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GPP DER NEE Make regression of night‐ time NEE ECOSYSTEM‐scale PROBLEM: What is GPP? EDDY COVARIANCE FROM FLUX TOWERS OBSERVES NET ECOSYSTEM‐ATMOSPHERE CO 2 EXCHANGE (NEE) We want photosynthesis (Gross primary production, or GPP) and Daytime Ecosystem Respiration (DER) Standard Solution (Reichstein et al., 2005): (1) DER = nighttime NEE regressed vs. Temperature

ECOSYSTEM‐scale PROBLEM: What is GPP? - kiss.caltech.edu

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Page 1: ECOSYSTEM‐scale PROBLEM: What is GPP? - kiss.caltech.edu

GPP DERNEE

Make regression of night‐time NEE

ECOSYSTEM‐scale PROBLEM:  What is GPP?EDDY COVARIANCE FROM FLUX TOWERS OBSERVES NET ECOSYSTEM‐ATMOSPHERE CO2 EXCHANGE (NEE)We want photosynthesis (Gross primary production, or GPP) and Daytime Ecosystem Respiration (DER)

Standard Solution (Reichstein et al., 2005):  (1) DER = nighttime NEE regressed vs. 

Temperature

Page 2: ECOSYSTEM‐scale PROBLEM: What is GPP? - kiss.caltech.edu

GPP DERNEE

EDDY COVARIANCE FROM FLUX TOWERS OBSERVES NET ECOSYSTEM‐ATMOSPHERE CO2 EXCHANGE (NEE)We want photosynthesis (Gross primary production, or GPP) and Daytime Ecosystem Respiration (DER)

Make regression of night‐time NEE

Standard Solution (Reichstein et al., 2005):  (1) DER = nighttime NEE regressed vs. 

Temperature  (& extrapolated to day)(for ~1‐2 week moving window)

(2) GPP = NEE – DER (residual)

Extrapolate from regression

ECOSYSTEM‐scale PROBLEM:  What is GPP?

Page 3: ECOSYSTEM‐scale PROBLEM: What is GPP? - kiss.caltech.edu

GPP DERNEE

Extrapolate from regression

Standard Solution (Reichstein et al., 2005):  (1) DER = nighttime NEE regressed vs. 

Temperature  (& extrapolated to day)(for ~1‐2 week moving window)

(2) GPP = NEE – DER (residual)

EDDY COVARIANCE FROM FLUX TOWERS OBSERVES NET ECOSYSTEM‐ATMOSPHERE CO2 EXCHANGE (NEE)We want photosynthesis (Gross primary production, or GPP) and Daytime Ecosystem Respiration (DER)

Make regression of night‐time NEE

Key Assumption:  Daytime physiology is like the nighttime

NOTE: DER is elevated

ECOSYSTEM‐scale PROBLEM:  What is GPP?

Page 4: ECOSYSTEM‐scale PROBLEM: What is GPP? - kiss.caltech.edu

Alternative:  Isotopic Scaling/Partitioning

One equations in two unknowns (GPP & Reco)

Key:  we need a second equation 

Page 5: ECOSYSTEM‐scale PROBLEM: What is GPP? - kiss.caltech.edu

Alternative:  Isotopic Scaling/Partitioning

Two equations in two unknowns (GPP & Reco)

δ13 is 13C content of sample relative to a standard 

Key:  Adding this second equation: 

Flux-weighted Isotopic composition of:

Net flux photosynthetic Respirationflux flux

Page 6: ECOSYSTEM‐scale PROBLEM: What is GPP? - kiss.caltech.edu

Alternative:  Isotopic Scaling/Partitioning

Two equations in two unknowns (GPP & Reco)

works by solving for GPP and DER (Reco during the day),given distinct stable isotopic signatures, δ13

Yakir & Wang 1996, Bowling et al. 2001, …

δ13 is 13C content of sample relative to a standard 

Key:  Adding this second equation: 

Page 7: ECOSYSTEM‐scale PROBLEM: What is GPP? - kiss.caltech.edu

Via eddy flux

Two equations in two unknowns (GPP & Reco)Requires:(1)  δ13

NEE to be measureable

δ13 is 13C content of sample relative to a standard 

Alternative:  Isotopic Scaling/Partitioning

Page 8: ECOSYSTEM‐scale PROBLEM: What is GPP? - kiss.caltech.edu

We measure the isotopic composition of respiration at night (using Keeling plots) 

