SHES 2303 MATHEMATICS IN BIOLOGY

1 Introduction
In this course, we will learn calculus methods that are useful for biological work. Much
of the (technical) material here will based on the textbook Thomas’ Calculus (hereafter
denoted as TC) by Finney, Weir and Giordano (2001). The examples, however, will be
drawn from biology. For now, let us look at a good example of how calculus can help
us understand biology better. Figure 1 shows a sigmoid (logistic) growth curve of S.
cerevisiae from Gause’s (1932) study of the growth of yeast in culture medium. Gause is
a Russian biologist who first demonstrated the classical result of the effect of competition
on the growth of two Paramecium species : caudatum and aurelia.

Estimated Growth Curve of S .cerevi si ae

12
10
8
Amount

6
4
2
0

0 10 20 30 40 50 60

Hours
Fig. 1 The growth profile of yeast after about 60 hours of culturing. Note the relatively slow rate of
growth in the beginning, followed by an approximately linear phase in the middle. The rate slows down
after 30 hours, and appears to reach a plateau (13) after 40 hours.

There are several interesting observations from Figure 1. First, we understand that the
slow rate of growth in the beginning is due to the small starting population. Binary cell
division, however, quickly allows the yeast population to build up a sufficiently large base,
which accelerates subsequent growth rates. Thus, from time 10 to 30, the population size
increases approximately six fold. After that, it reaches a limiting value of approximately
13, instead of increasing without bounds. The slow-down in growth (and its eventual
cessation) turns out to be partly caused by alcohol, which is its metabolic waste.

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2 Concept of Limit

Let us start with an intuitive understanding of limit. Suppose we have a function f (x)
that is well-defined on an open interval around x0 , except possibly at x0 itself. If for all
x sufficiently close to x0 , the function f (x) gets arbitrarily close to x0 , we shall say that
f (x) approaches the limit L as x approaches x0 . Mathematically, we write

lim f (x) = L.
x→x0

Figure 2 shows an example of a limit value.

x
−1 0 1

Fig. 2 The function f (x) = (x2 − 1)/(x − 1). Note that f (x) is not defined at x = 1 because the
denominator becomes 0 at that point.

The limit of f (x) in Figure 2 as x gets closer and closer to 1, is 2. It need not, however, be
defined at x = 1. The equality of limit and function value is special, and is an important
feature of a continuous function.
In order to make the concept of a limit clear, we shall adopt the following definition.

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DEFINITION 1 (TC). Let f (x) be defined on an open interval about x0 , except possibly
at x0 itself. We say that f (x) approaches the limit L as x approaches x0 and write

lim f (x) = L,
x→x0

if, for every number ǫ > 0, there exists a corresponding number δ > 0 such that for all
x, if
x0 − δ < x < x0 + δ,
then
L − ǫ < f (x) < L + ǫ.

EXAMPLE 1. Let f (x) = 5x − 3 be defined over all real x, except at x = 1. Show that

lim (5x − 3) = 2.
x→1

SOLUTION: A naive way is to replace x = 1 in 5x − 3. However, this cannot be

done because f (x) is not defined at that point. We therefore prove the result using the
definition of limit. Our proof is done if we can find a δ such that f (x) lies within ǫ away
from L = 2. Working backwards, we solve

5x − 3 = 2 − ǫ; 5x − 3 = 2 + ǫ,

which gives us the two lower and upper bounds of x as 1 − ǫ/5 and 1 + ǫ/5. Since x0 = 1,
we can just choose δ = ǫ/5 (or any other δ < ǫ/5, since this will guarantee that f (x)
remains within ǫ from L = 2. ♣

To make things concrete in the above example, let us set ǫ = 0.005, and then choose
δ = 0.001. This means that for any x between 0.999 and 1.001, f (x) correspondingly
moves between 1.995 and 2.005. The limit L is then possible by making ǫ as small as we
wish. This shows the superiority of the formal definition over a naive plug-in approach,
which fails if f (x) is not well-defined at x0 .

3 Finding Limits
It would be very tedious indeed if we have to apply the definition of the limit every time
we want to find a limit value. Fortunately, we can invoke the following theorem to help
us find limit values.

