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EBOOKS Emergent Properties of Individual
FOR THE Organisms

APPLIED BIOLOGY COLLECTION
Christopher J. Paradise A. Malcolm Campbell
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LIBRARY This book begins by describing what an individual organism is,
comparing preconceptions of the individual to non-standard
Create your own ways of thinking about individuals. Variation in what individuals
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hives as examples. Individuals are thus shown to be emergent
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properties. Other emergent properties of individuals are also
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described. Classic experiments that elucidated the source of
the greater your
Properties
emotions in humans and other mammals are described. Emo-
discount! tions arise from the actions of the nervous and endocrine system
and often include a variety of signals given to other individuals of
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fear and anger, two emotions that are closely related and often
confused, but that have been well studied. In one final example
of Individual
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of emergent properties of individuals, cooperative behavior is
analyzed. The behaviors displayed by individuals that facilitate
cooperation among individuals and why those individuals may
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Christopher J. Paradiseis professor of biology and environ-
mental studies at Davidson College. He teaches introductory
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A. Malcolm Campbell
Emergent Properties
of Individual Organisms
Emergent Properties
of Individual Organisms
Christopher J. Paradise, PhD

A. Malcolm Campbell, PhD
Emergent Properties of Individual Organisms
Copyright Christopher J. Paradise and A. Malcolm Campbell. 2016.
All rights reserved. No part of this publication may be reproduced, stored

in a retrieval system, or transmitted in any form or by any means
electronic, mechanical, photocopy, recording, or any other except for
brief quotations, not to exceed 250 words, without the prior permission
of the publisher.
First published in 2016 by

Momentum Press, LLC
222 East 46th Street, New York, NY 10017
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ISBN-13: 978-1-60650-963-0 (print)

ISBN-13: 978-1-60650-964-7 (e-book)
Momentum Press Biology Collection
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Printed in the United States of America

Abstract
This book begins by describing what an individual organism is, compar-
ing preconceptions of the individual to non-standard ways of thinking
about individuals. Variation in what individuals are is described, using
giant fungi, clonal trees and honey bee hives as examples. Individuals
are thus shown to be emergent properties. Other emergent properties
of individuals are also described. Classic experiments that elucidated the
source of emotions in humans and other mammals are described. Emo-
tions arise from the actions of the nervous and endocrine system and
often include a variety of signals given to other individuals of the same or
different species. In particular, this book focuses on fear and anger, two
emotions that are closely related and often confused, but that have been
well studied. In one final example of emergent properties of individuals,
cooperative behavior is analyzed. The behaviors displayed by individuals
that facilitate cooperation among individuals and why those individuals
may actually cooperate instead of compete when acquiring resources or
defending against predators are discussed.
Keywords
fungus, superorganism, behaviors, natural selection, fight-or-flight, divi-
sion of labor, dominance hierarchy, subordinate, fear, anger, individual,
colony, physiological response, endocrine system, nervous system, coop-
eration, mutual benefit
Contents
Preface...................................................................................................ix
Acknowledgments....................................................................................xi
Introduction.........................................................................................xiii
Chapter 1 The Definition of the Individual Can Be Stretched
Beyond the Classic Definition1
Chapter 2 The Nervous and Endocrine Systems Are the Sources
of Emotions.....................................................................17
Ethical, Legal, Social Implications: There Are Issues
With Using Prescription Drugs to Normalize
Behavior in Children26
Chapter 3 Individuals of Some Species Cooperate With Each
Other for Mutual Benefit.................................................29
Conclusion............................................................................................43
Glossary................................................................................................45
Index....................................................................................................47
Preface
This book about emergent properties of individual organisms 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 presented. Reading any of the thirty books by Paradise
and Campbell provides readers with biological context and comprehen-
sive 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: informa-
tion, 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 emergent properties of individu-
als 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 emergent properties of individuals the
way professional scientists dowith experimentation and data analysis.
In short, data are put back into the teaching of biological sciences.
Readers of this book who wish to see the textbook version of this
content can go to www.bio.davidson.edu/icb where they will find
pedagogically-designed and interactive Integrating Concepts in Biology
for introductory biology college courses or a high school AP Biology
course.
Acknowledgments
Publishing this book would not have been possible without the generous
gift of Dr. David Botstein who shared some of his Breakthrough Prize
with co-author AMC. Davids gift allowed us to hire talented artists (Tom
Webster and his staff at Lineworks, Inc.) and copyeditor Laura Loveall.
Thanks go to Kristen Mandava of Mandava Editorial Services for project
management and guidance. In particular, we are indebted to Katie Noble
and Melissa Hayban for their many hours and attention to detail.
Kristen Eshleman, Paul Brantley, Bill Hatfield and Olivia Booker
helped us with technology at Davidson College. We are grateful to ad-
ministrators Tom Ross, Clark Ross, Carol Quillen, Wendy Raymond,
Verna Case, and Barbara Lom who had confidence in us and encouraged
us to persist despite setbacks along the way.
Thanks to my wife Amy Brooks for her constant support during the
development of this textbook, and my daughter Evelyn for her endless
energy. Thanks to Malcolm Campbell for his steadfast resolve and opti-
mism. Without him, this book would not exist. Thanks to collaborator
Laurie Heyer for taking my sometimes half-baked math ideas and turn-
ing them into powerful and elegant Bio-Math Explorations. I learned a
lot from both of them. While the math is largely absent from this book,
our collaboration with her made this a better book. Nancy Stamp at
Binghamton University, and Bill Dunson and Richard Cyr at The
Pennsylvania State University influenced me greatly in how I think as
a scientist and approach my teaching. Finally, I thank my students in
Integrated Concepts in Biology II, who enthusiastically participated in
our experiment to redesign introductory biology, starting with the text
and ending with a new approach to teaching biology.
Introduction
Ren Descartess famous quote, I think, therefore I am, is a powerful
statement on emergent properties. A human exists because they have
consciousness and can think. The human brain can be described in terms
of cellular functions and biochemical reactions. Yet describing the brain
does not explain the emergent property of human consciousness. Brains
perform at a level that exceeds the functions of individual neurons. Truly,
who could have predicted that consciousness would arise from all the
components that make up the human brain? Many other aspects of in-
dividuals, including the individual itself, can be considered emergent
properties.
In this book, several ideas about properties of individuals that lead
to emergence of new and unpredictable properties will be explored. In
the first chapter, in line with Descartes, the definition of the individual
will be contemplated, especially in cases when the lines of individuals
are blurred. The second chapter addresses a topic very familiar in daily
life but rarely studied in biology classes. Data will be used to help un-
derstand the biological origins of emotions. The final chapter explores
another topic debated by biologists, which is the conditions under which
organisms cooperate with each other.
CHAPTER 1
The Definition of the

Individual Can Be Stretched
Beyond the Classic
Definition
A commonly held understanding of an individual organism may be clear:

All of the cells of mammals, for instance, define self by displaying pep-
tide fragments from every protein produced inside all of the cells of an
individual. We define an individual as a set of cells that display the same
message regarding what is the self. But we need a more general defini-
tion to apply to a wide range of organisms. As commonly used, the term
individual refers to an object that is indivisible (that is, a singular thing
separate from other singular things). All of the parts within the boundar-
ies of this singular object are indivisible. Multicellular organisms are indi-
viduals if the cells, tissues, and organs that make up the organism cannot
survive on their own. There may also be a genetic requirement in that an
individuals cells are all composed of the same genome.
Questions about what an individual organism is have been debated
by biologists for years. Considered from the standpoint of an animal,
it seems quite clear that an individual is composed of all the cells, tis-
sues, and organs contained within the epidermis of that animal. However,
there may be biological entities whose status as individual organisms is
far from obvious, making the question not so easy to answer. This may
challenge ones conception of just what is an individual. It also illuminates
the emergent property of the individual arising from a collection of cells.
James Anderson and his colleagues studied several species of fungus in
Michigan (Smith et al., 1992). One species is called Armillaria gallica, or
2 EMERGENT PROPERTIES OF INDIVIDUAL ORGANISMS
the honey mushroom. One of their goals was to determine the size lim-
its of individuals. This species of fungus, like many others, is a decom-
poser and survives on wood. Anderson and his colleagues sampled for the
fungus in a forest. To understand the results of the study, it is helpful to
know a few things about fungi. Many fungi live in the soil and are only
noticed when they produce fruiting bodies, also known as mushrooms
(Figure 1A). Mushrooms are the enlarged aboveground fleshy fruiting body
of certain types of fungi. The part that lives underground is composed of
microscopic thread-like structures called hyphae (Figure 1B). The branch-
ing growth pattern of hyphae leads to development of a mycelium, which
is an interconnected network of hyphae. An important aspect of the life
Figure 1 Fungi. A, Armillaria gallica mushrooms. B, Fungal hyphae.

Source: A, http://en.wikipedia.org/wiki/Armillaria_gallica#mediaviewer/File:Armillaria_
gallica_26659.jpg, Author: Dan Molter. This file is licensed under a Creative Commons 3.0
attribution license. B, http://en.wikipedia.org/wiki/Mycelium#mediaviewer/File:Hyphae.JPG,
Author: TheAlphaWolf. This file is licensed under the Creative Commons Attribution-Share
Alike 3.0 Unported license.
THE DEFINITION OF THE INDIVIDUAL CAN BE STRETCHED 3
cycle of Armillaria species is the formation of rhizomorphs, aggregations of

hyphae that transport resources and materials.
Fungi can reproduce sexually, but most fungi do not have readily
recognizable males and females. Basidiomycota, the group to which
Armillaria belongs, have haploid mycelia. Many fungi also have self-
incompatibility. The honey mushroom has two self-incompatibility
loci. Two individuals are reproductively compatible only if they have
different alleles at each self-incompatibility locus. If the mycelia of two
individuals are compatible, they fuse and swap nuclei. Two compatible
nuclei remain in pairs for a time. When nuclei do fuse a diploid cell is
formed, which quickly undergoes meiosis to produce four haploid nu-
clei that each migrate into a structure called a basidiospore, which are
often produced on mushrooms. The resulting haploid mycelia are called
mating types. Fungi can also reproduce asexually, through vegetative
growth of mycelia.
Anderson and his colleagues placed wooden tongue depressors in
the forest soil along two 1 kilometer transects, one oriented north/south
and the other east/west. Hyphae of the wood-decomposing fungi quickly
colonized these sticks. After a period of time, the researchers collected the
tongue depressors and cultured the fungi. In addition, they made cultures
from mushrooms and rhizomorphs that they collected from soil pits and
mycelia they collected from under the bark of trees. The scientists used
several methods to discriminate between honey mushroom individuals.
The scientists determined the self-incompatibility alleles of fungal
cultures. They also determined the patterns of DNA fragments that were
produced when mitochondrial DNA was chopped up using restriction
enzymes to produce fragments of varying length that were then separated
on gels. The self-incompatibility loci and mitochondrial DNA loci are
known to have multiple alleles in the honey mushroom. The researchers
reasoned that if the collected fungi were all from the same individual,
they would only see four mating types in haploid mycelia and mush-
room spores. This pattern would also be possible in an inbred population,
but in that case, the restriction enzyme data would show a low degree of
variation in the mitochondrial DNA. If more than two alleles were found
at a single locus among the fungal samples, then that would also indi-
cate more than one individual was present. Anderson and colleagues also
used random amplification of polymorphic DNA (RAPD) to look for

