Dog

On this page:

Dog - Aggressive Behavior in Dogs

Dog - Separation Anxiety in Dogs

Dog - Stress

Dog - Social Dominance

Dog - Issues Related to Females Aggression Increasing After Spaying

Dog - Domestication and Evolution of Dogs

Dog - Development

Dog - Barking

Dog - Compulsive Behavior

Dog - Bond

Dog - Shelter and Rescue

Dog - Other


Aggressive Behavior in Dogs

Alexander, S. (2004). Good dog a program to help dog owners address aggression problems in dogs. Animal Behavior Consulting: Theory and Practice, 1(1), 47-65.


American_Veterinary_Society_of_Animal_Behavior. (2007). Guidelines on the Use of Punishment for Dealing with Behavior Problems in Animals [Electronic Version]. AVSAB. Retrieved November 24, 2007 from http://www.avsabonline.org/avsabonline/images/stories/punishment%20guidelines-aversives%20effects-definitions.pdf.


Andy, O. J., & Velamati, S. (1978). Limbic system seizures and aggressive behavior (superkindling effects). Pavlov J Biol Sci, 13(4), 251-264.


    * Abstract: This study was done to further analyze the neural mechanisms underlying aggressive behavior associated with psychomotor or temporal lobe seizures. The studies revealed that superkindling the aggressive system by sequential stimulations at seizure-inducing thresholds, of two or more sites in the limbic, hypothalamic, and basal ganglia structures facilitated the production of aggressive seizures. Aggressive behavior in the freely moving cat was evaluated in relation to the occurrence of hissing and growling during stimulation, after-discharge and postictal period. The behavior was correlated with the frequency of the elicited seizures and the seizure durations. Aggression did develop as a component behavioral manifestation of the limbic (psychomotor) seizure. Development of aggressive seizures was facilitated by "priming" the aggressive system. Optimum levels of aggressive behavior occurred with seizures of medium duration. Catecholamine blockers tended to attentuate the occurrence of aggression, whereas the agonist tended to facilitate it. Once the aggressive system was rendered hyperexcitable, exteroceptive stimuli also evoked aggressive attack behavior. It was concluded that repeatedly recurring limbic system seizures through superkindling mechanisms can eventually render the limbic-basal ganglia-preoptico-hypothalamic aggressive system hyper-responsive to both recurring seizures and to exteroceptive stimuli with resulting aggressive behavior with or without an accompanying seizure.



Appleby, D. L., Bradshaw, J. W. S., and Casey, R. A. (2002). Relationship between aggressive and avoidance behaviour by dogs and their experience in the first six months of life. Veterinary Record, 150, 434-438.


Aronson, L. P., and Dodman, N.H. (1997). Thyroid Dysfunction as a Cause of Aggression in Dogs and Cats. Retrieved February 4, 2005, from http://www.beaconforhealth.org/Thyroid-Aggression.htm


    * Introductory paragraph: In human medicine, behavioral and psychological changes associated with thyroid dysfunction were noted in the nineteenth century. The behavioral abnormalities seen in the hyperthyroid cat have been well described in the literature, and mimic closely the restlessness, insomnia and irritability or aggression described in humans with thyrotoxicosis. Approximately 80% of hyperthyroid cats are hyperactive, while 10-25% are reported to be aggressive, Cats, as well as people, may experience the rarer manifestation of apathetic thyrotoxicosis, characterized by lethargy and depression. This is seen in approximately 10% of feline cases. Hyperthyroid cats are rarely presented to the veterinarian for behavioral signs, Perhaps because aggression is primarily wen when the cat is restrained, we as a profession are more likely to experience this aspect of the disease than the cats' owners, Treating the underlying thyroid problem generally resolves the behavioral problems however, and because onset is often insidious, it is only after the endocrine imbalance has been addressed that the owners appreciate the deterioration in their animal's behavior.



Balaban, M. T., Rhodes, D.L., and Neuringer, A. (1990). Orienting and defense responses to punishment: effects on learning. Biological Psychology, 30, 203-217.


    * Abstract: Two groups of students attempted to learn sequences of letter-number pairs. For both groups, a tone signalled each error. However, for aversive punishment subjects, a mildly painful electric shock followed the tone 20% of the time, whereas the informational punishment subjects received only the tone. Skin conductance responses (SCRs) and cardiac interbeat intervals indicated the presence of an orienting response to the tone in informational punishment subjects and a defense response to the tone in aversive punishment subjects. Accompanying these were significant differences in behavior: aversive punishment subjects completed fewer sequences and had higher error rates. The two groups did not differ in measures of tonic arousal. Session trends suggested that the cardiac orienting response developed in both groups as subjects learned to use the information in the punishment contingency. Defense responses to aversive punishers may complete with orienting responses necessary to the efficient learning of complex tasks.



Bandura, A., Ross, D., and Ross, S.A. (1961). Transmission of aggression through imitation of aggressive models. Journal of Abnormal and Social Psychology, 63, 575-582.


Beata, C. A. (2001). Diagnosis and treatment of aggression in dogs and cats. Retrieved February 3, 2005, from http://www.ivis.org/advances/Behavior_Houpt/beata/ivis.pdf


Beaver, B. V. et al. (2001). A community approach to dog bite prevention American Veterinary Medical Association Task Force on Canine Aggression and Human-Canine Interactions. JAVMA, 218(11), 1732-1749.


Beaver, B. V. (1993). Profiles of dogs presented for aggression. Journal of the American Animal Hospital Association, 29, 564-569.


Bernstein, I. S. (1981). Dominance: the baby and the bath water. The Behavioral and Brain Sciences, 4, 419-457.


Bettencourt, B. A., Talley, A., Benjamin, A. J., & Valentine, J. (2006). Personality and aggressive behavior under provoking and neutral conditions: a meta-analytic review. Psychol Bull, 132(5), 751-777.


Birdsall, T. C. (1998). 5-Hydroxytryptophan: A Clinically-Effective Serotonin Precursor. Alternative Medical Review, 3(4), 271-280.


    * Abstract: 5-Hydroxytryptophan (5-HTP) is the intermediate metabolite of the essential amino acid L-tryptophan (LT) in the biosynthesis of serotonin. Intestinal absorption of 5-HTP does not require the presence of a transport molecule, and is not affected by the presence of other amino acids; therefore it may be taken with meals without reducing its effectiveness. Unlike LT, 5-HTP cannot be shunted into niacin or protein production. Therapeutic use of 5-HTP bypasses the conversion of LT into 5-HTP by the enzyme tryptophan hydroxylase, which is the rate-limiting step in the synthesis of serotonin. 5-HTP is well absorbed from an oral dose, with about 70 percent ending up in the bloodstream. It easily crosses the blood-brain barrier and effectively increases central nervous system (CNS) synthesis of serotonin. In the CNS, serotonin levels have been implicated in the regulation of sleep, depression, anxiety, aggression, appetite, temperature, sexual behaviour, and pain sensation. Therapeutic administration of 5-HTP has been shown to be effective in treating a wide variety of conditions, including depression, fibromyalgia, binge eating associated with obesity, chronic headaches, and insomnia.



Blackshaw, J.K. (1991). An overview of types of aggressive behaviour in dogs and methods of treatment. Applied Animal Behaviour Science. 30(3/4): 351-361.


    * Abstract: In 223 cases of dogs presented to a specialist behavioural clinic in Brisbane, Australia, 87 (39%) were for severe aggression. The classes of aggression included dominance (31.6%), territorial (29%), predatory (12.3%), intermale (12.3%), sibling rivalry (7.9%), fear biting (6%) and idiopathic rage (0.9%). The breeds most represented which attacked humans were the Bull Terrier (16%), German Shepherd and crosses (15%), Cattle dog breeds (Blue Heeler and crosses, 9.2%), Terrier breeds (9.2%), Labrador (8%), Poodle and Cocker Spaniel (both 5.7%) and Rottweiler (4.6%). The dangerous dog list put out by the local Brisbane City Council includes the first three breeds mentioned and the Rottweiler as the top four breeds causing aggression problems. Hospital records in Victoria and Queensland confirm that most damage is caused to humans by Bull Terriers and German Shepherds. Many breeds similar to those in our study are also represented in American data on aggressive breeds. Treatments included obedience training only, restraint only, obedience and restraint, synthetic progestins and obedience, castration, progestins and obedience, castration and obedience, use of chlorpromazine and as a last resort, euthanasia (12.6%). Entire males formed the largest group (44%), followed by castrated males and females (both 21%) and spayed females (15%). Several breeds (Boxer, Briand, Samoyed and St. Bernard) only attacked other animals and birds. This study reinforces evidence that social disruption is caused by aggressive dogs, but it also indicates that many responsible clients seek advice on how to deal with this behavioural problem.



Blackshaw, J. (n/d). Meaningful temperament assessment for aggression in dogs - can it be done? Retrieved June 25, 2004, from https://secure.ava.com.au/content/confer/uam/proc99/blackshaw.htm


Borchelt, P. L., and Voith, V.L. (1996). Dominance Aggression in Dogs. In V. L. Voith, and Borchelt, P.L. (Ed.), Readings in Companion Animal Behavior (pp. 230-237). Trenton, NJ: Veterinary Learning Systems.


Cameron, D B. (1997). Canine dominance associated aggression: Concepts, incidence, and treatment in a private behavior practice. Applied Animal Behaviour Science. 52(3-4): 243-263.


    * Abstract: Dominance-associated aggression (DA) is a normal, natural, evolutionarily selected trait in many species including the canine species. A review of 35 DA cases presented to a small, private, behavior-only veterinary practice revealed that attention addiction was the most commonly associated (66%) secondary diagnosis. The diagnosis of DA was based on standard criteria. Treatments emphasized owner education and understanding of the problem in addition to common behavior modification, surgical, and pharmacological therapies. The necessity for the owners' gaining psychological leadership in relation to the dog was central to the suggested therapy for 34 of the 35 cases. A phone survey was successful in reaching 34 of the 35 cases. Owners' reports showed that 12% of their dogs showed excellent improvement, 44% reported good improvement, and 32% fair improvement. The owners reported themselves as 97% very or extremely pleased with the quality of information received. These figures and other aspects of canine DA were compared with a similar study done at a large, institutional behavior practice. The results were generally quite similar: DA is predominantly a male trait, found in a wide range of small to large purebred and mixed-bred dogs. DA signs are often apparent in quite young puppies, but do not become of significant concern to most pet owners until the dog is 6 to 24 months old. Both studies supported the concept that DA dogs are reliably responsive to treatment.



Chase, I. D., Bartolomeo, C., & Dugatkin, L., A. (1994). Aggressive interactions and inter-contest interval: how long do winners keep winning? Animal Behaviour, 48, 393-400.


Coccaro, E. F., Kavoussi, R. J., and Hauger, R. L. (1997). Serotonin function and antiaggressive response to fluoxetine: a pilot study. Biological Psychiatry, 42(7), 546-552.


    * Abstract: BACKGROUND: The reported inverse relationship between indices of central serotonin (5-HT) function and indices of impulsive aggression in human subjects suggests the possibility that enhancement of 5-HT activity will reduce impulsive aggressive behavior. Although evidence for this hypothesis is emerging, the relationship between baseline central 5-HT system function and antiaggressive responses to treatment with 5-HT agents has not yet been examined in human subjects. METHODS: In this pilot study, we examined the relationship between: a) pretreatment prolactin responses to d-fenfluramine (PRL[d-FEN]) challenge; and b) antiaggressive responses to 12 weeks of treatment with either fluoxetine or placebo in 15 impulsively aggressive personality disordered subjects as observed in a 12-week, double-blind, placebo-controlled trial. RESULTS: Among all subjects there were positive correlations between the pretreatment PRL[d-FEN] response and the percent improvement in Overt Aggression Scale-Modified scores for "Aggression" and "Irritability." These correlations were present in the fluoxetine (n10), but not in the placebo (n 5), treated subjects. CONCLUSIONS: These data suggest the possibility that the antiaggressive response to fluoxetine is directly, rather than inversely, dependent on the responsiveness of central 5-HT synapses in the brain of impulsive aggressive personality disordered subjects.



Cornell University College of Veterinary Medicine Animal Health Newsletter. (n/d). Early recognition and prevention of canine aggression. http://www.hilltopanimalhospital.com/aggression.htm


    * Introduction: It is difficult to find a book or an article about canine aggression that does not begin by stating two givens: Aggression is the most common and usually the most serious behavior problem in dogs. And more dogs are given up to shelters (and subsequently euthanatized) because of aggression rather than for any other reason. Veterinary behavior clinics and individual animal behaviorists throughout the United States report that canine aggression is the most common diagnosis for dogs they are asked to examine and treat. But as the euthanasia figures indicate, treatment for aggression is not always successful. Some behaviorists go as far as to say that aggression is a chronic problem that can be controlled in the majority of cases but can never be cured. The experts also say that owner compliance with treatment and early recognition of aggressive tendencies are critical to management of the problem. Management in these cases can be translated another and blunter way-critical to the life of the dog.



Dehasse, J., Braem, M., and Schroll, S. (n/d). Aggressive Behaviours in Dogs: A New Descriptive-Contextual Classification. Retrieved January 4, 2005, from http://joeldehasse.com/a-english/aggression-dogs-classification.html


    * Introduction: There are many different classifications of aggressive behaviours in dogs that do not seem to agree with each other. Our intention is to review and analyse these classifications and to propose a clinically operative classification based on the integration of several existing ones.



Delgado, J. M. (1976). Neurobiology of aggressive behavior. Boll Soc Ital Biol Sper, 52(18 Suppl), 1-19.


    * Abstract: Causality, neurological mechanisms, and behavioral manifestations may be heterogeneous in different forms of aggressive behavior, but some elements are shared by all forms of violence, including the necessity of sensory inputs, the coding and decoding of information according to acquired frames of reference, and the activation of pre-established patterns of response. Understanding and prevention of violence requires a simultaneous study of its social, cultural, and economic aspects, at parity with an investigation of its neurological mechanisms. Part of the latter information may be obtained through animal experimentation, preferably in non-human primates. Feline predatory behavior has no equivalent in man, and therefore its hypothalamic representation probably does not exist in the human brain. Codes of information, frames of reference for sensory perception, axis to evaluate threats, and formulas for aggressive performance are not established genetically but must be learned individually. We are born with the capacity to learn aggressive behavior, but not with established patterns of violence. Mechanisms for fighting which are acquired by individual experience may be triggered in a similar way by sensory cues, volition, and by electrical stimulation of specific cerebral areas. In monkeys, aggressive responses may be modified by changing the hierarchical position of the stimulated animal, indicating the physiological quality of the neurological mechanisms electrically activated.



DeNapoli, J. S., Dodman, Nicholas H., Shuster, Louis, Rand, William, Gross, Kathy. (2000). Effects of dietary protein content and tryptophan supplementation on dominance aggression, territorial aggression, and hyperactivity in dogs. Journal of the American Medical Association, 217(4), 504-508.


Dodman, N. H., Donnelly, R., Shuster, L., Mertens, P., Rand, W., and Miczek, K. (1996). Use of fluoxetine to treat dominance aggression in dogs. Journal of the American Veterinary Medical Association, 209(9), 1585-1587.


Dodman, N. H., Moon, R., and Zelin, M. (1996). Influence of owner personality type on expression and treatment outcome of dominance aggression in dogs. Journal of the American Veterinary Medical Association, 209(6), 1107-1109.


    * Abstract: OBJECTIVE: To determine the success rate of positive training methods and behavioral modification techniques in dogs with dominance aggression and to compare personality profiles between owners of dominant-aggressive and nondominant dogs. DESIGN: Prospective clinical study. ANIMALS: 10 dominant-aggressive dogs and 10 non-dominant, nonaggressive control dogs. PROCEDURE: Dominance aggression was quantified, using an aggression score, in the 10 dominant dogs before and after a nonconfrontational behavior modification program. The personality profile of the owners of dominant and control dogs, assessed by means of a Keirsey temperament sorter, was compared, as was the influence of owner personality on the outcome of behavioral modification in the dominant dogs. RESULTS: 9 of 10 dominant dogs responded to the nonconfrontational treatment program by a decrease in aggressive response to similar eliciting stimuli. Significant differences were not found between the personality of the owners of dominant versus control dogs, and owner personality did not significantly affect the outcome of behavior modification treatment. CLINICAL IMPLICATIONS: Nonconfrontational behavior modification programs are effective in reducing owner-directed dominance aggression in dogs. Owner personality does not necessarily predispose certain individuals to assaults by dominant dogs or profoundly affect their ability to engage in a successful behavioral modification program.



Dunbar, I. F. (1975). Behavior of castrated animals. Vet Rec, 96(4), 92-93.


Dodman, N. (n/d). Medical Causes of Aggression. http://www.petplace.com/articles/artShow.asp?artID=1807


    * Introduction: Aggression in dogs is defined as a threat of harmful behavior directed at another animal or person. It may involve snarling, growling, snapping, nipping, biting, or lunging. A dog may act aggressively for either behavioral or medical reasons, or a combination of both. Here are some of the medical conditions that may contribute to or cause canine aggression.



Edwards, D. E., and Kravitz, E.A. (1997). Serotonin, social status and aggression. Current Opinions in Neurobiology, 7, 812-819.


    * Abstract: Serotonin, social status and aggression appear to be linked in many animal species, including humans. The linkages are complex, and, for the most part, details relating the amine to the behavior remain obscure. During the past year, important advances have been made in a crustacean model system relating serotonin and aggression. The findings include the demonstration that serotonin injections will cause transient reversals in the unwillingness of subordinate animals to engage in agonistic encounters, and that at specific synaptic sites involved in activation of escape behavior, the direction of the modulation by serotonin depends on the social status of the animal.



Enquist, M. (1985). Communication during aggressive interactions with particular reference to variation in choice of behavior. Animal Behaviour, 33, 1152-1161.


Fatjó, J., & Manteca, X. (2002). Aggression Towards Unfamiliar People and Other Dogs: Diagnosis and Treatment. Retrieved October 18, 2005, from http://www.vin.com/proceedings/Proceedings.plx?CID=WSAVA2002&PID=2563


Giammanco, M., Tabacchi, G., Giammanco, S., Di Majo, D., and La Guardia, M. (2005). Testosterone and aggression [Electronic Version]. Med Sci Monit, 11, 136-145. Retrieved September 10, 2005 from http://www.medscimonit.com/pub/vol_11/no_4/4259.pdf.


Golab, G. C. (1998). New task force addresses canine aggression. J Am Vet Med Assoc, 213(8), 1097, 1108.


Goldberg, M. E., & Horovitz, Z. P. (1978). Antidepressants and aggressive behavior. Mod Probl Pharmacopsychiatry, 13, 29-52.


Goodwin, D., Bradshaw, J. W. S., & Wickens, S. M. (1997). Paedomorphosis affects agonistic visual signals of domestic dogs. Animal Behaviour, 53, 297-304.


Glancy, G. D., and Knott, T.F. (2003). Part III: The psychopharmacology of long-term aggression--toward an evidence-based algorithm. CPA Bulletin, February, 13-16.


    * Abstract: This is the third in a series of three articles. Taken together, these three papers review the published literature on the psychopharmacology of aggression. The objective is to guide clinicians toward a rational choice of pharmaceutical agents for the treatment of aggressive symptoms presented across the diagnostic spectrum. In the first two articles, we outlined a model for the classification of aggression, together with the method we used to search and review the literature (1,2). In this article, we focus on the use of beta blockers and antipsychotics (APs). We then come to some conclusions and attempt to use these to introduce what we believe is a useful algorithm to guide clinicians.



