Animal Behavior

Barron’s Chapter 18

PART I – Animal behavior

Animal Behavior

Read: “Intro to animal behavior”

Key points
• - Animal behavior includes all the ways animals interact with other organisms and the physical
environment.
•
• - Behavior can also be defined as a change in the activity of an organism in response to a stimulus, an
external or internal cue or combo of cues.
•
• - To fully understand a behavior, we want to know what causes it, how it develops in an individual, how it
benefits an organism, and how it evolved.
•
• - Some behaviors are innate, or genetically hardwired, while others are learned, or developed through
experience. In many cases, behaviors have both an innate component and a learned component.
•
• - Behavior is shaped by natural selection. Many behaviors directly increase an organism's fitness, that is,
they help it survive and reproduce.
•
Introduction
Do the squirrels in your neighborhood bury acorns underground? Does your cat start meowing around the
time you usually feed her? Do you start hanging around the kitchen when it’s close to dinnertime?

If you've noticed any of these things, congratulations—you've made your first observations in behavioral
biology! These are all examples of animal behaviors. Yep, you and I count as animals too. In fact, these
behaviors are just a tiny sampling of the amazing and diverse behaviors we can see in nature.

We could ask what behavior is used for, but it might be better to ask, what isn't it used for? Animals have
behaviors for almost every imaginable aspect of life, from finding food to wooing mates, from fighting off
rivals to raising offspring. Some of these behaviors are innate, or hardwired, in an organism's genes. For
instance, this is true of the squirrel and its acorn. Other behaviors are learned, such as your tendency to
hang around the kitchen at dinnertime or your ability to read the words on this screen.

In this article, we’ll take a closer look at animal behavior—how it’s studied, how it evolves, and how it
can run the gamut from hardwired to learned.

What is behavior?
Broadly speaking, animal behavior includes all the ways animals interact with other members of their
species, with organisms of other species, and with their environment.

Behavior can also be defined more narrowly as a change in the activity of an organism in response to
a stimulus, an external or internal cue or combination of cues.

For example, your dog might start drooling—a change in activity—in response to the sight of food—a
stimulus.

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 Behavioral biology is the study of the biological and evolutionary bases for behavior. Modern behavioral
biology draws on work from the related but distinct disciplines of ethology and comparative psychology.
•
• - Ethology is a field of basic biology, like ecology or genetics. It focuses on the behaviors of diverse
organisms in their natural environment.
•
• - Comparative psychology is an extension of work done in human psychology. It focuses largely on a few
species studied in a lab setting.
•
Behavioral biology also draws on many related areas of biology, including genetics, anatomy, physiology,
evolutionary biology, and, of course, neurobiology—which traces the neural circuits that underlie animal
behavior.

Four questions to understand a behavior

Nikolaas (Niko) Tinbergen was a Dutch ornithologist, or bird biologist, who studied behavior and is now
considered one of the founders of the field of ethology. Based on his own research, Tinbergen proposed
four basic questions helpful in understanding any animal behavior.

Let's look at these questions, using the production of song by the zebra finch—a common songbird—as an
example.

1.
2. 1. Causation—What causes the behavior? What triggers the behavior, and what body parts, functions,
and molecules are involved in carrying it out?
3.
Example: Singing is triggered in zebra finches by social cues, such as the proximity of a potential mate, as
well as the appropriate hormonal state. The ability to produce songs is influenced by male hormones and
occurs mainly in male birds. Songs are produced when air flows from air sacs in the bronchii through an
organ called the syrinx. Certain parts of the brain control song production and are well-developed in male
zebra finches.
4.
5. 2. Development—How does the behavior develop? Is the behavior present early in life? Does it change
over the course of the organism's lifetime? What experiences are necessary for its development?

Example: Young male zebra finches first listen to the songs of nearby males of their species, particularly
their fathers. Then, they start to practice singing. By adulthood, male zebra finches have learned to
produce their own songs, which are unique but often have similarities to those of their fathers. Once a
finch has perfected its song, the song remains fixed for life.

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6. 3. Function/adaptive value—How does the behavior affect fitness? How does the behavior affect an
organism's chances of survival and reproduction?
7.
Example: Singing helps male zebra finches attract mates, increasing the chances that they will reproduce.
Singing is part of an elaborate courtship ritual that entices the female to choose the male.
8.
9. 4. Phylogeny—How did the behavior evolve? How does the behavior compare to those of related species?
Why might it have evolved as it did?
10.
Example: Almost all species of birds can make vocal sounds, but only those in the suborder Passeri are
songbirds. Relative to the zebra finch, other songbird species differ in the timing of their listening and
practicing phases, the plasticity of song over their lifetimes, the extent to which the song is similar among
individuals of the species, and the way that singing is used—for example, for defense of territory vs.
courtship of mates.

