Limnologica33, 163-189 (2003)

© Urban& FischerVerlag
http://www.urbanfischer.deljournals/limno LIMNOLOGICA
Defensive reactions of freshwater ecosystems
against external influences
Wilfried Sch6nborn 1.

Friedrich-Schiller-University Jena, Institute of Ecology, Workgroup Limnology, Jena, Germany

Received October 29, 2002 • Accepted July 2, 2003

Abstract
Eutrophication and toxic loading of freshwater occurred even in early geological epochs as a
result of natural factors (e.g., large animals, volcanism), and nutrients and xenobiotics are
more quickly integrated in material cycling in aquatic than in terrestrial systems. Therefore,
aquatic ecosystems show many defensive mechanisms against organic and toxic loading.
Many other defensive reactions can be described in addition to the well-known example of
microbial self-purification.
Freshwater ecosystems possess compartments which cooperate towards the function and
protection of the whole system but, in opposition to these "euoecisms", there are also
"dysoecisms". The defensive reactions of an ecosystem are founded largely on species-ego-
istic adaptations that have an (accidental) system-altruistic effect. The whole ecosystem re-
acts only seldom, and it is not clear whether there are selection processes which favour water
bodies with a slow eutrophication and therefore slow silting-up, because the freshwaters are
important for the global water balance.
It is possible to compare organismic with ecosystemic defensive reactions but the origin of
both reactions is very different.

Key words: Freshwater ecosystems - defensive reactions - eutrophication - natural loadings

- euoecisms and dysoecisms

1. Introduction tems. Therefore, it was especially important for limnetic

ecosystems to develop defensive reactions against addi-
Aquatic ecosystems can be influenced by the input of tional load. In contrast to waters, heavy metal resistant
foreign substances - both degradable organic and inor- bacteria are absent in soils, because their contact with
ganic toxic substances - and by the immigration of for- heavy metals is relatively weak.
eign species, and they react more sensitively to these As a result of this additional high load of externally
loads than terrestrial systems. Whereas in terrestrial supplied nutrients, accelerating eutrophication occurs,
ecosystems, especially in soils, most of the nutrients and and aquatic systems can show signs of "diseases". The
dangerous substances are bound to, and stored on, soil main part of this external load is caused by the input of
particles, in aquatic ecosystems nutrients and xeno- degradable organic substances. However, the final prod-
biotics are diluted and suspended. In this way, they enter ucts of decomposition - especially nitrogen and phos-
the element cycling more quickly than in terrestrial sys- phorus - are also used by plants as nutrients. Nitrogen

*Corresponding author: Wilfried Sch6nborn, Friedrich-Schiller-University Jena, Institute of Ecology, Workgroup Limnology, Carl-Zeiss-
Promenade 10, D- 07745 Jena, Germany

0075-9511/03/33/03-163 $15.00/0 Limnologica (2003) 33, 163-189

164 " W. Sch6nborn

and phosphorus sources can alternatively contaminate blocked, resulting in the destabilization of the whole
water bodies directly from anthropogenic sources, e.g., system (RosENzwE~6 1971).
sewages and agrochemicals. This means of eutrophica- We can conclude that - together with its nutritional
tion has become highly important since human interfer- aspects - eutrophication is a real symptom of illness in a
ence with nature began, and is the result of urbanization limnetic ecosystem. Therefore, in limnetic systems, dif-
and of industrial and agricultural development. ferent defensive mechanisms are necessary against or-
In contrast to that of degradable organics, the role of ganic and toxic loads. Just as individuals react to stress
inorganic and toxic substances, either as organic or inor- situations and symptoms of diseases, ecosystems have
ganic compounds, is difficult to judge. The influence of also developed specific defence mechanisms, but these
these substances to ecosystems is very different, and it are frequently misunderstood.
can be supposed that such substances already affected The aim of the present paper is to give - after an intro-
aquatic environments before human interferences with duction to natural eutrophication - an overview of the
nature took place. different defence strategies, from the well-known self-
Of decisive importance for the existence of aquatic purification to the features and structures of ecosystem
ecosystems, however, is the process of eutrophication. compartments that cooperate towards the function and
Because ecosystems are complex systems - not merely a protection of the whole system.
group of individuals - it is not possible to equate exactly Another emphasis of this paper is the invasion of
nutrient-driven eutrophication with the feeding or over- water bodies by foreign species (neophyta and neozoa)
feeding of individuals. All individuals, whether they re- which may have occurred since freshwaters are existing.
ceive too little or too much food remain identifyable as These species are frequently introduced for anthro-
individuals until they die, and even then we can often pogenic purposes or promoted by anthropogenic im-
find their remains. It is otherwise with aquatic ecosys- pacts. The role these organisms play in limnetic ecosys-
tems since, with increasing eutrophication, they loose tems and also the systems' reactions to these interven-
their identity by changing trophic state. The changes re- tions will be described.
sult in a completely different ecosystem. In principal, we
can say that each stage of eutrophication results in the
"death" of the former system, but, unlike with individu- 2. The natural load of aquatic systems
als, we never find "corpses" - because a new ecosystem
develops instead. Allochthonous eutrophication and toxic loads have ex-
The process of accelerating eutrophication leads to isted ever since water has been available on Earth; even
the aging of lakes or streams, passing through several without anthropogenic impact these waters were sub-
ecosystem stages - one after the other - until the dissolu- jected to external stress situations. At the latest in the
tion of the final system. This means that the basin or the Mesozoic, with the advent of the first large land ani-
channel previously containing the ecosystem completely mals, polysaprobic waters have existed. High toxic
disappears. loads also occurred in connection with vulcanic activi-
In the following, some details of these processes are ties (i.e. acidification, high concentrations of heavy
given. Extreme eutrophication limits aquatic ecosystem metals). Human disturbance of water bodies also has a
functioning. If the hypertrophic state is exceeded (nitro- long history, however namely since the first permanent
gen >30 mg 1-1; phosphorus >3 mg 1-I) transparency settlements.
(Secchi depth) decreases and as a result of this light limi- One of the main factors contributing to the natural al-
tation aquatic plants cannot use the high nutrient concen- lochthonous eutrophication of waters is large land ani-
tration. Biodiversity consequently declines and benthic mals. Today, tropical lakes and rivers are eutrophied by
producers can disappear completly. Because the phyto- buffalo, elephant and antelope. The result of their occu-
plankton shades itself, its photosynthetic activity is re- pation of water bodies in tropical regions is an enor-
duced and, in consequence, the concentration of oxygen mous mass development of algae and macrophytes.
shows aperiodic oscillation; a high production of oxygen Also carnivores, for instance the tiger, frequently hunt
(_+20 mg 1-1 d 1 O2) is accompanied by high respiration their prey in water and crocodiles pull big game (graz-
rate (+30 mg 1-1 d-1 02) (U~tLMANN1966; KALBE1996). ers) into the water and tear them to pieces there. In addi-
Another characteristic of the instability of a hyper- tion, wallowing by big game and disturbance of bottom
trophic aquatic system is the extreme oscillation in the sediment by crocodiles severely clouds the water. These
density of planktonic algae (BARICAet al. 1992). The processes are accompanied on the one hand by a remo-
mass development of planktonic algae also influences bilization of nutrients from the sediments, in the direc-
the feeding behaviour of the zooplankton, first reducing tion of accelerating eutrophication, but on the other
the efficiency of filter feeders and then the rate of preda- hand, the growth of plants - and therefore also the
tion (BURNS 1968). Energy transfer may then be results of the eutrophication - will be inhibited.

Limnologica (2003) 33, 163-189

 Defensive reactions of freshwater ecosystems 165

This coupling of loading and relieving processes can In the Jurassic many grant dinosaurs also used waters.
be found frequently in ecosystems. One example is the Brontosaurus (with a length of 18 m) lived both on land
activities of buffalo. These big game eat land plants and and also in water, where it fed on aquatic plants as did
may deposit their excrements in water, but in contrast to Diplodocus (length 27 m). Another saurian, the 22 m
these eutrophication-activities they also eat great long and 11 m tall Brachiosaurus, used deep lakes and
amounts of aquatic macrophytes and then leave the fed upon benthic plants (Fig. 1). In the Upper Cretaceous
water, thus reducing eutrophication. Another example period the enormous Protoceratopsgrazed both on land
from tropical waters are the hippopotami. The bulls and water plants.
mark their territories by fanning large amounts of excre- Very important for water bodies during the Creta-
ment and often they prefere the banks, especially of ceous period were also the world-wide occurring
bights for this purpose. This leads to considerable fertil- Hadrosaur (length up to 11 m) which have had webs on
izing. By grazing aquatic plants hippopotami could the feet and a duck-like beak. They were plant feeding
counteract the eutrophication but they very often eat animals living close to water.
land plants. These big reptiles undoubtedly had the same effects
Waterfowl are another cause of eutrophication; not as big mammals. They would cause accelerating eu-
birds that live permanently or only in the breeding time trophication if they deposited their excrement into water.
on water and feed on aquatic organisms; but to high con- But simultaneously, they may have counteracted this nu-
centrations of waterfowl in autumn, winter and spring tritional load by grazing aquatic plants - one of these
which cause an immense nutrient supply. Geese, in par- sites has predominated and therefore decided on loading
ticular, contribute to accelerating eutrophication when or relief.
they eat land plants during the day but over-night on In the Tertiary, the explosive radiation of the Mam-
water. malia coincided with an extremly high density of big
The highest concentration of nitrogen is found in the mammals which in some cases populated lake shores in
excrement of grey herons, the smallest in the excrement such numbers that the lakes became hypertrophic. One
of ducks (including swans) and cormorants. Phosphorus example is the well-known Messel quarry, near Darm-
is concentrated in the excrement of grey herons and cor- stadt, Germany, where in the Eocene a permanent algae
morants, less in such of ducks. At present 36 species of bloom occurred. This resulted in the formation of oil
aquatic, fungi are known to grow on avian excrement and shale, a material which favours fossilization. The hyper-
mineralize them (CZECZUGA~; MAZALSKA2000). trophic Lake Messel produced methane in such high
Natural eutrophication must have been enormous also concentrations that it killed many bats flying over the
in former geological periods. In the Upper Carbonifer- lake that then fell in and were fossilized.
ous, already Edaphosaurus was living. In the Permian, Fossiliferous deposits are also known from other Ter-
Moschops, a big reptile fed on plants in desert rivers. tiary lakes. The fossils show for instance, that European

I"L;.
,, I.\

{,\ IJ'TL -:D .....,A! Fig. 1. The jurassic Brachiosaurus

grazing waterplants in a lake,
(Half-schematic drawing, in the
style of a pattern from the Czech
AcademicArtists, Z. BURIAN).

Limnologica (2003) 33,163-189

166 W. Sch6nborn

water bodies were then populated by hippopotami. Also fluences. Where nowadays are inland waters where big
tapirs, Mastodonts, the gigantic and monstrous hoofed animals are killed by high concentrations of methane or
animals of the genus Uintatherium and the Deinotheria - where animals coming to drink stick to oil shale?
animals similar to the elephants - brought a high nutrient The palaeontological examples of natural eutrophica-
load to the waters of the Tertiary swamp forests. All of tion given above should lead to a reorientation of the
these animals fed on submersed plants. field of limnolgy. The examples make obvious the role
Likewise, the sediments of shallow lakes from the of great Tetrapoda and birds as sources of a high envi-
Pleistocene (about 200.000 years ago) indicate strong ronmental load during the Earth's history.
eutrophication. In these deposits fallow-deer skeletons Because of these loads it was necessary for aquatic
were found, their backwards bent cervical vertebrae ecosystems to develop defence mechanisms comparable
pointing to an abnormal cause of death. The high with the reaction of individuals to diseases. Because, as
amounts of methane produced by hypertrophic lakes shown above, the natural loads persist for extended geo-
could also have been responsible in this Case (MANIA logical periods, the waters were well prepared for a new
1992). There is also a Pleistocene lake in California enormous challenge - the disturbances caused by human
which has been thoroughly profoundly investigated by impacts. Without the possibility of developing these de-
palaeontologists. Its high shores show thick layers of oil fence reactions through such long periods of time, the
shale containing the fossils of large mammals. When waters would presumably not have survived the anthro-
these animals came to drink they stuck in the oil shale pogenic loads.
where they were often attacked by carnivores, for in- In the following an overview about the different de-
stance by the big cat Smilodon. fence mechanisms of aquatic ecosystems is given.
But big animals were not only important for eutrophi-
cation in former periods. By their wallowing activities
they also changed the morphology of waters, especially 3. Defensive mechanisms
of streams and rivers, which influences of course also
the trophic state of the waters and also produces new 3.1. Microbial self-purification
pools. The dinosaurians uprooted great woodlands by
their long and strong tails, and this process was contin- The best investigated defensive reaction against eu-
ued in later periods by large mammals, using - as do ele- trophication is the well-known microbial self-purifica-
phants - their trunks and tusks. This uprooting thus de- tion. Primarily, microbial decomposition was generally
stroyed the nutrient-rich vegetation along lakes and the basis for a continuous evolution of organisms.
rivers with its high diversity of plant species and coinci- Without microbial decomposition there would have
dentely influenced the water. been an accumulation of organic remains on the Earth
The nutrient load introduced by heavy rain should which would have prevented any existence of organ-
also not be forgotten. In combination with erosion in the isms. This mechanism has been retained for the self-pu-
areas denuded of vegetation by animals rain could have rification of all habitats. The majority of microbial de-
been of decisive importance. composers does not live primarily in water bodies, but
All these examples show that high nutrient load of come into rivers and lakes together with allochthonous
aquatic ecosystems has occurred during the whole bio- matter or dead organisms. Therefore, microbial self-pu-
logical development of the Earth. Gigantic natural eu- rification may be only compared with reservation with
trophication processes led also in times past to the immune system of organisms (compare chapter
polysaprobic conditions. These conditions were outside 7.6.).
the range of the currently used limno-saprobity index The bacteria were one of the first groups of organisms
which would have to be expanded to include eu-saprobic on Earth and - despite the process of evolution - they
conditions, comparable to wastewater-saprobity. are also at present integrated in all ecosystems, and
One great ecological problem of our day - cattle graz- many of them inherited the function of decomposers of
ing, which affects water as well, especially rivers - is organic matter. This function, so to speak, was offered to
harmless in comparison to the eutrophication processes different ecosystems, which were thus not required to
of former times. In the inter-glacial periods, in present develop these systems themselves. This process was
day central Germany, enormous large mammals lived probably supported by selection, because without sapro-
and polluted the water. Only as a result of human activi- phytic bacteria all forms of higher life on earth are im-
ty, for instance the hunting of these animals, did the possible.
water quality probably improve (MANIA 1995). But microbial self-purification can not be regarded as
It can be concluded that natural eutrophication of wa- an universal remedy. The final products of protein de-
ters in the course of the Earth's history has greatly sur- composition are plant nutrients and therefore a potential
passed the eutrophication caused by anthropogenic in- of eutrophication. At a permanent load the microbial

