Ecology and Evoliution

3rd

6 Nov. 2012

A little advanced Ecology

Ecological Stoichiometry
and
Environmental Changes

Todays outline
1. Ecological Stoichiometry!
2. Light & Nutrient balance!
Theory/lab & field experiments!

3. CO2 and herbivore growth!

4. Function of algal diversity !

Light (energy)

Grazing food web

Detrital food web

CO2

Detritus

Producers
Nutrients

N, P

Environmental Disturbances
Eutrophication (Nutrients)

Global warming (CO2, Temperature)

Climate change (H2O, Light, Seasonality)

Changes in absolute (rate) and relative

(ratio) inputs of energy and materials.

http://www.city.sapporo.jp/kankyo/
gaikyo/taikiosen/taiki.htm

http://www.lbri.go.jp/omia/54/omia54-0.htm

http://www.kankyo.metro.tokyo.jp/sgw/page2.html

Stoichiometry? !
Numerical relationship in elements between
substrates and products of chemical reactions

An example from chemistry:

6CO2 + 6H2OC6H12O6 + 6O2
!
Each side has 6 Cs, 12 Hs and 18 Os

photosynthesis

Ecological
Stoichiometry? !
It examines how balance of elements in
organisms shapes ecological processes

Prey-predator interactions
(CX,PY)Predator + (CA,PB)Prey
Q(CX,PY) Predator + (Ca,Pb)Waste
!
Y+B=QY+b

Sterner & Elser (202) Ecological Stoichiometry, Princeton Univ

http://www.tokyo-med.ac.jp/genet/picts/cell.jpg

Ribosome: organelles where Proteins are synthesized

Cell with rich Ribosomehigh protein synthesis rate
Thus, P-rich cells are high in activity for material production
Heterotrophs with rich P is higher in their growth rate

http://www.hr-online.de/fs/schulfernsehen/genzeit.html

RNA and animal growth rate

Relative growth (per day)

Comparison among various animals and bacteria

%RNA in body (cells)

Plant (Algae) Stoichiometry

Redfield ratio

Favorable

P, N

6O2
6CO2!
6H2O

Nutrient !
deficient
6CO2!
6H2O

C6H12O6

6O2

P, N

6O2
C6H12O6

C-rich cells

C6H12O6

Light !
deficient
6CO2!
6H2O

C106N16P1

P, N

600

N30P1

Redfield ratio

C106N16P1

2-6

Redfield sea
106CO2+16NO3+H2PO4+122H2O+18H+
(C106H263O110N16P1) + 138O2

Alfred C. Redfield
November 15, 1890 March 17, 1983
By Roger Revelle

Homeostasis in elemental ratios

Autotrophs

Heterotrophs

Homeostasis

N:C Fungus

N:P algae

100

10

1
1

10

100

0.1
0.01
0.001

0.0001
0.0001 0.001

N:P medium

0.1

0.01

Daphnia P%

C:N bacteria

10

10

C:N substrate

N:C medium

10

1
0.1
0.01
0.01

0.1

Algal P%

10

Algae-Herbivore Stoichiometry
Herbivorous [Daphnia]

Algae
Variable P:C organisms

C
CP

High P:C organisms

P
P C

P
P

C
29

TER is C to element ratio below which animal growth is limited by the

content of that element (N, P, etc.) in food rather than food abundance (C).
In the case of P
P-net intake = [C-net intake] x ZP:C

I C FP: CP P = (I CC C )ZP: C

FP: C

kC ZP: C
P
=

P
I CP
(kC =

I CC C
)
IC

Minimal model (P=1, P=0)

TER

= F

P :C

* = kC Z P :C

(eq1)
(eq2)

ingestion rate
assimilation efficiency
loss rate

FC :P food C:P ratio

ZC :P animal C:P ratio

For Daphnia

gross growth efficiency

ZP:C ~0.03, kC ~0.3

TER P:C~ 9 gP/mgC

Urabe, J., and Y. Watanabe. 1992. Possibility of N or P limitation for planktonic cladocerans: an experimental test. Limnol. Oceanogr 37: 244-251.
Hessen D. O. 1992. Nutrient element limitation of zooplankton production. Am. Nat., 140: 799-814.