Cues that trigger behavior

At its core, an animal behavior is a response to an internal or external cue. Through behavior, animals can
act on the information they receive in ways that will, hopefully, favor their survival and reproductive
success.

What kinds of cues can trigger behavior? In some cases, the cue is largely external:
•
• - In hibernation, an animal goes into a den or burrow, reduces its metabolic rate, and enters a state of
inactivity during the winter, conserving resources while conditions are harsh and food is scarce.
Environmental cues often trigger hibernation behavior. For instance, brown bears enter their den and
hibernate when temperature drops to 0°C and snowfall begins.
•
• - Estivation is similar to hibernation, but it occurs during the summer months. Some desert animals
estivate in response to dry conditions. This shift helps them survive the harshest months of the year. The
snails in the photo below climb to the tops of fence posts to estivate.
•
• - Migration is a behavior in which animals move from one location to another in a seasonal pattern. For
instance, monarch butterflies living in the northern and central United States migrate to Mexico in the
autumn, where they spend the winter. Environmental cues that trigger the autumn migration include air
temperature, day length, and food availability.
•
In other cases, the cue for a behavior may be internal. For instance, some behaviors occur with
a circadian rhythm, meaning that they are triggered by the animal's internal body clock. You, for
example, tend to wake up and become active at roughly the same time each day. As you may have

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 discovered if you've ever taken a long flight, your body's alarm clock will still "go off" at the same time
even if the external cues change, which is what causes jet lag!

It's also common for behaviors to be triggered by a combination of internal and external cues interacting.
For instance, mating behaviors may be triggered in an animal only when it's in the right hormonal state,
an internal cue, and when it sees a member of the opposite sex, an external cue.

Innate vs. learned behaviors

When we are trying to understand how a behavior develops and how it arose evolutionarily, one important
question is whether the behavior is genetically preprogrammed or acquired through experience. Let's
consider some vocab:

• - Innate behaviors are genetically hardwired and are inherited by an organism from its parents.
•
• - Learned behaviors are not inherited. They develop during an organism's lifetime as the result of
experience and environmental influence.

Behavioral biologists have found that many behaviors have both an innate and a learned component. So,
it's generally most accurate for us to ask to what extent a behavior is innate or learned.

Mostly innate behaviors

There are some examples of behaviors that are really and truly hardwired. These behaviors take place in a
highly predictable way in response to the right stimulus, even if the organism has never before
encountered that stimulus.

For example, an adult salamander will swim perfectly if it's placed in water, even if it never saw water
when it was young and has never watched another salamander swim. In this case, the behavior of
swimming can only be explained as something genetically preprogrammed in the salamander.

Similarly, you—or any human—will rapidly jerk your hand away if you touch a very hot object. This
response is a reflex that's hardwired in the circuits of your sensory and motor neurons and doesn't even
involve your brain.

Partly innate, partly learned behaviors

In other cases, an organism is genetically programmed to develop a behavior, but the form the behavior
takes depends on the individual's experience.

One example is the learning of a song by a zebra finch or other songbird, as we saw above. All male zebra
finches will begin listening to and learning song at about the same age and practicing and producing song
at a slightly later age. Although this pattern is genetically determined, the exact features of the song a
bird sings will depend on the songs it hears during its learning period.

Another, more familiar example is language acquisition in humans. Babies are preprogrammed for
language learning, but which language they learn depends on what they're exposed to during their plastic,
or formative, period.

Mostly learned behaviors

In other cases, behaviors are largely dependent on experience—they're learned—and can't be fully
explained by genetic preprogramming.

For instance, if a rat receives a food reward each time it pushes a lever, it will quickly learn to push the
lever in order to get the food. Similarly, if a cow gets an electric shock each time it brushes up against an
electric fence, like the one below, it will rapidly learn to avoid the fence. Pushing a lever to get a reward
and avoiding electric fences are not hardwired in rats and cows but are, instead, learned behaviors the
animals develop through experience.
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 If a behavior is learned rather than innate, it isn't directly inherited. But it does still depend on genes. For
instance, not all types of animals could learn to push a lever to get a reward. The rat's capacity to learn
this behavior depends on how its brain is wired, and the construction, maintenance, and function of a rat
brain are all determined by genes in the rat genome.

Check your understanding

Beach mice dig burrows in the sand when they are living in their natural environment. A beach mouse
burrow has a specific shape with a long escape tunnel that is different from the burrows of closely related
mouse species.

