FY 77-78
Research Report
On The Impacts Of Ultra
Violet
Biological Systems:
A Study Related To
Stratospheric Ozone
tion
Submitted To:
The Stratospheric Impact Research
and Assessment Program (SIRA)
The L'.S. Environmental Protection Agency
Washington. D.C. 20604
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DISCLAIMER
THIS REPORT HAS NOT BEEN REVIEWED FOR APPROVAL BY THE
AGENCY AND HENCE ITS CONTENTS DO NOT REPRESENT THE VIEUS AND
POLICIES OF THE U.S. ENVIRONMENTAL PROTECTION AGENCY, NOR DOES
MENTION OF TRADE NAMES OR COMMERCIAL PRODUCTS CONSTITUTE ENDORSEMENT
OR RECOMMENDATION FOR USE.
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CONTENTS
SIRA FILE #
VOLUME I
1. Study Of Increase In Skin Cancer As A Function
Of Time And Age And Changing Stratospheric Ozone:
Need For Careful 1easure Of The Ultraviolet Dose 0 132.11
2. The Influence Of Age, Year Of Birth, And Date On
Mortality From Malignant Melanoma In The Populations
Of England & Wales, Canada And The White Population
Of The United States 0 132.31
3. Non Melanoma Skin Cancer Surveys In The United States
— An Environmental Epidemiologic Project 0 142.1].
4. Biological And Climatic Effects Research Terrestrial
Non—Human Organisms EXECUTIVE SUMMARY FOR 0 142.21 to 142.213
5. Biological Effects Of Ultraviolet Radiation On Plant
Growth And Function 0 142.21
6. Effects Of UV—B Radiation On Selected Leaf Pathogenic
Fungi And On Disease Severity 0 142.21g
7. The Effect Of Ultraviolet (UV—B) Radiation On Englemann
Spruce And Lodgepole Pine Seedlings 0 142.22
VOLUME II
8. UV—B Biological And Climate Effects Research 1/ 142.23
9. Ultraviolet Effects Of Physiological Activities Of -
Blud—Green Algage 11 142.24
10. Impact Of Solar UV—B Radiation On Crops And Crop
Canopies . . 0 142.25
II. High Altitude Studies Of Natural, Supplemental
And Deletion Of UV—B On Vegetables And Wheat 0 142.26
VOLUME III
12. UV—B Radiation Effects On Photosynthesis And
Plant Growth 0 142.27
13. Influence Of Broad Band UV—B On Physiology And
Behavior Of Beneficial And Harmful Insects 0 142.28
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SIRA FILE /1
14. A Study Of The Effects Of Increased UI/ —B
Irradiation On Environmental Dissipation Of
Agricultural Chemicals II 142.29
13 Biological Effects Of Ultraviolet Radiation
On Plant Growth And Development In Florist And
Nur e y Crops # 142.210
16. Biological Effects Of Ultraviolet Radiation On Cattle:
Bovine Ocular Squamous Cell Carcinoma 11 142.211
17. Radiation Sources And Related Environmental Control
For Biological And Climatic Effects UV Research (BACER) 11 142.212
18. Instrumentation For Measuring Irradiance In The
UI/—B Region . . . . . . . . . . . . . . . . . . . . . . . # 142. 213
1 Annual Report To EPA, Bacer Program For
Fiscal ‘lear .1978 . .... •*• tee .... ......s......,..a. .. . .••••, 142.34
20. Penetration Of UV— Into Natural Waters ... II 142.36
21 iligher Plant Responses To Elevated Ultraviolet
Irradiance /, 142.41
22 . Assessment Of The Impact Of Increased Solar
Ultraviolet Radiation Upon Marine Ecosystems ......... # 142.42
2 . UV—B lnstruinentationDevelopmcnt “.. #142.51
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IR’ FIL..& -
FINAL REPORT
UV—B RADIATION EFFECTS ON PHOTOSYNTHESIS AND PLANT GROWTH
W. B. Sisson
Jornada Experimental Range
U.S. Department of Agriculture
Las Cruces, New Mexico 88001
EPA—IAG—D6—O168
Project Officer:
R. J. McCracken
Agricultural Research, Science and Education Administration
U.S. Department of Agriculture
Washington, D.C. 20250
Prepared for
Environmental Protection Agency
BACER Program
Washington, D.C. 20460
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Introduction
A reduction of the atmospheric ozone layer through interactions with
oxides of nitrogen or halomethanes would result in increased levels of
terrestrial UV—B (280—320 mm) radiation (Green, Sawada and Shettle, 1974).
An increase in the !JV—B radiation component of the global irradiance could
result in significant biological effects since UV—B radiation is readily
absorbed by nucleic acid and protein chromophores (Giese, 1964).
The present study was initiated to evaluate plant responses to TJV—B
radiation as would occur under reduced atmospheric ozone concentrations.
Methods and Materials
All studies were completed in a fiberglass greenhouse. Four Westing-
house FS—40 ‘sunlamps’ were maintained approximately 1 M above the plant
surface. Each ‘sunlamp’ was filtered by either 5, 7.5 or 10 mu cellulose
acetate. The cellulose acetate filtered lamps were adjusted to produce a
gradient of UV-B radiation at the plant surface area. UV—B radiation mea-
surements were made daily with an IRL model 25 Radiometer (Optronics Lab.,
Silver Springs, Maryland). Spectral irradiance measurements were made
prior to treatment initiation and at least weekly thereafter with an Op—
tronics Laboratory model 721 Spectroradiometer. The control treatment
for all experiments consisted of four FS—40 ‘sunlamps’ filtered with 10
mu Mylar plastic film adjusted to the same configuration as the corres—
pondirtg UV—B radiation treatment. The cellulose acetate filters were re-
placed every third day and the mylar filteres were replaced every other
week.
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—2—
Net photosynthesis of single leaves was determined with a refrigerated
open gas circulation system with a model 365 Beckman IR gas analyzer.
Air and leaf temperatures within the curvette were measured with five—
wire (copper—constantan) thermocouples. A Cambridge model 880 dew—point
hygrometer measured changes in water—vapor concentrations. A constant
temperature of 27°C was maintained in the curvette during all photosyn-
thetic determinations. Total PAR quanta were monitored in the curvette by
a Lambda model LI—19OSR quantum sensor. One, 1500—W Westinghouse tunsten—
halogen lamp provided an irradiance level of 800 pe . m 2 . S 1 for all
photosynthetic determinations. Leaf areas were measured with a Lambda
model LI—3000 portable area meter. Photosynthetic rates are expressed on
a leaf area (one side) basis.
Indivudal shoots of alkali sacaton ( Sporobolus airoides ) were trans-
planted from the field into 10 cm pots. Upon initiation of the IJV radia-
tion treatment, all plants were defoliated to a one—inch height above the
soil’s surface. Thus, leaf growth as measured in this study is regrowth.
All other plant material was grown from seed in 10 cm pots in a t1 Jiffy—7
Plus” medium. The methods for acidic methanol—water extracts from Capsi —
cum frutescens follows the procedures outlined by Caidwell (1968). Absorb—
ance of the extract was measured with a Beckman Model DU spectrophotometer.
Quantification of the UV—B radiation dose was determined with the
equation:
r 91 4 [ 4 — (A/228.178)9]
UV—B 1 L 25 (X/228.178) J e (1)
Spectral irradiance of all UV—B radiation treatments employed in this
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—3—
study are presented in Figures 1 and 2.
Results
Leaf growth of alkali sacaton exposed to four UV—B radiation treat-
ments is shown in Figure 3. Each value represents the mean of 21 to 75
measurements. As shown in Figure 3, leaf growth was repressed as a func-
tion of the level of UV radiation. Table 1 presents the numerical values
and a statistical analysis of the data represented in Figure 3. The con-
trol treatment leaf length values were consistently different (P < .05)
from the UV radiation treated plants only at, and above a level of 7.5
mW . UV—B 1 . The 13.9 mW . m 2 UV—B 1 treatment resulted in repressed
leaf lengths statistically different (P < .05) from the other four treat-
ments.
Chile ( Capsicum frutescens ) responded to IJV—B radiation in a manner
similar to alkali sacaton through 22 days treatment (see Figure 4). That
is, total plant leaf area was repressed as a function of UV—B 1 . The values
represent the means of four or five plants and all treatment means differed
statistically (P < .05) except those plants exposed to 4.8 and 5.9 mW .
UV—B 1 . Similarly, after 55 days treatment, total leaf area, and wet and
dry weight of chile, differed between the control and five of the seven
UV—B radiation treatments (see Table 2). Variability of plants within
treatments C and E probably resulted in the lack of statistical differences
between these treatments and the control treatment. However, contrary to
the 22 day experiment (see Figure 4), there were no differences between
the iJV—B radiation treated plants for those parameters evaluated. Since
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Figure 1. Spectral irradiance of four Westinghouse FS—40 ‘sunlarnps’ each
filtered by 5 mu cellulose acetate yielding (A) 4.0, (B) 5.0,
(C) 6.8, (D) 9.1 and (E) 10.6 mW . 1J1 1 7_B 1 by equation 1..
Measurements were made in a fiberglass greenhouse with a model
721 Optronics Laboratory Spectroradiometer.
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100
.01
280
290
300
310
320
10
D
C
B
A
1
.1
WAVELENGTh (nm)
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FIgure 2. Spectral irradiance of four Westinghouse FS—40 ‘sunlamps’ each
filtered by 5 mu cellulose acetate yielding (A) 5.9, (B) 7.7,
(C) 9.8 and (D) 14.6 mW . UV—B 1 according to equation 1.
Measurements were made in a fiberglass greenhouse with a model
721 OptronIcs Laboratory Spectroradiometer.
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10
D
C
B
A
•0
100
.01
1
1
280
290 300 310 320
WAVELENGTh (nm)
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Figure 3. Leaf length (mm) of alkali sacaton ( Sporobolus airoides ) exposed
to four UV radiation (5.9 (s), 7.5 (A), 11.3 (X) and 13.9 (A)
—2
mW . m UV—B 1 ) and a control treatment (0). Each value repre—
sents the mean of 21 to 75 indivIdual leaves.
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1 2 3 4 5 6 7 8 9
80
70
40
n A tin an tnnvi A nnrn urn
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Table 1. Leaf growth of ALKALI sacaton ( Sporobolus airoides Torr.) exposed to four levels of UV—B
radiation (5.9, 7.5, 11.3 and 13.9 mW . m 2 UV—B 1 ) and a control treatment. Each value
represents the mean of 21 to 75 measurements.
UV—B 1 2
(mW.m)
Days
1
2
3
4
5
6
7
8
9
Control Treatment
255 a
391 a
515 a
636 a
748 a
853 a
935 a
988 a
1032 a
UV Treatments
A
59
176 b
314 b
448 ab
5 ab
6 78 ab
4 ab
8 62 ab
938 ab
995 ab
B
ab
307 b
384 b
473 b
562 b
648 b
727 b
800 b
846 ab
C
11.3
189 b
292 b
382 b
474 b
543 b
643 b
702 b
753 b
778 b
D
13.9
162 b
230 c
301 c
374 c
442 c
509 c
592 c
621 c
658 c
) 1
0
Values within the same column followed by different letters differ statistically (P < .05).
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Figure 4. Total plant leaf area of chile ( Capsicum frutenscens ) after 22
days exposure to 5 iSV radiation treatments (3.3, 4.8, 5.9 and
10.6 mW . n1 2 UV—B 1 ). Each value represents the mean of four
or five plants.
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I
3 5 7
30 -
I—
9
11
UV-B 1 (tnw’m 2 )
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—13—
Table 2. Total leaf area (cm 2 ), and wet and dry weight of chile ( Capsi —
cum frutescens ) exposed to 7 UV—B radiation doses (3.3, 5.1,
6.8, 7.7, 9.1, 10.0 and 10.6 mW . m 2 IJV—B 1 ) and a control
treatment. Each value represents the mean of three to six
replicates. A daily exposure period of 7 hours was used for
the 55 days of treatment.
UV—B 1
(mW . m 2 )
Wet
Weight (g)
Dry
Weight (g)
Total Leaf
Surface Area
Control Treatment
44.6
5.1
753.4
UV—B Treatments
A
3.3
32.7*
34*
582.6*
B
5.1
30.8*
3•3*
564.7*
C
6.8
35.7
3.9
765.3
D
7.7
28.2*
3.0*
528.8*
E
9.1
33.9
3.4
507.2
F
10.0
27.3*
2.9*
500.8*
C
10.6
33.1*
3 4*
613.6*
Values followed by asterisk (*) differ significantly (P < .05) from the
control treatment.
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—14—
it was only possible to irradite the topmost leaves of individual plants,
a shading effect may have been partially responsible for these results.
An alternative or additional factor responsible could be the absorption of
UV—B radiation prior to absorption by sensitive chromophores within the
leaves. Figures 5 through 12 represent the absorbance of methanol—water—
HCL extracts from the topmost leaves of the control and UV—B radiation
treated plants. All values represent relative absorbance since they were
normalized to Figure 12. Absorbance by this methanol—water—HCL extract
contains the flavonoids, some xanthophylls, cuticular waxes and other lip-
ids which represent efficient UV radiation absorbing compounds in plants
(Caidwell, 1968; Block, Durrum and Zweig, 1958; Geissman, 1955).
There were no apparent differences between the absorbance of the con-
trol treatment (Figure 5) extracts and those from plants exposed to 3.3
(Figure 6) and 6.8 (Figure 8) m . m 2 UV—B 1 . However, all other levels
of UV—B 1 resulted in increased absorbance within the UV—B radiation wave—
band. In addition, absorbarice tended to increase with increased levels of
UV—B 1 . These results tend to suggest that UV raliation absorbing compounds
are being produced somewhat according to the intensity of UV—B radiation
impinging on the exposed leaves. This increase in UV radiation absorbing
compounds would attenuate at least part of the UV—B radiation and, in part,
provide a protective screen to the more sensitive cell components.
A dose—response curve for photosynthesis of Cucurbita pepo f.s. early
summer crookneck exposed to four levels of UV—B radiation (5.1, 6.8, 7.7
aud 9.1 tiM . m 2 UV—B 1 ) is presented in Figure 13. The treatments were
initiated when the plants were 23 days old from the time of seed germina-
tion and all photosynthetic measurements were made on the first leaf after
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Figure 5. Relative absorbance of a inethanol—water—HCL extract from the
topmost leaves of chile ( Capsicum frutenscens ) in a control
treatment.
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1.0 -
0 I I
250 275 300 325 350 375 400
.75
.50 -
.25
I I — _I I
F
WAVELENGTH (run)
-------
Figure 6. Relative absorbance of a rnethanol—water—HCL extract from the
topmost leaves of chile ( Ca psicum frutescens ) exposed to 3.3
mW n 2 UV—B 1 for 7 hours daily for 55 days.
-------
j I I I
350 375 400
LO —
.75 — -
.50
.25 — —
0 —
250
275
I I
300
325
+
WAVELENGTH (nni)
-------
Figure 7. Relative absorbance of a methanol—water—HCL extract from the
topmost leaves of chile ( Capsicum frutescens ) exposed to 5.1
—2
niW • m UV—B 1 for 7 hours daily over 55 days
-------
1.0 - —
,,75 .
.50 -
.25 -
0 I I —+
250 275 300 325 350 375 400
WAVELENGTH (nm)
-------
Figure 8. Relative absorbance of a rnethanol—water—HCL extract from the
topmost leaves of chile ( Capsicuni frutescens ) exposed to 6.8
mW m 2 IJV—B 1 for 7 hours daily for 55 days.
-------
1.0 - -
.75 - —
.50 -
.25 —
0 I I
250 275 300 325 350 375 400
WAVELENGTh (rim)
-------
Figure 9. Relative absorbance of a methanol—water—HCL extract from the
topmost leaves of chile ( Capsicuin frutescens ) exposed to 7.7
—2
mW . m UV—B 1 for 7 hours daily over 55 days.
-------
I I I
I I I
I I I I I
300 325 350 375 400
1.0
.75
.50
.25
0
S
S
250
.,
275
WAVELENGTh (nni)
-------
Figure 10. Relative absorbance of a rnethauol—water—HCL extract from the
topmost leaves of chile ( Capsicuin frucescens ) exposed to 9.1
mW m 2 UV—B 1 for 7 hours daily over 55 days.
-------
250 275 300
I I I
I I I
325 350 375 400
a -
1.0
.75
.50
25
0
+
WAVELENGTH (urn)
-------
Figure 11. Relative absorbance of a methanol—water—HCL extract from the
topmost leaves of chile ( Capsicuin frutescens ) exposed to 10.0
—2
mW m TJV—B 1 for 7 hours daily over 55 days.
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1.0
.75 —
.50 -
.25 - -
0 I
250 275 300 325 350 375 400
WAVELENGTH (ruii)
-------
Figure 12. Realtive absorbance of a methano l—water—HCL extract from the
topmost leaves of chile ( C psicum frutescens ) exposed to 10.6
t i M . m 2 1W—B 1 for 7 hours daily over 55 days.
-------
1.0
I I I I I
I I I I I
250 275 300 325 350 375 400
.75 -
.50 - -
:25 - -
0—
WAVELENGTh (rIm)
I
-------
Figure 13. Net photosynthesis of Cucurbita pepo f.s. early summer crook—
neck exposed to four UV—B radiation treatments (5.1, 6.8, 7.7
and 9.1 mW . m 2 UV—B 1 ) expressed as a percent of the control
treatment plant rates.
-------
S
I I I I I I
2 4 6 8 10 12 1.4
E l
I
100 - -
90 -
80 —
70 -
6 0 -
50
S
S
UV-B 1 (mWm 2 * 10 )
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—33—
it had fully expanded.
Net photosynthetic rates of UV—B radiation treated plants appeared to
be repressed after exposure to approximately 6 * 1O mw . UV—Br and
continued to be reduced upon additional UV—B radiation exposure. This
same type of accumulative response in the depression of photosynthesis has
been shown earlier by Sisson and Caidwell (1976) for Rumex patlentia L.
Summary
The findings of this study are consistent with earlier biological
studies involving an enhanced terrestrial UV—B radiation component of the
global irradiance spectrum. That is, photosynthesis is depressed in an
accumulative manner and plant growth tends to be depressed as a function
of the level of UV—B radiation to which plants are exposed. However, it
is interesting to note that plants apparently possess an ability to res-
pond to UV—B radiation by synthesizing IJV—B radiation screening compounds
such as the flavonoids. To what extent this response would have in negat-
ing the IJV—B radiation induced damage cannot be determined at the present
time. Results of this study utilizing low levels of UV—B radiation, as
well as previous findings (Sission and Caidwell, 1977), would tend to sug-
gest that an intensified synthesis of UV radiation screening compounds
does not afford sufficient protection to sensitive target molecules within
plant cells of the more sensitive plants.
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-.34—
Literature Cited
Block, R. J., E. L. Durrum and C. Zweig. 1958. A manual of paper chro-
matography and paper electrophoresis. Academic Press, I’ ew York.
710 p.
Caldwell, H. N. 1968. Solar ultraviolet radiation as an ecological fac-
tor for alpine plants. Ecol. Mono. 38.243—268.
Geissman, T. A. 1955. Anthocyanins, chalcones, aurones, flavones and re-
lated water—soluble plant pigments. pp. 450—498. In: K. Paech and
M. V. Tracey (eds.). Monderne Methoden der Pflanzenanalyse, Vol. 3.
Springer—Verlag, Berlin.
Giese, A. C. 1964. Studies on ultraviolet radiation upon animal cells.
pp. 203—245. In: A. C. Ciese (ed.). Photophysiology. Academic
Press, I’ ew York.
Green, A. E. S., T. Sawada and E. P. Shettle. 1974. The middle ultraviolet
reaching the ground. Photochem. Photoblo. 19:251—259
Sisson, W. B. and N. N. Caidwell. 1976. Photosynthesis, dark respiration
and growth of Rumex patiencia L. exposed to ultraviolet irradiance
(280 to 315 join) simulating a reduced atmospheric ozone column. Plant
Physiol. 58:563—568.
Sisson, W. B. and H. M. Caidwell. 1977. Atmospheric ozone depletion: Re-
duction of photosynthesis and growth of a sensitive higher plant ex-
posed to enhanced U.V.—B radiation. J. Exp. Bot. 28:691—705.
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S3 n F/L / 0 2.
FINAL REPORT
INFLUENCE OF BROAD BAND Wi—B ON PHYSIOLOGY AND EHAV1OR
OF BENEFICIAL AND HARMFUL iNSECTS
Dora K. Hayes
Chemical and Biophysical Control Laboratory
Agricultural. Environmental Quality institute
Beltsville Agricultural Research Center
Beltsvllle, Maryland 20705
EPA—IAG—D6—0168
Pro - i ect Officer:
R. J. McCracken
Agricultural Research, Science and Education Administration
U.S. Department of Agricullure
Washington, D.C. 20250
Prepared for
Environmental Protection Agency
BACER Program
Washington, D.C. 20460
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DISC LA I ME R
This report has been reviewed by the Chemical and Biophysical Control
Laboratory, U.S 0 Environmental Protection Aqency, and approved for publication.
Approval does not signify that the contents necessarily reflect the views and
policies of the the U.S. Environmental Protection Agency, nor does mention of
trade names or commercial products constitute endorsement or recon iiendations for
use.
11
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FOREWORD
An increase in UV—B may be beneficial or harmful, dependinq upon the
effects of such an increase on pest species of insects and their host plants.
The task of the Chemical and Biophysical Control Laboratory was to investigate
the influence of broad band IJV-B on physiology and behavior of beneficial and
harmful insects in the following ways:
(a) Insects, where technology was available for rearing, were used to
determine the direct effect of UV-B on selected developmental staqes and on
interactions of insects with crops of economic importance.
(b) Effects of UV—B on metamorphosis, biological rhythms, and diapause
were investigated and the possibility that photorepair occurred was briefly
examined.
The investigators examined the direct effects of UV—B by conducting life
span determinations and by measuring egg hatch, life span and fucundity
in several generations of face flies and house flies. Pigment formation
after UV—B exposure indicated “sunburn. Host plant insect interaction studies
were conducted using soybeans and hush beans as hosts for the tobacco hudworm.
Cotton plants were reared to determine effects oF UV on pink boliworni hatch.
Effects of UV-B on life span of honey bees was measured, Other physiological
and biochemical effects could have been subtle. Effects of UV-B on metamorphosis
havealceady been mentioned. The capability for metamorphosis may have been
related to pigment formation and the capacity to diapause. Photorepair in vivo
was only touched upon; enzymatic activity and hiogenic amine levels were
examined; the rhythms in oxygen utilization provided a nondestructive technique
for examining effects of UV-B on insects.
1•11
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ABST R/\ cT
Our task was to study the influence of broad band UV—B on the physiology
and behavior of beneficial and harmful insects. This study was approached at
two levels: the first being to determine direct effects which could be observed
by determining mortality, observinq increased pigmentation or darkening, etc.,
and the second being to determine the physiological and biochemical effects
which require some further manipulation of whole or maccrated insects. To
determine the direct effects our laboraLory has investigated the following:
life span, egg hatch and adult emergence, pigment formation (by gross
observation), and host—plant in teracti on. The physi ol ogi cal and biochenii cal
effects investigated were as follows: metamorphosis, enzymatic activity,
pigment formation (detected biochemically), and rhythm using oxygen utilization
as an indicator.
Hatch of pink bollworm eggs, Pectinophora qossypiella (Saunders), as
decreased by at least 6 hr exposure to an irradiance level of 3,61 mWm’ BUV
(0.37 Wm 2 Abs) radidnt power between 280-320 nm. Life span oF the pink hollworms
which hatched from eqgs irradiated for 1 hr with UV-B was decreased. Exposure
throughout life of insects exposed during the egg stage to a regimen with no
darkness prolonged life of irradiated insects over those maintained in LD 16:8
in which the photoperiodic regime was manipulated. Diapause was potentiated -in
the pink bollworm i one test but not in a second after ii radiation of eggs for
1 hr with 3.61 mWm BUV (0.37 Abs) at 280-320 nm.
Eclosion of face fly, Musca autumnalis Dc Geer, pupae which have a calcereous
non—melanized puparium Was decreased by irradiating with UV-B 6 hr per day for
3-4 days. Fecundity and/or hatch of eggs from those face flies which emerged
was also decreased. No such effects were found on house fly pupae which are
melanized. Life span in young honey—bee, Apis mellifera L., workers was not
altered, nor were ultradian rhythms and level of °2 uptake altered by exposure
to UV-B.
Melanization in vivo and/or in vitro in larvae of the codling moth,
Laspeyresiap9mOflella (L.), face fly and tobacco budworn, Heliothis virescen (F.),
was increased by UV-B. No change in metabolism of 3rhydroxykynurenine was found.
Oxygen uptake of last instar codling moth larvae was increased by 1 hr of UV—B
irradiation, and ultradian rhythms were obliterated. In addition, in these
insects, release after treatment with triglycine or tetraglycine of material
absorbing at 3, 6.3, and 7.lpm was greatly enhanced by irradiation with UV B.
iv
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• . p . . 1
.2
.....p 4
.10
.15
• . • . . .23
9r
p I P • P
p .26
• 27
.30
• . . . • .a)
• . . . .36
CONTENTS
Foreword
Abstract . . . . . . • .
List of Illustrations
List of Abbreviations
Acknowledgments. . . . . • . . . .
iii
iv
• vi
vii
o . 0 P viii
1. Introduction
2. Conclusions and Recommendations •
3. Experimental Procedure and Results
Effects of UV—B on Mortality of Honey Bee.
Effects of UV .-B on Face Fly Pupae. •
Photorepair in vitro . •
Insect/Plant Interaction
Pigment Formation and Enzymatic Activity
UV—B and Changes in the Integument of
Lepidopteran Larvae • . . • . .
tJV—B and Oxygen Uptake . . . • • •
References . . • . • . • . . • . . • • . . . . • •
Bibliography . . .
V
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LIS1 OF ILLLJSTflJ\TIONS
Number
1 Top 3 photos illustrate differences in pigmentation observed after
exposure of face fly pupae to 8.014 mWm BLV (2.6 suns; 0.83 Wm 2 Abs)
compared to a control. The photo on bottom left shows portions of
3 control codling moth larvae. The larva in the center is not burned
dfld metamorphosis has occurred in the insects on the left and right.
The photo on the right illustrates pigmenLation of larvae after exposure
at a distance of 30 cm to a FS-40 lamp filtered with 5 mil cellulose
acetate. Exposures were for 6 hr/day for 4 days.
2a Oxygen upt ke after 10.02 iiiWm 2 BUV (3.3 suns; 1.04 Wm 2 Abs) to
13.36 mWm (.1.4 suns; L38 Wm 2 Abs) IJV—B irradiation by last instar
codling moth larvae with 02 uptake in 1 tl/g/hr plotted as a function of
time and calculated at 5 mi i i untervals
2b Similar to 2a 9 but determined at 15 miii intervals 0
2c Oxygen uptake by control codling moth larvae. Determined at 5 mm
intervals.
2d Similar to 2c. Determined at 15 mm intervals.
vi
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LIST OF /\BBREVJAT1ONS
A [ L ——Agricultural Equipment Ldboratory
DOPA --3 ,4-di hydroxyphenyl a] ani ne
IL ——Instrumentation Laboratory
3-OHK --3-hydroxykynurenine
PSL --Plant Stress Laboratory
UV-B ——Ultraviolet-B 280—320 nanometer portion oF electromagnetic
spectrum
hr -—hour
LD —-light: dark ratio, expressed in hours
PBW —-pink boliworm
vii
-------
ACKNOWLEDGMENTS
The assistance of the Plant Stress Laboratory, Plant Physiology Institute,
AR, SEA, in providing UV—fixtures, supports, and plants is grateful ty acknowledged.
Without the assistance of the Plant Physiology Institute Chairman, studies on
hatching of pink bollworm eggs could not have been done. The Aqricultural
Equipment Laboratory, Plant Physiology Institute, and the Instrumentation
Laboratory, Market Quality Research Institute, furnished assistance in
determining intensities and spectral distributions of UV light.
Insects were provided by AR, SEA, laboratories at Yakima, Washington,
Brownsville, Texas, and Plant Protection Institute, BARC, Beltsville, Maryland.
The assistance of Thor Lehnert, of the Bioenvironmental Bee Laboratory,
Plant Protection Institute, AR, SEA, was invaluable in determining the effects of
UV-B on bees.
viii
-------
I NTRODUCTION
The primary effort in the AR, SEA, laboratories at Beltsville was directed
toward developing techniques for irradiating biological samples iiith UV—B , which
is well characterized in terms of energy levels and spectral distribution.
Instrumentation Labordtory (IL) (Task X—XITJ), Agricultural Equipment
Laboratory (AEL) (Tasks XIV—XVJ), and the Plant Stress Laboratory (PSL)
(Tasks I=IV) have reported on equipment for determination of energy levels,
spectral distributions, and standardization of conceptual ways in which
exposures are expressed. The work reported here on insects suggests what
may or may not happen out—of—doors if levels of iJV—B radiation are increased.
In some instances, intensities are reported in “suns .” These values are
approximate numbers and in most cases, such intensities were provided to us
early in the testing process. Later data are reported in biologically
effective IJV-B in mWnr 2 BUV using the weightiug function A 9 described in the
AEL final report. Absolute values for UV are also given in t i n 2 .
1
-------
SECTION 2
CONCLUSIONS /\ND RECOMMENDATIONS
a. Hatch of pink boliworm eggs was decreased by at least 3 hr exposure
to UV—B. Eclosion of face fly pupae is decreased by three or four 6 hr
exposures to UV—B; eclosion of house fly pupae is not affected. As in
higher life forms, a pigmented exterior protects the organism from damage that
is easily discerned. No effects on fertility of house Flies were observed,
hut UV B irradiation of pupae decreased fertility and fucundity of face fly
adul ts.
b. Gross observations indicated increased pigment format-ion in larvae
of the codling moth and pupae of the face fly alter exposure to UV B. In
vitro increase in rates of inelanization in homogenates of irradiated insects
in the presence or absence of 3,4—dihydroxyphenylalanine might be paralleled
in vivo .
c. UV—B irradiation of tobacco budworm larvae on soy beans and bush beans
had little effect 0
d, Metamorphasis from larva to pupa was delayed in the codling moth by
irradiation with UV—B.
e. No effects on 3—hydroxykynurenine metabolism but some effects on
3,4—dioxyphenylalanine (DOPA) metabolism were observed after irradiation with
UV—B. This kind of activity may be related both to biogenic amine metabolism
and to the capacity to diapause. Aniines such as dopamine may be involved in
expression of intracellular effects of UV—B irradiation.
f. No effects on induction or prevention of diapause in the pink bollworm
as a result of exposure of eggs to UV-13 were noted.
g. Capacity for in vivo photoropair may exist in insects; more work should
be done.
h. Oxygen uptake in the codling moth is increased 1 hr after exposure to
UV-B. Oxygen uptake in young honey bee workers is not affected.
i. Infrared analysis of wash solutions from irradiated insects suggests that
UV—B may solubilize some cuticular components.
RECOMMENDATIONS
If increased, UV—B will damage some insects at those stages of development
during which these insects do not have adequate pigmentation to filter out
harmful wavelengths. Honey bees and some other beneficial insects probably
contain enough pigment to protect them. However, some eggs, larvae, pupae and
adults of both beneficial and pest insecs will be damaged if the cuticle
contains little pigmented material or material which will absorb MV-B.
2
-------
MILESTONES — EXTENT OMPLETION OF TASKS
REPORTING PERIOD JANUARY—DECEMBER 1977
Task VII. Influence of bread band UV—B on physiology and behavior of beneficial and harmful insects.
Oct Nov Dec Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec
a. Direct Effects
Life span determination _______ ________
Egg hatch ____________________
Life span — fecundity — not cOmpleted — insufficient time allowed by granting agency
2 to 3 generations
P1 gment formation - gross
observation
Host-insect-interaction — inadequate time to complete study _______________
Adaptability - inadequ te time
lItuIIJILIl1flhLII
b. Physiological and Biochemical Effects
Me tarno rph os i s ______________________ _______________________
Pigment formation — biochemistry - superficial observation
Capacity for diapause — one test, one species
Photorepair in vivo — incidental to life span study
Enzymatic activity — merged with pigment formation _________ _______
Biogenic amine levels A ________ ____
- LI I Ill
Rhythm - 02 utilization - continuing _________ _______
c. Final Report
A Begin Study (Projected Milestone) Revised Starting Time (Projected Revised Milestone)
L Complete Study (Completed Milestone) G Completed Revised Project (Completed Revised Milestone)
Completed Activity Projected Activity Revised Activity
-------
SECTION 3
EXPERIMENTAL PROCEDURES AND RESULTS
UV-B, EGG HATCHING AND LIFE SPAN IN THE PINK BOLLWORM
The effects of UV-B on the hatching of eggs of the pink boliworm supplied
by the Animal and Plant Health Inspection Service Laboratory, USDA, Phoenix,
Arizona, have been examined. Several different modes of exposure were employed.
A preliminary test to determine effects of UV on hatchability indicates that
FS-20 lamps filtered with either cellulose acetate or polystyrene inhibit egg
hatching; 8 out of 2 a minimum of 100 eggs hatched after exposu ’e to UV-B
between .349 mWm BUV (3.1 suns; 0.97 Wm Abs) — 13.34 mWm BUV (4.4 suns;
1.38 Wnr Abs). Other early studies summarized in Tables 1 and 2 showed that
high levels of UV—B reduced egg hatch. However, UV—B levels used were
unrealistic in light or further work by IL and R [ L,and the data shown in Table 3
indicate that continuous 2 exposure to UV—B for 6 hr results in some inhibition
of hatching at 9,319 mWm BU\J (3.1 suns; 0.97 Wnr 2 Abs).
Apparently, insects exposed at contro1” levels in the boxes used for UV
exposure were in some instances (Boxes 1, 2, 4, and 5) exposed to enouqh UV-B
so that hatching was inhibited.
Irradiation oF pink boliworm eggs with UV-B did not affect incidence of
diapause in larvae reared from these eggs. When larvae from one test were
irradiated with 3.61 niWm 2 BLJV (0.37 Wnr 2 Abs) energy in the 280—330 nm region
of the spectrum, adult survival in LD 16:8 as well as in two regimens in which
the photophase was shifted, was affected. A mortality of 50% in the controls
was observed 34 days after eggs were placed in the regimen LD 16:8 at 25 0 C.
In irradiated insects 50% mortality was observed 21 days after insects were
placed in LD 16:8 while in non—irradiated insects oniy 22% mortality was observed.
In the irradiated insects rate of death decreased so that at slightly less than
50% mortality in control differences were not significant, while at slightly
more than 50% mortality significant differences were observed. (See Tab t e 4).
4
-------
Table 1 l:ootnotes
a! Filtered through polystecene.
b/ Control held at 24° C L [ ) 16:8. Control was not exposed to UV-B.
C l Samples to be irradiated iere placed 30 cm from the FS-20 lamps.
Filters were used as indicated. Actual irradiance levels are not known.
6
-------
TABLE 2. EXPOSURE PBW EGGS ” TO UV ON 12/9/76 - 6 HR EXPOSURE.
Treatment
Live
larvae
Empty
vials
Dead
larvae
Total empty
vials +
dead larvae
%
live
early
death
late
death
Total
no.
Dark A
18
7
2
9
66.6
25.9
7.5
27
Dark C
15
6
1
7
68.2
27.2
4.5
22
Box 1 - FS-20
bare lamp
Box 2 - FS-2cW
cellulose acetate
filter
2
12
3
14
—
1
3
15
40.0
44.4
60.0
51.8
—
3.8
5
27
Box 3 - FS-2O
plexiglass filter
53
24
6
30
64.1
28.9
7.0
83
Box 4 — FS-20
lamp
Z
7
—
7
22.2
77.8
-
9
Egqs exposed 2 days after oviposition.
Energy level approximately 10.0 mWm 2 BUV (3.3 suns; 1.04 Wm 2 Abs) -
13.34 mtIm BUV (4.4 suns; 1.38 Wm 2 Abs) (see PSL report). Samples
to be irradiated were placed 30 cm from the FS—20 lamps. Filters
were used as indicated. Actual irradiance levels are not shown.
-------
TABLE 3. HATCHING OF PINK BOLLWORM EGGS AFTER EXPOSURE TO VARIOUS LEVELS OF UV-B LIGHT
Box
no.
Description
Irradiance
Set—up
No.
(3
hatched
hr) ’
No.
(6
hatched
hr) 1
mWm 2 BUV
SE 1
Wm 2 Abs
1
FS-2O bare lamp
(neutral density filter)
18.09
5.23
—
5.9
1.7
-
1.9
0.54
high
low
control
4
96
40
2
4
10
2
FS—2O — polystyrene
10.59
2.84
-
3.5
0.9
—
1.1
0.29
—
high
low
control
72
44
52
4
18
12
3
G 15—T8 lamp
254 line of W/ spectrum
22.52
4.80
—
7.4
1.6
-
2.3
0.5
—
high
low
control
8
36
60
0
2
30
4
FS—20 — cellulose acetate
aged 6 hr
5 mu
10.38
2.72
—
3.4
0.9
—
1.1
0.3
—
high
low
control
27
76
36
12
14
8
5
FS—20 — cellulose acetate
aged 6 hr
10 mu
9.64
2.17
-
3.2
0.7
-
1.0
0.2
—
high
low
control
54
44
28
4
8
4
Sun equivalents.
Numbers of eggs varied from 60-100. Pink bollworm females fix eggs to filter paper.
We ,judqed that it was deleterious to remove eggs from paper.
-------
TABLE 4. MORTALITY OF PINK BDLLWORM AFTER IRRADIATION ’ or [ GGS WITH UV-B
LIGHT FOLLOWED BY DELAYS IN ONSET OF TH [ PH0TOPHASE ’
Time from start to 50% mortality
Larvaeb/ Adults only Larvae, pupae,
found_— days adults - days
CONTROL (LD 16:8)
Not Irradiated 27 37 34
Irradiated 22 35
SHIFT-90° DELAY IN LD 16:8
EVERY
3 DAYS 27 31 27
5 29 22 ’ 2O ’
6 30 28 ! 24 ’
LL 28 34 33
3.61 mWm 2 BUV (0.37 Wn1 2 Abs) Energy - 280-320 nm; irradiated for 1 hr.
Temperature - 26°C; 40 vials per condition. Larvae found refers to larvae
which developed at least to 3rd instar and were observed after infestation
of medium.
After 50% mortality occurred, death rate decreased in irridiated insects so
that when 48% mortality occurred in control, difference was not significant
using chi—square but when 52% mortality :as observed in controls, difference
between control and irradiated was significant at <.025.
Significantly different from the control with P <.01 and .005 respectively
using chi—square when mortality oF 50% control mortality was determined.
9
-------
EFFECTS OF UV B ON MORTALITY OF THE HONEY BEE
Honey bee workers were e poscd at the time in the life SPdfl during which
they would normally first leave the hive.
The lamps were set up o that insects were irradiated at 7.32 mWrn 2 BUV
(0.76 Wm 2 Abs) with a 5 mu cellulose acetate filter and 2 FS—40 lamps 50 cm
from the top of the carton containing the insects. When suitable holding
conditions were obtained, in 3 tests, no differences in survival were observed
between irradiated and test insects. No differences were cietected when curves
describing oxyqen uptake as a function of time over a time span of 2 hr or
less or 24 hr were inspected visually.
Lamp set up : 2 FS-40 lamps, insects 50 cm from lamp filter: Cellulose acetate,
aged 6 hr 5 mu.
Expt 1 - single layer on lamps
Expt 2 — single layer on lamps
Expt 3 - single layer on lamps
Results are summarized in Thbles 5, 6 and 7.
10
-------
FIG. 1. CONTAINER FOR HONEY BEE TESTS
:1
- I
I
1
I ‘
. : .
• ; lt - ;A: ,r ‘ j .te . :
11
-------
TABLE 5. UV-B AND HONEY BEE SURVIVAL
Experiment 1 :
Honey bees (4-5 days) were received 6/27/77 and maintained with sugar cube
and water in dish with styrofoam on top of water 0 11 containers of bees
each - 9 for UV and 2 for control in rear of room 0 (See Fig 1 for container)
Single layer 6 hr old CA filter (5 mu) on the lamps.
Irradiation — 6/28/77, 6 hr 11:00 — 5:00
Irradiation - 6/29/77 — 6 hr 11:00 — 5:00
Total Death Counts :
Contai ncr
No 0 6/28 6/29 6/30 7/1 7/2 7/3 7/4
1 0 1 1 1 3 5 8
2 0 1 1 1 6 8 10
3 0 0 0 0 5 9 14
4 0 0 0 4 4 5 6
Irra- 5 0 0 1 3 4 5 5
diated 6 0 0 0 2 2 3 4
7 0 1 1 4 6 9 10
8 0 0 0 0 1 2 3
9 0 0 0 1 1 2 2
control (‘lO 1 3 5 5 6 7(mold in containers)
t. ii 0 0 0 0 4 6 9
Conclusions : Initial death rate as high as first 4 experiments. Could be due
to H?0 supply lasting longer. Note — even with lower mortality used only 1 layer
CA/2layersl
a/gummy with sugar.
12
-------
TABLE 6. UV-B ANI) HONEY BEE SURVIVAL
Experiment 2 :
Honey bees (4-5 days old) were received on 7/5/77 and maintained with sugar cube
and water in dish - styroFoam on top of 120. 11 containers — 20 bees each, 9
for UV, 2 for control in redr of room.
Single layer of 6 hr old CA filter (5 mil) on lamp.
Irradiation - 7/6/77, 6 hr 10:00 — 4:00
Irradiation — 7/7/77, 6 hr 10:00 — 4:00
Total Death Counts :
Container
No . 7/6 7/7 7/8 7/9 7/10 7/11 7/12
1 0 0 0 0 0
2 0 0 0 N N 0 0
3 0 0 0 0 0 0 0
4 0 0 0 T T 0 0
Irra- 5 0 0 0 1 4 (2 drowned)
diated 6 0 0 0 D D 2 3
7 0 0 1 0 0 2 2
8 0 0 0 N N 0 0
9 0 0 0 E E 0 0
/10 0 0 0 0 0
contro1 11 0 0 0 0 0
Conclusion : There was no initial death due to lamps.
13
-------
TABLE 7. UV—B AND HONEY BEE SURVIVAL
periment 3 :
Honey bees (4—5 days old) were received on 7/12/77 and maintained with sugar cube
and water dish with styrofoam on top. 11 containers of bees with 20 bees
each, 9 for UV-B and 2 for control in rear of room.
Single layer of 6 hr old CA (5 mil) on lamps
Irradiation - 7/12/77, 6 hr 11:00 - 5:00
Irradiation - 7/13/77, 6 hr 11:00 — 5:00
Total Death CounLs :
Control
No. 7/12 7/13 7/14 7/15 7/18
1 0 0 1 1 1
2 0 0 0 0 1
3 0 0 0 2 2
4 0 0 0 1 4
Irra- 5 0 0 0 3 5
diated 6 0 0 0 0 0
7 0 0 0 0 3
8 0 0 0 1 2
9 1 1 1 1 1
fio 0 0 0 1 3
control 0 0 1 3 5
Conclusion : There was no initial mortality due to UV—B.
14
-------
EFFEC1S or LJV—B ON FACE FLY PUPAL
The effects of ultraviolet light on eclosion of face fly and house fly
pupae 1—3 days after pupariurn formation were determined. Pupae wer exposed
to irradi nces of approximately 6.68 mWnr 2 I3UV (2.25 sun ; 0.69 Wm Abs), 2
8.01 mWm BUV (2.6 suns; 0 83 Win— 2 Abs), and 10.02 mWiii BUV (3.3 suns; 1.04 Wm
Abs). Light was supplied by 2 FS—40 tubes in one fixture fitted with a
reflector prepared by AEL. Irradiances were determined by the IL with the
Norris spectroradiometer; one large lot of polystyrene petri dishes was employed
for both the irradiance measurements and the subsequent tests, since these
dishes had been evaluated as filters during determinations of irradiance.
The UV—B light appeared to have no effect on eclosion of the house fly as
shown in Table 8. In fact, exposure to UV -B may have enhanced emergence. Data
such as those reported in Table 7 which indicate that UV—B had little effect on
house fly eclosion, were obtained at least in 3 more experiments. However, the
emergence 01 the face fly was affected by UV-B and perhaps the face fly could
serve as biological dosj,meter. When pupae of the,)face fly were exposed to
approximately 8.01 mWm BUV (2 25 suns; 0.69 Wm Abs), (Table 9) eclosion was
prevented in 30-50% of the insects 0 There was some variation from lol to lot,
with some lots showing almost no emergence. Burned face fly pupae are illustrated
in Figure 2. (Figure 2 also illustrated the pigmentption occurring when codljnq
moth larvae are exposed to approximately 10.02 mWm BIIV (3.3 suns; 1.04 Jm Abs
to 13.36 mWnr 2 BUV (4.4 suns; 1.38 Wnr 2 Abs) with UV—B fo a 6 hr span each
day for 4 days.)
In further tests newly formed face fly pupae were irradiated by 5 intensities
of UV—B to determine whether such exposures would increase pupal mortality or
decrease adult fecundity, longevity, or eqg viability. The tests were conducted
by placing 100 newly—formed (<1 day old) pupae at each of 5 spots which were 32 cm
beneath two 40-w BL fluorescent lamps which were covered by a sheet of 5 m thick
cellulose acetate. The cellulose acetate was aged for 6 hr prior to any test
of pupae and was replaced after every 18 hr of exp sure to UV. Radiance was
measured as fractions of a “standard sun” 3.06 mWm BUV in weighed units. The
pupae were held under 16 hr of cool white fluorescent lighting per day at 78°
± 2° F and 40% ± 5% RH. Pupae were exposed each day for 3 days for periods of
0.25, 0.5, 1, 3, and 6 hr per day. (Table 10).
Exposure of pupae to 6.73 mWm 2 BUV (0.69 Wm 2 Abs) or more of UV-B radiation
for periods of 1 hr per day for 3 days caused 77% pupal mortality and complete
loss of adult fecundity (Tables 10 and 11, Fig. 3),
Since pupal exposure to UV-B over a 3-day period resulted in a high mortality,
we exposed 4-day old pupae to UV-B for periods of 1, 2, 3, and 6 hr, during 1 day
only, to see whether the developing gonads could he affected without killing the
pupae. When the pupae were exposed for 3 hr or less, pupal mortality and adult
fecundity were unaffected, but egg hatch was reduced by 25-41% (Table 12),
Future work will be directed towards using UV as an additional method of
sterilizing male face flies.
15
-------
TABLE 8. HOUSE FLY ECLOSION ’ AFTER EXPOSURE TO UV-B ’
No. eclosed Avq.
Control 64 64
2.2 “suns” ’ 98 88
78
2.6 “suns” ” 71 75
78
3.3 “suns’s-” 85 85
at One hundred house fly pupae exposed in polystyrene
petri dishes.
b/ January 10, 1977 run.
C l Energy deterniined by AEL and IL.
16
-------
lADLE 9. EMERGENCE OF ADULTS OF THE FACE FLY, MUSCA AUTUMNALIS ,
AFTER EXPOSURE OF PUPAE ’ TO UV -B- ’
% adult
Irradiance ’ ccl osion
Test 1 (1/14/77 )
Control 43
(33—53)
2.2 suns 67
2.6 suns 47
3.3 suns 14
(0-27)
Test 2 (2/1/77 )
Control 65
2.2 suns 64
2.6 suns 45.8
3.3 suns 24
(1 8-3O) ’
Five pupae/group, Test 1; 25 pupae/dish, Test 2. Single
determinations run except where 2 values in parenthesis
indicate duplication of test.
UV-B supplied by 1 UV fixture equipped with a reflector
and t io FS-40 lamps. Cellulose acetate (5 mu).
Irradiance in suns as described in text of report.
Fifty insects in one dish.
17
-------
6 hr for
3 hr
1 hr
1/2 hr
114 hr
98 90 95
98 98 100
83 81 84
22 33
12 14 26
95 93
99 100
77 87
39 35
19 13
28 94.2
22 99
22 82.4
26 32.8
19 16 ,8
92
98
77
13
C
x
65.2 61.0 57.6 65.8 65.6 23.4
2.2 suns 6.73 mWm 2 BUV (0.69 Wm 2 Abs)
3.1 suns 9.35 mWm 2 BUV (0 97 Wm 2 Abs)
3.3 suns 10.02 mWm 2 BUV (1.04 Wrn 2 Abs)
3.5 suns 10.69 mWm 2 BUV (1.10 Wm 2 Abs)
3.7 suns 11.35 mWm 2 BUV (1 17 Wm 2 Abs)
TABLE 10. EFFECTS OF UV-B RADIATION
(>265
nni)
on face fly pupae
- 1977
Duration
of
Pupal mortality (%)
Corrected
mortality
Standard suns ’
exposure
2.2
3.1 3.5 3.5
3.7
check
3 days
II II
II II
II
II
-------
TABLE 11. EFFECTS OF UV-B RADIATION ON FACE FLY OVIPOSITION AND HATCH - 1977
No. of
eggs
per female
% Hatch
ndard
suns -’
Sta
Standa
rd suns - 1
Duration
of
exposure
1.6
3.1
3.3
3.5
check
0.75
1.4
1.5
1.6
1.7
check
6 hr for
each
of 3 days
0
0
0
0
2
75
3 hr
I’
0
0
0
0
3
83
1 hr
‘
1
o
0
0
0
2 1
91
1/2 hr
“
“
1 9
1.9
1 9
1 9
2
75
65
63
80
79
66
1/4 hr
“
“
1.8
1.8
1.9
2.0
1.9
59
60
35
74
95
93
a!
1 sun = 3.06 mWm
1.6 suns 5.0 niWm 2 ABS (0.52 Wm 2 Abs)
3.1 suns = 9.35 m4rn BUV (0.97 Wm 2 Abs)
3.3 suns = 10.02 rnWm 2 BUV (1.04 Wm° 2 Abs)
3.5 suns = 10.69 mWm 2 BUV (1.10 Wm 2 Abs)
-------
TABLE ia. UV-B AS A STERILANT FOR FACE FLIES ’
No. of each
1 hr
stage
2 hr
after
3 hr
exposure
6 hr
to UV-B for:
0 hr
Pupae
100
100
100
100
100
Adults
97
100
94
95
100
Eggs
91
96
100
62
100
% Hatch
45
56
44
43
75
Co rrc cte d
% mortality
40
25
41
43
a! Pupae were exposed to 1l.3mWm 2 BUV (3.7 suns; 1.17 Wm 2 Abs)
20
-------
Fig. 2
r iaf
a
‘5-
HL
S’,.c S
7 .
P P
&
Figure 1.
_____
• S
•
r
.—..
‘ :
. 5 5 5—S . 4
¶ V.,. . . ‘ . -
5,. 5/ • , , ,..
4 S ;‘ . •/• .(_.
I l
. 5 .—
1 ‘‘ .
I !
‘;. !
- - --
5/,Al/y is4 Itc d
I
4.
&g ;•f t ,
.:,: •..
, 5 i 5
) ‘ 5 .
j3. J( , }. ri e J ‘e O’ .’p3e
(‘i’tt i
\ J
.5.; ., .
. . ç L.
S : . , —; ‘
S. ,. .. 5 ,4 .. .
- M
, ,“
S J , . -.
: ‘ ‘. •.5, 5- “:
; ‘ d ‘ :I. .
S ... 5.
_ 55,• , s5_ s _ S . ‘
- :
1, 5. _ . ‘ ‘ - “ 5 .. .5 . .5
I . ..
5 ‘ • ‘ ,.. . , . .jr s —s - -:., -
(5 .5 . 2... : . ‘q . I
7 . J
/ (41/4.4 )
/
-------
Fig. 3
tOO
8o-
.2 ‘ S
I
I ’
4O
I
20
O i I:
2 4
Hours of exposure per day
Effects of W I B on face fly pupae
22
-------
PHOTOREPAIR IN VIVO
Possible occurrence of a repair enzyme was examined in one test, summarized
in Table 13. In this test, larvae of the codling moth, the pink boliworm, and
the European corn borer were divided into 3 groups, witr one group serving as
a control, A second group iias exposed at a di stance of 30 cm to two FS—40
lamps fitted with a mylar filter, and the third group to two FS—40 lamps
fitted with a 5 nil cellulose acetate filter for 6 hr. After e posure , a
portion of each group was held under room lights (about 4.5 uW/n ), and a
second portion was placed under two 15—watt daylight fluorescent lights at a
distance of 6 cm.
23
-------
TABLE 13. POSSIBLE REPAIR ENZYME ACTIVITY IN LARVAE OF TWO SPECIES OF LEPIDOPTERA
N.)
The European corn borer. Ostrinia nubilalis (H lbner), larvae had crawled under
had been provided as a moisture source and, thereby, escaped the UV radiation.
dampened filter paper which
Control
Cellulose acetate
Repair
Darkness—
Insect
LD 16:8
regimen
no light
Control
Repair
Dark
Pink bollworm
I pupa
1 larva
2 escaped
1 adult
1 pupa
3 larvae
I adult
1 pupa
2 larvae
1 dead
1
2
3
larva
pupae
adults
1
4
adult
pupae
2
2
1
larvae
pupae
escaped
Codling moth
6 adults
1 deformed
pupa
1 pupa
2 larvae
egg masses
present
5 adults
1 larva
egg masses
present
6 adults
3 deformed
pupae
1 escaped
egg masses
present
5
2
1
2
adults
pupae
black
larvae
dead
larvae
1
4
1
3
1
deformed
adult
adults
deformed
pupa
pupae
dead
larva
2
1
2
6
adults
egg mass
pupae
dead larvae
-------
INSECT/PLANT INTERACTIONS
The major aspect of this phase of Task VII has been learning how to conFine,
irradiate and recover insects used in studies of insect—plant interaction.
Trials with cages constructed of wire supports and nylon netting that completely
surrounded infested bean plants in pots showed that this . ,as not a satisfactory
method of simultaneous exposure of plants and insects, since the insects escaped.
At present the method of choice for small plants, such as soybeans and bush
beans, is to place the pest insect, in this case the tobacco budw’ rm, HelioLhis
virescens, on the plant in an aluminum and glass cage 26.5 x 26,5 x 26.5 mm
with galvanized aluminum screen on top and sides.
Two FS—40 tubes in one fixture were placed 31 cm from the screen top of
the cage. One thickness of 5 mil cellulose acetate was used as a filter. Ihe
screening, acting as a neutral density filLer, absorbed about one third of
the incident light 0 When 5 tobacco budworm larvae were placed on the one bean
plant in the cage and the cage and its contents were exposed to Wi—B during
the last larval instar (about 4 days) at 250 C three survived and pupated and
2 died. Since the tobacco budworm is cannibal istic the 2 deaths may have
resulted from injuries occurring during attacks of one insect on another. The
3 pupae were deformed and no adults emerged. In a control cage 2 larvae pupated
and emerged as adults and 3 died. It is anticipated that some ouLdoor exposures
can be conducted in these cages.
In further studies, when insects on plants were irradiaLed (5 plants) with
5 insects/plant) no differences were observed between control and test insect
survival and qualitative observations suggested that irradiated insects were
slightly more vigorous.
25
-------
PIGMENT FORMATION AND ENZYMATIC ACTIVITY
Tests to determine effects of irradiation with UV-13 of last instar codi ing
moth larvae on subsequent in vitro activity of polyphenoloxidases were
conducted y exposing test insects to 1JV B at irradiance l vels of approximately
9.349 mWm BUV (3.1 suns; 0.97 Wnr 2 Abs) or to 13.36 mWm BUV (4.4 suns;
1.38 Wm Abs) for 3, 2 , or 2 hr, homogenizing in (0.25W) sucrose in 0.lM
phosphate buffer, pH 7.0, that was deoxygenated with UHP nitrogen for 5 mm
prior to placing insects in the so:ution.
Enzyme activity was measured by determining time until browning of a
mixture of 0.1 m of homogenate (O..3 g codlinq moth larva in 5 ml bufFer) and
0.1 ml of 3, 4—dihydroxy-phenylalanine (3 mg/lU ml solution) as described by
Hayes, Johnson, and Schechter (1975). Control homogenates were prepared from
insects that were not irradiated.
Rates of browning differed on different days, but polyphenoloxidase
activity was increased by exposure to UV—B. For example, a mixture prepared
at 1552 hours from DOPA and homogenate from irradiated codling moths was
observed to be blackened by 1645; the homogenate prepared from control insects
required at least another hr to melanize.
