The Cognitive Dog

1. The Cognitive Dog • Class 6: Development

2. Agenda • Questions on paper • Plan for next couple of weeks • Genes and developmental environment • Development in the dog • Development: putting knowledge to work

3. Plan for next couple of weeks • 11/8: Temperament (assignment 1 due) • 11/15: The emotional dog, part 1. • 11/22: NO CLASS • 11/29: The emotional dog, part 2 • 12/6: Experimental design & analysis, Cue Use as case study • 12/13: Learning Theory & Social Learning • 12/20: Pharmacology (assignment 2 due)

4. The big idea for the evening...

5. Development: growing the shape of the brain • “...while the dog is in its first few weeks of life, and growing its brain, it is making the cell connections and re-arranging them in a specific way, according to the signals that are coming from outside. This development predetermines its adult behavior” - coppinger • “Each behavioral system — fear, submission, investigation, play — has its own rate of development, and varies among breeds. Each is dependent on glandular development and hormone secretions, as well as motor coordination and sensory perception. And each feeds back on the puppy to change not only the shape of the brain...” - coppinger Coppinger, R. and L. Coppinger (2001). Dogs: A Startling New Understanding of Canine Origin, Behavior, and Evolution. New York, NY, Scribner.

6. Growing the shape of the brain... • “Behavior is (always) epigenetic – above the genes – an interaction between the genes and the developmental environment” -Coppinger • Genes play a central role in the timing of developmental events, the rate of development, and duration of developmental periods. Small changes can have huge effects by altering when systems come on line relative to one another. • The resulting behavioral form of the animal emerges from the interaction of the genes and the developmental environment. • Evolution takes advantage of regularities in the developmental environment, e.g., mom, con-specifics, lack of mobility. The evolving genes rely on those regularities.

7. Growing the shape of the brain • “Without a focus on development, any observed variation in behavior within or across populations may be automatically attributed to genotypic differences, when, in fact, it may be a consequence of developmental processes.” -West • Example of a “social gateway” • “pattern of recurring social interactions... gating stimulation along different social pathways” • same population of genes, but reliably different behavioral trajectory c Jim Roetzel West, M. J., A. P. King, et al. (2003). "The case for developmental ecology." Animal Behaviour66(4): 617-622.

8. Growing the shape of the brain... • What you see is “the coaction of nature & nurture” • “evolution involves changes in the developmental system (of which the genes are an essential part) but not necessarily changes in the genes themselves... alterations in development may cause genes to become active in the developmental process that were heretofore quiescent...” -Gottlieb • In other words, systematic change in developmental context can produce systematic change in behavioral and/or morphological trajectory, without changing “genes” • Nature and Nurture: each is “on tap” as opposed to “on top Gottlieb, G. (1991). Individual Development and Evolution: The Genesis of Novel Behavior, Oxford University Press.

9. Growing the shape of the brain. • “Behavioral conformation here is a description of the behavioral shape — how the dog moves — when a dog is working” • “Behavioral conformation and physical conformation, in the final analysis, are one and the same.” • “It is not that border collies have genes for herding, but rather, because of gene action, they end up with a differently shaped brain than other breeds” Coppinger, R. and L. Coppinger (2001). Dogs: A Startling New Understanding of Canine Origin, Behavior, and Evolution. New York, NY, Scribner.

10. Overview of Developmental Periods

11. Scott & Fuller • Among the first to articulate clear periods in the development of dogs... • Neonatal • Transition • Period of socialization • Juvenile period

12. Lindsay, S. R. (2000). Applied Dog Behavior and Training. Ames, IA, Iowa State University Press. Neonatal Period • Mostly about Mom...

13. Lindsay, S. R. (2000). Applied Dog Behavior and Training. Ames, IA, Iowa State University Press. Transitional Period • Senses are coming on line & interactions with others becoming important

14. Lindsay, S. R. (2000). Applied Dog Behavior and Training. Ames, IA, Iowa State University Press. Primary & Secondary Socialization • Imprinting & working out mechanics of living in a social group

15. Lindsay, S. R. (2000). Applied Dog Behavior and Training. Ames, IA, Iowa State University Press. Integration

16. 4 Mo 8 Mo 12 Mo 16 Mo 98 70 14 28 42 56 84 Is this really my dog? period Secondary Fear period Flight Instinct period Seniority classification period Environmental awareness Fear imprint period Behavioral refinement period Curiosity period Hastings, P. and E. A. Rouse, Eds. (2004). another piece of the puzzle: Puppy Development. Aloha OR, Dogfolk Enterprises. More detailed view...

