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Biology Christopher J. Paradiseis professor of biology and environ-
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A. Malcolm Campbell
Organismal Homeostasis
Organismal Homeostasis

Christopher J. Paradise, PhD

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

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

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 Abstract
Organisms maintain homeostasis in a variety of ways. In the first part
of this book, mammals are shown to regulate their body temperatures
through homeostatic mechanisms. The data from thermoregulation ex-
periments that demonstrated the role of neurons in body temperature
homeostasis are examined. The second part of this book discusses how
organisms allocate the limited energy that is available to them for sur-
vival, growth, or reproduction. Excess energy in individuals can translate
to growth of populations: if enough remains after survival and growth, it
can be allocated to reproduction. However, even closely related organisms
may have different strategies for allocating resources that are dependent
upon the environmental conditions in which they exist.

Keywords
endotherms, body temperature, ambient temperature, seasonal dimor-
phism, phenotypes, hypothalamus, evaporative heat loss, principle of
allocation, mass budgets, assimilation, allocation, consumption, repro-
duction, biomass, foraging
 Contents
Preface...................................................................................................ix
Acknowledgments....................................................................................xi
Introductionxiii
Chapter 1 Mammals Possess Adaptations to Stay Warminthe
Winter and Cool inthe Summer........................................1
Ethical, Legal, Social Implications: Biologists Might
Consider Studying Males and Females Separately.........12
Chapter 2 An Individuals Foraging Can Affect the Entire
Population.......................................................................17
Ethical, Legal, Social Implications: Negative Birth
Rates in Human Societies Can Have Positive and
NegativeConsequences................................................27
Conclusion............................................................................................31
Glossary................................................................................................33
Index....................................................................................................35
 Preface
This book about organismal 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 some aspects of organismal homeo-
stasis and some of the supporting evidence behind our understanding.
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 organismal homeostasis the way profes-
sional 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 administrators
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 col-
laboration 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
In this book, two topics that relate to homeostasis at the organismal level
will be addressed. The first chapter is about maintaining a comfortable
body temperature, which many humans probably think about on a daily
basis. However, data will be examined that show how the body does much
of this work without the animal even knowing about it. In the second
chapter, how energy and nutrient resources affect homeostasis of indi-
viduals will be discussed. These resources obtained or not by individuals
then affect populations, to make the connection between acquisition of
energy and nutrients and the growth of populations. The case studies in
this book address energy flow, which is central to the homeostasis that
maintains the many physiological processes that permit all organisms to
survive and reproduce. All life requires energy; and if energy is limited,
individuals must make decisions about how to allocate their resources,
and this affects populations. By the end of the book, a better understand
how individual organisms maintain homeostasis will be obtained, as will
connections to homeostasis at the individual and population levels.
 CHAPTER 1

Mammals Possess
Adaptations to Stay
Warminthe Winter and
Cool inthe Summer

All organisms display some common properties regarding homeostasis.

They assess their current status for various physiological processes, they
collect information from the environment, and they integrate these indi-
vidual and environmental informational cues in order to maintain homeo
stasis. Finally, they often respond at the cellular level, and each cell contributes
to maintaining homeostasis of the entire organism.
In order to understand these common properties, scientists have in-
vestigated how animals maintain their internal body temperature. Mammals
and birds are called endotherms because they maintain a consistent internal
body temperature, compared to most insects, reptiles, most fish, and most
other animals. Endotherms are animals that maintain a consistent internal
core body temperature through physiological processes. Birds and mam-
mals can generate heat when they get cold, and they have mechanisms
todissipate heat when they are too warm. The core body temperature of
mammals varies by only a couple degrees for cats, rabbits, and kangaroo
rats, which are all placental mammals.
The core body temperatures of some less derived, more ancestral species
of mammals, such as the platypus and echidna, fluctuate more and drop
when the ambient temperature drops, but other mammals have much more
stable internal temperatures. Platypus and echidna vary their body tem-
peratures by as much as 10 when the ambient temperature ranges from 5 to
35C. Interestingly, platypus and echidna are both egg-laying mammals, and
2 ORGANISMAL HOMEOSTASIS

they evolved prior to the origin of placental mammals like cats, rabbits, and
kangaroo rats that give birth to live young. The original data that helped bi-
ologists understand how body temperature in endotherms is an example of
homeostasis at the organismal level will be the focus of this chapter.
Temperatures of extremities, such as arms and legs, will fluctuate more
with ambient temperatures even of endotherms. For instance, humans
vary their core temperatures no more than 1 C, whereas the skin tempera-
ture can vary by 4 C across the same range of ambient temperatures.
Reptiles, amphibians, insects and other animals that are ectotherms
have body temperatures that are highly correlated with, and essentially
the same as, ambient temperatures. As the ambient temperatures decline,
so do the body temperatures of ectotherms.
Scientists examining these temperature data wondered how endo-
therms maintain a constant body temperature across such a wide range of
ambient temperatures. Endotherms utilize homeostasis to regulate their
core body temperature more tightly than their surface temperature, al-
though the mechanism is unclear. It is well known that insulation is
crucial to efficiently heating and cooling a large space, such as a school.
The same turns out to be true for endothermic mammals and birds.
Endotherms have fat and fur or feathers that insulate their body.
What do animals do if they live in climates that have very large sea-
sonal changes in ambient temperatures? The arctic fox (Alopex lagopus), as
its name implies, lives in one of the coldest places on Earth. Arctic foxes
are reputed to have the warmest fur of any mammal, and the winter coat
looks different from the summer coat. The winter coat is visibly thicker and
white, whereas the summer coat is brown and hairs look shorter, at least
from a distance. Biologists use the term seasonal dimorphism to describe
these changes (dimorph means two forms). Seasonal dimorphism de-
scribes organisms with two phenotypes depending on the time of year.
Incontrast, humans do not have fur coats, so we cannot tolerate cold arctic
temperatures like the arctic fox.
Many mammals shed their hair in a gradual process when the seasons
change. The fox gradually loses its brown hairs, which are replaced by
white hairs. It would not be adaptive to lose all its summer hair quickly
and be bald when winter was approaching, so the change is gradual. Differ-
ent endotherm species use different mechanisms to maintain a consistent
body temperature. Consider the arctic fox as an example.
 MAMMALS POSSESS ADAPTATIONS TO MAINTAIN BODY TEMPERATURE 3

Scientists measured the length of arctic fox fur at 35 different sites on

the body of many individuals every month for 1 year (Underwood and
Reynolds, 1980). The investigators calculated the percent change in win-
ter fur length compared to the body position-specific average in August
when the fur was at its shortest length. The number of foxes measured
each month ranged from four to twelve, depending on how many they
could capture. The investigators also determined the overall average
monthly fur length (a single average from all 35body sites).
Foxes grow their fur significantly longer on most, but not all, parts of
their body. The researchers documented changes that ranged from 62%
(an actual shortening of fur length on the nose) to an almost 300% in-
crease in fur length (winter hairs almost 4 times as long as summer hairs)
in certain parts of the body. Statistically significant increases ranged from
132 to 275% longer. The insides of the legs, the ears and the nose were
the only places on the arctic fox where hairs were not significantly longer
in the winter than in the summer. It appears foxes grow thicker fur insula-
tion over most of their bodies, except where they need to maintain good
sensory perception (nose, mouth, and ears) and in areas that can radiate
heat centrally when they curl up for sleeping (inner legs). Seasonal dimor-
phic hair growth is one mechanism of homeostasis.
The researchers related fur length to the difference of body minus
ambient temperature to determine if they could detect whether changes
in temperatures provided a cue to the foxes. The researchers obtained
core body temperatures of the foxes using a rectal thermometer, which
averaged 38.4 C for the year. Monthly ambient temperatures were
averaged.
Throughout the year, foxes collect temperature information from the
environment and their bodies, and they respond to that information to
maintain body temperature homeostasis. Overall hair length has a positive
correlation with the temperature measurement (body minus ambient).
The temperature goes up not because the body temperature is going up,
but because the ambient temperature is decreasing rapidly with the
approach of winter (for the arctic fox and other placental mammals, core
body temperature remains relatively stable over time). Fur length lags
behind the change in ambient temperature, which could indicate the lag
time between foxes sensing the temperature change, their cells processing
that information, and the amount of time it takes for hair to grow
4 ORGANISMAL HOMEOSTASIS

