The Organization of

Behavior: Neurons and

Hormones
Nervous system controls
behavioral organization:
Flying moth

• Stimulus: ultrasound
form a predatory bat.

• Receptors firing,
neural events

• Response adaptive Pheromone –

releasing
female

• Filter out irrelevant

Risk (Bat) Vs
information Reproduction
(Female)
Different courtship displays of the male ring dove are under the control of different hormones

Hormones help animals keep their

behavioral options straight

A) Court: strutting and bowing:

Testosterone
AROMATASA

B) Nest soliciting: Estrogen

Mechanisms structure an
individual´s behavior
1. Neuronal command centers

2. Clocks the schedule the activity

3. Hormonal system
How neural command centers
organize behavior
Fact – All animals have many behaviors
that they could perform at a given
time in response to many different
stimuli.

Question: How do you avoid

maladaptive behavioral conflicts in
which two or more things are done at
once?
NS endowed with command center

1. Innate releasing
mechanism
2. Central pattern generators
3. Song control system

Nervous system organized in a

hierarchy of command
centers.

These command centers are in

neural contact, capable of
unified decision making.
Praying matins
Activities sorting by NS:
• Search for mates
• Sunbathe
• Copulate
• Fly
• Dive away from bat
• Stimulus
predatoryàEscape
response

StimulusàCerebroàExcitatory controlàmuscleàMovement
Inhibiting neural
activity
Command center send inhibitory messages
EXP1: Segment´s ganglio
(removed)àsegment´s muscle 3
failed.
4
EXP2: Segment´s ganglio with
electrical stimulusàcomplete
Movement (vigorous)
1 and 2
EXP3: Protocerebral ganglion (brain)
removedà involutary Movements
(walked and grasped)

EXP4: Protocerebral and

subesophageal gangliaàImmobile

Inhibiting the activities makes possible for the mantis to do just one thing at
a time.
Ganglia are command centers (inhibiting) and are controlled by other parts
of the NS.
But mature male manties do not always obey the
“rule” that complete removal of the head
eliminates behavior

http://www.youtube.com/watch?v=KYp_Xi4AtAQ
Blowflys
Studied in 1960 by V. Dethier
Detect rotting food by olfaction and taste with their
feet.
This promotes feeding
Have neural sensors in proboscis that senses sugar
concentration.
1. If sugar concentration > threshold, then feeding
command center starts to fire and sucking begins.
Command center fires for 90 seconds, then feeding
stops.
Repeat
Relationship: NS and feeding
blowfly
2. Feeding command 2
center is inhibited by 3
stretch receptors in
the foregut.

3. If the recurrent nerve

is cut, feeding
continues in 90
second intervals until 1
gut ruptures.
 Command center send inhibitory
messages: Animal´s behavioral priorities
A record of the activity of a
particular neuron in a
cricket’s brain that appears
to act as a command cell
controlling the male’s
chirping call, which is
produced when the male
rubs his wings together.
About 3 seconds into the
recording, the cricket was
Sensory
subjected to a sharp puff of
appendages air that struck its cerci, a
pair of sensory appendages
that project out from the
rear of the abdomen.

What adaptive value do you attach to the apparent proximate

mechanisms that help male crickets make the behavioral decisions
shown here?
Behavioral schedules: demands of
a changing environment
• Some activities are
better done at some
times of the day than
others.
• “calling” change
cyclically

Hide Move

Calling
Mechanisms that regulate the
changes
1. Animals change their priorities in
response to a timing mechanism with a
built-in schedule that acts independently
of any cues from the animal´s
surrounding (Time zones).

2. Animals alter the relationship among

command centers in their nervous
systems strictly on the basis of information
gathered by mechanisms that monitor the
surrounding environment.
Circadian Rhythms
24 hour cycles of behavior
change
Period of activity and inactivity
(often sleep).

Two hypotheses for controlling

circadian rhythms
1.Run by an internal clock
2. Response to external Fr:
environmental changes
Ex. Crickets calling/moving after Ultradian: <90 min
dark.
Circadian: 24 h

Infradian: 28 day
365 days
Circadian rhythms in cricket calling behavior

• Constant bright light: Crickets still continue to

callàBiological clock

desynchronized
hour
Free-running cycle: Days
period deviates.
1.Run by an internal
clock
Reset

Crickets use to
adjust their timing
Synchronized

mechanism:
ENTRAINED

• Enviroment-activated device sychronizes the clock

How do circadian mechanisms work?

