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Chapter 7: Energy and Metabolism

• Biological Work
o Energy: capacity to do work
 Express energy in kilojoules (units of work)
 Express energy in kilocalories (units of heat energy; thermal energy going from high temp. to low temp.)
 1 kcal = 4.184 kJ
o Kinetic energy: energy of motion
o Potential energy: capacity to do work as a result of position or state
o Chemical energy: potential energy stored in chemical bonds
o Mechanical energy: performs work by moving matter
• The Laws of Thermodynamics
o Thermodynamics: study of energy and its transformations
o Closed system: does not exchange energy with its surroundings
o Open system: exchanges energy with its surroundings (ie: biological systems)
o First law of thermodynamics: energy cannot be created or destroyed
 Organisms must capture energy from the environment and transform it to a form that can be used for biological work
o Second law of thermodynamics: when energy is converted from one form to another, some is converted into heat that disperses
into surroundings
o Heat: kinetic energy of randomly moving particles
 Total amount of energy that is available to do work is decreasing over time
o Entropy (S): measure of disorganized energy (unusable energy)
 Total entropy always increases over time
• Energy and Metabolism
o Metabolism: sum of all the chemical activities taking place in an organism
 Anabolism: various pathway sin which complex molecules are synthesized from simpler substances
 Catabolism: pathways in which larger molecules are broken down into smaller ones
o Bond energy: energy required to break that bond
o Enthalpy (H): total potential energy of the system (total energy of a system)
o Free energy (G): amount of energy available to do work under the conditions of a biochemical reaction (usable energy)
 H=G+TS [enthalpy = free energy + absolute temperature of system (K) * entropy]
 As entropy increases, amount of free energy decreases: ΔG=ΔH-TΔS
 As temperature increases, increase in entropy
o Exergonic reaction: releases energy and is spontaneous or a “downhill” reaction from high to low G (Gi>Gf) so G is (-)

 Free energy decreases during an exergonic reaction. Free energy increases during an endergonic reaction
o Endergonic reaction: gain of free energy (Gi<Gf) so G is (+)
 Must occur in a way that energy can be supplied from surroundings
o Concentration gradient: region of higher concentration and another region of lower concentration; orderly state; PE
o Free energy depends on concentrations of both reactants and products
o Dynamic equilibrium: rate of reverse reaction equals rate of forward reaction
o Coupled reactions exergonic reaction provides energy required to drive endergonic reaction
 Endergonic reaction can only proceed if it absorbs free energy released by exergonic reaction
 Product>reaction is endergonic reaction thus ∆G is (+), is not spontaneous, does not take place w/o energy source
 Reactant>product is exergonic reaction thus ∆G is (-), is spontaneous
• ATP, the Energy Currency of the Cell
o Adenosine triphosphate (ATP): holds readily available energy for very short periods; cell’s energy source like cash
 Nucleotide consisting of: adenine (nitrogen-containing organic base), ribose (5 C sugar), 3 phosphate (PO4) groups
o ATP donates energy through transfer of phosphate group
 Adenosine diphosphate (ADP): left when terminal phosphate is removed from ATP
 Inorganic phosphate (Pi): if phosphate group is not transferred to another molecule
 Exergonic reaction with a large -∆G
 Phosphorylation reaction: phosphate group is transferred to some other compound
o Adding phosphate group to AMP creates ADP and adding phosphate to ADP creates ATP
o ATP is link between exergonic reaction which is component of catabolic pathways
o ATP is link between endergonic reaction which is component of anabolic pathways
o Cell maintains ration of ATP to ADP far from equilibrium point, typically 10 ATP per ADP
 Makes hydrolysis reaction more strongly exergonic and more able to drive endergonic reactions to which it is coupled
• Energy Transfer in Redox Reactions
o Redox reaction: every oxidation reaction is accompanied by reduction reaction (OIL RIG)
 Oxidation reaction: substance loses electrons
 Reduction reaction: substance gains electrons
 Substance that becomes oxidized gives up energy as it releases electrons
 Substance that becomes reduced receives energy as it gains electrons
 Redox reactions usually involve transfer of hydrogen atom rather than just an electron
o When an electron is removed from organic compound, takes some energy stored in chemical bond that it was a part of. That
electron along with its energy is transferred to an acceptor molecule
o Electron progressively loses free energy as it is transferred from one acceptor to another
o Nicotinamide adenine dinucleotide (NAD+): frequently encountered acceptor molecules; when reduced it temporarily stores large
amounts of free energy
 NADH is produced when NAD+ + H2 thus NAD+ is oxidized and NADH is reduced
o Nicotinamide adenine dinucleotide phosphate (NADP+): hydrogen acceptor that is chemically similar to NAD+ but has extra PO4

