Population growth

Rates of population change

- Discrete vs. instantaneous
- Crude vs. specific

Birth, death, and survivorship
- Life tables

Geometric growth

Exponential growth

Logistic growth

Population growth (BIDE)

BIDE dynamics determine growth
- birth rate (b) - death rate (d)
- immigration rate (i) - emigration rate (e)

closed population: N
t+1
= N
t
+ N
t
(m d)
open population: N
t+1
= N
t
+ N
t
(m + i d e)

Population in
the future
Population
now

Population growth

Rate: an amount of change (in something) over time

Rate =

0
=

simpleright?

Discrete rate: average rate of change during t
Instantaneous rate: rate of change over a very short time (t 0)
N
0

N
1
t
0
time t
1

t
N
P
o
p
u
l
a
t
i
o
n


Population growth

Crude rate: total change in numbers over time, i.e.

Specific rate: rate scaled per organism, i.e.

(
0

)


Reproductive rate (natality):

Specific reproductive rate = # of offspring per individual per unit time

Population growth (BIDE)

Problem: reproductive rate often varies with age
every age group will have its own R
0

age-specific reproductive rate (m
x
)
# of offspring per female of age X per unit time
age age-specific
repro rate
(x) (m
x
)
0 m
0

1 m
1

2 m
2

. .
. .
w m
w

Onset of
reproduction
End of
reproduction
Death due to
senescence
mirth

Population growth

How many offspring can be produced for every female present at start?
= m
0
+ m
1
+ m
2
+ + m
w
= m
x


Gross reproductive rate
= # of offspring produced during a females life IF she lives to age w

Need to know what % of population survives or
dies at each age X

age-specific survivorship (l
x
)
= % of cohort alive at start of age X

l
x
=

0

age age-specific
repro rate
(x) (m
x
)
0 m
0

1 m
1

2 m
2

. .
. .
w m
w

m
x
= GRR

Population structure
(survival)

3 typical patterns
of survival

Type I

Type II

Type III

Population growth

How many offspring can be produced for every female present at the start?
How many offspring are produced? (i.e., accounting for those that die)


# +1
#
=

+
1

.

net reproductive rate = R
0
If R
0
> 1 population is growing
If R
0
= 1 population remains constant
If R
0
< 1 population is declining

Population growth

Life tables

Age
(x)

Cohort size
(nx)

Age-specific
survivorship (lx)

Age-specific
repro (mx)
0 l
0
m
0

1 l
1
m
1

2 l
2
m
2


w l

l
x
m
x

l
0
m
0

l
1
m
1

l
2
m
2


l

xl
x
m
x

0l
0
m
0

1l
1
m
1

2l
2
m
2


wl

Number surviving
to age X
Proportion
surviving to age X
m
x
= GRR l
x
m
x
= R
0
xl
x
m
x
R
0
= T

Population growth

T = generation time

The average age at which
a female gives birth

Population growth

At what rate would the population increase if
- R
0
remains constant and >1
- There are no limits on population growth?

+
1

= R
0
N
(t+1)
= N
t
R
0

So, after 1 generation N
1
= N
0
R
0

2 N
2
= N
1
R
0
= (N
0
R
0
)R
0
= N
0
R
0
2
3 N
3
= N
2
R
0
= (N
0
R
0
2
)R
0
= N
0
R
0
3



So after X generations N
x
= N
0
R
0
x



Population growth (Life table example)

Consider a mouse that has a one year lifespan. Over the winter, 90% of individuals
that were born the previous spring die. However, each one that overwinters
produces 12 babies.

If a population begins with 100 baby mice, how many mice are there each
subsequent spring?




Year (x)

Cohort size (n
x
)

Age-specific
survival (l
x
)

Age-specific
repro (m
x
)

l
x
m
x


xl
x
m
x

0 1.0
1
2
100 0 0 0
10 12 1.2 1.2 0.1
0 0 0 0 0
GRR = 12 R
0
= 1.2 T = 1
Next spring(s):
100 x 1.2 = 120
120 x 1.2 = 144

Density independent growth?

N

(
o
r

d
e
n
s
i
t
y
)

Time

Density dependent growth?

N

(
o
r

d
e
n
s
i
t
y
)

Time

Geometric growth

discrete, non-overlapping generations

N
t
= N
0

t

N
t
= number of individuals at time t
N
0
= initial number of individuals
= geometric rate of increase
t = number of time intervals or generations

Geometric growth

996 x 2.4177
2
=
480,924

Exponential growth

Continuous population growth can be modelled exponentially

= r
max
N

dN/dt = change in the number of individuals per unit time
r
max
= intrinsic rate of increase
max possible under deal conditions

Exponential growth

Example

= r
max
N

t N dN/dt
0 100 0.2 x 100 = 20
1 120 0.2 x 120 = 24
2 144 0.2 x 144 = 29
3 173

Exponential growth

Size of the population at any time can be calculated as

N
t
= N
0
e


N
t
= number of individuals at time t
N
0
= initial number of individuals
e = the base of natural logarithms
r
max
= intrinsic rate of increase
t = number of time intervals

r
max
t

Exponential growth

Examples

Exponential growth

Under constant, stable conditions r r
max


r
max
is reached and maintained only under ideal conditions:
- resources not limiting
- competitive vacuum

Malthus & Darwin: populations have a great growth potential
e.g. rapid growth when introduced into empty habitat


www.english-country-garden.com/a/i/birds/pheasant-1.jpg

Exponential growth

Under constant, stable conditions r r
max


But r
max
is balanced by extrinsic factors

As population density increases:
- Per capita resources decline
- Growth
- Age at maturity
- Fecundity
- Higher densities attract predators, so mortality
- Social strife within populations and disease

Environmental resistance (birth rate , or death rate , or both)

Exponential growth

In many populations, growth eventually slows or stops (

= 0)
results in an S-shaped (sigmoid) growth curve

Upper, equilibrium level
carrying capacity (K)

Populations below K grow
- Below inflection point, growth is accelerating
- Above inflection growth > 0, but decelerating
Populations above K decrease
Populations at K remain ~constant


K is a property of the environment: the max. sustainable population size

Density dependent growth

N alters birth and death
rates

Intraspecific competition

Carrying capacity = K



Logistic growth




Logistic population growth

dN/dt = r
max
N

r
max
= intrinsic rate of increase
K = carrying capacity

When N nears K, the right side of the equation nears zero
change in the population size approaches zero
When N is low, the right side of the equation is near 1
change in the population size is exponential



Logistic population growth

Size of the population at any time can be calculated as

N
t
=

1+

0
1

N
t
= number of individuals at time t
N
0
= initial number of individuals
e = the base of natural logarithms
r
max
= intrinsic rate of increase
t = number of time intervals
K = carrying capacity




Logistic population growth

What is the relationship between density and per capita rate of increase?


Logistic population growth

What is the relationship between density and change in population size?


Logistic population growth

Examples