Quality of UVR Exposure For Different Biological Systems Along A Latitudinal Gradient

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Quality of UVR exposure for different biological systems along a

latitudinal gradient
Maria Vernet,1 Susana Diaz,2,3 Humberto Fuenzalida, 4 Carolina Camilion,2 Charles R. Booth,5 Sergio
Cabrera,6 Claudio Casiccia,7 Guillermo Deferrari,2 Charlotte Lovengreen,8 Alejandro Paladini,3 Jorge
9
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5 Pedroni, Alejandro Rosales, and Horacio Zagarese
Received (in XXX, XXX) Xth XXXXXXXXX 200X, Accepted Xth XXXXXXXXX 200X
First published on the web Xth XXXXXXXXX 200X
DOI: 10.1039/b000000x
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Exposure of organisms to ultraviolet radiation (UVR) is characterized by the climatology (annual cycle) and the variance
(anomalies) of biologically-weighted irradiances at eight geographical locations in austral South America, from 1995-2002. Net
effect of UVR on biological systems is a result of the balance of damage and repair which depends on intensity and duration of
irradiance and is modulated by its variability. The emphasis in this study is on day-to-day variability, a time scale of importance
to adaptive strategies that counteract UVR damage. The irradiances were weighted with DNA- and phytoplankton photosynthesisaction spectra. Low latitude sites show high average UVR. For all sites, the frequency of days with above average irradiances is
higher than below average irradiances. Persistence in anomalies is generally low (< 0.36 autocorrelation coefficient), but higher
for DNA- than phytoplankton photosynthesis-weighted irradiances due to their higher correspondence to stratospheric ozone.
Cloudiness and other factors with small wavelength dependence (i.e., aerosols and albedo) are highly correlated with UVR
anomalies at low latitudes (24-33 S); ozone correlates higher at high latitudes (42-54.5 S). Our results show that organisms in
this region deal with several days of excess radiation and fewer, shorter and more intense periods of lower than average radiation.
Relief from UVR stress (or higher frequency of days below the climatology) is more prevalent at high latitudes (54.5 S). Thus,
lower latitudes are more stressful to organisms not only because of higher average UVR irradiance but also for the higher
frequency of days above the climatology.
Keywords: Ultraviolet radiation, latitudinal gradient, South America, DNA-weighted irradiance, phytoplankton-weighted
irradiance, UVR stress, UVR relief
Introduction
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The rapid ozone depletion in springtime over the Antarctic

continent has created the necessity of keeping track of
ultraviolet radiation (UVR) reaching the Earth surface,
particularly over austral latitudes.1 The southern cone of
South America is one of the few inhabited regions where
increased ultraviolet radiation could have a significant
impact on terrestrial life.2-5 At Ushuaia, Argentina (54.5 S)
a scanning spectroradiometer has been in operation since
1988 as part of the U.S. National Science Foundation UV
Radiation Monitoring Network.6-9 A Brewer spectrometer
belonging to INPE (National Institute for Space Research,
Brazil) is installed at Punta Arenas, Chile.10,11
Meteorological Services in Argentina and Chile also
maintain respective broadband UV networks.4 In this paper
we present results from a network of multi-channel filter
radiometers operating in Chile and Argentina since 1995.12
The time series from this network is still too short to detect
trends in UVR.13 Previous studies have shown the overall
UV irradiance levels in this region,3,4 the effect of latitude on
annual UVR,2 the influence of the Antarctic ozone hole in
springtime at high and mid latitudes2,14 and the relationship
between atmospheric conditions and UVR.14.15
Environmental variability in UVR is dependent on season,
location as well as local atmospheric conditions, aerosols
and gases. 16 Geometric factors (solar zenith angle and the
This journal is The Royal Society of Chemistry [year]
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Earth-Sun distance) are responsible for pronounced and

systematic variations in irradiance.17 Annual cycle in UVR at
different latitudes is mainly determined by these factors, as
well as the seasonal variability in stratospheric ozone.18
Short-term variability in UVR is attributed to atmospheric
conditions.19,20 Clouds attenuate UV irradiance except under
broken cloud conditions where an increase of up to 27%
above clear sky values can be observed.21,22 Spectral changes
in UVR are influenced by atmospheric conditions.23 The
thinning of the ozone layer translates into increases in UVB
(280 to 315 nm) radiation within the wavelength range of
295 to 320 nm, all other atmospheric parameters remaining
constant.6,24 UVA (315 to 400 nm) is more affected by
atmospheric processes other than ozone. In South America,
both UVA and UVB present a gradient with latitude.2,4,14
Furthermore, a higher cross correlation between 305 nm and
340 nm at lower latitudes and higher cross-correlation
between 305 nm irradiance and ozone at higher latitudes
indicated the importance of cloud cover and other factors
with relative small wavelength dependence (i.e., aerosols and
albedo) in explaining UVR variability at lower latitudes (24
S) and the importance of ozone at higher latitudes (54.5
S).14
Although UVB radiation represents less than 0.8% of the
total energy reaching the Earths surface, it is responsible for
almost half the photochemical reactions in aquatic
environments.25 UVB is known to affect a variety of cellular
Journal Name, [year], [vol], 0000 | 1
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processes and molecules in the marine environment and in

terrestrial systems that is manifested as reduced productivity,
habitats shifts or bio-geochemical cycle alterations.25-28 On
the other hand, although UVA may be damaging to cellular
process, such as photosynthesis, 29 it is also involved in repair
mechanisms.30 Thus, the ratio of UVB:UVA, driven by
changes in the atmosphere or by differential attenuation of
UVB and UVA in the water column31 has been identified as
key to our understanding of UV stress in aquatic ecosystems.
The net effect of UVR is a balance between photochemical
damage and biologically-driven processes of recovery and
repair.32 When considering UVR effects on organisms and
biological systems, it is critical to understand how the
environmental conditions affect the rate of UVR damage
(i.e., by changing exposure) as well as the rate of repair (i.e.,
by activating enzymatic processes). In addition to the
intensity and variability of surface radiation, UVR effects on
organisms are dependent on the sensitivity of the living
system to different wavelengths. Normalizing incident
radiation by the corresponding Biological Weighting
Function (BWF) allows for estimation of the potential
damage.33 This approach has shown that the sensitivity of a
living system can change by a factor of more than a 100
between 300 and 400 nm. 34 Variability in sensitivity to a
certain radiation energy among biological systems is also
large. The BWF for erythema changes by three orders of
magnitude between 300 and 340 nm. The BWF expressing
the sensitivity of fish larvae to UV radiation changes only by
1 order over the same wavelength range whereas unprotected
DNA is more than 4 orders of magnitude more sensitive at
300 nm. This high variability in sensitivity to UVR is
combined with variability in intensity and spectral
composition to estimate potential UVR damage to biological
systems.
Plankton organisms are particularly sensitive to UVR
variability due to their floating habitat.35 Mixing of aquatic
organisms from surface to depth (and vice versa) can
augment or reduce net damage by altering the balance in
damage and repair at different depths.36 Antarctic
phytoplankton incubated under sunlight showed a wide range
of inhibition under varying UVB.37 A reduced photosynthetic
performance was evident under full sunlight. They
neutralized net UVR damage when incident radiation was
not very high (i.e., a cloudy day) thus allowing repair
processes to be equivalent to damage; in addition, they
recovered photosynthetic efficiency when a cloudy day
followed a sunny day. Thus day-to-day UVR variability is
important in estimating net damage. In this study we
characterize the inter day changes in UVR for an eight year
series (1995 2002) at eight locations in South America.
The sites encompass a large diversity in latitude and climate
(24 54.5 S). In order to characterize exposure and to
ascertain the main environmental factors affecting it, we
present anomalies with respect to the annual mean. The
source of variability is analyzed with correlations to ozone
and 340 nm irradiance (representing clouds and other
atmospheric factors with relative small wavelength
dependence).38 The day-to-day variability is analyzed with
2 | Journal Name, [year], [vol], 0000
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Markov chains and the persistence of UVR is determined by

autocorrelations. Our results show that organisms in this
region deal with several days of excess radiation and fewer,
Site
Ushuaia
Punta
Arenas
Latitude
54.49 S
Longitude
68.19 W
Altitude
SL
53.09 S
70.55 W
SL
Trelew
43.25 S
65.30 W
SL
Bariloche
41.32 S
71.62 W
~300 m
Valdivia
39.48 S
73.14 W
SL
34.35 S
58.29 W
SL
INGEBI
33.27 S
70.40 W
543 m
24.10 S
65.01 W
1200 m
Univ. de Chile
Univ. Nac. de
Jujuy
Buenos
Aires
Santiago
Jujuy
65
Location
CADIC
Univ. de
Magallanes
Univ. de la
Patagonia
CRUB
Univ. Austral
de Chile
Table 1. The UVR network in South America, with eight multi-channel