noticeably longer. However, the length of day is also changing. It is im-

possible to know from these data whether light and/or temperature is the
environmental factor that stimulates hair growth in the fox. However, the
scientists could experimentally determine if one or both are required if
foxes were tested under laboratory conditions where light and tempera-
ture could be better controlled.
In contrast to the arctic fox, consider homeostatic body temperature
for a mammal that lives in one of the hottest places on Earth. The drom-
edary camel (Camelus dromedarius) also displays seasonal dimorphism
with thick, dark winter and thin, light colored summer coats (Figure 1).
Winters can be chilly in the northern desert, averaging around
13 C/55.4 F. But even during the summer, temperatures in the Sahara
Desert may range from cool at night to a blistering 54 C/129 F during
the day. In addition to the heat, the desert is an arid environment with
water sources separated by large distances. In extreme heat, humans sweat
and dogs pant excessively. Camels neither pant nor sweat in order to regu-
late their body temperature, unless their body temperature is exceedingly
high, unlike many other mammals.
The study of animal temperature homeostasis has involved many dif-
ferent biologists working with many different animals. The benefit of this
diversity is we can be more certain that broad principles apply to most if
not all endotherms. The problem though is that working with different
experimental conditions makes it challenging to synthesize all of the in-
formation into a cohesive working model. Despite the challenge of work-
ing with diverse data sets, one can deduce many generalized rules about
homeostatic thermoregulation in endotherms. One of the pioneers in
thermoregulation was Knut Schmidt-Nielsen, who researched animal
physiology from the 1950s through the 1970s. Among other species,
Schmidt-Nielsen studied camels because they endure extreme conditions
in the desert.
Biologists quickly explained many of the most obvious physical traits
that help camels survive desert conditions. Camel fur reflects solar heat
and provides a layer of cool, insulating air compared to the hot desert air.
Their long legs lift the animals off of the hot sand to minimize their expo-
sure to the radiating heat from the ground. The most obvious anatomical
feature, their hump, does not carry water. The humps are large fat deposits.
 MAMMALS POSSESS ADAPTATIONS TO MAINTAIN BODY TEMPERATURE 5

Figure 1 Dimorphism of camel fur. Winter (A) and summer (B)

coats of the camel Camelus dromedarius. These individuals live in
temperate European wildlife parks.
Source: A, From http://commons. wikimedia.org/wiki/File:Camelus_dromedarius_resting.jpg,
Author: LadyOfHats. Public domain. B, From http://commons. wikimedia.org/wiki/
File:Bactrian_Camel,_Whipsnade_-_geograph.org.uk_-_857721.jpg,Author: Stanley Howe.
This file is licensed under the Creative Commons Attribution-Share Alike 2.0 Generic license.
6 ORGANISMAL HOMEOSTASIS

The fat is not a delayed source of water, although fat metabolism does
produce a small amount of water. The amount of oxygen required to me-
tabolize the fat would result in more loss of water through breathing arid
desert air than would be gained through fat metabolism. For camels, fat
storage is used as a compact energy source but its location is an adaptation
to desert living. Fat is a very good thermal insulator, which means camels
are protected from the solar heat coming from above. Camels have very
little fat in other locations, which means they can radiate out excess heat
from the rest of their bodies.
Perhaps camels do not need to pant or sweat because they do not
gethot. Physiologists in the 1950s wanted to know if a camels body tem-
perature was as consistent as other placental mammals. To conduct their
research, the investigators measured the core body temperatures of camels
that were well hydrated and compared them to camels that had not con-
sumed water for several days. To measure core body temperatures, inves-
tigators inserted an appropriately sized thermometer in the rectum of the
individuals. Each camel was monitored for several days, and the results
from two individuals under the two hydration conditions were strikingly
different.
To the surprise of Schmidt-Nielsen and other biologists, camel body
temperature fluctuates substantially compared to other mammals. The
dehydrated camel had core body temperatures that fluctuated daily be-
tween about 35 C after midnight to as much as 41 C in the late after-
noon. The hydrated camel had core body temperatures that fluctuated
daily between about 37 C to only as high 39 C. Hydrated camels experi-
ence less fluctuation than dehydrated camels, which simply reflects the
larger mass of hydrated animals and waters chemical property of chang-
ing temperatures slowly.
Although camels appear to have an average body temperature near
37 C like humans, a human would risk brain damage with a core body
temperature that fluctuates as much as a camels, which can easily reach
40 C (104 F). By about 6 PM, before the sun begins to set near the equa-
torial horizon, camels routinely experience very high body temperatures,
but they dont begin to sweat until their core temperatures reach 41 C
(106 F). And in fact, camels maintain their brain temperature within
tighter limits than the rest of the body. A camel is better able to keep their
 MAMMALS POSSESS ADAPTATIONS TO MAINTAIN BODY TEMPERATURE 7

brains cool than most mammals, even when their body temperature is very
high. It turns out that many mammals can regulate both body and brain
temperature, and how this happens will be explored later in this chapter.
Prior to Schmidt-Nielsens research, other physiologists were trying to
understand where the homeostatic temperature regulation mechanism
was located within the body. Over time, philosophers argued that the
heart or even the liver regulated body temperature. By the 1930s, physi-
ologists focused their attention on the brain as the source of temperature
homeostasis (Figure 2). A team of physiologists working at the University
of Chicago refined the work of previous investigators and performed very
precise brain surgery on anesthetized cats and measured the consequences
(Clark et al., 1939).
In particular, these surgeries destroyed very small regions in the front,
or anterior, part of the hypothalamus, a region at the base of the brain near
the junction of the spinal cord and the brain that regulates homeostasis

8
temperature change from average (F)