Guy, N. C., Luescher, A., Dohoo, S. E., Spangler, E., Miller, J. B., Dohoo, I. R., et al. (2001). A case series of biting dogs: characteristics of the dogs, their behaviour, and their vistims. Applied Animal Behaviour Science, 74, 43-57.


Guy, N. C., Luescher, U. A., Dohoo, S. E., Spangler, E., Miller, J. B., Dohoo, I. R., et al. (2001). Demographic and aggressive characteristics of dogs in a general veterinary caseload. Applied Animal Behaviour Science, 74, 15-28.


Harper-Jaques, S., and Reimer, M. (1994). Management of Aggression and Violence. In A. J. Reiss, Miczek, K.A., and Roth, J.A. (Ed.), Understanding and Preventing Violence, Volume 2: Biobehavioral Influences (pp. 802-822).


Hart, B. J., and Hart, L. A. (1997). Selecting, raising, and caring for dogs to avoid problem aggression. Journal of the American Veterinary Medical Association, 210(8), 1129-1134.


Hart, B. L., & Eckstein, R. A. (1997). The role of gonadal hormones in the occurance of objectionable behaviours in dogs and cats. Applied Animal Behaviour Science, 52, 331-344.


Horwitz, D. F. (2001). Fears and fear aggression. Retrieved February 4, 2005, from http://www.vin.com/ACVC/2001/AuthorIndex.htm


Houpt, K. A., Honig, S. U., & Reisner, I. R. (1996). Breaking the human-companion animal bond. J Am Vet Med Assoc, 208(10), 1653-1659.


Insurance Information Institute. (2004). Dog Bite Liability. Retrieved February 3, 2005, from http://www.iii.org/media/hottopics/insurance/dogbite/


    * Introduction: In 2005 dog bites cost insurers $317.2 million, little changed from $321.6 million in 2003 but down 8 percent from $345.5 million in 2002. While the number of claims paid by insurers fell from approximately 20,800 in 2002 to 15,000 in 2005 -- a decrease of 28 percent -- the cost of the average dog bite claim rose sharply, from roughly $16,600 in 2002 to $21,200 in 2005. Liability claims account for approximately 4 percent of homeowners claims. Dog bite claims in 2005 accounted for about 15 percent of liability claims dollars paid under homeowners insurance policies. According to the Centers for Disease Control and Prevention, more than 4.7 million people are bitten by dogs annually, resulting in an estimated 800,000 injuries that require medical attention. With over 50 percent of the bites occurring on the dog owner’s property, the issue is a major source of concern for insurers. Over the years, many states have passed laws with stiff penalties for owners of dogs that cause serious injuries or deaths. In about one-third of states, owners are "strictly liable" for their dogs' behavior, while in the rest of the country they are liable only if they knew or should have known their dogs had a propensity to bite (known as the "one free bite" principle).



Jacobs, C., De Keuster, T., and Simoens, P. (2003). Assessing the pathological extent of aggressive behaviour in dogs. Vet Q., 25(2), 54-60.


    * Abstract: In this review the variety of parameters used for evaluating the pathological extent of aggressive behaviour is summarised and the practical usefulness of each parameter is discussed. The selected parameters are: the objective analytic description of the aggressive behaviour, the function of the aggression, the presence of the three phases of a normal aggression sequence, the number of bites per attack, the duration of the attack and the frequency of the aggressive behaviour. Other criteria such as the appropriateness of the aggression in relation to the context, the predictability of the aggression and the severity of the caused injury are biased because of the variation caused by numerous external factors. The relevance of the most suitable parameters will be assessed in a further study in which the distribution of aggression modulating neurotransmitter receptors will be determined.



Judge, P. G. (2000). Coping with Crowded Conditions. In F. Aureli, and de Waal, Frans B.M. (Ed.), Natural Conflict Resolution (pp. 129-154). Berkeley and Los Angeles: California University Press.


Katz, R. J. (1980). Role of serotonergic mechanisms in animal models of predation. Prog Neuropsychopharmacol, 4(3), 219-231.


Kenard, L. (1998). Reducing aggression and violence the serotonin connection. Retrieved September 6, 2005, from http://www.life-enhancement.com/article_print.asp?ID=208&type=


    * Introduction: obert Louis Stevenson's The Strange Case of Dr. Jekyll and Mr. Hyde has long been viewed as a dissection of the Good and Evil that can exist within a single human being. As the well-known story goes, Jekyll ingests his formula and is temporarily transformed from a conservative, well-respected English physician into a vain, uninhibited, terrifyingly violent criminal. Not only has Jekyll and Hyde stood the test of time as a morality tale, studies in brain chemistry and behavior more than a century later have shown Stevenson to have had remarkable prescience regarding the role of the neurotransmitter serotonin. Of course, Jekyll's formula was simply a fictional plot device. Other than somehow releasing some of man's baser instincts, Stevenson could have had no idea what was actually going on neurochemically. But the 100+ years of research on brain chemistry since Stevenson wrote his Victorian classic have revealed that anything that interferes with the actions of serotonin in the brain can bring about a syndrome that resembles Jekyll's transformation to Hyde. While certainly less dramatic than the transformation described by Stevenson, serotonin deficiency bears a striking resemblance in various manifestations as an increased tendency toward anxiety, depression, out-of-control disinhibition, and violence. Conversely, enhancing the activity of the serotonin system may have exactly the opposite effects in many people. Given our current knowledge of neurochemistry, there can be little doubt that if Stevenson were writing today, Jekyll's transforming formula would have been a potent anti-serotonergic agent.



Kim, H. H., Yeon, S. C., Houpt, K. A., Lee, H. C., Chang, H. H., & Lee, H. J. (2005). Effects of ovariohysterectomy on reactivity in German Shepherd dogs. The Veterinary Journal, xxx, xxx-xxx.


Kostowski, W. (1978). Effects of sedatives and major tranquilizers on aggressive behavior. Mod Probl Pharmacopsychiatry, 13, 1-12.


Kravitz, E. A. (2000). Serotonin and aggression: insights gained from a lobster model system and speculation on the role of amine neurons in complex behavior. Journal of Comparative Physiology, 186, 221-238.


    * Abstract: The amine serotonin has been suggested to play a key role in aggression in many species of animals, including man. Precisely how the amine functions, however, has remained a mystery. As with other important physiological questions, with their large uniquely identifiable neurons, invertebrate systems offer special advantages for the study of behavior. In this article we illustrate that principal with a description of our studies of the role of serotonin in aggression in a lobster model system. Aggression is a quantifiable behavior in crustaceans, the amine neuron systems believed to be important in that behavior have been completely mapped, and key physiological properties of an important subset of these netirons have been defined. These results are summarized here, including descriptions of the "gain-setter" role and "autoinhibition" shown by these neurons. Results of other investigations showing socially modulated changes in amine responsiveness at particular synaptic sites also are described. In addition, speculations are offered about how important developmental roles served by amines like serotonin, which have been well described by other investigators, may be related to the behaviors we are examining. These speculations draw heavily from the organizational/activational roles proposed for steroid hormones by Phoenix et al. (1959).



Kroll, T. L., Houpt, K.A., and Erb, H.N. (2004). The use of novel stimuli as indicators of aggressive behavior in dogs. Journal of the American Animal Hospital Association, 40, 13-19.


    * Abstract: To test the predictive value of a doll and an artificial hand, reactions of dogs (n=100) were compared to histories of behavior toward children. Each dog's reaction to the doll and the hand was categorized as normal, fearful, fearfully aggressive, or offensively aggressive. Sixty-five percent (n=37) of the dogs that had a normal or no reaction to the stimuli had a history of being good with children. Eighty-eight percent (n=34) of the dogs that had an aggressive reaction to the doll had a history of aggressive behavior toward a child. Dogs that were fearfully aggressive were significantly more likely to show fearful responses to the doll, and dogs that were either dominant or fearfully aggressive were more likely to exhibit aggression of the same type to the hand. The results of this study indicate that the doll and, to a lesser extent, the hand may be useful components in determining the aggressive tendencies of dogs. The results also point out the major limitations, because the false positives and false negatives are too frequent.



Kulkarni, A. S., & Plotnikoff, N. P. (1978). Effects of stimulants on aggressive behavior. Mod Probl Pharmacopsychiatry, 13, 69-81.


Lindell EM. (1997) Diagnosis and treatment of destructive behavior in dogs. Vet Clin North Am Small Anim Pract. 27(3): 533-47.


    * Abstract: Destructive behavior in dogs can be expensive for owners and life-threatening for dogs. The human-companion animal bond is jeopardized. A diagnostic plan should address both behavioral and medical causes of destructive behavior. Once a diagnosis has been established, a successful therapeutic plan can be formulated. Treatment includes modification of both behavior and environment and may incorporate the judicious use of psychotropic medication.


Manteca, X., & Fatjó, J. (2002). Difficulties in The Diagnosis of Dominance Aggression in Dogs. Retrieved October 18, 2005, from http://www.vin.com/proceedings/Proceedings.plx?CID=WSAVA2002&PID=2562


Marder AR.49: (1991) Animal bites: behavior modification of the offending animal. Semin Vet Med Surg (Small Anim). 6(3): 192-8.


Mazur, A., & Booth, A. (1998). Testosterone and Dominance in Men. Behavioural and Brain Sciences, 21, 353-363. [available at http://cogprints.org/663/00/bbs_mazur.html]


    * Abstract: In men, high levels of endogenous testosterone (T) seem to encourage behavior apparently intended to dominate -- to enhance one's status over -- other people. Sometimes dominant behavior is aggressive, its apparent intent being to inflict harm on another person, but often dominance is expressed nonaggressively. Sometimes dominant behavior takes the form of antisocial behavior, including rebellion against authority and law breaking. Measurement of T at a single point in time, presumably indicative of a man's basal T level, predicts many of these dominant or antisocial behaviors. T not only affects behavior but also responds to it. The act of competing for dominant status affects male T levels in two ways. First, T rises in the face of a challenge, as if it were an anticipatory response to impending competition. Second, after the competition, T rises in winners and declines in losers. Thus, there is a reciprocity between T and dominance behavior, each affecting the other. We contrast a reciprocal model, in which T level is variable, acting as both a cause and effect of behavior, with a basal model, in which T level is assumed to be a persistent trait that influences behavior. An unusual data set on Air Force veterans, in which data were collected four times over a decade, enables us to compare the basal and reciprocal models as explanations for the relationship between T and divorce. We discuss sociological implications of these models.



Miczek, K. A., Weerts, E., Haney, M., and Tidey, J. (1994). Neurobiological mechanisms controlling aggression: preclinical developments for pharmacotherapeutic interventions. Neuroscience and Biobehavioral Reviews, 18(1), 97-110.


    * Abstract: Current pharmacotherapeutic approaches to the management of violent and aggressive behavior rely mostly on agents that act as receptor agonists or antagonists at subtypes of brain dopaminergic, GABAergic, and serotonergic receptors. Ethological experimental studies in animals have shown that drugs may modulate aggression by inhibiting motor activity, by distorting aggression-provoking or -inhibiting signals, by fragmenting behavioral sequences or temporal patterning, or by increasing the rate and intensity of aggressive acts. Evidence from animal studies points to large changes in selected brain dopamine, serotonin, and GABA systems during and following aggressive and defensive behavior. However, the specificity of drugs that are currently used to control aggressive behavior through their action as agonists or antagonists at subtypes of dopamine, serotonin or GABA receptors continues to be of concern. Similar to the effects of widely used traditional neuroleptics that nonselectively antagonize dopamine receptors, the range of behaviors which is suppressed by either D1 or D2 receptor antagonists is pervasive. At present, systemic administration of dopamine receptor antagonists in animal preparations does not target aggression-specific mechanisms. The GABAA/Benzodiazepine/Chloride ionophore receptor complex is implicated in the aggression-heightening effects of alcohol and benzodiazepines. Although early reports focused on the "taming" effects of benzodiazepine anxiolytics, low doses may enhance aggression in both animals and humans. Benzodiazepine antagonists block heightened aggression after low doses of alcohol or benzodiazepines. Agonists at certain 5-HT1 receptor subtypes such as eltoprazine are potently effective in reducing aggressive behavior of males and females of various animal species under conditions that promote charging offensive-type aggression, without adversely affecting nonaggressive components of the behavioral repertoire. However, recent reports indicate that eltoprazine and related compounds may potentiate anxiety reactions in rodents, and question the behavioral specificity of these substances. Opioid receptor antagonists modulate primarily physiological and behavioral responses of defense and submission. Defeated animals show tolerance to opiate analgesia and withdrawal responses upon challenge with opioid receptor antagonists. Defensive and submissive vocalizations are potently blocked by opioid peptides. Substances that target specific receptor subtypes at serotonergic, GABAergic and opioidergic synapses are most promising for the selective modification of aggressive, defensive and submissive behavior patterns.



Monroe, K. (2005). PWD aggression Can it happen to you? The Courier, May/June, 95-105.


Moyer, K. E. (1968). Kinds of aggression and their physiological basis. Communications in Behavioral Biology, Part A, 2, 65-87.


Moynihan, M. H. (1998). The Social Regulation of Competition and Aggression in Animals. Washington: Smithsonian Institute.


Mueller, M., M., Wilczynski, S.M., Moore, J.W., Fusilier, I., and Trahant, D. (2001). Antecedent manipulations in a tangible condition: effects of stimulus preference on aggression. Journal of Applied Behavior Analysis, 34, 237-240. [available at http://www.pubmedcentral.nih.gov/picrender.fcgi?artid=1284319&blobtype=pdf]


    * Abstract: After a functional analysis indicated that aggression of an 8-year-old boy with autism was maintained by access to preferred items, antecedent manipulations involving the relative preference of restricted and noncontingently available stimuli were conducted. Restricting highly preferred items evoked the highest rates of aggression regardless of the preference level of the noncontingently available alternative items. Restricting less preferred stimuli was associated with moderate rates of aggression even when the alternative items were more preferred.



Netto, W. J., va der Borg, J.A., and Slegers, J.F. (1992). The establishment of dominance relationships in a dog pack and its relevance for the man-dog relationship. Tijdschr Diergeneeskd, 117, 51S-52S.


Netto, W. J., va der Borg, J.A., & Planta, D. J. U. (1997). Behavioural testing for aggression in the domestic dog. Applied Animal Behaviour Science, 52, 242-263.


Noll, D. E. (1999). The neuropsychology of human conflict. Retrieved from http://www.manageconflict.com/Neuropsy.htm.


O'Farrell, V., & Peachey, E. (1990). Behavioural effects of ovariohysterectomy on bitches. Journal of Small Animal Practice, 31, 595-598.


O'Heare, J. (2007). Aggressive Behavior in Dogs. Ottawa: DogPsych Publishing.


O'Heare, J. (2004). The Canine Aggression Workbook (3nd ed.). Ottawa: DogPsych Publishing.


Overall, K. L. (2007). Why electric shock is not behavior modification. Journal of Veterinary Behavior, 2, 1-4.


Overall, K. L. (2001). Aggression: triggers, flashpoints, and diagnoses. Retrieved February 4, 2005, from http://www.vin.com/VINDBPub/SearchPB/Proceedings/PR05000/PR00379.htm


Overall, K. L. (2001). Pharmacology and behavior: neurochemistry of anxiety and aggression. Retrieved February 4, 2005, from http://www.vin.com/ACVC/2001/AuthorIndex.htm


Overall, K. L. (2001). Aggression: treatment options. Retrieved February 4, 2005, from http://www.vin.com/ACVC/2001/AuthorIndex.htm


Overall, K. (n/d). Karen Overal's Behavior Modification Program Protocol for Relaxation. http://home.gci.net/~divs/behavior/bemod_relax.html


Overall. K. L. (2003). Suggest temporary boarding first when confronted with behavioral euthanasia. http://www.dvmnewsmagazine.com/dvm/article/articleDetail.jsp?id=76364


Overall. K. L. (2004). Behavior signals interpreted with body postures. http://www.dvmnewsmagazine.com/dvm/article/articleDetail.jsp?id=94404


Overall. K. L. (2004). Are you fluent in Dog? http://www.dvmnewsmagazine.com/dvm/article/articleDetail.jsp?id=143615


Pal, S. K., Ghosh, B., & Roy, S. (1998). Agonistic behaviour of free-ranging dogs (Canis familiaris) in relation to season, sex and age. Applied Animal Behaviour Science, 59, 331-348.


Pelios, L., Morren, J., Tesch, D., and Axelrod, S. (1999). The impact of functional analysis methodology on treatment choice for self-injurious and aggressive behavior. Journal of Applied Behavior Analysis, 32, 185-195. [available at: http://seab.envmed.rochester.edu/jaba/articles/1999/jaba-32-02-0185.pdf]


    * Abstract: Self-injurious behavior (SIB) and aggression have been the concern of researchers because of the serious impact these behaviors have on individuals’ lives. Despite the plethora of research on the treatment of SIB and aggressive behavior, the reported findings have been inconsistent regarding the effectiveness of reinforcement-based versus punishment-based procedures. We conducted a literature review to determine whether a trend could be detected in researchers’ selection of reinforcement-based procedures versus punishment-based procedures, particularly since the introduction of functional analysis to behavioral assessment. The data are consistent with predictions made in the past regarding the potential impact of functional analysis methodology. Specifically, the findings indicate that, once maintaining variables for problem behavior are identified, experimenters tend to choose reinforcement-based procedures rather than punishment-based procedures as treatment for both SIB and aggressive behavior. Results indicated an increased interest in studies on the treatment of SIB and aggressive behavior, particularly since 1988.



Penturk, S., & Yalcin, E. (2003). Hypocholesterolaemia in dogs with dominance aggression. J Vet Med A Physiol Pathol Clin Med, 50(7), 339-342.


Peremans, K., Audenaert, K., Coopman, F., Blanckaert, P., Jacobs, F., Otte, A., et al. (2003). Estimates of regional cerebral blood flow and 5-HT2A receptor density in impulsive, aggressive dogs with 99mTc-ECD and 123I-5-I-R91150. Eur J Nucl Med Mol Imaging, 30(11), 1538-1546.


Peremans, K., Audenaert, K., Hoybergs, Y., Otte, A., Goethals, I., Gielen, I., et al. (2005). The effect of citalopram hydrobromide on 5-HT2A receptors in the impulsive-aggressive dog, as measured with 123I-5-I-R91150 SPECT. Eur J Nucl Med Mol Imaging, 32(6), 708-716.


Perry, G. (n/d). Aggression in dogs: a complete review. Retrieved March 2004, from http://www.ava.com.au/UAM/proc92/15.htm


PetPlace.com. (n.d.). Tryptophan. Retrieved January 31, 2005, from http://www.petplace.com/articles/artShow.asp?artID=5187


Phillips, K. (n/d). Liability for dog bites and other looses. Retrieved February 3, 2005, from http://www.dogbitelaw.com/PAGES/liability.htm


Phillips, K. (n/d). Legal rights of a dog bite victim. Retrieved February 3, 2005, from http://www.dogbitelaw.com/PAGES/legal_ri.htm


Phillips, K. (n/d). Dog Bite Law Ontario. Retrieved February 3, 2005, from http://www.dogbitelaw.com/PAGES/Ontario.htm


Planta, D. J. U., and Netto, W.J. (1999). Behavioural testing for aggression in the domestic dog. Retrieved June 30, 2005, from http://wwwhome.cs.utwente.nl/~beijnum/fsk-evaluatie/Planta_proceedingslyon99.PDF


Planta, D. J. U. (2002). Testing dogs for aggressive biting behaviour and fear behaviour for breeding purposes. Retrieved from http://home.wanadoo.nl/rashonden-nederland/downloads/UK_Planta.pdf


Podberscek, A. L., & Serpell, J. A. (1998). Environmental influences on the expression of aggressive behaviour in Engligh Cocker Spaniels. Applied Animal Behaviour Science, 52(3-4), 215-227.