A young beach mouse is raised in captivity, without any access to dirt or sand or any chance to observe
burrowing adults. When given access to dirt, it immediately digs a burrow with the normal shape for its
species.
What type of behavior is burrow digging in the beach mouse?
Choose 1 answer:
• (Choice A) Innate
• (Choice B) Learned
• (Choice C) Mostly learned, with a small innate component

Natural selection shapes behavior.

To the extent that a behavior is genetically determined or relies on genes, it’s subject to evolutionary
forces, such as natural selection. In many cases, we can see how a behavior gives a survival or
reproduction benefit to an animal that performs it—in other words, the behavior increases fitness.

Here are some examples of behaviors that clearly increase fitness:

•
• - Baby birds of many species instinctively open their mouths for food when the mother returns to the nest.
Birds with this heritable behavior will tend to get fed more—and thus survive to adulthood more—than
those that don’t.
•
• - Mother greylag geese instinctively roll eggs back into the nest if they fall out. Geese with this heritable
behavior will tend to have more offspring that survive to hatch than geese without the behavior.
•
• - Zebra finch males learn songs while they are juveniles, young birds, and they use these songs in
courtship rituals. Birds with the heritable tendency to learn a song will obtain a mate more often than
those that don't.
•
An important point from the last example is that natural selection can act even when the behavior itself is
not inherited. A zebra finch doesn't inherit its song directly—it has to learn the song. But its capacity and
tendency to learn a song are genetically determined, so they can be subject to natural selection.

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 What about selfless or altruistic behaviors?

Some behaviors are—at least on the surface—hard to explain in terms of fitness and natural selection.

For instance, an organism that gives a warning call to other group members upon spotting a predator is
probably more likely to be eaten by the predator, not less. So, a genetic tendency to give warning calls
seems like it should reduce fitness. Yet, altruistic behavior—behavior that reduces an individual's fitness
while increasing that of other organisms—is seen in many species.

In some cases, altruistic behavior may persist because it increases the survival of the group in such a way
as to increase the genetic transmission of the behavior. For instance, if the individual who gives a warning
call is closely related to other members of the group, the warning call may cause its own genetic
determinants to be passed on—via the relatives, who share many genes with the individual—even if the
individual giving the warning call is killed. Supporting this idea, individuals of some species are more
likely to make warning calls when relatives are nearby.

This concept of inclusive fitness remains controversial in the evolutionary biology community. In fact,
some researchers think that the warning calls may instead increase individual fitness by scaring off the
predator, for example, or attracting other competing predators, or signaling the alertness of the prey.
There's evidence for this in some cases, but when it comes to the general verdict on altruism and inclusive
fitness, the jury is still out.

In other cases, selfless behavior may not actually be selfless; it may be reciprocated by the organism that
receives it. For example, monkeys may be more likely to groom other monkeys that have recently
groomed them. Social species often have tit-for-tat systems such as this in which one organism will help
out another that has recently done it a favor.

Read: “Innate behaviors”

Key points
• - Innate behavior is behavior that's genetically hardwired in an organism and can be performed in
response to a cue without prior experience.
•
• - Reflex actions, such as the knee-jerk reflex tested by doctors and the sucking reflex of human infants,
are very simple innate behaviors.
•
• - Some organisms perform innate kinesis, undirected change in movement, and taxis, directed change in
movement, behaviors in response to stimuli.
•
• - Fixed action patterns consist of a series of actions triggered by a key stimulus. The pattern will go to
completion even if the stimulus is removed.
•
• - Scientists can test if a behavior is innate by providing a stimulus to naive—untrained—animals and to see
if the behavior is automatically triggered.
•
Introduction
If you watch a herring gull caring for its chicks, you may notice a funny ritual around feeding time. The
parent gull has a red spot on its beak. When it taps its beak on the ground, the chick will peck at the spot
several times.

This pecking triggers a response in the parent: it throws up food for the chick. That may sound gross to us,
but to a herring gull chick, it's like a pizza dinner!

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The tapping behavior is innate, or genetically preprogrammed. Herring gull chicks will peck at the red
dots of their parents' beaks without any prior training. In fact, a baby herring gull can be tricked by a
yellow stick adorned with a red dot—it will peck at the stick just as eagerly as it would at a parent's beak.

This is just one example of an innate behavior, or behavior that's genetically hardwired in an organism.
Given the right cues, an organism will perform an innate behavior without the need for prior experience
or learning. Innate behaviors tend to be very predictable—like the herring gull tapping—and they are often
performed in a very similar way by all members of a species.

