintain temperature constant, triggering a rise in metabolic rate and body temperature as
environmental temperature rises

Upper lethal temperature, this is an extremely high environmental temperature at which increased metabolic rate
generates excessive heat which denatures enzymes and other structures, resulting into death of the organism.

Efficiency range (range of temperature neutrality), this is the external temperature range at which the body’s
physical mechanisms are capable of maintaining temperature constant. In man this is 27-31oC.The efficiency range
varies according to the environmental temperature in which the animal inhabits. This is because animals have the
ability to acclimatize. If the environmental temperature is high, acclimatization is by raising the upper critical
temperature and if low, acclimatization is by lowering the lower critical temperature.

(Fig 18.7 Roberts and Reiss page 311 and Fig 15.7 Roberts page 439)

Observations from the graph

a. the low critical temperature for animals living in cold places is much lower than for those which live in warm
places
b. the lower lethal temperature is much lower for cold-dwellers than for warm dwellers
c. below the lower critical temperature, the metabolic rate of warm-dwellers rises more sharply than in cold-
dwellers
d. the metabolic rate starts to rise at a much critical temperature for cold-dwellers than that for warm-dwellers

ADAPTATIONS TO EXTREME CLIMATES

Adaptations of life to low temperature

a. structural adaptations
 possession of thick fur for trapping a layer of air that is warmed and remains insulating the body against
heat loss e.g. polar bears

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 possession of a thick layer of subcutaneous fat for insulating against heat loss e.g. polar bears
 development of a large body size as compared to their counterparts in warmer climates to reduce the
surface area to volume ratio in order to reduce heat loss e.g. whales and polar bears i.e. Bergman’s rule
 extremities such as ear lobes are of reduced size than those of related species in warmer climates to reduce
surface area for heat loss i.e. Allen’s rule
b. other adaptations
 Development of a counter current heat exchange system in limbs to enable heat conservation by
minimizing its loss to the environment e.g. in ducks legs and dolphin flippers
The counter current heat exchange system is a heat conservation mechanism in limbs where there’s
effective heat transfer at all levels to the periphery of the limbs, by conduction from the incoming warmer
arterial blood to the outgoing colder venous blood
 Small sized animals hibernate e.g. bats, hamsters, hedgehogs and rodents like mice. Hibernation is a
seasonal response by animals to cold temperatures during which they become dormant, body temperature
and metabolic rate fall to the minimum required for maintaining the vital activities of the body. The
animals are said to be in a ‘deep sleep’ to reduce energy needs to survive the winter when food is scarce.
Brown fat is conserved and used up rapidly at the end of hibernation to quickly raise the metabolic heat.
Brown fat owes its colour to the numerous mitochondria it contains. The mitochondria generate heat and
not ATP. Animals moving out of hibernation break it down and it generates heat more quickly than
ordinary fat since it has good blood and nerve supply
 Some animals migrate to warmer places e.g. birds like swallows

(Fig 15.8 Roberts page 239 OR Fig 19.18 R. Soper page 664)

Adaptations of life to high temperature

a. Structural adaptations
 Bodies are thinly insulated with fat to increase heat loss
 Development of smaller body size than their counterparts in colder climates to increase surface area to
volume ratio in order to increase heat loss
 Extremities, such as ear lobes are large, thin with a rich blood supply to enable heat loss more easily e.g.
the elephant’s ears
 Having tissues that are tolerant to large temperature fluctuations between day and night e.g. the camel
b. Other adaptations
 Some animals aestivate. Aestivation is a seasonal response by animals to drought or excessive heat during
which they become dormant, and the metabolic rate decreases followed by a decrease in body temperature
to a minimum, required for maintaining the vital activities of the body. The African lungfish burrows into
mud until the dry season ends, earthworms and garden snails also aestivate.

(Fig 18.10 Roberts and Reiss page 312 & Fig 21.6 Chilton page 167)

Case study 1: adaptation to extreme heat, the camel

1. It faces the sun, therefore exposing a small surface area to the sun’s radiation
2. Its body is insulated by fat on top, which minimizes heat gain by radiation
3. The underpart of its body, which has much less insulation, radiates heat out to the ground cooled by the
animals’ shadow
4. Flattened nostrils which retard water loss
5. It excretes very concentrated urine
6. They can tolerate the loss of more than 25 percent of their body weight in water
7. They can go without drinking for as long as an entire week in the summer and three weeks in winter

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8. It can tolerate a fluctuation in internal body temperature of 5 to 6 oC i.e. it stores heat during day by letting
ist temperature rise up and then release it during the night and begins the following below its normal
temperature i.e. storing up coolness
9. The hump acts as an insulator and impedes heat flow into the body core

ROLE OF THE BRAIN IN TEMPERATURE CONTROL

The thermoregulatory centre the brain, the hypothalamus, is responsible for temperature regulation in the body.
Variation in body temperature is directly monitored by heat receptors in the hypothalamus and indirectly by
receptors in the skin. Receptors in the skin monitor variation in external temperature.

If the temperature of blood flowing through the hypothalamus decreases, the heat regain centre is stimulated to
send impulses to the liver and muscles to raise the metabolic rate so as to generate heat, and to the skin to cause
vasoconstriction, reduction in sweat production, contraction of the erector pili muscles and shivering. The overall
result is increased body temperature back to normal.

If the temperature of blood flowing through the hypothalamus rises, the heat loss centre is stimulated to send
impulses to the skin to cause vasodilation to enable more heat loss at the skin surface, increase sweat production to
enable more evaporation, relaxation of erector pili muscles to lower the hairs to avoid air insulation and to inhibit
shivering so as to minimise heat production by metabolic reactions. All these enable lowering of temperature to
normal.

Variation in external temperature stimulates receptors in the skin to send impulses to the skin and the animal’s
behaviour is modified accordingly e.g. if the skin heats up, the animal may move to a shade, while cooling of the
skin surface may give rise to increased metabolic activity.