Polsky, R. (2000). Can aggression in dogs be elicited through the use of electronic pet containment systems? Journal of Applied Animal Welfare Science, 3(4), 345-357.


    * Abstract: Five cases are described that involve severe attacks on humans by dogs who were being trained or maintained on an electronic pet containment system. The system is designed to boundary train a dog through the use of electric shock in an escape-avoidance conditioning paradigm. Data were collected from legal documents filed in personal injury lawsuits. Analysis of the findings show that all dogs lacked a marked history of aggressive responding, all were adult males, and most were reproductively intact. All attacks happened near the boundary of the property. In every case, the system was operational at the time of attack. Moreover, in most cases, the dog received shock. Findings lend themselves to possible interpretation in terms of unconditioned aggression as a result of a dog having received electronic shock and avoidance-motivated aggression mediated through fear reduction toward human stimuli.



Reich, M. (2001). Topics in veterinary Behavior: Canine fear aggression. http://www.southpaws.com/news/Newsletter-Spring%202001/page5.html


Reid, P. J., and Penny, N.J. (n/d). Canine aggression toward children: are simulations valid tools? Retrieved February 5, 2005, from http://www.animalbehaviour.ca/research.html


Reisner, I. (1998). Canine aggression: neurobiology, behavior and management. Retrieved from http://www.vetshow.com/friskies/cani.htm.


Reisner, I. R. (2003). Differential diagnosis and management of human-directed aggression in dogs. The Veterinary Clinics Small Animal Practice, 33, 303-320.


    * Abstract: Canine aggression directed to human beings is a common presenting complaint and requires attention to safety issues and behavior modification to minimize the risks of future aggression. Dogs may bite familiar people, including family members, or unfamiliar people for a variety of reasons. Anxiety plays an important role in aggression regardless of its target or circumstances. Effective management of aggression may include education and safety counseling for owners, lifestyle changes for dogs and owners, avoidance of provocations when possible, and behavior modification to minimize the risk of future bites. Drug therapy may be indicated to facilitate behavior modification or to reduce reactivity in the dog.



Reiss, A. J., Miczek, K.A., and Roth, J.A. (Ed.). (1994). Understanding and Preventing Violence, Volume 2: Biobehavioral influences (Free Executive Summary): National Academy of Sciences.


Rooney, N. J., and Bradshaw, J.W.S. (2003). Links between play and dominance and attachment dimensions of dog-human relationships. Journal of Applied Animal Welfare Science, 6(2), 67-94.


    * Abstract: It is often claimed that certain behavioral problems in domestic dogs can be triggered by the games played by dog and caregiver (owner). In this study, we examine possible links between the types of games played and dimensions of the dog-owner relationship that are generally considered to affect such problems. Fifty dog-owner partnerships were filmed during 3-min play sessions in which the owner was allowed to choose the games played. All partnerships then undertook a 1-hr test designed to measure elements of behavior commonly ascribed to "dominance" and "attachment." Principal components analysis of the data produced 2 dominance-related factors (Amenability and Confident Interactivity) and 4 factors describing aspects of attachment (Nonspecific Attention Seeking, Preference for Owner, Preference for Unfamiliar Person, and Separation-Related Behavior). Amenability, in particular, varied significantly between breeds. In the study, we then compared types of games played to each of these factors. Dogs playing rough-and-tumble scored higher for Amenability and lower on Separation-Related Behavior than did dogs playing other types of games. Dogs playing tug-of-war and fetch scored high on Confident Interactivity. Winning or losing these games had no consistent effect on their test scores. If the dog started the majority of the games, the dog was significantly less amenable and more likely to exhibit aggression. The results suggest that how dogs play reflects general attributes of their temperament and relationship with their owner. This study provides no evidence that games play a major deterministic role on dominance dimensions of dog-human relationships, but the results suggest that playing games involving considerable body contact may affect attachment dimensions.



Scott, J. P. (1948). Dominance and the frustration-aggression hypothesis. Physiological Zoology, 21, 31-39.


Selby, L. A., Rhoades, J. D., Irvin, J. A., Carey, G. E., & Wade, R. G. (1980). Values and limitations of pet ownership. J Am Vet Med Assoc, 176(11), 1274-1276.


    * Abstract: A questionnaire was used to collect information concerning the values and limitations of pet ownership. The major reasons given for nonownership were housing limitations, emotional dissatisfaction with animals, destructive habits of pets, and a transient household status. Various stages of the family's "life cycle" had a role in determining pet ownership. The pet's agreeable temperament (eg, intelligence, gentleness, and playfulness) was considered to be the major advantages of pet ownership. The major negative characteristics of pet ownership were the destructiveness and overaggressiveness of the pet.



Sherman, C.K. (1996). Characteristics, treatment, and outcome of 99 cases of aggression between dogs. Applied animal behaviour science. 47(1/2): 91-108.


Shyan, M. R., Fortune, K.A., and King, C. (2003). "Bark parks"--a study on interdog aggression in limited-control environments. Journal of Applied Animal Welfare Science, 6(1), 25-32.


Simon, N. G., Kaplan, J.R., Hu, S., Register, T.C., and Adams, M.R. (2004). Increased aggressive behavior and decreased affiliative behavior in adult male monkeys after long-term consumption of diets rich in soy protein and isoflavones (abstract only). Hormones and Behavior, 45(4), 278-284.


    * Abstract: Estrogen produced by aromatization of gonadal androgen has an important facilitative role in male-typical aggressive behavior that is mediated through its interaction with estrogen receptors (ER) in the brain. Isoflavones found in soybeans and soy-based dietary supplements bind ER and have dose- and tissue-dependent effects on estrogen-mediated responses. Yet, effects of isoflavone-rich diets on social and aggressive behavior have not been studied. We studied the effects of long-term (15 months) consumption of diets rich in soy isoflavones on spontaneous social behavior among adult male cynomolgus macaques (Macaca fascicularis) (n = 44) living in nine stable social groups. There were three experimental conditions which differed only by the source of dietary protein: casein and lactalbumin (no isoflavones), soy protein isolate containing 0.94 mg isoflavones/g protein, and soy protein isolate containing 1.88 mg isoflavones/g protein. In the monkeys fed the higher amount of isoflavones, frequencies of intense aggressive (67% higher) and submissive (203% higher) behavior were elevated relative to monkeys fed the control diet (P's < 0.05). In addition, the proportion of time spent by these monkeys in physical contact with other monkeys was reduced by 68%, time spent in proximity to other monkeys was reduced 50%, and time spent alone was increased 30% (P's < 0.02). There were no effects of treatment on serum testosterone or estradiol concentrations or the response of plasma testosterone to exogenous gonadotropin-releasing hormone (GnRH). The results indicate that long-term consumption of a diet rich in soy isoflavones can have marked influences on patterns of aggressive and social behavior.



Steiner, H., Saxena, K., and Chang, K. (2003). Psychopharmacologic strategies for the treatment of aggression in juveniles. CNS Spectrums, 8(4), 298-308.


    * Abstract: Maladaptive aggression in youth is one of the most common and troublesome reasons for referrals to child psychiatrists. It has a complex relationship with psychopathology. There are several syndromes, which are primary disturbances of clustered maladaptive aggression, most notably oppositional defiant disorder and conduct disorder. However, problems with aggression also appear in a wide range of other disturbances, such as bipolar disorder, posttraumatic stress disorder, and mood disorders. Additionally, aggression is normative, serves an adaptive purpose and can be situationally induced. These complexities need to be carefully addressed before targeting maladaptive aggression psychopharmacologically. We summarize the literature on the psychopharmacology of maladaptive aggression in youth, focusing on disorders without cognitive impairment. We delineate the subtypes of aggression which are most likely to respond to medication (reactive-affective-defensive-impulsive in their acute and chronic form) and conclude with a discussion of specific medication strategies which are supported by controlled clinical trials and clinical experience.



Torda, C. (1977). Compulsive aggressive or anxious behavior (postnatal preconditioning). Agressologie, 18(3), 149-160.


Tortora, D. F. (1983). Safety training: elimination of avoidance-motivated aggression in dogs. Journal of Experimental Psychology, 112(2), 176-214.


Uchida, Y., Dodman, N., DeNapoli, J., and Aronson, L. (1997). Characterization and treatment of 20 canine dominance aggression cases. J Vet Med Sci., 59(5), 397-399.


    * Abstract: This study was undertaken to characterize 20 cases of dominance aggression seen at Tufts University School of Veterinary Medicine and to investigate the efficacy of our non-confrontational behavior modification program for 8 weeks. The 20 cases included 18 pure breed and 2 mixed breed dogs. Thirteen of the dogs were male. The dogs' ages ranged from 7 to 84 months (mean 32.1 +/- 22.64 SE). There was no correlation between the severity of dominance aggression and the signalment of the dogs. At the conclusion of the eight week follow up period, 14 dogs (70%) were reported to have responded to the treatment to some degree. Six dogs did not demonstrate any noticeable reduction in aggressive behavior or became more aggressive. The results of the study is powerful evidence of the efficacy of the non-confrontational behavior modification program.



Ulrich, R. E., Hutchinson, R.R., and Azrin, N.H. (1965). Pain-elicited aggression. The Psychological Record, 15, 111-126. [available at http://www.pubmedcentral.gov/picrender.fcgi?artid=1338713&blobtype=pdf]


    * Abstract: Painful mechanical tail-pinch elicited aggressive responses in paired rats; response-contingent electric shock to either forepaws or hindpaws suppressed fighting and stereotyped aggressive postures, including those in which dominance was expressed. There was no evidence that aggression was facilitated by shock that was contingent on pain-elicited aggressive responses. Aggressive responding recovered when shock was discontinued.



Unshelm, A. R. J. (1997). Aggressive conflicts amongst dogs and factors affecting them. Applied Animal Behaviour Science, 52, 229-242.


van den Berg, L. S., M. B. H., and Knol, B. W. (2003). Behavior genetics of canine aggression: behavioral phenotyping of golden retrievers by means of aggression test. Behavior Genetics, 33(5), 469-483.


    * Abstrct: Molecular genetic analysis of complex traits such as aggression strongly depends on careful phenotyping of individuals. When studying canine aggression, the information provided by the owners of the dogs is often not detailed and reliable enough for this purpose. Therefore we subjected 83 golden retrievers, both aggressive and nonaggressive individuals, to a behavioral test. These tests were analyzed with help of an ethogram, resulting in a behavioral profile for each of the dogs. In this article three methods are described of converting these profiles into a measure of behavioral phenotype. The usefulness of the methods is evaluated by comparing the test results with information provided by owners. Moreover, the hypothesis underlying all these methods, that a lowered threshold for aggressive behavior in general is present in the dogs, is also evaluated. Future research will need to reveal whether the methods meet the high standards that are necessary for studying complex traits.


van Kerkhove, W. (2004). A fresh look at the wolf-pack theory of companion-animal dog social behavior. J Appl Anim Welf Sci, 7(4), 279-285; discussion 299-300.


Vas, J., Topal, J., Gacsi, M., Miklosi, A. P., R., and Csonji, T. V., & Csanyi, V. (2005). A friend or an enemy? Dogs' reaction to an unfamiliar person showing behavioural cues of threat and friendliness at different times. Applied Animal Behaviour Science, 94, 99-115.


Vasb, J., Topála, J., Gácsia, M., Miklósia, A., and Csányi, V. (2005). A friend or an enemy? Dogs’ reaction to an unfamiliar person showing behavioural cues of threat and friendliness at different times. Applied Animal Behaviour Science. 94(1-2), 99-115.


Virga, V., Houpt, K. A., and Scarlett, J. M. (2001). Efficacy of amitriptyline as a pahrmacological adjunct to behavioral modification in the management of aggressive behaviors in dogs. Journal of the American Animal Hospital Association, 37, 325-330.


Voith VL, Borchelt PL. (1982). Diagnosis and treatment of dominance aggression in dogs.Vet Clin North Am Small Anim Pract. 12(4): 655-63.


Voith, V. L. (1981). Profile of 100 animal behavior cases. Mod Vet Pract, 62(6), 483-484.


Voith, V. L. (1980). Play behavior interpreted as aggression or hyperactivity: case histories. Mod Vet Pract, 61(8), 707-709.


Voith, V. L. (1979). Multiple approaches to treating behavior problems. Mod Vet Pract, 60(8), 651-654.


White, M. M., Neilson, J. C., Hart, B. L., and Cliff, K. D. (1999). Effects of clomipramine hydrochloride on dominance-related aggression in dogs. Journal of the American Veterinary Medical Association, 215(9), 1288-1291.


    * Abstract: OBJECTIVE: To compare effects of the serotonergic drug clomipramine hydrochloride with those of placebo for treatment of dominance-related aggression in dogs. DESIGN: Randomized, placebo-controlled, double-blind clinical trial. ANIMALS: 28 neutered dogs > 1 year old with dominance-related aggression. PROCEDURE: Dogs displaying > or 3 aggressive episodes/wk toward > or 1 human family member in response to identifiable behavioral triggers were included in the study. Owners were instructed not to change patterns of interaction with their dogs during the study. After 2 weeks of baseline observations, dogs were treated for 6 weeks with clomipramine (1.5 mg/kg [0.7 mg/lb] of body weight, q 12 h; n = 15) or placebo (13). Responses to triggers were assigned the following aggression scores: no response, 0; growl or lip curl, 1; snap or bite, 2. Mean scores for responses to triggers were obtained during the 2-week pretreatment period (baseline) and during the first and second weeks, third and fourth weeks, and fifth and sixth weeks of treatment. At the end of the study, owners assigned a score designed to evaluate their overall perceived change in aggressiveness; this was referred to as the global score. RESULTS: Mean aggression scores decreased at the fifth and sixth week of treatment in both groups, compared with baseline scores. However, mean scores between groups were not different. Global scores, assigned by the owner, generally reflected changes in mean aggression scores. CONCLUSIONS AND CLINICAL RELEVANCE: Compared with placebo, clomipramine administered to dogs at the dosage recommended for treatment of separation anxiety did not reduce aggressiveness toward human family members.



Williams, N. G., and Borchelt, P.L. (2003). Full body restraint and rapid stimulus exposure as a treatment for dogs with defensive aggressive behavior: three case studies. International Journal of Comparative Psychology, 16, 226-236.


Yen, D.H. (1998). Learned Helplessness. http://www.noogenesis.com/malama/discouragement/helplessness.html


    * Introductory paragraph: In early 1965, Martin E. P. Seligman and his collegues, while studying the relationship between fear and learning, accidentally discovered an unexpected phenomenon while doing experiments on dogs using Pavlovian (classical conditioning). As you may observe in yourselves or a dog, when you are presented with food, you have a tendency to salivate. Pavlov discovered that if a ringing bell or tone is repeatedly paired with this presentation of food, the dog salivates. Later, all you have to do is ring the bell and the dog salivates. However, in Seligman's experiment, instead of pairing the tone with food, he paired it with a harmless shock, restraining the dog in a hammock during the learning phase. The idea, then, was that after the dog learned this, the dog would feel fear on the presentation of a tone, and would then run away or do some other behavior.



Young MS. (1982). Treatment of fear-induced aggression in dogs. Vet Clin North Am Small Anim Pract. 12 (4): 645-53.


Separation Anxiety in Dogs


Appleby, D., & Pluijmakers, J. (2004). Separation anxiety in dogs: The function of homeostasis in its development and treatment. Clinical Techniques in Small Animal Practice, 19(4), 205-215.


Beata, C. (2006). Anxiety versus fears versus phobias in adult dogs. Paper presented at the The North American Veterinary Conference - 2006.


Borchelt, P. L., & Voith, V. L. (1982). Diagnosis and treatment of separation-related behavior problems in dogs. Vet Clin North Am Small Anim Pract, 12(4), 625-635.


Burnum, J. F. (2000). The pet separation syndrome. Ann Intern Med, 133(4), 313-314.


Butchers, A. D. (2001). Assessing the efficacy of prescribing tricyclic antidepressants in conjunction with modifying behaviour therapy for the treatment of separation anxiety in dogs. Retrieved August 8, 2005, from http://vein.library.usyd.edu.au/links?Essays/2001/butchers.html


Cairns, R. B. (1966). Attachment behavior of mammals. Psychol Rev, 73(5), 409-426.


Cairns, R. B., & Werboff, J. (1967). Behavior development in the dog: an interspecific analysis. Science, 158(804), 1070-1072.


Casey, R. A. (1998). Use of clomipramine for separation anxiety in dogs. Vet Rec, 142(21), 587-588.


Davison, G. C. (1968). Systematic desensitization as a counter-conditioning process. J Abnorm Psychol, 73(2), 91-99.


Denenberg, V. H. (1964). Critical Periods, Stimulus Input, and Emotional Reactivity: a Theory of Infantile Stimulation. Psychol Rev, 71, 335-351.


Denenberg, V. H., & Morton, J. R. C. (1962). Effects of environmental complexity and social groupings upon modification of emotional behavior. Journal of Comparative and Physiological Psychology, 55(2), 242-246.


Eisenberger, N. I., Jarcho, J. M., Lieberman, M. D., & Naliboff, B. D. (2006). An experimental study of shared sensitivity to physical pain and social rejection. Pain, 126(1-3), 132-138.


Elliot, O., & Scott, J. P. (1961). The development of emotional distress reactions to separation, in puppies. J Genet Psychol, 99, 3-22.


Flannigan, G., & Dodman, N. H. (2001). Risk factors and behaviors associated with separation anxiety in dogs. J Am Vet Med Assoc, 219(4), 460-466.


    * Abstract: OBJECTIVES: To determine potential risk factors and behaviors associated with separation anxiety and develop a practical index to help in the diagnosis of separation anxiety in dogs. DESIGN: Case-control study. ANIMALS: 200 dogs with separation anxiety and 200 control dogs with other behavior problems. PROCEDURES: Medical records were reviewed for signalment, history of behavior problems, home environment, management, potentially associated behaviors, and concurrent problems. RESULTS: Dogs from a home with a single adult human were approximately 2.5 times as likely to have separation anxiety as dogs from multiple owner homes, and sexually intact dogs were a third as likely to have separation anxiety as neutered dogs. Several factors associated with hyperattachment to the owner were significantly associated with separation anxiety. Spoiling activities, sex of the dog, and the presence of other pets in the home were not associated with separation anxiety. CONCLUSIONS AND CLINICAL RELEVANCE: Results do not support the theory that early separation from the dam leads to future development of separation anxiety. Hyperattachment to the owner was significantly associated with separation anxiety; extreme following of the owner, departure cue anxiety, and excessive greeting may help clinicians distinguish between canine separation anxiety and other separation-related problems.



Gacsi, M., Topal, J., Miklosi, A., Doka, A., & Csanyi, V. (2001). Attachment behavior of adult dogs (Canis familiaris) living at rescue centers: forming new bonds. J Comp Psychol, 115(4), 423-431.