GUV-511 radiometers (Biospherical Instruments Inc., USA). CADIC:
Centro Austral de Investigaciones Cientficas; CRUB, Centro Regional
Universitario de Bariloche; INGEBI, Instituto Nacional de Gentica y
Biotecnologa.
shorter and more intense periods of lower than average

radiation. A latitudinal gradient in relief from UVR stress (or
higher frequency of days below the climatology) is observed.
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Materials and Methods

a. UV irradiances
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Irradiances were obtained from measurements provided by

the IAI Radiation Network (IAI RadNet). At present, this
network is composed of eight multi-channel radiometers
(GUV-511, Biospherical Instruments Inc., USA) located in
Chile and Argentina (Table 1). The instruments were
installed during the 1990s by different national and
international efforts and are still in operation. The stations
geographical distribution provides information on UVR at
the regional scale (sub-tropical to sub-Antarctic). Data from
1995 to 2002 are presented in this study.
The GUV-511 is a temperature stabilized multi-channel
radiometer, which measures downwelling irradiances with
moderately narrow bandwidth (near 10 nm) at approximately
305, 320, 340 and 380 nm, plus Photosynthetic Available
Radiation (PAR; 400-700 nm).39 Under a teflon diffuser, the
GUV-511 hosts the five filters and corresponding
photodiodes which are arranged with the 305 nm sensor at
the center and the other four around it. In normal operation
all channels can take a reading every 0.5 seconds and store
an average of 120 readings per minute. One advantage of this
type of instrument is that the four wavelengths in the UVR
are observed simultaneously so fast sky changes can be
tracked.
All instruments collected one-minute average, 24 hours a
day. Raw data were processed applying software provided by
Biospherical Instruments Inc., and modified by the
Laboratory of UV and Ozone, CADIC, Ushuaia. During data
processing, quality and consistency of the data were
checked, night values (instrument internal noise) were
subtracted, and calibration constants were applied. In cases
where the temperature controller failed or the operational
temperature of the instrument differed from the temperature
Ushuaia (6/28/2000 to 12/31/2000)
Erythema
(CIE)
DNA
(Setlow)
Plants
(Caldwell)
Fish
(Hunter)
Phyto
(Neal)
10
15
20
Ch 305 (x1)
Ch 320 (x2)
Ch 340 (x3)
Ch 380 (x4)
90-SZA (x5)
SZA 70
x1
a2e x22 + b2e x2
x3
x4
a5e x52 + b5e x5
SZA >70
ln x1
ln x2
ln x3
ln x4
k5e x53 + l5e x52 + m5e x5 + n5e
SZA 65
a1d x1 + b1d x1
a2d x2 + b2d x2
a3d x3 + b3d x3
a4d x4 + b4d x4
a5d x52 + b5d x5
SZA > 65
ln x1
ln x2
ln x3
ln x4
k5d x53 + l5d x52 + m5d x5 + n5d
SZA 70
a1p x1 + b1p x1
a2p x2 + b2p x2
a3p x3 + b3p x3
a4p x4 + b4p x4
a5p x52 + b5p x5
SZA > 70
ln x1
ln x2
ln x3
ln x4
k5p x53 + l5p x52 + m5p x5 + n5p
SZA 70
x1
a2f x2 + b2f x2
x3
x4
a5f x53 + b5f x52 + c5f x5
SZA > 70
ln x1
ln x2
ln x3
ln x4
k5f x53 + l5f x52 + m5f x5 + n5f
SZA 60
x1
x2
x3
x4
SZA > 60
ln x1
ln x2
at the moment of the calibration, temperature corrections

were applied to the calibration constants.40 Any observed
changes in radiometric data were assumed to be linear.
Therefore, calibration constants applied between two
calibrations dates were calculated by linear interpolation
between the two closest calibrations, unless an abrupt known
change had occurred (i.e. filter replacement).
The GUV-511 were periodically sun calibrated, usually
once a year, with a traveling reference GUV-511 (RGUV).39
The reference radiometer was calibrated, under solar light,
before and after traveling, against the spectroradiometer
SUV-100 of the U.S. National Science Foundation UV
Monitoring Network, installed in San Diego, California,
USA.41 Tests performed during the calibration in year 2000
indicated that the GUVs of the network had an error between
8 and 13%, for the 305 channel and Solar Zenith Angle
(SZA) smaller than 60 degrees with respect to the SUV-100
from San Diego. For the 340 nm channel, the error was
smaller than 5% for all SZA. Details of calibration can be
found in Diaz et al.12,14 The biologically weighted irradiances
presented in this paper were based on these measurements
and calculated using a multi-regression model (see below).
b. Ozone
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Daily total ozone column was obtained from satellites

Nimbus-7, Meteor-3 and Earth Probe, version 8, provided by
NASA Goddard Space Flight Centre.42,43 The time series
from July 1995 to December 2002 was used for this study.
ln x3
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c. Calculation of irradiance weighted by biological action

spectra
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Biologically weighted irradiances are easy to calculate from

spectral measurements, but calculation from multi-channel
radiometers is not direct. Dahlback44 proposes a method to
estimate the biologically weighted irradiances combining the
measurements of four UV channels with a radiative transfer
model. BSI45, 46 and later de La Casinire47 propose a multiregressive equation to derive the biologically weighted doses
from the GUV-511 irradiances, applying a look up table.
Here we use a similar approach but with some modifications.
In our method, we do not use a look up table, but include all
four UV channels in the multi-regressive equation and
determine a function that affects each channel. Also, we add
a term to take into account the influence of SZA. Indeed, we
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ln x4
a5ph x52 + b5ph x5

3
k5ph x5 + l5ph x52 + m5ph x5 + n5ph
Table 2. Function used to fit data from each channel in the GUV-511
and solar zenith angle with the weighted irradiance obtained from the
spectroradiometer. Eryth: erythema; CIE: Commission International de
lEclairage; DNA: Desoxyribo-nucleic acid; Fish: fish larvae; Phyto:
phytoplankton photosynthesis; SZA:solar zenith angle.
use the complement of the SZA in the equation to ensure the

condition that the weighted irradiance tends to zero when
each of the variables tend to zero. It has been observed that,
in calibrating multichannel instruments, ozone and SZA are
critical factors, mainly for the 305 channel44. By introducing
a multi-regressive method in calibrating the radiometers,
improvements were observed, mainly in the UVB channel.12
A similar approach is used here by adding the above
mentioned changes, thus including the effect of ozone and
SZA. Then, the equation we propose for weighted
irradiances is:
Iw = A f(x1) + B f(x2) + C f(x3) + D f(x4) + E f(x5)
where Iw is the weighted irradiance, A, B, C, D and, E are the

regression coefficients determined with least square methods
and f(xi) is a function of the irradiance measured by channel
i, except f(x5), which is a function of (90-SZA).
The function that affects each channel was determined as
the best fit of the scatter plot of the biologically weighted
irradiance obtained from the SUV-100 against the irradiance
measured by the GUV-511 channel under consideration. For
SZA, the scatter plot is done with the difference between the
weighted irradiance and irradiance channel 305, against the
complement of the SZA (90-SZA).
For larger SZA we used the logarithm of the irradiance to
improve the quality of the fitting, increasing the influence of
smaller irradiance values.
ln(Iw) = K f(x1)+L f(x2)+M f(x3)+N f(x4)+O f(x5)+P
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[1]
[2]
where K, L, M, N, O and P are the coefficients of the multiregressive equation determined with least square methods.
In order to develop and test the proposed method we used
simultaneous spectral and multichannel data from Ushuaia
(54o49 S, 68o19 W) and San Diego (32o45 N, 117o11 W).
The selected sites show important differences in the patterns
of ozone and cloud variability. Ushuaia experiences large
daily ozone variations during the spring because of the
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presence of the ozone hole, and is under rather

cloudy skies. San Diego presents a smaller variation in Coef
ozone and smaller SZA in the summer than Ushuaia
A
and preponderance of clear skies, although with more
B
C
influence of aerosols as consequence of its larger
D
population (about 3 millions versus
E
~ 50,000 for Ushuaia). At both sites, the spectral data
K
was obtained with spectroradiometers SUV-100, 48 and
L
M
the multichannel data with a GUV-511 radiometer.
N
Measurements at both sites were performed under all
O
weather conditions, which included different cloud
P
cover, large solar zenith angles, extreme ozone
conditions and various albedo situations.
The model was solved for Ushuaia combining
instantaneous spectral data weighted by a given action
spectrum and multichannel data, for the period July 1st to
December 31st, 2000, for all solar zenith angles smaller than
85 degrees. Once the equations and coefficients were
determined, the model was tested for San Diego, using data
for the period July 9th to December 20th, 1999. Data for the
radiometers GUV are obtained each minute, while the
spectroradiometer SUV-100 performs one scan each 15
minutes. Thus, 4 concurrent values per hour could be
obtained.
We solved the multi-regressive equation for four
biologically weighted irradiances: 1) parameterization of
DNA damage; 49 2) general plant response;50 3) erythema;51
and 4) phytoplankton photosynthesis.34 Table 2 shows the
fitting functions for each channel and for the complement of
SZA, where:
a2e = 0.005287, b2e = 0.39085
a5e = 0.000156, b5e = - 0.030537
k5e = 0.00003715, l5e = 0.00595,
m5e = - 0.34031, n5e = 5.40944
a1d = 0.002697, b1d = 0.057889
a2d = 0.000172, b2d = 0.003987
a3d = 0.00004022, b3d = 0.0019504
a4d = 0.0000224, b4d = 0.001884
a5d = -0.00000789, b5d = 0.000365
k5d = -0.000003612, l5d = 0.000746,
m5d = - 0.0451, n5d = -1.9213509
a1p = -0.0188, b1p = 1.238052
a2p = 0.0027, b2p = 0.081196
a3p = 0.000668, b3p = 0.03827
a4p = 0.0003675, b4p = 0.036437
a5p = -0.00007721, b5p = 0.003598
k5p = 0.000038, l5p= -0.006044,
m5p = 0.34984, n5p = -6.266525
a2f = 0.0027, b2f= 0.081196
a5f = 0.000025, b5f = - 0.00347, b5f = 0.022275
k5f = -0.00003136, l5f = 0.005166 ,
m5f = -0.30324, n5f = 1.5863
a5ph = -0.000868, b5ph= 0.028847
k5ph = 0.0000253, l5ph =-0.00371,
m5ph = 0.1971, n5ph=-3.43053
and where the first subscript corresponds to the term of the
equation and the second to the effect. The regression
coefficients corresponding to equation 1 and 2, which
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Erythema
(CIE)
DNA
(Setlow)
Plants
(Caldwell)
Fish
(Hunter)
2.788104
-0.182006
0.260082
-0.104722
0.072443
0.473879
-0.435139
1.0780513
-0.160834
-0.027122
-0.185785
1.351450
-1.474618
1.674240
-0.547163
0.938882
0.992873
-0.870759
1.054796
-0.236303
-0.129490
-3.497929
1.046271
-0.297090
0.512211
-0.258920
1.000786
0.790850
0.530770
-0.701140
0.335580
0.014770
0.035420
0.133200
-0.021230
0.006944
-0.002540
0.000955
0.507760
0.110230
0.346870
-0.008010
0.021980
-2.929690
Phytoplankton
(Neal)
0.020593
0.027934
0.054107
-0.023480
0.006420
0.053196
0.141762
0.770356
-0.003249
0.032851
-2.794240
Table 3. Coefficients obtained when solving the multi-regressive

