Animal Emotion – a Potpourri

1. Animal Emotion – a Potpourri • More on cow eye white! • Animal welfare and Mark Bekoff • Neurobiological perspectives • Love, lust – hormonal mechanisms • Do animals fall in love? • Pleasure centers in the brain • Baby faces

2. (2002) Eye white may indicate emotional state on a frustration–contentedness axis in dairy cows. (2004) A short note on effects of exposure to a novel stimulus (umbrella) on behaviour and percentage of eye-white in cows (2005) Effects of cow–calf separation on visible eye white and behaviour in dairy cows. (2006) The use of diazepam as a pharmacological validation of eye white as an indicator of emotional state in dairy cows (2006) Behaviour and percentage eye-white in cows waiting to be fed concentrate The bottom cow could see and smell grass, but not eat it

3. Cows and Umbrellas Fig. 1. Change in percentage of white in the eye over time as the cows were exposed to a suddenly opened umbrella. Each bar gives the (mean ± S.E.) for a 30 s observation period (N = 16). Before the test person entered (before); test person with closed umbrella in front of the cow (unopened); immediately after opening of the umbrella (opened1) and 30–60 s after opening of the umbrella (opened2).

4. Cows separated from calves Fig. 1. The calculated mean percentages of visible eye white (±SE) of 11 cows during separation and reunion with her calf. The percentages were calculated before the calf was taken out of the pen (c.i.p., calf in pen), 1–10 s after the calf was taken out of the pen, at several time spots during the rest of the 6 h when the calf was separated from the cow, when the cow saw the calf and it was about to be put back with its mother and at various time spots during the first hour

5. Cows waiting to be fed Fig. 1. Eye-white percentage during expectation of feed concentrate. The solid curve shows percentages of eye-white (±S.E.) of 12 cows before the stockman entered, up to nine times during 10 min waiting for the concentrate to be given, and once after they got it. As the stockman reached individual cows with different delays, N is lower in the last part of the figure. N included in each observation is given in the figure. Dotted and dot-hatched lines represent results from a previous experiment lasting 6 min; the upper, curve represents cows exposed to food deprivation (frustration), the lower, stippled represents cows given food (satisfaction). This experiment lasted only for 6 min; therefore these graphs are “stretched” to be more comparable with the graph representing positive expectation. The black, stippled line at 24.3% represents a “neutral” level of eye-white, when cows are not introduced to any stimuli. The “F” in the figure refers to when the cows were given the concentrate.

6. Cows on Diazepam Fig. 1. Percentage of eye white shown by the four experimental groups in Experiment 1 (left) and Experiment 2 (right). Exp. 1: solid, black line, thwarted feeding without diazepam; dotted, black line, thwarted feeding with diazepam; solid, grey line, normal feeding without diazepam; dotted, grey line, normal feeding with diazepam. Exp. 2: conditioned stimulus for concentrate feeding in all groups, dotted lines, with diazepam; solid lines, without diazepam. In Exp. 1, the first observation represents a base level. By the last observation in both experiments, all cows had received food or concentrate.

7. Animal Emotions: Exploring Passionate Natures Marc Bekoff

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

11. 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!

12. 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.:

13. 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.)

14. 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

15. 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

16. 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

17. Lust, Attraction & Attachment in Mammalian Reproduction Helen Fisher, Arthur Aron & Lucy Brown Principle sources and targets of hormones and neurotransmitters implicated in emotions of lust(androgens & estrogens, gonads), attraction (catecholamines, LC, VTA, SN), and attachment (oxytocin & vasopressin, PVN/SON of hypothalamus).

18. 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.

19. Pleasure Centers of the Brain Right here!

20. 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.

21. The nucleus accumbens, in the limbic system near the center of the brain, is often called the "pleasure center" of the brain, but the brain appears to have multiple pleasure centers in the limbic system. However, the nucleus accumbens is the among the most prominent. The nucleus accumbens mediates the release of the neurotransmitter dopamine, which underlies pleasure and relaxation. But the dopamine itself is released from the ventral tegmental area (VTA), another contender for the title of "pleasure center." The VTA releases dopamine to the nucleus accumbens, the prefrontal cortex, amygdala, and septum, all of which play an important role in what is called the reward circuit.

22. 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.

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

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

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