NEUR0014: Neural Basis of Motivation and
Learning
Lecture 1: Introduction
History of neuroscience of emotion
• Emotion – psychological states that are related to stimuli (rewarding or aversive, important
for animals survival or ability to reproduce)
Charles Darwin – ‘The Expression of Emotions in Man and Animals’
• Similarities between human and animal emotional displays
• Theses: Emotions can be reduced to a simplified set of universal emotions (6) –happiness,
sadness, fear, anger, surprise and disgust – universal across all human cultures (different
contexts and manners of expressions)
o These emotions can also be seen in other species as well as human cultures
o A universality of emotions
James-Lange Theory of Emotion
William James & Karl Lang. James (1884) What is an emotion?
Thesis: Emotions are not causal states, motivating animals to do a type of behaviour, but rather the
result of brain feedback in response to stimulus.
• Emotions are the psychological state that arises from feedback to the brain from the body in
response to positive or negative stimulus Commented [IM1]: You don’t run from the lion because you’re
afraid, you’re afraid because you ran from the lion
o E.g. the emotion of fear is our experience of what happens to our body in response
to a frightening negative stimulus (e.g. heart racing, muscle tensing, sweating)
• Negative stimulus à sympathetic nervous system à brain responds to internal bodily
signals à fear
o Emotion is a cortical interpretation of the physiological state of the body
• Different emotions à different sets of bodily changes Commented [IM2]: You are afraid because your body is telling
you there is something to be afraid of
You don’t run from the lion because you’re afraid, you’re afraid because you run from the lion
Cannon’s criticisms of James-Lange Theory
1. Bodily changes too slow and non-specific (e.g. sympathetic/parasympathetic NS vs many
emotions)
2. Experimental work shows that if you remove the PNS (para/symp), they can still display
emotional behaviour
a. Bodily changes not required for emotional behaviour
3. Philip Bard – cat decortications experiments; remove neocortex from rest of brain (b) does
not abolish emotional responses
, a. In direct opposition to J-L Theory
b. E.g. Sham Rage in cats, emotion responses still there but to incorrect ex. stimuli
4. Cut down line a (cortex, thalamus AND hypothalamus) – Then no emotional responses are
shown
a. Animals are still alive and show some behavioural responses e.g. simple motor
patterns (grooming, feeding); supported by motor patterns stored in brain stem and
spinal cord
b. No significant emotional response to positive or negative stimulus (e.g. sexual
response, run from threat; important for reproduction and survival)
c. Key brain area is believed to be hypothalamus
Cannon-Bard Theory of Emotion
• Hypothalamus is key node in network – produces appropriate bodily response AND signals
to cerebral cortex
o CC interprets hypothalamus activity à feeling
• Similar to J-L, the feeling of the emotion is NOT causative; other brain areas produce the Commented [IM3]: Emotional responses are not caused by
emotional experiences
behavioural response:
o J-L – cerebral cortex
o C-B – hypothalamus
• Emotion is interpreted by the CC; related to the relevant behavioural response
• Unlike J-L, the feeling of emotion is not caused by CC interpretation of bodily response, but
CC interpretation of hypothalamus activity
• J-L believes CC to produce and interpret bodily response
o C-B believes hypothalamus to produce bodily response, with bodily response not Commented [IM4]: Overcomes Cannon’s first criticism of J-L
Theory
directly influencing the feeling of emotion
Lecture 1: The Historical Emotional Brain, Part 2
The Limbic System or ‘Papez Circuit’ was thought to underlie emotion
• James Papez (1937)
• Papez was the first to introduce the idea that the limbic structure could mediate emotions
o Rabies à damage to hippocampus à extreme rage Commented [IM5]: Now believed to be a false interpretation.
