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This book will synthesize the concepts of selection against indi-
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Homeostasis
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populations of top predators, such as raptors, have suffered due
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mental studies at Davidson College. He teaches introductory
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A. Malcolm Campbell
Population Homeostasis
Population Homeostasis

Christopher J. Paradise, PhD


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

All rights reserved. No part of this publication may be reproduced, stored


in a retrieval system, or transmitted in any form or by any means
electronic, mechanical, photocopy, recording, or any other except for
brief quotations, not to exceed 250 words, without the prior permission
of the publisher.

First published in 2016 by


Momentum Press, LLC
222 East 46th Street, New York, NY 10017
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ISBN-13: 978-1-60650-975-3 (print)


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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
Darimont CT, Carlson SM, Kinnison MT, et al.: Human predators
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,
Tuscaloosa News.com (website): http://www.tuscaloosanews.com/
article/20070527/NEWS/705270427. Accessed June 14, 2010.
Gunnarsson B: Bird predation as a sex- and size-selective agent of the
arboreal spider Pityohyphantes phrygianus, Funct Ecol 12(3):453458,
1998.
Kettlewell HBD: Selection experiments on industrial melanism in the
Lepidoptera, Heredity 9:323342, 1955.
Kettlewell HBD: Further selection experiments on industrial melanism
in the Lepidoptera, Heredity 10:287301, 1956.
Kettlewell HBD: A survey of the frequencies of Biston betularia (L.) (lep.)
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-
sequences of predator removal in natural communities, Mol Ecol
17:97107, 2008.
Walsh MR, Reznick DN: Interactions between the direct and indirect
effects of predators determine life history evolution in a killifish, Proc
Nat Acad Sci 105(2):594599, 2008.
CHAPTER 2

Populations Are Regulated


Through Feedback
Mechanisms

Homeostasis implies balance and constancy, but when applied to popu-


lations, there is no such thing as a constantly-sized population. Popula-
tions are dynamic and grow and decline over time. For instance, death
of an individual may lead to an evolutionary response in the population,
and individuals foraging for limited resource allocate resources in such a
way as to attempt to maintain reproduction. Environmental factors are
the mechanisms that affect death and reproduction and, consequently,
population size. In neither case did the populations examined maintain
constant size or phenotypes.
Individuals can be limited by resources and that can potentially
regulate the growth of populations. To regulate a population is to control
its size in a manner that tends to achieve or return it to a size at equilib-
rium with the environment. As populations change size away from a size
optimal for a particular habitat, becoming either too large or too small,
feedback mechanisms may commence, causing the population to reverse
the growth or decline. To determine how populations are regulated Sally
Holbrook and Russell Schmitt (2002) examined juvenile mortality in two
tropical damselfishes, the yellowtail damselfish (Dascyllus flavicaudus) and
the three-spot damselfish (D. trimaculatus).
They conducted field observations and experiments to determine
whether mortality due to predation was dependent upon density. The
scientists also hoped to understand the mechanisms that caused the per
capita mortality rate to scale with density. The scientists performed their
16 POPULATION HOMEOSTASIS

experiments in lagoons in French Polynesia. The damselfish feed during


the day on plankton above habitats in which they shelter when threatened
and to rest at night. These fish spend the first few weeks of life swimming
in the open water, after which yellowtail settle on coral and the three-spot
settles on sea anemone, which are animals related to jellyfish and corals.
Once the fish settle, they do not usually relocate.
In their first experiment, the scientists tested the hypothesis that pred-
ators caused mortality of newly settled yellowtail and that mortality rates
depended upon yellowtail density. The researchers collected 24 corals of
about 20 30 centimeters and glued them to cinder blocks. They placed
the blocks in a grid of 6 4 in a lagoon. Predator exclusion cages, made
of nylon mesh with 5 millimeter openings were placed over half of the
corals. Half of the predator-free and half of the predator-exposed corals
were randomly assigned to receive either three or six newly settled yellow-
tails. The researchers dove to the grid twice each day for 3 days, counted
the number of yellowtail on each coral, and replaced missing fish so that
densities remained constant. The mean proportion of fish lost over the
3-day period was calculated for each combination of yellowtail density
and predator treatment.
A second experiment was conducted similar to the first, but this time
predator exclusion cages had a larger mesh size, which allowed smaller
predators to access the corals and hunt yellowtail but excluded large pred-
ators. They changed yellowtail densities so that corals in the grid con-
tained 4, 8, or 16 yellowtail. In this experiment, all corals were exposed to
predation, half were exposed to small predators, and half were exposed to
both large and small predators. Divers counted fish twice daily for 4 days,
but missing fish were not replaced.
The results showed that predation is a significant source of damselfish
mortality. More importantly, for regulating populations, the data showed
that mortality increased with increasing prey density. Other sources of
loss from a population (such as, emigration, starvation, and disease) were
ruled out because of the video evidence, but those are factors that could
very well regulate other species. The caging experiments of Holbrook and
Schmitt further revealed that small-bodied predators were responsible for
most damselfish mortality. At the highest damselfish density tested, there
was an additional component of mortality caused by large predators, but
Populations Are Regulated Through Feedback Mechanisms 17

