Definitions of emotions typically describe proximate aspects such as Physiology

1. Nesse & Ellsworth (2009) Evolution, Emotions & Mental Disorders American Psychologist 64: 129-139 • Definitions of emotions typically describe proximate aspects such as • Physiology • Subjective experience • Facial expression • An evolutionary approach defines what emotions are in terms of how they came to exist. • Emotions are modes of functioning, shaped by natural selection, that coordinate physiological, cognitive, motivational, behavioral and subjective responses in patterns that increase the ability to meet the adaptive challenges of situations that have recurred over evolutionary time (Nesse, 1990).

2. Nesse & Ellsworth (2009) Evolution, Emotions & Mental Disorders American Psychologist 64: 129-139 but only in certain situations. • They are adaptations that are useful • Like pain and sweating, they remain latent until an evolved mechanism detects cues associated with the situations in which they are advantageous. • Unlike simpler adaptations, however, emotions are not unimodal responses to specific situations, like sweating in response to overheating. Instead, emotions adjust multiple component processes to create an organized response to the adaptive challenges of a given situation. • Example: appraisals that indicate a nearby predator arouse an emergency response that adjusts and coordinates many aspects of physiology, cognition and behavior.

3. FIGHT OR FLIGHT OVERVIEW AMYGDALA HYPOTHALAMUS PITUITARY GLAND ADRENAL GLAND THROUGHOUT BODY

4. Nesse & Ellsworth (2009) Evolution, Emotions & Mental Disorders American Psychologist 64: 129-139 • Questions about an emotional trait: • What trait is like, how it works; • How it develops over the individual’s lifetime; • How it developed over time in the history of the species; • What evolutionary factors shaped the trait. • i.e., Tinbergen’s Four Questions

5. Nesse & Ellsworth (2009) Evolution, Emotions & Mental Disorders American Psychologist 64: 129-139 • How many emotions exist, and what are they? • Some theories: just two basic states – positive and negative • Other theories: small set of basic emotions • Still other theories: potentially infinite number • Agreement: valence is a necessary quality of emotion: pleasure & pain, approach & avoidance fundamental • Many models array the emotions on these dimensions, e.g., the circumplex model (Posner et al 2005)

6. Nesse & Ellsworth (2009) Evolution, Emotions & Mental Disorders American Psychologist 64: 129-139

7. Nesse & Ellsworth (2009) Evolution, Emotions & Mental Disorders American Psychologist 64: 129-139 • A phylogeny of emotions: specific emotions partially differentiated from more primal generic states because they improved ability to cope with specific kinds of threats and opportunities. • New emotional variations are useless unless they are expressed in situations where they are advantageous. • Emotions do not have clear boundaries. No clear taxonomy of emotions possible.

8. Nesse & Ellsworth (2009) Evolution, Emotions & Mental Disorders American Psychologist 64: 129-139

9. Animal Emotion

10. Inside the Animal Mind: Do Animals have Emotions? Jane Goodall versus Marian Dawkins Goodall just asserts that animals have emotions – its obvious! – and says that emotional similarity is most likely in animals closely related to us (like chimps).

11. Inside the Animal Mind: Do Animals have Emotions? Jane Goodall versus Marian Dawkins Dawkins emphasizes that emotions are likely to be different in different species – and so we should not anthropomorphize. But she also says that we should give animals the benefit of the doubt and assume they do have emotions unless we somehow get evidence otherwise. ☞Issue important because of concerns about animal welfare. General point: absence of evidence ≠ evidence of absence

12. Animal Emotion • Dawkins: Facial expressions are not a reliable signal of emotions in most other species. • Hard for us to see this because we expect animals to express emotions in the same way as we do. • Not expressing emotions for communication purposes is not the same thing as not having emotions.

13. Communicating Emotion – Homologies? 1. attention2. excitement3. narrow smile4. broad inviting grin5. laughing6. crying

14. Continuity of facial expressions in primates? From Jolly 1972

15. Continuity of facial expressions in primates? From Jolly 1972

16. Central Issues • Do we need to posit emotion to explain response (e.g., ‘fear’ to explain response to danger)? • What actually is the question we’re asking? • Do animals have emotions? • Do they have emotions similar to ours? • What does it take to answer these questions?

17. What is Emotion? Do we need “fear” to avoid the predator? Is “fear” just our perception of our internal physiological changes accompanying the innate flight or fight response? James-Lange theory

18. James-Lange Theory of Emotion “My theory ... is that the bodily changes follow directly the perception of the exciting fact, and that our feeling of the same changes as they occur is the emotion. Common sense says, we lose our fortune, are sorry and weep; we meet a bear, are frightened and run; we are insulted by a rival, are angry and strike. The hypothesis here to be defended says that this order of sequence is incorrect ... and that the more rational statement is that we feel sorry because we cry, angry because we strike, afraid because we tremble ... Without the bodily states following on the perception, the latter would be purely cognitive in form, pale, colorless, destitute of emotional warmth.” William James

19. Animal Emotions: Exploring Passionate Natures Marc Bekoff

20. Animal Emotions: Exploring Passionate Natures Marc Bekoff Marc & Bessie, a rescued dairy cow

23. Animal Emotions: Exploring Passionate Natures Marc Bekoff “Emotions can be broadly defined as psychological phenomena that help in behavioral management and control”. Well that’s a nice definition, but how exactly do the emotions do this? Sometimes they seem disruptive, not particularly helpful. For example, Spock→ seems to do better without emotions!

