PHOTOSYNTHESIS
Autotrophs
 producers of the biosphere, producing organic
molecules from CO2 and other inorganic molecules
 photoautotrophs - use the energy of sunlight to make
organic molecules from H2O and CO2
 feed not only themselves but also most of the living
world
Heterotrophs
 obtain their organic material from other organisms
 consumers of the biosphere
 almost all heterotrophs depend on photoautotrophs
for food and O2
 Decomposers: consume the remains of other
organisms by decomposing and feeding on organic
litter such as dead organisms, feces, and fallen
leaves (most fungi and many types of prokaryotes)
 spectrum used for photosynthesis
Photosynthesis
 process that converts solar energy into chemical
energy
 directly or indirectly nourishes almost the entire living
world
 occurs in plants, algae, certain other protists, and
some prokaryotes
Chloroplasts: The Sites of Photosynthesis in Plants
 Leaves - major locations of photosynthesis
 Chlorophyll - green pigment within chloroplasts
 Chloroplasts are found mainly in the cells of the
mesophyll, the tissue in the interior of the leaf.
 Light energy absorbed by chlorophyll drives the
synthesis of organic molecules in the chloroplast
 CO2 enters and O2 exits the leaf through microscopic
pores called stomata
 Chlorophyll is in the membranes of thylakoids
 Chloroplasts also contain stroma, a dense fluid
Summary Equation:
6 CO2 + 12 H2O + Light energy  C6H12O6 + 6 O2 + 6 H2O
Chloroplasts split H2O into hydrogen and oxygen,
incorporating the electrons of hydrogen into sugar molecules
Photosynthesis is a redox process in which H2O is oxidized
and CO2 is reduced.
Two Stages of Photosynthesis
 Light reactions (the photo part) and Calvin cycle (the
synthesis part)
 The light reactions (in the thylakoids): (1) split H2O (2)
release O2 (3) reduce NADP+ to NADPH and (4)
generate ATP from ADP by photophosphorylation
 Calvin cycle (in the stroma) forms sugar from CO2,
using ATP and NADPH
 Calvin cycle begins with carbon fixation, incorporating
CO2 into organic molecules
Nature of Sunlight:
 Light is a form of electromagnetic energy, also called
electromagnetic radiation (travels in rhythmic waves)
 Wavelength determines the type of electromagnetic
energy
 Visible light consists of wavelengths (including those
that drive photosynthesis) that produce colors we can
see
 Light also behaves as though it consists of discrete
particles, called photons
Photosynthetic Pigments:
 Substances that absorb visible light
 Different pigments absorb different wavelengths
 Chlorophyll a - main photosynthetic pigment
 Accessory pigments (chlorophyll b), broaden the
 Carotenoids absorb excessive light that would
damage chlorophyll (photoprotection)
The chlorophyll molecules of chloroplasts absorb vioIet-blue
and red light (the colors most effective in driving
photosynthesis) and reflect or transmit light. This is why leaves
appear green.
 An absorption spectrum is a graph plotting a
pigment’s light absorption versus wavelength
 The absorption spectrum of chlorophyll a suggests
that violet-blue and red light work best for
photosynthesis
Photosystem:
 consists of a reaction-center complex surrounded by
light-harvesting complexes
 light-harvesting complexes (pigment molecules bound
to proteins) funnel the energy of photons to the
reaction center
 A primary electron acceptor in the reaction center
accepts an excited electron from chlorophyll a
 PS II functions first and is best at absorbing a
wavelength of 680 nm
o The reaction-center chlorophyll a of PS II is
called P680
  PS I is best at absorbing a wavelength of 700 nm.
