• Energy

– Flujo de energía en el ecosistema

Light energy

ECOSYSTEM

Photosynthesis
in chloroplasts
Organic
CO2 + H2O +O
molecules 2
Cellular respiration
in mitochondria

ATP

powers most cellular work

Heat
Figure 9.2 energy

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 2 e – + 2 H+
2 e– + H+
NAD +
NADH
H
O Dehydrogenase
H H O
NH2 + 2[H] Reduction of NAD+ NH2 +
C C
(from food) Oxidation of NADH
N+ N Nicotinamide
Nicotinamide
O CH2 (oxidized form) (reduced form)
O
O P O –

O H H
O P O– HO OH NH2
HO
CH2
O N N
H
N N H
O

H H
HO OH Figure 9.4
 • Glicolisis
– Significa “Rompiento de azúcar”
(6-C a 3-C azúcar)
– Rompiento de la glucosa en piruvato
– Ocurre en el citoplasma de la célula

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 • La glicolisis consiste en 2 fases:
– Fase de inversión de energía
– Generación de energía
Glycolysis Citric Oxidative
acid
cycle phosphorylation

ATP ATP ATP

Energy investment phase

Glucose

2 ATP + 2 P 2 ATP used

Energy payoff phase

4 ADP + 4 P 4 ATP formed

2 NAD+ + 4 e- + 4 H 2 NADH + 2 H+
+

2 Pyruvate + 2 H2O

Glucose 2 Pyruvate + 2 H2O

4 ATP formed – 2 ATP used 2 ATP + 2 H+
2 NAD+ + 4 e– + 4 H 2 NADH
Figure 9.8 +

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 • Fase de inversión
CH2OH
Citric
HH H Glycolysis acid Oxidative
HO H cycle phosphorylation
HO OH
H OH
Glucose

ATP 1
Hexoquinasa
ADP
CH2OH P
HH OH
OH H
HO
H OH
Glucose-6-phosphate
2
Fosfoglucoisomerasa
CH2O P
O CH2OH
H HO
H HO
HO H
Fructose-6-phosphate
3
ATP
Fosfofructoquinasa
ADP
P O CH2 O CH2 O P
HO
H OH
HO H
Fructose-
1, 6-bisphosphate 4
aldolasa
5 H
P O CH2 Isomerase
C O
C O
CHOH
CH2OH
CH2 O P
Dihydroxyacetone Glyceraldehyde-
phosphate 3-phosphate
Figure 9.9 A
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 • Fase de Generación de energía
6
2 NAD+
Triosa fosfato deshidrogenasa
2 NADH 2 Pi
+ 2 H+
2
P O C O
CHOH
CH2 O P
1, 3-Bisphosphoglycerate
2 ADP
7
Fosfogliceroquinasa
2 ATP

2 O–
C
CHOH
CH2 O P
3-Phosphoglycerate
8
Fosfogliceromutasa
2 O–
C O
H C O P
CH2OH
2-Phosphoglycerate
9
2 H2O Enolasa
2 O–
C O
C O P
CH2
Phosphoenolpyruvate
2 ADP
10
Piruvato quinasa
2 ATP

2 O–
C O
C O
CH3
Figure 9.8 B Pyruvate

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 El ciclo del ácido cítrico completa la
oxidación de las moléculas orgánicas.
– Se lleva a cabo en la matriz de la mitocondria
– Completa la oxidación de la glucosa por
rompimiento del piruvato a CO2.

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 • Ante que comience el ciclo de Krebs
– El piruvato debe convertirse en Acetil-CoA,
lo que liga la glicolisis con el ciclo del
ácido cítrico.
CYTOSOL MITOCHONDRION

NAD+ NADH + H+
O–
S CoA
2
C O
C O

C O
CH3
1 3
CH3
Acetyle CoA
Pyruvate CO2 Coenzyme A

Transport protein

Figure 9.10

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 • Panorama general del ciclo del ácido cítrico

Pyruvate Glycolysis Citric

Oxidative
acid
(de la glicolisis, cycle phosphorylation

2 moleculas por glucosa)

ATP ATP ATP

CO2

CoA
NADH
+ 3 H+ Acetyle CoA
CoA
r cada ciclo, 2 carbonos salen como CO2,
se generan 3 NADH y 1 FADH2 y 1 ATP. CoA

Citric
acid 2 CO2
cycle
FADH2 3 NAD+

FAD 3 NADH
+ 3 H+
ADP + P i
ATP
Figure 9.11
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 Glycolysis Citric Oxidative
acid phosphorylation
cycle

