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HOMEOSTASIS

Teacher's Guide
The programs are broadcast by TVOntario, the television Note to Teachers
service of The Ontario Educational Communications The six programs in this series are arranged in order of
Contents
Authority. For broadcast dates consult the TVOntario increasing complexity, and should therefore be shown in
schedule in School Broadcasts, which is published in correct sequence. The series focuses on the principle of Program 1: COPING WITH CHANGE
September and distributed to all teachers in Ontario. The homeostasis at the level of the individual organism - the (An Introduction to Homeostasis). .................. 1
programs are available on videotape. Ordering information for original application of the term. Extensions of this principle Program 2: THE SEA WITHIN
videotapes and this publication appears on page 26. to all levels of life, from cell to biosphere, are mentioned in (Regulating the Body Fluids). ...................... 5
the last section of the guide, Program 6 : Hormonal Control.
Canadian Cataloguing in Publication Data Program 3: OSMOREGULATION
The guide suggests teaming activities to reinforce and (Fine-tuning the Control of the Body Fluids) ............ 9
Lang, Harold Murray ,1922 - complement each program, extending the teaming process
Homeostasis to make a complete teaching unit for senior biology. Program 4: THE FEEDBACK CYCLE
(A Controlling Principle for Homeostasis). ............ 15
To be used with the television program, Homeostasis.
Bibliography: p. The Series Program 5: BIOCHEMICAL BALANCES
Producer/Director: David Chamberlain (Regulation &f Body Chemistry). .................... 19
ISBN 0-88944-049-2
Writer: Alan Ritchie Program 6: HORMONAL CONTROL
1. Homeostasis (Television program) 2. Homeostasis. Narrator: Susan Copeland (The Coordination of Homeostasis). ................. 22
1. TVOntario. II. Title.
Consultant: H. Murray Lang
Ordering Information ............................... 26
QP90.4131984 574.1'88 84-093010-0 Animation: Groupe Imagination

The Guide
@Copyright 1984 by The Ontario Educational Communica-
tions Authority Project Leader. David Chamberlain
Writer' H. Murray Long
Editor: Elaine Aboud
Designer: Susan Mark

All rights reserved.

Printed in Canada
Coping With Change An Introduction to Homeostasis
Objectives year, Charles Darwin published On the Origin of Species by Means of
Natural Selection. Both of these principles have been equally productive for
Students should be able to: scientific work in their respective fields. Bernard showed that every organ-
i sm exists in two environments: an external one that is always changing, and
1. I dentify the need for organisms to control their internal environment.
an internal one, "le milieu interieur," that is carefully controlled to remain
2. Cite examples of controlled substances and conditions in organisms. nearly the same.
3. Recognize that a steady state is maintained by organisms within
Bernard: All the vital mechanisms, varied as they are, have only one
narrow limits through controlled fluctuations.
object, that of preserving constant the conditions of life in the internal
4. I dentify Claude Bernard as the originator of the concept of a steady environment. The independence conferred on an organism by its stable
state and a controlled internal environment (le milieu interieur), and internal environment sets it free to achieve its fullest development.
explain that he advanced this idea at the same time as Charles Darwin
The principle of the regulation of the internal environment within narrow
put forth a theory for natural selection in 1859. li mits was nameless until about 1930, when an American physiologist,
5. Define homeostasis, and identify Walter Cannon as the originator of
Walter B. Cannon, coined the term homeostasis. The word has Greek roots:
the term.
homoio meaning "the same," and stasis meaning "standing still:' Thus
6. Outline a model control system, and explain the interrelationships of
homeostasis may be defined as the preservation of constancy in the internal
its components: receptor, control centre, effector, feedback loop.
environment of an organism. It is often stated as the maintenance of a
steady state within an organism.
Program Description But the mechanisms of homeostasis do not maintain absolute and un-
changeable set-points. Usually there is a fluctuation above and below a set-
Since life began, every organism has had to struggle to maintain its life
point. For example, if you record your body temperature every two minutes you
against the fluctuation of environmental conditions. How do organisms cope might produce a graph as in Figure 1.
with extremes that may be life threatening? Homeostasis is about coping,
about the ways that organisms adjust their internal environments to com-
pensate for changes in the external environment. A sleeping dog is appar-
ently oblivious to its surroundings, yet internal receptors are constantly
monitoring conditions and making adjustments. If the temperature drops,
the muscles of the skin raise the hair to create more insulating air spaces.
Body
Shivering generates heat in the muscles below the skin. If the dog becomes
temperature
too hot, panting cools its body through evaporation of water from the lungs /OC\
and tongue. These automatic mechanisms help to regulate and maintain
body temperatures within narrow limits.
Besides temperature, organisms regulate many other factors within nar-
row limits. Consider blood sugar, for example. When a lion or tiger makes a
kill, it will gorge itself until filled. Once the prey has been consumed, it may
not make another kill for several days. Even humans are not always regular
i n their feeding habits: we may overeat on some occasion and skip meals at Figure 1
other times. Yet all organisms maintain the concentration of blood glucose
at much the same level at all times. Thus, when human body temperature is stated as 37°C, this figure repre-
The regulation of blood glucose and the regulation of body temperature sents the average, or mean, of many fluctuations. When 37*C is called
are just two examples of the principle of homeostasis. Claude Bernard, a
reknowned French physiologist, first suggested in 1859 the idea of a steady *From J. Olmstead and E.E. Olmstead, Claude Bernard and the Experimental Method
state: the maintenance of constant conditions within organisms. The same in Medicine (New York: Abelard Schuman Company Ltd., 1952), p. 224.

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"normal," it denotes a further averaging of many individual temperatures; receptor must do something to communicate what it has detected. Usually
yours may normally be above or below this mean. At different times of the it sends a message in the form of a chemical or nervous signal. This
day, and during different activities, human body temperature will differ When message reaches a control centre, which then selects an appropriate
we have a fever, and during ovulation, the set-point will be higher The response, and sends the right message to activate this response. This
analogy of the control of room temperatures by a thermostat may assist in message is transmitted to an effector, usually a gland or a muscle, or cilia or
understanding the fluctuations. flagella. I n an effective homeostatic mechanism, the effector, or its
Different kinds of organisms maintain different set-points for temperature response, must initiate a signal that will turn off the receptor or reset it for
control. Birds, for example, maintain their bodies about five degrees warmer further action. This step is called the feedback loop (Figure 4). In the
than mammals do due to increased metabolic rate. Birds and mammals program sych a control system is illustrated at the single cell level using the
maintain a nearly constant temperature, and an called homeothermic. The protist Euglena (Figure Q. Similar homeostatic mechanisms are effective for
graph of the body temperature of these organisms as the temperature of the
surroundings (Figure 2) is raised shows a flat part of the curve where
Control
homeostatic mechanisms can control temperature within narrow limits. Stress--* Receptor Effector ---* Response
centre
Beyond certain points the mechanisms can no longer protect the organism
from extreme temperatures. Feedback loop
Cold-blooded creatures show quite a different graph when the tempera-ture of their surroundings is raised (Figure 3). They are called poikilothermic
Figure 4
because their temperature varies with that of their surroundings.

Figure 3
Figure u

with change in a wide


Organisms cope range of factors in their surround-ings. If we plot graphs of these controlled entities against the changes in
the environment, in every case the curve wil resemble that of the homeo-thermic organism in a rising surrounding temperature. Thus, homeostatic

mechanisms control the total water content of the body, the levels of
glucose, sodium, calcium, hydrogen, and potassium ions in the blood, and
the arterial blood pressure. How do such mechanisms work?
For each control system, there must be a receptor that can detect the
change in or stress on the organism. It might also be called a sensor. The Figure 5. Euglena, a single-celled protoctist (protist)
organisms that function at the tissue level, such as fungi, sponges, and Activities 1, 2, and 3 before viewing, in order to better prepare the students
plants. Higher animals have more highly developed responses, based on for the program. If students have not studied Euglena in earlier grades, it
nervous systems, and these will be illustrated in later programs. would be a good idea to teach its structure, and to show how it can function
From single cells to large complex organisms with trillions of cells, as either an autotroph or a heterotroph.
homeostatic mechanisms are at work to maintain life. The thermostat analogy (Activity 3) reveals the whole basis of homeo-
stasis. It should be discussed prior to viewing Program 1.
Before Viewing
Program 1 can stand as its own introduction to the principle of homeostasis.
After Viewing
However, students will be in a better position to understand the concepts Have students discuss the concepts presented in order to consolidate their
involved if they have previously studied the physiology of several vertebrate learning. Then discuss the review questions (Activity 4).
(or mammalian or human) systems. It would be advisable to complete

