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A. Malcolm Campbell
Behavior and Information
Exchange
Behavior and Information
Exchange
Christopher J. Paradise, PhD

A. Malcolm Campbell, PhD
Behavior and Information Exchange
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
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brief quotations, not to exceed 250 words, without the prior permission
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First published in 2016 by

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Abstract
Animal behavior includes the exchange of non-heritable information be-
tween individuals of the same species. Animals exchange information for
a variety of reasons, including mating, defense, and cooperation, and all of
these situations will be discussed. This book will describe the functions of
communication and information transfer between organisms and explain
how animals communicate and find each other through use of different
signals. The costs and benefits of using various signals will be evaluated,
as will the costs and benefits of living in groups. Playback experiments
and the comparative method are approaches used for understanding and
interpreting signals used by organisms to communicate information to
other members of the same species. Plants also communicate information
between individuals, often for purposes of species identification during
mating. Female reproductive structures in plants recognize pollen from
members of the same species. Finally, the commonalities and differences
between animal and plant communication will be identified.
Keywords
communication, information, bioluminescence, vocalizations, playback
experiments, social animals, derived traits, comparative approach, sexual
reproduction, pollination, inbreeding, courtship behavior, mating
Contents
Preface...................................................................................................ix
Acknowledgments....................................................................................xi
Introduction.........................................................................................xiii
Chapter 1 Information is Transmitted Between Members
oftheSame Animal Species................................................1
Simple Communication in a Firefly...................................2
More Complex Communication in a Bird..........................7
Ethical, Legal, Social Implications: Love at First Sight
Has Biological Causes and Can Have Ethical and
Social Consequences.....................................................13
Chapter 2 Group Living Requires More Derived Mechanisms
ofInformation Transfer....................................................17
Chapter 3 Plants of the Same Species Recognize One Another
Through Pollen-Pistil Interactions....................................27
Conclusion............................................................................................37
Glossary................................................................................................39
Index....................................................................................................41
Preface
This book behavior and information exchange between individuals of the
same species 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 con-
text and comprehensive perspective so that readers can learn important
information from a single book with the potential to see how the major
themes span all size scales: molecular, cellular, organismal, population
and ecologic systems. The major themes of biology encapsulate the en-
tire discipline: information, evolution, cells, homeostasis, and emergent
properties.
In the twentieth century, biology was taught with a heavy emphasis
on long lists of terms and many specific details. All of these details were
presented in a way that obscured a more comprehensive understanding.
In this book, readers will learn about behavior and how behavior is part
of communication and information exchange between individuals, and
some of the supporting evidence behind our understanding of those phe-
nomena. The historic and more recent experiments and data will be ex-
plored. Instead of believing or simply accepting information, readers of
this book will learn about the science behind behavior and information
exchange 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 turning
them into powerful and elegant Bio-Math Explorations. I learned a lot
from both of them. While the math is largely absent from this book, our
collaboration with her made this a better book. Nancy Stamp at Bing-
hamton University, and Bill Dunson and Richard Cyr at The Pennsyl-
vania 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 experi-
ment to redesign introductory biology, starting with the text and ending
with a new approach to teaching biology.
Introduction
People often find it difficult to get their point across to other, or wonder
why no one told them about the important assignment they missed when
they were absent from class or missed a day of work. Communication
between individuals is a challenging part of everyday life for humans,
and miscommunication often leads to conflict and even wars. But with-
out communication, people would have no friends, and would eventually
find it hard just to survive. Many animals exchange information with
each other in response to their environment. People hear animals make
noises or see them display behaviors all the time in order to communicate
with other members of their species or with other species. However, what
people may not realize is that plants and even single-celled organisms
send signals to each other using simpler forms of communication, which
are hardwired into their DNA. As with humans, miscommunication can
lead to trouble for individuals, but it also leads to variation, which is a
key component of evolution. In this book, information at the population
level in the form of communication in animals and plants will be exam-
ined. Through careful observation and controlled experiments, scientists
can interpret the signals and understand these organisms better, and per-
haps even uncover a lesson for human communication.
When studying information at the organismal level in Chapter 3, the
focus was on information within organisms. In this chapter, information
exchanges between individuals within a population, which is a group of
individuals of a species living in a particular place at a particular time, will
be explored (Figure 1). Individuals of the same species exchange many
kinds of information, from chemical to auditory to visual. Figure 1 illus-
trates communication among animals, in this case birds, many of which
use visual displays, vocalizations, or both to convey information.
One of the four themes of the fundamental concept of information
is that non-heritable information is transmitted within and between bio-
logical systems. A bird that performs a courtship dance is transferring
information to a potential mate and that information is non-heritable.
xiv INTRODUCTION
receiver
sender of
visual or auditory
information
eavesdropper
(unintended
receiver)
Figure 1 A population of birds and information transfer between two

individuals with one other individual also receiving the information.
Source: C. Paradise.
If all of the information meant to be transferred is actually transferred

to the receiver, then another theme is fulfilled in that the information
is expressed and regulated by the sender without loss of content. If this
leads to mating and reproduction between the two individuals, then the
theme that heritable information provides for continuity of life comes
into play. However, readers should recognize that courtship displays are
not performed equally well by all individuals, and so information transfer
is imperfect and that leads to variation among individuals, which is an-
other of theme of information. In this book, information transfer among
individuals from a variety of species, including animals and plants, will
be explored.
CHAPTER 1
Information is Transmitted
Between Members of the
Same Animal Species
In 1950, the famous animal behaviorist Konrad Lorenz wrote, Animals

do not possess a language in the true sense of the word . . . every indi-
vidual has a certain number of innate movements and sounds for express-
ing feelings. Lorenz was rewarded with a Nobel Prize in Medicine or
Physiology for the contributions that he made to our understanding of
how animals communicate with each other and transmit information so
that it is satisfactorily received or understood.
Lorenz studied a species of highly social goose that has a system of sig-
nals produced and understood by every member without any previous ex-
perience. The coordination of social behavior through visual and auditory
signals gives the appearance that the birds are talking and understanding
a language of their own. Many of Lorenzs discoveries came from carefully
observing animals interacting with each other, which is something that
any biology student with a little patience can do. As Lorenz pointed out,
most animals understand the language of their species without any prior
experience; the ability to communicate is heritable information encoded
in DNA. Yet the act of communication represents non-heritable informa-
tion transmitted between individuals. Many animals interact with other
members of their own species, and the functions of their communication
fall into one of several categories.
Species recognition, mate/sex recognition, mate attraction, defense
against threats, facilitation of cooperation, and warning of danger are
some of the major functions of communication. For instance, mute swans
perform a courtship display, which functions to verify that the receiver
2 BEHAVIOR AND INFORMATION EXCHANGE
is a member of the same species and that precedes mating. A receiver is

an individual who detects a message, intentionally or accidentally. Com-
munication is used to defend against potential threats, as a cheetah does
when it raises its hackles, or used to facilitate cooperation, as when wolves
work together to bring down large prey, such as a bison. Finally, commu-
nication is used to warn other members of the same species of potential
danger, as a deer may do when it flashes the white underneath their tails.
There are costs and benefits associated with exhibiting behaviors. For
example, finding a mate consumes a lot of time and energy and may
be dangerous. A small expenditure of time and energy to assure that a
receiver is a member of the same species may offer significant selective
advantages in the long run. Similarly, displaying oneself with raised fur
represents a small cost with the potentially great benefit of removing the
threat. Cooperative behavior requires information transfer to more ef-
ficiently hunt or coordinate an escape. Behavioral scientists conduct re-
search to determine the selective advantage of various behaviors and how
signals are perceived by other members of the same species.
Simple Communication in a Firefly