equation for calculating weighted irradiances with the GUV-511. For
acronyms see Table 2 legend.
resulted of solving the multi-regressive equation for the four

action spectra are shown in Table 3. The coefficients of
determination (r2) for the regression and RMS errors
between the values of the biological weighted irradiances
calculated with the SUV and the GUV are shown in Table 4,
for hourly averaged values and SZA smaller than 85 degrees.
The multi-regressive model to calculate UV weighted
irradiances based on the multi-channel filter radiometers
presented here has universal application. It was developed
and tested at two sites that present very different atmospheric
and ground characteristics, for a period of near six month
(winter to summer), during which very extreme situations
were present (i.e., ozone hole overpass at Ushuaia, which
produces 60% ozone depletion, large SZA, snow cover in
Ushuaia during winter, etc.), and using two sets of
radiometer and spectro-radiometers, with very good results.
When comparing the results of the model proposed here with
a calculation of weighted irradiances based only on a
coefficient for each channel (instead of a function) and not
including the SZA term, we observed our method to give
better estimates, mainly for SZA larger than 50o, which is
particularly important at higher latitudes (e.g., for DNA the
RMS error diminished by half, for SZA between 50o and 85o,
data not shown). In this paper, we applied the abovementioned methodology to obtain biologically weighted
irradiances from the IAIRadNet.
d. Annual cycle
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To determine climatology we estimated the annual cycle

using a procedure described in Wilks52, which is a more
general version of the Fast Fourier Transform (FFT) spectral
analysis. Wilks approach has the advantage that equidistant
data are not needed because coefficients and phases
corresponding to the harmonic components are obtained via
multivariate analysis.14 Times series of noon values were
selected from the databases of biologically weighted
irradiances. Then, the time series average was calculated for
each Julian day and the annual cycle was inferred thereof.
The annual component of a time series may be represented
by a cosine function, as follows:
2t
[3]
y t = y + C1 cos
1
n
where yt is sampled value on day t; y is mean value of the

annual cycle; t is time; n is number of days in a cycle, in this

case 365; C1 is amplitude of the annual cycle; and 1 is
phase.
Equation [3] may be re-formulated taking into account
that:
[4]
cos( l ) = cos(l ) cos( ) + sin(l ) sin( )
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Given that
A1 = C1 cos(1)
B1 = C1 sin(1 )
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and that
x1 = cos()
x2 = sin()
equation [4] is transformed into a regression with two
predictors
[5]
y t = y + A1 x1 + B1 x 2
Usually, the annual cycle is not a pure cosine. It is rather
the summation of the first (n = 365), the second (n = 180)
and the third (n = 120) harmonics. Taking this into account,
and following the above-explained procedure, a multivariate
equation with six predictors is obtained which, when solved,
provides the annual cycle. The independent term of the
equation corresponds to the mean of the annual cycle or
continuous component in the Fast Fourier Transform. Tests
confirmed that this calculation of the annual cycle
represented well the data (not shown). The same procedure
was used to calculate the annual cycle for ozone.
Since the seasonal variability in UV radiation is very
pronounced, the annual cycle calculated for irradiance would
present large errors in winter, due to the small weight that
winter irradiance presents when solving the multivariate
equation using least squares. In order to avoid this problem,
logarithmic transformation was applied.14
e. Anomalies
35
Anomalies for total column ozone, irradiance at 340 nm and

biologically-weighted irradiance (DNA and
phytoplankton) were calculated as deviations
from the annual cycle as
V VJ
normalized_anomaly = d
[6]
VJ 40
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where Vd is the time series value at date d and

VJ is the annual cycle value at Julian Day J,
corresponding to date d. Gaps in data
collection were filled with the annual cycle.
Usually, the anomaly is normalized by
dividing it by the standard deviation.53 In this
case we preferred to use the value of the
annual cycle for normalization because, as
consequence of ozone depletion, the standard
deviations for DNA- and phytoplankton
photosynthesis weighted-irradiances and total
column ozone are perturbed at higher
latitudes.
Erythema
(CIE)
DNA
(Setlow)
Plants
(Caldwell)
Fish
(Hunter)
Phyto
(Neal)
Atmospheric variables often exhibit statistical dependence
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g. Markov chains - transition probabilities

To evaluate the damage-repair probabilities at each site, we
also estimated persistence in UVR by converting the time
series of anomalies into a Markov chain and calculating
transition probabilities.52,54 In order to perform the analysis,
the values of the anomaly time series were converted into
discrete values, representing a first-order Markov chain. A
Markov chain can be considered as being based on collection
of states of a model system. For each time period, the
Markov chain can either remain in the same state or change
to one of the other states. The behavior of a Markov chain is
governed by a set of probabilities for these transitions called
the transition probabilities. These probabilities specify
likelihoods for the system being in each of the possible states
during the next time period. For first-order Markov chain the
transition probabilities controlling the next state of the
system depend only on the current state of the system: they
are conditional probabilities of the type
ppt
today
Pr
ppt
yesterday 90
[7]
Ushuaia
(7/1/2000 to
12/31/2000)
San Diego
(7/9/1999 to
12/20/1999)
Hourly Average
Hourly Average
Error RMS
(%)
6.18
6.48
0.9973
8.82 (9.82)
6.45
0.9941
13.75
21.34 (35.64)
7.43
11.78
0.9949
16.72 (18.24)
7.48
0.9970
9.48
10.39 (11.05)
5.78
0.9932
8.35
8.27 (9.95)
R2
f. Correlations and autocorrelations

55
65
with their own past or future values. In the terminology of

atmospheric sciences, this dependence through time is
known as persistence. Then, persistence can be defined as
the existence of a positive statistical dependence among
successive values of a variable, or among successive
occurrences of a given event and can be characterized in
terms of serial correlations or temporal autocorrelations.22
We performed persistence studies based on time series of the
normalized anomalies for time lags of one to seven days.
Additionally, cross correlations of the normalized
anomalies of biologically weighted irradiances with the
normalized anomalies of ozone and 340 nm irradiance were
carried out with the aim of determining the influence of
different parameters on the weighted irradiances.
SZA 50
50<SZA<70
70<SZA<80 (85)
SZA 50
50<SZA<70
70<SZA<80 (85)
SZA 50
50<SZA<70
70<SZA<80 (85)
SZA 50
50<SZA<70
70<SZA<80 (85)
SZA 50
50<SZA<70
70<SZA<80 (85)
0.9944
0.9930
0.9953
0.9950
0.9901
Error RMS (%)

5.18
8.17
7.92 (9.66)
8.58
12.65
18.23 (23.38)
5.49
7.49
6.48 (8.27)
5.33
7.19
7.60 (9.07)
6.42
7.76
6.82 (7.49)
R2
Table 4. RMS error and determination coefficients between the hourly

averaged irradiances weighted by different action spectra calculated
from spectral measurements and using multi-channel instrument. Model
developed for Ushuaia and tested for San Diego.
50
55
10
15
20
25
30
35
Figure 1. Biological Weighting Functions (BWF) of key molecules

(Desoxyribo-nucleic acid or DNA), general plant response, human skin
(erythema), organisms (fish larvae) or ecosystem processes
(phytoplankton photosynthesis) showing the relative sensitivity to
ultraviolet radiation (UVR). The values are normalized to 1 at 300 nm.
DNA is the most sensitive to UVB (280 315 nm) with small sensitivity
to UVA (315 - 400 nm). Phytoplankton photosynthesis has the lowest
sensitivity to UVB and highest to UVA such that effective irradiance at
390 nm is only 100 times lower than at 300 nm. In contrast, DNA
effective irradiance at 340 nm is 600 times less effective than at 300 nm.
Other molecules or biological processes have intermediate sensitivity.
Analyses in this paper correspond to UVR weighted by the BWF
(calculated as Radiation * BWF) from DNA and phytoplankton
photosynthesis representing the two extremes in UVA and UVB
sensitivity.
where the probability Pr that a value be present today

