Acta Oecologica 24 (2003) S281–S288
www.elsevier.com/locate/actoec
Original article
Comparison of bacterial and phytoplankton productivity in extremely
acidic mining lakes and eutrophic hard water lakes
Brigitte Nixdorf *, Hartwig Krumbeck, Jörn Jander, Camilla Beulker
Brandenburg University of Technology Cottbus, Head of Water Protection, D-15526 Bad Saarow, Seestrasse 45, Germany
Abstract
Two types of standing waters are important in Brandenburg (Germany): eutrophic hard water lakes of glacial origin (about 2.800) and
several hundred mining lakes, mostly highlyacidic (pH between 2 and 3 and high concentrations of dissolved iron). A trophic gradient can be
used, describing the nutrient situation in these lakes from oligo- to mesotrophic mining lakes to eutrophic to polytrophic natural hard water
lakes. Bacterial and algal production in some typical lakes of these two types are compared. Whereas bacterial productivity in acidic mining
lakes is comparable with eutrophic hard water lakes, primary production is very low. It does not exceed the level of bacterial production
considering volume related daily carbon production. Mining lakes are thereby characterised as heterotrophic ecosystems. Reasons for the
production differences in both types of lakes can be looked for in the resource availability and top down control. Contrary to the eutrophic
natural lakes, phytoplankton productivity in mining lakes is controlled by carbon limitation. Consequences for the structural classification of
pelagic bacterial and phytoplankton communities are shown concerning the course of ecosystem succession from pioneer initiation in mining
lakes to high-biomass maturation stage in eutrophic hard water lakes.
© 2003 Published by Éditions scientifiques et médicales Elsevier SAS.
Keywords: Phytoplankton primary production; Bacterial production; Trophic cascade; Eutrophic lakes; Acidic waters; Resources
1. Introduction
About 2.800 eutrophic hard water lakes of glacial origin
and several hundred mining lakes, mostly highly acidic, exist
in Brandenburg (Germany). Especially, these newly formed
mining lakes offer a remarkable opportunity for limnologists
to investigate planktonic primary successions and productivity patterns in lakes. They are diverse in morphometry, mixis
regime and hydrochemistry. A typisation of acidic lakes is
given in Geller et al. (1998). They are among the largest and
most acidic lakes in Germany. The pH and total acidity (base
capacity KB4.3) range from 2.6 to 4 and 0 to 35 mmol l–1,
respectively.
The lake chemistry is the main determinant for the phytoplankton composition in the mining lakes (Nixdorf et al.,
1998a), but not in the natural lakes in the German lowlands.
For both types of standing waters, the supply of major nutrients mainly determines the primary productivity and thereby
the level of algal biomass. Productivity in mining lakes is
limited by phosphorus and—this is a paradox in coal mining
* Corresponding author.
E-mail address: [email protected] (B. Nixdorf).
© 2003 Published by Éditions scientifiques et médicales Elsevier SAS.
DOI: 1 0 . 1 0 1 6 / S 1 1 4 6 - 6 0 9 X ( 0 3 ) 0 0 0 3 1 - 6
lakes—by carbon (Lessmann and Nixdorf, 2002; Nixdorf
and Kapfer, 1998; Nixdorf et al., 2001). At pH 3, the atmospheric equilibrium concentration of CO2 is approximately
0.1 mg l–1 (Stumm and Morgan, 1996). The extremely acidic
limnetic ecosystems (pH < 3) are very sensitive and show
high fluctuations depending on the chemistry and hydrology
of the groundwater filling them.
The mining lakes are colonised by planktonic organisms
at an oligotrophic or mesotrophic level with dominant algal
taxa belonging to the Chrysophyceae, Chlorophyta and Dinophyceae (Beulker et al., 2003 (this volume); Lessmann and
Nixdorf, 2000; Nixdorf et al., 1998b). These taxa are sometimes found in considerable quantities and untypical vertical
patterns indicating a remarkable potential for primary production. They may respond to changes in a biotic conditions
with algal mass developments often in the hypolimnion or
near the sediment comparable with those of eutrophic conditions in neutral hard water lakes (Nixdorf and Hemm, 2001;
Steinberg et al., 1999). Eutrophic hard water lakes are more
influenced by morphometry and especially shallowness with
an intensive sediment water interaction resulting in eutrophy
or hypertrophy (Deneke and Nixdorf, 1999; Nixdorf and
Deneke, 1997). These conditions favour the mass develop-
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ment of cyanobacteria, some of them with remarkable toxin
production (Chorus, 2001; Mischke, 2003 (this volume);
Wiedner et al., 2001).
