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Structure and Function of Freshwater Ecosystems: Assessing the Potential

Impact of Pesticides

Article · January 1989

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pages 4-17, In J.R. Voshell, Jr. (ed.) Using mesocosms to
assess the aquatic ecological risk of pesticides: theory
and practice. Misc. Publ. No. 75 Entomol. Soc. Amer. Dec. 1989 Wallace: St

Structure and Function of Freshwater Ecosystems: observation

Assessing the Potential Impact of Pesticides quired to n
Ecosyste
]. BRUCE WALLACE component
Department of Entomology source, i.e.
University of Georgia as climate a
Athens, Georgia 30602 geochemist
trient avail,
autotrophs
ABSTRACT Animal communities play significant roles in In this article, I review some basic concepts carbon con
the processing of organic matter and nutrient cycling within relating to structure and function of ecosystems
lentic and lotic ecosystems. One of the most overwhelming required 1
tasks associated with assessing the impact of pesticides in with emphasis on the functional aspects of in- crobes). Ba
ecosystems is that of assessing the indirect effects that result vertebrates and some examples of the influence sumed, het<
from the direct effects of pesticides on the biota. Because of an insecticide on headwater streams, based herbivores
all functional groups do not respond similarly to pesticide on manipulative studies we have done at the biomass) ai
treatments, changes in functional community structure and
production can alter drastically ecosystem-level phenome- Coweeta Hydrologic Laboratory (U.S. Forest of other an
nona such as energy flow, food chains, nutrient cycles, and Service) in western North Carolina. I also ad- nivores, or
heterotrophic "regulation" (cybernetics) within the ecosys- dress the following question: Is our current plant prodi
tem. Some specific examples of indirect effects of pesticides knowledge of the structure and function of aquatic trital, or de<
on aquatic ecosystems are given. There are definite needs ecosystems sufficient to draw meaningful con-
for mesocosm and ecosystem-level testing of pesticides; how- processed c
ever, at the same time we need to ask the following question: clusions from toxicological tests? Kitchell et
How applicable are the results obtained from one freshwater Carpenter
system to another? Too few comparative studies of aquatic Ecosystem Structure and Function heterotropl
ecosystems exist to make broad sweeping generalizations their influe
between various regions, which differ in abiotic and biotic
Mclntosh (1985) has provided an entertaining
characteristics. More comparative studies of freshwater eco- ulation.
historical perspective of the development of the
systems are sorely needed. Manipulations of ecosystems, Studies <
which include applications of pesticides, can contribute sub-
ecosystem concept from its inception in the 1930s, properties
stantially to our knowledge of ecosystem structure and func-
through systems ecology, the International Bi- productivit
tion; however, the researchers must have a good working
ological Program, and systems analysis. Odum
knowledge of the particular ecosystem before such treat- proach circ
(1971) defined an ecosystem as follows: "Any unit
ments. Before system-level manipulations are made several
including all of the organisms (i.e., the "com-
questions should be asked, which should enable researchers
to formulate predictions and design experiments that key in
munity") in a given area interacting with the
on ecosystem processes that may be influenced by pesti-
physical environment so that a flow of energy
cides. leads to a clearly defined trophic structure, biotic INPUT
diversity, and material cycles (i.e., exchange of v
materials between living and non-living parts)
IN THE PAST DECADE, there has been an upsurge within the system is an ecological system or eco-
CO 2 , H 2
of interest in the influence of pesticides and other system." An ecosystem consists of any part of Other N
toxicants on ecosystems. Although there have the biosphere where inputs and exports of the NO , PO t
been many review articles concerned with the area selected for study can be measured (Neu-
influence of pesticides on ecosystems, many of hold 1986). As pointed out by Neuhold (1986)
these (e.g., Pimentel & Edwards 1982, Levin for aquatic systems, an ecosystem can range from
et al. 1984, Kimball & Levin 1985, Cairns 1986, between the banks of a pond, lake, or stream to
Neuhold 1986) stress the lack of knowledge of an entire catchment or watershed. Studies of \
the impact of toxicants on ecosystems. However, streams may be based upon a watershed ap-
Barnthouse et al. (1986) among others have proach, where small headwater streams are the
questioned the extent to which ecosystem theory main area of study (e.g., Fisher & Likens 1973,
is developed to be useful in all ecological risk Webster et. al. 1983), studies of individual reaches
assessments, especially with regard to providing or segments of streams (e.g., Fisher 1977), or
long-term predictions on the ecological conse- larger scale studies, which view streams as a
quences of the adding of toxic substances to eco- continuum of changes from headwaters to mouth Irig. I. Sim
systems. (Vannote et al. 1980, Minshall et al. 1983). The cycles as dasl
important criteria are that estimates of input and herbivore pal
organic math
output (either energy or nutrients) are made for llow. Further
© 1989 Entomological Society of America each study. Obviously, as the spatial scale of direction.
 Wallace: Structure and Function of Freshwater Ecosystems

