Chapter 10 - Photosynthesis & Respiration
Radiant Energy
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Living organisms ultimately depend upon food produced by photosynthesis.
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Autotrophs have ability to synthesize organic molecule from raw material;
heterotrophs must take in reformed organic molecules.
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Bodies of plants also become fossil fuels used for energy to drive modern
machinery and heat buildings.
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Sunlight
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a. Radiant energy is described by its wavelength; gamma rays are
shortest and radio waves are longest.
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b. The visible spectrum is a narrow band ranging from violet (shortest)
to red (longest).
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c. Energy content is also highest for shorter violet and lowest
for longer red light.
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About 42% of total solar radiation that hits the atmosphere reaches through
to the surface; higher energy wavelengths are screened out by ozone, lower
energy wavelengths are screened out by water vapor and carbon dioxide.
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Life is adapted in vision and photosynthesis to the middle wavelengths.
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Chlorophylls and carotenoids are pigments capable of absorbing portions
of the visible light spectrum.
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Chlorophyll absorbs far less green light; thus green is reflected and leaves
appear green.
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Carotenoids absorb violet-blue-green and reflect yellow-orange; when chlorophyll
breaks down in fall, these pigments remain to give some leaves their fall
color.
Structure and Function of Chloroplasts
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Chloroplasts are organelles found in plant cells that carry on photosynthesis.
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Water is both utilized and produced by photosynthesis.
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A generalized carbohydrate (CH2O) is also produced.
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The oxygen in the O2 produced by photosynthesis comes from the
input of water; this as shown by experiment where heavy oxygen (18C)
in water turns up as the total oxygen produced. The oxygen from CO2
therefore becomes part of the carbohydrate.
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Photosynthesis can also be represented as the reverse of cellular respiration;
water molecules are oxidized and CO2 is reduced.
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Anatomy of Chloroplasts
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a. Most chloroplasts are in leaves.
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b. Mesophyll cells receive water from vessels extending up from roots.
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c. CO2 enters and O2 exits a leaf through small pores.
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d. Chloroplasts are bounded by a double membrane.
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e. Inside these membranes is a large space called the stroma; the
stroma contains an energy-rich solution that reduces carbon dioxide (CO2),
converting it to an organic compound.
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f. Grana are stacks of flattened sacs called thylakoids that
contain the pigment chlorophyll.
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g. Chlorophyll and other pigments in the membranes absorb solar energy
which energizes electrons before reducing CO2 in the stroma.
Solar Energy Capture
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Photosynthesis involves two sets of reactions: the light-dependent reactions
that require light be present, and the light-independent reactions that
can take place in the dark.
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Generally, the light-dependent reactions remove low energy
electrons from water when chlorophyll absorbs energy; these electrons move
down an electron transport system to produce ATP from ADP and (P); energized
electrons are also taken up by NADP+1, which temporarily holds
energy to fuel upcoming CO2 reduction.
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Generally, the light-independent reactions use ATP and NADPH
formed in thylakoids to reduce CO2 in the stroma; the CO2
from the air is fixed by a substrate of the Calvin cycle to produce CH2O.
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The Calvin cycle is named for Melvin Calvin who used radioactive
carbon-14 to label the CO2 to discover the light-independent
reactions.
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Thylakoid membranes contain two light-gathering units, Photosystem I
(PS I) and Photosystem II (PS II), named in the order of their discovery.
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Pigment Complex Molecules
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a. Include chlorophyll a and chlorophyll b, and carotenoids.
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b. Serve as an "antenna" to gather sunlight energy until concentrated in
the reaction-center chlorophyll.
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c. Electrons become so excited that they escape to a nearby electron acceptor
molecule.
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Cyclic Electron Pathway
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a. This pathway begins after the PS I pigment complex absorbs solar energy.
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b. Energized electrons leave the reaction-center chlorophyll of PS I, pass
through the electron transport system (cytochrome system), release energy
used to produce ATP, and then return to the reaction-center again.
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c. This pathway only produces ATP.
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d. Some photosynthetic bacteria only utilize this pathway; it therefore
may have evolved early in the history of life.