Bacterial and phytoplankton production were measured in
selected artificial mining and natural lakes to compare the
intensity of production and to describe the relationship between the two main production processes within the microbial plankton across a trophic gradient.
2. Study site
Three of the investigated natural lakes are situated in the
Scharmützelsee region (57°20’N, 54°35’E) in Brandenburg
(Germany), about 60 km East from Berlin (see Table 1, for
more details) (Deneke and Nixdorf, 1999; Nixdorf and
Deneke, 1997). These lakes originate from the Weichsel
glacier and are located at the border of the southern plateau of
the Berlin glacial valley. The catchment area is dominated by
forests and agriculture with increasing importance of tourism. Lake Wolziger See (WOL) and lake Scharmützelsee
(RIE) with a mean depth of 5.5 m (WOL) and 9.0 m (RIE),
respectively, are dimictic. Lake Langer See (LAN) is
polymictic. All lakes are in a eutrophic to hypertrophic state
(Table 1; OECD, 1982).
The region of Lusatia comprises the south-eastern part of
Brandenburg and the north-eastern part of Saxony in East
Germany (Fig. 1). The standing waters in the region have
their origin in open-cast lignite mining activities since the
end of the 19th century. As a consequence of the intensive
exploitation and the drastic reduction of the brown coal
production after 1990, a number of holes were filled with
water (mainly re-rising groundwater and water from rivers).
Owing to the geogenic potential for acidification due to
pyrite oxidation, most of these lakes are extremely acidic (pH
< 3.5), rich in iron and sulphate. For our investigations, a
number of mining lakes were selected and measurements
were carried out sporadically for bacterial cell counts and
production in the following regions: Frankfurt/Oder (nonacidic lakes Helene and Katja), region Schlabendorf (Lichtenau, RL B, Seese RL 104), Grünewalde/Plessa (Lake Plessa
113, 107 and 111, Lake Grünewalde), Senftenberg (Koschen,
Sedlitz, Skado), Drebkau/Vetschau (Gräbendorf), Muskauer
Faltenbogen (Waldsee, Felixsee) region Hoyerswerda
(Lohsa II, Kortitzmühle, Dreiweibern). Lake Grünewalde
Fig. 1. Map of the investigated areas in Brandenburg and lake regions.
Natural hard water lakes belong to the Scharmützelseeregion (no. 8, near
Bad Saarow). Main investigations in the mining area were carried out in the
region Grünewalde/Plessa (no.3).
(Plessa 117) and Lake Plessa 111 in the region
Grünwalde/Plessa were regularly investigated. All mining
lakes are acidic and mainly oligo- or mesotrophic. Only Lake
113 is eutrophic due to some management measures based on
addition of substrates.
3. Materials and methods
Water samples for chemical and biological analyses (phytoplanktonic composition and biovolume as well as primary
production, chlorophyll a, bacterial abundance and production) were taken biweekly (natural lakes), monthly (mining
lakes in the Plessa region) or irregularly (other mining lakes)
as mixed samples from surface to bottom during mixing or as
vertically integrated samples from the epi- and meta/hypolimnion with a 2.3-l LIMNOS-sampler. The sampling
points were generally above the deepest points where the
Secchi depth also was estimated. Depth profiles of tempera-
Table 1
Morphometric and trophic data of three investigated lakes in the Scharmützelsee region and mining lake Grünewalder See (Plessa 117) in Lusatia (Zmax:
maximum depth, A: lake area, V: lake volume, CA: catchment area, SD: Secchi depth, Chl. a: chlorophyll a, TP: total phosphorus, TIC: total inorganic carbon).
SD, Chl a and TP concentrations are given as 3-year means (1998-2000) in the mixed water layer. Trophic state according to OECD (1982) (di: dimictic, poly:
polymictic, eu: eutrophic, hyper: hypertrophic)
Lake
Abbrv. Mixis
Trophic state
Zmax(m)
A (km2)
V(×106 m3)
CA(km2)
Scharmützelsee
Wolziger See
Langer See
Grünewalder See
(Plessa 117)
RIE
WOL
LAN
117
Eu
Hyper
Hyper
Oligo–meso
29.5
13.0
3.8
14
12.09
5.79
1.55
0.95
108.23
32.02
3.27
6.70
112
382
395
Di
Di (poly)
Poly
Di
SD
(m)
1.74
2.39
0.54
8.0
Chl a
(µg l–1)
25.1
20.1
99.0
3
TP
(µg l–1)
52
70
116
7
TIC
(mg l–1)
15–30
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B. Nixdorf et al. / Acta Oecologica 24 (2003) S281–S288
ture, oxygen (absolute and relative as saturation), pH, redox
potential, conductivityand since 1996, also chlorophyll fluorescence were measured at 0.5 m intervals by means of a
HYDROLAB 20 probe, a Haardt fluorescence probe and a
field computer (HUSKY Hunter).
