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 West, M. J., A. P. King, et al. (2003). "The case for developmental ecology." Animal Behaviour 66(4): 617-622. c Jim Roetzel

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

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

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

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

16. Curiosity period Behavioral refinement period Fear imprint period Environmental awareness Seniority classification period 14284256 70 84 98 4 Mo8 Mo12 Mo16 Mo Flight Instinct period Secondary Fear period Is this really my dog? period Hastings, P. and E. A. Rouse, Eds. (2004). another piece of the puzzle: Puppy Development. Aloha OR, Dogfolk Enterprises. More detailed view...

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

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... Bolhuis, J. and L. Giraldeau, Eds. (2005). The Behavior of Animals. London, UK, Blackwell Publishing. ‘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

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. An aside: gene action & developmental environment Coppinger, R. and L. Coppinger (2001). Dogs: A Startling New Understanding of Canine Origin, Behavior, and Evolution. New York, NY, Scribner. Predatory motor patterns get incorporated into social play and become a social behavior in BCs 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. Development of fear... Scott, J. P. and J. L. Fuller (1965). Genetics and the Social Behavior of the Dog. Chicago, Chicago University Press.

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

51. Puppy Development: Optimal Potential Once the genes are selected it is our job to take care of both mental and physical development –Our = breeder, shelter and then owner along with dog care professionals

52. Puppy Development: Optimal Potential Critical Periods –The window opens and closes for socialization –Once closed it locks

53. Puppy Development: Optimal Potential 0 to 12-15 days: Neonatal period The Beginning: Touch, taste Stacking the odds: –Whelping box in a quiet area near life –Den, spare room near household life –Private enough for mama dog but close enough for humans to hear pups and pups to hear household once hearing begins –Preferably not a basement or kennel

54. Puppy Development: Optimal Potential 0 to 12-15 days: Neonatal period Stacking the odds: –Create mild forms of stress on a daily basis through basic care Weigh Nail trim Hydration checks –Mild stress during this period has shown to have positive effects on learning and handling stress later Dr Michael Fox 1971 study

55. Puppy Development: Optimal Potential 2 to 3 weeks -Transition period: Begins with eyes opening at 10 to 15 days Changes that occur: –Lapping and Chewing begin –Self-elimination –Hearing 18-20 days – sound startle –Walk not crawl –Agonistic behavior begins –Conditioning can begin i.e. learning

56. Puppy Development: Optimal Potential 2 to 3 weeks -Transition period: Begins with eyes opening at 10 to 15 days Stacking the Odds: –Keep in a stable environment –Remain with mother –Provide new surfaces –Provide novel items & toys –Handle and Talk to daily –Nail trims –Weigh

57. Puppy Development: Optimal Potential 3 to 7 weeks - Canine Socialization begins Changes that occur: –Fear response begins –Teethcome in –Weaning begins –Learning to be a dog –and how to communicate

58. Puppy Development: Optimal Potential 3 to 7 weeks - Canine Socialization begins Stacking the Odds: –Move to larger as needed keeping in active area of house to hear household noises –Still with mom, –her choice –Handle & Talk to daily –Nails & Weights

59. Puppy Development: Optimal Potential 3 to 7 weeks - Canine Socialization begins Stacking the Odds: –Allow a surface to sleep and play on and a different area to toilet on –Add an open crate to the pen with soft bed –Increase individual human attention –

60. Puppy Development: Optimal Potential 3 to 7 weeks - Canine Socialization begins Stacking the Odds: –One on one human play time –New toys and items to chew on –Introduce novel objects –Take outdoors –as a group 1 st, –then 2 by 2, –then individually

61. Puppy Development: Optimal Potential 3 to 7 weeks - Canine Socialization begins Stacking the Odds: –Begin supplemental feeding use multiple bowls to prevent resource guarding –Introduce other safe household dogs and cats –Pups meet prospective –owners if possible –by 5 weeks

62. Puppy Development: Optimal Potential 3 to 7 weeks - Canine Socialization begins Stacking the Odds: –Have visitors of all –types, ages etc. Controllable Children –Car Rides –Vet visits, eye checks preferably before 1st fear period at 8 weeks –Grooming Intro

63. Puppy Development: Optimal Potential 7 -8 weeks – Human socialization period Stacking the Odds: –Separate to eat –Crate time 2 x 2 first –then alone a few days – before going to new home –Collar and leash training –Adoption can begin

64. Puppy Development: Optimal Potential 8 to 12 weeks – Continued human and canine socialization Changes that occur: –8 to 11 weeks: –First Fear Period –Social Dominance –begins around week 10

65. Puppy Development: Optimal Potential 8 to 12 weeks – Continued human and canine socialization Stacking the Odds –Avoid surgery, traumatic vet visits, groomer –Ideal adoption time 7-11 weeks –Pups staying with breeder beyond 12 weeks: send off for a weekend with friends prior to 12 weeks absolutely no later than 16 to avoid kennel syndrome (fear in new situations)

66. Puppy Development: Optimal Potential 8 to 12 weeks – Continued human and canine socialization Stacking the Odds –Begin training –Consistent expectations – jumping –Continue careful socializing, support any fear with treats and patience –Socialize with other safe pups and known safe adults

67. Puppy Development: Optimal Potential 8 to 12 weeks – Continued human and canine socialization Stacking the Odds –Management & Structure –Introduce to any –activity planned on as –an adult if possible –before 16 weeks Boating, Travel Shows, trials

68. Puppy Development: Optimal Potential 12 weeks to 16 weeks – Socialization continues Changes that occur: –Social dominance stage (begins around 10- 11 weeks) –Possible Increased Independence

69. Puppy Development: Optimal Potential 12 weeks to 16 weeks – Socialization continues Stacking the Odds: –Adoption should be done by 16 weeks –Pups staying with breeder beyond this time must go for a weekend with friends prior to 12 weeks absolutely no later than 16 to prevent shyness. –Visit many new places

70. Puppy Development: Optimal Potential 12 weeks to 16 weeks – Socialization continues Stacking the Odds: –Continue to socialize with other safe pups and known safe adults –Grooming –Training & Prevention –Consistent Expectations –Management & Structure

71. Puppy Development: Optimal Potential 4 months to 8 months – Changes that occur: –Flight instinct anytime during this stage: days to weeks –Some may challenge for leadership not come when called –Become more –independent –Chewing

72. Puppy Development: Optimal Potential 4 months to 8 months - Stacking the Odds : –Management –Training –Consistent Expectations - leadership –Reward behaviors you want manage or ignore behaviors you don’t want –Continued socialization with dogs, people, and new places

73. Puppy Development: Optimal Potential 6 to 18 months - Changes that occur: –Fear periods can come and go –Sexual maturity begins, possible increase in aggressive behaviors –Territorial, object –guarding

74. Puppy Development: Optimal Potential 6 to 18 months – Stacking the Odds: –Continued Training –Consistent Expectations - leadership –Management & Structure Exercise and a job for the dog –Agility, tracking, rally, obedience, breed related activities such as herding, earth dog, hunting –Continued socialization with dogs, people, and new places

75. Puppy Development: Optimal Potential 18 months to 4 years - Changes that occur: –Sexual maturity –Territorial behavior Can be earlier in some dogs –Object guarding Can be earlier in some dogs

76. Puppy Development: Optimal Potential 18 months to 4 years - Stacking the Odds: –Continued Training –Maintain Expectations - leadership –Management & Structure Exercise and a job for the dog –Agility, tracking, rally, obedience, breed related activities such as herding, earth dog, hunting –Maintain socialization with dogs, people, and new places

77. Puppy Development: Optimal Potential