When sensitivity to Uv—B as a function of time of day was Lested, we found
that “browning” or melanization occu red more rapidly dfter 1 hr of irradiation
(10.02 mWnr 2 BUV (3.3 suns; 1.04 Wm Abs)) at 1030 and 1130 than at 0830,
0930, 1230, and 1330 hr.
Analogous reactions may occur in tanning, or even sunburn in higher
animals. It is likely that this kind of redction in insects represents
activation of polyphenoloxidase by injury. Such reactions occur ‘ihen insects
are cut or homogenized and in the studies reported here, rate of damage and
possibly formation of autocatalytic products could occur in the intact,
irradiated insect, as evidenced by Figure 1, in which 3 sunburned cociling
moth larvae which failed to pupate are illustrated. The companion figure shows
portions of 2 pupae and 1 larva jith the control larva appearinq colorless and
the pupae appearing normal in reproduction.
Attempts are being made to quantitate activity of this enzyme. Since
oxygen is required for polyphenoloxidase activity, enhanced activity of this
enzyme could be partially responsible for increased 0 activity reported in some
insects in another section of this report.
26
-------
UV-B AND CHANGES IN THE INTEGUTIENT OF LEPIDOPFERAN L1\RVAE
Pink boliworm larvae, codling moth larvae and European corn borer larvae
were irr diated for 1 or 2 hr with IJV—B at 7.32 mWm BUV (2.4 suns;
0.76 Wm Abs), using a 5 mu cel lulose acetate filter and FS—40 lamps as
already described.
Using a microliter syringe, 3 microliters of 0.01 M triglycine or
tetraylycine was applied to the back of a larva to determine its effect
and the effect of UV—13 on the integrity of the ntegument. The droplet
stayed on for five minutes and then was washed off with 0.1 ml of distilled
water. Care was taken so that neither anterior nor posterior ends of the
insect became wet. The insect was removed and .2 ml of acetone was added to
stop the reaction. Distilled water served as a control.
This process was repeated three times using different larvae of the same
age and species and the final solution (9 microl iters triy]ycire, .3 ml
distilled water, .6 ml acetone) was placed in a l ) nil centrifuge tube and then
evaporated to dryness using a water bath (60 — 7Q) C) and a vacuum line, The
crystals left behind after evaporation of the solvents were removed and made
into a KBR disc for use in the Packard infrared spectrascope (IR).
The curves obtained by the controls showed that triglycirie removed
some substance (possibly fatty acid or wax) from the integument of each
species tested. After UV—B treatment for one hour, the irradiated insects
treated with distifled water showed a curve similar to that of a non.irradiated
insect treated with triglycine and even more integumental material appeared
in the wash solution 0 Typical IR curves are included. (Figs. 4a and 4h).
27
-------
Fig. 4a
WAVELENGTH (MICRONS)
6
7
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U 0.2
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4 5
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7
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-------
UV-B AND OXYGEN UPTAKE
In the determinations of the possible effects oF UV—B on oxygen uptake,
the insects used to date have been codlinq moth larvae, Laspeyresia pomonella
(L,). Thinning apples were infested with eggs and shipped to Beltsville by
the AR Laboratory, Yakinia, Washington. When the insects arrived, they were
maintained at 24° C ± 10 C under a lighting reqimen of 16 hr light, 8 hr dark
in a 24 hr day. The experiments were run with last instar larvae, 2 insects!
run. The irradiated larvae were exposed for 2 hr periods to ES O-4O watt
ultraviolet lamps which had been aged 100 hr 0 Insects were held in polystyreiie
petri dishes which had been shown by the IL to transmit tJV— equivalent to
9.35 inWm BUV (3.1 suns; 0.97 Urn- 2 Abs) to 13.34 mWnr 2 BUV (4.4 suns;
1.38 Wnr 2 Abs) when suspended 15 cm above the insects. The control larvae were
placed in the same room to receive the same background levels oF exposure to
white light (3 Wm 2 ). After eYposureô the larvae were placed in calibrated
chambers and submerged in iater at 24 C. The experimental design is described
by Hayes, Schechter, Mensins, and Horton (1968) and includes a polarographic
electrode to measure the oxygen concentration. Minute by minute readings were
collected from each larva and rate of oxygen uptake in jtl O /rng/hr was
calculated using an equation cited by Hayes. The Data System Application
Division, AR, SEA, developed the program for determining 02 uptake by insects and
supplied the computer interface. The insects .ere allowed to equilibrate for
the first hr; after that time, length of 02 uptake determinations varied in
time from 2 hr to 48 hr.
Figures 5a—d show that exposure For 1 hr to UV—B results in an initial
increase of 02 uptake and a subsequent damping of the short-term variations
(ultradian rhythms) in oxygen utilization. This has been a characteristic
pattern in the 02 uptake of codling moth larvae exposed to UV—B. Such a
pattern is probably also observed in other injured species. We can speculate
that it represents, in part, the increased metabolic activity required to
repair injuries.
30
-------
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-------
REFERENCES
1. Hayes, D. K., A. Johnson, and M. S. Schechtor. Inhibition of Melanization
in Homogenates of Larvae of the Tobacco Budworm, Heliothis vireccens F.
(Lepidoptera: Noctuidae). Comp. Biochem. Physiol. 50B:609-611, 1975.
2. Hayes, D. K., M. S. Schechter, F. Mensing, and J. Horton. Uptake of
Single Insects Determined with a Polarographft Electrode. Anal. Biochem.
26:51—60, 1968.
35
-------
BIBLIOGRAPHY
Cohen, S. H., J. A. Sousa, and J. F. Roach. 1973. Effects of UV Irradiation
on Nymphs of Five Species of Cockroaches. J. Econ. Entomol. 66:859—862.
Cohen, S. H., J. A. Sousa, F. J. Roach, and J. B. Gingrich. 1975. Effects of
UV Irradiation on Nymphs of Blattella qermanica and P jp1aneta americana.
1• Econ. Entomol. 68:687-693.
Depner, K. R. 1965. Ultraviolet Irradiation of Cattle in Relation to Diapause
in the Horn Fly, Haeinatobi i irritans (L.) (Diptera: Muscidae). mt. J.
Bionieteor. 9(2) 167—170.
Fujiwara, Y. 1975. Postreplication Repair of Ultraviolet Damage to DNA,
DNA—chain Elongation, and Effects of Metabolic Jnhibitors in Mouse L Cells.
Biophys. J. 15:403-415.
Fujiwara, Y., and Kondo, T. 1974. Postrepiication Repair of Ultraviolet
Damage to DNA in Xeroderma Piqmentosum, Other Human and Mouse Cells in
Culture. J. Radiat. Res. 15:81-89.
Hart, R. W., and R. B. Setlow. 1974. Correlation Between Deoxyribonucleic
Acid Excision—Repair and Life—Span in a Number of Mammalian Species.
Proc. Nat. Acad. Sci. 71:2169-2173.
Kaithoff, K. 1976. An Unusual Case of Photoreactivation Observed in an Insect
Egg ( Sniittia spp., Chironomidae: Diptera). Photochem. Photobiol. 23:93-101.
Kraemer, K. H., H. 6. Coon, R. A. Petinqa, S. F. Barrett, A. E. Rahe, and
R. H. Robbins. 1975. Genetic Heterogeneity in Xeroderma Pigmentosuni:
Coniplementation Groups and Their Relationship to DNA Repair Rates, Proc.
Nat. Acad. Sci. 72:59—63.
Lehmann. A. R., S. Kirk-Bell., C. F. I\rlett, M. C. Paterson, P. H. M. Lohman,
E. A. de Weerd-Kastelein, and D. Bootsma. 1975. Xeroderma Pigmeritosum
Cells With Normal Levels of Excision Repair Have a Defect in DNA Synthesis
After UV—Irradiation. Proc. Nat. Acad. Sci. 72:219—223.
Mennigmann, H. D. 1974. Caffeine Inhibition of Dark—Repair: Lack of
Reversal by Adenosine-(3’5’)-Monophosphate. mt. i. Radiat. Biol. 26:
101 —106.
36
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S lIM f/Lfr- /‘7 ,
FINAL REPORT
A STUDY OF THE EFFECTS OF INCREASED UV—B IRRADIATION ON
ENVIRON IENTAL DISSIPATION OF AGRICULTURAL CFIE IICALS
J. R. Plirnmer
Organic Chemical Synthesis Laboratory
Agricultural Environmental Quality Institute
Beltsville Agricultural Resoarch Center
Beltsville, Maryland 20705
EPA—IAC—D6—0l 68
Project Officer:
R. J. McCracken
Agricultural Research, Science and Education Administration
U.S. Department of Agriculture
Washington, 1).C. 20250
Prepared for
Environmental P4olection Agency
BACER Program
WashingLon, D.C. 20/460
-------
In trod tic t ion
Many synthetic compounds have been introduced into the biosphere
through hun-ian activities. Agricultural chemicals are widely dissipated
in the environment and exposed to many natural processes. Sunlight is
an extremely effective natural force in degrading or destroying many
synthetic chemicals and phenomena such as bleaching, fading of dye—
stuffs, and drying of paints are examples. Inorganic compounds are
generally quite resistant to the action of sunlight.
Many fertilizers and some of the older inorganic pesticides may
be included among resistant chemicals. however, the large—scale pro-
duction of synthetic organic compounds has changed the situation to a
considerable degree. Since the 1940’s, there has been continuous ex-
pansion in the production of synthetics derived from benez noid com-
pounds. Herbicides, insecticides arid fungicides of this group have
been used in increasing quantities.
The absorption spectrum of benzene and its derivatives is generally
characterized by intense absorption in the 250 — 260 nat region of the
ultraviolet spectrum. The absorption weakens or falls of f towards 300
nm. A necessary precondition for photochemical reaction is the absorp--
don of light, so that the possibility of chemical reaction increases
rapidly as the wavelength of incident light is decreased.
A small proportion of the solar spectrum falls below 300 rim, but
although this radiation is low in intensity, this radiation is much
more energetic than higher wavelengths. Energy of radiation is in-
versely proportional to wavelength, therefore the shorter wavelengths
in the near to middle ultraviolet (320 — 280 nat) are photochemically
active. They possess sufficient energy to disrupt many linkages be—
tween atoms in molecules. Thus photochemical reactions induced by
light below 300 nm generally involve bond breaking or rearrangement.
Photochemical products nay have altered toxicity, biological activity
or capacity to undergo degradation by comparison with the parent sub-
stance.
Secondary processes may also be responsible for photochemical
alteration of molecules. The process of sansiti ed photolysis in—
volves absorption of light by a molecular species which does not re-
act, but this T excited’ species brings about reaction in a non—absorbing
species. The process of photosensitization may be con-mon in the
environment, since natural waters contain unknown sensitizer molecules.
Rates of photodecomposition may often be greater in natural waters
than in distilled water. Reaction with oxygen under the action of
light or ‘photooxidation’ is also a con-mon phenomenon responsible for
thc dngradation of many organic molecules.
-------
—2--
Such phenomena alter the effectiveness of pesticide molecuies. For
example, the p hro ds are particularly susceptible to photochemical
breakdown, but a wide variety of molecules may be affected. Research
into the area of pesticide photodecomposition has been tarried out for
a number of years. The information developed is of potential value in
developing future strategies of pesticide use and maintaining pesticide
efficacy.
Development of reliable information on the hazard and toxicology of
1)hotOProducts is limited by (i) the ignorance of their chemical nature,
(ii) a lack of standardized procedures for inv sLigation of photochemical
reactions, (iii) a lack of information about the macro and micro environ—
men s to which pesticides are exposed in the field. This includes the
intensity and distribution of solar energy, effects of adsorption on soil,
dust, or foliar surfaces and the effects of sensitizers.
This information is needed if pesticides are to be used effectively,
since residual activity may be considerably reduced by irradiation. Pro-
tecting agents and formulation techniques must be developed. Conventional
pesticides have received some study and progress has been made in the
pyrethroid field toward the solution of the problem by incorporation of
new features in the molecule. Problems may also be encountered in the
successful utilization of potential alternative pest control methods such
as microbial agents, insect juvenile hormones, phenomones, chemosterilants,
etc.
If chemicals are to he effectively used in agriculture, it is important
that optimum conditions be defined for their effective use. Co .a.un.d.s..
wh-ie ntially labile in - ed. Methods of
protection against sunlig t should be sought and evaluated. Increased
solar radiation on the earth’s surface will result in increased photo—
alteration of organic molecules. There should be an effort to estimate
the quantitative significance of this increase in order to determine
whether it will affect farm production and economics.
-------
—3—
TilE PIIOTODEGRADAT.LON OF AGRICULTURAL CHEMICALS AND RELATED
ENVIRONMENTAL POLLUTANTS
The first year’s work was planned as the basis of a continuing
program in photochemistry. The most essential facilities of a photo—
chemical laboratory were installed, and the Physlcal and analytical
techniques for the measurement of photoehemical quantities were selected
and tested. The accumulated experience from this phase of the program
provides the background for its future expansion.
A number of pesticides, herbicides and related environmental
pollutants were examined for suitability for study with the available
apparatus in the B region of the ultraviolet spectrum. Finally, the
set of nineteen chlorinated phenol isomers was chosen for the first
large—scale comparison of a family of related compounds.
1) Apparatus for the Measurement of Quantum Yield
After a preliminary survey of the literature concerned with the
measurement of quantum yield, it was decided to make initial determina-
tions at 313 nanometers, because that is the only usable wavelength
of the medium pressure mercury arc that falls within the B region of
the ultraviolet.
A Itmerry_go_rOufldtl photolysis apparatus of the general type
described by Moses, Liu and Monroe (1) was built. The support and
drive mechanism were the standard components of the Ace Class Company
Model 6543 Turntable Photochemical Reactor. The sample holder and
window system similar to that described by Moses was designed and
constructed to our specifications. The aluminum parts were anodized
black, and the apparatus was assembled and installed in a temperature—
controlled water bath. To provide monochromatic radiation at 313
nanometers and to remove heat produced by the light source a cooled
circulating filtering system was constructed (2).
2) Operation of the Photochemical Reactor
The photolysis apparatus was installed in the small inner room
of a double darkroom. The power supply and controls were located in
the outer darkroom. For safety, the power connection for the mercury
arc was made through an electrical interlock on the door between them.
In tests with the filters covered by aluminum plates, no signifi-
cant stray light was detected by ferrioxa]ate actinometer tubes in the
sample holder.
-------
—4—
The light intensity was found to increase by a large factor during
a warm—up period. The extent to which this behavior depends upon the
characteristics of the power supply was left for further investigatinn
when time permits. For some of the earlier, shorter exposures, the
problem was avoided by allouirtg a sufficient warm—up period, followed
by a brief shutdown for sample insertion. For longer runs, the effect
was not disturbing. ike intcgrated light exposure, of course, depends
upon the actinometer analysis, not upon the clock.
3) Actinometry
The potassium ferrioxalate actinometer was used as described in
Nurov’s Handbook of Photochemistry (2). In that method, two separate
solutions of ferric sulfate in sulfuric acid solution and of potassium
oxalate are made up and standardized. The two solutions arC stable
indefinitely, and the actinometer is readily prepared by mixing and
diluting them. Some recrystallized potassium ferrioxalate was also
obtained for comparison. The extinction coefficient of the phenanthro—
line complex was determined for the available ultraviolet spectrophoto—
meter, a Beckman DB—GT, by preparing a series of concentrations from a
standardized ferrous sulfate solution, measuring the absorbances at
510 n.m and applying a least squares trearaent to the resulting data.
Although other workers have reported using very short exposures of
ferrioxalate actiiiometers before and after long runs, the dependence of
light intensity upon a warm—up period suggested using an actinometer
capable of integrating light exposure throughout the longer photolysis
times, which the dilute and sensitive ferrioxalatc actinometer cannot
do. The more concentrated uranyl oxalate actinometer was used, and
the oxalate was titrated with potassium pei anganate solution. Later,
one of the organic ones, such as the malachite green leucocyanide actino—
meter, will be used.
4) The Selection of Compounds for Photolysis at 313 rim
Ultraviolet absorption spectra of a number of pesticides and herbi-
cides were measured in water, ethanol and hexane solution, and a few
candidates were selected for initial experiments. Pcntachlorophenol was
chosen for a study of the dependence of quantum yield on con entration.
The choice of water as a solvent to simulate environmental conditions
may, in some cases, require the use of concentrations J.ower than those
usually used for quantum yield determinations. This problem has been
discussed by I . G. Zepp etal. (3). The analytical problems suggest the
use of compounds labeled with radioisotopes for counting or with stable
isotopes for mass spectrometric multiple ion detection methods.
5) Analysis of Phenols
In order to permit the use of a chromatograph that could be convenient-
ly obtained on loan, a method was developed for the determination by high
-------
—5—
pressure liquid chromatography of phenols recovered from their aqueous
salt solutions by acidification and extraction. It has been used for
the sodium salts of chlorinated phenols.
6) The Calculation of Quantum Yields
Each of the twenty tubes irradiated in the photochemical reactor
contained 4.00 ml of solution. Small losses of solvent by evaporation
need not be considered, because the entire contents of the tube ware
quantitatively transferred for analysis. Because the volume was snall,
it was often convenient to combine the contents of several tubes into
a single analytical sample. For example, ten tubes of uranyl oxalate,
in two group5 of five, provided two identical samples for permanganate
titration, and ten tubes of pentachiorophenol sodium salt solution, in
two groups of five, could be processed to give two extracts for chroma—
tographic analysis. This technique need not result in any accumulation
of volumetric errors, since, where it is justified, the original 4.00—mi
aliquots can be dispensed from a huret in such a way that only the initial
burnt reading for tl-’ first tube and the final buret reading for the last
tube are significant measurements for the group of tubes combined.
The oxalate in the uranyl oxalate actinometer was determined before
and after irradiation. The difference was used to calculate the number
of cinsteins absorbed by a tube. Let n be a number of moles/tube or a
number of cinsteins/tube, V be volume in liters ( 0.00400 liter/tube),
N be normality and M be molarity of solution. Let the subscripts I
denote initial values (before irradiation) and F denote final values
(after irradiation). The number of moles of oxalate ion destroyed per
tube is n = and the number of einsteins absorbed per tube is n
2 4
t n = ___ n —n,
c 2 0 4 I
V(N — 1 F
When the oxalate is titrated and calculated as oxalate ion concentration,
N = N
2
I (V t — blank) X N
+ 1/2 1 1n0 4 I 1n0 4
L V’oxalate
V’ is the volume of permanganate or combined oxalate sample in milliliters.
Since the quantum yield of the uranyl oxalate actinometer at 313 nm is
known (S. L. Murov, Handbook of Photochemistry ) to be
0.561 L n
C 2 0/ 4
-------
—6—
we find that the number of cinsteins absorbed per tube is
An
c 2 0 4 —
0.561
The number of einsteins absorbed pe tube is the same for all tubes
(actinometer and samples) because (1) the window size and shape are
the same, (2) the r tdial distance from the lamp, the light distribution,
etc., are averaged by the rotation and are therefore the same and (3)
in this method of operation the absorbance of the solution in each tube
is sufficient that essentially all of the acident light is absorbed.
If the volumes V ’ 1 are chromatographic injection volumes in micro—
liters, the molar concentrations of the PCP salt solutions before
extraction are calculated as
7 _ vu
Ill X PC?, std .
M =M XIPCP
PCP PCP, std.
L 11 PCI’, std.
in which II is the chromatographic peak height in chart divisions, pro-
vided that the volumes of the aqueous PC? sample and PCP standard solu-
tion are equal, and that the volumes of the corresponding extracts are
equal. The injected volumes are chosen to give approximately equal
chromatographic peak heights, in order to avoid nonlinearity problems.
The number of moles of PC? destroyed per tube if found by difference,
as in the case of the actinometer. (Both the unirradiated and irradiated
solutions are always extracted. The unirradiated one is the standard
solution.)
p p = -
Finally, the quantum yield for PC? disappearance at 313 nm is
= number of moles of PCI’ destroyed
313 number of einstcins of light absorbed
A. _An
‘ 3l3 — PCP
7) The Quantum Yield as a Function of Concentration
The apparatus and techniques were first applied in the measurement
of quantum yields for photodegradation by 313—nanometer radiation of
-------
—7—
penlachiorophenol, present as the salt in aqueous sodium bicarbonate
buffer solution (4) . The extension of this experiment to lower concen-
trations could be done with isotopically labeled material, as noted
above. Results for the higher concentrations of PCP are presented in
Table 1.
Table 1. Quantum Yield for Degradation of Pentachiorophenol Sodlu-ri Salt
Starting Irradiation PCP Padiation Quantum
Concentration Time in Destroyed Intensity Yield
of PCP Hours (%) (E/tube/min) at 313 nm
0.000939M 2.00 28.0 9.1 x 0.0096
0.000751 1.60 27.3 9.7 X o— 0.0088
0.000751 0.80 8.8 9.1 X 0.0061
In the first run for 2 hours, beginning with 0.000939M PCP of which
28.0% was destroyed, a quantum yield of 0.0096 was measured. In the
second run, beginning with 0.000751M PCP, the time was reduced to
1.60 hours, to give about the same depletion of the PCP (27.3%), as
expected. The quantum yield was a Little smaller, 0.0088. In the
third run, the time was reduced to 0.80 hour. The 0.00075lM initial
concentration was reduced by 8.8% by photolysis, with a quantum yield
of 0.0061.
8) Photodegradation of the Chlorinated Phenol Isomers
As the concluding step in the initial program, a comparative study
of the quantum yields for photodegradation of the complete set of
nineteen chlorinated phenol isomers has been begun. Some of them are
important as environmental pollutants (5). The scheduled completion
of this phase was interrupted by problems. The first was unavailahity
of analytical assistance at the most critical point. The second was
the inadequacy of the laboratory ventilation system to remove volatile
irritants in some of the runs. As a general safety measure, the
photolysis apparatus is being moved into an available small fume hood.
Therefore the examination of this set of compounds has been suspended
until after the reinstallation of the apparatus.
-------
—8—
References
(1) F. Guy Moses, Robert S. H. Liu and Bruce M. Monroe, Molecular
Photochemictry 1, 245—249 (1969)
(2) Steven L. Murov, Handbook of Photochemistry , Marcel Dekker, New
York, 1973.
(3) R. C. Zepp, Unpublished ork.
(4) C. W. Hiatt, W. T. Haskins and L. Olivier, The American Journal
of Tropical Medicine and Hygiene 9, 527—531 (1960).
(5) 1\kio Yasuhara, Akira Otsuki and Kelichiro Fuwa, Chemosphere 6,
659—664 (1977).
-------
,c,#e a
FINAL REPORT
BIOLOGICAL EFFECTS OF ULTRAVIOLET RADLATION ON PLANT
GROWTH AND DCVELOPI• NT IN FLORIST AND NURSERY CROPS
Peter Semeniuk
Florist and Nursery Crops Laboratory
Plant Genetics and GermpLasm Institute
Beltsville, Maryland 20705
EPA-’IAO—D6—Ol 68
Project Officer:
R. 3. McCracken
Agricultural Research, Science and Education Administration
U.S. Department of Agriculture
Washington, D.C. 20250
Prepared for
Environmental Protection Agency
BACER Program
Washington, D.C. 20460
-------
Biological Effrct , of 1JlLr iiiolet Radiation on Plant
c;rowtli and J)evclopnent n I or st an(l Nursery Crops
1/
Peter Semen 1
Science and Education Administration, Federal Research,
U.S. Deparlinent of Agriculture, Be]tsville, MD 20/05
Abstract. Under simulated UV—B enhancement conditions, florist and
nursery plants were irradiaLed in a glass greenhouse using Westinghouse
FS—40V fluorescent sun]a lnps filtered with 5 mu cellulose acetate
fillers for 6 hours (10:00 a.m.—/i :00 p.m.) each day for 2—12 weeks. A
simulated sun curve and a computer—generated action spectrum were used to
determine biologically effective dosage to provide a wide range of UV—B
irradiances from 50—400% increase in biologically effective UV (4—13
weighted mWm 2 ). Visible injury was noted in only 8 of 74 florist and
nursery species. Leaves of ‘C—I—White’ and ‘C—I—Red’ poinsettia showed
d istorLion, glazing and abnormal leaf cu ivature. Of all 16 Coleus culti—
vars, ‘Glory of Luxemburg’ was the most sensitive and ‘Pineapple’ the most
resistant to 50% or greater increase in biologically effective UV exposure
for 2 weeks. Breakdown in anthocyanin conicut ias observed in 15 of the
16 cultivars, with leaf extracts and fresh leaves of ‘Red Rainbow’ showing
a large increase in absorbance at 285 nm. Aster, Flollyhock, tmpatiens and
Vinca plants irradiated for 4 weeks developed slight chiorosis (degrada—
tion of chlorophyll). Stress response was noted i ii all 6 Browallia
species and cult-ivars. None of the 12 shrub or 6 tree species irradi aled
showed any sign of stress response to 12 weeks of UV—13.
1/
Research Ilorticul tui-i st, Florist and Nursery Crops Laboratory, Plant
Genetics nnd Germplrism Institute.
2/
Mention of a trademark name, proprieLary product or vendor does not con—
sti lute a guarantee or warranty of the product by the U. S. Depart:iii ’iir
of Agi-ictil Lure and (1005 not imply iL’-. approval to the excli ,c,o,i ol other
p rodiic L c or vendors t hat may ii so he ciii iii ’ e
-------
2
Biological effecLs of UV—A (320—4fl0 run) and T N—C ( I ess than 280 nm)
radLat ion in plants are well documented (2, 5, 6). A great vaiiely of
physiological and morphological plant responses to U ’ ] radiation has been
demonstrated over the past 75 years. However, most of these experiments
have employed ultraviolet lamps which emit radi ation unlike the radi ation
present in the normal terrestrial solar spectrum (3, 4).
The effects of IF /—B (280—320 nm) radiation on man have been studied
extensively but relatively little is known of the effects of UV—B
radiation on growth and development of plants U, 2, 6). The impact of
low level s of UV—B radiati on on plant growth and developmonL is largely
unknown since most UV—B studies were conducted with lamps that emitted
both I N— B and IN—C.
Previous biological studies in the UV—B region were greatly limited
by a lack of sui table rnonitoriTlg equipment and inadequate narrow and
broadband radiation sources. Under the aurpices of the Environmental
Protection Agency (EPA) interagency program on Biological and Climatic
Effects Research (BACER), the USDA Agricultural Equipment Laboratory (8)
and the Instrumentation Research Laboratory of Bellsville developed
improved experimental facilities for irradi aCing, measuring and moni Coring
1W—B radiation. My objective was to identify, through a screening program,
species and cultivars of economically imporLant florist and nursery plants
that aie scn i Live or i e s i ci ant to UV—B r:,dialiei; iind.’r giacc grreiiiinti i’
cond i t ions, oil I i ii ng standardi zed techniques and improved ins triimentat ion.
Observations i eic made on the extent of visual injury, change rn
pLgmenlation, leaf abscission, glazing, bronzing, chiorosis and other
plant stress responses.
-------
3
Materials ned Methodc
Experimental facil ides for UV—B i rrrtdi at. ion were developed n
coopera ti on with Lhc Bel tsville Agricul tural Equi pment Labora tory and
he Instrumentation Research Laboratory. The former developed Lhc lamp
configuration, weighting functions and irradiance levels at selected
locations; the laLter developed the instrumentation (spectroradiometer)
for measuring and monitoring UV—B radiation.
Westinghouse FS—40 fluorescent sunlamps were used in conjunction with
approprtatc plastic fil ters to provide increased levels of IJV—B i rradia—
t] on. Under simulated UV—J3 eiihancemen t condi ti ons, El oris t and nursery
plants were irradi ated in the greenhouse under 5 nil c cl J ilosc acetate
(UV—A & B) or 5 mu Mylar (UV—A only) for 6 hours continuously each day
for 2—12 weeks about solar noon (10:00 a.m.—4:00 p.m.). Minimum night
temperature was kept about 170 C and day temperatures did not exceed
24°.
The standard protocol established uas for FS—40 sunl amps to he aged
100 hours before use; cellulose acetate (CA) filters presolarized for 6
hours; filters changed twice weekly; and lamp height adjusted with each
filter change to maintain UV levels. PlanL were irradiated under li
fixtures, each containing 2 FS—Li0 sunlamps spaced 30 cm apart and 82 cm
above plants. A simulated sun curve and a computer—generated action
cpectrum were used to determi iw biologically effective UV—1l do .nge
Experimental p1 OUtS izere placed in selt’cted locations to provide a wide
range of UV—B irrad ances from 50—400% increase in hi ol ogi cal ly of foci lye
UV (4—13 weighted m Um 2 ). Irradiance levels were verified at the
beginning and end of experiments by measuring wi L ii an Optronic
Laboi .1 tom-ics inc. , Model 725 UV—B Radi nine Lcr .
-------
4
Results and Discussion
A list of plants tested for the relative sensitivity or resistance to
enhanced UV—B radiati on in gi ass greenhouse is shown in Table I . Vi s ihl e
injury w s noted in only 8 of 74 florist and nursery species (Table 2).
The remaining 6G species examined were found to tolerate in increase in
biologically effective levels of UV from 50—400%. The most common plant
response to high levels of UV—B irradiation (100% or greater increase in
biologically effecLive DV under FS—40 sunlaraps filtered with 5 ml] CA) was
breakdown of antliocyanins and chlorophyll, or more commonly, a glazing arid
bronzing of the tissue which is generally attributed to the presence of
oxidized, polymerzed, phenol ic compounds. This bronzing or darkening
1 e11omenon serves as a convenient indicator of DV absorption and damage.
Visible lesions, loss of chlorophyll and reduction or increase in piginen--
tation were not immediate but rather appeared slowly as reflections of
disturbance in the basic metabolic process. The stress responses did not
develop until 3—4 days after beginning UV exposuie.
‘C—I—White’ and ‘C—I--Red’ poinsettia planLs were particularly sensi-
tive to UV—B radiation and showed glazing, distortion and abnormal leaf
curvature that increased in severity with increased UV—B levels. Under
high levels of UV—B irradiation (100% or greater increase in b ological1y
effective DV), leaves of ‘Supreme Annette hlegg’ poinsettia formed
purplish— red anthocyanins, a typical stress response. It is possible
that ‘Supreme Annette Hegg’ has the capaci ty to prevent damage to
sensitive tissue from Dy—B radiation by producing materials in epiderinal
cells that absorb the radiation. Fl avonoi ds haVe been shown to absorb
effectively in the Dy—B
-------
5
radi ation regi on (7). Si [ I CC anthocyan is were identified ii the epi der—
hid I cells of p01 fl Ctti IS , 1 t Seems likely that these compouiick SCI VC 10
protect the inner cells of the icaveb agai nst further ultraviolet d ininge.
Pineapple’ was rue only one of 16 Coleus cu] tivars that showed no
visible pigment changes when exposed to artificially enhanced UV—B
irradiation.
Of the 16 cultivar-s, Clory of Luxemhurg’ as the mosi sensitive. It
was the only one that showed a slight loss of anthocyanin pigmentation at
50% increase in biologically effective 1W.
The remaining 14 cul ti vars of Coleus reqihi red at least 100% i nd-ease
in biologically effective UV before showing visible pigment breakdown.
Exposure of Coleus blumei ‘Red Ra inbow’ for 2 weeks to enhanced UV—B
radiation at ca 100% i.n biologically effective UV produced no significant
effect on dry matter accumulation, fresh weight, leaf area, or plant
height; however, the red pigment in young leaves was greatly reduced. A
small change in degradation of the anthocyanin pigment was evident alter 6
hours of exposure. This change was intensified with increased exposure up
to 24 hours (4—6 hr UV—B radja Lion). Both leaf extracts and fresh 1 eavc
showed a large increase in absorbance at 285 nm.
Visible breakdown in antluocyanin content was also observed in 13 oF
time other 14 cultivars exposed to 50% or greater increase in hiologi cally
cLfccLi ’e UV [ or 2 weeks. This decreise ni Llie aniliocyan ill Cotit iit of
Coleus Leaves suggests the usefulness of these plants as indicators of
UV—B damage.
-------
6
Aster, Ilollyhock, Tmpati ens and V uca p1 ants i rradi ated for /i weeks
showed on] y a s ibt1 e UV c tress response that was cha t -ac ten zed by
chiorosis (degradation of chlorophyll). The injurious effects of UV—B
radi ation were soon hia kcd by the growth of new 1 eaves which emerged from
the termi nal and lateral buds ía] lowing the cessation of l i v exposure. The
length of time necessaly for this regrooth was dependent upon the degree
of injury.
Fatsia japonira was the oniy one of the 19 tender perennial species
in which plant growth was slightly inhibi Led by 100% or great-cr
biologically effective UV. None of the 12 shrubs or 6 tree cpecios
irradiated showed any sign of stress response to ultraviolet radiation
after 12 weeks of exposure.
Stress response to cnha iced UV—B was noted in all 6 Browall i a species
and cultivars after 4 ieeks of irradiation. Leaves of Browallia tValijaI
‘Ultra’ , Panama’ , ‘Blue Be] is Improved’ , and ‘Blue Tro] 1’ were sina 11 Cr,
twisted and distorted. The only sign of stress in Browallia viscosa
‘Sapphire’ was the foitnation of purplish pigment in normally young green
flower buds.
-------
7
I.iterntui e Cited
1. Brodfuhrer, U. 1955. Dc r Einfuss H nor nbgestufien Dusicrung von
ul Ira viol etter Sonnestrahi ung auf das Wachstuni der PH an en. P1 auta
45, 1.
2. Caldwell, H. 14. 197]. Solar ultraviolet radiation and the growth
and development of higher p1 ants. Phot:opliysi ol ogy 6: 131—] 77.
Academic Press, New York (A. C. Giese, ed.).
3. Cline, H. C., and F. 8. Salisbury. 1966. Effects of ultraviolet
rathation on the leaves of higher plants. Rad. Bot. 6:151—163.
4. Dubrov, A. P. 1968. The genetic and physiologic effects of the
action of ultraviolet radiati on on hi gher plants. Akad. N.iuk . SSSR,
Moscow, 250 p., (Rev.)(Tn Russian).
5. Klein, P. M., Edsall, P. C., and 11. C. Centi]c. 1965. Effects of
near ultraviolet and green radi ation on plant growth. P1 ant Physi ol
40:903—906.
6. Lockhart, J. A., and U. Brodfuhrer—Fraazgrutc. 1961. The eFfects of
ultraviolet radiation on plants. Encyclopedia of Plant Pliysiol.
16:532—554. Springer—Verlag, Berlin.
7. Skelly, J. H., George, H. F., Ileggestad, 11. E., and U. T. Krizck.
1978. Air pollution and radiation stresses. ASAE Monograph
Modification of the Aerial Environment of Plants.
8. Tliimijian, R. V. , Campbel I, L. E . , and Ii. R . Cairns. 1978. Radi ati on
sources and related environmental control for biological and climatic
effects UV research (BACER).
-------
Table 1 . Rd aLive sensitivity and Lyp of in jury ol celect e1 speci pq nod
ciii Li vars of florist and nur ’i y p l ants in rcspon e 10 enhanced tJV—B
irradiance.
Type IJV—B Type
1/ Sensi- 7 , of
Common Name and Cii tivar Cenus_and_Species Plant— t ivi ty Injury —
African Violet Sairilpaulia ionnntha
Wendl.
Blue Fairy Tale TP IS
Ballet ‘VP IS
AgeraLum Ageratum
housLonianurn Mi]]
Biscaya A IS
North Star A IS
Blue Blazer A IS
Atlantic A IS
Agleonema Aglaonema commutaturn TP IS
Sc hot 1
Altl inea flibiscus syriacus L. A IS
Alyssum Lohular a maritime
(Li Desv.
Carpet of Snow A IS
Amaranthus Ameranthus
tricolor L.
Molten Fire A IS
Red Leaf A IS
Aphelandro Aphclandra TP IS
squarrosa Nees
Aster Astcr alpiiius L.
Totem Pole L i S—I C
-------
‘ 1
Arborvitae
Globosa
Pens a
Pyramide
Rheindiana
Balsam Fir
double—nd x
Begonia Roulette
Browallia
Sapphire
Browall a
Blue Bells
Blue Troll
Ultra
V an j a
Pan am a
13 ox w 00d
Jap. Green Beauty
Mic. Green Velvet
Type
of
P1 a ii 1—
S
S
S
S
‘F
IS
IS
LS
IS
Is
Is
rs
Type
of
1 n pi rv—
Common Name and Cultivar Ccnu md Speciec
U V—B
2 /
t I VI t\’—
Thuja occidentalis L.
Abie balsamea CL.)
I’Ii l l
Basil Red Leaf Ocimum bnsilic.j,n L. TI’
Begonia, tuberous roc’ed Bei c ’nia x tuberhybrida TP
Vos s
TP
TP
A
IS
IS
5—1 PE
Begonia rex Putz.
Browalija viscoca 1-113K
Browallia speciosa
Hook.
Buxtic microplivi in
Siebold & Zucc.
Tl
S—i
D,
T
TP
S—I
I),
T
rP
S—I
D,
T
TP
S—I
D,
T
S
IS
S
-------
Caladium candidum
Calendula Fiesta
Camellia sasanqua
S h owa
Cainellia Sho a
Canna Ambassador
Chinese Chestnut
Christmas Pepper
Chrysanthemu n
Classic
Elegant
Fred Shoesmjth
Flaming Sun
Gold tone
Iceberg
Imp. md. White
Jackpot
Minn. AuLumn
Pan c l i o
Penguin
Caniellia japonica L.
Canna x generalLs
L. U. Bailcy
Castanea mollissin ia
B luine
Capsicum annuuni L.
Chrysant-hcirum mori—
foli.uni Rainat.
Type
of
Plant —
P
P
P
P
P
P
P
P
1’
P
P
IS
IS
Is
IS
Is
Is
Is
Is
IS
Is
Typo
of
In jury —
Common Unme dnd Cul tiv r Genus and Species
]0
11 V —13
Sensi— 2
t 1 vi1 y —
Caladjuni x hortulanum
Bird & cy
Ca]endula officin—
aI’s L.
Camejlia sasanqun
Thunb.
TP IS
A IS
S
S
S
IS
IS
Is
IS
T IS
A IS
P IS
-------
Type
of
P1 rim L—
Sen ,i— 21
I V t\—
II
fl I
Commoit Name rind Cu! Li v.ir
Genus and Spec 1 CS
i
Is
Shoesmith
Sunburst
P
IS
Superchief
P
IS
Tinkerbell
IS
Coleus
Coleus blurnei Benth.
Pink Rainbow
A
S—i
Red Rainbow
A
S —i
Aetna
A
S1
Campfire
A
S—i
Carefree Red
A
S—i
Freckles
A
S—i
Fredericcj
A
S—i
Clory of Luxcmburg
A
S—2
Harlequin
A
S—I
Marty
A
S—i
Pineapple
A
IS
Saber Jade
A
S—i.
Saber Golden
A
S— I
Saber Pink Diagon
A
S— i
Saber Velvet
A
S— I
Saber Jade Paste]
A
S—i
Col. Blue Spruce
Picczi pungens Engeim.
I
IS
Cone Flower
Rudbeckia hit-ta L.
A
Is
PT.,
PL,
PL,
PL,
PL,
PL,
PL,
PL,
PL,
PL,
D, T
D, r
D
D
D
D
D
D
D
D
PL, D
PL, D
1’L, D
Ph, 1)
PL, I)
-------
17
Type Type
1/ s — 71 of
Connuon Name and C i1 Livni Ceiius nd Spcci ‘c P1 ant— t ivi ty— in lurv —
Crape Myrtle Lngerstrocmia S iS
n ica L.
Dusty Miller Centaurca maritima A IS
Duf our
Exacum Affine Exdcum affirie
Baif. 1.
Richly Frag. A IS
Blythe Spirit A IS
Midget A TS
False Aralia l)izygotheca ciey antJs — T1 IS
sinia (Ilort. Veitch)
R. Tig. & Gui ll um.
False Cypress CharnaccyNiris pisi — S IS
fera (Siehold &
Zuec.) Endi.
‘ Cyanovirid is ’
Fatsia Fatsia japonica Ti? S—I C
(Thiinb.) Decne. &
Planch.
Geranium Pelargoniurn x hortorum
L. F l. Bailey
Carefree White A IS
Scarlet Sprinter A IS
Glory Bush Tibouchina urvil I eana TP
(DC.) Cogn.
Glory Lilly Cl on osa roLlic — TI ? IS
chi.ldiaiia O’Brien
Gloxinia Emperor Wilhelm Sinrungia speciosa TI? IS
(Lodd.) ilicin
lIens & Clii ckens Sempervi viim tec 10mm IS
-------
13
Type IJV—ll Type
of , Sen i 7 I ol 3/
Comm on Ndm n tid Cu I L vi r Cciiii s 1(1 pccic” 1 Inn L— t i V L y— lii r y
1lcl otrope Marine Vnlerinnn officinnlis P Is
hal ly hex crcnn ta Thunb.
Glory S IS
Golden Gem s IS
hlollyhock Alcea rosen L. A IS
Majorette A IS
Powderpuffs A S—J C
Itnpatiens ImpaLiens wai]ernna
Uook. f.
Aflame A IS
A]oha A is
Ftitura Coral A IS
Futura Pink A IS
Futura Red A IS
Futura White A IS
ILUC A IS
White Imp P s—I C
Juniper Juniperus chinensis L.
Glanca S IS
Gold S IS
Old Gold S IS
Pftjzcr S IS
Skyrocket S IS
-------
Cracker Jack
Dwarf French Dib.
Early Gigantic
First Lady
Cold Rush
HawaiL
Honeybee
Orange Sherbet
Petite Gold
Petite Yellow
Spun Gold
Norway Spruce
Pansy Cold smi th Ci Ont
Lantana montevidensis
(K. Spreng.) Br]q.
Delphinium exaltaturn
Ait.
Leucothon axillaris
(LainTYD. Don
Tagetes patula L.
TageLes erecLa L.
Tagetes patula L.
Tagetes erecla L.
Tagetcs erccti L.
Tagetcs erecta L.
Tagetes erccta L.
Tagetes patula L.
Tagetes paLula L.
Tagetes patula L.
Tngctes 1)atlIla L.
Tagetes patula L.
PLCCa a]bies CL.)
Kars t.
Viola x witirockiana
Gaiiu
Pcpcromia ohtusifolia
(L.) A. Dictr.
Type
of
P1 ant—
S IS
A
A
A
A
A
IS
IS
Is
Is
Is
IS
Ty [ )(“
of 3/
I fl 1 tI T \ ‘ —
Common Name and Cu 1 t i var Gen i nd Spcc i es
IJV—B
Seus
t vi tv
IS
Lan tana
Larkspur Dk. Blue
Supreme
Leucothoc
Marigold Bolero
A IS
Is
IS
IS
IS
IS
A
A
A
A
A
A
A
A
Peperomi a
Pen winkle
Is
IS
TI
Vi nca minor L. 1’
S—I
-------
Pelunja
All Star
Comrnanche
Pink Cascdde
Pink Magic
Plum Pink
Silver Magic
Sugar 1)addy
Sugar Plum
Pieris
Plantain Lily
Poinsettia
Annettc Flegg
C—I—Red
C—I—White
Prof. Laurie
Super Star
Supreme Annette llcgg
V—lO (Amy)
1hite Anriette llcgg
of ,
Common Name dnd Culti var Genus and Speci C5 P1 nfL—
Sensi— 7 ,
t ivi
Type
ol
In jury —
A
A
A
A
A
A
A
A
S
IS
Is
IS
IS
IS
JS
IS
IS
Petunia x 1iybrLda Vi mi.
Picris japonica
(Thunb.) D. Don cx
C. Don
flosta sieboidji
(Paxt.) J. Ingram
Euphorbia puicherrima
W]lld ex Klot sch
15
D, C
D, C
P E
P Is
TP
TP
TP
TP
TP
TP
TP
TP
IS
S— 2
S— 2
iS
IS
S—i
rS
IS
-------
l’ortulaca, yellow
St. John’s Fire
Schefflera compacta
Snapdragon
Biocolor 1ajus Tet
Potomac White
Rocket. Whi te
Rose I’ixie
Spider Flower
Stokes Aster
Taxus
Portulaca grandi flora A
hook.
Priiuula obconica
hlance
Type
of
P1 an t —
UV — B
Sensi— 7
ti V i t_y .
Coimnori Name and Cultivar
16
Genus and Spec es
Ty J)C
ol
In jury —
Primrose
Red Improved Rose
Red Scarlet Maple
Rex Begonia
Salp]glossis Emperor
Salvia
Rodeo
IS
Is
Is
IS
______ ______________ Is
_______ _____ p is
Rosa x hybrida S
Acer rubrum L. T
Bc onia x rex—cultorum TP
L. H. Bailey
Salpiglossis s]nuata A
Ruiz & Pay.
Salvia splendens ello
cx R. & S.
A
A
l3rassaia actinophvl]a iT
Enchl.
Antirrhinurn majus L.
A IS
A IS
A IS
A IS
______ ________ A IS
______ ____ p is
S IS
Is
IS
IS
Cleome speciosa R if.
Stokccja lncvis
U. HiIU Greene
Taxus cuspiclata
Siebold & Zucc.
Dens a
S
Is
-------
17
Ty I ° l’v p’
oF SenSi—.,/ of - v
Common Name and Cul I ivar Conus and Spec es Plant— Liv i ty . in ur_y—
Intermedia S IS
Media J3rownii S 1S
Media iKelse)’ S IS
forenia TorenLa fournieri A IS
Linden cx E. Fowin.
Verbena Springtime Verbena x hybr da A IS
Vos s
Veronica Ilebe buxifojia P IS
(Bcnth. ) Cockaync &
Al] an
Vinca Trailing Varicgata Vinca major L. P IS
Zinnia Zinn]a clc’gans Jacq.
Carved Ivory A IS
Goddess A IS
tsabe]ina A IS
Peter Pan Pink A IS
White Dogwood Cornus florida L. T IS
1/
A = annual; P = peremial; TP = tender perennial; S = shrub; T tree.
2/
As indcated by plant expression: S = sensitive to either 100% (s—i) or 50%
(S—2) increase in biologically eftective UV; IS = insensitive.
3/
Visual injury to leaf: C = chlorosis; D distortion; G glazing,
PL pigment loss; FE = p1 gmeiit enhancement; T = twisted.
-------
18
fable 2. Relative seiisiL viLy of 7 1 i ‘j cie i ol Ilorist. arid riui ciy plnTit ii
response to cn1i jnced UV—13 irrndiancc.
Atinu a Is
____________________ Asters — S — I
Browallia — S—i
Coleus — S—i, 2
Ilollyhock — S—i
1/
S—i = Sensi tive to 100% or greater increase in biologically effective UV.
S-2 = Sensitive to 50% or greater increace in biologically effective UV.
Number Irradiated
Sensitive P1ant
Perenni ru
Shrubs
Trees
29. i
12
6
T .mpali ens —
S—i
None
None
Poinsettia —
S—i,
2
Fatsia — S—i
Vjnca — S—I
-------
SZIcUq FL9
FINAL REPORT
BiOLOGICAL EFF’LCT OF ULTRAVIOLET RADIATION ON CATTLE:
BOVINE OC uLAR SQUAMOUS CELL CARCINOMA
K. E. Kopecky
C. W. Pugh, Jr.
D. E, Hughes
Na tional Animal Disoa’.e Can i cr
Agricultural Research
Science and Education Adniinisriatjon
U S. Department of Agriculture
Ames, Iowa 50010
EPA-IAG. -D6—O168
Project Officer:
R. J, McCr acken
Agri.cultura) Research, Sconce and Education Administration
U.S. Department of Agriculture
Washington, D.C. 20250
Prepared for
Environmental Protection Agency
BACER Program
Washington, D.C. 20460
-------
S IJMMARY
This report summarizes the work done on the bovine ocular squamous
cell carcinoma—ultraviolet radiation project from late February 1977 to
Decenber 31, 1977. It includes a description of the experimental set up
and procedure used to irradiate cattle. Also included is a preliminary
report of a UV—induced pre—neoplastic ocular growth.
A second part of this report summarizes an epidamiological study
that shows that since 1950 the incidence of bovine ocular squamous cell
carcinoma seen at slaughter has increased. This increase was real and
not due to an increase in numbers of cattle.
-------
2
The bovine ocular squamous cell tarcinoma (cancer eye) study was
begun in an effort to develop an animal model for the study of the
biological effects of increased UV—B radiation that might follow modifications
of the atmosphere.
The relationship between increased fly—B radiation and cancer eye
has been assumed for many years. However, only in recent years has
enough epizootiological data has been collected to give some credence to
this hypothesis. 1 These data have been reviewed’ (Table 1) and s m marized. 2
The geographic location of the high—incidence area of cancer eye
involves two factors: time of exposure and intensity to solar radiation.
At lower latitudes the period of exposure to sunlight is longer; the
solar angle is greater, hence the solar radiation has less atmosphere to
penetrate. At higher altitudes the decreased density of the atmosphere
results in a greater intensity of solar radiation.
Lesions of cancer eye most frequently develop in unpigmented areas
of the ocular region. Apparently pigment has a significant inhibitory
effect against the development of lesion.
Cancer eye has been reported in almost every breed of cattle, but
it is found much more frequently in the Hereford breed than in others. 3
Within the Hereford breed, increased susceptibility to cancer eye has
been associated with certain strains of cattle which lack any pigment
around the eye, the scl.era and conjunctiva. Because the presence or
absence of this pigment is a genetic trait in the Hereford breed, we can
say that certain 1-lerefords are hereditarily susceptible to cancer eye.
The rate of occurrence of cancer eye in Herefords is based on
actual observation and data co] ection, but th relationship of sunlight
-------
3
or UV—B radiation to cancer eye is circumstantial. Therefore, it was
decided to obtain more evidence as to the relationship between UV—B and
cancer eye by actual irradiation studies in cattle.
In addition to the UV—B study, we stud cd existing data to determine
the incidence of cancer eye in the U.S. cattle population.
Materials and Methods
Cattle——Six Hereford cattle that lacked pi8ment around the eyes
were used. They were obtained from a purebred Hereford cattle herda
that had a history of cancer eye. The cattle were housed in 2 groups of
3 in an isolation barn that was air conditioned and temperature controlled.
in each group of 3, two animals received UV—B radiation, and one animal
served as the control. The controls were stanchioned in such a way that
they received no l iv radiation. The method of exposing the cattle to UV
irradiation is shown in Photo. 1. 0t e group of cattle was in a barn
w lth 3 windows and a north exposure; the other group as housed similarly
but had a south exposure. In addition to natural light, each barn was
lighted by four 150—watt overhead bulbs. When being irradiated, the
cattle faced away from the windows. The eyes of the cattle were
observed daily.
Ultraviolet (UV—B) Irradiation and Source——The UV lights were
Westinghouse FS—40 sunlamps mounted in a specular aluminum reflector
fixture (Photo. 1). Each fixture was covered by a sheet of 5 mil cellulose
acetate (CA)b which served to filter Out UV of wavelengths below 280 nm.
The CA sheets were pretreated for 6 hours with UV irradiation by FS—110
sunlamps, then uscd to cover tF’e lamps hich irradiated the cattle.
-------
4
The CA sheets were replaced at intervals < 30 ir r sheet. The
fixtures were hung about 102 cm from the plare c t cattle’s eye
(Photo. 1).
Initially a light fixture using 3 FS—40 bulbs was set up in a
single fixture covered by CA. This was hung 61 cm from the eyes of the
cattle. The change to the above described 4 lamp fixture was made early
in the experiment to comply with the standardization that was desired in
the Biological and Climatic Effect Research (BACER) program. At the
time of the change, the time of exposure was increased 3x. This increased
time of exposure was based on the relative dosage as determined by a IL
600 p1 otometer.C The time change of irradiation was due to a change in
distance and configuration of the lamp.
The cattle were exposed to the UV—B radiation daily (Table 3). The
time varied from 7 minutes at the start of the experiment (February 28,
1977) to 150 minutes on December 31, 1977. The time of exposure was
increased at irregular intervals but in such a way that the cornea was
not directly injured by the UV—B radiation.
Measurement of Ultraviolet Radiation——The intensity of the TJV
radiation was measured in milliwatts/sq. meter/nanometer by a single—
monochromator spectroradiometerd that was interfaced with a desk—top
Hewlett—Packard 9815A programmable calculator. The calculator controls
the operation of the spectroradiometer so that scanning is initiated on
command and read]ngs are recorded for each nanometer interval. The
calculator prints out the wavelength and irradiance for each wavelength
interval. The program was written by and is available from the
Instria-ientation Research Laboratory, USDA, Beltsville, MD.
-------
5
Bacteriologic dnd Virologic Examination——The eyes were cultured
periodically for Moraxella bovis and viruses; techniques previously
described were used. 4 The main virus considered was cytopathic herpes
virus because one group of investigators suggested that it might be
Involved In cancer eye of cattle.e Secretions from each eye of the 6
calves were collected with two sterilized cOttOfl—ti )ped applicators 4
times before UV irradiation treatment began and once a month for 12
months. The mares were also cultured 4 times before irradiation began
but were not cultured later. One applicator was used for the collection
of eye secretions for bacteriologic examination and w s placed in a
sterilized dry screw—capped tube. The other applicator was used for
virologic examination. Later, the secretions for bacteriologic examination
were streaked on the surface of 5% blood agar plates and the plates were
Incubated at 37 C for 24 hours and at 25 C for 24 additional hours.
Nasal secretions were handled in the same way as were the eye secretions.
After Incubation, the surface was observed for colonies of H. bovis .
The applicators with secretions for virologic examination were
placed in Earle’s balanced salt solution containing 0.25% lactalbumin
hydrolysate, antibiotics (dihyd ostreptomycin, 0.1 mglml; kanamycln, 0.1
mg/mi; and penicillIn, 100 units/ml) and 1O O% (by volume) calf serum
immediately after the sample was collected. Later, 0.1 ml of the suspension
was used to inoculate each of 2 tubes of secondary embryonic bovine
kidney (SEBK) cells. The cells were incubated at 3? C and observed each
day for cytopathic effects (CPE) for 7 days.
Hematologic Examination——Venous blood was taken from the calves on
their arrival at the .ation i1. ,nimal Disc ice Crnter (NADC) and 9 times
-------
6
during the study (Table 4). Blood was collec cd by venipuncture with
evacuated glass tubes containing EDTA for w’loie blood samples (used in
determining total white b]ood cells (WBC) and differential Wl C counts)
and evacuated glass tubes free of any anticoagulant for serum collection.
Total WBC counts were made with a Coulter counter. 5 The WBC differential
counts were made after the cells were fixed and stained by a modification
of the method 5 of May—Grundwald—Ciemsa staining.
Incidence ot Cancer Eye (Epithelioma) Determination——The data on
cancer eye incidence and slaughter data was provided by Meat and Poultry
Inspection (now Food Safet r and Quality Service) of the USDA. These data
were tabulated and then analyzed by 3RSSH method of Tukey. 6 For this
report any squamous cell carcinoma involving the ocular region is known
as an “epithelioma”; the two terms will be used interchangeably with
“cancer eye.”
Results
Bacteriologic Exam nation——Moraxe1la bovis was recovered from
either the eyes or nares or both of 3 of 6 calves after the first examination.
The infected calves were given oxytetracycline intravenously (I.V.) at a
dosage of 7 mg/kg of body weight. When cultured 1 week later, one eye
of each of 2 of the 3 previously infected calves was still infected;
therefore, the calves were given oxytetracycline I.V. at a dosage of 11
mg/kg of body weight. Moraxella bovis was not cultured from the eyes or
nares after the second treatment and the calves were presumed to be free
of H. bovis .
-------
7
V rologic Examination——Cytopathic changes were not observed in
tubes of SEBK cells during the study and the eyes were presumed to be
free of infection by cytopathic herpes virus.
Hematologic Data——Presented in Table 4.
Cross Anatonical Changes——Au mucosa surfaces of the eye were
highly inflamed. The sciera and conjunctiva of the cattle developed
various degrees of plaque formation and papilloma formation that appeared
after 16 weeks of radiation. The type lesions observed were considered
typical of early cancer eye (Photo. 2).
Histopathological Changes——A biopsy taken from one animal on October
31, 1977 (after 36 weeks of radiation) revealed metaplastic changes
consistent with preneoplasia including diffuse epilhelial hyperplasia
with acanthosis and presence of foci of clusters of typical squamous
cells in adjacent dermis.
The dermis was markedly edematous, and it vasculature and collagen
fibers were distorted and irrcgular. Many aggregates of small lymphocytes
were distributed throughout the dermis, and small foci of neutrophils
were present in many areas Accumulations of eosinophilic proteinaceous
material, representing intraepithelial edema (blister formation), were
present between the stratum germinativum and stratum spir’osurn in two
areas.