Limnologica (2003) 33, 163-189

 Defensive reactions of freshwater ecosystems 167

self-purification leads only to a mitigation of the eu- Cladophora-clumps were stored as detritus, so stop-
trophication processes in water bodies. In standing ping further decomposition (ScI~(SNBORN 1997). After
waters with a longer water residence time, nutrients can the total loss of Cladophora and reduction in the
be accumulated even at short-term loading in sediments discharge, the mass production of algae immediately
and algae and remain in this way available. resumed.
Other algae like Ulothrix and Oedogonium also show
3.2. Drift and throw-off of load the same drift mechanism, but it seems not to be as ef-
fective as in Cladophora. The diatom Melosira also de-
Apart from microbial self-purification, limnic ecosys- velops long, fragile filaments in brooks, covering stones
tems possess also other defensive mechanisms. Algae, and sediments at a high density. They break up with even
higher plants and, to a low degree, microorganisms as the slightest increase in the discharge, and are then car-
well can accumulate great quantities of nutrients and ried away. This mechanism also contributes to removing
transform them into biomass. Water bodies possess load from water bodies.
mechanisms to remove this biomass or a great part of it Among microorganisms, the polysaprobic filamen-
from the whole system or from a compartment of it. In tous bacteria Sphaerotilus, is involved in drift-events. It
running waters this mechanism is drift and in standing develops into clumps, somewhat more than a hand in
waters it is the throw-off of load by floatation ashore length, which become detached and show the well-
(flotsam). known fungal drift. Similar behaviour was registered in
the aquatic fungi Leptomitus and Fusarium. Of
• Drift Sphaerotilus it is also known that it can accumulate
In the drift of biomass from running waters the green heavy metals. Therefore, the micro-organisms given
alga Cladophora plays an important role. This algae oc- above affect unloading in a threefold way: by decompo-
curs even in oligosaprobic stream reaches, but with in- sition, by the drift of biomass and by the drift of xenobi-
creasing load mass production occurs. This mass pro- otics.
duction is caused by the high nutrient accumulation; the In running waters, small portions of Cladophora and
accumulation factor for N and P amounts to 5000 and Sphaerotilus may be continously removed, not only dur-
20 000 respectively. When eutrophication is advanced, ing high discharge, but also during low water, demon-
Cladophora clumps can grow up to 4 m within a few strating that biomass has a relatively short residence
days (ScHONBORN 1996). Nearly 16% of its fresh weight time in running water.
consist of seston (detritus), filtered by the filaments. For
optimal growth Cladophora needs a current velocity of • Throw-off of load (floatation ashore)
at least 0.5 m s-1. Clumps above 1 m length detach and The load throw-off by floating ashore (flotsam) is relat-
drift downstream if the current velocity exeeds 1 m s-1. A ed to the above described drift mechanisms, but occurs
reach may thus be freed from high amounts of biomass, in standing water. In eutrophic standing water, algae de-
but also from nutrients and xenobiotics, e.g. heavy met- tach from the sediment and float to the surface, where
als (ScHONBORN 1995). Depending on the roughness of they continue to grow and develop into algal mats. The
the stream bed the drifting Cladophora-clumps can be mats are drifted landwards by the wind and some will be
transported about different distances. Rough stream washed ashore. Important algae with this mechanism are
beds may catch and detain the drifting clumps thus hin- Cladophora fracta, Oedogonium, Spirogyra, Zygnema,
dering the drift mechanism, but with smooth beds the Mougeotia, Rhizoclonium and the blue-green alga
algae can reach the sea. Microspora. Cladophora fracta can also participate
As an example for the importance of the drift some in the purification process in running water, whereas
values should be given for the River Ilm, a low moun- Cladophora glomerata occurs only in running waters
tain range stream in Thuringia (Germany). The stretch and in the surf-zone of lakes.
investigated was a headwater with pebbles and stones. Important for the growth of algae on sediments are
The first 120 m of the stretch were unshaded and in this the light conditions at the bottom of the water. Light and
part 1.5 tons Cladophora-biomass were produced in a an increasing supply of nutrients lead to mass develop-
week. This weight even increased to 1.74 tons within ment of algae. But when algae-aufwuchs on sediments
one month by filtering seston. But after a high dis- reaches a certain density its surface becomes progres-
charge the whole stretch was freed from Cladophora sively rougher. If the laminar boundary layer of mat sur-
which was transported downstream. Within the next faces exeeds a certain roughness, turbulence occurs,
hundred metres or so nearly 70% of the drifting with the result that the aufwuchs detaches even at low
biomass were attached to protuberant stones and artifi- water movements.
cial elements, for instance piers and weirs. Shading by The algae mats can also drift into the reed-belts and a
bank trees led to rapid death; nearly 30% of the dead great part of the biomass of the whole lake can be "ban-

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168 W. SchOnborn

ished" into this zone. Within the plant belt, as a conse- bights which function as "sputum rooms". The more
quence of strong aeration, rapid decomposition is bights a lake has, the better its recovery mechanisms
favoured, particularly by the aeration of the rhizosphere seem to function. In heavily loaded Central European
and the released nutrients are quickly taken up by the lit- lakes, algae stranded along shores with many inlets are
toral vegetation. Therefore the pelagic eutrophic level is frequently up to 0.5 m wide. On 1 m of shore up to 1 kg
mostly lower than that of the littoral. fresh weight of algae can be found.
The wash-ashore-mechanism was already described The blue-green alga Aphanotheca on sediments pro-
by CEEB (1972) who recorded >2 kg m-3 (fresh weight) duces jelly-like masses the size of pigeon-eggs that drift
of blue-green algae washed ashore from a lake. Recent- on the water surface and then to the shore. They are
ly, Kms & DWORSKY(1.982) and SCH~I.Z-STEINERT& occupied by diatoms and small animals, and inside the
KIES (1996) have described these processes in Vaucheria jellies calcite precipitates and oxygen is produced,
sessilis which also detaches from the bottom and can which accelerates their decomposition.
drift ashore. In shallow waters with a drying period, the Even in the inlets of oligotrophic lakes masses of
algae mats frequently form paper-like layers over the dry Cladophora, Bulbochaete, Spirogyra and Mougeotia
sediment. These layers are converted by terrestrial or- drift onto the shores after they have been detached by
ganisms until the water body next fills. Vaucheria com- animals, or these days by bathers, in the water.
pacta, distributed from the fresh waters to mesohaline Not only filamentous algae and mats produced on the
zones of estuaries, also forms extensive mats on sedi- sediment can be drifted by wind ashore, but also wa-
ments. Like Cladophora, this species filters great terblooms. These consist mainly of algae which, be-
amounts of seston from the water and is a habitat for cause of embedded gas vacuoles, float on the surface. If
many animals. During storm surges Vaucheriadetaches the nutrient level is high these blooms can occur in
and is washed ashore (K()TTER 1961; SIMO~S 1974). The masses. An important bloom-forming green alga is
production amounted to ~16 g C week-~ m--2. Compara- Botryococcus braunii (Fig. 2). It forms colonies 0.5 mm
ble values for CIadophora in streams are 140 g (River in diameter and stores enormous amounts of oil in its
Saale, Thuringia; see SCI40NBORN1980) and 40 g (River cell walls thereby reducing the specific gravity. Large
Ilm, Thuringia; see SCI40NBORN 1996). Both species of quantities of starch are stored in the cells. This alga has
algae show the same behaviour and have similar func- existed since the Palaeozoic and it is a main component
tions in aquatic ecosystems. of oil pools. Stranding of this algae has probably been an
In Central Europe stranded algae are found mainly on important factor in unburdening water bodies through-
the east side of standing waters, because westwinds pre- out the Earth's history.
dominate. Frequently stranded algae are concentrated in The majority of waterbloom-forming algae are
Cyanophyceae. Their gas vacuoles are composed mostly
of molecular nitrogen (Microcystis, Anabaena, Gloeo-
trichia, Coelosphaerium, Gomphosphaera and Aphani-
zomenon). The drift of blue-green algal blooms, espe-
cially those of Microcystis, has been intensively investi-
gated (GEORCE & EDWARDS 1976; VAN DER VEER et al.
1993). Drifts are particularly concentrated in bights on
the lee side of water bodies and they rot quickly. VAN
DER VEER et al. (1993) have developed a model that de-
cribes the accumulation rate of drifting masses of blue-
green algae on the shore. The drift is dependent on the
quantitity of algae, lake surface area, wind course, and
wind velocity. In a shallow, hypertrophic, lake over 75%
of Microcystis blooms were washed ashore! But the
blooms must be distinguished from the other phyto-
plankton which colonize the whole light-exposed water-
body and is only seldom drifted by wind. In this case
other defensive mechanisms are effective that will be ex-
plained later in this paper.
Also the neuston participates in the wash-out effect.
If mass colonization of the boundary layer of the water
Fig. 2. The green alga Botryococcus braunii, an important storage or- surface occurs the wind can transport this neuston film
ganism which can be easyly removed by wind from lakes. (From STRE- shoreward. A particular role in this case is played by the
BLE& KRAUTER2002). green alga Ankyra (Characium) ancora which frequent-

Limnologica (2003) 33, 163-189

 Defensive reactions of freshwater ecosystems 169

ly occurs in high densities and has hydrophobic resting • Skimming

spores. The resting spores from Ankyra-films, driven Floating algal mats and pleuston plants are used by ter-
ashore by the wind, quickly reach the land and can be
restrial animals. Parts of consumed biomass will be de-
distributed to other waters (FOTT 1954). posited as faeces on land. This process may be designat-
The plenston, like the neuston, is integrated in the de- ed as skimming (Abschtpfen). A well-known case of
fensive mechanisms. Lemna and, in the tropics, also skimming in Europe is Lemna, which is used as food for
Salvinia molesta and Eichhornia crassipes spread car- ducks (Anatidae). Additionally, algal mats are also fre-
pet-like over the surface of lakes and slowly flowing quently used as food by ducks, above all because the
streams. Then they are drifted ashore, where they die. mats contain many small animals. Many aquatic birds
Especially in lakes with a strongly varying water level, eat filamentous algae especially in spring, and ducks
the removal of pleuston plants leads to significant im- also use the fungal drift previously described. They eat
mobilization of nutrients. During falling water level Sphaerotilus-clumps, including the microfauna inhabit-
pleuston plants dry out very quickly and will be metabo- ing the clumps. Drifting Sphaerotilus and fungal clumps
lized by terrestrial organisms (HARPER 1992; HARPERet are deposited by degrees and yield a fertile sediment.
al. 1993). This sediment is colonized by a high density of tubifi-
These wash-ashore mechanisms occur not only in cids. Ducks filter the tubificids from the sediment, and
standing waters but also in slowly flowing streams. In by this process consume great quantities of biomass, in-
such streams, filamentous algae can form large mats cluding nutrients.
which drift bankward, also Lemna-carpets are frequent-
ly pushed to the edge. Furthermore, macrophytes in the • Grazing
middle of slow and shallow streams catch great quanti-
tites of seston, drifting algal mats and Lemna, produc- Another terrestrial outlet-system of biomass and nutri-
ing enormous concentrations of biomass and other or- ents is the grazing of waterplants by terrestrial animals
ganic matter. These concentrations of organic matter, in which in addition to mammals birds (e.g., ducks,
representing in a hydrographic sense "transport-bod- coots) also partizipate. According to REICHHOLE(1993)
ies" will be drifted downstream by the next high dis- coots can consume up to 90% of the waterplants of lakes
charge. (e.g., reservoirs of the river Inn). The more the animals
Not only lakes and rivers have inherent disburdening defecate on land, the more intensive is the output of nu-
mechanisms, but so do periodic and episodic waters, and trients. Anatids can reduce the trophic and saprobic level
not only in the manner of drying algal mats mentioned enormously.
above. During dry periods deposited matter is oxydized But these relationships may be more complicated
and the bacteria get oxygen for decompositon (imitated than at first supposed. For instance, geese grazing in the
by rotating biological discs). When shallow waters daytime on fields and sleeping at night on the water are a
freeze, nutrients are probably released (DABORN& CLIF- significant factor in eutrophication (BAZELY& JEFFERIES
FORD 1974), which will mostly be washed out by heavy 1985; ZIEMANN1986). The faeces of herbivorous water-
rainfall in the next spring. fowl deposited in water produce rapid eutrophication,
because they make available nitrogen and phosphorus
previously bound in water plants. Coot (Fulica atra) and
3.3. Terrestrial outlet-systems also rudd (Scardinius erythrophthalmus) can consume
Systems of biomass release from waters that are based great stocks of Elodea. This gap is used by Ceratophyl-
on food chains, only have long-term effects. The unbur- lure that spreads quickly and takes up nitrogen and phos-
dening is delayed, for instance, because defecation by phorus by its leaf. Elodea, on the contrary, takes up nu-
terrestrial grazers and predators in water bodies is al- trients from the sediment (VANDONK et al. 1993). How-
ways possible. Respiration (metabolization) also be- ever, Ceratophyllum is not eaten by most animals and
longs to the biomass release systems, but this process this mechanism counteracts the availability of nutrients
will not be referred to in this paper. from the faeces.
Defensive reactions concern not only the reduction of There are many mammals that eat waterplants, for in-
nutrient levels, as will be shown in the following re- stance elk, elephant, rhinoceros and hippopotamus, but
marks, but also improvement of water quality. This also wild boar. Wild boar do not deposit faeces in water
means that the water becomes physiologically more tol- bodies but often emerge garlanded with floating algal
erable, for instance, through increased transparency and mats which are thus transported landward. A special role
decreased fluctuations in pH-values as well as CO2- and is played by seacows (Sirena), which can push up into
O2-concentrations. streams and consume great quantities of waterplants.
In the following the main terrestrial outlet-systems Because they remain in the water, they do not really be-
are explained. long to the terrestrial outlet system, but they keep even