(Fig 15.3 & 15.5 Roberts page 236 & 237)

TEMPERATURE CONTROL IN ECTOTHERMS

This is mainly achieved through modification of behaviour of the organism, which may include;

 Basking in the sun, at varying angles to the sun’s rays so as to gain heat e.g. lizards and crocodiles
 Hiding in burrows, holes or crevices in rocks away from sunlight reduces temperature e.g. lizards
 Panting and exposing the moist tissues of the mouth, by licking the body surface or by swallowing in
water, an animal can increase evaporation and so heat loss from the body
 Thermal gaping, opening the mouth to enable evaporation of moisture from the buccal cavity to cool blood
e.g. alligators
 Thermal dancing when it is hot i.e. lifting of opposite pairs of feet alternately so that they can cool in air
e.g. shovel-snout lizards
 Salivation over the neck and legs in tortoises to increase loss of heat as a result of water evaporating from
such surfaces

TEMPERATURE CONTROL IN PLANTS

Plants lose and gain heat by the same physical processes as animals i.e. radiation, convection, conduction and
evaporation

Plant tissues can tolerate wide fluctuations in temperature and are adapted to live in a variety of habitats, but still
they must regulate the temperature to avoid overheating which would denature enzymes, and freezing of tissues
which would slow down the metabolic processes.

Plants prevent overheating by;

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 Transpiration, as water evaporate into the atmosphere it cools down the body of the plant
 Wilting, parenchyma cells lose turgidity to reduce the surface area of leaves and stems exposed to the sun,
hence avoiding much heat gain
 Possession of a shiny cuticle on leaves to reflect heat (sun’s radiation) and avoid overheating
 Possession of small needle-like leaves in some plants also reduces excessive heat gain from the sun’s rays

Plants prevent freezing by;

 Producing spores or seeds which are very temperature resistant

 Losing the easily damaged leaves when external temperature is low e.g. during winter in temperate plants
 Orienting leaves to take maximum advantage of light at any one time so that they do not shade each other

EXCRETION AND OSMOREGULATION

Excretion is the expulsion of waste products of metabolism from the body. Examples of waste products of
metabolism include; carbon dioxide, urea, uric acid, ammonia, bile, excess water, excess mineral salts, oxygen
(plants) e.t.c.

Osmoregulation is the control of water and salt balance so that the concentration of dissolved substances in the
body fluids remains constant. It includes concentration of various ions e.g. Na +, K+, Cl- and water content.

Homeostasis is the maintenance of a constant internal environment within in a narrow range regardless of the
conditions in the external environment.

Secretion is the production of substances useful to the body by cells. For example release of hormones and digestive
juices.

Egestion is the removal from the body of undigested food and other substances, which have never been involved in
the metabolic activities of cells. For example, elimination of faeces from the gut (defeacation) and undigested food
from the food vacuole of amoeba.

Significance of excretion and osmoregulation

1. Enables removal of unwanted by-products of metabolic pathways to prevent unbalancing of the chemical
equilibrium of reactions
2. Removes toxic wastes, that if accumulated would affect the metabolic activities of organisms e.g. may act as
enzyme inhibitors
3. It regulates ionic concentration of the body fluids to facilitate efficiency of cell activities e.g. nervous
coordination
protein synthesis, hormone production, muscle contraction, enzyme activity e.t.c.
4. It regulates the water content of body fluids
5. Enables regulation of ions that have a major influence on the pH of body fluids e.g. H +, and HCO3-.
6. It removes unwanted substances that are taken in along with food.
7. Enables removal of excess nutrients which if allowed to accumulate would interfere will cell activities

EXCRETION AND OSMOREGULATION IN PLANTS

EXCRETION
Plants do not have complex/elaborate excretory systems as those in animals because of the following reasons;

 Toxic wastes do not accumulate because they are utilised by the plants e.g. carbon dioxide and water are
raw materials for photosynthesis while oxygen participates in respiration
 The rate and amount of catabolism is much slower and much less than that of animals of similar weight
and as a result, the waste products accumulate more slowly

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 Plant synthesise all their organic requirements according to demand, leaving no excess proteins hence very
little excretion of nitrogenous waste substances occurs.
 Organic acids which would be harmful to plants often combine with excess cations and precipitate as
insoluble crystals which can be safely stored in plant cells e.g. excess calcium ions combine with oxalic
and peptic acids to form the non-toxic calcium oxalate and calcium pectate.
 Extra gaseous wastes are removed from plant bodies by simple diffusion through the stomata and lenticels
 Most of the organic waste substances formed are non-harmful and can be stored in the plant tissues which
are removed periodically e.g. leaves and barks
 Some plants wastes such as resins in organs that later fall off e.g. leaves
 Excess water and dissolved gases are removed by exudation e.g. gums, resins, latex and rubber
 In some plants, excess water with dissolved salts oozes out through hydathodes, a process called guttation

Excretory products of plants include;

a. Carbon dioxide, water and oxygen from respiration and photosynthesis respectively
b. Anthocyanins stored in petals, leaves, fruits and barks.
c. Tannins deposited in dead tree tissues like barks and wood
d. Calcium oxalate, calcium carbonates and latex (rubber)
e. Alkaloids like quinine, cannabis, cocaine, caffeine, morphine e.t.c.

OSMOREGULATION
Plants are categorised into four major groups depending on the amount of water in their environment;

a. Hydrophytesare plants that live completely or partially submerged in fresh water. They have enough water and
therefore there’s no problem of obtaining it e.g. water lilies, water hyacinth, water lettuce e.t.c. They need to get
rid of excess water, absorb maximum sunlight, remain buoyant and absorb respiratory gases. Their
characteristics include;
i. Submerged plants
 The lamina is thin to allow water and dissolved oxygen to enter the leaves by osmosis through the
epidermis
 They have numerous chloroplasts to trap as much light as possible for photosynthesis
 They are broad to increase on the surface area for trapping sunlight and diffusion of respiratory gases
ii. Partially submerged plants
 The leaves have thick waxy cuticle on the upper layer to minimise water loss
 Stomata are located only on the upper surface to permit exchange of gases with the atmosphere
iii. Floating plants
 Some species bear numerous hairs on the upper surface to trap air and so make the plant buoyant e.g. the
water fern
 Water hyacinth has air filled tissues in the swollen base of the leaf petioles which makes the plant float
 The air filled tissues if the hyacinth also store oxygen
 Their roots are freely suspended in water from which they derive their mineral nutrients
 Their leaves have a large surface area (broad) to trap as much light as possible for photosynthesis
b. Mesophytes are plants inhabiting normal well-watered soils. They have the following characteristics;
 The leaf surface of some plants is hairy thus trapping air which forms a humid insulating layer that lowers
the rate of transpiration
 They have more stomata on the lower surface than on the upper surface to reduce on water loss through the
stomata
 Some plants have a milky latex which, being viscous, reduce transpiration
 A thick waxy transparent cuticle on the upper surface which minimises water loss through the upper
epidermis yet still permits light penetration into the leaf