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THEOREM 1 (TC). Let L, M, c and k be real numbers, and suppose

lim f (x) = L , lim g(x) = M.

x→c x→c

It follows that the following six rules are valid.

limx→c (f (x) + g(x)) = L+M (Sum Rule)

limx→c (f (x) − g(x)) = L−M (Difference Rule)

limx→c (f (x) · g(x)) = L·M (Product Rule)

limx→c (kf (x)) = k·L (Constant Multiple Rule)

limx→c fg(x)
(x)
= L
M
M 6= 0. (Quotient Rule)

limx→c (f (x))r/s = Lr/s s and r are integers , and s 6= 0. (Power Rule)

EXAMPLE 2. Let f (t) = (t2 +3t−10)/(t+5) be defined over all real t, except at t = −5.
Suppose we want to find (a) limt→5 f (t) and (b) limt→−5 f (t).
(a).

limt→5 (t2 + 3t − 10)

lim f (t) =
t→5 limt→5 (t + 5)
2
5 + 3(5) − 10
=
5+5
= 3.

(b). We first factorise the numerator to obtain (t + 5)(t − 2). Cancelling the common
factor in the numerator and denominator, we have

f (t) = t − 2 , t 6= −5.

Therefore,
lim f (t) = −7.
t→−5

It seems intuitive to cancel the common factor. However, if we try to apply the limit
quotient rule, we have to evaluate
t+5
lim .
t→−5 t + 5

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Clearly, substituting −5 into the numerator and denominator will result in 0/0, which is
undefined. To overcome this problem, we need the following result.

THEOREM 2 (TC). If the limit of quotients are of the form 0/0 or ∞/∞, then L’Hôpital’s
Rule states that  
f (x) f ′ (x)
lim = lim ′ ,
x→c g(x) x→c g (x)

where f ′ (x) and g ′ (x) denote the derivative of f and g with respect to x.

Note that we have not defined what a derivative is. For now, it is sufficient to
have an operational understanding of the latter (e.g. d(x2 )/dx = 2x, d(exp(x))/dx =
exp(x), d(sin(x))/dx = cos(x), d(log(x))/dx = 1/x, etc.)

EXAMPLE 3. In part (b) of EXAMPLE 2, the cancellation of the common factor under
limit operation is justified by L’Hôpital’s Rule, where we find that
t+5 1
lim = lim = 1.
x→−5 t + 5 x→−5 1

This means that we have

t+5
lim f (t) = lim (t − 2),
x→−5 x→−5 t + 5
t+5
= lim × lim (t − 2)
x→−5 t + 5 x→−5
= −7.

The limit rules in THEOREM 1 can also be applied in the case of infinite limits.

EXAMPLE 4. Find limx→∞ (1 + 1/x).

SOLUTION.  
1 1
lim 1+ = lim 1 + lim = 1.
x→∞ x x→∞ x→∞ x

We say that 1 is the horizontal asymptote of f (x). ♣

So far, we have been dealing with two-sided limits - that is, the limit of a function as x
approaches x0 from the values less than x0 as well as values greater than x0 . Occasionally,
a limit may fail to occur if the left-hand and right-hand limits are unequal.

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EXAMPLE 4. Consider the function,

0 :x<0
f (x) =
1 :x≥0

The left-hand limit of f (x) is

lim f (x) = 0,
x→0−

but its right-hand limit is

lim f (x) = 1.
x→0+

Since
lim f (x) 6= lim+ f (x),
x→0− x→0

limx→0 f (x) fails to exist. ♣

One-sided limits that take values ±∞ as x → x+ −

0 or x → x0 are known as vertical
asymptotes.

EXAMPLE 5. The function

1
f (x) = ,
2−x
has a vertical asymptote at x = 2, since
1
lim− = ∞,
x→2 2−x
and
1
lim+ = −∞.
x→2 2 − x

You should sketch f (x) to visualise the asymptote. ♣

Occasionally, a complicated function f (x) can be well-approximated by a simpler one

g(x) at the left end of f (x), if
f (x)
lim = 1;
x→−∞ g(x)

and at the right end of f (x), if

f (x)
lim = 1.
x→∞ g(x)

We say that g(x) is an end behaviour model for f (x).

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EXAMPLE 6. Find a simple function g(x) that can serve as an end behaviour model for
f (x) = x + log |x|.

SOLUTION. We first write f (x) in more detail,


x + log(x) :x>0
f (x) =
x + log(−x) : x < 0

If we choose g(x) = x, then the quotient

(
log(x)
f (x) 1+ x
:x>0
= log(−x)
g(x) 1+ x
:x<0

Taking limits, we have

 
log(x) 1/x
lim 1 + = 1 + lim = 1,
x→∞ x x→∞ 1
 
log(−x) 1/x
lim 1 + = 1 + lim = 1.
x→−∞ x x→−∞ 1

Note that we have used L’Hôpital’s Rule when evaluating limx→∞ log(x)/x. This shows
that g(x) = x is a reasonable end behaviour function for f (x). ♣

4 Continuity
Intuitively, we say that a function f (x) is continuous if it can be sketched over its domain
without pausing. The constant line f (x) = c, is an example of a continuous function
over the real line, since we can imagine drawing such a line without lifting our pen. To
be precise, however, we need the following definition.