Limnologica (2003) 33, 163-189

170 W. Sch6nborn

strongly eutrophic streams free of plants and therefore Campostoma, which is captured by perches and has been
they prevent the realization of the nutrient potency in used in attempts to control the eutrophication of running
waters. waters (PowER et al. 1985). Barbs and nases mainly are
captured by huchen. Crocodiles and other reptiles, e.g.
• Predator-prey-relarionships Python, hunt fish in various water bodies. Capybara
The predator-prey-relationships represent the most in- (Hydrochoerus capybara) in South America eat great
tensive outlet-system. In this mechanism the initial link quantities of waterplants therefore preventing weedi-
of the food-chain is a limnic organism, and the end of the ness. They are prey of the Anaconda, a top predator in
chain is a terrestrial predator. American waters. The snake eats the prey within the wa-
Widely spread is the system alga - fish - terrestrial ters and also defecates there. Therefore it only prevents
predator which should be described in the following. realization of the nutrient potential, but does not greatly
The majority of fish species eat algae and the typical fish reduce the nutrient burden.
smell comes from algae. In fresh waters we can find Water bodies are comparable with steppes or savan-
more herbivorous fish species than in the sea. nas since in both large grazers prevent, respectively,
Especially among the cyprinids many algal con- weediness or bushiness. In most cases large grazers
sumers are found. For instance, species of Varicorhinus, scarcely reduce the nutrient level, but its conversion to
an acient genus, living in South Asia and Africa, scrape phytomass is significantly reduced.
off the algal aufwuchs with the help of their under-stand- The next link within the terrestrial outlet-systems -
ing mouth and their pointed underlip. Their long intes- after algae and fish - are piscivorous terrestrial animals.
tine may also be an adaptation to this form of feeding. Because there are a lot of these animals, only a few ex-
The species V. capocta, occurring in the Sewan area (Ar- amples can be referred to.
menia), even eats the toxic green alga Botryococcus Among the families of birds Mergus-species, herons,
brauni, which forms blooms and is rich in nutritious storks and cormorants especially control fish densities.
matter. This alga plays a key role in the cleaning-sys- Goosander and red-breasted merganser (genus Mergus)
tems of many waters, as mentioned above. consumed nearly 3.5 tons salomonids per year in a
Other algae-consuming cyprinids are the nase Swedish stream, this amounts to 350 000 individuals if it
(Chondrostoma nasus), living in the same part of rivers is assumed that one individual weighs ~10 g. An adult
as graylings and barbs, many barb species, the chub
Mergus needs 400 g fish d-1 and so Mergus-species can
control salmonid populations (LINDROTIJ 1955). Data
(Leuciscus cephalus), the rudd (Scardinius erythroph-
from another area indicate that Mergus consumes simi-
thaImus), the black belly (Xenocypris), the genus
lar quantities of non-salmonids.
Discognathichthys and other species. In mountain Appropriate investigations are given for cormorants
brooks of South-East Asia species of Gyrinocheilus
by LEAI4 et al. (1980). In a hypertrophic shallow lake
feed on algae. The North-American minnow (Campos-
cormorants (PhaIacrocorax carbo) caught fish in such
toma anomalum) also eats filamentous algae (Spirogy- high quantities, that water transparency greatly in-
ra, Rhizostomum) and can significantly reduce the algal creased. This led to an increase of macro-zooplankton
density (POWER et al. 1985). Even the carp (Cyprinus species and also of macrophytes and zoobenthos. Of
carpio) eats filamentous algae if there is not enough course, the nutrient level remained constant, because a
zoobenthos. The silver carp, HypophthaImichthys long time is necessary before the influence of the birds
moIitrix, filters great quantities of phytoplankton. The greatly reduces nutrients. Furthermore, there should be a
Indonesian fish species Tilapia mossambica, Chanos long distance between the waterbody and breeding
chanos and Thynichthys species have also a high con- colonies and the re-solution of nutrients must be taken
sumption rate of microalgae. According to LAKSIJM~- into account, especially that of phosphorus from the sed-
NARAYANA(1965) the fish species Hilsa iliska, Gadusia iment. Cormorants can empty water bodies of fish and
chapra and Barbus stigma eat planktonic diatoms in are therefore regarded as a danger for inland fisheries.
the River Ganges. But it should be taken in account that, at present, cor-
Some of the fish species mentioned are used in algae morants are the most active and natural factor in the im-
control. The grass carp, Ctenopharyngodon, introduced provement of quality of standing waters in the Central
into Europe for waterplant control, eats macrophytes but European landscape.
also the macrophytic algae Spirogyra, Cladophora and In the same way, herons and storks can improve water
Chara. But it can not be regarded as a cleaning factor, quality. The Mycteria-storks of India and the African yel-
because nutrients are released in its excrements, which low-billed stork (Mycteria ibis) consume large quanti-
contain nearly 77% of the phosphorus-uptake, resulting rites of fish. One pair can catch >40 kg fish per breeding
in a strong development of phytoplankton. season, great colonies even some tons in the same time.
Frequently, the aquatic food-chain is lengthened by Tropical shallow waters and marsh areas support
predatory fish species. One example is the minnow, enormous densities of piscivorous birds. To this func-

Limnologica (2003) 33, 163-189

 Defensive reactions of freshwater ecosystems 171

tional group belongs also the hals (Alcedinidae) which predator) the food-chain Cladophora - Astacus - otter
mainly eat young fish that feed mainly on algae. can be used for biomanipulation and is thus a major fac-
Not only birds but many terrestrial mammals feed on tor in reducing of the trophic level. But it may also be
fish including some soricids living in water, bears, the important, that decapods detach more Cladophora than
water mongoose (Atilax paIudinosus) and the fishcat they consume (SCHMALZ 1999), and thus the two re-
(Felis chaus). Large cats like tiger and jaguar also fre- moval-systems are connected (Fig. 3).
quently catch their prey in water. To the terrestrial outlet-systems also belong Amphib-
First of all should be mentioned the family of otters ians, above all frogs and toads. Their tadpoles assimilate
(Lutrinae) because it plays an important role in the relief different matter from the water, which is, at their meta-
of all limnetic waters. It is distributed world-wide, most- morphosis, transported out of the water (DICKMAN1968;
ly in high densities, and occurs from the mountains to SEALE 1980). The tadpoles feed intensively on algae and
the plains and colonizes nearly all types of water bodies. take up also the animals which inhabit algae mats. The
Most important is that defecation takes place outside of tadpoles of Rana dalmatina, e.g., intensively consume
water - mostly on prominent points - and serves also for swimming mats of Spirogyra, Zygnema, Mougeotia,
the marking of territory. Cladophora and Oscillatoria. Because the tadpoles eat
All otters need great quantitites of crabs (Decapoda) some of their own proteinaceous faeces, release of nutri-
and fish. Especially the giant otter (Pteronura brasilien- ents to the environment is decreased as a result of repeat-
sis) of the Amazonian region needs decapods and fish in ed cycles of digestion (WARINGER-LOSCHENKOHL&
extremely high quantities. WARrNGER1990).
Under natural conditions, decapods occur in high Terrestrial outlet-mechanisms must have already ex-
densities in natural brooks, small rivers and shallow isted in the Mesozoic. Since this time eutrophication or
standing waters. Decapods consume great qua~titites of disburdening depend on the place of defecation and the
waterplants (Chara, seedlings of Potamogeton and other proportion of landplants in the food. Many saurians fed
macrophytes). According to NYSTROMet al. (1996a, b) on waterplants and it can be assumed that the hy-
they can significantly reduce the stock of waterplants drosaurians filtered swimming algae and pleuston from
and are regarded by these authors as key species in lim- the water. Pterodactylus and other flying saurians cap-
netic ecosystems. One group of Decapoda, the Astacura, tured great quantities of fish during flight. In this way,
feed frequently on Cladophora (LE SAGE & HARRISON these land animals may have relieved the water bodies
1980; HART 1992; ANWAND & VALENTIN 1996). This significantly.
species, which plays a very important role in the clean- In the Tertiary the otters occur in a high diversity,
ing system, removes biomass, nutrients and xenobiotics more than 10 genera were distributed world-wide. They
not only by a detachment-mechanism, but additionally may have disburdened the water bodies considerably
with the help of a food chain. Like the food-chain ex- which were polluted by many mammals living in high
plained above (algae - algivorous fish - terrestrial densities in this earth period.

/,
xC:7l Y ''

1/ l/ ,J
/'ff4'7

Fig. 3, Example of a terrestrial outlet-system: algae, fish, Astacusand otter. In the water body can be seen the cave of the crayfish, providing
the alder rhizosphere with water. Above the bank lies a food- and defecation-place of the otter. (Half-schematic drawing).

Limnologica (2003) 33, 163-189

172 W. Sch6nborn

In a completely different way hippopotamuses and The main benthic filter-mechanisms are large mussels
other large animals decrease eutrophication, especially and the role of Dreissena polymorpha has been especial-
in African lakes. The movements of their ponderous ly well investigated. During one hour, one mussel can
bodies continually destratify and therefore detoxify eu- filter 300 ml of water free from particles (RE~D~RS
trophic lakes. 1989); but there even higher values have been published.
Around 10 000 mussels can reduce by 69% the suspend-
ed matter of a middle-sized water body with an average
3.4. Marine outlet-systems load (VOO~T 1989; REEDERS 1990). In this way Dreisse-
Marine outlet-systems are represented by catadromous na can significantly decrease the degree of the trophic
fish, which migrate for spawning from fresh waters into status of a water body, it is, for instance, "the greatest
the sea. The most important group of this system are eels sewage treatment plant of Lake Constance" (KLEE
(Anguilla) which are extreme predators of decapods and 1971). The filter-system of Dreissena has probably been
fish and migrate fattened for spawning in the sea, from active since the Pliocene (KINZELBACH1992).
which they do not return. This output is probably not It may be important that the filter feeders are fre-
compensated for by the immigration of eel-larvae (fae- quently connected with terrestrial outlet-systems. This is
ces from marine food). possible because fish (rudd, bream, barb and especially
In contrast to catadromous fish, anadromous fish eel), diving ducks, coots and otters eat mussels. But de-
species, spawning in running waters, can be a significant capods are also great Dreissena predators, with the re-
eutrophication factor. A well-known example is the Pa- sult that they are integrated in this important food-chain
cific salmon, Oncorhynchus nerka, post-spawning for the relief of water (PmSIK 1974).
deaths of which can significantly increase the primary Other big mussels, e.g. unionids, also have a filtration
production in some stream reaches (R~crIZYet al. 1975), effect comparable to Dreissena but they are unfortunate-
and at some points also the saprobity. The existence of ly insufficiently investigated.
these two counteracting mechanisms means it may be The role of other benthic filter feeders like ciliates, ro-
difficult to estimate the result of marine outlet-systems. tifers, sponges, bryozoans, some trichopterans and
simuliids should not be underestimated. They metabo-
lize a part of their high quantities of food and are inte-
3.5. Filtration grated in outlet-systems.
Filtration of suspended matter occurs both mechanically
and by the activity of organisms. The most important me- 3.6. Precipitation (calcite precipitation)
chanical filter-system in running waters are large clumps
With the increase of the nutritional level the density of
of filamentous algae, especially Cladophora. Cladopho- phytoplankton in standing waters also rises. This leads
ra can clarify seston-loaded streams (ScHONBOm'~1997). to an increase in CO2-concentration with the result that
With the help of detachment-mechanisms, the filtered the Ca (HCO3) 2 can not remain in solution and precipi-
seston is transported downstream (see chapter 3.2). In tates as CaCO3, in the form of microscopical calcite
standing waters the stock of another plant, Myriophyl- crystals. Sand grains, shells of diatoms and probably
lure, store much detritus, but without drift. The effect of bacteria act as crystallization germs. Sinking calcite
this storage of suspended matter is unknown, but it may crystals transport nutrients from the pelagial into the
contribute a little to clearing the water. depth. This can happen mechanically by the crystals
Filtering organisms occur in standing as well as in pulling down of organic particles and algae, but also by
running waters, in the pelagic and benthic zone. The adsorption of dissolved phosphorus on crystal surfaces.
most important filter-system among pelagics are clado- This process, long since known as "biogenic decalcifica-
cerans, and there is an immense literature on their role in tion" or "whiting" was investigated anew by KOSCHEL
this respect. The biomanipulation programme developed (1990) who concluded that it is a cleaning reaction in
for lakes is based fundamentally on the filtering ability hardwater lakes. The effect of calcite precipitation on
of, above all, Daphnia. High densities of Daphnia- phosphorus content is given in Fig. 4.
species in lakes can produce a clearwater stage within a Integrated in this process is the phytoflagellate Pha-
short time. This process does not lead to a reduction in cotus (Fig. 5) since its empty calcareous shells sink
nutrients, but results in an improvement of the water (KosCHEL & RAIDT 1988). By this process Phacotus be-
quality in a biological sense. Biomanipulation imitates longs, like Cladophora or Botryococcus braunii, among
natural defensive mechanisms, but only incompletely, as the key groups for the relief of waters.
indeed, natural defensive mechanisms were also incom- More information about calcite precipitation as an
plete. The influence of cormorants and otters, as de- important defensive reaction of lakes will be given later
scribed above, comes closest in type to the biomanipula- in this paper, but the incomplete nature of these mecha-
tion applied. nisms will also be described.