Via eddy flux

Two equations in two unknowns (GPP & Reco)Requires:(1)  δ13

NEE to be measureable(2)  δ13

GPP to be different from δ13

R

Via nighttime

δ13 is 13C content of sample relative to a standard 

Alternative:  Isotopic Scaling/Partitioning

Page 9: ECOSYSTEM‐scale PROBLEM: What is GPP? - kiss.caltech.edu

Via eddy flux Via nighttime

Two equations in two unknowns (GPP & Reco)Requires:(1)  δ13

NEE to be measureable(2)  δ13

GPP to be different from δ13

R

Alternative:  Isotopic Scaling/Partitioning

Approach – scaling leaf physiology to the canopy:(1) leaf‐scale knowledge of physiology to constrain ecosystem‐scale iso‐

flux of GPP (depends on canopy‐scale stomatal conductance)→ (Farquhar et al. 1982 theory for leaf‐scale photosynthetic discrimination)

(2) Diffusion laws and ecosystem heat and water fluxes to constrain canopy scale stomatal conductance ‐‐ – now validated with flux measurements of carbonyl sulfide (OCS)  (Wehr, Commane et al. 2017)

Page 10: ECOSYSTEM‐scale PROBLEM: What is GPP? - kiss.caltech.edu

GPP

NEE Standard Daytime Ecosystem Respiration (DER)

NEE

Standard ‐GPP

Uptake

KEY RESULT: suppression of DER in light (& lower GPP)

Wehr et al.

DER

Page 11: ECOSYSTEM‐scale PROBLEM: What is GPP? - kiss.caltech.edu

GPP

NEE Standard Daytime Ecosystem Respiration (DER)

KEY RESULT: suppression of DER in light (& lower GPP)

Wehr et al.

DER

Uptake

Standard ‐GPP

NEE

DER

‐GPP

Page 12: ECOSYSTEM‐scale PROBLEM: What is GPP? - kiss.caltech.edu

GPP

NEE Standard Daytime Ecosystem Respiration (DER)

KEY RESULT: suppression of DER in light (& lower GPP)

Wehr et al.

DER

Uptake

Standard ‐GPP

NEE

DER

‐GPP

Page 13: ECOSYSTEM‐scale PROBLEM: What is GPP? - kiss.caltech.edu

WARNING! Forest HeterogeneitySeen in nighttime NEE

Wetland

Page 14: ECOSYSTEM‐scale PROBLEM: What is GPP? - kiss.caltech.edu

WARNING! Forest HeterogeneitySeen in nighttime NEE

Wetland

Page 15: ECOSYSTEM‐scale PROBLEM: What is GPP? - kiss.caltech.edu

GPP

NEE Standard Daytime Ecosystem Respiration (DER)

KEY RESULT: suppression of DER in light (& lower GPP)

Wehr et al.

DER

Uptake

Standard ‐GPP

NEE

DER

‐GPP

What is the mechanism?

Page 16: ECOSYSTEM‐scale PROBLEM: What is GPP? - kiss.caltech.edu

First detected ecosystem manifestation of the leaf‐level ‘Kok effect’ –inhibition of respiration by light?

RLIGHT

RDARK

light 

O2em

ission

Page 17: ECOSYSTEM‐scale PROBLEM: What is GPP? - kiss.caltech.edu

First detected ecosystem manifestation of the leaf‐level ‘Kok effect’ –inhibition of respiration by light?

O2em

ission

RLIGHT

RDARK

light 

Net photosynthesis

Net Respiration

Page 18: ECOSYSTEM‐scale PROBLEM: What is GPP? - kiss.caltech.edu

RLIGHT

RDARK

First detected ecosystem manifestation of the leaf‐level ‘Kok effect’ –inhibition of respiration by light?

Quercus rubra (Red Oak) Harvard Forest, 2015 

Mary Heskel et al.(Wood’s Hole MBL) 

RLIGHT

RDARK

light 

O2em

ission

CO2up

take

 (μmol m

‐2s‐1)

Net photosynthesis

Net Respiration

Photosynthetically active Radiation ( PAR )

3

Page 19: ECOSYSTEM‐scale PROBLEM: What is GPP? - kiss.caltech.edu

Slide 18

3 graph is getting out of the slideFlavia Costa,

Page 20: ECOSYSTEM‐scale PROBLEM: What is GPP? - kiss.caltech.edu

RESULT 2: Seasonal pattern

STANDARDISOTOPIC

Wehr et al, 2016, Nature

First half growing season

Page 21: ECOSYSTEM‐scale PROBLEM: What is GPP? - kiss.caltech.edu

Implications for Light-Use Efficiency (LUE) and Water Use Efficiency (WUE)

LUE important for estimating GPP from remote sensing: GPP = PAR fAPAR LUE

LUE = GPP / (APAR) (derived at eddy sites)

WUE important indicator of future C-water coupling:WUEintrinsic = (GPP / Transp) w = GPP / Gs(adjusted for water vapor deficit)

Important controversy about WUE trends (expected to increase with rising CO2):