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c. Halophytes are plants inhabiting areas of high salinity e.g. estuaries and salt marshes. They have a problem of
absorbing water from their highly saline surrounding and getting rid of excess salts. Their characteristics
include;
 They have special salt-secreting glands located on the upper epidermis. These extract and remove excess
salts absorbed into the tissues from the saline water
 They produce aerial stilt or prop roots which grow down into the soft mud and give the plant firm
anchorage and support
 They have special aerial roots called pneumatophores which project upwards from the soil so as to take up
oxygen through the numerous lenticels on their surface
 They are viviparous i.e. the seeds germinate while the fruit is still on the parent plant
 The seedlings are long and pointed to enable them penetrate the mud when they drop off from the parent
plant
d. Xerophytes are plants inhabiting dry areas e.g. deserts. Their characteristics include the following;
i. Modified leaves
 Some have very small leaves to reduce the surface area for transpiration
 Some shed their small leaves in the dry season
 The leaves of some are reduced to small spines
 The leaves may be small, rolled-up and needle-like
 There’s a waxy cuticle on the surface of the leaves to reduce on the transpiration rate
ii. Distribution of stomata
 They generally have few stomata, and these are mainly located on the lower leaf surface
 The stomata in cacti are located in grooves or sunken pits
 In cacti, stomata open at night
iii. Water storage in plant tissues. Some plants store water in leaves e.g. Bryophyllum and stems e.g. banana
iv. They have deep and extensive rooting systems
v. Some plants are very short-lived i.e. they grow, blossom and produce seeds in a short wet season
vi. Milky latex. Some plants have a white latex in their stems which reduces on their rate of transpiration and
also makes them distasteful to herbivores.

Adaptation of xerophytes for surviving unfavourable water balance i.e. more loss than uptake from the soil

Structural adaptations

i. Possession of extremely deep roots to obtain water from deep below the water table e.g. acacia
ii. Shallow root systems for absorbing moisture even slight showering e.g. cactus
iii. Possession of fleshy succulent stems and leaves that store water in large parenchyma cells e.g. brophyllum
and cactus
iv. Hairy epidermis for trapping humid air and reduce on the transpiration rate
v. Possession of stomata which are sunken in hairy leaf surfaces to trap air and reduce on the transpiration rate
vi. Rolling/folding of leaves to reduce on transpiration e.g. marram grass (Ammophila)
vii. Possession of a thick cuticle which is impermeable to water e.g. prickly pear (Opuntia)
viii. Reduction of surface area over which transpiration has to occur by having small leaves

Physiological adaptations

i. Reversal of the normal stomatal rhythms in some plants i.e. opening stomata at night and closing during
day time so as to reduce on water evaporation
ii. Increased levels of abscisic acid, which induces stomatal closure so as to reduce water loss
iii. Possession of tissues tolerant to dessication i.e. low solute potential of cytoplasm and production of
resistant enzymes
iv. Leaf fall in deciduous trees so as to cut down transpiration
v. Survival of drought as seeds or spores that are highly dehydrated and protected within a hard case

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EXCRETION AND OSMOREGULATION IN ANIMALS

Different animals use different organs to carry out excretory and osmoregulatory processes. The organs of some
animals are shown in the table below;

Animals Excretory and osmoregulatory structures

Unicellular organisms Cell surface membrane
Platyhelminthes Flame cells
Crustaceans Antennal glands
Annelids Nephridia
Arachnids Book lungs
Insects Malpighian tubules
Fish Gills and kidneys
Amphibians Lungs, kidneys, liver, gills and skin
Birds and reptiles Lungs, kidneys and liver
Mammals Lungs, kidneys, liver and skin
Animals excrete more than one product and the predominance of one over the others is determined by three factors;

a. The production of enzymes necessary to convert ammonia to either urea or uric acid
b. The availability of water in the habitat for the removal of the nitrogenous excretory material
c. The animal’s ability to control water loss or uptake by the body

The nitrogenous excretory products of animals include;

i. Ammonia which is highly soluble and readily diffusible, excreted by fresh water bony fish, protozoans, porifera
and cnidarians which live in abundance of water. Such animals are said to be ammoniotellic.
ii. Urea is less toxic than ammonia and very soluble hence easily diluted before elimination, so it is excreted by
some terrestrial animals like mammals and marine ones whose body fluids are hypotonic to seawater. Animals
that excrete urea as the main nitrogenous product are said to be ureotelic.
iii. Uric acid is almost nontoxic and highly insoluble, requiring very little water for its elimination so it is excreted
by animals living in very arid conditions e.g. birds, insects and reptiles, which live in areas of water shortage.
These animals are said to be uricotelic
iv. Trimethylamine oxide is soluble but nontoxic, requiring relatively less water for its elimination, so it is
excreted by marine bony fishes suffering from water shortage
v. Guanine is less soluble than uric acid and requires no water for its elimination hence it is excreted by terrestrial
spiders that live in scarcity of water.

Summary of the relationship between excretory produce and habitats of some animal groups

Animal Habitat Excretory product Nature of product

Mammals Terrestrial Urea
Birds and terrestrial insects Terrestrial Uric acid
Protozoa and fresh water bony fish Aquatic Ammonia Nitrogenous wastes
Spiders Terrestrial Guanine
Marine bony fish Aquatic Trimethylamine oxide
Mammals Terrestrial Bile salts
Mammals, birds and reptiles Terrestrial Excess water and mineral salts Non-nitrogenous wastes
Mammals, birds and protozoa Terrestrial Carbon dioxide

EXCRETION AND OSMOREGULATION IN FRESHWATER PROTOZOANS

Protozoans include amoeba and paramecium and they use the contractile vacuoles to carry out osmoregulation.
Since the cell contents are hypertonic to the surrounding, and the cell membrane is partially permeable, there is
constant influx of water into the cytoplasm by osmosis.

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Small vesicles in the cytoplasm fill up with fluid from the cytoplasm and pump salts back into the cytoplasm by
active transport, using energy provided by ATP from the numerous mitochondria surrounding the vesicles.

The vesicles, now containing water, fuse with the contractile vacuole which gradually expands. The impermeability
of the vacuolar membrane to water prevents osmotic outflow of water. On reaching a certain size, the contractile
vacuole fuses with the cell surface membrane, contracts suddenly and releases its water.

