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# Biological Science 5th Edition Freeman Quillin Allison Instructors Manual

ISBN-13: 978-0321841803 | ISBN-13: 978-0321841803 |

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CHAPTER

Population Ecology

54

Learning Objectives: Students should be able to…

• Use life tables to describe how likely it is that individuals in each age class in a population

will survive and reproduce.

• Calculate the growth rate of a population from life-table data or from direct observation of

changes in population size over time.

• Describe the variety of patterns in population size changes over time.

• Explain the link between understanding population dynamics and applying the data to

endangered species.

Lecture Outline

• A population is a group of individuals from the same species that live in the same area at

the same time.

• Population ecology is the study of how and why the number of individuals in a population

changes over

time.

I. Distribution and Abundance

1. Abiotic and biotic factors both determine the range of a species.

2. The range of a species can also be considered at different scales.

a. The population density of the common lizard varies throughout its range.

(Fig. 54.1)

3. In general individual organisms can be arranged in different patterns.

a. random—position of each individual is independent of the others

b. clumped—organisms associate in groups

c. uniform—even distribution usually associated with competition

4. A metapopulation is a population of populations connected by migration.

5. Immobile organisms are easy to assess by using quatrats or transects and simply

counting individuals.

6. Mobile animals are more difficult to assess. Mark-recapture studies are one method.

(Quantitative Methods 54.1)

a. The relationship between marked and unmarked individuals can be expressed as

m2/n2 = m1/N. (Eq. 54.1)

b. m1 is the number of animals marked in the first sample; m2 is the number of marked

individuals in the second sample; n2 is the total number of individuals in the second

sample; N is the total population size.

Copyright © 2014 Pearson Education, Inc.

555

7. Students should be able to complete this example: Suppose researchers marked 255

animals and later were able to trap a total of 162 individuals, of which 78 were

marked. What is the estimate for the total population size?

II. Demography

A. Demography is the study of factors that determine the size and structure of populations

through time.

1. Analyzing birth rates, death rates, immigration rates, and emigration rates is fundamental

to demography.

2.

A

generation is the average

time between

a mother’s

first offspring

and

her daughter’s

first

offspring.

B. Life tables

1. Formal demographic analyses of populations are based on a type of data set called a

life table, which summarizes the probability that an individual will survive and reproduce

in any

given year

over

the course of its lifetime.

2.

Lacerta vivipara:

a case study

a.

Researchers

set

out to estimate the life table

of a low-elevation population of

L.

vivipara

in the Netherlands with the goal

of

comparing

the results to data from

other

L.

vivipara

populations.

b.

The

data allowed them

to calculate the number

of individuals that survived each

year

as well as the number

of

individuals

produced per

female.

(Table

54.1)

(1)

An

age class is

a group of

individuals

of a

given age.

3.

Survivorship

a.

Survivorship

is

the proportion

of offspring

produced

that

survive, on

average, to a

particular

age (lx,

where x

represents the age

class

being considered).

b.

Survivorship

is

calculated by

dividing

the

number

of individuals

in

a given age

class

by

the number

of individuals that existed as offspring: lx

= Nx

/No.

(Quantitative

Methods 54.2,

Eq. 54.2)

c.

To recognize general patterns in survivorship and make comparisons among populations

or species, biologists

plot the logarithm

of the number

of survivors versus

age,

which is called a survivorship curve.

(Fig.

54.2;

BioSkills

6 in

Appendix B)

(1)

Type

I curve:

Survivorship

throughout

life

is high.

(2) Type II curve: Constant mortality occurs throughout life.

(3) Type III curve: The death rate is high early in life.

4. Fecundity

a. Fecundity is the number of female offspring produced by each female in the population.

b.

The

net reproductive

rate of a population

is calculated as:

R0

= Σ

lx

mx.

(Quantitative

Methods 54.2,

Eq. 54.3)

c.

Lifetime reproduction is a function of fecundity at each age (mx) and survivorship to

each age class (lx). If R0 is greater than 1, the population is increasing. If R0 is less

than 1, the population is declining.

d. Age-specific fecundity is the number of female offspring produced by a female in

age class x.

556 INSTRUCTOR GUIDE FOR BIOLOGICAL SCIENCE, 5e

Copyright © 2014 Pearson Education, Inc.

5. Students should be able to use the data in Table 54.1 to determine how many female

offspring an average L. vivipara female produces over the course of her lifetime,

and describe whether the population is growing, stable, or declining.

