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Marcos S. Buckeridge, 2008 Marcos S. Buckeridge Departamento de Botânica Instituto de Biociências Universidade de São Paulo Forests or ethanol: how plant physiology can be strategically used to reach the best choice? Instituto Nacional de Pesquisas Espaciais - INPE

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Page 1: Forests or ethanol: how plant physiology can be strategically … · 2008-04-18 · Forests or ethanol: how plant physiology can be strategically used to reach ... L O G I C A L S

Marcos S. Buckeridge, 2008

Marcos S. Buckeridge

Departamento de BotânicaInstituto de Biociências

Universidade de São Paulo

Forests or ethanol: how plant physiology can be

strategically used to reach the best choice?

Instituto Nacional de Pesquisas Espaciais - INPE

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Marcos S. Buckeridge, 2008

Air in samples of the AntarticContinent

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Marcos S. Buckeridge, 2008Marcos S. Buckeridge, 2008

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Marcos S. Buckeridge, 2008

Ruddiman, W.F. (2005) Scentific American n.35

8,000 years ago, carbon dioxide concentration

suddenly change from a tendency to

decrease to a tendency to increase

Natural

tendency

Observed tendency

Years ago

Car

bon

diox

ide

(par

ts p

er

mill

ion)

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Marcos S. Buckeridge, 2008

Ruddiman, W.F. (2005) Scentific American n.35

Gaciationlimiar

Real tendency

Natural initiation of the next glacial

period

Natural tendency of temperature changes

After de end of fossil fuels

Today

Lowest temperatures during glacial

periods

Higher temperatures

during interglacial

periods

Temperatures above the

possible level during glacial

eras

Primitive agricultureRapid industrializationFuture activities

Past and possible future effects of human activities on the average temperatures of

the planetMarcos S. Buckeridge, 2008

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Marcos S. Buckeridge, 2008Marcos S. Buckeridge, 2008

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Marcos S. Buckeridge, 2008

Global carbon stocks in vegetation and top 1 m of soils (based on WBGU, 1998).

Carbon Stocks (Gt C)Biome

Area(106 km2) Vegetation Soils Total

Tropical forests 17.6 212 216 428Temperate forests 10.4 59 100 159Boreal forests 13.7 88 471 559Tropical savannas 22.5 66 264 330Temperate grasslands 12.5 9 295 304Deserts and semideserts 45.5 8 191 199Tundra 9.5 6 121 127Wetlands 3.5 15 225 240Croplands 16.0 3 128 131

Total 151.2 466 2011 2477

Marcos S. Buckeridge, 2008

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Marcos S. Buckeridge, 2008

Why primary production is higher in the tropics?

Because climatic conditions afford coutinuous growth and C storage

What this has to do with plant functioning?

Higher productivity is directly related with how plants manage their nutrients, such as Carbon, Nitrogen, Phosphate etc

Can knowledge about plant functioning help to MITIGATE THE EFFECTS OF GLOBAL WARMING

?Probably, lets see some data......

Marcos S. Buckeridge, 2008

Page 9: Forests or ethanol: how plant physiology can be strategically … · 2008-04-18 · Forests or ethanol: how plant physiology can be strategically used to reach ... L O G I C A L S

Marcos S. Buckeridge, 2008

Light, Water & Nutrients CO2

PHOTOSYNTHESIS

SUCROSE STARCH

CELLULOSE

GROWTHMitigation of C emissions

Forests & Ecosystem Services

Crops that produce BIOFUELS

Marcos S. Buckeridge, 2008

Page 10: Forests or ethanol: how plant physiology can be strategically … · 2008-04-18 · Forests or ethanol: how plant physiology can be strategically used to reach ... L O G I C A L S

Marcos S. Buckeridge, 2008

How can we use plant science to help choosing

the best way to go?

Marcos S. Buckeridge, 2008

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Marcos S. Buckeridge, 2008

THE CASE OF TREESregenerating forests and reestablishing

ecosystem services

Marcos S. Buckeridge, 2008

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Marcos S. Buckeridge, 2008

Hymenaea in the future ?Hymenaea in the future ?

Experiments in open top chambers

2001 2001 -- 360 360 ppmppm COCO222050 2050 -- 720 720 ppmppm COCO22Marcos S. Buckeridge, 2008

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Marcos S. Buckeridge, 2008

Plantas jovens de jatobá crescendo por 90 dias com e sem os cotilédones

Marcos S. Buckeridge, 2008

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Marcos S. Buckeridge, 2008

Santos, Mercier, Purgato & Buckeridge, Plant Physiology, 2004, vol 135 p.