In this article, we'll see examples of behaviors that are largely or fully innate. Keep in mind, though, that
many behaviors in the real world are partly innate and partly learned. Zebra finches, for example, are
preprogrammed to learn a song, but which song they learn depends on their early experience.

Reflexes
Perhaps the simplest example of an innate behavior is a reflex action: an involuntary and rapid response
to a stimulus, or cue.

One example of a human reflex action is the knee-jerk reflex. To test this reflex, a doctor taps the
tendon below your kneecap with a rubber hammer. The tap activates nearby neurons, causing your lower
leg to kick involuntarily. This automatic response depends on circuits of neurons that run between the
knee and the spinal cord—it doesn't even involve your brain!

Simplified diagram of neural circuits involved in the knee-jerk reflex.

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Some reflexes are present in human babies but are lost or placed under conscious control as the baby
grows older. For instance, a newborn baby will suck at anything that touches the roof of its mouth. This
reflex helps the baby get food by ensuring it will suck at its mother's breast or a bottle placed in its
mouth.

Kinesis and taxis

Some organisms have innate behaviors in which they change their movement in response to a stimulus,
such as high temperature or a tasty food source.

In kinesis, an organism changes its movement in a non-directional way—e.g., speeding up or slowing

down—in response to a cue. For example, woodlice move faster in response to temperatures that are
higher or lower than their preferred range. The movement is random, but the higher speed increases the
chances that the woodlouse will make its way out of the bad environment.

Taxis is a form of movement behavior that involves movement towards or away from a stimulus. This
movement can be in response to light, known as phototaxis; chemical signals, known as chemotaxis; or
gravity, known as geotaxis—among other stimuli. It can also be directed towards, positive, or away from,
negative, the source of the stimulus.

For example, woodlice show negative phototaxis, meaning that they'll move away from a light source.
This behavior may be helpful because woodlice require a moist environment, and a sunny, light, spot is
more likely to be warm and dry.

Fixed action patterns

A fixed action pattern is a predictable series of actions triggered by a cue, sometimes called the key
stimulus. Though a fixed action pattern is more complex than a reflex, it's still automatic and involuntary.
Once triggered, it will go on to completion, even if the key stimulus is removed in the meantime.

We already saw one example of a fixed action pattern in the introduction of the article: spot-pecking
behavior in herring gulls. Let's look at a couple other examples that show how fixed action patterns work.

Case study: egg retrieval

A well-studied example of a fixed action pattern occurs in ground-nesting water birds, like greylag geese.
If a female greylag goose's egg rolls out of her nest, she will instinctively use her bill to push the egg back
into the nest in a series of very stereotyped, predictable, movements. The sight of an egg outside the nest
is the stimulus that triggers the retrieval behavior.

It's not too hard to imagine why this hardwired trait would be favored by natural selection. Goose mothers
that retrieve their lost eggs are likely to have more surviving offspring, on average, than those that don't.

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 However, this fixed action pattern can also occur under circumstances where it is not useful, in other
words, where it does not benefit the goose:

• - If the egg that rolls out of the nest is picked up and taken away, the goose will keep moving her head as
though pushing an imaginary egg.
•
• - The goose will try to push any egg-shaped object, such as a golf ball, if it is placed near the nest. In
fact, she'll even carry out the retrieval pattern in response to a much larger object, such as a volleyball!

This example illustrates the fixed aspect of a fixed action pattern. In the great majority of cases a goose
is likely to encounter in nature, the behavior of rolling any egg-like object near the nest back into the
nest will be beneficial. However, it's simply a biological program that runs in response to a stimulus and
can have unhelpful results under unusual circumstances.

Case study: male sticklebacks

Another classic example of a fixed action pattern comes from the three-spined stickleback, a small
freshwater fish. During the breeding season, male sticklebacks develop a red belly and display innate
aggressive behavior towards other males.

When a male stickleback spots another nearby male, he will launch into a fixed action pattern involving
aggressive displays designed to scare off the stranger. The specific stimulus that triggers this fixed action
pattern is the red belly coloration pattern characteristic of males during breeding season.

How do we know that this is the trigger? In the lab, researchers exposed male fish to objects that were
painted red on their lower halves but didn't otherwise look like a fish, see below. The male sticklebacks
responded aggressively to the objects just as if they were male sticklebacks. In contrast, no response was
triggered by lifelike male stickleback models that were painted white.

By some accounts, this fixed action pattern has even been triggered by a fire truck driving past a male
stickleback's tank!

How do we know if a behavior is innate?

By definition, an innate behavior is genetically built in to an organism rather than learned. But how do
biologists figure out if a behavior is innate?