Limnologica (2003) 33, 163-189

 Defensive reactions of freshwater ecosystems 173

1,4 -- -- 80

/
1.0 --
,/ /'\
-- 60
C~
E
// \\
t,,,.

¢O
o
t-
O 0°6 --
O -- 40 ¢o
t~ ¢-
o
O 13_

TotaI-P\ . . . . ~
0.2 -- -- 20

Fig. 4. The effect of calcite precipita-

tion on phosphorus content. (Modified
Vl l VII I VIII I Months after KOSCHELet al. 1988).

concentrated chironomid larvae in a high density, using

the filtered detritus as food and for building their tubes.
Simultaneously, the chironomid larvae tubes fix detritus
within the filamentous system, facilitating in this way
the quantitative detritus drift. The emergence period of
the larvae is adapted to this detachment-mechanism
(SCHONBORN 1997). Furthermore, in the Cladophora-
clumps the leech Erpobdella octoculata, a main predator
of the chironomid larvae, occurs in high density and thus
Fig. 5. The phytoflagellate Phacotus Cladophora also becomes a center of habitats.
lenticularis is integrated in the calcite But the process of concentration is not be finished yet.
precipitation, an important relief process Parallel to the increase of stream eutrophication, both
of the pelagial zone of lakes. (From the density of bacteria and of bacteriovorous animals
STREBLE& KRAUTER2002). rises, above all the density of ciliates. The ciliates, again,
are prey of various small metazoans which are absent
from oligotrophic stretches.
3.7. Concentration and centralization
Bacteriovorous animals similarly need predators if
We have seen that there are different processes in water their populations are to remain in the exponential growth
bodies which ease removal of biomass and solid matter. phase. The most important predator of ciliates is the
To these processes belong the formation of neuston, predaceous naidid oligochaete Chaetogaster diastro-
pleuston and rising algae, which develop expanded mats phus which, in turn, is the prey of the leech Erpobdella
near the water surface, but macrophytes can also store octoculata. Erpobdella also captures great numbers of
suspended matter. chironomid larvae and of the detritivorous naidid Nais
This applies particularly to the macrophytic green which frequently occurred in Cladophora-clumps
alga Cladophora in streams. Cladophora shows mass (ScHONBORN 1985a). ErpobdeIla is from 3 mm (emer-
production, accumulates nutrients and xenobiotics and gence from the cocoons) to nearly 60 mm and all devel-
filters great quantities of suspended matter and these opment-stages are predaceous, occurring in high densi-
masses can be transported away completely by a single ties and controlling the whole meio- and macrozooben-
high discharge. By their mass production Cladophora thos. In streams with severely shortened food-chains
can be regarded as a center of primary production. (e.g., lack of predaceous fish, piscivorous birds and
But Cladophora is also a habitat for a lot of animals mammals) Erpobdella, beside cyprinids and salmonids,
which play important roles in matter cycling. For in- can be a second top-predator and becomes a predation-
stance, in the filamentous system of Cladophora are center (SCHONBOPd,~1985b, 1992, 1995, 1996, 1998).

Limnologica (2003) 33, 163-189

 174 W. Sch6nbom

In summary, we can say that there are three centers The concentration leads simultaneously to a c e n t r a l -
which lead to a c o n c e n t r a t i o n of the ecosystem process- ization of the ecosystem, in response to the increase of
es in loaded streams, whereby the ecosystem relation- eutrophication. Therefore the ecosystem can react as an
ships are simplified. What does it mean in regard to the entirety against eutrophication. All the facilities that are
defence against eutrophication? involved in defensive reactions therefore act in concert.
- Parallel to the density of bacteria the density of bacte-
riovorous animals increases, stopping stationary 3.8. Accumulation
phases in bacteria growth and leading to permanent Accumulation can be interpreted as a defensive reaction
stimulation of bacterial production and consequently against toxic loading, initiated in the earliest geological
tO the decomposition of organic matter. periods in connection with increasing volcanism.
- Many bacteriovorous species are found in the group In the present paper, accumulation is defined as the
of ciliates, which play a key role in self-purification. enrichment of nutrients and xenobiotics in organisms.
Other groups, like rotifers, have also a great number The importance of nutrient enrichment is described
of bacteriovorous species. Especially important in this above (see chapter 3.2.). The accumulation of xenobi-
connection are the sediment feeders (oligochaetes, otics refers to heavy metals and today also to pesticides.
chironomid larvae), as explained above. These groups In waters of the temperate climate, primarily
increase with eutrophication, because of the enrich- Cladophora and water mosses accumulate xenobiotics.
ment of sediments with bacteria, the main food of the The accumulation factor can be more than 106. In adae-
larvae. In the River Saale (Germany) the oligochaete quate ecosystems these plants have a similar function to
worm Nais digests up about 280 g bacterial biomass the liver in higher organisms. In the case of CIadophora
m -2 a-1, whereas by ciliates only 182 g m -2 a-1 were the accumulated xenobiotics are removed from the
consumed in the same time (Scn&'mORN 1985a). stream reach concerned by a detachment-mechanism.
- The last link in this predator-chain is Erpobdella, Drifting Sphaerotilus also accumulates heavy metals,
which controls great parts of the macrozoobenthos however, especially copper. Mosses, indeed, are great
(SCHONBORN 1987; see also Fig. 6). accumulators, but very resistant against drift. They are

Other prey
Chironomid larvae
> 200 kJ m -2 a -1

,ncreaseof
Bacteria etogaster 900 kJ m a

-... " i...ji ."

200 kJ m"2a-1 ~ l ~ ~ I',~ .....",~i.;.'(

Storage of detritus in
Cladophora cMmps
~ 4000 kJ m -2 a -1

Cladophora
6 8 103 kJ m -2 8 -1
Fig. 6. C/adophoraclumps as a center of primary production, habitats and predation. The values give the production of the named organisms
or the storaged detritus, respectively. The detritus storage referred to a supposed vegetation period of Cladophoraof four months. The arrows
indicate the energy transfer. (Compiled from several papers, compare SCHONBORN1992).

Limnologica (2003) 33, 163-189

 Defensive reactions of freshwater ecosystems 175

scarcely used as food by animals, whereby the accumu- With eutrophication, standing waters in shore and lit-
lated matter is isolated from the water. toral areas expand their waterplant zone. This vegetation
The immense literature on aquatic toxicology has not zone has characteristic stratification. But only emersed
been taken into account in the present paper. waterplants are important as defense mechanisms be-
cause they can occupy wide zones of both standing wa-
3.9. Other relief processes ters and slowly running waters. Like organisms many
waters possess a "skin". In this case it is a two-fold strat-
• Resuspension ification: a tree-belt and an emerse belt. In mountain
Resuspension, the churning up of sediments, is caused brooks, the emersed plants are frequently replaced by
by bioturbation and in shallow waters additionally by sloping-plants, e.g. Petasites and Phalaris. This "skin"
wind. Oligochaetes, chironomid larvae and, especially, of water bodies takes an important protective function
benthivorous fish (by rooting up sediment) participate in against eutrophication.
the bioturbation, but also large mammals by bathing and
T h e a l d e r z o n e : Alders' system of dense filamentous
food seeking.
The churned up sediment increases water turbidity roots block the diffusion of matter into water. A 10 m
and therefore inhibits phytoplanktonic primary produc- broad strip of trees along running waters reduces nitro-
tion. This can partially suppress realization of the eu- gen input from the surrounding fields by 10-15% and
trophication potential. Resuspension is prevented fre- phosphorus input is reduced by about 20-30% (MANDER
quently by macrophytes, but on the other hand churned 1985). The alders' hydrophilic thick roots stabilize
up sediment is deposited on leaves and thus restricts stream- and river banks. Their filamentous roots, thin as
their production. a hair and swinging in water, serve as habitat for many
Resuspension also simultaneously leads to the remo- small animals which take part in decomposition. Land
bilization of phosphorus locked in the sediment. So it roots of alders possess tubercles with symbiotic bacteria
seems that two contrary effects occur: on the one hand, (Actinomycetes, especially Frankia), that fix the air ni-
realization of the eutrophication potential is reduced, trogen in the soil. Therefore, alders can grow in nutrient-
and on the other hand more phosphorus is available for poor soils. The high nitrogen content of alders is dis-
primary production. tributed up to the leaves. In contrast to many other tree
The storage of phosphorus, especially in sediments, species, prior to leaf abscission alders stop the removal
can also be interpreted as aimed against eutrophication, of nitrogen from the leaves. Therefore, the C/N-ratio of
and furthermore, its release from sediment by bioturba- alder leaves that fall into the water is relatively low and
tion brings it more quickly to outlet-systems than it may makes the leaves an important food source for aquatic
happen by degrees. But this is not supported by facts. invertebrates. The input of nutrients by alder leaves does
not cause an increase of eutrophication, because the de-
• Denitrification composition of the leaves by hyphomycetes, by bacteria
Another relief-system is denitrification which is inte- and by intestinal passages of shredders occurs slowly
grated into limnetic ecosystems and probably very effi- and successively. Animals in extremely nutrient poor
cient. Nitrate can be denitrified wherever an oxygen de- brooks feed on alder as well as willow leaves.
ficiency occurs. Most important is denitrification on the
T h e b e l t o f e m e r s e d p l a n t s : A belt of emersed plants,
surfaces of stones and other substrates. It can be ob-
served even in oxygen-rich headwaters because it takes especially the reed, Phragmites australis, provides an
place on the basis of PRANDTL'S boundary layer. Even effective protection against eutrophication. The main
some 100 pm above this layer oxygen occurs. Such oxy- mechanisms of this protection are basing on the follow-
gen-rich habitats represent an important cleaning-sys- ing principles:
tem if their area is great enough. In running waters - Emersed plants act as a dike against infiltrated matter.
epilithic denitrification rates of 16 mg m 2 d-1 have been - Shading by emersed plants counteracts the develop-
measured (NAKAJIMA1979). Animals can also take part ment of algae in shore areas.
in nitrate respiration and an important role in eutrophic - Algae washed ashore from the main body of water are
profundals is played by the ciliate Loxodes. caught by the emersed belt where they die from shad-
ing. Emersed plants function as a trap for matter. The
dead algae remain in the wind-protected belt, are de-
3.10. Morphological defensive systems posited and decompose quickly due to the high oxy-
The banks of running and standing water are frequently gen content.
colonized by trees. In Europe, these are alders (Alnus) - The sediment within the plant belt is sufficiently aired
and willows (Salix), in other geographical regions Ficus because the roots release oxygen (from the assimila-
species, gallery forests and mangroves. tion of the leaves and transported through the