(ppttoday) given a determined value yesterday (pptyesterday).
The simplest kind of discrete random variable pertains to the
situation of dichotomous (yes/no) events. In our case, we
will consider a two state first-order Markov chain.
The system will be in state 1 (event 1), if the anomaly is
bigger than zero (irradiance larger than climatology), and in
state 0 (event 0), if the anomaly is smaller or equal to zero
(irradiance smaller or equal to climatology). Three states
could have been considered: larger than 0, equal to 0 and
smaller than 0, but, in this analysis no anomaly is equal to 0,
then, the value 0 was included in state 0. Taken this into
account, the probability that a day with irradiances smaller
than, or equal to, the annual cycle (anomaly < 0) be followed
by a day with irradiance smaller than, or equal to, the annual
cycle is P00; and if followed by a day with irradiance higher
than annual cycle is P01. In the same way, the probability that
a day with irradiance larger than the annual cycle (anomaly >
0) be followed by a day with irradiance larger than the
annual cycle is P11; and if followed by a day with irradiance
smaller than or equal to the annual cycle is P10.
60
Site
Jujuy
Buenos
Aires
Trelew
Ushuaia
Santiago
Valdivia
Punta
Arenas
65
40
45
Climate is a key issue to understand the short-term and

seasonal variability in UVR. We summarize here the main
characteristics at each location. The study sites in South
America, between the Capricorn Tropic (24 S) and subAntarctic conditions (54.5 S) experience very different
climate conditions (Table 5). Jujuy, on the eastern slope of
the Andes, receives most of its rainfall during the summer
months by convective activity. Santiago and Valdivia belong
to the Mediterranean type of climate with a dry summer and
most rain falling in winter associated with frontal passages
Annual
rain
(mm)
615
Minimum
(mm) month
Maximum
(mm) month
Cloud
cover
(tenths)
n.a.
0 (July)
183 (January)
1010
59 (July)
118 (March)
4 even
156
549
339
2446
8 (August)
35 (October)
1 (January)
66 (February)
21 (May)
55 (March)
80 (June)
396 (June)
5 even
6 even
2 to 6
4 to 7
403
24 (October)
43 (May)
6 to 7
Table 5. Precipitation and cloud cover in the localities where GUV

radiometers are deployed.
b. UVR variability under clear sky conditions
70
75
80
Results
a. South American climate
that become more frequent the higher the latitude. The main
difference between these two sites is the annual
precipitation, almost an order of magnitude larger in
Valdivia, so that even in the driest month 66 mm are
received. Punta Arenas is part of a cold steppe that develops
on the eastern side of the Andes with a fairly scanty and
homogeneous precipitation regime throughout the year.
Ushuaia, although located inside the Andes shares this
condition. Trelew aridity is due to the rain shadow on the lee
(eastern) side of the Andes receiving its annual 156 mm in a
fairly well distributed manner through the year. Bariloche is
located at the foothills of the Andes on the lee side. Buenos
Aires has a fairly moist and mild climate with no major
rainfall season.
85
90
95
Organisms at any given location and living for at least one

year are exposed to a range of variability in effective UVR
of several orders of magnitude. Using clear sky weighted
irradiance at Ushuaia, 54.49 S, several characteristics in the
spectral UVR are of note. As expected, the difference in the
overall sensitivity in UVB vs. UVA of the different weighted
irradiances is similar to the shape of the Biological
Weighting Function (BWF, Fig. 1) given that incident
radiation is the same for all. In winter, the maximum
irradiance or the wavelength with highest effective
irradiance is 312 nm for DNA-weighted irradiances, 315 nm
for fish larvae, general plant-weighted irradiances, and 330
nm for phytoplankton photosynthesis (15 June; Fig. 2a).
Phytoplankton photosynthesis-weighted irradiances are an
order of magnitude higher than DNA-weighted irradiances in
the UVB and four orders of magnitude in the UVA (330 nm).
Although the relative sensitivity between UVB and UVA in
the different biological systems is maintained throughout the
seasons, there is a shift to lower maximum effective
wavelength in the summer. At this time, maximum
wavelength of DNA-weighted irradiance is 303 nm, 308 nm
for erythema- and general plant-weighted irradiances, and
315 nm for phytoplankton photosynthesis-weighted
irradiances (Fig. 2b). Also as expected, higher irradiances
are experienced in the summer,6-9 with an increase of two
orders of magnitude for DNA-weighted irradiances at its
maximum wavelength of 303 nm and an increase of one
order of magnitude for phytoplankton photosynthesisweighted irradiances at 315 nm (Fig. 2b). The other
35
exposure to organisms living in the southern cone of South

America. These two BWFs represent extremes in UVB:UVA
sensitivity, DNA mostly sensitive to UVB changes and
phytoplankton photosynthesis very sensitive to UVA.
c. UVR annual cycle
40
45
50
55
10
15
20
25
30
Figure 2. Spectral Irradiance at different seasons and for varying

stratospheric ozone concentrations. Clear sky spectral UVR between
295 and 390 nm weighted by BWF of different systems (DNA, open
squares; General Plant response, triangles; Fish Larvae survival, grey
squares; Erythema or skin sensitivity, black squares; Phytoplankton
Photosynthesis, diamond at a) austral winter; b) austral summer and c)
austral summer under 60% ozone reduction. All plots for Ushuaia,
Argentina at 54o49S, 68o19W. SZA depicts Solar Zenith Angle.
maximum sensitivity, 300 nm for DNA-weighted irradiances,

305 nm for erythema- and general plant-weighted
irradiances, and 310 nm for phytoplankton photosynthesis.
There is also an increase of almost one order of magnitude
for DNA effective irradiance with smaller increases for
biological systems experience intermediate increase in
effective irradiances. Finally, the effect of ozone is
experienced at the lowest UVB wavelengths, near 300 nm,
with no changes in the effective UVA radiation (Fig. 2c).
Under conditions similar to the ozone hole (60% ozone
reduction), there is a further decrease in the wavelength of
erythema-, general plant-, or phytoplankton photosynthesisweighted irradiances.
In summary, the sensitivity of the different biological
systems (Fig. 1) affects wavelength and intensity of exposure
as well as the balance between UVB and UVA (Fig. 2).
Seasonal changes in UVR affect overall intensity as well as
the wavelength of maximum effect, with lower maximum
effective wavelengths in the summer. Finally, ozone
influences mostly DNA effective irradiance. Lower ozone
decreases wavelength of maximum sensitivity and increases
irradiance, similar to the effect of smaller SZA (summer).
Based on these results we choose DNA- and phytoplankton
photosynthesis-weighted irradiances to characterize the UVR
60
65
70
75
80
85
The UVR climatology at each site is defined as the annual

cycle of the biologically weighted irradiances (smooth line in
Fig. 3). As expected, higher average DNA-weighted
irradiances are observed at lower latitudes year round.2 From
those, arid areas at mid latitudes (Santiago de Chile, Fig. 3c)
have higher average summer irradiances than wet forested
locations (Jujuy, Fig. 3a) in spite of the latter being further
north and experiencing higher UVR irradiance under clear
skies (smaller SZA) (Table 6). A large difference between
summer and winter is evident at all locations. This seasonal
contrast is more pronounced at higher latitudes (Table 6,
compare Jujuy with Ushuaia); furthermore, low latitude sites
present measurable UVR in the winter while no UVR is
detected south of 42 S (i.e., Trelew, Fig. 3f).
Daily changes in irradiance can be observed by comparing
DNA-weighted irradiances (thin line in Fig. 3) with the
annual cycle. Several patterns in irradiance variability are
apparent. Most sites show lower than average irradiances
(below the annual cycle) that are larger in magnitude and
shorter in duration than above average irradiances; in
contrast, above average irradiances usually last longer
(several days) and have moderate magnitude (Fig. 3b, 3e,
3f). The observed variability pattern, characteristic of the 8year time series, is attributed to the fact that the 365-day
annual cycle is based on average values. The bias towards
higher than average irradiances means that the integral of
above average irradiances is similar to the integral of below
average irradiances, which is sensitive not only to the
duration but also to the magnitude of irradiance events (see
also Anomalies section).
The presence of anomalous periods is also evident; for
example in Bariloche (Fig. 3e), consistent below average
irradiances lasted for several weeks in the springtime of
1998. Similarly, long sequences of days with above average
irradiances can occur at several locations, as seen in
Bariloche in January and February.
It is also of note a large variability in irradiance in Jujuy,
Buenos Aires, Valdivia and Trelew (Figs. 3a, 3b, 3d and 3f)
where days of very high (maximum) irradiance are followed
by days of very low irradiance; the resulting pattern is one of
large
oscillations.
For
example,
the
ratio
of
maximum/minimum irradiance in summer is 1238 for
Buenos Aires, in comparison to a minimum of 71 in Santiago
de Chile (Table 6). The maximum ratio is seen in Ushuaia
(1661). Such variability could be attributed to the high
irradiances found in summer during clear days combined
with characteristic thick cloudiness in overcast days.
Finally, high short peaks in irradiance during springtime,
previously attributed to the ozone hole, can be observed as
far north as Valdivia and Bariloche (39.5 S; Figs. 3d,
3e).2,14
a) DNA
Site
Jujuy
Buenos Aires
Santiago
Valdivia
Bariloche
Trelew
Punta Arenas
Ushuaia
b) Phytoplankton
Site
Jujuy
Buenos Aires
Santiago
Valdivia
Bariloche
Trelew
Punta Arenas
Ushuaia
10
15
20
25
30
35
40
Max
0.55
0.48
0.63
0.50
0.56
0.43
0.21
0.21
Annual Cycle
Min
0.102
0.034
0.033
0.016
0.013
0.017
0.003
0.002
Max/Min
5.41
14.04
18.68
30.14
40.56
25.40
73.00
90.34
Summer Irradiance
Max
Min
1.26
0.010
0.99
0.000
0.97
0.066
1.01
0.011
1.06
0.027
0.94
0.003
0.70
0.016
0.72
0.013
Winter Irradiance
Max
Min
0.38
0.0043
0.20
0.0007
0.15
0.0020
0.11
0.0006
0.92
0.0015
0.11
0.0017
0.02
0.0006
0.02
0.0001
Max
2.22
2.17
2.65
2.61
2.85
2.24
1.59
1.70
Annual Cycle
Min
0.96
0.59
0.56
0.44
0.47
0.54
0.21
0.20
Max/Min
2.30
3.68
4.65
5.93
5.94
4.13
7.43
8.33
Summer Irradiance
Max
Min
3.55
0.196
3.27
0.155
3.27
0.533
3.73
0.227
3.77
0.504
3.11
0.214
2.96
0.280
2.96
0.200
Winter Irradiance
Max
Min
2.21
0.076
1.39
0.043
1.39
0.030
1.29
0.047
1.33
0.056
1.10
0.066
0.66
0.055
0.63
0.033
Table 6. Irradiances in South America, weighted by DNA and

