40
Jaroslav Vrba Department of Ecosystem Biology Faculty of Science, University of South Bohemia, České Budějovice, Czech Republic Biology Centre ASCR, v.v.i. Institute of Hydrobiology Jaroslav Vrba: Ecological stoichiometry ALTER-net Summer School, Peyresq, 2008 Ecological stoichiometry – a bottleneck for biodiversity and ecosystem services

a bottleneck for biodiversity and ecosystem services · – a bottleneck for biodiversity and ecosystem services. ... • Biochemical and physiological constraints of life ... Our

  • Upload
    donhan

  • View
    221

  • Download
    0

Embed Size (px)

Citation preview

Page 1: a bottleneck for biodiversity and ecosystem services · – a bottleneck for biodiversity and ecosystem services. ... • Biochemical and physiological constraints of life ... Our

Jaroslav Vrba

Department of Ecosystem BiologyFaculty of Science, University of South Bohemia,

České Budějovice, Czech Republic

Biology Centre ASCR, v.v.i.Institute of Hydrobiology

Jaroslav Vrba: Ecological stoichiometryALTER-net Summer School, Peyresq, 2008

Ecological stoichiometry– a bottleneck for biodiversity and ecosystem services

Page 2: a bottleneck for biodiversity and ecosystem services · – a bottleneck for biodiversity and ecosystem services. ... • Biochemical and physiological constraints of life ... Our

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 servicesJaroslav Vrba: Ecological stoichiometryALTER-net Summer School, Peyresq, 2008

Ecological stoichiometry – outlines

Page 3: a bottleneck for biodiversity and ecosystem services · – a bottleneck for biodiversity and ecosystem services. ... • Biochemical and physiological constraints of life ... Our

Ecological stoichiometry – outlines

Stoichiometry: Law of definite proportion(or Law of constant composition)

Lotka (1925) –stoichiometry in biology

Liebig (1840) – Law of Minimum

Redfield (1934, 1958) –atomic C:N:P ratio = 106:16:1

Plankton ecology (>1990) –ecological stoichiometry

Conservation of mass and Conservation of energyin biology

Jaroslav Vrba: Ecological stoichiometryALTER-net Summer School, Peyresq, 2008

Page 4: a bottleneck for biodiversity and ecosystem services · – a bottleneck for biodiversity and ecosystem services. ... • Biochemical and physiological constraints of life ... Our
Page 5: a bottleneck for biodiversity and ecosystem services · – a bottleneck for biodiversity and ecosystem services. ... • Biochemical and physiological constraints of life ... Our

Ecological stoichiometry

0

20

40

60

80

100

120

140

160

1993

1994

1995

1996

1997

1998

1999

2000

2001

2002

2003

2004

2005

2006

2007

2008

No. o

f pap

ers

on W

oS

Ecological stoichiometry at WoS

Web of Science ~ World of Stoichiometry… ☺

Jaroslav Vrba: Ecological stoichiometryALTER-net Summer School, Peyresq, 2008

Page 6: a bottleneck for biodiversity and ecosystem services · – a bottleneck for biodiversity and ecosystem services. ... • Biochemical and physiological constraints of life ... Our

Biotic interactions!

ProductionRespiration

Ecological stoichiometry ?

Synthesis of production ecology and population ecology…

6 CO2 + 6 H2O + 2802 kJ C6H12O6 + 6 O2

= phytoplankton biomass

106 CO2 + 16 NO3– + HPO4

2– + 122 H2O + 18 H+

+ trace elements + energy

C106H263O110N16P1 + 138 O2

H375000000 O1320000000 C85700000 N6430000 Ca1500000 P1020000

S206000 Na183000 K177000 Cl127000 Mg40000 Si38600 Fe2680

Zn2110 Cu76 I14 Mn13 F13 Cr7 Se4 Mo3 Co1

human body =

4

Jaroslav Vrba: Ecological stoichiometryALTER-net Summer School, Peyresq, 2008

Page 7: a bottleneck for biodiversity and ecosystem services · – a bottleneck for biodiversity and ecosystem services. ... • Biochemical and physiological constraints of life ... Our

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 stoichiometryALTER-net Summer School, Peyresq, 2008

Page 8: a bottleneck for biodiversity and ecosystem services · – a bottleneck for biodiversity and ecosystem services. ... • Biochemical and physiological constraints of life ... Our

Stoichiometry of cells – cell chemistry

Composition of biomolecules – biochemical stoichiometry

Selection for C, N & P in biochemical evolution

Jaroslav Vrba: Ecological stoichiometryALTER-net Summer School, Peyresq, 2008

Growth Rate Hypothesis (GRH) = ribosomes

C(energy)=saccharides & lipids

C+N=proteins

Page 9: a bottleneck for biodiversity and ecosystem services · – a bottleneck for biodiversity and ecosystem services. ... • Biochemical and physiological constraints of life ... Our