observations increases, so does the effort re- taxa and may have certain advantages (i.e., pri-
quired to make meaningful measurements. marily the reduced expense of measurements)
Ecosystems are composed of abiotic and biotic for measuring long-term changes in ecosystem
components and some ultimate driving energy function in that it does not involve measure-
source, i.e., sunlight. Abiotic components, such ments of individual taxa or focus on internal pro-
as climate and geology (including local lithology, cesses (e.g., O'Neill & Richie 1979).
geochemistry, geomorphology), influence nu- A simple summary of energy and nutrient flow
trient availability and the overall rate at which through an ecosystem is outlined in Fig. 1. In
autotrophs (plants) fix carbon dioxide to organic lotic and lentic ecosystems, the flow of energy
ncepts carbon compounds (plant biomass), which are through the detrital pathway of heterotrophs may
/stems required by heterotrophs (animals and mi- far exceed that of the herbivore pathway. In many
of in- crobes). Based on the kinds of compounds con- aquatic ecosystems, such as headwater streams,
Suence sumed, heterotrophs can be broken down further: most of the plant biomass that drives the system
based herbivores (primary consumers that feed on live is produced outside of the stream (allochthon-
at the biomass) and carnivores (secondary consumers ous) and enters the stream as leaf material from
Forest of other animals whether herbivores, other car- the surrounding forest. Herbivores, or animals
so ad- nivores, or detritivores) (Fig. 1). However, most that feed directly on living plant tissue, gener-
urrent plant productivity is probably used in the de- ally represent a minute portion of the hetero-
iquatic trital, or decomposer, food chain rather than being trophs in shaded, headwater streams; however,
il con- processed directly by herbivores. O'Neill (1976), the relative importance of allochthonous versus
Kitchell et al. (1979), O'Neill & Richie (1979), autochthonous production (and the herbivore
Carpenter et al. (1985), and others have viewed versus detritivore pathways) may vary between
heterotrophs as key factors in ecosystems through streams and may differ by order (size), degree
their influence on nutrient cycling and rate reg- of shading, nutrients, biomes, etc. (Vannote et
aining ulation. al. 1980, Minshall et al. 1983). Streams, more
of the Studies of ecosystems may emphasize certain so than many lakes and terrestrial ecosystems,
1930s, properties of the system, such as total biomass, are characterized by strong unidirectional flow.
ml Bi- productivity, and nutrient cycling. This ap- Thus, in streams the predominant flux of nu-
Odum proach circumvents consideration of individual trients and energy is in a downstream direction.
iv unit
"com-
th the SUNLIGHT
:nergy
INPUT
biotic
nge of lie at
parts) heat
>r eco- Photosynthesis eat
CO 2 , H 2 O Plant
part of Other Nutrients
NO,, PO,., K + ect. Biomaas
of the
1 (Neu-
(1986) \ V t- - j \\
^e from \
earn to \ \ >^ 1 I
lies of Detritus k
EXPORT Plus Fung:
;d ap- - £ Bacteria • 1 1
heat
re the ^
1973,
^
heat \
>v
caches ENERGY FLOW
•7), or NUTRIENT CYCLES X
s as a EXPORT
mouth Fig. 1. Simple energy flow and nutrient cycling diagram lor an ecosystem with energy flow as solid lines and nutrient
.). The cycles as dashed lines. In lentic and lotic systems, the flow of energy through the detrital pathway may exceed that of the
ut and herbivore pathway and in heavily shaded, forested, headwater streams, most of the energy .source consists ofallochtlionous
organic matter inputs from the surrounding forest, and, hence, the detrital pathway is the dominant pathway of energy
ide for flow. Furthermore, because of the unidirectional flow of water in streams, energy flow and nutrient cycling is in a downstream
•ale of direction.

|§^x,:/l/;;:r>,;;

