Lecture 4

Question 1

Paradox of emotion

Answer 1

On the one hand, they seem self-evident and obvious when examined introspectively; on the other hand, they have been extremely difficult to define in objective scientific terms.

Question 2

Conventional view of emotion

Answer 2

Emotions incorporate many components that need to be coordinated and often synchronized for the experience of emotion. Cognitive, behavioral, and somatic responses are all part of the emotion. The subjective feelings that rise from the emotional experience allow for verbal report in humans.

Question 3

Anderson and Adolphs view on emotion

Answer 3

They believe that the observed behavior, subjective reports, psychophysiology, cognitive changes, and somatic responses that were thought to be part of the emotion are actually only associated with the central emotional state. They disagree with the conventional view, and argue that emotions involve all the same components but these are not part of the emotion, rather they are caused by the emotion state.

Question 4

Darwin’s view on emotion

Answer 4

Charles Darwin considered emotional expression from a functional and evolutionary standpoint. He asserted that emotion homologous (with similar function) to our own is easily recognizable in humans and other animals and can even be observed in insects. Darwin did not provide objective criteria for identifying emotion.

Question 5

Anderson and Adolphs vs. Darwin

Answer 5

Anderson & Adolphs agree with Darwin: even invertebrates have primitive emotion states, but they differ from Darwin in that they argue that these states are not necessarily homologous to specific categories that have been used to define human emotions (e.g. fear, anger, happiness). They assert that emotion states in all animals share certain fundamental properties: ‘emotion primitives’. Emotion primitives are the evolutionary building blocks of emotion that are shared across species. Species-typical behaviours that arise from these emotion primitive are not necessarily shared.

Question 6

Animals and emotions

Answer 6

The key question is not if animals share a specific emotion (e.g. fear) but if animals have central states that share features of emotions in general. We can investigate general features of emotion in animal models without using anthropocentric labels like ‘fear’, ’anger’, or ‘sadness’. For this to work, we need operational criteria for emotion across species. The goal of Adolphs and Anderson is to propose a way of thinking about emotion.

Question 7

4 points in studying emotions

Answer 7

1) The causal relationship between emotions and observable behaviour
2) The relationship between emotion states and subjective feelings
3) The characteristic features of emotion states shared by specific emotions
4) If there are uniquely human features of emotion

Question 8

Causal relationship between emotions and behaviour

Answer 8

Is it behavior that causes the emotion or the emotion causing the behavior? There is not yet universal agreement about the direction of
causality between emotion and behaviour. Part of this disagreement comes from lack of data.
Purely observational/correlational approaches studying the link between emotion and behavior cannot test the directionality of causality. Studying the neural basis of emotion states in model animals allows us to directly and rigorously test causality by manipulating neural activity with modern neuroscience tools.

Question 9

Behaviorist view on the relationship between central emotion states and subjective feelings

Answer 9

Emotional stimuli directly evoke behaviour in animals. In humans, the conscious awareness of behavioural and somatic responses evoked by the emotional stimulus gives rise to subjective feelings.

Question 10

Anderson & Adolphs view on the relationship between central emotion states and subjective feelings

Answer 10

Emotional stimuli evoke central emotion states. In animals and humans these states give rise to behavioural and other responses, including subjective feelings (in humans). This allows to study emotions in animals. Subjective feelings may happen in animals, but they can’t be studied.

Question 11

Animals and feelings

Answer 11

Feelings can only be assessed by verbal report so we can only study them in humans. If we equate emotions with subjective feelings, we cannot study emotion in animals. There is no reason a priori to conclude that animals do not have central emotion states. There is a connection between ability to communicate and what we assume that animals can feel. Is it that we think they don’t feel just because they can’t express it?

Question 12

Koko the gorilla - do animals have subjective feelings?

Answer 12

Koko was a gorilla that was taught sign language (it was thought that gorillas would be capable of language, but not spoken one because of physiological and anatomical restrictions). She was able to understand around 2000 english words. One day, she asked for a kitten. Unfortunately, it died, which caused Koko to exhibit signs of sadness. She whimpered with grief and “discussed” the death of her pet kitten for several days after getting the bad news.

Question 13

How to study emotions in animals?

Answer 13

Affective neuroscience should look for evidence in animals of central emotion states with certain fundamental properties that are causal in responding to certain stimuli with specific behaviours and the corresponding neural mechanisms.

