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“Dynamics of stoichiometrical and metabolomical traits under climate change” Jordi Sardans, Albert Rivas-Ubach, Albert Gargallo-Garriga, Ifigenia Urbina, Jofre Carnicer, Marc Estiarte, Iolanda Filella, Joan Llusià, Romà Ogaya, Josep Peñuelas Global Ecology Unit CREAF – CSIC – CEAB Barcelona Stoichiometry-metabolomic studies Useful traits to detct and study plant responses to environmental change Climate change Plant Metabolism and stoichiometry shifts Sensitivity enough to detect the dynamics of plant responses Mechanisms and functions involved in the organism response Potential improvement of global climate and balance models Clues to global ecosystem impacts mainly by the effects on trophic webs Long-time field studies My first studies of Biogeochemistry in the Global Ecology Unit Climanani Garraf Site Prades site Drought impacts N and P concentrations and contents in Mediterraenan ecosystems 1. Decreases in soil enzyme and root-enzyme activity 2. Decreases of N and P stocks in aboveground biomass 3. Increases in total soil N and P stocks but decreses of Total Aboveground plant-available stocks Quercus ilex a 4. Changes in plant N:P stoichiometry 50 Control Four general results observed in all studies a 40 0.05 a Arbutus unedo 8 Runoff exclussion plus partial rainfall exclussion ab b 20 a a b 10 0 Protease -Glucosidase Data from Sardans & Peñuelas (2005) Soil Biology and Biochemistry (a) (a) 4 0 (b) -4 Leaves 0.1 a 0.05 Wood (b) Aboveground Biomass fraction Sardans & Peñuelas (2007) Functional Ecology a b Erica multiflora 350 0.4 0.3 a 0.2 ab b 280 Control 210 Drought Warming 70 C D Treatments DD Sardans and Peñuelas (2007) Functional Ecology a 140 a 0.1 0 N/P a 30 Soil Soluble Pi/Po Runoff exclussion b b 50 -1 P increment ab b (kg ha ) (2005-1999) 0-15 cm soil depth 0.1 -1 Control -1 -1 Enzyme activity (g substrate hydrolized g soil h ) After four years of drought (Mediterranean forest) Soil Soluble Po (mg g ) Soil Soluble Ptotal (mg g-1) Drought 0 b b b b 0 Leaves Stem Leaf-Litter Sardans et al. (2008) Global Change Biology Changes in N/P ratio is not a minor question C/N/P ratio related to several ecosystems process In freshwater ecosystems Growth Rate Hypothesis (GR) Elser et al. (1996) In terrestrial ecosystems N/P RNA (P-rich) Growth rate No conclussive results of GR at this moment Related to species diversity Relatedto species style of life Related to soil trophic web structure But K-stocks were also afected by drought Plant biomass In soil 0-15 cm soil depth 0.1 10 5 b a ab b b a -1 (mg g ) 15 Soil soluble K a 0.05 ab b b b D DD 0 Leaves Stems Erica multiflora 10 8 a a 6 4 b 2 0 Leaves Stems 0 Soil soluble K/total K -1 Absolute aboveground K accumulation (kg ha ) (1999-2005) Globularia alypum 21 0.004 a 0.002 0 C Treatments Sardans and Peñuelas (2007) Functional Ecology Sardans et al. (2008) Plant and Soil Drought changes whole foliar elemental composition Soil stocks and biomass concentrations of other several elements (Mo, S or Ca) had been changed by drought Sardans et a. (2008) Biogeochemistry, Sardans et al. (2008) J. Geophys. Res. Globularia alypum Quercus ilex 1.0 1.0 Mg Fe Ca S Mg PC2 (19%) P 0.5 PC2 (19%) 0.5 N 0.0 Fe Na Mo Na Mo 0.0 PS C -0.5 -0.5 Ca N C K K -1.0 -1.0 -1.0 -0.5 0.0 0.5 -1.0 1.0 -0.5 0.0 PC1 (34%) 4 D D 4 1 D 0 D D C PC2 (29%) Drought 2 PC2 (19%) 1.0 6 3 C C -1 Drought 2 Warming W W W 0 D D C C Peñuelas et al. (2008) Pol J Ecol Control -2 -3 -4 0.5 PC1 (42%) C -2 Control C -4 -2 0 PC1 (34%) 2 4 -4 -2 0 2 PC1 (42%) 4 6 Climate change impact in terrestrial ecosystems stoichiometry: how can we advance? Experimental, observationals and review studies (reviews and meta-analysis) Sardans et al. (2012a) Biogeochemistry Sardans et al. (2012b) Pers Plant Ecol Evol Syst Sardans and Peñuelas (2012) Plant Physiology Capture our attention in current studies limitations How can improve stoichiometry studies to be a useful trait to detect dynamics shifts in terrestrial plant vegetation What dowe need? 1.Most studies have been focused on N and P (N/P ratio) 1.More elements such as K, but also Mg, S or Ca among others should be considered 2. Stoichiometric studies of terrestrial plants have been mailny focused on foliar tissues, specially in trees 2. Plants also allocate