Activities place the bulb of the thermometer under your


tongue in exactly the same position for exactly
Activity 2: Phototaxic Behavior of
60 s; then wait exactly 60 s before inserting the Green Flagellates
thermometer again. Plot the results on a graph
Activity 1: The Fluctuations in (Figure 6). Materials:
Human Body Temperature 2. Exercise moderately for 2 min by jogging "on 2 culture jars of living Euglena (or Chlamydomonas)
the spot:' Record your temperature immedi- black paper, scissors
Materials: ately after you stop exercising, and again after compound microscopes, well slides, cover slips
1 clinical thermometer per team 5 and 10 min. Plot these temperatures on a droppers, methyl cellulose
clock or watch that shows time in s graph similar to Figure 6.
3. Exercise vigorously for 2 min by running up
Method:
Method: and down stairs. Record your temperature 1. Several hours before class cover one of the
1. After sitting still for at least 5 min, record your i mmediately after you stop, and again after 5 culture jars with black paper except for a hole
temperature every 2 min for a period of 10 min. and 10 min. Plot these temperatures on a graph that you cut in the shape of an initial or a circle
For accuracy, standardize your method: shake similar to Figure 6. 1 cm in diameter. Leave -the other culture
the thermometer down each time until the fluid uncovered, as a control. Place both cultures in
reaches the same low point on the scale; then Questions: a fairly bright location.
What does the graph show about your resting 2. When the class is assembled remove the black
body temperature? What hypothesis can you paper. Have students compare the distribution
suggest to explain your observations? Do you of the organisms in the jars. Ask for
have enough data to calculate your "normal" suggestions of hypotheses to account for the
temperature? observations.
What are the immediate and long-term results 3. Students should examine some of the
of moderate and vigorous exercise? Are these organisms through a microscope by placing a
results consistent with those of other small drop of the culture in a well slide. Have
members of the class? Suggest what is them make a sketch of a typical organism.
happening in your body that would explain Encourage them to observe the organisms for
your observations. several minutes to find out how they move. If
the organisms move too quickly for study, add
Time (s)
Figure 6. Graph of changes in body temperature

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a small drop of methyl cellulose solution to 5. What function might be performed by the
slow them down. Can students detect any contractile vacuole?
Activity 4: Review Questions
other changes in the organisms? 1. Why do organisms need to control their internal
Questions: Activity 3: Thermostat Analogy environment?
2. What entities (conditions and substances) are
1. How do the organisms respond to light? • How does a thermostat work in controlling room regulated by organisms? (State one condition, two
2. What advantage would this behavior give the temperature? molecules, and two ions that are controlled.)
organism? • In what ways might the operation of the thermostat 3. Define homeostasis.
3. What structures do the organisms have that be analogous to temperature control in the body 4. Who first suggested the idea of a controlled internal
might detect light? (Figure 7)? Conduct a class discussion in the environment? What advantage did he see for the
4. What structures do the organisms have that pattern set by Joseph Schwab in "Invitation to organism if it controlled its internal environment?
would effect a response to light? How do these I nquiry No. 38" in Biology Teacher's Handbook (see How was his idea a useful principle for physiology?
structures function? Mayer, Further Reading). 5. Outline a model control system and explain how
each part of it would function in maintaining
homeostasis for an organism.
Room temperature rises (Electrical signals) Room temperature drops
above set-point Furnace turns off to set-point Further Reading
Berry, Gordon S. Biology of Ourselves - A Study of
Human Biology. Toronto: John Wiley and Sons,
Canada, Ltd., 1982.
Mayer, Wm V., ed. Biology Teacher's Handbook. 3rd ed.
Thermostat set New York: John Wiley and Sons, Ltd., 1978.
Nelson, G.E.; Robinson, G.G.; and Boolootian, R.A.
at 20°C Fundamental Concepts of Biology. 2nd ed.
New York: John Wiley and Sons, Ltd., 1970.
Olmstead, J., and Olmstead, E.E. Claude Bernard and
the Experimental Method in Medicine. New York:
Abelard Schuman Company Ltd., 1952.
Roddie, l. Physiology for Practioners. Edinburgh:
Churchill Livingston Company, 1975.
Sherman, I.W, and Sherman, VG. Biology - A Human
Approach. New York: Oxford University Press, 1979.
Weisz, Paul B. The Science of Biology. 4th ed.
Figure 7. Chalkboard discussion of thermostat model New York: McGraw-Hill Book Company, 1971.
The Sea Within Regulating the Body Fluids
Objectives I n contrast to the single cell, multi-cellular organisms maintain two
different internal environments:
Students should be able to: • the fluid within cells - the intracellular fluid (IGF); and
• the fluid that surrounds and bathes the cells - the extracellular fluid
1. Identify the importance of the control of water within an organism, and
(ECF).
the "compartments" in which water occurs: the extracellutar fluids
(ECF) and the intracellular fluids (ICF).
2. Identify MacCallum's hypothesis: the similarity in composition of the
body fluids of diverse organisms suggests that life must have
originated in the sea, and has maintained fluids resembling that sea
ever since.
3. I dentify three ions and three molecules in the body fluids that are
maintained homeostatically within narrow limits.
4. Explain the processess of osmosis, diffusion, exocytosis, and endocy-
tosis as homeostatic mechanisms at the cellular level.
5. Explain the hypothesis of the mechanisms ("pumps") that maintain the
concentrations of sodium and potassium in the ICE

Program Description
Program 2 deals with Claude Bernard's "le milieu infrieur," the internal
medium within the organism. Bernard's idea was that the organism
maintained a steady state within itself as protection against extremes of
change in the external environment. Water forms a large proportion of every
organism, but its maintenance within controlled limits poses problems.
Consider, for example, the single-celled amoeba that lives in freshwater
ponds. Within its cell are a number of solutes, including sugars and the ions
of several salts. By osmosis, water is constantly entering through the cell
membrane from outside, where there are more water molecules per unit
volume of solution. To keep from becoming too dilute,or from bursting (as
red blood cells do in pure water), the amoeba has developed a pump that
gets rid of the excess water. This is the organelle - the contractile vacuole.
It gradually fills with water, then contracts to expel its contents. Then it
repeats the process. Diffusion, the process by which substances move by
molecular action from where the concentration is high to where it is lower,
accounts for the movement of oxygen and carbon dioxide across the cell
membrane of the amoeba Ions needed for life, such as potassium and
calcium, will be pumped across the cell membrane into the cytoplasm of the
amoeba by molecular pumps that use energy to accomplish this active
transport. Thus, at the level of the single cell, the amoeba regulates its cell Figure 1. A comparison of the concentrations of ions in sea water and in the body
content to maintain life. fluids (ECF) of several organisms, using sodium as a standard (100 units)

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Thus the ECF is analogous to the pond in which the amoeba lives in that it receptors: dilution receptors, volume receptors, and pressure receptors.
provides a watery medium for every cell, bringing the substances needed for Homeostatic controls are set in motion, and within three hours most of the
life and carrying away the wastes. excess load is removed, excreted as urine (Figure 2e).
A comparison of the chemical compositions of the ECF of quite different
organisms reveals the surprising fact that the ions are very similar in kind
and in concentration (see Figure 1). In fact, their concentrations are quite
similar to those in sea water. This prompted a Canadian physiologist, A.B.
MacCallum, to suggest about 60 years ago that these similarities are
evidence that life orginated in the sea Did homeostatic mechanisms evolvp
so that organisms could maintain their internal environments much like that Figure 2d Figure 2e
of the early seas? After about three billion years of evolution, do we still
carry around within us an environment resembling that ancient sea? Today A comparison of the concentrations of the various ions in the body fluids
the sea has changed somewhat; all the rivers have been carrying salts and (see Activity 2) reveals that there is little change between the plasma and the
i ons from the land, which make the ocean saltier that when life began. fluid between the cells. Apparently, fluids pass readily from the blood
On average, about two-thirds of the human body is water, although this is vessels to the body spaces and vice versa because of the porous nature of
higher in newborn infants, and lower in women, because more fats are capillary walls. There is, however, a marked difference between the ECF and
stored. Water is the solvent and vehicle of the body fluids. The water in the the ICE The cell membranes evidently exert an active control on what gets
body is distributed in three main "compartments" or regions. Two-thirds of i nto or out of cells. Cell membranes must use energy in the form of ATP to
the water is found in the ICE, that is, inside the body cells (Figure 2a). Of the pump the needed ions in, and the undesirable ions out. `
remaining one-third, one-quarter makes up the blood plasma, while the rest Potassium is a very important ion within cells. It helps to maintain the
i s in the fluids surrounding cells-, mostly as lymph (Figure 2b). There is a intercellular osmotic pressure and to regulate pH. It promotes reactions
dynamic equilibrium between the three compartments; by osmosis and necessary in carbohydrate and protein metabolism and plays a vital role in
other mechanical processes, water flows readily from one region to another membrane polarization involved in the conduction of nerve impulses and the
(Figure 2c). contraction of muscle fibres. Ninety-eight percent of the body's potassium is
i n the cells, only two percent is outside. Most of the potassium is bound to
ECF I CF
proteins in the cells; the rest of it is attracted to inorganic phosphate ions:

1/3 2/3

Figure 2a Figure 2b

Figure 2c

" When we drink a large quantity of some beverage, we readily absorb The average human adult consumes from 2 to 3.5 g of potassium per day
much of the water through the linings of our stomach and intestines. For a i n food. About the same amount is excreted every day. This amounts to a
ti me, the volume of blood increases and our body fluids become more dilute daily turnover of from 1.5 to five percent of the body's total potassium. The
(Figure 2d). The extra stress on our system is detected by three kinds of l osses are greater when we suffer disease, trauma, or surgery. The body
i s hypothesized that there are active transport pumps through the cell
membrane. Perhaps the pump works by expelling sodium ions as it takes in
potassium ions: each of these ions is about 30 times more concentrated on
the opposite side of the membrane. Figure 3 shows a possible mechanism
to explain the action of the pump.

