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Homeostasis

All cells in the body live in essentially the same environment - the
extracellular fluid. This accounts for one-third of all fluid in the body, or
about 20% of the entire body by weight
The extracellular fluid is rapidly transported throughout the body by the
circulation of the blood, and then mixed between the blood and tissue fluids
by diffusion.
No cell in the body is located more than 25-50 m from a capillary, therefore
there is rapid diffusion of substances between the blood plasma and the fluid
surrounding the cell
Because of this rapid mixing, the physical and chemical composition of the
extracellular fluid is effectively the same throughout the entire body - a
principle first recognised by Claude Bernard

Interstitial fluid + cererebrospinal


fluid + intraocular fluid etc.
Extracellular
fluid

Blood plasma

Blood

Total volume
approx. 40
litres

Blood cells

Intracellular
fluid

The term homeostasis (first used by Walter Cannon, 1920s) means


the maintenance of the physical and chemical conditions of the
internal environment i.e. extracellular fluid.
This ensures that all cells receive an adequate supply of nutrients and
oxygen, that there is no build-up of metabolites, and that the
temperature, pH, and electrolyte composition of the extracellular fluid
is maintained within the narrow limits that are essential for normal
metabolism

Some important constituents of extracellular fluid

Variable
Oxygen
Carbon dioxide
Acid-base
Sodium ions
Glucose
Temperature

Normal value

Normal range

40
40
7.4
142
0.85
37.0

35-45
35-45
7.3-7.5
138-146
0.75-0.95
36.7-37.3

Units
mmHg
mmHg
pH units
mmol/L
mg/mL
degrees C

Figs 1-1 and 1-2 in Guyton AC (1976)


Textbook of Medical Physiology
Saunders, Philadelphia.

Control systems maintaining homeostasis

To maintain homeostasis, the body must detect deviations in the


factors within the internal environment that need to be held within
narrow limits
For example, to maintain the concentration of oxygen at an acceptable
level, there must be sensors that detect and signal changes in oxygen
level, which then leads to appropriate adjustments in the respiratory
activity.
In this case, there are specialised receptors that respond to changes in
oxygen concentration within the carotid body, a small organ near the
carotid bifurcation in the neck.
A fall in oxygen concentration in the blood reflexly stimulates an
increase in the rate and depth of breathing, thus restoring the oxygen
concentration

input
(desired
output)

error
detector

+_

Controller

Controlled
system

output

Sensor
feedback
pathway

Types of control systems: intrinsic and extrinsic systems


Intrinsic controls are built in to an organ. Examples:
(1) Metabolic autoregulation of blood flow to exercising muscle.
This is a local mechanism that tends to maintain a relatively constant
level of oxygen and carbon dioxide in the extracellular fluid in the
muscle, despite changes in the metabolic activity of the muscle.

Autoregulation of blood flow to exercising muscle


(intrinsic control system)
Increase in muscle metabolic activity (in specific region)
Increase in consumption of O2 and production of CO2
Decrease in local O2 concn in and increase in CO2 concn in
extracellular fluid
Vasodilation of arterioles in local region

(2) Increased force of contraction of cardiac muscle in response


to increased stretch. This mechanism prevents over-filling of the
heart in circumstances when the rate of return of blood to the heart
increases e.g. during exercise.

Increase in local blood flow


Increase in rate of supply of O2 and removal of CO2
O2 and CO2 levels in extracellular fluid return to normal

Feed-back control

Entrinsic control systems


Extrinsic controls are more common. These are neural and hormonal
mechanisms, located outside the organ being regulated. Extrinsic
control permits coordinated regulation of several organs. Examples:

Restoration of blood pressure when it decreases, e.g. upon standing up.


Sensors are baroreceptors, and the autonomic nervous system elicits
coordinated effects on the heart and blood vessels.
Restoration of blood volume when it decreases. Sensors are located in large
blood vessels and heart. Both autonomic and hormonal systems are involved
(i.e. renal vasoconstriction, release of antidiuretic hormone).
Maintenance of body temperature. Sensors are located in skin and brain,
especially hypothalamus. Both autonomic and somatic motor systems are
involved; e.g. cold causes constriction of blood vessels in skin, piloerection
and shivering. Heat stress has the opposite effect i.e. dilation of blood vessels
in skin, sweating, panting.
All above cases are examples of negative feedback i.e. the controlling system
brings about an adjustment which is in the opposite direction to the initial
disturbance.

140

140

Blood
Pressur
e
(mmHg)

120
100
80

80

8080
80

Heart
Rate
(beats/min)

Subject stands up

70

60
6050

Reflex increase
In heart rate

10 sec
10 sec

Normal

Fig 58-2 in Boron WF & Boulpaep EL


(2003) Medical Physiology
Saunders, Philadelphia.

No baroreceptors

Figure 2

Fig. 21-6 in Guyton AC (1976)


Textbook of Medical Physiology
Saunders, Philadelphia.

Fig 58-4 in Boron WF & Boulpaep EL


(2003) Medical Physiology
Saunders, Philadelphia.

Fig 58-6 in Boron WF & Boulpaep EL


(2003) Medical Physiology
Saunders, Philadelphia.

Figure 4

Gain of a control system

This is a measure of the degree of effectiveness with which a control


system maintains constant conditions following a disturbance.
To illustrate, consider the case of the body temperature control
system.
Under normal steady-state conditions, body temperature in a person is 37. Suppose the outside

temperature increases by 20 (e.g. from 15 to 35).


If there were no compensatory mechanisms, the body temperature would be expected to rise by the
same amount.
However, as soon as the body temperature rises, this is detected by receptors in the body, and
compensatory mechanisms are activated so that the body temperature rises by a much smaller amount,
say 0.5.
Normal value of controlled variable
= 37
Disturbance = 20
Actual value of controlled variable
= 37.5
Error = 0.5

Figure 6

Gain of a control system (continued)

This is a high gain, reflecting the effectiveness of this control system.


Other physiological control systems often have much less gain e.g. the blood
pressure control system has a gain of only approximately 3. This is why the
blood pressure shows much greater fluctuations than body temperature.
What are the advantages and disadvantages of a high-gain system?

Compensation = -19.5
The gain of a control system is defined as follows:
Gain = Compensation/Error
Thus, in this case the gain is -19.5/0.5 = -39

Set-point of a control system

This is the level about which the physiological variable is controlled

the set point often varies, according to physiological state. For


example, during exercise, blood pressure and body temperature both
rises, but are still regulated about their new levels

Positive feedback
This occurs when a change in the variable results in a
response that, rather than compensating for the initial
change, results in a further change in the same direction.
Example: the nerve action potential
Depolarisation of nerve cell
Opens voltage-sensitive Na+ channels

Na+ ions flow into cell, down concentration gradient


0
threshold
mV
-90

Feed-forward control
EXAMPLE: exercise
motor cortex

Conscious, paralysed, artificially


ventilated human subject

150
130

hypothalamic and
brainstem centres

110
200
150

increase in
sympathetic activity

100

50

Figure 9

increase in
blood
pressure

100%

50%

25%

0%

(numbers indicate % maximum effort)

0
0

100

200

300

Time (s)

400

500

From Gandevia et al. (1993) J. Physiol. 470: 85-107