4>EPA
                           United States
                           Environmental Protection
                           Agency
                                                    Environmental Research
                                                    Laboratory
                                                    Gulf Breeze FL 32561
                           Research and Development
                                                    EPA/600/M-86/004 Feb. 1986
                           ENVIRONMENTAL
                           RESEARCH    BRIEF
                                 Critical Responses of
                       Populations of Crustacea to Toxicants
                                   Charles L. McKenney, Jr.
Introduction
To adequately assess the biotic hazard of contaminants
entering the marine environment, suitable criteria must be
determined for detecting the ecological damage resulting
from long-term exposure to low concentrations of contami-
nants. Considering the close phylogenetic relationship
between Crustacea and the insect target for which pesti-
cides are directed, it is not surprising that crustaceans often
are more sensitive to pesticides than are other marine
organisms (Williams and Duke, 1979; Costlow, 1982).
Research at the Environmental Research Laboratory (ERL)
Gulf Breeze is  directed toward determining appropriate
measurements  for assessing the long-term  effects of
pesticides on estuarine crustacean populations based on
both laboratory and field experimentation.

Toxicity tests in the last decade have established the mysid
shrimp, Mysidopsis bahia, as one of the most sensitive
membersof the estuarine community to pesticide exposure
(for a review see Nimmo and Hamaker, 1982) when simple
mortality is viewed as the major criterion. For the majority
of pesticides examined in life-cycle toxicity tests using this
estuarine crustacean, a sublethal reduction in reproductive
potential has proven to be the most sensitive criterion yet
determined for chronic biological effects in this zooplankton
species (Nimmo eta/., 1977, 1979, 1980, 1981).

Mysid  reproduction, culminating in the release of fully
developed juveniles by brooding females, is, however, far
different from the reproductive process seen  in benthic
crustacean populations in the estuarine community These
benthic populations, including  both commercially  and
ecologically important crabs  and shrimp, reproduce by
releasing free-swimming pelagic larvae which undergo a
complex metamorphic process prior to  settling into the
parental benthic population as young juveniles. Further-
                                               more, larvae of estuarine crustaceans have been shown to
                                               be particularly sensitive to environmental stress.

                                               It is generally recognized that lower levels of biological
                                               organization, the cellular and organismal levels, respond to
                                               environmental stress long before higher  levels such as
                                               community and ecosystem levels (Figure 1) (National
                                               Academy of Science, 1971; Waldichuk, 1979; Bayneef a/.,
                                               1980; Kinne, 1980). Furthermore, ecologists have for many
                                               years examined the productive processes of communities
                                               and ecosystems by studying the underlying physiological
                                               patterns of response of a species to environmental variables
                                               throughout its  life cycle (Mann, 1969; Winberg,  1971;
                                               Steel, 1973, 1974; Grodzinski et a/., 1975).

                                               The objective of the research summarized herein was to
                                               provide information  necessary to determine appropriate
                                               responses for assessing the long-term effects of various
                                               classes of pesticides on estuarine crustacean populations.
                                               Dose-response relationships  of pesticide toxicity and
                                               individual physiological functions  were  examined and
                                               compared for various life stages  of estuarine mysids
                                               (Mysidopsis bahia), grass shrimp (Palaemonetes pugio),
                                               and mud crabs (Eurypanopeus depressus). Correlations
                                               between physiological dysfunction of discrete life stages
                                               and alterations in the ecological fitness of the population
                                               should aid in the selection of sensitive, rapid, and inexpen-
                                               sive monitoring tools for predicting chronic effects of
                                               pesticides on pesticide-sensitive estuarine populations.

                                               This research represents an ongoing effort by the ERL-Gulf
                                               Breeze to identify particularly sensitive members of the
                                               estuarine community to potential microcontamination by
                                               specific classes of compounds and to provide the U.S.
                                               Environmental Protection Agency (EPA) with ecologically-
                                               sound test methods for assessing the ecotoxicological
                                               hazard of these compounds to this vulnerable community.