Limnologica (2003) 33, 163-189

176 W. Sch6nborn

aerenchyma). This is primarily an adaptation to oxy- stance, accumulate copper and iron (FEZ+),but frequent-
gen-poor sediments, but it leads simultaneously to an ly the heavy metals stick together with detritus at the
increased decomposition within the belts and there- reed stocks. Following OSTERKAMPet al. (1999) reed
fore also to nitrification and sulphurization that leads stocks retain:
finally to detoxification. - 29-91% of filterable matter and
- Dead leaves of emersed plants can be found frequent- - 16% lead, frequently connected with filterable matter
ly on the sediment surface. Immediately below this (see Fig. 7).
layer of leaves are anaerobic conditions. In these flat From studies of wastewater treatments using aquatic
areas nitrates and sulphates are denitrified or desul- plants, it is established that mineral-oil hydrocarbons
phurified, respectively. The mosaic-like distribution (MHC) and polycyclic aromatic hydrocarbons (PAH)
of oxygen-rich and oxygen-poor patches in the belt of can be enriched in sediment between reed stalks. There-
emersed plants creates a detoxification- and disbur- fore, tree-belts and belts of emersed plants provide water
dening-system (B6a~R 1992). bodies with an effective protection against a wide spec-
- In anaerobic zones of water, anaerobic decomposition trum of allochthonous matter. If groundwater in flood-
occurs that results in development of H2S, NH4+ and plains reaches the surface of soils (stressed conditions),
C Q . But these processes do not damage the reed marshlands will be formed including swamp forests or,
plants (HORLIMANN 1951)because the anaerobic as in the Southern Africa, gigantic Cyperus areas. In
reaches will by bioturbation promptly be aerated. spite of increased nutrient input by big game, the marsh
Thus the patch dynamics between aerobic and anaero- soils and their vegetation near the water bodies accumu-
bic reaches function as an effective protection against late great quantities of nutrients and thereby prevent a
eutrophication. hypertrophication. In this way, the wet soils of flood-
- The plants accumulate nitrogen and phosphorus (per plains result also in increased denitrification.
reed stalk ~150-250 mg N and 10.5-18.5 mg P) that
is partly released in the summer months (SCHIEFER-
STEIN 1999). Therefore, the accumulation of nitrogen 4. Euoecismsanddysoecisms
and phosphorus does not seem to be an effective pro-
tection against eutrophication. 4.1. Euoecisms
The belt of emersed plants, especially the reed pro- Euoecisms are defined as a co-operation of various fea-
tects against not only organic loading, but also against tures of one or some partial systems to the benefit and
unorganic and toxic matter. Roots of the reed, for in- protection of the whole or greater parts of the ecosystem.

AFS
100

80

60'

¢,0
LL 40

20

0
3 5 7 9 11 13 15 17 19 21 23 2 5 2 7 2 9 31 33 35 37 39 41 43 45 47
Sample-Number
Pb
50

o'J 3O

..o 20
g.
10 . . . . . - . . . . . . . . . . . . . . . . . . .
Fig. 7. Detention of particulate matter (AFS) and
lead (Pb) within a waste water treatment using
0 waterplants. Measured in inflow, middle and outflow
3 5 7 g 11 13 15 17 19 21 23 25 27 29 31 33 35 37 39 41 43 45 47
during a storm-runoff-event. (After OSTERKAMPet al.
Sample-Number 1999).

Limnologica(2003) 33, 163-189

 Defensive reactions of freshwater ecosystems 177

We have just described such an euoecism with the tubificids. The food-seeking of bream influences not
alder belt and the belt of emersed plants. The function of only the oxidation of sediments and the stimulation of
alders, for instance, can be completed by forming a prey species, but also the removal of biomass from the
canopy. This cools the water and reduces primary pro- profundal zone. Furthermore, through the oxidation of
duction, leading finally to the protection of greatly load- sediment, more phosphorus is bound in chemical com-
ed brooks against weediness. The alder leaves can also pounds.
regarded as a pharmacy for crayfish (Astacus), because These euoecisms seem to be widely spread. Many eu-
the consumption of alder leaves is a protection against oecisms occur only in small spheres of activity without
fungus diseases. The great importance of crayfish for the influencing the whole system. Some examples are:
healthy of water bodies is decribed above (see chapter - Floating features of planktonic forms protect also
3.3). against predators.
When leaves of alders are transported downstream in - The suppression of secondary leaves of some flower-
temperate regions the spring winds blow the deciduous ing plants in the water current - mechanically or by
leaves of the hardwood vegetation from along the river- loss of assimilates - allows 0nly primary leaves which
side into the stream. These leaves are heavy decompos- are adapted excellently to current water.
able, but in the course of their decomposition the C/N-
ratio decreases and the leaves become edible for inhabi-
4.2. Dyseocisms and the "egoism"
tants of the stream.
in limnetic ecosystem compartments
Beside the harmonizing features of the belt of
emersed plants in regard to the wellbeing of the whole In contrast to the euoecisms, dysoecisms also exist.
system, their function as habitat for many birds, espe- Dysoecisms are defined as defensive reactions of a com-
cially for herons, is also of interest, because the birds partment of an ecosystem to the detriment of one or sev-
further remove biomass from the system. eral other compartments of the same system (SCHON-
Another euoecism begins with such planktonic di- BORN 1995). From an anthropomorphic point of view,
atoms which are only moderatley adapted because they the different compartments can react "egoistically", in
sink to the ground relatively quickly and serve as food regard to other parts of the system.
for profundal animals. Other sinking plankton also in- Such behaviour is exhibited, for instance, in the de-
creased the nutritive value of the sediments. Many ani- fensive system of Cladophora. Their mass production,
mals that feed on sunken living phytoplankton, or on which is presumed to be its defensive reaction, impairs
sediment generally contribute to bioturbation and to the the whole stream ecosystem. By its relatively coarse-fil-
shift of sediment. This leads to the oxidation of sedi- amentous system, many invertebrates are excluded from
ments and therefore to the stimulation of the bacterial the Cladophora clumps. These clumps are dominated
production, decomposition, nitrification, sulphurization mainly by chironomid larvae, leeches and other worms.
and to an increasing release of methane and molecular Young fish, for example, are frequently "caught" in the
nitrogen enclosed in the sediment. long filaments (FOTT 1971) and die.
Benthivorous fish also contribute to bioturbation in The mass production of Cladophora reduces the den-
the profundal zone. In Central Europe it is primarily sity of many algae and animals that otherwise would be
bream (Abramis brama) that feed on important sedi- typical species of the given bed structure or water quality.
ment-shifting animals, such as chironomid larvae and This refers especially to Ulotrichales and Vaucheria and,

3O

u
Ill

E 10
Fig. 8. Number of speciesof Trichoptera
(white columns) and Ephemeroptera
(hatched columns)in the longitudinal
profile in a stream dominated by Clado-
phora(llm, Germany)in comparison with
2 3 4 5 6 Mouth into the number of speciesexpected to occur.
Stations the River Saale (After SCHONBORN1996).

Limnologica (2003) 33, 163-189

178 W. Sch6nborn

in the case of animals, the larvae of ephemeropterans remaining in the water. Furthermore, in the course of
and trichopterans (ScNONBORN 1996; see also Fig. 8). decomposition, it can be released toxic substances. In
The chemical conditions within large Cladophora this case, defensive reactions against eutrophication can
clumps also change. Nitrate uptake, high oxygen pro- be directed against the ecosystem itself. Such secondary
duction and an increase of pH value (up to 10) are effects of decomposition are absent in terrestrial habi-
caused by the CO2 decline resulting from the high assim- tats.
ilation intensity (FOTT 1971). Similar dyseocisms occur in waterblooms. Before
The detachment and transport of Cladophora masses they wash ashore, relieving the water, they cause ex-
benefit only a particular stretch, not the whole stream. treme day-night-alterations of CO2, 02 und pH values.
On the contrary, in structure-rich and shaded down- Such dysoecisms can also indicate a transition to or-
stream stretches, Cladophora masses are attached on ganic behaviour. It is well known that the content of
overhanging stones and rotten. The mass production of prussic acid of waterplants inhibits their defence against
CIadophora is an over-reaction of the system against pests, instead to protect the plants.
early stages of pollution. The defensive reaction is di-
rected against the ecosystem itself.
A similar case of dysoecism can be observed in cal- 5. The aging of limnetic ecosystems
cite precipitation (SCnONBORN 1995). On the one hand,
the sinking calcite crystals clean the pelagial but to the Limnetic ecosystems age and at any time this aging can
detriment of the benthic communities. The crystals lead to the dissolution of the system. Especially for
block the interstitial system of sediments and the fila- lakes, this process of aging and siltation has been well
mentous aufwuchs on stones and macrophytes. This re- researched. A lake becomes shallow with dead plankton-
sults in high losses of microflora and microfauna, living ic and littoral organisms, especially plants. The morpho-
in the interstitials. In Lake Stechlin, Germany, for in- logical alterations in lakes lead to an increasing eutroph-
stance, 140 taxa of testacean rhizopods were registered ication and thus to an acceleration of siltation. Eutrophi-
within the algal aufwuchs. But during increasing calcite cation in the course of aging results in the debilitation of
precipitation caused by phytoplankton, these taxa were the system and a tendency toward pessimism (e.g., de-
reduced to about 80. crease of species diversity, increase of extreme life con-
After calcite precipitation in the aufwuchs of filamen- ditions). In this regard, the aging of lakes is frequently
tous algae, many calcite crystals were found. Littoral compared with organismic aging.
sediments and algal layers on surfaces of stones were Aging can be accelerated or retarded. The accelera-
covered by calcite precipitation with a yellowish-white, tion is caused, for instance, by clear cutting of border
quickly resuspensable coat. It consisted of felted flocks trees, by a permanent nutrient input or when lakes are
of detritus, in which crystals are included. Most calcite situated in regions with a nutrient-rich geological under-
deposits can be found at the north and east shores of ground. Extant oligotrophic lakes can be regarded as
lakes, because the wind drifts the crystals to the littoral, perennial young stages. The system cannot reject aging
and in particular to the surf zone. Whereas by this pro- because aging is an irreversible process whereby mor-
cess the transparency in the pelagial increases, the ben- phometric relationships are changing. But this process
thic zone is covered by quickly resuspensable sedi- can be retarded by defensive reactions.
ments. Shells of phytoplankton and zooplankton are fre- Similarly running waters also age. In rivers, the troph-
quently found within these deposits. ic state increases downstream, also under natural condi-
To a lesser degree, filamentous algae mats also have tions. As a consequence of stream continuity, the age
biogenic decalcification, especially if blue-green algae stages of running waters remain connected beginning
occur in high density. But this does not impair coloniza- with the brook (young stage), followed by the middle
tion of the mats. course with meanders (maturity) and ending with the
We can draw the conclusion that a lake does not react lower course and mouth (old age). Running waters flow
as an entirety to the impact of loadings, but that the de- into standing waters (for instance a lake, mostly a sea)
fensive reactions negatively impact other system com- and end their existence. A shallow, lowland stream with
partments. They "overshoot the mark", comparable with a slow current has little chance to defend against eu-
allergic reactions of organisms. trophication, because it can silt up.
Another example are bacteria, which evolved as de- In the tropics, the main factor of siltation is the water
struents and which are of fundamental importance to the hyacinth Eichhornia. Algal mats also contribute, rising
existence of life on earth. This notwithstanding, their from the stream bed and drifting to the bank where they
function in water bodies can have disastrous secondary are deposited by their slow current. This deposition is a
effects. By their consumption of oxygen, they can en- siltation process that simultaneously creates a habitat for
danger the ecosystem more than undecomposed matter Eichhornia. Unlike as standing water, a water current

