Population Homeostasis

Population Homeostasis

Christopher J. Paradise, PhD

A. Malcolm Campbell, PhD
Population Homeostasis
Copyright Christopher J. Paradise and A. Malcolm Campbell. 2016.

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 Abstract
This book will synthesize the concepts of selection against individuals in
response to environmental change to illustrate how selection against indi-
viduals results in homeostasis at the population level. For instance, selec-
tion against the light phenotype of the peppered moth during the early
part of the industrial revolution led to an increase of the dark phenotype,
which was better camouflaged against the soot that accumulated on tree
bark as a result of burning coal. Populations are shown to be regulated by
feedback mechanisms, several of which are discussed here. Populations
are regulated by extrinsic factors, such as competition and predation, and
that lead to changes in intrinsic factors, such as reproduction. Changes in
population density often lead to initiation of feedback mechanisms, such
as changes in birth or death rates. In a final example, pollutants are shown
to be a factor that can disrupt homeostasis of populations. In particular,
populations of top predators, such as raptors, have suffered due to bio-
magnification of toxins.

Keywords
predator, energy budget, populations, homeostasis, feedback mechanism,
camouflage, mortality, density-independent, pollutant, biomagnification,
density-dependent, predation, evolution, natural selection, birth rate,
death rate, density, dispersion
 Contents
Preface...................................................................................................ix
Acknowledgments....................................................................................xi
Introduction.........................................................................................xiii
Chapter 1 Death of a Single Individual Affects a Population...............1
Ethical, Legal, Social Implications: Saving Individuals
May Not Help to Save an Entire Population.................11
Chapter 2 Populations Are Regulated Through Feedback
Mechanisms.....................................................................15
Chapter 3 Biomagnification of DDT Affects Raptor
Populations......................................................................31
Conclusion............................................................................................37
Glossary................................................................................................39
Index....................................................................................................41
 Preface
This book about population homeostasis is part of a thirty book series
that collectively surveys all of the major themes in biology. Rather than
just present information as a collection of facts, the reader is treated more
like a scientist, which means the data behind the major themes are pre-
sented. Reading any of the thirty books by Paradise and Campbell pro-
vides readers with biological context and comprehensive perspective so
that readers can learn important information from a single book with
the potential to see how the major themes span all size scales: molecular,
cellular, organismal, population and ecologic systems. The major themes
of biology encapsulate the entire discipline: information, evolution, cells,
homeostasis and emergent properties.
In the twentieth century, biology was taught with a heavy emphasis
on long lists of terms and many specific details. All of these details were
presented in a way that obscured a more comprehensive understanding.
In this book, readers will learn about several examples of population ho-
meostasis and some of the supporting evidence behind our understand-
ing. The historic and more recent experiments and data will be explored.
Instead of believing or simply accepting information, readers of this book
will learn about the science behind population homeostasis the way pro-
fessional scientists dowith experimentation and data analysis. In short,
data are put back into the teaching of biological sciences.
Readers of this book who wish to see the textbook version of this
content can go to www.bio.davidson.edu/icb where they will find
pedagogically-designed and interactive Integrating Concepts in Biology
for introductory biology college courses or a high school AP Biology
course.
 Acknowledgments
Publishing this book would not have been possible without the generous
gift of Dr. David Botstein who shared some of his Breakthrough Prize
with co-author AMC. Davids gift allowed us to hire talented artists (Tom
Webster and his staff at Lineworks, Inc.) and copyeditor Laura Loveall.
Thanks go to Kristen Mandava of Mandava Editorial Services for project
management and guidance. In particular, we are indebted to Katie Noble
and Melissa Hayban for their many hours and attention to detail.
Kristen Eshleman, Paul Brantley, Bill Hatfield and Olivia Booker
helped us with technology at Davidson College. We are grateful to ad-
ministrators Tom Ross, Clark Ross, Carol Quillen, Wendy Raymond,
Verna Case, and Barbara Lom who had confidence in us and encouraged
us to persist despite setbacks along the way.
Thanks to my wife Amy Brooks for her constant support during the
development of this textbook, and my daughter Evelyn for her endless
energy. Thanks to Malcolm Campbell for his steadfast resolve and opti-
mism. Without him, this book would not exist. Thanks to collaborator
Laurie Heyer for taking my sometimes half-baked math ideas and turning
them into powerful and elegant Bio-Math Explorations. I learned a lot
from both of them. While the math is largely absent from this book,
our collaboration with her made this a better book. Nancy Stamp at
Binghamton University, and Bill Dunson and Richard Cyr at The
Pennsylvania State University influenced me greatly in how I think as
a scientist and approach my teaching. Finally, I thank my students in
Integrated Concepts in Biology II, who enthusiastically participated in
our experiment to redesign introductory biology, starting with the text
and ending with a new approach to teaching biology.
 Introduction
What happens to a colony of bacteria growing on a Petri dish? Can the
human population continue to expand on a finite planet? What causes
a locust population to be extremely dense one year and small the next?
How does a population exposed to a pollutant respond when homeostasis
is disrupted? Homeostasis at the population level involves a couple of
ideas. One is that natural selection maintains the fit of the organism to its
environment. But the environment is constantly changing, so feedback
from the environment changes the genetics of the population through
natural selection, which lags behind changes in environment. Homeosta-
sis involves regulating population sizes through negative feedback mecha-
nisms. If a population grows too large, its growth may slow down or
even reverse. All life requires energy; and if energy is limited, individuals
must make decisions about how to allocate their resources and this affects
populations. In this book, the mechanisms that maintain homeostasis
in populations of animals and plants will be explored. By the end of the
book, readers will better understand how populations are maintained and
regulated in changing environments.
 CHAPTER 1

