Ecological stoichiometry – a bottleneck for biodiversity and ecosystem services Jaroslav Vrba Department of Ecosystem Biology Faculty of Science, University of South Bohemia, Č eské Bud ě jovice, Czech Republic Biology Centre ASCR, v.v.i. Institute of Hydrobiology Jaroslav Vrba: Ecological stoichiometry ALTER-net Summer School, Peyresq, 2008
Ecological stoichiometry – outlines ES studies a balance of energy and particular chemical elements in ecological interactions • Historical outlines • Framework of evolutionary biology • Biochemical and physiological constraints of life • Population and community dynamics • Ecosystem structure and functioning • Sustainable ecosystem services Jaroslav Vrba: Ecological stoichiometry ALTER-net Summer School, Peyresq, 2008
Ecological stoichiometry – outlines Stoichiometry: Law of definite proportion (or Law of constant composition) Liebig (1840) – Law of Minimum Lotka (1925) – stoichiometry in biology Redfield (1934, 1958) – atomic C:N:P ratio = 106:16:1 Plankton ecology (>1990) – ecological stoichiometry Conservation of mass and Conservation of energy in biology Jaroslav Vrba: Ecological stoichiometry ALTER-net Summer School, Peyresq, 2008
Ecological stoichiometry at WoS 160 Ecological stoichiometry 140 No. of papers on WoS 120 100 80 60 40 20 0 1993 1994 1995 1996 1997 1998 1999 2000 2001 2002 2003 2004 2005 2006 2007 2008 Web of Science ~ World of Stoichiometry … ☺ Jaroslav Vrba: Ecological stoichiometry ALTER-net Summer School, Peyresq, 2008
Ecological stoichiometry ? Synthesis of production ecology and population ecology… 6 CO 2 + 6 H 2 O + 2802 kJ C 6 H 12 O 6 + 6 O 2 4 – + HPO 4 2– + 122 H 2 O + 18 H + 106 CO 2 + 16 NO 3 + trace elements + energy Biotic interactions! Production C 106 H 263 O 110 N 16 P 1 + 138 O 2 Respiration = phytoplankton biomass H 375000000 O 1320000000 C 85700000 N 6430000 Ca 1500000 P 1020000 human body = S 206000 Na 183000 K 177000 Cl 127000 Mg 40000 Si 38600 Fe 2680 Zn 2110 Cu 76 I 14 Mn 13 F 13 Cr 7 Se 4 Mo 3 Co 1 Jaroslav Vrba: Ecological stoichiometry ALTER-net Summer School, Peyresq, 2008
Biogenic elements are non-homeostatic ! Element Resource (A > 10 -2 > B> 10 -6 > C > 10 -9 > D) Earth crust Oceans Vertebrates (proxy for) (terrestrial ecos.) (aquatic ecosyst.) (heterotr. consumer) hydrogen D << A A H carbon B B < A C nitrogen B > C << A N oxygen A A A O sodium A A > B Na magnesium A > B B Mg silica A > B B Si phosphorus B > C << A P sulphur B B B S kalium A > B B K calcium A B < A Ca manganese B >> D < C Mn iron A >> D << B Fe Jaroslav Vrba: Ecological stoichiometry ALTER-net Summer School, Peyresq, 2008
Stoichiometry of cells – cell chemistry Composition of biomolecules – biochemical stoichiometry Selection for C, N & P in biochemical evolution G rowth R ate H ypothesis ( GRH ) = ribosomes C+N =proteins C ( energy )=saccharides & lipids Jaroslav Vrba: Ecological stoichiometry ALTER-net Summer School, Peyresq, 2008
Growth Rate Hypothesis (GHR) N 2 fixers K-strategists (N:P>40) r-strategists = ideal phytoplankton Jaroslav Vrba: Ecological stoichiometry Arrigo (2005) Nature 437 ALTER-net Summer School, Peyresq, 2008
Growth Rate Hypothesis (GHR) r-strategists (Cladocera) = high need in P ! Jaroslav Vrba: Ecological stoichiometry ALTER-net Summer School, Peyresq, 2008
Homeostasis of heterotrophic consumers Growth (resource utilization) may change stoichiometry Are you what you eat? Autotrophs: rather YES Heterotrophs: mostly NOT Jaroslav Vrba: Ecological stoichiometry ALTER-net Summer School, Peyresq, 2008
Homeostasis of vertebrates Structural investment = skeleton changes fundamentally needs in resource stoichiometry great need in P & Ca ! Jaroslav Vrba: Ecological stoichiometry ALTER-net Summer School, Peyresq, 2008
Homeostasis of vertebrates Structural investment = skeleton changes fundamentally needs in resource stoichiometry even during ontogenesis great need in P & Ca ! Jaroslav Vrba: Ecological stoichiometry Pilati & Vanni (2007) Oikos 116 ALTER-net Summer School, Peyresq, 2008
Stoichiometry of populations & communities 1. Conservation of mass holds for each element 2. Nutrient availability controls population growth & dynamics 3. Nutrient use efficiencies determine (species) competitiveness 4. Resources’ imbalance controls (particular) nutrient regeneration 5. Resource stoichiometry determines biotic interactions 6. Stoichiometry determines structure of food webs 7. Stoichiometry does control biodiversity Jaroslav Vrba: Ecological stoichiometry ALTER-net Summer School, Peyresq, 2008
