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Journal of Biological Education

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Teaching the Hardy-Weinberg Law

a
B. A. C. Dudley
a
Centre for Science Education, Chelsea College, University of
London
Published online: 13 Dec 2010.

To cite this article: B. A. C. Dudley (1972) Teaching the Hardy-Weinberg Law, Journal of Biological
Education, 6:6, 359-367, DOI: 10.1080/00219266.1972.9653803

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 Journal of Biological Education (1972) 6, 359-367
Teaching the Hardy-Weinberg Law
B. A. C. DUDLEY
Centre for Science Education,
Chelsea College, University of London*
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1. Introduction
One of the developments in the Nuffield
O-level Biology Project was to introduce Men-
delian genetics and population genetics into
the O-level course at year five. Although appar­
ently largely successful, a survey of the Nuffield
Biology O-level Project, undertaken by Miss
Grace Monger of the Centre for Science Educa­
tion, has shown that the teaching of population
genetics (Chapter 10) and in particular the teach­
ing of the Hardy-Weinberg Law is not altogether
satisfactory. It was because pupils found the
Hardy-Weinberg Law to be uninteresting and/or
difficult that an alternative scheme for teaching
this important topic has been devised. It is this
which is presented here in an extended form so
that teachers may use it as a source of material
and may select from it. It has been tried out
amongst pupils and also amongst teachers where
it was received with enthusiasm, the latter freely
admitting to it imparting to them a deeper
appreciation of the validity and the value of this
Law and hence a greater confidence of being able
to cope with it in class.
In the Nuffield O-level Biology Project itself
the Hardy-Weinberg Law is suddenly presented as
a mathematical model of a gene pool and in the text
2. Card One
(1) Use the black and the white beads avail-
able to represent the genetic cross between
Genetic cross : Dominant parent X recessive parent
Genotypes : AA
Bead model: 10 black beads
♦Present address: Department of Education, University of Keele, Keele, North Staffordshire.
359
rapidly and algebraically reaches the statement
"The frequency of AA in the population =pl
The frequency of aa in the population =q*
The frequency of Aa in the population =2pq"
The equation p'+2pq+qs = l is then used to
establish the frequency of the genotypes for
Tasters and Non-Tasters, for Tongue-rollers and
non Tongue-rollers and a little later for the
frequency in a population of the positive and
the negative rhesus factor of the blood.
At no point is the pupil given an opportunity
to appreciate that the law is a generalization, nor
that, as a generalization it is valid, nor that its
contribution to our knowledge of the genetic
structure of a population is of value. However,
each of these matters can be developed by care­
fully structuring the sequence of learning experi­
ences, starting, as does the Nuffield O-level
Biology Project, with the use of beads to represent
alleles, and sets of beads to represent first the
population of alleles in the parental generation
and then the population of alleles in the filial
generations. The class material, presented in the
form of work cards was as follows, with infor­
mation in the square brackets available only to
the teacher.
a pure dominant and a pure recessive
parent. Set up the system as follows.
aa
10 white beads
 360 B. A. C . D U D L E Y

Diagram of the model representing one pure dominant

and one pure recessive parent.

Exactly what does each bead represent ? What does each pair of beads represent ?
[Each bead represents an allele of the gene in [Each pair represents the geneotype of one
question]. offspring.]
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(2) Pick out one bead from each box and
arrange the beads in pairs on the bench.
Continue the process until all the beads
have been used. Arrange the pairs in rows.
[Result (2)
What does the set of pairs of beads repre­
sent?
[The set of pairs of beads represents the
genotypes of the F, generation.]

ao ©o
t o o®
TABLE 1
Genetic composition of the populations of successive generations
of offspring according to the bead model typical result
Offspring
Black, Black „, , w h j White, White
Generation (Homozygous m,,, ' „ n l K l (Homozygous
dominant) (Heterozygous) recessjve)

F2 2 6 2
/:: 3 4 3
/', 2 6 2
/, 2 6 2
l\; 2 6 2
1-, 3 4 3
l\ 3 4 3
1, 2 ft 2
/',n 2 6 2
Fu 4 2 4