Death of a Single Individual

Affects a Population

At some point in life, readers have probably watched a nature show that
depicted a predator, perhaps a lion, taking down its prey. Others may
have even witnessed a predator-prey interaction, such as a hawk attacking
a rabbit or a cat killing a mouse. Compassionate humans may have hoped
that a zebra in the clutches of a lion would escape and live to tell the tale.
But lions must eat too in order to maintain a positive energy budget, and
some might have had compassion for the hungry lion. An energy bud-
get is a measure of the energy entering and leaving a biological system.
Energy is transformed in cells and organisms to maintain homeostasis. In
this book, the concept of homeostasis will be applied to populations and
the processes that maintain such homeostasis. The death of a zebra may
affect the zebra population if that death leads to changes in the biological
system (i.e., the population) in the context of the environment.
The death of an individual may affect populations in different ways.
Many birds are predators on various small animals, including spiders and
moths, and the effects of bird predation on these populations have been
well studied. In a classic study on the effects of bird predation on arthro-
pods, Bernard Kettlewell examined predation on peppered moths, Biston
betularia, in the United Kingdom. There are several color morphs, or
phenotypes, in populations of peppered moths with individuals being
either whitish grey with a sprinkling of black dots (called typica), or being
of one of two melanic forms, called carbonaria and insularia. Carbonaria
individuals are completely dark brown except for small white dots on
2 POPULATION HOMEOSTASIS

