Brain Mechanisms for Offense, Defense, and Submission
Author's Response:
Specific questions concerning offense, defense, and submission
Page 54

Title/Abstract page

Pages 1 - 2

Defense: motivational mechanism
Page 3

Defense: motivating stimuli
Pages 4 - 5

Defense: motor patterning mechanism
Page 6

Defense: releasing & directing stimuli
Page 7

Pages 8 - 9 - 10

Pages 11 - 12

Primitive mammals & primates
Page 13

Pages 14 - 15 - 16

Figure 1: Defense
Page 17

Figure 2: Submission
Page 18

Figure 3: Interaction
Page 19

Figure 4: Offense
Page 20

Figure 5: Composite
Page 21

Open Peer Commentary
Pages 22-49

Author's Response:
motivational systems

Pages 50 - 51 - 52

Author's Response:
alternative analyses

Page 53

Author's Response:
specific questions

Pages 54 - 55 - 56

Author's Response:

Page 57

References A-E
Page 58

References F-M
Page 59

References N-Z
Page 60


Page 61

Specific questions concerning offense, defense, and submission. Several commentators agree that the distinction between offense and defense is a legitimate one, and the Blanchards and Brain provide new observations that strengthen the basis for the distinction. The Blanchards note that offense and defense are quite distinct along a number of dimensions and that in a fight between a dominant rat and a strange intruder, there is virtually no overlap between the offense of the former and the defense of the latter. Brain adds new data to the effect that offensive and defensive attacks may be distinguished on the basis of bite targets in mice.

Other commentators continue to doubt that offense and defense may be clearly distinguished. Leyhausen, for whom I originally learned the distinction between offense and defense, seems to consider that 1 have overdrawn the distinction. Panksepp wants to consider aggressive behavior as the result of "nonspecific" systems in contrast to my "specific" systems. He is right that I consider that offense and defense may be activated simultaneously, as in the case of the lactating female. Yet, even so, the result is not a hybrid behavior, but an alternation of offense and defense motor patterns. Panksepp also suggests that one can see a combination of offense and defense in cats during brain stimulation, but I am doubtful of this. As Miczek points out, most of the experimental work on neural mechanisms of aggression in cats actually involves defense, and Flynn (1976) also points out that offense is not normally seen in response to brain stimulation in cats. In my own experience with brain stimulation in cats (Adams and Flynn 1966; Adams 1968; Adams, Bacelli, Mancia, and Zanchetti 1969), I have never seen behavior corresponding to the "Angriffsreaktion" described by Leyhausen, which I have translated as offense. With this in mind, I cannot agree with Decsi & Nagy that "offense" characterizes the behavior of cats in a desperate situation. Rather, I think they refer to what I am calling defense. Wiepkema also does not believe that offense and defense are separate systems, but considers that they may be "intermingled" in different situations. As I mention above, his example of offense and defense by a lactating female does not in my opinion demonstrate such intermingling.

The distinction between defense and submission is particularly important in the present analysis. Therefore, I will take some care to respond to objections by a number of commentators, and especially by the Blanchards, who have contributed greatly to the literature on the behaviors of these systems.

The ultimate criterion for distinguishing defense and submission must be their neural circuitry, but since that is still in dispute, I must rest my claim on five types of less direct evidence. Because they are indirect they are open to the charges by Waldbillig and Brain that the definition is circular. No one of these types of evidence is indispensable. For example, contrary to Ursin's interpretation, I did not intend to base the distinction upon only one criterion such as the difference in behavior between wild and tame animals, and it is a misinterpretation when Fentress says that submission "apparently evolved in the lab." Rather, defense is observed more often in wild animals and submission more often in tame or laboratory animals. The distinction is based on five types of evidence: (1) differences between wild and tame animals; (2) an evolutionary argument on differential responding to conspecifics and to predators; (3) distinctions in the cat between affective defense (defense) and flight (submission); (4) an analysis of the function of forebrain structures in the emotional behavior of the rat; and (5) differential hormonal effects. Unfortunately, the hormonal evidence, which I think is most convincing of all, could be presented only briefly in the target article. As mentioned above, a more extensive treatment has been submitted for publication.

The Blanchards doubt that wild and laboratory rats differ in the relative strengths of defense and submission. They cite new unpublished evidence that wild and domestic rats have "consistent similarities" in defense and submission. My own unpublished observations are quite different; wild rats were much more likely than domestic rats to show a lunge-and-bite defense when attacked by another rat, and they were much less likely to show the full submissive posture. Ursin (1964) has reported that in response to provocation tame cats show only flight, which I interpret as submission, while wild cats show a lunge-and-bite attack and flight, which I interpret as defense. The situation in rats needs further clarification.

As to the evolutionary argument that submission inhibits conspecific but not predator attacks, the Blanchards doubt that ultrasound and full submissive posture inhibit conspecific bite-and-kick attack in the rat. They doubt the experimental evidence that ultrasound inhibits attack. In reply, I point to my own data (Lehman and Adams 1977) as well as those of others (Lore, Flanelly, and Farina 1976). At one point the Blanchards concede that ultrasound might inhibit conspecific attack; if so, would they expect it to inhibit the attack of a predator as well? As part of the same discussion, they raise what I consider to be a straw-man objection as to the way that the full submissive posture inhibits bite-and-kick attack by a conspecific. Their interpretation is no different from my own, as implied in the target article and as explicitly stated in an earlier paper (Adams 1976). In other, words, contrary to their statement, I also consider that the full submissive posture inhibits the bite-and-kick by denying the opponent access to the dorsal surface, which is a necessary releasing stimulus for the bite-and-kick attack. Returning to the original question, would they expect the same posture to inhibit the attack of a predator?

(Section continued on next page)

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