The reaction-center chlorophyll a of PS I is called
P700
Cyclic Electron Flow
 uses only photosystem I and produces ATP, but not
NADPH
 generates surplus ATP, satisfying the higher demand
in the Calvin cycle
Chemiosmosis: Chloroplasts vs Mitochondria
 Chemiosmosis - generation of ATP in chloroplasts
and mitochondria
 Mitochondria transfer chemical energy from food to
ATP; chloroplasts transform light energy into the
chemical energy of ATP
 Spatial organization of chemiosmosis differs between
chloroplasts and mitochondria but also shows
similarities
 Comparison of chemiosmosis in mitochondria and
chloroplasts. In both kinds of organelles, electron
transport chains pump protons (H+) across a
membrane from a region of low H+ concentration to
one of high H+ concentration. The protons then
diffuse back across the membrane through ATP
synthase, driving the synthesis of ATP.
Calvin Cycle
 Builds sugar from smaller molecules by using ATP
and the reducing power of electrons carried by
NADPH
 Carbon enters the cycle as CO2 and leaves as a
sugar named glyceraldehyde-3-phospate (G3P)
 Three phases: (1) Carbon fixation (catalyzed by
rubisco) (2) Reduction (3) Regeneration of the CO2
acceptor (RuBP)
(1) Incorporation of each CO2 molecule, one at a time, by
attaching it to a five carbon sugar named ribulose
bisphosphate (abbreviated RuBP).RuBP carboxylase,
or rubisco catalyzes this first step. Product of the
reaction is a six-carbon intermediate that splits in half,
forming two molecules of 3- phosphoglycerate
(2) Each molecule of 3- phosphoglycerate receives an
additional phosphate group from ATP, becoming
1,3·bisphosphoglycerate. Next, a pair of electrons
donated from NADPH reduces 1,3-
bisphosphoglycerate, which also loses a phosphate
group, becoming G3P. One molecule exits the cycle
to be used by the plant cell, but the other five
molecules must be recycled to regenerate the three
molecules of RuBP.
(3) Carbon skeletons of five molecules of G3P are
rearranged by the last steps of the Calvin cycle into
three molecules of RuBP. To accomplish this, the
cycle spends three more molecules of ATP. The
RuBP is now prepared to receive CO2 again, and the
cycle continues.
For the net synthesis of one G3P molecule, the Calvin recycle
consumes nine ATP and six NAPDH. The G3P from the Calvin
cycle is the starting material for metabolic pathways that
synthesize other organic compounds, including glucose and
other carbohydrates.
Alternative mechanisms of carbon fixation have evolved in hot,
arid climates
 One of the major problems facing terrestrial plants is
dehydration.
 The stomata are not only the major route for gas
exchange (CO2 in and O2 out), but also for the
evaporative loss of water.
 On hot, dry days plants close the stomata to conserve
water, but this causes problems for photosynthesis
In most plants (C3 plants) initial fixation of CO2 occurs via
rubisco and results in a three-carbon compound, 3-
phosphoglycerate.
These plants include rice, wheat, and soybeans.
 While rubisco normally accepts CO2, when the
O2/CO2 ratio increases (on a hot, dry day with closed
stomata), rubisco can add O2 to RuBP.
 When rubisco adds O2 to RuBP, RuBP splits into a
three-carbon piece and a two-carbon piece in a
process called photorespiration.
o The two-carbon fragment is exported from
the chloroplast and degraded to CO2 by
mitochondria and peroxisomes.
o Unlike normal respiration, this process
produces no ATP, nor additional organic
molecules.
 Photorespiration decreases photosynthetic output by
siphoning organic material from the Calvin cycle.
The C4 plants fix CO2 first in a four-carbon compound.
Several thousand plants, including sugarcane and corn, use
this pathway.
 In C4 plants, mesophyll cells incorporate CO2 into
organic molecules.
o The key enzyme, phosphoenolpyruvate
carboxylase, adds CO2 to
phosphoenolpyruvate (PEP) to form
oxaloacetetate.
o PEP carboxylase has a very high affinity for
CO2 and can fix CO2 efficiently when
rubisco cannot - on hot, dry days with the
stomata closed
 In effect, the mesophyll cells pump CO2 into the
bundle sheath cells, keeping CO2 levels high enough
for rubisco to accept CO2 and not O2
 C4 photosynthesis minimizes photorespiration and
enhances sugar production.