In this section, the signals involved in mating and how scientists decode
these signals will be studied. One example of such signals is in the light-
ning bug, or firefly. Fireflies of the Photinus genus have yellow segments
on their abdomensthose are the segments that glow through a bio-
chemical reaction that consumes energy. It will be instructive to examine
this species first, because they have very simple signals that they use to
communicate, and the function of their communication is to find mates.
Fireflies are beetles, which are insects, and they have evolved chemi-
cal and visual ways to communicate with flashing lights. On the surface,
bioluminescence, which is light production as a result of a chemical re-
action, may seem dangerous to the individual because it might attract
predators and be selected against. The signals might also cost time and
energy if they attract members of a different firefly species. There are
many species of fireflies, and many overlap in where they live. In order for
flashing bioluminescence to have evolved, the evolutionary benefits must
outweigh the risks, and the signals must be specific for each species.
INFORMATION TRANSMITTED WITHIN SPECIES 3
Male Photinus greeni fireflies live in the eastern United States and ac-
tively signal to females shortly after sunset during the summer. Males fly
about 1 meter above the ground and flash a signal to any female that may
be within 10 meters. The bioluminescent signal is a double pulse of light
that is separated by an interval of time. They repeat this combination
every 5 seconds or so, depending upon the species and, as will be seen,
other environmental factors.
Sara Lewis and her colleagues wanted to determine the variation in
the signal pattern produced by males when attempting to communicate
with a female. Using a digital camcorder, the investigators recorded 221
interpulse intervals (IPIs) from thirty males in Massachusetts. An IPI is
the amount of time between pulses of light. The flashes and the intervals
were short, but their camcorder recorded at a rate fast enough to capture
all of the action. Observations were made on different nights and there-
fore at several different temperatures. The scientists analyzed the videos to
measure the IPI for each male, and used regression, a statistical technique
of fitting an equation to real data points, to determine the best fit curve
for the data. They discovered a polynomial relationship between IPI and
temperature (IPI = 19.65 2.69x + 0.13x2 0.0022x3). The flash in-
terval, or interpulse interval, declines as temperature increases, but not
linearly.
Lewis and her colleagues quantified the variation among individuals.
Although the variation in time between light pulses due to temperature
was interesting, the investigators wanted to understand variation within
individual males, regardless of temperature fluctuations. Therefore, the
investigators used this regression curve to adjust each IPI to what it would
have been if it had been measured at 21.1o C. In other words, if a par-
ticular male firefly waited 3 seconds between pulses at 12.8o C, how long
would he have waited between pulses at 21.1o C? Part of the variation,
it was found, was due to temperature, because insects are ectotherms,
which are animals that regulate their body temperature using behavioral
mechanisms and heat from the environment. The rate of male signaling,
therefore, is partly dependent upon body temperature, which affects the
rate of chemical reactions inside an individual. Anyone who has watched
a bee or a butterfly on a cool spring morning would notice how slowly
they move compared to later in the day after the sun has warmed them
up. Low temperatures, below 15.6o C, appear to really slow down the
male fireflies and their signaling, and high temperatures shorten the IPI.
After adjusting for differences in ambient temperature, the scientists
produced a distribution of male IPIs that would be expected if all data
were collected at 21.1o C. In fact, there is a range of temperatures be-
tween 15.6o C and 21.1o C where IPI does not seem to be affected by
temperature; thus, the variation in IPIs is not due to temperature. Even
at a constant temperature, there is variation in pulse interval. Variation
among males, or even among signals produced by a single male, repre-
sents imperfect information transfer, which is one of the themes of infor-
mation transfer. The variation among males may be important to females
assessing information, or it may just be a consequence of an imperfect
biochemical pathway.
It can be seen from this example that fireflies transfer information
using visual communication. The data illustrate the relevant theme that
variation is inherent in information transfer and communication. After
the biologists discovered the variation in signaling by males at different
temperatures, they sought to determine how females would respond to
varying IPIs at different temperatures. Lewis and her colleagues tested
female responsiveness to simulated male signals at three different tem-
peratures: 15.6o C, 18o C, and 18.8 to 21.7o C. The highest temperature
tested was a range of temperatures, due to a small number of observations
at several higher temperatures. Male signals were double pulses of light-
emitting diode (LED) light of equal duration separated by five different
IPIs. The intervals ranged from 0.6 seconds less than the mean interval
to 0.2 degrees more than the mean interval for that temperature. For
instance, the mean IPI of males at 15.6o C was 1.8 seconds, so the IPIs
tested were 1.2, 1.4, 1.6, 1.8, and 2.0 seconds. Intervals were adjusted
downward for higher temperatures. The female response is a single flash
of light when exposed to the simulated male signal and the percentage of
females who responded was determined.
Just as there is variation in the IPIs of males signaling at a particular
temperature, there is variation in the female response to different IPIs. As
predicted, the greatest percentage of females respond at the average IPI
for a particular temperature (~90% of females tested at that IPI), except
at the highest temperature, when there is a slightly greater percentage of
females that respond at a slightly shorter than average IPI (~68% of fe-
males tested vs. ~65%). Males that have too short or too long an IPI for a
particular temperature may still induce a female to respond, although the
percentage of the females in a population that responds to one of those
signals is not as large as the average IPI percentage. In no case did 100%
of females respond. Even a correctly performed signal may be missed by
a female, or a female may misread the communication, causing her to fail
to respond.
When a female responds by flashing back to a male, he will flash again
in response. This leads to a dialogue of bioluminescent flashes, which is
an exchange of information much like when two humans have a conversa-
tion or dialogue. When a female responds, the male orients toward her and
flies closer. Dialogue continues until he reaches her, at which time mat-
ing begins. Different species vary in their pattern and timing of mating.
Photinus greeni courts in the early summer for about 90 minutes after sun-
set each night. Both males and females may mate on multiple evenings
over the course of the mating season. A higher percentage of females re-
spond to males whose pulse intervals are shorter or longer than the mean
pulse interval for a particular temperature when the number of available
males begins to decrease over the summer. Advertising their location with
light pulses increases the males risk due to predation, and over time, the
number of available males decreases. Although light may attract predators,
it is thought that predators have a harder time finding a flashing light than
one that is continuous. This also makes it more difficult for children chas-
ing fireflies at night and losing them during the dark period of their flash-
ing cycle. As a result of greater risk of predation for males than females,
females become less choosy as mates become more difficult to find.
Scientists have hypothesized that species recognition is involved in the
evolution of the flash pattern. In another test, this time of three closely
related species, including P. greeni, another researcher, J. Lloyd, tested this
hypothesis by measuring the response of P. greeni females to biolumines-
cent signals of males from the same species, along with two other closely
related species (Figure 2). Males of all three species signaled to females
with a double pulse of light, and the IPIs are shown in the x-axis.
As shown in Figure 2, P. greeni females did not respond to signals de-
signed to simulate the flash pattern of males from the other two species.
100
female P. greeni exhibiting response (%)
90 pulse interval
of male P. greeni
80
70
60
50
40
30
20 pulse interval pulse interval
of males of of males of
10 species A species B
0
0 0.2 0.4 0.6 0.8 1 1.2 1.4 1.6 1.8 2 2.2 2.4
tested interpulse interval (IPI, sec)
Figure 2 Female P. greeni were tested for their responsiveness to
pulse intervals of varying duration that were known to be found in
males of three different species (sample size 5 1 to 3 for each IPI
tested). The range of pulse intervals for males of each species is
indicated by the three different brackets.
Source: Data from Lloyd, 1969.
The function of firefly communication seems to be twofold. First, the

flash pattern is used to verify that the receiver and sender are members
of the same speciesfemales do not reply if the proper signal is not sent.
The consequences to females that respond to a male of another species are
that she wastes her time and energy dialoging and possibly mating with
an inappropriate male and that she risks missing an opportunity to mate
with a male of her species. The second function of firefly communication
is for a male to locate a receptive female through the visual dialogue.
Fireflies do not learn the language of fireflies. It is encoded in their
DNA. Without a dialogue, mating would be very difficult because males
and females of the same species would rarely find each other by crawling
and flying around aimlessly. Information transfer at the population level
allows individuals to communicate their identity to each other. Com-
munication provides for the continuity of their species. Although there
is variation in the signals, scientists have shown that over 70% of the
variability observed in male IPIs appears to be caused by differences in
temperature. Further, a proportion of the female population does not

reply to every males signal. Females either fail to see the pulse, or they do
not recognize it as the signal of a potential mate. The females biochem-
istry is also affected by temperature, which adjusts her understanding of
the pulse interval to account for changes in temperature. And it turns
out that the signals of the three species in Figure 2 have evolved enough
distinction to eliminate any overlap and miscommunication. Information
may be lost or an attempt at communication may fail because of inher-
ent variation within a species. But females of one species typically do not
respond to signals of males of other species.
More Complex Communication in a Bird

