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Wolves, Dogs, Rearing and Reinforcement: Complex Interactions Underlying Species Differences in Training and Problem-Solving Performance

An Erratum to this article was published on 14 July 2011


Frank and Frank et al. (1982–1987) administered a series of age-graded training and problem-solving tasks to samples of Eastern timber wolf (C. lupus lycaon) and Alaskan Malamute (C. familiaris) pups to test Frank’s (Zeitschrift für Tierpsychologie 53:389–399, 1980) model of the evolution of information processing under conditions of natural and artificial selection. Results confirmed the model’s prediction that wolves should perform better than dogs on problem-solving tasks and that dogs should perform better than wolves on training tasks. Further data collected at the University of Connecticut in 1983 revealed a more complex and refined picture, indicating that species differences can be mediated by a number of factors influencing wolf performance, including socialization regimen (hand-rearing vs. mother-rearing), interactive effects of socialization on the efficacy of both rewards and punishments, and the flexibility to select learning strategies that experimenters might not anticipate.

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  1. 1.

    The testing facility was a rehabilitated chicken coop located in Mansfield Depot on the site of an experimental chicken farm operated by the University of Connecticut in the 1950s.

  2. 2.

    Various alternative punishment media were considered. Similar studies have used electric grids surrounding the platform. However, our experience suggested that wolf pups would very quickly learn to jump beyond any grid that was small enough in area to be economically feasible. By chance, one the project’s volunteer assistants (Dawn Littleton) was a nationally ranked air-rifle competitor and suggested we use felt pellets, which are ordinary shot through an air rifle to clean the bore of lead fouling. A test of the pellets produced only a mild, though startling, sting when the pellets struck an unprotected hand at point-blank range. It is worthy of note that in several hundred shots, Ms. Littleton, shooting from a distance of approximately 4 m and with the front sight removed from the rifle, never failed to hit a called target (paw, tail, etc.).

  3. 3.

    In the original study, food was smeared into a groove in the underside of the unrewarded pair of blocks to prevent olfactory location. In the replication study, this precaution was refined: The grooves were replaced by countersunk 35-mm film containers with wire mesh lids, and the scent barrier shown in Fig. 3 was added to the WGTA.

  4. 4.

    Coppinger and Coppinger (2001) propose that selection for the ability to eat in proximity to people (and the consequent capacity to exploit a new niche, the village dump) produced a distinct breeding population of wolves from which domestic dogs evolved. This scenario suggests explanations for the unsocialized wolves’ indifference to food not considered by Frank, Frank, and Hasselbach (1989). For several weeks after weaning, the mother-reared pups were fed individually with the experimenters in attendance, both to ensure equal rationing of medication (incorporated into their food) and to increase daily contact with humans. Rather than habituating them to human presence, it is possible that the stress associated with forced human proximity may have established a conditioned inappetence. In the experimental setting, the mere presence of humans may therefore have averted the unsocialized pups from food reward.

  5. 5.

    The comparisons reported in Frank, Frank, and Hasselbach (1989) were based on a greater-than-chance threshold of 9 correct responses, the critical value for α = 0.01 (p ≅ 0.0085), which the first author mistakenly believed to be the critical value for α = 0.05.


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Correspondence to Harry Frank.

Additional information

Edited by Stephen Maxson.

The paper originated in a festschrift, Nurturing the Genome, to honor Benson E. Ginsburg on June 2, 2002 and is part of a special issue of Behavior Genetics based on that festschrift.

An erratum to this article can be found at

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Frank, H. Wolves, Dogs, Rearing and Reinforcement: Complex Interactions Underlying Species Differences in Training and Problem-Solving Performance. Behav Genet 41, 830–839 (2011).

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  • Wolves
  • Dogs
  • Socialization
  • Cognition
  • Problem-solving
  • Training
  • Reinforcement