Total 25 50 25

Average 2-5 50 2-5

[Under these conditions only random change can occur after

the F2 generation—the system is stable with the incidence of
Black, Black: Black, White: White, White in the ratio of
1:2: 1. It is important to ensure the distribution of beads in
each box and the selection of pairs of beads is random. Even
so, with only 20 beads in the system fluctuations about this
ratio are to be expected but the results of many individuals,
and average results of the class, closely approximate to it.]
 T E A C H I N G THE H A R D Y - W E I N B E R G LAW
What is the composition of each pair of
beads?
[Each is made of 1 black and 1 white bead.]
What is the genetic composition of the
individuals of the Fx generation ?
[All the offspring are heterozygous.]
(3) Return the first five pairs of beads to one
box and the second five pairs of beads to
the second box.
The two boxes represent the two sexes.
Label one box "Male" and the other box
"Female".
Note that each box contains beads repre­
senting the genotype of five individuals of
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361
pairs of beads now represent the next
generation, the F, generation.
Count the number of pairs of beads in the
row which are:
(i) black, black
(ii) black, white
(iii) white, white.
Copy the table below (Table 1) and record
the numbers.
(5) Return the first five pairs of beads to one
box and the other five pairs of beads to the
other. Without looking at the boxes pick out
one at a time from each box and so get a

the Fi generation. That is, one box con-
t a b s the genotypes of gene A of the male
population and the other the genotypes of
gene A of the female population.
(4) Gently shake each box to mix up the beads.
Without looking at the boxes pick out one
bead from each box and arrange them in
pairs on the bench as before. The rows of
row of pairs that represents the F, genera­
tion. Record the number of pairs that are
black, black; black, white; white white.
(6) Repeat the exercise until the table, Table
1, is complete.
Attention can be drawn to the fact that no
systematic change occurs between F, and F n
generations by drawing a graph of the results.

3. Card Two
Resume
There is no systematic change in the composi­
tion of the population through ten generations.
Setting aside the changes due to chance, this
must mean the numbers of the black and the
white beads in each box remain unchanged
throughout the experiment as does the number
of each of the three kinds of pairs of beads
selected from them.
1. Copy and complete the following table,
(Table 2).

TABLE 2
Composition of the population of beads and of the pairs of beads in
the model of the genetic system being studied

Number of beads in each box is 10

Total number of beads in the system is 20
Number of Proportion of beads in each box
Colour of beads in ' V u l e a r
beads Decimal
each box f r „ t T,, n Percentage form

Black 5 i 50 0-5
White 5 i 50 0-5
Pairs of beads Average
produced number
Black, Black 2-5 Ï 25 0-25
Black, White 50 i 50 0-50
White, White 2-5 i 25 0-25

[The completed table is presented on this occasion.]

 362 B. A. C. DUDLEY

4. Card Three
The pattern of events practised in Card One can be represented by the following :

TABLE 3

Dominant Recessive
Box one allele allele
Box two (male) (Black bead) (White bead)
(Female)
Frequency
Dominant
allele Black, Black Black, White
(Black bead)
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Recessive
Black, White White, White
(White bead)

(1) Copy out this table and add to your copy the
frequency of the black and the white beads in
each box.
(2) What is the frequency of the Black, Black;
the Black, White; and the White, White
pairs to be expected according to the table?
(3) What was the frequency of each of the three
types of pairs obtained in the experiment ?
(4) Does the table on this card represent accur­
ately the bead model system?
The bead experiment itself models (represents)
the significant events of a prolonged genetic
experiment that starts with a classical Mendelian
cross. From pure-bred parents, one pure domin­
ant and one pure recessive, this model shows the
phenotype ratio arising in the F, generation and
also shows that no change occurs in that ratio
(other than by chance) in succeeding generations.
(5) What is the phenotype ratio that persists
from the Ft generation? [3:1]
(6) What is the frequency of the dominant allele
and the recessive allele amongst the male and
the female parental populations? In the
model this is represented by the frequency of
the black and the white beads in each box.
[h i or 0-5, 0-5]
(7) Is the phenotype ratio dependent upon the
frequency of the two alleles ? [Yes]
The following tests will act as a further
check upon your answer to question 7.
Copy the table at the start of this card.
Write into it the frequency of the alleles for
the following crosses.

