MODIFIABILITY OF BEHAVIOR IN HYDROIDES

DIANTHUS V.
BT

ADA WATTERSON YERKES.

T h e sensitiveness of the tubicolous annelid Hydroides dianthus

V., manifested by its prompt withdrawal into its tube when jarred
or when a shadow falls upon it, suggested this worm as a good
object for a study in modifiability of behavior. Such a study was
carried on for a few weeks a t the Marine Biological Laboratory a t
Woods Hole during the past summer. Unfortunately the time
was too short for extensive experiments but the results obtained
thus far seem to be of sufficient value to warrant presentation.
The reactions of Hydroides to light have been studied by
HARGITT.’ who found that “under varying degrees of light intensity
* * * the results of all experiments involving increased intensity of
light were uniformly negative. O n the other hand, experiments
involving a sudden decrease of light intensity gave results as uni-
formly positive” (p. 3 1 9 ) . This confirms a statement made by
LOEB’(p. 103) concerning H. uncinata to the effect that only a
decrease in the intensity of light acts as a stimulus upon the ani-
mals: In the present experiments this stimulus was, therefore,
one of those selected for use and the decrease in intensity of the
light was obtained by bringing down rapidly a black cardboard
screen between the window and the dish which contained the
worms. After one or two seconds the screen was removed. For
convenience this decrease in light intensity will be referred to as a
shadow. Though the window faced south, the sunlight did not
fall directly upon the worms during the experiments. T h e
response to the stimulus was usually a quick contraction of the
animal into its tube. Occasionally this contraction occurred

C. W. Experhento on the Behavior of Tubidous Annelids. F u r n d of Expcrimntrrl

~HARGZTT,
zoo lo^, Vol. 3, pp. z g s - p ~ ,Jdy. I+.
?Lots, J. Ucber kiinstlichcUrnwandlung positiv heliotropischu Thierc in ncgrtiv Ldotropischc und
umgelehrt. P$iign’r drchiv. Vol. s+, pp. 81-108. 1893.
442 Journal of Comparative Neurology and Psychology.

more slowly and the tips of the circle of branchiae remained pro-
jecting a trifle from the mouth of the tube. T h e time during
which the animals remained contracted was noted in each case and
was found to vary in a way which will be described later.
A considerable number of tests were made to determine the sensi-
tiveness to shadow alone. In most of these the shadows were
given a t unequal intervals; for instance, if the animal contracted
in response to the first shadow, the second was given as soon as the
worm was again fully expanded. T h e time thus varied between
ten seconds and a minute or, in a few cases, two or three minutes.
If the animal did not respond, the shadows were repeated a t short
intervals, from five to ten seconds. Ten such trials were given in
succession. In the greater number of these tests (16 out of 27)
with different specimens the animals responded only the first time,
or possibly from one to three times, and then gave no further
response throughout the ten trials. This fact seems to be com-
parable to the results obtained by JENNINGS‘ (p. 172) with attached
infusoria which reacted the first time to a jet of water striking
against the disk or to other faint stimuli, or a few times to a
stronger stimulus, but did not respond to later repetitions of the
stimulus. This behavior JENNINGS interprets as due, not to
fatigue, since the number of reactions is few, but to a change in the
organism itself (p. I 73), a change which has distinctly a regulatory
character in the behavior of the animal. HARGIIT,on the con-
trary (p. 3 0 1 ) , who observed that when shadows fell rhythmically
on Hydroides and “the experiments were repeated with any con-
siderable frequency, specimens sooner or later became somewhat
irresponsive,” is inclined to regard this as the result of fatigue.
Evidence, however, from a later series of the present experiments,
tends to show that fatigue is not the cause, since the animals are
capable under stronger stimulation of contracting a large number
of times in rapid succession without apparent fatigue.
In ten cases of the twenty-seven mentioned above, the worms did
not respond the first time nor later during the ten trials. Whether
this is due to individual variation in sensitiveness or whether
the animals had responded previous to the experiments to the
shadows which fell upon them by chance while still in the aqua-