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Noncyclic Electron Pathway
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a. Noncyclic electron pathway results in both ATP and NADPH.
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b. Pigment complex of PS II absorbs solar energy.
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c. Excited electrons leave reaction-center chlorophyll a molecule.
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d. PS II takes replacement electrons from water, which splits releasing
oxygen.
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e. High energy electrons that leave PS II are captured by an acceptor molecule.
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f. Electrons pass from one carrier to the next in the electron transport
system as the released energy is used for ATP production.
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g. When PS I pigment complex absorbs solar energy, excited electrons leave
the reaction-center chlorophyll a and are captured by an acceptor
molecule that passes electrons on to NADP+, and NADP+
picks up H+ from stroma to form NADPH.
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h. Results of noncyclic electron flow: water is split, yielding electrons,
oxygen and hydrogen ions; ATP is produced; and NADP+ becomes
NADPH.
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Chemiosmotic ATP Synthesis
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a. The membranous system within the stroma forms flattened sacs called
thylakoids, in some places they are stacked to form grana.
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b. The thylakoid space within acts as a reservoir for hydrogen ions.
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c. Each time a water is split, two H+ remain in the thylakoid
space.
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d. Compared to the stroma, the large number of hydrogen ions in the thylakoid
space create an electrochemical gradient.
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e. The flow of H+ from high to low concentration across the
thylakoid membrane provides energy that allows ATP synthase enzyme to produce
ATP from ADP + (P).
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The Thylakoid Membrane is Organized
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a. Biochemical and structural studies show intact complexes in the thylakoid
membrane.
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b. PS II is the light-gathering complex that splits water and produces
oxygen.
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c. The cytochrome complex transports electrons between PS II and PS I.
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d. PS I is a light-gathering pigment associated with the enzyme that reduces
NADP+ to NADPH.
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e. ATP synthase complex has a H+ channel and a protruding ATP
synthase; H+ flows down this channel and ATP is produced from
ADP + (P).
Carbohydrate Synthesis
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The light-independent reactions are the second stage of photosynthesis.
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a. NADPH and ATP from the light-dependent reactions reduce carbon dioxide
to form a carbohydrate.
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b. Reduction of CO2 within the stroma of the chloroplast occurs
by the Calvin cycle.
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The Calvin Cycle
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a. Summary: carbon dioxide combines with a five-carbon sugar; the six-carbon
molecule breaks down to form two PGA (three-carbon) molecules; PGA is reduced
to PGAL by using NADPH and ATP; regenerating the five carbon sugar. PGAL,
the end product of the Calvin cycle, is converted to many other molecules
(glucose phosphate, fatty acids, amino acids, etc.) especially by algae
and plants.
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b. Details of the Calvin cycle can be divided into CO2 fixation,
CO2 reduction, and regeneration of RuBP.
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c. Fixing Carbon Dioxide
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i. Carbon dioxide combines with RuBP, a five-carbon molecule due to the
enzyme RuBP carboxylase.
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ii. This protein makes up 20-50% of protein content of chloroplasts.
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d. Reducing Carbon Dioxide
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i. The six-carbon molecule from CO2 fixation immediately breaks
down to form two PGA (phosphoglycerate) three-carbon molecules.
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ii. PGA is reduced to PGAL (phosphoglyceraldehyde) by using NADPH and ATP
from the light- dependent reaction in two steps.
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e. Regenerating RuBP
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i. For every three turns of the Calvin cycle, five molecules of PGAL are
used to re-form three molecules of RuBP.
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ii. The net gain of three turns of the Calvin cycle is one PGAL molecule.
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iii. This reaction utilizes some ATP produced by the light-dependent reactions.
Other Aspects of Photosynthesis
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C3 Versus C4 Photosynthesis
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a. C3 photosynthesis is described as above because a C3
molecule is detected immediately following CO2 fixation.
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b. In C4 photosynthesis, a C3 molecule is detected
following CO2 fixation.