the forewings and head, and insularia individuals are dark with a sprin-
kling of white scales. These nocturnal moths spend the day resting on tree
trunks and branches.
Prior to the Industrial Revolution, most individuals were of the
light-colored morph, but coal burning in England caused the trees on
which peppered moths rested to become blackened by soot. Scientists
observed that the dark phenotype in peppered moth populations began
to increase in frequency. This suggests a feedback mechanism operat-
ing to bring the system to homeostasis in a changed environment. Ket-
tlewell surveyed populations of peppered moths throughout the United
Kingdom in the 1950s and found variation in the proportions of the
three morphs that correlated with industrial centers. Higher proportions
of the carbonaria form were found in populations closer to or downwind
from industrial centers in England, whereas the typica form was the only
morph found in several populations well away from or upwind of indus-
try and coal burning.
Kettlewell hypothesized that predation by birds was the mechanism
acting on moth populations in environments where the phenotypes did
not have equal susceptibility to predation. Depending upon the back-
ground on which moths rested, moths were either more or less con-
spicuous to predators. Camouflage, which is concealment by means of
disguise or protective coloration, only works if preys are indistinguishable
from the background. If coloration does not help an individual blend in,
then it stands out to predators. In an environment where resting places
of moths have changed color, certain phenotypes may have a selective
disadvantage. To test this hypothesis, Kettlewell performed several experi-
ments. He defined conspicuousness of moths as visible to the human eye
at a distance of 9 meters or more and inconspicuousness as not visible to
the human eye at distances less than 9 meters. Kettlewell and a colleague
then released moths into forests and determined their conspicuousness
(Figure 1). The first forest contained approximately 90% oak trees with
dark bark and 10% birch trees with light-colored bark. The second had all
the trees and rocks covered by lichens, which gave the surfaces a mottled
light-colored look. Lichens are composite organisms made up of a fun-
gus that grows symbiotically with algae that forms a crust-like growth on
rocks or trees.
 Death of a Single Individual Affects a Population 3

100
= typica
80 = carbonaria
= insularia
percentage
60

40

20

0 0 0 n/a
0
oak (dark bark) birch (light bark) lichen-covered
surface type

Figure 1 Percentages of peppered moths deemed conspicuous by

researchers in one of two forests, one of which contained oaks and
birch and the other that contained lichen-covered trees and rocks.
Percentages of 0 are noted; n/a indicates that insularia morphs were
not released in the second forest.
Source: From Kettlewell, 1955, Tables 2 and 3.

Kettlewell conducted a mark-release-recapture experiment in which

large numbers of males were released into two forests, one exposed to
soot pollution, and one in a forest not exposed to pollution and with
lichen-covered trees. To increase the probability of recapturing marked
individuals and reduce the probability of escapees leaving the forests, Ket-
tlewell selected forests that were surrounded by open fields, gardens, and
water, which the moths are reluctant to cross. Moths were reared in a
laboratory, and males were released daily for almost 2 weeks. Each moth
was marked on the underside with a small dab of paint. Males were used
because they could be recaptured more easily than females using baited
traps. Each night Kettlewell and his assistants collected moths from traps
baited with a pheromone emitted by females to attract males or with light.
The traps were set around the perimeter of each forest. All peppered moths
caught were counted and examined for their sex and whether or not they
were marked (Figure 2). The biologist also determined the percentage of
each phenotype in the unmarked population (see Figure 2B).
Kettlewell devised an experiment and made observations to ensure
that birds were indeed preying on moths. Equal numbers of typica and
4 POPULATION HOMEOSTASIS

60 recaptured males
= typica
50 = carbonaria
= insularia
percentage

40

30

20

10

0
A polluted-1953 polluted-1955 lichen-covered

100 natural population

80
percentage

60

40

20

0
polluted-1953 polluted-1955 lichen-covered
B forest

Figure 2 Results of studies of peppered moths in two forests.

A, Percentage of each color morph recaptured in mark-release-
recapture experiments. B, Percentage of each color morph in natural
populations in same forests used for recapture experiments.
Source: Data from Kettlewell, 1956, Tables 1 and 5.

carbonaria individuals were placed on surfaces in an aviary, a flight cage for

birds. Half of each type was on surfaces that made them conspicuous and
half was on surfaces that made them inconspicuous. Two birds were then
allowed access to the aviary. Within 3 hours, all conspicuous moths had
been eaten, and 40% of the inconspicuous moths had been eaten. In ex-
periments where moths were released into forests, observations were made
of tree trunks where peppered moths rested during the day, and it was
confirmed on numerous occasions that birds were preying on the moths.
 Death of a Single Individual Affects a Population 5