Gaultier, E., Bonnafous, L., Bougrat, L., Lafont, C., & Pageat, P. (2005). Comparison of the efficacy of a synthetic dog-appeasing pheromone with clomipramine for the treatment of separation-related disorders in dogs. Vet Rec, 156(17), 533-538.


    * Abstract: Sixty-seven dogs that showed signs of distress when separated from their owners (destructiveness, excessive vocalisation and house soiling) and hyperattachment were used in a randomised, blind trial to assess the potential value of a dog-appeasing pheromone in reducing the unacceptable behaviours. For ethical reasons, there was no placebo group and the effects of the pheromone were compared with the effects of clomipramine which is regularly used to treat this type of problem. The undesirable behaviours decreased in both groups, but the overall assessment by the owners indicated that there was no significant difference between the two treatments, although there were fewer undesirable events in the dogs treated with the pheromone, and the administration of the pheromone appeared to be more convenient.



Gewirtz, J. L. (1972). Attachment, dependence, and a distinction in terms of stimulus control. In (pp. 139-177). Washington: V. H. Winston & Sons.


Gewirtz, J. L. (1960s). A learning analysis of the effects of normal stimulation, privation and deprivation on the acquisition of social motivation and attachment. In (pp. 213-303). London: Methuen.


Gazzano, A., Mariti, C., Notari, L., Sighiere, C., & McBride, E. A. (2007). Effects od early gentling and early environment on emotional development of puppies. Applied Animal Behaviour Science, xxx(xxx), xxx-xxx. in press.


Hewson, C. J. (2000). Clomipramine and behavioural therapy in the treatment of separation-related problems in dogs. Vet Rec, 146(4), 111-112.


Hewson, C. J., Luescher, U. A., & Ball, R. O. (1999). The use of chance-crrected agreement to diagnose canine compulsive disorder: An approach to behavioral disgnosis in the absence of a 'gold standard'. Can J Vet Res, 63, 201-206.


Horwitz, D. F. (2000). Diagnosis and treatment of canine separation anxiety and the use of clomipramine hydrochloride (clomicalm). Journal of the American Animal Hospital Association, 36(2), 107-109.


Horwitz, D. F. (2001). Separation anxiety in dogs. Retrieved February 4, 2005, from http://www.vin.com/ACVC/2001/AuthorIndex.htm


    * Introductory paragraph: Because domestic dogs usually consider the human family to be their social group, they become bonded to family members. When separated from family members dogs may experience distress and engage in problem behaviors related to the anxiety of separation. These behaviors include destruction, vocalization, elimination of urine and/or stool, anorexia, drooling, attempts at escape and/or behavioral depression. Treatment protocols include independence training, habituation, counter-conditioning and desensitization to owner departure and absence.



King, J. N., Simpson, B. S., Overall, K. L., Appleby, D., Pageat, P., Ross, C., Chaurand, J. P., Heath, S., Beata, C., Weiss, A. B., Muller, G., Paris, T., Bataille, B. G., Parker, J., Petit, S., and Wren, J. (2000). Treatment of separation anxiety in dogs with clomipramine: results from a prospective, randomized, double-blind, placebo-controlled, parallel-group, multicenter clinical trial. Applied Animal Behaviour Science, 67(4), 255-275.


    * Abstract: The efficacy and tolerability of clomipramine in the treatment of separation anxiety in dogs was tested in a prospective, randomized, double-blind, placebo-controlled, parallel-group, international multicenter clinical trial. For a diagnosis of separation anxiety, dogs had to exhibit at least one of the following signs in the absence of their owner: destruction, defecation, urination and/or vocalization, as well as the behaviour suggestive of "hyper-attachment" to their owner. A total of 95 dogs were randomized to receive one of the three treatments for 2-3 months: "standard-dose" clomipramine (1 to <2 mg/kg, PO, q. 12 h); "low-dose" clomipramine (0.5 to <1 mg/kg, PO, q. 12 h); and placebo (PO, q. 12 h). All dogs received behavioural therapy. Dogs were examined at four time points (days 0, 28, 56 and 84) after the initiation of therapy. Improvement in each dog's behaviour at days 28, 56 and 84 was evaluated in comparison to its behaviour at day 0.The results showed that, compared to placebo, dogs receiving standard-dose clomipramine were rated improved at least three times faster for the signs destruction, defecation and urination. At most time points, more dogs in the standard-dose clomipramine group were rated improved for the signs destruction, defecation and urination, and in an owner's global assessment of the dog's overall behaviour (p<0.05 at certain time points). However, there were no statistically significant differences at any time point between the standard dose and the placebo groups in the sign vocalization. The low-dose clomipramine group produced no statistically significant effect when compared with placebo. Mild and transient vomiting was noted as a side effect of clomipramine in a small number of dogs.It is concluded that addition of standard-dose (1 to <2 mg/kg, PO, q. 12 h) clomipramine to conventional behavioural therapy for 2-3 months ameliorated the signs of separation anxiety in dogs.



King, J. N. (2000). Pharmacological management of separation anxiety. In: Recent Advances in Companion Animal Behavior Problems, Houpt K.A. (Ed.)


International Veterinary Information Service, Ithaca NY, 2000. Retrieved February 4, 2005, from [available at http://www.ivis.org/advances/Behavior_Houpt/king/ivis.pdf]


    * Summary: The treatment of separation anxiety in dogs involves the combination of behaviour modification therapy, envienvironmental management and pharmacotherapy. Tricyclic antidepressants with strong serotonin-reuptake inhibiting properties appear to be the first choice for drug therapy, with additional administration of benzodiazepines in severe cases. With proper management, cases of separation anxiety carry a good prognosis.



Landsberg, G. M. (2001). Clomipramine--beyond separation anxiety. J Am Anim Hosp Assoc, 37(4), 313-318.


Lem, M. (2002). Behavior modification and pharmacotherapy for separation anxiety in a 2-year-old pointer cross. Canadian Veterinary Journal, 43, 220-222. [available at http://www.pubmedcentral.nih.gov/picrender.fcgi?artid=339210&blobtype=pdf]


    * Abstract: Separation anxiety is a common behavioral problem in dogs. Treatment is based on developing a behavior modification protocol that gradually desensitizes and counter-conditions the dog to being left alone, by rewarding calm, relaxed behavior. Judicious use of pharmacotherapy can be a useful adjunct to a behavior modification program.



Lund, J. D., & Jorgensen, M. C. (1999). Behaviour patterns and time course of activity in dogs with separation problems. Applied Animal Behaviour Science, 63(3), 219-236.


McCrave, E. A. (1991). Diagnostic criteria for separation anxiety in dogs. Veterinary Clinics of North America: Small Animal Practice, 21(2), 247-255.


McGreevy, P. D., & Masters, A. M. (2007). Risk factors for separation-related distress and feed-releated aggression in dogs: Additional findings from a survey of Australian dog owners. Applied Animal Behaviour Science, xxx(xxx), xxx-xxx. in press.


McMillan, F. D. (2001). More thoughts on separation anxiety. J Am Vet Med Assoc, 219(12), 1674-1675.


Newberry, R. C., & Swanson, J. C. (2007). Implications of breaking mother-young social bonds. Applied Animal Behaviour Science, xxx(xxx), xxx-xxx. in press.


O'Heare, J. (2003). The Canine Separation Anxiety Workbook (4th ed.). Ottawa: Dogpsych Publishing.


Overall, K. L. (2001). Screen for separation anxiety and noise phobias in dogs. Retrieved February 4, 2005, from http://www.vin.com/ACVC/2001/AuthorIndex.htm


    * Introduction: The first set of these questions deals with an "actual absence" - e.g., the client actually leaves the house and the dog is either alone or totally without the client. The second set deals with "virtual absence" - e.g., the client is home, but not accessible because the door is closed or the dog is barricaded in another room. The questions are the same for each but please answer both.



Overall, K. L., & Dunham, A. E. (2001). Disagrees with alternative view of separation anxiety. J Am Vet Med Assoc, 219(11), 1520, 1522.


Overall, K. L. (1998). Animal behavior case of the month. Stereotypical motor behavior that manifested when the owner departed or was out of the dog's sight. J Am Vet Med Assoc, 213(1), 34-36.


Overall, K. (n/d). Karen Overal's Behavior Modification Program Protocol for Relaxation. http://home.gci.net/~divs/behavior/bemod_relax.html


Overall. K. L. (2003). Separation anxiety: Not all dogs crated or kenneled successfully. http://www.dvmnewsmagazine.com/dvm/article/articleDetail.jsp?id=62241


Overall. K. L. (2003). Treating anxiety is different than 'managing' the problem. http://www.dvmnewsmagazine.com/dvm/article/articleDetail.jsp?id=43492


Overall. K. L. (2003). Separation anxiety: Early drug intervention can be beneficial. http://www.dvmnewsmagazine.com/dvm/article/articleDetail.jsp?id=58642


Overall, K. L., Hamilton, S. P., & Chang, M. L. (2006). Understanding the genetic basis of canine anxiety: Phenotyping dogs for behavioral, neurochemical, and genetic assessment. Journal of Veterinary Behavior, 1, 124-141.


Panksepp, J., Herman, B., Conner, R., Bishop, P., & Scott, J. P. (1978). The biology of social attachments: opiates alleviate separation distress. Biol Psychiatry, 13(5), 607-618.


    * Abstract: The possibility that brain opiate systems participate in the control of social affect was assessed by determining capacity of low doses of exogenous opiates (0.125-0.50 mg/kg oxymorphone, and 0.10-0.50 mg/kg morphine sulfate) to reduce distress vocalizations of socially isolated puppies. Low doses of opiates were capable of profoundly reducing crying as well as the motor agitation they exhibit during brief periods of social isolation. Since reductions in crying could be obtained with morphine in the absence of any gross behavioral disturbances, the possibility is entertained that brain opiates may function to control the intensity of emotions arising from social separation. Possible parallels between the biological nature of narcotic addiction and the formation of social bonds are discussed.


Papurt, M. L. (2001). An alternative look at separation anxiety. J Am Vet Med Assoc, 219(7), 910.


Pettijohn, T. F., Wong, T. W., Ebert, & Scott, J. P. (1977). Alleviation of separation distress in 3 breeds of young dogs. Developmental Psychobiology, 10(4), 373-381.


    * Abstract: Twenty-four puppies (8 each of Shetland sheepdogs, Telomians, and beagles) served as subjects in this experiment which was designed to examine the effectiveness of 12 stimulus conditions (food, toys, canine contact, and human contact) on alleviation of separation-induced distress vocalization. Testing consisted of a prestimulus trial, stimulus trial, and post-stimulus trial each session 3 times a week from 4 to 8 weeks after birth. Overall results showed human contact to be the most effective, followed by canine contact, toys, and food. Breed differences were significant in only 2 of the stimulus conditions. In the typical case of effective treatment, the vocalization rate declined from the prestimulus trial to a low point in the stimulus trial and then increased again in the post-stimulus trial.



Podberscek, A. L., Hsu, Y., Serpell, J. A. (1999). Evaluation of clomipramine as an adjunct to behavioural therapy in the treatment of separation-related problems in dogs. The Veterinary Record, 145(13), 365-369.


    * Abstract: Forty-nine dogs showing signs of separation-related problems were randomly assigned to one of three groups: group A (15 dogs) received a placebo twice daily; group B (17 dogs) received clomipramine at 0.5 to 1.0 mg/kg twice daily; and group C (17 dogs) received clomipramine at 1.0 to 2.0 mg/kg twice daily. All the dogs also received behavioural therapy. Their owners were required to complete questionnaires about their dog's behaviour initially, and one, four and eight weeks after the treatment with clomipramine began. Bipolar ratings scales were used to monitor the frequencies of 'general', 'attachment-related' and 'separation-related' behaviours. Kruskal-Wallis tests and Kendall Rank correlations were used to determine any initial differences between the treatment groups, and the association between the initial scores and behavioural changes after one week of treatment with clomipramine. Extended Mantel-Haenszel statistics were used to evaluate the effects of clomipramine treatment versus the placebo, and Page's test was used to assess the effectiveness of behavioural therapy on its own. There were no significant differences in the demographic characteristics of the owners of the dogs assigned to the three groups. The dogs differed slightly in age between groups, and the dogs in the two clomipramine-treated groups were reported as showing problems at a significantly earlier age than those in the placebo group. Clomipramine treatment had a sustained suppressive effect on the dogs' general activity levels, and a more modest suppressive effect on their attachment-related tendency to want much physical contact with their owners. The typical signs of separation-related behaviour problems were not significantly affected by treatment with clomipramine, but behavioural therapy on its own was highly effective in reducing behavioural problems.



Roen, D. T. (2000). Thoughts on separation anxiety in dogs. J Am Vet Med Assoc, 217(6), 818.


Schroll, S., Dehasse, J., Palme, R., Sommerfeld-Stur, I., & Löwenstein, G. (n/d). The use of DAP collar to reduce stress during training of police dogs A preliminary study. Retrieved June 4, 2006, from http://www.vet-magazin.com/.../DAP-collar.html


Schwartz, S. (2003). Separation anxiety syndrome in dogs and cats. JAVMA, 222(11), 1526-1532.


Seksel, K., and Lindeman, M. J. (2001). Use of clomipramine in treatment of obsessive-compulsive disorder, separation anxiety and noise phobia in dogs: a preliminary, clinical study. Autralian Veterinary Journal, 79(4), 252-256.


    * Abstract: OBJECTIVE: To evaluate the efficacy and tolerance of a treatment protocol for obsessive-compulsive disorder, separation anxiety and noise phobia in dogs. DESIGN: A study was undertaken to assess clinical responses in 24 dogs diagnosed with one or more of three behavioural disorders stated above to a treatment regimen that included clomipramine and behaviour modification. PROCEDURE: A detailed behavioural and clinical history was obtained for each dog. Obsessive-compulsive disorder was diagnosed in nine cases: primary presenting complaints were tail-chasing, shadow-chasing, circling and chewing; one case was diagnosed with concurrent separation anxiety. Separation anxiety was diagnosed in 14 cases: presenting complaints included destruction, vocalisation and escaping in the absence of the owner; four cases also exhibited noise phobia. The study also included one dog diagnosed with noise phobia only and another with inappropriate fear responses. Clomipramine was administered orally twice daily. The starting dose was 1 to 2 mg/kg bodyweight. The dose was increased incrementally to a maximum of 4 mg/kg if needed. A behaviour modification program was designed and the owner instructed on its implementation. Dogs continued medication for at least 1 month after clinical signs disappeared or were acceptably reduced, then withdrawal of medication was attempted by decreasing drug dosage at weekly intervals while behaviour modification continued. RESULTS: The presenting clinical sign was largely improved or disappeared in 16 dogs, 5 demonstrated slight to moderate improvement and the behaviour was unchanged in 3. Clomipramine withdrawal was attempted in nine cases: this was successful in five. CONCLUSION: Clomipramine was effective and well-tolerated in controlling signs of obsessive-compulsive disorder and/or separation anxiety and/or noise phobia in 16 of 24 assessable cases, when used in combination with behaviour modification, and improvement in clinical signs was noted in 5 others.



Shore, E. R., Douglas, D. K., & Riley, M. L. (2005). What's in it for the companion animal? Pet attachment and college students' behaviors toward pets. J Appl Anim Welf Sci, 8(1), 1-11.


Slabbert, J. M., & Rasa, O. A. (1993). The effect of early separation from the mother on pups in bonding to humans and pup health. J S Afr Vet Assoc, 64(1), 4-8.


Takeuchi, Y., Houpt, K. A., & Scarlett, J. M. (2000). Evaluation of treatments for separation anxiety in dogs. J Am Vet Med Assoc, 217(3), 342-345.


    * Abstract: Objective—To evaluate treatment outcome in dogs with separation anxiety and owner compliance with and perception of effectiveness of discharge instructions. Design—Cohort study. Animals—52 dogs with separation anxiety. Procedure—Sex, age at which the owner obtained the dog, age at which separation anxiety was first noticed, age at behavioral examination, and discharge instructions were obtained from medical records of each dog. Between 6 and 64 months after the behavioral examination, owners were contacted by telephone and questioned about the outcome of treatment, their compliance with discharge instructions, and their perception of the effectiveness of each instruction. Results—Thirty-two (62%) dogs had improved, whereas 20 were the same, were worse, or had been euthanatized or given away. Mixed-breed dogs were significantly less likely to improve than purebred dogs. Compliance varied according to discharge instruction. Significantly fewer dogs with owners that were given > 5 instructions improved or were cured, compared with those with owners given fewer instructions. Twenty-seven dogs were also treated with amitriptyline or other medication; 15 (56%) improved. Conclusions and Clinical Relevance—Owners complied with instructions that involved little time such as omitting punishment and providing a chew toy at the time of departure. Owners were also willing to increase the dog's exercise but were not willing to uncouple the cues of departure from real departures or desensitize the dog to impending departure. Administration of psychoactive medication may be necessary to augment behavior modification techniques designed to reduce separation anxiety in dogs



Topal, J., Miklosi, A., Csanyi, V., & Doka, A. (1998). Attachment behavior in dogs (Canis familiaris): a new application of Ainsworth's (1969) Strange Situation Test. J Comp Psychol, 112(3), 219-229.


    * Abstract: Fifty-one owner-dog pairs were observed in a modified version of M. D. S. Ainsworth's (1969) Strange Situation Test. The results demonstrate that adult dogs (Canis familiaris) show patterns of attachment behavior toward the owner. Although there was considerable variability in dogs' attachment behavior to humans, the authors did not find any effect of gender, age, living conditions, or breed on most of the behavioral variables. The human-dog relationship was described by means of a factor analysis in a 3-dimensional factor space: Anxiety, Acceptance, and Attachment. A cluster analysis revealed 5 substantially different classes of dogs, and dogs could be categorized along the secure-insecure attached dimensions of Ainsworth's original test. A dog's relationship to humans is analogous to child-parent and chimpanzee-human attachment behavior because the observed behavioral phenomena and the classification are similar to those described in mother-infant interactions.



Tuber, D. S., Sanders, S., Hennessy, M. B., & Miller, J. A. (1996). Behavioral and glucocorticoid responses of adult domestic dogs (Canis familiaris) to companionship and social separation. J Comp Psychol, 110(1), 103-108.


Dog Stress


? (n/d). General Adaptation Syndrome [Electronic Version]. Retrieved September 2, 2005 from http://library.thinkquest.org/C0123421/gas.htm.


Ahrens, F., Knies, K., Schneider, M., Kohler, F., & Erhard, M. H. (2005). Influence of different training and outdoor conditions on plasma histamine and cortisol concentrations in search-and-rescue dogs. Inflamm Res, 54 Suppl 1, S34-35.


al'Absi, M., Hugdahl, K., & Lovallo, W. R. (2002). Adrenocortical stress responses and altered working memory performance. Psychophysiology, 39(1), 95-99.


Beata, C. (2006). Anxiety versus fears versus phobias in adult dogs. Paper presented at the The North American Veterinary Conference - 2006.


Beera, B., Schilder, M.B.H., van Hoof, J.A.R.A.M., and de Vries, H.W. (1997). Manifestations of chronic and acute stress in dogs. Applied Animal Behaviour Science, 52, 307-319.