Stoichiometry of populations & communities In particular effects of P supply should impinge on fitness & drive evolutionary change Jaroslav Vrba: Ecological stoichiometry Jeyasingh & Weider (2007) Mol. Ecol. 16 ALTER-net Summer School, Peyresq, 2008
Food web stoichiometry Bottom-up: soil nutrient availability (i.e. rain) in a desert controls both producer ’s and consumer ’s stoichiometry Sabinia setosa (Curculionidea) Prosopis velutina Jaroslav Vrba: Ecological stoichiometry Schade et al. (2003) Ecol. Lett. 6 (Fabaceae) ALTER-net Summer School, Peyresq, 2008
Food web stoichiometry Top-down: cascading effect of predation and resource stoichiometry � Jaroslav Vrba: Ecological stoichiometry ALTER-net Summer School, Peyresq, 2008
Food web stoichiometry Top-down: cascading effect of predation and resource stoichiometry determine un/successful biomanipulation ☺☺ � � � � � � Jaroslav Vrba: Ecological stoichiometry ALTER-net Summer School, Peyresq, 2008
Food web stoichiometry Nutrient regeneration is species specific (26 vertebrates) Vanni et al. (2002) Ecol. Lett. 5 Jaroslav Vrba: Ecological stoichiometry ALTER-net Summer School, Peyresq, 2008
Food web stoichiometry Consumer and resource stoichiometry controls efficiency ( GGE C ) = carbon + energy dissipation ! Redfield Jaroslav Vrba: Ecological stoichiometry ALTER-net Summer School, Peyresq, 2008
Ecosystem stoichiometry Experimental eutrophication of wetlands (Belize: +P ) Vegetation Eleocharis Eleocharis Typha (control) +P +P plant biomass C:P 4865 712 1555 plant biomass N:P 74 12.1 15.1 Sediment microb. biom. C:P 97.2 17.7 69.2 Plants microb. biom. N:P 3.1 0.5 3.1 interstic. SRP (µg/l) 0.8 7.3 16.1 Soil P Soil Jaroslav Vrba: Ecological stoichiometry microbes ALTER-net Summer School, Peyresq, 2008
Ecosystem stoichiometry Experimental eutrophication of wetlands (Belize: +P ) = distinct stoichiometry of producers / detritus (litter) Rejmankova & Houdkova (2006) BGC 80, Šantr ůč ková et al. (unpubl.) Vegetation Eleocharis Eleocharis Typha (control) +P +P plant biomass C:P 4865 712 1555 plant biomass N:P 74 12.1 15.1 Sediment microb. biom. C:P 97.2 17.7 69.2 Plants microb. biom. N:P 3.1 0.5 3.1 interstic. SRP (µg/l) 0.8 7.3 16.1 Soil P Soil Jaroslav Vrba: Ecological stoichiometry microbes ALTER-net Summer School, Peyresq, 2008
Ecosystem stoichiometry Experimental eutrophication of wetlands (Belize: +P ) = distinct stoichiometry of producers / detritus (litter) + occurrence of mosquitoes (Anopheles spp.) causing a serious health hazard = malaria Grieco et al. (2005) J. Vector Ecol. 30 Grieco et al. (2006) J. Med. Entomol. 43 Grieco et al. (2007) J. Vector Ecol. 32 Jaroslav Vrba: Ecological stoichiometry ALTER-net Summer School, Peyresq, 2008
Ecosystem stoichiometry vs. productivity ? Distinct stoichiometry of terrestrial and aquatic producers Terrestrial ecosystems: high C:N:P = high (structural !) biomass Aquatic ecosystems: low C:N:P = low biomass, high production ! Jaroslav Vrba: Ecological stoichiometry ALTER-net Summer School, Peyresq, 2008
Ecosystem stoichiometry vs. productivity ? Distinct stoichiometry of terrestrial and aquatic producers Terrestrial ecosystems: high C:N:P = high (structural !) biomass Aquatic ecosystems: low C:N:P = low biomass, high production ! Jaroslav Vrba: Ecological stoichiometry ALTER-net Summer School, Peyresq, 2008
Ecosystem stoichiometry OECD Model (Vollenweider) : seston chlorophyl–TP relationship Jaroslav Vrba: Ecological stoichiometry ALTER-net Summer School, Peyresq, 2008
Ecosystem stoichiometry Anthropogenic impacts = deposition, fertilisers, eutrophication… � Human activity turns both landscape and the planet in “a large-scale excrement”… � Jaroslav Vrba: Ecological stoichiometry ALTER-net Summer School, Peyresq, 2008
Ecosystem stoichiometry Anthropogenic impacts = deposition, fertilisers, eutrophication… coastal ecosystems lakes Jaroslav Vrba: Ecological stoichiometry ALTER-net Summer School, Peyresq, 2008
Ecosystem stoichiometry Ecosystem services – e.g., of coastal ecosystems Si:N Jaroslav Vrba: Ecological stoichiometry ALTER-net Summer School, Peyresq, 2008
Ecosystem stoichiometry Ecosystem services – e.g., of coastal ecosystems Si:N Si:N Jaroslav Vrba: Ecological stoichiometry Ptacnik et al. (2005) Oikos 109 ALTER-net Summer School, Peyresq, 2008
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