recordings of attacks supported the notion that small predators made


most of the attacks.
Predation rate that varies with prey density is an example of a feed-
back mechanism that acts on demographic rates in populations. In situ-
ations of high prey density, predators consume a higher proportion of
individuals than individuals in low density populations. Similarly, if
densities drop too low, other feedback mechanisms might act to raise
birth or immigration rates or reduce sources of mortality. The end result
is that populations fluctuate constantly but are bounded by highs and
lows. This homeostasis at the population level contributes to persistence
of populations.
Holbrook and Schmitt next established a set of 24 uncaged anemo-
nes and transplanted three-spot damselfish to each anemone, with eight
anemones each receiving 5, 10, or 20 three-spots. The researchers set
up a similar experiment where yellowtails were transplanted to a grid
of 24uncaged corals. Eight corals in each treatment contained 3, 6, or
12 yellowtails. Divers counted fish on each anemone twice per day for
3days: just after dawn and just before dusk. The researchers determined
the number of fish lost at each census and replaced missing fish to main-
tain densities. Holbrook and Schmitt determined the proportion lost
during the night or day across the three time periods and found that for
both species, most of mortality was occurring at night, and the propor-
tion lost at night increased with increasing damselfish density.
Holbrook and Schmitt used a remote, underwater infrared video
system to record events continuously and observe fish without altering
their behavior. The cameras were mounted on a pole 1 meter above a
coral or anemone. Infrared lights provided illumination. The researchers
examined recordings of predator attacks to determine success of attacks
in relation to a damselfishs location. The scientists classified the position
of a damselfish as being on the edge, in the center, at the base, or in the
water column above their home coral or anemone. Most of the success-
ful attacks occurred at the edge of a shelter or in the water column above
it. In addition, the scientists set up experiments where damselfish were
transplanted at one of three densities to a coral or anemone and the num-
ber of individuals located in each position at regular time intervals was
determinedfrom which they determined the proportion of individuals
18 POPULATION HOMEOSTASIS

in vulnerable positions at each density. The proportion of individuals that


were in the vulnerable positions where successful predator attacks often
occur rose steadily as damselfish density increased.
As with homeostasis or any other biological phenomenon, scientists
are interested in the mechanisms that underlie the process. Although
Holbrook and Schmitt could have simply concluded that predation was
one of the feedback mechanisms that regulated populations of damselfish,
they went further to discover how prey behaviors interact with predation
to cause the variation observed at different densities. Predators may search
many corals or anemones each night looking for vulnerable prey, but in
order for negative feedback to work, there has to be some mechanism that
causes higher exposure of prey to the predator in high density popula-
tions. The researchers discovered a behavioral mechanism within the prey
that caused a higher proportion of damselfish to inhabit vulnerable loca-
tions when population densities were high.
Damselfish swim above their refuges to forage during the day. When
they detect a threat, they retreat to shelter. Shortly before sunset, dam-
selfish move to their shelters where they rest until morning. Predators
typically are less efficient at detecting and capturing prey in refuges. The
biologists found that mortality due to predation occurred primarily at
night. This might be counter to what one might have predicted if one
thought that fish are more vulnerable and more exposed while they are
feeding during the day.
In addition to time of day, the fish are more vulnerable in certain
locations within their refuge. Damselfish located at the edge or just off
the perimeter of their refuge were at greater risk than those in the center
or at the base. Damselfish are not foraging at night, so it is unlikely to
explain why damselfish leave their shelter. The data showed that higher
proportions of fish are in vulnerable positions as density increases, and it
turns out that fish are interacting with each other in an aggressive manner
to push one another into positions of vulnerability. Stronger or more ag-
gressive damselfish win these aggressive interactions and take positions of
least vulnerability. Weaker individuals are displaced into riskier locations.
Individuals compete for refuges within the corals and anemones, and this
causes per capita rates of aggression and the proportion displaced to risky
positions to increase with density. Similar studies in both freshwater and
Populations Are Regulated Through Feedback Mechanisms 19

terrestrial habitats suggest that the mechanisms described here are preva-
lent and can be important in maintaining homeostasis of populations.
In addition to feedback mechanisms that act to affect mortality rates
(either increasing when densities are high, or decreasing when densities
are low), feedback mechanisms may affect birth rates. Most people have
no doubt heard the expression breeding like rabbits, suggesting that rab-
bits have high birth rates and produce many offspring. But do they always
produce many offspring, or are there conditions that cause variation in
birth and death rates to maintain homeostasis? Heiko Rdel and his col-
leagues studied a population of European rabbits (Oryctolagus cuniculus)
in an enclosed area for 11 years (Rdel et al., 2004). As with Holbrook
and Schmitt, the scientists were interested in the feedback mechanisms
that maintained population homeostasis.
The annual rabbit population in the enclosure ranged in density from
18 to 44 rabbits per hectare and they lived in burrows and artificial con-
crete warrens, which had interconnecting chambers and removable tops.
Although rabbits could not exit the enclosure, their predators (such as,
martins, hawks, and cats) could enter. Rabbits have a discrete breeding
season. Individuals are born and then reach sexual maturity at the next
reproductive season. European rabbits live in social groups with separate
social hierarchies among the males and females. Each social group occu-
pies a distinct territory.
For any particular year, the researchers analyzed all individuals that
survived to the end of that reproductive season, but they used population
density at the beginning of the season as their independent variable. The
beginning of a reproductive season was defined as the date that the first
litter was found minus 30 days of gestation, usually from mid-February
to late March. During each reproductive season females can produce mul-
tiple litters, so the researchers checked all warrens and burrows (which
had covers installed on them so the researchers could peek inside) each
morning to monitor breeding. The date and litter size, which ranges from
2 to 12 offspring, of each litter discovered was recorded. Each adult had
an ear-tag so that individuals could be identified. From regular observa-
tions made during each reproductive season of each social group, the re-
searchers knew to which group each female belonged and her social rank,
which could change from year to year. Social rank was determined by
20 POPULATION HOMEOSTASIS