2
0 = 101.4 F
0
70 75 80 85 90 95
2

10
temperature of room (F)
Figure 2 Thermoregulation of cats after surgical manipulation of
hypothalamus. The eleven lines represent the core body temperatures
for experimental cats when placed in a room of the indicated
temperatures. The cats temperature was subtracted from the average
of 101.4o F before graphing. Each line represents a different cat,
and the length of the line indicates how long the cat lived.
Source: Data from Clark et al., 1939, Table 1.
8 ORGANISMAL HOMEOSTASIS

ofmany functions. When the cats recovered from surgery, they were
placed in rooms with different temperatures, and the investigators mea-
sured the core body temperatures of the cats as the room temperature
changed. The typical core body temperature of cats is 101.4 F +/ 1.5.
The team of physiologists measured the temperatures of eleven cats after
destroying the anterior hypothalamus, but five of the eleven cats died in
less than 10 days after the surgery. All five animals experienced either
abnormally high (maximum of 108.5 F) or low (minimum of 94.2 F)
core body temperatures before dying. Subsequent experiments by different
investigators have confirmed the results in Figure 2, so the data should be
considered valid even in the absence of data from the control animals.
Although previous publications implicated the hypothalamus in
body temperature homeostasis, the 1939 publication of Figure 2 dem-
onstrated what happened when a small area of the hypothalamus was
damaged. Scientists might have expected body temperatures to correlate
with the room temperatures like an ectotherm, but the data indicate
that body temperatures were no longer linked to the animals surroundings.
Rather than drifting up in warm rooms the way reptiles do, mammals
lacking a functional hypothalamus have no control over thermoregula-
tion, which might have caused the deaths of five animals in the study.
Rather than coasting to a standstill like a sailboat and passively riding
the waves of ambient temperature up and down, the heating and cooling
mechanisms were still functioning but they lacked any feedback to know
when to stop. The data indicate that mammals without functional tem-
perature regulation behave like a speedboat moving without any steer-
ing and can generate more heat despite being in a warm room, and vice
versa. In mathematical terms, the correlation coefficient for these data
would be very near zero.
The next set of experiments also focused on the cat hypothalamus,
butthey were published in 1961 by another team of physiologists
(
Nakayamaetal., 1961). In this experiment, the investigators gently
warmed individual neurons in the hypothalamus of anesthetized cats and
measured the neurons rate of depolarization and the animals breathing rate.
The data from this experiment indicate that individual neurons can
sense their local temperature and respond to changes by altering the rate
of their action potentials and thus cell-to-cell communication. After the
 MAMMALS POSSESS ADAPTATIONS TO MAINTAIN BODY TEMPERATURE 9

brain begins information processing, the anesthetized animal altered its

breathing rate, which helped maintain the normal body temperature.
However, it is important to note that first action potentials begin in to
increase dramatically. There is a delay before respiration rate increases.
After respiration rate begins to increase, neuronal activity declines, such
that the rate of neuronal activity is negatively correlated with respiration
rate; the temperature of the hypothalamus remains high, while the activ-
ity declines. The slight disconnect between neuronal temperature and its
activity could be caused by one or more mechanisms, none of which are
evident in this experiment. However, it appears that an increase in hypo-
thalamus temperature leads to a slightly delayed organismal response of
increased breathing and perhaps other physiological responses not re-
corded in this experiment.
The next set of experiments to consider used horses (McConaghy et al.,
1995). In order to make sense of the data, one needs to understand basic
horse head anatomy. The brain is located at the end of a long nasal cavity,
and the hypothalamus sits immediately above a small air pocket called the
cavernous sinus despite its relatively tiny size. The investigators placed
adult horses on treadmills where the speed of walking could be regulated
experimentally. The treadmill rate was gradually increased until the horses
were trotting at 7 meters per second (m/s). After 30 minutes, the horses
were allowed to walk at their own paces.
Throughout the experiment, the veterinarians measured the horses
temperatures in four locations: the pulmonary artery (representing core
body temperature), the hypothalamus, the cavernous sinus, and the rectum.
After analyzing the data, the investigators surgically altered the airflow by
connecting the trachea directly to the outside air, which meant the horse
could no longer use its nose or nasal sinus for breathing. The horse was
forced to accelerate its speed to 6 meters per second and then allowed to
slow down after 18 minutes of exercise, because the investigators did not
want the individual to overheat or suffer. Once again, the investigators
measured the temperatures of the experimental horse in the same four
anatomical locations.
Under natural, unmanipulated conditions, the horse maintained a
cooler cavernous sinus compared to the other three areas measured. Ini-
tially, the core and hypothalamus temperatures are identical; but once the
10 ORGANISMAL HOMEOSTASIS

horse reaches 7 m/s, the hypothalamus temperature does not rise as rapidly
as the core temperature, even though both rose steadily as the horse ran
the treadmill. The hypothalamus never got as hot as the core temperature.
Rectal temperatures rose also, but lagged behind those two temperatures.
Concurrently, as the hypothalamus temperature rose, the temperature of
the cavernous sinus decreased.
The cavernous sinus is adjacent to the hypothalamus, which protects
the brain region from overheating. If the horse is going to process its core
body temperature, neurons in the hypothalamus must detect an increase
in temperature so that they can increase their rate of action potentials and
initiate physiological responses to cool the animal. One physiological re-
sponse to exercise and hypothalamic heating is increased breathing, which
brings in more air that can reduce the temperature of the cavernous sinus
located at the far end of the nasal cavity. When the horse stops trotting,
the cavernous sinus briefly warmed up, but it gradually cooled off again
as the animal continued to regulate its body temperature through a vari-
ety of mechanisms including sweat evaporation. The core body tempera-
ture cooled faster than the hypothalamus until the two regions approached
their original 38 C.
When air was prevented from entering the horses nasal cavity, the
cavernous sinus heated up quickly, instead of staying stable or even de-
creasing as it did in the normal horse. This made the hypothalamus get
nearly as hot and rise as quickly as the core body temperature until the
experiment was stopped out of concern for the animal. The experimental
horse experienced a rise of about 3 C in the cavernous sinus compared to
the control animal. The hypothalamus cooled off very slowly, in fact more
slowly than the core body temperature.
From these data and those of previous experiments, it appears mammals
allow the hypothalamus to warm a little bit, which stimulates neurons to
increase their depolarization rate. The subsequent release of neurotrans-
mitter initiates cooling responses to maintain thermal homeostasis. Rapid
breathing is one cooling response that brings in cool air and maintains the
cavernous sinus to remain cooler than many other parts of the body. The
cooler temperature in the cavernous sinus protects the hypothalamus from
getting too warm, which could lead to brain damage. If the hypothalamus
were damaged, then the organisms ability to regulate body temperature
 MAMMALS POSSESS ADAPTATIONS TO MAINTAIN BODY TEMPERATURE 11

would be disconnected from its ability to increase or decrease body tem-

perature, as shown in Figure 2 when the cats lost hypothalamus function.
In short, endotherms sense their current body temperature and the ambient
temperature, integrate this information, and use neuronal responses to
maintain organismal thermal homeostasis.
In a final experiment, a human volunteer was used to examine the
relationship between heat generation and cooling capacity when this person
was exposed to different temperatures (Benzinger et al., 1961). Investiga-
tors ran water of a fixed temperature over the arms of one healthy young
man intermittently over the course of 3 months. The goal was to expose
him to different external temperatures ranging from 20 to 31o C. Once
his skin temperature reached the indicated value, investigators monitored
the cranial temperature, via the eardrum, of the young man as well as the
amount of heat generated by cellular respiration or heat lost by the volun-
teers sweating, which was measured by evaporative heat loss.
At all experimental skin temperatures, when internal cranial tempera-
ture was below 37.1o C, there was no evaporative heat loss. But there was
heat production at those lower internal cranial temperatures and heat pro-
duction was higher at lower skin temperatures. Once the internal cranial
temperature rose above 37.1o C, heat production dropped to 20 calories
per second, which is likely to be the basal metabolic rate of heat produc-
tion, and evaporative heat loss increased dramatically from 0 to almost
80calories per second at 37.6o C.
If the animal studies in this chapter can be generalized to all endo-
therms, then scientists should be able to predict the outcome when a human
volunteer is subjected to different skin temperatures. The man studied in
this experiment showed a clear trend that the colder the skin, the more
heat he generated through cellular respiration. This outcome indicates that
the hypothalamus must be able to influence overall cellular respiration
throughout the body. As the skin approached average human body tem-
perature of 37 C, the man generated less heat, because he was not as cold.
Once the core temperature reached this mans specific set point of 37.1 C,
he began to sweat and lose heat through water evaporation on various
parts of his body. He continued to sweat as long as his core body tempera-
ture exceeded his specific set temperature of 37.1 C. When the core tem-
perature overshoots 37.1 C, the body sweats and the heat production
12 ORGANISMAL HOMEOSTASIS