Rabies affected other brain regions.
o Hippocampus at centre of his theory
• Feeling of emotion produced via cingulated cortex (subset of neocortex)
• As in C-B, Hypothalamus theorised to produce bodily response to emotional stimulus,
before entering the Papez Circuit
Kluver-Bucy Syndrome
(1937) – Bilateral temporal lobectomy Rhesus monkeys displayed:
1. Visual defects
2. Oral tendencies
3. Changes in emotional behaviour (hypersexuality, hypo-emotionality)
, a. Inappropriate emotional response to neutral stimuli; dysregulation of emotional
behaviour
b. Due to damage to Amygdala Commented [IM6]: Introduces amygdala to Papez circuit/
emotional circuit of brain
MacLean (1949) – Defining the limbic system
Broca’s ‘lobe limbique’ + Papez’s emotional brain circuit à MacLean’s ‘emotional keyboard’ (1949)
• Stimulating key subregions of the hippocampus connect to specific ‘emotional circuits’
• Hippocampus is key organiser of limbic system
Generally believed to be incorrect; although the idea that different regions of the hippocampus have
different functions may hold merit (ventral/dorsal) [later lectures]
The Triune Brain (MacLean, 1970)
The brain can be classified into three primary systems:
1. Neo-mammalian brain - Neocortex (declarative, thinking)
2. Paleo-mammalian brain - Limbic system (affective, elaborating social emotions)
3. Reptilian brain - Basal Ganglia, Cortical nuclei, brainstem, mid-brain (innate behaviour, fixed
motor plans, mediating expressions of fear, aggression, sex etc.)
a. b/c animals can still produce basic motor commands with thalamus/hypothalamus
removed
b. Reptiles only have this brain – homologous CC
• With evolution, progressively more complex brain systems are added, resulting in more
complex behaviours
Limits of the limbic system concept
• Expanded to encompass many more brain regions than MacLean and Papez first suggested,
thus the concept has lost specificity
• MacLean’s evolutionary concept has been discredited; oversimplification. The brain is not
made up of different layers, some more primitive than others.
o Mammals have mammalian brain; reptilian brain is a reptilian brain
o Dentate gyrus (limbic system) has been discovered as a part of the hippocampus in
mammals but not birds and reptiles
§ New brain area that is not a part of the neocortex
• The limbic brain is not exclusively or specially connected to the hypothalamus; the
hypothalamus is connected to most of the brain
o Hypothalamus underlies emotion-motivated behaviour
• Some regions of the limbic systems appear to be involved in distinct psychological processes
rather than emotional processing; memory and spatial navigation (anterior thalamus,
mammillary bodies, hippocampus)
The limbic system (2006) – modern components
Heimer and Van Hoesen (2006)
, Basic anatomical components:
• Parahippocampal gyrus (neocortex)
• Olfactory cortex
• Hippocampus
• Cingulated gyrus
• Caudal orbital and medial prefrontal cortex
o Key for motivated behaviour
• Amygdaloid nuclear complex
• Antero-ventral insula
Closely associated areas:
• Basal forebrain nuclei
• Hypothalamus
• Nucleus accumbens and ventral pallidum (DA reward system)
Still key areas for emotional responses
Lecture 1: Modern neuroscience of the limbic system and the ‘Emotional Brain’, Part 3
Evidence for a memory role in:
• mammillary bodies (and anterior thalamus) – found in Korsakoff’s syndrome (alcohol-
associated mammillary body atrophy à memory loss)
• hippocampus (Bilateral temporal lobectomy of patient HM à anterograde amnesia
(Scoville and Milner, 1957))
o Hippocampus necessary to make long-term memories of experiences we undergo
Hippocampus acts as a navigational neural map. Hippocampal role in allocentric spatial cognition.
• Individual hippocampal neurons show functional tuning for spatial correlates for what the
animal is doing.
• Where the animal is (place cells)/ Where they are facing (head direction cells)/ Distance
estimation (grid cells).
• Roles in memory and space are intimately related; innate to mammalian navigation and
universal to link memories with navigation.
Inferior parietal cortex role in egocentric spatial cognition
Damage to the left posterior parietal cortex results in hemispatial neglect – a deficit in attention and
awareness of one side of space.
Interaction with hippocampus is important. All information coming into body is egocentrically
organised. When we want to make a behavioural response via motor system, this is also
egocentrically organised. Thus, there needs to be a translation system between the hippocampus
allocentric (universal) map and the egocentric reality. This is believed to occur in the parietal cortex.
