Adv. mar. Biol., Vol. 3, 1966, pp. 1-62
LEARNING BY MARINE INVERTEBRATES
I.
11.
111.
IV.
V.
VI.
VII.
VIII.
IX.
X.
M. J. WELLS
Department of Zoology, Downing Xtreet, Cambridge, England
Introduction
. . . . . . .. .. ..
A.
Experiments on Sensory Integration
. .
Annelids
..
.. .. .. .. ..
Cephalopod Molluscs . . .. .. .. ..
B.
Brain Lesions and Learning . . .. ..
A.
Habituation to Repeated Stimuli . . ..
B.
C.
Brain Lesions and Polychaete Learning . .
Crustaceans-and Limulus
. . .. .. ..
A.
B.
Other Evidence that Crustaceans Learn . .
Further Experiments with Errant Polychaetes
The Results of Formal Training Experiments
C.
Learning by Limulw polyphemus . . ..
Gastropods and Lamellibranchs
. . ..
..
A.
B.
Homing by Limpets . . .. .. ..
C.
Other Evidence of Learning by Molluscs . .
Platyhelminthes . .
.. .. .. .. ..
Echinoderms
.. .. .. .. .. ,.
Coelenterates . . .. .. .. ,. ..
Discussion
.. .. .. .. .. ..
References
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.. ..
Formal Training Experiments with Gastropods
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.. 1
.. 2
.. 3
.. 15
.. 20
.. 21
.. 26
.. 28
. . 29
.. 29
.. 33
.. 37
. . 37
.. 37
.. 38
.. 41
.. 42
.. 44
.. 47
.. 51
.. 85
I. INTRODUCTION
It would clearly be ridiculous to limit a review of invertebrate
learning to marine forms were the object to assess the relevance of
research on invertebrates to the study of learning generally. This
review does not attempt to do anything so ambitious. Its more limited
objectives are to show how much marine forms have to offer for work
on learning and how very little has been done with most of them. The
hope in doing this is to persuade zoologists and psychologists that the
existing literature is more than a mere anecdotage and that enough is
now understood to justify much more intensive research ; the range of
material is vast, and much of it is very promising indeed.
In what follows the experiments surveyed are dealt with group by
animal group, starting with the cephalopod molluscs which perform
remarkably like the higher vertebrates in learning tests, and working
through to the echinoderms and coelenterates where the very occurA.Y.B.-3
1
B
LEARNING BY MARINE INVERTEBRATES
I.
11.
111.
IV.
V.
VI.
VII.
VIII.
IX.
X.
M. J. WELLS
Department of Zoology, Downing Xtreet, Cambridge, England
Introduction
. . . . . . .. .. ..
A.
Experiments on Sensory Integration
. .
Annelids
..
.. .. .. .. ..
Cephalopod Molluscs . . .. .. .. ..
B.
Brain Lesions and Learning . . .. ..
A.
Habituation to Repeated Stimuli . . ..
B.
C.
Brain Lesions and Polychaete Learning . .
Crustaceans-and Limulus
. . .. .. ..
A.
B.
Other Evidence that Crustaceans Learn . .
Further Experiments with Errant Polychaetes
The Results of Formal Training Experiments
C.
Learning by Limulw polyphemus . . ..
Gastropods and Lamellibranchs
. . ..
..
A.
B.
Homing by Limpets . . .. .. ..
C.
Other Evidence of Learning by Molluscs . .
Platyhelminthes . .
.. .. .. .. ..
Echinoderms
.. .. .. .. .. ,.
Coelenterates . . .. .. .. ,. ..
Discussion
.. .. .. .. .. ..
References
. . .. ..
..
.. ..
Formal Training Experiments with Gastropods
..
..
..
..
..
..
..
..
..
..
..
..
..
..
..
..
..
..
..
. .
..
..
..
..
..
..
..
..
..
..
..
..
..
..
..
..
..
..
..
..
..
.. 1
.. 2
.. 3
.. 15
.. 20
.. 21
.. 26
.. 28
. . 29
.. 29
.. 33
.. 37
. . 37
.. 37
.. 38
.. 41
.. 42
.. 44
.. 47
.. 51
.. 85
I. INTRODUCTION
It would clearly be ridiculous to limit a review of invertebrate
learning to marine forms were the object to assess the relevance of
research on invertebrates to the study of learning generally. This
review does not attempt to do anything so ambitious. Its more limited
objectives are to show how much marine forms have to offer for work
on learning and how very little has been done with most of them. The
hope in doing this is to persuade zoologists and psychologists that the
existing literature is more than a mere anecdotage and that enough is
now understood to justify much more intensive research ; the range of
material is vast, and much of it is very promising indeed.
In what follows the experiments surveyed are dealt with group by
animal group, starting with the cephalopod molluscs which perform
remarkably like the higher vertebrates in learning tests, and working
through to the echinoderms and coelenterates where the very occurA.Y.B.-3
1
B