    * Abstract: Poor housing conditions, harsh training sessions and uncontrollable or unpredictable social environments are examples of the situations that may lead to reduced welfare status in dogs. Individuals that suffer from poor welfare presumably experience stress and may consequently exhibit stress responses. In order to evaluate stress responses as potential indicators of poor welfare in dogs, we review studies dealing with dogs subjected to stressors. The reported stress responses are categorized as being behavioural, physiological or immunological, and demonstrate the various ways stress is manifested in the dog. Stressors such as noise, immobilization, training, novelty, transport or restricted housing conditions have been reported to elicit responses in behavioural, cardiovascular, endocrine, renal, gastro-intestinal, and haematological parameters. These and other parameters that change during stress may thus be indicative of poor welfare. However, several sources of misinterpretation have to be considered before stress responses may be used as valid indicators of welfare. Although analogous to the human situation, especially chronic stress may impair welfare, most studies deal with acute stress and do not address chronic stress and related phenomena. Adaptation may counteract the initial stress response and render parameters of acute stress useless for assessing chronic stress. Adaptations to stress are thus in themselves indicative of reduced welfare. Such adaptations may be discovered by challenging a stress responsive system. Additional studies are recommended to investigate acute stress parameters as possible indicators of chronic stress. Differences in stressor properties and in individual characteristics of dogs introduce variability in stress responses. Such variability will complicate a valid interpretation of stress responses with regard to welfare. Obtaining and applying fundamental knowledge of stress responses in dogs and measuring more than one stress parameter are proposed to minimize the risk of misinterpreting measurements of stress.



Beerda, B., Schilder, M. B. H., Van Hoof, J. A. R. A. M., De Vries, H., and Mol, J. A. (1999). Chronic stress in dogs subjected to social and spatial restriction. I. Behavioral responses. Physiology & Behavior, 66(2), 233-242.


    * Abstract: Six weeks of social and spatial restriction were used as a model to induce chronic stress in Beagles. Behavioral and physiological measurements were performed during a period of enriched spacious outdoor housing in groups (GH) and during a subsequent period of solitary housing in small indoor kennels (IH). Behavioral parameters that may indicate chronic stress in dogs are reported. During IH, the dogs showed significantly (comparison-wise error rate <0.05) lower postures than during GH. IH induced enduring increments in frequencies of autogrooming, paw lifting, and vocalizing, and was associated with incidents of coprophagy and repetitive behavior. So far, we interpret the behavioral changes as signs of chronic stress. Relatively low levels of walking, digging, intentions to change from one state of locomotion to another, and increments in circling are conceived as obvious adaptations to the specific features of the IH system. By challenging the dogs outside their home kennel we tested whether the dogs' coping abilities were affected by IH. Dogs that were challenged were introduced into a novel environment, given the opportunity to escape from their home kennel, restrained, walked down an unfamiliar corridor, presented a novel object, exposed to loud noise, given food, or confronted with a conspecific. During IH, challenged dogs exhibited higher postures, showed more tail wagging, nosing, circling, urinating, and defecating, and changed more often from one state of locomotion (or posture) to another than during GH. These behavioral changes were observed across the different types of challenges, with the exception of the noise administration test. In the presence of conspecifics, the socially and spatially restricted male dogs behaved more dominantly and aggressive than during the time that they were kept in groups. Such behavior manifested as increased performances of raised hairs, growling, paw laying, and standing over. Both sexes showed increases in paw lifting, body shaking, ambivalent postures, intentions to change from one state of locomotion to another, and trembling in any of the challenges, excluding the walking down the corridor test. In short, during a variety of challenges, socially and spatially restricted dogs exhibited a heightened state of aggression, excitement, and uncertainty. Behavioral differences between dogs that had experienced pleasant and bad weather conditions during GH, suggested that "pleasant-weather individuals" had experienced early stress during the control period, and, as a result, responded to the subsequent period of IH differently. Regardless of the housing conditions, challenged bitches showed stronger indications of acute stress than male dogs. Gender did not affect the chronic stress responses to social and spatial restriction. A low posture and increased auto-grooming, paw lifting, vocalizing, repetitive behavior, and coprophagy may indicate chronic stress in dogs, and as such, can help to identify poor welfare. When challenged, chronically stressed dogs may show increased excitement, aggression, and uncertainty, but the nonspecificity of such emotional behavior will complicate its practical use with regard to the assessment of stress.



Blackshaw, J. (2000). Chronic stress in housed dogs. Retrieved August 8, 2005, from http://vein.library.usyd.edu.au/links/Essays/2000blackshaw.html


    * Introduction: Housing of production animals has long been considered an animal welfare issue. In the United Kingdom the concept of the 'Five Freedoms' has raised the standard of animal welfare for production animals. Has the time come to apply the same framework to companion animals?



Boxall, J., Heath, S., Bate, S., & Brautigam, J. (2004). Modern concepts of socialisation for dogs: implications for their behaviour, welfare and use in scientific procedures. Altern Lab Anim, 32 Suppl 2, 81-93.


Brown, J. D. (1991). Staying fit and staying well: Physical fitness as a moderator of life stress. Journal of

Personality and Social Psychology, 60(4), 555-561.


    * Abstract: Previous research suggests that physical fitness moderates the adverse effects of stressful life events. However, a reliance on self-reports of fitness and health may limit the validity of prior investigations. The present research tested the stress-buffering effect of fitness with subjective and objective indicators of exercise, fitness, and physical well-being. For self-reports of health, both self-reports of exercise and objective measures of fitness showed the buffering effect; however, only objective fitness levels buffered stress when visits to a health facility were considered. Additional evidence indicated that this effect was largely independent of measures of psychological distress. Implications for understanding the link between fitness, stress, and health status are discussed.



Clark, J. D., rager, D.R., Crowell-Davis, S., and Evans, D.L. (1997). Housing and exercise of dogs: Effects on behavior, immune function, and cortisol concentration. Laboratory Animal Science, 47(5), 500-510.


Cook, C. J. (2002). Glucocorticoid feedback increases the sensitivity of the limbic system to stress. Physiol Behav, 75(4), 455-464.


Dreschel, N. A., & Granger, D. A. (2005). Physiological and behavioral reactivity to stress in thunderstorm-phobic dogs and their caregivers. Applied Animal Behaviour Science.


Drugan, R. C., Basile, A. S., Ha, J. H., Healy, D., & Ferland, R. J. (1997). Analysis of the importance of controllable versus uncontrollable stress on subsequent behavioral and physiological functioning. Brain Res Brain Res Protoc, 2(1), 69-74.


Eisenberger, N. I., Jarcho, J. M., Lieberman, M. D., & Naliboff, B. D. (2006). An experimental study of shared sensitivity to physical pain and social rejection. Pain, 126(1-3), 132-138.


Garnier, F., Benoit, E., Virat, M., Ochoa, R., and Delatour, P. (1990). Adrenal cortical response in clinically normal dogs before and after adaptation to a housing environment. Laboratory Animals, 24, 40-43.


    * Abstract: 58 dogs (29 males and 29 females) selected as healthy on clinical and biochemical evaluations were subjected to an ACTH adrenal function test 2 days after their admission to a veterinary hospital (t + 0). Basal female serum cortisol concentrations were significantly higher than concentrations in males (77 nmol/l versus 43 nmol/l; P less than 0.01). Concentrations post stimulation were not statistically different (P greater than 0.05) between males and females: 306 (+/- 69) nmol/l versus 291 (+/- 73) nmol/l, respectively. Twelve dogs (6 males and 6 females), randomly selected from the 58, were subjected to the same test 5 weeks later (t + 5) and 12 weeks later (t + 12). Basal cortisol concentrations were lower at t + 5 or at t + 12 than at t + 0. Post stimulation mean cortisol concentrations were lower in males than in females at t + 5 (162 versus 232 nmol/l; P less than 0.05) but not at t + 0 (262 versus 320 nmol/l; P greater than 0.05) and t + 12 (188 versus 233 nmol/l; P greater than 0.05). These findings are indicating an increased susceptibility of bitches to environmental stress.



Hennessy, M. B., Davis, H. N., Williams, M. T., Mellott, C., & Douglas, C. W. (1997). Plasma cortisol levels of dogs at a county animal shelter. Physiol Behav, 62(3), 485-490.


    * Abstract: Plasmacortisol levels were examined to assess the stress of dogs in a county animal shelter. Groups of dogs confined in the shelter for their 1st, 2nd, or 3rd day had higher cortisol levels than did a group maintained in the shelter for more than 9 days. Dogs in the shelter for an intermediate period (Day 4-9) had intermediate levels of cortisol. The cortisol concentrations of dogs during their first day in the shelter were greater than either those of the same dogs on Day 4/5 in the shelter or those of a group of pet dogs sampled in their own homes. There was no overall effect of 20 min of social interaction with a human (e.g., petting) on the plasma cortisol levels of dogs in the shelter on Day 1-3. However, the gender of the petter did affect cortisol levels. Those dogs interacting with a female had lower cortisol concentrations at the end of the session than did dogs interacting with a male. The results suggest that confinement in a public animal shelter produces a prolonged activation of the hypothalamic-pituitary-adrenal axis. Further, it appears that some subtle aspect of interaction with a human may be capable of moderating this response. Possible implications for the welfare of confined dogs, and for the development of behavior problems in dogs obtained from shelters, are discussed.


Hennessy, M. B., Voith, V. L., Mazzei, S. J., Buttram, J., Miller, D. D., & Linden, F. (2001). Behavior and cortisol levels of dogs in a public animal shelter, and an exploration of the ability of these measures to predict problem behavior after adoption. Appl Anim Behav Sci, 73(3), 217-233.


Hydbring-Sandberg, E., von Walter, L. W., Hoglund, K., Svartberg, K., Swenson, L., & Forkman, B. (2004). Physiological reactions to fear provocation in dogs. J Endocrinol, 180(3), 439-448.


    * Abstract: Fear is a common behavioral problem in dogs. In this paper, we studied the association between behavioral and physiological responses in two potentially fear-eliciting situations. The aim was to establish whether it is possible to separate dogs of the collie breed that are fearful of floors and gunshots from those that are not by studying changes in heart rate and hematocrit, plasma cortisol, progesterone, testosterone, vasopressin, and beta-endorphin concentrations. Thirteen privately owned male dogs of the collie breed were studied during a floor test, using different types of floors, and a subsequent gunshot test. Seven of the dogs were identified as being fearful of floors and six were declared as fearless. Out of the 13 dogs, seven were fearful of gunshots and six were fearless of gunshots. Since fear of floors did not always occur concomitantly with fear of gunshots, there were consequently four different groups of dogs. The heart rate increased during the floor test in all groups, but dogs that were fearful of floors had higher heart rates than dogs that were fearless of floors. Dogs that were fearful of gunshots had higher heart rates, higher hematocrit levels and higher plasma concentrations of cortisol, progesterone, vasopressin, and beta-endorphins during the gunshot test than did dogs that were found to be fearless of gunshots. Plasma cortisol and progesterone increased drastically during the gunshot test in dogs identified as being fearful of gunshots. In fearful dogs, the testosterone concentration increased after completion of the floor test and before the gunshot test started, but there were no significant differences in testosterone between the groups. Since dogs fearful of gunshots had increased levels of several physiological parameters, the results demonstrated that this fear is a serious stress for the individual, a fear which it is possible to register with physiological variables.



Knies, K., Erhard, M. H., & Ahrens, F. (2005). Effect of moderate stress on plasma histamine concentration in laboratory dogs. Inflamm Res, 54 Suppl 1, S32-33.


Kobelt, A. J., Hemsworth, P. H., Barnett, J. L., & Butler, K. L. (2003). Sources of sampling variation in saliva cortisol in dogs. Res Vet Sci, 75(2), 157-161.


    * Abstract: The main advantage of collecting saliva cortisol as opposed to plasma cortisol is that it is non-invasive and therefore it is now widely used in stress measurement studies on farm animals and dogs. Although a plasma cortisol response to handling associated with blood collection generally occurs at 3 min from the commencement of handling, there is no information in the literature on the time course of the response of salivary cortisol concentration to handling. The aims of these experiments were to (1). determine if there is a response to up to 4 min handling that affects cortisol concentration in saliva and (2). determine the main causes of variation in saliva cortisol in dogs over time. In experiment 1, saliva was collected from six Kelpies at 0 min then 2, 3 or 4 min after the commencement of restraint. There was no handling effect found in up to 4 min sampling time. In experiment 2, saliva was collected from six Labrador Retrievers five times in 2 h (14:00-16:00), three days a week for four weeks. Some of the sources of variation in saliva cortisol over time included between dog variation that varied over a period of days and variation between occasions that affected the group of dogs as a whole.



Lund, J. D., & Jorgensen, M. C. (1999). Behaviour patterns and time course of activity in dogs with separation problems. Applied Animal Behaviour Science, 63(3), 219-236.


    * Abstract: An analysis of video-recordings of 20 dogs with separation problems suggested that separation behaviour may be divided into: (1) exploratory behaviour, (2) object play including elements of predatory behaviour, (3) destructive behaviour, and (4) vocalization. Elimination behaviour reported by other authors was found in one case only. Separation behaviour was related to the level of arousal. A clear distinction between ‘destructive' dogs and ‘howlers' was not justified. Object play seemed to be closely related to destructive behaviour. A model for the time course of activity from the owner's departure was developed. The model includes two components: (1) a cyclic component having a period of 23–28 min and controlled by internal factors, and (2) a long-term exponential decrease, which may be influenced by external factors arousing the dog. The results supported the view that separation problems are caused by frustration related to the dependency on the owner, whereas they are not caused by disobedience or boredom. The frustration in turn may lead to arousal, increased fear and the disinhibition of play or predatory behaviour and leading to destructive behaviour. The results also indicated that barking was caused by arousal, whereas howling and whining may reflect the presence of fear.



Nesse, R. M., and Young, E.A. (2000). Evolutionary orginins and functions of the stress response [Electronic Version]. Encyclopedia of Stress, 2, 79-84. Retrieved August 31, 2005 from http://www-personal.umich.edu/nesse/Articles/Stress&Evolution-2000.pdf.


Overall, K. L. (2001). How to deal with anxiety and distress responses: dogs. Retrieved February 4, 2005, from http://www.vin.com/ACVC/2001/AuthorIndex.htm


Overall, K. L., Hamilton, S. P., & Chang, M. L. (2006). Understanding the genetic basis of canine anxiety: Phenotyping dogs for behavioral, neurochemical, and genetic assessment. Journal of Veterinary Behavior, 1, 124-141.


Rooney, N. J., Gaines, S. A., & Bradshaw, J. W. (2007). Behavioural and glucocorticoid responses of dogs (Canis familiaris) to kennelling: Investigating mitigation of stress by prior habituation. Physiol Behav.


Rushen, J. (2000). Some issues in the interpretation of behavioural responses to stress. In G. P. Moberg, and Mench, J.A. (Ed.), The Biology of Stress Basic Principles and Implications for Animal Welfare (pp. 23-42). Davis: CABI Publishing.


Schroll, S., Dehasse, J., Palme, R., Sommerfeld-Stur, I., & Löwenstein, G. (n/d). The use of DAP collar to reduce stress during training of police dogs A preliminary study. Retrieved June 4, 2006, from http://www.vet-magazin.com/.../DAP-collar.html


Tinbergen, N. (1974). Ethology and stress diseases. Science, 185, 20-27.


Tod, E., Brander, D., & Waran, N. (2005). Efficacy of dog appeasing pheromone in reducing stress and fear related behaviour in shelter dogs. Applied Animal Behaviour Science, 93, 295-308.


Wikipedia.org. (n/d). Stress (medicine). Retrieved September 6, 2005, from http://en.wikipedia.org/wiki/General_adaptation_syndrome


Wikipedia.org. (n/d). Fight-or-flight response. Retrieved September 6, 2005, from http://en.wikipedia.org/wiki/Fight-or-flight_response


Wilson, S. D., and Keddy, Paul A. (1986). Species competitive ability and position along a natural stress/disturbance gradient. Ecology, 67(5), 1236-1242.


    * Abstract: We tested the prediction that plant species that grow in undisturbed, nutrient—rich habitats tend to have higher competitive abilities than those found in disturbed or nutrient—poor habitats. The distributions of seven species (Eriocaulon septangulare, Rhynchospora fusca, Hypericum ellipticum, Juncus pelocarpus, Lysimachia terrestris, Dulichium arundinaceum, and Drosera intermedia) were measured along a gradient of exposure to wave action on the shore of Axe Lake, Ontario. The exposure gradient incorporates disturbance, through the removal of plant biomass, and stress, through the creation of a gradient in sediment organic content, nutrient concentrations, and fine particle sizes. Species distributions on the exposure gradient were quantified by determining the mean sediment organic content of the quadrats containing each species. Competitive abilities were measured as relative increase in dry mass per plant, in a field experiment in which species were grown together in all pairwise combinations (N = 10 replicates). Species had significantly heterogeneous competitive abilities (P < .01). Species found on exposed, nutrient—poor shores (e.g., E. septangulare) had low competitive abilities, while those growing on sheltered, nutrient—rich shores (e.g., D. arundinaceum) had high competitive abilities. Competitive ability was significantly correlated with mean position on the exposure gradient.


Dog Social Dominance


(this section in particular has many articles not related directly to dogs, although understanding social dominance really means understanding it in many species. I will leave many dog dog citations in for those who wish to access the literature for other species as well). I will focus on providing abstracts for those related most to dogs.


Abernethy, V. (1981). Dominance, feminist hierarchies, and heterosexual dyads (commentary on Berntein (1981)). Behavioral and Brain Sciences, 4, 429-430.


Adams, D. B. (1979). Brain mechanisms for offense, defense, and submission. The Behavioral and Brain Sciences, 2, 201-241.


Allee, W. C. (1942). Social dominance and subordination among vertebrates. Biological Symposia, 8, 139-162.


Altman, S. A. (1981). Doiminance relationships: The Cheshire cat's grin (commentary on Berntein (1981)). Behavioral and Brain Sciences, 4, 430-431.


American_Veterinary_Society_of_Animal_Behavior. (2007). Guidelines on the Use of Punishment for Dealing with Behavior Problems in Animals [Electronic Version]. AVSAB. Retrieved November 24, 2007 from http://www.avsabonline.org/avsabonline/images/stories/punishment%20guidelines-aversives%20effects-definitions.pdf.


Appleby, M. C. (1979). The probability of linearity in hierarchies. The behavioral and brain sciences, 2, 201-241.


Aureli, F., Cords, Marina, and Van Schaik, Carel P. (2002). Conflict resolution following aggression in gregarious animals: a predictive framework. Animal Behaviour, 64, 325-343.


Baenniger, R. (1981). Dominance: On distinguishing the baby from the bathwater (commentary on Berntein (1981)). Behavioral and Brain Sciences, 4, 431-432.


Banks, E. M. (1981). Dominance and behavioral primatologist: A case of typological thinking? (commentary on Berntein (1981)). Behavioral and Brain Sciences, 4, 432-433.


Barrette, C. (1993). The 'inheritance of dominance', or of an aptitude to dominate. Animal Behaviour, 46, 591-593.


Barrette, C., & Vandal, D. (1986). Social rank, dominance, antler size, and access to food in snow-bound wild woodland caribou. Behaviour, 97(118-146).


Beaugrand, J. P. (1997). Relative importance of initial individual differences, agonistic experience, and assessment accuracy during hierarchy formation: a simulation study. Behavioral Processes, 41, 177-192.