Each species of firefly has a particular way of communicating its identity,
but its vocabulary is limited. Other animals, especially those with more
complex communication patterns, vary in what they say as well as how
they say it to one another. For example, the vocalizations, or acoustic
signals, of birds and other animals have a variety of functions. People will
not be able to understand what birds are communicating through their
singing, but most people have no doubt eavesdropped on the dialogues
of birds. In this section, the understanding of animal communication will
be expanded by examining a bird that uses vocalizations for more than
just finding a mate. As will be seen, this type of language is slightly more
complex than the fireflys language, but the species to be studied has an
instinctive and fairly simple language relative to many other birds.
There are over 8,500 species of birds, and most use vocalizations to
transfer information to other members of their species. Animals such as
fireflies and birds that are very mobile and need to find partners of the
same species to mate need a mechanism of communication that can func-
tion across large distances. Birds use vocalizations to communicate with
each other, and communication has the same functions discussed earlier,
including species recognition, mate/sex recognition, mate attraction, de-
fense against threats, facilitation of cooperation, and warning of danger.
Each species of bird may have unique vocalizations for each of these func-
tions. Birds also may use visual signals such as courtship dances during
mating, and individuals often combine visual signals with vocalizations.
There are about twenty species of birds called storm petrels, which live
most of their lives over the oceans. They get the name storm from the
fact that they often hide behind ships during storms. The name petrel
(derived from the Christian disciple Saint Peter) comes from their habit
of skimming across the water with their legs hanging down, which makes
the birds appear as if theyre walking on water. While skimming, these
sea-going birds feed on crustaceans and small fish, which they pluck out
of the water.
The breeding season is the only time that the storm petrels come to
land. They breed in colonies, mostly on islands, and dig burrows or use
small crevices on rocky slopes for nests. When they have a chick in the
nest, they only visit at night to feed their one offspring (females lay a
single egg each breeding season). These birds are monogamous and pairs
form long-term bonds that last for many years.
Wilsons storm petrel (Oceanites oceanicus) is a common and widely
distributed species of storm petrel. There may be thousands of birds in a
single colony, so one can imagine the cacophony of sound produced by all
those birds vocalizing. Because islands have a limited number of good bur-
row sites and a breeding colony can contain many birds, Wilsons storm
petrels must be able to find their nest and defend it against others who
may want to steal it for their own use. Competition for nest sites might
be intense. How can an individual bird find its mate within a breeding
colony while all the other Wilsons storm petrels are also squawking in a
cacophony of sound?
Vincent Bretagnolle studied a colony of Wilsons storm petrels on the
Kerguelen Islands in the southern Indian Ocean. He recorded acoustic sig-
nals from hundreds of birds to determine the full range of vocalizations, or
calls, of the birds. There were many petrels at the colony, so he placed the
microphone at two strategic points: at the mouth of a burrow to capture
calls from within, and on the ground away from burrows to capture calls
of petrels standing near the entrance of the burrow. Acoustic signals were
transformed into sonograms, which are images of sound frequencies over
time. Sounds produce pressure waves, and waves will have higher or lower
frequencies, depending on whether they are high or low pitched.
From the sonograms, Bretagnolle described several different types
of calls, which varied in total duration, number of syllables, duration of
syllables, and duration of silent periods. Periods of silence are analogous

to the IPIs in male firefly signals or pauses in human speech. There were
two types of grating calls, a short version and a long version, which were
the most common vocalizations used by both males and females. Grating
calls were usually made from within the burrow, and each type had a vary-
ing number of syllables, depending on the individual and situation. Male
storm petrels also produce two types of chattering calls outside the bur-
row, and chattering calls also have a variable number of syllables. If a male
made a chattering call from within a burrow, he also gave the grating call.
Bretagnolle also made note of the situations the caller was in when
making the call. Along with each call, he recorded whether the caller was
in its burrow, whether another petrel was flying overhead when the call
was made, whether it was face to face with another petrel, and the sex
of the other petrel with which the caller was interacting. Because males
produced a greater diversity of calls, Bretagnolle analyzed the frequencies
of types of male calls in different situations.
When a bird is alone it was found to utter the chattering call 40% of
the time, the chattering and grating calls 36% of the time, and the grat-
ing call 24% of the time. When a bird is flown over by another bird, it
utters the chattering call 48% of the time, the chattering and grating calls
26% of the time, and the grating call 26% of the time. When a bird is
facing another bird, it utters the chattering call only 9% of the time, the
chattering and grating calls 17% of the time, and the grating call 74% of
the time. For these data, the scientist did not note whether the caller was
a male or a female. But for another set of data in which he distinguished
between the two grating calls, Bretagnolle found that when males were
facing males, 64% of the time they used grating call #1 and 36% of the
time they used grating call #2. When a male faced a female, only 11%
of the time did they use grating call #1, and 89% of the time they used
grating call #2.
Bretagnolle wanted to determine if the calls used by a petrel depend
on the situation in which the bird finds itself. The investigator wanted to
know if the percentages change significantly when the petrel was alone,
flown over by another bird or faced another bird, or whether the sex of
the receiver made a difference. He used a statistical technique called a
chi-squared test (denoted by 2, the Greek letter chi [pronounced ki,
as in kite] raised to the power of 2) to compare each pair of situations.

The chi-squared test is a statistical method to decide whether or not two
distributions are significantly different.
When a male is not facing another bird, he frequently is chatter-
ing or chattering and grating. These calls may function to advertise the
males presence to females within hearing range, or to announce to other
males that the caller is a male with a burrow. Bretagnolle found that there
was no statistical difference in the calls uttered between a bird being alone
or being flown over by another bird (X 2; p = 0.36). However, there was
a strong statistical difference when comparing a bird facing another bird
and either of the other two situations (p < 0.01 for both situations).
A small p-value, or probability, means there is a statistically significant
difference between two situations. In other words, facing another bird
leads to a caller using calls in different frequencies than when it is alone
or being flown over by another bird. When a male faces another bird in
front of his burrow, he most often uses one of the grating calls. When the
scientist analyzed the frequency of the two grating calls, he found that
there was a statistically significant difference in grating call #1 was used sig-
nificantly more times when facing a male, while grating call #2 was used
more when facing a female. Bretagnolle concluded that grating call#1 is
used to defend the burrow when facing a male, and grating call#2 is used
for courtship when facing a female.
After recording all of these vocalizations, Bretagnolle performed a
series of playback experiments, where taped vocalizations were played
through loudspeakers near the colony or a burrow entrance. Playback
experiments record signals such as light flashes or vocalizations and play
them back to animals in controlled conditions to study their responses.
The playback experiment was designed to test the response of individual
petrels to vocalizations made by other individuals, in a manner similar to
the firefly experiments where researchers flashed simulated light patterns.
Think of these playback experiments like a game of Marco Polo. The
scientist plays Marco and determines whether any birds respond with
Polo. The difference here is that the scientist records the responding
birds behavior, whether it is another vocalization or whether it is some
non-acoustic behavior. By repeating the playback many times in many
situations, scientists can to decode the language of the subject species.
4.5
mean # of petrels responding ( 1 s.e.)

4
= direct flight
3.5 = circle flight
= landed
3
2.5
1.5
0.5
0
chattering grating chattering no call
and grating
playback call
Figure 3 Mean number of Wilsons storm petrels responding to
different acoustic signals during a playback experiment. Error bars
represent the standard error (SE) of the mean.
Source: Data from Bretagnolle (1989) Table 1.
Inthis case, Bretagnolle recorded any vocalizations and described in

writing any non-vocal responses after playback of chattering, grating, or
chattering plus grating. He calculated the average number of petrels re-
sponding during a playback with data compiled over 20 days of perform-
ing trials (Figure 3).
Additionally, he recorded individual birds vocalization responses
when subjected to chattering, male grating, or female grating. When ex-
posed to chattering or the male grating call, males mostly responded with
the male grating call, but when exposed to the female grating call, no
male responded. Females did not show a strong preference for any call,
although about 37% responded with the female grating call when they
heard chattering, and only about 20% responded with grating to either
male or female grating calls.
Male Wilsons storm petrels make two types of chattering calls and use
grating call #1, but females do not. Both males and females use grating
call #2. The chattering calls appear to be attractive to some females, and
may be a male sexual advertising call. If a male has a nest, he may call to
advertise that fact, using the chattering call outside the burrow and both
a chattering and a grating call from within the burrow. Remember that
there are many birds in the colony, and repeating a call many times can
help a female find a male in spite of other potential callers in the area. If
a female already has a mate, she will ignore the chattering even though
the sound is intended to recruit females; she may also miss a call in all the
noise. It seems that Wilsons storm petrels use vocalizations as a means of
gender recognition.
Males respond with the grating call to both types of male vocaliza-
tions. Grating sounds may be a signal to other males to keep away, per-
haps because burrow space and females are limited resources that force
males to compete. Bretagnolle concluded that Wilsons storm petrels used
the grating calls to advertise sexual identity, whereas males used the long
grating call (#1) when they had detected another male in the vicinity.
For many bird species, vocalizations are instinctual. Other studies have
determined that this is true for storm petrels. For some bird species, like
mynah birds and many songbirds, research has shown that young birds
have to learn the songs of the species.
The observations of individual interactions and the playback experi-
ments helped Bretagnolle decode this species vocalization signals. There
are other calls petrels make that were not analyzed, but this and other
studies reveal that communication is used by this species to transfer in-
formation about gender identification and possession of resources. Petrel
vocalizations convey important information for receivers to allow deter-
mination of the sex of senders, occupation of a burrow, hunger status, or
level of distress. The reception and ability to understand this information
is encoded in their DNA and could cause a change in a receivers behavior.
The instinctual aspect of communication in petrels is important. Calls
varied enough from each other, despite bird-to-bird variation, that receiv-
ers frequently understood the signal, even if they did not always respond
to it. The variation in petrel vocalization is analogous to humans with
different accents speaking the same language. However, with our limited
understanding of the languages of other species, we cannot determine
whether variations in vocalization represent a loss of information content,
or understanding by a receiver who chooses not to respond.
The range of information signals varies among speciessome have
more complex languages than others. Based on data from experiments
with fireflies and storm petrels, it can be concluded that storm petrels
communicate more information to each other than fireflies. Information
at the population level is illustrated by animal communication. In ani-
mals that have more complex living situations, individuals often perform
behaviors that, at first glance, look to have costs associated with them
with little apparent benefit. A large body of research has decoded the sig-
nals of animals living in herds, schools, or extended family groups. In the
next chapter, the complexities of information transfer in a group living
situation will be examined.
Ethical, Legal, Social Implications: Love at First

