Chapter 4 – Control of Behavior

Neural Mechanisms
Overview

• Focus of this chapter is :

The interaction of proximate and ultimate

analyses of behavioral control systems
 Neural systems of reflexive behaviors
Konrad Lorenz: The Fixed Action Pattern (FAP)
•Graylag goose & egg rolling
•Herring gull & feeding
FAPs appear to be completely innate:
•Reproductive behaviors
•Pheromone guided flight
•Courtship dances/rituals
•Copulation
•Escape responses
•Sensory and Motor Reflexes
Advantage:
•Does not have to be learned.
•Does not change (works the first time every time)
Disadvantage: Mimicry & Parasitism
•Cuckoo, a nest parasite
•Rover beetle nest parasite
•Orchid female insect “decoys” attract unsuspecting
males
•Predatory firefly females “mimic” the call of other
species females and lure males to “dinner”
4.1 A complex response to simple stimuli
4.2 A simple rule of thumb governs this beetle’s mating behavior
4.3 Pioneers in the study of animal behavior

Pioneers in the study of animal behavior: Niko Tinbergen, Konrad Lorenz and
Karl Von Frisch shared Nobel prize in medicine in 1973. These ethologists received
the prize in recognition of their contributions to an understanding of the proximate
causes of behavior
4.4 Begging behavior by a gull chick
A silver gull chick is being fed regurgitated food by its parent after pecking at the
adult’s bill
4.5 Effectiveness of different visual stimuli in triggering the begging behavior of herring gull chicks
4.6 Instinct theory
Code Breaking

• The simple relationship between an innate

releasing mechanism, sign stimulus, and
FAP is highlighted by the ability of some
species to exploit the FAPs of other
species- a tactic known as code breaking
4.7 A code breaker

This young cuckoo begs for

food from its foster parent,
a reed warbler, which provides
for the cuckoo at great cost to
itself and its own offspring
4.8 Bolas spider deception

The bolas spiders release mimetic sex pheromones in order to attract moths to
attack range. When sexually motivated moth comes flying upwind toward the
scent-releasing spider, the predator
swings a sticky ball of glue at
the insect.

here spider exploits moth FAP

(fly upward along a gradient of
pheromone, which usually takes
it to the female of his species
4.9 Complex code breaking by a wasp (Part 1)

Wasps lay eggs into butterfly larvae

Butterfly larvae
are protected
by colony of
red ants who
live on
butterfly
secretions
and in return
protect
butterfly
caterpillars
against any
invading
ants
4.9 Complex code breaking by a wasp (Part 1)

• (A) A parasitic wasp on top of a butterfly larva, that

is its victim
• (B) To get to the larva, which was being protected by
a colony of red ants, the wasp produced a bouquet
of volatile hydrocarbons. These chemicals release a
sequence of responses by red ants, whose reactions
are listed on the left. The responses of ants to wasp
dummies coated with a mixture of these chemicals
and with each component individually are shown in
the five graphs. Note that the result of the wasp’s
multiple mimicry of ant chemical signals is to induce
ants to flee from the wasp and instead to assault
their fellow colony members. The data are presented
as the ratio of the number of ants responding to the
hydrocarbon bouquet (or its individual components)
relative to a control solution composed of the
solvents used to extract the pheromones from the
wasps
4.9 Complex code breaking by a wasp (Part 2)
Moths and Bats

• Classic example of neural control of

behavior
– Fact – Bats eat bugs in flight
• Selection should favor anti-bat behavior in
a night flying insect.
• How do moths avoid bats?
Bat Feeding

• Bats use sonar to detect prey

• High frequency ultrasonic pulses
• Bat feeding buzzes announce their
presence to those that can detect high
frequency sounds.
Moth Hearing

• Moths have “ears” on the sides of the

thorax
• These structures are sensitive to high
frequency sounds and deaf to others.
4.11 Noctuid moth ears
Basic Neurophysiology

• Neuron – nerve cell

• Sensory neuron – carry information from
sensory organ to the central nervous
system (CNS)
• CNS – brain and spinal cord
– Most complex processing of information is
done here
• Motor neuron – carry information from
CNS to muscle groups
 Route of Transmission

• Ear Sensory neuron Interneuron

Ganglion Motor neuron Muscles
4.12 Neurons and their operation

Electric activity in neuron

originates after effects
of stimuli on dendrites.