 C4 plants thrive in hot regions with intense sunlight.

A second strategy to minimize photorespiration is found in
succulent plants, cacti, pineapples, and several other plant
families. These plants, known as CAM plants for crassulacean
acid metabolism (CAM), open stomata during the night and
close them during the day.
 During the night, these plants fix CO2 into a variety of
organic acids in mesophyll cells.
 During the day, the light reactions supply ATP and
NADPH to the Calvin cycle and CO2 is released from
the organic acids.
Photosynthesis is the biosphere’s metabolic foundation
In photosynthesis, the energy that enters the chloroplasts as
sunlight becomes stored as chemical energy in organic
compounds.
CELLULAR RESPIRATON
(1) Energy flows into an ecosystem as sunlight and
leaves as heat
(2) Photosynthesis generates O2 and organic molecules,
which are used in cellular respiration
(3) Cells use chemical energy stored in organic
molecules to regenerate ATP, which powers work
Catabolic Pathways and Production of ATP
 Fermentation - partial degradation of sugars that
occurs without O2
 Aerobic respiration consumes organic molecules and
O2 and yields ATP
 Anaerobic respiration consumes compounds other
than O2
Cellular Respiration
 Includes both aerobic and anaerobic respiration but is
often used to refer to aerobic respiration
 Although carbohydrates, fats, and proteins are all
consumed as fuel, it is helpful to trace cellular
respiration with the sugar glucose:
C6H12O6 + 6 O2  6 CO2 + 6 H2O + Energy (ATP
+ heat)
 NAD+ - Nicotinamide adenine dinucleotide; a
coenzyme occurring in most living cells and used as
an oxidizing or reducing agent in various metabolic
processes.
 NADH - The reduced form of NAD.
 Photophosphorylation - addition of a phosphate group
 Reduction - addition of hydrogen or electrons or
removal of oxygen
 Oxidation - removal of hydrogen or electrons or
addition of oxygen.
Oxidation of Organic Fuel Molecules
During cellular respiration, the fuel (such as glucose) is
oxidized, and O2 is reduced:
Stepwise Energy Harvest via NAD+ and the Electron Transport
Chain
(1) In cellular respiration, glucose and other organic
molecules are broken down in a series of steps
(2) Electrons from organic compounds are usually first
transferred to NAD+ , a coenzyme
(3) NAD+ functions as an oxidizing agent during cellular
respiration
(4) Each NADH (the reduced form of NAD+ ) represents
stored energy that is tapped to synthesize ATP
NADH+ as an electron shuttle
The full name for NAD+, nicotinamide adenine dinucleotide,
describes its structure: the molecule consists of two
nucleotides joined together at their phosphate groups (shown
in yellow). Nicotinamide is a nitrogenous base. In oxidation
reactions, each electron (e– ) travels with a proton (H+ )-thus,
as a hydrogen atom. The hydrogen atoms are not transferred
directly to oxygen, but instead are usually passed first to an
electron carrier, a coenzyme NAD+ . As an electron acceptor,
NAD+ functions as an oxidizing agent during respiration.
Enzyme called dehydrogenase removes a pair of hydrogen
atoms (2 electrons and 2 protons) from the substrate (glucose,
in this example), thereby oxidizing it. The enzyme delivers the
2 electrons along with 1 proton to its coenzyme, NAD+. The
other proton is released as a hydrogen ion (H+) into the
surrounding solution:
The Stages of Cellular Respiration
Three stages:
(1) Glycolysis (breaks down glucose into two molecules
of pyruvate)
- Glycolysis, which occurs in the cytosol, begins
chemiosmosis the degradation process by
breaking glucose into two molecules of a
compound called pyruvate
- Pyruvate enters the mitochondrion. where the
citric acid cycle oxidizes it to carbon dioxide.