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 In general, scientists test whether a behavior is innate by seeing whether it is performed correctly by
naive animals, animals that have not had a chance to learn the behavior by experience. For instance, this
might involve raising young animals separate from adults or without stimuli that trigger the behavior.

As an example, let's consider digging behavior in the deer mouse and the beach mouse. These species are
closely related and can interbreed, but they live in different natural environments and have different
burrow-digging behaviors:
•
• - The deer mouse digs a small, short burrow.
•
• - The beach mouse digs a long burrow with an escape tunnel or "back door" to get away from predators.
•

Is this difference in burrow digging innate? To ask this question, researchers raised mice of both species in
the lab with no exposure to sand or opportunity to burrow. Then, they provided them with sand, a cue for
burrow construction.

Given sand, each naive mouse dug exactly the type of burrow made by its species in the wild. That is,
beach mice dug a long burrow with an escape tunnel, while deer mice dug a short burrow without an
escape tunnel. The ability of the mice to construct their normal tunnels, without ever having seen such a
tunnel before, showed that the burrowing behavior was indeed innate.

Check your understanding

Which of the following best describes a fixed action pattern?
Choose 1 answer:
• (Choice A) A reflex behavior that occurs when an organism is very young
• (Choice B) An innate behavior that runs to completion once it's started
• (Choice C) A type of learned behavior found primarily in humans
• (Choice D) A type of innate behavior found only in parent birds and hatchlings

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 Read: “Learned behaviors”

Key points
• - Habituation is a simple learned behavior in which an animal gradually stops responding to a repeated
stimulus.
•
• - Imprinting is a specialized form of learning that occurs during a brief period in young animals—e.g.,
ducks imprinting on their mother.
•
• - In classical conditioning, a new stimulus is associated with a pre-existing response through repeated
pairing of new and previously known stimuli.
•
• - In operant conditioning, an animal learns to perform a behavior more or less frequently through a
reward or punishment that follows the behavior.
•
• - Some animals, especially primates, are capable of more complex forms of learning, such as problem-
solving and the construction of mental maps.

Introduction
If you own a dog—or have a friend who owns a dog—you probably know that dogs can be trained to do
things like sit, beg, roll over, and play dead. These are examples of learned behaviors, and dogs can be
capable of significant learning. By some estimates, a very clever dog has cognitive abilities on par with a
two-and-a-half-year-old human!

In general, a learned behavior is one that an organism develops as a result of experience. Learned
behaviors contrast with innate behaviors, which are genetically hardwired and can be performed without
any prior experience or training. Of course, some behaviors have both learned and innate elements. For
instance, zebra finches are genetically preprogrammed to learn a song, but the song they sing depends on
what they hear from their fathers.

In this article, we'll take a look at some examples of learned behaviors in animals. We'll start with simple
ones like habituation and imprinting, then work our way up to complex cases like operant conditioning
and cognitive learning.

Simple learned behaviors

Learned behaviors, even though they may have innate components or underpinnings, allow an individual
organism to adapt to changes in the environment. Learned behaviors are modified by previous
experiences; examples of simple learned behaviors include habituation and imprinting.

Habituation
Habituation is a simple form of learning in which an animal stops responding to a stimulus, or cue, after a
period of repeated exposure. This is a form of non-associative learning, meaning that the stimulus is not
linked with any punishment or reward.

For example, prairie dogs typically sound an alarm call when threatened by a predator. At first, they will
give this alarm call in response to hearing human steps, which indicate the presence of a large and
potentially hungry animal.

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However, the prairie dogs gradually become habituated to the sound of human footsteps, as they
repeatedly experience the sound without anything bad happening. Eventually, they stop giving the alarm
call in response to footsteps. In this example, habituation is specific to the sound of human footsteps, as
the animals still respond to the sounds of potential predators.

Imprinting
Imprinting is a simple and highly specific type of learning that occurs at a particular age or life stage
during the development of certain animals, such as ducks and geese. When ducklings hatch, they imprint
on the first adult animal they see, typically their mother. Once a duckling has imprinted on its mother,
the sight of the mother acts as a cue to trigger a suite of survival-promoting behaviors, such as following
the mother around and imitating her.

How do we know this is not an innate behavior, in which the duckling is hardwired to follow around a
female duck? That is, how do we know imprinting is a learning process conditioned by experience? If
newborn ducks or geese see a human before they see their mother, they will imprint on the human and
follow it around just as they would follow their real mother.