Learning
Lecture 1: Introduction
History of neuroscience of emotion
• Emotion – psychological states that are related to stimuli (rewarding or aversive, important
for animals survival or ability to reproduce)
Charles Darwin – ‘The Expression of Emotions in Man and Animals’
• Similarities between human and animal emotional displays
• Theses: Emotions can be reduced to a simplified set of universal emotions (6) –happiness,
sadness, fear, anger, surprise and disgust – universal across all human cultures (different
contexts and manners of expressions)
o These emotions can also be seen in other species as well as human cultures
o A universality of emotions
James-Lange Theory of Emotion
William James & Karl Lang. James (1884) What is an emotion?
Thesis: Emotions are not causal states, motivating animals to do a type of behaviour, but rather the
result of brain feedback in response to stimulus.
• Emotions are the psychological state that arises from feedback to the brain from the body in
response to positive or negative stimulus Commented [IM1]: You don’t run from the lion because you’re
afraid, you’re afraid because you ran from the lion
o E.g. the emotion of fear is our experience of what happens to our body in response
to a frightening negative stimulus (e.g. heart racing, muscle tensing, sweating)
• Negative stimulus à sympathetic nervous system à brain responds to internal bodily
signals à fear
o Emotion is a cortical interpretation of the physiological state of the body
• Different emotions à different sets of bodily changes Commented [IM2]: You are afraid because your body is telling
you there is something to be afraid of
You don’t run from the lion because you’re afraid, you’re afraid because you run from the lion
Cannon’s criticisms of James-Lange Theory
1. Bodily changes too slow and non-specific (e.g. sympathetic/parasympathetic NS vs many
emotions)
2. Experimental work shows that if you remove the PNS (para/symp), they can still display
emotional behaviour
a. Bodily changes not required for emotional behaviour
3. Philip Bard – cat decortications experiments; remove neocortex from rest of brain (b) does
not abolish emotional responses
, a. In direct opposition to J-L Theory
b. E.g. Sham Rage in cats, emotion responses still there but to incorrect ex. stimuli
4. Cut down line a (cortex, thalamus AND hypothalamus) – Then no emotional responses are
shown
a. Animals are still alive and show some behavioural responses e.g. simple motor
patterns (grooming, feeding); supported by motor patterns stored in brain stem and
spinal cord
b. No significant emotional response to positive or negative stimulus (e.g. sexual
response, run from threat; important for reproduction and survival)
c. Key brain area is believed to be hypothalamus
Cannon-Bard Theory of Emotion
• Hypothalamus is key node in network – produces appropriate bodily response AND signals
to cerebral cortex
o CC interprets hypothalamus activity à feeling
• Similar to J-L, the feeling of the emotion is NOT causative; other brain areas produce the Commented [IM3]: Emotional responses are not caused by
emotional experiences
behavioural response:
o J-L – cerebral cortex
o C-B – hypothalamus
• Emotion is interpreted by the CC; related to the relevant behavioural response
• Unlike J-L, the feeling of emotion is not caused by CC interpretation of bodily response, but
CC interpretation of hypothalamus activity
• J-L believes CC to produce and interpret bodily response
o C-B believes hypothalamus to produce bodily response, with bodily response not Commented [IM4]: Overcomes Cannon’s first criticism of J-L
Theory
directly influencing the feeling of emotion
Lecture 1: The Historical Emotional Brain, Part 2
The Limbic System or ‘Papez Circuit’ was thought to underlie emotion
• James Papez (1937)
• Papez was the first to introduce the idea that the limbic structure could mediate emotions
o Rabies à damage to hippocampus à extreme rage Commented [IM5]: Now believed to be a false interpretation.
