Motivational Systems of Agonistic Behavior in Muroid Rodents
Motor Patterns of Offense Page 5



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...motor patterns
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...motivating stimuli
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Four principal motor patterns have been associated consistently with offense in most muroid rodent species in which fighting behavior has been thoroughly described: Approach, bite-and-kick attack, offensive upright posture, and offensive sideways posture. These behaviors, and their variants which depend upon the extent to which they are combined with approach locomotor components, have been given a confusing variety of names, as catalogued in Table I. In this paper I will follow the terminology of Grant and Mackintosh [1963].

The bite-and-kick attack has been described in detail from motion picture analysis in the mouse [Banks, 1962] and rat [Miczek, quoted by Adams, 1976]. It consists of an approach to the opponent which culminates in the full aggressive posture with the attacker surmounting his victim at right angles across the back; from this posture, a bite on the opposite flank and simultaneous kick with the hind legs is usually delivered. An equivalent attack may also be delivered in some cases from an offensive sideways posture, in which case it may consist of a bite upon the flank of the opponent which faces the attacker [Rlanchard and Rlanchard, 1977]. When combined with chasing of a fleeing opponent, it may consist of a bite upon the rump of the fleeing victim or an attack leap onto it without any preliminary full aggressive posture.

The offensive upright posture and offensive sideways posture were described and illustrated for laboratory rodents by Grant and Mackintosh [1963]. In the former "the animals stand on their hind legs, normally presenting their ventral aspects to the other animal (and) occasionally when in this posture the animals push at each other with their fore-paws"[See Figure 2]. In the latter, "the animals orientate themselves broadside on to their opponents." Grant and Mackintosh [1963] distinguished between offense and defense versions of these postures. In our own work we have found that the offense postures involve locomotor initiation and approach, whereas the defense postures involve locomotor reaction and withdrawal [Lehman and Adams, 1977]. It has also been noted that during boxing of two animals in upright postures, the offensive animal tends to push forward, while the defensive animal tends to be more upright or leaning backwards [Davis, 1972].

I have suggested previously that the sideways and upright postures may be low and high intensity motor patterns, respectively, produced by a single motor patterning mechanism which may be called a "high posture" motor patterning mechanism [Lehman and Adams, 1977].

Whereas approach, bite-and-kick attack, and offensive upright posture have been described in every well-studied genus of muroid rodents, the offensive sideways posture is not always found. In addition to its apparent absence in some species of both murid and cricetid rodents (Notomys alexis, Apodemus sylvaticus, Lemmus lemmus, L trimucronatus, Sigmodon sp. and D groenlandicus), it is also less commonly observed in species of Microtus and Peromyscus. The study of various Microtus sp by Colvin [1973] failed to report this motor pattern, although it was described by Clarke [1956] in M agrestis. The studies of various Peromyscus sp by Eisenberg [1962, 1963] failed to find offensive sideways posture, although it had been described by Ralph and Stokes [1960] in P maniculatus, and was included in Eisenberg's 1968 review.

In some species without offensive sideways posture, it is possible that its place is taken by a similar posture which we may call an offensive quadrupedal posture. Under the name "stalking" it has been described as follows in N alexis by Stanley [1971]: " A furtive slow quadrupedal approach towards another animal, made by the aggressor. During stalking the eyes are open and the ears are pricked forward. The neck is stretched out, and the fur on the nape may be raised." A similar behavior has been called "threat" or "moving threat" in Peromyscus [Balph and Stokes, 1960; Eisenberg, 1962] and Apodemus [Montgomery, 1978].

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