Growth Rate Hypothesis (GHR)

r-strategists

= ideal phytoplankton

K-strategists N2 fixers(N:P>40)

Jaroslav Vrba: Ecological stoichiometryALTER-net Summer School, Peyresq, 2008

Arrigo (2005) Nature 437

Page 10: a bottleneck for biodiversity and ecosystem services · – a bottleneck for biodiversity and ecosystem services. ... • Biochemical and physiological constraints of life ... Our

Growth Rate Hypothesis (GHR)

r-strategists (Cladocera) = high need in P !

Jaroslav Vrba: Ecological stoichiometryALTER-net Summer School, Peyresq, 2008

Page 11: a bottleneck for biodiversity and ecosystem services · – a bottleneck for biodiversity and ecosystem services. ... • Biochemical and physiological constraints of life ... Our

Homeostasis of heterotrophic consumers

Growth (resource utilization) may change stoichiometry

Are you what you eat?

Autotrophs: rather YES Heterotrophs: mostly NOT

Jaroslav Vrba: Ecological stoichiometryALTER-net Summer School, Peyresq, 2008

Page 12: a bottleneck for biodiversity and ecosystem services · – a bottleneck for biodiversity and ecosystem services. ... • Biochemical and physiological constraints of life ... Our

Homeostasis of vertebrates

Structural investment = skeleton

changes fundamentally needs in resource stoichiometry

great need in P & Ca !

Jaroslav Vrba: Ecological stoichiometryALTER-net Summer School, Peyresq, 2008

Page 13: a bottleneck for biodiversity and ecosystem services · – a bottleneck for biodiversity and ecosystem services. ... • Biochemical and physiological constraints of life ... Our

great need in P & Ca !

Jaroslav Vrba: Ecological stoichiometryALTER-net Summer School, Peyresq, 2008

Homeostasis of vertebrates

Structural investment = skeleton

changes fundamentally needs in resource stoichiometry

even during ontogenesis

Pilati & Vanni (2007) Oikos 116

Page 14: a bottleneck for biodiversity and ecosystem services · – a bottleneck for biodiversity and ecosystem services. ... • Biochemical and physiological constraints of life ... Our

Stoichiometry of populations & communities

Jaroslav Vrba: Ecological stoichiometryALTER-net Summer School, Peyresq, 2008

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

Page 15: a bottleneck for biodiversity and ecosystem services · – a bottleneck for biodiversity and ecosystem services. ... • Biochemical and physiological constraints of life ... Our

Stoichiometry of populations & communities

Jaroslav Vrba: Ecological stoichiometryALTER-net Summer School, Peyresq, 2008

In particular effects of P supply should impinge on fitness

& drive evolutionary change

Jeyasingh & Weider (2007) Mol. Ecol. 16

Page 16: a bottleneck for biodiversity and ecosystem services · – a bottleneck for biodiversity and ecosystem services. ... • Biochemical and physiological constraints of life ... Our

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(Fabaceae)

Jaroslav Vrba: Ecological stoichiometryALTER-net Summer School, Peyresq, 2008

Schade et al. (2003) Ecol. Lett. 6

Page 17: a bottleneck for biodiversity and ecosystem services · – a bottleneck for biodiversity and ecosystem services. ... • Biochemical and physiological constraints of life ... Our

Jaroslav Vrba: Ecological stoichiometryALTER-net Summer School, Peyresq, 2008

Food web stoichiometry

Top-down: cascading effect of predation and resource

stoichiometry

Page 18: a bottleneck for biodiversity and ecosystem services · – a bottleneck for biodiversity and ecosystem services. ... • Biochemical and physiological constraints of life ... Our

Top-down: cascading effect of predation and resource

stoichiometry determine un/successful biomanipulation

☺☺

Jaroslav Vrba: Ecological stoichiometryALTER-net Summer School, Peyresq, 2008

Food web stoichiometry

Page 19: a bottleneck for biodiversity and ecosystem services · – a bottleneck for biodiversity and ecosystem services. ... • Biochemical and physiological constraints of life ... Our

Nutrient regeneration is species specific (26 vertebrates)

Jaroslav Vrba: Ecological stoichiometryALTER-net Summer School, Peyresq, 2008

Food web stoichiometry

Vanni et al. (2002) Ecol. Lett. 5

Page 20: a bottleneck for biodiversity and ecosystem services · – a bottleneck for biodiversity and ecosystem services. ... • Biochemical and physiological constraints of life ... Our

Consumer and resource stoichiometry controls efficiency

(GGEC) = carbon + energy dissipation !