Phytoplankton photosynthesis action spectra, in units of W cm-2.
Maxima (Max) in the Annual Cycle are observed between 21 December
and 13 January; minima (Min) between 22 and 30 June. Annual maxima
irradiances are measured between 2 December and 17 January, with the
exception of Jujuy where there is an early maximum in 6 November
1997; the minima in irradiance are between 7 June and 7 July with the
exception of Jujuy where there is a late minimum in 29 September 1998.
45
Phytoplankton photosynthesis-weighted irradiances at the

eight localities show very similar annual cycles and 50
variability than for DNA-weighted irradiances (Fig. 4). The
resulting regional pattern is one where summer irradiances
are highest at mid latitude locations (Figs. 4c, 4d, 4e),
intermediate at low or mid latitudes (Figs. 4a, 4b and 4f) and
lowest at higher latitudes (Figs. 4g and 4h). The
55
characteristic feature of several days with higher than
average irradiances interspersed with few days of very low
irradiance is also present here and is even more
a) DNA
pronounced for the phytoplankton- than for
DNA-weighted irradiances, in particular at mid
Site
latitudes (Figs. 4b, 4d, 4e and 4f).
Jujuy
Several features are characteristic of this
Buenos Aires
irradiance which is more heavily weighted by
Santiago
Valdivia
UVA (compare with Figs. 1 and 2). In general,
Bariloche
the
magnitude
of
the
phytoplankton
Trelew
photosynthesis weighted irradiances is three to
Punta Arenas
four times higher than DNA-weighted
Ushuaia
b) Phytoirradiances. This change is most obvious in the
plankton
winter when higher-latitude sites show
measurable
phytoplankton-weighted
UVR
Site
(Figs. 4e, 4f, 4g and 4h) but not DNA-weighted
Jujuy
irradiances (Figs. 3e, 3f, 3g and 3h). The
Buenos Aires
Santiago
overall effect is one of reduced seasonality (see
Valdivia
Annual Cycle Max/Min in Table 6).
Bariloche
Another major difference is the lower
Trelew
latitudinal gradient in the annual cycle
Punta Arenas
Ushuaia
observed
in
phytoplankton-weighted
irradiances. While for irradiances weighted with the DNA
action spectrum the average summer irradiance between
lowest and highest latitudes has a factor of ~2.5 (0.55 W
cm-2 in Jujuy and 0.21 W cm-2 in Ushuaia, Figs. 3a and 3h; 60
Table 6), it is ~1 for irradiances weighted with the

phytoplankton photosynthesis action spectrum (2.2 W cm-2
in Jujuy and 1.7 W cm-2 in Ushuaia, Figs. 4a and 4h). In
addition, irradiance variability is lower (oscillations of dayto-day irradiance are less pronounced) for phytoplankton
photosytnthesis-weighted irradiance than for DNA-weighted
irradiances (Table 6). A similar latitudinal gradient is
observed when comparing 305 nm and 340 nm irradiances.14
d. UVR anomalies
Anomalies can further describe the degree of day-to-day
variability of weighted irradiances (Figs. 5 and 6). Only one
year is shown in the figures to allow enough detail in the
graph to understand the differences among sites while
subsequent statistical analyses are based on the anomalies of
the entire time series (1995-2002). In contrast to deviations
Mean +/- Standard Deviation
Positive
Negative
Anomalies
Anomalies
0.155 +/- 0.137
-0.138 +/- 0.115
0.098 +/- 0.094
-0.087 +/- 0.100
0.059 +/- 0.055
-0.061 +/- 0.086
0.072 +/- 0.076
-0.061 +/- 0.083
0.070 +/- 0.078
-0.059 +/- 0.081
0.063 +/- 0.069
-0.057 +/- 0.077
0.043 +/- 0.065
-0.027 +/- 0.034
0.043 +/- 0.063
-0.028 +/- 0.034
Percentage of Total
Positive
Negative
Anomalies
Anomalies
66%
34%
68%
32%
65%
35%
63%
37%
61%
39%
64%
36%
55%
45%
55%
45%
Mean +/- Dev
Percentage of Total
Positive
Anomalies
0.594 +/- 0.477
0.466 +/- 0.258
0.284 +/- 0.187
0.474 +/- 0.260
0.363 +/- 0.219
0.294 +/- 0.156
0.327 +/- 0.292
0.300 +/- 0.293
Negative
Anomalies
-0.661 +/- 0.423
-0.531 +/- 0.421
-0.377 +/- 0.375
-0.479 +/- 0.428
-0.488 +/- 0.439
-0.435 +/- 0.392
-0.291 +/- 0.262
-0.279 +/- 0.261
Positive
Anomalies
66%
69%
70%
65%
70%
71%
60%
60%
Negative
Anomalies
34%
31%
30%
35%
30%
29%
40%
40%
Table 7. Statistics of the positive and negative UVR anomalies (19952002), a) for DNA- and b) for phytoplankton photosynthesis-weighted
irradiances.
10
15
20
25
30
35
40
45
Figure 3. Seasonal variability in UVR-weighted irradiances in South

America Daily noon DNA-weighted UVR (295 - 390 nm) at different
localities from 28 to 55 S, in units of W cm-2, from 1 July 1998 to 30
June 1999. The thin straight line depicts the climatology or annual
cycle. Jujuy 24.10o S, 65.01o W; Buenos Aires 34.35o S, 58.29o W;
Santiago 33.27o S, 70.40o W; Valdivia 39.48o S, 73.1o 4 W; Bariloche
41.32o S, 71.62o W; Trelew 43.35 o S, 65.05o W; Punta Arenas 53.09o S,
70.55o W; Ushuaia 54.49o S, 68.19o W.
of weighted irradiances to the annual cycle, the overall

magnitude of the anomalies of DNA- and phytoplankton
photosynthesis weighted irradiances is more similar, with the
exception of high-latitude sites (Figs. 5g and 5h). The
sequence of positive and negative anomalies in time follows
the pattern already described for irradiances (Figs. 3 and 4)
with several days of positive anomalies interspersed with
short, pronounced negative anomaly periods (see Figs. 5e
and 5f). On average for the 8-year time series, ~60% of the
anomalies are positive and 40% are negative (Table 7). This
is not an artifact of the annual cycle calculation as the
average and standard deviation of the positive and negative
anomalies is similar at each locality, with the exception of
Punta Arenas and Ushuaia where the positive anomalies for
DNA-weighted irradiances present higher values and
variability, as might be expected by the increased UVB
during periods of influence of the Antarctic ozone hole.6,7
Statistics for the period 1998-1999 indicate this is a
representative year (data not shown).
The presence of extended periods with positive or negative
anomalies, mentioned already for the irradiances (Figs. 3 and
4), are seen for most localities, with the exception of Buenos
Aires. These periods break up the temporal development of
positive and negative anomalies described above, can last
from 6 to 100 days and had an average extension of 20 + 9
days for positive anomalies and 30 + 15 days for negative
Site
Jujuy
Buenos Aires
Santiago
Valdivia
Bariloche
Trelew
Punta Arenas
Ushuaia
P00
0.52
0.42
0.51
0.45
0.38
0.37
0.43
0.42
Site
Jujuy
Buenos Aires
Santiago
Valdivia
Bariloche
Trelew
Punta Arenas
Ushuaia
P00
0.51
0.44
0.51
0.48
0.56
0.50
0.58
0.56
P01
0.48
0.58
0.49
0.55
0.62
0.63
0.57
0.58
a) DNA
P11
0.74
0.74
0.78
0.70
0.74
0.75
0.63
0.62
P10
0.26
0.26
0.22
0.30
0.26
0.25
0.37
0.38
P01
P11
P10
0.49
0.74
0.26
0.56
0.73
0.27
0.49
0.74
0.26
0.52
0.70
0.30
0.44
0.73
0.27
0.50
0.71
0.29
0.42
0.66
0.34
0.44
0.64
0.36
b) Phytoplankton
Table 8. UVR Anomaly Transition Probabilities a) for DNAweighted irradiance anomalies and b) phytoplankton photosynthesisweighted irradiance anomalies. The system will be in state 1 (event 1), if
the anomaly is bigger than zero (irradiance larger than the climatology),
and in state 0 (event 0), if the anomaly is smaller or equal to zero
(irradiance smaller or equal to the climatology). All transition
probabilities are significant at = 0.05.
10
15
20
25
Figure 4. Seasonal variability in UVR-weighted irradiances in South