+2.7%/yr (eddy flux network) vs +0.5%/yr (tree ring isotopes)(Keenan et al. 2013) (Frank et al. 2015)

and atmospheric 13C? (Keeling... Sing, et al. 2017)

Page 22: ECOSYSTEM‐scale PROBLEM: What is GPP? - kiss.caltech.edu

HF RESULTS: Seasonal Patterns

Absorbed Photosynthetically Active Radiation (APAR) (uE m-2 s-1)

GPP

(um

olm

-2s-

1 )

Page 23: ECOSYSTEM‐scale PROBLEM: What is GPP? - kiss.caltech.edu

HF RESULTS: Seasonal Patterns

Absorbed Photosynthetically Active Radiation (APAR) (uE m-2 s-1)

GPP

(um

olm

-2s-

1 )

Page 24: ECOSYSTEM‐scale PROBLEM: What is GPP? - kiss.caltech.edu

HF RESULTS: Seasonal Patterns

Absorbed Photosynthetically Active Radiation (APAR) (uE m-2 s-1)

GPP

(um

olm

-2s-

1 )

Standard partitioning shows declining photosynthetic capacity over the season

Page 25: ECOSYSTEM‐scale PROBLEM: What is GPP? - kiss.caltech.edu

Standard partitioning shows declining photosynthetic capacity over the season

Isotopic partitioning suggests that canopy photosynthetic capacity is more stable

HF RESULTS: Seasonal Patterns

Absorbed Photosynthetically Active Radiation (APAR) (uE m-2 s-1)

GPP

(um

olm

-2s-

1 )

Page 26: ECOSYSTEM‐scale PROBLEM: What is GPP? - kiss.caltech.edu

HF RESULT 2: Seasonal patternN

ote inverted scale!

STANDARDISOTOPIC

Wehr et al, 2016, Nature

Isotopic LUE better anti-correlated with APAR

APAR

Isotopic WUE better anti-correlated with gs

Page 27: ECOSYSTEM‐scale PROBLEM: What is GPP? - kiss.caltech.edu

Can we support or falsify GPP derivations (isotopic or conventional) with OCS or SIF?

Wehr et al, 2016, Nature

At Harvard Forest, Not yet! – broadly follows both GPPs, can’t distinguish between them

Page 28: ECOSYSTEM‐scale PROBLEM: What is GPP? - kiss.caltech.edu

Can we support or falsify GPP derivations (isotopic or conventional) with OCS or SIF?

(A GPP model integrating front end (OCS) and back-end (SIF) constraints)

NEEDED: Gs (OCS) …. Photochemistry (SIF)

GPP

Page 29: ECOSYSTEM‐scale PROBLEM: What is GPP? - kiss.caltech.edu

Canopy scale conductance via OCS flux measurements

Simple model

Boundary layer

Mesophyll(empirical

Temp function)

“biochemical conductance”(constant or T-dependent)

OCS flux[OCS]

stomatal conductance

(Wehr, Commane et al. 2017)

Page 30: ECOSYSTEM‐scale PROBLEM: What is GPP? - kiss.caltech.edu

Canopy scale conductance via OCS flux measurements

(Wehr, Commane et al. 2017)

Fit b0, b1, b2 to measured

Transpiration when evap negligible

Page 31: ECOSYSTEM‐scale PROBLEM: What is GPP? - kiss.caltech.edu

Canopy scale conductance via OCS flux measurements

(Wehr, Commane et al. 2017)

Fit b0, b1, b2 to measured

Transpiration when evap negligible

Stomatal conductance from OCS

from model

Page 32: ECOSYSTEM‐scale PROBLEM: What is GPP? - kiss.caltech.edu

Canopy scale conductance via OCS flux measurements

(Wehr, Commane et al. 2017)

Fit b0, b1, b2 to measured

Transpiration when evap negligible

OCS fluxes a very good tracer of stomatal & total canopy conductance

Page 33: ECOSYSTEM‐scale PROBLEM: What is GPP? - kiss.caltech.edu

Application: allows transpiration-Evaporation partitioning via OCS flux measurements

Page 34: ECOSYSTEM‐scale PROBLEM: What is GPP? - kiss.caltech.edu

1.  Light inhibition of daytime Respiration (Reco) in leaves has ecosystem consequences. (Apparently this is the first direct ecosystem‐level detection of the leaf‐level Kok effect)

2.  This could make a big difference:  Standard methods overestimate GPP by 25%, and overestimate daytime respiration (Reco ) by 100% at Harvard Forest.  Does this also apply at hundreds of eddy flux sites around the world?

3.  OCS a very good tracer of Gs, but Gs is not GPP.   Let’s use Gs(OCS) in a ‘better’ model incorporating tracers of 

different parts of the photosynthetic process.

Summary of What is GPP?