nutrients to other tissues. We need when possible, take into account whole plant stoichiometry 3. N/P stoichiometry of higher plants not only depends of the allocation to growth, several other functions can be important sinks for nutrients 3. To gain a global knowledgment of the functional causes underlie stoichiometric shifts in plant responses to climate change We propose to solve these constrains and advance in the frame of climate change impacts on terrestrial plant communities Plant functional response Make the next reasonig Climate change Changes in gene expresion Wholel plant stoichiometrical change 1 Changes in metabolism and molecular structure Change in element use 1. Ecometabolomic studies 2. Include other elements such as K. Study the elemental composition and its shifts as a whole Soil resources 2 Impacts on biotic relationships (competition, herbivorism….) 2 We began to use metabolomics analyses in our studies We developed a method to conduct metabolomic analyses with field sampling Rivas-Ubach et al. (2013) Methods in Ecology and Evolution Plants under different climate conditions shift their metabolism. The corresponding metabolomic analysis informs on the molecular causes underlying the shifts in elemental composition and stoichiometrical ratios Elemental Stoichiometry S Spring Season N/P Sugars Amino acids Rivas-Ubach et al. (2012) PNAS Metabolome Preliminary Results Metabolomic profiling of Quercus ilex. Seasonal PCA Drought: Polyphenolics (antioxidants) Potassium Oposite responses of roots and leaves Ecometabolomics : a tool for several ecological estudies Great sensitivity to detect plant responses • When the plant is wounded the metabolome changes No-wounded Wounded Sardans et al. (2013) Plant Biology Current ecometabolomic-stoichiometric studies Climate change effects on the populations of the south border of the distribution Area of important European forest species Fagus sylvatica Quercus ilex Pinus uncinata Stoichiometry and Metabolism on plant-herbivore relationship Pine processionary moths Thaumetopoea pityocampa Until this moment the results show that metabolomics can be an useful tool: 1. To give a global view of plant functions involved in plant responses 2.To explain the causes of plant elemental compostion and stoichiometrical shifts under drought 3. To improve the kowledge of the metabolic pathways up– and down-regulated under drought . 4.Is sensitive enough to detect plant molecular and elemental shifts under different environmental conditions through time 5.To know plant responses to herbivore attack Second: Whole plant elemental composition 2 The role of potassium Data from Catalan National Forestal Inventory Precipitation (L m-2 yr-1) 400-500 900-1000 500-600 1000-1100 600-700 1100-1200 700-800 > 1200 50 Km Foliar K concentration (mg g-1) Foliar K content (kg ha-1) Foliar K:C content ratio Foliar K:N content ratio 30 0.007- 4.31 8.2 – 72.7 0.5897 – 1.6434 3.41 – 4.52 4.31 – 7.65 72.7 – 92.2 1.6434 – 2.0491 4.52 – 5.67 7.65 – 12.66 92.2 – 114.2 2.0491 – 2.5288 5.67 – 7.62 12.66 – 22.43 114.2 – 152.7 2.5288 – 3.4242 > 7.62 > 22.43 > 152.7 Autumn a 20 ab aa b ab ab b 10 0 1.0 – 3.41 Summer Winter Spring -1 800-900 K content in foliar biomass (kg ha ) < 400 Evergreens Conifers Deciduous > 3.4242 K content is related with MAP Sardans et al. (2012) Functional Ecology Species adapted to dry climate have higher capacity to allocate more K to foliar biomass during summer Have we neglegted the K limiting role in terrestrial ecosystems? Review of published data 40 Number of studies No-limiting Limiting 33 75% of field studies have observed that K limits growth of terrestrial plants in field conditions 30 20 11 10 3 1 0 Grasslands Forests Vegetation type Sardans et al. (2012) Global Change Biology (in preparation) Climate change effects on global plant elemental composition Different elements plays different functional and structural functions C (structure,..) N (growth, light capture, metabolism functioning,..) P (growth, energy transfer,…) K (water economy, internal transport,…) Mg (light capture,…) “Biogeochemical niche” Each species, as a singular evolutionary product, should have an optimum elemental composition as consequence of optimum function Peñuelas et al. (2010) Global Change Biology Consequence of