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c. C3 plants:
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-include wheat, rice, oats and Kentucky blue grass
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-have mesophyll cells with well-formed chloroplasts, cells are arranged
in parallel layers
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d. C4 plants:
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-include sugarcane, corn and crabgrass
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-CO2 is taken up in mesophyll cells and then a C4
molecule (oxaloacetate) is pumped into the bundle sheath cells where it
releases CO2 to the Calvin cycle
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-have chlorophyll in both the bundle sheath cells and the mesophyll cells
that are arranged around the sheath.
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v. C3 plants have an advantage in moderate climates, C4
plants are more sheltered from drying in hot and dry climates because oxygen
accumulates when stomates close to protect the plant.
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CAM Photosynthesis
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a. CAM plants fix some CO2 at night, forming C4 molecule.
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b. CAM from "crassulacean-acid metabolism"; Crassulaceae is family of warm,
arid region flowers.
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c. With stomates closed in daytime, these plants conserve water.
Aerobic Cellular Respiration
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Cellular respiration includes all metabolic pathways where carbohydrates
and other metabolites are broken down to build up ATP.
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Aerobic cellular respiration includes pathways that require oxygen.
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Breaking glucose (a high-energy molecule) into CO2 and H2O
(low-energy molecules) is an exergonic process.
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Upon breakdown, electrons are removed from glucose and eventually received
by O2.
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Glucose is oxidized and O2 is reduced; glucose breakdown is
therefore an oxidation-reduction reaction.
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The buildup of ATP is an endergonic reaction, it requires energy.
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The breakdown of one glucose results in 36 to 38 ATP molecules being formed;
this is under 40% of the potential energy within a glucose molecule, over
60% is lost as heat.
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The Steps of Aerobic Respiration
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a. Aerobic cellular respiration is a gradual process that prevents energy
loss as heat.
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b. Glycolysis is the breakdown of glucose to two molecules of pyruvate;
occurs outside the mitochondria.
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c. During the transition reaction, pyruvate is oxidized to acetyl CoA and
CO2 is removed; the transition reaction occurs twice per glucose
molecule.
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d. The Krebs cycle is cyclical series of oxidation reactions that give
off CO2 and produce one ATP per cycle; it turns twice per glucose
and produces two ATP.
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e. The electron transport system is a series of carriers that accept electrons
removed from glucose and eventually pass then to oxygen; release of energy
along this electron transport chain results in ATP buildup.
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NAD is a coenzyme of oxidation-reduction.
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a. NAD+ picks up two electrons and one hydrogen ion; the substrate
is oxidized and NAD+ is reduced.
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b. The electrons received by NAD+ are used by the cell to produce
ATP.
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c. Like an enzyme, the coenzyme NAD+ is used over and over again;
only a small amount is therefore present in a cell.
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d. After NAD+ accepts electrons and is reduced to NADH, NADH
passes the electrons to another carrier and becomes oxidized to NAD+
again.
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e. FAD is sometimes used instead of NAD+ to oxidize substrates;
FAD accepts two electrons and two hydrogen ions and becomes FADH2.
Outside the Mitochondria: Glycolysis
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Glycolysis breaks down glucose to two molecules of pyruvate outside
the mitochondria.
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Found in all organisms, glycolysis probably evolved before the Krebs cycle
and electron transport system and probably is why it occurs in the cytoplasm
and does not require oxygen.
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The Energy Investment Steps:
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a. Two ATP are used to activate glucose (a six-carbon molecule).
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b. The resulting molecule is phosphorylated (phosphate groups are added).
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c. The C6 molecule splits into two C3 molecules,
each of which is phosphorylated.
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The Energy Generation Steps:
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a. Oxidation of substrates is carried out by NAD+ twice producing
two NADH.
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b. Energy released allows formation of four ATP by substrate-level phosphorylation.
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c. During substrate-level phosphorylation, a substrate passes a high-energy
phosphate to ADP forming ATP.
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d. Subtracting two ATP used to get the reaction started, there is a net
gain of two ATP.
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Glycolysis is not just an aerobic process but also occurs in anaerobic
fermentation.
Inside the Mitochondria
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The transition reaction, the Krebs cycle and the electron transport system
all take place inside the mitochondria.
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Enzymes for the Krebs cycle are located in the fluid-filled matrix
of the mitochondria.