Sight Has Biological Causes and Can Have Ethical
andSocialConsequences
Many people may have experienced love at first sight. When love at first
sight occurs, information is exchanged between two people in the form
of both visual and chemical cues. That information, conveyed through
visual and olfactory senses of another person, may be sufficient to cause
the receiver to fall head-over-heels in love. These changes in emotional
state may well lead to some social and ethical issues surrounding this type
of information transfer.
When two people meet, both individuals are simultaneously senders and
receivers of information. Both may react with alterations in brain chemistry
and body functioning. Flushed cheeks, sweaty palms, and a rapidly beating
heart are all signs that a human may be experiencing intense emotion. Other
signs are not as easy to observe. Human behavior can change dramatically
in response to hormones and neurotransmitters, including dopamine, sero-
tonin, and adrenaline, which can fluctuate during this encounter.
When infatuation occurs, the object of affection is often all a person
thinks about. The social implications of intense emotions could lead to
obvious adverse health effects and poor job or academic performance. Sev-
eral chemicals may play an important role in altered behavior. Dopamine,
which is both a hormone (released by the hypothalamus) and neurotrans-
mitter (produced in several parts of the brain), creates strong connections
between pleasure and a persons love interest. Dopamine acts very much
like an addictive drug.
Dopamine increases during infatuation while another neurotransmit-

ter, serotonin, decreases. Serotonin moderates aggression, body tempera-
ture, appetite, sleep, and sexuality. Low levels of serotonin are associated
with high sexual activity but are also associated with anxiety and depres-
sion. Adrenaline, another hormone, is released by the adrenal glands,
which sit atop the kidneys. Adrenaline makes us sweat and causes our
heart to beat fastit is the hormone involved in the fight or flight
response. But the sweaty palms and fast heart rate are also signs of love
at first sight.
Most of these hormonal effects last for a relatively short time. How-
ever, they could last long enough for a relationship to develop and ma-
ture. Serotonin levels may not return to normal for a year. In the early
stages of a relationship, individuals who formed strong bonds and were
fixated on their mate may have had more success evolutionarily, pass-
ing their genes on to more offspring than individuals who did not have
these traits. Strong bonding behavior would have spread throughout the
human population.
These are just some of the chemical changes that take place in people
who fall in love at first sight. Several hypotheses have been put forth re-
garding the initial attraction that one feels toward another human. If a
person is primed for a sexual encounter, or mating, they may assess the
quality of the other person and the likelihood that they have good genes.
A persons stature, physical appearance, and social status are all potential
clues as to the quality of his or her genes. Symmetry may also be an
important clue to the health and genetic fitness of the signaler. Men,
especially, are drawn to women who have symmetrical faces. Physical ap-
pearance is thus one signal that a receiver detects.
Another hypothesis supported by evidence is one in which humans
and other animals are attracted to individuals who have both looks and
smell similar to the receivers parents. One scientist found that human
subjects prefer faces similar to their own and suggests a preference for a re-
semblance to the parents. In one study, it was discovered that individuals
preferred the smell of others whose immune systems were different from
their own. In another study, women were found to prefer the smell of a
man who is similar to their father. It was speculated by scientists that the
immune system would still be different enough from her own to provide
a wide range of genes for immunity, but similar enough to her own to be
a known, successful quantity.
Finally, there are certain areas of the brain that become active when
subjects are shown pictures of the object of their love. One active area also
responds to euphoria-inducing drugs, such as heroin, and another area
is responsible for instinctual feelings. Conversely, the prefrontal cortex,
known to be overactive in depressed patients, becomes less active when
seeing the object of ones affection.
Now that we have an understanding of love at first sight, we can explore
some of the social implications. Both smell and looks may be important
information to a receiver, but that information is often sent unwillingly or
unknowingly. Although infatuations can be mutual, in other cases love is
not reciprocated. This could have strong social implications, which could
lead to obsessions, depression, stalking, and rapeall caused, in part, by
internal chemical signals altered by information from another person.
In some societies, it may not be permissible for people to marry out-
side their social group. Infatuation could lead to great difficulties in these
situations. If a relationship begins from infatuation, what are the chances
that it will last? This type of romance does not automatically mean
love forever. Release of another hormone, oxytocin, has been shown to
strengthen the bond between a couple, but people do not continue to
secrete high levels of this hormone after being in a long-term relationship.
Second, if parents or a wider social group do not approve of a relation-
ship, lovers could be ostracized or harmed.
Are there other ethical considerations? Consider Paris, prince of Troy,
who became infatuated with Helen of Troy. They ran away together and
this started a war. Helen was married to the king of Sparta at the time.
Her husband was not happy with his wifes affair and launched an ar-
mada, which led to a 10-year war. Although all infatuations may not lead
to international incidents, if infatuation occurs between individuals mar-
ried to other people, extramarital affairs could occur. Besides the obvious
ethical quandary, other harmful social interactions are bound to follow,
including divorce, estrangement from children, and bitter legal proceed-
ings. However, being complex social animals with high cognitive abili-
ties, humans, when armed with a biological understanding of love at first
sight, should be able to avoid such entanglements in theory.
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sexual recognitions and geographic variation, Behaviour 111:98112,
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Bretagnolle V, Robisson P: Species-specific recognition in birds: an exper-
imental investigation in the Wilsons storm petrel (Procellariiformes,
Hydrobatidae) using digitalized signals, Can J Zoolog 69:16691673,
1991.
Lloyd JE: Flashes, behavior and additional species of Nearctic Photinus
fireflies (Coleoptera: Lampyridae), Coleops Bull 23(2):2940, 1969.
Michaelidis C, Demary K, Lewis SM: Male courtship signals and female
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Behav 28:248252, 2007.
CHAPTER 2
Group Living Requires More

Derived Mechanisms of
Information Transfer
Fireflies and petrels communicate to identify and attract potential mates,

and in addition petrels communicate to protect personal space and de-
fend burrows or territories. The question at the heart of this chapter is
whether social animals, which live in groups and interact extensively
with others in the group, have evolved specialized communication sys-
tems. In other words, do animals living in groups have derived traits,
adaptations unique to that species and not found even in closely related
species, for information transfer? What additional messages are sent and
received by social animals? Any trait, whether its behavioral or physi-
cal, can be derived. For example, zebras have striped coloration that is a
derived trait when compared to other closely related horse-like animals.
One group of researchers, led by Marta Manser, investigated com-
munication among meerkats (Suricata suricata), which is a social species
of mongoose. Mongooses are small, slender mammals that feed on small
vertebrates and invertebrates, and they use a variety of vocalizations to
communicate with each other. Of the 30-odd species of mongooses living
in Asia, Europe, and Africa, most of them are solitary, although a few are
social.
In order to address whether the evolution of derived traits offers a
selective advantage, scientists often use a comparative approach, which
is a methodology where multiple species are studied to compare traits and
their functions in order to understand the evolution of those traits. To in-
vestigate the evolution of information transfer in social mongooses using
the comparative approach, a scientist would ideally want to have data on
the vocalizations, ecology and behavior of all mongoose species. Although