The different fractions of anions (among them, the nutrients such as phosphate, nitrate, silicate) and cations, dissolved or total inorganic carbon (DIC or TIC) and dissolved
organic carbon (DOC) and metals (Fe, Al, Cu, Cd, Mn, Co,
Ni) were determined according to the recommendations
given in the “Limnological method for investigation of mining lakes” (Schultze et al., 1994) and according to the standard methods (Deutsche Einheitsverfahren zur Wasser, 19761998). For the estimation of the base capacity (KB) or acid
capacity (KS), samples were titrated with 0.02 mol NaOH or
0.01 mol HCl.
Cell numbers of bacteria were estimated from a subsample of the production measurements after Porter and Feig
(1980). The samples were fixed with formaldehyde (final
concentration 10%) and 1–2 or 5–10 ml were filtrated on
black Nuclepore PC filters (pore size 0.2 µm) for neutral or
acidic lakes, respectively. After rinsing and drying, the filters
were imbedded and stained in a DAPI-buffer-suspension
(after Hentschke, personal communication) and counted by
epifluorescence technique. For this study, only the total number of bacteria were considered.
Bacterial production was estimated by measuring the incorporation of methyl[3H]thymidine (740 GBq mmol–1) into
cellular macromolecules at a concentration of 83 nmol. Duplicates of 3 ml and one formaldehyde fixed control (2.3%
final concentration) were incubated in the dark for 1 h in an
incubator at in situ temperature in polycarbonate or in glass
vials, respectively. In acidic lakes, we found an overestimation of thymidine incorporation in polycarbonate vials by
more than 50%. A preliminary conversion factor of 0.65 was
used for the correction of the former results. Thymidine
incorporation was stopped by the addition of 0.2 ml formaldehyde and 3 ml of ice-cold 10% TCA. The radioactivity of
the precipitate (2 ml were filtrated) retained on 0.2 µm PC
membrane filters was estimated in a Liquid Scintillation
Analyser (Packard, Tri-CARB 2100) using 1 ml Soluene 350
and 3 ml Hionic Flour as scintillation cocktails. Carbon
conversion factor (CCF = 18 fgC cell–1) was used according
to Bell (1986).
Samples for the estimation of primary production from
different depths of the water column or from the mixed layer
were taken and incubated in situ or in an incubator simulating
the natural temperature and underwater light conditions. In
the case of WOL, RIE and LAN, the mean underwater light
intensity of the mixed zone (Imix, Behrendt and Nixdorf,
1993) was chosen for laboratory incubation. For comparison
of primary production in natural and mining lakes, the results
of the incubation at the saturation light intensity (498 µE
m–2s–1) were used. The incorporation of 14C into the particulate fraction was measured by the standard procedure (Vollenweider, 1974). After filling samples into 25 ml glass
S283
bottles and inoculation with 0.05-0.125 ml NaH14CO3 (specific activity 0.185 MBq ml–1), the samples were incubated
for 3 h at around noon. After exposure, 10-20 ml were filtered
immediately through 0.2 µm membrane filters (Whatman).
Particulate 14C-incorporation into algae was measured by
LSC-counting (Packard Tri-CARB 2100). The surface integral of daily primary production was calculated from the
results of short-term incubation using the relationship between global radiation during the exposition time (or Imix, in
case of WOL, RIE and LAN) and the entire day.