heterozygosity in nuclear genetic loci. In RAPD, segments of DNA are
randomly amplified using polymerase chain reaction (PCR) primers. The
scientists used three primers, and each amplified more than one fragment
of DNA, which was either present or absent in different haploid cultures.
Again, a high degree of similarity among haploid cultures sampled from
different locations would be indicative of inbreeding, as opposed to a
single individual.
Anderson and his colleagues found only two alleles in each of five
loci digested by restriction enzymes in all fungal samples from one large
area of the forest, which they denoted as individual 1. Each of the fungal
samples from this same area contained all of the 11 RAPD sequences,
although not all 11 were evident in each haploid sample. Twenty more
RAPD and 27 RFLP from samples of diploid tissues in the individual 1
area were tested further and found to be identical across samples. Samples
from what the scientists denoted as individual 2 were found to have
only one allele at each of two restriction enzyme loci and lacked five of
the RAPD fragments found in individual 1. From the distribution of
the samples that showed similar genetic structures from multiple analy-
ses, Anderson and colleagues estimated the boundaries of these different
individuals.
One individual fungus can produce only four possible mating types.
Because there are two loci and each is always heterozygous, meiosis pro-
duces four haploid cells from one diploid cell and each contains one of
two alleles at each locus. However, with more than two alleles, many
compatible mating types are possible. Anderson and colleagues only
found two alleles at each self-incompatibility locus from samples over a
wide area, leading to the possibility that these samples were from the same
individual. The RFLP and RAPD data helped support the conclusion
that these samples were all from the same individual. Only two alleles
were found in each restriction enzyme locus in all samples found within
the individual 1 boundary, further supporting the conclusion that only
one individual was present.
The genetic signature of samples from the individual 2 area were dif-
ferent from those of individual 1, which allowed Anderson and colleagues
to clearly demarcate the southwestern boundary of individual 1. Had they
sampled further north and east, they might have discovered the full extent
of individual 1s boundary. Although Anderson and colleagues discov-
ered several large A. gallica individuals, the largest was estimated to cover
15 hectares. All of the fungal samples collected from this area had the
same mating type, similar mitochondrial DNA restriction patterns, even
if individual haploid cultures were not exactly the same, and the same
11 RAPD fragments. Because of the variation among individual haploid
cultures, the researchers knew that each of these loci were heterozygous in
the diploid condition, allowing them to reject the hypothesis that these
samples were from different but closely related individuals.
Anderson and colleagues next set out to estimate the biomass and
age of the largest individual, individual 1. To estimate age, the scientists
had to estimate the rate of expansive growth outward from the center.
They assumed that the rate of growth was constant, which allowed them
to back-calculate to determine how long it took individual 1 to expand
to its current size. They used three methods to estimate the growth rate
(Table 1). Anderson and colleagues measured the maximum linear growth
of rhizomorphs along wooden stakes during one growing season. The sci-
entists also measured the maximum growth rate of cultures in the labora-
tory at different temperatures. To estimate mass, the scientists collected
soil samples, carefully separated fungal rhizomorphs from roots and soil,
dried the material, and weighed it to obtain the biomass per unit area of
soil. The wet mass was also estimated.
Anderson and colleagues knew that soil temperatures in the forest
ranged from 14o C to 16o C for about 90 days per year; and the rest of
Table 1 Estimates of growth and size of large

A. gallica individual fungus.
method maximum growth rate
wooden stakes in forest soil
19.7 3.1 cm/year
(average of ten longest out of 45)
laboratory cultures at 12C 0.0 mm/day
biomass (dry mass) 6.25 g/m2
wet mass 64.4 g/m2
Source: From Smith et al., 1992, Figure 1 legend.

the year, temperatures were below 14o C. They then calculated the an-
nual growth rate of laboratory cultures using the daily growth at 15o C
for 90 days, which is 0.189 m/year (2.1 mm/day * 90 days * 1 m/100 mm).
This is quite comparable to the annual growth on wooden stakes in the
forest soil (0.197 m/year). If this individual fungus grows in all direc-
tions at the same rate from a central point, it would be circular, and the
maximum distance across would be the diameter of the circle. The ra-
dius is half of that, which is the maximum distance it grew during its
life. If it grew steadily at 0.193 m/year (the average of the two estimates
made by the scientists), then this fungus is 1,645 years old [(635 m/2)/
(0.193 m/year)]! Anderson and colleagues argued that this may actually
be an underestimate of the age of the fungus, because competition with
other honey mushroom individuals would slow growth, as would adverse
climate conditions (such as, cold growing seasons).
The wet mass of the honey mushroom in the Michigan forest was
calculated based on the average wet mass estimate of 64.4 g/m2. First
convert the area to hectares, using the conversion of 10,000 m2/hectare.
Second, convert grams to kilograms by dividing by 1,000. Finally, multi-
ply by 15 hectares, which is the estimated area occupied by individual 1,
to obtain 9,660 kg. That is 9.66 metric tons, and when Anderson and
colleagues multiplied that by 10 to obtain the estimated total fungal mass,
they obtained 96.6 metric tons, which is close to the mass of an adult blue
whale! This may well alter perception of what an individual is. This one
individual fungus likely grew vegetatively for centuries to become a very
old and large individual.
Not all fungi grow to this size; in fact, most are much smaller and
often multiple individuals of the same species live in the same soil, un-
like the situation with the honey mushroom. But individuals of other
species of fungi have been shown to achieve sizes of the magnitude of the
honey mushroom individual, so it is not uncommon. A phenomenon
about fungal growth is that it is dynamican individual can lose mass
in one area and regrow it later. Fungal tissue is not as highly integrated as
the tissues of a blue whale. The property of the individual emerges from
the hyphae, mycelia, and rhizomorphs that make up the individual, and
in this instance that individual occupies space that no other honey mush-
room can occupy.
Some scientists have questioned whether this enormous individual

that Anderson and colleagues found is actually an individual. They
claim that more empirical information is necessary, including whether
or not rhizomorphs in one part of the forest are actually connected to
rhizomorphs in another part of the forest. This may reveal something
about the concept of the individual; if the fungus is an individual, then
the boundaries of an individual do not necessarily have to be contained
within a single outer layer that we can perceive. The property of being
an individual emerges as more than the sum of the parts. Indeed, when
studying plants ones perception of the boundaries of an individual may
be biased by what is seen aboveground. But the patterns in aboveground
structures may suggest clues. For instance, quaking aspen (Populus tremu-
loides) often grows in distinct stands, with trunks in the middle larger
and taller than trunks towards the exterior. This pattern may result from
growth of a single individual or production of seeds that do not disperse
far from the parent.
Karen Mock and her colleagues tested the hypothesis that the trunks
in stands of quaking aspen belonged to a single individual (DeWoody
etal., 2008). They sampled stems from aspens in an aspen forest in central
Utah (Figure 2). Each of 209 sampled trunks was the closest to the center
point of a square in a grid of 50 m 50 m squares. If no stem was within
5 meters of the center point, no sample was taken in that square. This
sampling scheme allowed them to systematically sample the entire extent
of the forest but not have to sample every tree.
At each trunk, the researchers collected a tissue sample. DNA was
isolated and assayed for seven microsatellite loci (Table 2). Microsatellite
DNA is a repeating sequence of one to six base pairs of DNA. For in-
stance, the sequence cytosine-adenine (CA) may repeat, and the number
of times it repeats varies among different alleles. The high number of al-
leles present at microsatellite loci leads to many possible genotypes within
a species, making them very useful for population genetics studies.
At each locus in each sample, the scientists determined the size of the
repeated microsatellite DNA sequence, which let them identify the allele.
CACACACA could be distinguished from CACA, for instance, based on
size. The genotype of a particular stem is the set of alleles present at the
seven microsatellite loci. The researchers mapped the genotypes on an
Figure 2 Aerial photograph of aspen forest sampled by Mock and

colleagues (circled area).
Source: Image from Google Earth.
Table 2 Genetic analysis of quaking aspen stand in Utah.

Sample size was 209.
variable value
number of distinct genotypes 46
number of samples containing the most common genotype 141
number of samples containing genotypes differing from
5
most common genotype by a single allele
number of samples containing genotypes differing from
63
most common genotype at between 4 and 7 alleles
estimated area covered by most common genotype 43.6 hectares
Source: From De Woody et al., 2008, text.
aerial photograph of the site. One genotype was found in a majority of

samples. The extent of the distribution of this genotype was compared to
a boundary of trunks that had the same phenotype, determined from a
different study.
Mock and her colleagues concluded that there were up to 41 individ-

ual aspen in this forest, which is far fewer than the 209 stems sampled.
Most of the forest was made up of just one individual. Although they did
not sample the entire forest and could not determine the extent of the
other individuals, none of them could be as large as this one, based on
the total size of the aspen forest. The researchers concluded that all 141
trunks with the same genotype plus the five trunks that differed from the
dominant genotype at only one allele belonged to the same individual,
because the probability of seeing identical genotypes in two unrelated
individuals is very small. In the case of microsatellites, it is reasonable
to assume that the researchers picked loci that are on different chromo-
somes, and thus the probability of an individual getting a particular mic-
rosatellite allele is independent of getting a particular allele at a different
genetic locus.
Even if two loci are on the same chromosome, they would try to pick
microsatellites that are separated by enough DNA that the two loci are
rarely inherited together. They also knew that there were more than 2 al-
leles at each locus, up to 20 in some cases. While this makes the determi-
nation of the probability of obtaining the same alleles at each locus more
difficult, if one were to obtain the same alleles at each locus, it would be
highly likely that the two samples would be from the same individual.
If one assumed there were only two alleles at each locus, and each
one occurs with probability , the probability that two individuals have
thesame genotype at a particular locus is the probability of matching on
one chromosome times the probability of matching on the other chromo-
some, which is 1/2 1/2 = 1/4. They then raised that probability to the
power of 7 because there were 7 microsatellite loci tested. This yields a
probability of (1/4)7 6 1025 in the case of two equally likely alleles
at each locus.
However, because the researchers knew there were multiple alleles at
each locus, their probability estimates were much lower than that, on the
order of 1 10210. Because most microsatellites have at least five alleles,
the chance of identical genotypes in two randomly selected unrelated in-
dividuals is extremely small.
Based on a similar probability calculation, Mock and her colleagues
further concluded that the five trunks that differed by one allele were part
of the same individual but had undergone a mutation while growing

