C
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NATURAL LEVELS OF DDE AND THE UPTAKE OF C
BY DIFFERENT SIZE CLASSES OF THE MUSSEL
MYTILUS CALIFORNIANUS (CONRAD, 1837)
Karen K. Davis
Hopkins Marine Station:"
Stanford University
-DDT
FOOTNOTES
1Present address:
This work was supported in part by the Undergraduate Research
Participation Program of the National Science Foundation,
Grant + G4-5878.
O

RUNNING HEAD
DDT Uptake by !
tilu
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INTRODUCTION
Slightly over twenty years ago the introduction of DDT to
the world environment was received as the great panacea for the
control of insect pests of man's food crops. Furthermore, its
success in controlling malaria and other human diseases carried
by insects was undeniable. Only recently, however, have studies
of the long-term effects of DDT on non-target organisms begun
to cause alarm. Concern is growing now over the discovery that
although the actual DDT concentration in the oceans is only
in the order of 5-15 parts per trillion, DDT is taken up by
phytoplankton and then passed along the food chain through
zooplankton, fish, and birds where it may reach toxic levels
(Risebrough, Reiche, Peakall, Herman, and Kirven 1968; Risebrough,
Menzel, Martin, and Olcott 1967; Hickey and Anderson 1968).
The organism studied, Mytilus californianus, is a common
species of the intertidal region of the Monterey Bay area.
As a filter feeder it transports large quantities of water
through the mantle cavity. Many suspended nonliving particles
and small living organisms also enter with the water, come into
contact with the ctenidia and may or may not be picked up by
the palps and passed through the alimentary tract. It was sus-
pected that DDT might be accumulated from contact with water
passing through the mantle cavity as well as from particles of
page 2
food passing through the entire digestive system. The varia-
bility of background DDT levels occurring in natural populations
from different areas, especially in respect to size, was also
considered important because these levels could affect further
DDT uptake. Furthermore, because of the local variation in
pesticide levels, it would be interesting to know to what
extent the immediate environment of the animal might affect
its concentration of DDT.
The aim of this research was to determine background levels
of DDT and its residues in natural populations of Mytilus
californianus and to compare the uptake of C-labeled DDT by
the mussel through different modes of entrance: directly from
the surrounding sea water, and from phytoplankton, part of its
natural food supply. The results of this study show that DDT
does not only enter the food web via the phytoplankton, but may
also be taken up in substantial quantities by organisms higher
in the food web directly from the surrounding water.
MATERIALS AND METHODS
The experimental animals were collected in April and May
of 1969 from four different localities around the Monterey
peninsula. These were Cabgrillo Point, Point Pinos, Point Joe,
and Seal Rock in Monterey County, California. The areas were
chosen to represent populations from typical bay and outer coast
localities. The animals were all collected from approximately
the same tidal level to minimize the difference in growth
7
page 3
conditions related to differential exposure to the water.
The mussels for each area were kept in separate aerated
aquariums and supplied with a continual flow of sea water at
ambient temperature. They were allowed several days to adjust
to the new situation and to begin normal feeding activity
(Macginitie 1941).
The organisms used in the experiments were divided into
three size groups. Size I organisms were 0.8-1.2 cm. in length,
size II organisms 2.8-3.2 cm., and size III organisms 4.8-5.2
cm. Measurement was made at the widest point of the shell
parallel to the ventral edge. These young organisms were chosen
for the experiments to decrease the probability of spawning
during the experiment, thereby minimizing the possibility of
a large change in lipid content and possibly also DDT content
during the experiments. Though M. californianus has been observed
to be sexually mature at a length of 25 mm. and an age of
approximately 4 months, it does not appear to spawn until the
age of about 1 year (Coe and Fox 1942) at a length of approxi¬
mately 80 mm. (Fox and Coe 1943). Young animals were also
chosen for the experiments because of their ability to adjust
more quickly to experimental situations (Jørgensen 1960, 1966;
Macginitie 1941; Coe and Fox 1942).
