i* \ V Genetic aspects of dog behaviour 4 with particular reference to working ability M. B. WILLIS genetic factors, they will also be modified through Introduction interactions with the environment. Anyone using dogs as working animals has an obvi- Humphrey & Warner (1934) calculated pheno- ous interest in knowing: (a) the degree to which typic correlations among 42 physical and 9 behav- specific features are inherited; (b) the magnitude of ioural traits in GSDs, and found that 15 of the 378 the differences which exist between breeds; and (c) possible correlations reached statistical significance. the relationship, if any, between specific traits. It is probable that the underlying genetic correlations Although the dog has had a longer association, and were quite different and that the project was over- enjoys closer contacts, with humans than any other simplified in seeking Mendelian explanations. Never- species, genetic studies on the domestic dog are not theless, the Fortunate Fields study (Humphrey & extensive. Most genetic work has been directed Warner, 1934) did achieve success in producing towards understanding the mode of inheritance of superior animals for guide dog/police work using specific physical anomalies, whereas advanced studies this simplified system. on the inheritance of behaviour are relatively recent The present paper looks at behaviour in guide and stem largely from the pioneering work that led to dogs, hunting dogs, livestock guarding dogs and the publication of Scott & Fuller (1965). Mackenzie, police/service dogs. The importance to man of herd- Oltenacu & Houpt (1986) provided a broad review ing dogs is without question, but so few genetic data of behaviour genetics of the dog, but the present exist in this field that herding dogs are deliberately review focuses specifically on the genetics of working excluded from the present discussion. behaviour, since it is in this field that behaviour stud- ies are likely to have the greatest practical impact. Guide dogs for the blind The role of the guide dog must rank as one of the Background most useful modern occupations for a working dog. As Mackenzie et al. (1986) have shown, studies on For a long time the GSD was the breed of choice, the inheritance of behaviour began around the turn but in more recent years Labrador and golden of the century, but much of the early work was con- retrievers, and their crosses, have been commonly fined to well-established traits with a clear intention used (male GSDs being slightly too large for the of finding Mendelian explanations. Thus we see task). Many guide dog organizations now breed their Whitney (1929/?) suggesting that, in foxhounds, vocal own stock (that in Britain being the largest dog 7 trailing is dominant to mute trailing. Similarly, owning body in the country ), and studies from these Humphrey & Warner (1934) suggested that gunshy- organizations are now coming to fruition. ness in German Shepherd dogs (GSDs) was con- Most guide dog failures arise from character faults, trolled by a simple gene series with two alleles: N particularly fearfulness (Scott & Bielfelt, 1976; God- causing under-sensitivity, and n causing over sensi- dard & Beilharz, 1982). Clear sex differences have tivity. Thus NN animals were largely insensitive to also been noted: females, for example, show more sound, Nn were medium sensitive and nn oversensi- suspicion and fearfulness, while males show more tive. In addition to auditory sensitivity, these work- initiative. When assessing the degree to which a ers also looked at sensitivity to touch and postulated character or trait might be inherited an important a similar theory with SS being undersensitive, Ss consideration is its heritability. Strictly, heritability medium sensitive and ss oversensitive. They con- is the proportion of the total variance observed in a sidered that ss and nn animals were too sensitive to trait that is due to additive effects. In lay terms, it train effectively, and that the best animals to train might be best defined as that proportion of the par- were Nn/Ss. While most trainers would agree with ental superiority (over the population average) which the difficulties experienced in trying to train over or is transmitted to the offspring. Thus, a heritability of undersensitive animals, it is highly unlikely that such 40% would suggest that only 40% of any parental complex features as ear and body sensitivity are superiority (or inferiority) would be passed on to the going to be inherited as simple Mendelian traits. Not progeny. Heritability studies are usually based upon only will such sensitivities be influenced by complex paternal half-sib correlations, i.e. correlations Genetic aspects of behaviour 53 Table 4.1. Heritability estimates for guide dog traits (Australia) Trait Sire Dam Combined h2 SE h 2 SE h 2 SE Success 0.46 0.19 0.42 0.18 0.44 0.13 Fear 0.67 0.22 0.25 0.15 0.46 0.13 Dog distraction -0.04 0.08 0.23 0.14 0.09 