title: Humic substances. Part 2: Interactions with organisms.
abstract: OBJECTIVE
  Freshwater bodies which chemistry is dominated by dissolved humic substances (HS) seem to be the major type on Earth, due to huge non-calcareous geological formations in the Northern Hemisphere and in the tropics. Based on the paradigm of the inertness of being organic, direct interactions of dissolved HS with freshwater organisms are mostly neglected. However, dissolved organic carbon, the majority of which being HS, are natural environmental chemicals and should therefore directly interact with organisms. Major results that widened our perspective on humic substance ecology come from experiments with the compost nematode, Caenorhabditis elegans, which behaved contradictorily to textbook knowledge and provoked an in-depth re-consideration of some paradigms.
  
  
  METHODS
  To overcome old paradigms on HS and their potential interactions with organisms, we reviewed recent international literature, as well as 'grey' literature. We also include results from own ongoing studies.
  
  
  RESULTS
  This review focuses on direct interactions of dissolved HS with freshwater organisms and disregards indirect effects, such as under-water light quenching. Instead we show with some macrophyte and algal species that HS adversely interfere with photosynthesis and growth, whereby closely related algal species show different response patterns. In addition to this, HS suppress cyanobacteria more than eukaryotic algae. Quinones in the HS appear to be the effective structure. Furthermore, HS can modulate the offspring numbers in the nematode C. elegans and cause feminization of fish and amphibians--they possess hormone-like properties. The ecological consequences of this potential remain obscure at present. HS also have the potential to act as chemical attractants as shown with C. elegans and exert a mild chemical stress upon aquatic organisms in many ways: induction of molecular chaperons (stress proteins), induction and modulation of biotransformation and anti-oxidant enzymes. Furthermore, they produce an oxidative stress with lipidperoxidation as one clear symptom or even stress defense strategy. Stronger chemical stresses by HS may even lead to teratogenic effects as shown with fish embryos; all physiological responses to HS-mediated stress require energy, which were compensated on the expense of yolk as shown with zebra fish embryos. One Finnish field survey supports the view of a strong chemical stress, as the weight yield in fish species decreases with increasing HS content in the lakes.
  
  
  CONCLUSIONS
  HS exert a variety of stress symptoms in aquatic and compost organisms. According to current paradigms of ecotoxicology, these symptoms have to be considered adverse, because their compensation consumes energy which is deducted from the main metabolism. However, the nematode C. elegans looks actively for such stressful environments, and this behavior is only understandable in the light of new paradigms of aging mechanisms, particularly the Green Theory of Aging. In this respect, we discuss the mild HS-mediated stress to aquatic and compost organisms. New empirical findings with HS themselves and HS building blocks appear to be consistent with this emerging paradigm and show that the individual lifespan may be expanded. At present the ecological consequences of these findings remain obscure. However, a multiple-stress resistance may be acquired which improves the individual fitness in a fluctuating environment.
  
  
  CONCLUSIONS
  It appears that dissolved HS have to be considered abiotic ecological driving forces, somewhat less obvious than temperature, nutrients, or light.
  
  
  CONCLUSIONS
  The understanding of the ecological control by dissolved humic substances is still fragmentary and needs to be studied in more details.
authors: Humboldt University at Berlin, Institute of Biology, Freshwater and Stress Ecology, Germany. christian_ew_steinberg@web.de
authors: Christian E W
authors: CE
authors: Steinberg
authors: Thomas
authors: T
authors: Meinelt
authors: Maxim A
authors: MA
authors: Timofeyev
authors: Michal
authors: M
authors: Bittner
authors: Ralph
authors: R
authors: Menzel
mesh_terms: D018435:ATP Binding Cassette Transporter, Subfamily B / Q000378:metabolism; D000818:Animals; D002462:Cell Membrane / Q000187:drug effects; D052244:Endocrine Disruptors / Q000737:chemistry / Q000633:toxicity; D056890:Eukaryota / Q000187:drug effects / Q000378:metabolism; D005618:Fresh Water; D005786:Gene Expression Regulation / Q000187:drug effects; D006360:Heat-Shock Proteins / Q000378:metabolism; D006812:Humic Substances; D010100:Oxygen / Q000378:metabolism; D010675:Pheromones / Q000737:chemistry; D010788:Photosynthesis / Q000187:drug effects; D010944:Plants / Q000187:drug effects / Q000378:metabolism; D012964:Sodium / Q000378:metabolism; D014873:Water Pollutants / Q000737:chemistry / Q000633:toxicity*
chemical_list: D018435:ATP Binding Cassette Transporter, Subfamily B; D052244:Endocrine Disruptors; D006360:Heat-Shock Proteins; D006812:Humic Substances; D010675:Pheromones; D014873:Water Pollutants; D012964:Sodium; D010100:Oxygen