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Page 1: PowerPoint Presentation · Photosynthesis requires nitrogen C 55 H 72 O 5 N 4 Mg . Oxygenic photosynthesis and N fixation evolve in an anoxic world, but are so important that they

H375O132C88N6Ca1P1

Page 2: PowerPoint Presentation · Photosynthesis requires nitrogen C 55 H 72 O 5 N 4 Mg . Oxygenic photosynthesis and N fixation evolve in an anoxic world, but are so important that they

H263O110C106N16P1

http://www.smithsonianmag.com/

Page 3: PowerPoint Presentation · Photosynthesis requires nitrogen C 55 H 72 O 5 N 4 Mg . Oxygenic photosynthesis and N fixation evolve in an anoxic world, but are so important that they

H375O132C88N6Ca1P1

H263O110C106N16P1

Page 4: PowerPoint Presentation · Photosynthesis requires nitrogen C 55 H 72 O 5 N 4 Mg . Oxygenic photosynthesis and N fixation evolve in an anoxic world, but are so important that they
Page 5: PowerPoint Presentation · Photosynthesis requires nitrogen C 55 H 72 O 5 N 4 Mg . Oxygenic photosynthesis and N fixation evolve in an anoxic world, but are so important that they

N Fixation?

Page 6: PowerPoint Presentation · Photosynthesis requires nitrogen C 55 H 72 O 5 N 4 Mg . Oxygenic photosynthesis and N fixation evolve in an anoxic world, but are so important that they

Evolved ≈3.5 billion years ago

Converts N2 into NH3

Energetically expensive

(16ATP for 1 N)

Poisoned by oxygen

Biological Nitrogen Fixation

Page 7: PowerPoint Presentation · Photosynthesis requires nitrogen C 55 H 72 O 5 N 4 Mg . Oxygenic photosynthesis and N fixation evolve in an anoxic world, but are so important that they

Photosynthesis requires nitrogen

C55H72O5N4Mg

Page 8: PowerPoint Presentation · Photosynthesis requires nitrogen C 55 H 72 O 5 N 4 Mg . Oxygenic photosynthesis and N fixation evolve in an anoxic world, but are so important that they

Oxygenic photosynthesis and N fixation evolve in

an anoxic world, but are so important that they

remain basically unchanged for 3 billion years.

Page 9: PowerPoint Presentation · Photosynthesis requires nitrogen C 55 H 72 O 5 N 4 Mg . Oxygenic photosynthesis and N fixation evolve in an anoxic world, but are so important that they

Today photosynthesis and

other key processes are

limited by nitrogen

Page 10: PowerPoint Presentation · Photosynthesis requires nitrogen C 55 H 72 O 5 N 4 Mg . Oxygenic photosynthesis and N fixation evolve in an anoxic world, but are so important that they

Nitrogen Cycle Basics Transformations mediated by

microbes (auto and

heterotrophic)

Inputs via fixation + deposition

Outputs via leaching and gas

losses (denitrification) +

Mineral forms used by plants

(NO3- and NH4

+) + little DON

Losses of nitrate discriminate

against 15N, leaving it behind.

The N cycle, like life, is a redox

driven process.

Page 11: PowerPoint Presentation · Photosynthesis requires nitrogen C 55 H 72 O 5 N 4 Mg . Oxygenic photosynthesis and N fixation evolve in an anoxic world, but are so important that they

The N Cycle is Driven by Redox

Page 12: PowerPoint Presentation · Photosynthesis requires nitrogen C 55 H 72 O 5 N 4 Mg . Oxygenic photosynthesis and N fixation evolve in an anoxic world, but are so important that they

Ecosystem Mass Balance

IF Inputs > Outputs

THEN N pools grow

Losses grow

Page 13: PowerPoint Presentation · Photosynthesis requires nitrogen C 55 H 72 O 5 N 4 Mg . Oxygenic photosynthesis and N fixation evolve in an anoxic world, but are so important that they

Ecosystem Mass Balance

IF Inputs > Outputs

THEN N pools grow

Losses grow

IF Outputs > Inputs

THEN N pools shrink

Losses shrink

Page 14: PowerPoint Presentation · Photosynthesis requires nitrogen C 55 H 72 O 5 N 4 Mg . Oxygenic photosynthesis and N fixation evolve in an anoxic world, but are so important that they

Ecosystem Mass Balance

IF Inputs > Outputs

THEN N pools grow

Losses grow

IF Outputs > Inputs

THEN N pools shrink

Losses shrink

IF N availability >

demand

THEN Available N losses high

Page 15: PowerPoint Presentation · Photosynthesis requires nitrogen C 55 H 72 O 5 N 4 Mg . Oxygenic photosynthesis and N fixation evolve in an anoxic world, but are so important that they