Before Viewing
Students should do Activities 1 and 3 in order to understand the difficulties
that cells face in a watery environment and how unicellular organisms cope
with them. Decide also whether it would be better to have students consider
the data in Activity 2: the relative concentrations of different ions in body
fluids. The data will go by so quickly on the screen that advance
consideration may be useful. The units "milliequivalents" may need to be
explained (see footnote to Activity 2).

After Viewing
Figure 3 Students should complete the activities and discuss the points raised in the
program. They may write notes based on the concepts by answering the
does not store potassium, but requires a daily intake to supply its needs. In
review questions.
order to keep 30 times as much potassium inside the cells as in the ECF, it

Activities Activity 2: A Comparison of the Ion Concentrations in the Fluids of the


Human Body
Activity 1: The Effects of the Method: fluids. Compare and contrast the concentration
Environment on Cells 1. of each ion to determine what mechanisms are
Examine the following table of ion concentra-
at work to cause the differences.
tions in the three "compartments" of the body
This activity, described in Inquiry 18, The Functioning
Animal, p.43 (see Further Reading), asks students to CONCENTRATIONS OF IONS IN THE FLUIDS OF THE HUMAN BODY
i nvestigate the effects of different surrounding fluids i
(expressed n relative units, see P. a-)
on red blood cells. Students find that the red blood
cells, if they are to remain normal, must be surround- EXTRACELLULAR FLUID (ECF) I NTRACELLULAR FLUID (ICF)
ed by plasma that is close to a 0.9 percent solution of IONS Blood Plasma Between Cells Within Cells
sodium chloride in molarity. In pure water, the cells
absorb too much water by osmosis and burst, a condi- Sodium 142 145 5
tion called hemolysis. On the other hand, cells Potassium 5 4 150
surrounded by a hypertonic solution, such as ten Calcium 5 3 1
percent sodium chloride, become plasmolyzed and Magnesium 3 2 40
shrink, a condition called crenation. Chloride 104 116 5
Bicarbonate 27 27 10
Hydrogen Phosphate 2 3 110

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Questions: 2. Endocytosis: the engulfing of food into a cell. 6. Using the fluid mosaic model of a cell membrane,
1. Why is there a close similarity between the Add a drop of a smaller organism, such as explain how potassium and sodium ions might be
figures in the first and second columns? Colpidium or Paramecium, to the slide contain maintained in suitable concentrations by pumps in
2. What must the cell membranes be doing to i ng amoeba Observe carefully for several the membrane.
maintain the low concentration of sodium ions minutes, watching the behavior of the amoeba

3.
within cells?
Are there any other ions that show a pattern
When the smaller organism touches or
approaches the amoeba, how does the cell Further Reading
similar to that shown by the sodium ions? membrane react? What behavior of the inner Lang, H.M; Palfery, E.G.; and Van Nieuwenhove, E.L.R.
What relationship might there be between' protoplasm follows the stimulus to the The Functioning Animal, Toronto: Gage Educational
these and the sodium ions? membrane? How quickly are pseudopods
4. Contrast the concentrations of the potassium formed? How do they form a food vacuole? Publishing, 1978. McElroy, W13.; Swanson, C.P; and Macey, R.I. Biology
ions with those of the sodium ions. What must What happens to the organisms inside the and Man. Englewood Cliffs, N.J.: Prentice-Hall Inc.,
the cell membranes do to maintain the concen food vacuole? 1975.
tration of potassium ions within the cells? Note: If living cultures cannot be obtained, show a Mikal, Stanley. Homeostasis in Man: Fluids, Electro-
What other ions are similarly concentrated with short film of the functioning of the amoeba Several lytes, Proteins, Vitamins and Minerals in Clinical
cells? fil ms are available: Amoeba (Gaumont-British); The Care. Boston: Little Brown and Co., 1967.
Amoeba (Educational Pictures); Response in a Simple Wallace, Robert A. Biology, The World of Life. 2nd ed.
'The units are not important here; the concept is the relative Animal (BFA Educational Media); The Biology of the (Chapter 9) Santa Monica, Calif.: Goodyear Pub. Co.
concentration. The measurements given are expressed in Amoeba (Statens). 1978.
milliequivalents (mEq.). These are calculated by multiplying Weisz, Paul B. The Science of Biology. 4th ed. (Part 6:
the mass of ions (mg) per L by the valence, and dividing by Steady States) New York: McGraw-Hill, 1971.
the molar mass. This gives a measure of the number of Activity 4: Review Questions
ionic charges per L of solution.
1. Why do organisms need to control the amount of
water they contain? (What are the consequences of
Activity 3: Examination of Amoeba too much or too little water?)
to Observe Exocytosis and 2. In a multicellular organism, where is water located?
Endocytosis Name the various "compartments" in which water
occurs.
Materials: 3. What ions are similar in their concentrations in
culture of living Amoeba different organisms and in the seas? What hypo-
culture of living Colpidium or Paramecium thesis is suggested by these similarities?
compound microscopes 4. For each of the following terms (i) define the term,
microscope slides (or well slides), cover slips (ii) give one example of a substance that would
enter or leave a Gel! by the process and (iii)explain
droppers how the process contributes to homeostasis:
Method: (a) osmosis
1. Exocytosis: the expulsion of wastes from the (b) diffusion
cell. (c) exocytosis
Examine the amoeba. Locate the contractile (d) endocytosis
vacuole, and observe it steadily for several 5. What ions does a cell membrane:
minutes. How long does it take for the contrac- (a) concentrate within the cell?
tile vacuole to fill? How long does it take to (b) pump out of a cell?
empty? How soon does it begin to fill again?
Osmoregulation Fine-tuning the Control of the Body Fluids
Objectives nephrons flows together on its way to be excreted. In turn, the convoluted
tubule has three parts: the proximal tubule, nearest to the capsule; the loop
Students should be able to: of Henle, and the distal tubule that connects to the collecting region.
1. Define osmoregulation and identify the parts of the body involved in
the process and its control.
2. Describe the structure, location, and function of each part of the
kidney and each part of its unit of structure - the nephron.
3. Explain the role of osmosis in the reabsorption of useful fluids.
4. Explain the role of hormones in the control of reabsorption.
5. Explain the adaptations of the excretory systems of animals that live in
quite different environments: marine invertebrates, marine fish, fresh-
water fish, and mammals.

Program Description
An important aspect of homeostasis is the careful regulation of the
composition of the body fluids, a process called osmoregulation because it
involves the movement of water across membranes. Osmosis is determined
by the concentration of solutes on both sides of a membrane, and is a
passive process. This means that osmosis is a movement of water based
entirely on the behavior of molecules. This is quite different from active
transport, which depends on the expenditure of energy, in the form of ATP
Both osmosis and active transport are involved in osmoregulation.
I n vertebrates, the kidneys are the main organs of osmoregulation. Fifteen
percent of the blood being pumped out of the heart is directed into the
kidneys. Here the pressure of the blood forces 20 percent of the plasma to Figure 1. A nephron
l eave the blood capillaries. This creates a filtering action, because the blood
cells are kept in the capillaries when some of the plasma leaves. The fluid I n the proximal tubule, about 80 percent of the filtrate is reabsorbed: the
that leaves the blood amounts to 180 L per day. This would amount to a glucose, amino acids, hormones, vitamins, and ions. By active transport, the
-t because the body cells lining e u e move these materials into the cells and return them to
needs: water, many kinds of ions, glucose, and amino acids. However, as the the blood in the capillaries surrounding the tubule. In addition, water reab-
filtrate enters the kidney tubules, the processes of reabsorption begin, and sorption by osmosis occurs in response to the reabsorption of the solutes
by the time it has passed completely through the nephron, 99 percent of the by active transport. Next, in the loop of Henle, the lining cells pump sodium
fluid will have been returned to the blood. The remaining one percent will be i ons into the tubule by active transport. This occurs because of the high
excreted as urine. This process of reabsorption by the cells lining the concentration of sodium ions in the medulla region of the kidney. This
nephrons is selective; it regulates the concentration of the extracellular fluid. changes the osmotic relationship between the filtrate and the ECF surround-
Each human kidney contains about a million nephrons. A nephron (Figure i ng the tubules and capillaries, making it possible to keep reabsorbing water
1) consists of three main regions: Bowman's capsule where the filtrate from the filtrate. The action is called the counter-current model, since the
leaves the knot of capillaries; the convoluted tubule where selective reab- two arms of Henle's loop run counter to one another, and work in opposite
sorption occurs; and the collecting region where the urine from many . ways (Figure 2). In the first part (the descending arm), water diffuses out of