Question 14

Building blocks vs. features of emotions

Answer 14

Building blocks:
- essential, basic properties of emotion
- shared by all or most specific emotions
- present in precursors to emotion states in simpler organisms
Features:
- more elaborated and variable properties of emotions
- not shared by all emotions

Question 15

Building blocks vs. features example

Answer 15

For example, in a car, wheels are building blocks and air conditioning is a feature (only some cars have AC, but all cars have wheels)
* Emotion building block: valence. All emotion states share an intrinsic quality of evaluating good or bad, pleasant or unpleasant, approach or avoid.
* Emotion feature: social communication. It’s very prominent in mammals but likely recently evolved and not present in all animals

Question 16

A provisional list of emotion properties

Answer 16

The division of building blocks and features is not black and white. It is instructive, not absolute. Emotion properties are the processing features that define emotion states (i.e. the things that we look for in the brain to discover an emotion states). We can put together a provisional* list of operating characteristics of emotion states that are essential to carrying out the functional role of emotion to begin to illustrate how we can investigate emotions in general.
*(i.e. not complete, there could be others, some could be removed, this is not ‘truth’)

Question 17

Emotion properties list

Answer 17

• Scalability
• Valence
• Persistence
• Generalization
• Global coordination
• Automaticity
• Social communication
  The higher on the list (ex. scalability), the more the property acts like a building block, and the lower on the list (ex. communication), the more it acts as a feature.

Question 18

Scalability

Answer 18

An emotion state can scale in intensity. Importantly, parametric scaling can result in
discontinuous behaviours, such as the transition from hiding to fleeing during the approach of a predator. Intensity is often conceptualized as arousal, although these two are not the same thing.Scalability differentiates from stimulus-response reflexes which tend to be all or nothing responses.

Question 19

Valence

Answer 19

Valence is thought by many psychological theories to be a necessary feature of emotion experience (or ‘affect). It corresponds to the psychological dimensions of pleasantness/unpleasantness, or the stimulus-response dimension of appetitive vs. aversive (but again, these two are not the same thing).

Question 20

Persistence

Answer 20

An emotion state outlasts its eliciting stimulus, unlike reflexes, and so can integrate information over time, and can influence cognition and behaviour for some time. Different emotions have different persistence (ex. sadness lasts longer than surprise). Emotions typically persist for seconds to minutes. For example, fear has a long-lasting effect on behavior: heart rate, stress hormone levels, breathing rate etc. remain elevated for some time after encountering a threat.

Question 21

Generalization

Answer 21

Emotions can generalize over stimuli and behaviour, much of which depends on learning. This creates something like a “fan-in”/ “fan-out” architecture: many different stimuli link to one emotion state, which in turn causes many different behaviours, depending on context. Persistence and generalization underlie the flexibility of emotion states.

Question 22

Global coordination

Answer 22

Related to the property of generalization is the broader feature that emotion states orchestrate a very dense causal web of effects in the body and the brain: they engage the whole organism. In this respect, they are once again differentiated from reflexes.

Question 23

Automaticity

Answer 23

Emotions have greater priority over behavioural control than does volitional deliberation, and it requires effort to regulate them) a property that appears disproportionate, or even unique, in humans). Still, emotions are not as direct as reflexes.

Question 24

Social communication

Answer 24

In good part as a consequence of their priority over behavioural control, emotion states are pre-adapted to serve as social communicative signals. They can function as honest signals that predict another animal’s behavior, a property taken advantage of not only by conspecifics, but also predators and prey.

Question 25

Why make a list of emotion properties?

Answer 25

We can use these properties to differentiate emotion states from reflexes. We can also use these properties to characterize a specific emotion state and differentiate it from another emotion state. We can think of emotions in dimensional space.

Question 26

A dimensional space for emotions

Answer 26

‘Core affect’ is commonly represented in 2 dimensional space with axes defined by arousal and valence (e.g. Russel, 1980). We can locate any emotion within this 2D space . The proximity/distance between two emotions is interpreted as an indication of similarity/distance. We can think of this as a ‘similarity structure’ for emotions. This 2D representation is common to many
psychological theories. The two dimensions of valence and arousal correspond well to human ratings of emotions. This dimensional representation is based on human subjective emotional experience rather than behavioural or neural data.

Question 27

Edmund Rolls dimensions of emotions

Answer 27

Other theories propose different dimensions e.g. Edmund Rolls proposed that emotions can be defined as states elicited by administration or withholding of reward or punishment. It is based on behavioral responses (to obtain a reward or avoid smth negative).

Question 28

Problems with dimensional representations of emotions

Answer 28

In both models, althought the axes are different, the dimensionality remains low. For example, fear is a high-arousal, negative-valence state or a state caused by administration of a negative reinforcer (i.e. the anticipation of something bad). Any two dimensions are unlikely to be sufficient to capture all the variance in emotions. Specific emotions have been given labels (e.g. anger, fear, disgust) based on English words for emotion categories and located in 2D space. A dimensional approach could also be used to classify emotions without needing to classify under specific labels.

Question 29

3D representation of emotions

Answer 29

Emotion states could be categorized based on their location within multidimensional space, for example intensity, valence, and persistence (but more can be added). We can then observe how emotion states associated with similar or different behaviours cluster (or don’t) in this space. It is still not a solution, but it’s intended to encourage thinking about that subject (moving away from our dependence on language).

