Brain Research, 294 (1984) 333-339 333

Elsevier

The Influence of Different Light Spectra on the Suppression of Pineal Melatonin Content
in the Syrian Hamster

GEORGE C. BRAINARD*, BRUCE A. RICHARDSON**, THOMAS S. KING and RUSSEL J. REITER***

Department of Anatomy, The University of Texas Health Science Center at San Antonio, 7703 Floyd Curl Drive,
San Antonio, TX 78284 (U.S.A.)
(Accepted July 12th, 1983)

Key words: pineal gland - - melatonin - - light spectra - - blue light - - green light

The purpose of this study was to test the capacity of different visible wavelengths of light to suppress nocturnal levels of pineal mel-
atonin in hamsters. It was found that the visible wavelengths vary in their ability to perturb pineal melatonin. During the period of
peak pineal melatonin production, animals were exposed to fluorescent light sources having half-peak bandwidths of 339-371 nm
(near-ultraviolet), 435-51)0 nm (blue), 510-550 nm (green), 558-636 nm (yellow) and 653-668 nm (red). In each experiment, animals
were exposed to equal irradiances of each light source. The different irradiances used were 0.928, 0.200, 0.186, 0.074 and 0.019
uW/cmL The resultant data demonstrated that blue fluorescent light was the most efficient in suppressing pineal melatonin. Green flu-
orescent light was found to be the next most efficient light for inhibiting pineal melatonin followed by yellow fluorescent light. Near-ul-
traviolet and red light were the least capable of suppressing pineal melatonin. These observations suggest that the retinal photopig-
ment responsible for mediating the pineal gland's response to light in the hamster may be either rhodopsin or another blue-sensitive
chromophore.

INTRODUCTION primarily by p h o t o p e r i o d 7,14,15. A l t h o u g h light does

Animals living in natural habitats are exposed to a
variety of environmental light sources including the
sun, moon, stars and, in some cases, artificial light
from civilization. The sun itself produces a steady
output of spectral irradiance but the motion and at-
mosphere of the earth cause great modulations in the
spectrum and irradiance of light which reaches the
surface of the planet. Sunlight, at its peak, reaches an
irradiance of m o r e than 50,000 ktW/cm2 and falls as
low as 0.005/~W/cm 2 during the twilight periods of
dusk and dawn 24. The varied direct and indirect influ-
ences of light on animal species have been reviewed
elsewhere~,22-2L The following study was designed to
determine how light spectrum interacts with the
physiology of the pineal gland of the Syrian hamster
(Mesocricetus auratus). The hamster was chosen be-
cause it is a highly photosensitive species which has
daily and annual physiological rhythms that are cued
not impinge directly on the pineal, light influences this
gland by an indirect, multisynaptic neural pathway
projecting from the retina to the pineal glandlL This
neural pathway synchronizes the metabolism of pine-
al indoles to the daily environmental light:dark cy-
cle 11. Melatonin is one of the products of the neural-
ly-regulated pineal indole metabolism. Melatonin,
and perhaps other constituents secreted from the pi-
neal, influence the neuroendocrine-reproductive
axis14,16.
The formation of melatonin in the hamster pineal
gland is synchronized by the daily and annual
changes in the environmental light:dark cy-
cle 1,13,16,23. F u r t h e r m o r e , the unexpected exposure
of hamsters to light during the night causes a rapid
depression of pineal melatonin content 21. This light-
induced suppression of melatonin in hamsters is de-
pendent on the intensity of the light 2. The aim of the
following study was to d e t e r m i n e how the spectral

* Present address: Department of Neurology, Jefferson Medical College, 1025 Walnut St., Philadelphia, PA 19107. U.S.A.
** Present adress: Department of Anatomy, American University of the Caribbean School of Medicine, P.O. Box 400, Plymouth,
Montserrat, British West Indies.
*** To whom correspondence and reprint requests should be addressed.

0006-8993/84/$03.00 1984 Elsevier Science Publishers B.V.