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the tubule, and sodium is pumped into it. The changes make the filtrate Another hormone, ADH (anti-diuretic hormone), produced in the pituitary
increasingly hypertonic, or more concentrated with respect to the ECF In gland, acts on the membranes of the cells in the distal tubule to maintain
the second part (the ascending arm) of the loop, sodium ions are actively the osmotic pressure of the ECF If the body fluids become too dilute, then
pumped out of the filtrate. Now, hormones from the adrenal cortex, chiefly the secretion of ADH is inhibited, and more water is excreted in the form of
aldosterone, affect the membranes of the cells lining the tubule, making dilute urine. If, on the other hand, the ECF becomes too concentrated, then
them more permeable to absorb water from the filtrate." the pituitary releases more ADH into the blood. When the ADH reaches the
cells of the distal tubule, it increases the reabsorption of water, reducing the
volume of urine. Perhaps you have experienced the action of caffeine or
' alcohol: both inhibit the release of ADH, resulting in diuresis, the production
of large quantities of dilute urine.
The hormones that control the action of the nephron are triggered by
several different receptors. There are osmoreceptors in certain centres of
the hypothalamus of the brain and in the wall of the carotid artery; blood
volume receptors next to the glomerulus within the Bowman's capsules of
the nephrons; and blood pressure receptors in the atria of the heart. In
Program 6 of this series, you will learn how such receptors interact within
the endocrine system to maintain homeostatic control. It appears as if
several alternate systems have evolved to protect life through fine-tuning
osmoregulation. Figure 3 summarizes the interactions of the different
controls and responses.
If we compare the process of osmoregulation in several organisms from
different environments (Figure 4), we see evidence of the evolution of
increasing adaptations. Such minor adjustments of structure and function
have made possible the exploitation of a wide variety of difficult environ-
ments. In many marine invertebrates, such as jellyfish and sea cucumbers,
the body fluids tend to match the sea water in composition and osmotic
properties. In this isotonic state, they can maintain osmoregulation without
much expenditure of energy. They neither gain nor lose water and salts by
osmosis or diffusion.
Many marine invertebrates have adapted to life in estuaries or river
mouths where fresh water mingles with salt water. Because these organisms
Figure 2. Reabsorption in the uriniferous tubule of the nephron
contain more salts in their body fluids, fresh water poses a problem for them
because by osmosis they constantly take in water from their surroundings.
I n the last or distal part of the tubule, the action that began in the ascend-
To cope with this problem many different adaptations have evolved. For example, barnacles and mussels close up tightly when th eyaresurounde
i ng loop of Henle continues. Here, in addition to the continued reabsoption
of water, the same hormone - aldosterone - regulates the balance of by fresh water, and open up only when salt water returns. Some crabs have
several ions: hydrogen, potassium, ammonium, and magnesium. Thus the
developed special organs at the bases of their antennae for excreting the
function of aldosterone is to protect the volume and composition of the
excess water.
body fluid. The net result of all the reabsorption is that the urine becomes You might expect marine fish to resemble marine invertebrates in having
increasingly concentrated in solutes (hypertonic) as water is reclaimed.
i sotonic body fluids, but this is not the case. Actually, marine fish have body
fluids with ion concentrations more like those of the freshwater fish and
`Although aldosterone is not mentioned in the program because of time limitations, l and vertebrates. Evolutionists interpret this as evidence that the ancestors
we discuss it briefly here in order to provide further background information. of marine fish must have evolved in fresh water and later adapted to life in
Figure 3. Controls and processes involved in osmoregulation

the sea This adaptation required them to solve the problem of the continual Land-dwelling vertebrates face quite a different problem: how to retain
loss of water to the sea by osmosis. They would also tend to absorb salts enough water in their ECF to keep from drying up. They lose water by
too readily from the sea To cope with the problem of hypotonic fluids, evaporation from the body surfaces, from the breathing organs, and through
marine fish have to drink a lot of water and excrete excess salt through their the elimination of wastes. They must also maintain the balance of salts in
gills. In their kidneys, he nephrons lack the glomerull (knotsof capilaries) the within the proper limits for life . The adaptations of the human
for filtering, but have many capillaries surrounding the tubules for reabsorp- osmoregulation system apply to most land vertebrates, although there are
tion of water. Marine fish excrete isotonic urine, scanty and concentrated in i nteresting modifications for life in desert animals, such as the camel, the
wastes and salts. kangaroo rat, and the gerbil.
Freshwater fish have body fluids that are hypertonic to water, that is, there You may wonder how changes in the structure and functioning of a
are higher levels of solutes in their ECF than in their surroundings. Their nephron might come about. It is hypothesized that particular hormones,
problem is that water will constantly be entering their ECF by osmosis. They such as ADH, affect only target cells that have specific protein molecules in
rarely drink water. Their body coverings of scales limits the entry of water to their membranes. In this way, the hormone affects only the permeability of
the membranes of their mouths and gills. They have developed efficient the target cells, and has no apparent effect on other cells in the body. ADH
kidneys with large glomeruli for excreting the excess water. They excrete causes a change in permeability of the target cells, making them more
l arge quantities of dilute (hypotonic) urine. Their gill membranes take in the permeable to reabsorb water and specific ions. This change is reversible,
ions of salts from the water by active transport. adjusting for changing conditions of the ECF The change in permeability

11
SEA CUCUMBER 1. FRESHWATER FISH

2 MARINE FISH

3_ AMPHIBIANS, REPTILES
Scanty, concentrated urine

4. MAMMALS

MAMMAL

Figure 4. Osmoregulation in different environments Figure 5


may be due to the action of a specific enzyme, succinic dehydrogenase, an increasingly hostile environments. The remaining programs will examine in
enzyme that removes hydrogen from succinic acid. This action operates the more detail some of the ways that homeostatic mechanisms became
sodium pump, moving sodium ions across the cell membrane. Russian fine-tuned through biochemical controls.
physiologists have suggested that this hormone-enzyme-permeability
system may account for the evolution of the function of reabsorption for
different environments. In freshwater fish the enzyme is only active in the
Before Viewing
distal part of the uriniferous tubule, reabsorbing very little water (Figure 5). In Students should do the first two activities before viewing Program 3, in order
marine fish the distal tubules are reduced, and the activity of the enzyme is to gain a thorough understanding of the details of kidney structure and
very low. Here sodium is reabsorbed only when the filtrate is isotonic with functioning.
the blood. In amphibians and reptiles the main activity of the enzyme is still
in the distal tubules, but there is some activity in the proximal tubules as
well, enhancing the retention of water for a land environment. In birds and After Viewing
mammals there is increased activity of the enzyme in the proximal tubules,
There are a number of good activities related to osmoregulation; two are
but the strongest action is in the ascending part of the loop of Henle, and in
described in Activities 3 and 4. In addition, you might enjoy using the inquiry
the distal tubules. Thus we see how adaptation for life out of the water has fil mloop of the Biological Sciences Curriculum Study: The Kidney and
been made possible by minor changes in molecular activity that increased Homeostasis (available from Gage Educational Publishing or Longman
the efficiency of the control of the water and salts in the body fluids.
Canada Limited). A similar form of inquiry in print is the Nuffield Biology
Thus as organisms adapted to a wide range of changing environments, Program, Book 2: Living Things in Action, Chapter 15 (see Further Reading).
they continued to achieve homeostasis by fine-tuning of systems that were
already present. The net result was an even greater freedom to exploit

Activities Activity 2: Active Transport in Activity 4: A Comparison of


Living Cells Adaptations in Kidneys of
Activity 1: A Study of the Structure I n Benson et al., Investigations in Biology, I nvestiga-
Different Organisms
of the Kidney tion 13 asks students to investigate the uptake of
Aspects of the evolution of kidneys for life in different
congo red dye by the membranes of yeast cells, both
Many laboratory manuals explain how to dissect fresh environments are well described in Galbraith and
alive and dead.
Wilson, Biological Science: Principles and Patterns of
kidneys (pork or lamb are most like humans') from a
Life, pp. 317-19 and 449-51 (see Further Reading).
slaughterhouse or meat counter. Some teachers use
preserved kidneys from a biological supply company.
Activity 3: The Composition of Some of the same ideas may emerge in Crawford,
Some sources of laboratory instruction are: Urine Patterns in Biology, p. 387, Inquiry Investigation: "Is
Benson et al., Investigations in Biology, I nvestiga- Homeostasis -Common to All Vertebrates?" - and p. 390,
tion 37. Almost identical instructions are given in the following I nquiry Activity: "Variation in Kidney Tubules in
Crawford, Patterns in Biology, p. 389, Laboratory books, but the first tests for two additional ions: Vertebrates." There is also an enrichment column on
Activities 2 and 3. Benson et al., Investigations in Biology, I nvestiga- the camel. See also the textbooks by Volpe and by
Galbraith, Lab Manual for Biological Science tion 38. Wallace in the reading list.
(revised), Investigation 32. Crawford, Patterns in Biology, p. 397, "The Physical
(See Further Reading, Program 3) and Chemical Analysis of Urine."
Lang et al., The Functioning Animal, I nquiry 19. Activity 5: Review Questions
(See Further Reading, Program 2)
1. Define osmoregulation. In what organs of the
human body are the processes involved in
osmoregulation and its control carried on?