An interesting case of imprinting being used for good comes from efforts to rehabilitate the endangered
whooping crane by raising chicks in captivity. Biologists dress up in full whooping crane costume while
caring for the young birds, ensuring that they don't imprint on humans but rather on the bird dummies
that are part of the costume. Eventually, they teach the birds to migrate using an ultralight aircraft,
preparing them for release into the wild.

Conditioned behaviors
Conditioned behaviors are the result of associative learning, which takes two forms: classical
conditioning and operant conditioning.

Classical conditioning
In classical conditioning, a response already associated with one stimulus is associated with a second
stimulus to which it had no previous connection. The most famous example of classical conditioning comes
from Ivan Pavlov’s experiments in which dogs were conditioned to drool—a response previously associated
with food—upon hearing the sound of a bell.
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As Pavlov observed, and as you may have noticed too, dogs salivate, or drool, in response to the sight or
smell of food. This is something dogs do innately, without any need for learning. In the language of
classical conditioning, this existing stimulus-response pair can be broken into an unconditioned stimulus,
the sight or smell of food, and an unconditioned response, drooling.

In Pavlov's experiments, every time a dog was given food, another stimulus was provided alongside the
unconditioned stimulus. Specifically, a bell was rung at the same time the dog received food. This ringing
of the bell, paired with food, is an example of a conditioning stimulus—a new stimulus delivered in
parallel with the unconditioned stimulus.

Over time, the dogs learned to associate the ringing of the bell with food and to respond by drooling.
Eventually, they would respond with drool when the bell was rung, even when the unconditioned
stimulus, the food, was absent. This new, artificially formed stimulus-response pair consists of
a conditioned stimulus, the bell ringing, and a conditioned response, drooling.

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Is the unconditioned response, drooling in response to food, exactly identical to the conditioned response,
drooling in response to the bell? Not necessarily. Pavlov discovered that the saliva in the conditioned dogs
was actually different in composition than the saliva of unconditioned dogs.

Operant conditioning
Operant conditioning is a bit different than classical conditioning in that it does not rely on an existing
stimulus-response pair. Instead, whenever an organism performs a behavior—or an intermediate step on
the way to the complete behavior—it is given a reward or a punishment. At first, the organism may
perform the behavior—e.g., pressing a lever—purely by chance. Through reinforcement, the organism is
induced to perform the behavior more or less frequently.

One prominent early investigator of operant conditioning was the psychologist B. F. Skinner, the inventor
of the Skinner box, see image below. Skinner put rats in boxes containing a lever that would dispense food
when pushed by the rat. The rat would initially push the lever a few times by accident, and would then
begin to associate pushing the lever with getting the food. Over time, the rat would push the lever more
and more frequently in order to obtain the food.

Not all of Skinner's experiments involved pleasant treats. The bottom of the box consisted of a metal grid
that could deliver an electric shock to rats as a punishment. When the rat got an electric shock each time
it performed a certain behavior, it quickly learned to stop performing the behavior. As these examples
show, both positive and negative reinforcement can be used to shape an organism's behavior in operant
conditioning. Ouch! Poor rats!

Operant conditioning is the basis of most animal training. For instance, you might give your dog a biscuit
or a "Good dog!" every time it sits, rolls over, or refrains from barking. On the other hand, cows in a field
surrounded by an electrified fence will quickly learn to avoid brushing up against the fence.

As these examples illustrate, operant conditioning through reinforcement can cause animals to engage in
behaviors they would not have naturally performed or to avoid behaviors that are normally part of their
repertoire.

Learning and cognition

Humans, other primates, and some non-primate animals are capable of sophisticated learning that does
not fit under the heading of classical or operant conditioning. Let's look at some examples of problem-
solving and complex spatial learning in nonhuman animals.

Problem-solving in chimpanzees
The German scientist Wolfgang Köhler did some of the earliest studies on problem-solving in chimpanzees.
He found that the chimps were capable of abstract thought and could think their way through possible
solutions to a puzzle, envisioning the result of a solution even before they carried it out.

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 For example, in one experiment, Köhler hung a banana in the chimpanzees' cage, too high for them to
reach. Several boxes were also placed randomly on the floor. Faced with this dilemma, some of the
chimps—after a few false starts and some frustration—stacked the boxes one on top of the other, climbed
on top of them, and got the banana. This behavior suggests they could visualize the result of stacking the
boxes before they actually carried out the action.

Spatial learning in rats

Learning that extends beyond simple association is not limited to primates. For instance, maze-running
experiments done in the 1920s—maze shown below—demonstrated that rats were capable of complex
spatial learning.

In these experiments, rats were divided into three groups:

•
• - Group I: Rats got food at the end of the maze from day one.
•
• - Group II: Rats were placed in the maze on six consecutive days before receiving food at the end of the
maze.
•
• - Group III: Rats were placed in the maze for three consecutive days before receiving food at the end of
the maze.