Threshold Elemental Ratio

Threshold elemental ratio (TER)

below which animal growth is limited by nutrient
content (P, N) in food rather than food abundance (C)
Minimum model (P=1, bP=0)
In the case of P
TER= Body P:C ratio

x Gross Growth Efficiency for C
For Daphnia

TERP:C ~ 8.6 gP/mgC or C:P ~300 atomic ratio
30

Urabe and Watanabe (1992) Limnology and Oceanography, 37: 244-251

Light-Nutrient Balance and

Herbivore growth

Algal P:C ratio

Light intensity received each flask

Growth rate

Ingestion rate

(Carbon/day)

PPPP

Algal Abundance

(mgC/L)

A theory

Light intensity received each flask

Experiment
P 10 M (high nutrient)

High light

Low light
P

Low light

1.6 M (low nutrient)

High light

Results"

10 mmol/L

75

50
5
25

TER

0
10

100

Growth rate

(gDW/ind)

30

20

10

0
10

100

Light intensity ( E/m2/s)

Urabe & Sterner (1996) PNAS, 93:8465-8469

Algal P:C ratio

(gP/mgP)

10

Algal Abundance

(mgC/L)

High Nutrients

Results"
Low Nutrients

75

50

5
25

TER

0
10

100

Growth rate

(gDW/ind)

30

20

10

0
10

100

Light intensity ( E/m2/s)

Urabe & Sterner (1996) PNAS, 93:8465-8469

Algal P:C ratio

(gP/mgP)

P
1.6 mmol/L

Algal Abundance

(mgC/L)

10

In general.
Stephen Elser

I like fine
weather, but if
this makes my
food bad.

Background
The atmospheric CO2 level is
expected to double or treble
by 2100 years.

(IPCC 4th assessment report, 2007)

Most lakes are CO2

saturated
5.0
4.5

140
45

145

log[pCO2, ppm]

4.0

58 lakes

3.5
3.0

370ppm

2.5

40

2.0

19 lakes

1.5
1.0
1.0
35

20

Number of lakes

Elevation of atmospheric CO2

likely rises pCO2 in aquatics
much more via terrestrial input
CO2

CO2

DOC

CO2
DOC

Stoichiometric impacts
of rising pCO2 on a
plankton herbivore

Algal culture experiments

pCO2; 360 ppm or 2000 ppm

Semi-continuous culture (30% dilution per 2 days: 20C)

Nutrients: 1.5 M P, N:P=80; Light: 150 moles m-2 s-1

Abundance

!
Algae



Scenedesmus (green algae)



Cyclotella (diatom)



Synechococcus (cyanobacteria)

Steady state

biomass

Daphnia growth

experiment

Exponential

growth rate

Body mass changes for 5 days

days

Daphnia

Growth rate

(/d)

Algal P:C ratio

Algal biomass

(x 103)
(mgC/L)

Results
20

Green algae

20

16

16

12

12

0
6

CO2

TER

2
0

0.4

CO2

TER

CO2

0.2

Diatoms

0.4

CO2

CO2

0.2

360

CO2

2000

360

2000

CO2 treatment (ppm)

Daphnia

Growth rate

(/d)

Algal P:C ratio

Algal biomass

(x 103)
(mgC/L)

Results
20

Green algae

20

Diatoms

20

16

16

16

12

12

12

0
6

CO2

TER

2
0

0.4

CO2

CO2

0.4

360

CO2

2000

CO2

n.s

TER

CO2

0.4

0.2

0.2

n.s

TER

0.2

Cyanobacteria

360

CO2

2000

CO2 treatment (ppm)

Urabe et al (2003) GCB 9:818-825, Urabe & Waki (2009) GCB 15:523-531

2/6

CO2

n.s

0/6

CO2

360

2000

A pitfall
!
Various algal species co-occur in nature.

