Animal Behaviour - Emotions

Oatley and Jenkins 1996 - “An emotion is usually caused by a person consciously or unconsciously evaluating an event as relevant to a concern (goal) that is important; the emotion is felt as positive when the concern is advanced and negative when the concern is impeded.”

Ortony et al., 1988 - “Emotion may be understood as the outcome of an evaluation of the extent to which ones goals are being met.”

Rolls (2005) - Emotions are states elicited by rewards and punishers

Paul et al., 2005 - Emotions are processes concerned with achieving goals (seeking valuable resources/rewards or avoiding harm/punishment)

"See Bear" -> "Feel Fear" -> "Run"

William James (1900s) and Carl Lange proposed "See Bear" -> Pounding heart, sweaty palms, run away -> "Feel fear"

ie. That we feel emotions based on peripheral changes in the body

Emotions are felt due to sensing peripheral changes in our own body

Antonio Demasio (Modern Theorist) Agrees with this view.

Walter Cannon (1900s)

major body changes appear similar for a range of reported emotions

disruption of feedback from the periphery did not abolish emotional responses

subjective emotions appear to happen faster than visceral changes suggested to cause them.

humans injected with adrenaline do not report feeling a particular emotion

1962 - cognitive interpretation of peripheral body changes required to determine which emotion is experienced

Subjects injected with adrenaline, but half told it was just a vitamin. Both exposed to happy/playful confederate, half exposed to angry/rude confederate. Asked emotions afterwards - those who thought the drug was a vitamin attributed the feelings to emotion and said they felt happy/angry, those who knew the effects were due to the drug said they felt minimal emotion (cognitive dampening down of emotion)

Stimulus -> Central appraisal of stimulus (may NOT be conscious) -> Emotion

Detection of ones own physiological state eg. heart beat. People who are good at detecting their own heartbeats are more emotionally expressive and have more intense emotions

Subjective, Behavioural, Physiological, Cognitive

Something an animal will work for or learn an operant response to obtain.

Something an animal will work to escape or avoid

Unlearned reinforcers or punishers eg. predators, nice tastes, pain. Specified by genetics.

Learnt in association with a primary reinforcer - eg. sound/light

A sate one is in when working to gain a specific reward or avoid a specific punisher

Presentation of a reward - Pleasure.. ecstasy

Omission or termination of a reward - Sadness.. Rage

Presentation of a punisher - fear.. terror

Omission or termination of a punisher - relief

Orbitofrontal cortex and Amygdala allow learning of associations between primary and secondary reinforcers, tagged with reward information

Areas that control autonomic function eg. hypothalamus

Areas that control learning and behavioural responses eg. cingulate cortex and basal ganglia

Sensory processing areas like thalamus, primary taste cortex - no emotion associated

eg. seeing a dominant conspecific. Coded to allow recognition of WHAT they are but without any reward information at sensory processing structures eg. inferior temporal visual cortex

Enable a system in which survival goals (rewards and punishers) are coded by genes, and behaviour is guided by emotional states

Prepare the body for action by changes in physiological state

motivate specific action (eg. running away )

may influence cognitive function

Emotions - short term states elicited by stimuli

Moods - long term emotional or affective states that persist in the absence of any stimuli

Sensations - feed directly from sensory apparatus to the CNS

Emotions - no sensory apparatus, involve interpretation of stimuli

Primary areas will keep firing in the presence of a stimulus - you are eating cake, cake tastes like x

Secondary areas firing will drop off as value added by stimulus decreases - cake doesn’t taste good anymore

Arousal and Valence

High Arousal High Valence = Excitement

High Arousal Low Valence = Fear

Low Arousal Low Valence = Sadness

Low Arousal High Valence = Contentment

Gold- Standard self report

Rolls, Macphail

Panksepp, Baars

Physiological, behavioural, cognitive, subjective

| Subjective most difficult as requires self report

Yes - can be trained to work to access a reward, therefore it does feel emotion when trying to obtain the reward and when denied it

Emotional states will be elicited if animals can work (learn arbitrary operant responses to obtain rewards/avoid punishers)

The neural apparatus underpinning emotional processes in primates is present in majority of other mammalian species at least

primitive emotional circuitry eg. fear, rage ,panic exist in many species - But that does not necessarily mean they have a subjective component

Brain states and bodily responses are fundamental facets of emotion and occur in other species