vegetatively. The samples that differed by four to seven alleles were from
40 different individuals found in 63 trunks. Some of these individuals
were single stems, but some were made up of several trunks.
Aspen can grow vegetatively by sending up new trunks from roots
that spread out along the ground. They can also reproduce sexually, but
this type of asexual reproduction leads to many trunks that are all identi-
cal to each other genetically and phenotypically. The trunks are all con-
nected by an extensive root system. According to researchers, the large
aspen clone in central Utah is the largest individual organism on the
planet, at least the largest that has been measured. The estimated mass
of this one individual plant is 5,900 metric tons, an estimated 59 times
the mass of the honey mushroom fungus or an adult blue whale! Like the
individual honey mushroom fungus, aspen clones can occupy a large area.
Thousands of trunks are part of this one individual organism, and they
occupy over 100 acres. Multiple trunks of a single individual make that
individual more than just the sum of the parts. The parts can cooperate
together to transduce more energy from photosynthesis and better com-
pete for space and light.
Although the trunks in an aspen individual are all connected by their
roots, there may be parts that are not connected physically and yet may be
considered an individual. This possibility was considered with the honey
mushroom fungus. This may stretch someones conception of the indi-
vidual, yet the emergent property that arises is analogous to that which
was discovered for fungi and plants. A part cannot survive by itself, the
individual is greater than the sum of the parts, and parts create properties
that do not exist within the parts. Consider a species of animal where
individuals live together in a colony. Honey bee (Apis mellifera) colonies
have been studied by many researchers. Honey bees and some other
insects can produce heat metabolically; most insects cannot. Anton
Stabentheiner and his colleagues examined body temperatures of honey
bees and temperature regulation of honey bee colonies under varying
temperatures using infrared thermal imaging (Stabentheiner et al., 2010).
The researchers used three experimental hives covered with a plastic
film transparent to infrared radiation. Each hive had a tube connected
to the hive entrance through which bees came and went. Approximately
150 workers were marked with colored paint that indicated the date and
transferred to experimental hives every other day until colonies contained
4,000 to 5,000 worker bees, which took about 4 weeks. Once a hive was
of a certain age, the queen began producing her own workers, and those
bees were unmarked.
The scientists manipulated temperature by placing hives in a stream
of air at one of five temperatures. The scientists measured air temperature
near bees using thermocouples in various places within the hive, includ-
ing near sealed brood cells (where larvae developed), open brood cells
(where larvae are fed by workers), empty cells, pollen cells (where pollen
is stored), and sealed or open honey cells (where honey is stored).
The scientists measured the head, thorax, and abdomen temperatures
of marked bees of varying ages in different positions in the hive (Figure 3).
The scientists recorded the bees position, its age, its temperature, and the
hive temperature where the bee was located (Figure 4).
The researchers calculated the percentage of bees of each age in
each location at each ambient temperature with thorax temperatures
36, 0C
36
34
32
30
28
26
24
23, 0C
Figure 3 Infrared thermogram of honey bees. On the left and right

are two light gray bees whose thoracic temperatures are ~25o C; on
the top is a bee whose thoracic temperature is 35.6o C, and on the
bottom is a bee whose thoracic temperature is 32.4o C.
Source: Figure 1 from Stabentheiner et al., 2010.
= Tthorax
= Tcell rim
47
1183
= Ta
436
487
1410
836
234
496
540
370
1079
364
711
345
284
533
270
171
334
56
560
250
temperature (C)
450
63
60
404
387
37
174
74
74
42
27
17
Texp (C)
15
20
25
30
34
15
20
25
30
34
15
20
25
30
34
15
20
25
30
34
15
20
25
30
34
15
20
25
30
34
sealed brood open brood empty cells pollen open honey sealed honey
Figure 4 Thorax temperatures of worker honey bees in different hive

locations at each experimental temperature. Circles and squares
represent median comb cell rim and air temperatures (Ta) at a bees
location, respectively. The thorax temperatures are represented
as a box plot. The center line is the median, the upper and lower
boundaries of each box are the first and third quartile measurements,
and the ends of the bars are the maximum and minimum. Numbers
above each box plot are sample sizes.
Source: Figure 2 from Stabentheiner et al., 2010.
higher than their head and abdomen. Percentages of workers of differ-

ent ages with such thorax temperatures around brood cells are shown
in Figure 5.
When a honey bee hive is stressed by cold temperatures, workers re-
spond by generating heat. At low experimental temperatures, all hive air
and cell rim temperatures were above the experimental temperature, al-
though cells that contained developing larvae and pupae were kept the
warmest. Although there was a wide range of temperatures in worker bees
around the brood, empty, and pollen cells, the median thorax tempera-
tures were higher by several degrees in and around brood cells. As a result,
brood cells were generally kept above 30o C. Cells with honey were not
kept as far above the low experimental temperatures as brood cells were.
At high experimental temperatures worker bee body temperatures were
not that variable and most were close to or only a couple of degrees above
the experimental temperatures.
The generation of body heat is primarily accomplished using large
thoracic flight muscles. Worker bees older than 2 days are much more
likely to be involved in metabolic heat production than bees less than
100 brood
percentage of worker bees

= 0-2d
75 > 2-7d
> 7-12d
> 12d
50
25
0
15 20 25 30 34
A experimental temperature (C)
100 non-brood
percentage of worker bees
75
50
25
0
15 20 25 30 34
B experimental temperature (C)
Figure 5 Percentage of bees with thorax temperatures 0.2o C above

head and 1.0o C above abdomen temperature at five experimental hive
temperatures. A, Bees on brood cells. B, Bees in all other locations
(honey, pollen, and empty cells).
Source: Figure 5b from Stabentheiner et al., 2010.
2 days old. Insect development is temperature dependent. Developing

larvae and pupae cannot produce metabolic heat as worker bees can, and
young workers may not be as good at it as older bees. Workers near the
brood cells are generating more heat than workers in other areas, and
these are primarily older workers. But not all workers are generating heat
at any one time. This indicates division of labor among the members of
the hive. The generation of metabolic heat represents an energetic cost, so
individual bees may be generating heat only for short periods of time and
taking turns with the task. Additionally, honey bees of different ages ap-
pear to be performing different tasks, which is something that individual
cells in a multicellular organism do.
Although a colony of bees is a collection of individuals, the entire
hive may act like an individual, which is called a superorganism by some
scientists. The hive is a property that emerges from a collection of honey
bees, each a part of the whole, and the properties of the hive are greater
than the sum of the parts. Honey bees produce metabolic heat to main-
tain the temperature of the hive within certain limits but only at or near
brood cells. This is analogous to the maintenance of constant body tem-
perature in an endothermic animal. Thermal homeostasis of a honey bee
colony is the result of the cooperation of thousands of individuals. Our
concept of the individual can be expanded to include biological entities
like a honey bee hive. The study presented here explores but one aspect of
honey bee hives as individuals. In fact, the hive is a highly integrated en-
tity, and bees exhibit communication to more efficiently forage for food,
defend the hive, and grow. The cells and tissues in an individual animal
also display these properties.
Multicellular organisms display division of labor. Within an indi-
vidual organism, a variety of cells perform different tasks, and all of the
cells are integrated into a whole organism. The individual is a property
that emerges from this collection of parts. An individual organism can be
more than that, and the boundary of an individual is not always clear. An
organism may only be part of a much larger whole with the parts com-
municating and cooperating to produce a biological entity that has more
flexible responses to varying environmental conditions than any single
part has.
Bibliography
DeWoody J, Rowe CA, Hipkins VD, et al.: Pando lives: molecular ge-
netic evidence of a giant aspen clone in central Utah, West N Am
Naturalist 68(4):493497, 2008.
Smith ML, Bruhn JN, Anderson JB: The fungus Armillaria bulbosa is
among the largest and oldest living organisms, Nature 356:428433,
1992.
Stabentheiner A, Kovac H, Brodschneider R: Honey bee colony

thermoregulationregulatory mechanisms and contribution of indi-
viduals in dependence on age, location and thermal stress, PLoS One
5(1):e8967, 2010.
Volk T: The humongous fungusten years later, Department of Biology,
University of Wisconsin-La Crosse (website): http://botit.botany.wisc
.edu/toms_fungi/apr2002.html. Accessed July 15, 2014.
Wilson RA: The biological notion of individual, Stanford Encyclopedia
of Philosophy (website): http://plato.stanford.edu/entries/biology-
individual/. Accessed July 15, 2014.
CHAPTER 2
The Nervous and Endocrine

Systems Are the Sources
of Emotions
A young girl takes a test for which she is not well-prepared. While taking
the test, her pulse increases, she gets flushed, and her hands tremble. She
felt terrified during the test; her grade hinged on doing well on this test.
Afterward she felt weak and shaky, and later when she thought about
her performance, she again felt nervous and shaky. After a day or so, the
strong nervous feelings began to wear off, and she began to feel anger
toward herself for not preparing well. It turned out that the girl did better
than she expected, and when she got her test back, she felt another rush
of strong feelings; this time they were positive. During all of these strong
feelings (negative and positive), the girls mind and body were experienc-
ing a strong emotional reaction to a situation.
Here is an emergent property with which most people are very fa-
miliar. Almost everyone displays emotions ranging from happiness to
rage with everything in between. Emotions are specific reactions to a
particular event that are usually of a fairly short duration. Emotions are
emergent properties that arise at the level of the individual. In 1872
Charles D arwin published a book on his decades-long research called
The Expression of The Emotions in Man and Animals. Darwin described
the expression of behaviors in animals and humans, and he attempted
to relate these behaviors to emotions (Table 3). Darwins work detailed
observations that he and others had made; only a small sampling of his
data is shown. Although he ascribed various emotions to the expressive
behaviors, he could not know what emotion animals were feeling at the
time a particular behavior was expressed.
Table 3 A sampling of expressions in animals studied by Charles

