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Dec 31 / Lucas Brands

A Critical Analysis of the Canine Predatory Motor Sequence

Lucas Brand

In their book ‘Dogs: a new understanding of canine origin, behavior and evolution’ Coppinger and Coppinger (2001) explain that motor patterns are innate complex reflexive behaviours evoked by a releaser and have a species specific, stereotypic, shape. They state that the distinctiveness of these patterns means biologists can determine which predator killed the prey by looking at the carcass and argue that motor patterns don’t differ in kind amongst species or breeds, but rather differ in the frequency of expression. The full wolf predatory motor sequence (PMS) is said to be; orient > eye > stalk > chase > grab-bite > kill-bite > dissect > consume. This sequence is used as template for that of the domestic dog. But is the sequence as reflexive as they state? And is this PMS template useful for us as dog trainers and behaviourists?
The predatory motor sequence, how about dogs?
According to Coppinger and Coppinger (2001) there is one predatory motor sequence for general carnivores, which is adapted by species as different niches. The wolf-PMS, mainly based on groups hunting for ungulates, is an adapted, niche, version of that of a general carnivore.

But wolves also hunt for smaller animals such as birds or hare. How would this affect the predatory motor sequence? Within the species-specific PMS, patterns can be substituted. For example the chase pattern could be substituted by a fore-foot stab, used to hunt small critter, creating an altogether different sequence for a different type of hunting-purpose. They also explain that they can start the sequence at any pattern, meaning a wolf can start the sequence at stalk or chase, without first going through the motions of orient and eye.
Mainly through the works of Coppinger and studies based on their work, the wolf predatory motor sequence is used as a template for the PMS in the modern domestic dog. The PMS in dogs is said to be atrophied or hypertrophied through artificial selection, making them specialists, as it were, in certain patterns within the sequence. For example Border Collies have exaggerated eye>stalk>chase and Anatolian Shepherds an inhibited sequence of (orient) (eye) (stalk) (chase) (Coppinger, 2001., Udell, 2014).
But wolves and domestic dogs are not one and the same. All dogs share a common ancestry which is distinct from modern wolves, though they do have a common ancestor which is now extinct (Bergstrom et al, 2020). When looking at wolves, we are not looking at our domestic dog’s ancestor, which means we can’t say the wolf PMS is the ancestral PMS in dogs as Coppinger and Coppinger (2001) claim it to be. Modern wolves not being domesticated does not indicate they are more like their ancestor, the environment to which they have adapted might have changed as much as that of the dog.As Miklosi (2007) states, a wolf raised and socialized in a dog’s environment does not become dog-like and feral dogs do not show wolf-like behaviours. One of the differences between modern wolves and domestic dogs lies in their early development during the critical period of socialization, which impacts hunting and hazard avoidance, amongst other factors, and is likely to impact adult behaviour profoundly (Lord, 2012). I believe it is safe to assume that these factors indicate that comparing the PMS of wolves with that of dogs might be problematic. How can we separate the differences, both genetic and environmental, between dogs and wolves if we don’t have enough data about their common ancestor?

One way to overcome this issue is through the use of a ethocognitive model, as this can help us observe and research the role and interaction between genetic and environmental factors. The model looks at how genetic and environmental components occur and are intertwined, from the perceptual information input, via a referential system, to elicit an action system (behavioural output) such as a modal action pattern. This system is continuously updated and modified by exploring and monitoring the environment. Looking at which genetic and environmental factors at what stage within the ethocognitive model shape the occurrence of motor patterns might help us design experiments to find out differences and similarities in dog and wolves (Miklosi, 2007).

It is about working with our dogs, in environments where the releaser is present and teaching them to only engage in their breed specific motor patterns on our cue. 

Shaping the Predatory Sequence

Lord et al (2016) explain that training is about directing the dog to the environment where the PMS is elicited and that the emergence and intensity of displaying these patterns varies in different breeds. As an example they say that it is difficult to train a corgi not to nip at hocks. They argue that the only way to prevent a dog from showing their breed-specific patterns, is to remove them from the stimuli which elicit it. In a study by Howell et al (2020) aimed at measuring expert opinions on the management and expression of predatory behaviour in retired greyhounds it is said that predatory behaviour is difficult to prevent due to its self-reinforcing nature. They also indicate that never giving the dog the opportunity to engage in the behaviour is the best preventative method. This seems contrary to dog training and behaviour practice.

Training is not about either directing the dog to the environment that elicits the PMS, or preventing them from it. It is about working with our dogs, in environments where the releaser is present and teaching them to only engage in their breed specific motor patterns on our cue. A Spaniel’s PMS, for example, does not include ‘don’t flush the pheasant on left to prevent walking in shot range’.