Rabies affected other brain regions.
o Hippocampus at centre of his theory
• Feeling of emotion produced via cingulated cortex (subset of neocortex)
• As in C-B, Hypothalamus theorised to produce bodily response to emotional stimulus,
before entering the Papez Circuit
Kluver-Bucy Syndrome
(1937) – Bilateral temporal lobectomy Rhesus monkeys displayed:
1. Visual defects
2. Oral tendencies
3. Changes in emotional behaviour (hypersexuality, hypo-emotionality)
, a. Inappropriate emotional response to neutral stimuli; dysregulation of emotional
behaviour
b. Due to damage to Amygdala Commented [IM6]: Introduces amygdala to Papez circuit/
emotional circuit of brain
MacLean (1949) – Defining the limbic system
Broca’s ‘lobe limbique’ + Papez’s emotional brain circuit à MacLean’s ‘emotional keyboard’ (1949)
• Stimulating key subregions of the hippocampus connect to specific ‘emotional circuits’
• Hippocampus is key organiser of limbic system
Generally believed to be incorrect; although the idea that different regions of the hippocampus have
different functions may hold merit (ventral/dorsal) [later lectures]
The Triune Brain (MacLean, 1970)
The brain can be classified into three primary systems:
1. Neo-mammalian brain - Neocortex (declarative, thinking)
2. Paleo-mammalian brain - Limbic system (affective, elaborating social emotions)
3. Reptilian brain - Basal Ganglia, Cortical nuclei, brainstem, mid-brain (innate behaviour, fixed
motor plans, mediating expressions of fear, aggression, sex etc.)
a. b/c animals can still produce basic motor commands with thalamus/hypothalamus
removed
b. Reptiles only have this brain – homologous CC
• With evolution, progressively more complex brain systems are added, resulting in more
complex behaviours
Limits of the limbic system concept
• Expanded to encompass many more brain regions than MacLean and Papez first suggested,
thus the concept has lost specificity
• MacLean’s evolutionary concept has been discredited; oversimplification. The brain is not
made up of different layers, some more primitive than others.
o Mammals have mammalian brain; reptilian brain is a reptilian brain
o Dentate gyrus (limbic system) has been discovered as a part of the hippocampus in
mammals but not birds and reptiles
§ New brain area that is not a part of the neocortex
• The limbic brain is not exclusively or specially connected to the hypothalamus; the
hypothalamus is connected to most of the brain
o Hypothalamus underlies emotion-motivated behaviour
• Some regions of the limbic systems appear to be involved in distinct psychological processes
rather than emotional processing; memory and spatial navigation (anterior thalamus,
mammillary bodies, hippocampus)
The limbic system (2006) – modern components
Heimer and Van Hoesen (2006)
, Basic anatomical components:
• Parahippocampal gyrus (neocortex)
• Olfactory cortex
• Hippocampus
• Cingulated gyrus
• Caudal orbital and medial prefrontal cortex
o Key for motivated behaviour
• Amygdaloid nuclear complex
• Antero-ventral insula
Closely associated areas:
• Basal forebrain nuclei
• Hypothalamus
• Nucleus accumbens and ventral pallidum (DA reward system)
Still key areas for emotional responses
Lecture 1: Modern neuroscience of the limbic system and the ‘Emotional Brain’, Part 3
Evidence for a memory role in:
• mammillary bodies (and anterior thalamus) – found in Korsakoff’s syndrome (alcohol-
associated mammillary body atrophy à memory loss)
• hippocampus (Bilateral temporal lobectomy of patient HM à anterograde amnesia
(Scoville and Milner, 1957))
o Hippocampus necessary to make long-term memories of experiences we undergo
Hippocampus acts as a navigational neural map. Hippocampal role in allocentric spatial cognition.
• Individual hippocampal neurons show functional tuning for spatial correlates for what the
animal is doing.
• Where the animal is (place cells)/ Where they are facing (head direction cells)/ Distance
estimation (grid cells).
• Roles in memory and space are intimately related; innate to mammalian navigation and
universal to link memories with navigation.
Inferior parietal cortex role in egocentric spatial cognition
Damage to the left posterior parietal cortex results in hemispatial neglect – a deficit in attention and
awareness of one side of space.
Interaction with hippocampus is important. All information coming into body is egocentrically
organised. When we want to make a behavioural response via motor system, this is also
egocentrically organised. Thus, there needs to be a translation system between the hippocampus
allocentric (universal) map and the egocentric reality. This is believed to occur in the parietal cortex.