Darwin to which he ascribed various emotions, some of which are
shown.
expressions animals basic emotion(s)
trembling many vertebrate animals and humans fear, joy, sadness
sweating several mammals, including humans fear, anger
face blushing many vertebrate animals and humans anger, surprise
vocalizations many animals and humans fear, anger, joy
facial expressions many mammals, including humans fear, anger, joy
erect hairs or feathers many mammals, including humans, and birds fear, anger
Source: From Darwin, 1872, text.
Darwin inferred that expressive behaviors are outward signs that emo-
tion is being experienced, and they are an easily measurable component
of emotion. Expressive behaviors are also signals to other individuals of
the same or different species. Darwin showed that expressive behaviors
are widespread. This suggests, as he argued, that expressive behaviors and
emotions are adaptations that have evolved through natural selection.
The idea that many animals display similar behaviors when expressing an
emotion indicates a common evolutionary history and facilitates signal-
ing between species. It also suggests that expressive behaviors were se-
lected for by natural selection and that they have adaptive value. The
value relates to the senders ability to convey information to a receiving
animal. The function of a particular behavior may be to convey informa-
tion to another individual, to warn them of potential danger, to repel
an attack, or to express dissatisfaction with some act performed by the
receiving individual.
Although Darwin ascribed various emotions to expressive behaviors,
especially in the case of animals, he could not know what emotion they
were feeling, if any, when a particular behavior was expressed. However,
by carefully observing the circumstances in which an animal found itself,
Darwin inferred the emotional state of the animal. This is a reasonable
approach when one uses the comparative approach and collects data on
unique sets of behaviors expressed during circumstances that might point
to a particular emotion. For instance, fear is indicated by the combina-
tion of trembling, sweating, vocalizations, facial expressions, and rising
of the hair. Many animals use the same or similar combinations when
confronted by a predator or a dominant individual of the same species.
THE SOURCES OF EMOTIONS 19
Although vocalizations and facial expressions also indicate other emo-

tions, it is the set of vocalizations and facial expressions that are important
in deciding on or interpreting a particular emotion.
Emotions are displayed outwardly by expressive behaviors, but not all
changes may be visible. Albert Ax studied physiological responses to fear
and anger (Ax 1953). Ax used a polygraph, also known as a lie-detector,
to measure heart rate, respiration rate, skin conductance, and muscle
tension. Detectors were placed on the body and connected to the poly-
graph. Skin conductance is a measure of sweating, and muscle potential
is a measure of the activity or twitching of muscles. In addition, Ax mea-
sured blood pressure every minute during the tests.
Ax recruited 22 men and 21 women between the ages of 21 and 55
who were free of illness and had blood pressures below 140/90 for the
study. The subjects were informed that they would be part of a study
of the physiological differences between people with hypertension and
those without hypertension, and that their task was to relax and listen to
music for about an hour. After 25 minutes each patient received either a
fear or an anger stimulation. Each subject received both stimulations,
in random order, and separated by 10 to 15 minutes. Shortly after each
stimulation, the subject was interviewed regarding their memory and
feelings about the interruption to ensure that they had felt the emotion
that was being measured physiologically.
The fear stimulus consisted of an intermittent electrical shock to the
little finger, which never reached intensity sufficient to cause pain. When
the subject reported the shock, the researcher acted with surprise and
pressed a button that caused sparks to jump. The researcher then acted
alarmed and warned the subject that the sparks were from a high-voltage
short circuit. After 5 minutes, the shock wire was removed, and the sub-
ject was reassured that all was well.
The anger stimulus was presented as a polygraph operator who had
been fired for incompetence and then called back to cover for illness. Well
into the test, the operator entered the room, made the researcher leave
the room to monitor the polygraph, shut off the music, and criticized
the nurse, who was on duty to monitor blood pressure. The operator
then criticized the subject for being late and roughly checked the elec-
trodes on the subject. This continued for 5 minutes; after which time the
operator left the room and the researcher returned, apologizing for the
rude behavior.
For all of the variables, the maximum values recorded for 7 minutes
(during the stimulus [5 minutes] and right after [2 minutes]) were calcu-
lated as deviations from the resting level recorded just prior to the stimu-
lus (Figure 6). Diastole and systole are the two measurements in blood
pressure. Skin conductance is related to sweating. For muscle tension, Ax
measured the maximum change in muscle potential and the number of
muscle potential peaks per unit timea peak being defined as an incre-
mental doubling of size in a three second period.
There were strong similarities in physiological responses to fear and
anger. There were increases above the resting values in each of the vari-
ables measured. Although there were some similarities in physiological re-
sponse between fear and anger, Ax found that increases in diastolic blood
pressure were greater in the anger response than in the fear response. The
number of muscle tension peaks and increases in respiration rate and skin
35
change from resting value (units vary)
30 = anger
= fear
25
20
15
10
0
e e e te ce n s
ol ol at ra io ak
st st tr y tan ns pe
di
a sy ea
r
t or uc te e
h ira nd cl
e cl
p co us us
res in m m
sk
variable
Figure 6 Changes in physiological variables in humans in response to

the emotions of fear and anger.
Source: From Ax, 1953, Table 1.
conductance were greater in the fear response than in the anger response.
Measurable physiological responses resulted from stimuli that evoked fear
and anger.
As Darwin inferred with expressive behaviors, physiological responses
may aid in survival. When someone is afraid or angry, their heart races,
their palms get sweaty, and their respiration increases. These changes are
adaptive if individuals that experience these changes during stressful times
have a higher likelihood of surviving the stressful experience and ulti-
mately producing more offspring relative to individuals that do not expe-
rience these changes as strongly. It might be evolutionarily advantageous
during a stressful experience (such as, a predator attack or a fight among
males for access to a female) to have an increased heart rate, increased
breathing, and higher muscle tension. These changes might prepare an
animal for escape or a fight.
The detection of a potentially dangerous situation leads to expression
of the emotions of fear or anger; other situations lead to expression of
other emotions. A signal is detected in the environment and that leads
to physiological and behavioral changes. To determine the causes of the
physiological changes and the expressive behaviors associated with emo-
tion, William Cannon performed an experiment on cats. Based on evi-
dence that a hormone called adrenalin could increase blood pressure and
heart rate, Cannon looked for evidence that adrenalin was released into
the blood when an animal was frightened or angry. The scientist used
strips of intestinal smooth muscle obtained from other animals as an
indicator of adrenalin. Smooth muscle is a contractile type of muscle tis-
sue controlled by the involuntary nervous system, occurring in the walls
of the stomach, intestines, and blood vessels. Intestinal muscle, when
extracted and placed in a salt solution, spontaneously contracts. When
blood without adrenalin is added to the muscle, contractions continue;
but when blood with adrenalin is added, the muscle relaxes.
Cannon surgically inserted catheters in cats to collect blood from
the femoral vein and from the vena cava between the adrenal gland and
the liver. The femoral artery and vein are in the legs of humans and the
hind legs of cats. Part of the vena cava, the vein that returns blood to the
heart, is anterior to the heart (also called superior vena cava) and part is
posterior (also called inferior vena cava).
The adrenal glands sit atop the kidneys, and the adrenal veins lead
from the glands to the vena cava. The renal arteries and veins lead to and
from the kidneys, so one can visualize that the adrenal vessels run close
by the renal vessels.
Cannon (1911) hypothesized that the adrenal glands were the source
of adrenalin. He collected blood before and after the cat was exposed to
the sound of a loud dog barking and placed the blood onto the intestinal
muscle (Table 4). Before the exposure to the disturbance, cats were kept
quiet; and after the disturbance, they showed expressive behaviors of ei-
ther fear or anger. Cannon also removed the adrenal glands in some cats.
Finally, Cannon injected adrenalin directly into some cats that were not
exposed to any disturbance, after which he collected their blood.
The response of a cat to the bark of a dog might be fear (the cat runs
away or freezes to avoid detection) or anger (the cat gets ready to fight).
This kind of response is called the fight-or-flight response. Cannon
showed that adrenalin is not present in the blood of quiet, undisturbed
cats. However, upon being exposed to the barking of a dog, adrenalin is
released into the blood. It is released by the adrenal glands, because blood
collected immediately from the femoral vein does not have adrenalin and
cats without adrenal glands do not produce adrenalin under any circum-
stances. The role of adrenalin in fight-or-flight is well-known. Fight-
or-flight might also be called the anger-or-fear response, because these are
the emotions displayed. These are examples of behaviors and physical re-
sponses to emotions that improve the probability of survival.
It is now clear that emotions are produced by various physiological
responses and lead to expressive behaviors. Although only the specific
Table 4 Results from experiments investigating adrenalin in cats.

condition source of blood response of intestinal muscle
quiet cat femoral vein none
quiet cat vena cava near liver none
frightened/angry cat femoral vein none
frightened/angry cat vena cava near liver inhibition
adrenalectomy vena cava near liver none
adrenalin added to blood femoral vein inhibition
adrenalin added to blood vena cava near liver inhibition
Source: From Cannon, 1911, text.

effects of adrenalin are discussed here, a variety of other hormones af-

fect emotions in a similar manner. Adrenalin causes specific physiological
responses when released from the adrenal gland. Some external or inter-
nal stimulus must cause the release of adrenalin from the adrenal gland.
George Dreyer studied dogs to determine if the nervous system caused
the release of adrenalin (Dreyer 1899). Dreyer anesthetized and cut open
living dogs to expose the abdominal cavity. The scientist exposed the left
adrenal gland. He tied off the adrenal vein as close to the gland as possible
to prevent blood from other tissues mixing with the blood exiting the
adrenal gland, and he inserted a glass tube between the adrenal gland and
the ligature. Blood was collected from the open end of this tube. Dreyer
stimulated the splanchnic nerve, which connects to the adrenal gland, by
connecting it to a device that discharged electricity. The scientist moni-
tored blood pressure of the dog, which was found to rise each time the
device was turned on. Dreyer collected blood from the femoral vein and
the adrenal vein. From the adrenal, he collected blood prior to, during,
and sometimes after electrical stimulation.
Other dogs were anesthetized and injected with atropine, which in-
hibits certain nerves that have an inhibitory effect. Dreyer wanted to in-
hibit the vagus nerve so that it did not interfere with his experiment,
which when it fires has an effect of slowing the heart rate. He connected
the second set of dogs to a device that monitored blood pressure and in-
jected either blood from a dog from the first set, saline, or an extract from
adrenal glands that was removed from a third set of dogs into the jugular
vein of the test dog (Figure 7). In addition to monitoring blood pressure,
Dreyer also monitored heart rate, before and after injection. Each dog was
tested with at least three types of blood (femoral vein, adrenal vein before
stimulation, and adrenal vein during stimulation) and adrenal extract, in
succession. Four of the six dogs were tested with saline, and three were
tested with adrenal vein blood collected after stimulation.
Adrenalin is released from the adrenal gland and causes changes in
physiology and expressive behavior. It is associated with the fight-or-flight
response in animals. Dreyers data shows that anesthetized dogs not expe-
riencing any emotions can be stimulated to release a substance from the
adrenal gland into the blood that causes increases in heart rate and blood
pressure, much like adrenalin does. This substance is, of course, adrenalin.
70 change in heart rate
60
rate at greatest change
rate prior to injection 50
40
30
20
10
0
A 10
110 change in blood pressure