Limnologica (2003) 33, 163-189

 Defensive reactions of freshwater ecosystems 179

erodes a new drain beside the old bed but this new species arrived in Europe without causing losses of the
stream represents a new ecosystem. indigenous flora and fauna.
Other main factors of the siltation in tropic waters are Such benign immigration notwithstanding, there are
Salvinia (water fern) and Cyperus grasses. Water bodies also examples of conflicts between the indigenous
that have a dry phase stop their aging process in this dry species and immigrants. The conflicts - eradication or
period because the organic remnants are metabolized by displacement - caused by new predators or competitors
the air at this time, providing a little relief to the aging (e.g., for food, life space and so on) can be partial or
process. complete. But in the most cases the conflicts are only
local, concerning only a certain lake or region (TITTIZER
6. Immigration of foreign species 1997). A well investigated example of eradication of a
into limnetic ecosystems species by an immigrant is the copepod Heterocope in
(neophytes, neozoa) Lake Constance. During a period of eutrophication in
Lake Constance, the predatory Cyclops vicinus immi-
It is sometimes believed that the immigration of foreign grated and has eradicated Heterocope by capturing its
species into an ecosystem causes defensive reactions, nauplius larvae within two years.
comparable to the reaction of organisms to an infection The most sensational case of the predatory eradica-
or comparable to defensive reactions against eutrophica- tion of species by a neozoa is from Lake Victoria, when
tion. But this is not the case. The immigration of foreign the predatory Nile perch (Lates niloticus) was intro-
species is a complex process and our knowledge con- duced into the lake. Nearly 200 of the formerly 300
cerning the mechanisms of the integration of the foreign species of endemic cichlids disappeared or are threat-
species is deficient. ened. The whole ecosystem changed dramatically. The
First, there are situations, in which the ecosystems disappearance of 13 waste-feeding cichlid species was
have enough space for immigrants and therefore no compensated by the shrimp Caridina nilotica. The
conflicts with the indigeneous species exist. Foreign shrimp was permanently present in the lake in low densi-
species penetrate frequently into water bodies which are ties but after the elimination of the cichlids it appeared in
greatly changed and therefore show significant losses of masses. Generally, the elimination of primary con-
original species. Recent changes are caused in most sumers among the fish species was substituted by pelag-
cases by anthropogenic influences, e.g. eutrophication ic crustaceans. The whole food chain was changed or
by wastes, destruction of banks and shores, installation strongly simplified (WITTE et al. 1992; GOLDSCHMIDT
of dams and reservoirs within streams, increased accu- et al. 1993).
mulation of fine sediments and so on. Foreign species Another example of conflict are the 1033 fish species,
can compensate species losses and colonize new struc- formerly living in Mexico, U.S.A. and Canada. From
tures precipitated by the anthropogenical changes. Fre- these species, nearly 68% have disappeared or are
quently the immigrants are very mobile euryoecious strongly threatened. One of the causes for this drastic
species that can quickly colonize the new habitats. One change may be the introduction of foreign species (WIn-
example is the Rhine River. It lost nearly 70% of its SON 1997). As in terrestrial systems, the naturalization of
zoobenthic species in the time from 1900 to 1970. By exotic species was an important factor in the extinction
about 1990, this species deficit was overcome by new of indigeneous species.
species among which 12 were neozoans (KINZELBACH Not only foreign predators, but also competitors can
1982, 1983). significantly effect ecosystemic changes. In Europe,
Natural events can also favour such immigrations Dreissena polymorpha is a well known neozoa. It colo-
(e.g., glacial periods, geological relief changes, high nized streams and lakes but has not impaired any species
water, long dry periods and volcanism). Particularly in or caused ecosystemic changes. This species occurs
streams, foreign species can immigrate after high water mainly as a space competitor on hard substrates that are
events with a strong wash-up effect and can anticipate apparently only weak settled. Stronger changes occurred
indigenous recolonisation (TITTIZER 1997). in Lake Erie, Canada, where Dreissena bugensis colo-
There are also water bodies - perhaps most - where nized fine sediments This immigrant has reduced the
ecological niches are occupied incompletely, for in- density of the tube-dwelling amphipod Diporeia hori and
stance as result of glacial periods. These niches can be some small mussels, but it has not eliminated them. Fur-
filled by foreign species. In all such cases, it does not re- thermore, the pseudofaeces of Dreissena have promoted
sult in ecosystemic changes. Examples are the neophyte some indigeneous species (DERMOTT& KERREC 1997).
Bunias orientalis in the Bj6rka-Kfivlinge River in Swe- When a foreign species establishes a key position a
den (KoHLERet al. 2000) and the North American turbel- total reconstruction of the ecosystem is possible. This
larian species Dugesia tigrina, today widely distributed applies, for instance, to the water hyacinth Eichhornia
in European lakes. In the last 100 years, many new crassipes. Originally, from the Amazonian region it was

Limnologica (2003) 33, 163-189

180 W. Sch6nborn

imported to Africa. In Africa it develops dense carpets These processes seem to respond automatically based
on the water surface, by darkening the whole water on existing chemical and biological potentials. Thus, for
body. The Amazonian region has antagonists of this instance, as nutrient contents increase, the density of
plant, inhibiting its expansion. algae is increased. This is connected with a CO 2 deficit
There are a lot of examples of foreign species immi- and - following chemical laws - leads to calcite precipi-
gration into water bodies, but no case of an ecosystemic tation, which takes algae and other particles downward
defence reaction which is directed against these species. and adsorbes phosphorus. There are dysoecisms which
If there are conflicts, two or more species are confront- show that not the whole system takes part in these pro-
ed, and the superior species wins the interaction. The cesses. In a way, denitrification in waters also can be re-
penetrating species will be eliminated if it is a weak garded as a protective reaction that takes place automati-
competitor or a preferred prey of indigenous predators. cally following bio-chemical laws. It is unknown
Another reaction is high densities of a foreign species whether the results of such reactions hide selective
in the early stages of an immigration, but later these den- mechanisms. Surely these mechanisms exist in micro-
sities retreat and remain at a low level. A well known ex- bial self-purification, but they originated in earliest bio-
ample is Elodea, the waterweed, today a normally logical evolution.
niched plant. The mechanism of this elastic reaction is Because it is necessary that life spaces (including ter-
unknown. restrial) are not filled with caracasses, parts of dead or-
Unlike eutrophication which is caused by abiotic fac- ganisms and other organic waste, natural selection has
tors and concerns all species, immigration is a conflict be- promoted those organisms that contribute to the preser-
tween two or some species. Eutrophication is a permanent vation of life spaces and thus to the basis of their own
danger for the whole ecosystem. Foreign species which life. Consequently, selection promoted the origin of
may lack general defensive mechanisms are unpre- saprophytic bacteria, fungi, and such animals as shred-
dictable. Evidently the mechanisms of the origin of defen- ders, coprophages, necrophages and detritivores.
sive reactions against eutrophication are not applicable to Without bacteriovorous animals, decomposition pro-
foreign species. Ecosystems rarely react as an entirety. ceeds very slowly. From existing predator-prey relation-
Mostly only single compartments react "egoistically", but ships, increasing eutrophication automatically stimu-
these reactions probably have an "altruistic effect". lates either the food-chain bacteria - ciliates - Chaeto-
Even more unpredictable than a single foreign species gaster - Erpobdella or analogous chains that effect de-
that causes only weak changes, is the total reconstruc- composition activity. It is unknown to what extent natu-
tion of an ecosystem by immigrating species. Further- ral selection has promoted decomposition and therefore
more, a defensive reaction against these immigrants the preservation of life spaces. The above named food
does not make any biological sense, because the geo- chains evolved with regard to their food, but it is not
graphic expansion of species is an important factor of clear if this benefits the whole ecosystem. It could also
evolution. be that automatism - free from selection - can have eu-
oecistic effects.

7. Origin and character of defensive systems. 7.2. Species-egoistic adaptations

Analogies and differences with system-altruistic effects
in regard to organisms
Species-egoistic adaptations with system-altruistic ef-
This chapter attempts to examine and theoretically dis- fects are defined as the adaptation of a species to its en-
cuss the statements about defensive reactions in limnetic vironment, which simultaneously promotes the preser-
ecosystems given in this paper and to make conclusions vation of the whole system. This happens in the follow-
based on this examination. Another aim is it to explain the ing ways.
possible origin of defensive systems. Additionally, the na- 1) Adaptation of a species with the aim to keep the en-
ture of defensive systems is made transparent by compar- vironmental conditions constant in their own inter-
ison with organismic defensive reactions to harmful ex- est, but thereby simultaneously benefit the whole
ternal influences, especially to morbific agents. system.
This form of adaptation could be found in algae, but
7.1. Ecosystemicimmunity also in the above mentioned saprophytic bacteria
and fungi. Cladophora glomerata accumulates nu-
Ecosystemic immunity is the activation of microbial and trients in a high degree and is therefore able to colo-
chemical processes (microbial self-purification, calcite nize oligotrophic stretches of streams. The increas-
precipitation and denitrification) by increasing organic ing nutrient level in the water promotes nutrient ac-
inputs. cumulation in connection with a mass production of

Limnologica (2003) 33, 163-189

 Defensive reactions of freshWaterecosystems 181

this algae so that the running water might become Plant belts can be considered analogous to the skin of
dammed. Therefore Cladophora has developed a organisms. This protective system seems to be devel-
detach mechanism that secures its further existence. oped by species-egoistic adaptations which have si-
Much of this detached biomass, including accumu- multaneous system-altruistic effects. Reed grows in
lated nutrients and xenobiotics, drifts downstream shallow waters, therefore in the littoral in large lakes.
so that the stretch will be relieved and the produc- Its protective effect against eutrophication is proba-
tion of this green alga will be newly stimulated. The bly favoured by selection (e.g. growth of densely
stability of the ecosystem increases, but, as showed lying rhizomes, high density of stalk root-aerating,
above, this increased stability is connected with and so on) because a quick eutrophication in connec-
dysoecisms. tion with siltation would eliminate the life space of
Benthic algae in standing waters show the same pat- reed.
tern. By mass production, they rise from the sedi- But the reed itself is important to siltation through its
ment and drift ashore. Mass production is not only mud rich with cellulose (high C/N-ratio) that is diffi-
stimulated by direct nutrient supply, but also the cult to decompose. The decomposers of cellulose,
CO2 produced by the decomposition. Drift, caused most of all Cytophaga, secrete slime which stabilizes
by wind, leads to a relief of the water body. The the mud of reed and thus favours siltation. During the
same applies for waterblooms and pleuston. In all siltation process, reeds must also compete against not
cases the species or communities, stopping the pro- only other macrophytes but also the development of a
cess of eutrophication, are useful to themselves. bog. Therefore, selection favours the protective func-
LINSENMAIR(1994) makes similar conclusions with tion of the reed to preserve the water body and simul-
reference to the stability of ecosystems. The stabili- taneously the inhibition of competeting vegetation.
ty originates indirectly by the selection of popula- The function of the alder zone along water's edge
tions that have positive effects on the persistence of can be interpreted similarly to the reed. The selec-
other populations. tion favours trees with hydrophilic roots, tops closed
2) Another adaptation mechanism is the filling of a in a canopy over small bodies of water and fallen
niche with positive effects to the whole system. Tubi- leaves with a low C/N-ratio. These mechanisms pre-
ficids in sediments, for instance, have filled such a vent breakage of banks and fill of brook beds or wa-
ecological niche. The worms are adapted to low oxy- terground. Therefore they prevent generally the loss
of the living area of trees. The combination of fallen
gen contents by intestinal respiration and possession
leaves, which are difficult to decompose and inten-
of haemoglobin. The uptake of sediment and diges-
sive solar radiation result in an eutrophication con-
tion of its microorganisms, as well as the aeration of
nected with weediness that can finish the existence
the sediment caused by its shifting, lead to a stimula-
of smaller running or standing waters relatively
tion of bacterial production, organic decomposition,
quickly.
and aeration of sediments.
The fungitoxic substances within the alder leaves
3) Many land animals, both herbivores and predators, protect the crayfish against fungus diseases. Cray-
are adapted to food seeking within water bodies. The fish have an important function in the preservation
utilization of water bodies as food and common of the ecosystem "brook", but it is difficult to under-
space by terrestrial animals leads - with many con- stand the selective co-evolution between alders and
trary processes - to a relief of the water ecosystem. crayfish. Therefore, the positive relationship be-
These animals relieve the water bodies because their tween alder and crayfish may be only accidentally.
defecation takes place completely or primarily on In this respect, the alder may be considered to be a
land. Probably selection has favoured this form of weak competitor. Alders are inferior in competition
defecation. In the case of defecation within the with other trees, primarily because they increase
water, the transparency would have been consider- dryness and aerate the soil. But crayfish dig long
ably decreased, thereby obstructing food seeking. caves, which they use as hiding places, into the
We can say that this relief system was brought into banks and between alder roots and thus provide the
the limnetic ecosystem but it was not developed in- alder rhizosphere with water (see Fig. 3).
ternally.
4) Euoecisms are also a form of adaptation, although it 7.3. Accumulation of xenobiotics
may be very difficult to understand their origin.
Alder zones and zones of emersed plants are poly- The accumulation of xenobiotics is an acquired defen-
functional, complicated compartments serving the sive reaction against toxic substances and is useful for
whole system. In the same sense deltas and estuaries the preservation of the system in accordance with the
are also considered to be gigantic protective systems principle formulated in chapter 7.2. A toxic substance is
for the sea. removed from the water by accumulation without harm-