S CoA
C O
CH3

Acetyl CoA
CoA SH

NADH O C COO–
+H + CH2 1 COO– H2O
COO – CH2 COO–
NAD+
8 Oxaloacetate HO C COO –
CH2
CH2
2
COO – HC COO–
COO– HO CH
HO CH
Malate Citrate
CH2 COO–
Isocitrate
COO–
Figure 9.12
Citric
CO2
3
7
acid
NAD+
H2O cycle
COO–
COO–
NADH
CH + H+
Fumarate CoA SH CH2
HC
CH2 a-Ketoglutarate
COO–
C O
6 4
CoA SH COO– COO–
COO–
CH2 CH2
FADH2 5
CH2
CO2
CH2
FAD NAD+
COO– C O

Succinate Pi S CoA NADH

GTP GDP Succinyl + H+
CoA
ADP

ATP

Figure 9.12
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 • Concept 9.4: During oxidative
phosphorylation, chemiosmosis couples
electron transport to ATP synthesis
• NADH and FADH2

– Donate electrons to the electron transport chain,

which powers ATP synthesis via oxidative
phosphorylation

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 The Pathway of Electron Transport

– Electrons from NADH and FADH2

lose energy in several steps
– Couples this exergonic slide of
electrons to ATP synthesis or
oxidative phosphorylation
– The electron transport chain is
made of electron carrier
molecules embedded in the inner
membrane of mitochondria.
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 – Each carrier in the chain has a higher
electronegativity than the carrier
before it, so electrons are pulled
downhill towards oxygen
– Most carriers are protein molecules
except for ubiquinone (Q)
– At the end of the chain, electrons are
passed to oxygen, forming water

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 NADH is oxidized and flavoprotein is reduced as high
NADH energy electrons from NADH are transferred to FMN
50
↓
FADH2 flavoprotein is oxidized as it passes electrons to an iron
Multiprotein
sulfur protein, FeS.
40 I
↓
FMN FAD complexes
Free energy (G) relative to O2 (kcl/mol)

Fe•S Fe•S II

O FeS is oxidized as it pass electrons to ubiquinone Q

III

30
Cyt b
Fe•S
↓
Cyt c1 Q passes electrons on to a succession of electron carriers,
IV
Cyt c most of which are cytochromes.
Cyt a

20
Cyt a3 ↓
cyt a3 , the last cytochrome passes electrons to oxygen.

10

0 2 H + + 12 O2

Figure 9.13 H2O

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 Chemiosmosis: The Energy-Coupling Mechanism
• ATP synthase
– Is the enzyme that actually makes ATP
INTERMEMBRANE SPACE A rotor within the
H+
membrane spins
H+ H+ clockwise when
H+ H+ flows past
H+
it down the H+
gradient.
H+
H+
A stator anchored
in the membrane
holds the knob
stationary.

A rod (for “stalk”)

extending into
the knob also
spins, activating
catalytic sites in
H+ the knob.
Three catalytic
sites in the
ADP stationary knob
+ join inorganic
Pi ATP
Phosphate to ADP
to make ATP.
MITOCHONDRIAL MATRIX
Figure 9.14
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 • At certain steps along the electron transport
chain
– Electron transfer causes protein complexes
to pump H+ from the mitochondrial matrix
to the intermembrane space

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 • Chemiosmosis
– Is an energy-coupling mechanism that uses
energy in the form of a H+ gradient across a
membrane to drive cellular work

• The resulting H+ gradient

– Stores energy
– Drives chemiosmosis in ATP synthase
– Is referred to as a proton-motive force

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 • Chemiosmosis and the electron transport chain

Inner
Mitochondrial
Glycolysis
Oxidative
phosphorylation. membrane
electron transport
and chemiosmosis

ATP ATP ATP

H+
H +

H+
H+
Cyt c
Protein complex
Intermembrane of electron
space carners
Q IV
I III
ATP
Inner II synthase
mitochondrial FADH2 H2O
membrane FAD+ 2 H+ + 1/2 O2
NADH+
NAD+ ADP + Pi ATP
(Carrying electrons
from, food) H+
Mitochondrial Chemiosmosis
Electron transport chain
matrix Electron transport and pumping of protons (H ),
+ ATP synthesis powered by the flow
which create an H gradient across the membrane Of H back across the membrane
+ +

Figure 9.15 Oxidative phosphorylation

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 An Accounting of ATP Production by Cellular
Respiration
• During respiration, most energy flows in
this sequence
– Glucose to NADH to electron transport
chain to proton-motive force to ATP

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 • There are three main processes in this
metabolic enterprise
Electron shuttles MITOCHONDRION
CYTOSOL 2 NADH
span membrane
or
2 FADH2