!
!
!
!

Herbivores response may differ between

feeding on single algal species and
feeding on multiple algal species.

Mixed culture experiments

pCO2; 360 ppm or 2000 ppm

Semi-continuous culture (30% dilution per 2 days: 20C)

Nutrients: 1.5 M P, N:P=80; Light: 150 moles m-2 s-1

!
Algae



Scenedesmus (green algae)



Cyclotella (diatom)



Synechococcus (cyanobacteria)

Abundance

Mixed culture
Steady state

biomass

Daphnia growth

experiment

Exponential

growth rate
Body mass changes for 5 days

48

days

Results: two species

Green algae +

Cyanobacteria

Daphnia

Growth rate

(/d)

Algal P:C ratio

Algal biomass

(x 103)
(mgC/L)

20
16
12
8
4
0
6

CO2

TER

2
0
0.4

CO2

CO2

360

2000

0.2

CO2 treatment (ppm)

51

Urabe & Waki (2009) Global Change Biology 15:523-531

Results: two species

Daphnia

Growth rate

(/d)

Algal P:C ratio

Algal biomass

(x 103)
(mgC/L)

Green algae +

Cyanobacteria

Diatoms +

Green algae

20

20

16

16

12

12

0
6

CO2

TER

2
0
0.4

CO2

TER

CO2

0.2

0
0.4

CO2

CO2

0.2

CO2

360

2000

360

2000

CO2 treatment (ppm)

51

Urabe & Waki (2009) Global Change Biology 15:523-531

Results: two species

Daphnia

Growth rate

(/d)

Algal P:C ratio

Algal biomass

(x 103)
(mgC/L)

Green algae +

Cyanobacteria

Diatoms +

Green algae

Cyanobacteria +

Diatoms

20

20

20

16

16

16

12

12

12

0
6

CO2

TER

2
0
0.4

CO2

TER

CO2

0.2

0
0.4

CO2

360

2000

CO2

CO2

0
0.4

TER

0.2

360

CO2

2000

CO2 treatment (ppm)

51

0.2

Urabe & Waki (2009) Global Change Biology 15:523-531

CO2

n.s

CO2

360

2000

Green algae +

Diatoms +

Cyanobacteria

Algal P:C ratio

(x 103)

Algal biomass

(mgC/L)

Results: three species

20
16
12
8
4
0
6
4

CO2

Threshold P:C

2
0

Daphnia

Growth rate

(/d)

0.4

CO2

0.2

n.s
0

CO2

360

2000

CO2 treatment (ppm)

Urabe & Waki (2009) Global Change Biology 15:523-531

Results: three species

16
12
8
4

CO2

Threshold P:C

0.4

CO2

0.2

n.s
0

20
16
12
8
4

CO2

360

2000

CO2 treatment (ppm)

Algal P:C ratio

(x 103)

Daphnia

Growth rate

(/d)

Algal biomass

(mgC/L)

20

Green algae

Daphnia

Growth rate

(/d)

Algal P:C ratio

(x 103)

Algal biomass

(mgC/L)

Green algae +

Diatoms +

Cyanobacteria

6
4

CO2

Threshold P:C

2
0

0.4

CO2

CO2

0.2

360

2000

CO2 treatment (ppm)

Urabe & Waki (2009) Global Change Biology 15:523-531

!!!
Adverse eects of rising CO2 on herbivore
occur in single food environments but not
in multiple food environments.
!

Numerically minor algae play nutritionally

important roles.

Nutritionally

unimportant food become

beneficial in high CO2.

Algal

diversity serves to mitigate the

adverse effects of rising CO2.

20
16
12
8
4

6
4

CO2

Threshold P:C

2
0

Daphnia

Growth rate

(/d)

0.4

CO2

0.2

n.s
0

CO2

360

2000

CO2 treatment (ppm)

Algal P:C ratio

(x 103)

Green algae
20
16
12
8
4

Daphnia

Growth rate

(/d)

Algal P:C ratio

(x 103)

Algal biomass

(mgC/L)

Green algae +

Diatoms +

Cyanobacteria

Algal biomass

(mgC/L)

Why does this occur?