334

characteristics of light influence the light-induced

suppression of pineal melatonin. 200 l Red
I
MATERIALS AND METHODS I00 -

l \
Adult, male Syrian hamsters (100-120 g), aged 2-3
O.
months, were used in these experiments. Hamsters
were received from Lakeview Colony, Newfield. N J.
housed 6-8 per cage and maintained on a 10:14 Ye//ow
light:dark cycle (lights on 07.00 h) for a minimum of 3
weeks prior to experimentation. Animals were sup-
plied food and water ad libitum and bedding was
changed twice weekly. The daily light cycle was pro- O~ i

duced by overhead cool white fluorescent light

sources (Sylvania. F40/CW/RS/SS) which provided 200 ] Green
an irradiance of 200 _ 50 #W/em 2. The following ex-

L
periments were performed between 02.00 and 05.30 .. I00
h. Exposure of animals to experimental light sources
took place in the chamber shown in Fig. 1. This " 1
0 I

chamber consisted of a 47 x 34 x 28 cm box painted

black (Flat Black no. 920. Yankin Majestic Paint 0

Corp.) on the inside. The different experimental

~" t00 Blue
light sources had one-half peak bandwidths of
339-371 nm (near ultraviolet), 435-500 nm (blue). 50
510-550 nm (green), 558-636 nm (yellow) and 653-
668 nm (red). These fluorescent lights were manufac- 0
tured and generously supplied by Durotest (North
Bergen, N J). The spectral power of lights used in the
I00 Near U/trav/ole!

r ~ LiFgluhtoS
reosucrceent]
\
/ \ o/,,'--,
300 400
. .
500
. .
600
.
700
, ,
800
/ \ Wavelength (nm)
/ \
/ "\ Fig. 2. Spectral power distributions of the experimental light
SOUrCes.

/ \ following experiments were individually measured at

Fig. 1. Chamber for exposing hamsters to restricted bandwidth
fluorescent light. In this chamber, animals were able to free-
roam a 34 x 28 cm area. The animals were exposed to direct il-
lumination from the fluorescent light sources.
5 nm intervals with a high precision automatic re-
cording spectroradiometert0 by the manufacturer
(Fig. 2). Radiometric measurements of light irradi-
anees were taken with a J 16 Photometer/Radiometer
and a remote J6512 Irradianee Probe (Tektronix,
Beaverton, OR). Since ammals could free-roam the
exposure chamber, multiple-irradiance measure-
ments were taken across the entire surface over
which animals moved during light exposure. Exten-
 335

sive effort was made to ensure that the irradiance Experiment 3

over this surface was homogenous. Changes in light Jr- Three sets of 21 animals each were used in this ex-
radiance were produced by masking the fluorescent periment. In each set, 7 animals were exposed to ei-
lamp with opaque black plastic at regular intervals ther green light, blue light or darkness. The irradi-
along the lamp's length. ance used for each set was 0.186, 0.074 or 0.019
pW/cm 2.
Experiment 1 Immediately following the 20-min exposure to ex-
Groups of 8 hamsters each were exposed to the perimental light, pineal glands were rapidly collected
near ultraviolet, blue, green, yellow or red light from hamsters, placed in glass tubes containing 1.0
source for 20 rain. Each bulb was adjusted to produce ml phosphate-buffered saline (pH 7.0), disrupted by
an irradiance of 0.928 pW/cm 2. After 20 min, pineals sonication and stored at - - 2 0 C until thawed for ra-
were removed from the hamsters exposed to light dioimmunoassay. Pineal glands from unexposed con-
and from corresponding control groups of 8 hamsters trol animals were collected and processed under dim
not exposed to light. (0.5-1.5 pW/cm2) red light (Kodak Safe-light with a
15 W incandescent bulb behind a no. 1A red filter).
Experiment 2 Melatonin contents in duplicate 200 pl samples were
This experiment was identical to the first experi- measured by a single antibody technique 20. This tech-
ment except the irradiance of each light source was nique has been validated for quantifying melatonin
adjusted to 0.200pW/cm 2. content of hamster pineal homogenates 13. Control
samples estimated by radioimmunoassay to contain
0, 59,275, and 591 pg/ml melatonin had the respec-
tive interassay coefficients of variation: 4c4, 18%,
d'Syrian Hamsters 15% and 19%.
F~ Control Analysis of the data was performed with the aid of
~4oo] - ~ Exposed to 0 928 uW/cm ~ a Monroe 1860 Programmable Calculator and a Sal
p< 0 001 vs. Terminal Computer (Model 20). The data of each ex-
Unexposed Group
i 200 [ periment was initially subjected to a one-way analy-
sis of variance ( A N O V A ) . Significant differences be-
I0OO] tween means were determined by a Newman-Keuls
z.
test.
.E
a_
8oo! U~5
RESULTS
r-