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 Figure 1.    Rationale for the usefulness of monitoring sublethal physiological parameters as short-term predictive indicators of
            long-term ecological damage by potential marine pollutants.
    Natural Environmental Factors
    Man-Made Microcontaminants
            Detoxification
            Disease Defense
            Regulation
            Adaptation
                                                                                      Energy
                                                                                   • Demanding
                                                                                     Processes
                          Hours
                          Days
                         Months
                          Years
                         Decades
               Structural & Functional Changes

                     Biochemical
                     Cellular
                     Tissue
                     Organ
                     Organism

            Reduction in Physiological Performance
            (Reduced Stress Resistance, Growth, Reproduction)

            Reduction in Ecological Potential
                                                     Consequences
               Structural & Functional Changes


                     Population
                     Community
                     Ecosystem
Earlier efforts at this laboratory produced the first marine
invertebrate  life-cycle toxicity  test using the estuarine
mysid, Mysidopsis bahia (Nimmo et a/., 1977, 1979, 1 980,
1981; Nimmo and Hamaker, 1982). Recent emphasis has
shifted toward an appreciation of the  functional role of
ecological indicator species and examination of the impact
of pollutant stress on community processes, resulting from
dysfunction among populations of these species both in
laboratory  and field  experimentation.  Furthermore,  an
increased awareness of environmental modification in the
toxic expression of potential  microcontaminants has re-
sulted in laboratory examination of the mediation of toxicity
by physicochemical environmental factors.

Measurements of Chronic Toxicity to Endrin in
Mysidopsis bahia Populations

During a  life-cycle toxicity test with Mysidopsis bahia
exposed  to the organochlorinated insecticide,  endrin,
various life stages were sub-sampled and measurements
made of metabolic and growth functions (Figure 2). These
individual physiological measurements were  combined
into bioenergetic and physiological indices and compared
with alterations in survival and reproduction during the
chronic exposure period.

Concentrations of endrin  that  were acutely lethal (120
ng/L)  stimulated  respiration  rates  of newly  released
juveniles after only one day of exposure (Figure 3). Exposure
to chronically lethal concentrations (60 ng/L) similarly
resulted in higher metabolic rates, but to a lesser extent.
Sublethal endrin exposure significantly reduced  growth
rates of rapidly growing juveniles after four days exposure
to concentrations which were lethal after  nearly three
weeks exposure (60 ng/L). In accordance with lower net
growth efficiencies (K2 values), increased metabolic de-
mands accompanying endrin exposure reduced the amount
of assimilated  energy available for production of new
tissue.
Energy metabolism during juvenile stages of M. bahia was
based  primarily  on utilization  of  lipid substrates (as
indicated by the  ratio of atoms of oxygen consumed to
atoms of nitrogen excreted). During the maturation period,
higher ammonia  excretion  rates and  lower O:N ratios
suggested increased amounts of protein used as an energy
source. Reduced young production  following  chronic
exposure to endrin  concentrations of 30 ng/L or greater
was correlated with  increased lipid  catabolism during
maturation of young mysids into adults  (Figure 3). It was
postulated that enhanced energy demands, associated with
higher metabolic  rates during sublethal  endrin exposure,
favored usage of  energy-rich lipid material at the cost of
shunting lipids away from gametogenesis and preparation
for reproduction.

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 Figure 2.    Life cycle  of the estuarine mysid shrimp, Mysidopsis bah/a, depicting  the various life stages subsampled for
            physiological measurements during chronic pesticide exposure.
                          Multiple Broods
                                for
                            2-3 Months
     First Brood Release
          Day 20
               Day 16
             Young Adult
                         I    .
       Maturation
     2° Sex Characteristics
           Day 12
                               Day 10
                          Advanced Juvenile
                                        Day 1
                                    Early Juvenile
                        Day 4
                       Juvenile
 Measurements of Chronic Toxicity to
 Thiobencarb in Mysidopsis bahia Populations