Ecosystem Mass Balance

IF Inputs > Outputs

THEN N pools grow

Losses grow

IF Outputs > Inputs

THEN N pools shrink

Losses shrink

IF N availability >

demand

THEN Available N losses high

IF N availability <

demand

THEN Available N losses low

Page 16: PowerPoint Presentation · Photosynthesis requires nitrogen C 55 H 72 O 5 N 4 Mg . Oxygenic photosynthesis and N fixation evolve in an anoxic world, but are so important that they

Ecosystem Mass Balance

IF Inputs > Outputs

THEN N pools grow

Losses grow

IF Outputs > Inputs

THEN N pools shrink

Losses shrink

IF N availability >

demand

THEN Available N losses high

IF N availability <

demand

THEN Available N losses low

Page 17: PowerPoint Presentation · Photosynthesis requires nitrogen C 55 H 72 O 5 N 4 Mg . Oxygenic photosynthesis and N fixation evolve in an anoxic world, but are so important that they

Global patterns in the N cycle

Page 18: PowerPoint Presentation · Photosynthesis requires nitrogen C 55 H 72 O 5 N 4 Mg . Oxygenic photosynthesis and N fixation evolve in an anoxic world, but are so important that they

Fixation highest in tropics

Cleveland et al., 1999

Page 19: PowerPoint Presentation · Photosynthesis requires nitrogen C 55 H 72 O 5 N 4 Mg . Oxygenic photosynthesis and N fixation evolve in an anoxic world, but are so important that they

NUE lower in tropics

Vitousek 1982

Page 20: PowerPoint Presentation · Photosynthesis requires nitrogen C 55 H 72 O 5 N 4 Mg . Oxygenic photosynthesis and N fixation evolve in an anoxic world, but are so important that they

Brookshire et al 2012

NO3- losses higher in tropics

Page 21: PowerPoint Presentation · Photosynthesis requires nitrogen C 55 H 72 O 5 N 4 Mg . Oxygenic photosynthesis and N fixation evolve in an anoxic world, but are so important that they

Craine et al., 2015

Soil δ15N higher in tropics

MAT (°C) MAP (mm/yr)

-10 0 10 20 30 100 1,000 10,000

Page 22: PowerPoint Presentation · Photosynthesis requires nitrogen C 55 H 72 O 5 N 4 Mg . Oxygenic photosynthesis and N fixation evolve in an anoxic world, but are so important that they

Houlton and Bai 2009

Soil δ15N higher in tropics

Page 23: PowerPoint Presentation · Photosynthesis requires nitrogen C 55 H 72 O 5 N 4 Mg . Oxygenic photosynthesis and N fixation evolve in an anoxic world, but are so important that they

When, where and why does

N matter in intact

tropical ecosystems?

Page 24: PowerPoint Presentation · Photosynthesis requires nitrogen C 55 H 72 O 5 N 4 Mg . Oxygenic photosynthesis and N fixation evolve in an anoxic world, but are so important that they

“The tropics” are not one place!!!

Townsend et al 2008

Page 25: PowerPoint Presentation · Photosynthesis requires nitrogen C 55 H 72 O 5 N 4 Mg . Oxygenic photosynthesis and N fixation evolve in an anoxic world, but are so important that they

LeBauer and Tresider, 2008

Fertilization suggests N matters, but there aren’t

enough data to suggest when, where or why.

Page 26: PowerPoint Presentation · Photosynthesis requires nitrogen C 55 H 72 O 5 N 4 Mg . Oxygenic photosynthesis and N fixation evolve in an anoxic world, but are so important that they

Inputs: Fixation, Deposition

Outputs: Gas losses, leaching

Internal cycling: mineralization,

nitrification, immobilization,

DNRA, FEAMMOX…

Controls of tropical N availability

Page 27: PowerPoint Presentation · Photosynthesis requires nitrogen C 55 H 72 O 5 N 4 Mg . Oxygenic photosynthesis and N fixation evolve in an anoxic world, but are so important that they

Inputs: Fixation, Deposition

Outputs: Gas losses, leaching

Internal cycling: mineralization,

immobilization…

Controls of tropical N availability

Page 28: PowerPoint Presentation · Photosynthesis requires nitrogen C 55 H 72 O 5 N 4 Mg . Oxygenic photosynthesis and N fixation evolve in an anoxic world, but are so important that they

Hedin 2009

More fixers in the tropics

Page 29: PowerPoint Presentation · Photosynthesis requires nitrogen C 55 H 72 O 5 N 4 Mg . Oxygenic photosynthesis and N fixation evolve in an anoxic world, but are so important that they

Sullivan et al, 2014

But are they fixing?