Squamous cells in dermal clusters and its overlying epitheliurn
appeared atypical and moderately anaplastic, e.g., they were large, and
pleomorphic and had large, irregular nuclei with large nucleoli. The
dermis surrounding these squamous cell clusters (or epithelial pearls)
was edematous and contained fibrin and neutro’ iisJ (Photo. 3 and 4.)
-------
8
Incidence of Cancer Eye (Epitheliotnas)——The preliminary results of
the survey are shown (Table 5). There has been a 5x increase in total
epitheliomas recorded at the slaughter plants. When the total epitheliomas
were adjusted to consider the increase in number of animals slaughtered,
the incidence of epitheliomas increased about 2x (Table 5).
The incidences of epitheliomas were compared for 3 years with 3
other causes for condemnation (Table 6). When only animals that were
condemned on the “kill floor” at the abattoir were considered, condemnation
due to ephtheliomas increased greater than those due to other causes.
After the number condemned was adjusted for the increase in total
slaughter from 1955 to 1975, only condemnations due to epitFieliomas
increased; condemnations due to other causes decreased.
Discussion
The experimental results in the present study represent the first
successful attempt at producing neoplastic changes in a short period by
UV—B irradiation and normal animals exclusively. Previous work 7 with
UV—B irradiation involved the use of either genetic variants or various
carcinogenic chemicals or both. Various forms of skin cancer were
produced in these animals, but it was difficult to know whether the UV—B
or the chemical produced the lesion.
Most work in which UV—B irradiation was used involved the use of
mercury lamps that have a very strong emission at 254 nm and a damaging
effect on DNA. In this present study this strong wavelength emission
was eliminated by the use of 5 nil of CA uh ch adLtr bed wavelcngLhs
below 280 on. This situation is thought to be much closer to the natural
-------
9
IJV—B which penetrates the atmosphere and reaches the surface of the
earth. This filter also climinntes the strong UV band at 254 mm which
in itself is very damaging to cells because of effect on DNA. Further
evidence that the UV—B was the sole inducer of the lesions in the present
study is the fact that cytopathic herpes ‘viral agents and M. bovis were
not isolated.
Although epithalioma at the abattoir seemed to increase, more
information is needed because this apparent increase might be due 10 an
increased awareness by the meat inspectors, resulting in a spurious
increase in cancer eye incidence. It might be due to a combination of
both an actual increase and increased awareness by the inspectors. We
favor the hypothesis that there is a real increase because our investi-
gation indicated that epitheliomas were the only cause for condemnation
that actually increased Out of the several diseases studied. We think
that the increased awareness was responsible for the large increase in
total epitheliomas detected at slaughter but that the increase in the
incidence of condemned animals was due to a real increase in severe
cases of epithelioma. Further evidence supporting this hypothesis is
the fact that when we were looking for a cause of increased condemnations,
we related the incidence of epitheliomas with solar activity. 8 We
plotted with a 5 year latent period (from time of first exposure to time
of clinical signs) epitheliomas against solar activity; they matched
quite well (Fig. 2). Because cancer eye and solar radiation appeared to
be interrelated, this finding should not be so surprising. A delay in
appearance would correspond to the time from first exposure to develo ent
of clinical signs.
-------
10
This study to date has showu that UV—B ir: i Lo ,s a probable
cause of cancer eye and that increased UV—3 ra.i:ati i c uLd be expected
to increase both the total cases of cancer eye in Lsceptible animals
as well as an increase in the severity of this disease.
-------
Footnotes
aMeat Animal Research Center, USDA, Clay Center, Nebraska.
b eianese Plastics, 26 Main Str., Chatham, New Jersey.
°International Light, Inc., New Buryport, ‘iassachusctts.
dModel 1741, Optronic Lab., Inc., 7676 Fenton Street, Silver Spring,
Maryland.
eAnSOfl M: Bovine ocular squamous cell carcinoma: In vitro investi-
gation of a viral etiology. Abstracts No. 83, page 15, 57th Ann. Mtg.
of Research Workers in Animal Diseases. Chicago, Illinois, Nov. 29—30,
1976.
acutainer, Becton—Dickinson, Div. of Bectan, DickinL3n and
Company, Rutherford, New Jersey.
8 Co dter Counter, Model ZB 1 , Coulter Electronic, Inc ., Hialiah,
Florida.
hDescription and interpretation by N. F. Cheville, Pathological
Laboratory, NADC, Ames, Iowa.
1 Statistical operations performed by G. D. Booth, NADC, Ames, Iowa.
-------
Re fe ten c e s
1. Anderson DE: Cancer eye in cattle. Mod Vet Pract , 51:43—47, 1970.
2. Kopecky KE: Ozone depletion: Implication to the veterinarian.
J M i Vet Med A ’soc (in press), 1978.
3. Monlux AW, Anderson WA, Davis CL: The diagnosis of squamous cell
carcinoma of the eye (cancer eye) in cattle. AmJVetRes 18:5—34, 1957.
4. Pugh OW, Jr, Hughes DE, Packer PA: Bovine infectious keratocon—
junctivitis: Interactions of Moraxella bovi and infectious bovine rhino—
tracheitis virus. AmJ Vet Res 31:653—662, April 1970.
5. Winters H: Comparison of hematological stains in sheep and man.
Aust VetJ 41:14—16, January 1965.
6. Tukey JW: Exploratory Data Analysis. Addison Wesley Pubi. Co.,
Reading, Ma, 688 pp, 1977.
7. Suskind RR: Ultraviolet radiation carcinogenesis. In Sunlight and
Man. Thomas B. Fitzpatrick, consult. ed., Univ. of Tokyo Press, 285—298, 1974.
8. Special Committee for Solar Terrestrial Physics, National
Academy of Science: Solar—Terrestrial Physics and Meteorology: A
working document. Washington, DC, 1975.
-------
TABLE 1—Epidemiological Evidence for Relationship of Cancer
Eye and Sunlight (Ajiderson, 1970)
Level
of
No.
of
Age adjusted
frequency of
Critecion sunlight
animals
cancer eye
(U
Latitude Low 3445 3.6
Med 361 7.8
High 1154 9.2
Altitude Low 670 4.8
Med 3909 5.1
High 381 9.7
Hours of Low 3445 3.6
sunlight Med 823 5.6
High 692 11.7
-------
TABLE 2—Irradiance from Four FS—40 Lamps at 100 cm With arid
Without Cellu]ose Acetate Filters per Nanometer
Irradiance
Filter
Condition
filter
of
280—320
milliwatts/sq.
meter
None
—
11.3346
5 mu CA
New
6.5262
5 mu CA
Used 5 hr
5.4734
5 mu CA
Used 25 hr
4.3178
-------
TABLE 3—Exposure Time to UV—B Irradiation
Length
of
exposure
Number
of
days
Total
time
(minutes)
7
2
14
10
5
50
A* 15
2
30
20
26
520
25
18
450
30
28
840
1940
90
56
5040
105
42
4410
B** 120
42
5040
135
70
9450
150
15
2250
*
A 3 FS—40 bulbs in single fixture, 60 cm from plane of eyes;
Irradiance was not measured by spectroradiometer but was approxi-
mately 3x greater than B when measured by IR—600.
**
B 2 FS—40 bulbs, 2 fixtures, 60 cm apart, 100 cm from plane of
eyes; Irradiance 4.8956 milliwatts/sq. meter/nm.
-------
‘1 LE 4—Total White Blood Cell (NBC) Counts and Selected (Eosinophils, Neutrophils, Lymphocytes,
aid Monocytes) Differential WBC Counts of UV—B Irradiated and Nonirradiated Calves During Nine 12
Months Study*
2—28
3—23
5—5
6—15
7—13
8—] 8
9—28
11—16
12—2 1
2—8
3—23
5—5
6—15
7—13
8—18
9—28
11—16
12—2 1
2—8
3—23
5—5
6—15
7—13
8—18
9—28
11—16
12—2 1
2—28
3—23
5—5
6—15
7—13
8—18
9—28
11—16
12—21
2—8
3—23
5—5
6—15
7—13
8—18
9—28
11—16
12—2 1
2—8
3—23
5—5
6—15
7—13
8—18
9—28
11—16
12—2 1
4 21 68 3 8754
2 11 84 3 10600
4 14 79 3 12400
3 12 83 2 11000
4 22 68 6 11200
5 22 70 3 12700
6 17 75 2 13200
5 21 70 4 10800
2 15 80 3 9614
4 27 66 4 6234
3 16 82 2 6789
7 28 60 5 9467
4 22 70 3 7939
5 12 75 6 8435
5 12 79 5 8607
6 18 73 3 9422
3 16 76 5 6683
6 13 79 2 7818
9 17 71 3 12200
5 17 75 4 7499
8 18 71 4 9491
8 16 74 2 7663
6 29 59 6 9184
8 13 73 6 8965
4 12 79 5 9537
5 22 61 6 9795
12 9 77 2 11100
C,ilf No. Date Eosin Neut Lymph Mono Total WBC Calf No. Date Eosin Neut Lymph Mono Total WBC
**
2042
2369
2623
2698
2709
2713
1 13 85 1 8542
5 11 81 2 10400
1 13 84 2 10300
2 15 81 2 11600
4 18 72 7 11100
3 9 64 4 10600
3 12 83 2 11000
2 7 87 4 10300
3 4 91 2 6909
2 34 66 5 5999
2 22 73 4 6968
1 13 84 2 6497
4 10 85 1 6742
4 9 85 4 6629
4 20 71 5 7980
9 14 70 7 8038
4 13 79 4 8297
1 13 80 6 6053
3 26 64 6 8303
5 16 77 2 9991
4 20 74 2 10800
4 28 67 1 11700
7 16 71 6 11300
3 17 77 3 13800
4 13 78 5 12400
5 17 76 2 11000
3 13 82 2 9490
*
l3asophils and myelocytes Counts were determined to be less than 1% and therefore are not recorded.
* ;
Noriirradiated control.
-------
TABLE 5—Cattle Slaughtered .iith Epitheliornas
Epithel ioma
‘total
slaughtered
Year x i0 6
Total
epithel iona
3
as 1/100 of
percentage
total
si aught e red
1950 13.1 24.5 19
1951 12.6 24.7 20
1952 12.1 28.0 23
1953 15.2 30.0 20
1954 18.5 36.0 20
1955 18.7 35.4 19
1956 19.7 38.1 19
1957 20.1 46.9 23
1958 18.6 47.4 26
1959 17.3 39.7 23
1960 18.5 59.1 32
1961 19.9 69.6 35
1962 20.2 65.1 32
1963 20.9 75.0 36
1964 23.2 79.8 34
1965 25.8 85.8 33
1966 27.4 85.2 31
1967 27.9 77.3 28
1968 28.1 87.7 31
1969 30.2 109.9 36
1970 30.9 136.0 44
1971 31.0 102.7 33
1972 31.7 109.0 34
1973 31.6 123.8 39
1974 30.9 147.3 48
1975 34.8 141.7 41
1976 36.8 131.1 36
-------
Causes of
Condemnation 1955 1965 1975
T
12
,407(100*)
61,692(497)
152,063(1226)
**
Adj.
45,031(362)
81,754(659)
C
12
,407(100)
9,654(78)
12,134(98)
Adj.
7,047(57)
6,524(53)
Abscesses
Lyrnphona
T
Adj.
C
Adj.
T& C
Adj.
29,158(100)
9,675(100)
2,680(100)
415, 709(1425)
303,437(1040)
9,154(95)
6, 682 (69)
4,616(172)
3,369(126)
508,475(1744)
2 73,373(934)
11,445(118)
6, 153 (63)
4,160(158)
2,237(83)
Epitheliotna
T
Adj.
C
Ad j.
34,394(100)
4,077(100)
85, 84 1(243)
62,658(177)
7, 139(175)
5,211(128)
141,727(400)
76, 197 (215)
21,491(527)
11,554(283)
Total
Slaughter
18,728,579
(100)
25,803,948
(137)
34,825,463
(186)
T = Total cases condemned + retained.
C = Total condemned.
*
Percent (7.) of 19)) value.
TABLE 6——Comparison of 3 Condemnation Causes and Epithelioma for 3 Different
Years
Pneumonia
**
Adjusted to rate of increase from 1955 to 1975 for total slaughter.
-------
Photographs
Photo. 1——Arrangement of cattle and UV lamps.
Photo. 2——Early gross changes on 3rd eyelid and corneoscieral
junction after UV—B irradiation.
Photo. 3——Tissue section through 3rd eyelid. 25 X.
Photo. 4——Same as Photo. 3, but 100 X.
-------
d
LU
26
24
22
_.1
LL
44
40
EPITHELIOMA AS i % TOTAL SLAUGHTERED (3 RSSH)
32
30
20
95O
54
58
ARS
66 70 1974
-------
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FINAL REPORT
RADIATiON SOURCES AND RELATED ENVIRONMENTAL CONTROL FOR
BIOLOGICAL AND CLIMATIC EFFECTS UV RESEARCI-i (BACER)
Richard W. Thimijan
Harry R. Cams
Lowell B. Campbell
Agricultura] Equipment Laboratory
Plant Physiology institute
Beltsville Agricultural Research Center
Beltsville, Maryland 20705
EPA—IAG—D6--0168
Project Officer;
R. J. McCracken
Agricultural Research, Science and Education Administration
U.S. Department of Agriculture
Washington, D.C. 20250
Prepared for
Environmental Protection Agency
BACER Program
Washington, D.C. 20460
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RADIATION SOURCES AND RELATED ENVIRONMENTAL CONTROL FOR
BIOLOGICAL AND CLIMATIC EFFECTS UV RESEAP.CH (BACER)
CONTENTS
I INTRODUCTION
II SELECTION OF AVERAGE SUNLIGHT
III ARTIFICIAL UV-B RADIATION SOURCES
IV ERROR SOURCES
V WEIGHTING FUNCTIONS
VI BIOLOGICAL EVALUATION OF WEIGHTING FUNCTIONS
VII SUMMARY AND CONCLUSIONS
Vii i REFERENCES
IX TABLES
X FIGURES
XI APPENDIX I — PREDICTION OF IRRADIANCE
XII APPENDIX II — CONVERSION OF WEIGHTED UNITS, KNOWN RELATIVE
SPECTRAL POWER DISTRIBUTION
XIII APPENDIX III - CONVERSION IRLHETER RADIOMETER AND OPTRONICS
RADIOMETER READINGS TO ABSOLUTE mW.m 2 .nm SUMMED FOR 5 nm
BANDWIDTH FROM 280-320 rim
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BACER-AUG-78
SEA—PPIII—AEL
RADIATION SOURCES AND RELATED ENVIRONMENTAL CONTROL FOR
BIOLOGICAL AND CLIMATIC EFFECTS UV RESEARCH (BAcER)
This report describes sources arid filters for ultraviolet (UV)
radiation. Weighting functions were developed, evaluated and used to
describe practical sources and filters for UV relative to solar radiation
at the earth’s surface, sunlight.
Any stratosphere ozone reduction is expected to enhance the UV
radiation at the earth’s surface. Determination of the effects of such
enhanced UV on plant life and vegetation requires UV sources to
supplement sunlight or simulate sunlight with an enhanced UV component.
We divide solar radiation into several wavelength (A) regions:
Ultraviolet (UV ) 10—380 rim
UV—C 220—?8O rim
UV—B 280—320 mn
UV—A 320-380 mu
Visible 380—780 rim
Infrared 78O—lO rim
The radiant flux or power can be described by energy or number of
photons. However, for sources other than monochromatic (single
frequency), energy units are used, because they require no correction for
variation in absolute energy at different wavelengths, and because most
primary and secondary standards are calibrated in watts per wavelength
interval. In either system, source and filter descriptions must include
both total radiant power and spectral radiant power distribution.
Trade names are used to identify products and do not imply the product
used was better or worse than another similar product not mentioned.
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—2—
No artificial sources exactly simulate sunlight. For UV enhancement
simulation, sources must be selected that provide biological organisms’
basic light requirements as well as specific UV wavelengths.
Much existing literature on sources and plant response contains
uncertainties of absolute irradiance, source stability, spectral power
distribution, variation of plant response, spectral response of plants,
and other environmental conditions. Some of the difficulties arise from
uncertainties of UV radiation standards which are +5 percent. Inadequate
and misleading results have been reported due to th use of measuring
instruments with little or no attention to calibration.
Investigations were conducted by the Agricultural Equipment Labora-
tory in close cooperation with the Instrumentation Research Laboratory
(IRL) which had responsibility for developing portable equipment to
measure quantity and spectral power distribution of UV radiation.
The work was also cooperative with Florist and Nursery Crops
Laboratory, Plant Stress Laboratory, and Chemical and Biophysical Control
Laboratory for whom radiation sources were developed to provide given
levels of UV—B irradiance.
Simulation Problems in UI /—B Radiation
To simulate enhanced jill—B radiation, it is necessary to:
(1) Establish an ordinary or normal level of UI/—B radiation (280-
320 nm) upon which enhanced radiation can be imposed; and,
(2) Determine the biological response of plants to UV wavelengths.
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—3—
II SELECTION OF AVERAGE SUNLIGHT
Sunlight outside the earth’s atmosphere has been measured (Fig. 1,
Thekaekara, 1974). The earth’s atmosphere changes the direction,
quantity and spectral power distribution of sunlight before it reaches
the earth’s surface. Changes occur in the atmosphere with cycle lengths
of about 11 years (Abbott, 1963), 1 year, and 1 day. Much random
variation is superimposed on the-cyclical variation. All these changes
in the atmosphere affect the IN that reaches the earth’s surface.
Ozone is believed to be the principle absorber of ultraviolet at
280—320 nm, UV—B. Little radiant power of shorter wavelength, such as
260 tim, reaches the earth’s surface because it is absorbed by the earth’s
atmosphere. However, plants and animals respond to shorter wavelength
radiant power. The 280—320 tim definition of UV—B is a physical
definition and does not describe biological responses. There is much
evidence shorter wave length UV—B (280 t im) is biologically more active
than longer wavelength UV—1 (320 nm).
If ozone is reduced by 0.1 of its present quantity, the biologically
effective ultraviolet is expected to increase by 0.2 of its present
quantity. The exact ratio between reduction in ozone and expected
increase in biologically effective IJV depends on the present spectral
power distribution of ultraviolet &unlight and the spectral biological
effectiveness of ultraviolet; neither is well known. The absolute
quantities are also uncertain or unknown.
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—4—
The Radiation Biology Laboratory of the Smithsonian Institution
measured ultraviolet in 5—nm bands at Rockville, Maryland, 390 north
latitude. The daily total energy per am received on 49 days in June (+)
and July (x) 1976 when the equipment was working well are plotted in Fig.
2—6 for bands at 295, 300, 305, 315, 320 am. The daily totals were
ranked and plotted on the linear scale of extreme value probability paper.
The probability that daily total energy per nm in a particular
wavelength band would be as large as that read on the y—axis can be read
on the x—axis where lines parallel to the axes intersect the straight
line describing the data. An irradiance much in excess of the largest
value observed is very unlikely, according to this model, and there is a
real possibility that no irradiance in a particular band will be received
during a whole day. The data are skewed; the mode is larger than the
mean.
While days are not alike, we represented the data in each band by
its average. We then divided the average daily energy by the number of
seconds in 6 hours to obtain the average daily power; however, irradiance
would be higher or lower than average for most individual days.
Data that were missing or appeared to be unreliable (A = 310 am;
290 A > 325 am) were replaced. Published data and models were used
to replace data at short wavelengths. Data collected by IRL at Beltsville
served as a rough check on all data and supplied data at long wavelengths.
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—5—
Selected and processed data for the average U I? solar spectral
irradiance, mW.m 2 .nm 1 for a 6—hour day, June and July 1976, is
presented in Table 1, Fig. 7. The data resembles the calculated spectrum
presented by Caldwell (1977) who assumed 3.2 un of ozone at standard
temperature and pressure and a solar angle of 300 from zenith.
Nearly all natural 1JV—B is diffused before it reaches the earth’s
surface at sea level. Plants therefore receive natural UV—B from a
substantial portion of the sky, not only directly from the sun.
II I MITIFICL&L UV—B RADIATION SOURCES
Commercial ultraviolet sources are:
(1) Mercury Nigh—Pressure Lamps
(2) Xenon arc Lamps
(3) Mercury short arc Lamps
(4) Electric discharge lamps (low—pressure), fluorescent and
bactericidal
Of the commercial lamps, only xenon and fluorescent lamps have a
continuous spectrum in the UV—B region. The emission of the other lamps
is mainly mercury lines and is unsuited for tN—B simulation. Xenon arc
lamps have an emission spectrum somewhat similar to that of sunlight but
with excessive 1W radiation. Their efficiency is relatively low compared
to other discharge sources,-and cost, including power supplies, is
several times the cost for other light sources of similar power.
Filtering is difficult due to emission of radiant heat that amounts to
more than 70 percent of input power.
Prefabricated xenon solar simulators, including lamps, filtering,
cooling and required power supply are available with an area of coverage
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—6—
of O.01m 2 . For m 2 (one square meter) area coverage with solar
simulators, cost has been estimated at one million dollars.
Fluorescent UV lamps are suitable for horticultural use. The lamps
used in the BACER program are listed on page 8. They are linear sources
with low heat radiation. The spectral radiant power distribution does
not simulate sunlight but is primarily in the UV—B region. ‘Then filters
are used in combination with fluorescent UV lamps to supplement other
sources, the resulting emission can be used to simulate enhanced UV—B.
Fluorescent lamps. emit power equally along the length of the lamp
(x axis), but as the distance from the lamp parallel to the irradiated
surface increases (z axis), irradiance along the x axis and perpendicular
to the lamp (y axis) decreases rapidly from the center of the lamp
outwards.
Normalized irradiance levels, E, at varying distances, z, from a
bare.flourescent lamp may be computed with the following equation:
—l
(-1) E = cosO (4 it x M)
lTan’d 1 fM + d 1 M/(d + M 2 ) + Tan 1 d 2 /M + d 2 M/(d + 142))
2 2 2
cosO = z/M, H z + y , d 1 = x + x, d 2 x - x
Half the effective length is x and is 0.555 m for a 40—watt (F40)
lamp. Methods and conventions of the 1ES Lighting Handbook, (1972) for a
toroidal distribution from a line source were followed (Ref. 7).
The irradiance is also multiplied by 2 + 3(0.2 + 81 L5Ř)
cos 4 O for our aluminum reflector fixture containing 2 lamps (Fig. 8).
The normalized -irradiance is the multiple of total power output in any
system of units, from one lamp, per square meter.
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—7—
A computer program ’ was developed for ca]culation of normalized
irradianCe with x, y and z as variables for single lamps, for combinations
of lamps in parallel or positioned end to end. The computed normalized
irradiance at any x, y, z position (Fig. 9) multiplied by the total output
from one lamp and filter (Tables 2) 5) permits calculation of irradiance or
weighted irradiance for any position beneath a lamp or combinations of
lamps (Appendix 1). The equations also were extended, with more
complexity, to irradiance received from tilted lamps (not parallel to the
irradiated plane).
We calculated these normalized irradiances for the various lamp arrays
used at Beltsville (for example, Fig. 10, 11) and compared the calculated
values to measured values obtained from the IRLSpec—S single monochromator
spectroradiometer and the UV—B IRLMeter radiometer (Fig. 22). Linear
regression analysis confirmed the linear relationship between calculated
and measured irradiances (r 2 = 0.983). As distance increased along the x
and y axes, calculated values tended to exceed measured values. The cause
of the variation is not known. The calculated arrays were used in
selecting lamp configurations for our experiments. Calculated values
permitted us to take advantage of the unequal irradiance by reducing the
1/ Computer programs for varying lamp arrays were developed by J.H. Clark,
Hydrologic Data Laboratory. They may be obtained from Plant Physiology
Institute, BARC—West, Building 001, Beltsville, Maryland, 20705.
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—8—
r uobCr of opectroradiometer readings required. As a result, we were able
to obtain plant and animal response data in a single experiment over a
wide range of irradiance. These data were then subjected to regression
analyses to determine biological response.
Irradiance (E) under an array is closely correlated with both the
angle of incidence and the size of the solid angle the radiant power is
received from. Maximum irradiance under an array with tolerable shading
of sunlight in a greenhouse is roughly the equivalent total output of one
40-watt (F40) lamp per square meter.
Westinghouse FS40 (Pig. 12, 13), FS2O, FBZS4O—CLG (Fig. 14, 15),
FBZS2O—CLG and Rayonet F8T5 RPR 3000A 0 (Fig. 16, 17) fluorescent lamps
have been used recently in the BACER program. A portion of the spectral
power distribution of one representative lamp is given in Table 2 and in
Fig. 12—17. Fig. l6wasdrawn toa different scale to suggest the output
of a 40—watt (F40) lamp. The scale is labeled for an 8—watt (P8) lamp.
UV—B fluorescent lamp sources use one of two available phosphors.
Different lamp envelopes can be desctibed as various optical thicknesses
of the same kind of glass.
Little more irradiance (E) than the total output of one 40—watt
(F40) lamp per square meter can be obtained with an array of fluorescent
lamps that also permits a reasonable amount of light for photosynthesis
to reach plants. Therefore, the output of one 40—watt (F40) lamp may
reasonably be compared to solar irradiance on one square meter (Table 2,
Fig. 7, 12—19).
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—9—
1JV fluorescent lamps can be used to simulate greater than average
ur er UV solar irradiance at 305 am and shorter wavelengths. These
ln ps have roughly constant output from 305 to 320 am and less output at
longer wavelengths (Fig. 12—17). However, solar spectral irradiance
ncrcaSe5 rapidly from 305 to 330 am (Fig. 7). Fluorescent lamps fall
far short of the sun’s power at wavelengths much above 300 am and do not
pnr 11el the sun’s spectral radiant power at wavelengths shorter than 300
on (Fig. 18, 19).
Fluorescent lamp output can be filtered to remove short wavelength
radiant power and more nearly approximate the spectral power distri-
bution of sunlight. But the only practical wey to make up for the
missing long wavelength radiant power in the LW—B is to permit some
shorter wavelength radiant power to take its place. The radiant power
must be correctly weighted, however, No available filters wi]l exactly
meet the requirements. However, cellulose acetate is an acceptable
compromise. It removes practically all 254 am radiant power transmitted
by the lamp envelope and phosphor. An appropriate thickness transmits
enough lamp power for biological experiments. The rate at which
spectral transmission decreases with wavelength simulates the effect of
stratospheric ozone more closely than any lamp envelope glass.
We have expressed cellulose acetate spectral transmission in an
equation. Narrow bandpass data, dated December 1976, furnished by the
National Bureau of Standards were supplemented with data collected within
SEA. We wanted data for Lransmission by filters of any age and thickness
Lit any wavelength. Surface transmission through a cellulose acetate film
was taken to be 0.88 although higher transmission is sometimes observed.
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—10-
Fast aging apparently occurs at the filter surface independent of
thickness. For practical purposes, fast aging i . complete after 5 or 6
hours near FS4O lamps (time constant 2.2 hours). Time constants depend on
ypc of UV lamp and irradiance. Natural logarithm of transmission as a
function of wavelength, A, decreases with time, h, in hours as follows:
(2) 0.053 li-e _0.45h} (X/233.9702) 22-(X/233.9702) 55
The “primary” absorber does not change with age, but its natural
logarithm of transmission (minus) is proportional to thickness, t, in
0.01 inch units:
(3) t 48 x 1O 9 (X/87.61830) 12 e -(A/87.61830) 3
Another absorber is also present initially and changes slowly with
time, h. Its natural logarithm of transmission (minus) is believed
proportional to thickness, t:
(4) t (0.120 + 0.0025 h) (Xf144.8l24)Oe /144812 4 )
The negative of the logarithm of transmission is thus 0.128 plus the
sum of three wavelength—dependent tcrms, (2, 3, and 4 above) two of them
thickness dependent and two of them time dependent. But one of the time—
dependent teri is constant for practical purposes, and the other time—
dependent term does not change the spectral distribution of transmitted
power enough to matter. However, total power is reduced significantly SO
fillers are usually changed every 3 — 4 days o [ 6 hours use per day.
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—11—
Spectral power output is given in Table 2 for cellulose
Ł L.t rc_fi1ter d lamps. The cellulose acetate is presumed to have aged 6
t-iuro near FS4O lamps. The spectral radiant power output of any
o tnati0fl of lamp and any thickness of cellulose acetate can be
calculated with the above equations.
Filters are not identical from batch to batch, whether the filter is
Ł lamp envelope or a separate film. A single lamp’s output varies
I t5 urably, and the effect)Ve ultraviolet output of one lamp compared to
.nother varies by 0.1 of output due to variable glass envelope thickness
%.1 ch removes more or less UV of short wavelength . For one sample of
0.005- inch thick cellulose acetate from Peoria we obtained transmission
v I ueS similar to those for 0.003—inch thick cellulose acetate described
by the equation. All cellulose acetate—filtered lamp outputs tabulated
w rc calculated with equations 2, 3, and 4.
Mylar has been used to remove short wavelength UV in experiments
wk ere a minus UV—B treatment was desired. Spectral transmission of the
ueuetl thickness of Mylar, 0.005 inch, is similar to transmission of an
tqu.al thickness of cellulose acetate, but the spectral transmisSion is
6h1 [ ted about 24 pj to longer wavelengths. Detailed equations have not
-een developed, but would have the same features as the equation for
cellulose acetate. Aging is at least as drastic for Mylar as for
cr)lulose acetate, but replacement of the filter often was neglected.
was commonly changed at intervals of one or more weeks.
V ERROR SOURCES
Errors were introduced by summing at 5—nm intervals the rapidly
C g ng bpectral power distributions such as those from cellulose
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—12—
acetate—filtered lamps. When the spectrum changes rapidly with
wavelength, the sums are smaller by about 1 part in 20 than the sums at
1—nm intervals. Sums of output are more nearly correct for unfiltered
than for filtered emissions. Equations were developed that describe
phosphor spectral emission and envelope spectral tranlnsission, when sums
at intervals smaller then 5nm are required. These equations were used to
plot Fig. 12—19, but are not reported here.
In biological experiments, spectroradiometer m& asurements should be -
made whenever possible. They reduce possible errors due to a spectral
power distribution that is different than expected. Errors might be
caused by substitution of a different lamp or external filter or by
absence of a filter on a portion of the ]amp that was presumed covered.
However, even a well calibrated spcctroradioineter can introduce
errors of its own. Cosine response is never perfect. Slow response time
and fast scanning speed can reduce the apparent irradiance when the scan
is from smaller thru larger spectral irradiance. Realistic bandpass of
field spectroradiometers can overestimate effective radiant power by 1
part in 10 at specific wavelengths. However, the error due to bandwidth
greater than zero can be calculated as follows:
Describe signal with exponential equation;
In 14 (A—A )
(5) yy e o
where y is the true value at wavelength X and
0 0
N is the ratio of y, at A A + 1, to y.
Choose triangular bandpass, base is twice the bandwidth (BW).
Define a E in M times BW
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—13—
Manipu)ate equations and find that R, the multiple of y by which
the reading is too large, is:
(6) R = (e 8 + a 5 —2) a 2
If a = 2, R 1.381, about the case for 4—nm BW and UV solar
spectral irradiance at short wavelengths at the eartlYs surface.
The apparent wavelength offset, A—As is (in R) I (ln M)
which approaches the bandwidth for very large H (Table 3). Real
instruments have some sensitivity at wavelengths outside the simple
triangular bandpass, so they provide a reading when they are more than
one bandwidth away from any real signal.
Because global irradiance (from a hemisphere) on a horizontal
surface is usually measured, radiant power intercepted by a horizontal
surface is presumed to be of interest. But not all plant surfaces are
horizontal. Maximum leaf responses to UV, chiorotic lesions in cucumber
and anthocyanin disappearance in Coleus , are observed an leaves that are
perpendicular to the received LW (See PSL FNC Final Report). Irradiance
received per unit solid angle may be of some importance, but, direct
measurements are complex.
We found evidence that shorter wavelength UV, to about 265 mu, is
more effective than longer wavelength LW. Since we cannot afford to
duplicate the sun’s power, a weighting function describing an action
spectrum must be used. Errors introduced by use of a weighting function
that does not describe the action spectrum will be many times as large as
errors caused by most types of measurement errors.
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—14—
IV WEIGHTING FUNCTIONS
Biologically effective UV—B may be (incorrectly) defined as the
physical definition of UV—B which e%tends from 280 to 320 nm, each
wavelength rated equally (i.e. weighted by a factor of one). This
treatment was used for most biological results reported in CIA? Monograph
V. (Ref. 12), i.e., the need for a weighting function was ignored.
Several weighting functions have been reported to describe various action
spectra that describe the biological effectiveness of UV as a function of
wavelength.
Green and Moe (C—N) derived a weighting function baE d on the
standard erythemal curve as follows (Table 4, Fig. 20, 21):
(7) 4 exp (X - 296.5)12.692) l + exp (X 296.5)/2.692Jj 2
This description disregards responses at wavelengths shorter than 290 urn.
However, available lamps emit at these short wavelengths and the
emissions have been found to be biologically active.
Setlow (1974) reported a DNA action spectrum and Green and Miller
(1975) described it in mathematical terms as follows (Table 4, Fig. 20,
21):
(8) exp 13.82 1{l + exp 1(X -310)/9){ -iJ
This equation does not precisely represent Setlow’s data, but provides a
somewhat realistic weighting function to test.
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--15—
Caidwell (1968) presented a genera]ized action spectrum basically
derived from DNA response, but did not mathematically describe its
weighting function. At a BACER terrestrial workshop held at Beltaville
in September 1976 the question of a mathematically defined weighting
function was discussed. Scientists at Beltsville proposed the initial
use of the following weighting function (Ac21) that describes several
DNA action spectra for bactericidal response (Table 4, Fig. 20, 21):
(9) = exp — t (265—X)/2lJ 2
This weighting function also approximates the action spectrum described
by Caidwell if common broadband UV sources are evaluated.
We tested the appropriateness of these 4 and other weighting
functions in experiments with plants. We will describe our results later
in the report, but, they show that the empirically derived equation (10)
(AE9) provided the best fit to our plant data (Table 4, Fig. 20, 21):
(10) = 10.25(A/228.178) 9 ° exp 4-(XI228.l78) 90 J
This equation is one of a family of curves used to describe spectral
absorption of filters. The characteristic wavelength was derived from
-------
—16--
the Rydberg constant (CODATA, 1973); we selected for the equation,
quantized conetants that best fit the biological data. We believe the
Ag9 equation is a Letter description of Set1ow s data than the equation
of Green and Miller. Several other weighting functions that have been
proposed cannot be distinguished from those presented if only a limited
wavelength region is considered.
Some investigators report that the biological effectiveness of
ultraviolet is not negligible through much of UV—A. Response curves
reported are similar to the stray light passed by a filter, suggesting
the response is not real. flowever, some investigators are convinced that
the response is real, but that it differs from the response observed at
shorter wavelengths. Exposure time at different wavelengths should be
about equal for a valid comparison of responses observed at various wave --
lengths (Firiney, 1964). If UV—A does provide protection from UY—B
injury, it is possible that it might be injurious when given in large
doses in the absence of UV—B.
The relative biological effectiveness at 5—nm intervals as derived
from equations 7, 8, 9, and 10 with the maximum response adjusted to oue
are given in Table 4. The spectral radiant power emitted at each
- wavelength (Table 2) is multiplied by the weight for that wavelength.
For total weighted radiant power, the summed values (5—nm intervals) are
multiplied by 5. adiant power for each individual mercury emission line
is multiplied by its weighting factor and added directly to the total
weighted sum obtained above.
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—17—
By use of the weighting values per each 5 nni shown in Table 4 for the
various weighting functions and the average sunlight shown in Table 1.,
the weighted control sunlight becomes:
Ac9 3.06 mW.m 2 , Setlow 2.22 mW.m 2 , AE21 = 14.9 mW.tn 2 ,
and G—M = 51.3 mW.m 2
Parry Moon (1936) noted that weighted power is not radiant power or
irradiation. However, because the weighting function is dimensionless, the
weighted outputs have dimensions of power. or irradiation. Because the weighted
power has been evaluated by a mathematical process, it cannot be expressed in
customary physical units. Weighted power units or, as some people prefer,
qualifying names must be attached to the customary units (NBS 1977, Ref. 9).
U, • I I
We proposed use of weighted Wm
Performing the same calculations for each spectral radiant power value per
each 5—nm for the various UV sources, i.e., multiplying mWnm 1 (Table 2) by
the indicated weighting function weight (Table 4) the one lamp power provided
by each source for each weighting function in weighted mW is obtained.
The weighted mWm 2 divided by-the magnitude- of the appropriate ieighted
mWm 2 for the weighting function under consideration for average sunlight
yields a ratio (Beltsville Control Sunlight) that compares weighted lamp power
to weighted average sunlight. (Table 5). Various weighting functions give quite
different characterizations of the biologically effective radiation provided by
the different sources. If the control sunlight selected were of larger power
than. ours but of the same spectral -power distibution, the lamp—filter ratings
would be smaller by the same (inverse) proportion.
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—18—
In any case, these weighting functions yield quite different
evaluations of the effective radiant energy required to elicit a given
plant response. The best—fit weighting function would be expected to
provide for equal plant response for equal weighted radiant energy
regardless of lamp source spectral power distribution.
Which part of the ultraviolet spectrum is important? Many investi-
gators have neglected radiant power of wavelength shorter than 280 nm.
But many practical lampsources emit radiant power of wavelength shorter
than 280 nm, at least in small quantities. We stopped measuring at 250
rim, presuming that shorter wavelength power was not present in
significant quantities. We have measured power output at 254 mu from all
fluorescent lamps, sometimes in significant quantities from UV—B lamp
sources. The long wavelength limit of interest is usually specified by
where the AE9 weighting function approaches zero.
The spectral radiant power from several sources was multiplied by the
Ac9 weighting function. The weighted power in 5—nm bands and the power
contributed by individual mercury emission lines was divided by the total
sum. The proportion of the weighted sum in each 5—nm band or mercury
emission line is tabulated (Table 7).
If the lamp is designed to produce UV—B, the phosphor contributes
much more than the mercury emission lines. If speccral sampling
completely misses a mercury line, the error is not serious. (Bandwidth
of the instrument should be consistent with the sampling interval. With
a narrow bandwidth instrument, widely spaced samples would give strange
appearing spectra if a mercury emission line w&re sampled.) The 256—nm
-------
—19—
mercury emission line usually contributes little except with the Rayonet
F8T5 RPR 3000A° and the bactericidal or germicidal lamp G15TB.
None of the lamps have the same spectral power distribution as
average sunlight. Table 7 shows the variation, weighted by Ac9 and
normalized to unit effective power. A different thickness of cellulose
acetate makes only a small wavelength change in spectral power
distribution of the lamp and filter combination.
V BIOLOGICAL EVALUATION OF WEIGHTING FUNCTIONS
Because the accuracy of a weighting function may not be tested
directly with a broad—band UV source, we derived a constant (m) from each
of the 4 weighting functions discussed above and the spectral power
distribution of each lamp—filter combination.
These constants were used to interpret our plant response data. The
constants (Table 6) are derived by dividing the ratio, Beltsville Control
Sunlight, (Table 5, lines 3 to 6) by the absolute UV—B power (line 1) of
the appropriate source for each weighting function (see also Appendix
II). For each lamp source, each constant characterizes the absolute
watts and its biologically effective UV as defined by the selected
weighting function.
The goodness of fit of the various weighting functions may now be
examined. By use of the constants derived for each of the 4 weighting
functions from Table 6, the logarithm of the observed plant response
(UV—B to obtain half reduction in dry weight) may be plotted as a
function of the logarithm of the corresponding constant.
-------
—20—
The data plot as a straight line described in linear coordinates by:
(11) E/2 mb wliere,
E/2 required to reduce dry weight by half
m = constant
V geometric mean, if b = —l
b = exponent defining goodness of fit
E (Table 9, 10) and m (Table 6) are entered as data. V and b
0
(Table 11, 12) are fitted. If the weighting function fits well, the
exponent b —1. When we require b —1, V 1 is the geometric mean
(Column 1). The associated correlation coefficient, r , (Column 2)
received two contributions whichwe would like to separate: the inappro-
priateness of the weighting function and the variability of the data.
When we fit both V and b, the new correlation coefficient, r ,
(Column 3) may be nearly the same as r . If the calculated b is
nearly —1, (Column 4), we expect about the same correlation coefficient
since about the same equation is fitted. If the calculated b is not —l
(perhaps —2) and its associated correlation coefficient r 2 is closer to
-------
— 21 —
1 than was r 1 , we have confidence the weighting function tested is not
the correct one to describe the biological spectral response of the
response measured.
If r 1 and r 2 are both small and nearly the same the equation
tested may be entirely inappropriate. Or, if the previous statement is
true for several equations, there may be too much scatter in the data to
test any equation proposed as a weighting function to describe the
spectral biological responsc.
Any other equation substituted for equation 7, 8, 9, or 10 could be
tested using the irradiation and response data presented. An obvious new
equation is the “b” power of an equation already tested, such as At21,
which has r 2 closer to 1 than r 1 . Additional response data, perhaps
at 254 urn, would then be needed to distinguish between different
equations that fit about equally well. An example of an inappropriate
“equation’ which might be tested is to weight each wavelength equally
from 300 to 320 urn and weight all other wavelengths 0.
To provide information on the effectiveness of the four weighting
functions, discussed above, in evaluating the effect of various portions
of the UV region, plant e.speriinents were carried out by the Florist and
Nursery Crops Laboratory and the Plant Stress Laboratory, Beltsvi lle,
}laryland, and portions of the data obtained subjected to the above
analysis. Four spectrally different sources of UV were used, as follows:
-------
—22—
(1) unfiltered FS4O, (2) unfi)tered FBZS4O—CLG, (3) FS4O lamps filtered
with 0.005—inch thick (5 sill) cellulose acetate and (4) FS4O lamps
filtered with 0.010—inch thick cellulose acetate. Spectral differences
are shown in Table 2. A four—fixture configuration was used, Fig. 10. A
range of irradia ice was obtained by placing the test plants at varying
horizontal distances on the x and y surface from the source. We
recognize that this method provides an unavoidable correlation between
the horizontal irradiance and both the direction and size of the solid
angle from which it is obtained. The absolute UV—B power in Wm 2 for
maximum irradiance levels used in these experiments are shown in Table
8. Absolute levels varied about 10—fold under each set—up, depending on
plant position relative to the lamps. A typical experimental set—up is
shown in Fig. 11.
The species ae]ected for these experiments were the most sensitive to
UV—damage of the plants investigated at Beltsville. They were a cucumber
cultivar (Poinsett) and a Coleus cultivar (Frederici); Coleus was the
less sensitive of the two. In addition to plant growth responses, both
cultivars displayed a pigment change in response to UV. Frederici leaves
are deeply pigmented (anthocyanin). In response to enhanced UV—B,
pigment production is suppressed in newly emerging leaves. The Leaves of
Poinsett develop chiorotic lesions in the presence of UV—B of relatively
low irradiance levels. These pigment changes provided a means, in
8ddjtjo to growth responses, of measuring the effectiveness of the four
tleighting functions tested. Plants were grown either in a glass
greei hou that removed most of the UV—B from the sun or in a fiberglass—
Plastic greenhouse that removed, in addition, thc. TJV—A, but transmitted
blu 9 and longer wavelength visible light. Ultraviolet was measured at
-------
—23--
selected locations under each set—up with an IRLSpec spectroradiometer
and at all plant locations with the UV—B IRLMeter radiometer.
The detailed experimental methods and results are reported in the
Florist and Nursery Crop and Plant Stress Laboratories Final Reports.
Data obtained included: pigment response, plant shoot fresh and dry
weight and leaf area not including cotyledons for each plant. For our
analyses of weighting function effectiveness, only the dry weight data
were used. However, subjective evaluations of pigment supres5ion also
provided the means for an additional verification of the dry weight
results. Poinsett cucumbers were grown in May 1977 in both glass (H) and
plastic (L) greenhouses at 16 different UV—B irradiances under each array
(Table 8). Coleus (Frederici) were grown in June and July 1977 in the
glass (F,S) greenhouse at 26 to 29 UV—B irradiances. Two quarters of
each array were used and their plant responses are reported separately.
Cucumbers were grown again in July in the plastic (T) house at 26
different 1W irradiances. Differences between arrays for predicted
growth in the absence of UV may be due in part to temperature and
seasonal differences within the greenhouse (Tables 9, 10).
Results
Data were subjected to linear regression analysis and were found to
approximately fit a straight line characterized by:
(12) y/y 1 — E/E where
y = Observed growth at measured E
E = Absolute irradiance level
y = Predicted growth with zero 1W
E 0 = Absolute irradiance level predicted for zero growth
-------
—24-
The predicted growth in the absence of DV (y 0 ) and the predicted
absolute irradiance required to reduce growth by one—half (E/2) are
shown in Table 9.
The reciprocal of dry weight plotted as a function of UV—B power also
approximates a straight line described by:
(13) y/y = 1+E/E 5
where E is the absolute irradiance level predicted fcr one—half
growth. The other symbols are defined as in Equation 12. Regression
coefficients y and E are in Table 10.
o o.5
Both models predict growth reduction for very small exposures, but
actual reduction could not be observed directly because of variable plant
growth. Plant response was too small with respect to its variability for
us to adequately test the shape of the growth response curve. Therefore,
although we found (Table 4) that four times the present biologically
effective DV would be required to reduce the growth of a sensitive plant
by one—half, we cannot reliably predict from these experiments the effect
of a 20—40% enhancement of UV—B on plant growth and development. We
expect that the effect would not exceed that predicted by the regression
equations, but it might be less.
Plant top dry weight ranged over a multiple of 2 among groups of
plants in the same experiment, due to non—treatment causes. Effects of
-------
—25—
array position within the greenhouse could not be removed. Nevertheless,
within each group under an array, plant growth as a function of UV—B
could be described with either equation to predict the no—treatment
growth and the treatment needed to produce half the no— treatment growth
(Tables 9, 10).
Predicted treatment needed to produce half the no—treatment growth
was larger for Coleus (Frederici) than for the cultivar of cucumber
(Poinsett) grown (Table 9—12). It also was larger for the cucumber grown
in midsummer with higher temperatures and more sunlight than for the same
cultivar grown earlier and suggests the existence of a photorepair or
photoprotective mechanism in the test organisms.
Predicted IJV—B needed to produce half the no—treatment growth depends
on the spectral power distribution of the source including the filter.
Prediction varies by a factor of 4 to 8, depending on the source of UV—B
(Table 9,10). In the BACER program, except in certain special research
areas, the UV—B sources used were the FS4O lamp with appropriate plastic
filters or the FBZS4O—CLG lamp with the glass envelope acting as the
filter.
When we know the spectral power distribution of the lamp and filter
combination, and also the relative spectral effectiveness of ultraviolet
for the organism, we can use a practical source of ultraviolet to
simulate present or changed ultraviolet of sunlight, that is now incident
on the earth’s surface.
-------
—26—
VI SUMMARY AND CONCLUSIONS
Absolute UV—B does not describe plant response to ultraviolet. UV—B
can be used as a variable to evaluate other weighting functions.
The conversions for the unfiltered FS—40 are particularly interesting
because this lamp was used to simulate the sun’s UV—B in earlier experi-
ments. To convert from normalized IN—B to Ac9 weighting function units
we divide by 1.3 and multiply by 43 (Table 5, Appendix II), an overall
multiplication of 33. Converting from normalized LW—B to Setlow
weighting function units, divide by 1.3 and multiply by 67, an overall
multiplication of 52.
A comparison of the goodness of fit of the various weighting
functions to interpret dry weight data using the equation E = Vm 1 ’
is presented in Tables 11 and 12. Two treatments of the results are
presented; (1) by forcing b = —l for each weighting function, the
geometric means of E 0 (Beltsville Control Sunlight units) are derived
for the various weighting functions and their correlation coefficients
are compared as a measure of how well the weighting function describes
the observed plant data (columns 1 and 2); and, (2) the equation is
solved for b for each weighting function with correlation coefficients
indicating the accuracy of b (columns 3 and 4).
Plant response varies about as the second or third power of the
constants (m) calculated from the C—N weighting function (column 4).
Correlation coefficients were fairly good (co1uu i 3). However, when a
-------
—27—
linear relation was forced on the data (b —l) poor correlation
coefficients were obtained (column 2). Of the four weighting functions
tested) the erythemal provided the poorest fit to observed plant response.
Plant response varied about as the 1.5 power of the constants
calculated from our Aa21 weighting function. The correlation
coefficient was better, but also decreased when a linear relation was
forced. This weighting funclion also does not fit plant response well.
Plant response was nearly linearly inversely proportional to
constants calculated from either the equation respresenting Setlow’s data
or our Ac9 equation, as desired. The correlation coefficient was
usually better for constants calculated from the AE9 equation. Also,
leaf pigment response was more nearly the same for equal effective
irradiance calculated by the &9 equation. The Ac9 equation provides
the best fit to available data. It is suitable for interpreting results
of UV—B enhancement experiments.
We cannot reliably predict from these experiments what effect a
20—40% enhancement of 1W—B would have on plant grouth and development.
We expect it would be no greater than predicted by the regression
equations, but it might be less.
-------
—28—
REFERENCES
1. Abbott, C. C., Solar Variation and Weather, Smithsonian Misc.
Collection, Vol. 146, No. 3, pp 2—9. 18 Oct 1963.
2. Caidwell, N.M., Solar Ultraviolet Radiation as an Ecological
Factor in Alpine Plants, Ecol. Monogr., 38:243—268, 1968.
3. Caldwell, N. M., in “Research in Photobiologv” (A. Castellani, ed.)
p.600. Plenum Publishing Corporation, 1977.
4. CODATA Bulletin No. 11, International Council of Scientific Unions
CODATA Central Office, 19 Westendstrasse, 6 Frankfurt/Main, West
Germany (December 1973)
5. Finney, D.J., in “Statistical Methods in Biological Assay”, (2nd.
ed.), page 57, Charles Griffin & Co. Ltd., 1964.
6. Green, A.E.S., and J. H. Miller, “Impacts of Climatic Change on the
Biosphere, Part I: Ultraviolet Radiation Effects”, (Edited by
D.S. Nachtwey, M.M. Caidwell, and R. H. Biggs), pp. 2—60 to 2—70.
U.S. Dept. Transportation, DOT—TST—75—55, Washington D.C., 1975.
7. Kaufman, J.E., and J.F. Christensen, editors, IES Lighting Handbook,
The Standard Lighting Guide, pp. 9—56 to 9—58, Illuminating
Engineering Society, Fifth Edition, 1972.
8. Moon, P., “The Scientific Basis of Illuminating Engineering”, p. 38,
Electrical Engineering Texts, McGraw—Hill Book Company, Inc., New
York and London, First Edition, 1936.
9. Optical Radiation News, page 2, No. 22, US Department of Commerce,
National Bureau of Standards, October 1977.
10. Setlow, R.B., “Wavelengths in sunlight effective in producing skin
cancer: A theoretical analysis”, “Proc. Nat. Acad. Sd. USA”, 71:
3363—3366, (1974).
11. Thekaekara, M.P., “Extraterrestrial Solar Spectrum,, 3000—6100 A°
at 1 —A° Intervals”, Applied Optics, Vol. 13, pp. 518—522, March
1974. (Original data was collected 3—19 August 1967)
12. ClAP Monograph V, “Impacts of Climatic Change on the Biosphere, Part
I: Ultraviolet Radiation Effects”, (Edited by D.S. Nachtwey, M.M.
Calduell, and R. H. Biggs), U.S. Dept. Transportation, DOT—TST—75—55,
Washington, D.C., 1975.
-------
—29—
TABLES
Table 1 Average UV—B and UV—C Soldr Spectral Irradiance, June and
July, 1976, 6 hour day.
Table 2 Spectral UV—B and UV—C power output of one representative
lamp and filter of several types compared to sunlight.
Table 3 Estimation of error ratio (R) due to bandwidth (BW).
Table 4 UV Weighting Functions Tested, maximum adjusted to 1.
Table 5 Absolute and Weighted UV Power Output of One Representative
Lamp and Filter Compared to Control Sunlight.
Table 6 Constants, m, for Absolute UV Irradiance to Weighted Power,
Beltsville Control Sunlight.
Table 7 Effective UV Power Proportion, Ac9 Weighting Function.
Table 8 Maximum Irradiance in Experimental Setup, UV—B, 280—320 rim
Table 9 Predicted Irradiance to Reduce Dry Mass Growth One—Half of
Predicted Growth of Untreated Plants, UV—B, 280—320nm
Equation y/y 0 1 — E/E 0 .
Table 10 Predicted Irradiance to Reduce Dry Mass Growth One—Half of
Predicted Growth of Untreated Plants, UV—B, 280—320nm
Equation y 0 /y = 1 + EIE 0 5 .
Table 11 Correlation of Irradiances CE) to Reduce Dry Mass Growth
One—half with Multipliers for Four Spectral Power
Distributions. Multipliers (m) Derived from Four
Weighting Functions.
Equation y/y 0 = 1 — E/E 0 fitted.
Table 12 Correlation of Irradiances (E) to Reduce Dry Mass Growth
One—Half with Multipliers for Four Spectral Power
Distributions. Multipliers (m) Derived from Four
Weighting Functions.
Equation y 0 fy = 1 + E/E 0.5 fitted
Table 13 Weighted UV Power Output of One Representative Lamp
and Filter.
-------
—30—
TABLE 1
AVERAGE UV-B AND UV-C SOlAR SPECTRAL IRRADIANCE,
JUNE AND JULY 1976, 6-HOUR DAY.X/
Wavelength, nm Average Power, wm 2
320 264
315 179
310 93.7
305 29.5
300 3.13
295 0.136
290 O.OO0
!1 Used as standard average sunshine for presenting Beltsvij.le
Agriculture Research Center data in meaningful units similar to
those used for enhancement. Based on Smithsonian data, modified
with data from the literature and data collected by LRL at Beltsville ,
Maryland.
-------
31—
‘LE 2
SPECTRAL 1W—B AND 1W-C POWER OUTPUT!’ OF ONE REPRESENTATIVE LANP AND FILTER OF SEVERAL TYPES COMPARED TO SUNLIGHT
Column No. 1 2 3 4 5 6 7 8 9
Wavelength Average Astronaut FS4O FS4Q FBZS4O FS4O G15T8 RPR RPR
(nanometer) Sunlight on Sunlight on —CLG 3000 A° 3000 A°
one square one square FBT5 F8T5
meter meter CAI0 J CA5 ! no filter no no no filter CA5 ’
(Smithsonian) (Thekaekara) NBS Jan 77 filter filter
Phosphor milliwatt/nanometer watt/micrometer = megawatt/meter
320 264 830 55.4 61.5 41.8 94 0 9.3 6.1
15 179 764 58.2 65.5 54.0 104 0 12.6 7.9
10 93.7 689 55.7 63.8 59.1 105 0 16.2 9.8
05 29.5 603 46.5 55.7 52.3 97 0 18.3 10.5
300 3.13 514 24.6 36.3 35.0 78 0 17.8 8.28
95 .136 584 1.79 8.24 16.7 55 0 13.6 2.04
90 .0004 482 5.4(—6 il(—3 5.48 32 0 7.55 0
85 0 315 0 0 1.11 15 0 2.15 0
280 0 222 0 0 0.14 5.7 0 0.308 0
75 0 204 0 0 0 1.5 0 0.032 0
70 0 232 0 0 0 0.2. 0 0.003 0
65 0 185 0 0 0 0 0 (0.012 Hg lines)
260 0 130 0 0 0 0 0 0
mercury emmision line . ’, milliwatt
313 0 0 21.9 31 50 50 51.6 12.5 7.8
302.2 0 0 1.65 2.7 5 5 7.3 1.3 0.67
296.7 0 0 0.33 1.0 3.8 3.8 12.0 2.0 0.55
289.4 0 0 0 83(—6 1.2 1.2 3.0 1.0 66(—6
253.7 0 0 0 0 0 1.2 4300 32.6 0
1/ Zero (0) denotes quantities believed to be negligible.
2/ Cellulose acetate (CA) thickness in 0.001 inch increments, (aged for 6 hours near FS4O lamps).
3/ Phosphor power output is given per unit wavelength at 5 nm intervals.
2/ Mercury emission line power is given per lamp. These lines must be added to fully characterLze spectral
— radiant power, but these contributions are usually small.
-------
—32—
TABLE 3
ESTIMATION OF ERROR RATIO CR) DUE TO BAI DWIDTH (Bw)
Wavelength off etV
.1 1.000834 8.33 x 10
.2 1.00334 16.66 x 10
.5 1.0210 41.6 x 10
1 1.0862 82.6 x 10
2 1.381 161.4 x 10
5 5.86 354 ;c 10
10 220 539 x 10
1000 x 10
1/ a is band width (BW) times the natural logarithm of N. N is the
multiple of signal at unit wavelength from reference wavelength com-
pared to signal at reference wavelength (X 0 ).