America Daily noon phytoplankton photosynthesis-weighted UVR
(295 - 390 nm) at different localities from 28 to 55 S from 1 July 1998
to 30 June 1999. The thin straight line depicts the climatology or annual
cycle. Latitudes and Longitudes of each location can be found in Table 1
and Figure 3 legend.
anomalies during 1998-1999. For example, at Bariloche,

periods of positive anomalies were observed in the summer
between 12 January and 13 February 1999 and also in the
fall from 19 April to 15 May. Negative anomalies were
almost continuous in the spring, from 22 October to 3
December 1998 (Figs. 3e and 5e).
The magnitude and characteristic of the anomalies at each
site present interesting geographical differences. Most
noticeable, some locations have pronounced anomalies (i.e.,
Figs. 5b, 5g and 5h) while others are less variable (Fig. 5c).
Several mid-latitude locations present a seasonal cycle in the
phytoplankton-weighted anomalies: Buenos Aires, Santiago,
Valdivia and Bariloche show lower positive anomalies in
summer (Figs. 6b, 6c, 6d and 6e). There is also a difference
among locations with respect to negative anomalies:
Santiago de Chile and Bariloche often show a predominance
of several days of positive anomalies (Figs. 5c, 6c and 5e,
6e) whereas Jujuy, Valdivia and Ushuaia (Figs. 5a and 6a, 5d
and 6d, 5h and 6h) present more frequent negative
anomalies.
e. UVR anomaly autocorrelations
30
35
40
45
50
55
In order to characterize further the day-to-day variability in

weighted irradiance we used autocorrelation to estimate the
similarity in anomaly values in a seven day period (lag = 1 to
7). Autocorrelations of DNA-weighted anomalies show low
and consistent values (between 0.26 and 0.36, Fig. 7) at all
sites in consecutive days (lag = 1). Autocorrelations suggest
predictability in consecutive days is about 25-35%. For lags
larger than one, the values of the autocorrelations diminish,
more rapidly at mid-latitude (i.e., Buenos Aires, Fig. 7b)
than at higher-latitude sites (i.e., Ushuaia, Fig. 7h).
Autocorrelations of phytoplankton photosynthesis weighted
anomalies show a marked difference among sites, varying
between 0.04 and 0.27 (lag = 1); in general, there is a lower
autocorrelation than for DNA-weighted anomalies, in
agreement with the dependence of 305 nm irradiance on
ozone.14
The slope of the autocorrelation function can indicate the
degree of persistence in the system. In general the
persistence is low and lowest for phytoplankton
photosynthesis weighted anomalies (the latter presents a
steeper slope). At lower latitudes, the values for
phytoplankton- and DNA-weighted irradiance differ less
than for higher-latitude sites (the slope of the lines overlap,
i.e., Buenos Aires). As only DNA-weighted irradiances are
highly sensitive to stratospheric ozone concentration (as
shown in Fig. 2c), the higher similarity in persistence
between both weighted irradiances at lower-latitude sites can
be attributed to a higher influence of atmospheric conditions
on DNA-weighted irradiances. In contrast, only ozone
concentrations seem to have a higher influence on DNA-
Figure 5. Anomalies of DNA-weighted irradiances Normalized

anomalies for the eight studied sites in South America, from 1 July 1998
to 30 June 1999. Latitudes and Longitudes of each location can be found
in Table 1 and Figure 3 legend.
5
10
weighted irradiances further south and here the persistence

diverges.
In summary, a low persistence in UVR is characteristic of
this region, as implied by the see-saw pattern of irradiance
about the mean observed in Figs. 3 and 4. At higher
latitudes, the influence of ozone is associated with higher
persistence14 and as expected, DNA-weighted irradiances are
more sensitive to this influence.
f. UVR anomaly transition probabilities
15
20
25
30
35
40
45
50
55
Another way to characterize the irradiance anomalies is by

estimating the probability of finding a certain UVR anomaly
given an anomaly on the previous day, or transition
probabilities (Table 8). In general, both action spectra
weighted irradiances show similar transition probabilities at
all locations. The value of the probabilities indicates a very
interesting irradiance pattern for the region. P11 (the
probability of having two consecutive days with irradiance
larger than the annual cycle) is largest everywhere (~0.7). In
addition, P10 (the probability of having a day with irradiance
higher than the annual cycle followed by a day with
irradiance lower than the annual cycle) is lowest everywhere
(~0.3). These probabilities support the observation made
before that several days of higher than average UVR are
followed by shorter periods of lower than average irradiance
(Figs. 3 to 6 and Table 7). A latitudinal gradient is also
observed in these two transition probabilities, with P11 lower
and P10 higher at higher-latitude sites, implying a larger
probability of a day with high irradiance being followed by a
day of low irradiance towards the south (e.g., higher
frequency of cloudy or low ozone days).
On the other hand, P00 (the probability of finding two
consecutive days with irradiances equal or below the annual
cycle) and P01 (the probability of a day with irradiance equal
to or lower than the annual cycle followed by a day with
irradiance larger than the annual cycle) show similar values,
for both biologically-weighted irradiances at all locations
(~0.5). Thus, the chance of switching from low to high UVR
in two consecutive days is about the same as staying low.
Some latitudinal differences are observed between the action
spectra. At lower-latitude sites, P01 is higher for the DNAand lower for the phytoplankton-weighted irradiances; at
lower latitudes there is a higher probability of having a day
with above average UVR after a day with low UVR. The
opposite is observed for P00 where larger probabilities for
DNA-weighted irradiances are observed at lower latitudes,
i.e., two consecutive low-irradiance days are more common
at higher latitudes.
In summary for this region, 70% of the time a day with
high UVR is followed by another day with high UVR. And
this pattern is more prevalent at low latitudes. If a day has
low UVR, there is a 50% chance of having low UVR during
the next day. The chance of a second low UVR day is higher
at higher latitudes. Thus, we suggest that DNA would have
Figure 6. Anomalies of Phytoplankton Photosynthesis-weighted

irradiances - Normalized anomalies for the eight studied sites in South
America, from 1 July 1998 to 30 June 1999. Latitudes and Longitudes of
each location can be found in Table 1 and Figure 3 legend.
5
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25
30
35
40
more probability of repair at higher- than at lower-latitude

sites (P00 and P10 larger). Higher latitudes provide also more
relief from UVR stress for phytoplankton photosynthesis (P10
is highest at high latitudes) but the period of recovery is
shorter than for DNA (P00 is lower at higher latitudes).
g. Importance of Atmospheric Conditions and Ozone on
Irradiance Anomalies
Cross-correlation
of
biological-weighted
irradiance
anomalies can be used to ascertain importance of
environmental variables (Table 9). Influence of cloudiness
and other atmospheric processes (albedo, aerosols, etc.) on
UVR variability was estimated with 340 nm irradiances.20
The effect of stratospheric ozone on UVR was estimated
from ozone column height in Dobson units. In general,
weighted irradiances have a high correlation with 340 nm,
higher for phytoplankton (r = ~1.0) than for DNA (r = ~0.8)
and with decreasing influence with latitude for DNAweighted irradiance anomalies but high everywhere for
phytoplankton-weighted irradiance anomalies. In contrast,
influence of ozone is present everywhere for DNA (r = ~0.4),
and increases with latitude (r = 0.60), but not for
phytoplankton (r = ~0.01), as expected from the high UVA
sensitivity of phytoplankton seen in Fig. 2. Thus,` cloudiness
and other relatively wavelength independent factors are a
better predictor of DNA damage by UVR at lower latitudes
(24 S) and both ozone and 340-nm play an important role at
higher latitudes (54.5 S).14 Cloudiness is the only predictor
for phytoplankton photosynthesis UVR damage at all
latitudes.
There is a small but significant cross correlation between
340 nm irradiance and stratospheric ozone column (Table
10). The low correlation implies little overlap between these
two phenomena although it is known that 340 nm irradiance
and ozone column respond to atmospheric pressure (cyclonic
and anticyclonic systems).55 During a high-pressure system
(anticyclonic) there is good weather and clear skies resulting
in high irradiance, including high 340 nm irradiance. At the
same time, a high pressure system causes a decrease in ozone
Site
Jujuy
Buenos Aires
Santiago
Valdivia
Bariloche
Trelew
Punta Arenas
Ushuaia
45
DNA
340 nm
0.94
0.90
0.87
0.87
0.80
0.61
0.70
0.69
DNA
Ozone
0.24
0.33
0.28
0.37
0.46
0.47
0.48
0.60
Phytoplankton
340 nm
0.99
1.00
0.99
1.00
1.00
0.99
0.99
0.99
Phytoplankton
Ozone
0.06
ns
0.07
0.09
0.09
ns
0.05
0.05
Table 9. Cross-correlation coefficients (r) of biological-weighted

irradiances to ascertain importance of environmental variables, for lag =
0. Irradiance anomalies at 340 nm are an index of cloudiness (and
other wavelength independent atmospheric factors). Ozone column
anomalies were estimated from TOMS NASA satellite. All values are
statistically significant for = 0.01 unless otherwise indicated. (ns) not
statistically significant.
50
40
45
50
55
10
Figure 7. UVR Anomaly Autocorrelations Autocorrelations at lag 1