stops completely. The balance between heat production and heat loss at
the organismal level accomplishes the task of thermal homeostasis.
Thermal regulation at the organismal level is a classic example of ho-
meostasis. In the case of endotherms, the body utilizes feedback systems
to regulate and maintain the species-specific optimal body temperature.
Endotherms such as camels can tolerate a greater range of temperatures
during the day, but even they have feedback mechanisms to prevent their
bodies from getting too hot or too cold. Collecting and processing the
temperature information takes time, and the body responds with a slight
delay because it takes additional time to produce new heat or initiate
cooling mechanisms. As seen in the cat experiments, loss of information
processing produces inappropriate heat production or loss that is unre-
lated to ambient temperature, which can have fatal consequences.

Ethical, Legal, Social Implications: Biologists Might

Consider Studying Males and Females Separately
On July 4, 1776, the colonies of America officially ratified the Declara-
tion of Independence with one of the most famous quotes from the po-
litical realm, We hold these truths to be self-evident, that all men are
created equal, . . . . For thousands of years, people have accepted or rejected
the existence of physiological differences between men and women. No
one doubts the anatomical differences and the physiological consequences
associated with reproduction. But from that common starting place, the
spectrum of opinions varies substantially. Some people cite the Bible to
justify their opinions, whereas others reference scientific studies or statis-
tical analysis of standardized test scores. One of the most famous, or infa-
mous, public statements was made on January 14, 2005, by the president
of Harvard University, Dr. Larry Summers. Summers was reported to have
said or implied, the innate differences between men and women might
be one reason fewer women succeed in science and math careers. Regard-
less of his intentions or the context of his quote, Summers exposed a gap
in our understanding of the differences and similarities of males and females
within a species.
Biology research is supposed to understand how life works, but
research is funded and conducted by humans who have misconceptions
 MAMMALS POSSESS ADAPTATIONS TO MAINTAIN BODY TEMPERATURE 13

and prejudices. Behavioral neuroscientists, Irving Zucker and Annaliese

Beery, surveyed biology articles published in 2009 that used mammals for
research. The two investigators quantified the percentage of published
research papers that used only males, only females, both, or failed to de-
scribe the genders in their materials and methods. The discrepancy varies
among the different subcategories of biology. For example, behavioral bi-
ology studied both males and females in a majority of the papers, whereas
over half of the physiology, endocrinology, and pharmacology papers
studied only males. Reproductive biology studies used females only
slightly more than half the time. Immunology papers failed to describe
the gender of research subjects 60% of the time, although they often used
cell lines in tissue culture of unknown gender.
Perhaps gender does not really matter for many areas of biology, but
how can science know that if the hypothesis is not experimentally tested?
If gender does not matter, then finding exceptions would disprove the
hypothesis. Deborah Clegg from the University of Texas Southwestern
Medical School found major differences in gene regulation in males versus
female fat deposits. Women tend to store more fat just under their skin,
whereas men tend to accumulate fat in their abdomens. Neuroscientist
Rhonda Voskuhl recognized the gender differences in the disease multiple
sclerosis. Rather than ignoring the differences, Voskuhl has developed a
new medication that is in clinical trials to improve the health of men and
women suffering from multiple sclerosis. In 2001, the US General Ac-
counting Office found that eight of the ten drugs withdrawn from the
market between 1997 and 2000 had more severe negative effects on
women than men. Given these three examples of gender differences (not
counting reproduction), it seems clear that gender differences do exist,
but we do not understand when there are differences, nor the magnitude
of these differences.
Scientists could test the hypothesis that gender has an effect on a
physiological trait. For any experiment that tests the physiological re-
sponse to some factor, whether its internal (e.g., hormonal) or external
(e.g., a drug) to the organism, scientists should test both males and females
and then account for the factor of sex in statistical analysis. Experimental
studies should be designed to include both males and females routinely in
order to determine if there are sex effects. Other factors that may differ
14 ORGANISMAL HOMEOSTASIS

between males and females, body mass, core body temperature (which
also varies slightly among individuals), other drugs being taken, diet, nu-
tritional status, stress, and more are all factors that might affect the ac-
tions of drugs or the outcomes of disease. In fact, most drugs are prescribed
in specific doses, as if one dosage works for all adults, regardless of these
factors that vary from individual to individual. This may then cause more
or less response to the drug depending upon the individual. Biomedical
research typically does not take this variation into account. The implication
is that biomedical research should design studies that do take these factors
into account. It would be of use to at least know how much variation
there might be, even if dosages are not varied for different individuals.
In 2010, a group of biology journals agreed on a checklist of items
that should be included in the materials, and methods of all submitted
manuscripts and the gender of the study subjects was on that list. Fund-
ing agencies must also agree that gender differences should be considered
when allocating grant money. Unfortunately, very few graduate students
or medical students have any training in gender differences, so it will take
a long time for science and medicine to fully appreciate that men and
women are not created equal.

Bibliography
Benzinger TH, Pratt AW, Kitzinger C: Thermostatic control of human
metabolic heat production, Proc Nat Acad Sci 47(5):730739, 1961.
Bombardieri M: Harvard womens group rips Summers, Boston Globe
(website): http://www.boston.com/news/education/higher/articles/
2005/01/19/harvard_womens_group_rips_summers/. Accessed
November 26, 2010.
Cain JW, Krausman PR, et al.: Mechanisms of thermoregulation and water
balance in desert ungulates, Wildl Soc Bull 34(3):570581, 2006.
Clark GH, Magoun W, Ranson SW: Hypothalamic regulation of body
temperature, J Neurophysiology 2:6180, 1939.
Hardy JD, DuBois EF: Regulation of heat loss from man, Proc Nat Acad
Sci 23:624631, 1937.
McConaghy FF, Hales JRS, et al.: Selective brain cooling in the horse
during exercise and environmental heat stress, J Applied Physiol 79(6):
18491854, 1995.
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Nakayama T, Eisenman JS, Hardy JD: Single unit activity of anterior

hypothalamus during local heating, Science 134:560561, 1961.
Ramirez O III, Joregnsen J, Thrall DE: Imaging basilar skull fractures in
the horse: a review, Vet Radiol Ultrasound 39(5):391395, 2007.
Schmidt-Nielsen B, Schmidt-Nielsen K, Houpt TR, et al.: Water balance
of the camel, Am J Physiol 185:185194, 1956.
Schmidt-Nielsen K: Desert animals: physiological problems of heat and
water, 1965, Clarendon Press.
Sisson S: A textbook of veterinary anatomy, Philadelphia, 1910, W. B. Saunders
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327:15711572, 2010.
 CHAPTER 2