100
rate at greatest change
90
rate prior to injection
80
70
60
50
40
30
20
10
0
v io
r g te
r ct e
al pr rin af ra lin
or du ex
t sa
m lv v lv l
fe na al na na
e n e e
a dr re a dr dr
ad a
B treatment
Figure 7 Changes in heart rate (A) and blood pressure (B) in dogs after
injection with blood from various sources, saline, or extract of adrenal
glands. Treatments are presented in the order in which they were
injected into dogs. V, vein. Error bars are 1 standard error (SE).
Source: Data from Dreyer, 1899, Table p. 210.
Dreyer was the first to show that stimulation of the splanchnic nerve, part
of the autonomic nervous system, which regulates involuntary action in
vertebrates, caused release of adrenalin.
Of course, Dreyers experiment was not without some issues. For in-
stance, there was an increase in heart rate and blood pressure when dogs
were injected with blood from the adrenal vein of dogs collected prior
to electrical stimulation of the splanchnic nerve. This might indicate a

couple of things. The dogs were under stress prior to anesthetization, and
this could have caused release of adrenalin that continued even after being
anesthetized. Possibly some other substance secreted from other glands
or organs could have effects on heart rate and blood pressure. Despite
these possibilities, Dreyer was able to stimulate one nerve, which when
stimulated clearly caused release of some substance into the blood from
the adrenal gland. Blood from the femoral vein never caused any increases
in heart rate or blood pressure, nor did injection of saline solution, which
were negative controls. Those controls and the positive control of inject-
ing adrenalin in later injections were wise additions to the experiment,
despite some other problems.
The autonomic nervous system stimulates a gland that releases a
hormone into the blood. That hormone causes several of the responses
observed in humans by Ax. People have internal responses to the same
emotion similar to the ones observed in cats and dogs. For example, re-
gardless of age, ethnicity, or gender, when people are under stress, their
bodies release adrenaline; this hormone helps prepare the body to either
run away or fight, the so-called fight-or-flight reaction.
Other parts of the brain trigger other aspects of emotions. Although
many of those details are interesting, it is enough here to understand that
emotions are an emergent property of animals. Emotions have several
components. There is a genetic component; there are subjective feelings,
which are the ways animals experience emotions; there are measurable
physiological responses; and there are outward behaviors expressed.
Subjective feelings are difficult to measure in humans or other animals.
Physiological responses are easier to measure as shown here. The outward
expressions of an emotion are also relatively easy to measure, and they
provide other animals clues as to what an individual is experiencing and
consequently help to regulate social interactions. There are many parts to
emotionsfrom the neurons, the hormones, the expressive behaviors,
and the physiological changes. Behaviors are emergent properties that
arise from the actions of all of these parts. In the next chapter, one more
emergent property of the individual will be explored, and that is coop-
eration among individuals, which actually leads to emergent properties
above the individual level, in colonies and populations.
Ethical, Legal, Social Implications:

There Are Issues With Using Prescription Drugs
to Normalize Behavior in Children
Perhaps a child in a toy store is very active, jumping from one display to
another. Or another child is inattentive and easily distracted by external
stimuli. While these are a couple of symptoms of attention deficit hy-
peractivity disorder (ADHD), most children display these activities at
some point in their development. When the symptoms are not age ap-
propriate, then the child may be diagnosed with ADHD. Perhaps those
children are put on Ritalin or some other drug to treat these symptoms.
What is meant by age appropriate behavior? Age appropriate behavior
means the types of behavior that should be expected at any given age.
For example, if a two-year-old child walks up to a dog and hits it, this
might not surprise parents too much. It is not okay, but it isn't really that
unusual for a child at that age. The parental response may be to provide
appropriate guidance and boundaries so that the toddler learns hitting the
dog is not okay. If a four- or five-year-old child hits a dog, this might not
seem normal behavior for a child of this age. In this case, the parent or
teacher might want to investigate why the child is behaving in a manner
not appropriate for the age, which may include not having been around
dogs before or having been bitten by a dog in the past. Assessment of the
age-appropriateness of behaviors is one of the ways in which disorders are
diagnosed.
ADHD can be difficult to diagnose because of the wide array of
symptoms and resulting behaviors. Age-inappropriate behavior is only
one symptom. Some children with ADHD are hyperactive, whereas oth-
ers sit quietly. Some put too much focus on a task and have trouble shift-
ing it to something else. Others are only mildly inattentive but overly
impulsive. Three characteristics of ADHD are inattention, hyperactivity,
and impulsivity; and the symptoms that manifest depend upon which of
these predominates. An inattentive child may not follow directions well,
because he is easily distracted, and this could lead to careless mistakes and
poor performance in school. A hyperactive child may constantly fidget
and move around. She may talk excessively and is constantly on the go,
bouncing from one activity to another without completing any of them.
The impulsive child blurts out answers, has difficulty displaying patience,
and often interrupts others.
Despite these challenges, there are potential positive aspects of
ADHD. For instance, children with ADHD can be very creative and
imaginative. A child who daydreams or has many thoughts at once can
be an inventive artist or excel at brainstorming. Having many thoughts
at once may lead to flexibility and openness to different ideas. These chil-
dren are enthusiastic and interested in many different things. That gives
them a lot of energy and motivation, especially when they are involved in
a task that really interests them.
Parents and medical professionals may decide that disadvantages out-
weigh advantages, which leads to prescribing methylphenidate (Ritalin)
or some other drug to treat ADHD. A child on methylphenidate is under
a medication that changes the way his or her brain operates. That means
that ideas and impulses in a childs brain are sometimes suppressed. Be-
tween 1990 and 2000, methylphenidate production, which is closely
regulated by the Food and Drug Administration (FDA) in response to
demand from prescriptions, increased almost 750%, increasing by about
1,318 kg/yr. Methylphenidate prescriptions rose dramatically in the early
1990s and have since leveled off at approximately 11 million per year.
According to data, the vast majority of prescriptions for methylphenidate
are for children diagnosed with ADHD. The US produces and consumes
about 85% of the worlds methylphenidate. In comparison, prescriptions
of amphetamine, primarily Adderall (a mixture of amphetamine and dex-
troamphetamine), also used to treat ADHD, have increased dramatically,
from 1.3 million in 1996 to nearly 6 million in 1999.
Methylphenidate stimulates underactive neurons in the brain, which
allows the medicated person to focus and pay attention. Methylphenidate
also stimulates the release of dopamine, a chemical in the brain that is oneof
the messengers that helps one side of the brain communicate with the
other. The function and activity of the central nervous system (CNS) reacts
to the influences of dopamine. Methylphenidate improves attention span,
decreases distractibility, and reduces restlessness and fidgeting by enhancing
task-specific dopamine signals. Those with ADHD who take methylphe-
nidate see improved control of emotions and more focus in schoolwork,
which increases self-esteem and relationships with family and friends.
According to pediatricians, treatment with methylphenidate works

best when combined with behavioral interventions and lifestyle modifica-
tions. There are some, however, who advocate for use of behavioral inter-
ventions and lifestyle modifications without prescribing methylphenidate.
Adults can model behavior and encourage good behavior with healthy
praise, and they can negatively reinforce bad behavior with appropriate
consequences. Parents can provide a consistent routine, clear boundaries
and expectations, and use of activities that engage the mind. There are
others who argue that learning to live with ADHD through these non-
medicinal therapies prevents the loss of the positive effects of ADHD.
Bibliography
Ax AF: The physiological differentiation between fear and anger in
humans, Psychosom Med 15(5):433442, 1953.
Cannon WB: The stimulation of adrenal secretion by emotional excite-
ment, P Am Philos Soc 50:226227, 1911.
Collier B: The parental intelligence report on ADHD, ADHD-Report
.com (website): http://www.adhd-report.com/index.html. Accessed
April 22, 2010.
Darwin C: The Expression of Emotions in Man and Animals, London,
1872, John Murray.
Dreyer GP: On secretory nerves to the suprarenal capsules, Am J P
hysiology
2:203219, 1899.
Quinn P: ADHD in Children Health Center: behavioral techniques for
children with ADHD, WebMD (website): http://www.webmd.com/
add-adhd/guide/adhd-behavioral-techniques. Accessed April 22, 2010.
Smith M, Segal R: ADD/ADHD in children: signs and symptoms of
attention deficit disorder in kids, Helpguide.org (website): http://
helpguide.org/mental/adhd_add_signs_symptoms.htm. Accessed
April 22, 2010.
Weaver Dunne D: Statistics confirm rise in childhood ADHD and medi-
cation use, Education World (website): http://www.educationworld
.com/a_issues/issues148a.shtml. Accessed April 22, 2010.
CHAPTER 3
Individuals of Some Species

Cooperate With Each Other
for Mutual Benefit
Information transfer occurs frequently in groups of animals living to-