All cultures shows these

Happy

Sad

Fear

Anger

Suprise

Disgust

Play

Lust

Care

Fear

Rage

Panic

Seeking

Fear

Emotional state induced by the perception of actual danger

Motivation of appropriate defensive/avoidance reactions in a FLEXIBLE manner

Animals placed in contexts likely to induce a fear state, outputs recorded eg. spontaneous behaviour, responses to tests, physiological measures

The definition of a fear inducing stimulus is highly anthropomorphic

Species and situation specific

Freezing

Avoidance/hiding/escape eg. birds may mob a predator through fear

Defensive posture

Vocalisation eg. may only occur in presence of conspecifics (Audience effect)

Defeacation

When behaviour only occurs in the presence of conspecifics

Staying in enclosed areas

| Tonic immobility

Threat -> Thalamus -> Sensory Cortex/straight to amygdala -> Amygdala -> Nucleus Reticularis/Pontis Caudalis -> Potentiated startle response via spinal motor neurones

Emotional valence - reward state attenuates startle, fear state potentiates it

Against other tests of fear, use of anxiolytic drugs or other drugs known to elicit a particular emotional state in humans.

Behaviour difficult to interpret

Different tests disagree on their assessment of fearfulness

Different species respond differently

Fear (OR AROUSAL!) -> Stress ->

Activation of SAM (Sympathetic-adrenomedullary, ^BP, sweating, ^HR, vHR variability, ^ circulating catecholamines)

HPA (Hypothalamaic-Pituitary-Adrenal, elevated circulating ACTH and Cortisol

^PRL, ^b endorphin, ^central vasopressin and CRF

Sexual arousal/pleasure

Context

Spontaneous behaviour

Behavioural tests

Physiological measures

An emotional state induced by the perception of POTENTIAL danger.

similar adaptive value to fear (defines/avoidance) but is a longer lasting state of non-specific preparedness

outputs similar to fear, may be more vigilance rather than active escape

inputs differ - actual v potential danger

Central opioid activity

| Spontaneous behaviour - avoidance, bill wiping, facial expression looked at by Berridge and Robinson 2003

When behaviour is thwarted.

When attacked/challenged/thwarted by another individual

Redirected behaviour/stereotypy eg. feeding

Aggression

Sped up behaviour

Vocalisation

Eye white display

Sandem et al 2002, 2006

Thwarting or social confrontation leading to spontaneous anger behaviours

SAM and HPA axis

Elevated testosterone

Elevated central vasopressin

Diminshed central serotinergic activity

Decrease arousal (anxiolytic)

Approach, play, exploration, affiliative behaviour, vocalisation (eg. panksepps tickling rats)

Panksepp and Burgdorf 2000 - tickling rats

preference tests

place preference

attenuated startle

HPA activation - excitement or sexual pleasure

Central opioid - liking something

Dopamine - wanting something

Central oxytocin - social affiliation/pair-bonding

Omission, loss or threat of a rewarding stimulus

Attention to dead or dying conspecifics

Douglas-Hamilton et al 2006 - elephants

Specific emotional states

Tests involve exposing animals to situations researches presume to induce specific discrete emotions

Mostly negative states studied

Lacks an over arching framework or structure of emotions to integrate massive range of emotions and predict other states

Dimensional model of subjective emotion - arousal on y axis, valence on x.

Avoid false alarm behaviour if living in a low threat environment where stimuli are not likely to actually predict threats

Harding, Paul and Mendl 2004/2005

Rats trained OPERANTLY on a positive stimulus and negative stimulus.. Shown an ambiguous stimulus, see which response they perform.

Happy rats should judge ambiguous stimulus as positive stimulus

Rats housed in predictable/unpredictable housing.

Rats housed in unpredictable conditions more likely to treat probe as negative - pessimistic

Burman et al 2008 - placed pots with food on right or empty on left

When placed in middle did they expect to see food? Judged as latency to approach.

Mendl et al 2010, dogs with separation anxiety judge ambiguous locations negatively

Bateson et al 2007

| Enriched v unenriched starlings judging ambiguous lid colours

Incorporating discrete and dimensional approaches to emotion - predicts how changes in decision making (cognitive bias) reflect emotional valence

Cognitive bias can detect valence of emotion and possible arousal? (Not yet tested)

May reveal phenotypes vulnerable to emotional disorder

Do you have to have conscious reflection of emotional state to show cognitive bias?

Physiological, behavioural, cognitive, subjective