C. The role of life history

1. An organism’s life history consists of how it allocates resources to growth, reproduction,

and activities related to

survival.

2.

It

is not possible

for an organism

to have both high

fecundity

and high

survivorship.

A

trade-off

exists between survival

and reproduction.

3.

Fitness

trade-offs

occur because every

individual

has a restricted amount

of time and

energy

at its disposal;

that is,

its resources are limited.

4.

If

a female

lizard

devotes a great deal of energy

to feeding a large number

of offspring

as

they

develop,

it is not

possible

for her to devote that

same energy

to her immune

system,

growth,

nutrient stores, or other traits

that increase survival.

5.

In

almost

all

cases,

biologists find that life

history

is shaped by

natural

selection in a

way

that maximizes

an individual’s

fitness

in its environment.

6.

Students

should be able to (1) predict how

survivorship

and fecundity should compare

in L.

vivipara

in the

warmest versus

coldest parts

of their range, and (2) comment

on why females

in these

populations lay many eggs instead of giving birth to a

relatively

small

number of live young.

7.

In

general, organisms

with high fecundity

tend

to

grow

quickly,

reach

sexual maturity

at

a young

age, and produce many

small eggs or seeds.

8.

In

contrast, organisms

with high survivorship

tend to grow slowly

and

invest resources

in

traits that reduce damage

from

enemies

and increase their own ability

to

compete

for

water, sunlight, and food.

9.

An Arabidopsis

thaliana

plant and a coconut palm

represent

the two ends of a broad

continuum

of life-history

characteristics. (Fig.

54.3)

III. Population Growth

A. Quantifying the growth rate

1. Populations are affected by birth and immigration rates as well as death and emigration

rates.

2.

A

population’s

growth rate is the change in the number

of individuals

in the population

(ΔN)

per unit

time

(Δt).

3. If no immigration or emigration is occurring, then the growth rate is equal to the number

of individuals

(N)

in the

population

times

the difference between the birth

rate per

individual

(b)

and the death

rate per individual

(d).

4.

The

difference between the

birth rate and the

death rate per individual is called the

per-capita

rate

of increase and

is symbolized

r.

5.

A

species

with

a certain life history

has an

intrinsic rate of increase (rmax)

that does not

change.

But at any

time,

each population

of that species has an instantaneous growth

rate,

or per-capita

rate of increase,

symbolized

by

r.

B. Exponential growth

1. Exponential growth occurs when r does not change over time. It does not depend on

the number of individuals in the population. Therefore, it is density independent.

(Fig. 54.4)

Copyright © 2014 Pearson Education, Inc.

CHAPTER 54 Population Ecology 557

2. In nature, exponential growth has been observed in two circumstances:

a. A few individuals found a new population in a new habitat.

b. A population was devastated by a storm or other type of catastrophe and then began

to recover, starting with a few surviving individuals.

3. In reality, it is not possible for growth to continue indefinitely because the habitat

would not have enough resources for the number of individuals present.

4. When a population stabilizes at the maximum number that can be supported by the resources

available,

that population has reached

the habitat’s

carrying

capacity.

C.

Logistic

growth

1.

The

carrying

capacity

(K)

of a habitat depends

on a large

number

of

factors: food,

space,

water,

soil

quality,

resting or nesting

sites, and the

intensity

of

disease and

predation.

Carrying

capacity

may

change from

year

to year, depending

on conditions.

2.

The

logistic growth equation describes logistic

population

growth,

or changes in the

growth

rate that occur as a function of

population

size. Just as exponential growth

is

density

dependent, logistic growth

is density

dependent.

3.

Discrete

growth

(Quantitative

Methods 54.3)

a.

Biologists use

N

to symbolize

population

size. N0

is the population

size at time zero,

and

N1

is the population

size

one breeding interval later.

b.

The

growth rate is calculated

as λ

=

N1/N0,

where λ

is the finite rate of increase.

(Eqs.

54.4 &

54.5)

c.

The size of the population at the end of year t is calculated as Nt = N0λ

t

. (Eq. 54.6)

This equation summarizes how populations grow when breeding takes place seasonally.

The size

of the population

at time t

is equal to the

starting size times

the

finite

rate of increase multiplied

by

itself

t

times.

(1)

In

a sense, λ

works like the interest rate at a

bank. For species that breed

once

per

year,

the “interest”

on the population

is compounded

annually.

d.