17. Scott, J. P. and J. L. Fuller (1965). Genetics and the Social Behavior of the Dog. Chicago, Chicago University Press. Significant variation across breeds

18. Critical Periods & Imprinting(love of the familiar)

19. Critical (sensitive) periods • From Scott & Fuller... • “...we mean a special time in life when a small amount of experience will produce a great effect on later behavior” • “The difference between the amount of effort needed to produce the same effect at different periods determines just how critical the period is”

20. Example: gottlieb’s mallards • Experience tunes perceptual mechanism... • Mallards need to hear themselves or sibs vocalize in the shell in order to respond specifically to mom mallard’s call...

21. Example: Song bird learning... • ‘Innate’ coarse template • Sensitive period in which they tune template to dialect of surrounding con-specifics • Months later begin to practice song, matching production to template Bolhuis, J. and L. Giraldeau, Eds. (2005). The Behavior of Animals. London, UK, Blackwell Publishing.

22. Imprinting • Some species have a critical period in which they rapidly establish social bonds in a process called “imprinting” • Filial (mom & my buddies) and sexual • Often different periods for each • Surprisingly flexible (more about being in the right place at the right time)... • Probably tuned to certain features, and then use those to build more complete perceptual model • Probably a type of associative learning

23. Imprinting... • Bruce’s theory... • It is scaffolding that serves to quickly & coarsely identify potential social partners (gets you in the ballpark) • The world of a juvenile animal is such that even the coarsest mechanism may work most of the time... • Effect is to subsequently bias attention toward those partners • Biased attention makes it easier to learn more about characteristics of partners (refine model)

24. S&F on imprinting in dogs • “Whether rewarded, punished, or treated indifferently, the young animal of the proper age proceeds to form an emotional attachment to whatever is present in the environment at that time. The essential mechanism appears to be an internal process acting on the external environment. In this way it is indeed quite different from conditioning, which is directly dependent on outside circumstances” • “To state it more clearly: a young animal automatically becomes attached to individuals and objects to which it comes into contact during the critical period” Scott, J. P. and J. L. Fuller (1965). Genetics and the Social Behavior of the Dog. Chicago, Chicago University Press.

25. Is it all about food? • S&F and company did experiments in which the dogs were... • fed by machines, but presumably handled by people • fed by people, but minimal handling • punished for approaching people (don’t try this at home) • raised in field with minimal handling but daily exposure Scott, J. P. and J. L. Fuller (1965). Genetics and the Social Behavior of the Dog. Chicago, Chicago University Press.

26. Is it all about food? • “might be formed merely by having the person in sight at frequent intervals” • “feeding is not a necessary part of the development of the social bond...[indeed] feeding by itself does not produce a highly socialized animal” • “formed an attachment in spite of considerable punishment. We must remember that these were fox terrier puppies...” Scott, J. P. and J. L. Fuller (1965). Genetics and the Social Behavior of the Dog. Chicago, Chicago University Press.

27. When surrounded by other species dogs will imprint on them too... • Dogs will imprint on • Cats • People • Sheep • ... • Note: dogs can imprint on multiple species simultaneously if in an environment with multiple species...

28. Coppinger: How to grow an LGD vs. a Herding Dog • How to raise a live stock guarding dog... • Raise the pup among sheep for its first 16 weeks... • The effect is that the dog “treats sheep as its primary social companions.” • How to raise a herding dog... • Raise it among people & dogs for the first 4 months... • Introduce it to sheep at 4 months plus... • Sheep do not become primary social companions, people and dogs are

29. Motor Patterns & Critical Periods • An consistent pattern of movement displayed by animals of a species. • Innate (genes + development), sometimes referred to as “prefunctional” • You can observe motor patterns, but you can only infer function. • Examples: eye-stalk, chase, grab-bite, tongue flick...

30. Coppinger’s rule... • When motor patterns onset during a critical period, they become incorporated into the behavioral repertoire associated with that period... • Predatory motor patterns become part of social ‘play’ • In some instances, if motor patterns are never given an opportunity to be expressed during the critical period, they may drop out (permanently) from the repertoire... • Part of good training is managing the environment

31. Predatory motor patterns get incorporated into social play and become a social behavior in BCs Coppinger, R. and L. Coppinger (2001). Dogs: A Startling New Understanding of Canine Origin, Behavior, and Evolution. New York, NY, Scribner. An aside: gene action & developmental environment • It takes a difference in timing as well as in developmental environment to grow a BC vs. LGD

32. A wicked interesting idea to noodle on... • Maybe there is something about the imprinting phase of canids that makes it particularly susceptible to inter-species imprinting... • Could it be olfactory priming?