Redfield

Jaroslav Vrba: Ecological stoichiometryALTER-net Summer School, Peyresq, 2008

Food web stoichiometry

Page 21: a bottleneck for biodiversity and ecosystem services · – a bottleneck for biodiversity and ecosystem services. ... • Biochemical and physiological constraints of life ... Our

Vegetation Eleocharis (control)

Eleocharis +P

Typha +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 microb. biom. N:P 3.1 0.5 3.1 interstic. SRP (µg/l) 0.8 7.3 16.1

Experimental eutrophication of wetlands (Belize: +P)

Soil microbes

Soil P

Plants

Jaroslav Vrba: Ecological stoichiometryALTER-net Summer School, Peyresq, 2008

Ecosystem stoichiometry

Page 22: a bottleneck for biodiversity and ecosystem services · – a bottleneck for biodiversity and ecosystem services. ... • Biochemical and physiological constraints of life ... Our

Vegetation Eleocharis (control)

Eleocharis +P

Typha +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 microb. biom. N:P 3.1 0.5 3.1 interstic. SRP (µg/l) 0.8 7.3 16.1

Soil

microbesSoil P

Plants

Jaroslav Vrba: Ecological stoichiometryALTER-net Summer School, Peyresq, 2008

Experimental eutrophication of wetlands (Belize: +P)

= distinct stoichiometry of producers / detritus (litter)Rejmankova & Houdkova (2006) BGC 80, Šantrůčková et al. (unpubl.)

Ecosystem stoichiometry

Page 23: a bottleneck for biodiversity and ecosystem services · – a bottleneck for biodiversity and ecosystem services. ... • Biochemical and physiological constraints of life ... Our

Jaroslav Vrba: Ecological stoichiometryALTER-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 = malariaGrieco et al. (2005) J. Vector Ecol. 30Grieco et al. (2006) J. Med. Entomol. 43Grieco et al. (2007) J. Vector Ecol. 32

Page 24: a bottleneck for biodiversity and ecosystem services · – a bottleneck for biodiversity and ecosystem services. ... • Biochemical and physiological constraints of life ... Our

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 stoichiometryALTER-net Summer School, Peyresq, 2008

Ecosystem stoichiometry vs. productivity ?

Page 25: a bottleneck for biodiversity and ecosystem services · – a bottleneck for biodiversity and ecosystem services. ... • Biochemical and physiological constraints of life ... Our

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 stoichiometryALTER-net Summer School, Peyresq, 2008

Ecosystem stoichiometry vs. productivity ?

Page 26: a bottleneck for biodiversity and ecosystem services · – a bottleneck for biodiversity and ecosystem services. ... • Biochemical and physiological constraints of life ... Our

OECD Model (Vollenweider): seston chlorophyl–TP relationship

Jaroslav Vrba: Ecological stoichiometryALTER-net Summer School, Peyresq, 2008

Ecosystem stoichiometry

Page 27: a bottleneck for biodiversity and ecosystem services · – a bottleneck for biodiversity and ecosystem services. ... • Biochemical and physiological constraints of life ... Our

Anthropogenic impacts =

deposition, fertilisers, eutrophication…

Human activity turns

both landscape and the planet in “a large-scale excrement”…

Jaroslav Vrba: Ecological stoichiometryALTER-net Summer School, Peyresq, 2008

Ecosystem stoichiometry

Page 28: a bottleneck for biodiversity and ecosystem services · – a bottleneck for biodiversity and ecosystem services. ... • Biochemical and physiological constraints of life ... Our

lakes

coastal ecosystems

Jaroslav Vrba: Ecological stoichiometryALTER-net Summer School, Peyresq, 2008

Ecosystem stoichiometry

Anthropogenic impacts =

deposition, fertilisers, eutrophication…

Page 29: a bottleneck for biodiversity and ecosystem services · – a bottleneck for biodiversity and ecosystem services. ... • Biochemical and physiological constraints of life ... Our

Ecosystem services – e.g., of coastal ecosystems

Si:N

Jaroslav Vrba: Ecological stoichiometryALTER-net Summer School, Peyresq, 2008

Ecosystem stoichiometry

Page 30: a bottleneck for biodiversity and ecosystem services · – a bottleneck for biodiversity and ecosystem services. ... • Biochemical and physiological constraints of life ... Our

Si:N

Si:N

Jaroslav Vrba: Ecological stoichiometryALTER-net Summer School, Peyresq, 2008

Ecosystem services – e.g., of coastal ecosystems

Ecosystem stoichiometry

Ptacnik et al. (2005) Oikos 109

Page 31: a bottleneck for biodiversity and ecosystem services · – a bottleneck for biodiversity and ecosystem services. ... • Biochemical and physiological constraints of life ... Our

Ecosystem services – increase in [CO2] vs. production

Jaroslav Vrba: Ecological stoichiometryALTER-net Summer School, Peyresq, 2008

Ecosystem stoichiometry

Carbon sequestration ?Timber ?Biodiversity ?