(2) Citric acid cycle (completes the breakdown of
glucose)
- NADH and electron carrier coenzyme called
FADH transfer electrons derived from glucose to
electron transport chains which are built into the
inner mitochondrial membrane.
(3) Oxidative phosphorylation (accounts for most of the
ATP synthesis)
- During oxidative phosphorylation, electron
transport chains convert the chemical energy to a
form used for ATP synthesis in the process called
chemiosmosis.
Oxidative Phosphorylation: (6) Triose phosphate dehydrogenase catalyzes two
 Process that generates most of the ATP (powered by sequential reactions while it holds glyceraldehyde-3-
redox reactions) phosphate in its active site. Sugar is oxidized by the
 Accounts for almost 90% of the ATP generated by transfer of electrons and H+ to NAD+, forming NADH
cellular respiration (a redox reaction). Coefficient 2 precedes all
Substrate-Level Phosphorylation: molecules in the energy payoff phase; these steps
 Occurs when an enzyme transfers a phosphate group occur after glucose has been split into two three-
from a substrate molecule to ADP, rather than adding carbon sugars. This step yields 1,3-
an inorganic phosphate to ADP as in oxidative bisphosphoglycerate.
phosphorylation (7) This step produces 2 ATP, since every product after
 "Substrate molecule" here refers to an organic the sugarsplitting step is doubled. This ATP debt has
molecule generated as an intermediate during the now been repaid. Glucose has been converted to two
catabolism of glucose. molecules of 3- phosphoglycerate (which is not a
 forms a smaller amount of ATP in glycolysis and the sugar) by phosphyglerokinase
citric acid cycle (8) Phosphoglyceromutase relocates the remaining
phosphate group, preparing 2-phosphoglycerate for
Glycolysis the next step.
 breaks down glucose into two molecules of pyruvate (9) Enolase causes a double bond to form in the
(“splitting of sugar”) substrate by extracting a water molecule, yielding
phosphoenolpyruvate
 occurs in the cytoplasm and has two major phases:
(10) The last reaction of glycolysis produces more ATP by
(1) energy investment phase and (2) energy payoff
transferring the phosphate group from PEP to ADP, a
phase
second instance of substrate level phosphorylation.
(1) Glucose, a six-carbon sugar, is split into two three-
Catalyzed by pyruvate kinase.
carbon sugars. Smaller sugars are then oxidized and
their remaining atoms rearranged to form two
Pyruvate Oxidation
molecules of pyruvate. (Pyruvate is the ionized form
In the presence of O2, pyruvate enters the mitochondrion.
of pyruvic acid.)
Pyruvate must be converted to acetyl CoA, which links the
(2) Cell actually spends ATP
cycle to glycolysis.
(3) Investment is repaid with interest during the energy
Pyruvate is a charged molecule, so in eukaryotic cells it must
payoff phase, when ATP is produced by substrate-
enter the mitochondnon via active transport, with the help of a
level phosphorylation and NAD+ is reduced to NADH
transport protein. A complex of several enzymes catalyzes the
by electrons released from the oxidation of glucose.
three numbered steps
(4) Net energy yield from glycolysis, per glucose
(1) Pyruvate's carboxyl group (-COO- ) is removed and
molecule:
given off as a molecule of CO2
a. 2 Pyruvate + 2H2O
(2) The remaining two-carbon fragment is oxidized,
b. 2 ATP
forming a compound named acetate. An enzyme
c. 2NADH + 2H+
transfers the extracted electrons to NAD+ , storing
energy in the form of NADH.
Specific steps:
(3) Coenzyme A (CoA), a sulfur containing compound is
(1) Glucose enters the cell and is phosphorylated by the
attached to the acetate by an unstable bond that
enzyme hexokinase, which transfers a phosphate
makes the acetyl group very reactive.
group from ATP to the sugar.