repeated observations of interactions that revealed which rabbit in a pair


was dominant and which subordinate.
The researchers monitored the density of females each year and a ssessed
the relationship between the population density of females and the pro-
portion of 1-year-old females (Figure 4). The European rabbit population
fluctuated around an average with no long-term trend of either increasing
for more than 2 years in a row or decreasing to extinction. A growing
population grows because of higher reproduction, and the consequence
of this is that there should be a high proportion of 1-year olds when
population densities are high. The researchers found that the proportion
of 1-year-old females was positively correlated with the density of the
female population. Short-term population increases were caused by high
reproductive rates, leading to a high proportion of 1-year-old, first-season
breeders.
Rdel and his colleagues predicted that they would observe varia-
tion in reproduction that was dependent upon population density. They
examined the relationships between population density at the start of a
reproductive season and the number of offspring produced per year as

30
= density 1 yo females
25 = density older females
density (number/hectare)

20

15

10

0
92

93

94

95

96

97

98

99

00

01

02
19

19

19

19

19

19

19

19

20

20

20

year

Figure 4 Annual changes in densities of 1-year-old and older female


European rabbits.
Source: From Rdel et al., 2004, Figure 2.
Populations Are Regulated Through Feedback Mechanisms 21

well as the number of litters per female. Their annual sample sizes for
females producing litters ranged from 16 to 43 per year. The number
of offspring produced per year declined as female density increased, at
a rate of 0.41 per increase in 1 female per hectare. This means that as
female density increases by 1 hare per hectare, the number of offspring
per female declined by almost one-half a hare. This decline was statisti-
cally significant (P = 0.006). The number of litters produced per year
also declined as female density increased, at a rate of 0.07 per increase
in 1 female per hectare. This means that as female density increases by
10 hares per hectare, the number of litters per female declined by about
0.7. This decline was also statistically significant (P = 0.007).
Litter size can vary with the multiple breeding cycles within a repro-
ductive season, so Rdel and his colleagues examined density effects in
the first, second, and third breeding cycle separately, relating the mean
litter size of each cycle with the density of females at the beginning of that
cycle. Litter size did not vary with female density, suggesting that it was
the number of litters, not the size of individual litters in particular breed-
ing cycles that responded to changes in female density.
Rdel and his colleagues used female density as their measure of
density, because females compete for high quality burrows or warrens.
Competition for these sites should be more intense when the density of
females is higher. The scientists documented a strong inverse relationship
between the mean number of offspring produced in a year and the den-
sity of breeding females in that year. The annual number of offspring per
female decreased by about 65% when female density doubled. This was a
result of changes in the number of litters and total reproduction within a
reproductive season, not variation in actual litter size within a reproduc-
tive event. The number of litters produced per female declines not only
because of lowered reproductive output but also because there are fewer
females present later in a reproductive season, indicating presence of a
mortality effect.
Changes in the density of breeding adults that occur from one repro-
ductive season to the next are due to both adult deaths and juveniles that
enter the adult population as 1 year olds. Rdel and his colleagues ex-
amined differences in reproductive performance for 1-year-old and older
females. They compared the social rank of individuals in different age
22 POPULATION HOMEOSTASIS

classes to determine where 1 year olds were in the social hierarchy and
found that the mean social rank of 1-year-old females was lower by an
average of 1.7 rank positions than that of older females. The scientists also
tested for density-related changes in fecundity by comparing year-to-year
changes in fecundity to year-to-year changes in population density. Here
they tested for the effects of changes in age and changes in population
density from one reproductive season to the next on the changes in off-
spring produced from one year to the next.
The proportion of 1 year olds in a year, which is positively related
to total population density, is also related to the effect of density on re-
production. Rdel and his colleagues predicted that 1-year-old females
would have lower reproductive output than older females, and in fact,
1year olds produced fewer offspring per capita than older females. Age,
or time-dependence of reproduction, where young individuals have lower
reproductive performance is common in birds and small mammals.
One-year-old rabbits produce fewer litters their first year because of their
inexperience, low social rank, and small size, because they have not yet
achieved maximum size. A low social rank could cause more stress, which
could affect hormone levels and be expressed as lower reproductive output.
It also makes them inferior competitors for resources, such as warren sites.
As social rank and age increased so did reproductive output of females.
Despite this age effect, when the density of females increased between
the first and second breeding season, females showed a decrease in the
per capita number of offspring produced per season, suggesting that the
density effects, although related to age effects, were stronger than age.
Density dependent reproduction could be common in populations in-
habiting environments where fluctuations in density are common, and
it could operate through the mechanism of uneven reproductive perfor-
mance of newly matured individuals. When densities are high, reproduc-
tion may be more costly, and a lowered reproductive output might be a
trade-off between fecundity and survival. If an individual female produces
many litters in a reproductive season when densities are high, her survival
and survival of her offspring might be in jeopardy. The high density of the
population reduces availability of resources, which increases competition
and reduces survival. During a reproductive season when the population
density is high, a female might reproduce during the first breeding cycle
Populations Are Regulated Through Feedback Mechanisms 23