Körner (2006) New Phytol. 172

Page 32: a bottleneck for biodiversity and ecosystem services · – a bottleneck for biodiversity and ecosystem services. ... • Biochemical and physiological constraints of life ... Our

Ecosystem services – increase in [CO2] vs. production

Jaroslav Vrba: Ecological stoichiometryALTER-net Summer School, Peyresq, 2008

Ecosystem stoichiometry

Carbon sequestration ?Timber ?Biodiversity ?

Körner (2006) New Phytol. 172

Page 33: a bottleneck for biodiversity and ecosystem services · – a bottleneck for biodiversity and ecosystem services. ... • Biochemical and physiological constraints of life ... Our

rapid rotation = decrease in mean biomass storage

Jaroslav Vrba: Ecological stoichiometryALTER-net Summer School, Peyresq, 2008

Ecosystem services – increase in [CO2] vs. production

Biofuel plantation is no sustainable solution !

Ecosystem stoichiometry

Körner (2006) New Phytol. 172

Page 34: a bottleneck for biodiversity and ecosystem services · – a bottleneck for biodiversity and ecosystem services. ... • Biochemical and physiological constraints of life ... Our

Ecosystem services – increase in [CO2] vs. production

Food/crops planted at 2×[CO2] = micronutrient malnutrition

cerealsall crops

Jaroslav Vrba: Ecological stoichiometryALTER-net Summer School, Peyresq, 2008

Loladze (2002) Trends Ecol. Evol. 17

Ecosystem stoichiometry

Page 35: a bottleneck for biodiversity and ecosystem services · – a bottleneck for biodiversity and ecosystem services. ... • Biochemical and physiological constraints of life ... Our

Ecological stoichiometry – a synthesis

Jaroslav Vrba: Ecological stoichiometryALTER-net Summer School, Peyresq, 2008

Page 36: a bottleneck for biodiversity and ecosystem services · – a bottleneck for biodiversity and ecosystem services. ... • Biochemical and physiological constraints of life ... Our

Ecological stoichiometry – a synthesis

Jaroslav Vrba: Ecological stoichiometryALTER-net Summer School, Peyresq, 2008

Page 37: a bottleneck for biodiversity and ecosystem services · – a bottleneck for biodiversity and ecosystem services. ... • Biochemical and physiological constraints of life ... Our

Ecological stoichiometry – a synthesis

Jaroslav Vrba: Ecological stoichiometryALTER-net Summer School, Peyresq, 2008

Gaia, a global ecosystem? = homeostasis of the Ocean

Arrigo K.R. (2005) Marine microorganisms and global nutrient cycles. Nature 437, 349–355.

Page 38: a bottleneck for biodiversity and ecosystem services · – a bottleneck for biodiversity and ecosystem services. ... • Biochemical and physiological constraints of life ... Our

Jaroslav Vrba: Ecological stoichiometryALTER-net Summer School, Peyresq, 2008

Global homeostasis of soils

Cleveland C.C. & Liptzin D. (2007) C:N:P stoichiometry in soil: is there a ‘‘Redfield ratio’’ for the microbial biomass?. Biogeochemistry 85, 235–252.

Ecological stoichiometry – a synthesis

Our analysis indicates that, similar to marine phytoplankton, element con-centrations of individual phylogeneticgroups within the soil microbial community may vary, but on average, atomic C:N:P ratios in both the soil (186:13:1) and the soil microbial biomass (60:7:1) are well-constrained at the global scale.

Page 39: a bottleneck for biodiversity and ecosystem services · – a bottleneck for biodiversity and ecosystem services. ... • Biochemical and physiological constraints of life ... Our

… The issue is not that we must wait for a future biology to arrive, but that we should notice and take good stock of what is already underway.

Ecological stoichiometry – a challenge

Page 40: a bottleneck for biodiversity and ecosystem services · – a bottleneck for biodiversity and ecosystem services. ... • Biochemical and physiological constraints of life ... Our

Ecological stoichiometry – a challenge

Jaroslav Vrba: Ecological stoichiometryALTER-net Summer School, Peyresq, 2008

Thank you for considering ecological stoichiometry.

Take it as a homework !

Thank you for attention…

…and the ALTER-net Summer School conveners for invitation.