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Water transparency affects the thermal structure of lakes and within certain lake depth ranges
it can determine whether a lake mixes regularly (polymictic regime) or stratifies continuously (dimictic regime) from spring through summer
Phytoplankton biomass can influence transparency but the effect of its seasonal pattern on stratification is unknown
Therefore we analysed long term field data from two lakes of similar depth
transparency and climate but one polymictic and one dimictic and simulated a conceptual lake with a hydrodynamic model
Transparency in the study lakes was typically low during spring and summer blooms and high in between during the clear water phase (CWP)
caused when zooplankton graze the spring bloom
The effect of variability of transparency on thermal structure was stronger at intermediate transparency and stronger during a critical window in spring when the rate of lake warming is highest
Whereas the spring bloom strengthened stratification in spring
The presence or absence of the CWP influenced stratification duration and under some conditions determined the mixing regime
including biotic interactions that suppress the CWP
stratification duration and potentially also the mixing regime
They unequivocally conclude that a reduction in transparency decreases deep water temperatures
the thickness of the surface (mixed) layer and the overall heat content of the water body
The increased temperature difference between surface and deep water stabilises thermal stratification
the vast majority of these studies deals with deep
seasonally stratified lakes and relatively little is known about how transparency affects marginal lakes
the effect of the distinct seasonal pattern of transparency resulting from phytoplankton biomass on thermal structure and stratification duration
Our hydrodynamic model simulations were performed with an idealized seasonal phytoplankton pattern in a conceptual lake
where the only lake-specific parameters were light extinction coefficient (γ)
thus focusing on general phenomena rather than lake-specific detail
We hypothesize that seasonal phytoplankton dynamics and the CWP affect the stratification duration
thermal structure and potentially the mixing regime of temperate marginal lakes
Dependence of extinction (γ) on chlorophyll a and dissolved organic carbon (DOC) concentration in Müggelsee (a,b) and Heiligensee (c,d)
γ was calculated from Secchi transparency (Zsecchi) with equation 3 using c = 2.05 for Müggelsee and c = 2.13 for Heiligensee (see Methods)
chla is the chlorophyll concentration (μg L−1) and DOC is in mg L−1
light absorption through phytoplankton was the main driver of transparency in the study lakes
Mean seasonal pattern of chlorophyll a (Chla
(c,d)) and stratification strength (Ts − Tb
(e,f)) in Müggelsee (a,c,e) and Heiligensee (b,d,f)
lines with symbols show how the first (b,c,d,f) or second principal component (e) affect the mean seasonal pattern
Open symbols show a low score of the respective principal component (25th percentile in the time series) and closed symbols show a high score (75th percentile)
Seasonal patterns represented by like symbols in each lake are correlated
The scores for Chla in Müggelsee are not shown because they were not correlated with Zsecchi or Ts − Tb; instead the shaded region in (a) represents 1 standard deviation from the mean
Note reversed y-scale for Zsecchi reflecting depth measured downwards
Monthly loadings of the first two components (PC1
PC2) from the principal component analysis of inverse-transformed chlorophyll a (Chla−1
Both lakes were on average isothermal and unstratified from October to March as reflected in near zero values of the mean difference between surface and bottom temperatures (Ts − Tb, neglecting inverse stratification in winter and noting that negative values were truncated; Fig. 2e,f)
peaking in July at 1.6 °C and 8.1 °C in Müggelsee and Heiligensee
PC2 on the other hand had the highest absolute loading in April in both lakes and mainly represented variation in spring stratification relative to summer
We assessed synchrony between the two lakes by comparing the deseasonalised (centred) monthly means of Ts − Tb and Secchi transparency in the two lakes during the period of parallel measurements (1979–2000)
considering only the potentially stratified months of April to August
df = 93) were significantly correlated in the two lakes
Monthly chlorophyll concentrations did correlate between the lakes however (p = 0.008
at least during the years when parallel measurements existed (1991–2000)
We further checked the principal components for synchrony
PC1 for Secchi transparency was not correlated between the lakes (p = 0.10)
nor was PC2 for Ts − Tb in Müggelsee correlated with PC1 for Ts − Tb in Heiligensee (p = 0.55)
nor were the PCs for chlorophyll correlated (p = 0.72)
Thus transparency and stratification were correlated within each lake
but in a way that was not synchronous between the lakes
a stronger seasonal amplitude of transparency and a more intense CWP significantly weakened summer stratification and skewed temperature gradients away from summer towards spring in both lakes
This response was independent of whether the lake was polymictic or dimictic
Seasonal extinction scenarios used in model simulations
The base seasonal extinction pattern (black line in (a,b)) was defined based on long term measured extinction in Müggelsee (grey lines in (a)