DEFINITION 2 (TC). A function f (x) is continuous at an interior point c of its domain

if
lim f (x) = f (c).
x→c

It is continuous at a left endpoint a, or a right endpoint b of its domain, if

lim f (x) = f (a) , lim f (x) = f (b),

x→a+ x→b−

respectively.

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To check the continuity of a function f (x) at x = c, we need to ensure that the following
three conditions are satisfied:

1. f (c) exists .

2. limx→c f (x) exists .

3. limx→c f (x) = f (c).

EXAMPLE 7. The function

sin(x)

: x 6= 0
f (x) = x
1 :x=0

is continuous over the real line, including at c = 0. Condition 1 is obviously true, since
we have defined f (0). The second condition is also true, since by L’Hôpital’s Rule,

sin(x) cos(x)
lim = lim = 1.
x→0 x x→0 1
Finally, limx→0 f (x) = 1 = f (0), so Condition 3 is true as well. Note that if we did
not define f (0) = 1, the function would not have been continuous due to violation of
Condition 3.

Continuity of a function leads to the Intermediate Value Theorem (IVT).

THEOREM 3 (TC). Let y = f (x) be continuous on a closed interval [a, b]. It follows
that y takes on every value between f (a) and f (b).

Theorem 3 is very important for finding real roots, because it guarantees their existence.
Specifically, real roots are values of x that satisfy f (x) = 0. According to the IVT, if
f (x) is continuous in the interval [a, b], and that f changes sign in this interval, then
there must be at least one real root.

EXAMPLE 8. Why does cos(x) = x have at least one root?

SOLUTION. Rewriting the equality, we have cos(x) − x = 0. Denote the left-hand side
as f (x). Since f (x) is continuous for all real x, and that it changes sign in the interval
[0, π/2], it follows that the line 0 will cross f (x) at least once in that interval (the solution
is x = 0.739084).

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5 Rate of Change and Tangent Lines
Suppose we have a parabolic curve (y = x2 ), and we are interested in knowing the average
rate of change in y when x changes from 0 to 4. We find this by taking the ratio

δy 42 − 02
= = 4.
δx 4−0
This ratio corresponds to the slope of the secant line (the straight line passing through
(0,0) and (4,16) in Figure 3).

16 Q

δy

0 P x
4

δx

Fig. 3 The secant line passing through points P (0,0) and Q (4,16) on the function y = x2 .

If we are interested in finding the rate of change at the point x = 4 itself (the instanta-
neous rate of change), how should we go about doing it? We can start by choosing P to
be (4 − h, (4 − h)2 ), with h > 0, and then measure the slope of the resultant secant line,

δy 42 − (4 − h)2
= .
δx h
If we continue choosing P such that it becomes arbitrarily close to Q, then in the limit,
we have
δy 42 − (4 − h)2
lim = lim = 8.
h→0 δx h→0 h

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 We shall now give a formal definition of the slope and tangent line.

DEFINITION 3 (TC). The slope of the curve y = f (x) at the point Q(x0 , f (x0 )) is the
number
δy f (x0 + h) − f (x0 )
m = lim = lim ,
h→0 δx h→0 h
if the limit exists. We say that a line with slope m that goes through point Q at the
curve is tangent to Q.

EXAMPLE 9. The metabolism of drugs often leads to an exponential decay of the latter’s
concentration in the blood circulatory system. Let y = exp(−x) model such a process.
The half-life x1/2 of this process is the value of x where the concentration of drug y decays
to half of its original value at time x = 0. What is the rate of change at x1/2 ?

SOLUTION. In the beginning (x = 0), we have y = 1. At time x1/2 , this initial value
decreases to 1/2. Therefore, we solve for
1
= exp(−x1/2 ),
2
which gives x1/2 = log(2). Next, by definition, the secant line slope at x1/2 is given by

exp(−(x1/2 + h)) − exp(−x1/2 )

 
δy exp(−h) − 1
= = exp(−x1/2 ) .
δx h h

Taking limit, we have

 
δy − exp(−h)
lim = exp(−x1/2 ) lim
h→0 δx h→0 1
= − exp(−x1/2 )
1
= − .
2
♣

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6 References
1. Finney, R.L., Weir, M.D. and Giordano, F.R. (2001) Thomas’ Calculus. 10th ed.
Boston: Addison Wesley Longman.

2. Gause, G.F. (1932) Experimental studies on the struggle for existence. Journal of
Experimental Biology, 9: 389-402.
(A free copy is available at http://jeb.biologists.org/cgi/reprint/9/4/389.pdf)

KTF 2009/2010 (2)

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