Because of this downstream movement, there is
very little nutrient cycling in streams at a single
point. However, both nutrients and energy, which
are transported from upstream areas, may be
available to consumers in downstream reaches.
Hence, a considerable amount of nutrient cy-
cling may occur in streams, but because of its
spatial arrangement such movements have been
termed "spiralling" (Webster 1975, Wallace et
al. 1977, Newbold et al. 1982).
Although the scheme outlined in Fig. 1 has
some value in simplistic conceptualization of
ecosystems, in reality, few species fit exclusively
into one trophic level, because most aquatic in-
sects are somewhat omnivorous. For example,
many herbivores also consume small animals as
well as detritus while feeding. Conversely, de-
tritivores consume associated microbiota, which
may include fungi, bacteria, meiofauna, and dia-
toms as well as plant tissue. Hence, such clas-
sifications as shown in Fig. 1 generally provide
limited insight into mechanisms of food utiliza-
tion among aquatic organisms.
To address the problem of linking inverte-
brates to specific trophic level analysis, K. W.
Cummins devised a scheme of functional clas-
sification of animals based on morpho-behavioral
mechanisms used to acquire their food (Cum-
mins 1973, Merritt & Cummins 1984). These
functional groups are as follows: Scrapers, an-
imals adapted to graze or scrape materials
(periphyton, or attached algae, and its asso-
ciated microflora) from mineral and organic
substrates; Shredders, organisms that chew on
living vascular plants or consume decomposing
coarse vascular plant tissue (>1 mm diameter)
along with its associated microbiota; Collector-
gatherers, animals that feed primarily on de-
composing fine particulate organic matter
(<1 mm diameter) which is deposited within
lakes and streams; Collector-filterers, animals
with specialized anatomical structures (e.g.,
setae, mouthbrushes, or fans) or silk and silk-
like secretions that act as sieves to remove par-
ticulate matter from suspension (Jorgensen
1966, Wallace & Merritt 1980); and, Preda-
tors, those organisms that feed on animal tis-
sue by either engulfing their prey or piercing
prey and sucking their body contents. Al-
though the above groups cover most inverte-
brate feeding, Merritt & C u m m i n s (1984)
include two additional groups, Macrophyte-
piercers and Parasites.
These broad categories refer primarily to
modes of feeding and not to the type of food
Miscellaneous Publications No. 75
per se. For example, several filter-feeding
insects of high gradient streams are primarily
carnivores (e.g., Benke & Wallace 1980).
Scrapers also consume quantities of epilithon
(Locke 1980) and not solely periphyton.
Likewise, although shredders that chew leaf
litter in streams may select those leaves that
have been "microbially conditioned" by col-
onizing fungi and bacteria (e.g., Cummins &
Klug 1979), these shredders also ingest at-
tached algal cells, protozoans, and various
other components of the meiofauna during
feeding (Merritt & Cummins 1984). Some
shredders apparently obtain very little of their
assimilated energy directly from microbial
biomass (Cummins & Klug 1979, Lawson et
al. 1984, Findlay et al. 1986; however, see
Arsuffi & Suberkropp 1986) although micro-
bially derived enzymes from endosymbionts
or enzymes obtained from microbes ingested
with leaf tissue may be important in cellulose
hydrolysis (Sinsabaugh et al. 1985). The de-
scriptions of these mechanisms to obtain foods
are valid, and yet many questions remain
c o n c e r n i n g the sources of p r o t e i n ,
carbohydrates, fats, and assimilated energy
to each of these functional groups.
Functional Role of Animals in
Freshwater Ecosystems
Animal communities play significant roles in
the processing of organic matter and nutrient
cycling within lentic and lotic ecosystems. Their
feeding activities influence phytoplankton, per-
iphyton, vascular aquatic hydrophytes, coarse
particulate organic matter (CPOM = >1 mm
diameter), fine particulate organic matter (FPOM
= <1 mm diameter), microbial populations, other
animal populations, and nutrient cycles within
aquatic ecosystems.
Scrapers. Lamberti & Moore (1984) have
summarized some of the roles proposed for
scrapers in freshwater systems as follows: Graz-
ing reduces the number of cells and thickness
of the algae layer, hence, preventing nutrient
and light limitation to algae. Grazers may ingest,
dislodge, and thus remove dead and senescent
algal cells and thereby increase the turnover rates
(P/B = production/standing crop biomass) of the
algal population. Waste products from grazers
may provide nutrients for algal growth. Me-
chanical breakage of algal cells by grazers may
release nutrients and enhance nutrient availa-
bility to other cells. Grazers may prevent ac-
 ). 75 Wallace: Structure and Function of Freshwater Ecosystems
ding cumulation of certain waste products in algae
arily that would otherwise curtail growth of algae, and
)80). they may selectively remove less productive al-
. thon gal species, while enhancing fast-growing spe-
ton. cies of algae. Several of these roles remain
'leaf speculative, but they do indicate some of the
that mechanisms by which these herbivores affect
coi- their food resource.
ns & Shredders. Although some authors have as-
;t at- sumed that aquatic macrophytes are seldom con-
rious sumed by shredders before autumn senescence
Jring (e.g., Hutchinson 1981, Otto 1983), there is con-
jome siderable evidence that some consumers feed on
their many macrophytes throughout the growing sea-
obial son and it has been suggested that the emphasis
on et on macrophytes as only a source of detritus in
, see aquatic ecosystems is a simplification (Carpenter
licro- & Lodge 1986). Macrophytes in lentic (Carpen-
lionts ter & Lodge 1986, Lodge et al. 1985) and lotic
ested ecosystems (Wallace & O'Hop 1985) may be
ulose heavily grazed by consumers.
e de- Consumption of CPOM, or "shredding" by
foods invertebrates, increases the rate at which
:main CPOM is converted to FPOM (Cummins 1973,
tein, Petersen & Cummins 1974, Wallace et al.
nergy 1982b) and dissolved organic matter (DOM)
(Meyer & O'Hop 1983). FPOM is more ame-
nable to transport in the water column, which
may be of greater consequence than the shred-
ders' ability to degrade organic material di-
rectly by metabolic respiration, because FPOM
>les in egested as feces in upstream reaches is avail-
itrient able to downstream organisms.
Their Collector-Gatherers. Collector-gatherers are
i, per- adapted to feed primarily on FPOM that is de-
coarse posited on substrate surfaces. Despite their abun-
1 mm dance in freshwater habitats, the functional role
FPOM of collector-gatherers in streams has received little
s, other attention. Fisher & Gray (1983) provide an ex-
within cellent account of the role of collector-gatherers
in a Sonoran Desert stream (Arizona), where
t) have standing stock biomass of collectors was rather low
.ed for compared with their food resource (FPOM). The
: Graz- animals had high ingestion rates, consuming food
ickness equivalent to their own body weight every 4—6 h.
utrient Fisher & Gray suggest that egested feces (lower
ingest, food quality) were rapidly colonized by microbes,
icscent and on the average, were reingested every 2-3
sr rates d. Hence, rapid gut passage, low assimilation rates,
;)of the and reingestion of feces that were rapidly colo-
grazers nized by microbes, provided the energy base sup-
!i. Me- porting collector-gatherer production. However,
:rs may the extent to which collector-gatherers contribute
availa- to similar processes of FPOM turnover in cool,
ent ac- temperate streams is unknown.
In lake sediments, collectors may consume a
significant portion of the annual organic carbon
inputs that enter the system. For example, in
Mirror Lake, N.H., total zoobenthic production
was about 12% of the net input of organic carbon
inputs to the lake (Strayer & Likens 1986). In
shallow sediments of Mirror Lake, these authors
found that algae and detritus supported most
zoobenthos production, whereas in sediments of
deep waters, detritus was the main energy base.
Collector-Filterers. There is extensive lit-
erature on filter-feeding insects (see Wallace &
Merritt 1980), as well as other invertebrate fil-
ter-feeders (Jorgensen 1966). These constitute a
very heterogenous group with respect to trophic
level. Many of the Hydropsychidae and Brachy-
centridae (Trichoptera) in streams rely primarily
on animal drift (Wallace et al. 1977, Benke &
Wallace 1980, Georgian & Wallace 1981, Ross
& Wallace 1981, Parker & Voshell 1983). There-
fore, although these animals may be "filter feed-
ers" based on their mode of food capture, they
may also be carnivorous. Conversely, some taxa
such as the Philopotamidae (Wallace & Malas
1976), Simuliidae (Wotton 1977, Ross & Craig
1980), Chironomidae (Walsh 1950), and Sphae-
riidae (Pennak 1978), exploit extremely minute
particles suspended in the water column. Some
FPOM feeders, such as the Philopotamidae and
the Simuliidae, may actually increase particle
sizes by ingesting minute FPOM and egesting
compacted fecal particles larger than those orig-
inally consumed (Wallace & Malas 1976). Thus,
these animals may perform two functions; they
remove very fine particulate organic matter from
suspension, which may otherwise pass through
the stream segment, and defecate larger parti-
cles than consumed, which are then available to
a broad spectrum of larger particle-feeding de-
tritivores.
In lakes, excretion primarily by filter-feeding
zooplankton, which feed on planktonic algae (as
well as other particulate organic matter), rep-
resents an important source of nutrient regen-
eration in the water column. These released
nutrients are then used for additional algal growth
(Lehman 1980). Differences in size structure
among the herbivorous zooplankton community
can produce large differences in grazing and re-
cycling rates (see Carpenter et al. 1985).
Predators. Allan (1983) and Peckarsky (1984)
have reviewed a large body of literature asso-
ciated with predator-prey relationships in
streams. Although there is some experimental
evidence that predators can exert some influ-
 Miscellaneous Publications No. 75 Wallace: S