Beaugrand, J. P., & Goulet, C. (2000). Distinguishing kinds of prior dominance and subordination experiences in male Green swordtail fish (Xiphophorus helleri). Behavioural Processes, 50, 131-142.


Begin, J., Beaugrand, P. J., & Zayan, R. (1996). Selecting dominants and subordinates at conflict outcome can confound the effects of prior dominance or subordination experience. Behavioral Processes, 36, 219-226.


Bekoff, M. (1995). Play signals as punctuation: the structure of social play in canids. Behaviour, 132, 419-429.


Bekoff, M. (downloaded 12/02/03). Dominance in Animal Social Groups. MIT Encyclopedia of Cognitive Science at http://ai.ato.ms/MITECS/Entry/bekoff1.


Bernstein, I. S. (1981). Dominance: the baby and the bath water. The Behavioral and Brain Sciences, 4, 419-457.


Bernstein, I. S. (1981). Dominance relationships and rank: Explanations, and empirical challenges (response to commentary on Bernstein (1981)). Behavioral and Brain Sciences, 4, 449-457.


Bernstein, I. S., & T.P., G. (1980). The social component of dominance relationships in rhesus monkeys. Animal Behaviour, 28, 1033-1039.


Bolles, R. C. (1981). A parallel to dominance competition (commentary on Berntein (1981)). Behavioral and Brain Sciences, 4, 433-434.


Borchelt, P. L., and Voith, V.L. (1996). Dominance Aggression in Dogs. In V. L. Voith, and Borchelt, P.L. (Ed.), Readings in Companion Animal Behavior (pp. 230-237). Trenton, NJ: Veterinary Learning Systems.


Brain, P. F. (1981). The concept of dominance also has problems in studies on rodents (commentary on Berntein (1981)). Behavioral and Brain Sciences, 4, 434-435.


Bramblett, C. A. (1981). Dominance tabulation: Giving form to concepts (commentary on Berntein (1981)). Behavioral and Brain Sciences, 4, 435-436.


Brown, J. L. (1963). Aggressiveness dominance and social organization in the steller jay. Condor, 65, 460-484.


Candland, D. K., and Hoer, James B. (1981). The logical status of dominance (commentary on Berntein (1981)). Behavioral and Brain Sciences, 4, 436-437.


Capitanio, J. P. (1991). Levels of integration and the 'inheritance of dominance'. Animal Behaviour, 42, 495-496.


Chalmers, N. R. (1981). Dominance as part of a relationship (commentary on Berntein (1981)). Behavioral and Brain Sciences, 4, 437-438.


Chase, I. D. (1982). Behavioral sequences during dominance hierarchy formation in chickens. Science, 216, 439-440.


Chase, I. D. (1985). Explainations of hierarchy structure. Animal Behaviour, 34, 1265-1266.


Christian, J. (1970). Social subordination, population density and mammalian evolution. Science, 168, 84-90.


Christman, M. C., and Lewis, D. (2005). Spatial distribution of dominant animals within a grou: comparison of four staistical tests of location. Animal Behaviour, 70, 73-82.


Clothier, S. (n/d). Why not take candy from a baby (if he lets you?). at http://www.flyingdogpress.com/candy.html.


Collias, N. E. (1943). Statistical analysis of factors which make for success in initial encounters between hens. American Naturalist, 77, 519-538.


Conniff, R. (2000). I want to be boss? (the psychology of dominance). Discover, May.


de Waal, F. B. M. (1986). The interation of dominance and social bonding in primates. Quarterly Review of Biology, 61(4), 459-479.


DeNapoli, J. S., Dodman, Nicholas H., Shuster, Louis, Rand, William, Gross, Kathy. (2000). Effects of dietary protein content and tryptophan supplementation on dominance aggression, territorial aggression, and hyperactivity in dogs. Journal of the American Medical Association, 217(4), 504-508.


Dewsbury, D. A. (1990). Fathers and sons: genetic factors and social dominance in deer mice, Peromyscus maniculatus. Animal Behaviour, 39, 284-289.


Dodman, N. H., Donnelly, R., Shuster, L., Mertens, P., Rand, W., and Miczek, K. (1996). Use of fluoxetine to treat dominance aggression in dogs. Journal of the American Veterinary Medical Association, 209(9), 1585-1587.


    * Abstract: OBJECTIVE: To evaluate fluoxetine for the treatment of owner-directed dominance aggression in dogs. DESIGN: Prospective study. ANIMALS: 9 dogs of various breeds, ages, and either sex determined to have owner-directed dominance aggression. PROCEDURE: Placebo and fluoxetine (1 mg/kg of body weight) were compared for the treatment of owner-directed dominance aggression in a single-blind crossover study. Owners were instructed to record aggressive and nonaggressive responses of their dogs daily on a canine-overt aggression chart for the 5-week duration of the study. Total aggression scores (linear and geometric) were calculated for each week of the study. The frequency of individual responses was also analyzed independently. RESULTS: Fluoxetine resulted in a significant (P = 0.01) reduction in owner-directed dominance aggression after 3 weeks of treatment. No particular aggressive response accounted for the overall reduction in aggression. CLINICAL IMPLICATIONS: Fluoxetine may be useful in the management of dominance aggression in dogs.



Dodman, N. H., Moon, R., and Zelin, M. (1996). Influence of owner personality type on expression and treatment outcome of dominance aggression in dogs. Journal of the American Veterinary Medical Association, 209(6), 1107-1109.


    * Abstract: OBJECTIVE: To determine the success rate of positive training methods and behavioral modification techniques in dogs with dominance aggression and to compare personality profiles between owners of dominant-aggressive and nondominant dogs. DESIGN: Prospective clinical study. ANIMALS: 10 dominant-aggressive dogs and 10 non-dominant, nonaggressive control dogs. PROCEDURE: Dominance aggression was quantified, using an aggression score, in the 10 dominant dogs before and after a nonconfrontational behavior modification program. The personality profile of the owners of dominant and control dogs, assessed by means of a Keirsey temperament sorter, was compared, as was the influence of owner personality on the outcome of behavioral modification in the dominant dogs. RESULTS: 9 of 10 dominant dogs responded to the nonconfrontational treatment program by a decrease in aggressive response to similar eliciting stimuli. Significant differences were not found between the personality of the owners of dominant versus control dogs, and owner personality did not significantly affect the outcome of behavior modification treatment. CLINICAL IMPLICATIONS: Nonconfrontational behavior modification programs are effective in reducing owner-directed dominance aggression in dogs. Owner personality does not necessarily predispose certain individuals to assaults by dominant dogs or profoundly affect their ability to engage in a successful behavioral modification program.



Drews, C. (1993). The concept and definition of dominance in animal behaviour. Behaviour, 125(3-4), 283-313.


    * Abstract: The concept of dominance has contributed greatly to our understanding of social structure in animals. Over the past three decades, however, a variety of concepts and definitions of dominance have been introduced, leading to an ongoing debate about the usefulness and meaning of the concept. Criticisms aimed at one definition of dominance do not necessarilly apply to other definitions. Existing definitions can be structural or functional, refer to roles or to agonistic behaviour, regard dominance as a property of individuals or as an attribute of dyadic encounters, concentrate on aggression or on the lack of it, and be based either on theoretical constructs or on observable behaviour.



Dugatkin, L. A., and Ohlsen, Sandra R. (1989). Contrasting asymmetries in value expectation and resource holding power: effects on attack behaviour and dominance in the Pumpkinseed Sunfish, Lepomis gibbosus. Animal Behavior, 39(4), 802-804.


Eaton, G. G. (1981). Measurement and utility of dominance rankings (commentary on Berntein (1981)). Behavioral and Brain Sciences, 4, 438.


Edwards, D. E., and Kravitz, E.A. (1997). Serotonin, social status and aggression. Current Opinions in Neurobiology, 7, 812-819.


    * Abstract: Serotonin, social status and aggression appear to be linked in many animal species, including humans. The linkages are complex, and, for the most part, details relating the amine to the behavior remain obscure. During the past year, important advances have been made in a crustacean model system relating serotonin and aggression. The findings include the demonstration that serotonin injections will cause transient reversals in the unwillingness of subordinate animals to engage in agonistic encounters, and that at specific synaptic sites involved in activation of escape behavior, the direction of the modulation by serotonin depends on the social status of the animal.



Flannelly, K. J., and Blanchard, Robert J. (1981). Dominance: Cause or description of social relationships? (commentary on Berntein (1981)). Behavioral and Brain Sciences, 4, 438-440.


Gage, F. H. (1978). A multivariate approach to the analysis of social dominance. Behavioral Biology, 23, 38-51.


Gage, F. H. (1981). Dominance: Measure first and then define (commentary on Bernstein (1981)). Behavioral and Brain Sciences, 4, 440-441.


Gammell, M. P., De Vries, H., Jennings, D.J., Carlin, C.M., and Haydeb, T.J. (2003). David's score: a more appropriate dominance ranking method than Clutton_Brock et al.'s index. Animal Behaviour, 66, 601-605.


Gauthreaux Jr., S. A. (1981). Behavioral dominance from an ecological perspective (commentary on Bernstein (1981)). Behavioral and Brain Sciences, 4, 441.


Gese, E. M., & Mech, L. D. (1991). Dispersal of wolves (Canis lupus) in northeastern Minnesot 1969-1989. Cabadian Journal of Zoology, 69, 2946-2955.


Hand, J. L. (1986). Resolution of social conflict: dominance, egalitarianism, speres of dominance, and game theory. The Quarterly Review of Biology, 61(2), 201-220.


Hartmann, S. (2002). The Harmony Program version 2.1.


Hinde, R. A., and Datta, Saroj. (1981). Dominance: An intervening variable (commentary on Bernstein (1981)). Behavioral and Brain Sciences, 4, 442.


Hofer, H. a. E., Marion L. (2000). Conflict Management in Female-Dominated Spotted Hyenas. In F. Aureli, and de Waal, Frans B.M. (Ed.), Natural Conflict Resolution (pp. 232-234). Berkeley and Los Angeles: California University Press.


Hsu, Y., & Wolf, L. L. (2001). The winner and loser effect: what fighting behaviours are influenced? Animal Behaviour, 61, 777-786.


    * Abstract: We examined the effect of prior winning and losing experiences on the initiating and responding strategies of contestants in contests between individuals of Rivulus marmoratus (Cyprinodontidae). Each contestant was given a penultimate and a recent fighting experience approximately 48 and 24 h prior to the dyadic contests, respectively, through randomly selected procedures. Winning and losing experience appeared to influence different types of fighting behaviours. Losing experiences decreased the probability of an individual initiating a confrontation and thus increased its tendency to retreat immediately when challenged. Winning experiences did not affect the probability of initiation, but significantly increased the likelihood of an individual initiating with attacks that effectively deterred its opponents. A substantial proportion (59/153) of individuals retreated immediately when challenged and reduced the number of fights available for examining experience effects on responding strategies at later stages of a contest. None the less, winning experiences consistently increased the likelihood of an individual retaliating by attacking its opponent at various stages of a contest, and eventually increased its probability of escalating a confrontation into physical fights. However, the effects of losing experiences on these responding strategies were undetectable. Recent experiences significantly affected all fighting behaviours examined, but penultimate experiences significantly affected only the tendency to initiate a confrontation with attacks and the likelihood of escalation. These results indicated that prior experiences had the longest lasting effect on the potentially most costly fighting behaviour. Prior experiences influenced the outcome of nonescalated contests as well as the probability of escalation, but did not significantly affect the outcome of escalated contests. These results are consistent with the hypothesis that prior experiences modify the information that an individual has about its fighting ability but do not alter its actual fighting ability and that actual fighting ability becomes the more important influence on outcomes of escalated contests.



Jackson, W. M., and Winnegrad, R. L. (1988). Linearity in dominance hierarchy: a second look at the individual attribute model. Animal Behaviour, 36, 1237-1240.


James, W. T. (1955). Behaviors involved in expression of dominance among puppies. Psychological Reports, 1, 299-301.


Kaplan, J. R. (1981). A reexamination of dominance rank and hierarchy in primates. The Behavioral and Brain Sciences, 4, 442-443.


Kaplan, J. R. (1981). A reexamination of dominance rank and hierarchy in primates (commentary on Bernstein (1981)). Behavioral and Brain Sciences, 4, 442-443.


Kaufmann, J. H. (1983). On the definitions and functions of dominance and territoriality. Biological Review, 58, 1-20.


Lott, D. F. (1981). Circumstances in which exact dominance rank may be important (commentary on Bernstein (1981)). Behavioral and Brain Sciences, 4, 443-444.


Maxim, P. E. (1981). Dominance: A useful dimension of social communication (commentary on Bernstein (1981)). Behavioral and Brain Sciences, 4, 444-445.


Mazur, A., & Booth, A. (1998). Testosterone and Dominance in Men. Behavioural and Brain Sciences at, 21, 353-363.


    * Abstract: In men, high levels of endogenous testosterone (T) seem to encourage behavior intended to dominate – to enhance one's status over – other people. Sometimes dominant behavior is aggressive, its apparent intent being to inflict harm on another person, but often dominance is expressed nonaggressively. Sometimes dominant behavior takes the form of antisocial behavior, including rebellion against authority and law breaking. Measurement of T at a single point in time, presumably indicative of a man's basal T level, predicts many of these dominant or antisocial behaviors. T not only affects behavior but also responds to it. The act of competing for dominant status affects male T levels in two ways. First, T rises in the face of a challenge, as if it were an anticipatory response to impending competition. Second, after the competition, T rises in winners and declines in losers. Thus, there is a reciprocity between T and dominance behavior, each affecting the other. We contrast a reciprocal model, in which T level is variable, acting as both a cause and effect of behavior, with a basal model, in which T level is assumed to be a persistent trait that influences behavior. An unusual data set on Air Force veterans, in which data were collected four times over a decade, enables us to compare the basal and reciprocal models as explanations for the relationship between T and divorce. We discuss sociological implications of these models.



Mech, D. L. (1999). Alpha status, dominance, and division of labor in wolf packs. Canadian Journal of Zoology, 77, 1196-1203. [available at http://www.npwrc.usgs.gov/resource/mammals/alstat/alstat.htm]


    * Abstract: The prevailing view of a wolf (Canis lupus) pack is that of a group of individuals ever vying for dominance but held in check by the "alpha" pair, the alpha male and alpha female. Most research on the social dynamics of wolf packs, however, has been conducted on non-natural assortments of captive wolves. Here I describe the wolf-pack social order as it occurs in nature, discuss the alpha concept and social dominance and submission, and present data on the precise relationships among members in free-living packs, based on a literature review and 13 summers of observations of wolves on Ellesmere Island, Northwest Territories, Canada. I conclude that the typical wolf pack is a family, with the adult parents guiding the activities of the group in a division-of-labor system in which the female predominates primarily in such activities as pup care and defense and the male primarily during foraging and food-provisioning and the travels associated with them.



Mertens, P. A. (2004). The concept of dominance and the treatment of aggression in multidog homes: a comment on van Kerkhove's commentary. J Appl Anim Welf Sci, 7(4), 287-291; discussion 299-300.


Messier, F. (1985). Solitary living and extraterritorial movements of wolves in relation to social status and prey abundance. Canadian Journal ofr Zoology, 63, 239-245.


    * Abstract: The study examines three factors encouraging wolf (Canis lupus ) dispersal: prey abundnance, age of the animals, and their sex. Factors are related to the frequency at which wolves (n = 54) dissociated from their packs and (or) engaged in extraterritorial movements. A low prey base did not increase solitary living or excursion frequency of pups, did increase both traits in yearlings, and increased only solitary living among adults. Yearling and adult females were dissociated from their packs more frequently than males. Yearling females travelled more frequently outside their territories than did yearling males. On average, pup, yearling, and adult individuals made 1.1, 3.0, and 1.0 extraterritorial excursions per year, respectively. Such movements, mostly in winter, are interpreted as predispersal forays (failed dispersal), but also as an immediate action to survive through a temporary resource failure. Dispersal in wolves appears as a gradual and dynamic dissociation process extended over a period of a few months to a few years and beginning as early as 10 months of age.



Moore, A. J. (1990). The inheritance of social dominance, mating behaviour and attractiveness to mates in male Nauphoeta cinerea. Animal Behaviour, 39, 388-397.


Morse, D. H. (1974). Niche breadth as a function of social dominance. The American Naturalist, 108(964), 818-830.


    * Abstract: If one species is socially dominant to another, the subordinate usually narrows its niche when they occur together. When one species is dominant in some circumstances and a second in others, both narrow their niches when together. Subordinates usually have a larger fundamental niche than their dominants. The presence of dominants should result in selection for enlarged (or changed) fundamental niches by the subordinate species. Linear hierarchies of species should result in guilds whose members have different-sized fundamental and realized niches. Though large mobile species often have greater fundamental niches than small similar species, an inverse relationship between size and niche breadth usually occurs where clear dominance hierarchies exist, suggesting that social dominance has more than counteracted the effect of body size. The presence of dominants is a factor making conditions uncertain for subordinates. Limits to the niche breadth of dominants are in some cases directly set by physical factors but in others are unknown, though they may be set by one or more of a variety of biological factors. In some cases subordinates appear to avoid interactions with dominants. Unequivocal examples of dominance-mediated niche relationships are thus far confined to certain vertebrate and arthropod groups, although evidence available suggests that they are more widely spread.



Moynihan, M. H. (1998). The Social Regulation of Competition and Aggression in Animals. Washington: Smithsonian Institute.


Netto, W. J., va der Borg, J.A., and Slegers, J.F. (1992). The establishment of dominance relationships in a dog pack and its relevance for the man-dog relationship. Tijdschr Diergeneeskd, 117, 51S-52S.


O'Heare, J. (2003). Dominance Theory and Dogs (1st ed.). Ottawa: Dogpsych Publishing.


O'Heare, J. (2004). What is Social Dominance? Unpublished Ph.D. Dissertation, Breyer State University, Ottawa, Canada.


Parker, G. A. (1974). Assessment strategy and the evolution of fighting behaviour. Journal of Theoretical Biology, 47, 223 - 243.


Pereira, R. A. S., and Prado, A.P.D. (2005). Recognition of competitive asymmetries reduces the severity of fighting in male Idarnes fig wasps. Animal Behaviour, 70, 249-256.


Petraitis, P. S. (1981). Dominance rankings and problems of intransive relationships (commentary on Bernstein (1981)). Behavioral and Brain Sciences, 4, 445-446.


Plutchik, R. (1981). Dominance: A key ethological/sociological concept (commentary on Bernstein (1981)). Behavioral and Brain Sciences, 4, 446.


Preuschoft, S., and van Schaik, Carol P. (2000). Dominance and Communication Conflict Management in Various Social Settings. In F. Aureli, and de Waal, Frans B.M. (Ed.), Natural Conflict Resolution (pp. 77-105). Berkeley and Los Angeles: California University Press.


Pulliainen, E. (1967). A contribution to the study of the social behavior of the wolf. American Zoologist, 7, 313-317.