• Visual signals (Nerve

sensory information
1
from eyes to optic
lobes) are needed to
2
entrain the daily
rhythm to local
conditions.
• Rhythm persists in
the absence of this
information: Free-
running cycle
A master clock may regulate mechanisms controlling circadian rhythms within individuals

If optic lobes is separate from

the rest of the brain

Calling cycle break down

What is the Clock?
SCN – suprachiasmatic nucleus (Hypothalamus
vertebrates)
This contains the timing mechanism
If you oblate this region (electrically fry it), the brain loses
its rhythm.

Entrainment pathways differ across animals.

Mammals – phototransduction (light to brain) thru vision.
Eyes (retinal LD information) – SCN
A neural pathway

Birds and Reptiles – pineal gland detects light directly

A photo sensitive part of the brain that releases a hormonal signal
to the SCN.
Pineal Transplant Experiment
(Hamsters or birds receive transplants of SCN tissue)

Set 2 birds to have

pineal glands to
inverted light cycles.
A. L/D B. D/L
Put the glands in other
birds with removed
pineal glands
Now have cycle of A or
B donor,
respectively.
Mammalian Clock

SCN is the pacemaker of the clock.

A structure in hypothalamus
Eyes neurons SCN (entrainment pathway)
SCN is linked via neurons to the pineal gland.
Pineal gland secretes rhythmic pulses of
melatonin.
This is the messenger to the rest of the body.
SNC clock in mammals

SCN is a self-sustaining
oscillator.
A “clock” that can work in
isolation
Specific type of neuron in
SCN is the clock
Done with graphing
experiments

Destruction of SNC:
Arrhitmic patterns.
Hormone secretion,
locomotion, feeding
SNC clock operates via rhytmic
changes in gene activity
• PER production varies over 24 h
schedule in concer with tau gene
product

https://www.ibiology.org/genetics-and-gene-
regulation/circadian-clocks/#part-1
The genetics of biological clocks in mammals and fruit flies

1. PER production peak

2. PER
degradation by
Tau
The genetics of biological clocks in fruit flies

1. PER production peak

2. PER
degradation by
Tau

3. TIM
prduction
4. Tau =dbt
controls PER
localization
Mutations of the per gene affect the circadian rhythms of fruit flies
Multilevel regulation of circadian clock

Thalamus

The suprachiasmatic nucleus (SCN) circadian clock is affected by light (photic cues) through the
retino-hypothalamic tract (RHT) leading to use of glutamate (Glu) as a neurotransmitter.
Glutamate receptor triggering induces various intracellular responses, leading ultimately to gene
expression and phase shifts. Non-photic cues involve a variety of other neurotransmitter and
signalling pathways (three of which are shown here).
per gene is relatively inactive should behave in an arrhythmic fashion

PER decreased (care eggs PER increased (well

and nursing lavare, remain defined circadian
within hive) rhythms)
SCN master clock send out chemical
signals to target systems it controls

1. Molecules secreted by SCN are

regulate in a clock’s gene manner.
2. There should be receptor for the
molecules.
3. Experimental administration of the
chemical messenger should disrupt
the normal timing of an animal’s
behavior.
Expression of the gene that codes for PK2 (prokineticin 2) in the SCN (Rats)

PK2 decrease at night

Circadian control of wheel-running by white rats changes when the rats are injected with PK2

• PK2 (prokineticin): messenger that SCN uses to

communicate with target centers in brain (24 h schedule)
Adaptative value

1. Environment-independent
Result: a typical nocturnal mammal
component
To alter the timing of
behivoral cycles
2. Environment-independent
component
automatically becomes active at
about the right time each night,
while retaining the capacity to
shift its activity period gradually to
accommodate the chages in day
lenght that occur as spring
becomes summer, or summer
becommes fall.

To adjust their cycles in

keeping with local conditions
Naked mole-rats lack a circadian rhythm

Day-night cycle biologically

irrelevant:

Darkness, underground tunnels

Patterns of activity are shown for

six individuals from two captive
colonies held under constant low
light
Circannual Cycles: Long-term

Yearly cycles of behavior

Ultimate selection pressure is winter
Summer is transition
Tropics : annual precipitation cycle. Dry Wet.
Circannual rhythms are timed by a biological
clock of some sort.
Pineal? SCN? Not well understood.
Circannual rhythm of the golden-mantled ground squirrel

Hibernation: late fall and frigid winter in an underground

chamber

Constant darkness and Temperature with an abundance of food (Blinded)

Circannual rhythm in a stonechat (Part 1)

Kenyan (wild condition)àEurope (artifitial condition)

TropicàSeason

• Kenya: spring rainy season with insect abundance and

reproductive activity
Circannual rhythm in a stonechat (Part 2)

Annual reproductive cycle shifted out of phase

Circannual rhythm is generated in part by an internal, enviroment-

independent mechanism
The physical environment
influences Long-term cycles
Behavioral rhythms match with particular
features of the animal’s enviroment: sunrise,
sunset, rainy season, day lengths in spring.
Many nocturnal rodents follow this: Clear
avoidance of moonlight

Activity period reflects this fact

Is this run by a clock?