Sufficiently great electrical

changes in dendrite’s cell
membrane can trigger
an action potential.

The action potential

begins near the cell
body and travels along
the axon toward the
next cell in the network
4.13 Neural network of a moth

Receptors in the ear

relay information to
interneurons in the
thoracic ganglia

Thoracic ganglia
communicate with
motor neurons

Motor neurons
control wing muscles
4.14 Properties of the ultrasound-detecting auditory receptors of a noctuid moth (Part 1)

Sounds of low or moderate intensity do not generate action potentials in the A2 receptor
The A1 receptor fires sooner & more often as sound intensity increases
4.14 Properties of the ultrasound-detecting auditory receptors of a noctuid moth (Part 2)

The A1 receptor reacts strongly to pulses of ultrasound, but ceases to fire after a short
time if the stimulus is a constant sound
4.15 How moths might locate bats in space (Part 1)

The A1 receptor on the side closer to the bat fires sooner and more often
than the shielded A1 receptor in the other ear
4.15 How moths might locate bats in space (Part 2)

When a bat is directly behind the moth, both A1 receptors fire at same rate and time
4.15 How moths might locate bats in space (Part 3)

When bat is above the moth, activity in A1 receptors fluctuates

in synchrony with the moth’s wing beats
4.16 Bat ultrasonic cries trigger evasive behavior in a number of insects
Multiple-exposure photograph:
Normal flight- forelegs close to the body

When detects ultrasound rapidly extends When approached by hunting bat

its forelegs employs an anti-interception power dive
4.17 Is the A2 cell necessary for anti-interception behavior by moths? (Part 1)

• A: a sonogram of the ultrasonic pulses produced

by a bat approaching and then attacking a prey
• The bat’s ultrasonic cries speed up greatly and
become much more intense as it closes on the
prey (the terminal buzz phase)
• B: a sonogram of part of the approach portion of
the bat’s cry is shown over the response of the
A1 and B receptor cells in the noctuid moth’s ear
(the B cell is a nonauditory neuron)
4.17 Is the A2 cell necessary for anti-interception behavior by moths? (Part 1)
4.17 Is the A2 cell necessary for anti-interception behavior by moths? (Part 2)

• C: A sonogram of the initial part of the bat’s

terminal buzz is shown over the response of A1,
A2, and B cells in the moth’s ear
• The A1 receptor fires more often in response to
the terminal buzz than to the approach
component of bat’s cry
• D: During the latter part of the bat’s terminal
buzz, only the noctuid’s A1 cell is active, not the
A2 receptor
4.17 Is the A2 cell necessary for anti-interception behavior by moths? (Part 2)
What Does This Mean?

• A1 receptor – fires rapidly but slows after a bit

of a constant buzz (phasic)
• A2 receptor – fires only on high intensity calls
(bat near)
• A1 sensitive to both low and high intensity bat
feeding buzzes.
• Rate of A1 “firing” as buzz intensity
• A1 responses to pulsed sound
• Responds to 20-50 kHz sound
• A2 responds only to high intensity sound
Orientation

• Moth ears can tell

location of bat by
differences in
signal received on
left and right side
of the body.
• When the signal is
even on both sides
– bat is parallel.
• If bat is above,
detected during the
up wing beats with
a high firing rate.
Summary

• A1 is main bat detector

– Can detect a bat at 30 m.
– As A1 rate of fire increases, moth should turn
away from bat to reduce sonar echo.
• A2 is emergency system.
– Initiates erratic flight and last ditch effort to
evade capture
4.18 Avoidance of and attraction to different sound frequencies by crickets (Part 1)