Page 30: PowerPoint Presentation · Photosynthesis requires nitrogen C 55 H 72 O 5 N 4 Mg . Oxygenic photosynthesis and N fixation evolve in an anoxic world, but are so important that they

Sullivan et al, 2014

But are they fixing? Predicted from legume

abundance

Predicted from nodule

sampling

Page 31: PowerPoint Presentation · Photosynthesis requires nitrogen C 55 H 72 O 5 N 4 Mg . Oxygenic photosynthesis and N fixation evolve in an anoxic world, but are so important that they

0

2

4

6

8

10

11 24 40 Mature (>80 years)

Nitrogen Fixation In Bahia, Brazil (kg N/ha/yr)

But are they fixing?

Winbourne unpubl.

Forest Age (years)

Page 32: PowerPoint Presentation · Photosynthesis requires nitrogen C 55 H 72 O 5 N 4 Mg . Oxygenic photosynthesis and N fixation evolve in an anoxic world, but are so important that they

y = 21.061x + 1097.2 R² = 0.9622

0

20000

40000

60000

80000

100000

120000

140000

0 1000 2000 3000 4000 5000 6000

ngN

fix

ed

Nodule Biomass in a Plot (mg)

Counting legumes doesn’t work. Counting nodules does

Page 33: PowerPoint Presentation · Photosynthesis requires nitrogen C 55 H 72 O 5 N 4 Mg . Oxygenic photosynthesis and N fixation evolve in an anoxic world, but are so important that they

Sullivan et al, 2014

Our understanding of N inputs is poor

Page 34: PowerPoint Presentation · Photosynthesis requires nitrogen C 55 H 72 O 5 N 4 Mg . Oxygenic photosynthesis and N fixation evolve in an anoxic world, but are so important that they

Inputs: Fixation, Deposition

Outputs: Gas losses, leaching

Internal cycling: mineralization,

immobilization…

Controls of tropical N availability

Page 35: PowerPoint Presentation · Photosynthesis requires nitrogen C 55 H 72 O 5 N 4 Mg . Oxygenic photosynthesis and N fixation evolve in an anoxic world, but are so important that they

Outputs: Gas losses Highly variable in

space and time,

hard to measure

Highly variable in

space and time,

“easy”to measure

Impossible to

measure in the

field

Page 36: PowerPoint Presentation · Photosynthesis requires nitrogen C 55 H 72 O 5 N 4 Mg . Oxygenic photosynthesis and N fixation evolve in an anoxic world, but are so important that they

Zhuang et al. 2012

Tropical N gas losses: high but

poorly constrained, based on N2O

Page 37: PowerPoint Presentation · Photosynthesis requires nitrogen C 55 H 72 O 5 N 4 Mg . Oxygenic photosynthesis and N fixation evolve in an anoxic world, but are so important that they

N2 losses: Theory

Pilegaard 2013, from Davidson 2000

Page 38: PowerPoint Presentation · Photosynthesis requires nitrogen C 55 H 72 O 5 N 4 Mg . Oxygenic photosynthesis and N fixation evolve in an anoxic world, but are so important that they

“Direct” measurement of N2 emissions

N2

N2O

Slide from M. Almaraz

Page 39: PowerPoint Presentation · Photosynthesis requires nitrogen C 55 H 72 O 5 N 4 Mg . Oxygenic photosynthesis and N fixation evolve in an anoxic world, but are so important that they

N2 losses: Data From Puerto Rico

Almaraz unpubl.

Page 40: PowerPoint Presentation · Photosynthesis requires nitrogen C 55 H 72 O 5 N 4 Mg . Oxygenic photosynthesis and N fixation evolve in an anoxic world, but are so important that they

N2 losses ≠ N2O losses

Almaraz unpubl.

Page 41: PowerPoint Presentation · Photosynthesis requires nitrogen C 55 H 72 O 5 N 4 Mg . Oxygenic photosynthesis and N fixation evolve in an anoxic world, but are so important that they

Our understanding of N outputs is poor

Almaraz unpubl.

Page 42: PowerPoint Presentation · Photosynthesis requires nitrogen C 55 H 72 O 5 N 4 Mg . Oxygenic photosynthesis and N fixation evolve in an anoxic world, but are so important that they

Inputs: Fixation, Deposition

Outputs: Gas losses, leaching

Internal cycling: mineralization,

nitrification, immobilization…

Controls of tropical N availability

Page 43: PowerPoint Presentation · Photosynthesis requires nitrogen C 55 H 72 O 5 N 4 Mg . Oxygenic photosynthesis and N fixation evolve in an anoxic world, but are so important that they

“The tropics” are not one place!!!