(Fig 14.10 Roberts page 221 & Fig 20.2 R. Soper page 676 OR Fig 18.7 Clegg page 378)

NOTE;

1. The frequency of contractile vacuole discharge decreases with increase in salinity of surrounding water. In
fresh water, the frequency is high to expel water as fast as possible to avoid bursting of the cell.
2. Excretion of carbon dioxide and ammonia in protozoa is by simple diffusion across the cell surface
membrane

EXCRETION AND OSMOREGULATION IN TERRESTRIAL INSECTS

Excretion

The excretory system of an insect consists of malpighian tubules that extend throughout the body cavity. The ends
of the malpighian tubules are closed but the epithelial cells absorb nitrogenous wastes like sodium and potassium
urate from blood. Muscles in the tubule wall produce peristaltic movements that stir up the blood. Water and carbon
dioxide react with potassium urate in the tubule cells to form potassium hydrogen carbonate and uric acid.

Potassium hydrogen carbonate is absorbed back into blood while uric acid is deposited in the tubule lumen. As uric
acid moves from the distal to the proximal end of the malpighian tubule, water is vigorously taken back into blood
while solid crystals of uric acid are deposited in the lumen and later return to be passed out.

(Fig 14.14 A & B Roberts Page 226 OR Fig 18.8 Clegg page 378)

Osmoregulation in selected organisms

A. Shore crab (carcinus)

Antennal glands at the base of the antennae excrete excess water and nitrogenous wastes. Antennal glands are
incapable of holding back salts (they may eliminate salts and water alike)
B. Mitten crab (Eriocheir)
What happens in the shore crab also happens in the Mitten crab except that here the inward secretion of salts is
sufficient enough to enable the animal to flourish in fresh water.
C. Cray fish
Here antennal glands are capable of eliminating excess water but reabsorbs salts, resulting into production of
urine hypotonic to blood an internal osmotic pressure (OPi) higher than external osmotic pressure (OPe).
Reabsorption of salts occurs as urine flows along the coiled tubule.

Drawing of antennal glands of Cray fish and shore crab.

The graph below shows changes in the internal osmotic pressure (OPi) of blood with external osmotic
pressure in the surrounding medium (OPe) in three different genera of crabs

(Fig 14.9 Roberts page 220)

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Explanations of the variations in osmotic pressure of blood

 OPi of carcinus slightly decreases with slight lowering of OPe (little dilution of sea water) but osmoregulation
breaks down and OPi drops drastically when external medium (OPe) becomes too dilute. Carcinus in fresh
water experiences osmotic extraction of water from the body leading to dehydration of body tissue, hence
cannot exploit fresh water fully.
 OPi of Eriocheir slightly decreases throughout (in both sea and fresh water) because the animal is able to
osmoregulate in all concentrations of external medium. Eriocheir can tolerate much greater dilution and survive
in both fresh water and sea water
 A change in OPe results in a similar change in OPi of blood, which is an indicator that Maia crab cannot
osmoregulate

EXCRETION AND OSMOREGULATION IN FISH

Osmosis and Osmolarity
All animals-regardless ofphylogeny, habitat, ortype ofwaste produced-facethe same need for osmoregulation. Over
time, water uptake and loss must balance. If water uptake is excessive, animal cells swell and burst; if water loss is
substantial, they shrivel and die. Water enters and leaves cells by osmosis. Recall that osmosis is aspecial case
ofdiffusion, is the movement ofwater across aselectively permeable membrane. It occurs whenever two solutions
separated by the membrane differ in osmotic pressure or osmolarity (total solute concentration expressed as
molarity, or moles ofsolute per liter ofsolution). Iftwo solutions separated by aselectively permeable membranehave
the same osmolarity, they are said to be isoosmotic. Under these conditions, water molecules continually cross the
membrane but they do so at equal rates in both directions. In other words, there is nonet movement ofwaterby
osmosisbetween isoosmotic solutions. When two solutions differ in osmolarity, the one with the greater
concentration ofsolutes is said to be hyperosmotic, and the more dilute solution is said to be hypoosmotic. Water
therefore moves by osmosis from a hypoosmotic solution to a hyperosmotic one.

Osmotic Challenges

An animal can maintain water balance in two ways. One is to be an osmoconformerwhich is isoosmotic with its
surroundings i.e. its body fluids are in osmotic balance with its balance. The second is to be an
osmoregulator,which controls its internal osmolarity independent ofthat ofits environment.

All osmoconformersare marine animals. Because an osmoconformer's internal osmolarity is the same as that
ofitsenvironment, there is no tendency to gain or lose water. Many osmoconformers live in water that has a stable
composition and hence have aconstant internal osmolarity. Osmoregulation enables animals to live in environments
that are uninhabitable for osmoconformers, such as freshwater and terrestrial habitats. It also allows many marine
animals to maintain an internal osmolarity different from that of seawater. To survive in a hypoosmotic
environment, an osmoregulator must discharge excess water. In a hyperosmotic environment, an osmoregulator
must instead take in water to offset osmotic loss. Most animals, whether osmoconformers or osmoregulators,
cannottolerate substantial changes in external osmolarity and are said to be stenohaline (from the Greek stenos,
narrow, and halos, salt). In contrast, euryhaline animals (from the Greek eurys. broad), which include certain
osmoconformers and osmoregulators, can survive large fluctuations in external osmolarity. Many barnacles and
mussels covered and uncovered by ocean tides are euryhaline osmoconformers; familiar examples of euryhaline
osmoregulators are the various species of salmon

A. Fresh water teleosts (bony fish) e.g. tilapia, trout, stickle e.t.c. (OPi > OPe)
The excretory and osmoregulatory organs are gills and kidneys. The internal body fluids are hypertonic to the
surrounding water and therefore there is osmotic influx of water across the gills, lining of mouth and pharynx.
There is also efflux of solutes (ions and ammonia) into water by diffusion hence the kidneys are not important
in the excretion of nitrogenous wastes. The problem of intake of water and loss of ions is solved by;

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 Not drinking water

 Producing large volumes of dilute (hypotonic) urine
 Reabsorbing ions across the nephron tubules, from the glomerular filtrate back into blood. The high
glomerular filtration rate is enabled by numerous large glomeruli in the kidneys.
 Active uptake of salts from water by chloride secretory cells in the gills

B. Marine teleosts e.g. Cod, Mackerel (OPi > OPe)

The excretory and osmoregulatory organs are gills and kidneys. The internal body fluids being hypotonic to the
surrounding water, there is osmotic extraction of water from the body leading to dehydration of the tissues, a
situation described as physiological drought. This is overcome by;
 Drinking large volumes of sea water
 Having a kidney with a low filtration rate enabled by the few small sized glomeruli
 Excreting trimethylamine oxide, which is soluble but non-toxic and requires little water for
elimination
 The divalent ions (Ca2+, Mg2+, SO42-) in sea water that a marine fish drinks are eliminated through the
anus while the monovalent ions (K+, Na+, Cl-) are absorbed into blood and are actively transported out
of blood across the gills, reverse to the direction in fresh water fish. The divalent ions that enter blood
are secreted into the nephron tubules and excreted in urine. The waste products are selectively
extracted from blood by the tubule cells and ultrafiltration does not occur.