gether. A benefit of group living is a collective information transfer system
that leads to an early warning of danger. In this chapter, considerations
of group living will include plants and animals in order to explore the
emergent property of cooperation.
Cooperation is a complex concept from an evolutionary point of view.
Darwin noted the existence of cooperation within species. The reason
that cooperation evolved is considered in the context of it being an emer-
gent property of individual behavior and populations; and to consider
that, the mechanisms of cooperationhow cooperation works or how
individuals within a group actually do cooperate with each other, must
also be considered. The mechanism, or the how, of meerkat coopera-
tion is explored in another book in this series, Behavior and Information
Exchange, and here how individuals of some other species cooperate will
be examined.
If groups of bacteria and groups of animals cooperate, why not plants?
There might be several reasons why cooperation might be advantageous
in plants, which would lead to the evolution of cooperative behaviors.
A behavior in a plant is a rapid morphological or physiological response
to some event, such as attack by an herbivore, in contrast to animal be-
haviors. Richard Karban and his colleagues studied the mechanism of
cooperation in sagebrush (Artemisia tridentate) to test how plant behav-
ior affects populations of sagebrush (Karban et al., 2006). In the context
of cooperation among plants, individual plants may collect information
from other individuals in the population and respond with the same or
some other behavior.
The scientists set out to determine whether or not neighbors in a pop-
ulation of sagebrush plants can detect and respond to information sent
by other individuals affected by herbivores or when part of an individual
plant is experimentally clipped to simulate herbivory; and if so, how far
away signals can be detected by others in the population. The detection
of signals via information transfer could lead to increased expression of
natural defense mechanisms of plants. This emergent property within
the population would lead to increased production of chemical defenses
that could be effective against herbivores and lead to success of the entire
population of plants against a population of herbivores.
Sagebrush is a widespread plant of the Great Basin region of western
North America often making up more than 90% of the plant biomass at a
site. Populations are spatially structured to be very dense. There are many
invertebrate and vertebrate herbivores that feed on sagebrush. Many in-
sects feed on leaves in the spring, grasshoppers are the major herbivore in
the summer, and deer are common in the fall. Karban and his colleagues
conducted their experiments at two field sites dominated by sagebrush,
one near Convict Creek and the other near Sagahen Creek, both in Cali-
fornia. Convict Creek is walled in by steep slopes, and wind flows consis-
tently in a westerly direction down the slope of the canyon.
Karban and colleagues surveyed a sagebrush population to determine
how far apart individuals grew from each other. The researchers walked
a 2 kilometer transect, and every 20 meters they located the closest sage-
brush to where they stopped and measured the distance between it and its
nearest neighboring sagebrush plant.
The scientists found that 25% of sagebrush had branches of their
nearest neighbor 05 cm away, 38% had their nearest neighbor 510 cm
away, 20% had their nearest neighbor 1015 cm away, 16% had their
nearest neighbor 1535 cm away, and only 1% had a nearest neighbor
more than 50 cm away. Most sagebrush plants thus have their nearest
neighbor less than 35 cm away.
Then the researchers compared these data to the maximum distance
over which communication could be detected between individuals in a
separate experiment. The biologists selected 250 pairs of sagebrush plants
INDIVIDUALS OF SOME SPECIES COOPERATE FOR MUTUAL BENEFIT 31
that were within 1 meter of each other at Convict Creek. The pairs were
oriented east to west. The distances between pairs were recorded. Karban
and colleagues clipped the distal half of all leaves from one branch of the
upwind bush most adjacent to the downwind bush. They did this for 125
of the pairs. The upwind bush of the other pairs was not clipped. For
the next 2 months the researchers recorded the number of leaves with
herbivore damage on the adjacent branch of each downwind bush. Be-
cause branch size varied, they expressed the number of damaged leaves as
a percentage. Karban and his colleagues compared the mean percentage
of leaves damaged on branches of downwind bushes with clipped or un-
clipped upwind neighbors. The scientists examined this damage estimate
at three distance categories.
The scientists found that if the upwind neighbor had leaves clipped,
the downwind neighbor had significantly fewer leaves damaged than when
the upwind neighbor was unclipped. This was true for neighbors that were
020 cm apart and for neighbors 2160 cm apart, but that effect disap-
peared when neighbors were more than 60 cm away from each other. This
experiment suggests that there is a volatile chemical that is released by
plants that are clipped or consumed by herbivores that neighboring plants
can detect and respond to. Karban and his colleagues found that commu-
nication between sagebrush plants was detectable at distances up to 60 cm.
The biologists next selected 240 pairs of branches to test whether
volatile chemical cues are required for communication. One branch from
each pair was randomly selected and clipped for half the pairs, whereas for
the other half of the pairs the selected branch was an unclipped control.
Karban and his colleagues examined the other plant for damage, taking
note of whether the other plant was upwind or downwind of the selected
damaged/control branch. If the clipped branch was upwind, the down-
wind receiving branch on a neighboring plant experienced significantly
less, by about half, the damage that a neighbor that was downwind from
an unclipped neighbor. However, if the plant was upwind of a clipped
neighbor, it received the same level of herbivore damage as a neighbor an
unclipped plant.
Finally, 150 plants or pairs of plants within 20 cm were selected. Thirty
plants were randomly assigned to one of five treatments. There was an un-
clipped control, a treatment with a clipped branch and another branch
monitored on the same plant, a treatment similar to the previous but with
the clipped branch covered with a plastic bag, a treatment with a clipped
branch and another branch monitored on a different plant within 20cm,
and a treatment similar to the previous but with the clipped branch cov-
ered with a plastic bag. The researchers clipped leaves of each branch as-
signed to clipping treatments once per month for the summer. Monitored
branches were examined for herbivory in the autumn. Branches of nearby
clipped branches on the same plant had less damage than if the nearby
clipped branch was also bagged. This was also true for situations where
the clipped branch was on another plant.
Clipping treatments simulated attack by herbivores and caused release
of some chemical cue, resulting in less herbivore damage in individuals
receiving the chemical information. Herbivore damage was less in plants
that were downwind of unbagged, clipped plants, which suggests an air-
borne chemical is used for communication between sagebrush individu-
als and even between different branches on the same plant. The bagging
of clipped branches demonstrated that communication does not occur
through the vascular system of the same plant or through some non-
airborne mechanism between plants.
Branches of sagebrush plants are relatively independent of each other
and do not have strongly connected vascular structures. This means that
defensive compounds and chemical signals may not be able to travel eas-
ily from one branch to another. Sagebrush may have evolved airborne
chemicals to communicate among branches on the same plant. Natural
selection should favor the use of volatile cues if such releases benefit the
emitter. Once that adaptation evolved it would be very easy for neigh-
bors to eavesdrop on other plants. Entire populations could then become
aware and respond to herbivore attacks. This scenario could explain the
evolution of this information transfer mechanism and emergent property.
The close proximity of individual sagebrush plants to each other and
the large suite of herbivores that feed on sagebrush facilitated the emer-
gence of a property at the population level that evolved to protect indi-
viduals. A population-wide defense strategy would make it more difficult
for herbivores to survive, grow, and reproduce; and this would provide a
benefit to each individual plant. It is not known how common communi-
cation between plants is. Volatile communication between individuals has
been shown for sagebrush and another plant called alder, although there
is evidence suggestive of communication in other plants. Cooperation
among individuals is much more widespread in the animal kingdom and
is especially common in particular groups of social insects, which include
bees, wasps, ants, and termites.
Paper wasps (genus Polistes) are social insects. Among the 150 or so
species of Polistes there is a range of colony organization, from species
where one or both parents care only for their own young to highly social
species with multiple queens and a worker caste to rear offspring. The lat-
ter species live in colonies with one or several female queens who lay eggs
and workers involved in caring for the queens offspring but who do not
reproduce themselves. Colonies are established by one or a few females
and rarely last more than a year. Paper wasps create nests using chewed up
plant material, called pulp, which dries and has a paper-like texture. Nests
are small and open, which makes it possible to view the entire colony
simultaneously. Mary Jane West-Eberhard studied two species of Polistes:
P. fuscatus, a temperate species, and P. canadensis, a tropical species. She
spent many hours observing colonies and marked individuals of each spe-
cies. Individuals were marked with dots of paint as they emerged from
their pupal nest cells. This allowed West-Eberhard to uniquely identify
each individual in a colony and track its activities in the colony. Forty-
seven colonies of P. fuscatus were observed in southeastern Michigan, and
46 colonies of P. canadensis were observed in central Colombia.
Female P. fuscatus, who mated with males the previous summer, spend
the winter in protected places. They emerge in early spring, at which time
their ovaries begin to grow. West-Eberhard found that 97.4% (37/38) of
colonies were initiated by a single female, called a foundress, who will most
likely become the queen (the other was initiated by two foundresses).
After colony initiation, females who were born the previous year but did
not initiate a colony joined existing colonies; these wasps are also called
foundresses. Most colonies end up with one or two foundresses (Figure 8).
Female P. canadensis who have mated do not overwinter; they may
simply move on to try to initiate a new colony or join one that had been
initiated by another foundress. An even greater proportion of P. canaden-
sis colonies have more than one foundress, and colonies can have more
total foundresses than P. fuscatus (see Figure 8).
20
18
= P. fuscatus
16 = P. canadensis
14
number of colonies
12
10
2
0
1 2 3 4 5 6 7 8 9 10
number of foundresses
Figure 8 Frequency distribution of number of foundresses on colonies

of P. fuscatus and P. canadensis.
Source: Data from West-Eberhard, 1969, Tables 2 and 12.
There is a division of labor within these paper wasp colonies. Queens

are almost always the sole egg-layers, although the second most dominant
foundress is known to occasionally lay eggs. There is a strict dominance
hierarchy among foundresses with the queen being dominant in her in-
teractions with others in the colony. A dominance hierarchy is a form of
animal social structure in which a linear or nearly linear ranking exists with
each animal dominant over those below it and submissive to those above
it. Both queens and workers are involved in building the colony and caring
for the offspring. Workers are the subordinate foundresses, and have infe-
rior ranks in the dominance hierarchy, as are female offspring of the queen.
Building the colony entails collecting pulp, which becomes the paper of
paper wasp nests, and constructing cells in which larvae are reared. Trips
away from the colony can be to collect pulp, food, or water. West-Eberhard
found that subordinate foundresses and offspring had higher foraging rates
than the queen, bringing back close to 1 load per hour, whereas the queen
brought back only one load every 10 hours, on average. In addition, sub-
ordinates brought back pulp, water and food, whereas the queen never
foraged for food, but collected mostly pulp and some water.
The dominance of one wasp over the other leads to stimulation of

subordinates to forage for food for the colony. Dominance is exerted
through behaviors, such as leg or antenna biting. Individuals higher in the
dominance hierarchy are usually larger, and another scientist, L. Pardi,
showed that more dominant foundresses had more developed and larger
ovaries than subordinates.
Early on during the initiation of colonies and early construction of
nests, subordinate foundresses of P. fuscatus are known to sometimes
move from one nest to another. West-Eberhard marked individual
females from one colony that was built under the eave of a building.
The scientist tracked the females who overwintered and determined
where they ended up the following year and whether they initiated a
colony or joined one initiated by a sister. Forty-four percent of nests
with multiple foundresses were known to have sister foundresses. After
a short period of moving, female P. fuscatus usually settle in and stay at
a particular nest. The offspring produced first in the colony are sterile
workers, and this is also true for P. canadensis. These daughters of the
queen help, just as subordinate foundresses do, to rear more offspring.
Later in the growth of the colony, males and non-worker females are
produced.
West-Eberhard counted the number of cells constructed in each col-
ony. One individual offspring is reared in each cell, so the number of cells
is a good measure of total colony size. The biologist determined that the
total size of colonies varies with the number of foundresses, and she also
determined the per capita number of offspring per foundress in colonies
with variable numbers of foundresses (Figure 9).
Polistes canadensis tends to have fewer offspring in each colony but
has more foundresses than P. fuscatus. Only rarely did West-Eberhard find
colonies of the former with just one foundress, although that was more
common for P. fuscatus. West-Eberhard concluded that the differences
were climate or habitat related. Ecological interactions, such as competi-
tion or predation, could lead to situations where it is advantageous to
have more or fewer foundresses. More foundresses might be advantageous
in situations where there is a high probability that foundresses are related,
a large proportion do not have well-developed ovaries, or there is high
competition for resources.
140
120 = P. fuscatus
average cells/colony = P. canadensis
100
80
60
40
20
0
0 2 4 6 8 10
A number of foundresses
30
= P. fuscatus
average cells/foundresses
25 = P. canadensis
20
15
10
0
0 2 4 6 8 10
B number of foundresses
Figure 9 Relationship between colony size and number of foundresses

for two species of paper wasp. A, The number of cells in which larvae
are reared versus the number of foundresses in a colony. B, The
number of cells per foundress versus the number of foundresses in a
colony. Averages based on 1 to 40 colonies. Best-fit regression lines
are shown.
Source: Data from West-Eberhard, 1969, Tables 5 and 12.
More foundresses in P. fuscatus colonies led to a disadvantage in

that the number of offspring per foundress declined dramatically. Fewer
foundresses per P. canadensis colony produced fewer offspring per colony
than when P. fuscatus had only one or two foundresses, and the disadvan-
tage to having more foundresses was not as great as for P. fuscatus colonies.
This can be inferred by comparing the slopes in Figure 9B. Despite the
differences, there are similarities that lead to an emergent property in

