Sixteen Experimental Investigations from the Harvard Psychological Laboratory - Hugo Münsterberg (best life changing books txt) 📗
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beginning of comparative reaction-time studies by which it is hoped
important information may be gained concerning the significance and
modes of action of the nervous system. Comparative physiology has
already made clear that the time relations of neural processes deserve
careful study.
2. According to the strength of the stimulus, electric stimulation of
the frog causes three types of reaction: (1) A very weak or threshold
stimulus results in a deliberate or delayed reaction, the time of
which may be anywhere from 300[sigma] (thousandths of a second) to
2,000[sigma]. (2) A very strong stimulus causes a spinal reflex, whose
time is from 50 to 80[sigma]; and (3) a stimulus of intermediate
strength causes a quick instinctive reaction of from 150 to 170[sigma]
in duration.
3. The reaction time for electric stimuli whose relative values were
1, 2 and 4 were found to be 300.9[sigma], 231.5[sigma] and
103.1[sigma].
4. The reaction time of the frog to a tactual stimulus (contact of a
rubber point) is about 200[sigma].
5. The variability of reaction times of the frog is great, and
increases as the strength of the stimulus decreases.
6. When two kinds of stimuli (e.g., electrical and tactual) give
reaction times of equal variability, I consider them directly
comparable.
7. According to this criterion of comparability the reaction time to
electric stimulation which is comparable with that to tactual is
172.1[sigma]; and it is to be compared with 205.7[sigma]. Both of
these have a variability of approximately 34[sigma]. On this basis one
may say that the tactual reaction time is considerably longer than the
electrical.
PART III. AUDITORY REACTIONS OF FROGS.
X. HEARING IN THE FROG.
A. Influences of Sounds in the Laboratory.
After determining the simple reaction time of the green frog to
tactual and electrical stimulation, I attempted to do the same in case
of auditory stimuli. In this I was unsuccessful because of failure to
get the animal to give a motor response which could be recorded. The
animal was placed in an experimenting box with a string attached to
one hind leg as in the experiments described in Part II., and after it
had become accustomed to the situation a sound was made. A wide range
of sounds were tried, but to none except the croak of another frog was
a motor reaction frequently given. Even a loud noise, such as the
explosion of a large pistol cap, caused a visible motor reaction only
in rare cases. In fifty trials with this stimulus I succeeded in
getting three reactions, and since all of them measured between 230
and 240[sigma] it is perhaps worth while to record the result as
indicative of the auditory reaction time. As these were the only
measurements obtained, I have no satisfactory basis for the comparison
of auditory with other reaction times.
The remarkable inhibition of movement shown by the frog in the
presence of strong auditory stimulation, at least what is for the
human being a strong stimulus, led me to inquire concerning the limits
and delicacy of the sense of hearing in frogs. In the vast quantity of
literature on the structure and functions of the sense organs of the
animal I have been able to find only a few casual remarks concerning
hearing.
In approaching the problem of frog audition we may first examine the
structure of the ear for the purpose of ascertaining what sounds are
likely to affect the organ. There is no outer ear, but the membrana
tympani, or ear drum, covered with skin, appears as a flat disc from 5
to 10 mm. in diameter on the side of the head just back of the eye and
a little below it. In the middle ear there is but one bone, the
columella, forming the connecting link between the tympanum and the
internal ear. The inner ear, which contains the sense organs,
consists of a membranous bag, the chief parts of which are the
utriculus, the sacculus, the lagena, and the three semicircular
canals. The cavity of this membranous labyrinth is filled with a
fluid, the endolymph; and within the utriculus, sacculus and lagena
are masses of inorganic matter called the otoliths. The auditory nerve
terminates in eight sense organs, which contain hair cells. There is
no cochlea as in the mammalian ear. The assumption commonly made is
that vibrations in the water or air by direct contact cause the
tympanic membrane to vibrate; this in turn causes a movement of the
columella, which is transmitted to the perilymphatic fluid of the
inner ear. The sensory hair cells are disturbed by the movements of
the otoliths in the endolymph, and thus an impulse is originated in
the auditory nerve which results in a sensation more or less
resembling our auditory sensation. It is quite probable that the
frog’s sense of hearing is very different from ours, and that it is
affected only by gross air vibrations. This conclusion the anatomy of
the ear supports.
Although there does not seem to be a structural basis for a delicate
sense of hearing, one must examine the physiological facts at hand
before concluding that frogs do not possess a sense of hearing similar
to our own. First, the fact that frogs make vocal sounds is evidence
in favor of the hearing of such sounds at least, since it is difficult
to explain the origin of the ability to make a sound except through
its utility to the species. Granting, however, that a frog is able to
hear the croaks or pain-screams of its own species, the range of the
sense still remains very small, for although the race of frogs makes a
great variety of sounds, any one species croaks within a narrow range.