    * Abstract: Three wolf cubs ([female] and [male male]) taken blind from their den were studied in captivity. When the cubs were 20–22 weeks old, their behavior was studied in the presence of individuals of two breeds of dogs (German Shepherd and Samoyed). The wolf cubs had seen no canine individuals before the experiments and vice versa. The sexes and ages of the German Shepherds (they resemble European forest wolves in appearance) tested were as follows: female (3 years), male (17 months), female (21 weeks; the same height as the wolf cubs) and female (12 weeks). The two female Samoyeds were 2 years old. All the tests were performed in the pen of the wolf cubs and filmed. The main results are as follows: (1). Investigative behavior was observed during all the tests carried out. (2). No aggressiveness was observed between the wolf cubs and those German Shepherds which were as tall as the wolf cubs or taller. By contrast, the wolf cubs tried to kill the smallest German Shepherd (12 weeks old). (3). Great aggressiveness was observed between the wolf cubs and the Samoyeds from the moment of confrontation. (4). No difference was observed in the movements and behavior patterns of these two species. (5). The results are discussed.



Pusey, A. E., & Packer, C. (1997). The ecology of relationships. In J. R. Krebs & N. B. Davies (Eds.), Behavioural Ecology an evolutionary approach 4th. ed. Oxford: Blackwell Science.


Rabb, G. B. (1967). Social relationships in a group of captive wolves. American Zoologist, 7, 305-311.


    * Abstract: The social organization of a group of wolves in a large outdoor enclosure was followed through several breeding seasons. During the breeding season conflicts become more frequent and the social hierarchy obvious. The more dominant animals restrict courtship activities by inferior wolves of their own sex. However, apparently as a correlate of their position, two alpha males have shown less mating activity than other males. Mate preferences exhibited by animals of both sexes also limit the number of matings. The preferences appear related to the social hierarchy existing when an animal matures. Cultural transmission of social status is suggested by some changes in ranking of wolves raised in the woods at Brookfield. Temporary removal of the original alpha male and death of the original alpha female appear to have promoted changes in social order and an increase in actual mating combinations. The probable consanguineous nature of wolf groups and facets of the social behavior suggest that some form of group selection could be operative in the wild.



Rooney, N. J., and Bradshaw, J.W.S. (2003). Links between play and dominance and attachment dimensions of dog-human relationships. Journal of Applied Animal Welfare Science, 6(2), 67-94.


    * Abstract: It is often claimed that certain behavioral problems in domestic dogs can be triggered by the games played by dog and caregiver (owner). In this study, we examine possible links between the types of games played and dimensions of the dog-owner relationship that are generally considered to affect such problems. Fifty dog-owner partnerships were filmed during 3-min play sessions in which the owner was allowed to choose the games played. All partnerships then undertook a 1-hr test designed to measure elements of behavior commonly ascribed to "dominance" and "attachment." Principal components analysis of the data produced 2 dominance-related factors (Amenability and Confident Interactivity) and 4 factors describing aspects of attachment (Nonspecific Attention Seeking, Preference for Owner, Preference for Unfamiliar Person, and Separation-Related Behavior). Amenability, in particular, varied significantly between breeds. In the study, we then compared types of games played to each of these factors. Dogs playing rough-and-tumble scored higher for Amenability and lower on Separation-Related Behavior than did dogs playing other types of games. Dogs playing tug-of-war and fetch scored high on Confident Interactivity. Winning or losing these games had no consistent effect on their test scores. If the dog started the majority of the games, the dog was significantly less amenable and more likely to exhibit aggression. The results suggest that how dogs play reflects general attributes of their temperament and relationship with their owner. This study provides no evidence that games play a major deterministic role on dominance dimensions of dog-human relationships, but the results suggest that playing games involving considerable body contact may affect attachment dimensions.



Rooney, N. J., Bradshaw, J. W. S., & Robinson, I. H. (2000). A comparison of dog-dog and dog-human play behaviour. Applied Animal Behaviour Sciences, 66, 235-248.


Rooney, N. J., Bradshaw, J. W. S., & Robinson, I. H. (2001). Do dogs respond to play signals given by humans? Animal Behavior, 61, 715-722.


    * Abstract: Play signals are known to function in the solicitation and maintenance of intraspecific play, but their role in interspecific play is relatively unstudied. We carried out two studies to examine interspecific signalling when humans play with domestic dogs, Canis familiaris. In the first, we recorded dog–owner play sessions on video to identify actions used by 21 dog owners to initiate play with their dogs. Thirty-five actions were each used by three or more owners. These included postures, vocalizations and physical contact with the dog. The actions varied greatly in their apparent success at instigating play which was, surprisingly, unrelated to the frequency with which they were used. We then did an experiment to determine the effect of composites of commonly used signals upon the behaviour of 20 Labrador retrievers. The performance of both ‘Bow’ and ‘Lunge’ by a human altered the subsequent behaviour of the dogs. Both signals caused increases in play, and Lunge produced significant increases in play bout frequency and mean bout duration. The efficiency of both these postural signals was enhanced when they were accompanied by play vocalizations. Thus, specific actions used by humans do communicate a playful context to dogs and can be described as interspecific play signals.



Rosenberg, R. H. a. E., Magnus. (1991). Contest behaviour in Weidemeyer's admiral butterfly, Limenitis weidemeyerii (Nymphalidae): the effect of size and residency. Animal Behavior, 42, 805-811.


Rowell, T. E. (1974). The concept of dominance. Behavioral Biology, 11, 131-154.


Schenkel, R. (1967). Submission: its features and function in the wolf and dog. American Zoologist, 7, 319-329.


    * Abstract: Submission in the wolf and dog is defined on the basis ot its motivation: submission is the effort of the inferior to attain friendly or harmonic social integration. Submission functions as an appeal or a contribution to social integration, but only if it meets a corresponding attitude in the superior. The form of submissive behavior in wolf and dog is ritualized and symbolized cub-behavior. Two main forms of submissive behavior occur in wolf and dog: active submission, derived from begging for milk or food, and passive submission, derived from the posture which the cub adopts when cleaned by its mother. The definition of submission is generally applicable to vertebrates living in groups based on intimacy and a social hierarchical order. The concept of submission as the role of the defeated in the terminal phase of fight with the function to inhibit automatically aggression in the superior should be dismissed. In vertebrates at least three types of conflict with different terminal phases occur: (1). Severe fight based on intolerance; ends with flight by the inferior or with his death. (2). Ritualized fight over a privilege; ends with the "giving-up-the-claim ritual" of the inferior, which automatically blocks the aggression of the superior. (3). Minor conflict in closed groups; settled by submissive behavior of the inferior. In closed vertebrate groups, intermediate forms between (1) and (3) occur, depending on the proportion between activated intimacy and intolerance.



Scott, J. P. (1948). Dominance and the frustration-aggression hypothesis. Physiological Zoology, 21, 31-39.


Seyfarth, R. M. (1981). Do monkeys rank each other? (commentary on Bernstein (1981)). Behavioral and Brain Sciences, 4, 447-448.


Sidanius, J., and Pratto Felicia. (1999). Social Dominance. Cambridge, United Kingdom: University of Cambridge.


Smuts, B. (1981). Dominance: An alternative view (commentary on Bernstein (1981)). Behavioral and Brain Sciences, 4, 448-449.


Turner, G. F. (1994). The fighting tactics of male mouthbrooding cichlids: The effects of size and residency. Animal Behavior, 47, 655-662.


Uchida, Y., Dodman, N., DeNapoli, J., and Aronson, L. (1997). Characterization and treatment of 20 canine dominance aggression cases. J Vet Med Sci., 59(5), 397-399.


    * Abstract: This study was undertaken to characterize 20 cases of dominance aggression seen at Tufts University School of Veterinary Medicine and to investigate the efficacy of our non-confrontational behavior modification program for 8 weeks. The 20 cases included 18 pure breed and 2 mixed breed dogs. Thirteen of the dogs were male. The dogs' ages ranged from 7 to 84 months (mean 32.1 +/- 22.64 SE). There was no correlation between the severity of dominance aggression and the signalment of the dogs. At the conclusion of the eight week follow up period, 14 dogs (70%) were reported to have responded to the treatment to some degree. Six dogs did not demonstrate any noticeable reduction in aggressive behavior or became more aggressive. The results of the study is powerful evidence of the efficacy of the non-confrontational behavior modification program.



Uyeno, E. T., & White, M. (1967). Social Isolation and dominance behavior. Journal of Comparative and Psychiological Psychology, 63(1), 157-159.


van Kerkhove, W. (2004). A fresh look at the wolf-pack theory of companion-animal dog social behavior. J Appl Anim Welf Sci, 7(4), 279-285; discussion 299-300.


    * Abstract: A popular perspective on the social behavior of dogs in multiple-dog households sees the dogs' behavior as reflecting the sociobiological laws of the rigidly structured dominance hierarchy that has been described for wolf packs. This view suggests that aggression problems among dogs are natural expressions of conflict that arise whenever dominance status is in contention. One recommended solution has been for the owner to endorse and enforce a particular dominance hierarchy because, on the wolf pack model, aggression is minimized when the structure of the hierarchy is clear, strong, and stable. This article questions the validity of this perspective on 2 principal grounds. First, because it does not seem to occur in the wild, this article suggests the strong dominance hierarchy that has been described for wolves may be a by-product of captivity. If true, it implies that social behavior—even in wolves—may be a product more of environmental circumstances and contingencies than an instinctive directive. Second, because feral dogs do not exhibit the classic wolf-pack structure, the validity of the canid, social dominance hierarchy again comes into question. This article suggests that behavioral learning theory offers another perspective regarding the behavior of dogs and wolves in the wild or in captivity and offers an effective intervention for aggression problems.



van Schaik, C. P. a. A., Filippo. (2000). The Natural History of Valuable Relationships in Primates. In F. Aureli, and de Waal, Frans B.M. (Ed.), Natural Conflict Resolution (pp. 307-333). Berkeley and Los Angeles: California University Press.


Verbeek, P., Hartup, Willard W., and Collins W. Andrew. (2000). Conflict Management in Children and Adolescents. In F. Aureli, and de Waal, Frans B.M. (Ed.), Natural Conflict Resolution (pp. 34-53). Berkeley and Los Angeles: California University Press.


Verrell, P. A. (1986). Wrestling in red-spotted newt (Notophthalmus viridescens): resource value and contestant asymmetry determine contest duration and outcome. Animal Behavior, 34, 398-402.


Vessey, S. H. (1981). Dominance as control (commentary on Bernstein (1981). Behavioral and Brain Sciences, 4, 449.


Weibull, J. W. (1995). Evolutionary Game Theory. Massachusetts: MIT Press.


White, M. M., Neilson, J. C., Hart, B. L., and Cliff, K. D. (1999). Effects of clomipramine hydrochloride on dominance-related aggression in dogs. Journal of the American Veterinary Medical Association, 215(9), 1288-1291.


    * Abstract: OBJECTIVE: To compare effects of the serotonergic drug clomipramine hydrochloride with those of placebo for treatment of dominance-related aggression in dogs. DESIGN: Randomized, placebo-controlled, double-blind clinical trial. ANIMALS: 28 neutered dogs > 1 year old with dominance-related aggression. PROCEDURE: Dogs displaying > or =3 aggressive episodes/wk toward > or= 1 human family member in response to identifiable behavioral triggers were included in the study. Owners were instructed not to change patterns of interaction with their dogs during the study. After 2 weeks of baseline observations, dogs were treated for 6 weeks with clomipramine (1.5 mg/kg [0.7 mg/lb] of body weight, q 12 h; n = 15) or placebo (13). Responses to triggers were assigned the following aggression scores: no response, 0; growl or lip curl, 1; snap or bite, 2. Mean scores for responses to triggers were obtained during the 2-week pretreatment period (baseline) and during the first and second weeks, third and fourth weeks, and fifth and sixth weeks of treatment. At the end of the study, owners assigned a score designed to evaluate their overall perceived change in aggressiveness; this was referred to as the global score. RESULTS: Mean aggression scores decreased at the fifth and sixth week of treatment in both groups, compared with baseline scores. However, mean scores between groups were not different. Global scores, assigned by the owner, generally reflected changes in mean aggression scores. CONCLUSIONS AND CLINICAL RELEVANCE: Compared with placebo, clomipramine administered to dogs at the dosage recommended for treatment of separation anxiety did not reduce aggressiveness toward human family members.



Ydenberg, R. C., Giraldeau, L. A., and Falls, J. B. (1988). Neighbours, strangers, and asymmetric war of attrition. Animal Behavior, 36, 343-347.


Issues Related to Females Aggression Increasing After Spaying


de Jonge FH, Eerland EM, van de Poll NE. (1986). Sex-specific interactions between aggressive and sexual behavior in the rat: effects of testosterone and progesterone. Horm Behav. 20(4):432-44


    * Abstract: The influence of progesterone on sexual and aggressive behaviors during aggressive encounters was investigated in pairs of TP-treated male and female rats. Gonadectomized females, chronically injected with testosterone propionate (TP), showed low but consistent levels of feminine sexual behavior which alternated with aggression. Progesterone when given in addition to TP facilitated receptive and proceptive behaviors, but reduced levels of aggression. In TP-treated males, levels of aggression were the same as observed in TP-treated females. However, TP-treated males seldomly showed sexual behavior during aggressive encounters and additional treatment with progesterone did not affect their behavior. After the aggression tests, animals were tested in a social preference test in which an ovariectomized female cage mate and the opponent from the aggressive encounter served as incentives. Positive correlations between levels of aggression and social preference for an opponent were found in both sexes, although correlations only reached statistical significance when progesterone was given in addition to TP. These correlations were found in both sexes, despite the fact that group analysis revealed pronounced sex differences in social preference: males preferred to spend their time near ovariectomized female cage mates, whereas females divided their time equally among female cage mates and opponents.



Compaan JC, van Wattum G, de Ruiter AJ, van Oortmerssen GA, Koolhaas JM, Bohus B. (1993). Genetic differences in female house mice in aggressive response to sex steroid hormone treatment. Physiol Behav. 54(5):899-902


    * Abstract: Male mice, genetically selected for aggression, characterized by short attack latency (SAL) or long attack latency (LAL), differ on several testosterone (T)-related parameters during ontogeny and adult age. The variation in aggressive behavior at adult age may be due to differences in degree of androgenization prenatally. When exposed to T at prenatal, neonatal, and/or adult age, nonlactating females also display intraspecific fighting behavior. In the present study, we investigated in females of the SAL and LAL selection lines, whether the differentiation of aggression involves processes similar to ones seen in males. Therefore, we injected females with testosterone propionate (TP) or vehicle on the day of birth, treated them after ovariectomy at adult age with T, estradiol (E), or vehicle, and tested their aggressive response. We found that neonatally vehicle-treated SAL females show a higher aggressive response to chronic T treatment at adult age than LAL females receiving the same treatment. Females of both selection lines treated with vehicle or E as adults were not aggressive. Neonatal TP treatment did not influence the adult T sensitivity and difference between selection lines in response to T at adult age. However, neonatally TP-treated SAL females showed aggressive behavior when treated with E at adult age, whereas LAL females failed to do so. These results suggest a genetic difference in susceptibility to T and E, which plays a major role prenatally, in organizing the development of sex steroid-dependent neural systems.



Compaan JC, de Ruiter AJ, Koolhaas JM, van Oortmerssen GA, Bohus B. (1992). Differential effects of neonatal testosterone treatment on aggression in two selection lines of mice. Physiol Behav. 51(1):7-10


    * Abstract: Selection lines of mice, artificially selected for aggression based upon the attack latency score (ALS), were used. In order to determine the relative contribution of neonatal testosterone (T) in the development of aggression, we vary the plasma-T level in males of both selection lines on the day of birth. At 14 weeks the ALS was measured. Neonatal T treatment results in a reduction of aggression in the long attack latency (LAL) line, whereas aggressive behaviour of the short attack latency (SAL) line is not affected. Both selection lines show reduction in testicular weight, although the total amount of T-producing Leydig cells was not affected. Neonatal T may cause a permanent reduction in aggressive behaviour in in the LAL line only, probably due to differential appearance of critical periods. It is suggested that the difference in aggressive behaviour between SAL and LAL selection lines is due to a prenatally determined difference in neonatal T sensitivity of the brain.



O'Farrell, V., & Peachey, E. (1990). Behavioural effects of ovariohysterectomy on bitches. Journal of Small Animal Practice, 31, 595-598.


Overall, K.L. (1995). Sex and aggression. Canine Practice, 20(3): 16-18


van de Poll NE, van Zanten S, de Jonge FH. (1986). Effects of testosterone, estrogen, and dihydrotestosterone upon aggressive and sexual behavior of female rats. Horm Behav. 20(4):418-31.


    * Abstract: Groups of female TMD rats were treated either with estradiol benzoate (EB), dihydrotestosterone propionate (DHTP), testosterone propionate (TP), EB + DHTP (EB/DHTP), or with oil. These groups of females were tested for social aggression and for masculine and feminine sexual behavior. In addition, patterns of masculine and feminine sexual responses during the aggressive encounters, were investigated. TP-treated females of the same strain were used as opponents in the tests for aggression. In accordance with previous results, EB did not activate aggression whereas TP treatment resulted in a significant increase in aggression in females. Aggressive responses were activated by adding DHTP to EB, up to levels equal to those activated by TP. Sexual responses were observed in the tests for aggression as well as in tests for sexual behavior. The results indicated that feminine and masculine sexual responses were affected significantly by hormonal treatment. Mounting behavior in the test for aggression was activated by TP and by EB/DHTP. Lordosis and proceptive responses were inhibited in these groups as compared to EB-treated females, both in tests for aggression and in tests for sexual behavior. The results are consistent with the idea that dihydrotestosterone inhibits feminine and activates masculine sexual activity. The results also indicate that EB and DHTP synergistically activate aggression.



vom Saal, F.S. (1989). Sexual differentiation in litter-bearing mammals: influence of sex of adjacent fetuses in utero. Journal of Animal Science, 67:1824-1840


    * Abstract: In rodents and swine, individual differences in a broad range of characteristics correlate with intrauterine position during fetal life. By identifying the intrauterine position of mice at cesarean delivery, we can predict reliably postnatal reproductive traits such as genital morphology, timing of puberty, length of estrous cycles, timing of reproductive senescence, sexual attractiveness, sexual behavior, aggressiveness, daily activity level, body weight and tissue enzyme activity in females; in males we can predict genital and brain morphology, sexual behavior, aggressiveness, daily activity level, body weight, and tissue enzyme activity. In mice, as in all mammals, male fetuses have greater concentrations of testosterone than do females. In addition, female mouse fetuses have greater circulating concentrations of estradiol than do male fetuses, a condition not found in all mammals. A mouse fetus positioned between males has greater concentrations of testosterone than does a fetus of the same sex positioned between females, and a fetus positioned between females has greater concentrations of estradiol than does a fetus of the same sex positioned between males. Gonadal steroids regulate differentiation of secondary sexual characteristics. Studies in which the effects of intrauterine position have been eliminated by exposing fetuses to steroid receptor blockers reveal the critical role of steroids in mediating this phenomenon. The intrauterine position phenomenon provides the only mammalian model for relating postnatal traits to concentrations of endogenous hormones to which individuals are exposed during fetal life. Results from studies using this naturally occurring experimental system in litter-bearing species have given insights concerning the consequences of individual differences in steroid concentrations during sexual differentiation that likely apply to all mammals. One specific hypothesis is that circulating estradiol may interact with testosterone in mediating some aspects of sexual differentiation in rodents and, thus, possibly in other mammals.



vom Saal, F.S. (1981). Variation in phenotype due to random intrauterine positioning of male and female fetuses in rodents. Journal of Reproduction and Fertility, 62:633-650


    * Abstract: Rodents are polytocous mammals, and male and female fetuses can develop in utero contiguous to fetuses of the same or opposite sex. This paper describes experiments demonstrating that random intrauterine positioning of male and female fetuses results in within-sex variation in phenotype in mice and rats. This phenomenon provides a clear example of the degree to which the intrauterine environment can bias development in terms of effects on morphology, physiology and behaviour. I propose that individual differences in reproductively-related characteristics based on prior intrauterine position may play a role both in the regulation of population size in rodents and in the reproductive success of individuals as changes in population size occur.