2/ R is the ratio of the observed (theoretical) reading to the correct
reading.
3/ The multiple of bandwidth the (theoretical) reading appears to be
offset from the wavelength at which the observed reading should be
obtained.
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—33—
TABLE 4
UV WEIGHTING FUNCTIONS TESTED
Maximum responsc adjusted to 1
1/ Setlow summary data for DNA
equation.
response represented by Green and Miller
2/ Standard erythema response represented by Green and Moe equation.
Column No.
1
2 3 4 5
Ac 21
UV-B /
Wavelength
nanometer
25
320
15
10
05
300
95
90
85
280
75
70
65
260
55
50
45
240
313
302.2
296.7
289 .4
253.7
Ac9
.000002
• 000032
.00029
.00187
.00891
.0323
.09 17
.209
.391
.615
.827
.966
.997
.920
.769
.587
.414
.272
.00063
.0189
.0660
.227
.723
Set lowl’
.000009
.000031
.000 15
.00 100
.0065
.0326
.111
.259
.445
.621
.758
.852
.912
.948
.970
.983
.990
.994
.0003 2
.0167
.0767
.280
.974
.00105
.000647
0
to
.00345
.00414
.01013
.0262
.0266
.157
.0622
.673
.130
.926
.242
.301
.404
.0543
.600
.00867
0
to
.797
.00136
0
.945
.00021
0
1.000
.000033
0
.945
.000005
0
.797
.000001
0
.600
0
0
.404
0
.242
0
1
1
1
1
1
1
1
I
1
1
1
1
1
.00538
.0434
102
.259
.749
.00867
.384
.999
.249
5x10 7
0
3/ Definition.
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-.34-
LE S
ABSOLUTE AND WEIGHTED UV POWER OUTPUT OF ONE REPRESENTATIVE LANP AND FILTER
COMPARED TO CONTROL SUNLIGHT
157—B, watt 2.19
(absolute)
UV—B, I
normalized
Normalized weighted power
Ac9 I
SETLOW 1.
Ac21 1
1
1.10 1.34
0.50 0.61
to ratio/energy or
2.5 4.2
3.1 5.8
1.3 2.0
2.7 4.2
FBZS4O FS4O
-c LG
no filter
RPR
3000 A°
RPR
3000 A°
F8T 5
CA5I /
1/ Cellulose acetate (CA) thickness in 0.001 inch increments, (aged for 6 hours near FS4O lamps).
Column No. 1 2 3 4 5 6 7 8
Source Average
FS4O
FS4O
Sunlight
on one
square
CAb! ’
CA5 1 ,
.
filter
me ter
F8T5
no filter
G15T8
no filter
1.28 2.74 0.074
0.58 1.3 0.034
power units (Beltsville Control Sunlight.
8.3 43 1001)
12.5 67 1900
2.9 12 220
5.1 13 0 .31
0.48 0.22
0.22 0.10
0.95
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—35—
TABLE 6
CONSTANTS, tu, FOR ABSOLUTE UV IRRADIANCE TO WEICdTED PO YER,
BELTSVILLE CONTROL SUNLIGhT
Source FS4O F 540 FBZS4O FS4O
-CLG
CA1O!/ CA5V no filter no filter
Ac9 2.2 3.2 6.4 16
SETLOW 2.8 4.3 9.8 24
A 2l 1.2 1.5 2.3 4.3
G—M 2.4 3. 1 4.0 4.7
1/ Cellulose acetate (CA) thickness in 0.001 inch increments,
— (aged foi 6 hours near FS4O lamps).
-------
TABLE 7
EFFECTIVE UV POWER PROPORII , Ac9 WEIGHTING FUNCTION
Column No. 1 2 3 4 5 6 7 8
Wavelength Weighted FS4O FS4O FBZS4O FS4O G15T8 RPR RPR
Nanometer Control —CLG 3000A° 3000 A°
Sunshine F8T5 F8T5
CA b. ?! CA5 ’ no filter no filter no filter CA5.a’
proportion effective power in 5 tim band 2 ’
25 .0013 79(—6 50(—6 14(—6 7(—6 0 2(—6 lS(—6
320 .0137 .0012 775(—6 264(—6 113(—6 0 31(—6 334(—6
15 .085 .011 .007 .003 .001 0 386(—6 .0040
10 .286 .069 .046 .022 .007 0 .0032 .032
05 .429 .274 .191 .092 .033 0 .0173 .16].
300 .165 .525 .452 .224 .095 0 .061 .461
95 .020 .108 .291 .303 .191 0 .132 .322
90 136(—6 0.75(—6 .001 .227 .253 0 .167 933(—6
85 0 0 0 .086 .222 0 .089 0
280 0 0 0 .017 .132 0 .020 0
75 0 0 0 0 .047 0 .0028 0
70 0 0 0 0 .007 0 .0003 0
65 0 0 0 0 0 0 .0013 0
260 0 0 0 0 0 0 0 0
proportion effective power in mercury emission 1ine !
313 0 .0023 .0015 .0013 .0002 l1(—6 168(—6 .0017
302.2 0 .0052 .0039 .0038 .0007 44(—6 501(—6 .0043
296.7 0 .0036 .0053 .0098 .C019 255(—6 .0028 .0125
289.4 0 0 l(—6 .0113 .0022 219(—6 .0048 5 (—6
253.7 0 0 0 0 .0068 .999 .497 0
1! Sum of each column is one (1). Zero (0) denotes quantities believed to be negligibly small.
2/ Cellulose acetate (CA) thickness in 0.001 inch increments, (aged for 6 hours near FS 40 lamps).
3/ Phosphor power output is given per 5 nm band.
4/ Mercury emIssion line power is given per lamp. These lines must be added to fully characterize spectral
radiant power, but these contributions are usually small.
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—37—
TABLE 8
MAXIMUM IRRADIANCE IN EXPERIMENTAL SETUP
UV—B,280—320 nm
Experiment !/
H
L
T
F
S
Irradiance (E), Wm 2
1.60 0.66
1.19 0.59
1.84 1.19
2.02 1.60
1.92 1.77
0.29
0.18
0.53
0.84
0.84
Co lumnNo.
1
2
3
4
Spectrum
FS4 O
CA1O
FS4O
CA5
FBZS4O-CLG
no filter
FS 40
no filter
1.03
1.25
1.07
1.53
1.30
1/ H Poinsett cucumbers grown
L Poinsett cucumbers grown
T Poinsett cucumbers grown
F, S Coleus (Frederici) grown
in May in glass greenhouse
in May in plastic greenhouse
in July in p]astic greenhouse
in June and July in glass greenhouse
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—38—
TABLE 9
Predicted Irradiance to Reduce Dry Ifass Growth One—Half
of Predicted Growth of Untreated Plants
UV—B 280—320 rim
Equation y/y 0 = 1 — E/E
Column No. 1 2 3 4
Spectrum FS4O FS4O FBZS4O-CLG FS 40
CA 1O CA5 no filter no filter
Predicted Irradiance (E 0 12), W.tn 2
to Reduce Dry Mass Growth One—half
Experiment 1/
H 1.63 1.66 0.71 0.28
L 1.64 1.30 0.57 0.26
T 1.84 2.45 1.72 0.57
F 5.38 5.92 2.3k 1.04
S 5.33 5.06 2.25 0.81
Predicted Dry Mass(y 0 ), g in Absence of UV—B
H 0.60 0.66 0.55 0.56
L 0.79 1.04 0.76 1.25
T 0.79 0.66 0.62 0.58
F 2.04 2.19 2.62 2.67
S 2.06 2.18 2.47 2.67
1/ H Poinsett cucumbers grown in May in glass greenhouse
L Poinsett cucumbers grown in May in plastic greenhouse
T Poinsett cucumbers grown in July in plastic greenhouse
F, S Coleus (Frederici) grown in June and July in glass greenhouse
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—39—
TABLE 10
Predicted Irradtance to Reduce Dry Mass Crotith One—Half
of Predicted CroRth of Untreated Plants
UV—B 280—320 net
Equation y 0 fy = 1 +
H
L
T
F
S
Predicted Dry Mass(y 0 ), g
0.62 0.71
0.80 1.14
0.80 0.68
2.01 2.16
2.04 2.15
in Absence of UV—B
0.59
0.92
0.62
2.54
2.48
0.60
1.31
0.61
2.69
2.66
1/ H Poinsett cucumbers grown in May in glass greenhouse
L Poinsett cucumbers grown in May in plastic greenhouse
T Poinsett cucumbers grown in July in plastic greenhouse
F, S Coleus (Frederici) grown in June and JuJy in glass greenhouse
Column No. 1 2 3 4
Experiment 1 1
Spectrum FS4O
P 540 FBZS4O—CLG
PS 40
CA 1O
CA5 no filter
no filter
Predicted
Irradiance cLm 2
to Reduce
Dry Mass Growth One—half
N
2.09
1.59 0.74
0.28
L
2.15
1.26 0.43
0.32
T
2.56
2.97 2.31
0.60
F
8.20
9.99 3.41
1.23
S
8.79
8.53 2.85
0.89
-------
—40-
1ABLE 11
Correlation J’ of Irrodiances CE) to Reduce Dry Mass Growth One—half with
Multipliers for Four Spectral Power Distributions.
Multipliers (m) Derived from Four i eighting Functions.
Equation 3’IYo = 1 — E/E 0 fitted
Weighting
Func t ion
Experiment.?! geometric mean
fit E 0 /2 S/rob
21 11 Poinsett cucumbet.s grown in
L Poinsett cucumbers grown in
T Poinsett cucumbers grown in
F, S Coleus (Frederici) grown in
iay in glass greenhouse
May in plastic greenhouse
July in plastic greenhouse
June and July in glass greenhouse
Column No.
1 2 3
v 1 ’ r 2 r 2
4
-bY
H
4.4
.969
.970
.97
L
3.9
.993
.995
.97
T
7.5
.553
.772
.65
F
15.4
.948
.955
.92
S
13.7
.978
.978
1.01
H
6.3
.941
.962
.87
L
5.5
.975
.995
.88
T
10.6
.351
.71’2
.58
F
21.8
.911
.949
.83
S
19.4
.961
.969
.91
Ac 9
SEThOW
Ac 21
G-M
1/ Correlation coefficient r 2 shows goodness of fit of different
•multipliers derived from different weighting functicns in interpreting data
derived from different spectral power distribution UP sources.
H
1.74
.878
.976
1.46
L
1.52
.893
.990
1.46
T
2.95
.804
.804
1.01
F
6.03
.880
.957
1.40
S
5.38
.864
.985
1.54
B
2.97
.508
.839
2.69
L
2.60
.546
.929
2.79
T
5.03
.457
.537
1.63
F
S
10.3
9.18’
.518’
.495
.825
.850
2.57
2.83
31 Beltsville Control Sunshine to reduce growth one—ha]f calculated y
setting b —1.
4/ The weighting function that correctly describes the action spectrum
has b’ —1.
-------
—41--
TABLE 12
Correlatioj/ of Irradiances (E) to Reduce Dry Mass Growth One—half with
Multipliers for Four Spectral Power Distributions.
Multipliers (m) Derived from Four Weighting Functions.
Equation y 0 /y = 1 + E/E 05 fitted
Weigh ting
Function
Exper iment.a’geometric mean
fit E 05 Vmb
2/ H Poinsett cucumbers
L Poinsett cucumbers
T Poinsett cucumbers
F, S Coleus (Frederici)
gown
grown
grown
grown
3/ Beltsvil]e Control Sunshine to reduce growth one—half calculated by
setting b= —1.
4/ The weighting function which correctly describes the action spectrum
has b —1.
Column No.
1
2
v i 2./
3
r 2
4
r 2
L
4.0
.907
.907
1.00
T
9.4
.721
.797
.76
F
22.3
.935
.939
1.08
S
19.2
.938
.977
1.25
H
6.7
.991
.99S
.94
L
5.7
.919
.927
.91
T
13.2
.596
.766
.68
F
31.6
.929
.930
.97
S
27.2
.957
.970
1.13
H
1.9
.865
1.58
L
1.6
.788
1.49
T
3.7
.809
1.16
F
8.7
.804
1.63
S
7.5
.762
1.89
Ac9
SETLOW
A 21
C-H
1/ Correlation coefficient r 2 shows goodness of fit of different
multipliers derived from different weighting functions in interpreting data
derived from different spectral power distribution l iv sources.
.999
.882
.829
.946
.980
H
3.2
.510
.916
2.99
L
2.7
.527
.967
3.08
T
6.3
.440
.577
1.95
F
14.9
.444
.788
2.95
S
12.9
.419
.855
3.50
in May in glass greenhouse
in May in plastic greenhouse
in July in plastic greenhouse
in June and July in glass greenhouse
-------
—42—
TABLE 13
WEIGHTED UV POWER OUTPUT OF 0I E REPRESENTATIVE LAMP AND FILTER
Column No. 1 2 3 4 5 6
Source Control F 540 FS4O FBZS4O FS4O C15T8
Sunlight CLG
one CA1O.LI CA5! 1 no filter no filter no filter
sq uare
meter
UV—B, watt 2.19 1.10 1.34 1.28 2.74 0.074
Adjust multiplier in eac1- weighting function to normalize weighted power to
arbitrary energy or power units.
IJV—B 1 0.50 0.61 0.58 1.3 0.034
Ac9 1 2.5 4.2 8,3 43 )00 0
SETLOW 1 3.1 5.8 12.5 67 1900
AE21 1 1.3 2.0 2.9 12 220
G—M 1 2.7 4.2 5.1 13 0.31
Calibration constants for UV—B IRLMeter Radiometer, i0 scale
100 29 34 30 69 26
Multiply UV—B IRLNeter Radiometer 1O scale readings, to obtain UV--B, raW,
y l. at 280—320 cm, by
22 38 39 42 40 2.8
Cellulose acetate (CA) thickness is expre sed in 0.001 inch increments.
CAIO is a cellulose acetate filter 0.01 inch thick, aged for 6 hours
near FS 40 lamps.
-------
—43—
Daily UV Total Energy per nm for
Coefficient of Variatio.i = 0.4563
1976 Rockville, Maryland.
Daily UV Total Energy per nm for
Coefficient of Variation = 0.3936
1976 Rockville, Maryland.
Daily l iv Total Energy per nm for
Coefficient of Variation 0.3545
1976 Rockville, Maryland.
Daily UV Total Energy per nm for
Coefficient of Variation = 0.3188
1976 Rockville, Maryland.
Daily IJV Total Energy per nm for
Coefficient of Variation = 0.3009
1976 Rockville, Maryland.
FIGURES
Solar Spectral Irradiance, Earth, Outside Atmosphere,
(Thekaekara, Ref. 11).
Figure 1
Figure 2 Probability for Given
5 nra Band at 295 nra.
June (+) and July Cx)
Probability for Given
5 nm Band at 300 nra.
June (+) and July Cx)
Probability for Given
5 nra Band at 305 nra.
June (+) and July (x)
Probability for Given
5 nra Band at 315 nm.
June (+) and July (x)
Probability for Given
5 nra Band at 320 nra.
June (+) and July (x)
Figure 3
Figure 4
Figure 5
Figure 6
Figure 7
Figure 8
Figure 9
Figure 10
Figure 11
Figure 12
Figure 13
Figure 14
Figure 15
Average UV solar spectral irradiance, June and July, 1976,
6 hour day.
Reflector for UV Sunlamps.
Normalized Irradiance 0.82 meter under a typical UV—B Lamp
Array.
Typical UV—B Lamp
Typical UV—B Lamp
Spectral IJV power
Spectral UV power
FS4O lamp.
Spectral UV power
Spectral UV power
FBZS4O—CLG lamp.
Array.
Array in use.
output of one representative FS4O lamp.
output (logarithmic) of one representative
output of one representative F]IZS4O—CLG lamp.
output (logarithmic) of one representative
-------
-44—
FIGURES
Figure 16 Spectral UV power output of one representative F8T5 RPR
3000A° lamp.
Figure 17 Spectral UV power output (logarithmic) of one representative
F8T5 RPR 3000A° lamp.
Figure 18 UV power output of one 40—watt (F40) lamp compared to Average
Solar Irradiance.
Figure 19 l iv power output (logarithmic) of one 40—watt (F40) lamp compared
to Average Solar Spectral Irradiance.
Figure 20 Weighting Functions.
Figure 21 Weighting Functions (logarithmic).
Figure 22 Relative Response of UV—B IRLMeter Radiometer
-------
—45—
Figure 1. Solar Spectral Irradiance , Earth 1 Outside Atmoophere
(Thekeekara, Ref. 11).
2400
2000
600
1200
800
400
WAVELENGTH, p m
-------
—46-
Figure 2. Probability for Given Daily UV Total Energy per nm for 5 nni
Band at 295 tim. Coefficient of Variation = 0.4563
June (+) and July (x) 1976, Rockville, Maryland !‘
--
-
..—.- .l.
L
c
• — .
±
—4.
*
- t
_-j
-1
1
±1 LJ
1- 4 ._i
L1 L
I
1
I
I
I. 4
Li
t
I
All I i
L. . LII i14 ...
. .
:!JL UI:
:UJITh
±ft
II
-W i
_._ I
I . ’ —
U
LU
Li
1111
i
ii 1 J I’ j
iiI [ ft [ Iij= ‘
ft
I
i iLt ft
L 11 IL
Li
— I
:L .
- t I
I 4 .4
_ -_ h__ A .
I,
1.
1 t-ILi!
L
ITL
L I !
.990 .995 . 7
Cumulative Probability
1/ Based on Smithsonian data modified with data in literature and data
collected at Beltsville.
I l1
- :: : 1
Ii
I,
1,
! -
EL
—4
I i— - .
-f - -4
‘4,-
i
- - 4- .--
4- I 4
1..
—II 44
: i .
iT ’1iff
I
1
4-. 4
4 ii
44 .j 4.
- , .t 1 -
.1
I ,
I,
:::L..
LdLLL tL
I- - -I 4--
JJ1Tt._1:Lt : IL
-H— -— -1--i t-H-t-
‘I, 1144
I;
I it
tr -H
.4.4
ii
4- I -
lii:
I t-- I-
‘ -I
I I
II
.4 141
.I1 14 .l_
1 E-t -
j 1i -
41
U
II
it
4.1. 44 1
- . 1 Ii ...
441
ii1
i—I.
4.4
‘--4
14
j t4
-4 I I 4.
I_i I
I 4- 1
• 4
ft
• 4 .4 4..4.
411 A,
.4.. .
; r. L’::r
4.
411
-I-
!1 f
.1
lit
I.
— l
_ 1
4—
-I
:j :LTIIt
:1.
- —I--s
I i- I I 1T 4: itt
1Ii• 1 . 1 .j .L it
.1 IA
. 4 1_I 1 -
I 4 1_ .
. II.. ..4 .-A
I 4 I. 14 . I_4
114 I IJ.
1 ItLL
I 4.4 I . . I
I 4 I
A..
- I -
-4
I
I—
— I-—
• -4.
—4
——
if
4 I I4
—-- 4--.
I —
I L- I l 1 4-U
L LJIj .II
.J_i.iI .-1
—4.
-4-4
LI ,’
41.41 44
I ’ ll
1 -i
4 AJ . 2.94
•IXIOI Oft .0) .05 .10 .20 .)O .40 .S0 ,.bO JO .*) .90
.9 .97 .9.
-------
—47—
Figure 3. Probability for Given Daily UV Total Energy per nm for 5 rim
Band at 300 nm. Coefficient of Variation 0.3936
June (+) and July (x) 1976, Rockville, Maryland 1/
$ tt p p’ :
.05 .10 .20 . . O SO .bO .7
Cumulative Probability
!/ Based on Smithsonian data modified with data in literature and data-
N
. -1. ifl TTT fiji IJIP ftfll tt:f: 1 111 J? !flUflj: L 11I U ft JTT H—I -‘1 — •i —1 TN{
; t 1 T1 7 T7i 1 -
V#__44 .r i -ft J! J .i . JiJf 1
1 1 : .::’. i] ;: it J:
H b L 1 L4
L 1 1 3 _ i l . i 4--
i J__ .L i liii +i; iiu J L tt thL 1:L
NL ll ft 1 1 j lj i 1 JIIIJ± iL 1 hLi
i f 1 ; t.. tjt j.Jtt L1 J..J
± : xJ .t- i J i i L 44 U: ’. iLt ! I Ij_ -
lHi’
. 14 4.4 4 • 4 44. . . . — — _ . . . • - 4 1
I i 41 4 • 1 _i . .i I .4 .i —4.-I— I.—.. 4 . 4 .3 1,. 4 • • I . • —
—. 4.. - I 14. . 4 .i 4 :J .44 i i . 1li _. 41 . ._4 _4_I.4_ 4 I_i 1.4.. .41 • I -
2J _L . iL444 J± : ::, 1 J : :
- —- iji P i i
A_ I i LL .J • 11 - ç •_I L 1 s i i - . .. ± LJ • .. 41I. 4. LLLLJ :tLL!:
I ! IJI H 4 - i 1
J I J ’ rr 1 1 )i 1 tl II J j} _J J t
-j 1 itTh i it l’I itLLII J JJLL
:; L 1 i Lii ’ i J it 11 .1 IF lI’ 1
Hi 1! i L iL!i1 J1 U LJii JLLJLJ± LI’
1 — .4 144 . .,, .;.. • .4 4- . -4.4. 4 _+lJ44444 . jI . 4 — — i—T I —4 — 4 .i .4 ..4 - 1- 3 4. 1- - i • .
4 —— 4.4 •4 •,43 . i . Ł..._, ._44• .I._ 1_.4 f I - . 4_4_l .4_ —I— -4 -—-- .4 14 —.4—4-Jt —.. l .—’ I’l.. - 44 . 4 4—
- .. - I - ii •i i i . L 4 • • I - —I — - -- - 4 — - 44 -. - - 4-’- •“ r •
LL.
I . i - 1 [ t 114 ‘ iti 4 Tii ’1:,I . -L± iL IL
I- J i- f 1 :i t; 1 ± : hJ4 L i J ii j 1 i : : - 1: -
- ‘iJ_I ILJ: J: i b. rH ! -L.U..
. 4-4fl1 .001 .0
.,o .9 .97 .91
.9 1 40 •c c-.:
collected at Beltsville.
-------
—48-
Figure 4. Probability for Given Daily UV Total Energy per nm for 5 nm
Band at 305 nm. Coefficient of Variation = 0.3545
June (+) and July (x) 1976, Rockville, Maryland !‘
1J. iLjJJ H U iJ 11111
, ‘)I..
J j,
- L i± LL1
— It:;
_TL iIT. 1 . . ::: ; -‘1
4 4. J
‘ ‘ H t
1: H E’:ii i .
L-—-
-H---t
L 4
± ±
—4.4.4
H-r
.t 4 J
±!
ji j4
t ’ ’
ii t
ithL ii
‘444 .4
L14 , 14
• , i hI;
11t
,; -i
. ‘ I _4_ : ‘tI ;t i -1: -
1
t J-id i t I ’
-d— •-ii. .
. 4 -i . . ,4. , 1L4.
:
:2 1 L i f t i1 : ‘ : t:
____
-— -- - { 4 1 LA . 4 . - - .1 ‘‘u-
i 4
2
.=.L—Tf LJWJJi 1 :iT.
1i : : ;: ,
.
: ::i;
. 1.4.4 - .. - . . 4 . . —.
4 .. 4--.
4i -. ’i 1 ; H ;lt
14.4 , .4 .J, 4 4.,
4 4X- 4 JI 4 I ’I 1
— J A 4 -4- --41
t 1 :L 1 : nI i jH -
{ .4 - 1- - 1 -
1 iii!±L i .1jLt
:: ,;jj i 4j
$ i $-tj
i i
F-1ft J 1 t . i: I i I
>. .- ‘— ‘ —‘- 4 -f--i- I -4
ij iixT . 1 tt: h
- 1L. , Ł
- . . 1_1_L_ .L. . _ 4 — —
4
±. ± i t ‘4
Cumulative Probability
1/ Based on Smithsonian data modified with data in literature and data
collected at Beltsville.
4.1 .
r
- —--- - - : —
± j !
.4 . i4 4141 .j..
.4- .- ’ ! .4 4 .. 44 p• ,
j ii _ J
: :
4l i 1j -
tt . L L. .
:: 1t4 J :
1 ... 4 11.-I — ‘ — . 1 + _-1 1—
. 1.1 .4.. — . 1
- 1 ——4H - i- I--f-.- 1-4
,L4 4 —4
44111.... ‘4.4.,
;: ‘ jf f
—4-
—
If I l l
11
H
__tj ;p
1]
1 ;:: ;l 41 i— 1 1t1 1 . 4
44.4
4 . l . . . ‘.4 ‘ .l ’-4
4j
-.4- l .i 1l $ -t —
-i -1— .4 ...4-,1. , . .4 . .4 . ..1. 44 44444
• - 4 441 • - 1 . 4. 1 -,
± 1i I11ifl’ ±1-i1 iii4i i J:
F1j ±I jjb jJ jjiJ jfl-
—4-- 44-4-- • 1 - 4_ 4 - . 4W’ -
: jt ’
44 - ;Y4I jI
± tiii L±j ,i±EI
Hjkt 1 jft 4- 4-ti
. ji
L J1 - 4 _I J 1 ,H’
ui .i1’I ;’ittJ ”1 l1i I
d 11 i : 1II ] 1. i
i _ ’ ’ ‘i- -
4--f- .4 i 4-- - 1
n f
J
t 1,
: i -J’Ei
‘4-. I —4-— -
JJJ J
Ill
-
. fiij. . H -
114:1: IiIH
4 1_ ifli4fl - i
ftt 1 1
i±_-Hf i 44
4-1 1
l i
1
-44
it
4 - 4-.-
H-i -
—4 ,4_’ .1
41
ljj 1 sZ
144. 1.441:
LL
.4414.
1 ii
4411
.4 1.14 .
. 444 1-4 , . . 4+
4-4— rH4- - 1 1
4- -:
-4---
-J. :1: I
1it4IhJ -f . 4, - 1 4 — t 14--. — —-11 1111
. .44-41 . ‘ 4 -4 -_ - —4 1 .4 -4 -14 4 . .- ._ fl4.l
-if 4 jjj, 4.-4- I ±441 - i4 - -4J -Ij 1 4 -H--- J - H ’-
i fl i
t ? i t 1J: 1:1 1: - 1
ps Iqk . 3
0001 .001 .01 .05 .10 .20 .30 .40 .50 1.00 .70 .BD .90 .95 . 7 .98
I Ill -lIftil 4 Ii
r ti :4111 L : . .L: ’
j $4 L- + 4- ‘ - ---- ---1---
•f l4tkjJ iJ. l 4j _
iifflillIlli 1 - [ -i- i .
.990 .995 •99
-------
—49—
Figure 5. Probability for Given Daily 1W Total Energy per nm for 5 nm
Band at 315 rim. Coefficient of Variation 0.3188
June (+) and July Cx) 1976, Rockville, Maryland !‘
Cumulative Probability
!/ Based on Smithsonian data modified with data in literature and data
collected at Beltsville.
J11I T Tfj
1 4 i ttT i j1ij 1t 1 i U ’U I L
E ”
i-
—
1:
1 1 I J1
t • 1It t - t- i- ji n 1
ji ! ! t 1 J ‘
tIE f_L ±rli t
1
1 1UAiii ifl3ii i 4} jj4
11! t1 I
II I JII II
Ill
I .1 1 V i I J I Th 1 j1JJ
ii iJ ij 1 ttii i j F
—H i tll ftt j j . _1__ -
Lf i IffliJf
qi JT J}4. ;
- 4 H L
I i1 ll I
1i U111 tHI
1L 4 ± 4H4
4 t
fj
1 i -
1i1T L L
ji j I
IT -
Ii
T .
hr
J .LIL. Ht
I Th
4I
fi jI
-f
4.1.1
JjA 14 tL
j4 1
1 i44
L i
jjI iI I i -I -
11 . 11 f :114 if i J 4,I ‘l 1— 4 i
I i 4i i i t
1:
I
j f
ill
iii
I 44J
IiiJII1
- 4 H
j I
4 ‘i
4. .4
I -fl
itni
i - it
-1; - 1
1-
i
E f,g [ ftl
1-4
t
- 4-1-
1-
1
—1
11111
Ii
jI I T
1 E - M fl1 . 1fl W-m W I
______
MO lI
.0007 .001 .01 .OS .10 . 0 .30 .40 .50 .70
li41
j h
I -i±i1 i It
v - -- It_1ii t
I
4—
. .90 .?S .97 .91 .990 .995 .9
-------
—50-
Figure 6. Probability for Given Daily UV Total Energy per nm for 5 nm
Band at 320 nm. Coefficient of Variation 0.3009
June (+) and July (x) 1976, Rockville, Maryland
Cumulative Probability
!/ Based on Smithsonian data modified with data in literature and data
collected at Beltsville.
I
1
I
.31
w
LILI 4t
- -
t
I
ThI fflll
J j t ;IjLU j I
- : i . 41 4
T 1i
I .
I I
. . .±.
::1: L IJ iili fl tW ffl liii . ill
: JJp JJ J
: J1jT ±i i1jiJ; LiLf f II I 4__I 4t i i i jt
— 4 tj iJji
+b1 1L t 1 ; L N 1 I I
1 ll ui I
L*1IU. L :ith
3
I 1’ i ll]ft II : T
: llI J 1i .i i 11 J I II t11 111111 I ±IIltt I I 1 lilt :
“ ill t ffl 1 1111 :
iii ‘r i
111’ j 1 f
1 1 . t 4 44_ .
L1 i 1-J 011 J.4 j fl9
i1
—_ - — 4 t j J li li 4 - 4 J : - t 4
—:!4 p I1 ]lh tn I [ i ’ill1 fl JA .i
K
.0001 .001 .01 .OS .10 .20 .30 M SO r 60 °
.90
.9S .9? .91
.990 .,n . 9i
-------
—
F,igure 7.
9Z3
8 0
700
Average iiv Solar Spectr 1 lrred , ric . Juoc t nd July, 1976,
6 hour day.
E
U i
L)
‘—1
5 E;
400
100
0
250 300 352
I (“
7lJz)
%L\! rp’rTu
iU1 .LLI1uLFI fl
-------
—52—
Figure 8.
Reflector for I S V Sunlamps.
STOCK: 8 inch wide x 47 inch long to fit between lampholders
0.032 inch thick specular alumirn
ioa
*1
-------
—53—
Figure 9. Normalized Irradiance 0.82 meter under a typical 1JV—B Lamp Array
L FSLIO 1 .I1M LONG 6O 12O CM.
0.82 M F19OM LAMP SUf9FRCE
I •I 1 I I I 1 —I
a oz o.o 0.02 0.02 0.02 0.02 0.02 0.02 0.02 0.02 0.01 0.01 0.01 0.01 0.01 oioi
4 + ± ± + + + + + + + + + +
O D2 0.05 0.03 0.03 0.OS 0.03 0.02 0.02 0.02 0.02 0.02 0.01 0.01 0.01 0.01 0.01
1 + + + + + + + + + + + + + +
OJOS 0.03 0.05 0.05 0.03 0.05 0.05 0.03 0.05 0.02 0.02 0.02 0.01 0.01 001 0.01
+ + + ± + + + + + + + + + +
0 04 0.04 0.04 0.04 0.0 11 0.04 004 0.03 0.03 0.03 0.02 0.02 0.02 0.01 0.01 (LOl
+ 4- + + + + + + + + + + + +
0 oc a.os o.os a.cs 0.05 OOS 0.05 0.04 0.04 0.03 U.03 0.02 0.02 0.02 0.01 0.01
+ ± + ± + + + ± + + + + + + -
U 6.07 0.07 0.08 0.07 0.07 0.05 0.05 0.05 0.04 0.03 6.03 0.02 0.02 0.01 U. 01
4 + + + + + + + + + +
L.) oJo .10 0.10 0.11 0.10 OI Q 08 0.07 0.05 0.05 0.04 0.03 0.05 0.02 0.02 0.01
+ + ± + + + k ± + ± + + + +
D 0 12 0.13 0.15 0.15 0.15 0.13 0.12 0.1 0.00 0.05 0.05 0.04 0.05 0.02 0.02 0.01
0 0 20 50 20.20t7 0 \0 .08 0 5 0 4 0 3 Q 3 0 2 0. 2
I + ± + + ± + + + +
(•_) C. 0.30 . . . 0.30 25 0.}’Q 0.14 0 .1 1 0.07 0.05 0.04 0.03 0.02 ii 02
+ +\+ +\± + + + +
0 7 .47 . .55 . 0. 0. 7 26 0 18 0.12 .08 0.05 6.04 0.03 0.02 002
• 0.91 0 7 0 \O.1; ::: ::: ::: ::: ::: : :
0_B _52 0.96 1.01 0.95iU. a 0. 2 0. 2 .27 L17 0.1 6.07 0.05 0.03 0.02 0.02
- + + 1 H 4 + \+ + + + + +
0.5 0 82 0.95 1.00 0.9S 0. 2 D. 0 .4 0 2 ti .ID 0.12 0.08 6.05 0.04 0.03 0J02
- + + +f + + + + + + + -I -
0 6 0 aa 0.95 1.00 0.95 0. 2 . 4 0 30 0 .19 0.12 0.00 0.05 0.04 0.03 (1. 102
Iu ,I 1
- ± ?h. + 1 -H +1? 1+ + + + + +
002 0.95 .0 0.95 0. 2 0. 3c .1 c _29 1.18 0.1 0.08 0.05 0.04 0.05 0.02
± + I1i+I± + + + ± +
0.62 0. 2 .96 1.01 0.9 0 2 C.52 0. 2 .27 0.17 0. 0.07 0.05 0.05 0.02 0.02
6.87 0.91 Jj0 2/0 5 0 0 0 7 0 5 0 3 0 0242
0.49 0.65 0.76 0.79 0.75 0.55 0.49 0.34 0.22 0.14 0.10 0.05 0.05 0.03 0.02 0.02
)( IN o.a INC9. EJ9OM CENTE19
-------
Figure 10. Typical UV—B L imp Array.
—54—
-------
_55 -
Y 2
LI
Figure 11. Typical UV—B Lamp Array in use .
4 . - - — - 1 --- - 1 flj 1 t 4 A :;
(
r
‘H
— . ., . . : ; : ‘ 4r$ I ,
1 <
*4 1 1
).“ t /41
k
...i -( 1 r ee)ve / s
strb —t - CT; N
s’ ajw . , 4 ,
.1’ . .
4
115 I
.
I
/ .,
I ..
a-
)
I
/ .1
V
a
II /
‘ I
I
C
“ -S . ,
y I
7
-------
—56—
Figure 12.
2C0
iez
Spectral UV power output of one representative PS4O lamp.
I C 3
7 Z
—4
E
C
*
E
5ZZ
3 Z
0
L.J
r.
350
AVELE? TH
-------
—57-.
Figure 13. Spectral UV power output (logarithmic) of one representative FS4O lamp.
I I
-- ----‘- _ __J_________ --
-H — — -- -•
1.
I I
I I
— —— —
IL TI
1
-—-i - - l
I j
- _ •_i
I
I 1
! I
- i___i_ - - -_I—-i
•• HL - - -IH
p Iri ri •r’l, I
J1t LU ,Ifl n
1 z. 1 —. -- - - .-. - - - --. - - - .
FS4B
!
ia
MCD.
25
3 3Z 350
i g
-------
Figure 14.
—58--
Spectral UV power output of one repreaentative FBZS4O—CLC latrp.
-- --i-
FBZS Ř — CLG
1’ T •
1
_-j
— • _i . . — .
L .
H T
-
-. -,-- 1------
1 g ř
BgZ — -
7g0
E
C
41Z
3i
2 Z
1 Z
- .--t
0
25g
-------
-.59- .
Figure 15. Spectral IJV power output (logarithmic) of one representative
FBZS4O—CIA lamp.
- --.. ---——
FBZS4 —CLG
!1 1
AVELEt GTH nn
ia
—I
E
3 3z
35
‘C
-f
-------
—60—
Figure 16. Spectr i1 l i v power output of one r pre entative F8T5 RPR 3000A° lamp.
—4
*
AVELE CTH n
TIJ S
35Z
-------
Figure 17.
1 1. 7 Z
ir r
&
, 1q1.
—61—
Spcctral UI. ’ power output Clc,garithinic) of one repreaentative F8TS
RPR 3000A°
0 rr’
1). ) it5
AVELENCT n
-------
Figure 18.
—62—
TJV power Output of one 40—watt (F40) lau p compared to Average Solar
Irrad lance.
9zgI
8 Z
— -
7g
6Z0
5gz
4g
3Z
2 Z
FS .1 0
FBZS4 -CLC
I
1 E3
a
L .)
1.
r z
-------
Figure 19.
—63—
UV power output (logarithuiic) of one 40—watt (F40) lamp coi pared to
Average Solar Spectral Irradiance.
AVG. SU
1 z. 3Z
i Z.C7
1ř.řřk
uigc
I
35
I ( •
YAVELE CTH r
-------
Figure 20. Weighting Functions
—64—
2.Z
1.8
1 6
1. 4
1 ,2
413
— EThO JiS_ —— —
G EE RCE_AS__
0.6
0.2
35
AVELEt GTU
-------
—65—
Weighting Functions (1og rithmic).
1z z, zř - —
1 Z. CZC
ia za
g cz1
25
2L --
sErLo 1.5
r’ ’rtI I, —
L LI P. _______
Figure 21.
35
AVELD CTH ri i
-------
—66—
Figure 22. Relative Responoc of ISV—B IRLileter Radiometer.
>-
I-
>
U)
wr
C l )
PT-R403 - 0.0625 TEFLON
DIS C
.01
I
220
L
240
9
260 280 300
9
1_ __
Xnrn
-------
—67—
APPENDIX I
METHOD OF PREDICTING IRRADIANCE
Figure 9 displays a portion of the computed normalized irradiance at
a particular vertical distance from a Lamp array similar to that in
Figures 10 and 11, with the reflector cross—section in Figure 8. Four
fixtures, each containing two 40—watt (F40) fluorescent lamps were spaced
60 cm between the middle two fixtures and 120 cm between the outer,
lower, fixtures. Effective emitting length of the nominally 4—ft. long
lamp and lampholders is 1.11 meter. Irradiance was calculated on a plane
82 cm from the center of the lamp bulbs which is 80 cm from their lower
surface. The normalized irradiance is shown at grid intersections 0.2m
or 20 cm apart both parallel to the lamps (and x—axis), and at right
angler in the horizontal plane. There are four similar quadrants.
Expected spectral radiant power from each of several lamp and filter
combinations appears in Table 2, and weighted power in Table 5. Both
tables list the entire output of a single lamp.
For calculation of irradiance, multiply the output of one lamp and
filter by the normalLzed irradiance. The 8 lanips in the array are
expected to deliver the toLal output of one lamp per square meter near
the center of this plane. At x = 1.Om, y o, 0.73 of one lamp output
per square meter is expected. At x l.Om, y l.4m, 0.62 lamp output
per square meter is calculated. The npectral radiant power expected from
an unfiltered FS4O at 295 nm is 55 mW/nm (Table 2. column 6). The
-------
—68—
expected spectral irradiance at this last location ,nd wavelength is then
0.62 (55 mW.nm 1 ) = 34 mW.in nm’. The expected UV-B output
of an FS4O is 2.74 W (Table 5, column 5). The IJV--B irradiarice at x =
l.Om, y = l.4m is thus 0.62rn 2 (2.74W) = 1.7 The weighted
power output of an FS4O using Ac9 is 43 times fle1t viule Control
Sunlight on one square meter. If all the output. of one FS4O were used,
the lamp cou]d substitute, in UV biological effe tivene s, for 43m 2 of
average suimner sunshine. At this (x, y, z) location, 0.62 (43) 27
Beltsville Control Sunlight are expected. Similar calculations for an
FS4O filtered with 0.05 inch thick cellulose acctatc yield:
290nm (Table 2, Column 4)
0.62 m 2 (8.24 mW.nm l) 5.1 mW.m 2 .nm
UV—B (Table 5, Column 3)
0.62 m 2 (1.34 w) = 0.83 W.m 2
Ae9 (Table 5, Column 3)
0.62 (4.2 ni 2 ) 2.6 Beltsville Control Sunlight
-------
—69—
APPENDIX II
CONVERTING WEIGHTED UNITS, KNOWN RELATIVE SPECTRAL POWER DISTRIBUTION
Lamp ratings (Tables 3, 5, lines 2 thru 6, and 13) are used when
calculating irradiance or weighted irradiance under an array of lamps.
The lamp rating multiplies the calculated number of lamp outputs per
square meter. The ueighted irradiance is the expected multiple of
effective ultraviolet, Beltsville Control Sunlight, evaluated by the
specified weighting function.
There are other uses for lines 2 thru 6, Table 13. Line 7 is
provided for conversion of UV—B IRLMeter Radiometer readings to any
units. These calibration constants are for the UV—B IRLNeter Radiometer
lO scale. The constant in line 7 is the reading expected if one lamp
output per square meter reaches the UV—B IRLMeter Radiometer. Divide the
scale reading by the calibration constant to obtain the measured
number of lamp outputs per square meter. For example, if power received
from an FS4O lamp filtered with 0.005—inch thick cellulose acetate (FS4O
CM, Column 3) is measured, divide by 34. Then multiply the resulting
number of lamp outputs per square meter by. the lamp rating in the units
desired. For example, if watts UV—B are desired for the FS4O CA5
spectrum, multiply by 1.34W. if the number of lamp outputs per square
meter are of no interest, the two operations can be combined. Multiply
by 1,340/34 = 39 mW/rn 2 (Table 13, last line).
-------
—70—
* * *** * ** **** * **
The lamp ratings (Table 13, lines I thru 6) are used to form the
multipliers. To convert from units calculated in any weighting function,
divide by the tabulated rating for that weighting function and spectrum.
This yields number of lamp outputs. Then, multiply by the rating
tabulated for the same spectrum and for the weighting function with the
desired units. The procedure is the same as that used to convert
radiometer readings to UV—B.
Raw UV—B IRLMeter Radiometer readings are somet]-me6 mistakenly used
for comparison of UV with different spectral power distributions. The
meter spectral response is then the weighting [ unction, and it is
substantia) 1 -Y the R403 spectral response of the phototube, behind a
Teflon diffuser (Figure 22). Real sunhigl)t seldom has the spectral power
distribution of control sunlight, so the UV—B IRLMeter Radiometer should
not be used to measure sunlight.
-------
APPENDIX II I
Conversion of
UV—B IRL Meter Radiometer and Optronics Radiometer
Readings to Absolute mW.m 2 .nm-
Summed for 5—nm Bandwidth from
280 to 320 rim
-------
—73-.
Lamp and Filter
Action Spectra
Ac 9
Ac21
Table 2
WEIGHTED POWER OUTPUT OF ONE LAMP AND FILTER,
RATIO UNITS PER SQUARE METER
2.5
1.3
FS4O
CA 5 mu
4.2
2.0
8.3
2.9
A 9 To convert Beltsville Control Sunlight to weighted mW .m 2 , multiply
Beltsville Control Sunlight by 3.06 mW.m 2 .
A 2l. To convert Beltsvil]e Control Sunlight to weighted mW.m 2 , multiply
Beltsville Control Sunlight by 14.9 mW.if 2 .
Table 3
CONVERSION FACTORS UV—B IRL METER RADIOMETER READINGS TO
BELTSVILLE CONTROL SUNLIGHT
Multiply UV—B IRL Meter 1O 7 -sca1e readings by:
Action Spectra
Ac9 Ac21
Note: UV—B IRL Meter radiometer measurements of UVB enhancement in
the. presence ot.rad:iant energy from other surces should be made
in the absence of other sources or their contribution must be
measured and subtracted from the readings with all radiant
sources present.
F S40
CA 10 mu
BZS4O-
CLC
No filter
FS4O
No filter
Beltsville Control Sun ]
43
12
FS4O, 10 mil, CA, 6h
FS4O, 5 mu, CA, 6h
BZS4O—CLG, no filter
FS4O, no filter
0.086
0.123
0.271
0.628
0.046
0.057
0.096
O. 172
-------
—74-.
Table 4
CONVERSION FACTORS Ac21 BELTSVILLE CONTROL SUNLIGHT
TO A 9 BELTSVILLE CO FROL SUNLIGHT
FS4O FS4O BZS4O— FS4O
CLC
• p and Filter CA 10 mu CA 5 mu No filter No filter
.ultiply Ac21
c1tsville Control
sunlight by: 1.89 2.16 2.84 3.66
Table 5
CONVERSIONS: WEIGHTED mW.tn 2 Ac21 TO
WEIGHTED mW.m 2 Ac9
Lamp and Filter Ac21 Ac9
FS4O, 10 nil, CA, 6h 7.56(mW.m 2 ) = mW.m 2
19.5
FS4O, 5 nil, CA, 6h 13.O(mW.m 2 ) =
29.2
S4O—CLC, no filter 25.3(mW.nr 2 ) = mW.m 2
43.4
FS4O, no filter 132(mW.rn 2 ) = mW.m 2
176
EQUIVALENT POWER TERMS -
MW.in 3
w. m 2 . um
mW.m 2 . nm
11W.cm 2 . (lOnmY 1
erg. sec . cm 2 . nm
-To convert the above to W.if 2 .nm -. multiply by i0 3
-------
—75—
EXAMPLES
I. Reconstruction of absolute spectral radiant power distribution from
UV—B lRLMeter Radiometer or Optronic Radiometer readings mW.m 2 .nin ).
Action Spectrum: Ac9
Lamp and Filters: FS4O, 10 ml, CA, 6 h
Readings
Optronics Radiomeier
0.84
UV—B lRLMeter
Radiometer
(iO— scale)
49
Radiometer BeltsvilJe Control Sunlight
Optronics 0.84 x 5!” 4.2
UV—B IRLNeter 49 x 0.086. i 4.2
Obtain: Beltsville Control Sunlight of one lamp per
square meter from Table 2 to obtain ratio
Ratio: 4.2/2.5 1.68
1/ Optronics calibrated to full scale (1.0) equal to 5 Beltsville Control
Sunlight, for spectrum FS4O 10 mu, CA, 6h.
2/ From Table 3
-------
—76—
To reconstruct absolute radiant power distribution
multiply the spectral radiant power for one FS 40,
10 mu CA 6h filtered lariip per meter square (Table 1)
at 5nm intervals by the ratio as follows:
Power 0utput ! Unweighted
1ength (mW.m 2 .nm -) Ratio (mW.r& 2 nm 1 ) (nm) (mW.n 2 )
5nm bandwidt i
55.4 x 1.68 = 93 x 2.5 = 233 I
58.2 1.68 98 x 5 489
55.7 1.68 94 x 5 468
46.5 1.68 78 x 5 131
24.6 1.68 41 x 5 207
:95 1.79 1.68 3 x 5 15
:90 5.4 x l0 x 1.68 Ca 0 x 2.5 O i
:so 0
:75 0
:70 0
E 290—320 1411
Hg 1ines J
mW.m 2
313
21.9
1.68
37
302.2
1.65
1.68
3
296.7
0.33
1.68
1
289.4
Ca 0
0
253.7
0
0
E
290—320 1451
From Table 1
Beginning and ending net should be 1/2 bandwidth for summation
Hg lines are not per nanometer. Their sum is added directly to the
last column.
-------
—77-.
Reconstruction of absolute spectral radurni power from IJV—B
IRLNeter Radiometer readings (.m 2 .rn 1 )
Action Spectrum: AE21
Lamp and Filter: FS4O, no filter
Read ings
Optronic Radiometer
UV—B IRL Meter Radiometer
(io— scaJe)
Not Used
60
Radioineter
Optronics
Beltsville
Control
Sunlight
Not used
UV—B IRL Meter
60 x 0.172 1/
10
Obtain: Beltsville Control Sunlight of one lamp per square
meter from Teble 2 to obtain ratio
Ratio: 10/12.0 = 0.83
1/ From Table 3
-------
‘ro recon ;tr c io!u&e o e t dj L jbUtiOfl,
lTJUltIpIy the r ,o er for t,ni F540
no fi ter r I) at nm
1flterv 1a by e
Z 265--320
2243
Hg lines i
(mW.m 2 )
313
302.2
296.7
289.4
253.7
50 x
5
3.8
1.2
1.2 x
0.83
0.83
0.83
0 B3
0.83
.
42
4
3
1
1
E
265—320
2294
From Table
1
2/
3/ Beginning and ending nmshou!d be 1/2 bimdwidth for ;uinm tion
4/ Hg lines are not per nanometer. Their suez is added direetly to the
last column.
Wavelength
Power Unweighted
(mW.tz 2 .n .i) ( nm’ 1 ) (mn) mW.m
5 tim band . jd
320 -
.5
195 !
315
104
i
ij
5.0
432
310
105
0.83
g
5.0
436
305
5.0
403
300
78
o . s
6
5.0
324
295
55
o .
5.0
222
290
32
0
27
5.0
133
285
15
0.83
12
5.0
62
280
5.7
0.83
5
5.0
25
275
1.5
o..
1.
5.0
5
270
265
0.2
Ca 0
x
0.83
0J 3
0.2
x
5.0
2.5
1
Ca
-------
Si e F ”. ‘ /4 . /3
FINAL REPORT
INSTRUMENTATION FOR MEASURING IRRADIANCE IN THE UV—B REGION
J. D. Rowan
K. H. Norris
Instrumentation Research Laboratory
Agricultural Marketing Research Institute
Beltsville Agricultural Research Center
Beltsville, Maryland 20705
EPA—IAG--D6—Ol 68
Project Officer:
R. 3. McCracken
Agricultural Research, Science and Education Administration
U.S. Department of Agriculture
Washington, D.C. 20250
Prepared for
Environmental Protection Agency
BACER Program
Washington, D.C. 20460
-------
Annual Report 1977
EPA Interagency Program on Biological and Cliiratic Effects (BACER)
Instrumentation for easuring Irrathance in the UV—B Region
SUMMARY
The responsibility of the Instrumentation Research Laboratory (IRL) in
this program was developing portable instruments for use by biologists to measure
LJV-B irradiance in growth chambers, greenhouses, and field plots. A simple
I N—B radiometer and two UV-B spectroradiometers have been designed, constructed,
tested, and put into use in the (N—B research program. Each of these instru-
ments is now being manufactured by commercial firms.
The two spectroradioneters differ only in the monochromators: one has a
single holographic grating; the other, two holographic gratings for greater
stray light rejection. The spectroradiometers automatically scan the 250-400 nm
region in less than 5 minutes, printing a tape of the corrected irradiance as
a function of wavelength. The input is cosine corrected by a specially designed
teflon—bubble diffuser coupled to the input slits. The output of the menochro—
mator is measured with a multiplier-type phototube and a logarithmic response
amplifier. The amplifier output is digitized with a digital voltmeter, and the
digital output is interfaced with a desk-top programmable calculator.
The design provided for the spectroradiometer system response to be stored
in the calculator so that, as the spectrum is scanned, the calculator corrects
the measured signal for instrument calibration and outputs the true spectral
irradiance of the source being measured.
The programmable calculator controls the operation of the spectroradi—
ometer so that, on command, scanning is initiated and readings are recorded for
each nanometer interval. The calculator prints the wavelength and irradiance
for each wavelength interval and, at the end of the scan, reverses the wave-
length drive and returns the monochrontator to the starting wavelength. The
calculator is programmed to print an integral for a programmed action spectrum
at the end of each scan. At the completion of the scan, the data can be stored
on a magnetic tape, if desired, for future analyses.
Provision is included for a precise check of wavelength: the spectrum
of a miniature low-pressure mercury—arc lamp is scanned, and the calculator
computes the position of the 253.7 nm and the 296.7 nm lines to a precision of
+ 0.01 nni.
The performance specifications approach the requirements for TJV-B measure-
ments stated in NBS publication #20, “Optical Radiation News,” dated April 1977.
iii
-------
Annual Report 1977
EPA Interagency Program on Biological and Climatic Effects (BACER)
Instrumentation for Measuring Irradiance in the UV-B Region
Our UV—B measuring instruments developed for this project meet the
requested requirements. The broadband radiometer is small, hand—held,
battery operated, and has fast response for rapid measurements. The spectro-
radiometers have the following features:
(1) Fast, accurate, and reproducible.
(2) Convenient, automatic single—key—stroke operation.
(3) Programmable calculator-controlled scanning, Fig. 1.
(4) Logarith nic amplifier for wide dynamic range.
(5) Calculates and prints true spectral irradiance for each nm
wavelength, Fig. 2.
(6) Calculates and prints weighting function (A :9).
(7) Programmed—wavelength—calibration check.
(8) Double monochromator unit has very low stray light.
(9) Bandwidth of 2 nm.
(10) Temperature stable.
(11) Receptor has excellent cosine response.
(12) Spectra are stored on magnetic tape for efficient data processing.
(13) Portable-—operates on small lab cart.
(14) Minor disturbance of area by the “measuring head.”
Our spectroradiometer performance specifications approach the requirements
for UV-B measurements stated in NBS publication #20, “Optical Radiation News,”
dated April 1977. These requirements are cited in attached copy of “Making a
UV Measurement?,” Electro—Optical Systems Design, 9(6):17, 1977. (page 7)
Cosine Response for tJV-B
The spectroradiometers, as well as other UV-B radiation meters developed
in our program, incorporate a new design for cosine—corrected input optics.
We tested all available cosine—correction schemes, including integrating
‘Spheres, diffusing reflectors, and sintered—quartz diffusers. None of these
gave adequate performance for the 250 to 370 nm region, so we developed the
1
-------
teflon bubble diffuser. The spectral transmission of this teflon diffuser
is shown in Fig. 3. Fig. 4 is a drawing of the input optics for the spectro—
radiometers. We supplied a similar receptor to our cooperators that have a
Gamma Scientific monochromator, Fig. 5. The response of this type of diffuser
to radiation from different angles is compared with that of a commercial
instrument in Fig. 6. The teflon bubble receptor provides excellent cosine
correction, and has a stable surface which can be readily cleaned. This
type of diffuser is now used in commercial instruments.
Wavelength Accuracy
To check spectroradiometer wavelength accuracy, a low-pressure mercury—
arc lamp is scanned and the mercury line location computed. The wavelengths
of the 253.7 and the 296.7 nm lines are measured with a readout precision
of + 0.01 nm, so that wavelength shifts as low as 0.02 nm are readily detected,
Fig. 7. This field check can be made quickly and routinely between measurements
if desired.
Spe ctroradiome te rs
Our UV—B spectroradiometers are now used routinely by technicians to
measure UV-B irradiance in growth chambers, greenhouses, and field plots.
These instruments are ...dentified by the acronyms: IRLSpec-S for the single
monochromator version; LRLSpec-D for the double monochromator; and IRLSpec-SO
for the commercial model single monochromator. Numbers shown on the graphs——
for example, 17708l5.09——id ntify instrument (first digit) , year (next two
digits), month, day, and scan number for that day. The first digit may be
1,, 2, or 3 for IRLSpec—S, IRLSpec-D, or I1 LSpec-SO, respectively.
Measured performance specifications for the IRLSpec-S spectroradiometer
are as follows:
Wavelength range — 250 to 370 nm
Readout interval - 1 nm
Scanning speed — 0.5 nm/sec.
Spectral bandpass — 2 nm
Wavelength reproducibility — + 0.02 nm
Wavelength accuracy — + 0.05 rim at 296.7 nm
Radiometric reproducibility — ± 2%
Radiometrie accuracy - ± 5%
Radiometric range — 0.001 to 2000 mWm 2 nm
Stray light — less than 2 x l0 at 285 mu as tested
with a xenon arc lamp filtered with 0.5—mm-thick
cellulose acetate
Cosine correction — within + 10%
Output — printed tape direct reading in wavelength
and corrected irradiance at each nanometer interval
from 250 to 370 rim
Size of measuring head — less than 28 x 20 x 10 cm
Temperature stability less than 0.04 rim wavelength shift
for 25°C temperature change.
2
-------
The IRLSpec—D spectrorad3ometer, with double-grating monochromator,
provi 5 better stray light rejection. This unit incorporates all the
(eatur of the single moriochrornator unit although the measuring head is,
of necessity, slightly larger. Observed wavelength instability of the
double monoChrornator was caused by grating-syncdrive cable temperature
sensiti Y. Our technician, George Button, solved this temperature insta-
bility with his cable—stringing technique. The grating-drive cable spring
as eliminated, and the cables were crossed to provide automatic temperature
compensation and wavelength stability.
