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Organismal Homeostasis
10 9 8 7 6 5 4 3 2 1
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
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
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
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
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.
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.
MAMMALS POSSESS ADAPTATIONS TO MAINTAIN BODY TEMPERATURE 15
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