Domestication and Evolution of the Dog


Belyaev, D. K. (1979). The Wilhelmine E. Key 1978 invitational lecture. Destabilizing selection as a factor in domestication. J Hered, 70(5), 301-308. http://www.sbs.utexas.edu/genetics/Fall05/Handouts/Dogs/JHered-70-1979-destabilizing%20selection_silver%20fox%20experiment.pdf


Budiansky, S. (1999). The truth about dogs. The Atlantic Monthly at http://www.theatlantic.com/issues/99jul/9907dogs2.htm.


Clutton-Brock, J. (1995). Origins of the Dog: domestication and early history. In J. A. Serpell (Ed.), The Domestic Dog it's evolution, behaviour and interactions with people (pp. 7-20). Cambridge, United Kingdom: Cambridge University Press.


Cohn, J. (1997). How wild wolves became domestic dogs. Bioscience, 47(11), 725-729.


Coppinger, R., & Coopinger, L. (2001). Dogs A Startling New Understanding of Canine Origin, Behavior & Evolution. New York, NY: Scribner.


Coppinger, R., & Feinstein, M. (1991, January). ’Hark! hark! The dogs do bark...’ and bark and bark. Smithsonian,, 21(10), 119-125. [discussion of the domestic dogs' barking and how it compares to vocalizations in wolves. Interesting background and early reference to Coppinger's self-domestication theory.]


Drake, A. G., & Klingenberg, C. P. (2008). The pace of morphological change: historical transformation of skull shape in St Bernard dogs. Proc Biol Sci, 275(1630), 71-76.


Fox, M. W. (1978). The Dog Its Domestication and Behavior. New York: STMP Press.


Keegans, E. (n/d). The Evolution of Dogs and Dog Training. Retrieved June 19, 2006, from http://www.apdt.com/about/prog/scholarships/win/2002_winner_02.aspx


    * Introductory Paragraph: Knowledge regarding the evolution of dogs has played a large role in shaping the methods that dog trainers use. New information may affect us and cause us to reevaluate our current understanding of how to communicate with and train dogs. The science of tracking DNA is bringing us amazing discoveries about the origins of all life including human beings and our best friends, Canis familiaris. Studies of feral dogs, wolves, and pet and working dogs uncover new information as well. Dog trainers may find that this new knowledge is the fresh breeze that supports the new, gentler methods of dog training that are coming to the forefront of the industry.



Koler-Matznick, J. (2002). The origin of the dog revisited. Anthrozoos, 15(2), 98-118. http://www.canineworld.com/ngsdcs/Origin.of.the.Dog.pdf


    * Abstract. The most widely accepted hypothesis of the origin of the dog, Canis familiaris, is that the dog is a domesticated gray wolf, Canis lupus. This paper reviews the evidence for this conclusion, finds many unanswered questions and conceptual gaps in the wolf origin hypothesis, and explores the alternative hypothesis that the most likely ancestor of the domestic dog was a medium-size, generalist canid. Introduction. Currently, there is a general consensus that the sole ancestral species of the domestic dog, Canis familiaris L. 1758, is the gray wolf, Canis lupus L. 1758 (Coppinger & Smith 1983; Clutton-Brock 1984, 1995; Olsen 1985; Wayne 1986; Tchernov & Horowitz 1991; Morey 1992; Wilson and Reeder 1993; Cohn 1997; Vilà et al.1997; Budiansky 1999; Crockford 2000; Coppinger & Coppinger 2001). However, there is at least one competing hypothesis that is equally plausible, yet has not been given adequate evaluation in the published literature. The dog origin alternatives are: (1) an origin from golden jackal (Canis aureus) (Lorenz 1954); (2) an origin from hybrids of wild canids (Darwin 1875; Clutton-Brock 1977; Brisbin 1997); or (3) an origin from a wild Canis other than jackal or C. lupus (Epstein 1971; Zeuner 1963; Fox 1973; Manwell & Baker 1983). Jackal ancestry of the domestic dog (hereafter DD) can be dismissed due to the large genetic distance separating the two (Wayne & O'Brien, 1987; Wayne et al., 1991a). Although a hybrid origin (e.g., Canis simensis or Cuon alpinus with small subspecies of C. lupus), suggested by several authors to account for the sudden shift from wolf to primitive dog morphology, cannot be ruled out, currently no direct evidence supports this hypothesis. In this paper, I review selected behavioral, morphological, molecular, and fossil information relative to the competing hypotheses of wolf origin and wild canid origin. Appendix I provides my working definitions of terms with variable meanings.



Leonard, J. A., Wayne, R.K., Wheeler, J., Valadez, R., Guillen, S., and Vila, C. (2002). Ancient DNA evidence for old world origin of new world dogs. Science, 298, 1613-1616.


    * Abstract: Mitochondrial DNA sequences isolated from ancient dog remains from Latin America and Alaska showed that native American dogs originated from multiple Old World lineages of dogs that accompanied late Pleistocene humans across the Bering Strait. One clade of dog sequences was unique to the New World, which is consistent with a period of geographic isolation. This unique clade was absent from a large sample of modern dogs, which implies that European colonists systematically discouraged the breeding of native American dogs.



Lindblad-Toh, K., Wade, C. M., Mikkelsen, T. S., Karlsson, E. K., Jaffe, D. B., Kamal, M., et al. (2005). Genome sequence, comparative analysis and haplotype structure of the domestic dog. Nature, 438(7069), 803-819.


    * Abstract: Here we report a high-quality draft genome sequence of the domestic dog (Canis familiaris), together with a dense map of single nucleotide polymorphisms (SNPs) across breeds. The dog is of particular interest because it provides important evolutionary information and because existing breeds show great phenotypic diversity for morphological, physiological and behavioural traits. We use sequence comparison with the primate and rodent lineages to shed light on the structure and evolution of genomes and genes. Notably, the majority of the most highly conserved non-coding sequences in mammalian genomes are clustered near a small subset of genes with important roles in development. Analysis of SNPs reveals long-range haplotypes across the entire dog genome, and defines the nature of genetic diversity within and across breeds. The current SNP map now makes it possible for genome-wide association studies to identify genes responsible for diseases and traits, with important consequences for human and companion animal health.



Morell, V. (1997). The origin of the dog: running with wolves. Science, 276, 1647-1648.


    * Summary: From the pug to the St. Bernard, dogs today come in almost every size, shape, and color imaginable, but a genetic study on page 1687 of this issue shows that they all have the same forebear: the wolf. And although humans turned the wicked wolf into loyal Lassie at least twice, domestication was apparently a rare event, requiring special skill.



Morey, D. F. (1994). The Early evolution of the domestic dog. American Scientist, 82, 336-347.


Newby, J. (1999). The Animal Attraction Humans and their Animal Companions. Sydney, NSW, Australia: ABC Books. [out of print; difficult to find. try abebooks.com or half.com; excellent first 3 chapters evaluate current theories of domestication of the dog]


Olsen, S. J. (1). Origins of the domestic dog; the fossil record. Tucson, Arizona: The University of Arizona Press. [out of print, but can be found used online or in large public libraries; contains a bit more in depth information on the fossil record and puts the record into context. Not necessarily a need-to-have.]


Pugh, K. A. (2005). DNA analysis of the development and relatedness of domestic ndogs. Unpublished Review, University of Saskatoon, Saskatoon.


Savolainen, P., Zhang, Y.-P., Luo, J., Lundberg, J., & Leitner, T. (2002, November 22). Genetic Evidence for an East Asian Origin of Domestic Dogs. Science, 298, 1610-1613.


    * The origin of the domestic dog from wolves has been established, but the number of founding events, as well as where and when these occurred, is not known. To address these questions, we examined the mitochondrial DNA (mtDNA) sequence variation among 654 domestic dogs representing all major dog populations worldwide. Although our data indicate several maternal origins from wolf, >95% of all sequences belonged to three phylogenetic groups universally represented at similar frequencies, suggesting a common origin from a single gene pool for all dog populations. A larger genetic variation in East Asia than in other regions and the pattern of phylogeographic variation suggest an East Asian origin for the domestic dog, ~15,000 years ago.



Trut, L. N. (1999). Early Canid Domestication: The Farm-Fox Experiment. American Scientist, 87, 160-169. [available also at: http://www.floridalupine.org/publications/PDF/trut-fox-study.pdf]


Vila, C., Savolainen, P., Maldonado, J.E., Amorim, I.R., Rice, J.E., Honeycutt, R.L., Crandall, K.A., Lundeberg, J. and Wayne, R.K. (1997). Multiple and ancient origins of the domestic dog. Science, 276, 1687-1689. [also available at: http://www.kc.net/~wolf2dog/wayne1.htm]


    * Introduction: Mitochondrial DNA control region sequences were analyzed from 162 wolves at 27 localities worldwide and from 140 domestic dogs representing 67 breeds. Sequences from both dogs and wolves showed considerable diversity and supported the hypothesis that wolves were the ancestors of dogs. Most dog sequences belonged to a divergent monophyletic clade sharing no sequences with wolves. The sequence divergence within this clade suggested that dogs originated more than 100,000 years before the present. Associations of dog haplotypes with other wolf lineages indicated episodes of admixture between wolves and dogs. Repeated genetic exchange between dog and wolf populations may have been an important source of variation for artificial selection.



Wayne, R. K. (n/d). Molecular evolution of the dog family. Retrieved June 19, 2006, from http://www.kc.net/~wolf2dog/wayne2.htm


    * Introduction: Molecular genetic tools have been used to dissect the evolutionary relationships of the dog-like carnivores, revealing their place in the order Carnivora, the relationships of species within the family Canidae, and the genetic exchange that occurs among conspecific populations. High rates of gene flow among populations within some species, such as the coyote and gray wolf, have suppressed genetic divergence, and where these species hybridize, large hybrid zones have been formed. In fact, the phenotype of the endangered American red wolf may be strongly influenced by hybridization with coyotes and gray wolves. Hybridization and habitat fragmentation greatly complicate plans to conserve the genetic diversity of wild canids.



Wayne, R. K., & Ostrander, E. A. (1999). Origin, genetic diversity, and genome structure of the domestic dog. BioEssays, 21, 247-257.


    * Abstract: Comparative analysis of mammalian genomes provides important insight into the structure and function of genes. However, the comparative analysis of gene sequences from individuals of the same and different species also provides insight into the evolution of genes, populations, and species. We exemplify these two uses of genomic information. First, we document the evolutionary relationships of the domestic dog to other carnivores by using a variety of DNA-based information. A phylogenetic comparison of mitochondrial DNA sequences in dogs and gray wolves shows that dogs may have originated from multiple wolf populations at a time much earlier than suggested by the archaeologic record. We discuss previous theories about dog development and evolution in light of the new genetic data. Second, we review recent progress in dog genetic mapping due to the development of hypervariable markers and specific chromosome paints. Extensive genetic homology in gene order and function between humans and dogs has been discovered. The dog promises to be a valuable model for identifying genes that control morphologic differences between mammals as well as understanding genetically based disease.



Zgurski, J. (n/d). The Origin of the Domestic Dog, Canis familiaris. Retrieved June 19, 2006, from http://canidae.ca/dog.htm


    * Intro: The relationship between dogs and humans dates back at least 14 000 years, and during this time, the dog has evolved to become one of the most variable animal species. Today, anywhere from 300 - 400 distinct dog breeds exist, which vary dramatically in size from three to over 150 pounds and which display an astounding amount of variation in coat type, coat colour, and general morphology. The behaviours dogs display can often vary substantially from breed to breed as well and because of this, dogs are used by humans today to perform a large variety of tasks, including herding or guarding livestock, pulling sleds, tracking or retrieving game animals, destroying vermin, locating lost people or disaster victims, and assisting handicapped people. However, as early as 14 000 years ago, there were no domestic dogs. There were wolves, coyotes, and jackals - all potential ancestors of the dog - but no dogs. The questions as to where the dog originated, the identity of the wild canine that gave rise to the dog, and, most interestingly, why and how it became domesticated and associated with humans are all rather controversial topics. This essay will focus on several aspects of the evolution of the domestic dog, and will review recent studies that have focussed on identifying the dog's closest ancestor, determining the age of the dog and discovering its location of origin. The question as to whether the dog is a result of natural or artificial selection will also be addressed.


Dog Development


Denenberg, V. H. (1964). Critical Periods, Stimulus Input, and Emotional Reactivity: a Theory of Infantile Stimulation. Psychol Rev, 71, 335-351.


Denenberg, V. H., & Morton, J. R. C. (1962). Effects of environmental complexity and social groupings upon modification of emotional behavior. Journal of Comparative and Physiological Psychology, 55(2), 242-246.


Fox, M. W., & Stelzner, D. (1966). Behavioural effects of differential early experience in the dog. Animal Behaviour, 14, 273-281.


Dog Barking


Appleby, D. Constant barking can be avoided ~ Offering guidance to owners: The Pet Behaviour Centre.


Frisby, H. (n/d). Barking: a common dog behavior problem. Retrieved 2005, from at http://www.peteducation.com/behavior/barking.htm


Moffat, K. S., Landsberg, G. M., & Beaudet, R. (2003). Effectiveness and comparison of citronella and scentless spray bark collars for the control of barking in a veterinary hospital setting. J Am Anim Hosp Assoc, 39(4), 343-348.


Sales, G., Hubrecht, R., Peyvandi, A., Milligan, S., & Shield, B. (1997). Noise in dog kennelling: Is barking a welfare problem for dogs? Applied Animal Behaviour Science, 52, 321-329.


Salzinger, K., & Waller, M. B. (1962). The operant control of vocalization in the dog. Journal of the Experimental Analysis of Behavior, 5(3), 383-389.


Schroll, S., Dehasse, J., Palme, R., Sommerfeld-Stur, I., & Löwenstein, G. (n/d). The use of DAP collar to reduce stress during training of police dogs A preliminary study. Retrieved June 4, 2006, from http://www.vet-magazin.com/wissenschaft/.../DAP-collar.html


Wells, D. L., & Hepper, P. G. (2000). The influence of environmental change on the behaviour of sheltered dogs. Appl Anim Behav Sci, 68(2), 151-162.


    * Abstract: The majority of sheltered dogs are overlooked for purchase because they are considered undesirable by potential buyers. Many factors may determine a dog's appeal, although of interest here are the dog's behaviour and cage environment which can influence its desirability. People prefer dogs which are at the front rather than the back of the cage, quiet as opposed to barking, and alert rather than non-alert. Potential buyers also prefer dogs which are held in complex as opposed to barren environments. This study examined the behaviour of sheltered dogs in response to environmental change, to determine whether it influenced dog behaviour in ways that could be perceived as desirable to potential dog buyers, and/or had any effect upon the incidence of dogs purchased from the shelter. One hundred and twenty dogs sheltered by the Ulster Society for the Prevention of Cruelty to Animals were studied over a 4-h period. The dogs' position in the cage, vocalisation, and activity were investigated in response to increased human social stimulation, moving the dog's bed to the front of the cage, or suspending a toy from the front of the dog's cage. Social stimulation resulted in dogs spending more time at the front of the enclosure, more time standing, and slightly more time barking. Moving the bed to the front of the cage encouraged dogs to this position, but did not influence activity or vocalisation. Suspending a toy at the front of the pen exerted no effect on dog behaviour, although its presence in the pen may help to promote more positive perceptions of dog desirability. The incidence of dogs purchased from the rescue shelter increased whenever the dogs' cages were fitted with a bed at the front of the pen, whenever the dogs were subjected to increased regular human contact, and whenever a toy was placed at the front of the enclosure. Findings highlight the important role that cage environment can play in shaping the behaviour of sheltered dogs and influencing whether or not an animal will become purchased.



Yin, S. (2002). A new perspective on barking in dogs (Canis familiaris). Journal of Comparative Psychology, 116(2), 189-193.


    * Abstract: The disparity in bark frequency and context between dogs (Canis familiaris) and wolves (Canis lupus) has led some researchers to conclude that barking in the domestic dog is nonfunctional. This conclusion attributes the differences primarily to genetic variation caused by domestication rather than to the influence of social environment on ontogeny. Other researchers, however, have concluded that vocal usage and response to vocalizations in mammals are strongly guided by social interactions. Closer evaluation of dog vocalizations with respect to social environment reveals developmental factors that lead to both frequent barking and barking in many contexts. Additionally, spectrographic analysis indicates that bark structure varies predictably with context, suggesting that barks can be divided into contextual subtypes and may be a more complex form of communication than given credit.



Yin, S., & McCowan, B. (2004). Barking in domestic dogs: context specificity and individual identification. Animal Behaviour, 68, 343-355. [available at http://faculty.vetmed.ucdavis.edu/faculty/bjmccowan/Pubs/Yin&McCowan.2004.pdf]


    * Abstract: In this study we sought to determine whether dog barks could be divided into subtypes based on context. We recorded barking from 10 adult dogs, Canis familiaris, of six breeds in three different test situations: (1) a disturbance situation in which a stranger rang the doorbell, (2) an isolation situation in which the dog was locked outside or in a room isolated from its owner and (3) a play situation in which either two dogs or a human and a dog played together. We analysed spectrograms of 4672 barks using macros that took 60 sequential frequency measurements and 60 sequential amplitude measurements along the length of the call. Statistical analyses revealed that barks are graded vocalizations that range from harsh, low-frequency, unmodulated calls to harmonically rich, higher-frequency, modulated calls. The harsh, low-frequency, unmodulated barks were more commonly given in the disturbance situation, and the more tonal, higherpitch, modulated barks were more commonly given in the isolation and play situations. Disturbance barks were also longer in duration with more rapid repetition than the barks given in other contexts. Discriminant analysis revealed that dog barks can be divided into different subtypes based on context even within individual dogs, and that dogs can be identified by their bark spectrograms despite the context of the bark.


Dog - Compulsive Behavior in Dogs


Luescher, A. U. (2003). Diagnosis and management of compulsive disorders in dogs and cats. Vet Clin North Am Small Anim Pract, 33(2), 253-267, vi.

Dog - Bond

Jones, A. C., & Josephs, R. A. (2006). Interspecies hormonal interactions between man and the domestic dog (Canis familiaris). Horm Behav, 50(3), 393-400.

Dog - Shelter and Rescue

New, J. C., Salman, M. D., King, M., Scarlett, J. M., Kass, P. H., & Hutchison, J. M. (2000). Characteristics of Shelter-Relinquished Animals and Their Owners Compared With Animals and Their Owners in U.S. Pet-Owning Households. Journal of Applied Animal Welfare Science, 3(3), 179-201.

Dog - Other


Fox, M. W., & Stelzner, D. (1966). Behavioural effects of differential early experience in the dog. Animal Behaviour, 14, 273-281.


Gacsi M, Gyori B, Miklosi A, Viranyi Z, Kubinyi E, Topal J, Csanyi V. (2005). Species-specific differences and similarities in the behavior of hand-

raised dog and wolf pups in social situations with humans. Dev Psychobiol. 47(2):111-22.

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