(one day) to 7 (one week) for DNA- (closed circle) and phytoplankton
photosynthesis-(open triangle) weighted irradiances. All values
statistically significant for = 0.01, except for phytoplankton
photosynthesis at Jujuy lag 4, which is not statistically significant.
Values not included in the graphs are not statistically significant.
column56-58 and the opposite is true during a low pressure

system (cyclonic). Thus, although there is no cause and
effect between ozone column and 340 nm irradiance, there
can be a correlation as both variables are affected by
pressure systems. Our results suggest the correlation between
these atmospheric processes is found in austral South
America but it is not a dominant feature.
Discussion
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35
This study aimed at increasing our understanding of the

UVR conditions to which biological systems are exposed to
in order to better interpret existing data, extend our
knowledge of the factors important to the organisms and to
evaluate our present experimental design based largely on
short time exposures.59 An increased number of experiments
have addressed the importance of long-term UVR effects
(e.g., 48 h incubations37 or microcosms60). A better
understanding of the UVR conditions organisms experience
will help formulate hypotheses on the ability of biological
systems to handle UVR as well as what do organisms need in
order to live in a certain UVR environment. Our analysis has
emphasized UVR weighted by DNA- and phytoplankton
photosynthesis- biological weighting functions (BWFs).
DNA damage affects all living organisms; the BWF is of
widespread application allowing for comparisons among
studies.61,62 Phytoplankton photosynthesis represents an
important process in the Earths carbon cycle as they support
50% of the primary production on the planet.35 Thus, our
conclusions relate to representative biological systems.
It is known that the average UVR is higher at lower
latitudes (due to smaller SZA), both in summer and
winter.2,63 It has been shown also the importance of ozone in
60
65
southern localities and of tropospheric conditions (i.e.,

cloudiness) in mid- and lower latitudes due to the sensitivity
of DNA-weighted irradiances to UVB.14 What is new and
has been stressed in this paper is the importance of day-today variability in UVR, how this high-frequency variability
changes for UVB and UVA sensitive systems and how this
variability relates to atmospheric conditions. In addition,
phytoplankton effective UVR on a regional scale is
presented for the first time.
Several important findings will be considered here:
a. Phytoplankton (microalgae) need to deal with average
UVB and its temporal variability to protect and repair
DNA as well as with average UVA and its variability in
order to protect and maximize photosynthesis.
b. Large inter-site variability observed in this study, of
importance to regional or continental studies of UVR
effects, argues for independent studies at different
localities and against generalizations from one area to
another one, even at the same latitude.
c. An UVR environment with more frequent positive
anomalies (Figs. 5 and 6, Table 7) implies that the
organisms seem to live under chronic UVR stress.
d. High-latitude areas seem to provide phytoplankton cells
with more relief from UVR stress (Table 8), providing
more chances to repair DNA and recover photosynthesis
performance.
e. The effect of atmospheric conditions on UVA and UVAsensitive organisms and biological processes is of the
utmost importance, explaining most of the variability in
potential UVR damage to phytoplankton photosynthesis
(Table 9).
a. UVR variability in austral South America
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As shown in this study, organisms in austral South America

have to deal not only with average UVR but also with
different patterns of variability in different areas that will
affect direct and indirect effects, possibly the direction (sign)
of the UVR effect and the process of acclimation. For
example, the degree of day-to-day variability in DNAweighted irradiance observed in Jujuy (Figs. 3a) is as large
as the average irradiance (range of irradiance in January
from 0.1 to 1.0 W cm-2 with an average irradiance of 0.55
W cm-2), and larger than the difference in average
irradiance between summer and winter (0.55 to 0.10 W
Site
Jujuy
Buenos Aires
Santiago de Chile
Valdivia
Bariloche
Trelew
Punta Arenas
Ushuaia
85
Cross 340 nm Ozone

0.06
ns
0.09
0.06
0.06
0.06
0.10
ns
Table 10. Cross correlation of band 340 nm anomalies versus ozone

anomalies, for lag = 0. Lag of -1, +1 and +2 increased somewhat the
correlation coefficients (to a maximum of 0.13) suggesting some out of
phase response of each parameter to pressure systems. All values are
statistically significant for = 0.01 unless otherwise indicated. (ns) not
statistically significant.
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40
45
50
55
cm-2). Furthermore, the high variability in UVR anomalies

among sites, due mainly to the differences in atmospheric
conditions (Table 5) stresses the importance of taking into
account the general climate of the site when evaluating UVR
stress.
The extent to which UVR variability influences net
damage to biological systems has not been studied. A lack of
UVR characterization at different locations is in part
responsible, although relevant data from existing UVR
networks are becoming available.14,35,63 More studies based
on multi-day and multi-week experiments are needed.
Furthermore, an experimental design separating the effect of
average incident UVR to its variability will shed light on the
ultimate importance of considering UVR variability. The
effect of vertical mixing in modulating UVR damage in
phytoplankton is a good example of the importance of
considering variability in exposure to assess net UVR
damage.36 In general, studies are based on single biological
systems, as in the case of the BWFs used here. The system is
kept simple and the added complexity originates from the
light field. A different level of complexity is present when
experimenting on communities, as in the case of mesocosm
studies.64 It is difficult to predict a priori what short-term
surface UVR variability can add to these studies or even how
to separate the effect of average and day-to-day UVR. We
can speculate that the differential response of the various
components in the ecosystem could dampen the overall
effect of individual responses, in a complex matrix of
positive and negative feedback loops. Nevertheless, we can
argue that an organism living at high latitudes, with a higher
preponderance of lower than average irradiance days than
those living in sub-tropical regions, might need a different
rate of photosynthesis or DNA repair than those exposed to
extended periods of above average irradiance. As plants in
Ushuaia and the Antarctic Peninsula have a slower rate of
DNA repair28 which combined with low ambient
temperatures can increase a species sensitivity to UVR,65
any relief from UVR exposure, as shown in this study, might
be critical to their survival.
The observed difference in the effect of UVR variability
on the various biological systems argues also for a more
integral study of the response of different cellular component
or processes. In the case of phytoplankton, cellular DNA and
photosynthesis will be affected differently by UVB and
UVA. We need to consider both DNA- and photosynthesisweighted irradiances when studying overall UVR effects on
ecosystems as these two irradiances respond differently to
climate, ozone concentration, effect of latitude, and day-today changes in atmospheric conditions.66,67 Including this
knowledge can shed light on the balance of UVR damage to
different cellular processes at different latitudes. For
example, phytoplankton from high latitudes will be more or
less affected by UVR than cells from low latitudes
depending on what process is dominant, DNA damage or
photosynthetic inhibition. For any given experiment, higher
than average cloudy conditions at low latitudes could
decrease the chance of 2 or more consecutive days with
positive anomalies increasing the probability of two days
60
with negative anomalies (P00, Table 8). This will result in a

preferential increase in the ability of photosynthesis at low
latitudes to counteract UVR damage.
b. Chronic UVR Stress in austral South America
65
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80
85
90
95
100
105
110
If we assume phytoplankton and other biological organisms

or systems are adapted to average UVR (in this study
expressed as the climatology or annual cycle depicted in
Figs. 3 and 4), we can interpret the dominance of positive
irradiance anomalies as an environment conducive to chronic
UVR stress. More than 60% positive anomalies in irradiance
(Table 7) and an average 70% probability of two days with
above average UVR at all latitudes (Table 8) indicate that
cells damaged by UVR on a clear sunny day might not be
able to completely repair its cellular apparatus at night,
before the start of the next day. In this case, another day of
higher than average UV irradiance can further diminish the
cells ability to function at full capacity or even grow. A
very effective and efficient protective and repair mechanism
is needed for cells living in such environments. That was not
the case for Antarctic phytoplankton on long (18 hs) summer
days where cells could not recover its photosynthetic
capacity in the few dark hours. 37 A day of high UVR resulted
in a diminished photosynthetic performance at the beginning
of a second day with high irradiance. The damge from two
days of high-irradiance can be more important for
ecosystems dominated by picoplankton. Small cells are
particularly sensitive to DNA damage and photoproducts
accumulate from one day to the next.66 The pertinent
question that arises is, are organisms adapted to average or
daily maximum UVR? If they respond to average UVR, how
are their repair and protective mechanisms adapted to
preponderance of above average irradiances?
Aquatic organisms deal with excess UVR by protecting
themselves (i.e., screening), avoiding harmful radiation (i.e.
self shading or water column mixing) and by the various
repair mechanisms, in particular for DNA32,68 or D-protein in
the photosynthetic apparatus.69 If the adaptation of an
organism in the water is to maximum daily UVR then less
needs to be invested in short-term defense mechanisms. At
the same time, the cost of maintaining such an effective
system would be high. The emphasis in describing and
defining average UVR conditions (monthly averages63)
implies we believe biological systems respond to average
conditions. It is worth asking if organisms are adapted to
daily average or to daily maximum radiation, although the
answer might lie somewhere in the middle. We tend to
explain and interpret our results more on average values but
variability cannot be ignored and we propose here the ability
of biological systems (cell, organisms or ecosystems) to
handling variability could be intrinsic to survival in stressful
UVR conditions.
Conclusions
Organisms in South America are exposed to a latitudinal
gradient in UVR-weighted irradiances from subtropical to
subantarctic climates, with the magnitude of the
phytoplankton photosynthesis-weighted irradiances three to
10
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20
25
30
35
40
45
50
55
four times higher than DNA-weighted irradiances. In

general,
higher summer average weighted irradiances are

observed at lower latitudes (0.55 vs. 0.21 W cm-2
for DNA at Jujuy and Ushuaia, respectively);
the seasonal contrast in DNA-weighted irradiances

is more pronounced at higher latitudes (5.41
Max/Min irradiance at Jujuy vs. 90.34 Max/Min
irradiance at Ushuaia);
only latitudes lower than 42o S show measurable