As Miklosi (2007) pointed out, there is an interaction between genetic and environmental factors. Klein (2019) explains that even in early ethological research, Lorenz stated that only the pattern of consummatory behaviour is fixed, but that conditioning can alter the effectiveness or sensitivity to the releaser stimulus. New releasing stimuli and new behaviour patterns could even be conditioned, which increases an animal's adaptive success. This means the hard-wired behaviour patterns, such as the PMS, are at least partially shaped by experience, something which a study by Udell (2014) revealed as well. They compared three breeds with distinct predatory motor pattern sequences on their performance in human guided tasks and showed that the outcome is not only predicted by the predatory motor sequence, but also by lifetime experiences such as training. Although the PMS can be an important factor in the succes between different breeds in human guided tasks, their physical attributes such as size or visual acuity and lifetime experiences such as training or the lifestyle of the owners, serve to compound the effect. They state;“we demonstrated that explicit experience can be used to overcome breed-specific predispositions in some cases” – Udell et al, 2014.

FAPs, MAPs or non-random behaviour

Gadbois (2014) argues that early ethological models such as fixed or modal action patterns (FAP, MAP) are too simplistic to describe more complex behaviour patterns. According to them, modal action patterns have a restraint in expressive freedom due to their length, on average a MAP lasts 0.1 – 10 seconds and is triggered by simple sign stimulus (releaser). This means a lot of behaviour patterns we now call MAPs are not that at all. One of them being the predatory motor sequence in dogs, which is different to a modal action pattern in terms of duration and complexity. Even though highly recognizable and not random, there is a large degree of freedom to adapt to, for example, the movement of prey. According to Gadbois there is a realm of behaviour followed by a releasing stimulus possible, with significant variation and probability of occurrence. On one side there’s the 100% predictable reflexive behaviours, on the other the completely random behaviours with 0% predictability. Behaviour patterns can be short, long, simple, complex, deterministic or almost appear random. Fixed or modal action patterns are on the simple, short and deterministic side, but there are other kinds of non-random behaviour sequences which are less predictable. To look at predictability and freedom within sequences, they studied caching behaviour in canids. They found that caching sequences are not FAPs or MAPs due to their length and complexity, but that there is a dynamic intrinsic structure in the expression of the caching behaviour, which is species specific. According to Gadbois, looking at behaviour as a system of rules gives us the flexibility to look at sequences rather than strict or slightly adaptable patterns.

Apart from a new way of looking at complex and long behaviour patterns in terms of non-random sequences, they also unintentionally bring another point to the discussion around the validity of Coppinger and Coppinger’s template of the wolf PMS in dogs. 
Caching behaviour is not a simple substituting one pattern for another, but follows a whole different kind of sequence with different patterns which are still non-random, followed by a releaser and part of predation, but does not fit the PMS as widely used and described. But what drives an animal to respond to the releaser if the behaviour isn’t 100% reflexive?

Conscious choice, motivational control and affect
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Another variable which impacts behaviour is motivation. Motivation can vary over time and is influenced by the physical state of an animal. It’s responsible for setting high-level goals, such as chasing a certain prey, but the implementation of these goals is under control of lower-level processes, from reflexive actions to activation of motor neurons. Modal action patterns, amongst which the predatory motor sequence, are under motivational and emotional control. This control modulates the link between the releasing stimulus and the response (motor pattern). Motivations which are dominant inhibit the capacity of less dominant motivations (Toates, 2002). To give an example, if an animal has just eaten, the motivation to rest will abolish the motivation of predation in the presence of prey. In Applied Behaviour Analysis this concept is called a Motivating Operation (Langthorne and McGill, 2009).

Motivation can also be understood in terms of the SEEKING system, described by Wright and Panksepp (2012) as ‘psychomotor eagerness to obtain resources’. Though the PMS is likely under control of the SEEKING system, this system is said to allow for an adaptive variety of behaviour to engage with the environment. According to Howell (2020) this could possibly allow us to train alternative behaviours for the dog to perform when encountered with the releaser. Though motor patterns are self-reinforcing, and do not need an external reward to occur, the animal does need to make the conscious decision to act. They don’t just do the sequence, they make a cost-benefit analysis. This indicates that even though going through the motions of the PMS is automatically reinforcing, the dog does need to be willing to pay for the incentive (Spruijt, 2001). Knowing whether or not making a choice is worth it, relies on a learning history and with that its consequences.

Conclusion


The PMS in dogs as described by Coppinger and Coppinger (2001) seems to me to be too fixed in nature, based on the PMS of wolves, without much looking into the differences between wolves and dogs. It lacks the flexibility to include other type of predatory behaviours seen in canids, such as caching, and does not allow for the influence of environmental factors and how these elements are intertwined. Most work on the PMS in dogs refers to or is done by or with Coppinger, which means the same information is being re-used often. I was not surprised to find that studies like that of Lord et al (2016), which makes rather bold statements about trainability, include Coppinger as one of the authors. To me, training (working) dogs seems to be part of conditioning new releasers or decreasing their salience, which then becomes part of the cost-benefit analysis the dog makes before engaging in the behaviour. Another point of interest is that most studies concerned with the PMS in dogs, use working dogs which are bred for function. It raises the questions; What kind of socialization did these dogs have? The greyhounds in Howell’s (2020) study for example were not socialized with small animals. And how about the PMS in pet dogs? I also wonder how much of the selection for a hypertrophied or atrophied PMS is about temperament. For a lot of working breeds, cooperation with the handler is key. A collie or gundog who does not cooperate with the handler, is not a good working dog.