Measured performance specifications for the TRLSpec—D spectroradiometer
are as follows:
Wavelength range — 250 to 370 run
Readout interval - 1 nm
Scanning speed - 0.7 nm/sec.
Spectral bandpass — 2 rim
Wavelength reproducibility - + 0.02 nm
Wavelength accuracy — + 0.05 nm at 296.7 nm
Radiometric reproducibility - + 2%
Rad ometric accuracy - ±
Radiometric range 0.0001 to 1000 mW m 2 nm 1
Stray light — less than 5 x io—8 at 285 nni as tested
with a xenon arc lamp filtered with 0.5—mm—thick
cellulose acetate
Cosine correction — within + 10%
Output — printed tape direct reading in wavelength and
corrected irradiance at each nanometer interval from
250 to 370 nm
Size of measuring head — less than 28 x 25 x 10 cm
Temperature stability - less than 0.06 nm shift for
25°C temperature change.
The IRLSpec-D spectroradiometer was field tested at the University of
Colorado’s Snowmass, Colorado, high—altitude site during August. The instru-
ment was disassembled into components and securely packed into a regular
hard-side luggage as a shipping container and transported as ‘checked’ baggage
to and from Colorado. Spectral measurements were made at three elevations,
2377 m (7800 ft.), 2980 m (9777 ft.), and 3452 m (11326 ft.). This necessi-
tated disassembling and repacking the instrument and transporting it up and
down the mountain on unimproved roads to each site. Upon return to Beltsville,
the instrument continued to perform with only minor adjustments.
The commercial version of our IRLSpec-S, single monochromator spectro—
radiometer was manufactured to our specifications. This IRLSpec-S0 required
constant operator assistance, but our circuit modifications converted it to
automatic operation with single—key stroke to scan the spectrum, return, and
reset.
3
-------
Measured performance specifications for this commercial IRLSpec-SO
spectroradiometer are as follows:
Wavelength range — 250 to 370 run
Readout interval — 1 nm
Scanning speed - 1.0 nm/sec.
Spectral bandpass - 2 nm
Wavelength reproducibility - + 0.02 nm
Wavelength accuracy — + 0.05 nm at 296.7 rim
Radioinetric reproducibility — + 2%
Radiometric accuracy - ± -l
Radiometric range - 0.001 to 200 mW•m 2 rim
Stray light — less than 1 x io-- at 285 rim as tested
with a xenon arc lamp filtered with 0.5-mm-thick
cellulose acetate
Cosine correction - within + 10%
Output — printed tape direct reading in wavelength and
corrected irradiance at each nanometer interval from
250 to 370 nm
Size of measuring head - less than 25 x 21 x 13 cm
Temperature stability — less than 0.04 cm shift for
25°C temperature change.
Spectroradiometer Sensitivity and Dark Current
The photomultiplier—tube dark current, system correction, and standard—
lamp spectral irradiance are plotted in Fig. 8 for the IRLSpec—S with
Hamamatsu R166 phototube (solar blind response) and Corion SF-l.25 solar
blind filter. The IRLSpec—D photomultiplier dark current (Hamamatsu R212, S—S
response and Corion SF—l.25 solar blind filter), system correction, and
standard lamp spectral irradiance are plotted in Fig. 9. Plots of the IRLSpec-
SO photomultiplier-tube dark current (Haniamatsu R166, solar blind response,
and Corning 7—54 filter), system correction, and standard lamp spectral irradi-
ance are shown in Fig. 10.
The instruments developed in our program are now manufactured. A broad-
band UV—B radiometer is available from Optronic Labs, Inc. Spectroradiometers
similar to ours are advertised in Optical Spectra, l2(l):l3,5l, and 60, dated
January 1978, and are available from three firms:
EG&G, Inc.
35 Congress Street
Salem, Massachusetts 01970
Gamma Scientific, Inc.
3777 Ruffin Road
San Diego, California 92123
Optronic Labs, Inc.
7676 Fenton Street
Silver Spring, Maryland 20910
Irradiance Spectra
Typical spectra from our IRLSpec-D are plotted in Fig. 11, 12, 13, and 14;
from our IRLSpecS, Fig. 15 and 16; and from our IRLSpec—S0, Fig. 17.
4
-------
Irradiances of FS4O and FBZS4O lamps are compared in Fig. 11. A typical
curve for an FS4O lamp with and without a cellulose acetate (CA) filter is
shown in Fig. 12. CA filters with FS4O lamps are compared in Fig. 13. Measured
irradiances of four FS4O lamps used to enhance the sun’s UV-B at Snowmass,
Colorado’s 2980 m (9777 ft.) site, are plotted in Fig. 14. The fixtures were
two lamps mounted in pairs at a Z of 110 cm and filtered with 0.127-mm (0.005-in.)
CA. The change in irradiance of a FI3ZS2O lamp after aging 16.5 hr. is shown
in Fig. 15. The irradiance measurements of two Sylvania F15T8 CW lamps at
Z = 6 cm are compared for the IRLSpec-S, Fig. 16; the commercial IRLSpec-SO,
Fig. 17; and the IRLSpec-D, Fig. 18. Irradiance for a 15—watt daylight lamp
is shown in Fig. 19.
Spectral data of the sun at Beltsville, Maryland, during the year are
plotted in Fig. 20 and 21. The sun curve on a very clear day illustrates the
stray light at wavelengths below 289 nm for the IRLSpec-D, Fig. 22. The action
spectra of measured Beltsville sunshine with weighting function 9 are shown
in the region of 294 to 318 rim at the bottom of Fig. 20.
The weighting function plotted on the graphs is the AE9 equation developed
in cooperative research at BARC:
= [ o.25(A/228.l7e)9 .0] exp [ 4_(A/228.l78)9.O]
The equation was used to compute UV-B sun equivalent (UVSEB), indicated on the
graphs. Details of its development are presented in the Agricultural Equipment
Laboratory report.
Spectral irradiance data are now plotted automatically with the HP9815A
calculator interfaced directly and controlling a HP9872A plotter. After the
data are recorded on the cassette, selected scans are loaded into the calculator;
and the plotter draws and labels the graph to programmed dimensions, and then
automatically plots the data (Fig. 11, 12, 13, and 14). A curve is plotted from
250 nm to 370 rim in 25 seconds.
Narrow-band Radiometers
Twq narrow—band, probe-type radiometers were commercially manufactured to
our specifications. These radiometers have solar-blind photo-multiplier-type
detectors (Mfg. No. R166), a peak response at 294 nm and 299 nm, respectively,
and a half—band width of 5 nm. Irradiance readout is a 3 1/2 digit displa ’
located in the remote electronic unit with switched ranges from l0 to io 1
wattcm 2 nm . The small sensor head (5x5xl6 cm), with teflon dome receptor,
is attached by 2.75-rn cable for easy placement within growth chambers.
Broad-band UV-B Radiometers for Routine Monitoring
A small battery—powered radiometer, the IRLMeter, was designed and con-
structed for measurement of total UV-B radiation from artifical sources. The
instrument has a solar-blind vacuum photo—diode (Hamamatsu R403), an integrated
circuit amplifier, and a microammeter packaged in a meter case. The circuit,
as shown in Fig. 23, provides for tour decades of range switching. The relative
w avelength response, as shown in Fig. 24, peaks at 300 nm and is relatively flat
from 280 to 320 nm and the radiometer is not sensitive to radiation longer than
400 rim. Typical comparative data of the IRLNeter with the IRLSpec—S spectro—
radiometer are shown in Fig. 25.
5
-------
This broad—band UV—L3 radiometer sensitivity is adjusted to a full scale
reading of 2.0 UV-B sun equivalents, Beltsville (UVSEB) when measuring the
output of FS40 fluorescent lamps filtered with 0.127-mm (0.005 in.) CA.
Correction factors were developed for use with FS40 lamps with 0.254—mm
(0.010 in.) CA filter, and for use with BZS fluorescent lamps.
The IRLSun—meter radiometer is a modification of the IRLMeter. The new
calibration reads directly in sun equivalents, Beltsville (UvsEB AE9). Cali-
brations for two suns (tJVSEB A 9) full scale were developed for four sources:
(1) Sunshine for a clear day (6/8/77), 1:00 p.m. EDT
(2) FS4O lamps filtered with 0.127-mm (0.005 in.) CA
(3) FS4O lamps filtered with 0.254—nun (0.010 in.) CA
(4) FBZS4O CLO lamps (unfiltered)
A commercial version of the IRLMeter has been manufactured, the Optronic
Model 725. These Model 725 radiometers were calibrated by IRL for our coopera-
ting laboratories from IRLSpec—D irradiance values for five sun equivalents
full scale, with FS4O source filtered by 0.127-mm (0.005 in.) CA aged 6 hr.
A calibration scale factor was developed for 0.191—mm (0.0075 in.) CA, 0.254-nun
(0.010 in.) CA, 1.52—mm (0.036 in.) petri dish, and FBZS4O and FBZS2O lamps.
An Optronic Model 725 broad-band radiometer was calibrated from IRLSpec-D
irradiance values for 10 UVSEB full scale with a Rayonet F8T5 RPR3000A (8 watt)
lamp source filtered with 0.127-mm (0.005 in.) CA supplied by the Peoria coop-
erators and then aged 6 hr. This Peoria CA filter absorbed less at shorter
wavelength than Beltsville CA, Fig. 26, 27, and 28, significantly changing the
AE9 action integral.
Rayonet FBT5 RPR3000A (8 watt) lamps have strong energy in the 254 nm
region, the energy approaching the peak value at 313 nm (ratio 1.4). The peak
energy of the FS4O at 313 nm is approximately 700 times the energy at 254 nm.
This 254 run energy can be removed with CA filter, but it probably accelerates
the aging of the CA filter. The irradiance of the Rayonet RPR3000A with the
CA filter at 20 cm approximates the energy of the FS4O with CA filter at 50 cm
and 75.1 cm, Fig. 29, 30, 31, and 32.
Eye Protectors
Personal eye protectors were evaluated for their 1W-B attenuation, and
two FS4O lamps at 50 cm from the IRLSpec—D spectroradiorneter receptor were used.
An aperture through black cloth was necessary to prevent significant leakage
between lens arid receptor, even though they were placed as close as possible.
Plots of plastic goggles No. 6, 7, and 8 (dashed line) are at the 0.001 irradiance
level in Fig. 33. No. 5 was a pair of plastic goggles, and No. 2, a plastic
face shield. UV—B blocking of prescription eyeglasses (glass arid plastic) and
sunglasses are shown in Fig. 34.
6
-------
\J ’ L ’
%IILTI ’ S KIVi ’ .R
E ctro OP C Sy t ms Des n
Tb. F ,igiiiee.,ng Magazine of 1 icctro-OptieOl and Laser Technology
June 1977, Vol Q, ko 6
. , SKILNY I
j’ ,blishcr & l’ice President
News
R i I lAki) Ci \ iN(,H A l
WILLIAM S IIuDSl’LTIJ
Asa -rnte Editor
WILLIAM D AsiIMAN
lAoS ESMAIL
( ‘ .reiiiat iofl Manager
Gr ’ F BNIrSIOE
.Ifarketi ng Services
EDITORIAL ADVISORY BOARD
I Buisei, Consulting Physicist
I) Belforte, Area Everett Re8earch Lab
Buzzard, Le ci
kasasent, Carneg ir ’Meiion Univ
Hunter, Naval Research Lab
C Klauminzer, Molcet ran
It Lavin, General Electric
I) Locl
-------
LIST OF FIGURES
Fig. 1. - Spectroradiometer components
Fig. 2 — Typical UV spectra
Fig. 3 — Teflon spectral transmission
Fig. 4 - IRLSpec-S receptor optics
Fig. 5 — IRL receptor optics for gamma
Fig. 6 — Receptor cosine response
Fig. 7 — Hg line wavelength check — IRLSpec—D
Fig. 8 - IRLSpec-S sensitivity and dark current
Fig. 9 — IRLSpec—D sensitivity and dark current
Fig. 10 - IRLSpec-SO sensitivity and dark current
Fig. 11 — Lamp spectra (unfiltered) FS4O, FBZS4OCLG1O76 and weighting
function 9
Fig. 12 — Lamp spectra FS4O and cellulose acetate 6.5 hr. Z = 30 cm,
Z = 50 cm
Fig. 13 — Lamp spectra FS4O and 0.127 mm, 0.191 nun, 0.254 mm cellulose
acetate
Fig. 14 - Lamp spectra, Sno rmass, Colorado
Fig. 15 — Lamp aging - 13ZS20CLG0377
Fig. 16 — Sylvania F15T8 CW lamp spectra - IRLSpec- .S
Fig. 17 — Sylvania F15T8 CW lamp spectra - iRLSpec-S0
Fig. 18 - Sylvania F15T8 CW lamp spectra — IRLSpec-D
Fig. 19 - GE F15TBD lamp spectra — IRLSpec—D
Fig. 20 — Sun spectra — Beltsville, MD
Fig. 21 - Sun spectra — Beltsville, MD - IRLSpec-S
Fig. 22 - Sun - Beltsville, MD - IRLSpec-D
Fig. 23 — UV—B IP.LMeter circuit
Fig. 24 — IRLMeter spectral response
Fig. 25 - Regression IRLSpec-S vs. IRLMeter
8
-------
F8T5 RPR 3000A° spectra cellulose acetate aged 6 hr., Z =
F8T5 RPR 3000A° and sun (Beltsville) spectra
FS4O and Bayonet FBTS RPR 3000A°
FS4O and cellulose acetate aged 0.5 hr., Z = 50 cm
FS4O and cellulose acetate aged 6 hr., Z = 50 cm
— eye protective device
prescription eyeglasses and sunglasses
RPR 3000A 0 spectra new cellulose acetate, Z = 16.3 cm
RPR 3000A° spectra cellulose acetate aged 6 hr., Z =
Fig. 26 — Bayonet F8T5
Fig. 27 — Bayonet F8T5
16.3 cm
Fig. 28 — Bayonet
20 cm
Fig. 29 — Bayonet
Fig. 30 — Spectra
Fig. 31 — Spectra
Fig. 32 — Spectra
Fig. 33 — Spectra
Fig. 34 — Spectra
9
-------
MEASURING HEAD -
EFLON RECEPTOR
I I
1 MOTOR
I I
I I
I I
L PHOTOMETER J
MEASURINC HEAD
-TEFLON RECEPTOR
I I
I MOTOR
I I
I I
L PHOTOMETER
1
3-31-77
UV-B IRLSpec_D
3-31-77
-I
0
UV-B IRLSpec-S
Fig. 1 Spectroradjometer components
-------
FILE —17 3
TVF’E 2
USED 1208
MF I 1208
FS4O :$OCfl7
2780203. 09
250000.0039
251000.0048
252000.0305
253000. 1084
254000. 2177
255000. 1071
255000.0087
257000.0030
258000.0027
259000.0027
t ,L1UUU.UU
261000.0024
262000.0031
263000. 00 33
264000.0095
265000.0389
266000.0499
267000.0438
268000.0432
269000.0552
270 000. 1144
271000. 1838
272000.2473
273000.3516
274000.5030
275000.7620
275001.0823
277001.3 153
278001.5517
279002.1561
280003. 0536
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288014. 3190
289016.9874
290020. 5971
291023. 4209
292025. 7890
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29 4031. 45 99
295035.6576
2960 4 1 . 54 1 8
297044.7439
298047.0842
299049. :556
300052. 5190
301057.2056
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303063. 3780
304065. 5058
305067. 2258
307071.0456
308073. 4446
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310075. 3275
311078. 3118
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329054. 1393
310050. 1182
3 31046. 6747
332044.9549
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335040. :3120
336036.6655
33 70 3 4 . 95 3 3
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339032. 7506
340030.0634
341027. 7591
342026.5048
343025. 9462
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347019.5569
348019.0975
349018. 1677
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356011.0327
357010. 4509
35:3010. 2614
359009. 5725
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35 100:3. 05:35
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363007.5352
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365033.0470
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357012. 3363
358007. 0350
359005. 1119
l i ii. 9 FPOM
250
80. 245::
SUNS = 25.2240
‘3 LI N F F’ 011
TO
c t. ‘
MILL IL’ARTTS MS’:
3 42:3 . 0 34 1
SUM FR’Jt’i
TO
MILL I NAT IS (ISO
117. 1250
‘3 IJ 11 F P ‘J
TO
320
MILL I NAT IS (ISO
1 3 80 . 9 1:: 3
U N F RON
TO
340
MILLINATT’ ‘P150
I C1 47
s t1 ?
250
270
290
320
4
Fig. 2. Typical UV spectra
-------
‘ 2OO
Fig. 3. Teflon spectral transmission
FRONT
SURFACE
MIRROR
Fig. 4. RLSpec-S
RECEPTOR
OPTICS
3-31-77
1.4
0.020” TEFLON DISK
1.2
C,
0
-J
.1
300 400 500
Xnm
600
12
-------
Fi 9 . 5. IRL RECEPTOR OPTICS FOR GAMMA
..020” TEFLON
3-3 177
13
-------
I’
I
.1 %
0.9— I
C
I
a
0.9— 1
a
0I
I
0.7— U
I
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I
O.6
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‘ I i I I
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0.5— 1
w
> ‘ / GAMM-) ’
j04
• I
“TRUE C0SINE- t
I
I CURVE
0 ,
I t
I
II
0.2—
I
8
• I DATA POINTS () ARE
OJ I IRL COSINE RESPONSE
I TEFLON (O.O2O ) BUBBLE RECEPTOR’
VlItt;i
806040200 20406080
ANGLE - DEC.
U- 3-76
Fig. 6. Receptor cosine response
14
-------
i . Ui
2 40O02. 1 4
29 Ci O4 ?. :: :‘9
.4’cVi4 ‘rSS
!]•• •I i• J-4 •
1EFC L’F LI rE
&. H CAL.
!L .1J I j,.-
fl C
-
2 7C 04’. 44d
2*:L1UcH. :-:r4
Fig. 7. uq line wavelenqth check - IRLSpec—D
A F’UU OPE I TF A
rr pr
D PECOEP 0RT
E STOP
F FOPNAFD
O FEI nT [ AIR
I IflT
J 14. L.. CHEC 1
• I I l•
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U. L. CAL.
2 4C02.
: rç : ,, Rn: ::
29T0U- H 510
292fljiCi 015
M c’iiP HUE F
29t. 73
U. L. CAL.
F - 4 ic - ii- R
1 : 1 1:::
9Ci ,: ’ T. 533
I H
IIEF C UP I Li UE
4
U. H CAL.
[ H
2 40UC 3. 40
• •.l•: I Ii •1 —
:.. i;c4n4::
3970C’4’. 453
5CiU3 ‘54
MEFC,JS, LtIE AT
H. L. CAL.
)05fl0: .3 4
I E 0 0 4 4 1
392 03? 115
I1EFCUPI LP;E
fir
i c r ’ I . i’c: H I
H. L. CAL.
ciU UJ. ‘33
3940C10:. 155
4 1
3I ?_IU4 .51
•:
MERC UP’, L t iE A
I i3.
15
-------
1E 2T J T
F -2 flAw RI N J ]
I I I
——H -- -.- —- --— —— — — — •-
..4LJ Li..:4 J
- - -±+-- - \
I I J
I I I’II.
tE17 j__4 L
t L - 1-- - - - -t-
VAVBB Th r IRL I FR 1 OA
Fig. 8. IRLSpec—S senslt.vlty nd ddrk current
IE-01
1Ei4
uy I
16
-------
r
F —
AVELENCTh r IR .. A 14RI FR USCA
Fig. 9. IRLSpec_D sensitivity and dark current
17
I E
-------
AVELENG1H IRL AHRI FR USDA
Fig. 10. ZRLSpec—SO sensitivity and dark current
18
:1
.1.
a
w
C-,
-------
277 22 i_1_
277 223. 8
fi1
1.... .1.
I
fj1 1 FBZS4OCLG1O76
fi1e J 3 11S4L nfUJeT ed
21 1 W 1.gliting function :(9)
(J ,
I
/
I
I
/
/
I,
\
\
\
I
48.1
\
c o, —
( J t’J c’J C\I m m m cr
WAVELENGTH n IRL AHRI ARS
Fig. 11 Lamp spectra (unfiltered) FS4O, FBZS4OCLG1O76 and weighting
(Q N.
c m
USDA
function 9
1 ř
1 ř
75.1 cm
1
I
I
u 1
19
-------
UAVELE1 T4 r I A I FR USDA
Fig. 12 Lan p spectra FS4O and cellulose acetate 6.5 hr. Z = 30cm, Z 5 m
20
-------
2770923.09
2770923.07
2770923.05
2770923.04
0.00 fi1
ci1 3 1_ Qu !!tered :AE’=
f i1 4 1.. ..JS4.Q..J- 0. 10T CA-8h
fbjljj 5 + 0.0075” CA-7h
fi1 6 L_FS4O+O.005” CA-6h
21 1_ We g iflg function (9)
I I I
I
I
1
= 5.34
= 8.5
= 10.1
i4
CO t- co 0) — C\J ( ) L ) C o
C\J C J C C’J C\i C çr C T ) CT) CT) CT) (T) CT)
WAVELENGTH nm
0.127 mm, 0.191 mm,
IRL AHRI ARS USDA
0.254 mm cellulose acetate
= 50
= 50
= 50
1000
100
10
1
a 1
0.01
v- .1
E
C
11*
I
0.001
Fig. 13 Lamp spectra FS4O and
21
-------
Snowmass, Colorado
T1ZRI ua
FILEj1q-2 FS4O+O.005
22 81LAft.
Z2Vi 1i3-Fi2_FILE 2a-2
Fs40+0.oO5
Z7 RIŘJ F.IIF 27-?
77 1L3&.. . .EflS B—p
o-uw
FS4O+O.005
27E 1 Ř5J . F TI FJfl2- I e hting
CA—20] 4
‘CA-20
‘CA- 20
ered
CA - 26
uiiCti
!AVELENCTH r m IRL AMRI FR USDA
Fig. 14. Lamp spectra, Snowmass, Colorado
IP2
IP2
1P2
1 P2
1P2(
n9
E
c-’j
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w
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1řř
1ř
1.
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a
22
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+ Th : r fl: T :T: :!
iiiiiitti_ =1/--- Li iii_ ii
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r
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. - - — - -H - - —
-- - ‘1 - T
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i i iiiL- :; :: :i
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-
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ni
- :ii:T:-
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IIi
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i:’i:i :
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p
.:
9O 300 31° 3Z 30 4o jçô
6” 74
80 o
.. 4 - - -.4-- 4-- ... ,.— - 4 . i ii : 1- Z• I
. .4
4
3
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8
7
6
5
4
j. .1
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! ‘:;::: . : :
•4-; r;! ¶7A
3
4’
1 —
4—
2
( ±: ‘le’
Fig. is. Lamp aging — BZS 20 CLG0377
-------
Fig. 1.6 Sylvania F15T8 CW lamp spectra — lRLSpec -S
24
-
L r -
11 ITT
--+.- --- . . i
-4---,— --4--+----4—+-
1i i T I:
-
I
7.
IEEELiH 1
9
B
- 1
t iF11iT
iii i
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7L
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.01
9
a
7
6
5
4
25
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Fig. 10. Sylvania F15T8 CW lamp spectra IRLSpec—D
26
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8 c .r’iAf z1— 2.37
A CAL. Z9 .7
5- I’ -77
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c\J (‘.1 01 0) 0) 0) 0) 0) 01 01
WAVELENGTH nm IRL AMRI ARS USDA
119. 20. Sun spectra - Beltsville lID
1000
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10
1
0. 1
0.01
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‘a
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9
8
7
6
5
4
3
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30
to
9
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4
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Z0 30 40 350 60 10
-------
I000I
Fig. 23. UVB IRL METER CIRCUIT
RESPONSE OF UV-B IRL METER
2-7-77
Fig. 24. IRLMeter spectral response
I I I I _ _1___. I I __ t
R
20X
RE RD
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too—I- I I
>
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31
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20 X 2 3 PER INCH
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Fig. 26. Rayonet F8T5 RPR 3000A° ‘spectra new cellulose acetate
Z = 16.3 cm
222i a1a
222.1 2L.1.4.
2271 2L1 .L
. Efl F 1 -3 et
FILE GL 3
Flif #I& R yonet .
iřgo.
too.
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c J
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AVELENCTH rrnt IRL AHR1 FR USDA
Fig. 27. Rayonet FGT5 RPR 3000A° spectra cellulose acetate aged 6 hr.
Z = 16.3 cm
27711f 1 f A nr p7—1
22Z1103 05
27711Ř 1 Fi
Ravonet
FiLE ( 3flz3
FILF L 9-
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(T)
34.
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Fig. 28. Rayoeet FBT5 RPR 3000A° spectra ce11u osc acetate aged 6 hr.
Z = 20 nm
77lli l8 FTIF —2—a
ZL2I 1 FJLE -33
77711 7 FTLP -4—
R iyonet
Rayone t
R yonet
1000.
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1 a
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35
-------
277 . ri1 76 0 S n j svi11e 1:15 EDT
77 2 2 . 11 fi1o 14 1_01 . ay net Ill Unfiltered
770902410 f i1 # 15 1. Q .d aou J 0
0.00 fi1 21 1_ WeJ . ghU , g..junction (9)
in. CA—lOh
7,43
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c u c .i c i ci m m m
WAVELENGTH nm
Fig. 29. Rayonet F8T5 RPR 3000A° an 5 sun
IRL AMRI ARS USD ’
(Beltsville) spectra
36
-------
100
10
0. 1
0.01
00001
Lfl CD N cX O
(\i C’J C\J (‘J \i Cr)
\ —Wei htsrij fi’,,ct,crn (
-.4
CT ,
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F .g. 30. Spectra FS4O anl Rayonet E 8T5 RPR
r\J m It CD N
c cn
IRL AMRI ARS USDA
3000Pt
1000
2?7 J923..08 fj1ot 3 i unhlteréd i r= = ,‘ .i
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I
I
37
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1000.
I AVELENCTH nm 1RL AMRI FR USDA
Fig. 31. Spectra FS4O and cellulose acetate aged 0.5 hr. Z = 5Ocr
221122&2AflL 2 R j S40+Peo
221028.21 F540+Bel
22B 1O5 L FIIF J1 -1 Wei 3 htin
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1000.
100.
10.
1.
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C
c J
1AVELENCTH nm IRL AI4RI FR USDA
Fig. 32. Spectra FS4O and cellulose acetate aged 6 hr Z = 50 cm
39
-------
100
L - ‘• . . .
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rn 1 ’k
Fig. 33. Spectra - eye protective device
40
IRLSpec—D uv•-
BW:2 i’r
. -Th
2
1.
3 ,
2!
‘-4
N
— 4.7
SB
0.8001
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6°
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Fig. 34. Spectra prescription eyeglasses and sunglasses
- --f’l-!
- --r .
p.-
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• •1;
r
4].
-------
APPENDIX A
Ala — Ale — Operate Pro r ,vn, lRLSpec-D
A2 - Calibrate Program, IRL pec-D
A3a — A3b — OreraLe Program, IRLSpec.—S
A4a — A4d — Operate Program, rRLspec—so
Spectral Rcs nse Curves
- Standard Lamp Plotted Edch urn
— FS4O, FSZS4OCLG1O76, Sun, lRLSpec —S
— Filtered rS40 and FHZS4OCLG1O76, IRLSp c-S
— FS4O, F ZS4QCLGl076, Z = 73 cm, IRLSpec -D
- F iS4O WLG, FBZS2O WLG, Vitalite, IRLSpec-D
- F15TB
- F 1STB
- FiSTS
- F 15T8
CW Westinghouse, IRLSpoo—D
CW Westinghouse, IRISpec—5O
CW Sylvania, IRLSpec-D
GRO-LUX Sylvania F15TB NW
GE, IRLSpec-D
— F4OR GE, F4OIP Westinghouse, FISTBR GE, Z - 50 cm, JflLSpec
- F4OIR Westinghouse, Z = 20 cri, IRLSpec—D
— F15TBBL Sylvania, F15TSBLB GE, IRLSpec-D
- F40 Br. + FBZS4O CLG1O76, IRLSpec—D
— F20T12/2021 Sample No. 3175-2, rRLspec—D, IRLSpec-SO
— F2ORL Phillips phosphor, F4TSBL conventional phospher,
IRLSpec-S
AS
A7
AB
AS
AlO
All
Al 2
Al 3
A 14
Al 5
A 16
Al 7
AlS
Al 9
42
-------
AUTO 5 T APT
A—I PP. ‘: AL. C AU
8 —FE!
C—CAL. ULGTH
[ ‘—SC All PEC TEA
E —S TO P
F — F 0 P C l A F’ P
G—PPT. PATH
I—A’:T. TNT. 9
—NL’ITH. CHEU
L—ST t’. LAMP E l ?
i i C i’ ;
Il-C m i. PEAT’
ENTEP ‘P. 10.
2 7 8 01 2 C’. 0 0
PE ET
U U
ij i’
l i i i i i
O 0 : 1
U U
ti i — ; — :
O Ci: 4
An S
till -: -.
U U
ti ll -V4
01140
o 04 1
0042
004 1
0044
0 ( 145
004 5
0 U 47
hO 1 :1
C i C i 4 9
liii 5
Ci ‘351
A it S
A A
0054
nit 95
I i A S
0057
it n s
fl 5Ci
I I I r. Ii
nn, 1
U Ut
liii,-
0064
it A
11 1 r ,
U Ut 7
I lilt—,
111Dm ‘- I
L1U U
0071
0072
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0074
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L
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T
H
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C l
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P
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[ T ilE
E
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P
L I t IE
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F
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P
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L I HE
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T
0075
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0077 L’
W i ?::: H
0079 T
i_ i U : U H
0081 LINE
00 :12 1
— : —
(‘0:14 A
0085
0086 T
UU H
A ii;:: ;
0089 1
0090 H
0091 T
0094 9
0095 LINE
0095
uu’ 7 -
009:E: I i
0099 L
0100 G
0101 T
0102 H
0103
01 04
0105 C
0106 H
0107 E
0 1 O:: C
0109 1
0110 LINE
0111 L
0112 -
0111
0114 T
0115 B
0116
•0117
0118 L
0119 A
0120 H
01.21 P
012.2
0123 E
0124
(Cassette D4)
F iL E
it FE
USED
N A
till tIN
U U U
11 1111 —:
0004
liii 1 15
ti M lit—.
U U U
U U U:
ititti ‘a
0010
0011
0012
00 13
0014
00 1 5
0016
001 ?
00 1 8
0019
(111 —‘it
0021
U U .2
U U.2
0024
liii
U U .2
31
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74 5
1000
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H
I
F’
F’
H
L
H
ti
F’
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LINE
Fig. Ala. Operate Program IRLSpec—D
43
-------
Fig. Aib. Operate Program IRLSpec—D
(Cassette D4) (cont’d)
0125
LINE
017’ #PEGS
0126
N
0180 coic’
0127
—
0182 [ LEAF
0128
S
ill:::1 6
0129
U
0184 + —
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0185 ENTER;
01:1
S
I 0186 1
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c u :37 FRMT
0133
i i
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0 134
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N
I:
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0194 FRNT
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.
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0140
0197
0141
F
I 0198 L
0142
E
0199
0143
0144
0145
A
t’
LINE
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0201 1
0302 A
0146
E
0203 L
0147
0148
N
T
0204
0205 ENDc
0149
E
0306 5
0150
F!
0207 0
‘ J l S l
020r ITO
u15
1
0209 SFG
0153
P
0210 LE:L
0154
.
I ’
0155
H
0212 LE;L
0156
0
P
•
0157
.
0214 Fi7
0158
P
0215
0159
A
0217 0
J
0160
1
0218 1
0161
#
02v3 ST0÷
0162
END
0221 PLL
0153
STOP
0221 PRINT
‘
0164
Fl::
0224 Fl::
0166
2
0226 1
0167
EE
0227 STO
0i— 8
5
0228 IF IFG
0159
+
0225 GOT’:’
0170
STO
0*
02:1 1
0172
PRINT
03:2 2
0173
5
0331 0
0174
NASI
4
02:4 STO
0175
1
03 :5 3
0177
5
0316 9
0178
1
cc37 ITO
; C
F 000
F U U 0
4
I - i
F
0218 1
o3:: ENTEPT
0240 0
0241 + -
0243 LOAD
0243 5
0344 WRITE
0345 PEAT’
024:1 ENTEP
0249 4
fl3 1i + —
0351 IF ::=,
n :.c — —
0254 GOTO
113 r
0357 NE, ] E
025r FOP
0260 1
0361 STO
0363 PEI ID
0364 IF 0
0365 G0TO
0367 GOb
0369 :::,
0270 IF SF’:
0371 GO b
0371 STO
0274 5
0275 NE:ITE
0277 FOP
0278 READ
0380 ::
0381 5T0+
02S3 NE::T
113:::4 WRITE
03.15 P [ L
0387 P [ L I
—
0390 iot::
0391 STO I
0293 CFG
0394 P [ L
0295 3
0296 4
029:3 +
039 ENTERT
c i:: 1
o:.oi EE::
ii —: - , — :
ri
U :
44
-------
0303 + 0365 EH [ ic 0418 PCL
0304 + 0: 5 5 2 0419 Fl::
11305 PPIHT cl::57’ : 0421 PPIHT
0:05 IF ‘ FC 3 0:5:3 f 11422 EHTEPT
IrLi7 Gislil U it 0 - T’’F ‘ 4 -
0309 . 0171:1 FI:. 0 0424
0310 0 0372 PF1HT 0425 0
0:11 1 0:73 FHTEPT 0425 6
0312 510— F000 017’4 2 0427 —
0314 GOTO 0117 0175 4 0428 FF111.’
0315 HE::T A 0376 9 0430 5
0317’ 1 0377’ — 0431 U
0:18 HE:ITE 4 IH STO A 0432 i i
0:20 IF’; 0 . 7 ’ :3 04 :3 S
0321 F ’ : L A 03:30 0 04 14
0122 ::: I ;:31 To F 0435 =
0323 0 0382 CLEAF 0415 PRINT
c 1324 — 038:: STO j 0437’ EHDv’
0325 IF + 0384 FOP A F 04 3:3 COTO N
0326 ‘:01’:’ 1 1 ,355 Ffl A 0440 LE:L
032:3 IF SFG 2 0185 2 ———— A
0329 ‘ :OTO 03:37 4 0442
0:31 RETURN 11388 4 0441 0
0332 LE:L 0389 + 0444 1
F 0190 2 0445 nTO+ P000
0:34 3 0391 2 0447 FI 2
: HE TE 4 11 ‘i44 F’ L FL 0 ’ l
01:7’ PETUF II 0393 0451 PRINT
• 0338 LE:L 0394 1 0452 Fli
• ———— E 03’ 5 7 0454 1
0340 LLEAF’ 0 1 5 :3 0455 StO A
0341 HE:ITE 4 0397 — 0455 1
0343 PFTUPt1 0398 9 045 5
0:44 LE:L . 0399 ii:. 0458 0
E: 0400 510 H 045? 510 F
0345 1 0401 4 0450 4
0147 HEYTE 4 0402 — 0451 0
0349 1 0403 4 0452 ‘ITO
0350 RETURN 0404 11:: 0451 1
0351 LE:L 0405 810 0464 ‘ITO
I 0405 F’:L H 0455 5
0353 PPHTc” 0407’ 4 0456 HETE 4
0355 1 0408 — 045:3 PEAr’ 4
0355 H 0409 + — 0470 ENTEFt
0157’ 1 0410 el: 0471 4
035:3 0411 PIL 0472 + —
0159 9 0412 ÷ 047’:: IF ::=1
0:50 0411 P’Th I A 0474 ‘:oio 047:3
[ 1361 F [ 1415 + 0475 ‘:‘:‘TO 0
0:52 P 0415 ACC 0478 3
036:: 0 0417 HE::T A 0479 HEITE 4
0154 N 114:11 FOP
Fig. Mc. Operate Program IRLSpec—D (Cassette D4) (cont’d)
45
-------
04 F 9
I ’
c ::
4
-.
4
‘ S 54 1
H
O 54 1
0483
PEAt’
4
0484
IF 0
- 4 ’ ’C
.t
-- -
I -
0487
G’JTO
[ 1489
:: ‘‘
04’ZO
IF LFG
0491
GOlO
049::
[ LEAF
0494
‘ET’ ’
0495
5
0495
lIE:’’TE
0498
FOP
049’
PEAL’
0501
:: ‘‘
0503
ST [ ’+
[
050::
NE T
B
0504
3
0505
NE:’’TE
4
0507
1
0508
‘ ITO
F:
[ 1509
P [ L
[
0510
‘FlU-i - 1
A
0513
+ —
0513
P [ L
A
0514
3
0515
4
0515
9
0517’
+
0518
EN1 EF’t
osici
EE::
0530
4
0531
+
nc;33
t
0523
PPIHT
0534
IF SF’
-C-,c
i_ I — ‘ ,
-- -
i_,
05
NE::T
0538
GOTO
0530
FF111::
nc; :: :
n
nc;::::
\.‘
0534
E
0535
P
05:5
L
0537
0
053:3
A
0519
1’
0540
END’:’
0541
1
0543
NE:’’TE
4
0544
CFG
:3
1i545
[ FG
4
[ 1545
0547
0549
0551
- cc-- ,
I I I I
- C C - .
I I I I — .
0554
I i c;,:
- C C
4! I
Iic ; ;n
0551
I i c; R’
I i c;
0554
i i c ;
n c ; ; ;
0557
I i c -: F:
,: ci
- C--- ’ -
I_ I —
057 1
- C. -, - ,
0574
0575
- C — -,
I_I _I i
05 78
0579
c :: I i
05 :3 1
05::: 3
Ii c;FL:
0584
- c r c-
II I ,— , I
I i
I i C F:
I i C F: ci
Ii
I i C ci:’
0594
I i c; ci s
c
05 9 7
Ii ci ::
I_i i ci ci
I r, i i i
0501
L1h 11
F’ E I I I F’ I
L B L
L
L F: L
L
[ L E A F’
Lt’ U
L E: L
F’CL P0 47
P IL P0 48
L o’:
ElITE F’ I
4
÷
4-’
C
t
Fl:: 3
FF11 F’:”
LI NE
I i
II
E
H
I
PP TNT
E I Li
F l:: 4
[ FG 3
STOP
L E: L
l’l
F’ F’ NT c
11
F
F’
(Cassette D4)
E lit’ c
STOF’
F I::
F’ F’ I I IT
4
STO
PENT.’
T
EN I’ c
r T -
U
FE Il-IT
4
‘.4
‘F
‘:L E A F’
.rC
F ‘3 F’
F’ C L
4
‘-4
F’CL
STO+
NE::T
F l::
F’ C L
F’ F’ I I T
1 ’ l
I
L
L
I
4’
H
T
if
I lIT
E I I Li ’:’
11 ,—Li
050 4
i i ; C
i_ t’ Li
Li t—. 1:—:
11 4 -ti’- ’
0 5 1 0
0511
051 3
U 5 1::
051 5
051 5
05 1 7
[ i 5 1 F
05 1 9
L1 , -‘ I i
O 531
U t
l lt.
0534
Ii; -‘
U t’ ,. 7
hr il
05 31
i i :
ii 1- -:
05 3 4
11 , ,
Ub
Li r,
0540
054 1
054::
0544
0545
051 5
0547
054
0549
Ii ; ii
0551
Ii 3
1i c;t’
0554
‘ ; c
c -fr
0557
Ii ; ::
(cont’d)
H
i i
H
-I
Fig. Aid. Operate Program IRLSpec—D
46
-------
0659 COlD N 0703 P
0651 LEL 0704 E
1 1 ( 1705 S
0651 (LEAF 0705 E
0564 si ’: ‘: 0707 1
0655 4 0708 END:
0566 NE:ITE 4 0709 GOlD 0122
0668 4 0711 LE:L
0669 0 ———-
0670 ST ’: C 0713 LE:L
0571 1 ———- C
0672 ST’J E: (1715 P(L R0 0 0
0671 fl P E:- C 0717 F l:: 2
0674 READ 4 0719 PP 1111
067 6 :: r 0720 1
0577 si’:+ 0721 STO Es
0678 UE::T E: 0722 1
13579 P’:L C 0723 2
0580 PAU IE 0724 0
06:31 PAUSE 0725 STO F
0682 (LEAP 0726 F l:: 4
0683 S b 072: 1 FOP A F
0684 GOTO 0739 P(L I A
0586 LE:L 0731 P iTh A
0732 2
0588 LE:L 0713 4
O 0734 9
0690 . 0735 +
0691 0 0735 EN IEPI
0693 1 0717 1
0691 810— P000 0738 EE
0595 (LEAP 0739 3
0696 NE ’IE 4 0740 ÷
r,69s 1 0741 +
0599 NEH!TE 0742 PPINI
0701 PPNTc 0743 HE::l A
0744 STOP
0745 END
Fig. Ale. Operate Program IRLSpec—D (Cassette 1 )4) (cont’d)
47
-------
Fig. A2. Calibrate Program IRLSpec-D (Cassette D2)
FILE
I t
U SED
H A::
ii ii
H
F l
H
4
T
4
I - ,
F 0 F ’
F’ C L
4
( 1
+
‘: L
+
‘: L
4
1 0 1::
F: L
+
I 0 t
s i ’:
PC L
4
‘-1
+
F’ C L
E:
0051
:10 5 2
hA S
o o 54
It O S S
0055 A
[ 1057 N
i i i i s S B
S q
lilli- li —
0051 1
0051 LINE
0053 LINE
01: 154 LINE
A A ,- S
lilt 1—, I— ,
0057 N
l1lth
I iIth’ 4
L IL t Li
0071 F’
0072 U
0073 N
0074 5
0075 =
L ILt
0077 EHfIv
007:3 STOP
0079 910
00:30 1
0AF:1 910
00:33 1
00:13 5
00:14 0
0095 910
lilt:-: ,-,
Uu ’7 u
it
AAQA c;
0091 + —
0093 910
0093 1
0094
A hi’ S
0095 1
LIU’’7 7
0099 910
lilt’- ’’- ’
0100
0101 ::
0102 7
010:: r
I ’ifl 4 SICt
-F
0000
FF111
lilt H — ,
I_I LI Li
H
0004
L
0005
I
0005
B
0007
F’
L IL I ILI:::
H
0009
T
0010
I
OCt1 1
C’
0012
N
001::
L iNE
0014
C
0015
0
0015
F’
0017
F’
00 1:3
E
0019
C
0020
1
0021
1
lilt — .
0023
N
0024
0015
N
0025
I
0027
L
002:3
L
0019
LINE
0030
E:
[ to:: 1
E
U L I
003::
J 00:4
F’
E
hiA:S
I
it it’ •: -.
0037
P
003:1
B
0039
E
0040
Ii
004 1
ci Ct 4 3
1: 11: 14 ::
0044
0045
0045
0047
I 0049
0049
0050
N
LINE
F
I
L
E
0 1 05
0105
0 1 07
0 1 0
010(1
011 0
0 11 1
0112
0 113
0114
0 11 5
0115
0117
011 S
0 11
o 1 10
0131
01 12
0 1 2
0124
01 25
0 l IE
0127
0 1 3
0129
O 1:: 0
01:1
013
0 1::::
0134
0135
0135
01:7
1 31 3:3
0139
0 1 40
0141
0142
0 1 4 3
0144
0145
0146
0147
0 1 4:3
014:’
0 1 50
0151
0152
0 1 5 ::
0154
0 1 55
0156
0157
0 i 5:::
0150
H
01ri -
0152 LL
015:: LS
0154 -
0155
o15’ if
015:3 HE
015 1
0170 5
0171 0
01 71 EN
0173 1
0174 EH 1:
0175 0
0175 + -
0177 PCfH
017:1 1
0179 5
0190 0
01::: 1 E1 ITf
01:12 1
01:1:’ ENT:
01:34 1
01:35 +t-
01:35 F’’: p
i:t 1::: — F’F’H l
Al :3 ( 1 C
0190 A
0191 L
0192
01 93
0194 C
0195 0
0195 N
0157 9
01:1:3 1
01 ( 19
II —‘lilt
0201 1
0202 N
0:0:: LINE
0204 1
0205 1
U L1 LI
II I I :-:
0209 1
0210 5
0211 0
0212 END’”
0213 C LE li
0214 910
0215 (LEAF
0115 LD: ;
0217 END
II
H’
S
+
S I C’ —
F’: L
F’: L
+
EN TEP I
.4-
et::
+±—
+
5104
F [ L
F’: L
H
H
H
E
48
-------
[ ‘ATE
E LI’d SPE’:TRA
17:30210 - -1 OPERATE PROGRAM IRLSpec—S
0000 CHA 0051 CHA 0101 101 0151 RN llDkil mnj
0001 1’ 0053 i 0102 FL’3 0152 11 11202 SP
0002 A 1 1053 ‘ , ‘ 0103 00 0153 — cPli3 °b
0003 T o o s u 01011 JFER 01511 R i iu i
00011 E o o 5 U1W ul 0155 INE’ R
0005 ‘:HA 0056 p ulUt. Tub u15h RN RN
00C1t E 0057 E 0107 101 0157 00 ‘41
00u7 + I U 1Uc — Uljc. —
0008 1 t 109 RM 0159 021v ; L
0009 E P 0060 R 1j110 11 ulbO R 02111 j
0010 6 0061 A I:hhhl — ilbi IND fl211 112
0011 0 1t53 L1112 L L11b2 SN =
0012 EN CHA 0115 rr 01? 01
0013 10 00611 F1::5 UllU RN UlbLj IL’2 113111 - -
00111 E flO SS 02 0115 00 0165 FI::5 31S 1
0015 oP 0066 COL 0116 0165 03 2 1 =
0015 Be 00 57 10 0117 10 ’ 0167 C0L
--4— fl hlO 1 u 1110 ç- 1 c . - , — -
UUh LHH S SJL 021c: +
001:3 7 cicisq 0119 INC 0169 71 1
0019 0 flfl7O w 0120 SN 0170 01
0020 SN fl 71 L 0121 Ui ‘17i 1 21 1
0021 05 0122 SN 0172 Dl =
0022 FL I3 0123 03 0173 00 -
0023 10 01211 102 01711 Di : 5
002 1 1 :3 1i075 0125 RN 0175 01 fl22C 03
0025 0 R 0126 02 0176 :M fl3J
0026 SN 0127 Fr:-:s 0177 02 n337 ń7
0027 00 flci7P A 0128 00 0178 RN
0023 1 0-373 , 0123 COL U17 02 RN
0029 U. ulsw ki: Ij1:::w — 113: li R
0030 0 0131 RN 0181 RN I2j
0031 SN I lflA2 0132 03 0182 05 fl2 2 p
0052 0 1 0033 w 0fl3 US ui : : = 025 RN
U0 2 00:11 L U1:U 1 i. E’ l:U. 02311 E E
uu u 5 fl op s u1 ‘uL Ulc UU =
0035 0 0136 08 0186 FL’3 LO ,
0036 SN ç p - 0137 0187 01 - - -
0037 02 00.8 0139: 01 0188 lOB fl3 P, 3
01133 +B 0133 1 01.33 CHA =
0039 105 ncr;ct R U 1UL1 Dl U1 3ti c12u :,
00110 CHA A 01111 02 0191 CHA 62 1 1i
00111 2 ç 1 qt , ulU2 RN u132 1u2 ui3’43 711
00112 CHA I I V IZ 01113 02 0193 LF 02113 RN
00113 102 0C19U CHA U1ULL Fr: :s 0fl411 RN 021111
. 30 1 1 1 1 . — U1LLS UIO U115 U2 1i2 1 15 =
00115 0 Ul 01116 CUL 0136 — fl2U5 FISS
00116 1 0097 106 U1 1 17 U: U1’i, RN 02117 02
00117 D l 0099 CHA 0111:3 IUh i l l-i:-: ‘ fl2112. I OL
0011:3 10 ç 1 qc 1 01119 101 0199 =
— — — — — rr — — —
UUL’ RN 010(1 CHA U±L 1 — U UEi xr 4 0250 LF
0050 10
Fig. A3a
49
-------
OPERATE PROGRJU.I IRLSpec.S (cont’d}
0251 E 0301 FL’3 1i352 ‘:f-J R OLUI1 IND u 451
:1252 FLG 0302 02 035.3 — 0402 F M 01152 F
0253 1 14 0303 L 035t1 RN IIUIR 03 El115
0254 p @3011 1110 0355 15 c 1u.cuj . :m E145 11 0
fl2 5 : r 0305 RN 0356 = 0405 1i3 k’USS N
0256 C M U3Uh 00 @:!57 F i: :s 01 1 (16 Ft UUSh ‘HA
0257 02 03% 05 04.07 INt 0457 LE
ul2 IHA 0308 L 035,3 COL 01108 RN 01158 E
( 1259 0309 INC U36 1€ 04.03 fl3 UU5 EM
0260 1 0310 RN @361 LF ou lo :rn uLkb 1 08
0261 A 0311 01 0362 SP 0411 02 01161 (HA
0252 Ft 0312 = 0363 :3 0412 1 0452 T
0263 1 0313 N 0364 FL’3 0413 :rn 0463 0
0251 1 0314 02 0365 @7 0414 03 0464 (HA
0265 i 0315 Ft 0366 CHA 0415 RN 0465 E
0266 11 0316 IND 0367 5 0416 03 0466 LF
ci257 -r 0317 RN 0358 u out- — 01167 SN
0268 0318 00 0369 N 0412 1 ‘ 01168 09
0269 ‘3 ‘i319 - 0370 01113 5 0469 FL’3
0270 0320 Ft 0371 F 01120 = 01170 11
0271 A 0321 1110 0372 Ft 0421 0471 L
0272 - - 0322 RN 0373 0 01122 2 0472 IN C
0273 ç’ 0323 01 0374 N 0423 — 0473 RN
0274 (HA E1 2 4 = 0375 (HA 0424 RN € U7U 08
0275 E 25 34 €1376 E (1425 1111 (1475 FJ::s
0276 IFE u 26 02 0377 SM 01125 = 0476 03
0277 12 t 1337 1 0378 03 0427 IF— 0477 COL
0278! 3 E1328 :111 0379 ‘:HA 01128 03 Ot-tiS 12
0279 :3 ‘329 00 0380 1 0429 LF 0479 Ft
0280 0 UJ3O : i 0381 0 lnj3ci RN 0480 INC’
0281 ri W $1 01 03:32 (HA 0431 1t2 0U .1 RN
0232 FLG I RN 0E 3 E 0432 Fl 0L1C2
0283 12 ‘J$33 01 03811 9’l 01133 03 01183 F i:-:s
0284 — i 3 4 — 0385 O iL 0434 0W811 03
0395 1 0335 7 038b CM 1 1435 ‘ T 0485 COL
0286 0336 9 0387 02 11436 cr 048b 10
0287 0 0 37 0388 RN 0437 01187 LF
02E8 = IF— 0323 03 0439 2e OUtd 1
c1289 — Ui 9 02 0390 — 0439 3 cluEs :ii
0290 @ U0 RN 0391 2 @uu.ř ‘ a 0490 08
0291 0341 02 0392 + 0441 0 0491 RN
0292 SM 0342 F i: :s 0393 1 0442 CN 0492 0:3
0293 01 I l 43 @3 03911 5 04tH 116 0493 —
0294 + 03t14 CCL 0395 011L1-U ‘3T 0494 RN
0295 1 ‘ 5 08 0396 SN @1 143 10 01195 09
0396 0 ‘J346 LHA 13397 03 04115 CHA 0U 5
0297 0 W 47 03% FLG 0447 N 0497 IF—
0298 = S 0399 03 €1448 E 11
0299 SN @349 U 01100 L ńIUH N 0499 EP
0300 00 0350 f l 0500 00
0351 5 - - -
Fag. A3b
so
-------
AUTO START
A rPECTPA
E: REVERSE
C S L CAL.
E OTOP
F FORWARD
I: PPrNT DATA
I INTEGRAL
J PUN NLGTH
I NL’:TH EPUAT.
L lTD LAMP EOUH
I i S U H
0 OPERATE
ENTER P . NO. PA #
37SO 120, 00
PEAL ’ 1
HIT A € OF ’ J
0000 PPNTc’
IJIjLj, H
11 1111 — :
00 ( 14 3
0005 P
0005 E
UUL17 ,
00DB T
0009 P
0010 A
0011 LINE
001 E:
0013
0014 P
0015 E
oois v
0017 E
€1018 p
€1019 S
0020 E
0021
0022 LINE
L 1 L I — , — ;
0024
0027 L
i_ I U
11 11 -‘—4 I
L1LI;W H
oo::i L
0032
0033 LINE
0034 E
0035
Lii1 t.
00 :7 T
l L 1 —::—:
0019 r
(1U34€i L INE
0041 F
0042
0041 F
0044 0
0045 P
0046 N
0047 A
0048 P
0049 L i
0050 LINE
0051 C
00 52
0053 P
0054 P
0055 I
0056 N
0057 T
1111
0059 L i
H
0061 T
UUt H
0063 LINE
0064 I
fink;
0055 I
0057 N
006:3 T
006 E
uu7u ‘
0071 F
Uu7 H
0073 L
LINE €1123
0124
0 1 25
P 0126 0
0127 P
N 0128 E
0129 P
0i30 A
0131 T
0132 E
01:3
0134
NE 0135
0136
01 1 7
€11 18
0119
0140
0141
• 0142
0143
0144
0145
0145
0 1 48
0149
NE 0153
0151
0152
0151
0154
0155
0155
- I C -,
P_ I A
0 1 58
0160
0161
0163
0154
0155
0155
0157
015:3
016
0170
0171
0172
0 1 7 3
L I NE
E
N
T
E
P
1•1
I i
H
:1
L I NE
E I [ i
‘3 TO
E E
+
F l ::
F’ PINT
S T i:’
4
I I
STU
C
I I
STO
F l ::
Fig. Ma. Operate Program IRLSpec-SO
(Cassette B)
LINE
F iLE
USED
H A::
I I
-r
720
i i i i
0074
0075
IiL l7t
0077
007:3
k 1 ij 7 4
liii— LI
0081 N
0082 L
y U :
00:34 1
0085 H
00:35 LI
0087
i_ I I
0089 N
0090 L
0091 C
0092 T
0093 H
0094
0095 E
LILI 4fr I I
no 7 U
UU’fl: H
0099 T
o1i: io
ülüI Li
0102 L
O 1 0 ::
0104
0105
0106
0107
0108
0109
0110
0111
0112
oil:: E
J1l4 U
0115 U
0115 H
0117 LINE
131 iF: H
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52
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Fig. A4c. Operate Program IRLSpec—SO (Cassette B) (cont’d)
53
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Fig. A4d. Operate Program IRLSpec-SO (Cassette B) (cont’d)
54
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FINAL REPORT ‘
,q
i F
BIOLOGICAL EFFECTS OF ULTRAVIOLFT RADIATION
ON PLANT GROWTH A D FUNCTION
M. N. Christiansen
Plant Stress Laboratory
Plant Physiology Institute
Beltsville Agricultural Research Center
Beltsville, Maryland 20705
EPA—IAG—D6—0 168
Project Officer:
R. J. McCracken
Agricultural Research, Science and Education Administration
U.S. Department of Agriculture
Washington, D.C. 20250
Prepared for
Environmental Protection Agency
BACER Program
Washington, D.C. 20460
-------
/ ;f J 1 /
ANNUAL REPORT TO EPA, BAC [ R PROGRAM FOR FISCAL YEAR 1978
f 9ject Title :
Effect of UV-B Radiation on the Eggs and Larvae
of Marine Fishes
Principal Investigators :
John R. Hunter
National Marine Fisheries Service
Southwest Fisheries Center
La Jolla, California 92O 8
John II. Taylor
University of California San Diego
Department of Psychology
La Jolla, California 92093
October 18, 1978
-------
ANNUAL REPORT TO EPA, BACER PROGRAM FOR FISCAL YEAR 1978
Project Title: Effect of UV-B Radiation on the Eggs and Larvae of Marine
Fishes
Principal Investigators:
John R. Hunter John H. Taylor
National Marine Fisheries Serv. University of California San Diego
Southwest Fisheries Center Dept. of Psychology
La Jolla, California 92038 La Jolla, California 92093
INTRODUCTION AND OBJECTIVES
In the period October 1, 1977 to January 1, 1978, EPA provided funds
($12K) which, along with residual funds from FY77 ($1OK), provided salaries
for a reduced staff to analyze the data collected in the first year of the
project (FY77) and to complete a manuscript on that year’s work. This objec-
tive was met and the manuscript is now in press in Photochemistry and Photo -
biology (page proof attached). No additional funds were available in January
1978 and the staff was laid off. In April 1978 funds for 6 months’ work were
made available ($39K) and work began again in May 1978.
