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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
Marcos S. Buckeridge, 2008
Air in samples of the AntarticContinent
Marcos S. Buckeridge, 2008Marcos S. Buckeridge, 2008
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)
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
Marcos S. Buckeridge, 2008Marcos S. Buckeridge, 2008
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
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
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
Marcos S. Buckeridge, 2008
How can we use plant science to help choosing
the best way to go?
Marcos S. Buckeridge, 2008
Marcos S. Buckeridge, 2008
THE CASE OF TREESregenerating forests and reestablishing
ecosystem services
Marcos S. Buckeridge, 2008
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
Marcos S. Buckeridge, 2008
Plantas jovens de jatobá crescendo por 90 dias com e sem os cotilédones
Marcos S. Buckeridge, 2008
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
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
Marcos S. Buckeridge, 2008
CotyledonXG catabolism
Plantlet
Photosynthesis
sucrose
GROWTH
sucrose
IAA
CO2
Marcos S. Buckeridge, 2008
CotyledonXG catabolism
Plantlet
sucrose
GROWTH
sucrose
IAA
CO2
Photosynthesis
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
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
Marcos S. Buckeridge, 2008
OPEN TOP CHAMBER
CO2
4 m
1,5 m
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
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
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
Marcos S. Buckeridge, 2008
Plant organsPlant organs andand theirtheircarboncarbon metabolismmetabolism
Marcos S. Buckeridge, 2008
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
*
*
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
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
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
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
Marcos S. Buckeridge, 2008
Other native tree species of the Atlantic Forest
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
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
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
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
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
Marcos S. Buckeridge, 2008
Jatobá e açaí crescendo em alto CO2 e alta temperatura (+3oC)
Marcos S. Buckeridge, 2008
Marcos S. Buckeridge, 2008
NEXT – The combined effects of CO2and temperature
Marcos S. Buckeridge, 2008
SUGAR CANE
Physiological behaviour under elevated CO2
Marcos S. Buckeridge, 2008
Marcos S. Buckeridge, 2008Marcos S. Buckeridge, 2008
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
Marcos S. Buckeridge, 2008
Sugar cane in the open top chambers
360
360
720
720
Finantial support by Centro de Tecnologia Canavieira - Piracicaba
Marcos S. Buckeridge, 2008
Experimental design
CO2
4 m
1,5 m
Sugarcane variety SP80-3280
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
***
******
****
Marcos S. Buckeridge, 2008
ElevatedAmbient
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
**
***
******
**
***
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
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
Marcos S. Buckeridge, 2008
ElevatedAmbientProductivity
Marcos S. Buckeridge, 2008
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
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
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
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
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
Marcos S. Buckeridge, 2008
Marcos S. Buckeridge, 2008
-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
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
Marcos S. Buckeridge, 2008
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
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
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!
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
Marcos S. Buckeridge, 2008
Is the Midway
POSSIBLE?
FEASIBLE?
Marcos S. Buckeridge, 2008
Rodovia dos Bandeirantes – São PauloAgosto 2007
Marcos S. Buckeridge, 2008
Marcos S. Buckeridge, 2008
A tree of Schizolobium parayba
growing in a regenerating forest just beside a
sugarcane plantation
Rodovia dos BandeirantesSão Paulo
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
Marcos S. Buckeridge, 2008Marcos S. Buckeridge, 2008
Marcos S. Buckeridge, 2008
Thank You
A written version of this talk can be found in my article written for Comciencia/Labjorat
http://www.comciencia.br/comciencia
Marcos S. Buckeridge, 2008
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
Marcos S. Buckeridge, 2008
Marcos S. Buckeridge, 2008
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
Marcos S. Buckeridge, 2008
PROPRIEDADES DAS REDES
Marcos S. Buckeridge, 2008
Marcos S. Buckeridge, 2008Roger Guimerà & Luís A. Nunes Amaral NATURE vol. 433, 2005
Marcos S. Buckeridge, 2008
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)
Marcos S. Buckeridge, 2008
Pirâmide da universalidade (Oltvai, Z.N. & Barabási, A.L. 2002, Science 298: 763)
Marcos S. Buckeridge, 2008Marcos S. Buckeridge, 2008