TABLE 4
Table offrequencies of the cross A A x aa

Pure Pure dominant Dominant alleles Recessive alleles

recessive parent (Black beads) (White beads)
parent Frequency 1 0
Dominant Pure dominant Heterozygous
allele 0 (Black, Black) (White, Black)
(Black beads) 1x0=0 0x0=0
Recessive 1 Heterozygous Pure recessive
allele (Black, White) (White, White)
(White beads) 1x1 = 1 1x0=0

All the offspring are heterozygous, (Frequency of Black, White=l) as

is found to be the case both in Mendelian experiments and in the bead
model of such a system.
 T E A C H I N G THE H A R D Y - W E I N B E R G L A W
(i) Pure dominant x Pure recessive parents
Genetic cross: AA xaa, [see Table 4].
(ii) A backcross, between a double recessive
parent and a heterozygous parent.
On this occasion write a matrix of the
frequencies. There is no need to copy the
whole table.
5. Card Four
The phenotype ratio that occurs amongst off­
spring has been found to be dependent upon the
frequency of the alleles amongst the parental
generation. Whatever is the distribution of the
alleles amongst the parents, that distribution
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will determine the distribution of phenotypes
amongst the offspring. It must also determine the
composition of genotypes amongst the offspring.
Since this is always the case, it should be
possible to build a more general model.
Let the frequency of dominant alleles (black
beads) in a population be any fraction =q.
Since total frequency of all alleles (beads in a
population) = 1, then frequency of recessive
alleles (white beads) in the population = 1 —q.
The model then becomes
TABLE 5
Box one
Box two (males)
(Females)
Frequency
Dominant allele
(Black bead) <7
Recessive allele
(White bead) \-q
6. Card Five
According to the Hardy-Weinberg Law it is
possible to calculate, for any population, the
frequency of the three genotypes of a gene and
the frequency of its two alleles.
This indeed is the case when q=l—q=i, a
special case which results in the classical Mendel-
ian ratios.
Can the law really apply to any frequency of
alleles, and in any population?
The following experiment will serve to check
whether this is indeed the case.
[Make available a box of 50 beads, 20 of which
are white and 30 black].
You have been provided with a box of beads
which represents a population of unknown
363
[Genetic cross: Aa xaa]
Frequencies (i, i) /0>
©-GD
The result is a 1:1 ratio amongst the off­
spring, again a result which is found in
genetics experiments of this kind and in the
bead model of such a system.]
Copy the table and on your copy enter the
combinations of genotypes and their frequency.
List the frequency of
(a) Black, Black pairs of beads [q1];
(b) Black, White pairs of beads [2q (1 -q)] ;
(c) White, White pairs of beads [(1 -q) »].
What is the frequency in such a population of
(a) Homozygous dominant individuals [q']l
(b) Heterzyogous individuals [2q(l —q)] ?
(c) Homozygous recessive individuals
[(l-<7)8]?
Note that:
Mendel's ratios arise wheng=l — q = i, which
is a special case of this general rule.
The general rule is known as the Hardy-
Weinberg Law.
Dominant allele Recessive allele
(Black bead) (White bead)
q \-q
genetic composition. Use the Hardy-Weinberg
model to calculate the frequencies of the three
genotypes in this population and also the fre­
quencies of the dominant and the recessive
alleles within it. Proceed as follows:
(1) Without looking at the box select two beads
at a time and arrange them in pairs in one
long row. Be sure to arrange for the row
to be in a place where the beads will not be
disturbed for some time. This row of pairs
represents the individuals of the population.
(2) Identify and count those which would have
the recessive phenotype (White, White) since
these can be distinguished in a real popula­
tion.
 364 B . A. C. DUDLEY
(3) Count the number of individuals which [Incidence of pure
would have a dominant phenotype (those dominant types =036
with one or two black beads). Incidence of pure
Copy and complete the following table: récessives types =016
TABLE 6
Total =0-52
Number of
Phenotype individuals Incidence of
heterozygous types = 1 — 0-52=0-48]
Dominant (Black) [21] Express as a percentage each of the following,
Recessive (White) [ 4] as calculated on the basis of the Hardy-
Total [25] Weinberg model.
Incidence of pure dominant types (36 %)
(4) Express the incidence of the recessive (White) Incidence of pure recessive types : (16%)
phenotype as a percentage of the whole and
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Incidence of heterozygous types : (48 %)

then in decimal form. (8) This particular population is made up of
[Incidence of the recessive phenotype = 25 individuals. Calculate the numbers of
^xl00=16%=016] individuals of each genotype that must be
present in the population according to the
(5) According to the Hardy-Weinberg model if Hardy-Weinberg model.
the incidence of the dominant allele is q then [Numbers in the population that must be
the incidence of the double recessive must present according to the model:
be (1 -q)\
What, then, is q on this occasion, according (a) pure dominant =—— x 25 = 9 individuals
luu
to the model ? 48
[(1-<?)« = 016 (b) heterozygous ■x25 = 12 individuals
100
\-q = V016=0-4
q = 1-0-4 (c) pure recessive (given) = 4 individuals
= 0-6] Total 25 individuals]
(6) According to the model the frequency of (9) The incidence of the dominant allele (q) in
pure dominant individuals must be q'. the population, on this occasion has been
What, then, is their incidence on this occasion found to be 0-6. What, then, is the number of
according to the model ? dominant alleles present in this population.
[Incidence of double dominant =q' = (0-6)a = [Number of individuals in the population is 25
0-36] Number of alleles of this gene present is
(7) Sum the incidence of the pure dominant and 2x25=50
pure recessive types. Number of dominant alleles present is
The heterozygous types will form the re­ 0-6x50=30].
mainder. What is their incidence? (10) Complete the following table, returning to