C
0 600 In all 3 experiments, control animals which were
~D
unexposed to light during the night had typically el-
400] evated pineal melatonin contents. Hamsters which
were exposed to a 0.928 pW/cm 2 irradiance of red
and near ultra-violet fluorescent light for 20 min had
200
pineal melatonin contents which were not significantly
different from the pineal melatonin levels of unex-
0j 359-37 435-500 515-55o posed animals (Fig. 3). in contrast, animals exposed
558 636 653-668
[Near U.V ', (B ue) (Green) (Yellow) (Red)
to 0.928/~W/cm 2 of blue, green or yellow fluorescent
Peak Wavelengths-Nanometers light for 20 min had significantly suppressed pineal
Fig. 3. Influence of different light spectra at 0.928/~W/cm2 on melatonin contents compared to those of unexposed
pineal melatonin content. Bars indicate mean pineal melatonin animals.
contents and lines on top of bars indicate S.E. Pineals were col- In the second experiment, animals were exposed
lected from each group of light-exposed animals 20 min after
the lights were turned on. Pineals were collected from a corre- to only 0.200 pW/cm 2 of each light source. At this
sponding group of unexposed animals at the same time for each lower irradiance, the red, the near-ultraviolet and
group, n = 8 for each group. the yellow fluorescent light did not cause a significant
 336

suppression of pineal melatonin during a 20-min ex- 2400 O" Syrian Hamsters
posure (Fig. 4). In contrast, both the blue and green IBm]Exposed ro Ll~ht
light significantly (P < 0.001) inhibited pineal mela- x~ 2 0 0 0 1 ~ Not Exposed/o Light
tonin levels of exposed animals. 0
~ o <0001 v~ UnexPosedGroup
In the last experiment, hamsters exposed to either -~ 1600 ! e<OO/vs Unexposed Group
~c 0 < 0 0.5 v5 U n e x P o s e d G r o u p
blue or green fluorescent light at an irradiance of
0.186/~W/cm 2 for 20 min had significantly (P < 0.01
and P < 0.05, respectively) suppressed pineal mel-
atonin compared to those of unexposed animals o 800 TF
(Fig. 5). Though the animals exposed to blue light
had a lower mean melatonin content than the animals
409
exposed to green light, there was no statistically sig-
o
nificant difference between these two groups. Ani- BI Grn BI Grr~ BI Grn

0 86 0 r[ 7zL 0019
mals exposed to blue and green light at an irradiance
of 0.074 #W/cm 2 for 20 min had significantly lower pi- Light Irradlonce (pW/cm 2)

neal melatonin levels compared to the pineal melato- Fig. 5. Capacity of blue and green light for suppressing pineal
nin levels of unexposed animals (P < 0.001 and P < melatonin content. Bars indicate m e a n pineal melatonin con-
tents and the lines on top of the bars indicate S.E. Pineals were
0.05, respectively). Finally, animals exposed to an ir- collected from animals after 20 rnin exposure to each light
radiance of 0:019/~W/cm 2 of either blue or green light s o u r c e Pineals were collected from a group of unexposed ani-
mals at the same time. T h e m e a n melatonin level of animals ex-
posed to 0.019 flW/cm 2 of blue light was significantly lower (P
6 Syrian Hamsters < 0.01) than that of animals exposed to green light at the same
irradiance. For each group n = 7.
Control
r---I Exposed to 0.20 pW/cm z
for 20 min had pineal melatonin contents which were
1400] p<O.O01 vs. Unexposed Group
not significantly different from those of unexposed
T
animals: However. the mean melatonin level of ani-
,\\\x
12001 mals exposed to 0.019 uW/cm 2 of blue light was sig-
e-
nificantly lower (P < 0.01 ) than that of animals ex-
_o I000-
(.9 _L-- posed to green light at the same irradiances.
The raw data shown in Figs..3-5 then were con-
(P
800- verted. Each mean pineal melatonin content from
(3.. N\\N
-
animals exposed to a particular color of light was ex-
b,\\\',] pressed as a percentage of the corresponding mean
g 600-
N
hxxx\n
pineal melatonin content from animals unexposed to

400- BN light. These converted data were then plotted on the

Q. graphs shown in Figs. 6--8. When the irradiance of the
different colored light sources was set at 0.928
200- pW/cm 2 (Fig. 6) it was observed that green light pro-