social wasps. Group living, cooperation, and division of labor in social
insect colonies are emergent properties at the population level.
Worker and non-worker offspring were produced at different times
of the year for P. fuscatus, but this was not so for P. canadensis. Perhaps
because P. fuscatus females need to overwinter they, and the males, are not
produced until late summer, giving them a chance of survival through the
fall. There is no need to produce more workers late in the summer be-
cause there are enough to help rear males and non-worker females. Males
and non-worker females have potential to reproduce but workers do not,
so there is no reproductive advantage to a queen to continue producing
workers late in the season in temperate climates. Because P. canadensis
colonies can be initiated at any time of the year and there is no reason to
overwinter, a queen could have greater reproductive success if she pro-
duced both worker and non-worker females for as long as she lived. The
workers could stay and help rear offspring, whereas daughters who were
non-workers could disperse and initiate their own colonies.
Colony size increases with the number of foundresses, because they
can all work together to create a larger group. Recall that one advantage
to joining a colony and foregoing reproduction is if a joining foundress
is related to a queen, because she can help rear offspring that share her
genes. If the joining foundress has small ovaries, she would probably pro-
duce only a few offspring alone, but she could have higher reproduction
if she joined an existing nest and if the probability was reasonably high
that sheis related to the queen. Such is the case for these paper wasps,
and other research has shown that subordinates are allowed some repro-
duction in return for their help, which ensures that even if they are not
related to the queen they still pass on their genes to the next generation.
Animals use internal information, such as the level of hormones produced
by ovaries. Larger ovaries might produce more hormones, and the quan-
tity affects whether a female acts dominant and submissive. This informa-
tion, as well as external information gathered by a foundress when she
visits an already initiated nest, determines her behavioral strategy.
An evolved behavioral strategy might be to visit nests and determine
dominance rank in a colony. If it is low, a foundress might move on to an-
other nest, especially she might have high rank and be a queen at another
nest. Alternatively, females may be able to detect related females, although

it is not known whether individuals can detect kin in these species.
A P. fuscatus foundress that has initiated a colony is likely to have emerged
early in the spring, and West-Eberhard showed that early-emerging f emales
are more dominant. The queens strategy would be to allow foundresses to
join and help as long as they are subordinate. Each individual foundress
thus has a strategy based on her reproductive condition. But how does an
individual know if it is dominant or subordinate to another individual it
encounters?
When multiple foundresses cooperate to found a nest, they have in-
tense battles to establish dominance. These fights determine the ranking
within a colony. Elizabeth Tibbetts studied P. fuscatus with the goal to
determine why foundresses do not fight each time they encounter each
other. That is, do wasps have the ability to recognize colony-mates? If
wasps had that ability, it would reduce aggression, promote cooperation,
and would be another emergent property that arose from group living.
Tibbetts first observed variation among individuals. She accomplished
this by removing 259 wasps from 38 nests and cataloguing the markings
of yellow stripes, which may be present in various locations on the face
and abdomen. Tibbetts documented wide variation in the markings of
yellow stripes among individuals.
After characterizing wasps, most were weighed and all were labeled
with enamel paint before being returned to their colony. Tibbetts deter-
mined the rank of 42 of the foundresses from 19 nests by observing one-
on-one interactions between individuals in a colony. Tibbetts found that
the pattern of yellow markings was not correlated with dominance rank
in a colony or a foundress mass.
There is a large degree of variation in the facial and abdominal mark-
ings of P. fuscatus. There is more variation in the width and length of
yellow markings, and some wasps had other markings not shown. The
markings do not correlate with rank or size. Tibbetts hypothesized
that the markings were used as signals of individual identity. However,
Tibbetts identified several alternative functions that had to be ruled out.
Markings could have no function, or they could be used as signals of
dominance, membership in a colony, or relatedness among foundresses
in a colony.
To test these hypotheses, Tibbetts experimentally altered the mark-

ings of several wasps using yellow or black enamel paint. This was to
determine if foundresses treated altered foundresses as if they were outside
intruders and attacked them or if they could recognize altered individuals.
The scientist either added yellow stripes to wasps that did not have them
in a particular area or covered yellow stripes with black paint. Control
wasps were painted without altering their markings by using black paint
to cover black areas.
An experimental wasp was taken from her nest in the morning,
painted and returned to her nest. Tests were conducted on 11 foundresses,
one from each of 11 multiple-foundress colonies. The specific area painted
varied from wasp to wasp. In all cases, the same wasp was used for the con-
trol and treatment, although the order of presentation of control or treat-
ment varied. The biologist videotaped colony interactions for 2 hours.
On average, an experimentally altered foundress received significantly
more aggression in the first half hour after yellow facial markings were
added or obscured than in the first half hour in the control. Aggression
toward experimentally altered wasps declined over time; foundresses re-
ceived much more aggression in the first half hour than in the half hour
beginning 90 minutes after their return. The aggression that control
wasps received was lower and did not change over the 2 hour observation
period. Tibbetts concluded that the altered markings of the experimental
wasps were learned by their colony-mates within 90 minutes.
These results fail to support the four alternative hypotheses that Tib-
betts identified for the signal function of markings. First, there clearly is
a function, so one can easily reject the no-function hypothesis. There was
no correlation between the pattern of markings and either size or domi-
nance rank, which allow rejection of the hypothesis that the markings sig-
nal rank. The colony membership hypothesis can also be rejected because
it turns out that wasps use chemical cues to identify colony-mates, and
even though altered foundresses looked different, they were not driven
from the nest, which is the habit when foundresses are confronted by a
non-colony member. Aggression was higher, but the altered foundresses
were still accepted. Finally, if the function of the markings is to signal
relatedness, the level of aggression that a wasp receives should be the same
as long as her appearance is altered. This was not the case. The results
indicate that the new markings of a wasp caused the aggressive behavior
due to unfamiliarity and that the pattern of markings of P. fuscatus is used
for recognizing colony mates.
The dominance hierarchy is worked out among a group of foundresses
on a colony through intense physical interactions. If the dominance hier-
archy had to be worked out every time a foundress returned with water,
pulp, or food, the colony would not accomplish much. An adaptation
that evolved to facilitate cooperation among paper wasp foundresses is the
ability to recognize individuals. In that way, when an individual foundress
returns, the other foundresses know that she is a colony-member through
her odor and know her rank in the dominance hierarchy by recogniz-
ing her individually through her pattern of facial and body markings.
This ability is another emergent property that arises in populations where
group cooperation occurs.
This section began by considering cooperation as an emergent prop-
erty of populations. It was shown that sagebrush can exchange informa-
tion, which could lead to induction of anti-herbivore defenses among a
group of plants. It was then shown that social wasps cooperate in colonies
to rear offspring of primarily one individual. The value and evolutionary
advantage of cooperation was considered in each case, as was the mech-
anism of cooperation, which usually entails some kind of information
exchange.
Within a group, individuals perform different behaviors that lead to
flexibility of response by the group, lead to cooperation among group
members, and produce an entitythe group that exceeds the sum of its
parts, the individuals. Cooperation is shown to be an emergent property.
Emergent properties will be further examined at the population and eco-
logical system levels in other books in this series (Properties of Populations
and Ecological Interactions), including another example of cooperation
how a flock of birds or a school of fish can respond as a single individual.
Bibliography
Karban R, Shiojiri K, Huntzinger M, et al.: Damage-induced resistance in
sagebrush: volatiles are key to intra- and interplant communication,
Ecology 87(4): 922930, 2006.
Pardi L: Dominance order in Polistes wasps, Physiol Zool 21(1):113,

1948.
Tibbetts EA: Visual signals of individual identity in the wasp Polistes
fuscatus, Proc Biol Sci 269(1499):14231428, 2002.
West MJ: Foundress associations in polistine wasps: dominance hier-
archies and the evolution of social behavior, Science 157(3796):
15841585, 1967.
West-Eberhard MJ: The social biology of polistine wasps, Miscellaneous
Publications of the Museum of Zoology, University of Michigan 140:
1101, 1969.
Conclusion
Several remarkable cases of emergent properties at the level of the individ-
ual were examined in this book, with an attempt to understand why these
properties emerge and how they are greater than the sum of their parts.
The definition of the individual has been expanded. Emotions arise from
complex interactions of neurons, hormones, and behaviors. The coopera-
tion that exists among a group of individuals working together to secure
resources or defend against predators is a major theme of the concept of
emergent properties, and cooperation was examined in more depth in this
book. Finally, mechanisms exist at lower levels of biological organization,
specifically at the level of the individual, which cause emergent properties
in populations. In other books in this series, exploration of some of these
properties that emerge from lower levels, behaviors for instance, will show
how they lead to emergent properties in entire populations and ecological
systems.
Glossary
adrenalin. A hormone released by adrenal glands that increases heart rate, respira-
tion rate, and blood pressure.
age appropriate behavior. Behaviors displayed by a person that are typically dis-
played by people of that age, rather than people that are much younger or much
older.
alleles. A variant of a gene.
attention deficit hyperactivity disorder (ADHD). A condition that includes
persistent inattention, hyperactivity, and sometimes impulsivity.
autonomic nervous system. The autonomic nervous system regulates involuntary
action in vertebrates.
basidiospore. A structure often produced on mushrooms that contains spores.
behaviors. The way in which an animal acts in response to a stimulus.
biomass. The total amount of organic matter in an organism, population, or
other ecological system after water is removed.
colony. Several individual organisms (especially of the same species) living to-
gether in close association.
cooperation. Behavior that involves several individuals and is mutually beneficial.
diploid. Cells and organisms that carry two copies of almost every chromosome.
division of labor. The specialization of cooperative labor in specific tasks.
dominance hierarchy. A dominance hierarchy is a form of animal social structure
in which a linear or nearly linear ranking exists with each animal dominant over
those below it and submissive to those above it.
emergent properties. Characteristics that become apparent at a higher level of
biological complexity due to interactions among lower level components.
emotions. Emotions are specific reactions to a particular event that are usually of
a fairly short duration.
expressive behaviors. Expressed actions that indicate an individual is experiencing
emotion.
fight-or-flight. The physiological response to a threatening situation, which read-
ies one either to resist forcibly or to run away.
fungus. Fungi are multicellular heterotrophic organisms with chitinous cell walls.
haploid. The condition of having one copy of each chromosome.
heart rate. The number of heart beats per unit time.
herbivore. An herbivore is an animal that eats plants.
heterozygosity. The number or percentage of genes in an individuals or individu-
als in a population that are heterozygous.
46 GLOSSARY
hyphae. Hyphae are the filaments composing the vegetative part of a fungus,
often forming a dense mat.
loci. Locus is the position of a gene on a chromosome; loci is plural of locus.
mating types. Haploid mycelia in fungi have molecular mechanisms that regulate
compatibility in these sexually reproducing eukaryotes.
microsatellite DNA. A repeating sequence of one to six base pairs of DNA.
morphological. Having to do with the form or structure of an organism.
muscle tension. The condition where muscles remain semi-contracted for an
extended period of time.
mushrooms. The fruiting bodies of fungi.
mutation. A change in DNA sequence.
mycelium. An interconnected network of fungal hyphae.
natural selection. An adaptive mechanism by which evolution takes place and is
often summarized as differential survival and reproduction.
physiological. Having to do with the functions and activities of organisms and
their parts, including physical and chemical processes.
polygraph. A scientific instrument that measures a variety of physiological
responses, and often called a lie-detector.
respiration rate. The number of breaths per unit time.
rhizomorphs. Aggregations of hyphae that transport resources and nutrients in
many fungi.
self-incompatibility. A term for several genetic mechanisms in some organisms
that prevents self-fertilization and thus encourages outcrossing.
skin conductance. A physiological measure of sweating caused by the skin
momentarily becoming a better conductor of electricity when stimuli occur that
are physiologically arousing.
smooth muscle. Smooth muscle is a contractile type of muscle tissue controlled
by the involuntary nervous system, occurring in the walls of the stomach, intes-
tines, and blood vessels.
subordinate. A subordinate is an individual that has inferior rank in the domi-
nance hierarchy.
superorganism. An organism consisting of many organisms, often used to
describe a social unit of eusocial animals, with division of labor and where indi-
viduals cannot survive by themselves for extended periods.
vegetatively. A form of asexual reproduction, or growth of an organism that does
not involve the reproductive parts.
Index
Adrenal glands, 22 Dopamine, 27
Adrenalin, 2123 Dreyer, George, 23
Age appropriate behavior, 26
Airborne mechanism, 32 Emergent property, xiii
Allele, 3, 9 Emotions, 17
Anderson, James, 16 Expressive behaviors, 18
Anger-or-fear response, 22
Anger stimulus, 19 Fear, 18
Armillaria gallica (A. gallica), 12 stimulus, 19
growth rate, estimation of, 5 Fight-or-flight muscle, 22
Atropine, 23 Fungus, 12
Attention deficit hyperactivity growth of, 6
disorder (ADHD), 26
Autonomic nervous system, 24, 25 Growth rate, estimation of, 5
Ax, Albert, 19
Haploid mycelia, 3
Basidiospore, 3 Heart rate, 19
Behavior and Information Heterozygosity
Exchange, 29 in nuclear genetic loci, 4
Behaviors Honey bee (Apis mellifera), 10
age appropriate, 26 infrared thermogram of, 11
emergent property, 25 metabolic heat, production
expressive, 18 of, 14
Brood cells, 1112, 13 thermal homeostasis, 14
thorax temperatures, 12, 13
Cannon, William, 2122 Human brain, xiii
Central nervous system (CNS), 27 Hyphae, 2
Colonies, 33
size, 37 Individual organism, definition
Cooperation, 33 of,114
emergent property, 40
mechanisms of, 29 Karban, Richard, 2930
Darwin, Charles, 29 Lie-detector, 19