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C. Marine elasmobranchs (cartilaginous fish) e.g. dog fish, sharks, rays e.t.c.
Their tissue is slightly hypertonic to the seawater causing a slight influx of water, which is readily expelled by
the kidneys. Hypertonic tissue fluid results from retention of urea which is facilitated by;
 Impermeability of gills to urea
 Urea reabsorption from the nephron tubules, maintaining its concentration at over 100 times higher
than that in animals
 Tolerance of tissues and enzymes to high urea concentration
D. Migratory fish e.g. salmon and eels
Their ability to move from one extreme osmotic environment (fresh water) to another (sea water) is achieved
by;
 Changes in kidney filtration rates
 Reversal of the direction in which the chloride secretory cells transfer cells i.e. in fresh water, they
take in salt and they may move the salts out in sea water.

OSMOREGULATION IN TERRESTRIAL ANIMALS

Terrestrial animals lose water through evaporation from the permeable surface exposed to the atmosphere. They
exhibit different physiological, morphological and behavioural adaptations so as to minimise water loss.

Physiological adaptations

i. Reduction in glomerular filtration rate e.g. the desert frog (Chiroleptes) has few glomeruli than its relatives
living in moist temperate regions
ii. Production of non-toxic nitrogenous wastes e.g. insoluble uric acid (reptiles, birds and insects) and the relatively
less toxic urea (mammals and amphibians) that require little water for removal
iii. Extensive water reabsorption from glomerular filtrate (mammals and birds) and rectum (insects) e.g. the
kangaroo rat has an extra-long loop of Henle enabling it to produce hypertonic urine
iv. Use of metabolic water from fat through respiration. This explains why desert animals like the kangaroo rat tend
to metabolise fat which yield more water on oxidation than carbohydrates
v. Possession of tissues tolerant to dehydration e.g. a camel can survive for a long time without drinking water

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vi. Ability to sweat abnormally high temperatures e.g. a camel begins sweating at 41 oC
vii. Ability to reduce the need for nitrogenous excretion e.g. a camel secretes urea into the lumen of the alimentary
canal where bacteria convert it to protein which is then utilised as food

Behavioural adaptations

i. Change of habitat depending on the weather conditions

ii. Some animals aestivate e.g. the African lung fish
Aestivation is the seasonal response by animals to drought or excessive heat, during which they become
dormant, body temperature rate decrease to the minimum required for maintaining the vital activities of the
body. It is an adaptation for temperature regulation as well as water conservation. During aestivation, the
African lung fish burrows and encases in a cocoon of hard mud lined with mucus

Morphological/structural adaptations

Possession of water proof integuments which include the keratinous scales of reptile, cornfield epithelium of
mammals and the waxy cuticle of insects.

Case study 2: adaptation to lack of water, the Kangaroo rat

The kangaroo rat, a desert inhabitant, may spend its entire life without drinking water. It lives on seeds and other dry
plant materials.

1. It has no sweat glands, therefore no water lost as sweat

2. Being nocturnal, it searches for food only when the external temperature is relatively cool.
3. Its feaces have a very low water content
4. Its urine is highly concentrated
5. Some cooling of the expired air takes place in its long nose, with condensation of water

OSMOREGULATION IN AMPHIBIANS AND REPTILES

The kidneys of amphibians are identical to those of fresh water fish since the amphibians were the first terrestrial
animals. Reptiles on the other hand, have different kidney structures because they live in diverse habitats;

a. Those that are living mainly in fresh water e.g. some crocodiles, possess kidneys like those of fresh water fishes
and amphibians
b. Marine reptiles e.g. some crocodiles, turtles, sea snakes and some lizards e.g. the Iguana possess kidneys
identical to those of their fresh water relatives. However, since these kidneys reabsorb salt, marine reptiles
cannot excrete a great deal of salt in their urine. Instead they eliminate excess salt by means of salt secreting
glands located near the nose or the eye, hence ‘the turtle shedding tears’
c. Terrestrial reptiles reabsorb much of the salt and water in the nephron tubules of the kidneys, enabling the
reptiles to conserve blood volume in the dry environment

EXCETION AND OSMOREGULATION IN MAMMALS AND BIRDS

Mammals and birds are the only vertebrates able to produce urine that has a higher osmotic concentration than their
body fluids. This enables them to excrete waste products in a small volume of water so that more water can be
retained in the body. For example, human kidneys produce urine 4.2 times as concentrated as their blood plasma, the
camel, gerbil and pocket mouse can excrete urine 8, 14 and 22 times as concentrated as their blood plasma
respectively. Birds however have a relatively few or no nephrons with long loops, so they cannot produce urine that
is concentrated as that of mammals. Marine birds e.g. Penguins, Gulls and Cormorants drink salt water and then
excrete the excess salt from salt secreting nasal glands near the eyes, giving an impression that these birds have
runny noses.