Not surprisingly, rats given a food reward from day one appeared to learn faster—had a more rapid drop in
their number of errors while running the maze—than rats not given an initial reward. What was most
striking, however, was what happened after the Group II and III rats were given food.

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 In both groups, the day after the food had been provided, the rats showed a sharp drop in number of
errors, almost catching up to the Group I rats. This pattern suggested that the Group II and III rats had, in
fact, been learning efficiently, building a mental map, in the previous days. They just didn't have much
reason to demonstrate their learning until the food showed up!

These results show that rats are capable of complex spatial learning, even in the absence of a direct
reward, in other words, without reinforcement. Later experiments confirmed that the rats make a
representation of the maze in their minds—a cognitive map—rather than simply learning a conditioned
series of turns.

Watch “Animal communication”

Read: “Animal communication”

Key points
• - Communication is when one animal transmits information to another animal causing some kind of
change in the animal that gets the information.
•
• - Communication is usually between animals of a single species, but it can also happen between two
animals of different species.
•
• - Animals communicate using signals, which can include visual; auditory, or sound-based; chemical,
involving pheromones; or tactile, touch-based, cues.
•
• - Communication behaviors can help animals find mates, establish dominance, defend territory,
coordinate group behavior, and care for young.
•
Introduction
Have you ever wondered how ants follow what seem to be invisible trails leading to food? Why male dogs
mark their territory by peeing on bushes and lampposts when you take them for a walk? What birds are
saying to one another when they chirp outside your window?

If so, you're in the right place! In this article, we'll take a look at these—and many other—forms of
communication used in the animal kingdom.

Communication takes many forms

Communication—when we're talking about animal behavior—can be any process where information is
passed from one animal to another causing a change or response in the receiving animal.

Communication most often happens between members of a species, though it can also take place
between different species. For instance, your dog may bark at you to ask for a treat! Some species are
very social, living in groups and interacting all the time; communication is essential for keeping these
groups cohesive and organized. However, even animals that are relative loners usually have to
communicate at least a little, if only to find a mate.

What forms can communication behaviors take? Well, animal sensory systems vary quite a great deal. For
instance, a dog's sense of smell is 40 times more acute than ours! Because of this sensory diversity,
different animals communicate using a wide range of stimuli, known collectively as signals.

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 Below are some common types of signals:
• - Pheromones—chemicals
• - Auditory cues—sounds
• - Visual cues
• - Tactile cues—touch
•
In some cases, signals can even be electric!

Where does this diversity of communication behaviors come from? Like other traits, communication
behaviors—and/or the capacity for learning these behaviors—arise through natural selection. Heritable
communication behaviors that increase an organism's likelihood of surviving and reproducing will tend to
persist and become common in a population or species.

In the rest of the article, we'll look at some examples of the many ways that animals can communicate
with one another.

Pheromones
A pheromone is a secreted chemical signal used to trigger a response in another individual of the same
species. Pheromones are especially common among social insects, such as ants and bees. Pheromones may
attract the opposite sex, raise an alarm, mark a food trail, or trigger other, more complex behaviors.

The diagram below shows pheromone trails laid down by ants to direct others in the colony to sources of
food. When a food source is rich, ants will deposit pheromone on both the outgoing and return legs of
their trip, building up the trail and attracting more ants. When the food source is about to run out, the
ants will stop adding pheromone on the way back, letting the trail fade out.

Ants also use pheromones to communicate their social status, or role, in the colony, and ants of different
"castes" may respond differently to the same pheromone signals. A squashed ant will also release a burst
of pheromones that warns nearby ants of danger—and may incite them to swarm and sting.

Dogs also communicate using pheromones. They sniff each other to collect this chemical information, and
many of the chemicals are also released in their urine. By peeing on a bush or post, a dog leaves a mark of
its identity that can be read by other passing dogs and may stake its claim to nearby territory.

Auditory signals
Auditory communication—communication based on sound—is widely used in the animal kingdom.

Auditory communication is particularly important in birds, who use sounds to convey warnings, attract
mates, defend territories, and coordinate group behaviors. Some birds also produce birdsong,
vocalizations that are relatively long and melodic and tend to be similar among the members of a species.