0
duced the greatest percent inhibition (82%) of mean
pineal melatonin content. Blue and yellow light also
339"37t 435-500 515-550 558-636 653~668
(Near U,V.) (Blue) (Green) (Yellow) (Red) produced a high percent depression of mean pineal
melatonin content (68% and 71% respectively). In
Peak Wavelengths-Nanometers
contrast, near-ultraviolet light produced a lower per-
Fig. 4. Influence of different light spectra at 0.200/~W/cm= on
cent depression (39%) of mean pineal melatonin
pineal melatonin content. Bars indicate m e a n pineal melatonin
contents and the lines on top of bars indicate S,E. Pineals were content, while red light failed to inhibit pineal mel-
collected from each group of light-exposed animals 20 min after atonin levels. These results indicate that at an irradi-
the lights were turned oni Pineals were collected from a corre-
ance of 0.928 #W/cm=. green fluorescent light is the
sponding group of u n e x p o s e d animals at the same time. For
each group n = 8. most efficient light for suppressing nocturnal pineal
 337

~' Syrian Hamsters Syrian Hamsters

Exposed to 0.928 iJW/cm 2 Exposed to 0.20 lJWlcm z
~00 I00-
-,i--
C t-
o O
o (.D
t- 8 0 ._c 80-
t- t-
O O
..i,.-
0

60-
-6 B
t- C
n-- 4O 40-
c-
O C
o
20~ 20-
n i G.)
rm

20O 360 400 5o0 ebo 7ha 0

200 560 400 500 600 700
m m m m m
(NearU.V.) (Blue) (Yellow) m m m m m m
(Green) (Red) (Near U.V.) (Blue) (Yellow)
Wavelength : N a n o m e t e r s (Green) (Red)

