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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)