Question 30

Intensity of emotion (scalability pt.2)

Answer 30

It is not yet clear if the intensity of emotion is inherent to the mechanism of a specific emotion or if there might exist some kind of general arousal system for emotion. Differences in intensity could be graded or qualitatitive. A graded increase in intensity could be observed by increasing vigor of the same behaviour e.g running from threat. Gradations in emotional intensity can also have non-linear effects on behaviour. For example, threat imminence continuum of defensive behaviour in rodents and octopi. These behaviors are all related to the same emotion, but vary in intensity.

Question 31

Antithesis (valence pt.2)

Answer 31

Darwin believed in emotional ‘antithesis’: Emotions come in pairs of opposites which are expressed by physical opposite and complementary behaviours. This could be important for social communication functions. Emotions exist on this valence spectrum, from intensely positive to negative (described by valence). In truth, opposite behaviors could simply be a coincidence, not a proof of fundamental antithesis.

Question 32

Why are emotions persistent?

Answer 32

Persistence makes emotion states flexible. A persistent emotion state allows for integration of different kinds of sensory information over time which may be important for neural computation and action selection. Persistence also allows emotion states to interact with other other internal states and powerfully influence cognition and behavior.

Question 33

Emotion vs. mood

Answer 33

Emotions generally do not persist for long
after the situation that triggered the emotion has been resolved. Moods are emotion-like states that last much longer than emotions (hours-years). Moods may be more prominent in humans than animals. Emotions function to cope with present, acute situations; while moods may function to cope with events in the past or future. They’re probably not just emotions on a longer time scale. Moods often have no clear trigger. They often involve effects on cognition more than on behaviour. Moods are similar to emotions in having dimensions of persistence, scalability, valence, generalization and automaticity, but they’re distinct from emotions in not serving a clear social communication function.

Question 34

Persistence vs. memory

Answer 34

Persistence seems to be independent from memory and consciousness. Amnesic patients still experience persistent sadness after watching a sad film, even though they don’t remember having seen the film.

Question 35

Context generalization (trans-situationality)

Answer 35

Because of the property of persistence, an emotion state induced by one stimulus can generalize to a different context and influence responding to different stimuli. Context generalization allows emotions to bias cognition and behaviour. Applying this criterion can distinguish between a behavioural response mediated by a simple reflex and by a persistent internal state that generalizes to other context to influence subsequent behaviour.

Question 36

Generalization and honey bees

Answer 36

Stressed honeybees show a negative bias in a test of choosing between ambiguous odor cues. This suggests that the stress manipulation induced an internal state in honeybees that influenced their behaviour in other contexts.

Question 37

Stimulus generalization and pleiotropy

Answer 37

Stimulus generalization & pleiotropy contribute to other aspects of generalizability of emotion states. Many stimuli ‘fan-in’ (stimulus generalization) to cause an emotion state which can then ‘fan-out’ (pleiotropy) to cause many effects. The same behavioral expression can be triggered by many different stimuli, including those for which the behavior appears to serve no useful purpose, if those stimuli evoke the same internal emotion state. Emotion states are pleiotropic (they have multiple, parallel effects on behaviour, body, cognition). Simple reflex responses generally don’t induce multidimensional responses.

Question 38

Darwin on generalization

Answer 38

Darwin’s example : cats kneading their paws on a blanket. This behaviour serves to stimulate milk flow from a nursing mother but serves no purpose in the blanket example. Darwin argues that it became associated with the same state
(“pleasure”) either through habit (learning) or “inheritance”.

Question 39

Emotions and learning

Answer 39

Stimulus generalization is closely linked to learning. Most stimuli that cause emotions
gain this property through experience (i.e. associative emotional learning). The best understood example of this is Pavlovian conditioning: through presentation with an electric shock, a previously neutral cue comes to elicit the same behavioural response (freezing) as the shock itself. All species studied show associative emotional learning. Some species also show learning through observation without the need for direct experience (ex. humans can learn from being told about things). Learning is a key mechanism that increases stimulus generalization.

Question 40

Fear pleiotropy

Answer 40

Fear induces defensive behaviours (freezing,
fleeing) as well as endocrine changes (stress hormones release), autonomic changes (heart rate, blood pressure, sweating) and cognitive changes (attention & memory)

Question 41

Emotions and learning - domain specificity

Answer 41

There is a restricted range of stimuli or circumstances that can cause an emotion state and some stimuli can be more readily learned about than others. For example, tastes readily come to elicit disgust but is much less likely that a tone will come to elicit disgust. Domain specificity distinguishes emotions from volitional control. This also shows the limits of associative learning. For example, it’s easier to believe that snakes are scary rather than a flower (we learn snake-shock association faster than flower-shock in exmperiments).

Question 42

Uses of associative learning

Answer 42

Associative learning can be used to test if a stimulus induces an emotion state. For example, conditioned place preference pairs a neutral stimulus (one half of a box) with something potentially rewarding or aversive (e.g. an injection of a drug). In a later test session, where the animal spends time is interpreted an index of if the stimulus induced a rewarding or aversive internal state.