4. Results
4.1. Chemistry of the lakes and production resources
In Table 2, the most important parameters for characterising the lake chemistry and resources for pelagic production
are compared for mining lakes and natural lakes. It is obvious
that the two types of lakes differ considerably in their chemistry, mainly the production resources. The concentration of
Mg, Fe, Al and Mn as well as sulphate is one to three order of
magnitudes higher in acidic mining lakes. This difference is
also reflected by the conductivity reaching more than 4000
µS cm–1 in mining lakes. With respect to the dissolved inorganic nitrogen supply, we measured an excess of ammonia
and nitrate in mining lakes. Whereas eutrophic hard water
lakes have sufficient nutrients for a very high primary production (especially the very shallow lake Langer See, see
also Mischke, 2003, ( this volume), the mining lakes are very
poor in phosphorus and in inorganic carbon. Though we have
to consider the two types of lakes as two quite different
habitats for primary producers.
The range of DOC concentration in the mining lakes is
comparable with those of meso- or eutrophic natural lakes in
Table 2
Range of important hydrochemical parameters in Lusatian Fe buffered
mining lakes (after Nixdorf et al., 2001) and in natural lakes (Scharmützelsee region)
Number of lakes
Conductivity (µs cm–1)
pH
KS4.3(mmol l–1)
KB8.2(mmol l–1)
TIC (mg l–1)
Mg (mg l–1)
Ca (mg l–1)
SO4(mg l–1)
NH4-N (mg l–1)
NO3-N (mg l–1)
DIP (µg l–1)
TP (µg l–1)
Fe (mg l–1)
Al (mg l–1)
Mn (mg l–1)
DOC (mg l–1)
Fe buffered mining
lakes
15
520–4340
2.2–3.4
–0.7 to –38.0
1.1–46.2
<0.2–18.3
5.9–84
68–550
454–5890
0.12–14.2
0.19–10.8
1–30
4–53
8.0–513
0.4–60.5
0.3–10.1
0.4–10.1
Natural hard water
lakes
13
277–697
6.4–9.7
1.6–2.6
–
19.5–28.6
<0.1–12.3
26.9–104.4
17.2–114.1
<0.02–3.8
<0.01–1.4
<1.2–545
9–317
<0.01–1.4
<0.01–0.35
<0.01–0.7
2.8–18.5
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B. Nixdorf et al. / Acta Oecologica 24 (2003) S281–S288
the region. Whether and to which amount or portion of this
organic carbon is available for bacteria is unknown.
4.2. Bacterial cell numbers
Total bacterial cell numbers in natural lakes were high
during the investigation period 2000/2001 (Fig. 2). A range
between 2.7 and 14 × 106 cells ml–1 was detected with
maximum values of total cell numbers in late spring and early
summer in Lake Langer See, the system with the highest
trophic state. The variation of total bacterial cell numbers in
mining lakes is higher and up to now, no trend in colonisation
or in seasonal development is obvious. Taking the initial
results of bacterial cell counting into account (Köcher and
Nixdorf, 1993; Nixdorf and Arndt, 1993), in extremely acidic
mining lakes, we found very low abundances of bacteria,
whereas in lake Waldsee, a meromictic mining lake with
higher trophy, cell numbers are comparable with natural
eutrophic hard water lakes. Generally, the bacterial cell numbers in acidic mining lake are only about 10% compared with
natural hard water lakes. The range of observed bacterial cell
numbers is from 0.2 to 2 × 106 cells ml–1, dominated by
bacteria smaller than 1 µm. Contrary to natural hard water
lakes, the portion of large rods and bacterial filaments is
higher with great morphological diversity.
4.3. Bacterial production
For the calculation of bacterial production, two estimation
procedures were used: first, the direct conversion of the
values obtained from the measurement into the amount of
carbon produced per hour and volume (mg C m–3h–1) and
second, the conversion of these values into the daily production (mg C m-3 d–1) by a factor of 24 assuming a constant
production intensity during the day and night. The bacterial
production shows a distinct seasonal course in the eutrophic
hard water lakes with maximum values in summer and autumn (see Fig. 3). The highest production is reached in the
hypertrophic lake Langer See (94.6 mg C m–3 h–1). The
average of the volume related bacterial production in these
lakes varies from 23.1 (LAN), to 7.7 (WOL) and 3.5 mg C
m–3 h–1 (RIE), whereas the average of the bacterial production measured in all mining lakes is 7.8 mg C m–3 h–1
(Table 3). The daily bacterial production is on average between 76 and 503 mg C m–3 d–1 in natural lakes and 151 mg
C m–3 d–1 in mining lakes.
4.4. Phytoplankton primary production
Fig. 2. Bacterial cell numbers in natural lakes of the Scharmützelsee region
from spring 2000 to 2001.