The composition of phenotypes in populations of peppered moths is

correlated with the environment. In forests that have trees with dark bark
or trees covered in soot populations have a high proportion of carbonaria
morphs. Soot covered forests are more likely to be close to industrial cen-
ters where burning of coal was common in the eighteenth century. In fact,
many naturalists observed the increase in carbonaria morphs in popula-
tions during the industrial development of Great Britain. In forests with
lichen-covered trees, high proportion of birches, or low levels of pollution
populations have a high proportion of typica morphs. During the late
twentieth century when the UK enacted air pollution regulations, forests
covered in soot began to return to their previous character, and along with
changes in the trees and surfaces where moths rested came a change in
the frequencies of moth phenotypes. Changes in the background where
peppered moths rested determine the frequency of phenotypes in the
population.
Kettlewell showed that predation was the mechanism causing
negative feedback and maintaining homeostasis in peppered moths.

He reasoned correctly that light-colored typica moths on soot-covered

trees or dark-barked oaks were more conspicuous to bird predators than
carbonaria moths on the same surface. By first demonstrating moth con-
spicuousness for human observers with a visual system similar to birds,
Kettlewell could use the argument that bird predators might select moths
that are easier to see as prey before they find inconspicuous prey. His re-
sults showed light-colored or mottled moths on oak bark are much more
conspicuous than the solid dark-colored carbonaria moths. Typica and
insularia moths on birch bark, and typica moths on lichen-covered sur-
faces are completely inconspicuous to humans, as defined by Kettlewell.
Carbonaria, however, stands out.
Kettlewells mark-release-recapture experiments and his other ob-
servations showed that birds did prey upon peppered moths, and they
discovered conspicuous prey more easily than inconspicuous prey. This
predation was most likely the cause of the lower percentages of recap-
tured conspicuous phenotypes in a particular environment. The selection
against conspicuous phenotypes by a predator could lead to high propor-
tions of inconspicuous phenotype. Further, if the environment changes
in such a way that a previously inconspicuous phenotype becomes
6 POPULATION HOMEOSTASIS

conspicuous, selection would then act against that phenotype. This is

what was o bserved in Great Britainfirst during the Industrial Revolu-
tion, and then later as air pollution regulations were enacted. Humans
caused the change in the environment, and bird predators were the feed-
back mechanism that maintained homeostasis. Death of individuals leads
to changes in populations. Homeostasis mechanisms can cause changes
in a biological system if that system experiences environmental changes.
Homeostasis would only be expected to keep the system constant if the
environment remained constant.
In another experiment that examined how death of individuals affects
populations, David Reznick and Matthew Walsh examined the indirect
evolutionary effects of predation in a fish. The direct effect of predation
is to kill the individual prey and act as a selective agent on the rest of the
population. Indirect effects of predation include an increase in food avail-
ability to survivors, and changes in behavior when prey detects preda-
tors in the area. Reznick and Walsh were interested in the evolutionary
changes caused when populations of the same species were either exposed
to predation or not, how exposure to predation led to changes in food
availability, and whether those changes led to changes in populations that
were maintained through homeostasis.
Reznick and Walsh studied fish in rivers on the island of Trinidad.
These rivers often have stretches separated by small waterfalls that prevent
dispersal upriver. Scientists have observed that a killifish, Rivulus hartii,
is able to disperse upriver to colonize habitats not accessible to other fish.
Populations of this killifish are often found in stretches in which they
are the only species present. They are also found in sites that contain
two species of predator fish. Reznick and Walsh predicted that killifish
populations in predation-free sites would exhibit phenotypic differences
from populations living in sites exposed to high predation and that these
differences are heritable. In addition, the scientists predicted that indirect
effects of predation, mediated through food availability, would be impor-
tant in the evolution of phenotypes.
The biologists collected killifish from predator-free (above waterfalls)
and high predation sites (both predators present) in two rivers. The biolo-
gists established four laboratory populations from two rivers (two were
predation-free and two were high predation sites) that were kept isolated
 Death of a Single Individual Affects a Population 7