there are many studies on information transfer in social mongooses, soli-
tary mongooses such as the Cape grey mongoose (Galerella pulverulenta)
have not been well studied. Unfortunately, this is often the case in science,
where a lack of data can hinder hypothesis testing. Comparing two spe-
cies will still serve as a simple demonstration of the comparative approach
and will help determine if social mongooses have communication behav-
iors that are absent in solitary mongooses.
The Cape grey mongoose is diurnal, meaning it is active during the
day, and it lives in shrubby bush areas and feeds on small mammals and
insects. About 90% of Cape grey mongoose sightings are of a single in-
dividual. Most other observations are of pairs, particularly in July and
August. It is rare to see more than two mongooses together, although
groups of younger individuals are known to sleep together. This species
does not dig burrows but rather sleeps in areas of thick vegetation. In one
study, mongooses were trapped and fitted with radio collars. The loca-
tion of each mongoose was then determined multiple times over the next
several weeks to determine their home ranges within a national park in
South Africa, which is the area where an animal spends its time.
Home range boundary lines were drawn around the area where each
mongoose spent most of its time. Of 43 sightings of mongooses, 41 times
(95.3%) they were observed alone, and the other two times they were ob-
served with only one other mongoose. In another study, 89% of sightings
were of solitary animals, 10% of pairs, and 1% were in groups of three.
Pairs are more often observed in July and August.
Cape grey mongooses live mostly solitary lives, but there is a large
overlap in the home ranges of individuals; so the probability of one mon-
goose crossing paths with another is greater than if home ranges did not
overlap. It would be useful to know the behaviors exhibited by a mon-
goose upon seeing another mongoose, including behaviors, expressions,
and vocalizations. In fact, the Cape grey mongoose does have some vo-
calizations and that is not surprising. A solitary mongoose has very little
reason to vocalize, because there would be few friends around to hear it
and there are many foes who could intercept the message. Although no
research has been done on their calls, a scientist might hypothesize that
the function of vocalizations made between two mongooses are to greet
GROUP LIVING REQUIRES DERIVED MECHANISMS OF TRANSFER 19
one another, to determine breeding status, or to chase an intruder from a

territory. When a mongoose observes a potential predator, vocalizations
tend to be restricted to danger signals, which are to make the predator
think twice before striking and possibly give the mongoose time to es-
cape. The Cape grey lacks a large vocabulary or range of vocalizations,
so it is unlikely that they are able to warn other individuals of danger.
Now that a little information is known about the solitary Cape grey
mongoose, the social meerkat can be studied to identify differences and
possible derived traits. Like the Cape grey, meerkats are diurnal, but they
live in grasslands and more open areas than the Cape grey mongoose
and feed mostly on insects. Meerkats live and forage in family groups, or
clans, ranging from 3 to 33 individuals.
Typically, living in groups requires behaviors not exhibited by solitary
animals. For instance, individual meerkats who do not have any offspring
help raise the young of other adults. Because they live in family clans, the
helpers are related to the parents and the offspring (cousins, aunts, and
uncles in human terms). Behaviors that facilitate cooperation, including
group defense or hunting, communication to and from offspring should
be and are observed in meerkats. The watching, or guarding, behavior
that is commonly observed in meerkats, for instance, might be such a
behavior that would allow individuals to perform certain behaviors while
others stay on alert for danger.
To determine how group living influenced communication, Manser
and her colleagues studied meerkats in a national park in South A frica,
observing behavior and recording vocalizations. The researchers noted
the presence of potential predators in the area, including jackals, eagles,
hawks, and snakes. Sentinel behavior, where one individual stands guard
while others dig in the sand to hunt for insects to eat, was observed 55%
of the time when members of the clan were foraging for food. Sentinels
typically climb a small hill or tree near where clan members are foraging
and stand on their hind legs, using their tail to make a tripod. Meerkat
sentinels were found to vocalize about 80% of the time that they were
on duty (i.e., 8 out of every 10 minutes), and when no predator or
threat was visible, sentinels used 1 of 6 different calls, all relatively short
and similar to one another, which Manser and her colleagues called
sentinel calls.
Based on her observations, Manser concluded that the single, double,

triple, and multiple note calls functioned to let foraging meerkats know
that a sentinel was on duty. Foraging usually consists of digging in the dirt
to find insects, so foragers benefit greatly from the presence of a sentinel.
Obviously there would be no need for solitary mongooses that hunt for
prey aboveground to make such calls, so already a careful observer can see
a striking difference in the behavior and vocalizations of these two species
of mongoose. Manser observed that the four most common sentinel calls
were made when no danger was present.
Many other vocalizations are exhibited less frequently by meer-
kats when a potential threat is observed, whether acting as sentinels or
not. Each time she heard one of these less frequent calls, Manser noted
whether the caller was a sentinel or a forager, the presence and type of any
predator, and the approximate distance from the caller to the predator.
Manser also categorized the level of urgency by evaluating the responses
of the signaler and receivers and by the proximity of the danger. An eagle
swooping directly overhead, for example, would prompt a sentinel to
give what Manser named the high urgency aerial call. In addition, there
were generic panic and alert calls. When the sentinel stopped guarding
and returned to foraging, it gave a contact call to nearby meerkats.
In comparison to the solitary mongoose, the acoustic signals in
meerkats appear to be uniquely derived adaptations for living in groups.
However, Manser could not simply assume the functions of these meer-
kat signals; she needed to perform controlled experiments to determine
the function of each signal and conclude whether they really are adap-
tive behaviors derived to perform specific functions for communal liv-
ing. Manser performed a series of controlled playback experiments,
using an experimental design that is similar to the ones discussed in
the previous chapter when decoding the language of fireflies and storm
petrels.
Meerkats do not recognize humans as a potential threat, and once
they got used to her, the clans ignored the biologist; so she could walk
among them, record vocalizations, and observe behavior. Manser noted
the presence of a potential threat and its distance from the clan when a
particular call was made, but this only means that there is a correlation
between the occurrence of a behavior and a type of predator.
To help determine the function of the sentinel calls, Manser recorded

the time it took for a meerkat to go on duty under three conditions:
1) when there was no sentinel on duty, 2) when a sentinel was on duty
but not calling, and 3) when a sentinel was on duty and calling. She
found that the time interval between new meerkat sentinels taking over
on guard duty was much shorter, on the order of 10 minutes, when there
was no sentinel on duty. When a sentinel was on duty and not calling, a
new sentinel would take over after 3055 minutes and when a sentinel
was on duty and vocalizing a new sentinel would assume guard duty after
5090 minutes.
At times when no sentinels were on duty, Manser played back one of
the sentinel calls or a close contact call, which is a call emitted when a
sentinel comes off duty, or background noise with no calls to individuals
when no sentinels were actually on duty. Manser measured the time it
took for a meerkat to go on sentinel duty and the percent of time forag-
ing meerkats spent on alert instead of digging in the sand for prey in each
condition. She found that when background noise was played the time
interval between guards on duty ranged from about 1216 minutes and
foraging meerkats spent a median of 20 minutes on alert, which was sig-
nificantly longer than the median amount of time on alert (10 minutes)
for foraging meerkats that heard Mansers sentinel playback call. In ad-
dition, meerkats that were exposed to the sentinel playback call assumed
guard duty only after about 2226 minutes, far longer than when exposed
only to background noise.
Sentinels spend much of their time vocalizing when on guard duty.
But there are times when they are silent. When sentinels are vocalizing,
other meerkats are much less likely to assume sentinel duty, freeing them
up to continue foraging. Sentinels leave duty for a variety of reasons,
including hunger and fatigue. They may cease vocalizing when they are
ready to leave their post, and that itself is information to foraging meer-
kats. The silence has an effect on meerkats, causing one or more of them
to assume sentinel duty. Meerkats assume sentinel duty more often when
no sentinel calls are being made. When a sentinel is vocalizing, other
meerkats are less likely to join in that duty. Therefore, the sentinel and
close contact calls enable a coordinated system of group cooperation
and security.
Although there was no statistical difference in time spent on alert for