Seem to anticipate moonrise

Can’t locate or identify this clock yet.
5.16 Lunar cycle of banner-tailed kangaroo rats

1. Food dispenser/Time that

released millet seed at
hourly intervals.
2. Black mark represents a visit
made by a rat (timer).
3. Nov-March: active at nigth
only when the moon was not
shining.
4. A shortage of seeds later in
the year caused the animals
to feed throughout the night,
even when the moon was
up, and later still to forage
during all hours of the day.

Large cache of seedin

the fall

Without moonlight : Safe to predators (Coyotes)

Internal Circalunar Clock ?
A cycle of photosensitivity

Testicular growht:
Winter
Spring

Cycle is photosensitive

Birds possess a clock sensitive to light between hours 17 and 19 of each day:
birds sometimes experienced light during the time when their brains were
predicted to be highly photosensitive

Hormone chages leas to development of Reproductive

equipment
A hormonal response to light
Internal regulation of LH: Luteinizing
Darkness hormone ( a hormone released by the
anterior pituitary and carried to the
testes, where it stimulates the growth
of these tissues)

Complete darkness for

different lenghts of
time, then 8h light
exposure
Food intake and reproductive timing in the white-winged crossbill

Break in breeding, even

where food was plentiful

Food abundance?

Short day, long-night non

breeding
5.20 Photoperiod affects testis size in the red crossbill

Constant temperaute
Unlimited acces to food

Environment
Lond day, indep
increases in the
size of gonads
and LH in blood
Hormone
recovery

Explain:
1. The photoperiod-driven mechanism might be a noadaptative holdover from the past
2. Reproductive benefits from Retention of a physiological system
50-day clock

The Reproductive Cycle of Mus musculus

Copulation
Followed by male aggression and infanticide
Kill all young mice in home range (2-3 weeks)
Gradual shift to parental mode when own
offspring born
Weaning of young promotes copulation
again.
 7 weeks after: Weaning of
young promotes copulation
again and infanticide
advantageously.

3-6 weeks Gradual shift

to parental mode when
own offspring born

Kill all (invariably be

directed against a
rival male´s offspring
young mice in home
range
Regulation of infanticide by male house mice (Part 1)

• We know it’s a clock because we can mess with it.

• A timer (somewhere) counts 50 photoperiods after copulation in ♂.
• If you speed up day (24 to 19 hour light cycle)
• You can speed up the cycle to 50 short photoperiods.

• Two different condition

• Fast 11:11 L:D (22 h /day)
• Slow 13.5:13.5 (27 h/day)

• REAL DAYS

• Total number of LD cycles, not the number of 24 h periods, controlled the

infanticidal tendencies of males
Regulation of infanticide by male house mice (Part 2)

No Infanticide to
infanticide 3 “week”

Timing device register the number of LD cycles that have occurred since mating,
and that this information provides the proximate basis for the control of the
infanticidal response
A hormonal effect on infanticidal behavior in laboratory mice

• Relationship between progesterone and male aggresion (Infanticide)

Not detect
the
progesterone
in their
bodies

KO Mice males
that lacked
progesterone
receptors

Progesterone: supress parental behavior in female

Levels slowly fall after copulation, the parental capacity of the male slowly increase in
time for him to avoid attacking his own brood
Testosterone and the control of sexual motivation in male Japanese quail (Part 1)

Bringing the male together to female for yet another copulation is influenced by
the presence of hormone: Testosterone/Estrogen

Genomic
pahtway

Testes Brain
Active after Testosterone
mated

Lovelorn time
Testosterone and the control of sexual motivation in male Japanese quail (Part 2)

Three groups were castrated

Hormonal adjustments affect
reproductive behavior
Japanese quailà 17 B-Estrogen

Prairie voleàVasopresina.