genetically from each other. The researchers reared fish for two genera-
tions, and it was the second generation offspring that were used in the
experiment.
Reznick and Walsh grew the second generation offspring in groups
of eight fish per aquarium and fed them as much food as they could
eat. When the fish were 20 days old, individual fish were placed in their
own aquaria. Half of the second generation fish were randomly chosen to
receive a low level of food (liver paste and brine shrimp larvae) and the
other half were given a high level of food. The high level of food was based
on estimates of food available to fish in high predation sites, and the low
level of food was half of that. The biologists noted when fish matured and
measured their mass and age. As the fish approached the age at which
killifish matured, the scientists placed each individual into a tank with an
already mature individual of the opposite sex. If a viable egg was found
in the tank, the individual was deemed mature. If an individual was not
yet mature, the scientists repeated this procedure until it was found to
be mature. At that point, the individual was weighed and the number of
days since hatching was recorded.
Reznick and Walsh collected eggs from mature females for 2 weeks to
quantify the number of offspring produced, egg size, and size of hatch-
lings. The number of eggs produced per female was counted daily, eggs
were weighed, and ten eggs per female were allowed to hatch and these
individuals were weighed. The reproductive allotment was a measure that
Reznick and Walsh calculated as a measure of the allocation of individual
resources invested in reproduction. This is a measure of daily investment
in reproduction relative to body size, calculated as ([mean per day egg
production mean egg size]/mean size of female) 100.
The decline in the number of survivors due to predation could result
in greater availability of resources for survivors. These direct and indi-
rect effects could lead to evolutionary changes, and Reznick and Walsh
predicted that killifish from high predation sites would be younger and
smaller at maturity and produce more small offspring than fish from
predation-free populations. Reznick and Walsh showed that killifish from
populations exposed to high predation responded as predicted and that
these differences were heritable. Fish from high predation populations
also invested more of their energy to reproductive effort than fish in
8 POPULATION HOMEOSTASIS

predator-free sites. Descendants of predator-free fish never outperformed

descendants of high-predation fish by maturing earlier (in fact, they ma-
tured much later) or producing more eggs, at least within a food level.
A direct evolutionary response of a population to high mortality rates
would be earlier maturity and higher fecundity. If there is a high risk of
predation, individuals that matured earlier and produced more offspring
before they get eaten by a predator would have a selective advantage over
other individuals.
The scientists also showed that when more food was available fish
grew faster, matured earlier at a larger body size, and produced more
young, relative to individuals from the same population exposed to low
food. When killifish were reared under conditions simulating natural dif-
ferences in food availability caused by presence or absence of predators,
the biologists were able to determine how those two factors interacted
to affect adaptations. Fish from predation-free sites, which presumably
evolved to low levels of food available in the habitat, matured later than
fish from high predation sites. Killifish from high predation environments
were smaller and laid more eggs per day than fish without predators, espe-
cially at the high food level, and the researchers concluded that this is an
indirect evolutionary response to high mortality rates.
The differences in responses between populations from high preda-
tion sites and predator-free sites are less pronounced or absent when
food levels were low than when they were high. A possible explanation
for this is that fish from predator-free sites have adapted to low levels of
food availability and are not able to respond easily to high levels of food.
Killifish from high predation sites have evolved to exploit more efficiently
the consistently higher levels of food they routinely encounter due to the
indirect effects of predation.
The indirect effects of predation led to evolutionary changes. Each
population maintains homeostasis, but the parameters are maintained at
values dependent upon environmental conditions. This was also the case
for peppered moths. Organisms adapt to environmental conditions, such
as the amount of food available and the presence or absence of preda-
tion to best exploit their resources. When a predator is introduced into a
predator-free habitat, homeostasis of the prey population is likely to be
disrupted because the populations there have adapted to the predator-free
 Death of a Single Individual Affects a Population 9