Having satisfied myself that motor reactions for reaction-time
measurements could not be gotten to any ordinary sounds in the
laboratory, I tried the effect of the reflex croaking of another frog
of the same species. In attempting to get frogs to croak regularly, I
tested the effect of removing the brain. The animals are said to croak
reflexly after this operation whenever the back is stroked; but for
some reason I have never been successful in getting the reaction
uniformly. In many cases I was able to make normal animals croak by
rubbing the back or flanks, and to this sound the animals under
observation occasionally responded by taking what looked like an
attitude of attention. They straightened up and raised the head as if
listening. In no case have other motor responses been noticed; and the
above response was so rare that no reaction-time measurements could be
made.
Again, while working with the green frog on habit formation, I one day
placed two animals in a labyrinth from which they could escape by
jumping into a tank of water. Several times when one frog jumped into
the water I noticed the other one straighten up and hold the
‘listening’ or ‘attentive’ attitude for some seconds. As the animals
could not see one another this is good evidence of their ability to
hear the splash made by a frog when it strikes the water.
B. Influence of Sounds in Nature.
In order to learn how far fear and artificial conditions were causes
of the inhibition of response to sounds in the laboratory, and how far
the phenomenon was indicative of the animal’s inability to perceive
sounds, I observed frogs in their native haunts.
By approaching a pond quietly, it is easy to get within a few yards of
frogs sitting on the banks. In most cases they will not jump until
they have evidence of being noticed. Repeatedly I have noted that it
is never possible to get near to any frogs in the same region after
one has jumped in. In this we have additional proof that they hear the
splash-sound. To make sure that sight was not responsible for this
on-guard condition in which one finds the frogs after one of their
number has jumped into the water, I made observations on animals that
were hidden from one another. The results were the same. I therefore
conclude that the splash of a frog jumping into the water is not only
perceived by other frogs in the vicinity, but that it is a peculiarly
significant sound for them, since it is indicative of danger, and
serves to put them ‘on watch.’
A great variety of sounds, ranging in pitch from a low tone in
imitation of the bull frog’s croak to a shrill whistle, and in
loudness from the fall of a pebble to the report of a pistol, were
tried for the purpose of testing their effects upon the animals in
their natural environment. To no sound have I ever seen a motor
response given. One can approach to within a few feet of a green frog
or bull frog and make all sorts of noises without causing it to give
any signs of uneasiness. Just as soon, however, as a quick movement is
made by the observer the animal jumps. I have repeatedly crept up very
close to frogs, keeping myself screened from them by bushes or trees,
and made various sounds, but have never succeeded in scaring an animal
into a motor response so long as I was invisible. Apparently they
depend almost entirely upon vision for the avoidance of dangers.
Sounds like the splash of a plunging frog or the croak or pain-scream
of another member of the species serve as warnings, but the animals do
not jump into the water until they see some sign of an unusual or
dangerous object. On one occasion I was able to walk to a spot where a
large bull frog was sitting by the edge of the water, after the frogs
about it had plunged in. This individual, although it seemed to be on
the alert, let me approach close to it. I then saw that the eye turned
toward me was injured. The animal sat still, despite the noise I made,
simply because it was unable to see me; as soon as I brought myself
within the field of vision of the functional eye the frog was off like
a flash.
Many observers have told me that frogs could hear the human voice and
that slight sounds made by a passer-by would cause them to stop
croaking. In no case, however, have such observers been able to assert
that the animals were unaffected by visual stimuli at the same time. I
have myself many times noticed the croaking stop as I approached a
pond, but could never be certain that none of the frogs had seen me.
It is a noteworthy fact that when one frog in a pond begins to croak
the others soon join it. Likewise, when one member of such a chorus is
frightened and stops the others become silent. This indicates that the
cessation of croaking is a sign of danger and is imitated just as is
the croaking. There is in this fact conclusive evidence that the
animals hear one another, and the probability is very great that they
hear a wide range of sounds to which they give no motor reactions,
since they do not depend upon sound for escaping their enemies.
The phenomenon of inhibition of movement in response to sounds which
we have good reason to think the frogs hear, and to which such an
animal as a turtle or bird would react by trying to escape, is thus
shown to be common for frogs in nature as well as in the laboratory.
This inhibition is in itself not surprising, since many animals
habitually escape certain of their enemies by remaining motionless,
but it is an interesting phenomenon for the physiologist. We have to
inquire, for instance, what effects sounds which stimulate the
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