2 NADH 2 NADH 6 NADH 2 FADH2

Glycolysis Oxidative
2 Citric phosphorylation:
2 Acetyl acid electron transport
Glucose Pyruvate CoA cycle and
chemiosmosis

+ 2 ATP + 2 ATP + about 32 or 34 ATP

by substrate-level by substrate-level by oxidative phosphorylation, depending
phosphorylation phosphorylation on which shuttle transports electrons
from NADH in cytosol

About
Maximum per glucose: 36 or 38 ATP

Figure 9.16

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 • About 40% of the energy in a glucose molecule
– Is transferred to ATP during cellular
respiration, making approximately 38 ATP

1 ATP → - 7.3 kcal/mol

38 ATP X 7.3 / 686 = 40%

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 • Concept 9.5: Fermentation enables some
cells to produce ATP without the use of
oxygen
• Cellular respiration
– Relies on oxygen to produce ATP
• In the absence of oxygen
– Cells can still produce ATP through
fermentation

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 • Glycolysis
– Can produce ATP with or without oxygen, in
aerobic or anaerobic conditions
– Couples with fermentation to produce ATP

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 Types of Fermentation
• Fermentation consists of
– Glycolysis plus reactions that regenerate
NAD+, which can be reused by glyocolysis

• In alcohol fermentation
– Pyruvate is converted to ethanol in two
steps, one of which releases CO2
• During lactic acid fermentation
– Pyruvate is reduced directly to NADH to
form lactate as a waste product
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 2 ADP + 2 P1 2 ATP O–
C O
C O
Glucose Glycolysis
CH3
2 Pyruvate
2 NAD + 2 NADH 2 CO2

H H
H C OH C O
CH3 CH3
2 Ethanol 2 Acetaldehyde
(a) Alcohol fermentation

2 ADP + 2 P1 2 ATP

Glucose Glycolysis O–
C O
C O
2 NAD+ 2 NADH
O CH3
C O
H C OH
CH3
2 Lactate

Figure 9.17 (b) Lactic acid fermentation

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 Fermentation and Cellular Respiration Compared

• Both fermentation and cellular respiration

– Use glycolysis to oxidize glucose and other
organic fuels to pyruvate
• Fermentation and cellular respiration
– Differ in their final electron acceptor
• Cellular respiration
– Produces more ATP

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 • Pyruvate is a key juncture in catabolism

Glucose

CYTOSOL

Pyruvate
No O2 present O2 present
Fermentation Cellular respiration

MITOCHONDRION
Ethanol Acetyl CoA
or
lactate
Citric
acid
cycle

Figure 9.18

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 The Evolutionary Significance of Glycolysis
• Glycolysis
– Occurs in nearly all organisms
– Probably evolved in ancient prokaryotes
before there was oxygen in the atmosphere

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 • Concept 9.6: Glycolysis and the citric acid
cycle connect to many other metabolic
pathways

The Versatility of Catabolism

• Catabolic pathways
– Funnel electrons from many kinds of
organic molecules into cellular respiration

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 • The catabolism of various molecules from food
Proteins Carbohydrates Fats

Amino Sugars Glycerol Fatty

acids acids

Glycolysis
Glucose

Glyceraldehyde-3- P

NH3 Pyruvate

Acetyl CoA

Citric
acid
cycle

Oxidative
Figure 9.19 phosphorylation

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 Biosynthesis (Anabolic Pathways)
• The body
– Uses small molecules to build other
substances
• These small molecules
– May come directly from food or through
glycolysis or the citric acid cycle

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 Regulation of Cellular Respiration via Feedback
Mechanisms
• Cellular respiration
– Is controlled by allosteric enzymes at key
points in glycolysis and the citric acid cycle

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 • The control of cellular respiration
Glucose
AMP
Glycolysis

Fructose-6-phosphate Stimulates
+
Phosphofructokinase
– –
Fructose-1,6-bisphosphate
Inhibits Inhibits

Pyruvate

ATP Citrate
Acetyl CoA

Citric
acid
cycle

Oxidative
Figure 9.20 phosphorylation

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 • An overview of cellular respiration

Electrons Electrons carried

carried via NADH and
via NADH FADH2

Oxidative
Citric phosphorylation:
Glycolsis
Glucos acid electron
Pyruvate cycle
e transport and
chemiosmosis

Cytosol
Mitochondrion

ATP ATP ATP

Substrate-level Oxidative
Substrate-level
phosphorylation phosphorylation
Figure 9.6 phosphorylation

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 Respiración celular
• La respiración es la acumulaciónde 3 etapas
metabólicas
– Glicolisis does not require O2

– Ciclo del ácido cítrico o ciclo de Krebs

require O2
– Fosforilación oxidativa

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