TABLE 7
By calculation from By counting
Number of beads the Hardy-Weinberg the row of
model beads

Black (=dominant allele) 30 30

White (=recessive allele) 20 20
Number of pairs
Black, Black
(pure dominant individuals) 9 9
White, White
(=pure recessive individuals) 4 4
Black, White
(=heterozygous individuals) 12 12
[The completed table is presented on this occasion].
 T E A C H I N G THE H A R D Y - W E I N B E R G
your row of pairs of beads for the informa­
tion required in the final column.
[Caution: A perfect match between the
figures in the two columns of the above table
will result in most, but not in all cases.
When a discrepancy does arise, provided the
7. Card Six
Some uses of the Hardy- Weinberg model
Use the Hardy-Weinberg model to solve the
following problems.
Problem number one
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A poultry farmer has a flock of chickens and is
worried by the occasional appearance of an
undesirable phenotype, which occurs in 1% of
his flock. He has found by breeding experiments
that such individuals have a double recessive
genotype (aa).
The farmer must purify his flock for it to have
any economic value, and has to take one of two
choices. He can kill each of the double recessive
individuals as they appear or he can create a new
pure flock from homozygous dominant indivi­
duals within his present flock. Which method
will give him the quickest result?
What advice would you give him and what
would be your reasons for your advice ? Remem­
ber the farmer will have to be shown your
calculations before he can be expected to be
persuaded.
He wishes to maintain a flock of 1000 chickens.
(Hint: The most important fact to establish is
the number of carriers (hétérozygotes) in the
population).
[Answer
The incidence of carriers in the flock is calcu­
lated on the basis of the Hardy-Weinberg
model as follows:
The incidence of the double recessive genotype
(aa) is 1 % = 0 0 1 . According to the Hardy-Wein­
berg model, the incidence of genotype aa
is ( I - ? ) '
Hence (l-tf)«=0-01
\ - q =V001=01
=1-01=0-9.
q
Frequency of pure dominants=^ a =(0-9)' =
0-81 or 81%.
Frequency of pure récessives (given)=001 or
1%
Frequency of the hétérozygotes=1 00—0-82 =
018 or 18%.
In a flock of 1000 chickens the number of
poultry with the undesirable phenotype=
LAW 365
instructions have been followed with care,
the error is due to chance, a factor not taken
into account in the Hardy-Weinberg model
because there a population of infinite size is
envisaged, and not a finite one of only 50
beads].
1
xl000=10.
100
and the number of poultry that are carriers=
i i x 1000=180.
Killing the 10 will leave unaffected the
180 chickens that carry this undesirable charac­
ter. It is unlikely that killing the double recessive
phenotypes will have any significant effect upon
the incidence of this character, at least, not for
many generations. It is impractical to purify the
flock by this means alone because of the numbers
that are heterozygous.
It is necessary therefore to identify and elimi­
nate the hétérozygotes, and in order to do this
the farmer must employ the standard Mendelian
backcross.
Step 1. An individual with a normal phenotype
is crossed with one that has the undesirable
phenotype. If all the offspring are normal then
the poultry with the dominant phenotype will
be pure dominant. If it was a carrier only half
the offspring would have been of the dominant
phenotype as follows :
Parents : (1 ) AA x aa (2) Aa x aa
Offspring: Aa Aa, aa
Pure dominant parents identified in this way
would then have to be isolated and a flock built
up from them.]
Problem number two
In human populations about 1 in every 1500
babies borne has a condition known as cystic
fibrosis. It is a double recessive condition that
currently is fatal, usually before the child
reaches adolescence. Given this information
use the Hardy-Weinberg model to calculate how
many adults in each 100 can be expected to be
carriers and how many carriers there are to be
expected in your school.
[Answer
The incidence of cystic fibrosis = 1 in 1500 =
_ L =00007.
1500
It is a double recessive condition (aa).
 366 B. A . C.
According to the Hardy-Weinberg model—
Incidence of genotype aa=(l—^) 8 =0-0007.
Hence 1-<7=V00007=002569
? = 1 -0-02569=0-9744.