If

a population’s

age structure

is stable, then

its finite rate

of increase has a simple

relationship

to

its net

reproductive rate: λ

= R0/g

where

R0

is the net

reproductive

rate

and g

is the generation time. (Eq.

54.7)

4.

Continuous

growth

(Quantitative

Methods 54.3)

a.

A

population’s

per-capita increase is symbolized

r

and is defined as the per-capita

birth

rate minus

the per-capita death rate.

b. The relationship between the finite rate of increase and the per-capita growth rate is

λ = e

r

, where e is the natural logarithm, or about 2.72. (Eq. 54.8) (See BioSkills 6 in

Appendix B.)

c. The size of the population at the end of year t is calculated as Nt = N0e

rt

. (Eq. 54.9)

This equation summarizes how populations grow when they breed continuously.

Because r represents the growth rate at any given time and because r and λ are so

closely related, biologists routinely calculate r even for species that breed seasonally.

d.

The

instantaneous rate of increase,

r,

is also directly

related

to the net reproductive

rate: r = ln R0/g. (Eq. 54.10)

5. In summary, biologists have developed several ways of calculating and expressing a

population’s growth rate. λ has the advantage of being easy to understand, and R0 has

the advantage of being calculated directly from life-table data. Although r is slightly

558 INSTRUCTOR GUIDE FOR BIOLOGICAL SCIENCE, 5e

Copyright © 2014 Pearson Education, Inc.

more difficult conceptually, it is the most useful expression for the growth rate

because it is independent of generation time and is relevant for species that breed

either seasonally or continuously.

6. Applying the models

a. Students should be able to solve the problems at the end of Quantitative Methods

54.3.

7. Graphing logistic growth

a. Density-dependent graphs have three sections: (Fig. 54.5a)

(1) Initially growth is exponential, meaning that r is constant.

(2) N increases to the point at which competition for resources or other densitydependent

factors begins to

occur. Growth

rate begins to decline.

(3)

Eventually

the growth rate reaches

0; that

is, the habitat’s

carrying

capacity

is

reached.

b.

For

example,

both

species of Paramecia

exhibit logistic

growth, but

the carrying

capacities

differ. (Fig.

54.5b)

D. What limits growth rates and population sizes?

1. Density-independent factors change birth rates and death rates irrespective of the

number of individuals in a population.

2. Density-dependent factors are usually biotic and change in intensity as a function of

population size.

3. A closer look at density dependence

a. Researchers studied the bridled goby, a coral-reef fish. They stocked artificial reefs

with varying densities of adult gobies. After 2.5 months, they captured all the

gobies and computed the growth rate of individuals, the survival rate, and the

immigration rate.

b. Adult gobies survive better when population density is low. More juvenile gobies

immigrate successfully when population density is low. Higher rates of predation

and disease might occur in dense populations. (Fig. 54.6a)

c. There is a strong density-dependent relationship for clutch size in sparrows.

(Fig. 54.6b)

d. Density-dependent changes in survivorship and fecundity cause logistic population

growth. In this way, density-dependent factors define a particular habitat’s carrying

capacity.

4. Carrying capacity is not fixed. It varies in space and time because conditions in some

years are better than in others.

IV. Population Dynamics

A. Population dynamics are changes in populations over time.

1. Researchers have uncovered a wide array of patterns in natural populations in addition

to exponential and logistic growth.

B. How do metapopulations change through time?

1. Because humans are reducing large, contiguous areas of forest and grasslands to isolated

patches or

reserves, more

and more

species

are

being forced into

a metapopulation

structure⎯a

population

of populations.

(Fig.

54.8)

Copyright © 2014 Pearson Education, Inc.

CHAPTER 54 Population Ecology 559

2. Ilkka Hanski and colleagues showed that metapopulations exist in nature. They performed

a mark-recapture

study

on

the Glanville

fritillary

butterfly.

They

found a

migration

rate of 9%,

which was high enough

to recolonize

extinct populations.

They

also

confirmed that some

populations

had gone

extinct while others had been created.

3.

Given

enough

time,

each population

within

the

larger

metapopulation is expected to

go

extinct. The

cause could

be catastrophic, such as a storm

or oil spill, or a disease

outbreak

or a sudden

influx

of predators.

4.

Migration

from

nearby

populations can reestablish

populations in these

empty

habitat

fragments.

In this way,

there

is a balance

between extinction and recolonization.