Note

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1. The production of hypertonic urine is accomplished by a portion of the nephron called the loop of Henle
found only in the kidneys of mammals and birds
2. The ability to produce concentrated (hypertonic) urine enables the organism to conserve water and is a
useful adaptation to terrestrial life

STRUCTURE OF THE MAMMALIAN KIDNEY

(Fig 14.1 Roberts page 211)

EXCRETORY AND HOMEOSTATIC FUNCTIONS OF THE KIDNEYS

1. Excretion of metabolic waste products such as urea, excess water, uric acid, ammonia, creatine e.t.c.
2. Regulation of water and salute content of blood
3. Maintenance of pH of body fluids at 7.4 (acid-base balance by removing or neutralizing excess acidic or basic
ions)
4. Regulation of blood levels of ions as Na +, K+, Cl-, Ca2+ e.t.c.
5. Secretion of hormone, erythropoietin, which stimulates red blood cell production for transporting oxygen
6. Retention of important nutrients such as glucose and amino acids through reabsorption from the glomerular
filtrate into blood

EXCRETORY FUNCTION OF THE KIDNEY

The kidney accomplishes its excretory function by a number of processes which occur at different regions of the
nephron;

i. Ultrafiltration (pressure filtration) at the glomerulus of the Bowman’s capsule

ii. Selective reabsorption in the tubules
iii. Tubular secretion at the proximal and distal convoluted tubules
iv. Counter current multiplier effect in the loop of Henle
v. Water reabsorption in the distal convoluted tubule and collecting duct

ULTRAFILTRATION

This is the first stage of urine formation at the glomerular capillary wall of the kidney nephrons during which
hydrostatic pressure forces small molecules in blood of the glomerular capillaries to pass across the basement
membrane into the capsular space BUT large molecules are held back.

The substances that are forced by pressure to pass passively across the basement membrane filter include small
molecules like water, glucose, amino acids, vitamins, urea, uric acid, ions, creatine and some hormones while large
the large molecules retained in blood include red blood cells, platelets, white blood cells and large sized plasma
proteins.

Although filtration occurs through three layers of the glomerular capillary, the endothelium is a coarse screen
retaining only blood cells. The negatively charged basement membrane retains negatively charged large sized
protein molecules while the selective filtration occurs at the diaphragm of the slit pores formed by the foot-like
projections of supporting cells podocytes.

(Fig 17.5 Roberts and Reiss page 383)

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SELECTIVE REABSORPTION AT THE TUBULES

Because particle size and rather than their importance determines the substance to pass through the basement
membrane during ultrafiltration, useful substances such as glucose enter the capsular space to form the glomerular
filtrate and they have to be absorbed later.

As the glomerular filtrate (renal fluid) flows along the tubule of the nephron, all the glucose, 85% of water, Na +, Cl-,
amino acids, vitamins, hormones, 50% of urea are absorbed from the proximal convoluted tubule into the
surrounding blood capillaries.

Glucose, amino acids, Na+, H2PO4-, and HCO3-, diffuse into the proximal convoluted tubule and then actively
transported into the blood capillaries. The active uptake of Na + is followed by the passive uptake of Cl- which raises
the osmotic pressure in the cells, enabling entry of water into the capillaries by osmosis.

50% of urea is reabsorbed by diffusion but the small sized proteins in the renal filtrate are removed by pinocytosis.
As a result of this activity, the tubular filtrate is isotonic with the blood in the surrounding capillaries.

(Fig 17.7 Roberts and Reiss page 284)

TUBULAR SECRETION AT THE PROXIMAL CONVOLUTED TUBULE

Finally active secretion of unwanted substances like creatine, some urea, ammonia (active transport), uric acid,
H+(active transport), and K+ occurs from blood capillaries into the proximal convoluted tubule.

COUNTER CURRENT MULTIPLIER EFFECT IN THE LOOP OF HENLE

This is a mechanism which occurs in the loop of Henle, resulting into creation of a very high concentration gradient
between the tissue fluid and blood in the medulla, and the urine in the collecting duct.

The ascending limb of the loop of Henle is relatively impermeable to water while the descending limb is freely
permeable to water BUT impermeable to water and salts. Na+ and Cl- are actively pumped out of the upper part of
the ascending limb but diffuse from the lower part, raising the solute concentration in the interstitial region and
lowering the concentration in the ascending limb. Water is osmotically drawn from the descending limb and
collecting duct and carried away by blood in the vesa recta, resulting into a slightly higher concentration in the
descending limb than the adjacent ascending limb and hypertonic urine is formed.

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The concentration effect is multiplied such that the fluid in and around the loop of Henle becomes saltier with the
saltiest region being the hair pin bend. The glomerular filtrate becomes less salty as it goes up to the ascending limb.

(Fig 20.4 Kent page 149 and Roberts and Reiss page 285)

WATER REABSORPTION

This occurs in the distal convoluted tubule and collecting duct. It is under the influence of hormones which regulate
water content and solute content of blood.

As fluid flows down the collecting duct, water is drawn out of it osmotically into the interstitium, resulting in
hypertonic urine production.

Changes in the renal-plasma ratio of glucose, urea and chloride in the kidney of a frog. A renal plasma ratio of 1.0
means that the concentrations are the same in the renal fluid and plasma. If the ratio is greater than 1.0 the
concentration is greater in the renal fluid than in the plasma. If it is less than 1.0, the concentration is lower in the
renal fluid than in the plasma. Urea rises mainly because of the large amount of water reabsorbed. This data shows
whether a substance is reabsorbed or not.

(Fig 14.5 Roberts page 215)

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DETERMINANTS OF THE GLOMERULAR FILTARTION RATES

Glomerular Filtration Rate (GFR) is the net rate of formation of filtrate by the two kidneys. GFR is equal to the
Renal Plasma Flow (RPF) rate, the rate of plasma flow through the renal arteries, multiplied by the filtration of this
plasma flow that is filtered (Filtered Fraction, FF).

GFR = RPF + FF

FF is determined by three factors,

 Filtration pressure across the glomerular capillary walls

 Permeability of the renal filter to the fluid
 Total surface area available for filtration

The kidneys receive a large part of the cardiac output of 5.6litres per minute, the kidneys might receive about 1.2
litres per minute, about 20%.

GFR increases if the,

 Mean glomerular capillary pressure rises as a result of either dilation of the afferent arterioles of
constriction of the efferent arterioles
 Concentration of the plasma proteins decreases because this reduces the force favoring reabsorption.

GFR decreases if the,

 The hydrostatic pressure in the Bowman’s capsule rises e.g. if the ureter is occluded.
 Plasma protein escapes into the Bowman’s capsule, because proteins in the Bowman’s capsule make the
net osmotic force across the glomerular wall smaller.

As a consequence of net reabsorption of solute, the volume of fluid in the nephrons decreases, until only about 1%
or 2% of the original filtrate volume reaches the ureter as final urine.

The adaptive significance of formation of urine by filtration in the glomerular followed by reabsorption and
secretion in the later parts of the nephrons is to enable the kidney excrete soluble chemicals that might enter the
body, e.g. drugs and bacterial toxins but for which there are no specific tubular reabsorption pathways.