An Individuals Foraging Can

Affect the Entire Population

Both plants and animals use environmental information to forage for

resources. In this chapter, how energy and nutrient resources affect homeo
stasis of populations will be explored in order to make the connection
between acquisition of energy and nutrients and the growth of popula-
tions. The concept of homeostasis and how it operates at the population
level will thus be further examined.
Organisms are faced with a couple of basic problems. One is acquisi-
tion of resources, whether those resources are food, nutrients, water, mates,
or space. Another is how to allocate energy and nutrient resources among
growth and reproduction. To be successful evolutionarily, an organisms genes
must stay in the population, meaning it must reproduce and pass its genes
on to the next generation, which is fitness. The principle of allocation
states that evolution will lead to organisms that optimize partitioning of
resources to maximize fitness.
Humans face analogous problems of resource allocation in every day
life. Consider a student and how their time is a limited resource. There is
only so much time to study, eat, exercise, work, or relax. To maximize
success, a student needs to decide how best to allocate his or her time.
Lets assume on the average weekday 5 hours is spent traveling to and
from class. Then 3 hours each day is spent in the cafeteria eating and
7hours sleeping. Spending 8 hours each day playing video games leaves
1hour for studying, doing homework, exercising, or earning money. This
is not a successful strategy. Student thus has to trade-off time at the video
console for time spent studying in order to do well on a test. Similarly,
organisms might have to trade-off investment in one area when putting
energy into another area.
18 ORGANISMAL HOMEOSTASIS

Michael Fenner and his colleagues studied resource allocation in two

species of grass, Johnson grass (Sorghum halepense) and sorghum (S. bicolor).
Johnson grass is a perennial (lives for 3 or more years) weed that reproduces
and spreads through both vegetative and sexual reproduction. Sorghum is
an annual (completes its life cycle in a single growing season) crop that
reproduces only through seeds.
The scientists varied the levels of nutrients available to the plants in a
greenhouse experiment. Six to nine seeds per species per treatment were
germinated and grown in their own pots in a sand and vermiculite (a natu-
ral mineral containing magnesium, aluminum, iron, and silicon) mixture.
Plants were watered twice per week with one of three solutions, high,
medium, and low nutrient concentration. The high concentration con-
tained 0.03 g N, 0.06 g P, and 0.03 g K per 100 mL of solution, medium
was one-third of high, and low was one-ninth of high. This continued as
the plants grew.
At the end of 120 days, the approximate lifespan of Johnson grass,
those plants were harvested. The biologists counted inflorescences, tillers,
and the number of seeds per inflorescence (Figure 3A and B). An inflores-
cence is a group or cluster of flowers arranged on a stem composed of a
main branch or a complicated arrangement of branches. A tiller is a shoot
that springs from the root or bottom of a primary stalk. The scientists
separated out the different plant tissues, dried the material, and weighed
biomass (Figure 3C).
Fenner and his colleagues harvested sorghum at full maturity, at
135days. Because these plants dont have tillers or rhizomes and only one
inflorescence per plant, the scientists measured shoot and total seed mass,
as well as number of seeds per inflorescence (Figure 4). Rhizomes are hori-
zontal subterranean stems distinguished from true roots in possessing
buds, nodes, and scale-like leaves.
The scientists measured the allocation of nutrients to seeds, rhizomes,
and vegetative tissues (Figure 5). For Johnson grass, the scientists measured
the concentrations of nitrogen, phosphorus, and potassium in seeds, rhi-
zomes and shoots, and for sorghum they measured those same nutrients
in seeds and shoots.
Both grasses responded to higher nutrient levels by growing to a larger
size and producing more seeds and, for Johnson grass, more inflorescences,
 AN INDIVIDUALS FORAGING CAN AFFECT THE POPULATION 19

60 inflorescences and tillers per plant 250 seeds/inflorescences

50 = inforescences
200
= tillers
number ( 1 SE)

number ( 1 SE)
40
150
30
100
20

50
10

0 0
low medium high low medium high
A nutrient treatment B nutrient treatment

120 biomass of shoots, rhizomes and seeds

100 = shoot mass

dry mass (g 1 SE)

= rhizome mass
80 = total seed mass

60

40

20

0
low medium high
C nutrient treatment
Figure 3 Effects of nutrient level on Johnson grass. A, Numbers
of inflorescences and tillers per plant. B, Numbers of seeds per
inflorescence. C, Biomass of nonreproductive shoots, asexually
reproduced rhizomes, and sexually reproduced seeds. Height of
bars5averages; error bars 5 1 standard error (SE).
Source: From Benech Arnold et al., 1992, Table 1.

rhizomes, and tillers. Plants in the low nutrient level treatment were
smaller in each tissue category than plants in medium and high nutrient
levels. The number of seeds per inflorescence was not affected in Johnson
grass, but because there were more inflorescences in high nutrient plants,
the total number of seeds per plant increased with nutrient level. The
proportion of biomass Johnson grass allocated to seeds was about 0.05 to
0.07 and was much higher (from 0.30 to 0.37) for rhizomes. That pro-
portion remained fairly constant across nutrient levels. Thus, Johnson
20 ORGANISMAL HOMEOSTASIS

50 biomass of shoots and seeds 1400 seeds/inflorescence

= shoot mass 1200
40 = total seed mass
dry mass (g 1 SE)

number ( 1 SE)
1000
30 800

20 600

400
10
200

0 0
low medium high low medium high
A nutrient treatment B nutrient treatment
Figure 4 Effects of nutrient level on sorghum. A, Biomass of
nonreproductive shoots and sexually reproduced seeds. B, Numbers
ofseeds per inflorescence. Values are averages, and error bars
represent 1 SE.
Source: From Benech Arnold et al., 1992, Table 2.

grass allocated five to seven times more biomass to rhizomes than to seeds
at each nutrient level.
Nitrogen and phosphorus concentrations were highest in seeds com-
pared to shoots and rhizomes in Johnson grass. Rhizomes had higher con-
centrations of nitrogen than shoots. At high levels of nutrients, the shoot
phosphorus concentration was higher than in rhizomes, whereas shoots
and rhizomes had similar concentrations in low or medium nutrient lev-
els. A very different pattern emerged with potassium in Johnson grass.
Shoots and rhizomes had similar concentrations of potassium, much
higher than in seeds. The concentration of each nutrient in seeds in-
creased with increasing nutrient level. Nutrient concentrations in rhi-
zomes with increases in nutrient level either were not as dramatic or and
did not increase at all in potassium. This suggests priority to vegetative
reproduction at the expense of sexual reproduction; plants allocated high
proportions of nutrients to rhizomes no matter how low nutrient levels
were, and this caused constraints on allocation to other vegetative tissues.
Although Johnson grass individuals allocated higher concentrations of
some nutrients to seeds, the proportion of the absolute amount of nutri-
ents allocated to rhizomes was higher than that allocated to seeds due to
the higher biomass of rhizomes.
 AN INDIVIDUALS FORAGING CAN AFFECT THE POPULATION 21

concentration (mg/g dry mass 1 SE)

25 nitrogen in 8 phosphorus in 25 potassium in
Johnson grass Johnson grass Johnson grass
= shoot
20 = seed 20
6 = rhizome
15 15
4
10 10

2
5 5

0 0 0
low medium high low medium high low medium high
A B C
concentration (mg/g dry mass 1 SE)

20 nitrogen in sorghum 8 phosphorus in sorghum 35 potassium in sorghum

30
15 6
25

20
10 4
15

10
5 2
5

0 0 0
low medium high low medium high low medium high
D nutrient treatment E nutrient treatment F nutrient treatment

Figure 5 Concentration of three nutrients in Johnson grass and

sorghum. Concentrations are in mg per gram dry mass. A to C,
Nitrogen, phosphorus, and potassium in Johnson grass shoots,
seeds, and rhizomes, respectively. D to F, Nitrogen, phosphorus,
and potassium in sorghum shoots and seeds, respectively.
Source: From Benech Arnold et al., 1992, Tables 3 and 4.