(2) Glucose-6- phosphate is converted to its isomer,
Citric Acid Cycle
fructose6-phosphate.
(3) Phosphofructokinase enzyme transfers a phosphate  Takes place within the mitochondrial matrix
group from ATP to the sugar, investing another  Oxidizes organic fuel derived from pyruvate,
molecule of ATP in glycolysis. So far, 2 ATP have generating 1 ATP, 3 NADH, and 1 FADH2 per turn
been used. With phosphate groups on its opposite (1) Acetyl CoA adds its two-carbon acetyl group to
ends, the sugar is now ready to be split in half. This is oxaloacetate, producing citrate (citrate synthase)
a key step for regulation of glycolysis. (2) Citrate is converted to its isomer, isocitrate, by
(4) Aldolase cleaves the sugar molecule into two different removal of one water molecule and addition of
three carbon sugars: dihydroxyacetone phosphate another
and glyceraldehyde-3- phosphate. These two sugars (3) Isocitrate is oxidized, reducing NAD+ to NADH. Then
are isomers of each other. the resulting compound loses a CO2 molecule
(5) Isomerase catalyzes the reversible conversion (isocitrate dehydrogenase)
between the two three-carbon sugars.
 (4) Another CO2 is lost, and -Ketoglutarate is oxidized,
reducing NAD+ to NADH. The remaining molecule is
then attached to coenzyme A by an unstable bond.
(alpha-ketoglutarate dehydrogenase)
(5) CoA in succinyl CoA is displaced by a phosphate
group, which is transferred to GDP forming GTP, a
molecule with function similar to ATP that, in some
cases, is used to generate ATP. (succinyl CoA
dehydroganse)
(6) Two hydrogens are transferred to FAD (flavin adenine
dinucleotide), forming FADH2 and oxidizing succinate
to fumarase (succinate dehydrogenase)
(7) Addition of a water molecule rearranges bonds in the
substrate, turning fumarate to malate (fumarase)
(8) Malate is oxidized, reducing NAD+ to NADH and
regenerating oxaloacetate (malate dehydrogenase)
NADH and FADH2 produced by the cycle relay electrons
extracted from food to the electron transport chain ATP is also
produced.
Oxidative Phosphorylation
 NADH and FADH2 account for most of the energy
extracted from food. These two electron carriers
donate electrons to the electron transport chain,
which powers ATP synthesis via oxidative
phosphorylation.
 ETC: Electron transport and pumping of protons (H+),
which create an H+ gradient across the membrane
 ATP synthesis powered by the flow of H+ back across
the membrane
Electron Transport Chain:
Found in the cristae of mitochondrion
 The carriers alternate reduced and oxidized states as
they accept and donate electrons
 Electrons are passed through a number of proteins
including cytochromes
 Electrons are finally passed to O2 , forming H2O
(1) Electrons removed from glucose by NAD+, during
glycolysis and the citric acid cycle, are transferred
from NADH to the first molecule of the electron
transport chain in complex I. This molecule is a
flavoprotein, called flavin mononucleotide (FMN).
(2) Flavoprotein returns to its oxidized form as it passes
electrons to an iron-sulfur protein (Fe-S in complex I)
(3) Iron-sulfur protein then passes the electrons to a
compound called ubiquinone (Q).
(4) Another source of electrons for the transport chain is
FADH2, which adds its electrons to the electron
transport chain at complex II a lower energy level than
NADH does
(5) Most of the remaining electron carriers between
ubiquinone and oxygen are proteins called
cytochromes, each a different protein with a slightly
different electron-carrying heme group
a. In Complex III: Cyt b, FeS, Cyt C1
b. Cyt C connects III and IV
c. In Complex IV: Cyt a, Cyt a3
(6) The last cytochrome of the chain, cyt a3, passes its
electrons to oxygen, which is very electronegative
(7) Each oxygen atom also picks up a pair of hydrogen
ions from the aqueous solution, forming water.