only and put her energy and resources into caring for the first batch of off-
spring while storing energy for the next reproductive season. This might
be especially true for young females that do not have experience rearing
offspring and do not have the high social standing to compete well for
resources.
The density of animal populations, then, can initiate feedback mecha-
nisms that affect both death and birth rates. Other feedback mechanisms
(such as, dispersal or growth) may also respond to changes in population
density, and these may be important in other organisms, such as plants,
fungi, and bacteria. The population growth of bacteria, for instance, is
also regulated and exhibits a characteristic growth curve, at least under
controlled conditions. Reproduction, recruitment of seedlings into the
population, and mortality can be readily studied in long-lived plants,
such as trees. A population of organisms that cannot relocate, such as
plants, may be very sensitive to density and possess a variety of feedback
mechanisms to regulate populations and maintain homeostasis.
Norm Kenkel (1988) studied a population of jack pine (Pinus banksi-
ana) in a Canadian boreal forest. A boreal forest is a subarctic, evergreen
coniferous forest dominated by firs and spruces. Jack pine is a tree that
grows to 20 meters tall and 30 centimeters in diameter. This tree grows
rapidly during the first 50 years of life and then slows to almost no growth
after about 80 years. Individuals produce a large number of seeds in cones
that open after being exposed to the heat of fire, which is an adaptation
to living in fire-prone areas. Thus, large numbers of seeds are released and
germinate all at the same time, resulting in stands of trees that are almost
all the same age. Seedlings may grow next to other seedlings or next to
large mature trees that survived the fire. Densities of germinating seeds
after a fire are often very high, but an individual plant may be influenced
not so much by overall population density but rather by the proximity,
size, activity, and density of its immediate neighbors.
Within one population of jack pine, where other tree species did not
live, Kenkel randomly selected a 50 meter 50 meter area. As part of
a forest reserve, this population was protected from human disturbance.
The jack pines in this population averaged 15 centimeters in diameter
and 16 meters in height, and the stand was estimated to have been es-
tablished after a fire 50 years earlier. Kenkel mapped the position of each
24 POPULATION HOMEOSTASIS

individual living and standing dead jack pine and jack pine stumps. This
provided a good indication of the initial population with the exception of
seedlings and saplings that died in the early years of stand establishment.
Other research suggested that mortality of seedlings is not dependent
upon density, so he considered that his mapping method of living and
dead older trees would allow him to investigate mortality that did depend
upon density.
Kenkel used his map of the living and dead trees to determine the x,y
coordinates of each trees location relative to the southeastern corner of the
study area. Kenkel mapped 1,375 individuals; 459 living trees and 916
dead standing trees and stumps. The biologist analyzed the spatial patterns
of jack pine within the study area, analyzing living and dead trees together
(including stumps), living trees alone, and dead trees alone, using several
different statistical methods. For each of the three populations of trees
(living and dead, living alone, or dead alone), Kenkel determined the den-
sity of trees and nearest neighbor distance (NND) for each individual
in the population and computed the mean NND (Table 2). NND is the
distance, found using the coordinates in the grid, from one living tree to
the next nearest living tree in the living tree population. For the living plus
dead population Kenkel determined the distance from any one tree, living
or dead, to the next nearest tree, living or dead. In this way, the biologist
generated three densities and three mean NNDs.
Kenkel then calculated a value called the modified Clark-Evans statistic
(Table 2). For this method, Kenkel calculated the distance that would be
expected for the density of a population if the individuals had a random
dispersion. Random dispersion occurs when the probability of finding
an individual at any point in the area is the same for all points. If a popu-
lation of individuals is randomly distributed within an area, each will
have a random NND. When dispersion is clumped, the mean NND is

Table 2 Descriptive statistics and Clark-Evans (CE) statistics for a


jack pine forest.
population density (#/m2) mean NND (m) CE probability
all trees: living, dead, & stumps 0.55 0.554 1.567 0.117
living trees 0.18 1.107 5.56 <0.0001
dead trees 0.37 0.633 1.898 0.942

Source: From Kenkel, 1988, in text.


Populations Are Regulated Through Feedback Mechanisms 25

much smaller than the expected distance. Clumped dispersion occurs


when individuals are closer than expected by chance. Therefore, large
negative values of CE suggest clumped distributions. Large positive values
suggest regularly spaced, or uniform, distributions, and are caused by the
average NND being greater than the expected distance. When the disper-
sion of individuals is uniform, the mean NND is greater than expected
under the assumption of randomness, and the ratio is greater than one.
Uniform dispersion occurs when there is a regularity of distance between
individuals.
The CE statistic value and the probability of obtaining that value or
greater are in Table 2. The probability that CE is greater than a large
negative value is high, and if it was 0.975 or more, Kenkel concluded the
distribution was clumped. The probability that CE is larger than a large
positive value is low, and if it was 0.025 or less, he concluded a regularly
spaced distribution. All probabilities in between were judged to be from
a random pattern.
After a fire, thousands of jack pine seeds will be released and germi-
nate, leading to very high densities of seedlings and saplings. Any distur-
bance that occurs (such as, another fire, a drought, or even the presence of
herbivores) could cause mortality of seedlings and saplings irrespective
of density. Mortality of young individual trees could occur from a variety
of factors that are density-independent. A density-independent factor af-
fects the size of a population regardless of the population density. Older
trees have a much higher probability of survival, but mortality could still
occur in the population. Kenkel hypothesized that he could discern popu-
lation regulation by examining the distribution of living and dead trees in a
forest. A uniform pattern would be strong evidence for density-dependent
competition for resources, such as space, sunlight, and soil nutrients in
plants. Tracking the fate of individuals over time provided evidence for the
importance of population regulation feedback mechanisms.
Kenkel used the hypothesis of a random distribution of plants as his
null hypothesis, the hypothesis of no effect. A random distribution of
the initial population is likely, because the density-independent factors
that caused mortality in seedlings and saplings likely killed in a random
pattern. As trees grow larger, they need more space. If two trees grew close
together when young, one of them would have to die when it was older so
26 POPULATION HOMEOSTASIS