ADAPTATIONS OF THE MAMMALIAN NEPHRON TO ITS FUNCTIONS

a. The tubules
 the proximal convoluted tubule cells;
- Bear numerous microvilli at the free end to increase the surface area for reabsorption of substances like
glucose, amino acids, vitamins, sodium chloride and water.
- Contain numerous mitochondria to form ATP that provide energy required in active transport of
glucose, amino acids, Na+, H2PO4- and HCO3- into the blood capillaries.
- The cell surface membrane is indented to form a large area of intercellular spaces bathed with fluid
- Contain numerous pinocytic vesicles, which enable digestion of small protein molecules from the renal
filtrate
 The loop of Henle is U-shaped with parallel, opposite flows of tubular fluid in its limbs to provide a
multiplier effect that creates a concentration gradient which enables increased water reabsorption
 The capillaries of vesa recta from the loops are in close proximity with tubules to increase the reabsorption
of useful substances from the filtrate
 The distal convoluted tubule is long and coiled to increase the surface area for reabsorption of water and
mineral salts
 The distal and proximal convoluted tubules are coiled to slow down the movement of renal filtrate to allow
more time for efficient reabsorption of substances like water and mineral salts
b. Parts other than the tubule

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 Afferent arterial entering the Bowman’s capsule has a wider lumen than that of the efferent arterial leaving
it, resulting into high hydrostatic pressure that causes ultra-filtration to occur
 The Bowman’s capsule is funnel-shaped to direct the renal filtrate into the proximal convoluted tubule
 The structural arrangement of the three layers of the glomerular capillary enables the diaphragms of slit
pores formed by foot-like projections of podocytes to offer selective filtration while blood cells and the
negatively charged large plasma protein are retained by endothelium and basement membrane respectively

Why does urine production almost stop after serious bleeding?

The amount of urine produced is proportional to the amount of blood flowing through the kidneys. The total blood
volume in the body reduces if serious bleeding occurs, resulting into diversion of blood from other tissues (including
the kidneys) to brain to maintain life. Therefore the volume of blood flowing through the kidneys reduces greatly to
the extent that less ultra-filtration occurs leading to formation of less urine.

HOMEOSTATIC FUNCTIONS OF THE KIDNEY

The kidneys carries out it homeostatic functions with other organs of the body under the influence of several
hormones. The homeostatic functions of the kidney include;

i. Maintenance of pH of body fluids at 7.4 (acid base balance) to avoid denaturing of enzymes and other
proteins, which would result into death
ii. Regulation of blood levels of ions such as Na+, K+, Cl-, Ca2+ e.t.c.
iii. Regulation of water and solute content of blood (osmotic regulation)

MAINTENANCE OF pH OF BODY FLUIDS

The body produces more acids than bases, causing the blood pH to become low (acidic) from the normal pH of 7.4
due to the increase in the concentration of hydrogen ions (H +) that are produced by metabolic processes. In the cells
of the distal convoluted tubule, the carbon dioxide from aerobic respiration, catalyzed by carbonic anhydrase
enzyme, reacts with water to form carbonic acid, which dissociates into hydrogen carbonate ions. The hydrogen ions
are pumped into lumen where they are buffered by hydrogen phosphate (HPO 42-) as it takes up sodium ions to form
sodium hydrogen phosphate (NaH2PO4) which is excreted in urine while retaining the hydrogen carbonate ions.

Equation 1

Equation 2

Exceptional lowering of pH causes the cells lining the distal convoluted tubule to deaminate glutamine, amino acid,
to form ammonia. The ammonia then combines with hydrogen ions to form ammonium ions which are
excreted.Blood pH then rises (becomes less acidic) due to the absorption of the hydrogen carbonate ions that are
produced by the dissociation of carbonic acid. In order to control pH, the hydrogen carbonate ions are excreted
while the hydrogen ions are retained.

Diagram

Note;

 Within the plasma, hydrogen carbonate, proteins and hydrogen phosphate act as pH buffers by temporarily
taking up any excess hydrogen ions and at the same time keeping the pH constant
 The body maintains a constant pH by;

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i. Expulsion of carbon dioxide by lungs, which would accumulate and react with water to form carbonic
acid
ii. The buffering mechanisms involving plasma proteins in blood
iii. The kidneys expelling hydrogen ions and retaining hydrogen carbonate ions
 The acid-balance (pH) is maintained by the lungs, blood and thekidneys.

BUFFER SYSYTEMS

The internal pH ofmost living cells is close to 7. Even a slight change in pH can be harmful, because thechemical
processes ofthecell are very sensitive to the concentrationsofhydrogen and hydroxide ions. The pH of human blood
is very close to 7.4, which is slightly basic. A person cannot survive for more than a few minutes if the blood pH
drops to 7 or rises to 7.8, and a chemical system exists in the blood that maintains a stable pH.

If you add 0.01 mol of a strong acid to a litre ofpure water, the pH drops from 7.0 to 2.0. If the same amount of acid
is added to a litre ofblood, however, the pH decrease is only from 7.4 to 7.3. Why does the addition ofacid have so
much less ofan effect on the pH ofblood than it does on the pH of water? The presence ofsubstances called buffers
allows for a relatively constant pH in biological fluids despite the addition ofacids or bases. Buffers are substances
that minimise changes in the concentrations ofH+ and OH- in a solution. They do so by accepting hydrogen ions
from the solution when they are in excess and donating hydrogen ions to the solution when they have been depleted.
Most buffer solutions contain a weak acid and its corresponding base, which combine reversibly with hydrogen ions.
There are several buffers that contribute to pH stability in human blood and many other biological solutions. They
include;

a. The carbonic acid-bicarbonate buffer system

Carbonic acid (H2CO3) is formed when carbon dioxide (CO2) reacts with water in blood plasma, it then
dissociates to yield a bicarbonate ion (HCO3-) and a hydrogen ion (H+):

The chemical equilibrium between carbonic acid and bicarbonate acts as a pH regulator, the reaction
shifting left or right as other processes in the solution add or remove hydrogen ions.
 If the H+ concentration in blood begins to fall (that is, if pH rises), the reaction proceeds to the right
and more carbonic acid dissociates, replenishing hydrogen ions.
 When H+ concentration in blood begins to rise (when pH drops), the reaction proceeds to the left, with
HCO3- (the base) removing the hydrogen ions from the solution and forming H 2CO3