13
14

2. Make a labelled diagram to represent the structure


of the human kidney as seen in longitudinal section, Further Reading
and showing its connections to the circulatory Benson, Garth D., et al. Investigations in Biology. Don
system and to the urinary bladder. Beside each label Mills, Ontario: Addison-Wesley Publishers, 1977.
state the function of that part.
Borow, Maxwell. Fundamentals of Homeostasis. 2nd
3. What is a nephron? Make a labelled diagram to
ed. (Chapter 3) Flushing, N.Y: Medical Examination
show the parts of a nephron and their relationships
Publishing Company, 1977.
to the circulatory system.
Crawford, Ian. Patterns in Biology. Toronto: McGraw-
4. Describe in detail the functioning of each part of the
Hill Ryerson Ltd., 1983.
nephron.
Galbraith, Donald I. Lab Manual - Biological Science
5. Explain the role of osmosis in the reabsorption of
Principles and Patterns of Life. Revised ed. Toronto:
water in the nephron.
Holt, Rinehart and Winston of Canada, Ltd., 1976.
6. What is the role of hormones in the control of
Galbraith, D.I., and Wilson, D.G. Biological Science
osmoregulation?
Principles and Patterns of Life. 3rd ed. Toronto:
7. Explain the adaptations of excretory systems of the
Holt, Rinehart and Winston of Canada, Ltd., 1978.
following organisms that permit them to live in quite
Ginetzinsky, H. `The Role of Hyaluronic Structures in
different surroundings:
(a) marine invertebrates the Evolution of Water Excretion:' In The
Development of Homeostasis. London: Academic
(b) marine fish
Press,1960.
(c) freshwater fish
Nuffield Foundation. Living Things in Action. Revised
(d) land-dwelling vertebrates
ed. London: Longman,1974.
Volpe, E. Peter. Man, Nature and Society. An
Introduction to Biology. 2nd ed. Dubuque, Iowa:
Wm. C. Brown, 1979.
Wallace, Robert A. Biology, The World of Life. 2nd ed.
Santa Monica, Calif.: Goodyear Publishing Co., 1978.
The Feedback Cycle A Controlling Principle for Homeostasis

Objectives Negative feedback, on the other hand, maintains stability. It does this by
causing the product of the system to shut the system off. I n a fluctuating
Students should be able to: system negative feedback succeeds in keeping the swings within proper
li mits (Figure 2). Even if the environment adversely affects an organism with
1. Distinguish between positive and negative feedback, and identify the an original wide fluctuation from normal conditions, negative feedback will
consequences of each for an organism. restore equilibrium. Thus negative feedback is the adaptive mechanism that
2. Explain the role of negative feedback in maintaining homeostasis. maintains life i n every organism (Figure 3).
3. I dentify the location of temperature receptors in the body.
4. Describe five responses initiated by the hypothalamus when body
temperatures rise or fall appreciably.
5. Identify the advantages that a nervous system gives animals for coping
with change.

Program Description
The first program in this series showed a model of a basic control system,
consisting of a receptor, a control centre, and an effector To complete the
system, a feedback message was needed to inform the receptor that the
response had been completed and to stop sending its message. In this Figure 2
program feedback mechanisms are examined in more detail.
There are two kinds of feedback: positive and negative. Positive feedback
enhances the original signal, making a stronger response. It is like the howl
you hear in a sound amplifying system when a microphone picks up the
sound from its own speaker and magnifies it out of control. In a system that
fluctuates, positive feedback causes the increasing swings, leading to insta-
Figure 3
bility (Figure 1). In a living system, positive feedback leads to death, since
the organism can no longer control its environment within narrow limits. An
To illustrate the feedback principle this program reviews the homeostasis
example of positive feedback in an organism is severe shock.
of temperature control which was discussed in Program 1. The receptors for
heat and cold are sensory cells in the skin and in the hypothalamus. The
hypothalamus is located deep in the head at the base of the brain (see.
Figure 4). This i s the region where nerve fibres cross over from the spinal
cord to the brain, and from the eyes to the opposite sides of the brain. Thus
the hypothalamus is at a very important junction of many nerve fibres. It
contains 16 clusters of nerve cells that are concerned with many aspects of
homeostasis: controlling hunger, thirst, and the response to fright, as well as
regulating the temperature and composition of the blood. These centres
detect misalignment of the different components of the steady state and
respond by producing nerve impulses and hormones. Thus the hypothala-
mus contains many control centres, and allows the master control function
Figure 1
to shift from one centre to another as the need arises.

15
16

not effective. We notice the gooseflesh, the little bumps in the skin made by
the erector muscles at the base of each hair. This response might be called
a vestige of an evolutionary change, because it suggests that we likely are
descended from ancestors that had more hairy bodies. Both of the first two
responses are aimed at conserving the body's supply of heat by restricting
further heat loss.
The third effector of response is in the deeper muscles of the skin. Nerve
i mpulses make these muscles contract in waves that we call shivering.
The contraction of the muscles releases stored energy and thus generates
heat. Shivering helps warm the body because blood flowing through these
muscles will soon be on its way to the rest of the body.
There is still another response - a fourth way to keep warm. A cluster of
cells in the hypothalamus produces a releasing hormone. This enters the
blood capillaries and is carried the short distance down the stalk to the
anterior lobe of the pituitary gland. Here the releasing hormone stimulates
certain cells to produce another hormone: thyroid stimulating hormone (or
TSH). TSH enters the blood and is carried to the thyroid gland in the neck.
Here TSH stimulates the cells to produce yet another hormone, thyroxin.
This, too, is carried by the blood. Eventually it reaches every cell in the body.
One of its effects is to raise the basic metabolic rate, making every cell
Figure 4. Longitudinal median section of the human brain, showing the location of the generate more heat as it uses up stored materials. In effect, it sets up our
hypothalamus and pituitary "thermostat" and we get warm all over.
The fifth response is initiated by the brain. In the cortex of the cerebrum a
The pituitary is quite close to the hypothalamus and intimately connected thought forms: "I feel cold." The brain decides that we had better move
to it. The pituitary is suspended from the base of the brain by a short stalk around. We stamp our feet and swing our arms about or put on warmer
that contains both nerve fibres and blood capillaries. clothes. This action of the voluntary muscles (the muscles that are con-
The program then examines temperature control by looking first at the trolled by thought and that move the skeleton) generates still more heat, and
response to a cool environment. Cold detectors in the skin send a nerve warms up our blood.
i mpulse to the hypothalamus and to the cerebral cortex of the brain (Figure Feedback to the hypothalamus comes in two ways. First, the rise in
5). The initial response of the nerves is to dilate the blood vessels in the temperature of the blood stops the hypothalamus from emitting its nerve
skin, causing rosy cheeks. One of the nerve clusters in the hypothalamus is impulses. Second, when the thyroxin in the blood reaches the cells in the
also monitoring the temperature of the blood, ready to respond when the hypothalamus, it turns off the releasing hormone.
temperature deep in the body drops too low. Nerve impulses travel from the What happens when your body temperature rises? You may have noticed
hypothalamus to three different effectors. One of these is the set of muscles that your skin becomes redder when you become hot. This is caused by the
that control the size of the arterioles carrying blood to the skin. When the increased flow of blood to the skin, the body's heat radiator. Sensor cells in
skin becomes too cold, these muscles constrict the arterioles so that less the hypothalamus, responding to an increase in the temperature of the
blood will flow. This has the effect of preventing further heat loss to the blood, have inhibited the message to the muscles that control the arterioles
environment - it stops further cooling of the blood. But the disadvantage in the skin. These muscles have relaxed, letting the arterioles dilate to allow
now is that the skin will become still colder. In making this response the a much larger flow of blood to the skin; this cools the skin by transferring
body chooses to sacrifice some external cells in order to maintain the interi- heat to the environment.
or environment. The result of this decision may be frostbite. At the same time the change in the impulses from the hypothalamus has
The second effector of the response against cold is the set of muscles in signalled to the sweat glands in the skin to release perspiration. Evaporation
the skin that make the hair stand erect. In most mammals and birds this res- of water requires heat for the change of state. Vaporization of the sweat
ponse fluffs up the body covering, trapping more air and making it a better carries heat away from the skin, cooling the skin and the blood flowing
i nsulating layer. In humans, however, we have so little hair that the method is through it. In very hot weather perspiration may get rid of as much as 1.5 L
RECEPTORS

Muscles close
arterioles, reducing
blood flow to skin

Start exercising:
muscles get warm

Figure 5. The homeostasis of temperature control

of water in an hour If we lose too much water this way we can suffer heat pattems are evident when we look at the regulation of the many different
shock. Under normal conditions, however, the combination of sweating and entities that organisms regulate. The more altemate effectors an organism
i ncreased blood flow to the skin will result in enough cooling so that the can use to maintain homeostasis, the better are its chances of survival.
hypothalamus will shut off the emergency cooling measures. Organisms that developed nervous systems are better able to control
The regulation of body temperature has illustrated how intricate some of the stability of their internal environments because nerves make possible a
the mechanisms for homeostasis are. You can see that there are several quicker response to a stimulus, triggering rapid reflex regulatory
ways of increasing and decreasing body temperature. The same kinds of mechanisms.