Cichild fishà gonadotropin-releasing

hormone

Midshipman fishàarginine vasotocin

An associated reproductive pattern
• Hormonal organization of reproduction often involves coordinated
changes in gamete production and in sexual activity: associated
reproductive pattern

Court

Mature sperm

Testosterone

Red deer
EXPàCastrated
Digitally produced variation in male facial features

Ovulation in women find males with masculinized facial features especially attractive
The effects of castration followed by testosterone therapy on three male guinea pigs

• Testosterone: court and mating

• Testosterone: court song, not season pattern

Hormonal and behavioral cycles in single-brood and multiple-brood populations

High levels the testosterone are not essential for male sexual behavior in this species

Testosterone

=Mate

Testosterone
Testosterone and female aggression in the dunnock

Aggresion No aggresion

• Female try to keep another females away

from their partner (s)
The chemical structure of testosterone and its diverse effects on physiology and behavior

Survival
changes

Viruses
Bacteria
Parasites

Stting ducks
Fighting
Lower care

• Testosterone fall to almost nothing ouside the breeding

season or after territorial challenges.
• Testosterone interferences with immune system
A survival cost of testosterone: fighting, sitting duck

Testosterone:
• fighting with rivals
• Sitting duck for `redators or
parasites
Testosterone and territorial behavior (Part 1)
5.32 Testosterone and territorial behavior (Part 2)

Estrogen Territoriality?
Estrogen and territorial behavior

Estrogen decreased: sing less and stay father away from simulated intruding rival

Fadrozole decreases Estrogen

This socially driven change in brain steroid metabolism

1. Dehydroepiandrosterone(DHEA),
also known as androstenolone, is
an endogenous steroid hormone.
2. It is one of the most abundant
circulating steroids in humans, in
whom it is produced in the adrenal
glands, the gonads, and
the brain, where it functions as
a metabolic intermediate in
the biosynthesis of
the androgenand estrogen sex
steroids.
3. La 3β-hidroxiesteroide
deshidrogenasa, es
una enzima que cataliza la síntesis
de la progesterona desde
la pregnenolona.
Spring mating aggregation of red-sided garter snakes

• Dissociated
reproductive
pattern
• No circulating
testosterone
(role?)
• No aggresion
• Pineal gland:
Temperature and
court (following
spring)
5.35 Testosterone and the long-term maintenance of mating behavior

• Fall: concentration of testosterone (anticipation)

• Temperature increase cue the activation.

• Testosterone: organizational role
Summary

1. Because an animal´s environment provides various stimuli that

could trigger contradictory responses, and because its physical
and social environment often change over time, animals gain by
having mechanisms that set priorities for their different behavioral
options. One such proximate system includes behavioral command
centers with the capacity to inhibit one another, so that animals
do not try to do several things simultaneously.

2. As the environment changes, the nature of the inhibitory

relationships between neural command centers may also change.
Devices to achieve this end include the various pacemaker or clock
mechanisms that regulate nervous system functioning and
hormonal output in cycles that typically, last either 24 hours or 365
days. Circadian and circannual clocks have environment –
independent components, but they can also adjust their
performance by acquiring information from the environment about
local conditions, such as the time of sunrise or sunset.
Summary
3. Hormones function within those mechanisms that establish
behavioral priorities. In many animals, changes in the physical
environment (such as seasonal changes in photoperiod) and in the
social environment (such as the presence of potential mates) are
detected by neural mechanisms and translated into hormonal
messages. These chemical signals often set in motion a cascading
series of physiological and behavioral changes that make
reproductive activity the top priority at times when it is most likely to
translate into the production of surviving offspring.

4. The precise roles played by hormones in effecting behavioral

change vary from species to species. Male sexual behavior, for
example, may or may not be dependent on high testosterone
concentrations in the blood. Nevertheless, the proximate
mechanisms of behavioral organization of many different species
show similarities, such as a reliance of one sort o another on
testosterone and certain other widely distributed hormones for
organizing male reproductive behavior. This pattern reflects the
nature of evolutionary change, in which the attributes of today´s
species are modified versions of previous ones, not inventions that
have arisen out of thin air.
Memories are stored as alterations in the
strength of synaptic connections between
neurons in the CNS.
Ovarian Hormones,
Aging and Stress on
Hippocampal Synaptic
Plasticity

• Hormones estradiol
and progesterone
regulate a wide variety
of non-reproductive
functions in the central
nervous system by
interacting with
molecular and cellular
processes.

Time
• 17β-estradiol, the most potent of the biologically relevant
estrogens, enhances synaptic transmission and the
magnitude of long-term potentiation