ON SEEING IN THE DARK.

1
REMARKS ON THE EVOLUTION OF THE E Y E .
BY DR. OSKAR NAG EL.
In my investigations on certain properties of illuminant sub-
stances, I have had to study to some extent the anatomy of the
eye and the optical functions in colored light and in the dark
(beginning of brightness). I have arrived at certain conclusions
relative to distinct optical perception in the daylight and in the
dark, which possibly are of interest for comparative anatomy
and physiology, and also for the theory of evolution.
First of all I may mention the anatomical a.nd physiological
facts upon which my conclusions are based.
The fibers of the optic nerve end in cylindrical rods (about
120 millions) and flask-shaped cones (about 60,000). The light-
rays from an object, upon which our eyes are fixed, strike upon
the fovea centralis, which contains cones only and no rods.
The further we go away from the yellow spot, the more rods
are found, and at the circumference they are in the majority.
At the place where the optic nerve enters the eyeball, neither
rods nor cones are present, hence this point is entirely insensi-
tive to light.
When we fix our eyes upon an object, its image falls upon
the yellow spot (foveal vision); hence during the action of direct
(foveal) vision the rods are entirely out of action, while in in-
direct (peripheral) vision the rods come into action together with
the cones.
In 1887, H. F . Weber found2 during his investigations on
the relative economy of incandescent lamps, that a carbon fila-
ment emits a ghostly gray light before the red-glow starts. This
first trace of a misty gray light appears to the eye as something
unsteady and glimmering. As the temperature rises, the bright-
ness of this light rapidly increases, going over from gloomy
'The MS. of this article was received March 13, 1908. — E D .
'See Sitzber. d. Berliner A. d. IV., 28, p. 491, 1887 ; Wiedemann's Ann.,
32, p. 526, 1887.
250
 ON SEEING IN THE DARK. 351

gray into ash-gray, yellowish gray and finally into fire-red.

When the first trace of the red light appears, the last trace of
glimmering, trembling and vibrating, which was present in all
the stages of the gray-glow, disappears. According to H. F .
Weber and E. Emden l the first emission of light is visible with
gold at 423 0 C , with German silver at 403 0 C.
These observations led O. Lummer 2 to the following con-
ception relative to the nature of our eye, wherein he ascribes
following the theories of modern physiology, entirely different
functions to the two sensible components of the retina, namely
the cones and the rods, by considering same as two separate
optical apparatus.
If we observe in a dark room the gradual increase of the
temperature of a body, according to Lummer, our eye feels
twice a sudden change, first from darkness to gray-glow and
then from the latter to red-glow. In both cases this sudden
change or transition corresponds to the transgression of the
limit of sensibility of our optic nerve (the so-called threshold of
sensibility). The arising of the gray-glow corresponds to the
threshold of sensibility of the rods, the one of the red-glow
however, to the threshold of sensibility of the cones. Hence the
sensation of the gray-glow is effected by the rods, of the red-
glow by the cones.
" Based upon the new physiological researches relative to the
vision at low brightness and the influence of the purple pigment
in the retina, the function of our two retina-organs was gradu-
ally separated and their separate tasks ascertained."
J. v. Kries 8 solved the still existing difficulties and contra-
dictions by the hypothesis, that the cones form our color-capable
•bright-apparatus' and the rods our totally color-blind 'dark-
apparatus.' According to this theory the cones enable us to see
at great brightness and their irritations by the light-waves
causes in the brain the sensation of color, while the rods which
are totally color-blind, come into effect only at very low bright-
ness and have the faculty of intensely increasing their sensi-
1
Wiedemann's Ann., 36, pp. 214-236, 1889.
* Wiedemann's Ann., 62, pp. 14-29, 1897.
8
Zeiisckr. f. Psych, und Phys. d. Sinnesorgane, 9, pp. 81-123, 1894-
252 OSKAR NAGEL.