17
18

Before Viewing After Viewing


Ask students to do Activity 1 and to prepare a diagram (similar to Figure 4) Distribute the copies of Figure 5, and discuss the intricacies of the
of the median longitudinal section through the brain, so that they will homeostatic and feedback mechansims. Have students prepare notes based
understand the relationships between the hypothalamus, the brain stem, the on the review questions (Activity 3).
pituitary, and the cerebral hemispheres. Make copies of Figure 5 to give out
after the viewing.

Activities homeostatic system, showing the relationship of


negative feedback to the other components.
4. Where are the temperature receptors located in the
human body?
Activity 1: The Structure of the 5. Describe five responses initiated by the hypothala-
Mammalian Brain mus when body temperature rises or falls
appreciably.
Several laboratory manuals provide good instructions 6. How do plants, fungi, protoctists (protists), and
for students to explore the structure of the brain of a monerans deal with changes in their surroundings,
sheep. Preserved material is firmer and more suitable since they lack nervous systems?
than fresh material for this activity. Some useful 7. What advantages do animals have in coping with
sources are: changes in their surroundings in comparison with
Benson et al., Investigations in Biology, I nvestiga- organisms that lack nervous systems?
tion 4.
Cooley and Vanderwolf, The Sheep Brain: A Basic
Guide. Further Reading
Crawford, Patterns in Biology, p. 415.
(See Further Reading) Benson, Garth D., et al. Investigations in Biology. Don
Mills, Ontario: Addison-Wesley Publishers, 1977
Brown, G.D., and Creedy J. Experimental Biology
Activity 2: The Effect of Tempera- Manual. London: Heineman Educational Books,
ture on the Heartbeat of Daphnia Limited, 1970. '
Cooley, Richard K., and Vanderwolf, C.H. The Sheep
Here is a manageable investigation, using a small Brain. A Basic Guide. London, Ontario: A.J. Kirby
i nvertebrate: Company, 1979.
Brown and Creedy, Experimental Biology Manual, Crawford, Ian. Patterns in Biology. Toronto: McGraw-
p. 204. Hill Ryerson Co., 1983.
(See Further Reading) Kirk, David, et al. Biology Today. 2nd ed. Del Mar,
Calif.: Random House, 1975.
Sherman, I.W., and Sherman, V.G. Biology. A Human
Activity 3: Review Questions Approach. New York: Oxford University Press, 1979.
1. What is positive feedback? What is the effect of Wallace, Robert A. Biology, The World of Life. 2nd ed.
positive feedback in an organism? Santa Monica, Calif.: Goodyear Publishing Co., 1978.
2. What is negative feedback? What is the effect of Weisz, Paul B. The Science of Biology. 4th ed.
negative feedback in an organism? New York: McGraw-Hill Co., 1971.
3. Make a diagram to represent a simple model of a
Biochemical Balances Regulation of Body Chemistry
Objectives Now the Law of Mass Action says that if we have a large mass of carbon
dioxide present, the reaction will go more rapidly to the right. On the other
Students should be able to: hand, if there is little carbon dioxide present, but a larger mass of carbonic
acid, the reaction will proceed mainly to the left.
1. State the Law of Mass Action, and give an example of a reaction Carbon dioxide is produced in the cells of an organism when cellular
controlled by this principle.
respiration makes energy available. By diffusion, carbon dioxide molecules
2. Explain the interaction of carbon dioxide and oxygen in the tissues and will move out of the cells into the surrounding extracellular fluid (ECF) and
i n the lungs in terms of mass action. i nto the blood. Here, it will react with water to form carbonic acid. An
3. Write a word equation to represent the action of hemoglobin in the enzyme in the blood, carbonic anhydrase, speeds up this reaction 13 000
presence of surplus oxygen or surplus hydrogen ions. times. While five percent of the carbon dioxide is carried in simple solution,
4. Construct a diagram to explain how the level of carbon dioxide in the
just dissolved in the water, 12.5 times as much is carried in chemical combi-
blood is controlled homeostatically.
nation. But carbonic acid is not very stable in the presence of the many ions
5. Define competitive inhibition, and give an example of the principle.
carried in the plasma and the tissue fluids. So as fast as carbonic acid
6. Explain how competitive inhibition affects homeostasis at the cellular forms, it breaks up into ions:
l evel.

Program Description
The series, thus far, has shown several examples of homeostasis - the
ways that particular entities within the body fluids are regulated. The regu-
l ation of the composition and temperature of body fluids is the first step in
ensuring that chemical changes in the body are under control. This program
examines some phases of homeostasis that involve chemical reactions.
A first principle of chemical reactions is called the Law of Mass Action.
This law states that at a given temperature the rate of a chemical process is
Notice those reversible reactions; all of these substances are in a dyna-
directly proportional to the masses of the reacting substances. That is, if we
mic equilibrium which can shift in either direction to absorb an additional
have more of a particular substance that takes part in a reaction, then the load of materials from the cells or the environment. If you added up the
reaction will go faster to use up that substance, provided it is in a closed
percentages, you may have wondered why they only totalled 80 percent. The
system. An example is the reaction of carbon dioxide with water to produce
other 20 percent of carbon dioxide is carried in the blood bound to proteins,
carbonic acid:
i ncluding the hemoglobin of the red blood cells. Now the hydrogen ion that
i s set free in the above reaction can combine with hemoglobin as well.
When it does this it changes the shape of the hemoglobin molecule so that
oxygen, carried by the hemoglobin, is set free to diffuse into the cells. There
This reaction can proceed in either direction; if there is a lot of carbonic
it will be used in cellular respiration:
acid (as in a freshly opened bottle of carbonated beverage), then the reaction
will go to the left, producing carbon dioxide and water. We can show this
with arrows in both directions:

19
20

In this way, hydrogen ions can be tied up with bicarbonate, or with


hemoglobin, or with other proteins and ions, to prevent the blood from
becoming too acid. This action to control the pH of the blood is called
buffering. It is one of the aspects of homeostasis.
When the blood reaches the capillaries of the lungs, the Law of Mass
Action goes to work again to reverse these reactions. In the air within the
lungs there is a greater concentration of oxygen than in the blood. Thus
oxygen diffuses from the air into the blood. As it combines with the reduced
hemoglobin the molecule changes shape again, releasing the hydrogen ions:

At the same time, the carbon dioxide bound to a few of the hemoglobin
molecules will be set free. The hydrogen ions will then combine with
bicarbonate ions:

This reaction ultimately sets free more carbon dioxide. By diffusion, Figure 1. Summary of reactions involved in gas exchange in tissues and lungs
carbon dioxide will move from the blood, where its concentration is high,
i nto the air of the lung, where its concentration is low. When we next exhale, An example of competitive inhibition in a living system occurs in animal
some of the carbon dioxide will be excreted from the body (see Figure 1). cells during the production of one of the amino acids, isoleucine:
So far the program has dealt with the effects of mass action. A second
kind of chemical control of the rate of a reaction is one where the end
product combines with the enzyme that caused the change, thus competing
with the substance that started the reaction. For example, suppose a
particular enzyme converts substance X into substance Y: X enzyme . - Y.
Now as soon as Y is formed it combines with the enzyme. This means that
there will be very little of the enzyme left to continue changing substance X.
This kind of action is called competitive inhibition because the competition Thus isoleucine competes with the starting substance, threonine, to inhibit
of both X and Y for the enzyme limits or inhibits the reaction. It is an the rate of its own production in the cell. If no isoleucine is present, the cell
example of negative feedback: will make some. As soon as any is present, production stops. Thus we have
a homeostatic system where the product controls its own rate of production.
Before Viewing After Viewing
A review of cellular respiration and the ways that gases are carried in the Discuss the principles of mass action and competitive inhibition. Complete
blood would be a valuable introduction to this program. Inquiry-oriented the remaining activities. Ask students to make notes, based on the review
teachers may wish to have students investigate the first two activities before questions (Activity 4).
viewing the program. Others will prefer to leave them until after viewing.