 A number of vital life processes of the estuarine mysid,
 Mysidopsis bahia, were examined throughout its life cycle
 during exposure to the carbamate herbicide, thiobencarb
 (Figure4). Initial exposure of juvenile mysids to thiobencarb
 resulted in elevated respiration rates.  Concentrations of
 thiobencarb which produced significant reductions in
 population  survival  through  a  complete  life cycle in
 approximately 24 days (181 ng thiobencarb/L) significantly
 stimulated respiration rates of juveniles after only four days
 of exposure. Increased metabolic demands with sublethal
thiobencarb exposure (> 22 ng/L) reduced the  amount of
 assimilated energy available for production of newtissue by
juvenile mysids, resulting in retarded juvenile growth rates
after only four days of exposure. Modifications in the energy
 metabolism of individual mysids exposed to thiobencarb
were related  to  decreases in total  young production of
discrete  populations. Greater  usage  of  proteinaceous
substrates for energy metabolism during the maturation of
M. bahia juveniles was altered by exposure to high sublethal
concentrations of thiobencarb. Higher 0:N ratios during the
maturation of these thiobencarb-exposed mysids suggest a
greater reliance on the more energy-rich lipid substrates in
order to support the elevated rates of oxidative metabolism,
resulting in less lipid material being available for gamete
production.
From the responses of M. bahia to chronic thiobencarb
exposure through an entire life cycle, it may be concluded
that metabolic dysfunction in individual mysids preceded
population responses important in determining community
trophic patterns. Lower secondary production in a crus-
tacean population, as indicated in this study by retarded
mysid growth rates and inhibited reproductive rates, would
alter the  energy-flow patterns between connected trophic
levels in  the ecosystem. Although thiobencarb is directly
toxic to fish  in low ng/L concentrations (Johnson  and
Fenley, 1980; Schimmel et a/., 1983), the reduced second-
ary production of mysid populations with exposure to low
ng/L concentrations of thiobencarb could indirectly affect
fish populations dependent upon mysids as a food source
(Darnell,  1958; Odum,  1971;  Chao and  Musick,  1977;
Mauchline, 1980)  and disrupt the balance  of  estuarine
food webs. Short-term measurements of altered metabolic
patterns in contaminated zooplankton, therefore, may offer
the potential  of monitoring for and predicting ecological
disruptions in the estuarine ecosystem at higher levels of
biological organization, i.e., at the population or community
level.

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Figure 3.    Summary of the effects of endrin on Mysidopsis bah/a exposed through an entire life cycle. VO2 = weight-specific
            oxygen consumption rate; K2 = net growth efficiency (percentage of assimilated energy used for growth); O:N = atoms
            of oxygen consumed to atoms of nitrogen excreted; VNH3 = weight-specific ammonia excretion rate.
        130
        110
         90
  QJ S'
  g>    -JQ
  I
  o
  —I
         50
         30
         10
                    VOi
                \
      Acute Lethality
                                 Growth

                                 K2
                                1

                             +
                                  Growth
                                   Chronic
                                   Lethality
                                                                              Young
                                                                              Production
                                                                                 • 1
                                                        •   VNH3
              Early
            Juvenile
             (Day 1)
Juvenile
 (Day 4)
        Advanced                    Early
        Juvenile                    Adult
        (Day 10)                    (Day 16)

Life Stage (Day of Exposure)
                              Adult
                             (Day 20)
Figure 4.    Summary of the effects of thiobencarb on Mysidopsis bahia exposed through an entire life cycle. Terms defined in
            legend for Figure 3.
       250
       200
       150
       100
        50
                    1
                    | Growth
                                                V02
                      Growth
                                                 VOz
                                                                           O:N
                                    Chronic
                                    Lethality
                                  . r Young
                                 ~ T Production
              Juvenile
               (Day 4)
              Advanced
               Juvenile
               (Day 10)
 Early
 Adult
(Day 16)
                                                       Adult
                                                      (Day 23)
                                             Life Stage (Day of Exposure)

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Field-Study Confirmation of Toxicity
Measurements of Mysidopsis bahia to Fenthion