environment. Similarly, reintroduction of predators, such as wolves, might

have more consequences to prey populations than originally predicted.
Reintroductions are initiated by humans, but humans also act as
predators, removing individuals from populations. In a dramatic example
of the effects of humans as predators, Chris Darimont and his colleagues
analyzed and compiled published studies of 40 populations of 29 species
where humans act as predators. Hunting, fishing, and plant harvesting
are examples of humans as predators where humans are taking individual
prey with particular phenotypes, such as the largest or the ones with the
largest antlers. The species studied included fish, hooved mammals, in-
vertebrates, and plants. The researchers sought to determine the extent
of phenotypic change caused by humans acting as predators compared to
other published studies where phenotypic changes were documented in
systems where humans were not a selective factor and in systems where
humans were acting as a selective factor but not as a predator (e.g., intro-
ductions of species into new habitats).
Darimont and his colleagues extracted 475 estimates of changes in
form, structure, or reproductive biology from research on the 29 species
harvested by humans (Table 1). They calculated the percentage change in
a variable as the mean variable value measured at the initial time point
minus the mean variable value measured at the end of the study, divided
by the mean variable value measured at the initial time point ([x1 x2]/x1).
The scientists estimated the rate of phenotypic change for each
variable, along with rates of change for similar variables from other
human-influenced populations and populations not influenced by humans
(Figure 3). The rate of phenotypic change is calculated as the difference
between the natural logarithms of the trait means at the two time points
divided by time ([ln(x1) ln(x2)]/t). The scientists determined the average
mean and the maximum absolute value of the numerator of the phenotypic

Table 1 Cases of phenotypic change in form, structure, and

reproductive traits caused by human predation.
number of cases of total number total percent average percent
phenotypic change of cases of cases change in variable
form/structure 282 297 94.9 18.3 13.7
reproductive 173 178 97.2 24.9 22.3

Source: From Darimont et al., 2009, text.

10 POPULATION HOMEOSTASIS

0.6
a
= natural

phenotypic change ( 1 SE)

0.5
mean or maximum = other human
= human predator
0.4
ab
0.3 a

b b
0.2

0.1 c

0.0
mean change maximum change

Figure 3 Average mean and maximum phenotypic changes affected

by three categories of selection: humans as predators, other human
interference, and natural environmental changes. Within a change
category, bars with similar letters above them are not significantly
different from each other, whereas bars with different letters are
significantly different from each other.
Source: Slopes from Darimont et al., 2009, Figure 1.

rate change against the length of time between the two measurements.
They used absolute value because they were interested in the magnitude of
change, not whether a change was positive or negative. The mean change
represents what might typically be observed in a population exposed to the
particular selective factor, and the maximum estimated change in Figure 3
represents what could potentially occur in such a situation.
Humans have the capacity to harvest large proportions of individual
prey, and they target individuals of very specific ages, sizes, and types.
This capacity is greater than natural predators, and Darimont and his
colleagues showed that this can lead to rapid phenotypic changes in both
morphological and reproductive traits in prey populations. Humans as
predators caused a consistently higher phenotypic rate change, whether
measured as the mean or the maximum for a population, than either
natural populations not being exploited by humans or in populations af-
fected in some other way by humans. Statistical analysis revealed that the
mean phenotypic change was higher for humans as predators than for the
other two categories, and the maximum phenotypic change was higher
for humans as predators than for natural populations.
 Death of a Single Individual Affects a Population 11