that the other would have space to survive. This would cause non-random
mortality of mature trees to yield a more uniform distribution of trees
that are all about the same age and size. This supposes that competition is
the underlying mechanism that regulates population density.
The hypothesis of randomness supposes no such mechanism and is an
excellent null hypothesis. Scientists want to be sure they are not falsely
rejecting a null hypothesis, so they err on the side of conservativeness and
will only reject a null hypothesis if the data reveal a very strong reason for
doing so. Just by chance, a study could lead to rejection of a null hypoth-
esis, even when the null hypothesis is true. To reduce that probability, sci-
entists reduce the range of probabilities for rejection of null hypotheses.
In this case, 2.5% at the low end and 2.5% at the high end of the range
were where the probabilities were for null hypothesis rejection.
Kenkel concluded from his analysis that the initial population was
randomly distributed, in accordance with his prediction that prior to the
onset of population regulation mechanisms distribution would be ran-
dom. Two-thirds of the mature trees died after reaching maturity. Kenkel
could not reject the random distribution for the trees that died, but he
concluded that there was a trend toward a clumped distribution. Although
the distribution of dead trees is interesting, it is the distribution of living
trees that matters for the population, and Kenkel found that the survivors
were distributed uniformly. The density was lower, and the mean NND
was more than twice that of the initial population (living and dead trees
combined). This suggests that there was competition for resources, such
as light and soil nutrients. Trees that were too close together competed,
some died, and only ones a certain distance from others survived.
In a couple of examples in this chapter feedback mechanisms regulat-
ing animals and plants have been shown to lead to increased mortality
when population densities are high. Feedback mechanisms are involved
when reproduction in animals is sensitive to the density of a population.
Other organisms such as plants may also have feedback mechanisms
that regulate the density of populations through effects on reproduction.
Bill Kunin (1992) studied reproduction in populations of a plant called
white wallrocket (Diplotaxis erucoides) as a function of NND. This annual
plant grows to about 0.5 meters and is found throughout much of the
coastal area surrounding Mediterranean Sea. It often forms large dense
Populations Are Regulated Through Feedback Mechanisms 27

patches but can also live in scattered, sparse populations. These plants are
pollinated by insects. Once pollinated, flowers develop into papery seed
capsules with up to 60 small seeds. If not pollinated, flowers form small,
empty seed capsules.
Kunin studied two populations, both of which contained several
small clumps of closely spaced plants surrounded by areas with relatively
widely dispersed white wallrocket individuals. The scientist arbitrarily se-
lected 20 plants in each population, making sure that he selected plants
that had NNDs of more than 2 meters, as well as individuals that were
much closer together.
Kunin recorded the number of inflorescences on each selected plant,
which he used as an index of plant size, and the distance to the nearest
white wallrocket neighbor. He found no relationship between plant size
and NND. Kunin randomly chose up to six inflorescences from each
plant, and for each he counted the number of full and empty seed cap-
sules and flower buds that did not develop into flowers. The biologist
compared the proportion of flowers that set seed to the NND. For this
variable, he found a strong negative correlation between proportion of
flowers that set seed and NND.
He collected all of the seed capsules that contained seeds from these
inflorescences, five of which he chose randomly to open up and count the
number of seeds they contained, he and compared the number of seeds
per fruit to NND, and found that this variable was also strongly nega-
tively correlated with NND. Finally, Kunin estimated the total seeds pro-
duced by each plant and compared that to the NND, and again, found
that this variable was also strongly negatively correlated with NND.
When jack pine trees grow too closely together, one or more of them
dies, leading to greater NNDs. Competition for resources such as space
may cause mortality, but it may also cause individuals to grow more
slowly. If such were the case, Kunin would have observed a positive re-
lationship between plant size, as measured by number of inflorescences,
and NND. He did not observe such a relationship.
The proportion of capsules that set seed and the number of seeds per
capsule was both found to be inversely correlated with the NND. Plants
that had closer neighbors typically had greater reproduction, and some
plants that were very isolated came close to failing to reproduce at all.
28 POPULATION HOMEOSTASIS