bility in the dark. This property is called by Kries ' dark

adaptation.' Before the cones react upon colored light, the
rods cause in the brain the sensation of colorless brightness.
Hence we have at very low brightness a contest of the two
optical apparatus which, at sufficiently low brightness, is settled
in favor of the color-blind rods, so that then everything appears
gray in gray, i. e.t in colorless brightness.
We can now understand why we do not see distinctly in a
dark gray light (at low brightness); it is because, if we fix our
eyes upon an object (/. e., have an image produced on the fovea
centralis), we have no apparatus that is affected by such image
or light; the rods which are able to receive this light are absent
from the fovea centralis and thence only a glimmering restless
image is produced by an object in gray light. If we would
have a sufficient number of rods among the cones around the
fovea centralis, we would see much better in the dark than we
actually see. This enables us to explain why a large number
of animals see very distinctly in the dark. The cause of this
most probably is, that in their eyes, which are of nearly the
same construction as the human eyes, rods and cones are uni-
formly intermixed. It can be proven that a horse sees very
distinctly in the dark, by simply taking a ride on a very dark
night over country roads. The horse will trot and gallop as
safely as in daylight, while sometimes the rider will be afraid,
that the horse may stumble over a root or fall into a ditch.
But there are even animals in existence which we positively
know see much more distinctly at very low brightness than in
the daylight: owls, etc. Here we have a case where, very
probably, rods and cones have exchanged locations, as com-
pared to the human eye. The rods are around the fovea cen-
tralis and the cones at the periphery. Hence with these eyes
a distinct vision is effected at very low brightness, while at a
greater brightness only a restless, glimmering sensation will be
produced. While we see ghostly lights in the dark, the owl
probably sees ghostly lights in the daylight. For physically a
ghostly apparition is nothing else but a bright point which we
cannot get into the foveal vision and which whenever we try to
do so, is naturally vanishing, thereby causing the impression of
motion.
 ON SEEING IN THE DARK. 253

We can infer, that as soon as the eye of about the construc-

tion of the mammalia — or aves-eye — was developed in the
course of evolution, rods and cones were uniformly intermingled,
or rather became uniformly intermingled by natural selection.
Thereby the animals were adapted to the light in the daylight
and in the dark. In such animals that are now procuring their
food mainly during the night and in the dark, the cones were
gradually driven back by the rods. In the human eye the
opposite process took place; the rods were driven back by the
cones. We only see a restless glimmering light, a ghostly
apparition, where a horse will distinctly see an object.
Another question is whether we have simply lost the faculty
of seeing in the dark or whether we have exchanged same for
a higher faculty; we have lost one faculty which is of no use
to us now and have gained one that is extremely valuable for
our development. By driving back the rods we have concen-
trated and increased the strength and sensitiveness of our cones ;
we have developed and are still further developing and refining
our sense for color and for light, our capacity for distinguishing
the closest shades of colors. Whether we look at it practically
or symbolically, we find that the driving back of the rods makes
us masters of light and colors.
We can observe the development of the human eye within
the historical times. Homer does not distinguish black from
blue, but his heroes find their way as well in the dark night as
in daytime. And now take a modern silk-dyer or cotton-printer
who clearly distinguishes thousands of different shades, though
he is not able to find his way on a dark country-road.
Rivers states 1 that some Australian tribes have but three
expressions for colors, one for red, purple and orange, another
for white, yellow and green and the third for dark blue and
indigo. Some primitive races cannot distinguish blue from
green, nor blue from violet. Tintometric tests have proven that
in these races the threshold of sensibility is very low for red,
much higher for yellow and very high for blue. Lobsien 2 has
found that violet is frequently taken for brown by children ; he
1
Popular Science Monthly, 1901, p. 44.
'Zeitschriflf. Psych, und Phys. d. Sinnesorg., 1904, p. 29.
254 OSKAR NA GEL.

shows that at the age of thirteen to fourteen years violet, orange

and indigo are still struggling for their development.
We have reason to believe that this evolutionary contest be-
tween rods and cones does not require a long period and that
adaptation is effected in a comparatively short time. Generally
people that have lived in the country for generations and have
to use their eyes in the dark, are in this respect very much
superior to the inhabitants of the cities. In a few generations
of city-life, however, this faculty is lost or rather exchanged for
a higher faculty, which makes our eyes, from the standpoint of
evolution, superior to the eye of the eagle or owl.
If we would know as much about the work and action of the
different parts of the ear as we do of the eye, we would proba-
bly be enabled to form an idea about the limits of sounds per-
ceived by the different species of animals.