sentinel versus close contact calls, which was another comparison made,
Manser suggested that meerkats raised their heads out of the sand to de-
termine whether another meerkat was going on sentinel duty after the
caller of the close contact call announced that it was coming off duty.
Sentinels use their calls to inform other meerkats that there is a sentinel
on duty, which allows the foraging meerkats to spend less time on alert
and more time foraging for insects.
Manser observed that the alarm calls in were highly correlated with
the specific predator and urgency of the situation. For example, some calls
were emitted when there was an aerial threat, such as an eagle or a hawk,
with urgency ranging from low to high. Other calls were associated with a
terrestrial threat, such as a jackal or snake, again with increasing levels of
urgency. Finally, another set of calls preceded the clan gathering together
in a protective mob. There were also generic alarm calls that were used
during times when a particular threat was present but the calling meerkat
could not or did not identify the threat.
To further test the functions and benefits of all these diverse calls,
Manser and her colleagues performed a series of playbacks. Calls were
recorded during actual predator encounters or during simulated predator
encounters. For instance, a person walking with a dog on a leash was used
to simulate a jackal. The scientists conducted playback experiments in the
absence of real predators where different alarm calls, categorized by level
of urgency and type of predator, were played to several different groups of
meerkats and their responses were documented (Figure 4).
There were a number of behaviors observed during the playback ex-
periment. Upon hearing some calls, meerkats scanned the surrounding
area or looked up and scanned the sky. Sometimes they scanned as they
ran to the shelter of a burrow or simply ran to the nearest burrow. Under
some conditions, they gathered to form a group, after which they either
stayed in place or ran together to a burrow.
Manser and her colleagues used playback experiments to decode a
complex, derived set of vocalizations that meerkats use to reduce the suc-
cess of predator attacks. They found that the behavioral responses of for-
aging meerkats varied with the type of call played back (see Figure 4). If
a meerkat sees an aerial predator such as an eagle, its aerial alarm call led
erect fur
erect tail
approach caller
away together
gathering
run to shelter
shelter and scan
= low/aerial
scan sky = medium/aerial
= high/aerial
= low/terr
= medium/terr
= high/terr
scan area = low/recruit
= high/recruit
= low/alert
= low/animal
= high/panic
no response
0 20 40 60 80 100
percentage of meerkats showing behavior
Figure 4 The percentages of meerkats displaying one of nine
behaviors in response to recorded vocalizations when no predator was
within sight. Percentages in a vocalization category may add up to
more than 100%, because meerkats may exhibit multiple behaviors
after one call.
Source: From Manser et al. (2001) Table 2.
other meerkats to scan and seek shelter. If a terrestrial predator was identi-
fied, sentinels gave a terrestrial call with a level of urgency related to how
far away the threat was located, and meerkats responded to those alarms
by seeking shelter, gathering together, or running away together.
A sentinel gives a high recruit call if a snake was observed or a low
recruit call if a meerkat from a rival group came near. The clans respond
by gathering together with erect fur and tails to fend off the intruder. The
erect fur and tails and numerous individuals give the appearance of large
size, perhaps making a predator think twice before attacking. As with
other types of predators, meerkats have adapted to snakes with an alarm
call that sends specific information about the type of threat, and they
respond to that information appropriately.
The feeding strategies of predators that eat meerkats contributed to
the evolution of meerkat group behavior. However, living in a group
could be costly if the group is more noticeable to predators. If a benefit
of group living is a collective protection system with an early warning of
danger, there must be effective information transfer between individuals
within the group. The Cape grey mongoose would not exhibit such infor-
mation transfer, nor would they be expected to have such a complex suite
of vocalizations; there would be little to no selection pressure in an animal
that spends most of its time alone, does not dig for insects in the sand,
and is protected by shrubby habitat. Meerkats, who live in the open and
are exposed to predators while foraging, evolved a sentinel system with a
complex set of sentinel and alarm calls that carry specific information to
receivers. Meerkats transmit information via a signal representing data
that justifies a change, and receivers alter their behavior depending on the
data in the signals that they receive.
Storm petrels from the previous chapter have a limited range of
calls and the calls they use are innate (they do not learn them but know
them when they are born). While not discussed here, it is well-known
that other birds have more complex social structures or interactions than
storm petrels, and like the meerkat compared to the mongoose, that has
led to the evolution of more complex vocalizations than storm petrels.
Some birds, like meerkats, may live in colonies year round and cooperate
in foraging, flocking, or rearing of offspring, activities that storm petrels
do not engage in.
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2001.
CHAPTER 3
Plants of the Same Species

Recognize One Another
Through Pollen-Pistil
Interactions
Like animals, plants benefit from being able to transfer information be-
tween one another. Signals passed between plants are not like the visual
and auditory communications in animals because plants lack a nervous
system and, therefore, require completely different mechanisms for com-
munication. Any investigation of information at the population level
should include examination of information transfer in non-animal species.
The signals passed between plants are chemical in nature, such as the
pleasant smells flowers emit. As with other types of communication, plant
signals are non-heritable information transmitted between individuals, but
the ability to transmit that information is heritable. Animals and microbes
also use chemical signals, but the focus will be on plants in this chapter.
Chemical signals in plants initiate a series of biochemical reactions, trig-
gering changes in the receiver plants cells and providing for continuity of
life, illustrating two of the themes of information transfer. The categories
of information passed between plants of the same species will be familiar
to anyone who has studied information in animal species: two major cat-
egories of plant communication are identification of sex and of species.
Many plants reproduce asexually, but it is during sexual reproduc-
tion that information transfer at the population level takes place. For
sexually reproducing plants, it does not benefit the plant to waste its en-
ergy and gametes (sex cellseggs and sperm) on mating with a plant of
a different species. The union of an egg and sperm results in a fertilized
egg. In any organism that reproduces sexually, the act of sexual repro-
duction requires information and communication to identify members
or gametes of the same species. The reproductive cycle of a species,
which depends on several types of information, provides for the con-
tinuity of life.
For the incredibly diverse plants known as flowering plants, flowers
are the focal point of the reproductive strategy. Gametes are produced in
specialized structures on the flower. In order to understand the process of
sexual reproduction in flowering plants, it will be useful to closely exam-
ine flower anatomy (Figure 5). A model organism for plant biology and
genetic studies is Arabidopsis thaliana, called simply Arabidopsis by the
biologists who study it. Its flower anatomy is relatively simple. The pollen
grains develop on the anther, the male part of the flower that produces
pollen, and produce the sperm. The sperm must travel in pollen grains via
pollination from the anther to the stigma, which is the part of the pistil
upon which pollen lands and germinates. The stigma is the sticky surface
at the top of the pistil, which is the female part of the flower and contains
the ovules, where the eggs are produced.
Anthers generally sit atop stalks arranged around a central pistil,
which is composed of three partsthe stigma, the stalk-like style, and
the ovary itself, which contains the ovules where eggs are produced. The
number of each of these structures varies from species to species.
female parts
stigma
pistil style pollen sacs
ovary
petals
male parts sepals

anther
stamen
filament
Figure 5 Arabidopsis thaliana flower

anatomy. Pollen is released from the
anthers and travels to the stigma.
Plants of the Same Species Recognize One Another 29
Pollen must be transferred from one flower to another, and because

agents other than the plant itself transfer the pollen, pollination may be in-
efficient. Wind, water, animal (insects, birds, and bats), and self-pollination
are mechanisms of getting pollen from the anther to the stigma. With
wind and water, pollen is not directed but rather where the pollen ends
up is somewhat random and unpredictable. Thus, those types of plants
tend to produce a lot of pollen that they spread into the environment.
The cost is the tremendous amount of pollen that has to be produced,
but the benefit is that those plants do not have to rely on other species to
disperse their pollen.
For animal dispersed pollen, the benefit may be that pollinators are
often reliable and will visit the same type of flower, and thus may deliver
pollen to another individual of the same species. Not as much pollen
needs to be produced, but it can be utilized by pollinators that are not ef-
ficient at pollinating or visit many different flower species, so pollen may
still be wasted. Animal-dispersed pollen has a better chance of landing on
another flower, but it may not be a flower of the correct species.
Pollen, especially wind-dispersed pollen, may land in many places
other than on a flower belonging to the same species. Pollen can land on
another plant, on the ground, or in someones nose, making them sneeze.
These inefficiencies partially explain why plants produce so much pollen,
which is much to the sorrow of those of us with allergies. Finally, pollen
may be transferred through self-pollination, which is the transfer of pol-
len from anther to stigma on the same flower.
Lets return to Arabidopsis and determine some of the factors involved
in its pollination. M. Hoffman and colleagues observed Arabidopsis flow-
ers in a small natural population. They estimated the number of flowers
to open each day between the 7th and 17th of May, a period of intense
flowering. They also sampled and identified the insects that pollinated
Arabidopsis flowers and estimated the percentage of flowers that were vis-
ited by insects.
Insects, especially bees, flies, and the tiny insects called thrips, may
pollinate Arabidopsis. But even on the warm, sunny days that the insects
visited Arabidopsis flowers only a small proportion of flowers were vis-
ited, less than 2.5%. The absence of pollinators suggests that Arabidopsis
self-pollinates. One advantage to self-pollination is that pollen is not
dependent upon animals or wind. A disadvantage may be that inbreed-

