HMB200 Lecture 8: Motivated Behaviours and Falling in Love

HMB200 Lecture 8
Motivated Behaviours and

Falling in Love
1
Get to know my students
• Is the course helping you to become a pediatric psychiatrist?
• What literature do you love?
• How is your brain rewired when playing video games?
Guess where I’ve been…
2
Lecture 8 outline
Motivated and habitual behaviours, and falling in love
1. Homeostasis and goal directed behaviours
2. Effector Systems for goal directed and habit directed behaviours
3. How are goal directed behaviours shaped, and habits formed?
4. The role of dopamine in goal directed and habit directed behaviours (the error predictor)
5. Dopamine and the pair bond
6. Oxytocin and vasopressin and the pair bond
7. The partner engram associated with the reward system
Great Resources to Prepare
In Class Team UP! Activity based on the following video:
https://play.library.utoronto.ca/bjlgOSnj3FO6Links to an external site.
And the highlighted sections of this reading.
Young Wang_2004_The neurobiology of pair bonding 2019.pdf
The video and reading are required readings for the course and are testable.
3
The active maintenance of a relatively Homeostasis
constant internal environment is called
homeostasis. Constancy of the internal
environment is the basis of the freedom of
action we and other animals enjoy because
it partially decouples our physiology from
immediate external conditions and greatly
extends the range of available habitats. Stimulates
goal directed
As the internal environment moves out of behaviours
balance (e.g. low energy, low water),
behavioural effectors are stimulated to
trigger goal directed behaviours (return to
balance).
The more the imbalance, the stronger the

behavioural effectors are stimulated.
Low imbalance = low motivational state

Large imbalance = high motivational state
(stronger effector of goal directed behavour)
What are goal directed behaviours?
There are many different types of goal directed There behaviours are driven by motivational states.
behaviours. These can be internal (e.g.)
Some are involved in tightly regulated • Hunger
homeostatic mechanisms: • Thirst
• Eating • Temperature sensors
• Drinking
• Sexual arousal
• Finding warmth and comfort
• Curiosity to explore
Some are involved in seemingly non-regulated
but adaptive systems These can be external cues (e.g.)
• Foraging or simply exploring • Sight of food or water, or potential mate
• Reproductive behavior • Time of day
Neuroscientists discovered that there is

are strongly conserved neural pathways
that seem to be involved in triggering
many of these behaviours.
Example: Electrical stimulation of the
medial forebrain bundle.
5
Some Goal Directed Behaviours
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The human version of foraging or random exploration
7
Goal directed behaviours are regulated – homeostasis
Some states (e.g. hunger, thirst,
There behaviours are driven by motivational states. temp) are regulated
These can be internal (e.g.)
• Hunger
Example homeostatic
• Thirst
regulation feeding behviour
• Temperature sensors
• Sexual arousal
• Curiosity to explore
These can be external cues (e.g.) Satiety = Hunger =
• Sight of food or water No Goal directed
behaviour Behaviour
• Time of day
8
Other goal directed behaviours are non-regulated but
homeostasis-like
There behaviours are driven by motivational states. Example: Exploring or
These can be internal (e.g.) sexual behaviour
• Hunger
• Thirst
• Temperature sensors
• Sexual arousal
• Curiosity to explore “Satiety” “In the Mood”
These can be external cues (e.g.)
Possible
• Sight of food or water, or potential mate Possible
mechanism mechanism:
• Time of day, seasonal Higher Dopamine Low Dopamine
9
Conserved motivational states/behaviours
https://www.youtube.com/watch?v=MixNh9L7G5M
Almost 100 years ago, Cannon’s classic research revealed that motivation states are highly conserved and trigger greatly overlapping
behaviours. E.g “Hangry” phenomenon (MacCormack and Lindquist, 2018). Another startling example of this: controversial observation of
1,000+ judges hearing parole decisions in Israel—the decisions at the beginning of the day started 65% positive, steadily plummeting to less
than 10% right before a meal break, only to jump back up to 65% after the meal (Danziger et al., 2011).
10
https://tyelab.org/wp-content/uploads/tyereview2018.pdf
https://tinyurl.com/hmb200qa
11
Lecture 8 outline
12
Effector systems for goal directed behaviours
Recall: The Amygdala effector reflexive behaviour:

Hypothalamus (neural and hormonal)
AND
“The medial forebrain bundle” Brainstem circuits
Stimulation here triggers those
behaviours (very powerfully!)
Motivational Similar concept (and greatly overlap brain regions) for goal directed behaviours
state or cues
Frontal Cortex
Goal directed behaviours
Goal directed behavioural effectors
Ventral Tegmental Area Amygdala
Substantia Nigra Dorsolateral Striatum Hypothalamus (neural and hormonal)
Dorsomedial Striatum AND
Nucleus Accumbens (Ventral Striatum) Brainstem circuits
(Ventral Pallidum)
13
Habit memory systems form from cue associated goal
directed behaviours
N N
W E W E
S S
Recall: Starts as goal directed behaviour (foraging)…and then became a habit based behaviour.
RIGHT: (Notice how powerful the habit memory is: the animal does not notice a different starting location
and most still turned left)
Effector systems for goal directed and habit behaviours
Amygdala (BLA, CEA)
Dorsolateral Striatum
The Frontal Cortex Dorsomedial Striatum
Blue structures are involved in Nucleus Accumbens (Ventral Striatum)
input can modulate
motivational state directed action (Ventral Pallidum)
the outputs
Orange structures are involved in habitual

responding action
Ventral Tegmental Area

Substantia Nigra Behavioural pathways include
• motivational state directed
• habit memory system directed
Motivational
state or cues
Hypothalamus
Blue structures are involved in Brainstem mediated
motivational state directed action behaviours
BOTH behaviour systems involve
NO
Dopamine signaling GO
GO
https://www-sciencedirect-com.myaccess.library.utoronto.ca/science/article/pii/S0197018618306776#fig2 15
16
Lecture 8 outline
17
Operant conditioning
Consequences influence Behaviours
Classical conditioning
Cues trigger reflexes
Behaviour Consequence
Nose Poke I got a treat
Situation I was placed in a

blue box
Consequence
I was hurt
Behaviour
Fear
https://www.youtube.com/watch?v=WEWJH-W38ZQ
Operant Conditioning – it feels good so do it again
https://www.youtube.com/watch?v=WEWJH-W38ZQ 19
Classical conditioning Operant conditioning
Amygdala
Fear
Behaviour Consequence
Nose Poke I got a treat
Dopamine
reward
system
In the classical conditioning, the representation of a CUE (very valuable – it is a predictor) with the US
overlaps in time. The association can be easily explained through associativity principle, in which a response
to a neutral stimulus becomes strong (both at the neuronal and behavioural levels) due to temporal overlap
between the two stimuli. Note that the classical conditioning is passive. The situation and consequence happen
to the rat. The rat can form the association even without explicit awareness.
However, to survive, the animal must act and behave (e.g., find food, mate, shelter, protect young, escape
predators). In the operant conditioning, the rat is an active participant. It must act towards reaching a goal
and to learn the consequence of its behaviour.
Dopamine signals through the goal directed pathway likely strengthens that behavioural ouput. It is still
dependent on motivational state (ie. when the animal isn’t hungry, it will nose poke less)
Operant conditioning
Consequences influence Behaviours
AND
Classical conditioning like phenomenon can also occur:
Situation Behaviour Consequence

Light’s ON Nose Poke I got a treat
Dopamine (the reward) may ALSO strengthen the habit pathway (the CUE becomes like a CS,
with the dopamine signal acting like the US to strengthen that synapse).
This habit pathway is LESS dependent on behavioural state, and more simply CUE dependent
Everyday examples – operant conditioning
Operant conditioning: the consequence affects
the frequency of the behaviour that preceeded it.
I happen to walk I walk in and MMMMM

by this sign order fries delicious
Reward
I will walk in
more often,
especially when
I’m hungry
Operant conditioning increases the specific rewarded goal
directed behaviour I walk in I will walk in
and order more often,
fries especially when
I’m hungry
GO
Next time your motivational state directs

(e.g. you are hungry) you will more likely
repeat this behaviour compared to other
foraging/feeding behaviour.
E.g. Hunger
Operant conditioning will increase

GO THIS SPECIFIC motivational state directed behaviour
23
Everyday examples – operant conditioning and habits
Classical conditioning Operant conditioning: the consequence affects
the frequency of the behaviour that preceeded it.
THE SIGN!! I walk in and MMMMM

I’m walking in! order fries delicious
Reward
I will walk in
Classical conditioning – like more often,
especially when
The “CUE” becomes like a CS I’m hungry
to trigger the behaviour
Increase in the specific goal directed behaviour AND habit
I’m hungry, I
THE SIGN!!
walk in and
I’m walking in!
order fries
Operant conditioning will increase