24. Animal Emotions: Exploring Passionate Natures Marc Bekoff The study of play has always been an occasion to talk about animal emotions, perhaps because it seems to be an activity engaged in just for the fun of it. (Though contemporary thinking emphasizes its role in the development of necessary adult skills that may be used in hunting, fighting, etc.)

25. Animal Emotions: Exploring Passionate Natures Marc Bekoff Sadness & Grief Many of examples have accumulated showing apparently sad or grieving animals after loss of child or parent or mate, e.g., Goodall’s story of the chimp Flint’s (terminal) depression following the death of his mother Flo. video

26. Animal Emotions: Exploring Passionate Natures Marc Bekoff Sharing the burden of proof “In the future, skeptics should be required to mount serious defense of their position and share the burden of proof with those who accept that many animals do indeed experience myriad emotions.No longer will it be acceptable to claim that ‘yes, chimpanzees or ravens seem to love one another’ or that ‘elephants seem to field grief’’ and then present innumerable reasons – ‘we can never really know that animals feel emotions’ – why this cannot be so. Explanations about the existence of animal emotions often have as good a foundation as many other explanations that we readily accept (e.g., [some] claims about evolution…) Developing comparative evidence Taking off from the maxim “absence of evidence is not evidence of absence”, I would agree that the burden of proof is equally on skeptics (animals do not have emotions) and true believers (animals have human-like emotions). But the key will be to develop methods that can convincingly reveal the nature of emotional states (similarities and differences) in different animals. video

27. Animal Emotions: Exploring Passionate Natures Marc Bekoff Much of Bekoff’s argument seems to rely on a combination of an appeal to authority (Jane Goodall, Joyce Poole, et al) and compelling anecdotes or commentaries involving charismatic animals (e.g., chimps, elephants). e.g.:

28. Hormonal and Neural Mechanisms • Do animals fall in love? • Pleasure centers in the brain • Neural basis of the “cute response” (Lorenz)

29. Do Animals Fall in Love? • Peacock: no • Prairie vole: yes – life-long pair bonds • Oxytocin (& vasopressin) • “Their chemistry may be, well, just chemistry” • Oxytocin involved in parent-child bond too (along with prolactin)

30. Pleasure Centers of the Brain James Olds and Peter Milner in the 1950s probing the limbic systems of rats looking for pain centers, found instead “pleasure centers”: rats would respond at high rates for stimulation of their septal areas, located near the brain stem and among the oldest areas of the brain. Rats would cross electrified floors to press a self-stimulation switch, and would press it 1000s of times to the exclusion of food or water. Female rats would even abandon their unweaned pups to self-stimulate. Key centers were medial forebrain bundle (MFB) and ventral tegmentum (VT)Dopamine appears to be the most important neurotransmitter involved.

31. Pleasure Centers of the Brain Right here!

32. Pleasure Centers of the Brain Only a few experiments have been conducted involving the electrical stimulation of human pleasure centers. Generally these investigations are considered taboo. In the 1970s, Robert Heath, who believed he could "cure" homosexuality, wired up gay volunteers to an electrical apparatus that directly stimulated their nucleus accumbens, producing feelings of extreme pleasure. Given the choice, one man, code-named B-19, electrically self-stimulated his reward circuitry some 1,500 times in a 3-hour session (“he had to be disconnected, despite his vigorous protests”). Few experiments directly stimulating the human pleasure centers have been conducted since.

33. Baby schema modulates the brain reward system in nulliparous women Glocker et al PNAS 2009 Ethologist Konrad Lorenz defined the baby schema as a set of infantile physical features, such as round face, high forehead and big eyes, that is perceived as cute and motivates caretaking behavior in animals including humans, with the evolutionary function of enhancing offspring survival. “Using functional magnetic resonance imaging and controlled manipulation of the baby schema in infant faces, we found that the baby schema activates the nucleus accumbens, a key structure of the mesocorticolimbic system mediating reward processing and appetitive motivation, in nulliparous women. Our findings suggest that engagement of the mesocorticolimbic system is the neurophysiologic mechanism by which baby schema promotes human caregiving, regardless of kinship.”

34. Baby schema modulates the brain reward system in nulliparous women Glocker et al PNAS 2009

35. Baby schema modulates the brain reward system in nulliparous women Glocker et al PNAS 2009