The high magnitude of phenotypic change in human-exploited prey

populations indicate that evolution can proceed very rapidly especially in
the face of dramatic environmental changes, such as fishing. Large and
rapid phenotypic changes to highly exploited commercially harvested
populations, such as fish, could seriously disrupt a populations ability
to maintain homeostasis and persist. As with killifish, exposure to preda-
tion leads to evolutionary shifts in reproductive investment and time to
maturity. The same changes occur in populations when human predators
select large, mature individuals. Rapid shifts to early maturity at smaller
sizes can reduce overall fecundity and lead to declines in the total amount
of biomass available to humans for harvest. This could rapidly lead to
overfishing and possibly extinction of populations or even entire species.
This chapter has explored how predators affect the evolution of popu-
lations and how prey populations adapt to the presence or absence of
predators in the context of their environment. Populations utilize nega-
tive feedback mechanisms, such as natural selection, to regulate and
maintain optimal conditions in a time-dependent process. Individuals
that escape predation are presumably better fit for the environment, but if
the environment constantly changes, populations are constantly evolving
to catch up to changes. The themes of homeostasis have all been evident
as the answer to the question of how the death of one individual affects
an entire population has been explored. In the next chapter, more about
how the acquisition of energy by one individual affects an entire popula-
tion will be explored.

Ethical, Legal, Social Implications: Saving Individuals

May Not Help to Save an Entire Population
Many people have experienced or witnessed instances of individual ani-
mals in jeopardy and they have probably wished that they could or even
have actually tried to save them from harm. At the beginning of the first
part of this chapter, readers considered how they might feel if they were
witnessing a predator-prey interaction. Perhaps some might have wished
to save the zebra from the lions clutches. If animals have the same rights
as humans (as some humans argue), then each individual should be af-
forded the same protections and considerations as individual humans.
12 POPULATION HOMEOSTASIS

To these people, animals are non-human persons, to be treated kindly

and with respect, provided rights, and cared for if sick, wounded, or in
danger of being killed. Now that argument might not extend to saving a
zebra from a lion, because the lion has a right to eat and no one can fault
or deny an animal its right to exist simply because it is a predator. But in
situations where humans, human actions, or human structures are caus-
ative agents in the endangerment of an individual, then those that argue
this point of view will intervene to save the life if possible.
Through destruction of habitat, overharvesting, pollution, and intro-
duction of exotic species, humans have caused the endangerment not just
of individuals but entire species. Conservation biology strives to preserve
species in danger of extinction and is especially concerned with species
endangered through human actions. Following the previous argument,
one would conclude that a strategy to protect species is to attempt to
save every individual. If humanity can save the life of every individual
belonging to an endangered species, then that will surely save the spe-
cies. If the life of an animal can be saved, then it will be saved by those
conservation-minded humans. But does such an act actually help the
species or the goals of conservation biology?
Natural selection would say no. An animal that is disoriented and
wandering around the streets of a city got there perhaps because it was
confused by street lights or noise, infected by a parasite, or had some
genetic abnormality or mutation. Confusion, susceptibility to infection,
and genetic mutations are factors upon which natural selection may act
to eliminate this individual under normal circumstances. If it is saved and
it goes back to the population and mates with another individual, those
genes remain in the population. This might ultimately harm the entire
population. Selection against particular phenotypes leads to evolution of
populations in response to the selective factor. For this reason, it might
be prudent to not save the individual, especially if one does not know the
cause of its being in the situation in which it finds itself.
Additionally, some would argue that the loss of one individual in
such an instance will not affect the populationindividuals die all the
time, and it does not have a large detrimental effect on population size
to lose one individual. As will be discussed later in this book, one aspect
 Death of a Single Individual Affects a Population 13

of population homeostasis involves changes in birth and death rates in

response to population density. Fewer individuals in a population, all else
being equal, may have higher per capita birth rate. Homeostatic mecha-
nisms will regulate the population so that the population will rebound if
overall habitat conditions improve. It is the conservation of habitat that is
more important than saving one individual.
Decisions then may come down to financial resources. Whatever hu-
mans spend saving individual animals could be spent preserving habitat,
cleaning up polluted soils and waters, and controlling invasive species.
Many species, endangered or not, could live and thrive in cleaner environ-
ments, and this strategy protects threatened and non-threatened species
alike. The goal of conservation should be on saving populations, not indi-
viduals. Although it is true that individuals make up populations, the pres-
ervation of one individual may not help an endangered population much
and is not a wise use of limited financial resources. One benefit of saving
individuals that might help both strategies is the publicity and subsequent
increased awareness of the plight of endangered species in the public.