THIS SPECIFIC motivational state directed behaviour
Classical Conditioning will increase
CUE triggered goal directed behaviour (which is not so
dependent on “state”)
E.g. Hunger
GO
25
Reward in the brain
The Amygdala
Recent studies have demonstrated that the amygdala (which we know plays a role in “Defensive” behaviours –
through emotional/stress processing by classical conditioning) ALSO plays a role in reward based behaviours.
Neurons in the basolateral amygdala (BLA) respond to both rewarding and aversive stimuli, and are connected to
output systems for both defensive (through CeA) and appetitive (through NAc/striatum) behaviours.
Neurons with positive and negative valence seem to be intermixed withing the BLA. However, some studies have
demonstrated that activation of the BLA neurons whose axons are in the central amygdala result in defensive
behaviour. Activation of the BLA neurons that project to the nucleus accumbens biases animal towards an
appetitive response.
Reward in the brain
The Nucleus Accumbens
And Striatum
Hi dopamine stimulates behaviour
LO dopamine stimulates behaviour
The dopaminergic system in the brain is highly complex. Many debates about
its function remain. But there is no doubt that is plays the key role in motivated
learning and reward representation.
A very simplified view, that we will follow to learn some concepts:
In one pathway, dopamine signals “reward”, and lack of dopamine might lead
to activation of those behaviours, analogous to “empty stomach”.
In another pathway, dopamine seems to be important to learn about rewards

(i.e. the habit system) and high dopamine there may be activate those
behaviours (habits) to obtain the reward – e.g “turn left”, follow the
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Lecture 8 outline
29
Dopamine signals reward
At first (never seen the sign before):

dopamine surge when eating the fries
(same with any rewarding stimulus)
30
Dopamine changes to signal the predictor of reward
Seminal research that shaped our

understanding of dopaminergic system
And Dopamine will signal a prediction error
As predicted
Greater
Prediction error
Prediction error
Seminal research that shaped our

Wolfram Schultz et al. Science 1997;275:1593-1599 understanding of dopaminergic system
Let’s go back in time. This is your first time at McDonald’s. You see the logo for the first
time. You try their fries for the first time.
What is the likely DA signalling in the nucleus accumbens when you see the logo and
eat the fries (increase, decrease, no change)?
DA signalling
Time
First time
Now to the present day. You’ve been going to McDonald’s so many times!
You arrive, see the logo, eat the fries.
DA signalling
Time
100th time
One day you come to your local McDonald’s. You are regular there. The attendant
decides that as a reward you should be getting 2 for the prices of 1!
DA signalling
Time
2 for 1!!!
You keep coming and each time the kind attendant still serves you double…
After a long time…

DA signalling
Time
100th time
2 for 1
Next day you walk in, the kind attendant no longer works at McDonald’s.
The new attendant gives you just a regular serving.
DA signalling
Time
First time
back only 1
You keep going in, and only get a regular serving each time.
DA signalling
Time
100th time
only 1
You arrive at McDonalds. The attendant informs you that the fry machine is broken.
No fries today.
DA signalling
Time
Broken!
You’ve been coming here every day for the past two weeks and every time you hear
“the fry machine is broken”. Still no fries.
DA signalling
Time
15th time
Broken!
Dopaminergic signalling
Response to reward
Dopamine responses in rat nucleus

accumbens distinguish between a reward-
CS+ predicting conditioned stimulus (CS+) and
a non-reward-predicting conditioned
stimulus (CS−). The dopamine release
CS- comprises an initial indiscriminate
detection component and a subsequent
identification and value component
Da signalling is actually more complicated

than that – there is an initial detection
increase, then a valuation component
https://www-nature-com.myaccess.library.utoronto.ca/articles/nrn.2015.26.pdf?origin=ppub
Dopamine isn’t pleasure – it’s the anticipation of pleasure
Dopamine isn’t about happiness – it’s about the pursuit of happiness
https://www.youtube.com/watch?v=axrywDP9Ii0 42
Effector systems for motivated behaviours
Behavioural pathways include
The Frontal Cortex • motivational state directed
input can modulate • Habit memory system dependent
The reward
the outputs
GO
Conditioned Stimuli
(individual stimuli or
contexts)
Thalamus, sensory
The reward cortices,
hippocampus/
memory systems NO
GO
43
Supplemental: The brain is more complex than that :(
https://www.sciencedirect.com/science/article/pii/B9780128022061000350?via%3Dihub 44
45
LEFT: Images of NASA’s Perseverance and its photo of the Mars’
terrain on sol 18 after landing.
RIGHT: One of the first images that appeared on the web after the
Perseverance's landing.
GUESS WHAT WAS SEEN WHEN YOU ZOOMED IN JUST AT THE