Low-level exposure to fenthion, following ground  ULV
application of this organophosphate insecticide in various
mosquito control programs in  Florida, produced  both
increased mortality and sublethal growth  retardation  of
Mysidopsis bahia juveniles. Production of juvenile popula-
tions without continuous recruitment is the outcome of two
opposing processes: increased weight of individuals in the
population and decreased numbers of individuals in the
population due to mortality (Winberg, 1971).  Increased
mortality  and sublethal  growth retardation  of mysids
following low-level exposure to fenthion, therefore, would
result in reduced population production of this crustacean,
which  serves as an important  link in the estuarine  food
chain between primary producers and commercially im-
portant fish utilizing the estuary as a nursery.

Significantly higher rates  of oxygen consumption  of
fenthion-exposed  mysids eight days after the field spray
accompanied the reduced weights of the exposed mysids,
suggesting bioenergetic disruptions  in mysids exposed
sublethally to fenthion in  the field. Similar results have
been reported for M. bahia exposed to sublethal concentra-
tions  of  pesticides  through an  entire life cycle  in the
laboratory (McKenney, 1982, 1985). Increased metabolic
demands on mysids exposed in the laboratory to pesticides,
as indicated by higher respiration rates, reduced the
amount of assimilated energy available for production  of
new tissue, resulting in lower juvenile growth rates. These
field study results, therefore,  confirm those  of earlier
laboratory studies; short-term measurements of metabolic
dysfunction in mysids may predict altered production rates
in mysid populations.

Measurements of Chronic Toxicity to
Fenvalerate in Palaemonetes pugio Populations

Larvae of the estuarine grass shrimp, Palaemonetes pugio,
were reared in the laboratory from hatch through meta-
morphosis (Figure 5)  under optimal salinity conditions
(20 o/oo) in a range of lethal and sublethal concentrations
of the pyrethroid insecticide,  fenvalerate. Greater than 50
percent of the larvae exposed to measured concentrations
averaging 0.8 ng/L of fenvalerate died within four days of
continuous exposure.  This concentration  of fenvalerate
represents the lowest level of  fenvalerate  reported  to
produce  toxic effects  on  nontarget aquatic organisms.
Furthermore, since no mortality occurred for adult P. pugio
exposed to this same fenvalerate concentration, early larval
stages of estuarine crustaceans may be the  most sensitive
to pesticide toxicity.

Continual exposure to  the  sublethal  concentration  of
fenvalerate (a nominal concentration of 1.6 ng/L) delayed
completion of metamorphosis for developing grass shrimp
larvae  by  nearly two days. Extension of this particularly
vulnerable pelagic phase in the life cycle  may increase
predation pressure on the species. Increased predation on
larvae would reduce the number available for recruitment
into the parental benthic population or for dispersal  and
establishment of new populations in less severe environ-
ments.
Figure 5.   Life cycle of the ecologically important estuarine
           grass shrimp, Palaemonetes pugio. This species
           has been used  successfully at ERL-Gulf Breeze
           both in life-cycle and larval toxicity tests.
        Life Cycle of Palaemonetes Pugio
Spawning
   Juvenile Growth
     Maturation
   Larval
Development
                       Metamorphosis
                        to Postlarvae
Oxygen consumption rates of  newly-released  P. pugio
larvae were significantly higher after exposure for 24 hours
to fenvalerate concentrations which, after continual ex-
posure through the entire larval development, resulted in
significantly  fewer  larvae completing  metamorphosis.
These results suggest that  altered rates of respiration of
marine Crustacea may serve as rapid biological monitors of
detrimental effects of pesticide  exposure  to important
components of the estuarine community.

Upon completion of metamorphosis, postlarval P. pugio had
significantly higher rates of oxygen consumption in  sub-
lethal concentrations of fenvalerate, which previously had
not altered larval metabolism. This species shifts from a
free-swimming pelagic larvae to benthic postlarvae when
metamorphosis is complete. The highly adsorptive  nature
of fenvalerate (Schoor and  McKenney,  1983) could have
resulted  in adsorption  onto  the glass  surface of the
exposure beakers during this study. Presumably benthic
postlarvae  could have  been more readily  exposed  to
fenvalerate adsorbed to the glass surface than larvae in the
water column, thus affording greater  bioavailability and
increased metabolic sensitivity.