Experiments conducted in FY77 established for anchovy and mackerel
the dose-response relationship for mortality and histological and morpho-
logical effects for the 4—day embryonic period (egg and yolk-sac stages)
to the onset of feeding. A weighting technique was developed for relating
these effects to ozone depletion and the results indicated that significant
effects could occur near the surface in the sea after a 4-day exposure at
a 19% reduction in ozone. The objectives for the last 6 months of FY78 were:
(1) to extend the work on UV-B effects on larvae beyond the embryonic period
using artificial sources of UV—B; (2) to evaluate the weighting function
using solar UV-B; and (3) to monitor solar UV-B in La Jolla.
-------
EFFECT OF 12-DAY EXPOSUI E OF UV-13
i e t hod s
Apparatus and radioiietric methods iere essentially the same as described
in our publication (Hunter, Taylor and Moser, op. cit.). The methodology is
summarized below:
1. UV-B source was FS 40 lamps filtered by Cellulose Triacetate (CTA)
at the container surface.
2. Visible radiation provided by Chroma 50 lamps.
3. Lamp cycle, 12 hours daily visible radiation, and 6 hours UV—B, with
the 6-hour UV-B cycle centered in the 12-hour day.
4. Duration of each experiment was 12 days (1 day as eggs and 11 days as larvae)
5. Dosage for treatments was varied by use of perforated mylar filters,
placed on the top of treatment containers, raising and lo iering banks
of lamps, and placement of opaque aluminized mylar tape over ES 40 lamps.
6. Control containers had solid mylar covers.
7. UV-3 monitored continuously throughout an experiment.
A difference from the past study was that experiments ended after 12 days
while past work ended at the close of the embryonic period (4 days). As a
consequence, larger 1OL containers were used and live food was added and
monitored daily. Owing to the lower daily dosage required in these experi-
ments, we used mylar tape to reduce FS 40 lamp output as well as the perforated
filters used in past work. Thus, the distance oF lamps fran the containers
cannot be used as a measure of UV-B irradiation in our treatments. We provide
the DNA-weighted and unweighted spectral irradiance transmitted through CTA
2
-------
at the water level at the LD 50 (Table 1). This table can be used to
convert DNA—weighted dosage in Table 2 to unweighted dosage for all
aquarium work. We use the DNA—weighted dosage ui our discussion of results
because only through appi ication of a weiyhLincj function can the results
be related o ozone depletion.
Results
Fifty percent of anchovy larvae survived a cumulative UV-B dose of
675 (J.m 2 )DNA eff. (95% confidence intervals, 595-767 (J.m 2 )DNA eff)
over a 12-day period (Figure 1, Table 2). This cumulative dose was nearly
half that needed for a 50% mortality when the exposure was 4 days
(1150 (J.m 2 )DNA eff.’ Hunter etal., op. cit.). Thus direct reciprocity
of cumulative dosage between a 4-day exposure and a 12-day exposure did
not exist; the longer the exposure to UV-E3, the lower the total cumulative
dosage necessary to produce similar effects.
t nearly all doses, survivors of a 12-day UV-B exposure had grown
more slowly than the controls; mean length at 12 days was inversely propor-
tional to dosage (Figure 2). Surviving larvae had histological charac-
teristics usually associated with the arly stages of starvation, i.e., low
zymogen levels in the pancreas and low glycogen in the liver. Only at the
higher doses (dosage > 1000 (J.m 2 )DNA eff) were brain lesions detected
but the lesions were less well deFined than in larvae exposed for 4 days
at comparable doses.
In summary, extending the period of exposure from 4 to 12 days resulted
in reducing the cumulative dosage for 50% survival by nearly a factor of two.
3
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A reason for this deviation from reciprocity may be the necessity for
larvae at the end of the 4-day embryonic period to successfully initiate
feeding. Larvae in the 4—day experiment showed a significant incidence of
brain lesions even at the lowest dosage (760 eff)’ although 90%
survived the 4-day exposure. It seems doubtful that any of these damaged
larvae would be able to initiate feeding. That less than 50% survived
such a dosage over 12 days supports this contention.
These data also suggest that UV-B treatment over 12 days affected the
ability of larvae to capture sufficient numbers of prey even at the lower
dosages. This is indicated by the retardation of growth in nearly all
treatments (Figure 2, Table 2) and by the frequent occurrence of larvae
with histological features associated with the first stages of starvation.
This effect could be mediated by a direct damage to sensory or other systems
required for capturing live prey or by such indirect effects as higher
metabolic demand caused by the need to repair damage. Regardless of the
mechanism, the probability of survival in the sea for such slow-growing
larvae is low. Even robust larvae can withstand only a few days of starvation
arid the abundance of food in the sea is variable. In addition, slow growth
results in a longer exposure to predators while at a highly vulnerable size
and hence reduces survival.
Relation to ozone depletion
To relate the results of the 12-day exposure to UV- 13 under various ozone
levels, experimental results are expressed as the daily UV—B dose (Setlow
DNA weighted) rather than the total cumulative close. Daily UV doses for
various ozone levels are given for June, in our latitude, just below the
4
-------
surface ci clear ocean water. These data are based on the findings 01
Smith and Baker (In Press) I
Daily dose (J.m 2 )DNA eff. Event
44 Significant retardation of growth after
12 days exposure at this daily dose
56 50% larvae die after 12 day exposure at
this daily dose.
105 .32 cm ozone (“ambient”)
141 .28 cm ozone (13% reduction)
190 .24 cm ozone (25% reduction)
288 50% anchovy die after a 4-day exposure at
this daily level.
358 .16 cm ozone (50% reduction)
The above table indicates that most larval anchovy would die if they
existed for 12 days in June just below the water surface in clear ocean water.
Clearly, larval anchovy are highly sensitive to UV-B radiation. The comru-
tations of Smith and Baker cop. cit.) indicate that in our latitude, in June,
atp nt ozone levels , 50% of anchovy larvae would die if they existed
for 12 days at a depth of about 4 in oc less in clear ocean water, and
that 50% would die if they existed for the same period at about 0.8 in or
less in the most turbid water they studied. Lower ozone concentrations,
of course, would increase the depth range over which this mortality would
he effected and thus increase the proportion of the population damaged.
1 See attached galley proofs for reference.
5
-------
Anchovy larvae occur throughout the upper mixed layer from the surfdce 10
the thermocline, but the proportion of the population at various depths
v ii thin this region is unknown.
These estimatcs depend on the assumption that the Setlow DNA weighting
function adequately expresses the relation between wavelength and dosage
in anchovy and on statistical and radiometric uncertainties. Tests under
natural solar UV-B should eliminate many oF these uncertainties.
TESTS WITH SOLAR UV—B
The aquarium work discussed in the previous section indicates that all
larvae would be expected to die over a 12-day period in La Jolla at existant
June-August UV-B levels, if the DNA weighting procedure is correct. Thus
experiments with natural, unsupplemented UV-B are an essential confirniation
of our laboratory findings.
We have conducted to date six 12-day experiments using solar UV-B treat-
ments and mylar filtered controls on the roof of the Southwest Fisheries
Center. To reduce evaporation, treatment containers were covered with ACLAR,
which transmits solar UV-B. Solar UV-B was monitored continuously over the
12 days using an integrating UV-B Norris meter, with frequent checks made
with a spectroradiometer.
Our experiments have not been successful as yet because of low survival
in the controls. Low survival can be attributed to problems encountered
in working in an exposed, uncontrolled environment including: temperature
regulation, weather conditions, and the necessity of using containers with
6
-------
gradually sloping walls to avoid shadows. Nevertheless, the results arc
of interest. Although the survival in the controls was low (5—20% survival)
none of the larvde in the treatments survived in any of the six experiments.
There seems little doubt that unsuppleniented solar UV-B affected survival
but the extent cannot be evaluated quantitatively until we obtain higher
control survival. We place top priority on the this problem for FY79. We
anticipate we will be successful because control survival has improved in
roof-top apparatus over the course of the work.
SPECTRORADIOMETRY
The basic instrument used for spectroradiometry of both natural and
artificial sources was the Optronic. Laboratories Model 74lV, which covers
the range from 250 to 800 nanometers. Spectral resolution of this instru-
ment is one urn for the 250-360 band and two nm from 362 to 800 urn. Since
the device incorporates a single monochrornator, stray light inevitably con-
tarninates the data to a small degree at the lowest energy levels. Continuous
monitoring of UV energy during experiments was done by use of the UV-B meter
designed by Norris and made by Optronic Laboratories, which was calibrated
frequently against the 741V. Fundamental calibrations of the spectroradiometer
were made against a 1000W tungsten-halogen standard of known output. The
Norris meter outputs were recorded on strip charts, and these records were
planimetered to give total dosages for each day of the experiments.
Natural Global Ultraviolet
Approximately 300 scans of the natural global ultraviolet were made on
the roof of our laboratory at the site of the experimental apparatus. Output
7
-------
from the spectroradioneter was processed by its associated hewlett-Packard
98151\ programmable calculator and ancillary plotter. Records exist in
three forms: pri nted tapes; scm i 1 ogari thrni c plotted graphs; and magnet i c
tapes. Special programming tapes allow us to integrate the energy in any
desired band of wavelengths, to apply any selected weighting function, and
to convert to any projected optical ozone depth. The results of these scans
are given in Table 3. In most cases we nedsured only at local apparent
noon, but half—hourly or hourly scans were made on selected days throughout
the period in order to determine the seasonal changes in the energy envelope.
Weather notes were made on a brief and qualitative basis. Calibration of
the Norris UV-B meter was based upon the spectroradiometric data for local
apparent noon. [ it is evident that the conversion factor between the two
instruments changes somewhat with time of day, owing to changes in the spectral
content of the daylight and to the fact that the Norris meter has its own
weighting function. This effect is still under evaluation at this triting.]
Artificial Ultraviolet
Ultraviolet energy in the aquarium was supplied by cured FS.-40 fluorescent
sunlamps, with a very small contribution in the UV-A by the Chrcma -50 lamps.
The energy levels were measured for both tables using the spectroradiometer,
while the container-to-container differences were measured with the Norris
device. These measurements were made only occasionally, and did not vary
unless the physical arrangements were altered. Daily monitoring of both
visible and UV during experiments was done by use of a photometric deck cell
(cosine collector) and a second Norris UV-B meter, respectively. As in the
rooftop case, data recovery was by planiinetry of strip chart records. Such
8
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records, of course, are scarcely needed owing to the constancy of the art-
ficial irradiation regimen. They did serve, however, to detect lamp fiilure
or other difliculties with the apparatus.
RESULTS FROM THE SPECTRORADIOIILTRY OF GLOBAL UNTRAVIOLET
Table 3 summarizes the data from spectroradiometry oF natural energy
in the spectral range from 290 to 360 nanometers. More detailed data can he
recovered from our tapes if desired, with a resolution of 1 nm. Emphasis
has been given to the clear—day case, mainly because scattered clouds and
patchy low coastal fogs introduce variations in the data which contaminate
the integrals in an unpredictable manner. Since the spectroradiometer scan
in the UV takes just over 3 minutes, time varying nonuniforniities of ir-
radiance of that approximate time scale invalidate the data. [ The florris
strip chart records, however, follow such changes faithfully.] Solid over-
cast and uniform haze conditions do not present a problem.
Table 3 comprises part of the data concerning the natural global (sun
plus sky) irradiance measured on the roof of the La Jolla laboratory for
different days of the year, times of day, and weather conditions. Recto
pages consist of six to eight columns, giving:
1. Date
2. Clock standard time of measurement
3. Integrated energy (Wcm 2 ) from 290 to 320 nm
4. Integcated energy (W .cni 2 ) from 320-360 nm
5. Integrated energy (W.cm 2 ) from 290-360 nm
9
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6. BrieF weather notes
7. Solar zenith angle at local apparent noon
8. Magnetic tape file numbers, whero recorded.
Verso pages give, in two coluiins, the Setlow- ieightecl energy integrals
(W. cm 2 ) in the 290 320 waveband, and the dose rate (in J hr ) in the same
spectral range.
10
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2 Day E?:po w’
200 500
L L J
2,000 4 00O
C
Jo off.
i9 r!l. Percent survival (probit) scale of anchovy larvae as a function of
total cumulative dose ofUV-B, weighted by the DNA action spectrum of Setlow
(1974) using the analytical fit of Green and Miller (1975). Lines are the
regress ion of mortality probi t on log dosage for the present 12-day exposure
experiment and for the 4-day exposure in Hunter et al. (in press). Open
circles are the observed mean survival for 12 days exposure; solid circles,
the estimated LD 50 ; and bars, the 95% confidence intervals for the LD 50 .
4 D Exposure
BO
7O
60
50
40
20
5
2
oaI
>
0
LrJ
0
0
-------
7
y 5.83 -O.00ND
0 200 400
600
800 1,000 1,200 1,400 l,60
(jo ) i3 eff.
fj ure2. Relation between mean length and total cumulative dose weighted by
DNA for anchovy larvae after a 12-day exposure to UV—B. Points are means
for experiments and bars are two standard errors.
6
5
0
w
4
3
1—
U,
-------
Table 1. Daily dose of UV-B per nanometer producing a 50% mortal ty in
anchovy larvae after a 12-day exposui’e. Düi ly dose given in
J •rir 2 (unweightod) and in (J •n’ 2 )D A off. (weighted by
DNA action spectrum of Setlow, 1974)1. Energy source
was rS —40 lamps filtered by cellulose
triacetate (CT/ )
Wavelength
nm
Daily
dose
J•m 2
(J.rn 2 )A eff.
286 1.3 0.52
287 2.0 0.74
288 3.3 1.10
289 5.5 1.60
290 8.8 2.3
291 13.1 3.0
292 18.6 3.6
293 25.5 4.2
294 33.3 4.5
295 42.1 4.8
296 51.0 4.6
297 57.3 4.1
298 63.2 3.6
299 68.9 3.0
300 75.8 2.5
301 81.9 2.0
302 88.9 1.6
303 93.3 1.2
304 91.7 0.85
305 97.0 0.63
306 100 0.46
307 103 0.33
308 107 0.24
309 119 0.17
310 142 0.14
311 184 0.12
312 224 0.10
313 224 0.07
314 170 0.04
315 126 0.02
316 111 0.01
311 107 0.008
318 106 0.006
319. 105 0.004
320 103 0.002
2954 52.22
1 Ratio of the sums of unweighted to weighted can be used to convert data
in Table 2 and rigures 1 and 2 to urnioightod dosane since spectral ir—
radiance is the same for all experiments , i . e.
(J .m 2 )DNA efi. X 56.59 J.trH.
2 Suni is 0.7% less thai computer integration; difference caused by rounding
to 3 digit cctiracy.
-------
] able 2. Percent survival and mean length of anchovy larvae ex-
posed for 12 days to various doses of U\’—B froiii FS 40
lamps filtered by CIA.
TR [ IVrMLNT
SURVIVAL
L a r v a 1
lcnqLh
(mm)
SE 3
lotal hrs.
Exp. No. of UV—B
(date) exposure
Dose 1
2
(J.m D 1 ,\ eff.
The an ii
of larvae survival
per container normal ized
SE 3 to cotiUol
1 0
(5-25) 71.29
71.29
0
1141
1565
48.2 15.9
10.3 8.51
0.5 0.85
100.0
22.9
1.0
6.02 0.93
3.65k 0.82
2.91k 0.77
100
64
5
2 0
(6-7) 71.34
71.34
0
747
1374
55.8 20.6
28.3 16.9
0.10 0.32
100.0
47.9
0.2
6.34 0.98
5.3l 0.86
- -
96
84
-
3 0
(6-22) 70.51
70.51
0
532
698
71.5 21.1
58.0 26.9
33.2 10.7
100.0
81.1
46.4
6.07 1.20
4.96k 1.02
4.55k 0.94
100
100
100
4 0
(7-11) 68.51
68.51
0
587
788
70.0 11.3
47.3 21.0
24.1 7.53
100.0
67.6
344
5.65 1.61
5.28 1.34
453 l 1.00
100
100
100
5 0
(9-6) 71.22
71.22
0
575
799
49.4 15.3
31.3 7.44
1.89 1.69
100.0
63.4
3.8
5.17 1.15
4.32k 0.84
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hiS 115
tj.i.nr.ror’.tii, rS.’rji Sri 7 pp00tt ’nt
I rr 1 iiaR)rJ Pier I . 1 t i ? trio I iii U. ’
riot I it 5. 7? i.,irki (Sk it C ti’ttpQ
E U FECT OF ULTRAVIOLET IRRADIATiON ON EGGS
AN!) LAR\’AE OF TI-IF NORTI )Lf-tN A NCI-IOVY,
ENG R / I U / iS A’fO [ . i i IA L(flit and ii Ceo I SLY Mitsu is
N,rtrou,rI Oceanic tort Atnirr pirerie Ad.nrarsriaiion National Ma ci n c Fisurcrrts See s ice,
Sn’uirs toherics C ct:r I .r Juilti CA 92tfl I I S A
tUnrs’Lrait) of C.iiilortira, Sri ISrigo, Center for itr rn iii blot tuition F’i nCc’,Str’’’.
La Jotdi CA 92093 U S A
(R i .‘d S M ares it 1978 nrrrj ted 22 .Tttnr’ 1978)
Attsti ret —/tnchusy and rtt,ic.t o c T egcs aid nt,-sae linac we’re exposed in u. r,rdt (run ni ti m trioacrtr
hand cr1 wuseicimiha i’cts een ‘SO it’d 320 n m, tile U ’s -Li regmuri c’f tin sp:.Jrirli Tm i,idi,ririin lard-.
were fiascO opon predicted UV—B flirt s i sis th at would result from aiiluopo°atiic drinr:iutioti of Eart h
llrotcctmtc o’onc 51)5 1! ih?sr response el itionsh’ps fr’r nlo’i.mtr’) and it istoTogici ! and umor phiobogic.’t
e!fccts a. c drier ruined lot tao sti’iire,mt spectral cn.’r’es en npn;itrons trsrn’ Ea-4t) suiii.i r ips ‘rd
two fiRer wimbmr,rrioiis Ait ilioss v sri arn Ie rcnsiiti C Oman 15.101. r d to UV—i I i it,i fist .tltCitrfl)
were ariab aed ri Icrums of i)\ \ c ti cc im se doses, e tire l ois C l atec 4 spcetm .‘i fluent e (‘m J 1 m 2 /tini) wilt
the eri e C) at each rtnt a ct:’iitd by its e fleet is’CUCSm C Li’Iil c to Tic Sctioo ‘enet it , ‘cit 1) A act tori
speetrotri I rlt of .rni_imos’, ‘t.it stied a cnrnt’Trrtts s O k \ dIsc us e dos. 01 1150 ito — o s ci a 4-dat
period lii the nut is m g ijesac irradj,ttiorr rmdt’icd I.ioris in tire br.riii .ord e)c,cau-cd to ti let 1 dispet—
stott ci i pr ’ meet a (in n mci nroihorcs and retarded f’i ow t t i aird dis dopms’nt /st lb los Oct dosage
used, 7(0 (J UI i -i grossiim s ia n rt_taic.cd .ini1 brain lcsiorv. oo-eurrr_d in .int_ios C,rictdattoris
of Smith a rid t laker in tti a rs no) it dicate that in at no a is ss at cr a si 0 i; li i .tnt I nerric tree of lesions
aird ictaid,ttroit of gmo’..tlt iii anchor) co ul d cretin at the surface at .r 2S2 rcdrii tort iii oaone and
doss n to 35 at at a 50’ restilci ion Egg-. ,‘nd ui .rc of anchovy ut_cut at tirese c t cptirs
IN 1 ItODLCTIO\
Evrdonce exists for lethal or detrnncnt:rl eRects of LV
irradiat inn ott Cests iittd larvae ci lushes ‘Ibis Ittero—
tore, rem r”ss ed hay Luskr (1961). cicscr tires efl’eels in
ft esh water fis hes, pat 1 etulari) tire satr,ron r 1s, a nuT
lot Is itrforniatuon esr ts on pelagic mar ne ursircS
Mote recently Mat mar o artd )3t tat d (1966) arid
Ponirner wa (1974) showed t hot p rotc eggs of mar-
ine fishes may be quite sansil is t_ 10 nat ur .rh U V irr,r-
distron Rarltotnetu ic me-ssurremcnts in p.ss t studies,
how ver, .rrc t n.rdcqtu.tte for pred sting t mu. t flccts on
nt,irune tihes 01 (te l C,tsCcI solar LV radiation tesitli—,
ung ft cm 4 intl mutt ion of t me ornnc I a) cc The obje Oti s e
of Otis study was to dcscrtbu’ I lie offt_ets of LV-)) adt-
atton on c’ s ’. ott O i,irs ac of tss a pelagic nmartrte VisIt,
the noethcrtt ante Imnry (Limrjrnu!mm l Imit / i’s) mid thtc
Paciluc macfleet_I (Scoumtaur Jtiperntctr-.) and to c-it mite
tire potential hat_at d to these species of uicrcascd
hoveL of soi.tr IOV irr.ud r,ut .011
1 katlt species occur tim tht ’ c and 1.11 v ol stage ’ tnj
the tipper I in of liii, ss.flcr eoltuittii ( ‘slil-iteont, 1959,,
A ir I s h our and Ste - i cutS, 1976) a nO are. tke’rtforc,
csposed to sio,itiltc.ritt lesebs of LIV i .‘tlt,ttr srrt ii tltc
sen 1 hose species ore more stutrier aisle to the dir ccl
eRect of LV it i .tdmatton dorung the crsg and hor s .tb
t h _tn at a nv other tunic baec.tusc oft heir i ear_s or—
face dist rihittie,t, in tel iii. ), back of seaR s or other
Inteetitnent, .-nid I lie foe I html tire sensitive Pt OCc’55c5
ci cii g.mnogcncsrc ate Irk tug place
En Otis stud) ac esa’nincd IlIC eRects of LV urradta-
iron dnring t ue cutibty onto per mci, that is, dun itii the
Itrst 1—5 d ays of life ss’licut tire fish Oct-it as eggs and
sutist) lars’,ie (7 I mn5) subsist tag ott yolk At (its. etOac
of this per nil, ite C) is become jiugtr’cntcd and liurne—
ttonai, the jaw b,’ec oust_-s iemnctuo tab oem ly all the yolk
is e\haosbcd. aid feeding hcguns
ittCtt 1 51.51 5 At’cD Mt ittODS
I pptirammrc A tcmnpcratimre-enntett llcd wlttte fiberglass
st.’ien rabtt_, 360 cu liii’, 122 Ciii rrr,Te aid IS cl ii dce 1 ’,
w.us tmmstai!cit ott iT’t_ roof anti tiso sssre nt-,t.tlhcd a the.
itt ictite ai_jriar mini of tire Suit’rtincsr I tshcrrt s C cnmcr .‘t I
Jolt t CA t .reir tt’le r’ rc ailed into I it, Sent tn isa
tint .‘eti ti.ttc .ts-inznctt a Iliiirltld north r In di rttif 3 ii
fire ra 5 irormietrrr. c,rliSr.ororis n id .is’o;’ttmcnn Of t o-i COIl—
t.nrcrr Art alit ritiitmtt Ir.titiu’ttoit. enilosuitt’ tire nrirft’ats
siiicr r,ihts- u S c sri i, siiu’l putt lit_ti smrppcvt it_ic tutu i’ceitr
Ii ’ s I trips 1?’ sImi .tios r i tic st is r sitef,i ( I in ii 1 tic
fi ‘rite is scud,’ 0 tsiilr t cri,clt (‘ h r’ , 5 ,rmmri .rii uctiipeia—
im,ii it it tr it ii. s at_ I c ’ .iiu vu. r 5 i’irl.itt_ri urnh ut .tri e.ririti—
totter am a ] tir nil I ins St_ru, it f r i t i’ ti ,itt’tflitisd ,thrtittt 205i
• 5501 t stlpp’intr 0 in itri P 5 Conir ci fer mi tilL I a’ it i r it
tur’Imt,t )‘iOiisliolu i S t_s list, tirrt!tii_’ii_ i i .tiiO C luiii,rtic I t1 & is
kes.iri it I h\c_ I lIt i’rt’’t rat
-------
7
JIIII”t l lltr iii,, li it’,, I I ‘iA ’ iI(ii , t,l II ( ‘,iiit!iii’i i .1ocir
rs -40 LAMPS
TflLI 1MFNVI —
eli-I,
I lflSltC I Dia r ,ni (if ioof lop f,ici itv (soiti mi) I1iiKtlfl placemint of liedttflt It CofltauilC [ tirliliciti
UV—1-t souIcLS, .mnd IlOith-SO ltil (i l fll 1105 of tl( i ,.Iiit Libli Int’l chcic’ s esploded sic’s of i ,m o iduil
trcal huh contaiicr timid iscocititud lilturs
of thu colic U\’ It Intl ii ittsm t.tud 9 94 ’ of thc nt ’! urd
radia ion hct ’ i ’ iL cn 360 _ini 700 inc
The hai h of fluioiccuuiil I imps cited Iii (icc. t.qu.l’ nun
crperiict.ns s.c. cc mriiicrd on Iar’c rcinfurccd pi’. 55(01 !
5iiCCtc I’cninit_’d vitli 11th ror—poii hecI ilictinimcii (t\l k t ’’i
to pci ’ . dc opt unhlni rc. l . cli , e cfl ic iciti, for b ii 1 ‘. i’,ttsic
and LJV v depth’ lit the col.ti ILIflI, hc’iccscr, the UV-
scippk niCnt in t f S40 liii i ‘ i c c, t Itru ’ . ided ‘isitli I flcc-
tars, ii order to riiinimc,_ sh.idous oii the Ircatilknt tt’hit_
In tht icjii nih app ii utlis, visibli, ii’ht ‘ i ’ ..LS provid ci
by Clicoina 50 ’ fluii’i scd ilt iinip . ‘ i i nCh Cl’. i 1 Liii
appro’ini II lc.ci to ti sb of I h _ cc bk cot_il SNctrum,
und thc 1_tV csc.r b) I’S-tO fmuoruccu nt stint iinp liii’
tube3 scere nlouitcd aim S tO cm ccitt rs es Ii thud lamp
a IJ\’ source .iqti lIiilTfl hi Ihtln ’li uS \\ilC adjiisli_d to
cilh’’i 93cm or 61 (Ui abuse the ‘isiter curl_icc ci the tic_it—
multI CUPLhllILIS lt ,tIorc I cc .01 I.iiti ts ‘iscre tmed Ion
appu (is 200 Ic in ordui to nu_icli .j ccitt it LI) flit porlioti
of the en_lit) d id_i) CeCiL siippIi d h> the tn’tiiuf,’ctuii,r
I: r.dirunue of tb_i tic urneilt t cbie ‘i . i d fmsiii poiiui to
pOluct All hjiit’ii thi SO (I f Tc LnceS ‘iLl ( lilt It, ‘i , tilL) ‘ ii CCC
carc [ u t i) uc)Licurc 0, md liken ad’ _inl a_ic of ii .iIspi (iprlatc.
piiccmelil of IrL mticcent Coiit,iincrs on Illo t.ihl,’s I c tinplcs
ci tit non—uniform Ills dcci ihcitions I ii ,irulic i_cl Satiric
1110 .t’ill ill I it 2 sulitili ‘.hii’i’is couc mptit_r ‘ tiiLr itcd puts
of the COLCI for tb_i co’ii iiini ,iiid cct ‘l iii.. ,iquiriiit’C
tilsie. Addctionil s’ iioutlcuit ol LIio Contours Otis ii hi_s ed
by ciisering the c tIiciit intl w.’iisofth _iqii iciuni room
‘is ith hci ’ . > ,iitimmtiumn foil
[ tic. diii.ition .iiid uiltelIsci) of zcrtilii ,i ,ui irr.uit.itu ’i ‘isiS
uoiitu ili_’ci b cs ,itcii,iirt the Ittcotc c.cmit I In Ice
uc 1 ii.imiui’s clithics, tb. ( hcioinu SO cciii I ‘i—lU i_nips so_it
‘i ’ isutcli iil in .u,, tudimi_ic scitti I u 3 I tic r.itui,sn it_i liii this
p_illicit of ilicil and 0 tI i_I sltps i ,iL irucL ac.iiiiiil of
(Iii, umctui.iI he ,iicd fill of r.idj (it Liii ‘\ 11.1 .1 1)111(11,
(I_i). ,i’, ‘isehl i tin chsiir 1(1 ilicilnnu/( .ii lV ticiiltld ti tilL
tinim,ils I u itie 3 .il,o mdii it. tlit ippiti’ Un_ilL C li E nt
of Outs lit Illicit otis it n tim_il i l 0 liii i ‘I souncds Oil
conutiiic tt in t 1 i L. col it ‘miii All iii,itlt lion sth._iuii sscr i_
(CIII I on neil ci i minis
J ’ u iIuiicittrt , 31 111 ct’s, ond j f Ira V.u bus intc’iral n ’t In—
Sli ilniLull (Sit itiiirn Incter —Soi,ir I ight Comp_imt 1,m\_13
u.md iocmi_itm_r— Opiro iii Liho atori_is cluck ciii line’ a-
0111 iii) ‘icce tiscd for piioiumi’ti u_i’ and rchionic.mi IC deter-
liii lIon’ dnnin thc, fi r_i months of I is_i_itch bc.c_iui’u of
lit — lick of a U V—scnst ice sped 101 ,idmomn_iu. r It ‘is tic later
I ‘inc ‘ (‘ilnlptilur—t’i ii,,t_iui(I luiOt” ‘,hi,v nit u’oii—uiitfiir—
nmtV of iii iii hut sour tic_icu ili’nl lillis ‘ishcn till kV
lint, .iiu lit Fir nIh_i ml 1W—Il Sinitis i,il’ti ’ i ciii! ‘i’ is
9! tin in .icii ‘1111111 1mb ’,, nEd I ‘2 ciii lu coliriluim liii.
,iijii,liuiiiii 1 iii. ilut sliii\ts liii. t ti t il I bill) uiitl i_uilri t
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AQlJlJiiui, l
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00 ) 5 05 . 110 :503 i .Zi icc’a :150 twO
n OL I3S
t OLQ ( IUA
Uc.ti oico 0.15 ‘200 I 05 1153 4Ct’v
noun s
Figiti_ I rhcui die rcp,cs n zktion or b c ‘rad tio:i
iC iii iCii it , , aqilarlilill a.ixl T o cotat 111111 ‘ h.’.icd ,,nas s how
UV—}l tti IfiJ i (ii i elotic lii ih 1 cr1 ii ti ’ alt S aider (ur ic
e c u Its fii’’ii s i . Iclitag ci U t’— b I in ps to so p d.mc ii ;
ii.tui at (oiii oucitt (.Ica tindcr dashed cut ‘ci [ Lo ah
OftjiIi(Oi liii
poscibte (o coast sic I h test ti so a . 1 id to 1 ecatifir a C
ci’r 1ff i’.tS c’ It Hi Int l us Ut’’ S I ecol d ing spc c ii i ,ittioi actcr
(Ciptionie Isiodel i L— I ’) 1 Ins Iii ’ .tto’s lcl; ‘Ciii ‘cii aii,o-
ntattc-.ihly from 250 i i 8’JO itiu 0 tilt a I ri o c,iiioiil from
250 to t OO i ii cid 2 it o readout at loost I t’ cit I c i ’ ti c
Watch, iigii accoutre t sin; .i lots prcscliic hg inaip tot
rahbr’ucn ii t i ± 05 orn .1 id I he cpcurc’r.’itio:i1 5 i ii a cu-
rccy was ± 1 ‘ , I tic d’ taml i 1 .10 5C 5’ S t IC S ci dLC,I’IL’c
to uteii.i. oicii I5 to I he use cli a logo I dine .ili}ihfier
lot c i tit) c libra lii i I ii c it dc’r e b Ose of a I fiOj 59 to a:’-
itCn—Ii,iIuochi si.iodartt Icit; ot larat cri at S D Pi) ( — I 0 t2’ i
A on 2- tn p!lolotic ltic iieiicli ,tnd 0 tr.ie,_,’liIc to tt,c
Watioti,it ‘.ttircsu c i Ste ,da,ds UO,LS scare edict iii i’icd f.y
nicastir ios i ii : ait so lnic ri tdi filets (it t’’to i i i, —1) loft—
gtatU froro 255 to 100 111 ( 1 tiiuhiiphia by tiic do,,itioo cif
the Ci )1tiTctit it It d O ll rspiccscd ri 3 itt
c c ’s and larval it ch ‘, Lit. irrad iatcd in two is.ryc
ether iii’ti tha i Tinhit cuppienr.iirsd by a, u S c, d US’ i i ado
e l ton (sot iutiiin sp lcrn), 0, by .t couubii,at lull of US It and,
Viti I d e I “It I i oat atit ( ic ii Ut’: c’Ls (eq cat taut a, ste in)
Na ltui ii I tdc’iittii i i i, risun,’oricI, both ins ult and otisicte
the canopy of the ic’IOlislIpi and Its ip_Lht.d chin) d’sti
but ion ,i.tca tot’ ccl fo’ d Tic r i o , ii ass
Ooc-/ pot ‘c opyitnic is_ake, cc ’ tied u’Ii Ii lii , c i , tape
on the c’iqts,cL :iil cc’flt)iO,ii, SIJO ,ii of filierecl scz’it.utcr
were used as icc rotinu,urrs k wh , br.rl cr si.ic p c i :iicd
with: a cii ‘sitichi teLl t ue trci’tIac A ecsnrtoi fitter (or Pt
Iris) api’r.p-l ‘ti to a gis i n eoiidui,ctui (lie I)
V -i ric’us ‘‘ ste in cicri is is .rc usid :0 Iii a e qi In coit-
trot of tioc,otrs aid to present c’.’apc’t t,ioci iii 11w c cii-
talucis I by isc:e Pot>st)reic.—cte, r shect, 0 I i flint
flock, C i Iniose Iri,ictt.,ti_ —ci a, sticet 0 13 ni,:i, Aclar®—
clear sii.tl, U I I o lin Pot3 rl flu 658-if— niobdech discs,
097 coin, Si) ‘r—-eic.ir sheet, 0 3 Olin
thuthi t1cit i)rcni_c awl rtIlnL,’,e in cci i c sserc turd to
pro ide il l b rent st.ot 1 55,1 ic cur oil clii. ecu em it the
UV- I i sj ‘cetral rc”o’n IIic A riar”, is hwlt thin’ ill’s isc lt j
itt the U S it rc or,, si as t n t d tsa coon ni cc apoolto ‘‘ I s’- Iii
nct’hi,’iL’!e ,:itcfliatiisn of 1 19-it \it lii, tsb,i.h t_tia,iittl’,
butltt. 119 Ii ii is t Out 1 f l sod,t sti c’is to eoic’r hI., toiitri’l
coot turici ’ ( I it’ — i i the ,iiust’r k 1 I i in’.nti,taiuce o StyIir
in 1 1w U 5’- Il t’,itiit 285 ‘iJ ititi. n ito iii t7, of 1 ii it far
C iA wick: I 5—It) unit’, ,,o,l m ti n itch ‘hic 5 i tu, (tic St it’ll’
t3NA action spectruiii ( a s c ii ‘ml kIti1 1975) di 4 tIcs—
Inc tt.lfl’Oliiitatlcc of StyL,i is (IS’ of th w fit C IA
iii rspc-iuincnl i i do ’ . kitls (otticr h ut l i i i ’ ‘i tOO”,
of Intl .1 1 ni iii utitrii’.ii)) oLic tolutmiult, it by nt if circuit
St t i i lilIti- , that rent ‘u Ii,rattd I,, ln’ti cf s ‘onus site-.
.1% ‘,tcii li i I ii’ I lito ii,,’. ,it i,i tsf (Ii’ Sc liii ’ . s’ ‘‘, d iii—
by to’ 1’, il ‘in is it I i ti , t,,u,,u ‘ci ,‘f Ii ’ ‘I , and ccii
tcr—ii (cit r di.i,iiir,’, ( cit ttlitsi,ilti \\‘Ii, ii mid siidi ‘ii
Aria i,iti m ,ii LII I Lii i of lii. i,iuu,c .ini .t cist,l \hil
: 1 5 tite oiul,l ch ic wat’’c u I Icieclit ‘cs c it inet,icnt tJ\ II
tins ‘as 910 5PtS - iVY, 200 JtS, 59 iii !, - 0
liii o ’itat pce,iii, ii’ icsiidi’iy 1111111 usc of s_do lid
flitcis tinchi,ccc iili:i—’iiitf,’ ii lti (if irr.t,ti diii i losi It,c
c.,itte siirf,icc / Ic LILCt I. rit,ilts’ far liii t’i’tcl so!,::
cOtlipolic lit (itS it,uttir,ii li\’—ll lists I torn —ii ,iiisiui’li tic, lit-
ter’. hut coa’i:L,.,t Is L.s Lii t stciu’Ld stout, s (sk ihuii,c-
atid [ Turn: weal Ioo,ui I i is I .oiui , :‘ I it’lL —,iIes
15f5z 11’i-ilIc-? r/i’srqiu !y’’s of urichu,,ss and tim ii- 1’rJ Otis
01,’ ,oncd front hi ooul s,oet ifl 111112 ii’;rl iii IL p oa!iicm se
eo’ti.ttost throlQ’i.omti th u ie.a at lii. Souihiscc’,t l u—li. c s
C u i u ( [ tour ‘ i l l, I ‘ 7 7 I In mo I t spirimuc iii ’ . eg” . s_Hoc
is_Id fir 71 h u t tO C I’. fore ci 5 ’d, it to itifo ,’ in, i eric
of o t to st I’d r u ’ tEat Ii I lee n L’tt . 1 ad ruin I rol (ci’, fit Ii t
of iS t c ecu,rt,i:ui:rc cirmiket si Iii cit c ’ ‘, cit I-, Arc,’. of
sinujha , tiitictcnt UV—!t iii iii Is_ti cii cciii 55,1(11 tide Ot,uI
c L iii lied hi) 5 ( 4 11_it a ,s,tn,hcr .1 ttd 1 rcat hit ut .1 ti oriu c’i
Cuu.Ii,ruiCis V Cr .55 1’s_cl -i s5iier_ mu ] diii ihicce It-tub an
a r.unrlr’iit basis ‘spcc ific IL. I -dc Oil .deet-_ci for spcciIl.e
t,e.utineuiu; to hhucrsasr. iinilr’crt’iy 0 r ttos_i’’c 551111111 ,t trc.’l-
lctcuut conti51-. ss, rc iI iii failed elhihfttIt In c’ificii ‘it fwids
ira ‘.u iiui c. tcsis uiucnt -.cil hit .i (ic-Id cc is I uti,t,aiit A rc a ’
of lists ii ‘hiiti:j crick c s liDli _ It o crud-s ala! , irIs or
t,’bL. on is_ic ith 11w ,nrr 1 of lamps ta the s_cat aqtiaiututn
I ut ile sci: ci as mdc . 1
[ ‘p. imeiits cs_lcd at tic close of tIre moist) os_c pertod
Ati iuc’s-y s_ri c i s 1 ie’std ho, atrioi Ii tO ii i ii (tue egg ‘tige
md IS 151 I i , -, >t’,—s u i l,uttcc, rind i.iid,e.eI about 6—t6i
as c ’s and 2 — 3) I i as tars_a: 1 urea r spi I uncut aI Los_ha—
Iin uc 5’ cii usc-rI in t b lint. 5vL used the sc’t,l [ lsi.il eiuctos_d
I’ ) Sent ii Clcucs’ 0 , t I lt Soul Cc of (IV Lw rpj w.’s I S-tO
It: p s Shred h5 p a bst ,Iel10 cups ci , lii hc.iI,cns. and tire
Cc It t o ic tic ‘-dO etc of it i U k citci to (a Sc tot) t sps I and
I rititic 4 ‘ l5 ttr ii iiansilt’itaitr,_ rtii’.-cs fist clue pIistu fiitcr
niatcr, its intd it it t tst, hint c i ’ . I lie sti,udicch ri rap-
less_ut t ilt tcialtsc tint,’-. h Sen ito ity of IaN ’S is a [ tine-
lion tii tic t1uii’ uli — I
L4
it
I —
-ç
5 5
I—
ace ate ca-a mc i ace 30 320 a’o
Is-taint r oCttt (-‘c, ,ucu,atc,th
-------
P 540 Clirorna
50 665—U+
CTA
0 i S 4053
4,300 248 300 6 192
3 6 l e o ’ s
733 476
= My:-. PS = Poi s:vre- e, CTA CclIilcsa tr’.c_tate, 666—U = Polys:’.rene 6(6—U
rr PS c’cpcrlrcn:, :95—3 :0 a CT \ c-flcr mc:
1 = Mea t flJ C ? 5LY1’vQrs ‘cr eor\ircr, s = S,’’t’Jard dcvat,o
P 5 40 Sal : ’-
Table
I E rect
or uv
or’ zrci-ovy
d’i’-ng embryonic
s’r’gc
Tom!
To ’”
US’ lo t—p
L A
i ’ n t e
Mt ar
n.,r’aar
s irvi’. -i i
Eta’
“a
r)
UV
so a re
\‘isibte egg
source F i i er ’ h
ctposj-e
larvae
‘n
L a’p
etoosure
I’
oos ze
meter-c ‘idle
I i
Dosaw°
S n 2
Nt -rbcr
of
c c 1 ’cn
or
sur tttors 3
C s
.lor,r IA-ti
tO
cc- tro ’s
P 5 0
Solar
(5—2)
( 6 -7)
(7—2 5)
( 8—I)
5
( 9—22)
6
(12— 19)
F540 Ci’ro-na M
50 CTA
CTA
CTA
CTr \
FS40 Coromo ‘4
50 CTt\
CT A
CTA
CT’\
F540 Chrema ‘4
S ’ S CTA
CT ’.
CTA
‘4
PS
PS
PS
PS
0
000
000
1000
1000
0
1500
1500
1500
1503
0
2500
2503
2503
2500
2000
2000
:000
2000
12000
0
i3300
26900
4 1 ,,00
6 6C0
5
1 5
I S
3
5
2 167
1947
1(7
577
033
667
55 1
809
500
072
1000
39 5
539
435
5
‘4
PS
PS
PS
PS
0
1000
1000
l O GO
1000
0
1700
17C0
1700
1703
0
2700
2700
2700
2700
9 2:0
9:40
9240
9 J0
92 0
0
14200
294 ( 0
44900
91100
t O
15
IS
5
5
:375
30:6
19:0
650
O l3
759
s:o
972
326
062
t O D D
93 ’-
524
215
I I
0
620
620
6 :0
620
0
2 100
2j00
2100
2100
0
2 20
2723
2720
:720
4 -00
4 ,400
4”0
4’,00
3720
0
6 D0
6-350
12000
162000
iS
IS
14
15
15
2540
2310
957
373
03
551
5 i2
58!
5 9
052
1000
909
377
147
05
0
525
525
525
525
0
2100
2 i00
2 1C3
2100
0
2525
2 C5
2525
2625
‘,c o
.930
3010
7910
3 160
0
62500
90,-CO
uSD00
i’- 5000
IS
5
I S
5
15
t20
3907
2050
547
033
278
282
(‘07
603
13
1000
9:5
7—S
400
08
0
(-55
658
655
0
1975
975
1975
0
2633
26 3 1
2673
43”O
4 1(0
. 11( 1)
4,350
0
72000
109 C0
1u7,030
IS
i i
I S
IS
2293
6e5
‘- 7
113
SC ’ -
427
1 !
06
1000
255
64
‘-0
0
600
0
2053
0
2613
C
7
>
-------
Tabc 2 Efloot of V on rn c zcci d c-r! rvonc sn—c
• Mv: r, PS = Poi st ,cne CTA = C c l os rLxcct. c
75— 0ii PS c pcrri crs. 235—320 n-n CTA csoc-r’cn s
= \ cz’.t nu’ -ibcr ct r :vos pcr co a,cr, = Sn’djrd !cvja’ion
To ’1
Tout
Pc-ccf
UV Inrp
UV
vsbjc
Mczir n”nhc
su-.t a1
E pt
e7po rc
L ir
d s’ e
? i. rncc
‘0
UV
V s hic
1 r ac
c posi c
nicc- ‘-Uk
Dos’gc
of
surivors 3
to
(c u c)
o r c
so ,rcc
Fi l icr’
h Ii
!
1’
J n’ —
co L crs
S
c Or ’ irCIS
1S40
So -
M
0 0
0
H ‘00
0
5
291
2( 0
1000
(4—:5)
PS
PS
PS
PS
1000 1558
1005 1553
l000 155S
1000 1558
2553
255 8
2533
2558
11400
11400
11-00
H ‘ .00
25’C3
23400
65500
111030
15
iS
5
IS
093
4227
0 702
000
417
443
985
000
9 5-
525
00
2
FS4O
Chror a
M
0 0
0
57 0
0
iS
23
1 )15
1000
(7 _ 9)
cc)
( ‘T
u\
CIA
CIA
1000 1750
000 750
i’ (5 1 7 ()
000 1750
250
‘C 5:
2 c :
7 :50
S7 ’)
S 7 0
49—1)
57 .i)
27r 0
50(1110
I
2S 0i0
l
5
c
15
‘ 1 ) 7
7 7
027
000
:. -
u c j
000
1
‘ 7
1)1)
3
FS4O
Chror
\i
0 0
0
4.730
0
15
SOCO
714
000
3—9)
50
CTA
CTA
CT.\
CTA
775 725
775 1725
775 725
775 725
2500
2500
?S0 )
2i 1)
47 0
ir’ 0
477 1)
i7’0
(5(03
0 , ’0
0 41)0
)2 1
is
5
1 5
;\5;
H
47
213
2 9
0 -
; ,
980
1)06
1 i2 I
2
4
S40
Cironu
81
0 1 )
Ii
4 1U0
0
5
(‘ 20
276
11)11(1
( 30J
SO
C 2\
CTh
CT\
600 2100
600 2100
600 2100
2701)
2700
2700
4 —00
4,- . OC
00
91 700
125 0 10
77000
5
14
.5
3171
ICG O
(07
95
3 1
32
75
411
75
0
r
-------
S i
JOHN J . I)t. ’>ilLit, Joii’i I L IAYI.OR aid 11, (zt ti toY t\lo Ij(
‘ 2
• - r
—
3 3
“ I
F
1 i’ure 5, State of mehinosome di ’; ’cr ion in Mylar control :ind UV—1 ) iiu:tdiatcd larvae (Tr:t\ fl to
scala from phot oni u aphs. Shaded at k -as on lar ’ac ir.d kate areas on Lad) used in aticho y for
radinn. NumUci are !‘redes used to qilaiti Iv disp ’t sion: trade I ivie; tvpkal of controls:
2—3 occuri ed lit in ;tdiatcd sreciniens. R i ht and kit columns of fi iire indicate the range of di pcrsion
St it cs included W t Inn a grade.
ii
2, Tabk I: mackerel cxp. I, Table 2): in tlIC .conil ,
acluirium t;ilde were u ud, the source of UV CflClIv was
I - S—40 l:intp fdiercd by CCUIII&’e tri;ti ct:ite (CIA) and the
source of vi ihle enemy u,. Cliroma 51) Lin)ps (nnctio y
cxps. 3. 4. and 5: inach .crd esps. 2. 3, and 4): the I liii d,
the aqttarilt:n table was n . d with FS U l:tap fUtcied
by poi’. i ‘ne (66-U a : d (1 .-\ (anchovy cxp. (d.
Waici 1i ’illl; ’ratuJfc in the containers, and I ,JVI) and
ii%ible flux iere tnozlit( ’:ed cant muon dv, atal sit tittit
oxyi n ant a tint tia at t ..r t ie ’iii i in’ antI cut? ol cacti
C\jiCItflhi ,’ttt , ‘ : ,ihiiitt,• sari s: Itoi 33.0 to l.0 ’’, , O ) L i1
Ironi 973 to 71.2’,, s ,ttIl,tjcn, aln litana Lain 111)10 ta
O.0 )l lirtu l in&l teinperattirt’ from 16.3 It 17.1’ C. Sitrviviin’
kit tic were Cotink’d at the end of art cxperin)ent : sonic
6.’,ccl itt l otiin’s ,s ,ibit tin for ltistoIogic;tl analysis, and the
rest prc ciucd ii i 3’ , form attn.
1 ut mjlin -piesuived specimens Iron) C pcriments
mO C I’. ’\ ft tiers a t -i C çxaiiiii ’d to (IC(L’i flijflt efrects
of tiV-l1 Oi l tint: oh , i tal pignentaHon, V.’e nhi’astiroJ
the st,indard heath, tu !V de .th .it the (it h’ .in of the Nc—
total fin, maxinalin dcHh of the )I).—ne, C\’(’ pil :TIIeii—
it, it id c. tetit of IntrO di .rersion in nc ?tt )ajhor ’
of 32 S5 Iaiva ‘er In’ fan’it. ]ypk ;dk, fl ’ .c larvae pirr
C i It ,i ncr a it hut a ii i-it metit u ct -c nie;tst,rc j h) ta flu litters
,u :1 v ,tI t the ititinhet of 5ttrvjit , l.ruru:ic u’.erc as,i ’;ed
a t r ,idt ’ of I to 3 (‘ I i 1)1 ‘ Li; j , of c ’e pii:iitt -itt;tt ion • (Jrndc
1 rim
1
I
2
-------
i lb t t ii
I, otil i li,i(i lSf pi titi itt (ii [ ‘ii’iulCitt thu ihitit_il iaiiii_l
0111>’ tttt 0 ’i I tii tigIfl Cit tilt ( ( I. ‘,idC ‘ IiiCt nidi ‘ C
pi t ti tcnt. i i i i il. it ‘i d )t cni’ 1 ’kiil, pinictitid Situ it a!
(tIL of clisput soi l of iStc n ,clttii chit_s (ph i i il ‘ ‘ t’’ilhis)
l cttIii n ,iit t4tot v is si it J I ’ ) .L si ’lin ink
I to ihc cc,iijal> c c lid L) ’1ihiO}t il i.iJc 3 in
Ihc ic Ily tlispLc - i_n (I I ’’ In ,titLl,tiVi, 00 iC Itias Ill’
tIt cciit , ii I L 1 of ll’i. body to ‘r. d_’l, tiid it —
co_h, tiiL ,us Oi_> h.iJ nore mci iic:ihtoic’.. 1011r rc Oil’
cccic itsid ili’_ dot- I surf tLL of I I. hi . d ‘ il 1 cic 1110!
fit, tOll SLhtil fl tt ’iIl of dtc I
C oI u 1 h h2 tti ii c c , iii “i_Ct 01.5 i_it at ¶i (i/thu .2ic pie—
ptic t.I unit i i_It cpc_itiicl ptt_s_rc _i I C ’ lttsi i,Iit’ IL_I 101k
Slidic Irtini s ! t d cVxrititLI.1s ‘ ‘ic is.hiucitcd fir u k
it iin ,hcr of cicns I I I!ti_ Li tin ‘ti_I nucI ,i , ht>i_r of II , . ,
iCl iit ,i I aI c. IC _It_ c\.tmit l (1 V it I ’tii Ittoobdue of tlt_sa ’c
to coid tlttcrp ’cl.iilc_ prcjudic I i itn i’c li_ dclin__d ic
COiltiliii_ ’ of one Or mcli SphICI c .il p’ ct oLc ttii_kt sui —
rotindcd Sty iii evit i’ccIh ,iI.ir c.cool t,’d ri_ toii, fic i ’i_tiil’,
CoflhillIliiif’ cisiniipliilic cvlopl.tsiti. ililifis i 0 prOc i ii. iii
ifldex of i_I in ye .o i 1uiico ti of dis cse CCtliI’Ii ii tiC
nui’ibi_r ci lesiol c in tlt ,_ hr ,iit , n’I c in cci i i_ili0ht
of a I,irvi rind dii lcd ic tot.tl by lit. nittOtiLl of cei_I ,oi’c
cxdnltticd Iictdc ice ic di_uitted is ih’ ’ .icit,t _ niiiiitict
of kstons to . Is Ut_ r ccc i wi fIic rumltcr of si_c I m u
c anitncd pi_r l .i’’ i ii id, r ti cc b iniIioi IIiI
1 —40 , Sir ,iti, i— i ni_I 1o nmji_ t ,crci t_> is, 4 c—S’ l , -in,
4 —50 Typ’ 0) 12-la lii V ‘L irc iii ,cnt cct’rc t_ .ii,iitit_J
bitt feocu In Sic’ ‘uCl’tc 0m1 1 1 lCi) itmic Si”
Sitrci al it ilic ettutrols ci l,’! tn.on ’’ iidiocy c’p_i
1Oct05 11001 41 to and Iroi -i 1 Ii , ‘i l u,
hccausc of diI ’ienccs ifl CV Vi Shiv “ n c.u cpi\’ns
(1 thks I intl 2) To nuljii’ ,i lot lit i, (Ii 11 2 1CIu,t_’, sc._ do idd
li i i. pcrc nt Still va’ tn lIic ireaiiulcmi - Si > th ‘I of the i_nit—
trots ri tacti t c lct miilCnt 1 liese n ’i ,iii, d curc’ic ii dita
Still used to c - lril ‘In : dri’.c-rtspci’nc tote usim’y p
an .tlycic (iinii_c, t’)S ’ Adjit;ti.ti_ ii ,_ licatin lit dita uut
iliii . n initei uiIcc L:iscs liii sot coti ;troiialiliui:s, .titd ii’ ii_—
b> dccrcas,.s Ilic I it flCC Th util ri_3ii inttc ccas r romoI —
tioii_tl to tit: ticiprocil of lI’i_ pcici_it itirt oat of ili, ion-
trots and SIC II.i’t_ Ciii ni_lcd ilic ct.ind tuii d c tattOt , h> 110!—
S ti lt) tug b> Slits 1ucior
>
>
1
I —
(-I
I_I
LI
Li
0
10 50 (0 no tOO LO b c ) ?i_0 210 2 (X)
I’tcttrn 6 Sir ccitt sum cm ii of ito, tilt_Ill •IihLltO\) it tii ct id of Sic ittibi> C’ntc prriod (lii ichit cc
i ,ttI tot ii bull luic i si_ f U \ — I> (lit’ sc,ik I i i S _ I it hi lIt ii. p_i I I ii n_i -y it hut u,iiic
I ticti ’) S (iLi!i_t 5 I IC 5 “I fl I uli’fi , V ilit i_s buS’ hit _ i tilt fibi’i—. ( S ,\) pi_i 1 st 1 it I ii’ lilt_is (PS) iiitI
pitilsi) ciii 6ti(t—U t C_i A I i’i a dli it it. —‘‘us oI ullilil ihitc jnt.lii Ott iii’ i Io ’ i_ huh jitItlils dli. Itt.
itli_,tIl cii, i. il lie IS i 0, 1 1 itO r’
III ‘.111 IS
lit (i\fiCi itii.ii’il’ . co ,ilichi ii Iii the iiiit’i tim ltSItW
I S—’I0 li.t’p) iiiil LLhIith ’t 5 Ii iliC&’t.ite flb t_ rs (( IA)
co’ of lii iii I locy siiictvd .1 ( (u nIt d lv i. IJ\’ 11 (2 )iS
to 320 tutu ito-dc l 91,200 mm 2 (95°f .