HORIZON??
Break: Let’s turn our camera and show your favourite
pandemic “haircut”!
Lecture 8 outline
49
How do we fall in love?
Reward Pair bonding
Pair bonding is a term used in biology and behavioural sciences

to describe a strong social relationship between individuals in a
breeding pair in monogamous species.
Summary Team Up! Instructions - ONLINE
Make visible the participants panel. Slide this up

Instructor will place you into breakout rooms.
1. The first name in your room is the Leader (“Driver”) – Driver does step 2
2. Step 2—Driver clicks CREATE NEW GROUP and shares Group ID
with group members (in chat box) Other group members WAIT
3. Step 3 Other Group members fill in Group ID and click JOIN Group
(DO NOT CLICK Create New Group)
4. WAIT till all group members have joined (type in chat box when you have joined)
then DRIVER SELECTS Q1
5. Use the Chat box and/or microphone to discuss questions – each group member may select their
preferred answer (“Driver” will see these choices) –
6. Driver selects the FINAL ANSWER FOR THE GROUP – then move to next Q
7. When quiz is done DRIVER SELECTS SUBMIT → Instant feedback If Correct = 5 point
NOTE: We will use the same system for Stage 2 of your If wrong, try again → 3 points
midterm in tutorial. If wrong, try again → 2 points etc. 51
Step 3 Top name in breakout roomis
the leader (driver)
Step 4—Leader clicks CREATE NEW GROUP

and shares Group ID with group members
Step 1 Other group member WAIT
Step 5
Step 2 Other Group members fill in Group
Click GO number and click JOIN Group
TO TOOL (DO NOT CLICK Create New Group)
Step 6: DRIVER SELECTS Q1 to begin – Click Submit when you are done
Why do some species pair bond and other do not?
Prairie voles and meadow/montane voles do not appear distinct but they have several key
behavioural differences which makes their goals and motivations dramatically different.
Prairie voles form monogamous, long lasting bonds. Both male and female contribute to
raising the young. Meadow/montane voles spend most of their lives solitary. Only females
participate in caring for the young.
Neuroscientists took this opportunity to discover the underlying neurobiological mechanisms

that lead some to fall in love and stay together and others to always remain loners.
Partner preference test is a measure of pair bonding
Partner preference test is designed to quantify selective social

bonding behaviour.
Single male and female voles are placed together for 24 hours
(co-habitation). They may or may not mate during this time
(however mating facilitates pair bonding).
After co-habitation, a vole is placed in a chamber, where it can
make a choice to socialize with the partner, a stranger or neither.
Prairie voles spend much more time with the
partner than with a stranger vole or alone in
the middle chamber. (They pair bond!)
In contrast, montane voles prefer to spend
time alone in the middle chamber.
How do pair bonds form? Dopamine
In all species, DA signals reward, mating is a big trigger for DA release. By
simple operant conditioning, mating is reinforced.
However, it seems that only in pair bonding species, does mating (and DA
release) support pair bonding.
In female prairie voles, treatment with a D2R antagonist before co-
habitation blocks partner preference.
In female prairie voles, treatment with a D2R agonist facilitates partner
preference even in the absence of mating!
But why only in prairie and not meadow voles?

57
Lecture 8 outline
58
Oxytocin
Parental behaviour, empathy, social bonding, individual
discrimination, mother-infant bonding
In prairie voles In meadow voles,

Hi levels of OTR in In prairie voles: OTR antagonist in NAcc or
Lo levels of OTR in
NAcc, PFC and CP PFC blocks partner preference.
NAcc, PFC and CP
In females, a key difference between monogamous and non-monogamous voles is the

distribution of the oxytocin receptors (OTR). OTR density is significantly greater in the NAcc, PFC
and CP (caudate putamen) of the prairie voles compared to meadow voles.
NOTE: Most non-pair bonding species retain similar maternal/paternal infant bonding behaviours
Vasopressin
Social recognition, territorial marking,
aggression, mate-guarding
In prairie voles: AVPR antagonist in ventral

pallidum blocks partner preference.
Amazingly: Artificial expression of AVPR in

ventral pallidum of male meadow voles
(and mice!) makes them more likely to form
pair bond and show partner preference!
In males, a key difference between monogamous and non-monogamous voles is the