C. Why do some populations cycle?

1. Population cycles are regular fluctuations in size that some populations exhibit.

2. For example, the snowshoe hare and lynx populations in northern Canada exhibit

regular cycling. (Fig. 54.9)

3. Two hypotheses were proposed for an experiment: (Fig. 54.10)

a. Hares use up all of their resources when population density is high, and they starve.

In response, lynx also starve.

b. Lynx populations reach high density in response to an increase in hare density.

Lynx eat so many hares that the prey population crashes.

4. The data support the hypothesis that hare populations are limited by the availability of

food as well as by predation and that these two factors interact. This explains why

when hares are at high density, individuals are weakened by nutritional stress and

therefore more susceptible to predation.

V. Human Population Growth

A. Age structure in human populations

1. Analyzing an age pyramid can give biologists important information about a population’s

history.

2.

In

countries where

industrial and technological development

is advanced

and average

incomes

are

relatively

high, the age pyramid

of the population

tends

to be even. (Fig.

54.11a)

3.

The

age pyramid

predicted for developed

countries in

2050

highlights

their major public

policy

concern: How to care for

an increasingly

aged population.

4.

In

contrast, the age distribution is bottom-heavy

in

less-developed countries. (Fig.

54.11b)

5. The age pyramid predicted for less-developed countries in 2050 illustrates their major

public policy concern: providing education and jobs for an enormous influx of young

people who will want to be starting families.

6. Overall population size will increase dramatically in developing countries. Even

though the number of children per female (fecundity) will drop, there will be so many

women having children that the number of offspring born per year is expected to stay

high.

B. Analyzing change in the growth rate of human populations

1. The growth rate for humans has been increasing since about 1750, leading to a steeply

rising curve over the past few centuries. The highest values occurred between 1965

and 1970, when population growth averaged 2.04% per year. (Fig. 54.12)

560 INSTRUCTOR GUIDE FOR BIOLOGICAL SCIENCE, 5e

Copyright © 2014 Pearson Education, Inc.

2. It is not possible to overstate the consequences of recent and current increases in human

population.

It is the primary

cause of habitat loss, declines in living standards,

mass

movement

of people, political instability,

and acute

shortages of water and

other

resources.

3.

The

one encouraging

trend

in the data is that

the growth

rate of the human

population

has

already

peaked

and begun to

decline.

4.

Will

the size of the human

population

peak

in your

lifetime?

a.

Population

growth

has slowed over the past few years;

now it is 1.2%

per year.

Humans

are ending

a period

of rapid growth

that lasted well over 500

years.

How

quickly

growth

rates decline and the maximum

population size is ultimately

reached

will depend

on changes in fertility

rates and the course of the AIDS epidemic.

b.

Extrapolating

the world’s

population to the

year

2050 is

based on three different

fertility

rates: (Fig.

54.13)

(1)

High fertility⎯nearly

11 billion

(2)

Medium

fertility⎯about

9.3 billion

(3)

Low fertility⎯8

billion

c.

The future of the human population hinges on fertility rates—on how many children

each of the women living today decides to have.

VI. How Can Population Ecology Help Conserve Biodiversity?

A. Using life-table data

1. Population projections enable biologists to alter values for survivorship and fecundity

at particular ages and assess the consequences.

2. Using the life table data in Figure 54.14a, calculations are shown to predict how many

adults (of 1000 introduced lizards) will survive and how many offspring from each age

class will be produced. (Fig. 54.14b)

3. Figure 54.14c extends the calculations by predicting what will happen as the offspring

of the original females begin to breed.

4. Students should be able to continue the analysis into the third year and predict

whether the population of lizards will stay the same, decline, or increase over time.

5. Studies performed on life-table data support some general conclusions.

a. Whooping cranes, sea turtles, spotted owls, and many other endangered species

have high juvenile mortality, low adult mortality, and low fecundity. (Fig. 54.15)

b. Climate change has enormous implications for the life-history traits of populations.

Example: Temperature determined sex in reptiles—significant implications for

growth rates.

6. In some cases, projections made from life tables may be too simplistic to be useful.

B. Preserving metapopulations

1. Conservationists draw heavily on concepts and techniques from population ecology

when designing programs to save species threatened with extinction.

2. A small, isolated population—even one within a nature reserve—is unlikely to survive

over the long term. (Fig. 54.16)

3. Results on the Glanville fritillaries have important messages for conservation biologists.

Copyright © 2014 Pearson Education, Inc.