IAA XGMs

Sc

St

Sc

Sc

IAA

Hydrolases

+H

Grt

NPA

7

4 32

5 6

8

1+

-

+

Marcos S. Buckeridge, 2008

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Marcos S. Buckeridge, 2008

Xyloglucan

XGOs

Xyl

Sucrose

Degalactosylated XGOs

XTH

hcbetagal

beta glucosidase

Glc

alpha xylosidase

Gal

P-sugars ?

sucrose synthase

Auxin

DNA

mRNA

auxin-conjugate

LIGHT

NPA treatment

Shoot excision

Sucrose

GROWTH

Starch

P-sugars

sucrose synthase

invertase

Pentose P pathway ?

Starch

coty

ledo

n

hypo

coty

l

leaf

phy, cry ?

?

?

??

invertase

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Marcos S. Buckeridge, 2008

CotyledonXG catabolism

Plantlet

Photosynthesis

sucrose

GROWTH

sucrose

IAA

CO2

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Marcos S. Buckeridge, 2008

CotyledonXG catabolism

Plantlet

sucrose

GROWTH

sucrose

IAA

CO2

Photosynthesis

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Marcos S. Buckeridge, 2008

6 8 10 12 14 16 18-2

-1

0

1

2

3

4

5

6

7Net Photosynthesis - Training (Level 1)

Phot

osyn

thes

is (u

mol

CO

2m-2s-1

)

Time (Hours)

Measured Value estimated Value

Neural network to forecast photosynthesis in Hymenaea

Barriga et al. Submitted to Ecological ModellingMarcos S. Buckeridge, 2008

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Marcos S. Buckeridge, 2008

Plants in the future ?Plants in the future ?

Experiments in open top chambers

2001 2001 -- 360 360 ppmppm COCO222007 2007 -- 384 384 ppmppm CO2CO2

2050/2080 2050/2080 -- 720 720 ppmppm COCO22Marcos S. Buckeridge, 2008

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Marcos S. Buckeridge, 2008

OPEN TOP CHAMBER

CO2

4 m

1,5 m

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Marcos S. Buckeridge, 2008Aidar et al. 2002 V2 (2) (www.biotaneotropica.org.br)

Figure 9 – Responses of the light saturated net photosynthesis (Amax) for eophylls fromHymenaea courbaril seedlings with cotyledons to atmospheric CO2 concentrations. Values for 360 and 720 pmm CO2 concentrations were measured in our open top chambers; values for CO2 concentration of 120 and 1200 ppm were obtained through the A x Ci curves simulated by IRGA (Li-Cor 6400).

y = 6.2979Ln(x) - 26.932r = 0.989

0

2

4

6

8

10

12

14

16

18

20

0 360 720 1080 1440CO2 atmospheric concentration (ppm)

Amax

(µm

ol C

O2

m-2 s

-1)

Marcos S. Buckeridge, 2008

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Marcos S. Buckeridge, 2008

Stomata in Hymenaea are decreasing

5

10

15

20

25

30

1900 1950 2000 2050Time (years)

Stom

atal

Inde

x

Marcos S. Buckeridge, 2008

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Marcos S. Buckeridge, 2008Costa, Aidar, Viveiros Martinez and Buckeridge, unpublished

1919 – 280ppm 2002 – 360ppm 2075 – 720ppm

13

14

15

16

17

18

360 720 360 720

Cotyledons Without cotyledons

Stom

ato

inde

x

EophyllMetaphyll

10

12

14

16

18

20

22

24

26

1900 1950 2000 2050 2100

time (years)st

omat

o in

dex

? 1929 = 20

Marcos S. Buckeridge, 2008

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Marcos S. Buckeridge, 2008

Plant organsPlant organs andand theirtheircarboncarbon metabolismmetabolism

Marcos S. Buckeridge, 2008

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Marcos S. Buckeridge, 2008

DEVELOPMENTAL PARAMETERS

-20

-10

0

10

20

30

40

50

60

Ste

m le

ngth

(cm

)

Siz

e of

eoph

yls

Siz

e of

met

aphy

ls

Tota

l lea

f are

a(c

m2)

Rel

ative

leaf

area

Roo

t:Sho

otra

tio

Bio

mas

s (g

)

% o

f cha

nge

Storage

No Storage

*

*

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Marcos S. Buckeridge, 2008

0

10

20

30

40

50

60

70

80

90

Sucrose Starch Cellulose

% o

f inc

reas

e in

ele

vate

d C

O2

Surplus of “carbon” in leaves of Hymenaea (jatobá) in elevated CO2

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Marcos S. Buckeridge, 2008

Foto

s M

arce

lo M

acha

do &

Mar

cos

Buc

kerid

ge–

IB U

SP

200

7

Chloroplast

Starch

Starch in pallisadecells of jatobagrowing underelevated CO2

Plant obesity?