Thus, the carbonic acid-bicarbonate buffering system consists of an acid and a base in equilibrium with
each other. Most other buffers are also acid-base pairs.

b. Phosphate buffer system

This is a solution of HPO42- and H2PO4-
The dihydrogen phosphate (H2PO4-) dissociates into hydrogen phosphate (HPO42-) and hydrogen ions (H+).
Addition of small quantities of acid will cause the excess incoming hydrogen ions to react with hydrogen
phosphate ions forming dihydrogen phosphate.
Equation

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When an excess of the base is added, the excess incoming hydroxyl ions will react with hydrogen ions
forming water and leading to further dissociation of dihydrogen ions to replenish the hydrogen ions and
form hydrogen phosphate ions
Equation

c. Protein buffer system

This is due to the hydroxyl group and amino group attached to the amino acids that make up proteins.
When excess hydroxide ions are added to the solution containing amino acids, the carboxyl group of the
amino acid donates a proton which reacts with hydroxyl ions forming water thus lowering the pH. When
excess hydrogen ions are added, they react with the amino group forming ammonium ions thus raising the
pH to normal.

REGULATION OF BLOOD LEVELS OF IONS

The kidney closely regulates the concentration of ions such as Na +, K+, H+, Ca2+, Cl- and HCO3-. These ions are
important for their specific roles in various chemical processes, such as maintenance of protein structure, membrane
permeability, propagation of the nerve impulse and muscle contraction. Some are important in the regulation of the
pH of blood, which is also regulated, in part, by the kidneys. The concentration of particular types of ions in blood
and tissue fluid is regulated in three ways. Hormones control the;

i. Uptake of ions from the gut into the blood stream

ii. Removal of ions from the body by kidneys and elimination in the urine
iii. Release of ions into the blood stream from organs which contain them in high concentration

Regulation of calcium ions (Ca2+)

Low blood calcium level stimulate the parathyroid gland to secrete parathormone (parathyroid hormone) which
increases the calcium level and decreases the phosphate level by promoting;

 Bone break down by osteoclasts

 Calcium retention by the kidneys
 Excretion of hydrogen phosphate in the urine
 Activation of vitamin D, which in turn stimulates the absorption of calcium from the gut

High blood levels of calcium stimulate the thyroid gland to secrete calcitonin hormone, which increases bone build
u by osteoblasts so as to reduce calcium levels.

Note:

i. Calcium plays an important role in nerve conduction, muscle contraction and blood clotting
ii. Deficiency of the parathyroid hormone results in tetany i.e. shaking of the body due to continuous muscle
contraction caused by increase excitability of the nerves which fire spontaneously and without rest

Regulation of sodium ions (Na+)

A decrease in blood sodium leads to decreased blood volume and reduced blood pressure because less water is
drawn into blood by osmosis.

The juxtaglomerular complex, situated between the distal convoluted tubule and the afferent arteriole, is stimulated
to release the enzyme renin (not the digestive rennin). Renin catalyses the conversion of angiotensinogen, a plasma
protein into a hormone angiotensin, which stimulates the adrenal cortex to secrete aldosterone hormone.
Aldosterone has the following effects;

 Stimulates the active uptake of sodium ions from the glomerular filtrate into the plasma of capillaries
surrounding the nephron. This induces osmotic uptake of water into blood thus increasing blood volume and
sodium level back to the norm, accompanied by loss of potassium ions

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 Stimulates sodium absorption in the gut and decreases loss of sodium in sweat so as to raise sodium levels to
cause an osmotic in flow of water thus increasing the blood volume and pressure
 Stimulates the brain to increase the sensation of thirst

Increased sodium levels in blood cause increase blood volume and pressure less production of urine.

Note:

Low levelsof sodium in blood are detected by the hypothalamus, which stimulates the anterior pituitary gland to
secrete the andrenocorticotrophic hormone (ACTH), which also controls the activity of the adrenal cortex i.e.
secretion of aldosterone hormone.

OSMOTIC REGULATION

Increased concentration of solutes in blood (little water relative to salts) is detected by osmoreceptors in the
hypothalamus which stimulate the posterior pituitary gland to secrete antidiuretic hormone (ADH)/ vasopressin and
at the same time triggers the sensation of thirst, resulting into the drinking of water. ADH increases the permeability
of the distal convoluted tubule and the collecting duct to water, allowing the osmotic flow of water from the
glomerular filtrate into the cortex and medulla hence reducing the osmotic pressure of blood but increasing that of
urine. ADH also increases the permeability of the collecting duct to urea, enabling its diffusion into the medulla
tissue fluid where it increases the osmotic pressure, leading to the extraction of water from the descending limb.

Low solute concentration in blood (too much relative to salts) inhibits ADH release, tubule walls and collecting
ducts become impermeable to water, less water is reabsorbed from the glomerular filtrate into blood and a large
volume of dilute urine is passed out hence raising the osmotic pressure of blood.

Note:

 Diuresis is the production of copious dilute urine and antidiuresis is the opposite
 Insufficient production of ADH leads to a condition known as diabetes inspidus chacterised by frequent copious
urination
 Increase in blood osmotic pressure results from ingestion of little water, much sweating, ingestion of large
amounts of salt while a decrease in blood osmotic pressure may be due to little sweating, ingestion of large
volumes of water and little salt intake.

1. D.T.Taylor, N.P.O. Green, G.W. Stout and R. Soper. Biological Science, 3rd edition,
Cambridge University Press
2. M.B.V.Roberts, Biology a Functional approach, 4th edition, Nelson
3. C.J.Clegg with D.G.Mackean, ADVANCED BIOLOGY PRICIPLES AND
APPLICATIONS, 2nd EDITION, HODDER EDUCATION
4. Glenn and Susan Toole, NEW UNDERSTANDING BIOLOGY for advanced level, 2nd
edition, Nelson thornes
5. M.B.V.ROBERTS and T.J.KING, BIOLOGY a functional approach STUDENTS’
MANUAL, 2nd EDITION, Nelson
6. Michael Kent, Advanced BIOLOGY, OXFORD UNIVERSITY PRESS
7. J.Simpkins and J.I.williams, Advanced Human Biology, Collins Educational
8. Michael Roberts, Michael Reiss and Grace Monger, Advanced Biology, Nelson
9. Phillips W.D and Chilton T.J. A-level biology revised edition
10. Ann Fullick, Advanced Biology

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END

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