each line representing one year) and Heiligensee (grey lines in (b))
Extinction scenarios were then derived from the base extinction pattern by varying annual mean extinction with a clear water phase (CWP
(c)) and without a CWP (d) or by varying the timing of the spring bloom (e)
Seasonally variable extinction scenarios were compared with control scenarios of constant extinction (f)
Model validation against measured temperatures and stratification in Müggelsee (a,c) and Heiligensee (b,d)
open orange symbols show surface temperatures
closed blue symbols show surface-bottom temperature differences (a) or bottom temperatures (b)
Lines show the corresponding simulated temperatures
vertical bars show the observed (orange) and modelled (blue) total number of stratified days per year
Error bars in (d) were estimated from the sampling interval
We used the period 2004–2009 in (c) for Müggelsee because high frequency temperature profiles were available only after 2004
The model was parameterised for a conceptual lake similar to both Müggelsee and Heiligensee
It was run with the same meteorological forcing for both lakes with the exception of a 50% increase in wind speed for Müggelsee
Simulated effect of annual mean extinction,
on mean lake thermal properties under constant extinction
or variable extinction scenarios with and without clear water phase (CWP)
Grey dashed lines denote a transitional region between polymixis and dimixis at Dur = 120 days
The lake becomes dimictic at higher extinction when the CWP is present than when it is absent
Seasonally variable extinction produced striking effects on thermal structure in the transitional range of 0.5–1.5 m−1. Variable extinction with a CWP weakened thermal gradients and stratification compared to constant extinction within the range = 0.6–1.0 m−1, as evident from higher Tb and shorter stratification (Fig. 6)
temperature gradients and stratification duration were considerably higher at the same across the whole transitional range (0.5–1.5 m−1)
at = 0.7 m−1 mean Tb was 1 °C lower and stratification duration was on average 49 days longer when the CWP was absent than when it was present
the presence or absence of the CWP determined whether the lake was dimictic or polymictic
Seasonal development of mean simulated lake thermal characteristics
comparing constant extinction (dashed lines) and seasonally variable extinction scenarios with a clear water phase (CWP
= 0.7 m−1 (a,b,c) or 1.0 m−1 (d,e,f)) and without a CWP (dot-dashed lines
Scenarios with variable spring bloom timing showed that a delay in spring bloom timing from day 90 to day 120 (approximate range observed in Lake Müggelsee)
increased thermal stability of the water column slightly as evident in higher Ts − Tb beginning near the spring bloom and persisting as long as Ts − Tb was positive
Bloom timing had little effect on Ts or overall stratification duration
Effect of a short term deviation in extinction at different times of year on the annual mean difference between surface (Ts) and bottom temperature (Tb) (c,d)
The deviation was created by doubling (a) or halving (b) a constant arbitrary baseline extinction (1.0 m−1) for 20 days
Simulations were performed with the deviation midpoint shifted to different days of the year
Sensitivity of mixing regime to mean extinction ()
mean lake depth (Zmean) and the clear water phase (CWP) for a lake with 1000 m fetch similar to Heiligensee (a) and for a lake with 4000 m fetch and increased wind speed by 50%
The shaded area represents the region that is on average only polymictic when the CWP is present
delineating the transitional conditions under which lakes shifted between polymictic and dimictic regimes in at least 10% of simulated years
The circles marked “M” and “H” show the average observed conditions in Müggelsee and Heiligensee
We demonstrated that stratification duration and the mixing regime of marginal lakes may respond strongly to seasonal changes in phytoplankton biomass
Our study showed that cardinal planktonic events in spring
potentially have a large influence because they fall within a critical window during which transparency has a much stronger effect than at other times of the year
The model simulations indicated the existence of certain depth and extinction combinations where the presence or absence of the CWP altered the average mixing regime
The empirical results demonstrated that a stronger seasonal pattern associated with a more intense CWP decreased summer stratification relative to spring
We therefore confirm our hypothesis that seasonal phytoplankton dynamics and the CWP significantly affect the stratification duration
thermal structure and mixing regime of temperate marginal lakes
which does not seem to be the case in deep dimictic temperate lakes
Our results are in line with these conclusions and suggest that the effect of transparency is a lot stronger in marginal lakes because its seasonal variability has a greater effect in the range between stable polymixis and stable dimixis
Thus we conclude that not only the annual mean transparency
but also the seasonal variation in transparency and the CWP play an important role
This is not surprising for marginal lakes because by definition
the mean lake depth is similar to the thermocline depth
so there is most likely an interaction between stratification and bloom formation
This interaction makes it difficult to infer causality from field data
to help interpret whether phytoplankton-mediated changes in transparency also alter stratification