Vacuolecontainingsucrose

Elevated CO2

Current CO2

COLUNA NEOTRÓPICAS http://www.revistapesquisa.fapesp.br

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Marcos S. Buckeridge, 2008

CO2

STARCHPhotosynthesis

Respiration

Stomata

Glucose SUCROSE

CELL WALLHK

O2 SECONDARY METABOLITES

Respiratory chain

LIPIDS

hic CelluloseSynthesis

Housekeeping &

growth

Defence & metabolic

control

LONG TERM STORAGE CARBON

Amazon = 1 ton/ hectare/year

Savannah = 0.1 ton/ hectare/year

C sequestration in different Biomes

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Marcos S. Buckeridge, 2008

RUBISCOSTOMATA MITOCHONDRIA NUCLEUS

Water and carbohydrate status

in leaves

Carbonic anhydraseand RUBISCO

ABI4

Respiration

Photosynthesis

Con

duct

ance

-

+

-

Trasnpiration-

Stomata

l den

sity

e- transport+

C assimilation+

Starch, leaf area, biomass, root growth, stress tolerance, defence and fertility

+

N Assimilation and micorrizal association

-

Photosynthetic protein-

Cel

lLe

afP

lant

Page 30: Forests or ethanol: how plant physiology can be strategically … · 2008-04-18 · Forests or ethanol: how plant physiology can be strategically used to reach ... L O G I C A L S

Marcos S. Buckeridge, 2008

Other native tree species of the Atlantic Forest

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Marcos S. Buckeridge, 2008

Gramíneas, asteráceasSesbania, embaúba, solanaceas

t = ano zerot = 10 anos

Sucessão ecológica – o processo que forma as florestas

t = 30 anost = +40 anos

Guapuruvú, pau-jacaré, ipês, pau-brasilJatobá, jacarandá, copaíba

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Marcos S. Buckeridge, 2008

Gradient of physiological limitation in ecological succession

Water

Light

10 µmoles.m-2.s −2

2000 µmoles.m-2.s −2

High lightintensity andlowavailability of water

Optimalmicroclimaticconditions, ideal for growth anddevelopment

Low lightintensityand highavailabilityof water

Pioneer → Secondary → Late Secondary/Climax

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Marcos S. Buckeridge, 2008

Sesbania virgata

Schyzolobium parahyba

Piptadenia gonoacantha

Dalbergia nigra

Hymenaea courbarilECOLOGICAL SUCCESSION

25-30 anos

25-30 anos

50-100 anos

>100 anos

5 a 10 anos

19 Kg per Ton(70 Kg of CO2 per ton)

137 Kg per Ton(487 Kg of CO2 per ton)

23 Kg per Ton(84 Kg of CO2 per ton)

14 Kg per Ton(51 Kg of CO2 per ton)

In prep

Marcos S. Buckeridge, 2008

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Marcos S. Buckeridge, 2008

Performance de 5 spp de Leguminosae de diferentes estágios de sucessão em alto CO2. O percentual de diferença de biomassa foi dividido pelo percentual de diferença em assimilação fotossintética e multiplicado

pela eficiência do uso da água-valores médios

0,00

20,00

40,00

60,00

80,00

100,00

120,00

Sesb Schizo Pipta Dalber Hyme

Espécies

Perf

orm

ance

fisi

ológ

ica

em a

lto C

O2

(% m

s/%

A.E

UA

)

Species

Phys

iolo

gica

l per

form

ance

in h

igh

CO

2

Physiological performances of 5 tropical legume species in high CO2

Marcos S. Buckeridge, 2008

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Marcos S. Buckeridge, 2008

0

5

10

15

20

25

30

35

40

0 10 20 30 40 50

Tempo (anos)

Pote

ncia

l de

sequ

estro

de

C

pioneiras Secundárias Iniciais

Secundárias Tardias

Sequ

estro

de C

com

o

proc

esso

de su

cess

ão

Marcos S. Buckeridge, 2008

Page 36: Forests or ethanol: how plant physiology can be strategically … · 2008-04-18 · Forests or ethanol: how plant physiology can be strategically used to reach ... L O G I C A L S

