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Population
Bundlethe more
camouflaged against the soot that accumulated on tree bark as
books you buy,
a result of burning coal. Populations are shown to be regulated
the greater your
Homeostasis
by feedback mechanisms, several of which are discussed here.
discount!
Populations are regulated by extrinsic factors, such as competi-
tion and predation, and that lead to changes in intrinsic factors,
THE CONTENT such as reproduction. Changes in population density often lead
Energy Physics to initiation of feedback mechanisms, such as changes in birth
Engineering or death rates. In a final example, pollutants are shown to be a
factor that can disrupt homeostasis of populations. In particular,
Biotechnology
populations of top predators, such as raptors, have suffered due
Biology
to biomagnification of toxins.
Mathematics
Chemistry Christopher J. Paradiseis professor of biology and environ-
mental studies at Davidson College. He teaches introductory
biology, ecology, entomology, and topical seminars on ecotoxi-
THE TERMS
cology and renewable natural resources. He also occasionally
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leads a study abroad program in India. His research evaluates
for a one time fee anthropogenic factors that influence insect biodiversity at a
No subscriptions or variety of scales. His current research interests include effects of
access fees land use patterns on pollinator communities in parks.
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A. Malcolm Campbellteaches biology at Davidson College,
concurrent usage
NC. He received national and international education awards:
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Genetics Society of America (2013); American Association for
Free MARC records
the Advancement of Science (2012); and American Society for
Cell Biology (2006). He was the founding co-editor in chief of
For further information,
CBE Life Sciences Education; founding director of Genome
a free trial, or to order,
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sales@momentumpress.net American Society for Cell Biology governing council (20122014)
A. Malcolm Campbell
Population Homeostasis
Population Homeostasis
10 9 8 7 6 5 4 3 2 1
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
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
60 recaptured males
= typica
50 = carbonaria
= insularia
percentage
40
30
20
10
0
A polluted-1953 polluted-1955 lichen-covered
80
percentage
60
40
20
0
polluted-1953 polluted-1955 lichen-covered
B forest
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
0.6
a
= natural
b b
0.2
0.1 c
0.0
mean change maximum change
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
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
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
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
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
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
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
150 0.16
100 0.14
50 0.12
0 0.10
0 2 4 6 8 10
DDE concentration in diet (ppm)
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
Random dispersion, 24
Random distribution, of plants, 25
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