For comparison of bacterial and algal production, all primary production values are converted into the mass of as-
Fig. 3. Annual course of bacterial production in Brandenburg natural lakes from spring 2000 to 2001 (maximum value in Lake Langer See is 126 mg C m–3h–1)
and results of sporadical measurements of bacterial production in different mining lakes. The values are arranged according to the intensity of bacterial
production in mining lakes.
B. Nixdorf et al. / Acta Oecologica 24 (2003) S281–S288
S285
Table 3
Comparison of bacterial production (BP) and primary production (PP) in natural hard water lakes (LAN, WOL and RIE for abbrev. see study sites and Table 1)
and mining lakes (ML)
Lake
LAN
WOL
RIE
ML
PP per hour
(mg C m–3 h–1)
228.23
45.85
27.38
6.17
PP per day
(mg C m–3 d–1)
1141.15
229.25
136.90
30.85
BP per hour
(mg C m–3 h–1)
23.12
7.69
3.52
7.75
BP per day
(mg C m–3 d–1)
554.88
184.56
84.48
186.00
PP/BP per hour
BP/B per day
9.87
7.78
5.96
0.80
2.06
1.62
1.24
0.17
Fig. 4. Annual course of primary production in Brandenburg natural lakes from spring 2000 to 2001 and results of sporadical measurements in different mining
lakes from 2000 to 2001 measured at saturating light intensities (498 µE m–2s–1, grew area) or at the average light intensity in the mixed column (Imix, hatched
area) in natural lakes in 2000 and 2001.
similated carbon related to volume and hour or day, which
were obtained during incubation at saturating light intensities. Data which are shown in Fig. 4 show a similar seasonal
course of primary production in natural hard water lakes
comparable with bacterial production (Fig. 3), but at a higher
intensity of production. In lake Langer See, the average
primary production is highest (228 mg C m–3 h–1 or 1141 mg
C m–3 d–1, respectively). Lake Scharmützelsee and Wolziger
See produce only 10-20% of this amount (Table 3), whereas
the mining lakes on average reach only 3% of the production
in the highly eutrophic lake Langer See.
5. Discussion
5.1. General aspects of microbial productivity
In general, the obtained values of the natural hard water
lakes agree well with the data measured in other eutrophic
systems in this region (Nixdorf and Arndt, 1993) and collected for trophic classification approaches in the OECD
study (OECD, 1982). Areal production was not calculated for
this paper because of the number of indefinite conversion
procedures taking the whole water column into account.
Initial results related to these aspects of areal productivity in
the investigated lakes in the Scharmützelsee region are published in Krumbeck (2001) and Krumbeck et al. (1998).
Comparing both lake systems, the following trend for
algal and bacterial production is obvious: the trophic gradient
within the natural hard water lakes is reflected by the primary
and also the bacterial production. The very shallow lake
Langer See has the highest primary and bacterial production
reaching an annual primary productivity of about 1000 g C
m–2 a–1. These types of lowland lakes are described as the
most productive systems which are able to convert the same
mass of phosphorus into double the amount of phytoplankton
biomass compared with the less shallow or the dimictic lakes
in the region (Nixdorf and Deneke, 1997; Willén, 2000). The
very favourable underwater light climate in a water column
of about 2 m is the main reason for the very intense productivity of the very shallow lakes. Underwater light in dimictic
lakes is “diluted” in the sense of Reynolds (1997) in an
epilimnetic layer of 4–7 m or in the whole mixed layer during
circulation periods. This explains the lower production of the
lake Wolziger See although it has only lower TP concentration. Another reason for the lower productivity in the dimictic
lakes Wolziger See and Scharmützelsee is the threshold hy-
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pothesis (Chorus, 1995), which explains the biomass response of phytoplankton on nutrient reduction exceeding a
distinct level of about 50-60 mg P m–3. This threshold seems
to be dependent on the lake morphometry and mixing and
will be reached in WOL and RIE earlier and will be more
obvious in production parameters than in phytoplankton biomass.
5.2. Comparison of bacterial and algal productivity in
mining and natural lakes
For the assessment of bacterial and algal production, both
the processes are compared in Table 3. Here, two different
values are obtained considering hourly or daily volume related production. The differences are explained by the conversion of hourly values into daily productions: whereas
bacterial production is multiplied by 24, daily primary production is calculated by a light factor for the real light hours
and intensity received during the light day. Phytoplankton
production per hour during the light day is about 6- to 10-fold
higher than bacterial production in natural eutrophic lakes.