the optimum adaptation to maximize species fitness in determined abiotic and biotic circumstances, i. e. consequence of long-term genetic adaptation (Genotype) , but also of short-term capacity to respond under certain limits to life-time environmental competition shifts (Phenotype flexibility) Native and alien species in Hawai Principal component 2: 15.4% 6 Different forest types in Catalonia 1.0 (A) K PN 0.5 4 0.5 Cu 2 0.0 -2 Fe -0.5 -4 PC2 (22.7%) Ni Alien Native -1.0 -3 0.0 -2 NP PK NK -1 0 1 2 -1.0 3 -0.5 Peñuelas et al. (2010) Global Change Biology -1.0 -0.5 0.0 0.5 1.0 PC1 (28.6%) 1.0 8 (B) F. excelsior Q. petrea 4 Q. canariensis F. sylvatica C. sativa Q. faginea N/P N/K 0.0 Ca S -0.5 Q. suber K A. unedo Q. humilis N P Q. ilex P. sylvestris P. pinea P. halepensis P. pinaster Q. cerrioides -1.0 -1.0 P. nigra P. uncinata A. alba Gymnosperms -0.5 -4 -3 -2 0.0 0.5 1.0 PC1 (27.7%) -2 -5 Mg P/K Q. humilis x cerrioides 2 0 C 0.5 PC2 (20.7%) PC2 (22.7%) (A) Angiosperms 6 -1 0 1 bc Mediterranean species PC1 (28.6%) 15 3 (C) PC 2 = - 1.68 + 0.0021 MAP R = 0.26, P < 0.0001 Separated throughout PC4 (explaining 12% of variability) b (1,2,7,8) c (6) (4) (5) a (3) (B) 10 P. halepensis (3) P. pinaster (2) (1) a P. nigra (ab) (1) (2) ab (3) b (4) c P. pinea (4) 0 Gimnosperms PC2 (20.7%) PC 2 5 0 Q. ilex (5) A. unedo (6) (5) d (6) d (7) e Q. suber (7) -3 Angiosperms C. sativa (8) (8) f -5 -6 -10 200 400 600 800 1000 MAP (L m-2 yr-1) 1200 1400 -0.6 0 0.6 PC1 (27.7%) 0.0 0.5 Principal component 1: 35.6% Principal component 1: 35.6% C -0.5 Mediterranean Wet temperate Alpine Transition Med-Temp. K N -6 Mg -1.0 Zn 0 Ca S 1.0 Na 1.2 1.0 Principal component 2: 15.4% “Biogeochemical niche” in action Until this moment the results show that the use of more elements that N and P Give a more global view of the use of resources Improve the sensitivity in the use of plant elemental composition shift to detect responses to environmental change Changing from local to global scale N eutrophication threatens to shift the global stoichiometry Global N deposition ~ 114 Mtones N 400 N from industrial fertilizers N emissions from fuel combustion N fixation of rice and legume crops 350 51 63 2 Total anthropogenic N Total anthropogenic P (mineral fertilizers) 300 Tg year-1 250 yr-1 Global P deposition ~ 3.3 Mtones N yr-1 3 0.3 Natural N2 fixation from continents plus oceans Continents N:P ~ 47 200 Oceans N:P ~ 370 150 Natural N2 fixation from continents 100 50 16 35.4 12 26.4 Redfield’s ratio 8 17.7 4 N:P ratio (molar basis) N:P ratio (mass basis) 0 Terrestrial plants N:P ~ 22-30 8.8 1860s 1900s 1950s 1980s 1990s 2000s Peñuelas et al. (2012) Global Change Biology Plankton and open ocean waters N:P ~ 15-16 Human induced chsnges on N/P ratios are already altering ecosystems function and biodiversity by the impacts on N/P ratio is already occuring Number of “Web of Science” studies reporting effects of changes in N:P ratios and its effects on ecosystems species composition and function Peñuelas et al. (2013) Natre Communications Submitted . Global P-cycle is being altered by human activity Peñuelas et al. (2013) Nature Communicatios(Submitted) Global climate and C-balance models related to global change should include the changes in nutrients balances and stoichiometry Our first attempt...Projections under different scenarios of the P demands to fixing C emitted by human activities. The models used were TAXIS, HadCM3, IPSL-CM2 , IPSL-CM4-LOOP, CSM, MPI, LLNL, FRCGC, UMD, UVIC, CLIMBER, BERNCC Phosphorus and N/P ratio could be gaining role in global capacity to C fixation and consequently in the global climatic control Peñuelas et al. (2013) Nature Geoscience (submitted) A new tool to study plant responses to climate change Ecometabolomics-stoichiometric studies Whole elemental composition shifts Whole metabolome shifts Sensitive traits to detect dynamic plant responses Improvement of the knowledge of the functional mechanisms underliying plant responses (growth, storage, defense, antistress….) Clues on further consequences throughout trophic web Useful information to improve global elemental budgets (C, N, P,..) and climatic models Global Ecology Unit CREAF – CSIC – CEAB Barcelona From elements to global scale Thank you by your attention and…………by no sleepping (if is the case)