Sorghum also produced less biomass and fewer seeds at lower nutrient
levels. Recall that this species produces only one inflorescence per plant,
and the number of seeds per inflorescence responded strongly to nutrient
level. Thus, the total number of seeds produced per individual varies with
nutrient level, as it did for Johnson grass. Approximately 0.30 to 0.40 of
total biomass was allocated to seeds in sorghum. Biomass allocation to
reproductive structures was higher, however, when nutrient levels were
high as compared to low or medium. Even though it varied among nutri-
ent levels, that proportion was similar to the proportion of biomass allo-
cated to rhizomes in Johnson grass.
Sorghum showed a different nutrient allocation strategy in response
to changing nutrient levels. Nutrient concentrations were relatively
22 ORGANISMAL HOMEOSTASIS

constant in seeds regardless of nutrient level, although potassium was

lower at medium and low nutrient levels. The proportion of nutrients al-
located to seeds in sorghum was similar to the proportion allocated to
rhizomes in Johnson grass. The proportion of nitrogen and phosphorus
allocated to Johnson grass rhizomes and sorghum seeds both increased
with decreasing nutrient supply. This suggests that, as with rhizomes in
Johnson grass, seeds are a priority for allocating resources. With no alter-
native means of reproduction, the strategy of sorghum is to produce seeds
that have a certain nutrient composition; and as nutrient levels increase,
they devote more resources to producing more mass and more seeds.
In terms of allocation of limited resources, Johnson grass and sor-
ghum have priorities for vegetative reproduction and sexual reproduction,
respectively. By allocating sufficient resources for offspring to survive, par-
ent plants increase the chances of success of offspring, whether they are
produced sexually or asexually. Seed reproduction in Johnson grass might
be an alternative means of reproduction to increase genetic diversity and
allow for long-distance dispersal, but only when sufficient resources are
available. Individuals allocate limited resources to reproduction, whether
sexual or asexual, sometimes at the expense of growth, and this maintains
homeostasis at the population level. There are often trade-offs and con-
straints in the allocation of energy and nutrients; energy or nutrients al-
located to survival or growth are not available for reproduction, and this
may affect population growth.
Animals should also be subject to the principle of allocation. Despite
their mobility, their food may still be limited and animals must still allo-
cate resources among the physiological demands of survival, growth, and
reproduction. The principle of allocation states that trade-offs among
competing demands arise when organisms can process only a limited
amount of resources, even if organisms could increase consumption rates
when resource availability changes. Organisms that could compensate for
changes in resource availability could allocate resources to both growth
and reproduction.
David Rollo and Mark Hawryluk performed an experiment to exam-
ine trade-offs and ability to compensate for altered resource availability in
two species of freshwater snail, the marsh pondsnail Stagnicola elodes and
the tadpole physa Physella gyrina (Rollo and Hawryluk, 1988). The pondsnail
 AN INDIVIDUALS FORAGING CAN AFFECT THE POPULATION 23

is about twice as big as the physa, but the species have similar habitat
and food requirements. These snail species mostly feed on detritus and
microbes that grow on underwater substrates, but sometimes they also
feed on animal carcasses.
Rollo and Hawryluk collected snails from a pond after the breeding
season. The scientists maintained the snails in aquaria with warm tem-
peratures and 13.5 hours of light per day, which were conditions pre-
dicted to stimulate breeding. The snails were fed a diet of fish food and
agar, a jello-like material derived from marine algae that the snails ate and
also helped hold the fish food together. The scientists ground up the fish
food, mixed it with agar, and added hot water. When cooled, the agar had
solidified and the researchers could cut off consistently-sized pieces. After
2 weeks, reproduction was stimulated in both species and the experiment
began.
Rollo and Hawryluk selected 90 snails of each species and randomly
assigned them to one of three diets. Sets of ten snails were placed in large
glass dishes filled with water. While Fenner and his colleagues manipu-
lated nutrients, Rollo and Hawryluk manipulated protein content by pre-
paring three diets with different ratios of protein-rich nutrients (the fish
food) to low-quality filler (agar). Thus, snails were fed one of three diets:
1) high quality diet with fish food and agar, 2) medium quality diet with
50% of the fish food, and 3) low quality diet with 25% of the fish food.
The scientists provided food several times in excess of consumption to
each treatment.
Growth was measured weekly as shell length and wet mass, which was
measured after blotting off excess water. Growth rates were determined
using regression analysis with the slope used as the estimate of change in
size per unit time (Table 1). Every 3 days the researchers collected egg
clutches (sets of eggs laid by a female) in the dishes and recorded the
number of clutches and number of eggs per clutch (Figure 6). The experi-
ment was maintained for 35 days.
Growth rates as measured by shell length declined for each species as
protein in the diet decreased, but the decline in growth rate was much
more dramatic for the physa than for the pondsnail. The same trend was
evident in body mass growth rate estimates for the physa, but pondsnail
growth rate in the medium was slightly higher than in the high protein
24 ORGANISMAL HOMEOSTASIS

Table 1 Slopes of growth rates of two species of snails fed diets

containing different amounts of protein.
protein content shell growth body wet mass
species
in diet rate (m/day) growth rate (mg/day)
high 27.8 1.82
marsh pondsnail medium 23.3 2.14
low 14.4 1.41
high 28.1 1.11
tadpole physa medium 18.2 0.86
low 3.8 0.30
Source: From Rollo and Hawryluk, 1988, Tables 3 and 4.

120 clutches 40 eggs/clutch

= marsh pondsnail
total number of clutches

100 = tadpole physa

mean eggs/clutch

30
80

60 20

40
10
20

0 0
high medium low high medium low
A protein content B protein content
Figure 6 Reproduction in snails fed different protein content diets.
A, Mean total number of clutches produced. B, Mean number of
eggs per clutch for each species of snail.
Source: From Rollo and Hawryluk, 1988, Table 2, Figure 2.

diet. Rollo and Hawryluk predicted that if the protein content was halved
or quartered, growth rates would also be halved or quartered, respectively.
That did not happen for either species of snail, except for the physa on
low protein, where their body mass growth rate was very close to the ex-
pectation (27% of high protein). This suggests that both species are com-
pensating for lowered protein.
Physas produced many more eggs per gram of snail body mass than
pondsnails. Both species responded strongly to protein content in their
diets; and as protein decreased, so did the total number of clutches pro-
duced in the experiment, although only physas on the medium protein
diet and pondsnails on the low protein diet showed the expected 50%
and 25% drop, respectively, in number of clutches relative to the high
 AN INDIVIDUALS FORAGING CAN AFFECT THE POPULATION 25

0.5 consumption 0.18 assimilation

0.16

(mg/mg dry mass/day)

= marsh pondsnail
mass allocation 0.4 = tadpole physa 0.14
0.12
0.3
0.10
0.08
0.2
0.06
0.1 0.04
0.02
0 0
high medium low high medium low
A protein content B protein content
Figure 7 Consumption, assimilation and allocation in two snails
fed different protein content diets. A, Mass of food consumed
per milligram of snail dry mass per day. B, Assimilation, or
consumptionfecal matter.
Source: From Rollo and Hawryluk, 1988, Table 1.