DDE levels in natural populations were measured directly
by gas chromatography. Samples were extracted and prepared for
chromatographic analysis according to the method described by
Stanley and LeFavoure (1965). Analysis of the extracts was
page 4
performed on a Beckman GC-4 equipped with an electron capture
detector. The chromatograms were performed isothermally (20090)
on 3 per cent QF-1 on Chromosorb W, 80-100 mesh treated with
DMCS. The carrier gas was helium.
All of the experiments involving C uptake were run in
a similar manner. Prior to incubation of the mussels, the
shells were scraped free of barnacles, limpets, algae and other
attached organisms. The animals were then placed vertically
on a flat surface on absorbent paper to allow water in the mantle
cavity to drain out. This not only induced the mussels to take
in water soon after placement in the DDT solution (Fox, Sverdrup.
and Cunningham 1937), but it also helped clear any food or
inorganic particles suspended in the mantle cavity water.
All experimental organisms were incubated in closed jars
to minimize the loss of DDT from solution through codistillation
(Acree, Beroza, Bowman 1963). Since previous studies have shown
that Mytilus can filter particles as small as 1 u (Jørgensen
1960; Jorgensen and Goldberg 1953) and oxyhemoglobin particles
as small as 40-50 A (Fox, Oppenheimer, and Kittredge 1953),
the sea water to be used in the containers was millipore filtered
successively through 8 u, 0.45 u, and 0.22 u pore filters to
insure that no particles greater than 0.22 u remained in solution.
In early experiments one organism was placed in each con-
tainer. Size I organisms were incubated in 60 ml. water in a
half-pint jar, size II organisms in 180 ml. in a one-pint jar,
and size III organisms in 540 ml. in a one-quart jar. In later
2
page 5
experiments three organisms, one of each size class, were placed
in each one-quart jar with 780 ml. filtered sea water.
Animals were incubated in concentrations of 10 p.p.t.,
100 p.p.t., 1.0 p.p.b., and 1.4 p.p.b. C -DDT for 12 hours.
Some of the animals were assayed for uptake immediately at
the end of the 12 hours, while others were kept 23 hours in
normal sea water before being tested to show the level of DDI
retained.
The amount of C14 uptake was determined by dissecting the
animal, taking its wet weight, homogenizing it in 1,4-dioxane,
and counting in the scintillation counter. Size I organisms
were homogenized twice in 14 ml. dioxane, size II organisms
twice in 3 ml. dioxane, and size III organisms twice in 6 ml.
dioxane. The extracts from the two homogenates were combined,
and a 2 ml. aliquot was placed in toluene solution and counted
2 minutes in a Nuclear-Chicago Unilux II scintillation counter.
The primary foods of Mytilus are diatoms, dinoflagellates,
and detritus (Field 1921-1922; Buley 1936; Macginitie 1941;
Coe and Fox 1942). The diatom Nitzschia closterium minutissima
was chosen for the feeding experiments because it was readily
filtered out of suspension and digested by Mytilus (Schultis
1969). C+-DDT uptake from phytoplankton was tested in a similar
manner to earlier uptake experiments. A sample of Nitzschia
consisting of approximately 1.24 x 10 cells was taken from a
pure culture and incubated in 6.6 p.p.b. C-DDT for 64 hours
at room temperature in diffuse light. This concentration of
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page 6
DDT does not appear to harm the phytoplankton (Bailey 1969).
The bottles were periodically shaken to keep the phytoplankton
in a uniform suspension, and at the end of the incubation
period, the solution was centrifuged and rinsed twice with
filtered sea water to remove any DDT which might have adsorbed
to the surface of the phytoplankton. The phytoplankton were
then introduced to four experimental jars with 3 organisms each,
one from each size class. About 2.5 x 10' cells were provided
per container. Both the experimental conditions in the jars
and the treatment of the animals at the end of the 12-hour
incubation period were identical to those in the experiments
with C uptake from water alone.