This is opposite to what was observed as a density effect in rabbits, in


that rabbits that were in low density populations tended to have higher
birth rates. Taking both effects together with number of flowers per in-
florescence and number of inflorescences per plant, Kunin estimated the
total seed production of plants and found a strong inverse relationship to
NND. Total reproduction per plant dropped dramatically as the distances
between plants rose. This suggested to Kunin that reproductive success in
white wallrocket is dependent on density, although this is o pposite to what
might be expected. Kunin argued that plants in high density clumps were
visited by more pollinators and had a higher degree of cross-pollination,
which is essential for this plant. Plants in low density populations may be
limited by access to pollinators and other plants pollen in such a way that
their reproduction suffers.
If population density negatively affects reproduction in low density
plant populations, then density may also affect spatial patterns of popula-
tions. This may manifest itself as an emergent property leading to a meta-
population where many subpopulations are clumped and reside in locally
high density populations. Patchiness of populations may be enhanced.
Plants that live in clumps or high density populations may have a selective
advantage leading to the evolution of short-range dispersal. Very short
dispersal may lead to very high densities and intense competition result-
ing in mortality, smaller size, and lower reproduction. Selection against
dispersal distances that are too far and too short would lead to stabilizing
selection and dispersal distances that are intermediate but close enough
for clumping to occur.
Density-dependent effects are varied and that density-dependent reg-
ulation is widespread. Population homeostasis is not exhibited as constant
population density. Populations and communities are dynamic systems,
and changes occur over time. Negative feedback is necessary for the regu-
lation of a dynamic variable, whether it be a population or a community.
There is a long-term average population density, which is generally regu-
lated between two extremes. Environmental changes, whether episodic or
persistent, can alter the negative feedback process so that the long term
average density changes in a gradual or abrupt manner. If a new preda-
tor were introduced, for instance, the new optimal population density
for a prey population might decrease. A variety of feedback mechanisms
Populations Are Regulated Through Feedback Mechanisms 29

act on populations to maintain homeostasis relative to the environment


and changes in the environment. Environmental conditions can upset
homeostasis of the population to the extent that a population can grow
exponentially or decline to extinction. In the next chapter, such a change
in conditions that upset homeostasis to such a degree as to threaten
extinction of species will be explored.

Bibliography
Holbrook SJ, Schmitt RJ: Competition for shelter space causes
density-dependent predation mortality in damselfishes, Ecology 83(10):
28552868, 2002.
Kenkel NC: Pattern of self-thinning in jack pine: testing the random
mortality hypothesis, Ecology 69(4):10171024, 1988.
Kunin WE: Density and reproductive success in wild populations of
Diplotaxis erucoides (Brassicaceae), Oecologia 91(1):129133, 1992.
Rdel HG, Bora A, Kaiser J, et al.: Density-dependent reproduction
in the European rabbit: a consequence of individual response and
age-dependent reproductive performance, Oikos 104:529539, 2004.
CHAPTER 3

Biomagnification of DDT
Affects Raptor Populations

In this chapter, a case where homeostasis at the population level is affected


by a pollutant acting on individuals will be examined. A pollutant is a
chemical found in ecological systems due to human activities that causes
harmful effects on organisms. Populations can be regulated and that regu-
lation occurs through negative feedback mechanisms that act to increase
birth rate or decrease death rate when populations are small and to de-
crease birth rate and increase death rate when populations are large. What
happens when feedback mechanism fail to work properly is a question of
great interest to biologists.
Dichloro-diphenyl-trichloroethane (DDT) is a human-made chlori-
nated hydrocarbon. DDT possesses insecticidal properties and has led to
the evolution of pest populations resistant to the negative effects of DDT.
DDT is both stable and lipophilic, as are many chlorinated hydrocarbons,
which breaks down slowly in the environment and can be stored in animal
fat tissue. DDT is not particularly lethal to humans, so it was used in World
War II to control mosquito and louse populations for the protection of US
troops. DDT became the pesticide of choice after World War II, because
it was inexpensive, effective, and thought to be nonlethal to vertebrates.
In addition to its use as the main weapon to eradicate mosquitoes in the
global war against malaria in the 1960s, it was used heavily in agriculture
to eliminate crop pests. DDT was considered to be the solution to many
of our insect pest problems. Heavy use led to contamination of environ-
ments around the world; DDT can even be found in the fat tissue of Arctic
animals where it has never been sprayed. DDT is known to break down,
usually slowly, by microbes to dichloro-diphenyl-dichloroethylene (DDE).
32 POPULATION HOMEOSTASIS

DDE is commonly detected at concentrations in the environment that


often exceed those measured for DDT.
Reports of environmental contamination and observations of
eggshell-thinning and consequent reproductive failure of birds-of-prey
led scientists to suspect that environmental contamination was the cause
of population declines in certain bird species, including bald eagles
(Haliaeetus leucocephalus) and peregrine falcons (Falco peregrines). Jeffrey
Lincer, although not the first scientist to study the correlation between
raptor population declines and contamination by DDT and DDE, was
the first scientist to conduct a controlled study of captive raptors com-
bined with field observations of natural populations to determine if there
was a causative relationship (that is, to examine if DDE contamination
was causing eggshell-thinning). Lincer chose to work with a raptor closely
related to the peregrine falcon, whose populations were known to be in
decline. The American kestrel or sparrow hawk (F. sparverius) is more
common than peregrines, can be bred in captivity, and feeds on large
insects, lizards, and mice.
Lincer established 85 nest boxes in a 155 km2 area near Ithaca, New
York, to provide a source from which eggs could be collected for analysis
or for young to be used in experiments. Nest boxes that became occupied
by kestrels were checked two to three times per week for eggs, which were
counted and monitored for hatching, and sometimes taken back to the
laboratory. When taken to the laboratory, the biologist checked the eggs
for cracks, weighed them, and measured the breadth and length using cal-
ipers. Lincer broke each shell in half at the equator, removed the contents
of the eggs and measured the thickness of the shell at the equator. The
scientist took four measurements of each shell half, yielding eight total
measurements, threw out the highest and lowest measure, and used the
rest to calculate mean shell thickness. He also calculated Ratcliffes Index
(RI), which is equal to the weight of the dried eggshell in milligrams di-
vided by the length times the breadth in millimeters. The contents of eggs
were analyzed for concentrations of DDT, DDE, and other chlorinated
hydrocarbons, although only DDE was determined in each year of the
study. DDE was consistently high in the eggs.
Eggshell thickness measurements were compared to DDE concen-
trations in eggs. Eggshell thickness was expressed as a percentage of
Biomagnification of DDT Affects Raptor Populations 33