• A: In the absence of sound, a flying tethered

cricket holds its abdomen straight
• B: If it hears low-frequency sound, it turns
towards the source of sound
• C: If it hears high-frequency sounds it turns
away.
(Males of this species produce sounds in the 5
kHz range; some predatory bats produce high-
frequency call of about 40 kHz)
Sensory analysis
in humans
brain evolved in
response to
selection
pressures
associated
with
particular
physical and
social
environments
4.31 Sensory analysis in humans and naked mole-rats

Brains evolved
in response to
selection pressures
associated
with particular physical and social environments

The comparison shows

Two mammalian species; the smaller drawing
shows the actual anatomical proportions while
the larger
drawing shows
how the body is
proportionately
represented
in the somatosensory cortex
Adaptations and The Proximate Mechanisms of Behavior

• Individual cells and neural networks have

special operating rules that filter the
information an animal receives, relays,
and processes
• These rules adaptively change the ways in
which members of different species
perceive their environments.
• Ex- many bees can see UV light which we
cannot see.
Adaptations and The Proximate Mechanisms of Behavior

• Ultraviolet-reflecting patterns have great

biological significance for some species
• A set of photographs on the next slide:
• The image on the left shows the organism as it
appears to human
• While the image on the right shows the
organism’s UV-reflecting (pale) surfaces
• Only males (top specimens) of this sulphur
butterfly species have UV-reflecting patches on
their wings, which helps signal their sex to other
individuals of their species
4.32 Ultraviolet-reflecting patterns have great biological significance for some species
4.33 A bird that can sense ultraviolet light

The male bluethroat’s throat feathers appear

Purely blue to us, but not to bluethroats
Themselves, which also see the UV
Light reflecting from the feathers
4.34 The ultraviolet reflectance of the throat patch affects mate choice in the bluethroat

Males with experimentally reduced UV

reflectance were slower to attract mates
than control males

Males with reduced UV reflectance also

obtained fewer copulations with females
that were not their primary mates
Adaptive mechanisms of human perception

• Bluethroats and suphur butterflies have nervous

systems with perceptual capabilities to meet
their environment needs.
• Apply this to human beings and we could predict
that
• Natural selection should have endowed us with
proximate mechanisms suited to the ecological
conditions we face
• Evidence suggests that highly specialized
auditory skills have evolved to match our social
environment
• A dominant auditory skill is the language
• Our visual perception complements our
acoustical analysis of language
• Listener’s perception of spoken language is
heavily influenced by visual cues—like speaker’s
lip movement
Adaptive mechanisms of human perception

• Lip reading depends on specific neural

clusters within visual cortex
• Specifically a part of brain called superior
temporal sulcus becomes active when
we see moving mouths, hands, and eyes
4.35 Relevant movements activate neurons in different parts of the superior temporal sulcus

Socially relevant movements of lips, mouth, hands, and body activate neurons in
different parts of the superior temporal sulcus in human brian. The right and left
hemispheres are shown here. Each circle represents findings by a particular
research team

So natural selection has endowed us with remarkable ability to detect even subtle
movements of these body parts .
This enables an individual not only to read lips but also to deduce the intensions
of other people--- a very useful ability for members of a highly social species.
So we are equipped to predict the actions of those around us
Fusiform gyrus and our capacity to identify familiar faces

• A technique—functional magnetic resonance

imaging- used to identify the part of cerebral
cortex which becomes active when a person
looks at a picture of a face
• A part of brain on the underside of the cerebral
cortex-------fusiform gyrus
• Critical role in identifying familiar faces quickly
and accurately
• Damage to this brain part affects this capacity
• Still the person is able to recognize people by
their voices but not by faces…even his own face
4.37 Specialization of function in different parts of the visual cortex of humans

Different neural circuits perform different analyses of images in our environment

The ability to remember faces, for example, is dependent on a specialized site in
the fusiform gyrus

The sites that respond

strongly to images of entire
faces are different from those that react primarily to other parts of face,
like eyes, which in turn are different from those that are activated when the subject is
shown a picture of an object
Adaptive mechanisms of navigation

• Humans’ is a very social environment

• Recognizing faces and deducing
intentions of others around you has
adaptive value
• So, natural selection has endowed
humans with extraordinary capabilities
• Natural selection has shaped our nervous
system
Adaptive mechanisms of navigation