ence on the structure of lentic and intertidal to lentic and lotic ecosystems (see Hurlbert 1975, Insectic
communities, there is no strong evidence for R. R. Wallace & Hynes 1981); however, to my Streams.
predators exerting a significant influence on lotic knowledge, not one has examined changes in forested ca
communities (Allan 1983). Allan suggested sev- secondary productivity of invertebrates associ- in the fonr
eral reasons why benthic communities of streams ated with such additions. Secondary production autumn-sh
are not structured by predation. These include, corrects for differences in the turnover rates est (Fisher
the absence of a dominant predator, the pres- among animals and is more indicative of re- Webster et
ence of many refuges, cryptic coloration, and source use (e.g., Benke 1984). The reason is bris, comb
behavioral adaptations of prey. Very heteroge- obvious; the measurement of secondary produc- and relath
nous substrates, which may provide refuges for tivity is an expensive, time-consuming task. channels tl
benthic prey (Allan 1983), and the selection of One of the most overwhelming tasks associ- bris dams,
a broad spectrum of prey taxa by individual in- ated with studying the effects of pesticides in tion of th<
vertebrate predators would further dampen the ecosystems is the assessment of the indirect ef- streams is <
direct effects of predation in these small head- fects that result from the direct effects of pes- Naiman &
water streams. Knowledge of prey and predator ticides on the biota. A major problem with Wallace et
production, the relative availability of prey pro- pesticides and ecosystem studies is that pesti- shall et al.
duction to predators, and the bioenergetics of cides can produce a multitude of secondary, or materials c
predators are required before making any con- indirect, effects that are seldom documented. Thus, hea(
clusions about the influence of predation on lotic Hurlbert (1975) defined secondary effects of pes- transforma
communities. Assessments of predator—prey re- ticides as "changes that take place in an ecosys- and export
lationships based only on either numerical abun- tem following and as a result of direct pesticide ganic matt
dances or biomass are very tenuous. effects (primary) on the growth, survival, or re- What fac
In contrast to lotic environments, predation production of sensitive species." subsequen
in lentic waters has been shown to influence Hurlbert presented an excellent review of the headwater
community structure through alterations in both literature on secondary, or indirect, effects of that micro
community structure of vertebrate zooplankti- pesticides on ecosystems through the early 1970s. invertebra
vores and invertebrate zooplanktivores, and shifts However, I question his statement that "even Meyer & <
in zooplankton community structure, which in- the best-designed experiments on secondary ef- influence
fluences herbivory and standing crops and pro- fects of pesticides are of limited value and re- within he;
ductivity of algae (e.g., Carpenter et al. 1985). search efforts are more profitably focused on rect evide;
Such cascading trophic interactions may influ- primary effects." Drastic alterations of species a significar
ence energy flow and productivity at all trophic influence community structure and can severely cessing an<
levels as well as nutrient cycles within lentic alter functional group abundances, biomass, and Merritt et
ecosystems (Carpenter et al. 1985). production within ecosystems. The biota may a small ai
play important roles in the processing of energy tersheds v
Effects of Pesticides on Ecosystems and cycling of nutrients within aquatic ecosys- the main
tems. The influence of pesticides on overall eco- release. T
Direct Versus Indirect Effects. Direct (pri- system processes, such as detritus processing vertebrate
mary) effects of pesticides in an ecosystem are and nutrient cycling, may exceed that of direct headwater
considered to be those "that influence growth, effects on growth, survival, or reproduction of benthic in\
survival, or reproduction of organisms . . . ir- individual species, although such direct and in- of total cat(
respective of the time interval and ultimate ef- direct effects are not inseparable. Because all vertebrate.--
fect (e.g., immediate mortality on contact, egg functional groups do not respond similarily to several attr
mortality after years of gradual pesticide accu- pesticide treatments, changes in functional com- the ability t
mulation by parent birds, etc." (Hurlbert 1975). munity structure and production can drastically include tl
Obviously, such direct effects potentially can in- alter ecosystem-level functions such as energy ingestion r
fluence species diversity of ecosystems (Pimen- flow, food chains, nutrient cycles, and hetero- Anderson
tel & Edwards 1982), as well as primary and trophic "regulation" (cybernetics) within the low assimi
secondary productivity. Hence, such direct ef- ecosystem. Indirect effects of pesticides (or any tion/ingesti
fects can alter the structure and function of aquatic toxic substance) on processes such as processing etal. 1986
ecosystems. Although these direct effects are fairly organic matter and nutrient cycling deserve much have been
well documented in the literature, some are not. more ecosystem-level study (Levin et al. 1984, (Petersen
For example, numerous studies have docu- Pimentel & Edwards 1982). The following ex- & O'Hop
mented large-scale changes in community struc- amples document some of these effects on fresh- In 1980.
ture of invertebrates following pesticide additions water ecosystems. water stre
>ns No. 75 Wallace: Structure and Function of Freshwater Ecosystems

Ibert 1975, Insecticide Treatments on Headwater oratory in western North Carolina with the
ver, to my Streams. Small headwater streams that drain insecticide, methoxychlor, to reduce inverte-
changes in forested catchments receive most of their energy brate abundances (Wallace et al. 1982b, Cuffney
tes associ- in the form of allochthonous detritus, primarily et al. 1984). We addressed two primary ques-
production autumn-shed leaves, from the surrounding for- tions in this experiment: Would manipulation of
over rates est (Fisher & Likens 1973, Cummins et al. 1983, invertebrate abundances influence processing
:ive of re- Webster et al. 1983). Large inputs of woody de- rates of leaf litter? and Would manipulation of
• reason is bris, combined with shallow, narrow channels, invertebrate populations influence transport of
.ry produc- and relatively low discharge tend to result in FPOM to downstream areas?
ig task, channels that are characterized by numerous de- Treatment of the stream draining Watershed
.sks associ- bris dams. As a consequence, only a small frac- 53 resulted in several major changes in the stream
sticides in tion of the CPOM that annually enters such ecosystem. Initial treatment resulted in massive
ndirect ef- streams is exported as CPOM (Sedell et al. 1978, insect drift (>10,000 individuals/m3 of stream
cts of pes- Naiman & Sedell 1979, Webster & Patten 1979, discharge compared with pretreatment drift of
blem with Wallace et al. 1982a, Cummins et al. 1983, Min- <10 individuals/m3 discharge). The structure of
that pesti- shall et al. 1983, Cuffney et al. 1989). Exported the invertebrate community was shifted from
xmdary, or materials consist primarily of FPOM and DOM. domination by large insects to domination by
cumented. Thus, headwater streams are sites of retention, noninsects (primarily oligochaetes and cope-
ects of pes- transformation (CPOM to FPOM and DOM), pods) with few insects (primarily chironomids
an ecosys- and export (primarily FPOM and DOM) of or- and odonates) remaining. Treatment drastically
:t pesticide ganic matter. reduced insect shredders and resulted in sig-
•ival, or re- What factors influence the transformation and nificantly slower leaf processing rates compared
subsequent transport of organic matter within to those of the adjacent reference stream, Wa-
• view of the headwater streams? Laboratory studies suggest tershed 54 (Table 1). Consequently, concentra-
, effects "of that microbial flora (Suberkropp & Klug 1980), tion of FPOM and export to downstream reaches
sarly 1970s, invertebrates (e.g., Anderson & Sedell 1979, was severely reduced (Wallace et al. 1982b,
that "even Meyer & O'Hop 1983), and physical processing Cuffney et al. 1984).
condary ef- influence the transformation of organic matter Within two years (1982-1983) following the
lue and re- within headwater streams. However, little di- first treatment, there was no significant differ-
focused on rect evidence has been available that indicates ence in large insect shredder biomass (Wallace
; of species a significant role for invertebrates in CPOM pro- et al. 1986) or production (unpublished data) in
an severely cessing and FPOM generation in streams (e.g., litterbags (large meshed bags, >5-mm mesh
iomass, and Merritt et al. 1984). Although streams represent openings, used to confine known quantities of
biota may a small area compared with that of the wa- leaf litter) of the two streams. However, indi-
g of energy tersheds which they drain, they also represent vidual taxa of shredders were quite different
itic ecosys- the main route of organic matter and nutrient among streams. Despite differences in taxa con-
overall eco- release. The suggestion that benthic macroin- stituting the shredder guild, processing rates of
processing vertebrates can influence ecosystem functions in leaf litter had been largely restored in the treat-
nat of direct headwater streams may seem remote because ment stream (Table 1). Suspended concentration
reduction of benthic invertebrates represent a small portion of FPOM in the treatment stream also increased
irect and in- of total catchment biomass. However, macroin- from 4 to > 10 times compared with treatment
Because all vertebrates, in particular insect shredders have year levels (Wallace et al. 1986).
similarily to several attributes that suggest that they may have We have been repeating this experiment on
otional com- the ability to influence ecosystem function. These a much more intensive scale. From late 1984 to
i drastically include the following: shredders have high December 1985, we conducted a year of inten-
i as energy ingestion rates (e.g., McDiffett 1970, Grafius & sive pretreatment data on the above two streams
and hetero- Anderson 1979, Perry et al. 1986); they also have as well as a third stream draining Watershed 55
within the low assimilation efficiencies (A.E. = [assimila- at Coweeta. There are several reasons for re-
ides (or any tion/ingestion] * 100) (e.g. McDiffett 1970, Perry peating these ecosystem-level manipulations. In
s processing et al. 1986, Anderson & Cargill 1987); and they the first study, important data were not ob-
jserve much have been shown to comminute CPOM to FPOM tained, including measurements of input and
et al. 1984, (Petersen & Cummins 1974) and DOM (Meyer output of organic matter for these streams, con-
blowing ex- & O'Hop 1983) in laboratory studies. tinuous measurements of organic matter output
cts on fresh- In 1980, we treated one of two adjacent head- during storm and nonstorm periods, compari-
water streams at the Coweeta Hydrologic Lab- sons of storage of CPOM and FPOM in manip-

