Dissolved humic substances – ecological driving forces from the individual to the ecosystem level?

1. This review focuses on direct and indirect interactions between dissolved humic substances (HS) and freshwater organisms and presents novel opinions and hypotheses on their ecological significance. Despite their abundance in freshwaters, the role of HS is still inadequately understood. These substances have been considered too large to be taken up by freshwater organisms. On the contrary, here we present evidence that dissolved HS are indeed taken up and interact directly and/or indirectly with freshwater organisms. 2. We show that dissolved HS exert a mild chemical stress upon aquatic organisms in many ways; they induce molecular chaperones (stress shock proteins), induce and modulate biotransformation enzymes and modulate (mainly inhibiting) the photosynthetic release of oxygen by freshwater plants. Furthermore, they produce an oxidative stress, which may lead to membrane oxidation. HS modulate the multixenobiotic resistance activity and probably other membrane‐bound pumps. This property may lead to the increased bioaccumulation of xenobiotic chemicals. Furthermore, they can modulate the numbers of offspring in a nematode and feminise fish and amphibians. The ecological consequences of this potential remain obscure at present. HS also have the potential to act as chemical attractants (as shown with a nematode). 3. In some macrophytes and algae we show that HS interfere with photosynthesis and growth. For instance, the presence of HS suppresses cyanobacteria more than eukaryotic algae. By applying a quantitative structure activity relationship approach, we show that quinones in the HS interfere with photosynthetic electron transport. We show that even Phragmites leachate can act as a kind of phytotoxin. HS also have the potential to suppress fungal growth, as shown with the water mould Saprolegnia parasitica and force the fungus to respond by spore production. 4. In very soft, humic freshwaters, such as the Rio Negro, Brazil, HS stimulate the uptake of essential ions, such as Na and Ca, at extremely low pH (3.5–4.0) and prevent the ionoregulatory disturbance induced by acid waters, thereby enabling fish to survive in these environments. 5. We discuss whether or not HS are directly utilised by aquatic microorganisms or via exoenzymes, which may be washed in from the terrestrial catchment. There is accumulating evidence that the quality of the HS controls microbial growth. In total, net‐heterotrophy may result from HS‐mediated suppression of primary production by the quinone structures and/or from HS‐mediated support of microbial growth. As there is also evidence that HS have the potential to support photoautotrophic growth and suppress microbial growth, the opposite community effect could result. Consequently, dissolved organic carbon (DOC) has to be chemically characterised, rather than simply measuring bulk DOC concentration. 6. In sum, dissolved HS interact with freshwater organisms in a variety of ways in unenriched humic lakes. In addition to the well known effects of HS on light regime, for example, and the direct and indirect supply with carbon (energy), other interactions may be much more subtle. For instance, HS may induce internal biochemical stress defence systems and have the potential to cause acclimatisation and even adaptation. We are just at the beginning of understanding these interactions between dissolved HS and freshwater organisms.     

Planktonic bloom-forming Cyanobacteria and the eutrophication of lakes and rivers

SUMMARY. 1. Eutrophication of water bodies involves the enrichment of plant nutrients, often followed by significant shifts in the phytoplankton towards Cyanobacteria. When comparing different aquatic systems, even with similar nutrient contents and in the same climatic region, inverse deductions are not valid; i.e. (a) the presence of Cyanobacteria does not necessarily indicate eutrophic conditions, or (b) eutrophic or even poly-trophic conditions do not necessarily support cyanobacterial development.
2. Above a threshold of 10 μg 1⁻¹ total phosphorus, the development of Cyanobacteria can be described by physical factors, such as water column stability. By characterizing different forms of turbulence, the presence or absence of Cyanobacteria in lakes and rivers can be predicted.
3.When the turbulence of the water column is rather low, as it is in sheltered or meromictic lakes, Cyanobacteria can build up dense populations. In nutrient poor systems, species of Oscillatoria and (seldom) Aphanizomenon are dominating.
4.If the turbulence of the water column is high (mixing depth much greater than euphotic depth) or the mixing pattern is irregular, as in slowly flowing or regulated rivers, Cyanobacteria are outcompeted.
5. In the presence of frequent or permanent turbulence, but with mixing depths lower or not much greater than the euphotic zone (as it is the case in shallow, unstratified lakes, mostly eutrophic or even hypertrophic), Cyanobacteria can outgrow normally dominant r -strategists under conditions of low N:P ratios, high water temperatures, pH >9.0 or low light availabilities.
6. If turbulence is comparatively great (10 to $15 m) and stable for a longer period of time, some cyanobacteria are able to adapt.
7. Our statements are discussed on the basis of physiological characteristics.