Fig. 6. The mean percent depression of pineal melatonin con-

tent by light spectra (0.928 1~W/cm2). The raw data for this
Wavelength: Nanometers
graph are shown in Fig. 3. Fig. 7. The mean percent depression of pineal melatonin con-
tent by light spectra (0.200/~W/cm:). The raw data for this
graph are shown in Fig. 4.
melatonin content. However, no statistical signifi-
cance was determined between melatonin levels of
animals exposed to 0.928 uW/cm 2 of green, blue or DISCUSSION
yellow light.
When the irradiance of the different colored light The data from the 3 experiments detailed above
sources was set at 0.200 #W/cm 2 (Fig. 6), it was ob-
served that blue light produced the greatest percent O" Syrian Hamsters
depression (91%) of pineal melatonin; similarly, "E
c-
I00 ~
green light also produced a high percent inhibition O
(71%). In contrast, yellow, red and near-ultraviolet
light produced very low or no percent depression
(19%, 1% and 0%, respectively) of mean pineal mel-
atonin content. These results indicate that an irradi- coO Green
ance of 0.200 uW/cm 2, blue fluorescent light is the
most efficient light for suppressing nocturnal pineal
melatonin content. However, no statistical signifi- n-
cance was observed between the mean melatonin g
contents of animals exposed to 0.200/~W/cm 2 of blue
and green light.
In the final experiment when blue and green light
sources produced irradiances of 0.186, 0.074 or 0.019 o~ 0 0.04 0.08 0.12 016 0.20
/zW/cm2, the blue fluorescent light consistently pro-
Light Irradiunce (IJW/cm 2)
duced a 2 4 % - 2 5 % greater depression of mean pineal
Fig. 8. The mean percent depression of pineal melatonin con-
melatonin compared to that caused by the green light tent by light spectra (0.186, 0.074 and 0.019/xW/cm2). The raw
(Fig. 8). data for this graph are shown in Fig. 5.
338
demonstrate that the blue fluorescent light source
with a half-peak band width of 435-500 nm was the
most efficient light for suppressing pineal melatonin
Green fluorescent light (510-550 nm) was the next
most efficient light followed by yellow fluorescent
light (558-636 nm). Of the lights tested, near-ultra-
violet (339-371nm) and red (653-668 nm) were the
least capable of inhibiting pineal melatonin. Interest-
ingly, most or all the irradiances of the different light
sources in the above experiments are much higher
than the animals require for the sensory capacity of
vision 23,24. Under all the experimental light irradi-
ances, the investigators could easily see the hamsters
and read the protocol sheets. The hamsters also
seemed to be able to see under all levels of illumina-
tion as evidenced by their attempts at avoiding cap-
ture. Very low irradiances (0.000001 ~W/cm 2) are
capable of eliciting retinal activity in hamsters, but
are not capable of suppressing pineal melatonin pro-
ductionl7, TM. The above study and earlier experi-
ments 2 indicate that the neural pathways connecting
the eye to the pineal have a higher threshold response
to light than do the retinocortical tracts. Other recent
data support this idea, In rats and cats. threshold irra-
diances of light which stimulate electrical activity in
the suprachiasmatic nucleus ( S C N ) 9 a r e much higher
than irradiances needed for the visual sense T M
Cardinali and colleagues pioneered the field of
testing the impact of different visible wavelengths of
light on the pineal gland 3. in their experiment, albino
rats were kept in continuous darkness for 7 days and
then exposed to a red. yellow, green, blue or near-ul-
traviolet fluorescent light for 12-96 h. Green light
was observed to produce the most rapid and com-
plete suppression of H I O M T activity followed by
blue, yellow, near-ultraviolet and red light. The
lights used in that study were from the same manufac-
turer (Duro-test) as were those used in the 3 experi-
ments described above. The results of the study by
Cardinali and colleagues differ from the findings of
the experiments reported here. Their experiment
showed that green light (510-550 nm) was the most
potent suppressor of pineal H I O M T activity, where-
as our studies clearly demonstrate that blue light
(435-500 nm) is the most potent inhibitor of pineal
melatonin. The differences between the study by
Cardinali and colleagues and the experiments report-
ed here may be attributed to species differences (rats
vs hamsters), protocol differences ~chronic vs .lcutc
light exposures), or the measurement ot different
endpoints ( H I O M T activity vs metatonin content !
Since the rodent retina is dominated bv rod photo-
receptors, the major photopigment is thought to be
rhodopsin 1~. The peak visible wavelength in the rho-
dopsin absorpuon spectrum is about 500 nm. Hence,
the data showing maximal suppression of metatonin
by light in the 435-500 nm range are consistent with
the hypothesis that rhodopsin i~ the photopigment
which mediates the pineal gland's response to light.
However. it is known that there is a small populanon
of cone photoreceptors in the rat:' mouse 4, and ham-
ster (N. Buyukmikei, personal communication.
School of Veterinary Medicine, University of Cali-
fornia. Davis. CAl. It has also been demonstrated
that there are color-sensitive photopigments other
than rhodopsin in the rodent retina 5. Cvanolabe, a
blue sensitive photopigment found in some mamma-
lian cones, has a peak wavelength absorption around
440 nm xg. Thus, the above data showing maximal
melatonin suppression by wavelengths in the
435-500 nm range are also consistent with the hy-
pothesis that a blue-sensitive photopigment may me-
diate the pineal response to light. Further studies are
required to clarify what photopigments are present in
the hamster retina and which photopigment mediates
the response of the pineal to light.
in an earlier study, it was discovered that hamsters
exposed to photoperiods of natural light had signifi-
cant differences in their peak melatonin production
compared to animals exposed to equal length photo-
periods of artificial ligh0. It was speculated that
those observed differences may have been due to dif-
ferences in intensity and spectral composition of nat-
ural and artificial light. Photoperiods of artificial light
are typically non-varying m intensity and spectrum of
light. In sharp contrast, animals kept under natural
photoperiods are exposed to wide ranges of light in-
tensity from 50.000 ktW/cm 2 for peak sunlight to
0.00005 /~W/cm 2 for overcast starlight 24, Further-
more. the spectral characteristic of solar illumination
changes continuously from dusk to dawn. Specifical-
ly, light from the sun around sunset and sunrise em-
phasizes the longer wavelengths (red) of visible spec-
trumS. The data reported here indicate the red
wavelengths of light were the least effective in sup-
pressing nocturnal pineal melatonin. It is possible

that the varying spectral characteristics of sunlight
are biologically m e a n i n g f u l in t e r m s of the control of
pineal physiology in animals in their natural habitats.
H o w e v e r , caution should be e x e r c i s e d in e x t r a p o l a t -
ing findings with m o n o c h r o m a t i c light to effects of
b r o a d - s p e c t r u m lighting. F u r t h e r e x p e r i m e n t a t i o n is
r e q u i r e d to d e t e r m i n e the exact role that solar spec-
trum has in regulating m e l a t o n i n p r o d u c t i o n .
T h e studies r e p o r t e d h e r e are the first direct d e m -
onstration that different w a v e l e n g t h s of light h a v e
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ACKNOWLEDGEMENTS
This w o r k was s u p p o r t e d by N S F G r a n t PCM
8003441 to R. J. R. T h e a u t h o r s wish to t h a n k Valen-
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