Limnologica (2003) 33, 163-189

182 W. Sch6nborn

ing the accumulator. This makes ecological sense, par- 7.5. The importance of defensive reactions
ticularly in plants with a surplus of biomass that drifts in the preservation of limnetic ecosystems
downstream or ashore (Cladophora). But it applies only
during short-term contamination events because perma- It seems obvious that without microbial decomposition
nent loading makes accumulation meaningless in an of organic matter, no life on the earth would be possible,
ecological sense. due to the mass of dead bodies. Contrary to the other de-
It is possible that these mechanisms originally ap- fensive reactions described in this paper, only the micro-
peared during volcanic eruptions which influened the bial decomposition as a self-purification-mechanism is
surrounding waters. Different geological periods (e.g., well studied. These other defensive reactions probably
in the late Cretaceous) experienced high frequecies of do not reduce the degree of eutrophication, but they
eruptions. It is probable that most accumulation mecha- make its increase more difficult. This applies mainly to
nisms originated during these times. In this regard, the long-term loading processes, as they are commonly
eruptions of the volcano St. Helens (Washington, found in nature. But short-term loadings can be thor-
U.S.A.) may be considered as an example of the present oughly averted. Without these defensive reactions, many
period (WISSMARet al. 1982). water bodies would have a short existence. This applies
also to running waters, provided they are shallow and
flowing slowly. They can quickly develop weediness
7.4. Defensive reactions and stability and then silt up by depositing algal mats and pleuston
of ecosystems plants.
Defensive reactions against eutrophication can be inter- It may be possible to explain defensive reactions and
preted as stabilization mechanisms. In an ecological euoecisms by the help of a single species with an "altru-
sense, stability is difficult to define. In our case, stability istic" effect. DUNBAR(1972) published a theory accord-
refers exclusively to the preservation of the trophic level ing to which ecosystems are highly integrated unities
during increasing loading. that compete jointly and evolve as a unit. This may in-
After the cessation of nutrient input or the removal of deed play a role, but it could be also given another expla-
loading substances, waters remain on their trophic level nation. It may be that those water bodies have longer
for some time, beause different nutrients are stored dif- survived which have had protective features. Accelerat-
ferently (e.g., in sediments or organisms). This particu- ed siltation of standing and running water bodies would
larly applies to standing water with a long water-resi- have catastrophic consequences for the water balance of
dence time. This concerns the so-called "recovery time" the inland. Small standing waters - for instance pools
of the water, which cannot be used for a theoretical dis- and smaller water bodies - can be formed rapidly by
cussion of stability. Every eutrophication is a wear pro- changes in the relief of the earth surfaces or by large ani-
cess of the water body. mals. But it is doubtful whether this process will occur
The more defensive mechanisms a limnetic ecosys- in a shorter time than the physical extinction or ecosys-
tem has at its disposal, the more stable its state. But it temic destabilization when permanent eutrophication
could be shown that the defence against a disturbance takes place. When flowing water must arode a new way
factor simultaneously effects a detriment to other com- after siltation of a lowland stream, it needs time, but for
partments of the ecosystem, causing instability. As ex- the formation of really important water bodies, periods
plained above the whole system never reacts to distur- of geological folding processes or ice ages may be nec-
bance factors as an entirety, but only in single compart- essary.
ments, that is 'dysoecismically and therefore incom- Even the sea puts out by breakers great quantitites of
pletely'. Even normal microbial self-purification can algae and animals. But it is unknown to what extent the
impair the ecosystem by oxygen consumption and the wrack-masses relieve the seashore.
release of the toxic by-products of decomposition. De-
spite of the close integration of an ecosystem, only
those single populations react that have developed
7.6. Analogy with organismic defensive reactions
adaptation mechanisms. This is, however, sufficient for It may be a dubious attempt to draw parallels between
the general stabilization of the system. The conditions ecosystems and organisms, but it is helpful in under-
can be very complicated. According to KALBE(1996), standing defensive reactions in limnetic ecosystems.
the ranges of stability are broader with increasing troph- The comparison with the immune system has already
ic status, but the transitions will be instable: oligotrophy been demonstrated. In the case of aquatic ecosystems,
- eutrophy - hypertrophy are stable stages; mesotrophy there are latent potentialities that are mobilized automat-
- polytrophy - and all conditions more than hypertro- ically by eutrophication (chemical processes) and are se-
phy are instable stages until finally the ecosystem loses lectively developed (microbial processes). Another com-
its function. parison can be made to algal masses washed ashore, the

Limnologica (2003) 33, 163-189

 Defensive reactions of freshwater ecosystems 183

so-called "algal expectoration". This process is similar are defensive reactions, it is also possible to similarly
to a cough, a head cold, vomitings and expectoration - understand algal masses that wash ashore.
usually considered as animal or human diseases. These There are no parallelisms between the origin of the
are defensive reactions of the body to get rid of morbic defensive reactions of organisms and those of limnetic
agents. In the case of algae, these are high concentra- ecosystems. Both reactions have similar results but are
tions of nutrients and xenobiotics. systemspecific.
Like some organisms have a liver, a limnetic ecosys-
tem has "organs" of detoxification in the form of xenobi-
otics-accumulating plants, most frequently Cladophora 8. Human support of the defensive reactions
and water mosses, but also many other plants. of water bodies against eutrophication
Just as the skin or membranes of organisms serve to
protect against foreign influences, reed belts and border Human intervention can support the defensive reactions
trees perform a comparable function in limnetic ecosys- of water bodies against eutrophication. The most impor-
tems. Like skin that not only protects and demarcates or- tant measures are:
ganisms but also enables a selective exchange of matter, - Algal mats on water surfaces and algae washed ashore
the alder belt and belt of emersed plants control not only should be removed. Drifting CIadophora in streams
the uptake of CO2, the aerating of sediments and the fix- should be caught and removed. For this purpose, spe-
ation and transport of nitrogen into aquatic systems, but cial nets for skimming and catching should be con-
also release of nitrogen by denitrification. structed.
In the same manner as organisms possess special pro-
- Mass developments of neuston and pleuston (Lemna,
tective mechanisms for entryways into the body (e.g.,
Eichhornia and others) should be removed.
antibacterial substances in saliva, nose, tears and ears;
- Support terrestrial outlet systems (protection of otters,
gastric acid), entryways into limnetic ecosystems are
ecological management of cormorants; protection of
also protected. Springs, for instance, precipitate sub-
all animals which take part in defence mechanisms).
stances from the earth's interior (calcium, iron, man-
- Preserve eels and amphibians.
ganes), therefore these minerals reach only a low con-
- Induce calcite precipitation in such a way that the
centration in streams and lakes. Large lakes and oceans
sinking crystals do not reach the benthal. There are al-
are protected by deltas and estuaries which accumulate
ready artificial calcite treatments of lakes (MURPHY&
many dissolved and particulated substances transported
PREPAS 1990; DITTRICHet al. 1995), but without re-
by streams. The origin of such protective systems is dif-
gard to the benthic zone.
ficult to explain [lucky chance or selection in the sense
- Protect water mosses (xenobiotics accumulators with-
of DUNBAR(1972)?].
out drift mechanisms).
The most conspicuous parallelism between organisms
- Protection and renaturation of the alder belts and belts
and limnetic ecosystems is aging. This relates primarily
of emersed plants in the water bodies and along the
to lakes but running waters can also age, however in a
changed manner in consequence of their kinetic energy banks.
or their continuum. Although the aging mechanisms of - Reservoirs and other artificial water bodies should be
lakes are totally different in comparison to organisms, rich in bights and reed habitats.
over feeding leads, in both systems, to an acceleration of The removal of plants (mostly algae, neuston and
aging. Underfed organisms and oligotrophic lakes live pleuston) that underlie the detach-and-drift mecha-
longer than overfed bodies or hypertrophic waters, re- nisms mimics natural ecosystemic processes. But this
spectively. does not apply to other plants, for instance a removal of
As already described, defensive reactions of ecosys- plants from the littoral zone will damage the whole
tems can impair some compartments of the ecosystem. ecosystem.
These so-called dysoecisms also show conspicuous par-
allelisms to organisms. For instance, the immune system
of an organism frequently shows overreactions whereby Summary
parts of its own body can be attacked. Examples include
multiple sclerosis, rheumatism, arthritis, diabetes and an - Nutrients and xenobiotics reach the element cycle
acceleration of aging. Fever, as a defensive reaction, can more quickly in water bodies than in terrestrial sys-
harm the body. Allergies are defensive reactions against tems. Therefore, aquatic ecosystems develop particu-
foreign substances, but the reactions overshoot the mark lar defensive mechanisms against organic and toxic
and can even cause diseases. loadings.
Like a cough, sneezing and expectoration are fre- - Large animals (big game), since about the Mesozoic,
quently interpreted as autonomous diseases but in reality have been significant causes of eutrophication in

Limnologica (2003) 33, 163-189

 184 W. Sch6nborn

aquatic systems. Their influence has caused abundant - Simultaneously, the leech ErpobdelIa octoculata de-
hypertrophy and continues at present in regions with velops high densities in the clumps and becomes a
high densities of elephants, buffalos, hippopotamuses centre of predation. Its development stages (3-60
and crocodiles. ram) are all carnivorous and control nearly the whole
- Geological periods of increased volcanism may result meio- and macrozoobenthos. These processes lead to
in higher toxic impacts on water bodies. Freshwater a centralization of the ecosystem as a response to in-
ecosystems have had long time periods to develop de- creasing trophic state. This may lead to a bundling of
fensive reactions against organic and toxic loadings. reactions contributing to self-purification.
- A well known defensive reaction of aquatic ecosys- - Eutrophication promotes the density of bacteria. It
tems is microbial self-purification. comes to a linear energy-and-matter flow system
- Many benthic algae accumulate nutrients and xenobi- (e.g., bacteria - ciliates - Chaetogaster - Erpobdella),
otics and tend toward mass production. In running which keeps the exponential growth phase of the bac-
water, they can detach by high current velocities and terial populations as a stimulation to their production
drift downstream. In standing water, they rise to the and an intensification of the decomposition of organic
surface and drift ashore by wind. These processes un- matter. This process is a part of centralization, too.
burden (at least) one of the ecosystem compartments - The accumulation of heavy metals and other noxious
(a stream stretch or the open water zone). In standing matter is another form of defensive reactions. The ac-
water in these processes are also involved in the lake cumulated matter can be drifted together with the
bloomings, neuston and pleuston. plants or microorganisms (e.g., Cladophora, Sphaero-
- There are important terrestrial outlet mechanisms by tilus), or isolated, as in the case of the detach- and
food chains from freshwater organism to terrestrial consume-resistant water mosses.
consumer. Examples: Terrestrial consumers of mi- - Resuspension of sediments and denitrification in
croorganisms, algae, pleuston and small animals (e.g. water bodies are also part of the disburdening process.
ducks); consumers of waterplants (many birds and - The zone of alders and emersed plants along running
mammals); terrestrial predators capturing (primarily) and standing waters protects water bodies from eu-
fish and decapods. Fish and decapods eat a lot of trophication comparable to the skin of organisms that
algae. Their accumulated nutrients can be transported protects against penetrating xenobiotic matter. Like
out of the aquatic environment. The most important the skin, this double protection layer of aquatic sys-
predators are cormorants, certain species of storks and tems is also a multifunctional defence system.
otters. Amphibians also take many material outward - There are features and structures of ecosystem com-
after their metamorphosis. partments that contribute to the utility and protection
- There are also marine outlet systems, for example of the whole system. This phenomenon can be named
eels. But they seem to be not very effective. "euoecism". On the other hand, there are also "dysoe-
- Filtering mechanisms can also unburden water bodies cisms". Mostly they arise when defensive reactions
such as the mechanical filtering of seston in intersti- adversely affect other compartments. But inadequa-
tial spaces of plant clumps (e.g., Cladophora, Myrio- cies of defensive reactions are also dysoecisms.
phyllum) and the biological filtering of filter feeders - Aging of water bodies, connected with eutrophica-
(Cladocera, big mussels and other groups). Filter tion, is an irreversible process (like the aging of or-
feeders are particularly effective when combined with ganisms) and causes morphometric changes leading
one of the outlet systems. to the "death" of the water bodies. But defensive reac-
- Planktonic calcite precipitation also removes biomass tions can retard this process.
and nutrients (phosphorus) from the open-water zone - There are no defensive reactions against the introduc-
and seem to be a natural self-purification process. tion of foreign species (neophytes and neozoa) into
- Eutrophication leads to a concentration of vegetable water bodies. The intruder faces one or more resident
biomass and nutrients. This may facilitate their re- species. As opposed to situations of eutrophication,
moval. Means of removal are high water discharge (in most conflicts do not concern the whole ecosystem.
running water), wind (in standing water), and herbiv- But there are cases in which the invasive species
orous terrestrial animals (through "skimming" or catastrophically affect flora and fauna (e.g., Lake Vic-
grazing). The effect of this concentration is especially toria, Africa). Furthermore, the occurrence of foreign
evident in the green alga Cladophora. The mass pro- species is not foreseeable (eutrophication is a perma-
duction of Cladophora yields a centre of primary pro- nent danger), therefore evolution could not effectively
duction and habitats (many animals colonize the operate. An exception is the case in which the inva-
clumps, especially larvae of chironomids, which sion is stopped by indigenous predators.
transform the detritus, filtered by a filament network - Microorganisms in particular possess features or activi-
of the clumps in fixed tubes). ties as well as processes founded on chemical laws that