protein diet. The number of eggs per clutch declined slightly for physas
and increased slightly for marsh pondsnails.
During days 20 through 22, Rollo and Hawryluk estimated mass
budgets of snails, which is a concept similar to energy budgets. Mass
budgets are itemized summaries of matter consumption and allocation.
On day 20, each snail was weighed, and each population of ten snails
was placed into a clean dish. Pre-weighed pieces of the appropriate diet
were placed into each dish. At the end of day 22, the scientists removed
any remaining food and dried it to remove water. They estimated initial
dry mass of food by drying a set of fresh food pieces that were not given
to snails. In this way, the researchers estimated consumption of dry
mass of food (Figure 7A). Snails were reweighed; egg clutches counted,
dried, and weighed; and feces collected by filtering the water in the dish
through filter paper and drying the collected feces. Ten snails of each
species from high quality diet dishes were separated from their shells.
Shells and bodies were weighed and dried to obtain dry mass and mass
gain. Accounting for consumption and feces production allowed the
scientists to estimate assimilation of food (Figure 7B), and all of the
components of growth and reproduction were used to estimate mass
budgets. Assimilation is the process of incorporating consumed food
into the body after digestion.
26 ORGANISMAL HOMEOSTASIS

Physas ate much more of the high protein diet than the pondsnail,
despite being smaller. Both species showed compensation for lowered
protein in their diets, but the marsh pondsnail increased consumption
much more than the tadpole physa, eating more than expected on the
medium protein diet; very close to the expected four times the amount on
the low protein diet. Physas increased consumption, but it was much less
than expected based on the reduction in protein.
The mass budget analysis of Rollo and Hawryluk revealed that much
of what is consumed is either defecated or used for respiration. Respira-
tion is consumption unaccounted for by feces, growth and reproduction,
and includes matter excreted, used for energy or mucus production. The
analysis indicated that as protein content declined, relative allocation to
growth declined in both species. However, this was offset by the increased
consumption, which caused actual growth rates of pondsnails to be
greater in the medium protein diet than the high protein diet and about
90% of the high protein diet for pondsnails on the low protein diet. Thus,
pondsnails showed a strong ability to maintain growth rate. Physas were less
able to maintain growth rate than pondsnails and were less able to compen-
sate with increased consumption as protein declined. Thus, the pondsnail
had superior compensatory ability than the physa for consumption.
As protein content declined, relative allocation to reproduction also
declined in both species with the exception of the physa on the low pro-
tein diet. Here the scientists found that relative allocation was lower than
in the high protein diet, but it was higher than in the medium. Thus,
physas showed an ability to maintain reproduction despite resource limi-
tation. Rollo and Hawryluk concluded that pondsnails maintained
growth but reduced reproduction, whereas physas maintained reproduction
but reduced growth when protein decreased. Each species has an alloca-
tion strategy that maintains population homeostasis. Larger species, such
as the pondsnail, allocate more resources toward growth; larger individu-
als have greater reproduction later if they invest more resources earlier
into attaining a larger size. Smaller species might evolve a strategy to re-
produce at the expense of growth when resources are limiting. Both species
continued to grow and reproduce, but when resources were limited, there
was a shift in priority to one strategy or another, as discussed for grasses.
Differential support of one function over another during times of stress or
 AN INDIVIDUALS FORAGING CAN AFFECT THE POPULATION 27

resource limitation indicates that a trade-off is occurring between those

functions.
In this chapter, how resources affect homeostasis of populations has
been explored, and a connection has been made between populations and
acquisition of energy and nutrients, which are required for the organiza-
tion of life and the growth of populations. Homeostasis operates at the
population level through behaviors and actions of individuals. Each spe-
cies has a particular resource allocation strategy that has evolved to work
in a particular environment. The optimal allocation of resources within
an individual is necessary to maintain homeostasis at the population level.
The principle of allocation dictates that resources used for one func-
tion cannot be used for another, and perhaps readers can use this principle
to analyze their own allocation decisions. Although allocation strategies
may vary among species, depending on factors such as size, lifespan, and
time to maturity, all organisms may face constraints that cause them to
trade-off investment in one function to put resources into another.

Ethical, Legal, Social Implications: Negative Birth

Rates in Human Societies Can Have Positive and
NegativeConsequences
There are costs and benefits of population growth policies in countries
like China and India, which are both attempting to slow down the rate of
growth of their populations. Populations growing exponentially may ex-
ceed available resources, thus disrupting population homeostasis. China
is doing a better job of slowing its population growth rate than India, but
this comes at the expense of individual reproductive rights, although
China has recently relaxed even further its one child per family policy.
China, the United States, and an estimated 27 other, mostly developed,
countries have reduced fertility to replacement level or just below. Re-
placement level fertility in developed countries is 2.1 children per couple
(the 0.1 accounts for infant mortality). But because there are high pro-
portions of young people that have yet to move into their reproductive
years, these populations are still expanding. All else being equal, these
countries will achieve a stable population size or even begin to shrink
once these young people age beyond their high fertility years. What would
28 ORGANISMAL HOMEOSTASIS

happen to China if the average fertility rate were actually one child per
couple, as the Chinese government dictates? The growth rate of the popu-
lation would be negative, and one can examine the situation in some
other countries to gain perspective about what happens to countries in
such a situation.
Population growth estimates made by the United Nations identify
two groups of countries where populations are either already shrinking or
are projected to shrink soon. In one group, an estimated 33 countries
(mostly in Europe but also including Japan and Russia) populations are
either already stable or slowly shrinking due to declining birth rates. The
other group comprises countries with populations declining due to rising
death rate, and two countries in southern Africa belong to that group.
Although it may seem beneficial to achieve zero growth rate given the
demands on the worlds resources and thus achieve population homeo
stasis, an age structure that has changed due to changing birth or death
rates may lead to severe adverse effects on society.
Consider Japan, which has a rapidly aging population. As Japan de-
veloped economically, young people entered the workforce, married later
in life, and had fewer children. Birth rates declined, health improved, and
people lived longer. The age structure became inverted and is projected to
become even further inverted in the next 50 years. On the plus side, the
Japanese people have high life expectancy, but there are significant costs
to a society with a high proportion of the population over the age of 65
with healthcare needs, costs of pensions, and long periods of retirement.
Support of the elderly had been a strong traditional value in Japanese
society, but there has been a decline in the last few decades of parents liv-
ing with adult children. The Japanese government is now concerned that
society will have to take care of seniors who are unable to support them-
selves or who have no family to support them. This will be difficult as the
proportion of the working population shrinks and the retired proportion
grows. The government encourages employers to provide more employment
opportunities for the elderly at the same time that it is trying to raise the
eligible age for public pensions. This is potentially costly for employers
who retain long-time employees earning high wages, and thus costs are
shifted from society to the employer. The elderly may not even want or be
able to continue working.
 AN INDIVIDUALS FORAGING CAN AFFECT THE POPULATION 29