ing may result, lowering genetic diversity. The advantage to insect pol-
lination is that insects may transfer pollen fairly long distances.
In self-pollination there must be a mechanism for ensuring that the
pollen that lands on the stigma is from the self and not from another
individual or from another species. To ensure pollination by the correct
species, there are five stages of pollen germination and growth where in-
formation is exchanged between the pistil and the pollen: 1) pollen adhe-
sion and hydration (transfer of water and nutrients from the stigma to the
pollen grain), 2) pollen germination, 3) pollen tube growth (formed by
the pollen grain through which the sperm travel to the ovary), 4) pollen
tube elongation in the style, and 5) fertilization in the ovary. Adhesion
and hydration occur on the stigma. The pollen grain then germinates also
on top of the stigma and begins to form the pollen tube, which grows
down into the style, upon which the stigma sits. The tube elongates all
the way down the style and finds its way to the ovary, where the sperm are
deposited and the egg is fertilized.
While a pollen grain is made up of only a few cells, it is a complicated
structure, and each species is uniquely shaped and sized. Pollen grains of
closely related species may be somewhat similar in size, shape, and outer
surface features. Stigmas are also complex, again with features that are ex-
clusive to each species. As with pollen grains, stigmas and, for that matter,
the entire flower may also be similar in closely related species.
When pollen lands on a stigma of the same species, it must stick in
order for the pollen tube to eventually grow; this is adhesion. However,
pollen from other species can stick and prevent the correct pollen from
sticking. Therefore, plants that do not bind the pollen of other species
have a selective advantage. Scientists who study Arabidopsis have con-
ducted experiments to determine the communication that occurs be-
tween a pollen grain and the stigma, which is essential to successful pollen
adhesion and hydration.
In one study, Daphne Preuss and her colleagues tested pollen adhe-
sion on Arabidopsis stigmas using pollen from a variety of plants. Their
experiment included pollen from a set of closely related plants and pollen
from a set of plants more distantly related to Arabidopsis. For each sample,
the researchers covered pistils from Arabidopsis flowers with pollen grains
of each test species. Preuss colleagues then washed the pistils in one of
two chemical buffer solutions. The average number of pollen grains still
bound to stigmas was determined.
Pollen that adheres tightly to Arabidopsis stigmas ought to be from
plants closely related to Arabidopsis, because closely related species have
more similar pollen and stigmas. Arabidopsis pollen grains adhered more
tightly than most other species, with the exception of sycamore, west-
ern ragweed, and mugwort, all species known to be closely related to
Arabidopsis. Some closely related species did not have pollen grains that
adhered well, but none of the distantly related species adhered well at all.
Several species that showed relatively strong adhesion in the first test
were selected for a differential buffer test. In this test, stigmas were washed
with one buffer after they were covered with pollen grains, and then they
were washed with another buffer that removed all but the most tightly
bound pollen grains. The researchers found that pollen from Arabidopsis
adhered more tightly to Arabidopsis stigmas, even after exposure to the
stronger buffer, than pollen from other species did. This more convinc-
ingly demonstrates species-specificity of the pollen-stigma interaction;
Arabidopsis pollen and stigmas transfer information to recognize each
other, which causes quick and strong binding.
In related experiments, Preuss and her colleagues sought to determine
the source of the signalthat is, the part of the pollen grain recognized
by the stigma that led to adhesion. Pollen grains have an outer layer with
an external sculpted portion, which is a patterned, ridged wall that varies
from species to species. This wall protects DNA during dispersal, and the
specific pattern of ridges and valleys of the pollen grain might also play a
role in adhering to the appropriate stigma. In addition, the valleys often
contain sticky substances made of lipids and proteins. Keep in mind that
the pollen is the sender of a signal and the plant upon which it lands is
the receiver.
The researchers tested whether pollen coat lipids and proteins are
critical for adhesion by removing them from the outer layer of the pollen
grain. They used three methods to remove these lipids and proteins. The
researchers tested adhesion on wild-type pollen; mutant pollen (cer6-2)
that produced pollen with no lipids and only a few proteins; cyclohexane-
treated wild-type pollen, which removed lipids and proteins from the
pollen coat; and heat-treated wild-type pollen. In addition, they tested

normal pollen on cer6-2 stigmas.
The researchers had evidence from the first experiment that the initial
signals communicated between pollen and stigma, at least for Arabidopsis,
are important for species recognition. The results from this second ex-
periment revealed that initial adhesion does not depend on the lipids and
proteins in the pollen coat, because when those molecules were absent
from the cyclohexane-treated or cer6-2 mutant pollen, pollen was found
to still adhere with equal efficacy as normal-type pollen.
In nature, a few minutes after pollen adhesion, hydration begins. Be-
cause the pollen is dry and has few nutrients, pollen tube growth depends
upon hydration. The scientists further tested normal and cer6-2 pollen
to determine if binding affinity increased as hydration and germination
proceeded. A set of stigmas were pollinated with one or the other type of
pollen. At 5, 15, and 30 minutes after pollination, a subset of the stigmas
exposed to each type of pollen were washed with buffer to determine the
number of pollen grains that were tightly bound to stigmas at that time.
The result of this experiment suggests that lipid and/or protein signals
seem to be important after hydration occurs. After hydration, a pollen
tube begins to grow. After hydration and pollen tube germination, the
buffer still washes off the same number of pollen grains in the mutant,
but the number of pollen grains adhering in the wild-type pollen con-
tinues to increase as the time after washing increases. This indicates that
more pollen grains are adhering in a secondary process initiated during or
after hydration. Lipids and/or proteins, missing from the mutant, appear
to be important in that secondary process. Mutant Arabidopsis that lacked
lipids or protein in their pollen coats, but not both, would be useful in
further investigating the signals important in adhesion, hydration, and
pollen tube growth.
The evidence from the experiments suggested that pollen from the
wrong species can adhere to a stigma. Another scientist, A. Ray and his
colleagues investigated whether some pollen from an unrelated species
could hydrate and begin to grow pollen tubes. Pollen tubes grow down
the pistil from the stigma, and ultimately one pollen tube grows to each
ovule. If the sperm deposited by the pollen tube is of the same species,
fertilization takes place. The researchers wondered whether pollen tubes
are guided by signals from the ovule or egg and whether the signals are
species-specific. In their study, they discovered mutant Arabidopsis plants
that had normal tissue in the pistil, but in which half of the ovules failed
to produce eggs when the ovary underwent meiosis. Although some
plants have flowers with a single ovule in the ovary, many plants, like
Arabidopsis, have flowers with multiple ovules per ovary.
When flowers from the mutant plants had reached the appropri-
ate stage of development, the scientists dusted the stigmas with wild-
type pollen and then incubated the flowers. The ovaries were stained for
the chemical callose, which is a polysaccharide (a type of carbohydrate),
produced by pollen tubes, although other plant tissues produce it as well.
The staining procedure shows pollen tubes extending toward the ovaries,
as well as a bright spot right where the pollen tube enters a particular
ovule. If the scientists observed this in their samples, they recorded that
the pollen tube had arrived at the ovule.
Callose is also associated with certain cells within viable ovules. Using
just one staining technique, the scientists could track pollen tube growth
and determine which ovules in the mutant plant had produced eggs and
which had not. For each flower, the scientists determined the number
of each type of ovule as well as the number of pollen tubes that reached
each type of ovule. The pollen tubes that reached normal ovules were
considered a successful transfer of information between two individu-
als of the same species. They found that 124 out of 184 normal ovules
had apollen tube associated with it after 20 hours of growth. None of
the 189abnormal ovules had pollen tubes associated with them after 20
hours of incubation.
The researchers also found that pollen of two closely-related species
could grow in Arabidopsis pistils. In one of these species, only 3% of the
pollen tubes arrived near ovules, and none of the 13 pollen tubes that
reached an egg in an ovule successfully entered. In the other species, only
1.1% of pollen tubes made it to the vicinity of ovules, and as with the
other species, none were able to enter.
Plants that reproduce sexually must have a mechanism to ensure that
only species-specific sperm reach the eggs. The experiments of Preuss and
her colleagues show that for Arabidopsis tight binding of pollen is species-
specific, and the signal is in the outer layer of the pollen but is not the
lipid or protein components. Lipids and proteins on Arabidopsis pollen

play a role after initial adhesion; mutants without the lipids and pro-
teins do not show secondary binding or hydration. Ray and his colleagues
then demonstrated that although pollen tubes can grow in related species
styles, pollen tube guidance requires a signal from an ovule, and the signal
is specific to a species. Plant pollination is analogous to male and female
animals exchanging signals that help them recognize each other as a mem-
ber of the opposite sex and the same species. The data analyzed reveals the
specificity of information transfer in plants.
Communication and information transfer occurs at several different
stages in the pollination process. Plants mostly use chemical means of a
variety of sorts for species-recognition. Plants cannot use other mecha-
nisms that animals often use, such as vocal or visual cues. However, many
of the functions are similar, and include sex and species-recognition and
defense. Many plants and animals use chemical signals to communicate
information. However, animals, and not plants, have olfactory sensory
mechanisms and nervous systems to facilitate that information transfer.
Plants must use other means to detect and respond to chemical signals.
Animals have additional means of communication, including vocal and
visual, that plants do not use.
The principles of information transfer are both widespread and have
common functions. How information is transferred varies among differ-
ent organisms and can come in the form of acoustical, visual, or chemical
signals. The ability to transmit these signals is heritable, but the informa-
tion transmitted is non-heritable. Species recognition and bringing to-
gether two individuals for the purpose of reproduction are widespread
requirements for the continuity of life. In fact, information exchange oc-
curs between individuals of different species and between individuals and
their environment, and some species take advantage of other species and
their species-recognition signals.
Bibliography
Aarts MG, Keijzer CJ, Stiekema WJ, et al: Molecular characterization of
the CER1 gene of Arabidopsis involved in epicuticular wax biosynthe-
sis and pollen fertility, Plant Cell 7(12):21152127, 1995.
Hoffmann MH, Bremer M, Schneider K, et al: Flower visitors in a natural