C I — 0 ,i cY) - I ‘dd,fiOO I i, — ‘) oc cr a 4-it iv pci ad
(I’ict 6) , 5 5U , iii it t C i ‘ ttL\ c t.l _ çt imata—
tic e UV-i dctcai. oh 125.000 J (95 1 1
C I = 9S, O0 -1 -1S0(’() Jut ‘) OcCI a 4- I c 5-dc
liii uid (I i 75 \Vltctu Ihi, cmmct d 1c ftoni S— 10 l_iimtpi
cc is lilici cii by poli_styt cue (PS), liii. tt) tu.i tin cit
t1 luccct dii ag s Ii tlics c\ !tciiltmclilc 50 of thc
1 tnciuniv > suti i.d ,i c_iittiuthtticc doc ,t’c (:i7s_l’bn itn)
of 31.000 tilt 1 (95/, C I -= ?S 000 3S i)c) I ni
(Fig 6), ct md (I ll_ I from ii ci 111. C \ 11C1 t itCi it S iig x rsl
(toil SO ’ of wicket ci ii c tbk lo Sn I S ic C ii do’c of
1 iiio itt 6 5.500 5 -2 occr a ‘ -di>’ pci ion (T.ihdi_ 2, C\1)
1) Diliei 1CC’ . tui the clctcc—t cs cotise I clii toil heicci Co
c pcrttn( tIc cmplcil ‘.ti_ (_ IA fiSk_is attd hii_i ,c ttct’ nz
I ’S f’ili_oi wi> bi_ 1 itht h It’d ho difli_t CilIA’S lit thi spcc—
toil I ,ln ’ .’’iLcton of ttv lcd hiitats Thc PS Iilut_t Itinc—
Iii its abust 30’ cmi i hi. cnc’r > ti 2S0 nni c’ ci cas the
C (‘A Ititci Itismits less thin l , (I i - 4) 1 hoc, Itr—
sac and ci’ s ttt ,dt_i tioi 15 hltit i CCi.I\ i_li mate entry
itt tiuc sito’ tot Ohu_h noic ‘CIiO IC c’acelcttgtits of tIc
U\’ bttit.i Utidi_ i both CX [ iCt flhiu_tltll comi’h iltritt ,
ancitocy c’.i.r_. moic scitcit st ICI (JV ItI,uhtdtitltt thin
mack. ci 1 ht_ i_S )I i c ,iCc in tttc 51)50 cc is :tbottl 3 1, 00
1111 2 tb’ 1550 c\pct tmcuutoi otidi outs
The tiki,bihiiod of bictio icih cI1 c Is ma> bo sot cushy
011(11 ti_S 1 it,tit_cI i in- tistit nic ,ut of dose is b’scct on
itulcgtded I 1 ox jlont_ O Otic dtminlilo’i resulTs in a
dsspt opot I totmotci> Ii c ttic ic.a_sC in tad i, lS CO crpy in
70
(.0
I-
(0
0
Ii
45 a,
‘10
tO i S 70 30
30
-------
)iJtI\ tC ti ‘Ili, J u lio— It F,isiuu ot, t it (itilikI S ttlii’.iit
•0
I i ,
95
90
83
70
c c
Sc
so
30
20
i i ,
7
-J
0-
D
U )
2 0
‘U
I . ’
0,
-o —70
-(5
- 0 GO
— 0 - ‘ 5
N
oN
- N -
- MA C kE REL N\\ - 55
• -30
0 -25
I j _ J
I_ -i
C-, I
ti
1 .
70 ‘20 iOO ‘20 0,0 1 )0 7,0
1 1 3
it.iieicii’l ii I ill ctttit PS or CiA 1iitei f t 114 .1) 1.1
tho. c\JLrI;iictit, 177 9 (1,’, ci t ue ouciit.y L i I .10
stii t is td dos.iic if 24,000 F ii 2 ‘Flits cioo-,i c i
muc h tugitci Iii . 1 1 ttiii’c 0 odiicmIig Ci’lIlp ii bk stir—
VI ’ .il lit tic iiliu’i iITit.Ilt\ (Ii’ 6) hit u lien
i’.t igiited ii ) the DNA .iciioms -.iceti m lii i it i lt,isohl-
111113 citce to tiici-.e ittes ( [ ii: 5) 1 ii us, the Sct low
i)NA .,ct’ou ‘,tlcct mini It ‘ s .1 I chotiseij ‘ bi d nodei
fOr piecifttiilg 10 1 )12? tJV—lt tflceis iii atitiini ’ l Oll .12
nuder citfloi in tflC i liii t tt_tt’i dish [ ‘at uris Tins
iesu l l not only tui rities (lit, esiitts fmoii L\f’Oi i lilutils
cnipkui ii’ dIftit mit ctul’cFi ii cuer’by dish itli ltiolls hut
:iko in.ikc— possible C,iiciil.it üii t’i dose lutcis th it
V. Ot Itti ti( i_ill 11 ndei .iiidinS c ’iid a ions of 0/(11k’
d i llimnhi tiOm l
)I itti’loqui ru cj7c’cts
Lesions occui icd in the iritmum .ini1 the c e iii
.iiietsoi 3 and m acic , c i i _i iv.ie ccii saving e\tiOSfli c tO
UV i ii ,idm’tuoii (lug 9) 1 ticidetiec Of les;out-. o.—cs
tumuiter iii the PS esixutnicults th I ll Ii ) Flue (‘1 A cspcru—
mitemuts as could hr. ecpocicsi ti cm the d ific i LI1(C iii
cpst i ii energy con’posfiirihu of LI V IS it i ed Luum (\i
101 1 14) 1k tt IJi,c tile (t05,i C ti prtiduiee .mui cq iuivaiemit
rCSpObc’ using CT / S (liii is thin tc’heio PS uultt rs litre
uscd br t salop!e, t 1)2 iliCili t ulCid cc ’ of C) C i i SitiflS
in .iutc)iovv ‘i S 25 0 51 ± 0 15 at a do 1 .igc of 4 ,000
iiiu 2 liii.’ PS C\t)ci hut Ills IS itcre,is it i cqtm mred .t
dos,’gc of 9-1,000 J in 2 to produce in muierrienc e of
0 11 028 in [ ho C ] A c jiei hlneiuts Mom c spcuins is
ss crc cs,uinm no d Ii i to1i.o4Ic ,u1)y ill tile CiA 1’, PC runiciuts
T_ihit 3 Disc (icr urn oh he ic 5 7 ’ of uort (ucrui anotic’iy
I_itt_i: tot blc’tsct) re liC it’) rttiiitosr i ll _ice) ii i. 1111(1 2 iundcr
I -S--IC) t.rnups
Fmguu e 7 Pcrtcrl curt it ut e Pit ‘ ic riirickci I ‘ii IJIC ciii )
of the civtti ionic period ) 0 r 1 Th,’ sc_ I : ) md il_il tilinum);Ciit
dosc ci (tV—It (It’s soD ) in I tn— Lucre, Sdiiiicci 113 1
1 S- 0 I iitupc I l l’ ered h) ccliii tow hi ic_i ic ’ (C IA) p0 1 1 1k
lift moan suu ,)tal for 15 (c’I)tfl ilk
thu pau l of thc t.J\’—tS spee0 1 h_’nj Lu) sshmeh his ti
fishes , i e the tno4 scii tt is e Thus an efTrct ic uiose
must be based upon a tsoioiui tog function that toi l es
account of tho sc .is etc’iiot h uk p: idiocy of hiiutnc cii
action ttcotchnglt , S .C appi. Ci Sc’ Cr_ri C acting
weighting functions to the coiii’iuicd chit _i 1rci ii tire
CTA .nid PS e\)ichullCrits ‘( I u’:tcid.ii action st iect i nor
(Kruuftr.un and Christ a iccn, 1972) C aldut oil c cenem ,i-
Itied actuon spectrum (Cuteen and Stiltem, 1975), .ind
ScOot s S te l ion sprctrurn for DNA (Grcen wui Miller, __________
19 IS) I IL tpCI hicit c’leruil’ dust rihut ion lot etch treat-
ittent tinder CIA aird PS s as Ot ightect by tin it piyi ,i
the ineugy at eactu n,mmson’Jer hs the appropraic cool-
licient ft ouin (lie particohat ise nhtinc function 2,iti tic.
ticigiuted rircots For a p.ir( icni,ir t reahtrelut iI)tCtZi atcd
Tab le I go cs t he mintteiuitted enorgy p ci ni norneter
for tile L U 10 in the PS :‘nit C t A e\pcrinients ci Oim
which thiesc. c.uicc’ia’ ions are h, ‘ccd
The tseigtut I ho I ga’i C 1h 1 - host fit to It “i ess on of -
prohtt on limo i c-n ih cd do c’ for t ti cnmhi ncct ci ,mt.i
was tile Scticui’, I ON ‘ act man sp. ctu Cf’i (Fur h 1 01
example. the u ti 0 est ima tc s i c t ho CiA ,\nct PS e ’
prrmrnc ott, nh-_ti ceighteci hi the SoOcie. DNA ,ut tmo’i
spccti limo, c lmikrect ity nob S7 , i’ herems iti_ ’ tlmtti_ucth
by 47 , nheiu eceinhltud Ii > t he Citcicctt .uchmon s’ice—
(ruin As c_an be Scoit by CCIIfl 0i III)? F igs ( .UId 5,
the ScOnt i ttcigiit ii i : ftmIlLticlim proi clod .111 .‘ciectiimtr_
adJimstn leult for the ctul 1 crcnces cm ‘p eti.ii cOnlhiosmiiomu
between (tie itso c\pcmimu’cns 11 1 1 .0 for aimctuosy
using tb. DN \ s ciuctied do ‘igcs fuonu t ue two is-
pCrInmcI14s ‘isis 1150 Jni -2 (Y1,• C I -- 9’)fi-f tOO)
As .1 it ‘. 1 of this is ci’tii mlii.! I 0 1 ctlmli C, ‘ii c ciudnclcd
. htt Ct)ui’i mnicuut mmsi’iIo fl!t m criirit’ri [ cii) (jio)\.tyic Iii
hOG 1_i —t Cl It) tIu it pi i’di’i e.t .i intohi ii .i rniu’t I i i , 1 l L I I Ii me m
\ 1 .’q, ei_nguti
Dose (Sin 1)
\Vas ckn 14 t i i
Disc (tin 7)
(urn)
I ’S
C IA
(rum)
PS
C A
276
217
218
2 / 9
2 8 1)
28t
232
2 33
231
28 S
286
237
283
78’)
290
,9t
297
293
29 t
,iJ5
‘u9 s
1 Siint ot
&
I0
(4
20
30
4t
53
‘ it
9t
1(4
(-IS
t I lt
2 (8
-iso
‘F IJ I
lit
137
‘t t7
!d O
( r Ift
7 (0
781
22
27
26
3 5
$IJ
0 ’)
)05
1 1 ,9
912
Fit)
1610
I O i l)
299 839
300 90 1
301 939
30? JOlt)
301 1070
701 itt/f)
305 113))
106 tt6O
30/ 1(90
1Q3 1200
309 (7(0
310 1280
3it 13/0
312 1411)
3(1 1160
3(1 11ff)
3(5 (270
3(6 1200
3t7 tt7O
318 1150
119 1)10
1 - i0 (IOU
2320
2610
2 1090
3190
3-140
3 5’ !)
3750
3900
7980
41 )51)
‘1(80
4370
‘t 1) Itt
‘ :900
solo
4664)
‘1 130
4170
4010
3990
‘ s i in
3310
I 1ruts ‘i I ut’tc Itt, uv2 tium
2’ If iuitt’tlra) i_it
-------
UV tF i o
9
,—
z M’
70
IC,
I gun. 8 Icr cent ui i o of ioi i, ii o’’ hot) ‘i tIn cd nf ih Ombi) omc p i iivi (prulit cc,ds)
.,ii’l (ill nii,,,jt 110.0 d ,c of IJ\ It o H ’h 15 J I’) I) ’ .utiun s iCCiI u n or ‘‘ik’,s (I ’I’ I) iuiig
au it>ii tl (it -,f (JILCI) .JIlLi ‘st)li (I97 ) I ‘no k ibc i , ..,c ( ,In, , or mu i—duy p. ,ihii on lot d s,v
— foi tli (( ‘‘il )Iikd ocugliled d Li Ii ‘iii lit, CIA nd I ’S expi n,Oiiis
and the uuicid icc of Icsions v ,u jdc ,ttcd .ts find Oil
of doc , ’rc (I i I U) r Lii at hc IOV.L ’St do ,cc
cinp loyd in the CIA e \ i nlicnt (il,OrtO it n — , m ci-
deuce of i ’,’OuC in the hi nun ii at,, Iio y ic.i’, dullcrcnl
fuoti ; the toiL ul (/ tc’ ,t I’ 001) and hit in the
CyC % dC , Lick, .0 ( 1 the 5 ,iiid 10’,, itoh ihu!Jt) k ck
At 68,000 J in , inctd ice of Ic ,ioii ii bc’; Ii hi in
and c e in 1 u;chcvy ; erc d,ILicnt Iio’uu l ii; co ,,tiol
(P < 0(11 bc brim I’ C 0 OS lot C) c) 1 Inn., ‘u°ndi-
c,int d.ui iiacic to ho; Ii hi a iii and cyc occui ted after
otil> 4 d ,iyt, t ocui loo .sl doc.’to.c
‘rho iI icud filO of 1,_ciuto it I uiii did eve v. 1 is nuni
higher in ancluo’y that; in In,,, 1 tiJ \t dřs 1 o ’c . nt ir
tlic LDcC, for ,t,_h, spectz’c tic unt,idcnc,_ or C) C L ions
about Il ,it c hines .u,d biaiiu lesiots 1’ c limcs
ht 1tet iii .Iiii_l;O )’ ihati Hi tO cLot ci hyt_ ksions
r.iiigcd fronu to 4 8 ni ; (iiitt\iinum dinlensi)n) ii
anchovy w Lri’.it. it; iii i( kci ci I hey vet C sOme . I 1JI
sin,ui!cr, nsu.iiiy i .imiginr Ii oun 5 IL) I0 mn ; jid oC ,—
ion,ih hy to j’m l.estois aRc occurred iii (he o bc—
br) hitib i i i Lioth species but iei e not es ,ilu ,iicd flic
ii ealnicilts •iflectc’d tilt, r ,ilc of developineti; of
melaiiishic piement in lit’ cyc 1 hi pi upon ion of Iii —
ac cx iniocd Ititohogicilly i lii pom hilly pi mnentcd
om u n ,i’iiicniccl C) CS I ncr , _:isccl ni hotli Sj c vi ii
d c ,i’te I its citect is dLSU mb -cd ii iiorc dct ,’iI foi;
nu or pin iO ’l )_J I cc .10 n,i lion n i Is ti usctisscd ni tic
nC t cccl ton
Duu nip, the ) oIL i,ic p i iuii, (lie yoll supply dc—
C (L,ices .uc I ‘ii ic Hic,cu’,c ni ku ’ iii .tmid 1nih depth
(liii 1112 tic ,i’tt (I 1 i 5 , (hic C))’ I_L Colitis P i ’tC’ Hi I ) ’
ioon. puginL nt , _d, inilti .uI lie cud of the p__i od ,_
J th u ’CHt . 1ti iu 1I i CH111 t i 11 .iiiil tlic )ciIl hP 1 )’ i’
110111) i’i,hi_in ’l__d At ii, , , in_i I our C’p rim_’ik, ill
I.it\.ui iii tli toOti uds ;%c,u_ .ii hlit hit_ti ‘ t .tge ol de; , _l—
opnicut ‘I rL,it II its ii. tire ni,tHcr ni li_nIl, ,uiiti
Atom J iIiOIO/i( oh
tiefuihi, lid irio c you 5 aid less eye I5 tu1ieiltati0n than
the controls (I i cc ii tn_i 12) This UV—It ii,ilm’itic ,ui I
ictaided di_cck’pmcot to h,ii ..tc and (hi C’\lCit of
ictarda ion s’ us r T acd 10 dos,’’,. At thi io ’ Oct
dosi’t iii our c ’ p_rnri_nls (6’( 000 lti — ‘) Tanetli,
body d ‘tht, iyc. r ’ioent .l oil, und ulk diaiiti_r
t ,_re dill 5 ci , lioni thic cciuitiobc ct I’ < 0 OUt) in—
ilic, ,ti Ii’, .i so’ui ,Iucamtt i tic I of U \‘— 17 ri id it oil on
C,i 0511 h itt ,! dt sckipnie.il OSLO It the usc-h doc,i,e
l’hc in,iI ,ini -ci,n i ,_s ii th inittiop!toi i.s of uueliovy
and noel ii oh Iii Sd, s tIC di n ,ist’d iii ihtcn.u._ e\posLui
to UV I) i ,udi 1 ’t io u, s l,ei (cc (tie)’ 0010 1 ipgu et,ted ii
ihc COldiiui’s (I ig 13) TIto ,_stt’nt of rtkpcisicun .i nidi—
Cdlccl by o ’ gi id in’ 5) sIt. Ui iii_ e.is ,_d su ; Ii dosa ’e
i\ich,ii,o oi,,cc in [ Li ’ full3 disli,’iscd condition ( ;r.uI -__
3, Fi”, S) ( ‘ccii red con. moniy at dos. ;gcs at id ribot’.
90,000 I iii — ,nid Ic-cs co’nmonI3 at loss’. i ducj _t t ,s,
bit CS n 01 i li lots i_st du cge (63000 1 in ‘1 pigment
in ncltos V mc Iinophiorec ss is niorc d is n i ccd I hu in
( Ii,. coithiols (t lost, P 0001)
‘ [ lie uk r,ituii’ Oil i’utvcLs of UV i.idi,ition on pm’—
ni 5 nt inos’.mci ,t ss tint; in -lntoplu ;cc of lishis is
llmcmitLd IL) cflccis of c li Lne ) front g uiiucicl.ul lantp’
In cuil’rist to uui f’ituctntd’c, tin_si. stiucimec Ins t te ,ule that
1J\ ’ iii ,idnii m l d l 25— I tm utiduccs sonic au_ti (_‘ .tt Ion
rdtli , .r Ihi,iii dispcisiuott of ii. l.toosoitics (ruin. I ’) 73,
I Fujii c i i -il , 1973) In a-kh,m ion, I u n cm ii! (1973)
dctuonstt.iied t hut U\’ in idu,ittuii c,utn.cd ,t
stoti of (lie ri_-ponse of (tic inchinoptmoi c ho hi;mcnl
ad’ ’ nLp .uIii 1’t simti tiui , .es (nulL-pHi phnimic ,imiil inch i—
ton,ti) n bu ,_lt s. is piotturtisI_il to dos.igi_ Tlu ’ sug—
t’,’stC il th tJ\’—inluic_ , _d i’tI ,:hmt ion of ni ;’t,unosninio
inc ’luuliy t’ os uus _ct by It \‘—i din. d l’ iui , in the I
Pcu’o 111,1 toi .\sttnl it; ut ’ .tcinit _t if the cit,ni’t’, we I’
ii iii ii li_spoihe to (I \‘- 17 si_re i i’s’ciutd, or sscrc
nised In it t iii ii dcin,i 1 ’e s’.onlil require bum tli.-r t’’.—
let ill ,. ii i Ilicili .i nil ‘ii lii i LI’.). O i( 1 li_il) sis
0
0
0
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rr ritiin
PS ilLilOl
ii osu ciii
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JoIr4 U. I 1uai .lint: II. AVIAiI’. auti I I. (1xu m i v NIosu
ligure 9. 1)e (top kit) Il:L! br,o;i (top t i ht) of ItLFtIICIfl an.1:( vy I:trvj.’ atal eye (tiuttn:t 121 1) tucJ
bI tIII (I ’c tto:n tz . 1 it) if I :irafic i!i,I(,.eii! I i i sh(,Ľ I.’sitit (am pio 1u ci l iv I ‘v’ Fradi;tI mor
iIlt (‘1’,\ Pitems. lii,t!i:it izi . vcre LU) U .lnm 2 (top Ht), I 5k Jiii (tip ri1lmt). k)) I.. Jmi 2 (bottom
Ictt), J )o) U .lmit 2 (tatt na rt t1u). S t tial sectmon\ t:miiti’t! wttk I and F. loiN.
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EYE LESIOHS
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DR/ uN LESIONS
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Al Cl ITJVY
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DOSE nv-C t Urn ?
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DOS S DVII (Ci m ) DOSS U/-C (Ki rn• 2 1
liotre ID Moan acid /ne c of IN 1O I Jt lit IlK C i 0 3 id hr.’tr, of nor 1i:rn a,,elioi and Pat tIc ni,ck, rot
Iari.ic Il iaC tills ISLd i anon’ npOc’t’ hul dott of L\ II hi ith unit LV c’icr y coulcc Isac I’t—lO
laitlp fihcrcJ Li) C iA i m m t ). —0 LJV I I lid M I.ir filters, po ltIlt Ire nicli’s fo, (,t.ltltl, itt . tad
icr l teal It_i ,, .irc i-,’o st.nw’ard or ion
I
H
I rrr
70, CO 51) 103 110 ,I?0
!
it o
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MACC I ? EL
200
120 50 00 200
hISS liSStO\
Out .tppi n_tel i i , tl,ccc c’ psi n o .015 c c_i c to cuninlilo
na lni at conditions aid i’c did net C t .iliislc photo—
birt h Tiilg icf.iir of U\’—inlnc. ii it 1’t1i ’0 In f tc
aqutinni cyst ni o hi o hi’’l’ts tune tni,,cd On 5 I ,
1)1 101/ . 111(1 00110110, .1 to’ 2 5 Ii if Ic I I d V i C\tiiltttic
( II ’’ —1) Thit. fli i ii lii. c’’lOll lOt phuitin .,ict is it ion,
Iiou’.ci or, o_i so.nccshi it h.acr iti the oqti_ii in
(cii Il,,ni nU 1 or notiii,’h roinl i uion oil it , ,’. cmi ld
It.icc ,‘fl,rte,I t1. tolls lii i the JiNA sc u’hitcd
(I .tt rc-. s ,n l i , i,’,i ti I ’. .11 101 1 the one in 1 (I-
nicult Ut lii ’ i i oil 1 snl ii ni 4 i , !j it l iii ‘.nb)ht:c iiieiitccl
s et h IJ\’ (o”clios y PS C perirncnls) Its in lie :‘qn’r—
111111 e lpO l 10 1.011 5 51t4’llrcts tint i i i csnltt i,ere not
flecte il P 5 Iii.. Iot”.’n let ci of photcii cacti’ .iting eneqy
in the c t.n nail s stcffl
A so iL n,’ k it Ii C of I ic so t csi,hu ss a’. lie non ,, -
COt tt t11. 11 .d jid cit Ill) c lufined d.liil1g0 .11(0 . 11 1
cpo nic ol cn N I Cisc Tl’e it m i ni t ,,,.ii’dtd Ictions
it, die hi u n ,tii ,I ciii,,, iii bc ,t I ct,.iilrut,oii oF guci’..tl,
111(1 itti L cf’’’ in1 .inI m i mi_i ill_Il cli ’ j) 1,1011 c_il lrlcIIII(—
st ’nio Ste’nit,c.n,t d ‘ni,’t n’—ctnicil lot r.nk I I I those
ii’. 0 u 1, 11 ’ 1 inn, 1.1111) iler tint ii tin sor—
tois if ill d.i IL Ii .,, ititit nliliFcl) liii ,iilV
tii (11111 it’Ctl ‘-ill’. (iiit, I ’ ’I (II , 55 (f thi . ..,ic, ( 1(1 ,11( 1
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1’i ure II ‘ II.lnd, ,Id LIlZ ,Il, body dlrili (\C pi ’ .L (nI i ( (I I I ur ’ 1 k .n .1 oR ( 1 ,U1LICI oil, , , .11 (I II IIIL1 .
nL1dm ) su, ‘, ‘r ,c , ’, dc’ . s řf IJ\’ ( (I S It) I, ’II1 I., C I A ti’L ’rs) I\I I,l! f , ’’ i I COIllI ( II ’, sho’’, n
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I i ’iirc 13 I cL ’tton i ’ ’v c cii .i\t mc i lmtO3(’I ’IL ill per—
t,icit tr di. rl l uni i ’ iii.d (io .ie ’. in .iIIC IIc\) ,nit in ‘i —
cici ii u Sourec ol ii .td olin i 0. IS—’0 i ,’itt ‘ iii
C_I /I fiilcr i i\ U tn’,t tc ’Ic line. r ‘s -—1) co.c jloiris
aic means br Ii .,ielc rt mutt I i , .u,id.’id crior’.
(‘udilu. I i C1 ti Li) ,IL ’iL ’.. i’d CO ’’li ’tO Ii and ri,,d ,_
3 conpit . ly d ISiL ’i u.d ( .e . ri 2 5)
be ,mbli. to fced sucCr ; iiml i i m I ii t ion of de riop-
ment is anotimur potciittai soorce of niot t.’iiiy uutt cr
natuuii coui htion imccllco -it. it piolnact. the ct,i°c @1
hi !mesi s’ule i.hiimty I’4OtLii Ii 11101 tihity in the sca
dururir the ciiihrynnic pci mod (e ’ ; throi!:rI i yř!h-sic
ct .t i u eS) is gc ’n I .iI s i—JO limes hi ’ ler ti in ,titei lit.
larvae be i ie dio ’ (Joiie ,ind 1 L I II, 197”i)
\\F csl inc lcd [ rota hi r ii red i inc mci ’ iircmc nrc
madi.. mu Sepic iiitier- Not coil that tie UN \
sci Iited domi. br 4 c i ys in in i c’ iii F a Jolla is ould
tu 43 1 iui (I d.t ) - ,, . assuminp the d r !3
dose to i x abort ci \ I iuC 0€ cnc ,. ,it ü1ii i_on
/scsuuiring a 6 ’, i c liirou )i the Sc I siiif ,ic,_ cists
a s,lluu. of 1(3 L in 7 (i d.m 3 ) icr the DN\
ssc iphtcd do. just i ’ciaeathi tire stir fice I Inc cstir lr ,rIe
di lL_i id tv old) 2 ii (mm tire s .itue 122 [ mi (1
d .iy) , cil iilcricd by Sni’ti md iLtkci (in liii
issu ,_) usiri’ (Jicu.ii ci it’s (19 / 1) Sm.i IlI— 11)111 it ii ,i t i I —
lut i i ciltiJ ,i Lii i_(it ’Ili€ic’utt ’c Ii’ to iiit’Iiii ’c’ttt’i IIII_ c iii r
to u!iiOP lit’ r .ii Limit)’ cli ’sc ja bt_imcrliu thu. ste
Sot F i e ii iii u_i it it . .tI_a J it! it lot t !mc’ ,ii tutu itl
OIm)itC di.pili of 1) 12 ciii I loin to iii tie omit t spilt
niL uil ,ii t niI . ’ . tu i’Ificts (‘if (‘,ititu ’ ilepictiotu Ciii i_n
.tii ilyti I,’rt nil it_)t iicmsvnss ii i t ‘ . ‘ ic’L&l it ni tilt_i’, ,_
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in 1 n ii i,rui p Si_ic l\co \‘OFI
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ItFI jill \C 1”,
-------
SI/ ’4 ‘
PENETR&TION OF U\T-B INTO NATURAL WATERS
R. C. Smith and K. S. Baker
University of California, San Diego
Scripps Institution of Oceanography
Visibility Laboratory
San Diego, California 92152
Report for period 1 Hay to 30 September of work under Research Grant Award
U.S. Dept. of Coimiterce - NOAA No. 04-8-M01-125 for the U.S. Environmental
Protection Agency - Biological and Climatic Effects Research (BACER) program.
Principal Investigator:
Raymond C. Smith
Visibility Laboratory
Scripps Institution of Oceanography
University of California, San Diego
San Diego, California 92152
Approved:
_____ 2
\ I _____
/11 L. Harris, Sr., ircctor
‘ .Visibi] ity Laboratory
30 September ]978
-------
PENETRATION OF UV-B INTO NATURAL WATERS
ABSTRACT
An increase in the incidence of solar ultraviolet radiation upon
oceans and lakes, as a consequence of anthropogenic diminishing of the ozone
layer in the stratosphere, might well have a signif] cant effect upon primary
producers and other organisms in these natural waters. The primary ohj ective
of the research during this funding period (an abbreviated continuation of
research done under NOAA Grant No. 04-7-158-44039) was to obtain represen-
tative oceanographic data, during a cruise on the R/V KNORR, in order to
make preliminary estimates of the potential effects of increased ultraviolet
radiation incident upon aquatic environments.
Accurate data on the penetration of ultraviolet radiation into a
variety of ocean waters types have been obtained on a cruise between
Kwajalein and Samoa in the central equatorial Pacific. As correlary infor-
mation, important chemical and biological characteristics of these waters
were also measured. Further, primary productivity of natural phytoplankton
populations, and the influence of UV-B radiation on this productivity, was
measured for a variety of ocean waters ranging in chlorophyll a concentrations
from 0.02 to 0.3 mg•m 3 chi a. In addition, spectral, total and selected
narrow band UV irradiance measurements were made of above water incident
irradiance as the RV/ ORR moved from 9°N to 14°S.
The research reported herein was limited to field work at sea.
Current funding did not include funds for the analysis, interpretation and
publication of these data. Thus, this report briefly outlines experimental
methods and lists the data obtained, but includes no discussion of the data
or results.
Progress during the past yenr, prior to the field work reported herein,
has been completely described in a report for fiscal year 1977 of work under
-------
-2-
Research Grant Award, U.S. Department of Commerce - NOAA No. O4-7-158- 14039
for the U.S. Environmental Protection Agency - Biologica] and Climate Effects
Research (BACER) program, submitted 15 January 1978. In addition, a manu-
script “Penetration of LW-B into Natural Waters” by R.C. Smith and K.S. Baker
has been accepted for publication in the Journal of Photochemistry and
Photobiology.
RESEARCh AND METHODS
Primary objectives of this research are to determine the penetration
of UV-B into various natural waters, to correlate this penetration with the
dissolved organic material (D ) and pigment concentration in these waters,
and to assess the potential effects of increased ultraviolet radiation on
primary productivity of natural phytoplankton and other aquatic organisms.
To obtain the necessary optical data, a submersible spectroradionieter
(modeled after the Scripps Spectroradiometer, Tyler and Smith 1970) capable
of measuring underwater spectral irradiance, including the IJV-B portion of
the spectrum, has been constructed. This instri nnent was briefly described
in the Report for FY 1977. A more complete description of the instrument
is anticipated.
The submersible spectroradiometer was used to measure above water
spectral irradiance and below water spectral irradiance as a function of
depth. The submerged instrument was used to obtain data in two different
ways. First, at a fixed depth, the spectral irradiance was measured over
a range of wavelengths. This gives a measure of the downward spectral
irradiance at a fixed depth and data obtained in this manner is designated
Ed(z) in Table 1 (0+ indicates above surface spectral irradiance measurement).
Second, for a fixed wavelength, data was obtained as a function of depth.
This gives an accurate determination of the diffuse attenuation coefficient
for irradiance at the fixed wavelength. Data obtained in this manner is
designated K(A) in Table 1. These spectral irradiance measurements were
carried out for the full range of sun zenith angles and for water types
ranging from 0.02 mgn1 3 chi a to 0.3 mg m 3 chi a.
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-3-
Primary productivi ty, for this same range of water types, was measured
using the “simulated-in-situ” techniques where “IC was used to incubate
samples from selected depths on the deck of the ship (Strickland and Parsons,
1968, Part V). Primary productivity was measured using both ambient solar
radiation and solar radiation appropriately enhanced with UV-B radiation
from Westinghouse FS4O sun lamps. Stations at winch productivity was deter-
mined are indicated in Table 1, where the depths [ meters] from whidi samples
are obtained are listed.
Surface chlorophyll and phaeopigrient concentrations were measured
along the ship track and as a function of depth at selected stations using
a Turner designs “flow thru” fluorometer, for continuous measurements, and
a Turner 111 fluorometer, for discrete chlorophyll measurements. Our
chlorophy] 1 measurement technique followed the methods described by Strick-
land and Parsons (1968, Part IV.3). “Average” surface values for chi a at
each station are i.ndicated in Table 1.
Above water irradiance was measured using a nimiber of instruments, in
addition to spectral irradiance (250-800 nm) using the Scripps 1W-B submersible
spectroradiometer: (1) total irradiance using an Eppley pyrheliometer, (2)
total irradiance using a Sol-a-meter, (3) irradiance at 340 nm using an Eppley
ultra-violet radiometer, (4) irradiance at 365 nm using an International Light
calibrated photodetector. These instruments were monitored throughout the
day using a Hewlett Packard digital voltmeter/clock-timer/relay activator
system connected directly to an HP9825A calculator. Days for which these
data were obtained are indicated in Table 1, under “Surface Irradiance”,
according to which instruments were monitored.
The Turner fluorometer, with the 1-IP VTVIVI/Relay/Clock system, was used
to obtain several tracks for the purpose of obtaining the variance spectra of
chlorophyll. These data will be used for the study of phytoplankton patchiness
in the equatorial Pacific waters. These data are indicated in Table 1
(column 10) by their data file number.
In addition to our own research efforts, the above data were collected
as part of collaborative work with other research groups. In cooperation with
-------
-4-
Dr. 0. Zafiriou (Woods Hole Oceanographic Institution), photochemical re-
actions in seawater (other than photosynthesis) with respect to attenuation
of U\T radiation were investigated. This work was also carried out on the
RV/1 ORR (W.H.O.I.) during a major transect across the equator, proceeding
from about 9°N to 14°S in the central equatorial Pacific. Principal par-
ticipating groups and interets were: Dr. 0. Zafiriou W.H.O.I.), seawater
photocheniistry and marine d e]nistry; Dr. R.C. Smith (S.r.O.), above surface
and underwater spectral irradiance chlorophyll and DQM concentrations and
beam transmittance; Dr. Mack McFarland from NOAA/Boulder Aeronomy Laboratory,
measuring atmospheric NO, NO 2 , ozone and selected freons. The cruise track
for this collaboration work was ideal for gathering a unique, highly inter -
esting set of surface and underwater ultraviolet radiation data (along with
corresponding atmospheric data) from this remote maritime region. The equa-
torial region itself, according to Green’s tables, simulates an increase of
25 6 above the ozone levels found at higher latitudes. The cruise track
crossed a variety of environments in the northern and southern hemispheres,
including the edges of the tradewind belts, the doldrums, the inter-tropical
covergence zone, and the oceanic equatorial current/countercurrent system
with associated high biological productivity, flanked by oligotrophic water
to the north and south.
Our field data outlined in Table 1, will be used to determine the
diffuse attenuation c efficient for irradiance in the UV-B region of the
spectrum for the range of water types encountered on tins cruise. These K
data will then be correlated with pigment concentrations in order to provide
a technique to characterizing ocean optical properties in the UV region of
the spectrum by the pigment concentration. This would allow estunates of
UV penetration into natural waters to be made from a loiowledge of chlorophyll
concentrations. In addition, an effort will be made to quant]tatively relate
the penetration of IJV-B into the various water types on the primary produc-
tivity in these waters. This information is necessary in order to estimate
the potential effects of UV-B on natural phytoplankton populations.
-------
5-
RESULTS
Cruise results are summarized in Table 1. A station labeled with an
asterisk ( ) indicates data taken while the ship was underway. For example,
at station K (SlO Station #11) on 4 August the ship position was 00 1.1’S,
1710 4. Y ’W. At this station: the surface chlorophyll a concentration was
0.20 mg m 3 ; productivity studies were made using water incubated from
0,8,16 and 32 meters; irradiance above the surface was recorded using all
four sensors listed above; downwelling spectral irradiance underwater was
measured at 5,10,15 and 20 meters; the diffuse attenuation coefficient for
irradiance was nieasured at 305,335,350,365,380 and 410 nni; and a run to
measure the chlorophyll variance spectra was made.
REFERENCES
Strickland, J.D.H. and T.R. Parsons. 1968. “A Practical Handbook of Seawater
Analysis.” Fisheries Research Board of Canada. Bulletin 167. Ottawa,
Canada.
-------
L 12 7 2° 25.4’S 168° 3 OW
*L 12 3 3° 27.5 S 166° 55 0 W
0 18 1.2,3,4
0 25
0,10,20,40 1,2,3,4 Ed(0°)
‘4 13 9 3° 39.1’S 166° 40 lW
0 iS
0,9,13,36 1,2,3,4 E (0,3,S,10)
305,320,350,365,410
‘4 14 (0 4° 33 2 ’S 164° 50 O’W
0.16
1,2,3,4 E (0,20,15,10,S)
305,320, 330,365, 560,410,450
Q 17 13 ‘° SI 1’S 165° 23 OW
13 14 9° (0 l’s loo’ 3 OW
U 15
0 [ 2
0,9,13,36 1,2,5,4
0,10,20,40 1.2,4 E (S. ’).1S ,2o)
UNDF2 kTb ( 1RR40 [ A CE O!LO F1rnL
FRRiDL3 , . CE K [ ,r(’) at X(rail VAI4 [ A’rE SPtt1R
TABLE
1
SThTION
i 1L)1 Sb
1978
(lATE
LATIS1JUE
O’4CITUDE
CII I a
( . 1
P XXJCTIVITY
9JRF CE
IRJ4ADIA3CC
°B
D
E
F
F
6
2
4
5
6
6
7
JULY
20
22
23
24
23
26
7° 42.0 N
7° 33 8’N
7° 26.5 N
7° 23 O’N
1° 27 O’N
7° 19 1 ‘i
172° 0 O’E
176° 01 8’E
178° 9 2’S
179° 33 6 l
178° 42 9’W
177° 12.7 W
0 015
0.015
0.013
0.05
0 06
0 09
0
0,15,30,45
3
3
Eo( 0 , 1 5)
Ed(0°) vs sun
Ed(0°) - r or
0 d ’’ 5 ’ 6 5,11.5,16.5)
305,320
305,320
U
8
27
5° 7 5 N
175° 34 OW
0 10
2,3
EJ(l.S. 3 .S) vs sun
¶1
9
23
5° 37 SN
175° 34 0”n
0 Is
2
1
9
29
3° 39 4’N
174° 57 2’W
0 12
0,15,35,55
2,3
EJ(0,1.S,6 5)
J
10
30
1° 48 5 N
173° 34 0 W
0.15
1,2,3,4
EJ(l 5,3,5,10,15)
J
10
31
1° 45.2 N
173° 3.4 5 N
0 25
0,8,16,32
1,2,3,4
E 1 (0,5,10,15,20)
J
10
A1JJJST
1
1° 10 Z’N
172° 44.] .’W
0 25
0,8.16,32
1,2,3,4
E 4 (0°,5,l0,15)
*K
11
2
0° S.O’N
171° 15 O’W
0 30
0
1,2,3,4
E (0 ) overcast
I ’.
11
3
0° 1 3’S
171° 5 4’W
1,Z,3,4
Ea(0°)
K
[ 1
4
0° 1 1’S
171° 4 9’U
0 20
0,8,16,32
1,2,3,4
E (S.lO lS . 2 O)
K
11
5
0° 47 6’S
170° 9’h
0 25
0 ,8,lb,32
1,2,3,4
EdiO ,l .5,5,19)
•)
11
0
1° 41 2’S
101° 1 S’W
1,2,3,1
E (0)
Fl 07 26
F2 07 26
F10728
F10731
Fl 0804
Ft 0505
305,320,335,350,365,380
30S
305
305
305,320,335,350,365,380,410
305,320,325,350,365,380,410
P
16
11
5° 55.0’S
164° 0 O’W 0.17
1,2,3,4 E (O°,5)
°P
[ 0
U
5° 55 0’S
164° 0 O’W
1,2,3 Ea(0’) VS lUll
305 .320,335,550,365,380, 110,450
..‘ Ii IS (U’ II .1’’, bl ——
I’2
‘ ‘ ‘5 3’ (l 3 3, 33(I,3 15,330 U I), ISO I-i .’ i 2
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/To ,. ‘/
ANNUAL REPORT Jctober 15, 1977
Contract NAS—9—14871
NASA
Higher Plant Responses to Elevated Ultraviolet Irradiance
Dept. Range Science & Ecology Center
Uta.a State University
Martyn H. Caldwell, Project Leader
Susan J. Lindoo, Postdoctoral Fellow
Ron Rohberecht, Graduate Research Assistant
Fred N. Fox, Graduate Research Assistant
Judy Dickson, Graduate Research Assistant
Steve Holman, Graduate Research Assistant
Harvey L. Neuber, Technician
Lee B. Canp, echnician
Much of the material contained in this report is preliminary in
nature and is subject to revision and reintaLprctat on. The
authors request that it not be cited without expressed permission.
-------
This report will relate the progress made in our laboratory during
the 1976—77 fiscal year toward determining the effects on plant growth of
enhanced ultraviolet irradiation. The report will describe work done
in the following areas:
1. Variations in leaf epidermal transmittance of UV—B radiation of
Utah and Alaskan plant species.
2. Measurements of a natural solar UV—B gradient at sites located
along a North and South An erican transect (includes measurements
of optical properties of plants located along this gradient).
3. AnaLysis of UV—B radiation induced inhibition of young leaf
expansion.
A. Possible involvement of the low irradiance phytochrome
system in UV—B induced inhibition of leaf expansion and
promotion of anthocyanin production.
B. Determination of levels of the growth inhibitor, abscisic
acid, in leaves of plants exposed to enhanced and control
UV—B radiation.
C. Examination of UV—B radiation effects on cell division and
cell expansion.
4. Competitive interaction in plant populations exposed to enhanced
UV—B radiation.
5. Interactions between enhanced UV—B radiation stress and water
stress on plant growth.
Throughout the project enhanced ISV—B irradiation has been simulated
through the use of Westinghouse ‘ FS—40 sunlamps covered with one or two
layers of Kodacel TA—40l (5 mil) plastic film. (Work now under way will
also use sunlamps covered with cellulose acetate plastic film or
Westinghouse BZS lamps. These two new systems permit a wider range of
UV—B irradiation regimes.) Control conditions were maintained by using
FS—40 lamps covered with one layer of Mylar Type A (5 or 10 mil) plastic
film. These lamps were used in growth chambers (main light source in
chambers was a 6000—Watt Osram Co. xenon arc covered with plastic or glass
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—2—
filters), in greenhouses, and in field situations. The xenon arc light
source used in the growth chambers contributes a low but significant
level of UV—B radiation. The spectral regimes used in the various
research efforts will be described in each part of the report.
In order to provide a description of integrated IJV—B irradiation
being used, the UV—B radiation regimes are also expressed in terms of
biologically effective radiation (UV_BBE radiation). The weighting for
biological effectiveness is taken from a relationship recognized by
Caldwell (1971) and mathematically described by Green et al. (1974),
and roughly follows the action spectra of effects mediated by nucleic
acids and proteins. This is normalized at 280 rim.
Much of the research conducted in our lab is done with one plant
species, Rumex 2 ientia. This permits research findings on one aspect
of the plant’s response to enhanced UV—B irradiation to be integrated
with findings from other areas to allow a more complete model of
adaptation and response. However, other species have been examined in
field s.udies on epiderrnal transmittance of UV—B radiation, and in studies
on the effects of enhanced UV—B on competitive interactions.
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—3—
1. Variations in leaf epidermal transmittance of UV—B radiation.
(Robberecht)
The paper by Robberecht and Caidwell which is appended to this report
summarizes the work done in this area. This paper will be submitted to
the journal Oecologia .
2. Measurements of a natural UV—B gradient at sites located along a
North and South American transect. (Caidwell and Robberecht)
(Funding for this project is also being supplied by a graht from
the National Science Foundation).
Studies are now underway involving a natural gradient of solar UV—B
radiation extending from low latitude sites at high elevations above sea
level, to the northern arctic. This natural gradient of Ladiation was chosen
as a background in which to study plant adaptation, specifically the tundra
plant life form, to different UV—B radiation regimes. The present phase
of this work involves three facets.
First, it has been necessary to measure the 1W—B spectral irradiance
along this gradient in order to determine the applicability of atmospheric
transmittance models such as that of Green et al. (1974). These spectral
irradiance measurements were made during 1977 under various conditions of
ozone concentration, atmospheric turbidity, solar angle, and elevation
above sea level at several sites. The low latitude sites include several
locations in the Peruvian Andes, in the Venezuelan Andes, and on the
island of Maui in the Hawaiian Islands. The high latitude sites include
Barrow, Meade River, and Fairbanks, Alaska. Measurements were also taken
at mid—latitude sites at both high and low elevations in Utah. A final
calculation of these data is dependent on a “slit function” for the
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—4—
instrumentation, a Fourier transform, which is now being calculated by
the National Bureau of Standards. Preliminary data are, however, shown
in Figure 1. Curve A of this figure is spectral irradiance at solar
noon at Meade River, Alaska (70°N, 15 in elevation) close to the summer
solstice. Curve B represents the spectral irradiance at solar nuon on
Macabaji Pass in the Venezuelan Andes (8°N) in April when the sun is
only 2° from the zenith. This site is at 3560 in elevation. The
instantaneous biologically effective UV—B global solar irradiance (sun
plus sky) differs by a factor of 6 between these two extremes. The
peak daily dose of biologically effective UV—B radiation differs by a
magnitude somewhat greater than 6. Comparing the Andean sites with a
mid—latitude site, there is approximately a three—fold d...fference in both
instantaneous noon irradiarice at peak solar angles and also in the total
daily dose during the period of the year of maximum UV—B radiation. This
natural geographical gradient is steeper than predicted by the model of
Green et al. (1974).
A econd phase of the gradient research program involves investigation
of plant optical properties in the field at selected locations. Although
plants studied at all locations along this latitudinal gradient generally
t
had rather low UV—B epidermal transmittance, species at mid— and high—
latitude sites exhibited greater variabilit , with some species having
rather high apidermal UV transmittance, whereas the species at lower
latitude, high elevation sites consistently exhibited very low epidermal
UV—B transmittance. This was found for agricultural as well as wildland
species. The reflectance of UV—B radiation from leaf surfaces was generally
quite low for all plants investigated. Attenuation of the radiation thus
appears to be primarily caused by absorption in the epidermis itself.
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—5 --
A third phase of this gradient research has only recently been
initiated. This involves the comparative sensitivity of closely related
plants, from selected locations along this latitudinal gradient, when
grown in controlled environment facilities.
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3. Analysis of tJV—B radiation induced inhibition of young leaf expansion.
Previous work by Sisson and Caidwell (1976) has demonstrated a
decreased growth rate of newly emerging Rumex patientia leaves under
enhanced UV—B irradiation as compared to rates for leaves on control plants
under low UV—B irradiation (Figure 2). This decreased growth rate is most
pronounced for the first 3—4 days (the period of most rapid leaf expansion)
after which it more closely resembles the rate for control leaves. As a
result the final size of UV—B treated leaves is less than control leaves.
Inhibition of young leaf expansion, even though it may last only during
the first few days of growth, is still of critical importance since the
end result is a leaf smaller than its potential size. This smaller
leaf size translates into a smaller plant capable of less photosynthesis.
The better understanding of how enhanced UV—B radiation results in
decreased young leaf growth may lead to methods of counteracting or
minimizing these effects.
Various aspects of this decrease in young leaf expansion are being
studied. One portion of the work is to determine if this decrease is a
result primarily of inhibition of cell division, or of cell expansion.
This work will be discussed in a following subsection. Another aspect
of the problem is to determine how enhanced UV—B radiation interacts
with factors which control young leaf growth with the view of eventually
discovering the mechanism of UV—B inhibition. The possibilities of UV—B
radiation acting through two or these factors, the low irradiance phyto—
chrome system and altered hormone levels (specifically the growth inhibitor,
abscisic acid) are discussed in the following two subsections.
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3. A. Possible involvement of the low irradiance phytochrome system in
UV—B induced inhibiUon of leaf expansion and promotion of
anthocyanin production.(Lindoo)
A summary of the work done in this area is found in the paper by Lindoo
and Caidwell appended to this report. The paper has been submitted to
Plant fl siology.
3. B. Determination of levels of the growth inhibitor, abscisic acid,
in the leaves of plants exposed to enhanced and control IJV—B
radiation. (Lindoo)
Plant growth is controlled and coordinated by endogenous plant
hormones. The decreased growth rate observed in young leaves exposed to
enhanced UV—B may result from increased levels of growth inhibitors,
specifically abscisic acid (ABA). The results of an expe’iment described
in last year’s annual report indicate that exogenously applied ABA can
inhibit young leaf growth in Rumex patientia . In that experiment, an
aqueous solution of M ABA was applied to leaves of Rumex plants
beginning when the leaf tip first emerged from the sheath, and the
applications were repeated daily. Leaf length was measured as the distance
from the leaf tip to the base of the plant. The relative growth rate of
leaves treated with ABA was less than the rate of control leaves during
the first few days of most rapid growth.
Further, increases in ABA levels have been observed in plants placed
under different types of stress. Wright and Hiron (1972) reported that
subjecting plants to such water balance—related stresses as water—logging
the roots, dehydrating the plants under a stream of warm air, or with—
holdin water, resulted in high ABA levels. Boussiba et al. (1975)
found an increase in ABA concentration in plants subjected to mineral
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—8—
deprivation, salination or B0 3 toxicity. Enhanced UV—B radiation has
been demonstrated to induce typical responses to stress in Rumex patientla
(decreased rate of photosynthesis, increased dark respiration, increased
leaf residual resistance to CO 2 movement, cellular damage). Therefore,
we have hypothesized that UV—B stress may result in increased levels
of A.BA and have been examining free ABA levels in leaves of Rumex plants
subjected to UV—B stress or control conditions for varying periods of
times. We are also interested in ABA levels in UV—B stressed leaves of
different ages since leaves at various physiological stages of development
might have different degrees of response to the stress.
Two experiments have been completed which measured the levels of
ABA in the three youngest leaves of plants held under control and enhanced
UV—B conditions for 1, 3 and 5 days. (A preliminary experiment has also
been done which measured ABA levels in the two youngest leaves after
2, 3, 5 or 8 days of control or UV—B conditions.)
Rumex plants used in the two experiments were 4—5 weeks old with the
6th true leaf just emerging from the sheath. Plant pairs were selected
on the basis of similarity in lengths of the 6th, 5th and 4th leaves.
One member of each pair was randomly assigned to the control group and the
other member to the UV—B stressed group. The treatments were done in
growth chambers equipped with xenon burners and FS—40 sunlamps covered
with Mylar (control) or Kodacel plastic film (UV—B stressed). In both
experiments the control plants received about 100 J•m 2 UV_BBE per 7 hr
day. In the first experiment) the IJV—B stressed plants received about
1550 J m 2 UV_BBE per day, and in the second experiment about 2250 Jm 2
UV_BBE per day. The lengths of the 6th, 5th and 4th leaves were measured
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daily. (A graph of the increase in leaf lengths of Rumex plants held
under control and enhanced UV—B conditions is shown in Figure 4.)
There were 12 plants in each treatment and 1; plants per treatment
were harvested at the end of the 1st, 3rd and 5th days of exposure. The
harvested leaves (blade and petiole) were quickly weighed, frozen in dry
ice, and kept on dry ice until extracted. All samples, except the 6th
leaves, were extracted separately. The 6th leaves in each treatment
iere pooled into 2 groups of 2 leaves each (this was done for each harvest
date in experiment 1 and for the 1st harvest date in experiment 2). The
tissue was extracted in 90% methanol and put through a partition
purification procedure. The samples were methylated with diazomethane
gas (to produce a methyl ebter of ABA) and further purified on thin layer
chromatography (TLC). ABA levels in the samples were measured on a gas—
liquid chromatograph equipped with an electron capture detector. (Samples
in the 8—day experiment mentioned above were not purified by TLC before
being gas chromatographed. Extraneous peaks from contaminants in the
samples may have inflated the ABA peaks and made accurate measurem [ at
of the peaks difficult. Therefore, the results of this experiment are
considered preliminary.) The measured ABA levels were expressed as ng
ABA/g fresh weight of leaf tissue.
ABA levels in control and UV-B stressed leaves from the two five—
day experiments are compared in Table 1. There was, in general, a
decrease in ABA level with leaf age. This phenomenon has been reported
in the literature (Weinbaum and Powell 1975). (Although this may seem
paradoxical since ABA is a growth inhibitor, it may be related to the fact
that the youngest growing tissues, as a rule, also have much higher levels
of growth promoters than somewhat more mature tissues.)
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—10—
Table 1. ABA concentrations (ng ABA/g fresh weight) in Rumex patientia
leaves exposed to control (100 J .m 2 UV—BBE per day) or enhanced
IJV—B conditions (eicpt. 1 1550 J•m 2 UVBE per day, expt. 2
2250 J•m 2 UV—BBE per day). The 6th, 5th and 4th true leaves
were extracted from plants harvested after 1, 3, or 5 days
of exposure.
Leaf
Day
harvested
6th (youngest)
5th
4th
Control
UV—B
Control
—
UV—B
Control
UV—J
1st
expt.
expt.
1
2
20
105
12
36
8
41
7
20
3
19
6
8
3rd
expt.
expt.
1
2
11
19
12
13
7
10
8
9
5
7
5
7
5th
expt.
expt.
1
2
12
23
17
28
8
11
8
12
7
9
6
12
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—11—
Since the plants, and therefore the leaves, In the UV—B and control
treatments were paired, analyses of the results were done with a paired t
statistic. In no case was the concentration of ABA in the UV—B stressed
leaves significantly greater than in the control leaves. In experiment.
2, at the end of the 1st day, all 3 sets of control leaves had considerably
greater ABA levels than the comparable UV—B stressed leaves. Since these
levels were much higher than on other harvest dates, and since this response
was not found in experiment 1, it may be that the control plants in the
second experiment had been water stressed during the first day (ABA levels
are very sensitive to water stress).
The similarity in ABA concentrations in control and UV—B stressed
leaves suggests that enhanc d UV—B radiation stress is not mediated
through higher levels of abscisic acid, at least in young leaves. Therefore,
it appears that the decreased rate of young leaf growth in plants exposed
to enhanced UV—B is not due to inhibition from high levels of ABA.
However, the work just discussed was done with young leaves given enhanced
UV—B radiation for a maximum of 5 days. There is another aspect of plant
growth in which increased levels of ABA may mediate the observed effects
of UV—B stress. Sisson and Caldwell (1977) reported that the photosyntheticalLy
productive life of Rumex leaves ‘held under enhanced UV—B radiation is
shortened compared to control leaves. Further, as discussed in last year’s
annual report, total protein and RNA appear to decline more rapidly in
older leaves held under enhanced UV—B than in equal—aged control leaves.
This suggests that UV—B stress given to older leaves, particularly if the
exposure is prolonged, way result in an early senescence. Therefore, it
is possible that UV—B stress, given to older leaves or given for prolonged
time periods, may resu].t in cumulative damage leading to increased ABA
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—l2
levels which may precipitate an earlier leaf senescence. In the
preliminary experiment mentioned above, leaves held under enhanced UV—B
for 8 days had a considerably higher ABA level than control leaves.
However, due to the purification problems, these results are considered
preliminary. An experiment is now under way which will examine ABA
levels in the 3 youngest leaves held under enhanced UV—13 and control
conditions for 8 days.
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3. C. Examination of UV--B radiation effects on cell division and cell
expansion. (Dickson)
This project Is now completed and a manuscript Is being prepared
for publication which will report on the work discussed below.
Leaf ontogeny and ultimate leaf size are determined by three factors:
(1) the rate and duration of cell expansion, (2) the rate and duration of
cell division, and (3) the number of cells in the leaf primordiutn. The
reduction of leaf growth by UV—B radiation may result from a direct effect
of UV—B radiation on any of these processes. It is possible by
experimental design to eliminate differences in the number of cells in
the leaf prirnordiuin. Therefore, this project was designed to examine
the pattern of cell division and cell expansion in leaves exposed to
control or enhanced UV—B radiation regimes. The following hypotheses
were tested:
(1) Cell division proceeds normally in both treatments, but the
rate or duration of cell expansion is reduced by UV—B radiation.
and (2) Cell expansion proceeds normally in both treatments, but the
rate or duration of cell division is depressed by UV—B radiation.
METHODS . Seedlings of Rumex patientia L. were planted into 10 x 10 cm
pots and placed in a growth chamber where pretreatment conditions were
Identical to those under which the experiment would be conducted (excluding
UV—B supplement). Based on the uniformity of growth prior to the initiation
of the fifth leaf, 24 plants were chosen and paired for the experiment.
One of each pair was randomly selected for the control or UV—B treatments.
Pairs were maintained identically regarding pot position and azimuth
angle and leaf angle of the fifth leaf in the chamber In order to reduce
variation in growth due to small differences in microclimate.
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Growth Chambers . The controlled environment chambers were equipped
with a 6000—W Osram Co. xenon arc burner which was maintained at an
average output of 750 pe1nsteinsm 2 •s (400—700 nm) as measured by
a Lambda Co. Model LI—l90—SR quantum sensor. The xenon arc was enclosed
in 2—mm Schott Co. WG—320 glass filters which effectively absorbed
radiation of wave]engths less than 300 ma. In addition, three Westinghouse
FS—40 “slaps” were set parallel in a frame oriented lengthwise in the
chamber. The frame was suspended 27 cm above the pots which were set in.
two rows corresponding to the spaces between the three lamps. Supple-
mental UV—B radiation was supplied by the sunlamps filtered with Kodacel
TA—40l (10 mu) plastic film which was replaced daily to maintain the
desired radiation environm ’nt. In the control chamber the sunlamps were
filtered by Mylar Type ‘A’ (10 mu) plastic film. A general description
of this lamp—filter system was reported by Sisson and Caidwell (1975).
The spectral irradiance from these lamp systems is shown in Figure 3.
Due to the greater visible and UV—A (320—400 rim) irradiance in the
natural environment, a comparison of spectra from growth chambers dnd
those predicted for global irradiance at various ozone reductions is
difficult. The growth chamber and predicted regimes can, however, be
related in terms of the biologically effective UV—B irradiance UV_BBE.