~JENNINGS,H.S. Behavior of the Lower Organisms. New Tork, Macmillan. Pp. 366, figs. IM.
I!+
 YERKES,
Behavior of Hydroides. 4'43
rium, it is impossible to say. T h e former seems probable, since
one animal was tested which responded every one of the ten
times, although the conditions were apparently exactly similar to
those of another worm which was tried simultaneously and
reacted only four times out of ten.
After some experiments of a different kind, to be described later,
had been given, some of the specimens used in them and a few new
ones were tested again for the effect of shadow alone. T h e
shadows this time were given at regular intervals of one or two
minutes. T h e number of responses in these cases is larger than in
the preceding, for out of twenty series of ten trials each, i. e., a
possible two hundred reactions, there were ninety-five responses to
the shadow. T h e longer interval between shadows seems, there-
fore, to interfere with the process of getting accustomed to the
stimulus which occurs when the intervals are only a few seconds in
length. I n Table I are shown the records of two series with one
specimen. In the first series the intervals between shadows were
fifteen seconds. In the second series, given four days later but
without any intervening training, the intervals are two minutes.

TABLE I.
SSRIISI. SEurs 2.
No. Shndmu. Time. NO. Shadow. Time.
1 0
- I
rc 18'

2 0 - a 'I
30.
3 0 - 3
I6
1s'
4 0 - 4
II
48'
5 0 - 5
6r
4'
6 0 - 6 II 28"
7 0 - 7
"
3=
8 0 - 8
I#
38"
9 0 - 9
"
33"
10 0 - I0 " 16'
In all of the tables o means no reaction, and '' means reaction.

A series of experiments was now started in which two different

stimuli were given in rapid succession. T h e response to light, it
has been shown, is variable, since the animals sometimes respond
and sometimes do not. The response to mechanical stimulation
caused by jarring the animals or by touching the branchial fila-
ments, with a glass rod is much more regular. In fact, the worms
 Journal of Comparative Neurology and Psychology.

respond immediately and invariably except in certain instances to

be mentioned later.
In these experiments three or more individuals in the same dish,
usually growing on the same piece of shell or stone, were used
simultaneously. At the beginning all were given ten trials with
shadow alone with the usual result. Then the shadow was given
and if the animals did not respond to this the mechanical stimu-
lation caused by touching the filaments gently with a glass rod
followed immediately. T h e period of retraction was noted and
when the animals were again expanded the experiment was
repeated. T h e trials were recorded in groups of ten each and
from one to six of these groups were given daily. A number of
specimens were tested thus for several days each. Some of these,
having been in the laboratory for some time, then became inactive
and refused to come out of the tube after being placed in the experi-
menting dish. Two animals, however, Nos. 40 and 41,obtained
from fresh material were tried for ten days and the records of these
two, which show a number of interesting points, will be discussed
in detail.
In the trial experiments in which shadow alone was given a t
intervals of 5 to 10 seconds No. 40 responded not at all and No. 41
only the first time. After a short interval the shadow and touch
series was given. In this No. 40 reacted to shadow alone three
times during the first ten trials, No. 41 only once and neither of
them again throughout the forty trials given that morning. T h e
fact that when responses to shadow alone occurred they almost
always came within the first twenty trials soon became evident and
the records will, therefore, be given for these twenty alone. Th e
results for the ten days can best be represented by the following
curves in which the number of responses to shadow alone for each
of the successive days, as indicated on the base line, is marked on
the ordinates.
In the case of No. 40 there is a marked increase in the number of
responses to shadow from the first to the eighth day and those of
the later days are much higher than a t first.' For No. 41 the
increase is not so great, though the rise in the curve from the
second to the fifth day is pronounced. Even a t the beginning, the
responses of this individual were not as numerous as for No. 40
and after the tenth day it became so inactive, sulking at the mouth
of the tube or remaining retracted into it so long, that it became
 YERKEG,
Behavior bf Hydroides. 445