Marcos S. Buckeridge, 2008

Jatobá e açaí crescendo em alto CO2 e alta temperatura (+3oC)

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Marcos S. Buckeridge, 2008

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Marcos S. Buckeridge, 2008

NEXT – The combined effects of CO2and temperature

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Marcos S. Buckeridge, 2008

SUGAR CANE

Physiological behaviour under elevated CO2

Marcos S. Buckeridge, 2008

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Marcos S. Buckeridge, 2008Marcos S. Buckeridge, 2008

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Marcos S. Buckeridge, 2008

Comparisons between C3 and C4 plants in elevated CO2

Meta analytical data

Wand et al. (1999) Poorter & Navas (2003)

C4 Plants

contribute with ca18% for world productivity24% x 29%

33% x 25%There are more

results with C4 fromtemperate climate

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Marcos S. Buckeridge, 2008

Sugar cane in the open top chambers

360

360

720

720

Finantial support by Centro de Tecnologia Canavieira - Piracicaba

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Marcos S. Buckeridge, 2008

Experimental design

CO2

4 m

1,5 m

Sugarcane variety SP80-3280

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Marcos S. Buckeridge, 2008

Photosynthesis

-10

0

10

20

30

40

50

60

70

6 10 13 18 21 26 31 50

Weeks of CO2

% c

hang

e in

ele

vate

d C

O 2

B

0

5

10

15

20

25

30

35

40

0 5 10 15 20 25 30 35 40 45 50

Weeks of CO2

CO 2 a

ssim

ilatio

n (µ

mol

CO 2 m

-2 s

-1)

Ambient

Elevated

***

******

****

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Marcos S. Buckeridge, 2008

ElevatedAmbient

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Marcos S. Buckeridge, 2008

Plant height under elevated CO2

0

0,5

1

1,5

2

2,5

3

3,5

0 5 10 15 20 25 30 35 40 45 50

Weeks of CO2

Hei

ght (

m)

Ambient

Elevated

**

***

******

**

***

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Marcos S. Buckeridge, 2008

0

100

200

300

400

500

600

700

13 22 26 31 50

Leaf

bio

mas

s (g

)

*

*

B

0

200

400

600

800

1000

1200

1400

1600

1800

13 22 26 31 50

Weeks of CO2

Cul

m b

iom

ass

(g)

* **

**C

0

50

100

150

200

250

300

13 22 26 31 50

Weeks of CO2

Roo

t bio

mas

s (g

)

D

0

500

1000

1500

2000

2500

3000

13 22 26 31 50

Tota

l bio

mas

s (g

)

**

**

***

A

+ 50%

Marcos S. Buckeridge, 2008

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Marcos S. Buckeridge, 2008

Roots of sugar cane

ASPECT AFTER 3 MONTHS -Note that the comparison is between 3 plants from 360ppm agains two plants from720ppm of CO2

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Marcos S. Buckeridge, 2008

ElevatedAmbientProductivity

Marcos S. Buckeridge, 2008

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Marcos S. Buckeridge, 2008

Sugar and Fibre in elevated CO2

0

1

2

3

4

5

6

7

8

Fiber Sucrose

% F

resh

wei

ght

Ambient Elevated

Marcos S. Buckeridge, 2008

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Marcos S. Buckeridge, 2008

Microarrays

0

5

10

15

20

25

30

35

40

6 10 13 18 21 26 31 50

Weeks after CO2

A (µ

mol

CO 2

m-2

s-1

)Ambiente

Elevado ******

***

*

(a)AmbientElevated

Microarray analyses

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Marcos S. Buckeridge, 2008

0 1 2 3 4 5

Carbon metabolism

Cell cycle

Development

Lipid metabolism

No match

Acid nucleic metabolism

Photosynthesis

Protein metabolism

Receptor

Secondary metabolism

Stress response

Transcription factors

Transport

Func

tiona

l cat

egor

ies

Number of genes

Repressed

Induced

Pattern of gene expression in

sugarcaneunder normal and elevated

CO2

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Marcos S. Buckeridge, 2008

Microarray analysis of the CO2 experiments

-1,606translational initiation factor eIF-4AProtein metabolism-1,232

putative glucose-6-phosphate dehydrogenaseCarbohydrate metabolism

-2,189beta-glucosidase isozyme 2 precursorCarbohydrate metabolism3,59AE9 stearoyl-ACP desaturase