0.08 Excitability 0.00 0.09 0.17 0.13 0.09 0.08 Source: Goddard & Beilharz (1982). between the performances of different progeny sired females, were produced by Bartlett (1976) and are by a particular father. For such studies a series of given in Table 4.2. In many cases values did not differ sires needs to be used but the performance of the sire significantly from zero, but some aspects of sensi- is not required, only that of his progeny. Alterna- tivity were moderately heritable. Studies on Cali- tively, offspring-parent regressions can be made, in fornian guide dogs (mainly GSD) have been pub- which case the performance of both offspring and lished by Scott & Biefelt (1976) and are shown in parents needs to be known. These offspring-parent Table 4.3. In 11 of the 13 traits, dam components regressions can be based upon one parent (usually attained higher heritabilities than sire components the sire) or the average of both parents (termed off- emphasizing the importance of maternal effects. spring-mid-parent regressions). A difficulty with However, most of the heritabilities were not signifi- such methods is that parents and offspring are, of cantly different from zero. necessity, assessed in different years, even if assessed Both American studies suggest lower heritabilities at comparable ages. for major traits than do the more recent Australian Heritability studies are, strictly speaking, only rel- studies, but the reasons for this are unclear. The Aus- evant to the population from which they were tralian figures derive from more sophisticated statisti- derived and for the period of time when they were cal analyses than the American ones, but one cannot assessed. Nevertheless, they can provide a broad guide to inheritance. Because heritabilities are ratios they are susceptible to variation caused by environ- Table 4.2. Heritability estimates for guide dog traits mental features. Failure to reduce environmental (USA) variation or to assess animals in a consistent fashion will tend to reduce heritability estimates and thus Trait Male Female Both produce values that may be lower than the true fig- 1 Body sensitivity 0.26 0.05 0.10 ures. Despite these limitations, heritabilities are Ear sensitivity 1 0.49 0.14 0.25 important because they give guidelines to the conse- 1 Fighting instinct -0.05 -0.08 -0.04 quences of various selection procedures. Highly heri- Protective instinct* -0.21 -0.13 -0.12 table traits should respond well to direct selection Nose acuityn 0.30 0.05 0.12 " testing, low values may 1 1 and performance whereas Intelligence -0.17 -0.07 -0.06 or even the ' necessitate selection through progeny, Willingness11 -0.14 -0.04 -0.03 " genetic advances. 1 use of crossbreeding, to produce Energy 1 -0.03 0.06 0.05 394 Australian " Heritability estimates derived from Confidence1 1 0.04 0.26 0.16 guide retrievers) are shown in Table dogs (Labrador Self rightm 0.15 0.25 0.22 4.1. They were generally high for 'success at becom- ing a guide dog' and for 'fear' (the principal cause f low score least effect; ft l°w score greatest effect; of culling) (Goddard & Beilharz, 1982). Heritability ttt l°w score indicates most eager to submit to estimates from American guide dogs (various another. breeds), based on over 700 males and over 1000 Source: Barlett (1976) 54 Part II: Behaviour and behaviour problems Table 4.3. Heritability estimates for guide dog traits 4.5. The Australian work suggests high heritabilities (California) for fear or 'nervousness', and strong positive corre- lations between this and 'sound shyness' and negative 2 Trait Feature h correlations with 'willingness'. Most practical breed- ers believe nervousness to be relatively strongly Sit Forced sit with 3 0.06 inherited, and there is empirical evidence in many repetitions breeds that breeding from nervous dogs leads to the Come Called with 5 repetitions 0.14 production of increased proportions of nervous pro- Fetch Playful retrieving 3 0.24 geny (see below and Serpell & Jagoe, Chapter 6). repetitions Trained response Complex excitability, 0.08 nervousness Hunting Willingness Responsiveness to tester 0.12 Body sensitivity Complex reaction to pain 0.16 Considering that man probably first used the dog as Ear sensitivity Complex reaction to sound 0.00 an aid to hunting (see Clutton-Brock, 1984 and New experience Response to novel stimuli 0.06 Chapter 2), we have learned very little about hunting Traffic Reaction to moving cart 0.12 attributes in genetic terms. Whitney's (\929a, b) early Footing crossing Ability to identify surface 0.06 work on hunting traits was largely confined to rela- underfoot tively simple behavioural features, whereas cross- Closeness How close passed to 0.04 breeding work by Marchlewski (cited in Burns & obstructions Frazer, 1966) showed that pointing behaviour was a Heel Acceptance of leash 0.10 complex trait with no indication that the progeny of training superior field-trialists were any better than average.