ESULTS
Assays with the gas chromatograph revealed a distinct
difference in DDE levels in organisms of different sizes but
a minimal difference between organisms from different areas
(Table 1, Figure 1). Total DDE content was seen consistently
to decrease with increasing size of the organisms. The concen¬
tration of DDE is expressed in terms of wet weight of all the
body tissues combined. Table II shows the correlation between
wet and dry weight for the three size classes. However, the
relationship between size class and DDE level appears to be
identical whether plotted against wet weight or dry weight.
The experiments with the uptake of C+-DDT directly from
the water revealed a similar trend of DDT accumulation with size.
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page 7
The preliminary experiments showed that the size I organisms
(0.8-1.2 cm.) concentrated DDT 20 to 70 times greater than did
the size III organisms (4.8-5.2 cm.). This level did not
significantly decrease 23 hours after termination of the incu¬
bations which suggests retention of virtually all DDT taken up
during the experiment. However, correction of these values
was necessary to account for the small organisms having available
a greater volume of water per unit body weight than the large
ones. The correction factor reduced the relative uptake of
size I organisms to approximately 5 tines the uptake by size
III organisms (Table 3). For further verification of this
prediction, three organisms, one of each size class, were tested
in the same container such that each would initially have available
the same total amount of water. Five replicates of the experi-
ment were made and the initial level of C-DDT in the water
was the same for each container, 1 p.p.b. After incubation for
12 hours, the animals were removed and assayed. The results
are shown in Table 4 and Figure 2. The same trends noted earlier
were found, and the earlier corrected values were found to be
valid, as size I organisms concentrated C+-DDT approximately
5 times more than size III organisms.
This same tendency was again evident in the experiments
with the uptake of C-DDT from Nitzschia (Table 5, Figure 3).
The smaller Mytilus concentrated the DDT 4 to 4 times greater
than did the larger ones. However, note that while the same
trend is apparent, the total uptake was far less than the uptake
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page 8
directly from the water. Furthermore, it is distinctly possible
that this small degree of C+ uptake from a potential food
source was a result of both uptake from the food itself and
from C which had been released into the water by the labeled
Nitzschia.
DISCUSSION
There are several possible explanations for the variation
in DDE accumulation between large and small Mytilus. This
trend could easily be understood if the lipid content of the
mussels showed a similar decrease with size, but instead an
increase occurs. Much of the lipid in a mussel is stored in its
gonads (Rodegker and Nevenzel 1964), and as the animal grows,
this organ greatly increases in size, extending into the mantle
and surrounding many of the tissues. Shortly before the mussel
spawns, the dry weight of the gonads may be equal to or greater
than that of all the other body tissues combined (Coe and Fox
1942). Fat is deposited in the connective tissues of the gonads,
causing a direct relationship between total fat content and the
size of the reproductive organs. In these experiments it was
noted that the gonads of the larger organisms were very exten¬
sive, and there was no evidence that the animals had recently
spawned thereby lowering their lipid content. One would predict
from the increase in lipid content of larger mussels that they
would take up more C-DDT than the smaller ones. However, this
was not the case.
7
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page 9
The relative level of DDE could be higher in the smaller
organisms if only the larger ones had the capability of metabo-
lizing and excreting the pesticide. However, other studies
have shown no such breakdown occurring in either small or large
M. californianus (Davis and Comrey 1969).
The findings of this study may be related to the fact that
young Mytilus grow, filter, and adjust more rapidly than older
ones when disturbed (Coe and Fox 1942). The average amount of
water filtered by M. californianus decreases with age (Fox,
Sverdrup, and Cunningham 1937) and as a possible consequence,
the amount of DDT taken up from the environment via both the
water and phytoplankton may decrease. Mussels may also cease
filtering when disturbed. Jørgensen (1960, 1966) was able to
obtain normal filtering rates in the lab only from very small
mussels. Furthermore, though a mussel normally feeds 97-99%,
of the time (Loosanoff 1942), it may stop eating when disrupted,
the younger ones adjusting more quickly to experimental situations.