The quantity thus obtained for UV_BBE during mid-May (40°N latitude
and an ozone concentration of 0.34 atm-cm) is approximately 1000 Jm 2 .day 1 ,
using the model of Green et al. (1974). In the growth chambers, the
UV_BBE dose was calculated to be 2200 and 100 Jm 2 .day for enhanced UV—B
and control treatments, respectively. This UV—B treatment corresponds
approximately to a 40% decrease in ozone concentration (0.20 atm—cm)
(Green et al. 1974).
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—15—
The simulated day In the growth chambers was 11 hours with the
FS—40 lamps on during the middle 7 hours. The temperature and humidity
cycles represented a mid—M ’y condition at which time Rumex Patientia
begins growth in the field. The daily maximum was 29 C, 20% RH and
minimum was 13 C, 40% RH.
Leaf Measurements . The length of the fifth leaf was recorded daily
before the sunlamps came on. At this time and at mid—day, pots were
rotated 90° and advanced one position in the clockwise direction to minimize
effecEs of small irregularities in the radiation field. On the second
day of the experiment, the fifth leaf was sufficiently long to be tied
back with white string in order to expose the abaxial surface to the over-
head source of UV radiation. This angle of inclination was similar for
each pair of plants. As the leaf grew, it was gently trained to a
horizontal position.
On the 3rd, 4th, 6th and 16th days, three pairs of plants were
chosen at random and the fifth leaf was sampled. Recordings were made
of total leaf weight, and weight of the blade and petiole separateiy.
The leaf was then cut in half 1eng hwise. One half was sectioned and
fixed in formal acetic acid for tissue preservation. This tissue was
later dehydrated in a tertiary—butyl alcohol series, embedded in paraffin,
sectioned at lOu or 2O i, and stained with toluidine blue—0 (Sass 1958).
The slides thus obtained were used for cell size determination. Fifty
measurements were made near the midrib on the horizontal (parallel to
the plane of the blade) and vertical dimensions of both the upper epidermis
and the upper palisade mesophyll. The cell types were chosen because
of their characteristic shapes, resulting from expansion in perpendicular
planes, and their developmental relationship as adjacent tissues.
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—16--
The other halE of the leaf was weighed, cut into thirds by length,
and reweighed. The area of each section was determined from tracings
with a Lambda fIodel LI—3000 portable area meter. For eventual volume
calculations, the thickness of each section was determined with a micro—
meter (Zeus Co.). Thickness of sections that were too small for measure-
ment was established based on a correlation between thickness as measured
on fixed and fresh tissue.
These pieces were then placed in 2—3 ml of 57. chromic acid.
Maceration of the tissue was complete in 3—5 days at room temperature
with occasional shaking on a variable speed whirl mixer. No evidence
of cell wall destruction nor loss of intracellular components was
observed. The cell suspension was diluted with 97, NaC1 to between
10,000—40,000 cell/.5 ml and counted by a Model B Coulter Counter
(Haileah Electronics). Tissue density was calculated subsequently.
RESULTS . The pattern of leaf growth for the duration of the
experiment is shown in Figure 4. A logistic curve:
1 where:
f(x) =
1 - (1 — A \e 1 = maximal length
1 m t\lm 11) 1. initial length
r = growth rale
was fitted to both sets of data. It is seen that under the enhanced UV—B
radiation regime, plants had significantly shorter blades (p
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—17—
provided by Sisson and Caidwell (1976). In addition, the fitted logistic
curve predicts that control leaves grew significantly faster (p
-------
—18—
implies that there is possibly an earlier introduction of the expansion
phase of leaf growth in this treatment.
The cell density data are shown in Figure 6. This again is a
composite graph using data from the three areas of leaf tissue. A
power curve
bx
f(x) = ae where:
a maximal intercept
b slope
was fitted to each set of data yielding estimates for intercept and
slope. Use of the t—test statistic showed that though the intercepts
are not significantly different (p>O.lO), the slopes are (p
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—19—
to depress photosynthesis, UV radiation may indirectly influence the
structure of the leaf through limiting the supply of energy for these
processes. Ultraviolet radiation may also affect the normal activity
of auxin since it is a UV—chromatophore (Giese 1964, Curry et al. 1956).
If it is assumed that with UV—B irradiation the rate and duration
of cell division proceed normally, but that the rate (but not the
ultimate extent) of cell expansion is reduced, one might expect patterns
of growth similar to those shown in Figure 7A. The increase in cell
size and decrease in tissue cell density would lag behind that of the
control plant growth, instead of proceeding as the present data suggest.
In addition, the hypothesized pattern of blade growth (Figure 7A), which
predicts that mature leaves from both treatments reach tli same size,
is not consistent with the observed pattern (Figure 4) in which mature
size differs significantly.
If, rather than the rate, the duration or extent of cell expansion
was reduced by UV—B radiation while all other processes were unchanged,
one would expect to observe data as suggested in Figure 7B. Cells
in mature leaveswould be smaller, on the average, and tissue would,
therefore, have a greater cell density in the UV—B treatment plants.
This prediction is not in agreement with the pattern observed, where
mature cell size and tissue cell density are not found to differ
significantly between treatments (Figures 5 and 6).
The rate at which cell division occurs is determined generally
by the sequence of DNA replication, spindle formation and cell expansion.
As it is known that D ’JA, RNA and proteins are UV—B chromatophores
(Ciese 1964), these are possible targets for UV radiation. The inhibitioxi
of cell division by UV radiation has been demonstrated (Cleaver 1965,
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—20—
1967, Domon and Rauth 1968, Bootsma and Humphrey 1968, Han et al. 1971,
Carison 1976a,b). These researchers observed that tne rate of cell
division was most sensitive to UV radiation during the replication phase.
Thus, if the rate at which cell division occurs is reduced, the
growth pattern would likely be that pictured in Figure 7C. When the
period of division is prolonged, fewer cells are in the process of
dividing at any given time. As normal leaf unfoldment proceeds, more
cells cease dividing and undergo expansion and differentiation. Therefore,
the average cell size would begin increasing before that in control plants.
Accordingly, tissue cell density would decrease earlier. The result
would be fewer cells in the leaf and, therefore, a smaller mature leaf
size. The model predicted by this hypothesis is in agrecm nt with the
data presented (Figures 4, 5 and 6).
The duration of the cell division phase in leaves is a variable
character and differs between species. Cessation occurs when leaves
are 1/5—1/6, 1/4—1/3, 1/3—1/2 or 1/5 final size for tobacco (Avery 1933),
cucumber (Hilthorpe and Newton 1963), spinach (Sauer and Possingham
1970), and cocklebur (Maksymowych and Erickson 1960), respectively.
Division following unfoldment is independent of the intensity of photo-
synthetically active radiation (Milthorpe and Ne’ .iton 1963). However,
there e
-------
-density would decrease earlier in leaf development. Again, the data fit
this hypothesized pattern (Figures 4, 5 and 6).
When con paring the -obsirved di a uitFi those predidted by the models
of leaf growth in terms of cell size,- tissue cell density, and blade
lengt , it is reasonable to refute the possibility that some modification
of the processes of cell expansion, either in its rate or duration, is
responsible for reduced leaf growth in UV irradiated plants. On the other
hand, the data strongly indicate that the decreased rate of leaf growth -
observed under enhanced UV—B radiation is influenced primarily by an
alteration in the cell divfsion process.
A corroboration of the conclusion may be gleaned f1’óin the pattern of
leaf development normally expected in plants. The numbčrof cells in
the leaf primordium affects uftimate leaf size. It has been shown that
this number is dependent upon the intensity of visible r adiation (cell
number increases with increasing intensity) and the poSition of the leaf
along the vegetative axis (Milthorpe and Newton 1963). Typically, the
first tiue leaves of a plant (e.g. Xanthium, Helianthus ; and Rumex ) show
a slower relative growth r Le and reach a smaller matu -e size than later
leaves on the same plant. This pattern is a function of’a slower cell
division rate which results in fewercells in the prima dium (Milthorpe
andNewton 1963). The cells in these firsttrue leave eventually attain
a size and shape sirnilar to those in the later leaves, but due to fewer
cells initially, the leaves are smaller. -
Plants exposed to enhanced UV—B radiation exhibit a similar pattern of
growth. Cell size and shapeare comparable in mature leaves of both tJV—B
and control treatments (Figures 4, 5 and 6). If the analogy holds,
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—22—
one can thus conclude that leaves of plants exposed to enhanced IJV—B
radiation, like the first true leaves of normal plants not subjected to
UV—B radiation, are limited in size and in growth rate by a smaller
total number of cells due to fewer cell divisions. By experimental design.
the number of cells in the leaf primordium for both treatments should have
been the same, therefore, it is assumed that the difference between
treatments was a product of fewer division in the UV—B treatment plants
during unfoldment.
Further experimentation may yield evidence which would enable us to
determine whether the UV radiation acts to reduce the rate or the
duration of cell division. Preliminary observation of the frequency of
mitosis in leaves of both UV—B and control treatments indicates that the
duration of the cell division phasedoes not differ. Mitotic frequencies
were similar in both control and irradiated leaves, i.e., 32%, 24%
(control)—25% (liv), 7% and 0%, approximately, for harvest days 3, 4, 6
and 16, respectively.
The ecological implication of rcduced leaf growth by enhanced
UV—B radiation appears clear. Reduced leaf area would reduce carbon
fixation by the plant, which in turn reduces plant biomass and may alter
competitive effectiveness. An inderstanding of the mechanism involved
in UV—B radiant impairment of leaf growth should greatly facilitate
prediction of how this stress may interact with other stresses to which
plants are normally subjected.
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—23—
4. Competitive interactions in plant populations exposed to enhanced
UV—]3 radiation. (Fox)
This work has now been completed and a manuscript for submission
to the Journal Oecologia is being prepared which will encompass the
work discussed below.
Damage caused by moderate UV—B irradiation is subtle and often
difficult to detect in single plants grown in individual pots. Irradiating
plants grown under conditions of competitive stress may augment the
expression of the radiation damage as well as providing conditions for growth
which are more representative of natural conditions. Further, plants
differ in their susceptibility to UV—B radiation, probably due to
differences in leaf geometry, morphology, and repair capabilities. Because
of this differential tolerance to UV—B irradiance, plant species might
differ in the amount of injury or growth impairment which they incur and
competitive ability could be influenced. Therefore, this approach permits
a better assessment of potential shifts in competitive balance in natural
communities which might occur under increased UV—B irradiance. Based on
these premises, the following hypothesis was proposed.
1. The competitive balance between species is altered by a moderate
increase of UI/_B radiation.
Abiotic stress factors such as nutrient deficiency, water stress, and
extreme temperatures might be expected to alter the expression of UV—B--
induced damage or growth impairment. Herbivory, competition, and other
biotic factors could also influence the expression of UI/—B damage in
plants.
The test this thesis, the following hypothesis was set forth:
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—24—
2. If the UV—B insult is exacerbated by’thiterspecific competitive
stress, then elevated UV—B irradiance should have a greater effect under
conditions of most severe interspecific competition.
If changes in competitive response actually occur under enhanced UV—B
irradiation, they are probably the result of altered physiological
processes. This should also be reflected in altered growth parameters
of individual plants. Therefore, the third hypothesis to be tested was;
3. Enhanced UV—B irradiance will result in altered individual
plant growth parameters.
If hypotheses I and 3 are supported, then a corollary hypothesis
can be developed.
4. Since UV—B radiation is generally detrimental to plants, the
growth of a species can appear to be enhanced, only as a result of UV—D
radiation growth impairment of the competing species.
Eight plant species pairs were chosen to represent competitive
Interactions encounted in actual field systems. Several criteria were
used in the selection of the species pairs. Each competing pair represented
a biologically meaningful association. Some attempt was also made to
pair a species which might be sensitive to UV—B radiation with a species
which might be resistant. Sensitivity to radiation at 245 rim (Cline
and Salisbury 1965) was used to predict sensitivity to IJV—B radiation.
(There are drawbacks associated with this procedure since the effects
on plant growth of enhanced ISV—B radiation may not be the same as the
effects with UV—C radiation.) Table 2 is a list of the competing pairs
of species studied which comprised three general ecological categories:
agricultural crops and their associated weeds, montane summer range species,
and disturbed—area weedy associations. The timing of growth was considered
-------
Table 2. Species pairs grown in competition arranged by type of plant association.- - ’
Plant Association
Competing_Species_Pair
Species 1
Species 2
Agricultural crops and
associated weeds
A]yssur alyssoides L.
Arnaranthus retroflexus L.
Brassica nigra (L.) Koch
Aniarantlius retroflexus L.
Setaria glauca (L.) Beauv.
Pistim sativurn L.
Nedica o sativa L.
Iledicago sativa L.
Al1 urn cepa L.
Trifolium pratense
L.
Montane forage associates
(summer range)
Disturbed area weedy associates
Poa pratensis L.
Brornus tectorum L.
Plantago patagonica Jacq.
Ceum mzicrophyllum Willd.
Alyssum alyssoides L.
Lepidium perfoliatum L.
nomenclature follows llolmgren and ReveaL, 1966.
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—26—
and each species was paired with another species which normally grows at
the sane time of the season. The findings from five of these pairs will
be discussed in detail under Results.
METHODS . Seeds from each competing pair ware randomly sown in 10 cm x
10 cm pots using a modified DeWit replacement series (Dewit 1960). A
constant total density of 50 seeds per pot was maintained, but the relative
densities of the competitive species were varied. Three relative sowing
frequency classes were used with five replicates of each. The classes
were:
Class
1 5 seeds cospecies 1: 45 seeds cospecies 2
2 25 seeds cospecies 1: 25 seeds cospecies 2
3 45 seeds cospecies 1: 5 seeds cospecies 2
Each pot was partially filled with a coarse screened potting soil mixture
over which was placed 1—2 cm of finely screened mixture. The potting
mixture contained two parts screened clay—loam and one part commercial
patting soil. The mixture was then screened and sterilized. The seeds
were randoinly sown and covered with 1 cm of the fine screened potting
mixture. Immediately after sowing in late July the pots were placed in
field plots for the duration of the study. The pots were watered each
morning to field capacity with an automatic individual pot watering
system.
For seven hours each day centered about solar noon. UV—B radiation
was supplemented with five filtered Westinghouse FS—40 “sunlamps” situated
43 cm above the pot rims. These fixtures directly irradiated a 120 cm x
120 cm area in which the pots were arranged in a 10 x 10 pot grid. The
outer row of pots on each side was sown with a mixture of Medicago sativa
-------
—27—
and Amarantlius retroflexus and left as an unsampled border. Since the
outside row of pots had no neignbors on at Jeast one side, an edge effect
could deve]op due to differences in irradiation and wind movement. The
unsampled border prevented the development of an edge effect on experimental
plants. The remaining test pots were randomly reassigned new positions
every three days to help reduce the effects of uneven irradiation and
other microenvironmental differences.
The surviving density of each cospecies was determined at intervals
during the growing season. At harvest in September and October all plants
were individually clipped at ground level and shoot dry weight was
determined after drying at 60°C. Shoot height and leaf area were determined
for those species pairs for which it was feasible. Leaf areas were
measured with a Lambda Model 11—3000 area meter.
Two UV—B irradiance treatments were provided for all density classes
of each competing species pair by using identical lamp fixtures with
different plastic film filters. The lamps in the supplemented UV—B
irradiance treatment were covered with one layer of Kodacel TA 401 (5 mu;
0.13 mm) (Kodak Co.) polyester film. Since this film solarizes and
becomes more opaque to UV—B radiation, these filters were changed every
three to five days. The lamps in the control treatment were covered with
one layer of Mylar Type A (10 mu; 0.25 mm) .(Dupont Co.) polyester film.
Figure 8 shows the spectral irradiance of these two treatments for
August 16 when the sun was at an angle from the zenith, 0, of 370 and
stratospheric ozone concentration was about 0.295 atrn•cm (Hering and
Borden 1967). At this time of year, 0 is 28° at solar noon. At 0 of
370, the IJV—B supplemented irradiance (curve B) corresponded to a 0.08
atm cm decrease in stratospheric ozone calculated from the equation of
-------
—28—
Sisson and Caidwell (1975). This represented a 27 percent decrease in
ozone from a base level of 0.295 atm cni.
Calculation of the competition parameters . The Dewit model of
competition (DeWit 1960) evaluates competitive interactions by determination
of the relative crowding coefficient, k 12 , which is characterized by:
0 Z M
k (1)
12 Z °2 N 1
where Z 1 , 22 are the number of seeds sow-n for species 1 and
species 2, respectively, and + = constant,
N , N 2 are the respective yields (e.g. dry matter, number
of seeds produced) when species 1 and 2 are grown
in monoculture,
and 01, 02 are the respective yields then species 1 and 2
are grown in mixed stands.
By substituting the experimental values of N 1 and N 2 and 01 and 02
for each combination of seed sowing densities 21 and Z 2 , one or more
values of can be obtained. This value is substituted into the
following equations
A 1 = k 12 z 1 (k 12 Z 1 + Z 1 ] 1 and A 2 z 2 [ k 12 Z 1 + Z 2 ] (2)
where z 1 , z 2 are the respective relative seed frequencies for species
1 and 2 and range from 0 to 1 s .ich that the sum of both is 1.
21 z
and
A 1 , A 2 are the respective areas available to species 1 and 2 and A 3 + A 2
or a constant. These values of A 1 and A 2 are plotted on the abscissa
and the values of 01 and 02 for the corresponding values of and
are plotted on the ordinate. If two straight lines, one for each species,
-------
—29—
do not result, then these values of k 12 , N 1 , ond N 2 arc adjusted in
equations 1 and 2 until straight lines are derived. By such an iterative
method, a single estimate of k 12 is obtained.
Changes in competitive interaction under different conditions are
often evaluated by comparing crowding coefficients estimated using this
method (e.g., Friedman and Elberse 1g76). However, since only a single
estimate of k 12 is obtained, differences between treatments cannot be
evaluated iith statistical confidence.
In order to treat differences of k 12 statistically, a different method
of calculation was dev. sed. Multiple estimates of the were derived
for both the control and enhanced UV—B radiation plots by substituting
values of 01 and 02 at l Z 2 25 seeds into equation 1. Since
and Z 2 were equal and thus cancel in the equation, the calculation of
the k 12 depends only on the ratio of the yields °1’°2 for each replicate
pot. The mean values of all replicate tnonoculture yields of species 1
and 2 were used to derive a constant value of N 2 /N 1 . Since an estimate
of the variance of each treatment population can then be made, standard
statistical tests can be performed. In this study nonoculture yields
(at z 1 = 1.0, = 1.0) were not determined. Yields under nearly
monospecific stands (at z = 0.9, z = 0.9) were determined arid the ratio
1 2
of these yields M /N was substituted for the constant M 2 /N 1 in equatIon 1.
This was not substantially different from the k 12 calculated by
DeWit’s method. The values of k 12 reported in three different
competition studies in the literature were correlated with the calculated
values of k 12 and k 2 using the method desctibed above. To determine k 2 .
values of were interpolated from the reported data. The correlation
indicates that in those cases where the reported k 12 differed from unity,
-------
—30-
if k 2 differed from k 12 it was always biased to a more conservative
estimate of the competitive imbalance than k 12 (i.e., k 2 was closer to
1.0 than was k 12 ). Therefore, calculating the k 2 by this method gave
an accurate and resonable estimate of the k 12 and if k 2 differed from
k 12 , it would tend to always under stimate rather than overestimate the
departure of k 12 from unity.
When k 2 equals 1.0, neither species is considered to have a
competitive advantage. If k 2 is greater than 1.0 species 1 has the
competitive advantage whereis if k 2 is less than 1.0, species 2 is in the
position of competitive advantage.
RESULTS. Amaranthus retroflexus: Medicago sativa . The density at
four sampling dates of Amaranthus retroflexus (species 1) and Medicago
sativa (species 2) grown in competition is shown in Figure 9. At 13 days
after sowing, lower densities of both N. sativa and A. retroflexus
occurred under the enhanced UV—B treatment when initially sown at 0.1/0.9
ratio (Figure 9a). This difference was not maintained at subsequent
sampling dates, however. The density of N. sativa grown under enhanced
1W—B radiation at z 1 22 0.5 increased during the season and at the
time of harvest was greater than the density under the control conditions
(Figure 9d). The increase in shoot biomass (Figure l0 for N. sativa
at the intermediate seeding frequency (z 2 = 0.5) was due to greater
numbers of plants (Figure 9a) and was not the result of a change in mean
plant weight (Figure lOd), while the decrease in shoot biomass at higher
seeding frequency (z 2 0.9) was due to a decrease in the mean weight
of each individual and not to a change in the number of plants. The
decrease in shoot biomass at = 0.5 for A. retroflexus was due to an
-------
—31—
apparent decrease in the weight of individual, plants. This decrease
in plant size was obscured by the presence of flower and seed heads on
some of the A. retroflexus individuals which contributed to a large
variance in the mean shoot weights. However, the size difference is
clearly illustrated by the reduction of plant height under the enhanced
UV—B conditions (Figure 11).
Plant response to competitive stress is not limited to fluctuations
in mortality but may also include considerable change in individual
plant size. Paimblad (1968) reported that Capsella bursa—pastori
responded to increasing interspecific competition by maintaining a
relatively high density of smaller individual plants. increased UV—B
radiation did not cause higher mortality to 14. sativa grown at high
density (z 2 = 0.9) but did result in the same density of smaller individual
plants when compared to the control. This appareatly was also the case
for A. retroflexus grown at z 1 = 0.9 and 0.5. At somewhat higher levels of
interspecific competition frornA. retroflexus (z 1 = = 0.5) M. satlva
had higher mortality in the control group compared to the group rec ivir1g
enhanced UV- ’B radiation, while the mean plant weight of the two groups
remained the same.
The largest mean individual shoot weight of H. sativa occurred under
conditions of lowest interspecific competitiye stress (z 2 = 0.9) and the
lesser UV—B radiation load. In the control plants the reduction in plant
weight at the other two seeding frequencies from this maximum value
could only be the result of increased competition from A. retroflexus .
However, in the UV—B treatment group the reduction of individual plant
weight occurred under the lowest level of interspecific (z 2 = 0.9)
competition, and therefore was apparently the result of UV—B induced damage.
-------
—32--
Since growth of the M. sativa population appears to be impaired by UV-13
radiation and interspecific competition did not exacerbate the depression
of individual plant growth, the higher shoot biomass produced under enhanced
TJV—B radiation at 0.5 is initially perplexing. There was a reciprocal
adjustment in biomass between the two species in which N. sativa succeeded
at the expense of A. retroflexus . This indicated that the competitive
ability of N. sativa was less adversely affected by UV—B radiation than
was the competitiveness of A. retroflexus . The competitiveness of Medicag
under elevated UV—B radiation was clearly shown by the value of b 1 12 = 0.73
(Figure 12). Since it was less than 1.0, this indicates that M. sativa
had a competitive advantage over A. retroflexus . The situation under the
control conditions was reversed, with Amaranthus being tde more competitive
species as indicated by a bki 2 greater than 1.0 (3.56). Therefore,
although both species were affected by UV—B radiation, the competitive
balance shifted in favor of Medicago under enhanced UV—B radiation. The
change in dkl 2 from 1.84 to 0.71 also indicated that competitive inter—
action had changed.
The d 1 12 is probably not as good an indicator of competitive balance
as is the bkl 2 . Except for vegetatively reproducing species, the density
of shoots is likely not as indi ative of competitive ability as is the
total biomass produced by the species. Forannual species the production
of seeds would likely be the most meaningful indication. Since
biomass is a reflection of the amount of resources utilized by a species,
the competitive success of that species would seem to be better expressed
by total biomass, rather than the number of Individuals in the population.
Imagine the hypothetical situation of two species, each grown in competition
with a third species which maintains the same biomass and density when
-------
—33—
grown with either competitor. If the first species produces a few
individuals of large size then it would appear to be as successful
against the competitor as would the second species which produced the
same total biomass but from many individuals of smaller size. Thus the
bki 2 which is calculated from biomass measurements would seem to be the
better indicator of competitive balance. However, the manner in which
species interact competitivelywith each other may change independent of
the competitive balance as indicated by total biomass. In another hypothetical
situation, the bionass produced by each of two competitors might not change
under two treatments, while the number and size of individuals resulting
in that constant biomass might differ between the treatments. Thus, the
competitive interaction would have apparently changed while competitive
balance, taken as total biomass, would not. Computation of the dki 2
depends on density and would thus seem to reflect competitive interaction.
Van et a]. (1976) measured photosynthesis and biomass of thirteen
crop species and noted that none of the four species having the C 4
pathway of carbon assimilation were susceptible to elevated UV—B
irradiance. Amaranthus retroflexus , a C 4 species (Welkie and Caldwell
1970) showed sensitivity to tJV—B radiation in this study.
Plantago patagonica: Lepid um perfoliatum . The density of four
sampling dates of Plantago patagonica (species 1) and Lepidium perfoliatum
(species 2) are shown in Figure 13. At all sampling dates there were
greater numbers of L. perfoliatun at the relative seeding frequency
z 2 = 0.9 when grown under enhanced tJV—B radiation. The density of P.
patagonica at z 1 = 0.5 was higher under conditions of increased UV—]3
radiation at 13 and 89 days (Figure 13a and d). The increased density of
-------
L. perfoliaturn (z 2 0.9) caused the apparently higher (albeit
statistically insignificant) shoot biomass (Figure l4 since the weight
of individual shoots did not change. An increase in individual L.
perfoliatum shoot weight, however, accounted for increased shoot biomass
at 0.5. There was no change in the shoot biomass produced by P.
patagonica (z 1 0.5) due to increased IJV—B radiation. The constant
bloinass was the result of a greater number of individuals of smaller size
surviving in the UV-B treatment group. The differences in density of
survivors which occurred in both species are reflected in the change in
d 1 l2 (Figure lsaand b). The values of d 1 l2 indicate that the competitive
interactions of L. perfoliaturn and P. patagonica were changed under a
higher iSV—B radiation environment. The competitive balance did not change,
however, as shown by the values of bki 2 which indicated that L. perfoliatum
had the advanLage in both cases.
Amaranthus retroflexus: Allium cepa . Another species which showed
plasticity in its response was A. when grown with A. retroflexus
(Table 3). At high intraspecific density the shoot biomass remained
unaffected. Although the mean plant size was smaller (both height and
weight) this was compensated by a greater plant density.
Poa pratensis: Ceum macrophyllurn. The density of Poa pratensis
(species 1) at z 1 0.5 and 0.9-was lower under enhanced UV—B radiation
when sampled 13 days after sowing. At z 2 0.5 Geum macrophyllum (species
2) exhibited lower density at the same sampling date (Figure 16). At
later sampling dates the densities of both species increased and there
were no differences between treatments.
-------
TaL1e 3• Vc; 7 .nst of cot7ot ng spocien pair; to rwci leve l ; of UV-lI radlntton.
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-------
—36—
A significant shift in the competitive balance between P. pratensis
and C. macrophyllum occurred under enhanced UV—B radiation (Figure 17a and d).
The values of bki 2 indicate that C. macropliyllum had a competitive
advantage in the control group and P. pratensis had an advantage under
enhanced UV—B radiation. There were no significant differences in the
density of surviving plants at the time of harvest for both groups, which
is also reflected in the lack of change of the dkl 2 (Figure l7a and b).
Therefore, the shift in competitive balance had to be the result of
differences in individual shoot biomass. The apparent reciprocal adjustment
in the shoot biomass (Figure 18a) indicated that when mutual interspecific
competition was most severe (z 1 = z 2 = 0.5) the combined effects of IJV—]3
radiation damage and competitive stress on G. macrophyli m allowed the
apparently more resistant P. pratensis to utilize a larger share of the
resources. At a high density of C. macrophyllum (z 2 = 0.9) there was no
difference between treatments in shoot biomass, indicating that Ti l l —B
radiation apparently did not have a direct effect on shoot biomass nor
even an indirect effect mediated through intraspecific competitior . However,
at z 1 0.5 the individual shoot weight decreased in the treatment
group below the value for z 2 = 0.9, but was not less than this value in
the control group. The reduct?on was apparently the result of increased
competitive stress from P. pratensis presumably because Geum was somewhat
impaired by the Till radiation. In the control group, the individual shoot
weight of C. macrophyllum (z 2 0.1) was reduced by increased competition
from P. iratens such that there was no difference between the treatments
(Figure 181)). The reduction of both total shoot
biomass and mean individual, shoot weight in C. macrophylium in response to
-------
—27—
increasing competitive ctress of P. pratens s became apparent at a lower
density of P. pratensis (z 1 0.5) for plants grown under a high UV—B
radiation environment than for those plants grown under the control
conditions.
yssum alyssoides: Pisurn sativum . Figure 19 illustrates the density
of Alyssum alyssoides (species 1) and Pisum sativum (species 2) at three
sampling dates after sowing. The density of A. alyssoides when sown at
0.9 was lower under conditions of enhanced UV—B radiation at 13 and
42 days after sowing (Figures l9a and c). At = 0.5 A. alyssoides
density was also lower at 13 and 37 days (Figures 19a and b). The
number of P. sativum individuals at = 0.1 was lower in the UV—B
treatment group at 13, 37 and 42 days. The density of P. sativum
through the growing season at z 2 0.5 and 0.9 is shown in Figure l9d.
The density of P. sativum increased to some extent from 13 to 37 days
and then decreased by 42 days for both treatment and control groups at
z 2 = 0.9 and for the treatment group at a 2 = 0.5. The density of the
control group at = 0.5 remained constant at all sample dates. T -e
increase and decrease in density was most pronounced for the enhanced
UV—B gro’ ps at both z 2 = 0.5 and 0.9. The reduced density in the treatment
groups was significant at 13 arid 42 days but not at 37 days.
Evaluation of the competitive status of A. alyssoides and P. sativum
indicates that there was no change in interaction or balance as a consequence
of increasing the UV—B radiation dose, and that P. sativum had the advantage
in both experimental groups (Figure 20c and d). These interpretations
must be regarded critically, however, due to some bias included in the
calculations of the relative cro ding coefficients for this particular
-------
—38—
species pair. The size of individual P. sativum plants and the consequent
shoot bioinass was substantially larger than that of A. alyssoides .
Consequently H/Mi probably overestimated and the crowding co—
efficicnts may not reflect the actual balance or change in competitive
interac don.
The development of the P. sativum leaf canopy was an important factor
in the response of this species to UV—B radiation. Pisum sativum formed
dense canopies with leaf area indices ranging from approximately 1 at
z 2 0.1 to 10 at z 2 = 0.9 at the time of harvest (Figure 2l . In the
control groups, P. sativum density was close to the maximum value even
at the earliest sampling date (Figure 19 ), and did not decrease until
leaf canopies were well developed at which time thinning occurred probably
due to increased intraspecific competition. In the treatment group the
density of plants was considerably lower at the first sampling date
(z 2 = 0.5 and 0.9). As the leaf area increased, the number of plants
also increased. Leaves are essentially opaque to UV—B radiation. Thus
a well developed canopy probably afforded some protection for developing
seedlings against the UV—B radiation. The result was that no significant
difference in density existed between the two groups by 37 days. Since, at
this time, the number of surviving plants in the enhanced UV—B group
represented most of the seeds sown (35 plants from 45 seeds for z 2
0.9 and 20 plants from 25 seeds for z 2 = 0.5) the large differences observed
earlier could not have been the result of seedling mortality. Inhibited
germination or arrested seedling emergence appear to be the cause of
this difference early in the experiment. The inhibition was removed and
development continued when the leaf canopy increased reducing the amount
of IN—B radiation reaching the soil surface. It was also observed that
-------
—39—
the vigorous germination of P. sativum disrupted the soil surface causing
seed coats to become exposed to UV.-B radiation.
Enhanced UV—B radiation reduced the density of A. alyssoides and P.
sativum for several seeding frequencies on many sampling dates (Figure 19).
This indicated that both species were damaged by exposure to UV—B radiation.
Total leaf area and shoot biomass (Figures 21a and 22a) which like
density, are characteristics of populations and hot of individuals, were
reduced in many instances under the increased lW—B radiation environment.
With the exception of individual leaf area at z 2 0.1, neither the weight
nor the leaf area of individual plants were affected. The differences in
total shoot biomass at the population level between treatments were the
result of differences in density. Although leaf area and weight of
individual plants were riot affected, shoot height of P. sativum was
sensitive to elevated UV—B radiation at higher intraspecific densities
(Figure 23).
Pisum sativum plants had the highest individual leaf area under
conditions of lower UV—B radiation and lower intraspecific density (z 2 -
0.1). As the seeding frequency in the control group increased (z 2 = 0.5)
mean leaf area decreased from the higher value at = O.i and remained
at this low value as the densit increased still higher (z 2 0.9) . In
the U 1 /—B treatment leaf areas was depressed to approximately this same
low level even at the lowest relative seeding frequency at = 0.1.
Thus, for Pisum plants grown in control conditions, leaf area was
apparently reduced as intraspecific competitive stress increased. However,
the leaf area of plants raised under enhanced UV—B was low at low levels
of intraspecific competition. This may have occurred becduse either
-------
—40—
leaf area was affected at lower levels of intraspecific competition when
plants were grown under higher IJV—B irradiance than when grown under lower
UV—B irradiance, or because leaf area was affected directly by UV—B
radiation regardless of intraspecific stress.
DISCUSSION . In addition to P. sativum whIch exhibited sensitivity
to UV—B radiation at germination or seedling emergence, several other
species appeared to be sensitive at the seedling stage. Lower densities
were often observed at early sampling dates, but the differential
disappeared by later dates (Figures 9 and 16, Table 3). This would
have been due to other plants replacing those that suffered mortality
as seedlings or to the renewed germination which was previously arrested.
Species which exhibited the phenomenon included P. pratensis , C. marcophyllum ,
T. pratense , both A. retroflexus and N. sativa when grown together, and
A. alyssoides , when grown competitively with B. tectorum . Density was
the only parameter measured for seedlings.
Although the seedling stage appears to be a time in phenological
development when several species are sensitive to TJV—B radiation, how
they are affected is unclear. Studies which follow individuals of the
population through frequent censuses, and furnish information on individual
plant parameters such as seedling height and biomass, would do much to
elucidate the effect. Seedling mortality of plants in natural systems might
be expected to be more severely influenced by IJV—B radiation since water
stress and competition from other species generally would be greater
than. in systems of well watered pot—grown plants. Plant species which
germinate in June when the solar altitude is high would be exposed to
-------
—41—
greater UV—F3 radiation loads and usually also greater water stress than
species which germinate earlier in the spring.
The concept that other strcss factors may influence the expression of
growth impairment by UV—B radiation has significance in the understanding
of the impact of enhanced UV—B irradiance on natural systems. Specifically,
the hypothesis that growth is more impaired by TJV—B radiation under the
most severe interspecific competition was examined in this study. The
relative seeding frequency where the densities of the competitors were
equivalent (z 1 = = 0.5) should exhibit the greatest effect on growth
(except when one species of the pair has large seeds). The hypothesis
was supported in the case of Plantago patagonica, Geum macrophyllum,
Trifolium pretense and Brassica nig (Figures 13, 14, 16, and 18,
Table 3). The growth of these species was affected only at z 1 0.5.
Three other species, Bromus tectorurn, Setaria and Amaranthus
retroflexus when sown with Allium cepa showed effects on growth only at
= 0.9 (Table 3). These results do not support the hypothesis. This
suggests that for these species, intraspecific competitive stress
apparently exacerbates UV—B radiation insult more than does interspecific
competitions. To test the impact of intraspecific competition stress,
these species would have to be grow-n at various densities in monoculture.
Pisuni sativum appeared very sensitive to UV—B irradiance and exhibited
impairment of growth regardless of the degree of interspecific competition.
The renviining species were apparently affected by both types of competitive
stress and exhibited changes in growth at intermediate and high relative
seeding frequencies in most cases.
In addition to the influences of competitive presence on the expression
of UV—B damage, competitive balance itself may be affected by increased
-------
—42—
UV—B irradiance. This hypothesis was supported by the altered competitive
balance observed in three of the species pairs studied. The competitive
interactions of Plcntago patagonica and Lepidium perfoliatum changed in
response to supplemented UV—13 irradiance, although Lepidium maintained
the competitive advantage under both UV—B irradiance treatments (Figure 15).
Amaranthus retroflexus and liedicago sativa responded to TJV—3 irradiation
by changing both competitive interaction and balance. Amaranthus was more
competitive in the control group and Medicago was the better competitor
in the enhanced UV—B treatment group (Figure 12). Under the control
conditions Geum macrophyllum had the competitive advantage over Poa
pratensis but the competitive balance was reversed under the elevated
UV—B radiation conditions (Figure 17). No alterations in competitive
balance were observed for the remaining species pairs.
The hypothesis that UV—B radiation influences plant growth parameters
was tested. There is much evidence to support this hypothesis. Leaf
area of individual plants was reduced far Fisum sativum under an enhanced
UV—B radiation regime (Figure 16). shoot height decreased for Amaranchus
retroflexus , Allium cepa and Pisum sativum (Figures hand 23). tndividual
shoot weight was sElected for Setaria glauca, Alliun cepa, Alyssum
alyssoide , Bromus tectorum 1 Lepidiun perfoliatum, Plantago patagonica,
Geum macrophyllum , and Nedicago sativa at various relative seeding
frequencies (Figures 10, 14, 18, and Table 3 ). Mortality of individuals
was affected and consequently at the population level the density at
different sampling dates for various relative seeding frequencies was
altered from Bromus tectorum , alyssoides, Alliurn cepa, Brassica
nigra, Trifohium pratense, Plantago patagonica, Lepidium perfoliatum , Poa
pratensis, Geum macrophyllum, Amatanthus retroflexus, Medicago sativa ,
and Pisum sativum (Figures 9, 13, 16 , and 19, Table 3).
-------
—43—
The final hypothesis tested in this study was that since the effect
of UV—B radiation is considered to be generally detrimental to plants,
when growth of a species does appear to be enhanced, then this occurs
only as a result of UV radiation damage to the competing species. For
the species pairs A. retroflexus vs. A. cepa , B. nigra vs. f. sativa ,
A. alyssoides vs. P. sativurn , and S. glauca vs. T. ratense the changes
which did occur in response to UV—B irradiance were always detrimental
to growth (Figures 19, 21, 22, and 23, Table 3). Poa pratensis when
grown with Geurn macrophyllum and Nedicago sativa when grown with Amaranthus
retroflcxus showed increased growth under ISV—B radiation, but there were
concomitant decreases in the growth of their competitors. This change
in competitive balance was also reflected in the change in bkj 2 for each
pair, and thus supports the hypothesis (Figures 10, 12, 17, and 18).
Bromus tectorum when grown with Alyssum alyssoides and L pidium perfoliatuin
when grown with Pl ’ ntago patagonica exhibited increased shoot biomass at
a single seeding frequency, while their competitors exhibited no decrease
(Figures l4and 15, Table 3). However, there were not changes in competitive
balance (as indicated by bkl 2 ) , as one would expect as a result of an
increase in shoot biomass in one species of a competing pair. There is
at least one other reason why an increase in shoot biomass of a single
species night occur. If a pathogen which restricted plant growth of
one species was sensitive to UV—]3 radiation then increased irradiance
would increase the production of the plant species by reducing the
effectiveness of the pathogen.
-------
—44—
5. Interactions between enhanced UV—B radiation Stress and Water Stress
on Plant Growth (1-jolman).
Many aspects of the natural environment must be considered in
accurately predicting the impact of an increase in UV—B irradition on
terrestrial ecosystems. It is of primary importance to assess the potential
interactive effect on plants of an enhanced UV—B radiation regime in
combination with other natural stress factors. Stress combinations may
affect plants differently than would any one stress alone and, as a natural
stress factor, UV—B irradiat ion should be examined with this in mind.
Some work has been done on this interaction. Fox (Section 4 of this
report) examined the combination of enhanced UV—B irradiation and intra—
and inter-specific competitive stress and found that, for some species, an
interaction does exist. Tyrelle (1976) working with UV—A (334—365 mm)
irradiation and mild heat stress, demonstrated a synergistic interaction on
survivorship in 1. coil .
One of the most common and widespread natural stress factors is water
stress. In most terrestrial ecosystems abundant and easily available water
is uncommon. Even in well—watered soil, plants may suffer water deficit
if transpiration rates are high. Plants which would be subjected to an
enhanced UV-B radiation stress due to atmospheric ozone depletion might
already be exposed to frequent water stress. Thus information is needed on
the potential effects of an interation between these 2 stresses.
Whole plant responses to a moderate increase in UV—B irradiation or
to a mild water stress are quite similar. Under either condition, photo-
synthesis is reduced (Slatyar 1967, Van et al. 1976) dark respiration may
increase (Sisson and Caidwell 1976, Brix 1962) and protein synthesis is
reduced (CaidweLl et al. 1976, Sullivan 1975). These common plant responses
-------
—45.-
indicate that a factor interaction is likely.
Growth and biornass production seem to be the best indicators of whole
plant response to the 2 stresses since each of the effects of water stress
in UV—B stress listed above will result in a reduction in plant growth.
Therefore, plant growth and biomass production will be used to assess the
interaction. Three hypotheses seem likely:
1) The interaction of enhanced UV—B irradiation and mild water stress
will be synergistic.
2) The interaction of the two stress factors will be negatively syner-
gistic.
3) The interaction of the two stress factors will be strictly additive.
If one stress weakens or damages the plant so that its sensitivity to
damage from the other stress is enhanced, the first hypothesis will be
supported. If one stress depresses a particular process or reaction to the
point that it becomes rate limiting, regardless of the effect of the other
stress, a negative synergisim will be observed. Finally, if the effect of
one stress is simply to augment the damage done by the other then an add-
itive interaction will be observed.
The rate of leaf and stem growth and biomass production of seedlings
exposed to simultaneous water stress and enhanced UV—B irradiation will be
used to assess the interaction. Seedlings are well suited to this research
since they are relatively stress sensitive, quick growing, and small. All
experiments will be carried out in controlled environment chambers. Three
species will be tested; Rumex 2 ientia L., Balsamorhiza sagittata (Pursh)
Nutt., and a grass species that has not yet been chosen.
Two levels of UV-B irradiation and 2 levels of water stress will be
imposed on the plants. Enhanced UV—B irradiation will be supplied by
-------
—46—
Westinghouse FS—40 sun lamps filtered with plastic films. The 2 UV—B
treatments will be; 1) no supplemental UV—B irradiation below 315 and 2)
enchance UV—B irradiation. The enhanced UV—B treatment dosage has yet to
be determined. The basis for choosing the dosage is discussed below.
Superimposed on the UV—B treatments will be 2 levels of water stress.
Soil will be maintained at field capacity for one treatment and at some level
of mild water deficit for the other. Soil water content will be controlled
using the method of Tingey and Stockwell (1977). Individual plants will be
grown in plastic soil frames wrapped in cellulose acetate semipermeable
u mbrane. When immersed in solutions of known osmotic potential, water will
pass across the membrane, between the soil and the solution and result in the
soil water potential coming to equilibrium with the solution osmotic potential.
The osmoticum will be Polyethylene Clycol 20,000 and solutions will be prepared
based on the data Tingey (personal communication).
Before the interactions can be assessed, the UV—B and water stress
dosages must be determined. The whole plant response to each treatment alone
must be approximately equal for both treatments to avoid overwhelming the
plant with one stress when both are applied simultaneously. Work is presently
being done on Ruinex patientia to determine what stress levels will cause
equivalent, measurable plant response. It is important that these stress
levels are easy to obtain and control, and that they are realistic levels to
expect under natural conditions.
A preliminary growth response curve for R. patientia under enhanced uv—B
irradiation is presented in Figure 24 . Eight pots, each containing 40 seedlings
of approximately equal size and stage of growth, were used in the study. After
the seedlings had entered the first leaf stage, 4 pots were placed in a
-------
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-------
l.0
0 .6-
Q2-
0.I
r cv ’
—
F
c .J
Q02
U,
C)
0.01 -
0
0.006 -
0.002 -
0.001 -
290 320
Wavelength (nm)
Figure 1. Spcctrnl irradiaricc ncar solar noon at Mucabaji Pass,
Verle2uela and Heade River, Alaska. (Data are prelirnin ry
and subject to reevaluation.)
Meade River
Mucabaji Pass
300 310
-------
Figure 2.
>.
U
0
E
U i
cc
-7
C ’
‘I
-J
U i
I i_
UJ
-j
0
0 -cDL . C
- — ____J ._____ __..__.____J—_.____ ._ I ____.,_ — ___ — —. ___ __ J
6 0 0 12 i’1 16 13
LEAF AGE (DAYS)
Leaf elongation rates (mm .day ) of the first leaf of Rumex
patientia L. during l’8 days of UV—irradiance (0) (simulating
a 0.18 atm .cm ozone column at solar altitude of 60°) and
control (0) treatment in controlied—environmei t studies.
The curves represent least squares fits of the IJV irradiance
(—-——--) and control treatment ( —) elongation rates.
the dashed line (——--) represents the ex ectcd leaf elongation
rate of Ihe liv irradiated leaf if the reduction in elongation
rate were solely a function of the depressed photosynthetic
rates of that leaf.
Of,
0
4
3
0
‘ - I
C.
0/
/ \
\
\o
\
\
\
0
0
-------
C)
c
0
-D
C
1..
Figure 3. Spectroradiometer measurements ot irradiance in controlled
environment chambers. Irradiation is suppled by a 6000—W xenon
arc lamp with Schott Co. WC—320 glass filters and 1 estinghouSe
Co. FS—40 lamps filtered with 1ylar Type ‘A’ (10 mu) plastic
film (CONTROL treatment) and Kodacel TA 401 (10 mu) plastic
film (UV —Pi ENHANCED treatment).
260 290 300 3 0
Wuvelength (nm)
320 330 340
-------
120
E
E
I
F-
0
z
L i i
-J
1i
I i i
-J
40
Figure 4. Leaf length for the fifth leaf of Rumex atientia L., during 16 days of UV irradiation
(Q ——— ) (equivalent to solar UV—B radiation with an atmospheric ozone concentration of
0.20 atm—cm at 60° solar altitude) and control (o — o) treatment. Irradiation between
400 and 300 nm was 750 Jeinsteinsm 2 s .
80
**
CO NT R 0 L
* * **
UV-B
/
/
20
2
4
6 8 10 2
DURATION OF EXPOSURE (days)
14
-------
Figure 5.
U
NJ
c i )
-J
-J
U
U
0 50 100
BLADE
Cell size of the palisade mesophyll and upper epidermis of
the fifth leaf of Rurnex patientia L. exposed to UV—B irradiation
( ——— G) (equivalent to solar IJV—B radiation with an atmospheric
ozone concentration of 0.20 atm—cm at 600 solar altitude) and
control (o —o) treatment. Cell length is measured perpendicular
to the piano of the blade, while width Is measured parallel
to the p]ane of the blade.
0 50 100
LENGTH (mm)
-------
7
Figure 6.
6
5
4
3
2
0
E
U
>-
1—
U,
LU
C D
-J
LU
0
LU
C ,)
(I)
Tissue cell density of the fifth Leaf of Rumex j ntin L.
exposed to UV-13 irradiation ( —— o — Q) (equivalent to
solar UV-B radiation with an atmospheric o onc concentration
of 0.20 atm—cm at 60° solar altitude) and control (t — o — )
treatments. The symbols , o and o refer to data collected
from tissue in the tip, middle and basal portions of the
leaf, respectively.
60 80 (00
LENGTh (rnrn
0
2 - 40
BLADE
-------
A B
Figure 7. Models of hypothesized growth patterns as expressed in cell
size, tissue cell density, and total blade length for leaves
exposed to UV—13 irradiation ( ) and control ( )
treatments. (For an explanation of hypotheses A, B, and
C, see tCxt.)
C
— TIME > —TIME > TIME >
-------
Wave’ength (nm)
Figure 8.
Spectral irradiance for control (line A) and enhanced
UV—B (line B) treatment for August 16 at 1:50 p.m. local
time (0 = 37°) with an approxiulate ozone concentration of
0.295 atm cm. The biologically effective UV—B irradiance
wa 0.0332 and 0.0799 J•m 2 for the control and enhanced
UV—B treatment, respectively.
w
0
C
c3
-a
a
290 300 3 O
320
-------
40
Relative Seedrng
sot va
Control : Enhanced UV B
retroflcxus
Frequency of Medicago sotiva/Amoranthus retroflexus
Figure 9.
Density (plantsdm 2 ) of Amaranthus retrofIexus and Nedicago
sativa grown in competition at A) 13 days, B) 37 days, C)
50 days, and D) 89 days, after sowing at three relative
seeding frequency classes, under two levels of UV—B radiation.
Values are means of 5 replicates ±1 S.E. (*p<.O 5 ).
30
20
(0
I
0
40
30
20
I0
0
.11.9 .5/.5 .9/.1 .1/.9 .5/.5 .9/.1
-------
Relative Seeding Frequency of
Medicacjo sotiva / Amaranthus retroflexus
Medicago sativa
Arnoronthus
retroflexus
Figure 10.
A) Shoot bioniass (gdm 2 ) and B) individual plant weight (g)
of Amaranthus retroflexus and Medic sativa at time of
harvest after sowing at three relative seeding frequency
classes, under two levels of UV—B radiation. The means are
reported ±1 S.E. (*p<,Q5) .
Csj
SE
0
CO
E
0
0
0
U)
Enhanced UVB
4-
4-
0
0
U)
a
>
-o
.1 /.9 .5/.5 .9/. 1
-------
E
E
-c
C)
0
0
(I )
40
2.0
Figure 11.
Re lative Seeding Frequency
Medicaqo saliva / Amoranthus retroflexus
Control.: Enhanced UV—V
Amaranthus retroflexus
Shoot height (mm) of Amaranthus retroflexus at time of
harvest after sowing with He4 go sativn at three
relative seeding frequency classes, under two levels of
UV—B radiation. The means are reported ± L S.E.
(*p<. 05).
8
60
.9/. 1
-------
Control Enhanced UV-B
_____________________________________________________________________________________________ _______________________________________________________________________________________________
E
-o A B
• 30
Cr)
.4 —
C
.2
c i 20 1
/ ____o
/
>‘
4 -
—
—
U)
C
,
___ x x
o o_—_
0—
C%4 125 0 o D
/
/
1.00- /
U, /
U) /
0
E0.75 / 0
— /
/ ,0
- 0.50
o /
o x ‘F
/
-c
0.25 X /
o— _
0
0 Amre 0 l.0 mre 0
OMesa” i.0 0Mesa l.0
Relative Seeding Frequency
Figure 12. Replacement diagrams of density (plants•drn 2 ) (A) and (B)
and shoot bionass (g•dm 2 ) (C) and (1)) for Amaranthus
retroflexus (Amre, species 1) and Mcdi c g sativa (riesa,
species 2) at time of harvest when grown under two levels
of UV—B radiation. The dkj2 are J.84 and 0.71 (p<.O5) for
control (A) and enhanced UV—B conditions (B) and the bkj2
are 3.56 and 0.73 (p<.05) for control (C) and enhanced IJV—B
radiation (D).
-------
Control: Enhanced UV—13
Lepidiurn perfoliatum
Plantago atagonica
Figure 13.
Density (plantsdm 2 ) of Plantago patagon ca and Lepidium
pcrfoliaturn grown in competition at A) 13 days, B) 37 days,
C) 50 days, and D) 89 days, after sowing at three relative
seeding frequency classes, under two levels of UV—B radiation.
Values are means of 5 replicates ±1 S.E. ( p<.OS).
c’4
U)
-a
>..
4-
(1)
C
.I/.9 .5/.5 .91.1
Relotive Seeding Frequency of
.1/.9 .51.5 .91.1
Lepidium perfoliafum/ Plantago potagonica
-------
U ,
U,
E
C )
4 .-
0
0
C l )
0)
4 -
0)
C)
4-
0
0
-C
U)
C )
• 0
Con trol:
fl :
Lepidium perfoliat:urn
Plantago patagonica
Figure 14.
A) Shoot biomass (g.dm 2 ) nd B) individual plant weight (g)
of Plantago Th onica and Lepidium perfoliaturn it time of
harvest after sowing at three relative seeding frequency
classes, under two levels of UV-B radiation. The means are
reported ±1 S.E. (*p< . 35 ) .
E
0
S..
. 1/.9 .5/.5 .9/.1
Relative Seeding Frequency of
Lepidium perfoliotum / P anfago rx tcgonica
Enhanced UV—B
-------
Control Enhanced UV—B
40
I:
0.30
U,
0.20- -o
0.10 -
°LO P Ipa 0
— — — Lepe — ——-*1.0
Relative
FIgure 15.
B
p
1
/
/
/
/
/
/
/
x
/
/
0 •
D
—
N
N
I.0 PIpa 0
0———— Lepe ———— I.0
Seeding Frequency
Replacement diagrams of density (plantsdm 2 ) (A) and (B)
and shoot biomass (gdni 2 ) (C) and (D) for Plantago
patagonica (Pipa, species 1) and Lepidium perfoliatum (Lepe,
species 2) at time of harvest when grown under two levels of
UV-B radiation. The dkjz are 0.70 and 1.30 (p<.O5) for
control (A) and enhanced UV—B conditions (1 ) and the bki2
are 0.75 and 0.68 (no significant difference) for control
(C) and enhanced UV—B conditions (D).
A
—
x x
C
0
x
-o
0
0
-------
40
20
10
0
Relative Seeding Frequency of
Control: Enhanced UV—B
Geum macrophyllum/ Poa pratensis
Geum macropyllurn
Poa pratensis
Figure 16.
Density (plants .dm 2 ) of Poa pratensis and Geum macrophyllum
grown in competition at A) 13 days, B) 37 days, C) 50 days, and
D) 86 days, after sowing at three relative seeding frequency
classes, under two ]evels of UV—B radiation. Values are
means of 5 replicates ±1 S.E. (*p<.OS).
40
30
20
I0
0
(I )
C
a
30
.1/.9 .5/.5 .9/.L .1/.9 .5/.5 .9/.1
-------
c’J
E
U,
C
0
0
>
.4-
C’)
C
C)
C
E
-
C ’)
(1)
0
E
0
-4-
0
0
-c
40
30
20
l0
0
o: o
0.20
ojo
0
Figure 17.
Control
A
—0
/
xN
/
/
/
/
/
/
/
I
I
0
C
0
-0
l .0 Papr 0
0—-- -——- Gemo——-- -l.0
/
0
x
0
/
,
/
/
B
‘,-
,
D
0
/
/
Popr 0
0-- ——-Gema- ——— L0
Relative Seeding Frequency
Replacement diagrams of density (plarits.drn 2 ) (A) and (B) and
shoot hioinass (g-dm 2 ) (C) and (D) for Poa pratensis (Papr, species 1)
and Ceuin macrophyllum (Cema, species 2) at time of harvest when
grown under two levels of UV—B radiation. The dkj2 are 0.72 and
1.32 (no significant difference) for control (A) and enhanced UV—B
conditions (B) and the bki 7 are 0.85 and 2.28 (p<.05) for control
(C) and enhanced UV—B cona3itions (D).
/
Enhanced UV—B
x
/
/
/
/
0
/
x /
/
/
/
/
/
/
/
x
/
/
-------
.c’J
o.ao
(I )
In
0
E
0
0
o
U)
4 -
-c
C)
4 -
0
0
.c
C l)
ti
0
• 0
C
.1/.9 .51.5 .9/.1
Relative Seeding Frequency of
Ceurn mcicrophytlum / Poc pratensis
Control: Enhanced UV—B
Geum macrophyllum
Poa pratensis
Figure 18. A) Shoot biomass (gdm 2 ) and B) individual plant weight (g)
of Poa pratensis and Ceum ma phy11um at time of harvest
after sowing at three relative seeding.frequency classes,
under two levels of UV—B radiation. The means are reported
±1 S.E. (*p(.05).
-------
40
30
20
l0
0
40
30
20
I0
0
Relc iv Se2ciing Fr qu ncy of Piz rri s f!vurn I Alyssum &‘!ssoides
Control: Enhanced 1W—B
[ 1 i Pisuin ________
Figure 19.
yssum alyssoides
( ‘3
E
0
C
I,
a.
U,
C
.1 ,
0
D
z = 0.9 Control
40 2
30- z 2 Q9 WI-B
: -
z 2 -0.5 UV-13
0 10 20 30 40 50
Doys of UV-5 Exposure
Density (plantsdin 2 ) of Alyssum âlyssoides and Piaüm sativutti grown in competition at A)
13 days, B) 37 days and C) 42 days, after sowing at three relative seeding frequency classes,
under two levels of UV—B. radiation. Values are means of 5 replicates ±1 S.E. D) change in
density (plantsdm 2 ) of Pisum sativum through the season for two relative seeding
frequencies (*p<.O 5 ) .
C
I .,
c .
U,
C
C,
0
\
•l/.9 .5/.5 .9/.1
------- |