Considering the daily volume related production, this relation is diminished to 2.1 for lake Langer See and about
1.2–1.6 for the other hard water lakes (WOL and RIE). These
results confirm the classification approach for these lakes
characterising them as hypertrophic and eutrophic systems,
respectively. Autotrophic processes dominate the metabolism in the hypertrophic ecosystems and a compensation of
bacterial heterotrophic and phytoplankton autotrophic production was found in lakes with reduced nutrient loads and
trophy, like WOL and RIE. The amount of the primary
production that covers the bacterial production in eutrophic
lakes in the Scharmützelsee region was not measured. It is
known from eutrophic and hypertrophic lakes that exudation
as a main source of organic carbon supply for bacteria is
relatively low compared with the production of particulate
organic matter. For Lake Müggelsee, a shallow polymictic
and eutrophic lake in the Berlin/Brandenburg region, phytoplankton exudation was estimated covering 11–88% of the
bacterial production (Nixdorf and Arndt, 1993). Assuming a
similar low exudation rate in the investigated lakes of the
Scharmützelsee region, and in fact of the higher bacterial
production, other DOC sources for bacteria must be assumed
like autolysis of cyanobacteria, decomposition of detritus,
sloppy feeding and release of DOC due to grazing processes
as well as external inputs of allochthonous DOC.
A somewhat different relation between the microbial production components is obtained from the results of mining
lakes. Here, volume related primary production per hour is
only 80% of the bacterial production and only 17% of the
daily production. Although there are a number of uncertain
methodological problems; the trend of the relation between
primary and bacterial production characterises the mining
lakes as heterotrophic ecosystems. This trend is confirmed by
the oxygen budget of mining lakes, which shows in most
investigations under saturation between 85% and 95%. Mining lakes were described in the ecosystem characterisation
concerning energy and resources as main forcing elements
after Reynolds (1997) as “void” ecosystems (Nixdorf et al.,
2001). These lakes are poor in nutrients as the main production resources for autotrophic organisms and poor in energy
supply. This will be true for chemical energy as the main
resource for chemolithotrophic organisms and also for underwater light supply. It was shown that the most important
production resources like inorganic carbon und dissolved
inorganic phosphorus tends to accumulate near the bottom
due to groundwater inflow and mineralisation processes
(Nixdorf and Kapfer, 1998).
5.3. Control and utilisation of resources for primary
production
In Table 4, a comparison of the influence of single production resources on the phytoplankton production response is
given. It describes the control and utilisation of the main
primary production resources like under water light, inorganic carbon and nitrogen and total phosphorus in natural
and mining lakes. Three ecological aspects are considered:
fluctuations, supply and availability as a complex characterising the limitation conditions and the calculation of the
amount of resources to fill the carrying capacity. Underwater
light climate in natural eutrophic lakes is mainly influenced
by vegetation processes within the phytoplankton and
favours the development of cyanobacteria by self-shading.
The relatively clear mining lakes depress algal development
in surface layers due to over- or supersaturation of light
supporting deep Chl. a-maxima (DCM). Inorganic carbon
will never limit the primary production in natural hard water
lakes by the amount of TIC, but by the pH dependent availability of different inorganic carbon species. Contrary to
these lakes, TIC-limitation in acidic mining lakes is assumed
since primary production increases strongly after carbon
addition experiments (Beulker et al., 2003 (this volume);
Goldman et al., 1974; Lessmann and Nixdorf, 2000). Strategies avoided are mixotrophy (Beulker et al., 2003 (this volume) improvement of migration abilities by flagella and low
grazing and sedimentation losses. In mining lakes, naked
chrysophytes and chlamydophytes dominate with mixotrophic and migration abilities. The irregular increase of
TIC in deeper water layers supports the development of
DCM. Phosphorus (as TP) will limit the primary production
in both lake systems, whereas dissolved inorganic nitrogen
can be an important control for phytoplankton succession in
natural lakes (Mischke, 2003 (this volume) but will never
reach the carrying capacity in mining lakes.