In addition, a mussel may transport water without feeding if
the sheet of mucus normally covering the gills is not present
(Macginitie 1941). As a result, a mussel could take up more
DDT directly from the water than through feeding on phytoplankton.
The results of this study show that substantial quantities
of DDT are obtained directly from the sea water. Furthermore,
they suggest that this mode of entrance may be even more impor¬
tant in the uptake of DDT than the accumulation from feeding on
contaminated food materials. However, more information is
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page 10
needed to fully evaluate these findings. If indeed Mytilus
is typical of marine organisms in its ability to take up so
much insecticide directly from the sea water (Kaplan 1969;
King 1969; Phillips 1969; Sutton 1969), then the marine organisms
could be in even greater danger from DDT pollution than originally
thought.
78
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FIGURE LEGENDS
Figure 1. The solid colored bars represent organisms
from Seal Rock, the horizontally crossed bar an organism from
Point Joe, and the cross-hatched bar an organism from Cabarillo
Point. The first three bars show mean DDE levels for 50, 10.
and 6 mussels, respectively, while the last three show levels
for individual animals.
Figure 2. Each graph shows the uptake over a 12-hour
period of C-DDT directly from the water by a l cm., a 3 cm..
and a 5 cm. Mytilus incubated together in the same jar. The
initial C+-DDT concentration was 1.0 p.p.b. in 780 ml. filtered
sea water.
Figure 3. Each line represents the uptake over a 12-hour
period of C-DDT from Nitzschia by a l cm., a 3 cm., and a
5 cm. Mytilus incubated together in 780 ml. filtered sea water.
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SUMMARY
1. In natural populations Mytilus californianus of lengths
1, 3, and 5 cm. have been found to have concentrations of DDE
of 120, 96, and 62 p.p.b., respectively, per wet weight of body
tissues.
2. Mussels can take up substantial amounts of DDT directly
from the surrounding sea water.
3. Over a 12 hour period mussels 1 cm. in length take up
5 times as much C+-DDT directly from the water as animals
5 cm. in length.
4. The uptake of C+-DDT from phytoplankton by the smaller
mussels is 4-44 times greater than the uptake by the larger
animals over the same time period.
5. The amount of water filtered by a mussel decreases
with age and may account for the more rapid uptake of DDT by
younger organisms.
6. The ability of younger mussels to adjust to experimental
conditions more readily than older ones may also partially explain
the differential uptake rates.
7. It is suggested that if Mytilus is typical of marine
organisms in its ability to concentrate DDT directly from the
surrounding sea water, then the marine pesticide pollution
problem may be greater than originally thought.
REFERENCES
Acree, F., M. Beroza, and M. C. Bowman. 1963. Codistillation
of DDT With Water. Agricultural and Food Chemistry, 11:
278-280.
Bailey, S. 1969. Personal Communication.
Buley, H. M. 1936. Consumption of Diatoms and Dinoflagellates
by the Mussel. Bulletin. Scripps Institution of Ocean-
ography. Technical Series, 4:19-27.
Coe, W. R., and D. L. Fox. 1942. Biology of the California
Sea-Mussel (Mytilus californianus) I. Influence of Temp-
erature, Food Supply, Sex and Age on the Rate of Growth.
Journal of Experimental Zoology, 90:1-30.
Davis, J. D., and C. Comrey. 1969. Personal Communication.
Field, I. A. 1921-22. Biology and Economic Value of the Sea
Mussel, Mytilus edulis. Bulletin of the United States
Bureau of Fisheries, 38:125-259.
Fox, D. L., and W. R. Coe. 1943. Biology of the California
Sea-Mussel (Mytilus californianus) II. Nutrition, Meta-
bolism, Growth and Calcium Deposition. Journal of Experi-
mental Zoology, 93:205-249.
C. H. Oppenheimer, and J. S. Kittredge. 1953. Micro-
filtration in Oceanographic Research II. Retention of
Colloidal Micelles by Adsorptive Filters and by Filter-
page 2
feeding Invertebrates; Proportions of Dispersed Organic
to Dispersed Inorganic Matter and to Organic Solutes.