Activities Activity 3: The Use of an Enzyme Further Reading


Inhibitor to And Out the Individual Baker, J.J.W, and Allen, G.E. The Study of Biology. 4th
Activity 1: To Study Gas Exchange Steps in the Pathway of a Reaction ed. Reading, Mass.: Addison-Wesley Publishing
Company, 1982.
in Humans This laboratory activity makes use of a four-step Benson, Garth D., et al. Investigations in Biology. Don
pathway from the Krebs Cycle. It focusses on the Mills, Ontario: Addison-Wesley Publishers, 1977.
Refer to Benson et al., Investigation in Biology, conversion of succinic acid to fumaric acid, using Borow, Maxwell. Fundamentals of Homeostasis. 2nd
I nvestigations 33 and 34 (see Further Reading). The germinating bean seeds or the kidney of a freshly ed. (Chapter 6) Flushing, N.Y: Medical Examination
first investigation demonstrates the effects of carbon killed rat or mouse. Refer to Brown and Creedy, Publishing Company, 1977.
dioxide on the rate of breathing: normally after exer- Experimental Biology Manual, p.133 (see Further Brown, G.D., and Creedy, J. Experimental Biology
cising, during hyperventilation, and while breathing Reading). Manual. London: Heinemann Educational Books,
i nto a plastic bag. The second involves the chemistry
Limited, 1970.
of exhaled gas. An alternate version of the latter is
Activity 4: Review Questions Galbraith, Donald I. Lab Manual - Biological Science
found in Galbraith, Lab Manual - Biological Science,
Principles and Patterns of Life. Revised ed. Toronto:
I nvestigation 23 (see Further Reading).
1. What is the Law of Mass Action? State an example Holt, Rinehart and Winston of Canada, 1976.
of a chemical reaction that is controlled by this law. Mikal, Stanley. Homeostasis in Man - Fluids,
Activity 2: The Mechanism of 2. How does the Law of Mass Action relate to the Electrolytes, Proteins, Vitamins and Minerals in
Stomatal Movement interchange of oxygen and carbon dioxide: Clinical Care. Boston: Little, Brown and Co., 1967.
(a) between a tissue cell and the blood in a Nelson, G.E.; Robinson, G.G.; Boolootian, R.A.
Mass action determines many of the responses of capillary? Fundamental Concepts of Biology. 2nd ed.
plants to changing conditions, and is thus important (b) between the blood in a capillary and the New York: John Wiley and Sons Ltd., 1970.
i n maintaining homeostasis. This laboratory activity, alveolus?
described fully in Brown and Creedy, Experimental 3. Write a word equation to show how hemoglobin
Biology Manual, p.177 (see Further Reading), uses the reacts:
spiderwort Tradescanda, a hardy plant for the (a) in the presence of excess hydrogen ions
classroom. It is motivating for students because of its (b) in the presence of excess oxygen.
purple epidermal cells, setting off the green guard 4. Make a diagram to represent the homeostatic
cells. The leaves are placed in four different solutions, control of the level of carbon dioxide in the human
i n darkness and in light, and after 15 minutes students body.
count 25 stomata and record how many of them are 5. Define competitive inhibition, and give an example
open and how many are closed. From the data they of the principle.
deduce the conditions that determine stomatal 6. Explain how competitive inhibition affects homeo-
opening. stasis in a cell.

21
22

Hormonal Control The Coordination of Homeostasis


caused by the action of a particular enzyme, succinic dehydrogenase, that
Objectives removes hydrogen from succinic acid.
Students should be able to:
1. Define hormone and endocrine gland.
2. Identify common hormones (such as thyroxin, ADH, insulin, and
glucagon) and state where they are produced and where they produce
their effects.
3. Describe the location and relationship between the hypothalamus and
the pituitary gland.
4. Describe the role of the hypothalamus and the pituitary gland in the
control of the reabsorption of water. They should also be able to
explain how feedback operates in this system.
5. Outline the two-hormone system for the control of blood glucose.
6. Name the different kinds of cells in the pancreas and identify the
secretions produced by each.
7. Outline the hormonal control system that determines the level of
thyroxin in the body.
Figure 1. Hypothetical model to account for the action of a hormone only on specific
target cells
Program Description I n the example above, the need for the hormone ADH stimulates its pro-
Many of the homeostatic mechanisms that have been illustrated in earlier duction. When it is no longer needed the production of the hormone ceases.
programs have been controlled by hormones. The glands of the body may be This is the simplest of the hormonal control mechanisms. In this program
divided into two classes: the exocrine glands that pour their secretions we see examples of two more complex mechanisms. In the first example,
directly into ducts that deliver them where they are needed; and the endo- two hormones are produced that have opposite actions. Thus they are called
crine glands, whose secretions are called hormones, which are delivered by antagonistic. Their combined effect is to control the level of blood glucose
the blood stream to all parts of the body. Hormones are secreted in small to within the very narrow limits of 0.1 percent. The second example is a more
amounts, and they produce their effects at a distance from their source. complex mechanism involving four hormones and a very finely tuned control
Specific target cells are affected by hormones, while other cells are of metabolism.
apparently. unaffected. The program uses the control of blood glucose as an example of a home-
What might account for the difference in action upon the cells? One ostatic control in which two hormones act antagonistically. Program 1
hypothesis is that target cells must have specific receptor modules, prob- mentioned that when we eat at rather odd intervals, or even if we fast, it is
ably proteins, located in their cell membranes. When the right hormone i mportant that the level of glucose in the blood, be regulated. Glucose must
reaches these receptors, the protein might produce a secondary messenger reach all of our cells at a fairly constant rate to supply energy for their
molecule inside the cell, one that would activate a particular enzyme and various activities. It is especially important for the brain cells to be nour-
initiate a series of reactions (Figure 1). For example, in Program 3 it was ished, as they are the first to malfunction when the level of glucose
explained that the anti-diuretic hormone (ADH) from the pituitary gland becomes too low. How is the homeostasis of glucose accomplished?
controlled the permeability of the cell membranes in the distal portion of the Everyone knows of the disease sugar diabetes (diabetes mellitus), involving
uriniferous tubules in the kidney, making them reabsorb more water and a misalignment of this control system. Diabetics must consciously regulate
specific ions. This change in permeability is reversible, adjusting for varied their glucose level by controlling their intake of sugars and hormone.
conditions of the body fluid (ECF). The change in permeability may be The rest of us are lucky in that our homeostatic controls are automatic.
Sugar diabetes is caused by the failure of the body to produce sufficient The second hormone, glucagon, is also a very small protein, consisting of
i nsulin. Sir Frederick Banting was a Canadian doctor who established, in only 29 amino acids. It is produced in the alpha cells of the islets of
1922, the relationship of insulin from the pancreas to the control of blood Langerhans when the level of glucose in the blood falls below 0.1 percent.
sugar. In the pancreas there are about a million small clusters of cells called Glucagon acts on the liver to take glucose out of storage, converting
the islets of Langerhans. They are distinct from other cells of the pancreas glycogen to glucose-1 phosphate:
that secrete digestive juices. The islets pour their secretions directly into the
blood rather than into the pancreatic duct. There are two kinds of cells in the
islets of Langerhans. designated aloha The beta cells produce
insulin Ti ne alpha ceps produce anotner hormone, glucagon, that acts
antagonistically to insulin. In other tissues, glucagon accelerates the breakdown of glucose, producing
energy. In the kidneys, glucagon changes the rates of excretion of sodium,
potassium, and phosphate ions. We can now complete our homeostasis
diagram:

REGULATED RECEPTOR MESSAGE EFFECTORS RESULTS


ENTITY

I nsulin is a small protein consisting of 51 amino acids. Its action is to


reduce the level of glucose in the blood. It does this by converting glucose
to glycogen in muscle and liver cells, and by converting glucose to fats and
proteins in other body cells. These actions effectively store glucose in larger
molecules for future use. Insulin also accelerates the breakdown of glucose
in most cells, releasing energy. There are two hypotheses about how insulin
accomplishes these actions. First, as discussed in Program 4, insulin may
affect cell membranes, making the membranes facilitate the movement of
glucose into the cells. Or, second, insulin may accelerate the phosphoryla- Figure 3
tion of glucose to make it more chemically active so that it can react faster.
I n the antagonistic pair, the second hormone succeeds in turning off the
action of the first, before large swings can occur in the levels of the
regulated entities. If we consider only the first hormone, insulin, a graph of
Thus the homeostatic diagram for the action of insulin in the control of
the levels of the regulated entity would look like this:
glucose level looks like this:
REGULATED RECEPTOR MESSAGE EFFECTORS RESULTS
ENTITY