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Pathways: oxygen and glucose diffuse into cells from bloodstream, pyruvate
(as an end product of glycolysis) diffuses into mitochondria; CO2
and ATP diffuse back out of mitochondria into cytoplasm and CO2
further diffuses back to bloodstream. Water can remain in mitochondria,
in cytoplasm, or enter bloodstream for excretion. ATP remains as a source
of energy for the cell to do work.
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Since most of ATP is produced in mitochondria, mitochondria are often called
the powerhouses of the cell.
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The transition reaction:
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a. Connects glycolysis to Krebs cycle.
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b. Occurs within matrix of mitochondria.
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c. From glucose, two molecules of pyruvate are converted to a two-carbon
acetyl group attached to coenzyme A (CoA).
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d. CO2 is given off.
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The Krebs cycle:
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a. Occurs in matrix of mitochondria.
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b. Takes up acetyl group (acetyl CoA) from transition reaction and oxidizes
it to two CO2 molecules.
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c. During the process, most of the electrons are accepted by NAD+
but in one instance they are taken by FAD.
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The Electron Transport System:
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a. Is located on cristae (projections of inner membrane).
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b. Consists of a series of carriers that pass electrons; some of protein
carriers are cytochrome molecules so system is also called the cytochrome
system.
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c. Accounts for most of the ATP produced.
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d. When NADH gives up electrons, it becomes NAD+ while an electron
carrier gains electrons and is reduced.
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e. Each sequential carrier becomes reduced and then oxidized as electrons
move down the system.
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f. As electrons pass from carrier to carrier, energy is released and used
to form ATP molecules.
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g. When NADH delivers electrons to the first carrier, enough energy is
released by time electrons reach the O2 to produce three ATPs.
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h. When FADH2 delivers electrons to electron transport system,
only two ATPs result.
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The Cristae:
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a. Contain carriers of electron transport system arranged in a functional
manner.
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b. The carriers use released energy to pump the surplus hydrogen ions carried
by NADH and FADH2 into intermembrane space of mitochondria.
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c. Cristae also contain ATP synthase complex:
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i. Hydrogen ions flow from high to low concentration from intermembrane
space across to matrix.
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ii. Resulting H+ flow drive enzyme ATP synthase to synthesize
ATP from ADP + (P) .
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iii. Process is called chemiosmosis because ATP production is tied
to an electrochemical gradient.
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Calculating Energy Yield from Glucose Metabolism (Fig. 7.9)
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a. Per glucose, four ATP are formed by substrate-level phosphorylation,
two during glycolysis and two during two turns of Krebs cycle.
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b. Per glucose, ten NADH and two FADH2 take electrons to the
electron transport system.
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c. For each NADH formed inside the mitochondria by Krebs cycle, three ATP
result; for each FADH2, only two ATP are produced.
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d. The glycolytic pathway outside the mitochondria produces only two ATP
when the electrons are shuttled to the electron transport system inside
the mitochondrion.
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How Efficient is Aerobic Respiration?
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a. Difference in energy content between glucose and O2, and
products CO2 and H2O is 686 kilocalories.
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b. The ATP third phosphate bond has energy content of 7.3 kilocalories,
36 ATP are produced per glucose breakdown totaling 263 kilocalories.
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c. Efficiency is 263/686 or 39%.
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d. Sixty-one percent is lost as heat; in birds and mammals, this heat assists
in maintaining body temperature.
Metabolic Pool and Biosynthesis
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Catabolic Reactions
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a. Catabolic reactions break down molecules, driving anabolic reactions
which synthesize molecules.
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b. Aerobic cellular respiration has already shown the catabolism of glucose.
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c. Fats are broken down:
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i. Glycerol is converted to PGAL in glycolytic pathway.
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ii. Fatty acids are converted to acetyl-CoA, which enters the Krebs cycle.
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iii. Fats are an efficient form of stored energy, an 18-carbon fatty acid
results in nine acetyl-CoA molecules that produce 216 ATP molecules via
respiration.
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d. Amino acids are broken down:
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i. Amino acids must undergo deamination (removal of amino group; this occurs
in the liver).