The Expression of The Emotions in
Man and Animals, 17, 18 Mass, estimation of, 5, 10
Diastole, 20 Mating types, in haploid
Diploid cell, 3 mycelia, 3
Division of labor, 34 Methylphenidate, 2728
DNA fragments, patterns of, 3 Microsatellite DNA, 7
Dominance hierarchy, 34, 35, 40 Mock, Karen, 710
48 INDEX
Multicellular organisms, 14 Random amplification of

Muscle tension, 19 polymorphic DNA
Mushrooms, 2 (RAPD), 4
wet mass of, 6 and RFLP data, 4
Mutation, 10 Respiration rate, 19
Mycelium, 2 RFLP
and RAPD data, 4
Natural selection, 18, 32 Rhizomorphs, 3
Nervous and endocrine systems,
1728 Sagebrush (Artemisia tridentate),
2930, 40
Paper wasps, 33 Sampling scheme, 7
P. Canadensis, 33, 35, 37 Self-incompatibility
P. fuscatus, 33 of fungi, 3
Physiological response, 19 Skin conductance, 19
changes in, 20 Smooth muscle, 21
Polistes, 33 Stem, genotype of, 78
Polygraph, 19 Subordinate, workers, 34
Polymerase chain reaction (PCR) Superorganism, 14
primers, 4 Systole, 20
Quaking aspen Tibbetts, Elizabeth, 3839

(Populus tremuloides), 7
genetic analysis of, 8 West-Eberhard, Mary Jane, 33, 35
OTHER TITLES IN OUR BIOLOGY
COLLECTION
Behavior and Information Exchangeby Christopher J. Paradise and A. Malcolm Campbell

Cells in Tissuesby Christopher J. Paradise and A. Malcolm Campbell
Ecological Dynamicsby Christopher J. Paradise and A. Malcolm Campbell
Evolution of Interactions in Communitiesby Christopher J. Paradise and
A.Malcolm Campbell
Evolutionary Historyby Christopher J. Paradise and A. Malcolm Campbell
Effects of Genetic and Pathogenic Diseases on Cellsby Christopher J.Paradise and
A.Malcolm Campbell
Information in the Environmentby Christopher J. Paradise and A. Malcolm Campbell
Mechanisms of Evolutionby Christopher J. Paradise and A. Malcolm Campbell
Properties in and of Populationsby Christopher J. Paradise and A. Malcolm Campbell
Variation and Population Geneticsby Christopher J. Paradise and A. Malcolm Campbell
Ecological Homeostasisby Christopher J. Paradise and A. Malcolm Campbell
Ecological Interactionsby Christopher J. Paradise and A. Malcolm Campbell
Organismal Homeostasisby Christopher J. Paradise and A. Malcolm Campbell
Population Homeostasisby Christopher J. Paradise and A. Malcolm Campbell
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EBOOKS Emergent Properties of Individual
FOR THE Organisms
APPLIED BIOLOGY COLLECTION
Christopher J. Paradise A. Malcolm Campbell
SCIENCES
LIBRARY This book begins by describing what an individual organism is,
comparing preconceptions of the individual to non-standard
Create your own ways of thinking about individuals. Variation in what individuals
Customized Content are is described, using giant fungi, clonal trees and honey bee
hives as examples. Individuals are thus shown to be emergent
Emergent
Bundlethe more
properties. Other emergent properties of individuals are also
books you buy,
described. Classic experiments that elucidated the source of
the greater your
Properties
emotions in humans and other mammals are described. Emo-
discount! tions arise from the actions of the nervous and endocrine system
and often include a variety of signals given to other individuals of
THE CONTENT
Energy Physics
the same or different species. In particular, this book focuses on
fear and anger, two emotions that are closely related and often
confused, but that have been well studied. In one final example
of Individual
Engineering
Biotechnology
of emergent properties of individuals, cooperative behavior is
analyzed. The behaviors displayed by individuals that facilitate
cooperation among individuals and why those individuals may
Organisms
Biology
actually cooperate instead of compete when acquiring resources
Mathematics
or defending against predators are discussed.
Chemistry
Christopher J. Paradiseis professor of biology and environ-
mental studies at Davidson College. He teaches introductory
THE TERMS
biology, ecology, entomology, and topical seminars on ecotoxi-
Perpetual access cology and renewable natural resources. He also occasionally
for a one time fee leads a study abroad program in India. His research evaluates
No subscriptions or anthropogenic factors that influence insect biodiversity at a
access fees variety of scales. His current research interests include effects of
land use patterns on pollinator communities in parks.
Unlimited
concurrent usage A. Malcolm Campbellteaches biology at Davidson College,
Downloadable PDFs NC. He received national and international education awards:
Free MARC records Genetics Society of America (2013); American Association for
the Advancement of Science (2012); and American Society for
For further information, Cell Biology (2006). He was the founding co-editor in chief of
CBE Life Sciences Education; founding director of Genome
a free trial, or to order,
contact: Consortium for Active Teaching (GCAT); and member of the Christopher J. Paradise
sales@momentumpress.net American Society for Cell Biology governing council (20122014)
A. Malcolm Campbell

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EBOOKS Emergent Properties of Individual

FOR THE Organisms

Christopher J. Paradise, PhD

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

First published in 2016 by

ISBN-13: 978-1-60650-963-0 (print)

Momentum Press Biology Collection

Cover and interior design by S4Carlisle Publishing Services Private Ltd.,

Printed in the United States of America

The Definition of the

A commonly held understanding of an individual organism may be clear:

Figure 1 Fungi. A, Armillaria gallica mushrooms. B, Fungal hyphae.

cycle of Armillaria species is the formation of rhizomorphs, aggregations of

used random amplification of polymorphic DNA (RAPD) to look for

Table 1 Estimates of growth and size of large

Source: From Smith et al., 1992, Figure 1 legend.

Some scientists have questioned whether this enormous individual

Figure 2 Aerial photograph of aspen forest sampled by Mock and

Table 2 Genetic analysis of quaking aspen stand in Utah.

Source: From De Woody et al., 2008, text.

aerial photograph of the site. One genotype was found in a majority of

Mock and her colleagues concluded that there were up to 41 individ-

of the same individual but had undergone a mutation while growing

Figure 3 Infrared thermogram of honey bees. On the left and right

Figure 4 Thorax temperatures of worker honey bees in different hive

higher than their head and abdomen. Percentages of workers of differ-

percentage of worker bees

Figure 5 Percentage of bees with thorax temperatures 0.2o C above

2 days old. Insect development is temperature dependent. Developing

Stabentheiner A, Kovac H, Brodschneider R: Honey bee colony

The Nervous and Endocrine

Table 3 A sampling of expressions in animals studied by Charles

Source: From Darwin, 1872, text.

Although vocalizations and facial expressions also indicate other emo-

Figure 6 Changes in physiological variables in humans in response to

Table 4 Results from experiments investigating adrenalin in cats.

Source: From Cannon, 1911, text.

effects of adrenalin are discussed here, a variety of other hormones af-

70 change in heart rate

110 change in blood pressure

to electrical stimulation of the splanchnic nerve. This might indicate a

Ethical, Legal, Social Implications:

According to pediatricians, treatment with methylphenidate works

Individuals of Some Species

Information transfer occurs frequently in groups of animals living to-

Figure 8 Frequency distribution of number of foundresses on colonies

There is a division of labor within these paper wasp colonies. Queens

The dominance of one wasp over the other leads to stimulation of

Figure 9 Relationship between colony size and number of foundresses

More foundresses in P. fuscatus colonies led to a disadvantage in

differences, there are similarities that lead to an emergent property in

nest. Alternatively, females may be able to detect related females, although

To test these hypotheses, Tibbetts experimentally altered the mark-

Pardi L: Dominance order in Polistes wasps, Physiol Zool 21(1):113,

Darwin, Charles, 29 Lie-detector, 19

Multicellular organisms, 14 Random amplification of

Quaking aspen Tibbetts, Elizabeth, 3839

Behavior and Information Exchangeby Christopher J. Paradise and A. Malcolm Campbell

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