Figure 4

23
24

The second hormone refines the control so that the regulated entity is But it is not quite that simple. The detectors of the concentration of thyroxin
kept within even narrower limits: are not in the pituitary, but in the hypothalamus above it, in the brain. Here
are located the control centres for many of the regulated entities. A group of
chemoreceptors with a good blood supply is constantly monitoring the com-
position of the blood. When these cells detect an insufficiency of thyroxin,
they secrete a short polypeptide, called thyrotropin releasing factor (TRF),
i nto the blood. There is a special portal system of veins that carry this
hormone directly to the anterior lobe of the pituitary, where it causes the
cells to release TSH. When thyroxin levels rise, another group of control
cells, also in the hypothalamus, secretes another short polypeptide, somato-
Figure 6 tropin release inhibiting factor (SRIF). SRIF inhibits the production of several
hormones from the pituitary: TSF, human growth hormone, insulin, and
Thus, homeostasis, the regulation of the internal environment within
glucagon. Thus, our diagram of the interactions (Figure 7) involves three
narrow limits, is fine-tuned by the interaction of antagonistic pairs of
glands, four hormones, and a very fine-tuned homeostatic control.
hormones.
A third kind of homeostatic control is the interaction of sets of hormones,
where one hormone stimulates the production of a second that suppresses
the first. This kind of interaction maintains a steady level of hormone in the
blood. One example of such a system is the production of thyroxin, which
controls the rate of metabolism in the body. Extra thyroxin makes each cell
release more energy, using up more oxygen and producing more heat.
Thyroxin is produced in the thyroid gland in the neck. Its production is
stimulated by TSH (thyroid stimulating hormone) from the anterior lobe of
the pituitary gland at the base of the brain. When the level of thyroxin falls
too low, the pituitary causes more TSH to flow. This in turn stimulates the
thyroid to produce more thyroxin. As blood containing more thyroxin flows
through the brain, the level of TSH is suppressed. So we have a feedback
l oop (Figure 6) where two hormones produced by two glands turn each other
on and off, regulating the level of thyroxin in the blood, and thus controlling
our rate of metabolism.

Figure 7. Homeostatic control involves three glands and four hormones

Further Study
There has not been time in this series to show all the aspects of homeo-
stasis. Students can extend this study by researching other fascinating
aspects of hormonal control, such as the antagonistic pair of hormones that
controls the level of calcium ions in the blood (parathormone and calcitonin),
and the interacting feedback control mechanisms that determine the
menstrual cycle of the human female. How many hormones are involved,
where are they produced, and where do they exert their effects? How does
Figure 6. The interaction of two glands and two hormones an understanding of this cycle enable women to plan their pregnancies?
The interrelationship between genetics and homeostasis has not been homeostatic control to account for the balance in predator-prey relation-
discussed in this series, again, because of time limitations. How is the ships, or the co-evolution of insects and flowering plants, or the mainte-
ability to develop homeostasis inherited? How can the same hormones, FSH nance of the level of the elements involved in the bio-geo-chemical cycles?
and LH, have such different results in the two sexes? How do genes affect Self-regulation within narrow limits is thus a characteristic of life at all levels,
the functioning of hormones and vice versa? Do certain substances block from the individual cell to the entire biosphere.
the flow of genetic information?
Some authors extend the meaning of homeostasis beyond the limits of
the individual organism. Certainly, feedback systems appear to control the
Before Viewing
life and functioning of larger biological units: the species, the population, The students could view Program 6 without any prior introduction, however,
the community, the ecosystem, the biome, indeed the whole biosphere. Can Program 3 introduces the concept more simply. Some of the activities
the stability of such large systems be considered homeostasis? How does a related to this program could be used to prepare students for viewing.
species cope with variations in the environment, yet maintain itself genera-
tion after generation? How do homeostatic mechanisms reduce errors and
damage in genes and repair DNA molecules to ensure genetic stability?
After Viewing
How does the Hardy-Weinberg Law relate to the stability of genes in a Students should do several of the related activities. It would be useful to
population? In what way is evolution a kind of homeostatic adaptation of teach the additional examples that are discussed in Further Study: the
species? Dynamic equilibrium, the establishment of controls that keep control of calcium ion concentration and the regulation of the menstrual
fluctuations within limits, seems to be as much a characteristic of the larger cycle of the human female. Complete the discussion by asking students to
biological entities as of the individual organism. Can we use our model of prepare notes based on the review questions (Activity 5).

Activities the small crustacean. Refer to Lang et al., The


Functioning Animal, I nquiry 21 (see Further Reading).
Activity 5: Review Questions
1. (a) Define an endocrine gland and tell how it differs
Activity 1: Invitations to Inquiry Activity 3: An Investigation of the from an exocrine gland.
(b) Define hormone and explain the hypothesis of
If you like the teacher-centred inquiry mode of teach- Effect of Adrenalin and Acetylcho- how a hormone affects certain target cells
ing, you may wish to adapt some of the six Invitations line on the Heart of a Frog without having an effect on most cells of the
that Joseph J. Schwab has provided on this topic in body.
the Biology Teacher's Handbook, pp. 462-482 (see Not every teacher will want to pith a live frog for this 2. Construct a table with the following headings:
Mayer, Further Reading). They are: activity, but those who do can refer to Brown and HORMONE WHERE PRODUCED WHERE IT ACTS
I nvitation 39: Control of Blood Sugar Creedy, Experimental Biology Manual, page 254 (see NATURE OF THE ACTION
I nvitation 40: Blood Sugar and the Internal Further Reading). Complete the table for the following hormones:
Environment thyroxine, ADH, insulin, glucagon, parathormone,
I nvitation 41: Blood Sugar and Insulin Activity 4: The Importance of calcitonin, estrogen, progesterone.
I nvitation 42: Blood Sugar and Hunger 3. Describe the location of the hypothalamus and of
I nvitation 43: Basal Metabolic Rate Homeostasis the pituitary gland. What is the nature of the
I nvitation 44: The Stress Reaction: Adrenaline connection between them? How does the hypothal-
This is a series of questions dealing with osmotic
amus control the functioning of the pituitary?
pressure and the level of calcium ions - a "dry" lab,
Activity 2: Effects of a Hormone on 4. Describe the control of the reabsorption of water in
or cerebrational inquiry. It can be found in Benson et
the human body. How does feedback work in this
the Heart Rate of Daphnia al, Investigations in Biology, I nvestigation 10 (see
system?
Further Reading).
5. Make a diagram to explain the homeostatic control
This laboratory-oriented inquiry makes use of of the level of the glucose in the blood.
ephinephrine (adrenalin) to stimulate the heart rate of

25
26

6. Describe the structure of the pancreas and state


exactly where each of its secretions are produced.
7. Outline the hormonal control of the level of thyroxin Ordering
in the body.
8. In what ways is homeostasis a characteristic of life
at all levels? In your answer give examples of the
Information
application of control systems at the population,
species, community, biome, and biosphere levels. To order this publication or videotapes of the
programs in the series Homeostasis, or for additional
Further Reading information, please contact the following:

Baker, J.J.W., and Allen, G.E. The Study of Biology. Ontario


4th ed. Reading, Mass.: Addison-Wesley Publication TVOntario Sales and Licensing
Co.,1982. Box 200, Station Q
Benson, Garth D., et al. Investigations in Biology. Don Toronto, Ontario M4T 2T1
Mills, Ontario: Addison-Wesley Publishing Co.,1977. (416) 484-2613
Brown, G.D., and Creedy, J. Experimental Biology
Manual. London: Heineman Educational Publishing,
1970.
Emmel, Thomas Co. Worlds Within Worlds. New York:
Harcourt Brace Jovanovich, Inc., 1977.
Jones, Kenneth C., and Gaudin, A.J. Introductory
Biology. New York: John Wiley and Sons, Ltd., 1977.
Kimball, John W Biology. 5th ed. Reading, Mass.:
Addison-Wesley Publishing Co., 19B3. Videotapes BPN
Lang, H.M.; Palfery, E.G.; and Van Nieuwenhove, E.L.R. Program 1: Coping with Change 226401
The Functioning Animal. Toronto: Gage Educational Program 2: The Sea Within 226402
Publishing, 1978. Program 3: Osmoregulation 226403
Mayer, Wm. V., ed. Biology Teacher's Handbook. Program 4: The Feedback Cycle 226404
3rd ed. New York: John Wiley and Sons, Ltd., 1978. Program 5: Biochemical Balances 226405
Program 6: Honnonal Control 226406
McElroy, WD.; Swanson, C.P; and Macey, R .I. Biology
and Man. Englewood Cliffs, N.J.: Prentice Hall Inc.,
1975.
Volpe, E. Peter. Man, Nature, and Society. An
Introduction to Biology. 2nd ed. Dubuque, Iowa:
Wm. C. Brown, Ltd., 1979.

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