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ii. The amino group becomes ammonia (NH3) which becomes urea
via the urea cycle.
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iii. The carbon skeleton produced by deamination can then enter Krebs cycle
depending on the number of carbons left after deamination.
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Anabolic Reactions
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a. Anabolic reactions use ATP produced during catabolism to synthesize
molecules.
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b. The cell's metabolic pool consists of large molecules that can
be converted to other molecules without buildup from small molecular units;
for example, PGAL can be converted to glycerol and acetyl groups can be
joined to form fatty acids.
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c. Some metabolites are converted to amino acids through transamination,
transfer of an amino group from one amino acid to another.
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d. Plants can synthesize all the amino acids they need.
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e. Animals lack some enzymes to synthesize some amino acids; therefore
must secure these from diet; lack of sufficient amino acids in diet results
in protein deficiency disease.
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f. Humans can synthesize 11 of 20 common, necessary amino acids; therefore
must secure other nine from diet.
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g. All the reactions involved in cellular respiration are part of a metabolic
pool in which one type of molecule can be converted to another in catabolism
or anabolism.
Fermentation
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Cellular respiration includes both aerobic cellular respiration and fermentation.
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Fermentation is a series of enzymatic reactions where glucose is incompletely
metabolized into lactate or CO2 and alcohol.
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Fermentation is anaerobic; it does not require O2.
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During fermentation, there is a net gain of only two ATPs..
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Fermentation consists of glycolysis plus the reduction of pyruvate.
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a. Pathway is anaerobic because after NADH transfers its electrons to pyruvate,
it is free to return and pick up more electrons during earlier glycolysis
reactions.
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b. Lactic acid bacteria reduce pyruvate to lactic acid; other bacteria
produce important industrial chemicals: isopropanol, butyric acid, propionic
acid, and acetic acid.
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c. Yeasts represent organisms that reduce pyruvate to alcohol and carbon
dioxide.
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d. Animals reduce pyruvate to lactate when pyruvate is produced faster
than it can be oxidized.
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Advantages and Disadvantages of Fermentation
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a. In spite of low yield and toxicity, fermentation provides rapid burst
of ATP.
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b. In muscle cells of animals, fermentation provides two ATP molecules
while oxygen is temporarily in limited supply while exercising.
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c. When blood cannot carry away lactate fast enough, lactate builds up
in muscles, changes the pH, and causes muscle fatigue and preventing contraction.
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d. When we stop exercising, we are in oxygen debt and continue breathing
heavily until lactate is transported to the liver to be reconverted to
pyruvate, or pyruvate is respired completely.
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e. Efficiency of Fermentation
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i. Energy content of last ATP bond is 7.3 kilocalories and two are produced
per glucose fermentation for a total of 14.6 kilocalories.
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ii. Complete glucose breakdown to CO2 and H2O yields
686 kilocalories.
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iii. Efficiency of fermentation is therefore 14.6/686 or 2.1%.
Photosynthesis Versus Aerobic Cellular Respiration
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Both plants and animals carry out respiration, only plants carry on photosynthesis.
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Cell organelle for aerobic respiration is the mitochondrion; the organelle
for photosynthesis is the chloroplast.
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Overall equation for aerobic cellular respiration is the opposite of that
for photosynthesis:
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ATP energy + 6 CO2 + 6 H2O --> C6H12O6
+ 6 O2 (cellular respiration)
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solar energy + 6 CO2 + 6 H2O <-- C6H12O6
+ 6 O2 (photosynthesis)
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Cellular respiration requires oxygen, breaks down carbon dioxide, and occurs
in both plants and animals, day or night.
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Photosynthesis requires carbon dioxide, releases oxygen, involves reduction,
and stores energy.
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Both photosynthesis and cellular respiration are metabolic pathways.
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Both make use of an electron transport system located in a membrane to
produce ATP.
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Both use a hydrogen carrier (NAD+ in respiration, NADP+
in photosynthesis).
Photosynthesis occurs only during daytime in plants; during daylight hours,
the rate of photosynthesis exceeds the rate of aerobic cellular respiration.