';*•.£?. jji&V-
J ^i'-j-WiWi-
...'*$!&$*&$••
10 Miscellaneous Publications No. 75 Wallace: ,'

Table 1. Influence of invertebrate reduction by insecticide treatment on leaf processing rates in streams draining Table 2. C
Watershed 53 (treated stream, TS) and Watershed 54 (reference stream, RS) in 1980-1981 (initial treatment year for 55 during tl
TS) and 1982-1983 (2nd-year recovery of TS)
1980-1981 1982-1983 Stream
Leaf Time for 95% loss Time for 95% loss
Species Stream (95% CI) (95% CI) WS53
VVS 54
Dogwood RS 144 (130-161) 95 (82-112) WS55
Dogwood TS 285 (246-340) 131 (118-147)
Red maple RS 217 (195-246) 212 (182-256)
Red maple TS 555 (468-681) 252 (209-315) WS53
White oak RS 283 (260-309) 285 (258-319) WS54
White oak TS 810 (697-966) 272 (244-309) WS55
Rhododendron RS 611 (517-749) 526 (410-731)
Rhododendron TS 2,304 (1,997-2,723) 348 (270-491) 1984-1985 =
of the insect
Data are recalculated from exponential breakdown rates (k, day ', ± 95% CI) of Wallace et al. (1986) to express processing
rates as time in days for 95% breakdown, ± 95% CI in days.
The res
the impo
ulated and unmanipulated streams, and large rivers. Obviously, the view that streams stream pi
measurements of microbial abundance and res- are more resilient to pesticides because the run- FPOM ge
piration. Finally, because no two streams are ning water flushes pesticide pollutants (e.g., Pi- level treai
identical, one is never certain if results of such mentel & Edwards 1982) should be tempered 1985 (Wa
manipulations will be similar. This study, as do for small, retentive streams, which have high drift, subs
most ecosystem-level studies, suffers from the substrate sediment/water ratios. structure,
problem of pseudoreplication, and, thus, differ- During and following treatment of Watershed processin;
ences cannot be attributed strictly to treatment 54 in December 1985, we observed changes sim- stream re
effects (Hurlbert 1984). ilar to those seen in the earlier treatment of Wa- reference
In December 1985, the former reference tershed 53 in 1980. These include: massive Cuffney e
stream used in the initial study, Watershed 54, invertebrate drift during initial treatment, and the studies d
was treated with methoxychlor and Watershed elimination of most invertebrates belonging to the vertebrat<
55 was used as a new reference stream. Despite shredder and collector-filterer functional groups particles tl
an application rate of 10 mg/liter methoxychlor (Wallace et al. 1989); and elimination of the shred- Material <
for 4 h, based on stream discharge, the maxi- der functional group from Watershed 54 has been not be lo
mum concentration in stream water at the flume accompanied by significantly reduced processing stream co
was only 124 |ig/liter, or 0.008% of the appli- rates of leaf litter in this stream compared with ported m;
cation rate. Within 31 h after the treatment ended, those of Watershed 53 (fifth and sixth year recov- to downst
the methoxychlor concentration had dropped to ery) and Watershed 55 (reference stream) (Cuftney Anderson
1.8 jig/liter in stream water. Only 1.6% of the et al. in press). No significant differences have Our ne
methoxychlor applied to the stream was ex- been observed for either bacterial abundances or influence
ported during treatment and for a 31-h period microbial respiration on leaf and wood detritus in downstrea
following treatment. In contrast to low concen- the treated stream (Watershed 54) compared to elemental
trations in stream water, residues in stream sed- those of Watershed 53 or Watershed 55 (Cuffriey gen in nea
iments of Watershed 54 averaged 2.8 (Jig et al. in press). Furthermore, concentrations of Golladay
methoxychlor/g dry wt of sediments in June 1986 FPOM and export to downstream reaches have cation). Tl
(following a second 2-h treatment of methoxy- been reduced compared to Watershed 53 and Wa- to downs-
chlor to the stream in March 1986) (Wallace et tershed 55 (Cufihey et al. in press). Table 2 shows manipulat
al. 1989). These results are in striking contrast only concentrations of FPOM in stream water, but addition t<
with those obtained for a large river (Fredeen total export of FPOM (discharge X concentration) vertebrate
et al. 1975), where concentrations 6.5 km down- to downstream reaches has also been greatly re- particular
stream from the point of injection of methoxy- duced in Watershed 54 compared with that of enhanced
chlor were half that of the upstream application Watershed 53 and Watershed 55 (Cufihey et al. Indirect
rate. Thus, the high retention and low export of in press). In summary, the ecosystem-level re- above ex;
methoxychlor from small headwater streams, such sponses following treatment of Watershed 54 were concernec
as those at Coweeta, is in striking contrast with similar to those observed 5-6 yr earlier for Wa- streams,
the high entrainment and low retention seen in tershed 53. grazer fo(