population of Arabidopsis thaliana, Plant Biology 5:491494, 2003.
Ray SM, Park SS, Ray A: Pollen tube guidance by the female gameto-
phyte, Development 124(12):24892498, 1997.
Shimizu KK, Okada K: Attractive and repulsive interactions between
female and male gametophytes in Arabidopsis pollen tube guidance,
Development 127(20):45114518, 2000.
Wheeler MJ, Franklin-Tong VE, Franklin FCH: Tansley Review No.128.
The molecular and genetic basis of pollen-pistil interactions, New
Phytol 151(3):565584, 2001.
Zinkl GM, Zwiebel BI, Grier DG, et al: Pollen-stigma adhesion in Arabi-
dopsis: a species-specific interaction mediated by lipophilic molecules
in the pollen exine, Development 126(23):54315440, 1999.
Conclusion
Animals and plants use communication to transfer information so that
they can recognize other members of the same species. Recognition al-
lows individuals to cooperate for defense and finding food, and it also fa-
cilitates mating and sexual reproduction. Although the mechanisms that
organisms use to communicate differ widely, the principles of communi-
cation are similar across all of these groups. Regardless of the mechanism
or the reason for the communication, information transfer is not always
100% perfect, which leads to variation in the response of signal receivers,
either in how they respond or at what point they respond.
In addition to the costs that might be associated with either incom-
plete or error-free information transfer, there are other costs associated
with sending and receiving signals, but the benefits must outweigh the
costs. When benefits of a behavior outweigh costs, there is selective ad-
vantage to individuals that exhibit the behavior: they may have greater
survival and reproductive success than individuals that do not exhibit
the behavior. Over evolutionary time, however, species have evolved to
manipulate or intercept signals between individuals of another species
in order to more successfully prey on them or to better compete for a
resource. The signals that pass between individuals of two different spe-
cies are explored elsewhere in this collection, in the book Information
in the Environment, along with other types of information in ecological
systems.
Glossary
adrenaline. A hormone secreted by the adrenal glands under stress that increases
heart rate, respiration rate, and carbohydrate metabolism.
anther. The part of the flower that produces pollen.
bioluminescence. Light production as a result of a chemical reaction.
chi-squared test. The chi-squared test is a statistical method to decide whether or
not two distributions are significantly different.
communication. Transmission of information between two individuals so that it
is satisfactorily received or understood.
comparative approach. A methodology where multiple species are studied to
compare traits and their functions, in order to understand the evolution of those
traits.
competition. The demand by two or more organisms for limited environmental
resources.
derived traits. An adaptation unique to a particular species or group of species,
which is evolved in one species but is not found in other closely related species.
dopamine. A neurotransmitter formed during the synthesis of norepinephrine
essential to the normal functioning of the central nervous system.
ectotherms. Animals that regulate its body temperature using behavioral mecha-
nisms and heat from the environment.
gametes. The egg and sperm are haploid sex cells and are required for sexual
reproduction.
inbreeding. Inbreeding is reproduction between close relatives.
information. That which is conveyed or represented. Information is stored, com-
municated, and implemented or interpreted.
median. The median of a set of observations is a value such that less than half of
the observations are smaller than the median, and less than half of the observa-
tions are larger than the median.
oxytocin. A hormone released by the pituitary gland that also acts as a neu-
rotransmitter and plays a role in childbirth and is associated with feelings of love
or attachment.
pistil. The female part of a flower, where eggs are produced.
playback experiments. Experiments where signals such as light flashes or vocal-
izations are recorded and played back to animals in controlled conditions to study
their responses.
pollination. A process that transfers pollen.
population. A population is a group of individuals of the same species living in
the same place at the same time.
40 GLOSSARY
receiver. An individual who detects a message, intentionally or accidentally.

self-pollination. The transfer of pollen from anther to stigma on the same flower.
serotonin. Serotonin is a neurotransmitter that helps to regulate emotional
moods and gastric functions.
sexual reproduction. Reproduction of offspring involving fusion of haploid
gametes.
social animals. Animals that live in groups, usually of the same species, interact
extensively with others in the group, and have a recognizable society.
stigma. The part of the pistil upon which pollen lands and germinates.
vocalizations. The communication sounds made by animals.
Index
Adrenaline, 13, 14 in non-animal species, 27
Anther, 28, 29 during sexual reproduction,
Arabidopsis, 2834 2728
Interpulse intervals
Bioluminescence, 2 (IPIs), 35
Calls, types of, 89 Lorenz, Konrad, 1

Cape grey mongoose, 18
Chattering calls, 912 Meerkats
Chi-squared test, 910 behavior, 1920
Close contact calls, 21, 22 duty conditions, 21
Communication vocalizations, 23
in bird, 713 Mutant Arabidopsis, 32, 33
in firefly, 27 Mutant pollen, 31, 32
functions of, 1
and information transfer, 34 Oxytocin, 15
instinctual aspect of, 12
plant, 27 Periods of silence, 9
Comparative approach, 1718 Photinus, 2
Cooperative behavior, 2 Photinus greeni, 5
Pistil, 28, 30, 31, 32, 33
Dopamine, 13, 14 Plant
communication categories, 27
Ectotherms, 3 pollination, 34
Playback experiments, 10, 12
Fireflies Pollen
simple communication in, 27 adhesion and hydration, 30
transfer information using visual germination, stages
communication of, 30
Flowering plants, 28 grains, 28, 3032
Pollination, 2830, 32, 34
Galerella pulverulenta. See Cape grey
mongoose Receiver, 2, 6, 9
Gametes, 27, 28
Grating calls, 912 Self-pollination, 29, 30
Sentinel calls
Imperfect information transfer, 4 versus close contact
Inbreeding, 30 calls, 22
Information transfer definition of, 19
communication and, 34 function of, 21
mechanisms of, 1721 as potential threat, 20
42 INDEX
Serotonin, 13, 14 Vocalizations

Signal pattern, variations in, 3 of birds, 78
Sonograms, 8 as means of gender recognition, 12
Stigma, 2833 by meerkats, 20
Storm petrels, 813
Wilsons storm petrel (Oceanites
THrips, 29 oceanicus), 8
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First published in 2016 by

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

Printed in the United States of America.

Figure 1 A population of birds and information transfer between two

If all of the information meant to be transferred is actually transferred

In 1950, the famous animal behaviorist Konrad Lorenz wrote, Animals

is a member of the same species and that precedes mating. A receiver is

Simple Communication in a Firefly

The function of firefly communication seems to be twofold. First, the

temperature. Further, a proportion of the female population does not

More Complex Communication in a Bird

syllables, and duration of silent periods. Periods of silence are analogous

as in kite] raised to the power of 2) to compare each pair of situations.

mean # of petrels responding ( 1 s.e.)

Inthis case, Bretagnolle recorded any vocalizations and described in

Ethical, Legal, Social Implications: Love at First

Dopamine increases during infatuation while another neurotransmit-

Group Living Requires More

Fireflies and petrels communicate to identify and attract potential mates,

the vocalizations, ecology and behavior of all mongoose species. Although

one another, to determine breeding status, or to chase an intruder from a

Based on her observations, Manser concluded that the single, double,

To help determine the function of the sentinel calls, Manser recorded

Although there was no statistical difference in time spent on alert for

shelter and scan

Plants of the Same Species

male parts sepals

Figure 5 Arabidopsis thaliana flower

Pollen must be transferred from one flower to another, and because

dependent upon animals or wind. A disadvantage may be that inbreed-

pollen coat; and heat-treated wild-type pollen. In addition, they tested

lipid or protein components. Lipids and proteins on Arabidopsis pollen

Hoffmann MH, Bremer M, Schneider K, et al: Flower visitors in a natural

receiver. An individual who detects a message, intentionally or accidentally.

Calls, types of, 89 Lorenz, Konrad, 1

Serotonin, 13, 14 Vocalizations

Cells in Tissuesby Christopher J. Paradise and A. Malcolm Campbell

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