6
4

CO2

Threshold P:C

2
0

0.4

CO2

CO2

0.2

360

2000

CO2 treatment (ppm)

Why high growth is maintained at high CO2 when fed multple algae?

Possible mechanisms
Feeding

compensation

Stimulate feeding
activities

Assimilation

enhancement

Functionally increase
assimilation efficiency

Urabe & Waki (2009) Global Change Biology 15:523-531

54

Nutritional

complementarity

Complement

deficient nutrients
each other, or
ameliorate some
harmful substances.

Lab Experiment
again

Measurements for
G.I.A. rates

Growth rate

Ingestion rate (32P)

Assimilation rate (32P)

360ppm

Low CO2

2000 ppm
Food abundance 1mgC/L

1 species
treatment

2-3 species
treatment

High CO2

Food abundance 1mgC/L

1 species
treatment

2-3 species
treatment

Results

Growth rat ( /d)

0.4

Growth rate vs Algal P:C ratio

Low
CO2

0.3

High

CO2

0.2

Single algae!

0.1

Dual algae!

r 2 = 0.109
0
0

2.5

7.5

Algal P (gP/L) or P:C ratio (gP/mgC)

10

Treble algae

Results

Growth rat ( /d)

0.4

Growth rate vs Algal P:C ratio

Low
CO2

0.3
0.2

Single algae!

0.1

Dual algae!

r 2 = 0.109
0
0

2.5

7.5

10

Algal P (gP/L) or P:C ratio (gP/mgC)

0.4

Growth rate ( /d)

High

CO2

Growth rate vs Ingested P

0.3
0.2
0.1
0

r 2 = 0.444
0

10

Ingestion for P (ngP/h/ind)

15

20

Treble algae

Algal diversity and feeding performance

Feeding rate (ml/h/ind)

Low CO2 (high P content algae)

High CO2 (low P content algae)
2.0
1.5
1.0
0.5
0.0

2-3

Results
Growth rate ( /d)

0.4

Growth rate vs Ingested P

Low
CO2

0.3

Single algae!

0.2

0.1
0

High

CO2

Dual algae!

r 2 = 0.444
0

10

Ingestion for P (ngP/h/ind)

15

!
20

Treble algae

Results
Growth rate ( /d)

0.4

Growth rate vs Ingested P

Low
CO2

0.3

Single algae!

0.2

0.1
0

Dual algae!

r 2 = 0.444
0

10

15

!
20

Ingestion for P (ngP/h/ind)

Growth rate (/d)

0.4

Growth rate vs Assimilated P

0.3
0.2
0.1
r 2 = 0.726
0

High

CO2

Assimilation rate for P (ngP/h/ind)

12

Treble algae

Assimilation efficiency for P (%)

Algal diversity and digestive performance

low CO2 (high P content algae)
High CO2 (low P content algae)
0.8

0.7

0.6

0.5

2-3

Why high growth is maintained at high CO2 when fed multiple algae?

Possible mechanisms
Feeding

compensation

Stimulate feeding
activities

54

Assimilation

enhancement

Functionally increase
assimilation efficiency

Nutritional

complementarity

Complement

deficient nutrients
each other, or
ameliorate some
harmful substances.

Summary (I)
Ecological Stoichiometry (ES) examines the
balance of elements in ecological interactions.
It is useful to understand the responses of
ecosystems to environmental changes (light/CO2/
nutrients) via quantity and quality of plants.

Putative increase in pCO2 likely increase plant
(algal) abundance but reduce the nutrient contents.

Thus, algal species beneficial at present may not
be beneficial for herbivore in future (high CO2
world).

Summary (II)

However, adverse effects of rising CO2 on

herbivores can be mitigated by algal diversity

!
Algal

diversity is certainly important for

sustaining herbivores especially at high CO2
environment in future.

Finally, we need to study more on assimilation

activities of herbivores to understand their
response to changes in food condition.