Lipid, fatty-acid and isoprenoid metabolism

1,735ASR-likeStress response1,508Chlorophyll A-B binding proteinPhotosynthesis2,582

xyloglucan endo-transglycosylase/hydrolaseCell wall metabolism

1,583photosystem I reaction centre subunit n,

chloroplast precursorPhotosynthesis

1,26Ferredoxin I; chloroplast precursorPhotosynthesis1,315photosystem II protein K; psbKPhotosynthesis1,194light-induced proteinDevelopment

Ratio (elevated/am

bient)Gene descriptionCategories

3 months

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Marcos S. Buckeridge, 2008-1,632cyclin H-1Cell cycle

-1,541kelch repeat-containing F-box family proteinCell cycle

-1,494Lateral organ boudaries proteinDevelopment

-1,352auxin response factor 2Transcription

-1,367ThaumatinPathogenicity

-1,342C2 domain-containing protein-likeProtein metabolism

-2,768ferritinStress response

-1,615cell wall invertaseCarbohydrate metabolism

-1,354caffeoyl-CoA 3-O-methyltransferase 1Secondary metabolism

-1,398unknow

-1,42dehydrinStress response

0,17chromodomain-helicase-DNA-binding proteinNucleic acid metabolism

1,541pre-mRNA splicing factorTranscription

1,909putative auxin-independent growth promoterDevelopment

1,397putative nucleostemin (GTPase of unknown function)

1,229Aldo/keto reductase; Sigma-54 factorProtein metabolism

1,395cathepsin B-like cysteine proteaseProtein metabolism

1,504putative glutamate-tRNA ligaseProtein metabolism

1,271unknow

1,706serine/threonine-protein kinase NAKReceptors

1,252Sugar transporterTransporters

2,299dormancy-associated proteinDevelopment

1,349cathepsin B-like cysteine proteaseProtein metabolism

1,37Alpha-L-arabinofuranosidaseCell wall metabolism

1,245phosphoenolpyruvate carboxylaseCarbohydr. metabolism/Photosynthesis

1,454large ribosomal protein 2Protein metabolism

5 months

Marcos S. Buckeridge, 2008

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Marcos S. Buckeridge, 2008

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-505

1015202530354045

0 500 1000 1500 2000 2500

PPDF (µmol m-2 s-1)

A (µ

mol

m-2

s-1

)

-5

0

5

10

15

20

25

0 500 1000 1500 2000 2500

PPDF (µmol m-2 s-1)A

(µm

ol m

-2 s

-1)

-5

0

5

10

15

20

25

30

0 200 400 600 800 1000 1200

Ci (µmol mol-1)

A (µ

mol

m-2

s-1

)

22 weeks

50 weeks

A x Ci

Photosyntheticbehaviour of sugarcane in elevated CO2

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Marcos S. Buckeridge, 2008

0,800 ± 0,004 *0,786 ± 0,00526

0,802 ± 0,003 *0,786 ± 0,00818

0,778 ± 0,0090,780 ± 0,00210

ElevatedAmbient

Fv/ FmWeeks in elevated [CO2]

Table 2. Chlorophyl fluorescence of leaves of sugarcane plants growing under ambient (370ppm and elevated (720ppm) of CO2. * P<0.1

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FSII

CCL

Transportede elétrons NADPH

Gradiente de pH no tilacóide

ATP

Ciclode

Calvin

4H2O

4H + O2

Fluorescência

CO2

Carboidratos

Calor PEPc

Crescimento

Figura 1Esquema mostrando os principais passos do processo de

fotossíntese e suas interrelações. (CCL= centro de captação de luz, fsII=fotossistema II, atp=adenosina trifosfato, nadph=nicotinamida

adenosina difosfato reduzida. Note que na captação de gáscarbônico há duas vias, a C3 e a via C4. Todos as vias levam ao

mesmo lugar, que é produzir carboidratos que serão utilizados parao crescimento da planta

Ácido com 4 carbonos

Via C3

Via C4

Celula do mesofilo

Celula daBainha Vascular

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Marcos S. Buckeridge, 2008

What all this means in respect to BIOMASS?

2) Only in the forest of South America, we have 70 billion tons of C

Brazil - 2005-2006•Total production – 400 millions of tons

•But cane biomass has approx. 90% of water!•This means that 40 million is the dry mass

•Considering carbon as 40% of the dry massApprox. 16 million of sugarcane biomass is carbon

THUS, our sugarcane represents c.a. 0.01% of the C in South American forests

3) The C emission from burning the Amazon amounts to 3 billion tons, which means that all sugarcane production

represents 0.0004% of what is burnt

Then, what should we do?