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 Many non-bird species also communicate using sound:
•
• - Monkeys cry out a warning when a predator is near, giving the other members of the troop a chance to
escape. Vervet monkeys even have different calls to indicate different predators.
•
• - Bullfrogs croak to attract female frogs as mates. In some frog species, the sounds can be heard up to a
mile away!
•
• - Gibbons use calls to mark their territory, keeping potential competitors away. A paired male and
female, and even their offspring, may make the calls together.
•
Water, like air, can carry sound waves, and marine animals also use sound to communicate. Dolphins, for
instance, produce various noises—including whistles, chirps, and clicks—and arrange them in complex
patterns. The idea that this might represent a form of language is intriguing but controversial.
Visual signals
Visual communication involves signals that can be seen. Examples of these signals include gestures, facial
expressions, body postures, and coloration.

Gesture and posture are widely used visual signals. For instance, chimpanzees communicate a threat by
raising their arms, slapping the ground, or staring directly at another chimpanzee. Gestures and postures
are commonly used in mating rituals and may place other signals—such as bright coloring—on display.

Facial expressions are also used to convey information in some species. For instance, what is known as the
fear grin—shown on the face of the young chimpanzee below—signals submission. This expression is used
by young chimpanzees when approaching a dominant male in their troop to indicate they accept the
male's dominance.

Changes in coloration also serve as visual signals. For instance, in some species of monkeys, the skin
around a female’s reproductive organs becomes brightly colored when the female is in the fertile stage of
her reproductive cycle. The color change signals that the female can be approached by suitors.

An organism's general coloration—rather than a change in color—may also act as a visual signal. For
instance, the bright coloration of some toxic species, such as the poison dart frog, acts as a do-not-eat
warning signal to predators.

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 Tactile signals—touch
Tactile signals are more limited in range than the other types of signals, as two organisms must be right
next to each other in order to touch. Still, these signals are an important part of the communication
repertoire of many species.

Tactile signals are fairly common in insects. For instance, a honeybee forager that's found a food source
will perform an intricate series of motions called a waggle dance to indicate the location of the food.
Since this dance is done in darkness inside the nest, the other bees interpret it largely through touch.

Tactile signals also play an important role in social relationships. For instance, in many primate species,
members of a group will groom one another—removing parasites and performing other hygiene tasks. This
largely tactile behavior reinforces cooperation and social bonds among group members.

Tactile stimuli also play a role in the survival of very young organisms. For instance, newborn puppies will
instinctively knead at their mother's mammary glands, causing the release of the hormone oxytocin and
production of milk.

What is communication used for?

As the examples above illustrate, animals communicate using many different types of signals, and they
also use these signals in a wide range of contexts. Here are some of the most common functions of
communication:

• - Obtaining mates. Many animals have elaborate communication behaviors surrounding mating, which may
involve attracting a mate or competing with other potential suitors for access to mates.
•
• Communication behaviors surrounding mating are often highly ritualized. For instance, a male may
perform an intricate dance, show off decorative features—such as bright patches or elaborate patterns—or
perform a characteristic song to attract a female. Similarly, males may compete with each for mates
other using ritualized display behaviors, which usually involve posturing and gestural or vocal "threats"
rather than actual fighting.

• - Establishing dominance or defending territory. In many species, communication behaviors are

important in establishing dominance in a social hierarchy or defending territory.

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• Communication, for example, may allow disputes over status or territory to be settled without the need
for fighting. By posturing, vocalizing, or making aggressive gestures, both participants make a relatively
honest advertisement of their ability and willingness to fight. This allows both parties to size each other
up, and the weaker may voluntarily back down.

• - Coordinating group behaviors. In social species, communication is key in coordinating the activities of
the group, such as food acquisition and defense, and in maintaining group cohesion.

Communication may be used, for example, to direct other group members to a food source. Honeybee
foragers use the waggle dance for this purpose, and ants use pheromone trails. Pack-hunting predators,
such as wolves, also communicate to capture prey as a group.

Group members may signal to coordinate defensive behaviors. For example, this is the case when a
crushed ant incites other ants to swarm, or when a monkey gives an alarm call upon spotting a predator.

Communication behaviors can also maintain cohesion within a group or establish social bonds and
relationships. For instance, grooming among primates fosters cooperation and cohesion among group
members.

• - Caring for young. Among species that provide parental care to offspring, communication coordinates
parent and offspring behaviors to help ensure that the offspring will survive.

• Tactile signals exchanged between newborn animals and their mothers, for example, trigger the mother
to provide food and may also stimulate the formation of parent-child bonds through hormone release.

Gull chicks tapping on the red spots on their parents' beaks—see article on innate behavior—is another
example of a communication behavior that favors the survival of offspring.

As these examples show, communication helps organisms interact to carry out basic life functions, such as
surviving, obtaining mates, and caring for young.

Watch “Animal behavior: foraging”

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