\>--.:vr..;;v-v* : '^'"*'-"'--~ :: Vi ? '--S' l '>^^5'w-. : :'- • " • • • •

^
 Wallace: Structure and Function of Freshwater Ecosystems 11

Table 2. Organic and inorganic seston concentrations (± 95% CI) measured at the flumes of Watersheds 53, 54, and
55 during the period of December 1984 through early May 1987
% Change
Stream 1984-1985 1985-1987 1985-1987
Organic seston (mg AFDM/liter)
WS 53 0.854 (±0.095) 0.844 (±0.179) -1.2
WS54 1.545 (±0.301) 0.270 (±0.180) -82.5
WS55 1.427 (±0.318) 1.179 (±0.286) -17.4
Inorganic seston (mg ASH/liter)
VVS53 0.328 (±0.069) 0.499 (±0.111) + 52.1
WS54 2. 673 (±1.020) 0.350 (±0.085) -81.9
WS55 1.837 (±0.520) 1.650 (±0.413) -10.2
1984-1985 = pretreatment of Watershed 54 and 1985-1987 = posttreatment period of Watershed 54 following application
of the insecticide, methoxychlor, in December 1985.

The results of these studies have established
the importance of invertebrates in headwater
stream processes such as CPOM processing,
FPOM generation and export. Both ecosystem-
l- level treatments in 1980 (Watershed 53) and in
d 1985 (Watershed 54) resulted in massive insect
h drift, subsequent changes in benthic community
structure, and significant reductions in leaf-litter
d processing rates and export of FPOM to down-
1- stream reaches compared to adjacent or nearby
l- reference streams (Wallace et al. 1982b, 1986;
e Cuffney et al. 1984, in press). Together, these
ie studies demonstrate an important role for in-
ie vertebrates in contributing to transport of small
is particles that are exported to downstream reaches.
1- Material exported from upstream reaches may
not be lost from the stream when the entire
stream continuum is considered because the ex-
ported material is an important source of energy
-Jl
V- to downstream fauna (e.g., Short & Maslin 1977,
Anderson & Sedell 1979, Wallace & Merritt 1980).
-y
/e Our next effort will be to concentrate on the
or influence of invertebrates on nutrient export to
in downstream reaches. A large portion of the total
to elemental transport of phosphorous and nitro-
ey gen in nearby Coweeta streams is particulate (S.
of Golladay and J. Webster, personal communi-
ve cation). The large differences in particle export
'a- to downstream areas in manipulated and un-
A'S manipulated streams certainly .suggest that, in
•ut addition to promoting organic matter export, in-
vertebrates also may promote nutrient export,
,n)
•e- particularly nitrogen and phosphorous, through
of enhanced particle export.
al. Indirect Effects on Grazer Food Chains. The
above example of indirect effects of pesticides
•e-
•re concerned the detritus food chain in headwater
/a- Streams. Similar indirect effects are known for
grazer food chains. For example, Yasuno et al.
(1982) treated a stream in Japan with the insec-
ticide temephos at the rate of 2 ing/liter in late
December, which resulted in massive insect drift
of most of the benthic insects, including algal-
grazing herbivores (scrapers). Removal of scrap-
ers was accompanied by large increases in algal
standing crops, as measured by the chlorophyll
a content of tiles placed in upstream (untreated)
and downstream (treated) segments (Fig. 2).
Furthermore, overall abundance of many spe-
cies of algae increased in treated regions com-
pared to untreated segments (Yasuno et al. 1982).
Standing crop biomass of algae decreased in the
treated segment as grazing herbivores recolon-
ized the treated reaches. The observed effects
D = UNTREATED AREA
250 -i
• = STATION 2 (TREATED)
STATION 3 (TREATED)
DATE
Fig. 2. Influence of temeplios on algal standing crop in a
headwater stream (data from Yasuno et al. 1982). Temephos
treatment in mid-December eliminated many herbivores
(scrapers), which resulted in large increases in algal standing
crops (as measured by chlorophyll a) in the treatment reaches
compared with the upstream untreated reach. By late win-
ter, herbivores were recolonizinK the treated reaches that
reduced algal standing stocks.
12 Miscellaneous Publications No. 75 Wallace: Struc