CHAPTER 54 Population Ecology 561

a. Areas being protected need to be large enough in area to maintain large populations.

b. If that is not possible, smaller tracts of land should be connected by corridors of

habitat so that migration is possible.

c. Land where populations have emigrated should be preserved so that immigration is

possible.

Chapter Vocabulary

To emphasize the functional meanings of these terms, the list is organized by topic rather

than by first mention in the chapter. It includes terms that may have been introduced in earlier

chapters but are important to the current chapter as well. It also includes terms other than

those highlighted in bold type in the chapter text.

population

per-capita rate of

increase (r)

quadrats

evolutionary

ecology

intrinsic rate of

increase (rmax)

mark-recapture

study

immigration

exponential growth

population

cycles

demography

population density

age pyramid

generation

density-dependent

replacement rate

life table

carrying capacity (K)

zero population

growth (ZPG)

cohort

logistic growth equation

survivorship curve

logistic population growth

fecundity

finite rate of

increase (λ)

population viability

analysis (PVA)

age-specific fecundity

density-independent factors

age class

density-dependent factors

fitness trade-offs

life history

population dynamics

net reproductive rate

metapopulation

emigration

density-independent

age

structure

Lecture Activities

Population Exercises

Activity Ideas for Any Class Size

Estimated duration of activity: 5 to 10 minutes

• The human population has grown at a nearly exponential rate during the past several centuries

and, although it has shown recent signs of slowing,

continues to increase at an

impressive

rate. Is the

human

population close

to reaching

Earth’s

carrying

capacity,

or

have we

already

reached it? What would

happen

if

the human

population surpassed

Earth’s

carrying

capacity?

Is it

possible for any

population

to do

this?

How

and when would you

go

about

calculating

the carrying

capacity

of

the environment

for any

given species

in its particular

habitat?

What

is the carrying

capacity

of Earth

most

dependent on,

and how

might

it be

calculated?

• At high numbers, population growth becomes density dependent and is limited by factors

that operate better under high-density situations, such as disease, competition, and predation.

But ecologists

know that

population

growth

at all

levels is also limited

by

densityindependent

factors.

What is meant

by

the

term

density-independent?

How would

densityindependent

factors

affect a

population

and

work

to inhibit

or

even

stop population

562 INSTRUCTOR GUIDE FOR BIOLOGICAL SCIENCE, 5e

Copyright © 2014 Pearson Education, Inc.

growth? What are some examples of density-independent factors that operate in this

capacity?

• It is the year 2525, and the human race can travel quickly and easily between the stars and

planets. While out exploring the universe one day, you discover a new planet that is essentially

identical to Earth, except it has no humans,

cities, or constructs of any

kind.

Otherwise,

the planet has all the same

wildlife,

plants,

geographical features,

and climate.

A few

years

later your

superiors tell

you that

they

have

found

thousands

of

volunteers

who are

eager

to inhabit this new planet. It is your job

to

organize

the initial colonization of this

new

planet. Where on this

“earth” would

you

start your

colony?

Why

have you

chosen

your

particular

location? Would

you

colonize

in

several different locations or just one?

What

factors enter into your

decision? What

kind of

age

structure and sex ratio would you

want

to start with? Why?

What

is likely to happen

to

the human

population

once

it becomes

established

in your

location?

What kind of

growth

and growth

rate will the new

population

exhibit?

After several years,

how will

its growth rate, survivorship, and age

structure

change?

• Try an exercise related to the survivorship/fecundity graph. First, choose some animals

with high fecundity, such as an octopus. Lead the students to discover that this animal lays

an incredibly large number of eggs per reproductive event. Then ask the students to guess

the life span of this species. After a few of these examples, see whether students can see

the relationship between massive reproductive output and short life expectancy. Now, if

time permits, switch to high-survivorship, low-fecundity species—like an elephant. Continue

the same

exercise

with

these types of species, ending

with humans.

At this point, students

should realize

that humans

have low fecundity

but

high

survivorship.

Does

this

change

with different regions of the world,

or

is it all

the same?

How would this

change if

most

women

on

Earth had 20

children over

the course

of their lifetime

and this continued

for

500 years?

Can you

find a correlation within

humans in the literature that supports the

survivorship/fecundity

relationship,

or doesn’t

this

occur within species?

Copyright © 2014 Pearson Education, Inc.

CHAPTER 54 Population Ecology 563