• Ancient man was a hunter-gatherer who had to

travel great distances away from his home base
in search of food and other resources
• We should be expecting this legacy in our brain
structure as well
• Natural selection has shaped certain areas in
our brain for navigation purposes
• Like food storing birds have enlarged
hippocampus
• Magnetic resonance imaging has been used to
monitor human hippocampus activity
4.39 The hippocampus is essential for navigation by humans (Part 1)

An aerial view map of the virtual town through which experimental

subjects navigated. Three examples of attempts to get from A to B
are shown. The shortest route in yellow was considered the
most accurate
4.39 The hippocampus is essential for navigation by humans (Part 2)

The location of right hippocampus

in a brain viewed from the underside

Average magnetic resonance image for ten

subjects while performing the navigation task,
showing the location of peak neural activity
(bright yellow), which lies in the right hippocampus
4.39 The hippocampus is essential for navigation by humans (Part 3)

The accuracy of the ten navigators was a function of the intensity of the
Activity in their right hippocampus
Navigation: human versus some animals

• Humans without a map and a compass

are handicapped when it comes to
navigating across unfamiliar areas.
• But many other animals show no such
disability because they possess an internal
map sense(knowing location of home or
some other goal) and an internal compass
sense (knowing in which direction to
move)…..like honey bee and homing
pigeon
4.40 The ability to navigate unfamiliar terrain requires a compass sense and map sense (Part 1)

Compass sense: knowing in which direction to move

Map sense: knowing location of a goal

The flight path taken by a wandering albatross on a foraging journey of over 4000
kms from its nest in the Crozet Island in the southern Indian Ocean, north of
Antarctica, and back again
4.40 The ability to navigate unfamiliar terrain requires a compass sense and map sense (Part 2)

A trip of 592 meters by a foraging ant out from its nest (at the large open circle at the
bottom) and then directly back home after capturing a prey (at the spot marked with
a large red circle), 140 meters away from home
4.41 Clock shifting and altered navigation in homing pigeons

The results of an experiment in which

pigeons were released at Marathon,
New York, about 30 kms east of Ithaca,
where their home loft was located. On
sunny days, birds in the control group
generally headed west, back toward the
home loft. But pigeons whose biological
clocks had been shifted by 6 hours,
usually misoriented, so that, on average,
they headed north
4.42 The fall migration route of monarch butterflies takes some butterflies from Canada to Mexico

Monarchs cluster in a few forest patches high in the central Mexican mountains
Experimental manipulation of the biological clock

• Individuals were tested in an outdoor flight cage after:

• one group had been held indoors under an artificial
light-dark cycle with lights on at 7.00 am and off at 7.00
pm. This group tried to fly in southwesterly direction
• A second group of butterflies that had been held with
lights on at 1.00 am and lights off at 1.00 pm. Flew in a
southeasterly direction
• Clock-shift changed the orientation of migrating
monarchs

• An evidence of the importance of a sun compass in

helping monarchs stay on course.
4.43 Manipulation of the biological clock changes the orientation of migrating monarchs
Polarized light affects the orientation of monarch butterflies

• What wave lengths of sunlight are critical for the navigational

orientation of monarch butterflies?
• Experimental evidence: sun’s UV radiation is essential for monarch
navigation
• UV light helps monarchs get started flying in the right direction
• But once airborne, they navigate with the help of cues provided by
the polarized skylight pattern
• We can’t see polarized light, which is produced when sunlight enters
the Earth’s atmosphere and the light scatters so that some light
waves are vibrating perpendicular to the direction of actual sunrays.
• The three-dimensional pattern of polarized light in the sky created in
this fashion depends upon the position of the sun relative to the
Earth so this pattern changes as the sun moves across the sky.
• If an animal on ground is able to perceive the pattern of polarized
light in the sky, this stimulus can serve as a proxy for the position of
the sun at any given time.
4.44 Polarized light affects the orientation of monarch butterflies
Skylight e-vector: angle of polarized light reaching monarchs from the sky

When horizontal filter shifted

the angle of polarized light
monarchs altered their flight orientation accordingly by 90 degrees