on algal standing crop are similar to those ob- elude reduced food resources for herbivores, al- Branch (EEB)
served when grazing insects were experimen- tered substrate availability for those animals that the EEB prog
tally excluded from tiles placed in a California rely on plant surfaces, reduction in overall sub- lication is ext
stream (Lamberti & Resh 1983). strate complexity that may influence predation, (p. 53) acknow
Hansen & Carton (1982) documented similar alterations in food chains with the addition of and terrestria
increases in algal biomass following application of plant detritus, increased nutrient concentrations problems cen
a pesticide that eliminated herbivores in labora- in the water column that can stimulate plankton effects of pesti
tory streams. However, they noted that a decrease populations, and dissolved oxygen depletion and are what mak
in algal diversity accompanied the increase in bio- increases in concentration of free carbon dioxide ficult at the e<
mass, which was attributed to increased algal com- as a result of increases in decomposing plant system appro
petition for limited attachment sites. Hansen & detritus (see Brooker & Edwards 1975, Hurlbert streams, cann
Carton also report that it would not be possible 1975). The reduction in biomass and abundances facing registr;
to forecast changes accurately in algal biomass from of herbivorous zooplankton following application and ethical dr
laboratory toxicity tests without prior knowledge of atrazine appears to be related indirectly to ties in such ;
that the primary production was herbivore limited reduction of the zooplankter's food resource, never sure ho-
and that release from this herbivore control would phytoplankton (deNoyelles et al. 1982). Studies one pond, laki
produce competition for space. of the effects of atrazine in small ponds have system, and, t
Changes in algal biomass following pesticide shown that abundances of nonpredatory insects ofpseudorepli
applications are not limited to streams. For ex- were more severely influenced than predatory these problem
ample, elimination of herbivorous arthropods in species (Dewey 1986). Furthermore, the lowest by adequate j
ponds following treatment with a synthetic pyr- concentration at which atrazine affected aquatic comparisons ^
ethroid, cypermethrin, has also been observed insects was an order of magnitude below that at ecosystems,
to result in increased biomass of filamentous al- which direct toxic effects previously were mea- treatments an.
gae (Grassland 1982). A dense algal mat devel- sured in the laboratory. Dewey (1986) suggested expensive for i
oped over the surface of the treated ponds, which that the main effect of atrazine may have been where the foe
effectively limited light penetration and photo- through indirect mechanisms which influence and outputs.
synthesis in the underlying water column. This food availability to herbivores, or altered insect Several of
led to a serious depletion of dissolved oxygen in habitats (macrophytes). Lugthart (1987b) re- symposium ad
the water of the treatment ponds. ported that the effects of the herbicide, diquat, methods assoc
Grazers in lotic (e.g., Newbold et al. 1982) on invertebrate populations appeared to be in- desirability of
and lentic ecosystems (e.g., Lehman 1980, Car- direct, resulting from the destruction of macro- problems asso
penter et al. 1985) may contribute significantly phyte habitat, rather than direct, toxic effects. expense, and e
to nutrient cycling. Thus, any disturbance that whole-system
influences abundances, biomass, or production Conclusions routine testin;
of grazers may influence nutrient cycling in ecologists (e.g
freshwater systems. Furthermore, the exclusion When this symposium was organized, I was nents of meso
or reduction of grazers in streams may contrib- asked to address the following question: "Is our However, s
ute to the development of dense growths of fi- knowledge of structure and function of aquatic mesocosms sh
lamentous algae, which retain and retard the ecosystems sufficiently developed to draw the following:
downstream transport of FPOM. meaningful conclusions from laboratory and me- proach the nat
Other Indirect Effects. Other indirect ef- socosm toxicological tests?" My answer is yes in streams? Ar
fects of insecticides in freshwater ecosystems in- and no. Certainly, in the past two decades a large littoral enclosu
clude those which influence predator—prey amount of information has been gleaned from vironments? Is
relationships among the animal community. For ecosystem approaches, but many questions re- be directly app
example, elimination of some predaceous taxa main. In some situations, the use of pesticides the problem o
of Plecoptera and Trichoptera, following treat- may help ecologists address some key questions such as mesc
ment with DDT, appears to have resulted in a about various ecosystem processes and the role enough to ac
resurgence of larval black flies in some cases (see of animals in those processes, as Moore (1967) ranges that ex
Hurlbert 1975, R. R. Wallace & Hynes 1981). originally suggested. addition, do
Elimination or reduction of prey may also shift The U.S. Environmental Protection Agency's rionsofthe ea
diets of invertebrate predators (Wallace et al. publication on ecological risk assessment, which resent, raise p
1987), salamanders (Lugthart 1987a), and fish is supposed to represent "state-of-the-art tech- as currents an
(Hurlbert 1975, R. R. Wallace & Hynes 1981). niques for ecological risk assessment!,]. • -pre- I am extren
Herbicides may also produce a multitude of sents procedures, philosophies, and extrapolative suits from first-
indirect effects on aquatic ecosystems. These in- techniques as used in the Ecological Effects second- and tli
 No. 75 Wallace: Structure and Function of Freshwater Ecosystems 13

ores, al- Branch (EEB)" (Urban & Cook 1986). Although eeta Basin. There are too many differences—
nals that the EEB program that is described in this pub- stream discharge regimes, inputs and retention
rail sub- lication is extremely detailed, Urban & Cook of organic matter, and localized geomorphology,
edation, (p. 53) acknowledge many weaknesses for aquatic which influences depositional versus erosional
lition of and terrestrial risk assessment. Most of these habitats, as well as functional feeding groups and
itrations problems center around assessing the indirect organic matter retention (e.g., Huryn & Wallace
plankton effects of pesticides. Indirect effects of pesticides 1987)—to make such comparisons. Further-
:tion and are what make predictions of their impact dif- more, any attempt to extrapolate results from
i dioxide ficult at the ecosystem level. Obviously, whole first-order Coweeta streams to similar streams
ng plant system approaches, that is ponds, lakes, or in other geographical regions, e.g., Rocky
Hurlbert streams, cannot be required for every pesticide Mountain, boreal forest, or prairie, would be
mdances facing registration. There are severe financial futile. Do the same problems exist when results
plication and ethical drawbacks, and scientific uncertain- are extrapolated from mesocosms, which "mimic"
rectly to ties in such approaches. For example, one is lentic conditions, to ponds and lakes that differ
•esource, never sure how applicable results obtained from greatly in their basin geomorphology, nutrient
. Studies one pond, lake, or stream are to another similar inputs, flora, fauna and primary and secondary
ids have system, and, therefore, suffer from the problem productivity? These are disturbing questions,
y insects of pseudoreplication (Hurlbert 1984). However, which are not easily answered; yet, they are im-
>redatory these problems can be improved to some extent portant questions.
ic lowest by adequate pretreatment data, and treatment Laboratory microcosms, mesocosms, and eco-
d aquatic comparisons with nearby reference ("control") system-level studies are all important in solving
.w that at ecosystems. True replication (i.e., multiple problems associated with toxicants. Single- and
ere mea- treatments and controls) would be prohibitively multiple-species microcosms are of tremendous
uggested expensive for most ecosystem studies, especially value in providing information on toxicity to in-
ave been where the focus is on inputs, biota, processes, dividual species, toxicant transfer, storage, and
influence and outputs. detoxification. This information allows those
ed insect Several of the subsequent chapters in this working with mesocosms to focus on certain pop-
)87b) re- symposium address the potential, problems, and ulations, functional groups, or ecosystem-level
j, diquat, methods associated with mesocosms. Despite the processes such as detritus processing or mineral
to be in- desirability of whole-system approaches and the cycling, which may be influenced by a particular
of macro- problems associated with mesocosms, the labor, functional group.
c effects. expense, and ethical considerations involved with Mesocosms can provide valuable links be-
whole-system studies make them unsuitable for tween single- or multiple-species' effects and
routine testing of insecticides. Indeed, several ecosystem-level effects. For example, does a tox-
ecologists (e.g., Odum 1984) have been propo- icant produce changes in community structure
ed, I was nents of mesocosm studies. or functional group composition that can influ-
n: "Is our However, several disturbing questions about ence energy flow or nutrient cycling? Does a
of aquatic mesocosms should be addressed. These include toxicant eliminate predatory taxa, which may re-
i to draw the following: Are there any mesocosms that ap- sult in increased abundances and productivity
ry and me- proach die natural variablility or conditions found of certain prey? Do these prey possess the ability
wer is yes in streams? Are results derived from ponds and to alter ecosystem-level processes that influence
ides a large littoral enclosures directly applicable to lotic en- primary and secondary productivity, decompo-
aned from vironments? Is there any reason why they should sition of organic matter, and mineral cycling in
estions re- be directly applicable? Neuhold (1986) also raised a particular ecosystem (e.g., Oberndorfer et al.
pesticides the problem of scale: Are experimental systems 1984)? Mesocosms have the potential of framing
questions such as mesocosms and "limnocorals" large questions to address many exciting ecological
d the role enough to accomodate those organisms with problems and, thus, can provide an important
ore (1967) ranges that exceed the limits of mesocosms? In framework for ecological studies. However, based
addition, do "limnocorals", or penned off por- on points raised above, mesocosms cannot be
i Agency's tions of the ecosystems they are intended to rep- expected to duplicate the range of conditions
3nt, which resent, raise problems with physical factors such found in natural ecosystems. Indeed, one could
3-art tech- as currents and temperature? argue strongly that no one ecosystem is com-
] . . . pre- I am extremely leary of extrapolating our re- pletely similar to another.
irapolative sults from first-order streams at Coweeta to larger Many scientists (e.g., Kimball & Levin 1984,
:al Effects second- and third-order streams within the Cow- Levin et al. 1984, Cairns 1986) have recognized
14 Miscellaneous Publications No. 75 Wallace: <