5.4. Methodological aspects and open questions
Limnological experiments in extremely acidic waters, especially process measurements, require a number of tests
over a wide range of different types of acidic waters before a
method can be established (Herzsprung et al., 1998; Woelfl
and Whitton, 2000). Therefore, a critical view of our results
seems to be necessary. Production measurements in acidic
B. Nixdorf et al. / Acta Oecologica 24 (2003) S281–S288
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Table 4
Control and utilisation of resources for primary production in natural and in mining lakes and phytoplankton or population response
Resource
Underwater light (PFD)
Natural lakes
Mining lakes
Fluctuations and Seasonal
Turbidity by
influence on
turbidity by
chemical
temporal
plankton mass precipitation
changes
developments more than by
vegetation
Carbon (TIC)
Natural lakes
Constant,
carbonate
buffer, TIC
species
depending on
pH
Mining lakes
Fluctuating, at
pH > 5.6
expanding, not
known in
detaila
Supply and
availability,
consequences
for productivity
Limitation
assumed below
1 m, favouring
Cyanobacteria
Limitation by
availability of
single TIC
species only
Limitation by
amount,
bottom up
control
Filling the
available
carrying
capacity?
In the upper
part of the
column
? (Not by
amount)
In mixed
column
Optimum in
the water
column, DCM,
limitation at
the sediment
surface
In hypolimnion
or at the
sediment only
Phosphorus (TP)
Natural lakes
Mining lakes
Fluctuations by Fluctuating,
external and
precipitation
internal
by Fe and Al,
(sediments)
expanding by
inputs and
matter input
primary
and
production
eutrophication
Limitation in
Limitation
the production assumed
layer
Nitrogen (DIN)
Natural lakes
Fluctuations by
external and
internal
(sediments)
inputs and
primary
production
Limitation in
the production
layer, favouring
N-fixing
Cyanobacteria
In mixed
column
In mixed
column
In mixed
column
Mining lakes
Fluctuations due
to high
NH4concentration
of groundwater,
nitrification
limited
More than
needed
Never reached
a
Moderately acidic, pH 5–6, this is the transition range of the carbonate buffer system, where the equilibrium with atmospheric CO2 starts to determine the
buffer capacity. Below a pH value of ~5.6, the sum of inorganic carbon species approaches 10–5 mol l–1, and the system will be pH-stabilised again, when the
aluminium buffer system will be reached (see Nixdorf et al., 2001).
mining lakes are rare (Gyure et al., 1987; Kapfer, 1998;
Kapfer et al., 1997, 1999; Kwiatkowski and Roff, 1976;
Schindler, 1994; Schindler and Holmgren, 1971) and our
values have to be considered as the initial results. We measured, in some cases, very high rates of TIC dark fixation and
exudation in acidic mining lakes (results not published yet).
Whether these high dark fixation rates indicate a high activity
of chemolithotrophic microorganisms like Fe-, S- and ammonia oxidisers or even of autotrophic phytoplankton is not
known up to now. The initial results of high exudation rates
do not explain the high bacterial production in mining lakes.
The amount of released DOC during photosynthesis and
biomass synthesis does not cover the amount of bacterial
carbon requirements for production.
Further uncertain aspects involve the application of conversion factors calculating the bacterial production on a base
of thymidine incorporation. These conversion factors are
obtained from cultivated bacteria or freshwater bacteria from
non-acidic systems. They have to be checked for bacteria in
acidic waters and for a possible influence of the nano- and
picoalgal components for thymidine incorporation.
6. Conclusion
1. Primary production in eutrophic to hypertrophic natural
lakes reflects the nutrient state and biomass development more sensitive compared with the classical
trophic parameters.
2. The shallow lake Langer See has the highest phytoplankton and bacterial productivity (algal production
about 1000 g C m–2a–1). The relation of volume related
primaryand bacterial production is relatively constant
in eutrophic lakes, indicating the dominance of autotrophic processes.
3. Mining lakes are identified as heterotrophic systems
considering the relation between algal and bacterial
production.
4. The main resources for primary production interact
quite differently in both lake systems: good underwater
light supply and carbon limitation favour the development of DCM of pioneer phytoplankton in mining
lakes, whereas self-shading by cyanobacteria in TP and
DIN limited natural systems supports the planktonic
succession to the association S (sensu Reynolds
(1997)).
Acknowledgements
We would like to thank Jörg Koebcke, Remo Ender, Mike
Hemm, Simone Petersohn and Erwin Banscher for supporting sampling in mining lakes and chemical analyses. Bacterial and primary production measures were partly carried out
by Gudrun Lippert, Angelika Striemann and Ute Abel. We
also want to thank Ingo Henschke and Wolfgang Terlinden
for sampling in the Scharmützelseeregion. This study was
financially supported by the DFG (SFB 565), BMBF
(0339746) and LMBV (45016514).
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