These factors taken together are why I personally tend to lean towards Gadbois's (2014) argument that ethological models, that of FAP and MAP, are too simplistic and find the theory of more flexible natural sequences with varying degrees of freedom more compelling. Taking into account other factors such as temperament, SEEKING, motivation and learning history seems to me more in line with what we experience training and living with dogs.

References

  1. Bergström, A., Frantz, L., Schmidt, R., Ersmark, E., Lebrasseur, O., Girdland-Flink, L., Lin, A., Storå, J., Sjögren, K., Anthony, D., Antipina, E., Amiri, S., Bar-Oz, G., Bazaliiskii, V., Bulatović, J., Brown, D., Carmagnini, A., Davy, T., Fedorov, S., Fiore, I., Fulton, D., Germonpré, M., Haile, J., Irving-Pease, E., Jamieson, A., Janssens, L., Kirillova, I., Horwitz, L., Kuzmanovic-Cvetković, J., Kuzmin, Y., Losey, R., Dizdar, D., Mashkour, M., Novak, M., Onar, V., Orton, D., Pasarić, M., Radivojević, M., Rajković, D., Roberts, B., Ryan, H., Sablin, M., Shidlovskiy, F., Stojanović, I., Tagliacozzo, A., Trantalidou, K., Ullén, I., Villaluenga, A., Wapnish, P., Dobney, K., Götherström, A., Linderholm, A., Dalén, L., Pinhasi, R., Larson, G. and Skoglund, P., (2020) Origins and genetic legacy of prehistoric dogs. Science, 370(6516), pp.557-564

    Coppinger, R. and Coppinger, L. (2001) Dogs; A new understanding of canine origin, behavior, and evolution. Chicago: University of Chicago Press, Chapter 6.

    Jensen, P., (2007) The behavioural biology of dogs. Wallingford, Oxfordshire: CABI International, pp.61-75.

    Gadbois, S., Sievert, O., Reeve, C., Harrington, F. and Fentress, J., (2015) Revisiting the concept of behavior patterns in animal behavior with an example from food-caching sequences in Wolves (Canis lupus), Coyotes (Canis latrans), and Red Foxes (Vulpes vulpes). Behavioural Processes, 110, pp.3-14.

    Howell, T. and Bennett, P. (2020) Preventing predatory behaviour in greyhounds retired from the racing industry: Expert opinions collected using a survey and interviews. Applied Animal Behaviour Science, 226, p.104988.

    Klein, S. (2019) Learning: Principles and Applications. 8th ed. Sage Publications Inc, Chapter 2.

    Lord, K., 2012. A Comparison of the Sensory Development of Wolves (Canis lupus lupus) and Dogs (Canis lupus familiaris). Ethology, 119(2), pp.110-120.

    Langthorne, P. and McGill, P. (2009) ‘A Tutorial on the Concept of the Motivating Operation and its Importance to Application’, Behavior Analysis in Practice, 2(2), pp. 22–31.

    Lord, K., Schneider, R. and Coppinger, R. (2016) Evolution of working dogs. The Domestic Dog; Its Evolution, Behavior and Interactions with People, pp.42-66.

    Miklosi, A. (2007) Dog behaviour, evolution, and cognition. Oxford University Press, Chapter 1.

    Shan, S., Xu, F. and Brenig, B. (2021) Genome-Wide Association Studies Reveal Neurological Genes for Dog Herding, Predation, Temperament, and Trainability Traits. Frontiers in Veterinary Science, 8.

    Spruijt, B., van den Bos, R. and Pijlman, F. (2001) A concept of welfare based on reward evaluating mechanisms in the brain: anticipatory behaviour as an indicator for the state of reward systems. Applied Animal Behaviour Science, 72(2), pp.145-171.

    Toates, Frederick (2002). Physiology, motivation and the organization of behaviour. In: Jensen, P. ed. The Ethology of domestic animals: an introductory text. Wallingford, UK: CABI Publishing, pp. 31–50.

    Udell, M., Ewald, M., Dorey, N. and Wynne, C. (2014) Exploring breed differences in dogs (Canis familiaris): does exaggeration or inhibition of predatory response predict performance on human-guided tasks?. Animal Behaviour, 89, pp.99-105.

    Wright, J. and Panksepp, J.,(2012) An Evolutionary Framework to Understand Foraging, Wanting, and Desire: The Neuropsychology of the SEEKING System. Neuropsychoanalysis, 14(1), pp.5-39.
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