The model simulations of the conceptual lake indicated that a more intense CWP compared to constant extinction weakens summer stratification relative to spring under both weak stratification (summer Ts − Tb ~ 0–5 °C) and stronger stratification (summer Ts − Tb ~ 5–10 °C) as observed in Müggelsee and Heiligensee
we did not detect any significant synchrony in the correlated principal components between the two lakes
suggesting that internal dynamics might have a stronger influence than regional climate
in addition to the well-known effects of stratification on bloom formation
seasonal variability of transparency through phytoplankton also significantly influences stratification in marginal lakes regardless of whether predominantly polymictic or seasonally stratified
which explains why the CWP and spring bloom have such a strong effect on thermal structure
This was clearly evident in the statistical analysis of Müggelsee and Heiligensee
where a smaller spring bloom and clearer CWP weakened summer stratification relative to spring
The simulations with the conceptual lake reproduced this stratification response to seasonal transparency
delivering an explanation for this behaviour
Whereas the spring bloom initially strengthens stratification
the CWP reduces vertical temperature differences by allowing radiation to penetrate to and heat deeper water layers and thus potentially alter the mixing regime
The importance of the CWP and the weather conditions during the spring critical window also explains the anomaly in the model validation against the stratification duration observed in Müggelsee in 2008 (Fig. 5c)
2008 had the highest mean global radiation
the third highest mean air temperature and the lowest mean wind speed in the 30 year data period
using a standard extinction scenario in the validation
the model predicted strong stratification in Müggelsee beginning in late April
which was stable enough to persist into summer
Whereas this led to a long period of stratification in the simulation
the observed stratification duration in Müggelsee was considerably shorter
strong stratification also developed in late April but had disappeared again by mid June
The reason is likely that the CWP in May 2008 (mean Secchi depth 4.1 m) was one of the most intense in the 35 year record
being only exceeded in May 1998 (mean Secchi depth 4.5 m)
Therefore biotic interactions may accelerate the mixing regime shifts in temperate marginal lakes expected due to climate change
DOC was measured by non-dispersive infrared sensing after combustion with an N/C analyser (Shimadzu in Heiligensee
or a Jena Analytics multi N/C 2100 in Müggelsee)
γ was reconstructed for 1975–2001 from the longer Secchi depth time series
FLake indirectly assumes the lake to have a constant cross-sectional area and flat bottom (maximum depth is equal to the mean depth)
the bottom temperature in FLake is the temperature in a hypothetical lake of regular form with the same area and volume as the real lake
The result of this simplification is the possibility to apply the self-similarity approach to the temperature profile and an enormous increase of computational speed (FLake is at least 102 times faster than any existing one-dimensional lake model)
The side effect is a divergence of the simulated near-bottom temperatures from the observed ones
The inconsistencies in the bottom temperature are however comparable to those found in other lake models and can be reduced by fitting the model to a certain lake
In this study we intentionally avoid any fitting
using the model as a process-based mechanistic representation of heat transport in an ‘ideal lake’ capturing the general dynamics and mean behaviour observed in two real lakes
Model simulations with FLake were forced using meteorological data (3-hourly resolution) from 1980–2010 from a weather station in Potsdam
Some missing cloud cover data and other small data gaps were filled with data from nearby stations
The model forcing variables include solar radiation
humidity and long wave radiation estimated from cloud cover
where t is the independent variable (time, day of the year) and a, b, c, d and e are free parameters. The extinction during the summer phytoplankton maximum was described by a Gauss (bell) function, g(t) (equation 5):
Equations (4) were parameterized with the values in Table 1 in addition to σ = 35
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We thank Patrick Neale for assistance with statistical analyses
Ursula Newen and Helgard Täuscher for sampling and sample processing and the Leibniz-Institute of Freshwater Ecology and Inland Fisheries (IGB) for the long-term ecological program
Meteorological data were kindly provided by the German Weather Service
Schmidt and Torsten Seltmann provided valuable input
This research was funded by a German Science Foundation (DFG) grant (No
was additionally supported by the DFG grant IceBound (KI-853/11-1)
was supported by the DFG grant LakeRisk (AD 91/13-1) and the EU-project LIMNOTIP funded under the FP7 ERA-Net Scheme (Biodiversa
Leibniz-Institute of Freshwater Ecology and Inland Fisheries Department of Ecohydrology
Leibniz-Institute of Freshwater Ecology and Inland Fisheries Department of Ecosystem Research
Freie Universität Berlin Department of Biology
performed the modelling and statistical analyses and wrote the manuscript
All authors contributed to interpreting the results and editing the manuscript
The authors declare no competing financial interests
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