But........1) Half of the sugarcane is used for production of alcohol, thus 8 million

tons of C equivalent

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Marcos S. Buckeridge, 2008

Cane X Forest a Brazilian dilema for the XXI Century

0

200

400

600

800

1000

1200

1400

1600

1800

2000

0 10 20 30 40Time (years)

Suga

r can

e (m

illio

ns o

f ton

s)

0

50

100

150

200

250

300

350

400

450

Fore

sts

(Gto

ns)

C sequestration potentital in crops and forest

In 100 years, the sugarcane production can cover only 0.1% ofa 10% burning of the Amazon!

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Marcos S. Buckeridge, 2008

THE MIDWAY“O caminho do meio”

Cane aloneOnly biofuel production

Cane with forest corridorsMore ethanol production

More C sequestration, plus ecosystem services

1) Increase in cane productivity2) Regeneration of forests and cerrado

BUCKERIDGE, M.S. (2007) Seqüestro de carbono, cana-de-açúcar e o efeito Cinderela. Comciência - LabJorhttp://www.comciencia.br/comciencia/?section=8&edicao=23&id=258

Environmental Friendly Ethanol

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Marcos S. Buckeridge, 2008

Is the Midway

POSSIBLE?

FEASIBLE?

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Marcos S. Buckeridge, 2008

Rodovia dos Bandeirantes – São PauloAgosto 2007

Marcos S. Buckeridge, 2008

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A tree of Schizolobium parayba

growing in a regenerating forest just beside a

sugarcane plantation

Rodovia dos BandeirantesSão Paulo

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Marcos S. Buckeridge, 2008

Inst. De BotânicaMarcos Aidar, Marilia Gaspar

Marco TinéEmerson Silva

&Sonia Dietrich

USP-São PauloGlaucia Souza

Alessandro WacloviskyCarlos Martinez

USP-RP

Purdue UniversityNick Carpita

Mureen MacCann

Post docs, PhD and MSc students and

Technicians

Amanda P. SouzaAna Maria da SilvaPaula Feilx CostaMarcelo MachadoMauro Marabesi

COLLABORATORS

Climatic Change Research [email protected]

FAPESPCentro de Tecnologia Canavieira – Piracicaba

Ministério da Ciência e TecnologiaNature Conservancy

Marcos S. Buckeridge, 2008

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Marcos S. Buckeridge, 2008Marcos S. Buckeridge, 2008

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[email protected]

Thank You

A written version of this talk can be found in my article written for Comciencia/Labjorat

http://www.comciencia.br/comciencia

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Mecanismo de Conecção e disparo

Um mecanismo básico para a semióseambiental

AmbienteMetabolism

o

W1W2

W3

W4

Σw ≅ Σy

Y1

Y1

Y1

Y1

LuzTemperaturaÁguaCO2

Expressão gênicaAção enzimática

Interações moleculares

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Número de links (k)

Núm

ero

de n

ós c

om k

links

Núm

ero

de n

ós c

om k

links

Número de links (k)

Por quê os controladores de vôo de Brasília derrubaram a rede aérea brasileira, mas o acidente com o vôo 1907 não?

Rede hierárquica

Rede a

o aca

so

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PROPRIEDADES DAS REDES

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Marcos S. Buckeridge, 2008Roger Guimerà & Luís A. Nunes Amaral NATURE vol. 433, 2005

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Fot

ossí

nte

se

ENERGIA

CO2

H O2

HEXOSES

Reservas

Importação

Via

das

pent

oses

(NA

DPH

+H) +

Glic

ólise

Piruvato(Via anaeróbica)

FermentaçãoLactatoEtanol

(Via

aer

óbic

a)

Ciclo do ácidotricarboxílico

(redução de NAD)

Esqueleto deCarbono

MANUTENÇÃOCRESCIMENTO

CO2

Cadeia de Transportede elétrons

(oxidação de NAD namembrana interna)

H O2

(via ATPase) (ATP)

respiração insensível ao Cianeto

Lipídeos

ProteínasÁcidos nucléicosLipídeosCompostos sec.

β-oxidação

NAD

H+H

+

Fluxo de elétronsa favor do gradiente ENERGIA

(calor)

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Pirâmide da universalidade (Oltvai, Z.N. & Barabási, A.L. 2002, Science 298: 763)

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Marcos S. Buckeridge, 2008Marcos S. Buckeridge, 2008