pre-DDT thickness from measurements made in an earlier study. Shell


thickness declined as concentration of DDE in eggs increased. These field
concentration data indicate a correlation between DDE concentrations
and eggshell thickness. Correlation does not equal causation, and other
chlorinated hydrocarbons (even in low concentrations) could be the cause
of the observed effect. Other substances for which the scientist did not
analyze could also be the cause. PCBs are one candidate chemical that
might be causing the observed effect, although Lincers data on PCBs is
quite meager.
Very often, supporting evidence comes from a comparative approach
in which multiple related species are examined for an observed correla-
tion. Lincer collected data from other studies that examined DDE con-
centrations in eggs and the percentage decrease in eggshell thickness from
pre-DDT levels. These data can be used to help support the conclusion
that DDE is causing eggshell thinning. In fact, there was a greater de-
crease in eggshell thickness as DDE concentration increased across spe-
cies, and several populations of raptors that had the greatest percentage
decrease in thickness were known to be in decline. However, even with all
of these data, the relationship is largely correlative, and an experimental
manipulation was necessary to draw a conclusion about the effect of a
particular independent variable on a dependent variable.
Lincer knew that he had to perform a manipulative experiment. Birds
for the breeding experiment were trapped during autumn migration and
kept in cages large enough to allow for short flights with one male and
one female in each cage. Each kestrel was fed 30 grams of meat daily from
day-old chicks. The following spring kestrels began to be fed meat dosed
with one of five concentrations of DDE, and this continued through
the summer. Each bird was randomly selected to receive food injected
with either a dose of DDE or a control. Lincer made solutions of four
DDE concentrations: 0.3, 3.0, 6.0, or 10.0 parts per million (ppm),
based on wet weight. Parts per million (ppm) is concentration of a sub-
stance in media and equals mg per liter (mg/L) of water or per kilogram
(mg/kg). The fifth concentration was 0 ppm, the control, which was
sesame oil, the solvent. Lincer injected 0.2 mL of each solution into the
meat just prior to feeding. The birds were observed during that time, and
any eggs produced were removed. Eggs were collected 5 days after the
34 POPULATION HOMEOSTASIS

effects of DDE in diet of kestrels


250 0.20
= shell thickness
DDE concentration in eggs (ppm)

eggshell thickness (mm 1 SD)


= shell thickness regression
= [DDE] in eggs 0.18
200 = egg DDE regression

150 0.16

100 0.14

50 0.12

0 0.10
0 2 4 6 8 10
DDE concentration in diet (ppm)

Figure 5 Relationship between dietary DDE fed to American kestrels


and DDE in eggs produced and eggshell thickness.
Source: Data from Lincer, 1975, Figure 2 and Table 2.

last egg of a clutch was laid to allow development to occur to determine


embryo viability. The scientist collected the same data on eggs from the
laboratory population as he did from wild populations (Figure 5).
Lincers laboratory experiment demonstrated that increased con-
sumption of DDE led to decreases in eggshell thickness. The magni-
tude of eggshell thinning corresponded well with the field data, further
supporting the conclusion that DDE was a causative agent in eggshell
thinning. Notice that even the control eggs had some DDE, which was
likely due to a birds diet prior to being captured. It is possible that birds
were contaminated by other chemicals, and testing for other chemicals
in the blood of captive adults would have helped rule out other causative
agents. Unknown contamination could have led to the results obtained,
but it is unlikely because of the random assignment of birds to treatments
and the replication of the experiment.
Despite the potential flaws, the conclusion from the manipulative
experiment combined with field observations is strong. The weight of
evidence that DDE was the causative factor in eggshell thinning was
mounting from field studies, comparative studies, and manipulative ex-
periments. The eggshell thinning data, along with the fact that mosquitoes
Biomagnification of DDT Affects Raptor Populations 35

and other pests were rapidly evolving resistance to DDT, ultimately led to
the banning of DDT in many countries. Chlorinated hydrocarbons last
a long time in the environment and biomagnify in food chains and food
webs. Biomagnification is the increase in concentration of persistent pol-
lutants that occurs in a food chain.
Small amounts in organisms at the base of food webs, such as plants,
accumulate in higher trophic levels because of the lipophilic nature of
these compounds, their persistence, and the fact that each predator eats
many prey items, each of which may have the contaminant. Ingested
contaminants are not excreted but stored, accumulating over time to
concentrations that may cause harm to the organism. In the case of
birds-of-prey and other predators at the top of food webs, concentrations
increase to levels that disrupted their ability to reproduce. This translates
at the population level to a disruption in population homeostasis. Even
though banned in many countries, DDT is still used in some countries,
and DDT sprayed decades ago is still in the environment in the form of
DDT, DDE, and other byproducts.

