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Organismal Homeostasis

Organismal Homeostasis

Christopher J. Paradise, PhD

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

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First published in 2016 by

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ISBN-13: 978-1-60650-973-9 (print)

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Printed in the United States of America.

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.

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
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
Ethical, Legal, Social Implications: Negative Birth
Rates in Human Societies Can Have Positive and
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 where they will find pedagogically-
designed and interactive Integrating Concepts in Biology for introductory
biology college courses or a high school AP Biology course.
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.
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.

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

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.

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
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

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.

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.,
Author: LadyOfHats. Public domain. B, From http://commons.
File:Bactrian_Camel,,Author: Stanley Howe.
This file is licensed under the Creative Commons Attribution-Share Alike 2.0 Generic license.

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
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

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

temperature change from average (F)

0 = 101.4 F
70 75 80 85 90 95

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.

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

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

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

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

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

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

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.
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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.

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Acquired immune deficiency Johnson grass (Sorghum halepense), 18
syndrome (AIDS), 29 concentration of nutrients in, 21
Agar, 23 nutrient level effects on, 19
Alopex lagopus, 2 seed reproduction in, 22
Ambient temperature, 1
Ancestral species, Mammals, 114
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Assimilation, 25 ancestral species of, 1
placental, origin of, 2
Basal metabolic rate, 11 functional hypothalamus, lacking, 8
Beery, Annaliese, 13 Mass budgets, 25
Behavioral biology, 13
Biomedical research, 14 Phenotypes, 2
Body temperature, xiii, 114 Physella gyrina, 22
Population growth, 28
Camelus dromedaries, 4 Population Reference Bureau, 29
Cavernous sinus, 9, 10 Principle of allocation, 17, 22
Clegg, Deborah, 13 Pulmonary artery, 9

Ectotherms, 2 Rapid breathing, 10

Endotherms, 1, 2, 1112 Rectum, 9, 10
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Energy flow, xiii Rhizomes, 18
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Fat metabolism, 6 Schmidt-Nielsen, Knut, 4

Feedback mechanisms, 12 Seasonal dimorphism, 2
Fenner, Michael, 18 Sorghum (S. bicolor), 18
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Gender differences, 1214 nutrient level effects on, 20
Stagnicola elodes, 22
Hawryluk, Mark, 2223, 26 Summers, Larry, 12
Homeostasis, 2
animal temperature, study of, 4 Thermal regulation, 12
Human immunodeficiency virus Tiller, 18
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Hypothalamus, 711 Voskuhl, Rhonda, 13

Inflorescence, 18 Zucker, Irving, 13


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