DNA-weighted irradiances in winter.
UVR variability is biased towards above average
irradiances (and positive anomalies) where organisms are
exposed to several days of intermediate positive anomalies
followed by fewer days with negative, more intense,
anomalies. This condition can be labeled chronic damaging
UVR exposure and applies to both UVB- and UVA-sensitive
biological systems:
sixty two percent of DNA-weighted anomalies and

66.37% of phtoplankton-weighted anomalies are
positive, for the 1995-2002 time series.
Different patterns of day-to-day UVR variability are
observed:
variability in irradiance is maximum in Jujuy,

Buenos Aires, Valdivia and Trelew: for example,
the ratio of maximum/minimum irradiance in
summer is 1238 for Buenos Aires, in comparison to
a minimum of 71 in Santiago de Chile;
the presence of anomalous periods is present

everywhere, with consistent below (or above)
average irradiances lasting for several weeks (20 +
9 days for positive anomalies and 30 + 15 days for
negative anomalies for all locations with the
exception of Buenos Aires).
The gradient in average irradiance and its variablity is
modulated by climate, most importantly cloudiness. Clouds
affect irradiance intensity, spectral UVR and short-term
variability at each locality. For example,
maximum summer average irradiance is lower at

Jujuy (24.10 S) than at Santiago (33.27 S) due to
cloudiness (0.55 vs. 0.63 W cm-2);
seventy percent of the time a day with high UVR is

followed by another day with high UVR. And this
pattern is more prevalent at low latitudes (Table 8).
A low persistence in UVR is characteristic of this region
with a 4%-36% predictibility from day to day (lag = 1):
low and consistent autocorrelation of DNAweighted anomalies (0.26 - 0.36) at all sites in
consecutive days;
autocorrelations for phytoplankton photosynthesis

weighted anomalies show a large difference among
sites, varying between 0.04 and 0.27.
Potential net damage by UVR will differ depending on the
spectral sensitivity of cellular processes. DNA- and
phytoplankton photosynthesis-weighted irradiances, used
here to evaluate possible damage to biological systems
sensitive only to UVB or to both UVB and UVA, indicate
that the latitudinal gradient of UVB-effective irradiance
60
65
70
75
80
85
between 24 S and 54.5 S is more pronounced than the

UVA-effective irradiance:
weighted irradiances have a high correlation with

340 nm, higher for phytoplankton (r = ~1.0) than
for DNA (r = ~0.8);
340-nm has a decreasing influence with latitude for

DNA-weighted irradiance anomalies (r = 0.90 at
34.10 S and 0.69 at 54.5 S) but high everywhere
for phytoplankton-weighted irradiance anomalies;
influence of ozone is present everywhere for DNA

(r = ~0.4), and increases with latitude (r = 0.60 at
Ushuaia vs. 0.24 at Jujuy)
Higher latitudes have lower average UVB radiation and
higher frequency of below average irradiance providing
more relief from UVR damaging radiation than lower
latitudes. Transition probabilities from first-order Markov
chains indicate that:
DNA would have more probability of repair at

higher than at lower latitudes (P00 and P10 10% 40% higher at Ushuaia than Jujuy);
higher latitudes provide also more relief from UVR

stress for phytoplankton photosynthesis (P10 = 0.38
at high latitudes vs 0.26 at lower latitudes);
the period of recovery from UVR stress is shorter

for phytoplankton than for DNA as two consecutive
days of lower than average irradiance is lower at
higher latitudes (P00 is 0.42 in Ushuaia vs. 0.52 in
Jujuy).
Acknowledgements
90
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110
This work was supported in part by the Inter-American

Institute for Global Change Research (IAI) ISP and CRN026 projects (1996-2004). The authors wish to thank Mr.
Rodrigo Sanchez for UVR data processing data and graphics;
Dr. Luis Orce for setting up the UV network in Argentina,
Drs. San Romn, Buitrago, Labraga, Helbling and Diguez,
for their collaboration in data collecting at Ushuaia, Jujuy,
Puerto Madyn, Playa Unin and Bariloche, respectively. The
authors wish to acknowledge Drs. McPeters and Herman,
NASA/GSFC for providing total column ozone data; Mr. Jim
Ehramjian, James Robertson and the UV group (Biospherical
Instruments Inc.) for the calibration of the GUVs and
reference GUV and to Dr. Germar Bernhard for
spectroradiometer (SUV) data from San Diego and Ushuaia.
Ms. W. Kozlowski and L. Yarmey contributed to manuscript
preparation. Thanks also to Dr. Sharon Stammerjohn for
discussions on anomalies. Additional funding for data
collection and analysis was provided by the National Science
Foundation,
Argentinas
Consejo
Nacional
de
Investigaciones Cientificas y Tecnicas (CONICET), Chiles
Consejo Nacional de Ciencia y Tecnologia (CONICYT) and
Centro Austral de Investigaciones Cientificas (CADIC) in
Ushuaia, Argentina.
Notes and References
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25
1
Integrative Oceanography Division, Scripps Institution of
Oceanography, University of California San Diego, La Jolla, CA 920930218, USA, mvernet@ucsd.edu
2
Centro Austral de Investigaciones Cientficas (CADIC), B. Houssay
200, 9410, Ushuaia, Argentina, subediaz@fibertel.com.ar ,
carocamilion@hotmail.com
3
Instituto de Gentica y Biologa Molecular, Obligado 2490, 1428
Buenos Aires, Argentina, paladini@dna.uba.ar
4
Universidad de Chile, Departamento de Geofsica, Casilla 2777,
Santiago, Chile, hfuenzal@dgf.uchile.cl
5
Biospherical Instruments Inc., 5340 Riley St, San Diego CA 92110,
USA, booth@biospherical.com
6
Universidad de Chile, Casilla 70061 Correo 7, Santiago, Chile,
scabrera@conductor.med.uchile.cl
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Universidad de Magallanes, Laboratorio de Ozono y RUV, Casilla
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Universidad Austral de Chile, Facultad de Ciencias, Ed. Emilio Pugin,
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Universidad de la Patagonia S. J. Bosco, Dep. Fsica, Gales 50, 9100
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Quality of UVR Exposure For Different Biological Systems Along A Latitudinal Gradient

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CREATED USING THE RSC ARTICLE TEMPLATE (VER. 3.1) - SEE WWW.RSC.

ORG/ELECTRONICFILES FOR DETAILS

Quality of UVR exposure for different biological systems along a

The rapid ozone depletion in springtime over the Antarctic

This journal is The Royal Society of Chemistry [year]

Earth-Sun distance) are responsible for pronounced and

Journal Name, [year], [vol], 0000 | 1

processes and molecules in the marine environment and in

Markov chains and the persistence of UVR is determined by

Table 1. The UVR network in South America, with eight multi-channel

shorter and more intense periods of lower than average

Materials and Methods

Irradiances were obtained from measurements provided by

Ushuaia (6/28/2000 to 12/31/2000)

a2e x22 + b2e x2

a5e x52 + b5e x5

k5e x53 + l5e x52 + m5e x5 + n5e

a5d x52 + b5d x5

k5d x53 + l5d x52 + m5d x5 + n5d

a5p x52 + b5p x5

k5p x53 + l5p x52 + m5p x5 + n5p

a5f x53 + b5f x52 + c5f x5

k5f x53 + l5f x52 + m5f x5 + n5f

at the moment of the calibration, temperature corrections

Daily total ozone column was obtained from satellites

c. Calculation of irradiance weighted by biological action

Biologically weighted irradiances are easy to calculate from

This journal is The Royal Society of Chemistry [year]

a5ph x52 + b5ph x5

k5ph x5 + l5ph x52 + m5ph x5 + n5ph

use the complement of the SZA in the equation to ensure the

where Iw is the weighted irradiance, A, B, C, D and, E are the

Journal Name, [year], [vol], 0000 | 3

presence of the ozone hole, and is under rather

Table 3. Coefficients obtained when solving the multi-regressive

resulted of solving the multi-regressive equation for the four

To determine climatology we estimated the annual cycle

where yt is sampled value on day t; y is mean value of the

annual cycle; t is time; n is number of days in a cycle, in this

Anomalies for total column ozone, irradiance at 340 nm and

where Vd is the time series value at date d and

Atmospheric variables often exhibit statistical dependence

This journal is The Royal Society of Chemistry [year]

g. Markov chains - transition probabilities

f. Correlations and autocorrelations

with their own past or future values. In the terminology of

Error RMS (%)

Table 4. RMS error and determination coefficients between the hourly

Journal Name, [year], [vol], 0000 | 5

Figure 1. Biological Weighting Functions (BWF) of key molecules

where the probability Pr that a value be present today

Climate is a key issue to understand the short-term and

Table 5. Precipitation and cloud cover in the localities where GUV

b. UVR variability under clear sky conditions

Organisms at any given location and living for at least one

exposure to organisms living in the southern cone of South

Figure 2. Spectral Irradiance at different seasons and for varying

maximum sensitivity, 300 nm for DNA-weighted irradiances,

This journal is The Royal Society of Chemistry [year]

The UVR climatology at each site is defined as the annual

Journal Name, [year], [vol], 0000 | 7

Table 6. Irradiances in South America, weighted by DNA and

Phytoplankton photosynthesis-weighted irradiances at the