Chemiosmosis: The Energy-Coupling Mechanism
 ATP synthase, the enzyme that actually makes ATP
from ADP and inorganic phosphate.
 ATP synthase uses the energy of an existing ion
gradient to power ATP synthesis.
 ATP synthase uses the exergonic flow of H+ to drive
phosphorylation of ATP
 This is an example of chemiosmosis, the use of
energy in a H+ gradient to drive cellular work
Chemiosmosis: The Energy-Coupling Mechanism
 The energy stored in a H+ gradient across a
membrane couples the redox reactions of the electron
transport chain to ATP synthesis
 The H+ gradient is referred to as a proton-motive
force, emphasizing its capacity to do work
 NADH and FADH2 shuttle high-energy electrons
extracted from food during glycolysis and the citric
acid cycle to an electron transport chain built into the
inner mitochondrial membrane. The gold arrows trace
the transport of electrons, which finally pass to
oxygen at the "downhill" end of the chain, forming
water.
 Most of the electron carriers of the chain are grouped
into four complexes. Two mobile carriers, ubiquinone
(Q) and cytochrome c (Cyt c),move rapidly, ferrying
electrons between the large complexes. As
complexes I, III, and IV accept and then donate
electrons, they pump protons from the mitochondrial
matrix into the intermembrane space.
 During chemiosmosis, the protons flow back down
their gradient via ATP synthase, which is built into the
membrane nearby. The ATP synthase harnesses the
protonmotive force to phosphorylate ADP forming
ATP.
Per molecule of Glucose
Substrate-level phosphorylation: 2 ATP from glycolysis + 2
ATP (directly GTP) from Krebs cycle
Oxidative phosphorylation:
2 NADH+H+ from glycolysis: 2 × 1.5 ATP (if glycerol
phosphate shuttle transfers hydrogen atoms) or 2 ×
2.5 ATP (malate-aspartate shuttle)
2 NADH+H+ from the oxidative decarboxylation of
pyruvate and 6 from Krebs cycle: 8 × 2.5 ATP 2
FADH2 from the Krebs cycle: 2 × 1.5 ATP
Altogether this gives 4 + 3 (or 5) + 20 + 3 = 30 (or 32) ATP per Glycolysis and the citric acid cycle connect to many other
molecule of glucose. metabolic pathways.
(1) Carbohydrates, fats, and proteins can all be used as
fuel for cellular respiration. Monomers of these
Fermentation and anaerobic respiration molecules enter glycolysis or the citric acid cycle at
 Glycolysis can produce ATP with or without O2 (in various points.
aerobic or anaerobic conditions (2) Glycolysis and the citric acid cycle are catabolic
 Anaerobic respiration uses an electron transport chain funnels through which electrons from all kinds of
with an electron acceptor other than O2 , for example organic molecules flow on their exergonic fall to
sulfate oxygen.
 Fermentation uses phosphorylation instead of an
electron transport chain to generate ATP
Fermentation: consists of glycolysis plus reactions that
regenerate NAD+ , which can be reused by glycolysis. Two
common types:  alcohol fermentation  lactic acid
fermentation
Alchohol fermentation:
 Pyruvate is converted to ethanol in two steps, with the
first releasing CO2
 used in brewing, winemaking, and baking (yeast)

Lactic acid fermentation:

 Pyruvate is reduced to NADH, forming lactate as an
end product, with no release of CO2
 Used to make cheese and yogurt (fungi and bacteria)
•
 Used by human muscle to generate ATP when O2 is
scarce

Fermentation vs Aerobic Respiration

(1) Both use glycolysis to oxidize glucose and other
organic fuels to pyruvate
(2) Have different final electron acceptors: an organic
molecule (such as pyruvate or acetaldehyde) in
fermentation and O2 in cellular respiration
(3) Cellular respiration produces 38 ATP per glucose
molecule; fermentation produces 2 ATP per glucose
molecule

The

catabolism of various molecules from food.