33. Fear

34. Interplay of attraction & fear... • “The balanced interplay of attraction and fear is fundamental to bonding and socialization in the broadest sense” • “Early experience with novelty is crucial for the development of normal exploratory behavior”

35. Fear develops after imprinting • Fear begins to appear at 5 weeks and peaks at 8-9 weeks • At its height (8-9 weeks), fearful experiences may result in lifetime fears (almost like negative imprinting) • Not the time to go to your neighborhood dog park... • Fear of the novel (non-fiction ok...) • novel in novel environments less scary than novel objects in familiar environment

36. Scott, J. P. and J. L. Fuller (1965). Genetics and the Social Behavior of the Dog. Chicago, Chicago University Press. Development of fear...

37. Fear in opposition to imprinting • Imprinting and fear work hand-in-hand in the socialization process. • Fear of novelty develops once imprinting has anchored the familiar. • It makes sense that the development of fear would trail imprinting • Learn who your brother is before you learn who your enemy is...

38. Is novelty bad? • Animals attend to novelty because it can signal a biologically significant object or event, good or bad... • Approach (exploration & learning) • Withdrawal (fear & avoidance) • One of the great developmental challenges is learning how to balance exploration vs. fear, vis-a-vis novelty • See an ebb and flow of attraction & withdrawal throughout development

39. Primary & secondary fear periods • Primary is at 8-9 weeks • Life long implications • Secondary is more diffuse (6 - 14 months) • comes on suddenly, lasts for a week or two, disappears • less likely to have long term effects • cookies and a deep breath: this too will pass..

40. Fear kicks in much earlier in wolves... • Coppinger argues onset of “hazard avoidance” motor patterns occurs at 19 days in wolves vs. 8-9 weeks in dogs • Much shorter window for socialization • Intensity and length of fear period may be longer • Wild canids have pronounced secondary fear period around time of sexual maturity Coppinger, R. and L. Coppinger (2001). Dogs: A Startling New Understanding of Canine Origin, Behavior, and Evolution. New York, NY, Scribner.

41. Fox on Socialization & Fear • “... young wolves that are well socialized at three months will lose their attachment if subsequently isolated. It seems essential, therefore, that social behavior must continue to be reinforced. In the absence of such reinforcement, fear and escape reactions overshadow approach and active submission or “greeting” responses towards humans... Fear of the unfamiliar is the primary obstacle to wolf socialization” Fox, M. W. (1971). Behaviour of Wolves, Dogs and Related Canids. Malabar, FL, Krieger Publishing Company.

42. The emerging picture

43. Co-action of genes and developmental context • The diversity of motor patterns (frequency, sequence, context in which they are displayed) that marks different breeds arises from diversely “shaped” brains caused by a diversity of gene action... • genes interacting with developmental environment • the diversity is a product of natural AND artificial selection • Exaggerated and suppressed so as to achieve goal...

44. Imprinting... • Canids imprint shortly after the transition period and this allows them to imprint across species and thus form social bonds across species • One consequence is that the imprinting process biases what they attend to, and this scaffolds inter-species learning • Unclear what the impact of domestication was on imprinting...

45. Socialization... • Imprinting scaffolds social learning, but socialization is more than just imprinting • During secondary socialization, the animal works out the details of living in a social group (made up of those animals on which it has imprinted.) • learn reliable cues that predict behavior of members of social group • refine strategies for avoiding conflict • The longer the period, perhaps the greater the opportunity to form flexible social bonds.

46. Fear... • Fear comes on line after the imprinting & socialization process has commenced and is underway • At some point, it limits the ability to form social bonds • The later the onset of fear, and the lower the level of fear, the greater the opportunity to form social bonds.

47. Fear • Domestication seems to have had 2 effects vis-a-vis wolves with respect to fear... • The onset of the primary fear period is significantly later, and the period ‘seems’ to be shorter. • The secondary fear period is significantly later, more diffuse and significantly less intense.

48. So... • Imprinting, increased length of the primary and secondary socialization periods, and the dramatic shift and diminished strength of the primary and secondary fear stages, all combine to set the stage for an animal who could... • easily form interspecies bonds • easily attend to and learn to use inter-species cues to guide its own behavior

49. Puppy Development: Stacking the Odds for Optimal Potential • Cognitive Dog - Class 7

50. Puppy Development: Optimal Potential • Temperament is formed by • Nature (genes) AND • Nurture (socializing) • One is often blamed • disregarding the other