Frequency of pure dominant phenotypes
(AA)=? 2 =0-9744*=0-9495.
Frequency of pure recessive phenotypes
(given)=00007
Frequency of hétérozygotes (carriers) (Aa) =
1-0-9502=0-05 (approx.).
According to the model approximately 5%,
5 people in every 100, are carriers for the
congenital error (a ratio of approximately 1 in
20).
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A simple calculation serves as a check on this
result. An infant with cystic fibrosis, being a
double recessive condition, can be produced only
if both parents are carriers; no cystic fibrosis
individual has yet survived to adulthood to
become a parent.
The chance of one parent being a carrier is —
The chance of both parents being carriers is
1 1-—
20X20_400
The chance of such parents having a cystic
fibrosis child is - (the classical Mendelian ratio
4
in a different form). Hence the total chance of a
cystic fibrosis child being borne is —■ x-; = 77nn'
1 ,
a figure that compares with the incidence of
1500
given initially. The discrepancy is due to the
approximation involved in the statement that 1
adult in 20 is a carrier, the incidence is marginally
higher than 1 in 20.
The number of carriers in your school can be
expected to be approximately— x n, where n is
the number of pupils in the school.]
Problem number three
The incidence of a number of recessive con­
genital errors in humans are known to be as
follows :
Phenylketonuria 1 in 25,000 births
Albino 1 in 20,000 births
Amaurotic idiocy 1 in 200,000 births
What is the incidence in the human population
of the carriers of each of these errors ? Express
your answers as a ratio of one in so many, as
the number per 100, and as the number per
1000.
DUDLEY
[Answer
These calculations can be simplified by taking
the general case and applying the Hardy-
Weinberg model.
Given the incidence of the double recessive
condition is a%. In decimal form a%=b.
From the Hardy-Weinberg model,
b ={\-qy
\-q=Vb
q=l-Vb
The frequency of the homozygous dominant
condition is
q*=(l-VW
= l-2Vb+b.
The frequency of the homozygous recessive
condition is b. So the frequency of the hetero­
zygous dominant condition is
l-(b + l-2Vb+b)
= \-2b-\+2Vb
=2Vb-2b
=2(Vb-b).
Hence, if the incidence of the homozygous
recessive condition is b, then the incidence of
the homozygous dominant condition is
1—2 Vb+b and the incidence of the hetero­
zygous dominant condition is 2(Vb—b).
These expressions can be checked by using
them first to recalculate the incidence of cystic
fibrosis and also the problem of the poultry flock.
Having demonstrated their validity they can then
be used with confidence to answer the questions
posed on the work card, for example, the inci­
dence of carriers of phenylketonuria.
Incidence of phenylketonuria is 1 in 25,000
births. A ratio of in decimal form is
0-00004=6, and V0O0O04=0-00633= Vb.
According to the Hardy-Weinberg model, the
incidence of carriers (hétérozygotes)
=2(Vb-b)
=2(0-00633-0-00004)
=2(0-00629)
=001358 or 1-358 in 100
, . 100 . . _.
= 1 m —TTg , approx. 1 in 75.
The incidence in the human population of
carriers of phenylketonuria is approximately
1 in 75.
In the school population the number of pupils
that can be expected to be carriers of this con-
. 1 0 0
genital error is x n, where n is the num­
ber of pupils in the school (some 13 or 14 carriers of
phenylketonuria per 1000 head of the population).]
 TEACHING THE HARDY-WEINBERG LAW
8. Card Seven
Calculate for your own class the incidence of
(a) inability to roll the tongue (recessive) and
the number of tongue rollers in the class
that are (i) homozygous and (ii) hetero­
zygous for this character.
(b) non-tasters of PTC (recessive) and similar
calculations to those in (a) above for those
who can taste it.
What is the incidence of individuals in your
class who are
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367
(i) non-tasters of PTC and unable to roll the
tongue ?
(ii) homozygous for both characteristics ?
(iii) heterozygous for both ?
[The data that is required for the gene­
tic analyses of this population has to be
gathered by the pupils themselves while
previously in this work scheme calculations
were based upon the second hand data
provided.]

Reference
NUFFIELD O-LEVEL BIOLOGY PROJECT (1966-67). Student's Text Year V. London : Penguin/Longman.