Bibliography
Cleveland CJ, Weis JS: DDT, The Encyclopedia of Earth (website): http://
www.eoearth.org/view/article/151631. Accessed August 31, 2010.
Lincer JL: DDE-induced eggshell-thinning in the American kestrel: a
comparison of the field situation and laboratory results, J Applied Ecol
12(3):781793, 1975.
Conclusion
Homeostasis does not necessarily mean that populations do not change.
As with any biological system, when the environment changes, the system
responds. Responses often occur through negative feedback mechanisms,
which act to reverse change in density. As the environment changes, natu-
ral selection maintains the fit of the organism to that environment. Given
that the environment constantly changes, homeostasis leads to changes in
the genetics of populations. Homeostasis also involves regulating popula-
tion sizes through negative feedback. If a population grows too large, its
growth may slow down or even reverse. Population densities vary, and
changes in the environment may lead to altered densities best suited for
the new conditions through the action of feedback mechanisms. Survival,
growth, and reproduction all require energy, and organisms make deci-
sions about how to allocate their resources, which affects populations.
Populations interact with their environment, as discussed when studying
the effects of predators on populations. Populations interact with each
other and with the nonliving components of the environment, and these
interactions can also lead to feedback and homeostasis.
Glossary
biomagnification. Biomagnification is the increase in concentration of persistent
pollutants that occurs in a food chain.
boreal forest. A subarctic, evergreen coniferous forest dominated by firs and spruces.
camouflage. Camouflage is concealment by means of disguise or protective
coloration.
clumped dispersion. Clumped dispersion occurs when individuals are closer than
expected by chance.
density. The quantity of a substance or population per unit volume, unit area, or
unit length.
density-independent. A density-independent factor affects the size of a popula-
tion regardless of the population density.
energy budget. An energy budget is a measure of the energy entering and leaving
a biological system.
feedback mechanism. A process that uses the conditions of one component to
regulate the function of the other. It can increase (positive feedback) or dampen
(negative feedback) the change in the system.
hierarchy. A social ranking system in many animals, where individuals are ranked
one above the other according to status or authority.
homeostasis. Maintenance of internal conditions within a range of acceptable
extremes.
lichens. Lichens are composite organisms made up of a fungus that grows symbi-
otically with algae that forms a crust-like growth on rocks or trees.
mortality. The relative frequency of deaths in a population.
nearest neighbor distance. In ecology, the distance to the nearest individual or-
ganism, usually of the same species.
parts per million. Parts per million (ppm) is concentration of a substance in
media and equals mg per liter (mg/L) of water or per kilogram (mg/kg).
pollutant. A pollutant is a chemical found in ecological systems due to human
activities that causes harmful effects on organisms.
populations. A population is a group of individuals of the same species living in
the same place at the same time.
predator. Predators are organisms that obtain energy by consuming, and usually
killing, other organisms.
random dispersion. Random dispersion occurs when the probability of finding
an individual at any point in the area is the same for all points.
regulate. To regulate is to control the size of a population with a tendency to
achieve or return to a size at equilibrium with the environment.
uniform dispersion. Uniform dispersion occurs when there is a regularity of dis-
tance between individuals.
Index
American kestrel, 3233 Haliaeetus leucocephalus, 32
Aviary, 4 Holbrook, Sally, 1518
Homeostasis, 1, 6, 13, 15
Behavioral mechanism, in prey, 18 in populations, xiii
Biomagnification, 35
Biston betularia, 1 Insularia, 2, 5
Boreal forest, 23
Jack pine, 23
Camouflage, 2 clark-evans (CE) statistics for, 24
Carbonaria, 1, 2 mapping method of living and dead
Clumped dispersion, 25 trees in, 24
Conservation biology, 12
Kenkel, Norm, 2326
Damselfish, 18 Kettlewell, Bernard, 15
mortality rates in, 1516 Kunin, bill, 2628
underwater study of, 17
Darimont, Chris, 910 Lichens, 2
Dascyllus flavicaudus, 15 Lincer, Jeffrey, 3234
Density-independent, 25
Dichloro-diphenyl-dichloroethylene Mark-release-recapture
(DDE), 3132, 33 experiment, 3
eggshell thinning, 3334 Modified Clark-Evans statistic, 24
Dichloro-diphenyl-trichloroethane Mortality rate, 8
(DDT) Moths
biomagnification of, 3135 conspicuousness of, 2, 3
Diplotaxis erucoides, 26 inconspicuousness of, 2
D. trimaculatus, 15
Natural selection, xiii
Energy budget, 1 Nearest neighbor distance
Environmental change, xiii (NND), 24
Environmental factors, 15 white wallrocket, function
European rabbits of,2627
annual changes in densities of, 20 Negative feedback mechanisms, 11
birth rates, 19 environmental changes, 28
number of litters per female, 21
reproductive season, beginning Oryctolagus cuniculus, 19
of,19
Parts per million (ppm), 33
Falco peregrines, 32 Phenotypic change, rate of, 910
Feedback mechanism, 2, 17, 23, 26 Pheromone, 3
birth rates, affecting, 19 Pinus banksiana, 23
F. sparverius, 32 Pollutant, 31
42 INDEX

Populations Ratcliffes Index (RI), 32


death of individual, affects, 113 Regulate, population, 15
of european rabbits, 19, 20 Reznick, David, 67
female density, 2022 Rivulus hartii, 6
regulate, 15 Rdel, Heiko, 1922
regulated through feedback
mechanisms, 1529 Saving individuals, 1113
Predation, 1, 16, 18 Schmitt, Russell, 1518
by birds, 2 Social hierarchy, 22
direct effects of, 6
indirect effects of, 6, 8 Typica, 1, 2, 5
Predator, 1
humans as, 10 Uniform dispersion, 25
reintroduction of, 9
Prey density, 17 Walsh, Matthew, 67

Random dispersion, 24
Random distribution, of plants, 25
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