Limnologica (2003) 33, 163-189

 Defensive reactions of freshwater ecosystems 185

automatically counteract eutrophication (e.g., self-pu- Examples:

rification, calcite precipitation, denitrification). These • Self-purification, denitrification and calcite precipi-
processes are comparable to the immune systems of or- tation are comparable to the immune system.
ganisms, but can function only in combination with mi- • Concentration of algal masses and their removal (on
croorganisms within a food chain (predator-prey-rela- banks and shores and in the bights as "sputum
tionships). These processes can be the result of selec- rooms") is comparable to phlegm and expectora-
tion in the sense that water bodies with such systems tion.
have slower siltation and are therefore, less prone to ex- • Accumulation of noxious matter by plants is com-
tinction. But this is generally difficult to establish, parable to organs of decontamination (e.g., liver).
exept in the case of saprophytic bacteria which are sure- • Processes in springs and river mouths are compara-
ly promoted by selection to save the aquatic system ble to protection of the entrance portals of the body.
from being filled by dead organic material. • Zones of alders and emersed plants are comparable
- Many defensive reactions of water bodies are founded to skin.
on species-egoistic adaptations with an (accidentally) • Disturbances in the matter cycle by strong eutrophi-
system-altruistic effect. This applies especially to de- cation (hypertrophy) are comparable to the increase
fensive systems based on algae, but also to the self- of diseases by over nutrition.
purification by saprophytic microorganisms in earli- • Dysoecisms are comparable to overreactions and
est times and also to euoecisms. The adaptation of inadequacies of organismic defense mechanisms.
species is egoistic because it counteracts the eutrophi- The origin and mechanisms of defensive reactions are
cation only to keep the species' own favourable living completely different from those of organisms.
conditions, or in the case of saprophytic microorgan- - Human intervention can support the natural defensive
isms to keep their living space. There are also annida- reactions in freshwaters with the help of relatively
tions of species with positive effects on the whole sys- simple techniques.
tem (e.g., Tubificidae).
- In most cases, the described defensive reactions could
not be developed from the ecosystem, but it seems Zusammenfassung
that they have been "caught" by the water bodies,
which is especially clear in the case of terrestrial out- - N~ihrstoffe und Schadstoffe gelangen in Gewfissern
let systems. schneller in den Stoffkreislauf als in terrestrischen
- The accumulation of noxious matter (e.g., heavy met- Systemen. Daher haben Gew~isser-Okosysteme gegen
als) of certain plant species, which itself is probably eine l~lberlastung besondere Abwehrreaktionen ausge-
an acquired defensive reaction against toxic loadings, bildet.
is primarily an egoistic phenomenon that nevertheless - Grogtiere waren und sind extreme Eutrophierungs-
serves the preservation of the entire ecosystem. The faktoren fiir die Gew~isser, zumindest seit dem Meso-
reaction may have originated in geological periods of zoikum. Ihr Einflug ftihrt noch heute zur Gewfisser-
high volcanism. Hypertrophie in Gebieten mit einer hohen Dichte an
- Every eutrophic event wears down the ecosystem. In Elefanten, Btiffeln, Flusspferden und Krokodilen. In
the course of eutrophication, usually several ecosys- geologischen Zeiten eines erh6hten Vulkanismus
tems disappear, but without leaving behind corpses. wird es auch zur stiirkeren toxischen Belastung der
- The more a freshwater ecosystem possesses defensive Gewiisser gekommen sein. Die Gewfisser-Okosys-
mechanisms, the more stable is its state with regard to teme hatten also lange Zeitr~iume zur Verftigung, um
trophism. An ecosystem does not react as a unit, but Abwehrreaktionen gegen die Eutrophierung und
by single species (populations) that have a stabilized anorganische Belastung zu entwickeln.
efficacy that supports the whole system. - D i e bekannteste Abwehrreaktion der Gew~isser-
- For the global water balance, it is important that fresh- Okosysteme ist die mikrobielle Selbstreinigung.
waters are not too quickly silted up by sediments - Viele benthische Algen speichern N~ihr- und Schad-
(caused mostly by decomposed waterplants). There- stoffe und neigen zur Massenentwicklung. Sie werden
fore, defensive reactions against eutrophication have in Fliel3gew~issernbei erh6hten Durchfltissen abgeris-
an existential importance for a heritable earth surface, sen und abgetrieben oder sammeln sich in Stand-
because these defenses restrain siltation and lead to a gew~issern an der Oberfl~iche und werden dann vom
selective advantage. Wind an das Ufer oder dartiber hinaus gesp01t. Hier-
- A n a l o g i e s of defensive reactions of freshwater durch wird zumindest ein Kompartiment (z. B. Fluss-
ecosystems with those of organisms are quite possi- strecke, Freiwasser in Standgew~issern) entlastet. In
ble, but only in regard to their effects, not to their Standgewfissern nehmen daran auch Wasserbltiten,
manner of origin. Neuston und Pleuston tell.

Limnologica (2003) 33, 163-189

 186 W. Sch6nborn

- Von groger Bedeutung sind auch die terrestrischen Ciliaten - Chaetogaster - Erpobdella) in der expo-
Ausleitsysteme mittels Nahrungsketten: Wasser- nentiellen Wachstumsphase ihrer Populationen,
organismus ~ terrestrischer Konsument. Zu nennen wodurch ihre Produktion und Abbauleistung stim-
sind Mikroorganismen, Algen- und Pleustonfresser uliert wird. Auch diese Vorg~nge geh6ren zur Zentral-
(z.B. Enten), Wasserpflanzenfresser generell (viele isation des Okosystems.
Vogel- und Sfiugerarten) und terrestrische Prfidatoren, - Die Akkumulation von Schadstoffen durch Pflanzen
die vor allem Fische erbeuten. Fische und auch Grog- ist ebenfalls eine Abwehrreaktion. Die akkumulierten
krebse konsumieren groBe Mengen an Algen, deren Schadstoffe, z.B. Schwermetalle, werden entweder
gespeicherte Stoffe durch die Pr~idatoren nach augen mit den abdriftenden Pflanzen bzw. Mikroorganismen
gelangen. Die wichtigsten derartigen Prfidatoren sind (Cladophora, Sphaerotilus) entfernt oder wie im Falle
Kormorane, bestimmte Storcharten und Otterarten. der abflussresistenten und kaum konsumierbaren
Auch Lurche nehmen tiber ihre wasserbewohnenden Wassermoose, isoliert.
Larven nach der Metamorphose viele Stoffe mit nach - Auch Resuspension von Sediment und Denitrifikation
augen. in Gew~issern sind entlastende Vorg~inge.
- Weniger ergiebig als terrestrische scheinen marine - Der Erlengt~rtel an FlieB- und Standgew~issern und die
Ausleitsysteme zu sein, repr~isentiert vor allem durch Zone emerser Pflanzen (vor allem Schilf) scht~tzen
die Aale. Gewfisser enorm vor der Eutrophierung (vergleichbar
- Auch Filtriersysteme entlasten die Gewgsser: mecha- der Haut von Organismen gegen eindringende Stoffe).
nische durch Filtration von Seston in interstitial- Diese doppelte morphologische Schutzschicht ist ein
reichen Pflanzenbtischeln (z.B. Cladophora und multifunktionelles Abwehrsystem gegen eindrin-
MyriophyIlum) wie auch organismische durch Auf- gende Stoffe.
nahme partikul~irer Nahrung aus dem Freiwasser - Es gibt viele Eigenschaften yon Teilsystemen, die sich
(Cladoceren, Grogmuscheln und Vertreter vieler an- zum Schutz und Nutzen des Gesamtsystems erg~in-
derer Gruppen). Filtrierer sind besonders leistungs- zen. Sie werden als Eu6kien bezeichnet. Dem-
stark far die Entlastung, wenn sie an die genannten gegenfiber gibt es auch Dys6kien. Die meisten yon
Ausleitsysteme gekoppelt sind. ihnen entstehen, wenn die Abwehrreaktionen eines
- Bei der planktischen Kalzitf~illung wird ebenfalls Kompartiments des Okosystems zu Lasten eines an-
Biomasse aus dem Freiwasser entfernt, indem sie mit deren gehen. Zu den Dys6kien geh6ren auch viele
den sinkenden Kristallen abw~rts gerissen wird. Auch Unvollkommenheiten der Abwehrreaktionen.
der sich an den Kristallen adsorbierende Phosphor - Die Alterung der Gew~isser, bei der die Eutrophie zu-
wirkt ft~r das Pelagial entlastend. nimmt, ist, wie bei den Organismen, ein irreversibler
- Die Eutrophierung bewirkt eine Konzentration der Vorgang, bei dem sich im Falle der Seen die mor-
pflanzlichen Biomasse und der N~ihrstoffe und macht phometrischen Relationen ~indern. Abwehrreaktionen
sie somit zugleich leichter entfernbar. Medien sind die verz6gern diesen Vorgang.
steigende Wasserft~hrung (FlieBgewfisser), der Wind - Gegen das Eindringen von Fremdarten (Neophyten,
(Standgew~isser) und die pflanzenfressenden Tiere Neozoen) in ein Gew~isser gibt es keine Abwehrreak-
(Absch6pfer, Ab~iser). Der Konzentrationseffekt ist tionen. Hierbei stehen sich zun~chst immer zwei oder
besonders infolge ihrer Massenentwicklung an mehrere Arten konfliktm~iBig gegent~ber. Ein Ein-
Cladophora ausgebildet. Sie wird zu einem Zentrum dringling ist zun~ichst kein das Gesamtsystem betref-
der Primfirproduktion und auch zu einem Habitat- fender Faktor wie die Eutrophierung. Uberdies sind
zentrum (viele Tiere, vor allem Chironomidenlarven, Eindringlinge stets unvorhersehbar gewesen, so dass
die den yon den Thalli gefilterten Detritus zu festen sich die Evolution nicht darauf einstellen konnte. Eine
R6hren umbauen). Gleichzeitig entwickelt sich der Ausnahme machen einheimische R~iuber, die die Ein-
Egel Erpobdella octoculata dutch enorme Dichte- dringlinge vernichten. Das Gesamtsystem reagiert
zunahme zu einem Pr~dationszentrum. Alle seine Sta- fast hie (auch nicht bei der Eutrophierung), sondern es
dien (3-60 ram) sind r~tuberisch und kontrollieren fast sind immer nur Teilsysteme oder einzelne Arten, die
das gesamte Meio- und Makrozoobenthos. Diese dann (vermutlich meist zuffillig) eine "altruistische"
Vorg~inge ftihren zu einer Zentralisation des Okosys- Wirkung haben.
terns als Antwort auf die Trophiezunahme, zugleich - Mikroorganismen besitzen seit langen geologischen
zu einer Bt~ndelung der an der Selbstreinigung Perioden Eigenschaften, die der Eutrophierung (au-
beteiligten Krfifte. tomatisch) entgegenwirken (Selbstreinigung). Ahn-
- Mit der Eutrophierung steigt infolge der Konzentra- liches gilt ftir Prozesse, die auf chemischen Gesetz-
tion der organischen Stoffe die Dichte der Bakterien m~iBigkeiten beruhen, z.B. Kalzitffillung und Denitri-
an. Sie verbleiben dank der Ausbildung eines linearen fikation. Sie wirken wie ein Immunsystem, das yon
Energie- und Stoff-FluB-Systems (z.B. Bakterien - den Gew~issern "eingefangen" wurde. Ftir das Funk-

Limnologica (2003) 33, 163-189

 Defensive reactions of freshwater ecosystems 187

tionieren der Selbstreinigung ist allerdings noch eine Beispiele ftir solche Analogien:
an die Mikroorganismen sich anschliegende Nah- • Die Selbstreinigung, Kalzitffillung und Denitrifika-
rungskette (R~iuber-Beute-Verhfiltnisse) verantwort- tion sind mit dem Immunsystem vergleichbar.
lich. Dies kann das Ergebnis einer Selektion sein, • Die Ansammlung von Algenmassen und deren
indem Gew~isser mit derartigen Systemen weniger Ausstof3 - wobei Gew~isserbuchten und Ufer als
schnell verlandeten, ist aber nicht eindeutig nachzu- "Sputumkammern" fungieren - sind mit der
vollziehen. DaB jedoch die saprophytischen Bakterien Schleimbildung und dem Auswurf vergleichbar.
in unseren Gew~issern selektiv gef6rdert wurden, ist • Schadstoffakkumulierende Pflanzen k6nnen mit
stark zu vermuten. Entgiftungsorganen verglichen werden.
- In vielen Ffillen beruhen die Abwehrsysteme der • Vorg~ingein Quellen und Flussmtindungen sind mit
Gew~isser auf artegoistischen Anpassungen mit zuf~il- dem Schutz der Eintrittswege in den K6rper ver-
ligen system-altruistischen Auswirkungen. Dies gilt gleichbar.
vor allem neben den genannten saprophytischen Bak- • Eden- und Emersengtirtel lassen sich mit der Haut
terien auch fiir die auf Algen basierenden Abwehrsys- vergleichen.
teme. Diese Anpassungen einer Art sind ,,egoistisch", • St6rungen im Stoffhaushalt durch Eutrophierung
weil sie der Eutrophierung nur entgegenwirken, um (--+ Hypertrophie) k6nnen mit der Zunahme von
die ft~r die Art gtinstigen Lebensbedingungen zu er- Krankheiten durch Oberern~hrung verglichen wer-
halten. Es gibt auch Anpassungen einer Art in einer den
Nische mit positiven Auswirkungen far das Gesamt- • Dys6kien sind mit der Uberreaktionen und Unvoll-
system (z.B. Tubificiden). In vielen F~illen scheinen kommenheiten organismischer Abwehrmechanis-
derartige Verh~iltnisse aber nicht vom CIkosystem men vergleichbar.
selbst entwickelt, sondern von ihm nur "eingefangen" - Der Mensch kann durch relativ einfache Verfahren die
zu sein. Auch die Eu6kien sind vermutlich durch eine nattirlichen Abwehrreaktionen der limnischen ()ko-
artegoistische Selektion mit (zun~ichst zuf~illiger) sys- systeme untersttitzen.
tem-altruistischer Wirkung entstanden.
- Die Akkumulation yon Schadstoffen (z.B. Schwer-
metallen) ist ebenfalls eine erworbene Abwehrreak- Acknowledgements
tion gegen toxische Stoffe, die dem Erhalt des
Okosystems nutzt, die prim~ir aber artegoistischer I thank Andrew Davis and David McFarland for improving the
Natur ist. Diese Reaktion wird in frt~heren geologi- English style and helpful comments. Special thanks must be
schen Zeiten mit erh6htem Vulkanismus entstanden given to Anita Lange for their constructive discussions and
sein. close cooperation in preparation and improvement of this
- Jede Eutrophierung ist ein Verbrauchsprozeg des manuscript. Many colleagues who have given valuable sug-
Gew~ssers. Im Verlauf der Eutrophierung verschwin- gestions and remarks is gratefully acknowledged.
den meist mehrere 0kosysteme, aber ohne Leichen-
bildung. Je mehr ein limnisches (Jkosystem tiber Ab-
wehrmechanismen verfiigt, um so stabiler ist es, bezo- References
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