Responses of an estuarine organism to a toxicant are
dictated by the simultaneous influences of a number of
endogenous and exogenous variables (Figure 6). Tolerance
of osmotic stress is the most essential adaptation required
for a population  to  succeed in the  fluctuating salinity
conditions of an estuary (for reviews see Lockwood, 1976;
Gilles and Jeuniaux, 1979). Moreover, metabolic compen-
sation to salinity by developing larvae of P. pugio may  be
modified by sublethal toxicant exposure (McKenney and
Neff,  1981). Therefore, as an indication of the ecological
fitness of larval P. pugio, a secondary objective of this study
was to measure larval metabolism during osmotic stress to

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Figure 6.   Responses of an estuarine organism to a toxicant
           are dictated by the simultaneous influences of a
           number of endogenous and exogenous variables,
           including fluctuating salinity conditions.
                   Exogenous Factors
     (i.e., temperature, food levels, dissolved oxygen, etc.)
     Sublethal
     Toxicant
      Stress
Figure 7.    Complete larval development  of the estuarine
            mud  crab, Eurypanopeus  depressus, through
            metamorphosis.
                  Endogenous Factors
        (size, sexual maturity, stage in life cycle, etc.)
 determine if the ability of larvae to adapt to fluctuating
 salinities was altered by sublethal exposure to fenvalerate.

 Acute osmotic stress modified the metabolism of larval P.
 pugio reared in sublethal, nominal fenvalerate concentra-
 tions of 0.1 and 0.2 ng/L and these metabolic responses
 varied with  stage  of  development. After eight days of
 exposure to fenvalerate, oxygen consumption rates were
 elevated when larvae were acutely exposed to hypoosmotic
 stress (1 Oo/oo S). Metabolic responses of premetamorphic
 larvae to  hyperosmotic stress (30 o/oo S)  were  also
 modified by sublethal fenvalerate exposure. Alterations in
 metabolic-salinity patterns of larval grass shrimp devel-
 oping under sublethal fenvalerate concentrations suggest a
 reduction in the ecological fitness of this sensitive life stage
 manifested as a limitation in their capacity to adapt to the
 fluctuating salinity conditions of estuarine waters.

 Measurements of Chronic Toxicity to Lindane  in
 Eurypanopeus depressus Populations

 The various life stages of an  estuarine  mud crab (Eury-
 panopeus depressus),  including the zoea, megalopa,  and
 adult stages (Figure 7), exhibited different response patterns
 to lindane exposure, hypoosmotic stress, and interactions
 between the pesticide and salinity stress. Larval stages of £.
 depressus were more  sensitive to lindane exposure  than
 adults. The larval 96-hour LC50 value for lindane exposure
 was  0.66 IJQ/L as opposed to 25  /jg/L for adult crabs.
 Long-term  exposure to  sublethal concentrations of the
 organochlorinated insecticide, lindane, caused alterations
 in ionic  and osmotic regulatory ability  and related com-
 pensatory metabolic  mechanisms  in  E. depressus.  A
 lindane exposure concentration of 1.45 uq/L reduced the
 hemolymph osmotic concentrations m adult crabs. Chloride
 ion regulation, however, was  a more sensitive criterion,
 being disrupted at a lindane exposure concentration of 0.70
 /ug/L. A  lindane  exposure concentration of 0.01  ug/L
 increased larval mortality and altered larval respiration and
 ammonia excretion rates directly and in combination with
 salinity  stress.  Increased larval sensitivity to pesticide
exposure may reduce larval survival/recruitment in pesti-
cide contaminated areas, resulting in altered distributional
patterns in adult benthic populations. Similarly, disruptions
of osmo-regulatory  mechanisms  may  limit  the  natural
distribution of this  species  to  areas with less  salinity
variability.