The government is also attempting to alter the decline in birth rates

by creating conditions and incentives for families to give birth and rear
children. Any effects these changes have on age structure, however, will
not be seen for decades.
The Population Reference Bureau in 2008 identified two countries in
southern Africa, Lesotho and Swaziland, with high human immunodefi-
ciency virus (HIV) infection rates and widespread hunger, which belong
to the rising death rate group. Each year, millions of people become newly
infected with HIV, and 95% of them are in sub-Saharan Africa, Eastern
Europe, and Asia. In Lesotho, a tiny country completely surrounded by
South Africa, it is estimated that 25% of adults have HIV, and the ac-
quired immune deficiency syndrome (AIDS) epidemic has led to high
rates of mortality and a drop in life expectancy (the number of years an
individual is predicted to live) from 60 to 34 years. However, high fertility
has kept the population from declining until recently.
The number of countries in this unfortunate group could grow as
populations in low income countries continue to fight the HIV pandemic
and other density-dependent effects. In populations suffering from high
HIV infection rates, AIDS-related deaths alter the age structure because
the highest mortality rates occur among young and middle-aged adults,
rather than infants or the elderly. When parents and breadwinners are
affected in such a manner, there are serious adverse consequences for fam-
ilies and national economies. Families suffer because of loss of income,
healthcare costs, and loss of parents. Farms, schools, industry, healthcare
systems, and governments all suffer from loss of workers, absenteeism, and
payment of healthcare and death benefits. Loss of agricultural production
is also being caused by high HIV infection rates, and this has serious con-
sequences for a countrys ability to feed its people.
There are tangible benefits to a shrinking population. Resources per
capita will be higher, or at least not continue to decline, and thus quality
of life should be higher. Longevity may increase, and if the cause of a
shrinking population is lower fertility rates caused by increased empower-
ment and education of women, that may benefit and advance the society
as a whole. Other benefits may include less stress and strain on the environ-
mental support systems that provide us with ecosystem services. Fewer young
men may lead to lower unemployment and a lower crime rate. It also means
30 ORGANISMAL HOMEOSTASIS

less traffic, pollution, and less crowded highways. Yet a rapidly shrinking
population could mean trouble as indicated above. Policies may be imple-
mented to reduce population in a controlled and slower manner and
prepare for a rising population of older people, to avoid costs of rapid
shrinkage.

Bibliography
Benech Arnold RL, Fenner M, Edwards PJ: Mineral allocation to repro-
duction in Sorghum bicolor and Sorghum halepense in relation to pa-
rental nutrient supply, Oecologia 92(1):138144, 1992.
Brown LR: Plan B 4.0: Mobilizing to save civilization, ed 1, New York,
2009, W. W. Norton & Company.
Martin LG: The graying of Japan, Popul Bull 44(2):142, 1989.
Ministry of Education, Culture, Sports, Science and TechnologyJapan:
Challenge for building a future society: the role of science and tech-
nology in an aging society with fewer children. Part 1.1 Current status
of an aging society with fewer children and challenges for science and
technology, MEXT (website): http://www.mext.go.jp/english/news/
2007/03/07022214/001/003.htm. Accessed August 12, 2010.
Population Reference Bureau (website): http://www.prb.org/. Accessed
August 9, 2010.
Rollo CD, Hawryluk MD: Compensatory scope and resource allocation
in two species of aquatic snails, Ecology 69(1):146156, 1988.
United Nations, Department of Economic and Social Affairs, Population
Division: World population prospects: the 2010 revision: Volume I:
comprehensive tables, 2011, (website): http://esa.un.org/wpp/
documentation/pdf/WPP2010_Volume-I_Comprehensive-Tables.pdf.
Accessed August 1, 2014.
 Conclusion
As with any biological system, when the environment changes, the system
responds. In this book, homeostasis at the organismal level was examined,
although many connections to the population level were made. Mammals
expend energy to maintain a consistent body temperature. They use feed-
back mechanisms to keep their body temperatures within a narrow range.
Natural selection has provided different species with unique solutions,
such as the camels hump of fat, the arctic foxs seasonal dimorphic fur,
and the horses skull morphology. Survival, growth, and reproduction all
require energy, and organisms make decisions about how to allocate their
resources, which affects populations. Homeostasis mechanisms are inter-
related and constantly adjust to changes in the environment.
 Glossary
ambient temperature. Temperature of the external environment.
ancestral species. An older species that existed in the past and serves as an ances-
tor to a more recently living species.
assimilation. Assimilation is the process of absorbing nutrients and incorporating
them into the body.
basal metabolic rate. The rate at which an organism uses energy while at rest to
keep vital functions going, including breathing and keeping warm.
biomass. The total amount of organic matter in an organism, population, or other
ecological system after water is removed.
cavernous sinus. A small air pocket between the nasal cavity and brain of some
mammals.
core body temperature. The temperature of the internal environment of an ani-
mal, but not of appendages, such as legs.
ectotherms. Ectotherms are animals that regulate their body temperatures using
behavioral mechanisms and heat from the environment.
endotherms. Endotherms are animals (specifically birds and mammals) that
maintain a consistent internal body temperature through physiological processes.
evaporative heat loss. Heat lost by sweating or otherwise evaporating water from
the surface of a body.
fertility. The quality of an organisms ability to produce offspring, which is depen-
dent on age, health, and other factors.
hypothalamus. A region of the brain below the thalamus that coordinates both
the autonomic nervous system and the activity of the pituitary, controlling body
temperature, thirst, hunger, and other functions.
inflorescences. A group or cluster of flowers arranged on a stem composed of a
main branch or a complicated arrangement of branches.
tillers. A shoot that springs from the root or bottom of a primary stalk.
life expectancy. The number of years an individual of a certain age is expected to live.
mass budgets. Itemized summaries of inputs and outputs of matter, including
consumption and allocation
phenotypes. The ways an organism appears or behaves due to its genetic makeup
and environmental influences.
principle of allocation. The principle of allocation states that natural selection
results in organisms optimizing partitioning of resources to maximize fitness.
rhizomes. Horizontal subterranean stems distinguished from true roots in
possessing buds, nodes, and scale-like leaves.
seasonal dimorphism. Seasonal dimorphism describes organisms with two pheno
types depending on the time of year.
 Index
Acquired immune deficiency
syndrome (AIDS), 29
Agar, 23
Alopex lagopus, 2
Ambient temperature, 1
Ancestral species,
of mammals, 1
Assimilation, 25
Basal metabolic rate, 11
Beery, Annaliese, 13
Behavioral biology, 13
Biomedical research, 14
Body temperature, xiii, 114
Camelus dromedaries, 4
Cavernous sinus, 9, 10
Clegg, Deborah, 13
Ectotherms, 2
Endotherms, 1, 2, 1112
Energy and nutrient
resources, xiii
Energy flow, xiii
Evaporative heat loss, 11
Fat metabolism, 6
Feedback mechanisms, 12
Fenner, Michael, 18
Gender differences, 1214
Hawryluk, Mark, 2223, 26
Homeostasis, 2
animal temperature, study of, 4
Human immunodeficiency virus
(HIV), infection rates, 29
Hypothalamus, 711
Inflorescence, 18
Johnson grass (Sorghum halepense), 18
concentration of nutrients in, 21
nutrient level effects on, 19
seed reproduction in, 22
Mammals, 114
body temperature of, 1
ancestral species of, 1
placental, origin of, 2
functional hypothalamus, lacking, 8
Mass budgets, 25
Phenotypes, 2
Physella gyrina, 22
Population growth, 28
Population Reference Bureau, 29
Principle of allocation, 17, 22
Pulmonary artery, 9
Rapid breathing, 10
Rectum, 9, 10
Reproductive biology, 13
Respiration, 26
Rhizomes, 18
Rollo, David, 2223, 26
Schmidt-Nielsen, Knut, 4
Seasonal dimorphism, 2
Sorghum (S. bicolor), 18
concentration of nutrients in, 21
nutrient level effects on, 20
Stagnicola elodes, 22
Summers, Larry, 12
Thermal regulation, 12
Tiller, 18
Voskuhl, Rhonda, 13
Zucker, Irving, 13
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