Townsend et al 2008

Page 44: PowerPoint Presentation · Photosynthesis requires nitrogen C 55 H 72 O 5 N 4 Mg . Oxygenic photosynthesis and N fixation evolve in an anoxic world, but are so important that they

N availability

Time Climate Parent

material

Topograph

y Organisms

Heterogeneity is challenging

Page 45: PowerPoint Presentation · Photosynthesis requires nitrogen C 55 H 72 O 5 N 4 Mg . Oxygenic photosynthesis and N fixation evolve in an anoxic world, but are so important that they

N availability

Time Climate Parent

material Topography Organisms

Heterogeneity is challenging

Page 46: PowerPoint Presentation · Photosynthesis requires nitrogen C 55 H 72 O 5 N 4 Mg . Oxygenic photosynthesis and N fixation evolve in an anoxic world, but are so important that they

How do topography, rainfall, and

foliar N influence N availability?

Page 47: PowerPoint Presentation · Photosynthesis requires nitrogen C 55 H 72 O 5 N 4 Mg . Oxygenic photosynthesis and N fixation evolve in an anoxic world, but are so important that they

The Osa Peninsula, Costa Rica

Page 48: PowerPoint Presentation · Photosynthesis requires nitrogen C 55 H 72 O 5 N 4 Mg . Oxygenic photosynthesis and N fixation evolve in an anoxic world, but are so important that they

Hyperspectral-derived canopy N

Lidar derived topography

Digital

Elevation

Models

Foliar

Nitrogen

Maps

Page 49: PowerPoint Presentation · Photosynthesis requires nitrogen C 55 H 72 O 5 N 4 Mg . Oxygenic photosynthesis and N fixation evolve in an anoxic world, but are so important that they

Topography

Climate

Page 50: PowerPoint Presentation · Photosynthesis requires nitrogen C 55 H 72 O 5 N 4 Mg . Oxygenic photosynthesis and N fixation evolve in an anoxic world, but are so important that they

Variation in topography, climate.

Drake Bay

Piro North

Piro South

Page 51: PowerPoint Presentation · Photosynthesis requires nitrogen C 55 H 72 O 5 N 4 Mg . Oxygenic photosynthesis and N fixation evolve in an anoxic world, but are so important that they

Piro South Piro North Drake Bay

Topography

Broad, flat ridge Narrow ridge Narrow ridge

Page 52: PowerPoint Presentation · Photosynthesis requires nitrogen C 55 H 72 O 5 N 4 Mg . Oxygenic photosynthesis and N fixation evolve in an anoxic world, but are so important that they

Piro South Piro North Drake Bay

Broad, flat ridge Narrow ridge Narrow ridge

Topography

Page 53: PowerPoint Presentation · Photosynthesis requires nitrogen C 55 H 72 O 5 N 4 Mg . Oxygenic photosynthesis and N fixation evolve in an anoxic world, but are so important that they

Piro South Piro North Drake Bay

Climate

Page 54: PowerPoint Presentation · Photosynthesis requires nitrogen C 55 H 72 O 5 N 4 Mg . Oxygenic photosynthesis and N fixation evolve in an anoxic world, but are so important that they

Drake Bay

MAP ~3000 mm MAP ~6000 mm

Piro South Piro North

Climate

Page 55: PowerPoint Presentation · Photosynthesis requires nitrogen C 55 H 72 O 5 N 4 Mg . Oxygenic photosynthesis and N fixation evolve in an anoxic world, but are so important that they

Sample collection & analyses

Piro South Piro North Drake Bay

Top Shoulder

Middle

Bottom

• 3 regions x 4 catenas x 4 transects x 2 seasons

Page 56: PowerPoint Presentation · Photosynthesis requires nitrogen C 55 H 72 O 5 N 4 Mg . Oxygenic photosynthesis and N fixation evolve in an anoxic world, but are so important that they

N metrics measured Piro South Piro North Drake Bay

NO3--N and NH4

+-N

Net nitrification

Net N mineralization

δ15N

Instantaneous

5 days

100s – 1000s of years

Page 57: PowerPoint Presentation · Photosynthesis requires nitrogen C 55 H 72 O 5 N 4 Mg . Oxygenic photosynthesis and N fixation evolve in an anoxic world, but are so important that they

Piro South Piro North Drake Bay

Topography (δ15N o/oo)

Top Shoulder Middle Bottom

01

23

45

67

xcoord

δ1

5N

(o

/oo)

a b b

b δ15N was

elevated on

broad, flat

ridges.