Bibliography
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outpace other agents of trait change in the wild, Proc Nat Acad Sci
106(3):952954, 2009.
Gibbons W: Ecoviews: saving species outweighs saving two whales,
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article/20070527/NEWS/705270427. Accessed June 14, 2010.
Gunnarsson B: Bird predation as a sex- and size-selective agent of the
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and its melanic forms in Great Britain, Heredity 12:5172, 1958.
14 POPULATION HOMEOSTASIS

Reznick DN, Ghalambor CK, Crooks K: Experimental studies of evo-

lution in guppies: a model for understanding the evolutionary con-
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 Index
American kestrel, 3233
Aviary, 4
Behavioral mechanism, in prey, 18
Biomagnification, 35
Biston betularia, 1
Boreal forest, 23
Camouflage, 2
Carbonaria, 1, 2
Clumped dispersion, 25
Conservation biology, 12
Damselfish, 18
mortality rates in, 1516
underwater study of, 17
Darimont, Chris, 910
Dascyllus flavicaudus, 15
Density-independent, 25
Dichloro-diphenyl-dichloroethylene
(DDE), 3132, 33
eggshell thinning, 3334
Dichloro-diphenyl-trichloroethane
(DDT)
biomagnification of, 3135
Diplotaxis erucoides, 26
D. trimaculatus, 15
Energy budget, 1
Environmental change, xiii
Environmental factors, 15
European rabbits
annual changes in densities of, 20
birth rates, 19
number of litters per female, 21
reproductive season, beginning
of,19
Falco peregrines, 32
Feedback mechanism, 2, 17, 23, 26
birth rates, affecting, 19
F. sparverius, 32
Haliaeetus leucocephalus, 32
Holbrook, Sally, 1518
Homeostasis, 1, 6, 13, 15
in populations, xiii
Insularia, 2, 5
Jack pine, 23
clark-evans (CE) statistics for, 24
mapping method of living and dead
trees in, 24
Kenkel, Norm, 2326
Kettlewell, Bernard, 15
Kunin, bill, 2628
Lichens, 2
Lincer, Jeffrey, 3234
Mark-release-recapture
experiment, 3
Modified Clark-Evans statistic, 24
Mortality rate, 8
Moths
conspicuousness of, 2, 3
inconspicuousness of, 2
Natural selection, xiii
Nearest neighbor distance
(NND), 24
white wallrocket, function
of,2627
Negative feedback mechanisms, 11
environmental changes, 28
Oryctolagus cuniculus, 19
Parts per million (ppm), 33
Phenotypic change, rate of, 910
Pheromone, 3
Pinus banksiana, 23
Pollutant, 31
42 INDEX
Populations
death of individual, affects, 113
of european rabbits, 19, 20
female density, 2022
regulate, 15
regulated through feedback
mechanisms, 1529
Predation, 1, 16, 18
by birds, 2
direct effects of, 6
indirect effects of, 6, 8
Predator, 1
humans as, 10
reintroduction of, 9
Prey density, 17
Random dispersion, 24
Random distribution, of plants, 25
Ratcliffes Index (RI), 32
Regulate, population, 15
Reznick, David, 67
Rivulus hartii, 6
Rdel, Heiko, 1922
Saving individuals, 1113
Schmitt, Russell, 1518
Social hierarchy, 22
Typica, 1, 2, 5
Uniform dispersion, 25
Walsh, Matthew, 67
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