Reply to Makous
Reply to Makous 1
GERALD S. WASSERMAN, University
Wisconsin, Madison, Wis.53706
01
Makous (1970) makes several comments
about our earlier paper (Wasserman &
Patton, 1969) that oversimplify the
situation and require a reply. First, Makous
is persuaded that Limulus reliably moves
i t stall down ward in response to
stimulation. This is not the case: In our
work, the tail moved downward in
response to light and upwards in response
to shock. The upwards movements
occurred for weak (0.1 mA) as weil as
strong (50 mA) shocks. We used a powerful
shock simply because rapid habituation
was observed for weak shocks. In Smith &
Baker's (1960) work, the tail always moved
upward in response to both light and shock
(Baker, personal communication). So the
direction of tail movement is not invariant.
Second, and more importantly, there is
the question of the probability of a tall
movement, irrespective of its direction.
Makous gives the impression that this
"natural part of the animal's escape
reaction" is a well·known and invariant
consequence of stimulation. Had this been
the case, it is very hard to understand why
any of us would have tried to use tail
movement as the response in a
conditioning experiment. However, at the
time of our report, the experimental
evidence was that this response does not
occur with any great frequency. In Smith
an d B aker's controls, the response
frequency was about 30%. In Makous's
184
(1969) second classical conditioning
experiment, the response frequency was
about 2.5%. In our first set of sensitization
and pseudoconditioning controls, the
response frequency was also about 30%.
These first control experiments were
completed prior to the appearance of our
re port in this journal, although they were
not mentioned in that report. Had our
control experiments turned out otherwise,
we would have withdrawn our report prior
to publication.
Subsequen tly, however, and in
contradiction to both the earlier reports
and our own experience, we have found
that some, but not all, naive Limulus
exhibit a tail movement in response to light
alone, that this response can occur with
frequencies weil above 75%, and that it
persists without habituation for periods of
several months. This behavior seems to be
more profitably viewed as a
Hxed-action-pattern (F AP) that may or
may not be expressed, depending on
conditions that are not yet understood. In
another paper (Wasserman & Patton, in preparation), we have described these results in
detail, as weil as the use of the FAP to
obtain a visual threshold. There is nothing
in the experimental literature that might
have suggested that such an obvious and
persistent F AP involving tai! movement
could exist; however, analogous FAPs
involving leg movements have been
recently reported by Corning & von Burg
(1968), although they report rapid
habituation of these leg-movement FAPs.
Makous (1970), then, is certainly correct in
questioning our report of successful
conditioning, but for reasons other than
those expressed. The essential point here is
not that our work lacked the relevant
controls, but that the relevant control
procedures have given us different results
with different groups of animals. The fact
that some animals exhibit a high
probability of tail movement to light alone
renders it quite likely that the effect of
pairing light with shock potentiates an
otherwise unexpressed tail movement in
animals that exhibit a low prob ability of
tall movements to light alone. At this time,
it is our belief that the existence of this
F AP involving tail movement controverts
all prior reports of associative conditioning
in Limulus, and requires that we all change
our ideas on this subject.
REFERENCES
CORNING, w. C., & von BURG, R. Behavioral
and neurophysiological investigations of
LimuluJ polyphemus. In J. Salanki (Ed.),
Neurobiology of invertebrates. New York:
Plenum, 1968.
MAKOUS, W. L. Conditioning in the horseshoe
crab. Psychonomic Science, 1969, 14,4-6.
MAKOUS, W. L. Conditioning in Limulus:
Comment on Wasserman and Patton.
Psychonomic Science, 1970, 00, 000-000.
SMITH, J. c., & BAKER, H. D. Conditioning in
the horsehoe crab. Journal of Comparative and
Phsyiological Psychology, 1960, 53,279-281.
WASSERMAN, G. S., & PATTON, D. G.
Avoidance conditioning in Limulus.
Psychonomic Science, 1969, 15, 143.
NOTES
1. This research was supported in part by the
Wisconsin Alumni Research Foundation, in part
by the Biomedical Sciences Support Grant
administered by the University Graduate
Research Committee, and in part by a grant from
the National Science Foundation (GB-8581).
2. Wasserman, G. S., & Patton, D. G. Visually
guided behavior in Limulu:: A fIXed-action
pattern elicited by light. In preparation.
psyehon. Sei., 1970, Vol. 19 (3)
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