Conclusions

Short-term measurements of altered metabolic patterns in
contaminated zooplankton offer the potential for monitoring
and predicting disruptions in  estuarine ecosystems. Bio-
energetic events at the organismal level precede secondary
production rate changes at the population level. Laboratory
responses to both an organochlorinated pesticide (endrin)
and a carbamate pesticide (thiobencarb) were characterized
by modification of energy metabolism in individual mysids,
and these results preceded  lower secondary production
rates caused by the same compounds. Sublethal exposure
of estuarine mysids to concentrations of these pesticides,
which  initially  elevated mysid respiration, eventually
inhibited growth  and  reproductive capacity  of  isolated
mysid  populations in  the laboratory.  Results  of these
laboratory studies were confirmed in a field study, indicating
that physiological measurements of metabolic dysfunction
in mysids exposed sublethally to pesticides may be used to
predict altered production rates in  mysid populations.

Larvae of estuarine crabs and shrimp were more sensitive
to pesticide exposure than adults. Larvae of the estuarine
grass shrimp, Palaemonetes pugio, died upon exposure to
the lowest concentrations of a synthetic pyrethroid insecti-
cide (fenvalerate) found to produce toxic effects  on non-
target aquatic organisms. Extremely low, sublethal levels of
fenvalerate (below the limit of analytical detection) reduced
the ecological fitness  of larval estuarine grass  shrimp
 (Palaemonetes pugio) by limiting their capacity to adapt to
 naturally occurring salinity fluctuations in estuarine waters.
 Concentrations of the organochlorinated insecticide, lin-
 dane, which were toxic to larval mud crabs (Eurypanopeus
 depressus), were several orders of magnitude below those

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toxic to adult crabs. Moreover, chronic exposure to sublethal
concentrations of this pesticide altered ionic and osmoregu-
latory abilities and related compensatory metabolic mech-
anisms in larvae  and  adults of this estuarine crab, with
larvae being more sensitive than adults.

The research is described in the following publications:

McKenney,  C. L., Jr.  1982.  Interrelationships between
  energy metabolism,  growth dynamics, and reproduction
  during the life cycle of Mysidopsisbah/a as influenced by
  sublethal endrin exposure. In: Physiological Mechanisms
  of Marine Pollutant Toxicity. W. B. Vernberg, A. Cala-
  brese, F. P. Thurberg, and F. J. Vernberg (eds.). Academic
  Press, New York, p. 447-476.
McKenney, C. L, Jr. and D. B. Hamaker. 1984. Effects of
  fenvalerateon larval  development of Palaemonetespugio
  (Holthuis) and  on  larval metabolism during  osmotic
  stress. Aquatic Toxicology,  5:343-355.
McKenney, C. L., Jr. 1985. Associations between  physio-
  logical alterations and population changes in an estuarine
  mysid during chronic exposure to a pesticide. In:  Marine
  Pollution and Physiology. F. J. Vernberg, F. P. Thurberg,
  A. Calabrese, and W.  B. Vernberg (eds.). University of
  South Carolina Press, Columbia, SC, p. 397-418.
McKenney, C. L., Jr., E. Matthews, D. A. Lawrence, and M.
  A. Shirley. 1985. Effects of ground ULV application of
  fenthion  on  estuarine biota. IV. Lethal  and sublethal
  responses of an estuarine mysid. Journal ofFlorida Anti-
  Mosquito Association,  in press.
Shirley, M. A. and C. L. McKenney, Jr.  1986. Influence of
  lindane on  survival,  osmoregulatory and metabolic
  responses of the larvae and adults of the estuarine crab,
  Eurypanopeus depressus.  In: Physiological Effects of
  Pollutants in Estuarine and Coastal  Organisms. W. B.
  Vernberg, A. Calabrese, F. P. Thurberg, and F. J. Vernberg
  (eds.). University of South Carolina Press,  Columbia, SC.
  in press.
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