distribution of the arginine vasopressin receptors (AVPR). AVPR density is significantly
greater in the ventral pallidum of the prairie voles compared to meadow voles.
D2R support bonding, D1R also support bonding
Male and female prairie voles that are sexually naïve show
affiliative behaviours towards stranger conspecifics.
AFTER MATING (and bonding) prairie voles display aggression
towards stranger males and females. This actually promotes the pair
bond with the mated partner.
Male aggression (mate guarding) and female aggression (monogamy
maintenance) depend on D1 receptor activity in the NAcc .
Pair-bonded males have greater density of D1 receptors compared to
bachelors. Blocking D1 receptor activity in the NAcc abolishes
aggressive behaviour toward strangers.
D1R activity seems to support pair bonding by promoting aggression
to everyone else, besides your mate!
62
Lecture 8 outline
63
The partner engram and the pair bond
Pair bonding forms when
Partner engram The partner engram becomes CSs
for social affiliation
(through the VP)
and become rewarding themselves
Pro-social outputs
Social recognition is crucial for pair bonding. Olfactory cues are critical for social recognition in many mammalian species (even
humans). Olfactory bulb and Anterior olfactory nucleus (AON) express OXTR, necessary for social recognition. Amygdala forms social
traces. Hippocampus forms a social engram for the partner.
In all species, dopamine reward signals reinforce mating.
In pair bonding species, the partner engram is activated at the same time as DA reward system. Partner engram becomes rewarding
and this leads to intense pro-social behaviour toward the partner.
Also, partner engram inputs to the ventral pallidum are activated at the same time as reward inputs. Associative plasticity strengthens that
connection and the partner engram comes to continuously activates the VP over long periods of time creating an enduring pair bond since
motor output from the VP is pro-social behaviours.
https://www-nature-com.myaccess.library.utoronto.ca/articles/s41583-018-0072-6/figures/2
The simple view

Cues of a conspecific Reward: DA release
Social approach + mating
Social affiliation
Partner engram Social Aggression to others
Dopamine Dopamine
+
OXT/AVP
The reward system is strongly activated in response to mating in monogamous and non-monogamous
species. In non-monogamous species mating does not lead to pair bonding while in monogamous
species mating facilitates pair-bonding (but is not necessary).
In monogamous species, pair bonding involves a specific interaction or overlap of the reward system
with the partner recognition system (operant conditioning). A partner (the social engram) comes to
evoke rewarding feelings (by classical conditioning) and evoke pro-social behaviour (through the VP) and
attraction over long period of time. Pair bonding may be thought of as a conditioned partner preference.
The “Red Car” partner engram?
66
Pair Bonding and Love - summary
We propose that pair bond formation is essentially the result of neural plasticity that links
the neural encoding of partner cues with the brain’s reward system such that a partner
becomes persistently and inherently reinforced, leading to selective affiliative behaviour.
We propose that positive reinforcement facilitates the formation of a pair bond, while
negative reinforcement plays a role in maintaining the bond.
What about humans?
We propose that pair bonding is the evolutionary antecedent of romantic love and that the
pair bond is an essential element of romantic love. The pair bond is the innate,
subconscious motivating force forging a strong attraction between partners. Romantic love
is the emergent property of the integration of the pair bond mechanisms described here…
Larry Young, 2018
You don't love someone for their looks, or their clothes, or for their fancy car, but because they sing a song only
you can hear. Oscar Wilde
https://www-nature-com.myaccess.library.utoronto.ca/articles/s41583-018-0072-6#Fig2 67
68

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HMB200 Lecture 8

Motivated Behaviours and

Guess where I’ve been…

The more the imbalance, the stronger the

Low imbalance = low motivational state

Neuroscientists discovered that there is

Recall: The Amygdala effector reflexive behaviour:

Orange structures are involved in habitual

Ventral Tegmental Area

Situation I was placed in a

Situation Behaviour Consequence

Situation Behaviour Consequence

I happen to walk I walk in and MMMMM

Next time your motivational state directs

Operant conditioning will increase

Situation Behaviour Consequence

THE SIGN!! I walk in and MMMMM

Operant conditioning will increase

LO dopamine stimulates behaviour

A very simplified view, that we will follow to learn some concepts:

In another pathway, dopamine seems to be important to learn about rewards

At first (never seen the sign before):

Seminal research that shaped our

Seminal research that shaped our

After a long time…

Dopamine responses in rat nucleus

Da signalling is actually more complicated

GUESS WHAT WAS SEEN WHEN YOU ZOOMED IN JUST AT THE

Reward Pair bonding

Pair bonding is a term used in biology and behavioural sciences

Make visible the participants panel. Slide this up

Step 4—Leader clicks CREATE NEW GROUP

Neuroscientists took this opportunity to discover the underlying neurobiological mechanisms

Partner preference test is designed to quantify selective social

But why only in prairie and not meadow voles?

In prairie voles In meadow voles,

In females, a key difference between monogamous and non-monogamous voles is the

In prairie voles: AVPR antagonist in ventral

Amazingly: Artificial expression of AVPR in

In males, a key difference between monogamous and non-monogamous voles is the

Situation Behaviour Consequence

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