Top Shoulder Middle Bottom

01

23

45

67

xcoord

δ1

5N

(o

/oo) b

b b b

Osborne et al, in revision

Page 58: PowerPoint Presentation · Photosynthesis requires nitrogen C 55 H 72 O 5 N 4 Mg . Oxygenic photosynthesis and N fixation evolve in an anoxic world, but are so important that they

Top Shoulder Middle Bottom

01

23

45

67

xcoord

δ1

5N

(o

/oo)

Top Shoulder Middle Bottom

01

23

45

67

xcoord

δ1

5N

(o

/oo)

Climate (δ15N o/oo) Drake Bay

MAP ~3000 mm MAP ~6000 mm

Piro South Piro North

δ15N was

elevated

under drier

conditions.

mean = 4.34* mean = 3.46

prob > F 0.0363*

Osborne et al, in revision

Page 59: PowerPoint Presentation · Photosynthesis requires nitrogen C 55 H 72 O 5 N 4 Mg . Oxygenic photosynthesis and N fixation evolve in an anoxic world, but are so important that they

Piro South Piro North Drake Bay

Topography and climate summary

NO3--N and NH4

+-N

Climate

effect Topography

effect

✗ ✗

δ15N

NO3--N and NH4

+-N

Osborne et al, in revision

Page 60: PowerPoint Presentation · Photosynthesis requires nitrogen C 55 H 72 O 5 N 4 Mg . Oxygenic photosynthesis and N fixation evolve in an anoxic world, but are so important that they

Organisms

Climate

Topography

Page 61: PowerPoint Presentation · Photosynthesis requires nitrogen C 55 H 72 O 5 N 4 Mg . Oxygenic photosynthesis and N fixation evolve in an anoxic world, but are so important that they

Organisms

Low N

High N

Osborne et al, in revision

Page 62: PowerPoint Presentation · Photosynthesis requires nitrogen C 55 H 72 O 5 N 4 Mg . Oxygenic photosynthesis and N fixation evolve in an anoxic world, but are so important that they

Osborne et al, in revision

Organisms

Page 63: PowerPoint Presentation · Photosynthesis requires nitrogen C 55 H 72 O 5 N 4 Mg . Oxygenic photosynthesis and N fixation evolve in an anoxic world, but are so important that they

scale here

Osborne et al, in revision

Organisms

Page 64: PowerPoint Presentation · Photosynthesis requires nitrogen C 55 H 72 O 5 N 4 Mg . Oxygenic photosynthesis and N fixation evolve in an anoxic world, but are so important that they

High

Foliar N

Low

Foliar N

Hot Cold

01

23

45

67

NO3-N (mg kg-1)

Transect

NO

3-N

(m

g k

g-1

)

Hot Cold

01

23

45

67

NH4-N (mg kg-1)

Transect

NH

4-N

(m

g k

g-1

)

Hot Cold

01

23

45

67

Net Nitrification (mg kg-1 d-1)

Transect

Net N

itrifica

tio

n (

mg

kg

-1 d

-1)

Hot Cold

01

23

45

67

Net Mineralization mg kg-1 d-1

Transect

Net

Min

era

lizatio

n (

mg

kg

-1 d

-1)

Osborne et al, in revision

Organisms – link to soil N

Page 65: PowerPoint Presentation · Photosynthesis requires nitrogen C 55 H 72 O 5 N 4 Mg . Oxygenic photosynthesis and N fixation evolve in an anoxic world, but are so important that they

Airborne mapping may help deal

with heterogeneity at large scales

Osborne et al, in revision

Page 66: PowerPoint Presentation · Photosynthesis requires nitrogen C 55 H 72 O 5 N 4 Mg . Oxygenic photosynthesis and N fixation evolve in an anoxic world, but are so important that they

The role of trees in driving the N cycling

may be more important than we know.

Asner et al., 2014

Page 67: PowerPoint Presentation · Photosynthesis requires nitrogen C 55 H 72 O 5 N 4 Mg . Oxygenic photosynthesis and N fixation evolve in an anoxic world, but are so important that they

“The tropics” are not one place!!!

Osborne et al, in revision

Page 68: PowerPoint Presentation · Photosynthesis requires nitrogen C 55 H 72 O 5 N 4 Mg . Oxygenic photosynthesis and N fixation evolve in an anoxic world, but are so important that they

Questions?