NO.of reachoms

L u m b e r 43

.... .............. Number 41

446 Journal o f Comparative Neurology and Psychology.

impossible to use it further for the experiments. This seems to

indicate a good deal of individual variation and unfortunately the
specimens tried were not numerous enough to provide averages.
After the tenth day No. 40 was used for the experiments where
shadows alone were given at regular two minute intervals and
these series render further work with combined stimuli incompar-
able.
TABLE II.
OF No. 40.
REACIIONS
No. S h h . Touch. Time. No. Shadow.
I II - 34" 3' 0

z ir - I 9" 32 0

3
#I - 29" 33 0
II
4 0 235" 34 0

5
II - 50" 35 0
I6
6 0 93" 36 0

7
II - 34" 37 0
II
8 0 59" 38 0

9
11 - 27. 39 0
I1
I0 0 41" 40 0
II
I1 0 52" 41 0
ir
Ia 0 43# 42 0
ll
'3 0 27" 43 0
II
14 0 26" 44 0
#I
1.5 0 18" 45 0
II
16 0 25" 46 0
0 II
17 18' 47 0
18 0
I1
136" 48 0

'9
II - 35" 49 0
'I
20 0 37. 50 0
I#
21 0 24# 5' 0
za IS - 35" 52 0
I1
23 0 2s' 53 0
II
14 0 21" 54 0
11
25 0 18" 55 0
26 II
0 zo" 56 0
I1
27 0 2 I" 57 0
z8 0 II
I 6' 58 0
I1
29 0 I 6' 59 0
II 60
30 0 171 0

In explanation of this increase in responsiveness to shadow alone

it seems possible that the shadow and touch have become so asso-
ciated that the occurrence of the one intimates the probability that
the other will follow. T he response to touch is fairly uniform and
 YERKES,
Behavior of Hydroides. 447
after a few such experiences, the shadow, cast before, as it were, by
the coming event, more frequently produces a reaction. T h e
repetition Gf both stimuli many times in succession causes this
effect to wear away temporarily, since the touch is not sufficiently
injurious to force reaction.
Two other interesting facts have appeared in the course of these
experiments. One is shown clearly in the record of one day’s
experiments given in Table 11. In this both stimuli were given;
the responses to shadow are shown in the first column, those to
touch in the second and the length of time during which the
animal remained retracted into its tube in the third.
The period of retraction is short the first three times-rgto 34ff-
but the fourth time it is nearly four minutes. For the next
thirteen times . it ranges from eighteen to ninety-three seconds;
then comes another period of nearly four minutes followed by
nineteen contractions which last from twelve to eighty-five seconds
each and then a contraction of nearly twelve minutes’ duration.
Thus after the fourth, eighteenth, thirty-eighth and sixtieth trials
the animal remained contracted for a relativelv long period, vary-
ing from four to twelve minutes, whereas the’intervening contrac-
tions seldom lasted more than one and a half minutes and are
usually less than thirty seconds. This rhythm is very marked in
all the series which were carried on long enough each day to show
it, and in the short series there is usually one period of retraction
noticeably longer than any of the others.
T h e occurrence of unusually long periods of retraction in the
case of another individual, No. 41,for the first five and the tenth
days of the experimentation is shown in Table 111. It is worthy
of note that the frequency of the prolonged retraction periods is
much greater on the tenth day than on the first.
These periodic “rests” might be attributed to fatigue, but they
do not seem necessarily due to that. Being a tube-dwelling
animal, Hydroides is narrowly limited in the kind of reactions
possible to it. It responds to mechanical stimulation repeated a t
frequent intervals in the most apparent way, by a contraction into
the tube. If the behavior of the animal were modifiable we might
expect after a number of repetitions of this stimulus a change in
reaction comparable to that of Stentor‘ when, after repeated faint

~JENNINGS,H.
S. Loc. cit., pp. 173-175,
448 gournal of Comparative Neurology and Psychology.
stimuli, it ceases to contract and begins to bend in different direc-
tions, or, when after increase in the stimulus it breaks loose and
TABLE III.
Prromc O c c n a n ~ c ror PROLONGED
RETRACTION
IN No.41.

ra day. zd day. 3d d.r.