and advocated mesocosm and ecosystem-level propriate situations, pesticides can be very
testing of pesticides. I concur with the need for valuable in assessing the influence of flora and
ecosystem-level testing; however, at the same fauna on ecosystem processes. Wetzel (1983, T. F. »
time we need to ask the following question: How 752), Likens (1985), Carpenter & Lodge (1986) Gretchen
applicable are the results obtained from one have emphasized the need for ecosystem ex- assisted vv
freshwater system to another? Too few compar- periments that compare manipulated and ref- D. Huryn
ative studies of aquatic ecosystems exist to make erence (or "control") ecosystems. Two decades icism on e
broad sweeping generalizations between various ago, Moore (1967) suggested that in some cases of the resi
regions, which differ in climate, geomorphol- pesticides can serve as useful tools in ecolog- ported by
ogy, basin morphometry, nutrients, flow re- ical research. Pesticides could be used to "dis- BSR-87-U
gimes, community structure, primary and sect" ecosystems to study direct and indirect
secondary production, etc. More comparative effects and could be of great value in studying
studies of freshwater ecosystems are sorely organisms and processes that influence energy
needed. flow and nutrient cycling within ecosystems.
It may be unfair to expect the chemical in- This has been true for a limited number of Allan, }. D.
dustry to invest large sums of money into eco- aquatic and terrestrial studies. pp.1!
system-level testing, because the results so Strea
Ecosystem manipulations can contribute sub- ecolo
obtained may have little applicability to other stantially to our knowledge of ecosystem struc- Anderson, >
untested ecosystems. However, the chemical ture and function; however, the researcher must ogy o
industry also has an obligation to the environ- have a good working knowledge of the particular F. SI;
ment, society, and government to minimize ecosystem before such treatments. The ap- ecolo
tebra
environmental disturbances. This obligation proach must not be to simply apply pesticides Anderson, >
requires efforts that extend beyond the profit and "see what develops." In addition to studying by m
motive. To foster a more productive relation- the movement, fate, and detoxification of pes- Rev.
ship between ecologists and the chemical in- ticides in aquatic ecosystems, prior testing, or Arsuffi, T.
dustry, it seems necessary that the chemical knowledge of a particular ecosystem is strongly streai
by di
industry be responsive and cooperative in advised to delineate the following: What taxa are Soc.
working toward the longer term needs of the present and may be potentially affected? What Barnthouse,
environment, society, and government. Con- is the major energy base of the ecosystem? What Suter
versely, ecologists must recognize that until are the potential effects on the microbial com- in eo
Posto
suitable alternatives are found, chemical pest munity? How and to what extent, either directly envir
control is an integral part of the general eco- or indirectly, can functional feeding groups of Amer
nomic well-being of society and accept that the animal community be altered? Are the func- delpl
some environmental degradation is inevitable tional feeding groups that use the major energy Benke, A. C
based on current pest control methods. sects,
base going to be severely influenced, either di- berg
Together, ecologists, toxicologists, and the rectly or indirectly, by treatment?; If so, the New
chemical industry should work in concert to study researchers should formulate predictions and Benlce, A. C
the impact of pesticides on ecosystem structure design experiments that key in on ecosystem prodi
and function, develop pesticides that minimize processes, such as energy flow (including pri- ern A
Brooker, M.
direct and indirect effects on communities and mary production), food chain effects, and nu- bicide
ecosystems, and promote the mutual exchange trient cycling. What are some of the potential 9: 1-1
of ideas and developments. Ideally, ecologists indirect effects of the above items on community Cairns, }., }\
can provide new ideas and approaches to toxi- structure, ecosystem structure and function? and dictioi
cologists, and, vice versa, toxicologists should What are the relative short- and long-term ef- biolog
Carpenter, '
be able to stimulate ecological thinking. For ex- fects of these items on the ecosystem that is mers<
ample, the stream studies described under "Di- proposed for study, and what factors influence Bot. :
rect and Indirect Effects," above were not fluctuations in similar unmanipulated systems? Carpenter,
designed to test the impact of an insecticide on Obviously, if the answers to all of these ques- Case:
BioSc
headwater streams but rather, to use an insec- tions were known beforehand, there would be Crossland, *
ticide as a "tool" to examine the role of inver- no need to conduct the research. However, sim- rin Ij
tebrates in ecosystem-level processes of ply thinking about such questions will alert one ment
headwater streams. to expect the unsuspected before any system- CufTney, T.
Pesti.
Certainly, widespread use of pesticides to level manipulation with pesticides and to design verte
manipulate flora or fauna of large-scale eco- experiments that can cope with a multitude of system
systems cannot be condoned; however, in ap- direct and indirect effects. 153-
Wallace: Structure and Function of Freshwater Ecosystems 15

Acknowledgment Cuffney, T. F., J. B. Wallace & G. J. Lugthart. In press.

Experimental evidence of quantifying the role of
benthic invertebrates in organic matter dynamics of
T. F. Cuffney, A. Lingle, John Lugthart, headwater streams. Freshwater Biol.
Gretchen Schurr, and Francie Smith-Cuffney Cummins, K. W. 1973. Trophic relations of aquatic in-
sects. Annu. Rev. Entomol. 18: 183-206.
assisted with various aspects of the project. A. Cummins, K. W. & M. J. Klug. 1979. Feeding ecology
D. Huryn and J. Lugthart provided helpful crit- of stream invertebrates. Annu. Rev. Ecol. Syst. 10:
icism on early drafts of the manuscript. Portions 147-172.
of the results reported in this article were sup- Cummins, K. W., J. R. Sedell, F. J. Swanson, G. W.
ported by NSF Grants BSR-8316082 and NSF Minshall, S. C. Fisher, C. E. Gushing, R. C. Pe-
ter sen fie R. L. Vannote. 1983. Organic matter
BSR-87-18005. budgets for stream ecosystems: problems with their
evaluation, pp. 299-353. In J. R. Barnes & G. W.
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