Journal of Marine Research, 12:233-243.
H. U. Sverdrup, and J. P. Cunningham. 1937. The Rate
of Water Propulsion by the California Mussel. Biological
Bulletin, 72:417-438.
Hickey, J. J., and D. W. Anderson. 1968. Chlorinated Hydro-
carbons and Eggshell Changes in Raptorial and Fish-Eating
Birds. Science, 162:271-273.
Jørgensen, C. B. 1960. Efficiency of Particle Retention and
Rate of Water Transport in Undisturbed Lamellibranchs.
Journal du Conseil. Conseil Permanent International
pour l'Exploration de la Mer, 26:94-116.
1966. Biology of Suspension Feeding. Pergamon Press,
New York, N. Y. 357 p.
, and E. D. Goldberg. 1953. Particle Filtration in Some
Ascidians and Lamellibranchs. Biological Bulletin,
105:477-489.
Kaplan, P. 1969. Personal Communication.
King, R. 1969. Personal Communication.
Loosanoff, V. L. 1942. Shell Movements of the Edible Mussel,
Mytilus edulis (L.) in Relation to Temperature. Ecology,
23:231-234.
Macginitie, G. E. 1941. On the Method of Feeding of Four
Pelecypods. Biological Bulletin, 80(1):18-25.
Ae
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page 3
Phillips, G. F. 1969. Personal Communication.
Risebrough, R. W., D. B. Menzel, D. J. Martin, and H. S. Olcott.
1967. DDT Residues in Pacific Sea Birds: a Persistent
Insecticide in Marine Food Chains. Nature, 216:589-591.
Risebrough, R. W., P. Reiche, D. B. Peakall, S. G. Herman, and
M. N. Kirven. 1968. Polychlorinated Biphenyls in the
Global Ecosystem. Nature, 220:1098-1102.
Rodegker, W., and J. C. Nevenzel. 1964. The Fatty Acid
Composition of Three Marine Invertebrates. Comparative
Biochemistry and Physiology, 11:53-60.
Schultis, S. 1969. Personal Communication.
Stanley, R. L., and H. T. LeFavoure. 1965. Rapid Digestion and
Cleanup of Animal Tissues for Pesticide Residue Analysis.
Journal of the Association of Official Agricultural Chemists,
18:666-667.
Sutton, J. 1969. Personal Communication.

86
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Range of
Length
(mm.)
8.0-12.0
29.0-31.0
19.0-50.9
- -
Mean
Length
(mm.)
9.1
29.9
19.7
19.3
50.5
52.0
TABLE 1. DDE LEVELS IN MYTILUS

Mean
Collection
Wet Wgt.
Indi¬
Site
viduals
(mgm.)
50
Seal Rock
22.5
851
10
Seal Rock
Seal Rock
3580
3780
Point Joe
1150
Seal Rock
4810
Cabarillo Point
p.p.b. DDE
(per wet
wgt.)
120
96
62
16
31
C
9
C
Length
(mm.)
8.2
8.9
10.5
10.5
11.0
28.8
29.2
9.8
31.4
31.5
49.1
50.5
51.0
51.0
51.
TABLE 2. WET WEIGHT VS. DRY WEIGHI
-
Wet Wgt.
Mean Dry
Dry Wgt.
Wgt. (mgm.)
(mgm.)
(mgm.)
6.9
3.3
12.4
4.9
1.5
7.9
28.0
27.
10.1
6.9
719
116
839
187
235
176
255
191
1072
165
625
3607
609
3906
894
5232
4413
750
2808
109
651
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TABLE 5. CH-DDT UPTAKE FROM NITZSCHIA
S OF MYTILUS
BY 3 SIZ
—
-DDT
Mean C
Size
Mean Wet
Range (mm.)
Wgt. (mgm.)
uptake (p.p.b.)
22.
8.0-12.0
8.70
28.0-32.0
1010
3.03
3957
18.0-52.0
2.01