No. of Reaction. No. of Reaction. Time. No. of Reaction. T i m r .
4 2 312" 8 18S.
7 6 267" '7 w
W 15 384" 23 I so'
'3 18 160" '9 161.

59 3' 610" 40 573.

37 38 447.
+h day. 5th day. rcnk day.
No. of Reaction. Time. No. of Reaction. Time. No. of Reaction. Timr.
I I 84' 7 281" 5 1

3 18s. 11 370" 7 210"

'3 305" 35 381. I0 170"

33 5 4 50 470" 11 191'
40 w' '5 zoo.
58 148' 16
I8 1 ~ 3 ~

swims away. This prolonged contraction iri Hydroides may be

simply a difference in the response to stimulation as the result of
past experience. This instance seems, in fact, an illustration of
one of the conclusions stated by JENNINGS (p. 445)' in a recent
paper on the modifiability in behavior of the earth-worm, to the
effect that "the reaction to a given stimulus depends partly on
previous stimuli received ."
A second fact which may perhaps be explained by the same
hypothesis is brought out in some of the longer series. In the
record given above in Table 11, for instance, after some thirty-
three trials theanimal did not respond to the shadow nor to the
first touch from the glass rod. Two, three, or five strokes were
required to bring about a reaction. T h e number of strokes needed
each time is recorded in parenthesis. This tendency increased
throughout the series until in the fifty-fifth trial the filaments
were stroked 152 times in rapid succession before they were with-
drawn. T h e touch at first was quick and slight, but producing
no reaction it was made heavier until the filaments were stroked
IJennings, H. S. Modifiability in behavior. II. Factors determining direction and character
of movement in the earth-worm. JOY^. of Expcr. Zool., Vol. 3, pp. 435-55. I$.
 YERKE s, Behavior of Hydroides. 449
from base to summit and the reverse so forcibly as to turn the
whole crown of filaments over to one side or the other in a horizon-
tal position. T h e contraction after such treatment this particular
time lasted only ten seconds. This change in behavior could
certainly not be due to fatigue since the resting condition is surely
that of withdrawal into the tube and in this case the animal not
only endured the stimulus repeatedly but after a contraction which
lasted a remarkably short time was ready for it again. This
occurred many times with several specimens when series of twenty
or more trials were given in succession. I n many other cases there
was only a partial retraction into the tube and the crown of branchiz
would remain half out of the tube until the next expansion took
place.
SUMMARY

I . To a photic stimulus consisting of a decrease in the intensity

of light, repeated at short and irregular intervals, Hydroides re-
sponds at first by contraction but later gives no reaction.
2. When the stimulus occurs at longer intervals the responses
are more frequent.
3. T o tactual stimulation the animal responds almost uniformly
by a contraction into the tube.
4. When the photic stimulus is followed immediately by the
tactual the worms gradually respond more frequently to the former
alone than they did previous to this training; i.e., they learn to
react to the shadow.
5. T h e time of retraction after the tactual stimulus is usually
short but in any long series a retraction of much longer dura-
tion occurs periodically as a possible variation in the method of
response to repeated stimulation.
6. After many repetitions of shadow .and tactual stimuli the
animals no longer react normally to either but require frequently a
large number of tactual stimulations to induce contraction.
7. The behavior of Hydroides is thus. eminently modifiable
since it varies with repetitions of a stimulus or when two stimuli of
different reactive value are repeatedly given in succession.

Woods Hole, Mass.,

September, 1 9 6 .