 NADPH: reduced form of NADP+; not involved in ATP synthesis; used to provide energy for certain reactions
o Flavin adenine dinucleotide (FAD): nucleotide that accepts hydrogen atoms and their electrons
 FADH2: reduced form of FAD
o Cytochromes: proteins that contain iron; iron component accepts e- from H atoms and then transfers e- to other compound
o NAD+, NADP+, FAD, cytochromes are electron transfer agents
 Exists in reduced state which has more free energy or oxidized state which has less free energy
• Enzymes
o Enzymes: biological catalysts that increase the speed of chemical reactions without being consumed by the reaction
 Catalase: highest catalytic rate; protects cells by destroying hydrogen peroxide (H2O2)
o Energy of activation (EA) or activation energy: energy required to break existing bonds and begin reaction
o Enzyme lowers activation energy thus allowing the reaction to proceed faster
 Enzyme has no effect on overall free-energy change
o Substrate: substance on which enzyme forms unstable intermediate complex with
o Active site: region to which the substrate binds, to form enzyme-substrate complex (ES complex)
 Grooves or cavities in enzyme formed by amino acid side chains; located close to surface
o Induced fit: change in shape of enzyme from binding of substrate to enzyme
o Shape of substrate changes slightly which distorts its chemical bonds facilitating breakage of old bonds and formation of new ones;
thus substrate is now a product which diffuses away from enzyme and enzyme is free to catalyze other reactions
o Enzymes are specific in bonding to a certain shape or to a certain chemical bond
o Cofactors of enzyme: separately two have no catalytic activity but together it allows the enzyme to function
 Apoenzyme: protein component of enzyme
 Cofactor: chemical component that is either inorganic or organic
o Coenzyme: organic, nonpolypeptide compound that binds to apoenzyme and serves as cofactor; transfers e-
o Coenzyme A: involved in transfer of groups derived from organic acids
o Enzymes work best under certain optimal conditions: temperature, pH, ion concentration; any deviation from optimal condition
adversely affects enzyme activity
 Temperature: 35˚C-40˚C; as T increases, molecular collisions inc.; high T denatures enzymes so unable to metabolize
 pH: 6 – 8; when very acidic or basic, most enzymes become inactive and irreversibly denatured
o Metabolic pathway: series of chemical reactions in which product of one reaction becomes substrate of next reaction
o If pH and T are kept constant, concentration of substrate or enzyme affects rate of reaction
 If excess of substrate present, enzyme concentration is rate-limiting factor; initial rate is proportional to enzyme conc.
 If enzyme conc. kept constant, rate of enzymatic reaction is proportional to substrate conc.
 Substrate conc. is rate limiting factor at lower concentrations, rate of reaction is proportional to substrate conc.
 At higher substrate conc. enzyme molecules are saturated with substrate (substrate bound to all available active sites of
enzyme molecules); increasing substrate conc. does not increase net reaction rate
o Feedback inhibition: formation of product inhibits an earlier reaction in sequence
o Enzymatic control: in inactive form, active sites misshaped; factors influencing are pH, conc. of ions, addition of PO4 to AA
o Allosteric site: receptor site other than active site; when substrate binds, active site changes, modifying enzyme’s activity
o Allosteric regulators: substances that affect enzyme activity by binding to allosteric sites; some keep enzyme inactive
o Reversible inhibition: when inhibitor forms weak chemical bonds with enzyme
 Competitive inhibition: inhibitor competes with normal substrate for binding to active site
• Fits into active site temporarily, thus not allowing actual substrate to bind
• Increasing substrate concentration reverses competitive inhibition
 Noncompetitive inhibition: inhibitor binds to allosteric site
• Inactivates enzyme by altering shape so active site cannot bind with substrate
 Irreversible inhibition: inhibitor permanently inactivates or destroys an enzyme when the inhibitor combines with one of
enzyme’s functional groups, either at active site or allosteric site
Chapter 8: How Cells Make ATP: Energy-Releasing Pathways
• Redox Reactions
o Aerobic respiration: form of cellular respiration requiring molecular oxygen (O2)
 Nutrients are catabolized to CO2 and H2O
 Used to obtain energy from glucose
• Glucose becomes oxidized and oxygen becomes reduced
• C6H12O6 + 6 O2 + 6 H2O ( 6 CO2 + 12 H2O + energy (in chemical bonds of ATP) [oxidation and reduction]
• The Four Stages of Aerobic Respiration
o Glycolysis (takes place in cytosol)
 Six carbon glucose molecule is converted to two three carbon molecules of pyruvate
 C6H12O6 + 2 ATP + 2 ADP + 2 Pi + 2 NAD+ ( 2 pyruvate + 4 ATP + 2 NADH + H2O
o Formation of acetyl coenzyme A (takes place in mitochondria)
 Each pyruvate enters a mitochondrion and is oxidized to a two-carbon group (acetate) that combines with coenzyme A, forming
acetyl conenzyme A
 NADPH is produced, CO2 is released as waste product
 2 pyruvate + 2 coenzyme A + 2 NAD+  2 acetyl CoA + 2 CO2 + 2 NADH
o The citric acid cycle (takes place in mitochondria)
 Acetate group of acetyl coenzyme A combines with four carbon molecule (oxaloacetate) to form six carbon molecule (citrate)
 Citrate is recycled to oxaloacetate and CO2 is released as waste product. Energy captured as ATP and reduced NADH and
FADH2
 2 acetyl CoA + 6 NAD+ + 2 FAD + 2 ADP + 2 Pi + 2 H2O  4 CO2 + 6 NADH + 2 FADH2 + 2 ATP + 2 CoA
o Electron transport and chemiosmosis (takes place in mitochondria)
 Electrons removed from glucose during preceding stages are transferred from NADH and FADH2 to chain of electron acceptor
compounds
 As electrons are passed from one electron acceptor to another, some of energy is used to transport hydrogen ions across inner
mitochondrial membrane forming proton gradient
 In process known as chemiosmosis, energy of proton gradient used to produce ATP
 NADH + 3 ADP + 3 Pi + ½ O2  NAD+ + 3 ATP + H2O
FADH2 + 2 ADP + 2 Pi + ½ O2  FAD+ + 2 ATP + H2O
o Types of reactions involved in aerobic respirations are:
 Dehydrogenations: two hydrogen atoms are removed from substrate and transferred to NAD+ or FAD
 Decarboxylations: part of carboxyl group (–COOH) is removed from substrate as molecule of CO2
 Preparation reactions: molecules undergo rearrangements and other changes that can undergo further dehydrogenation or
decarboxylations
o Glycolysis: sugar splitting; sugar glucose is metabolized
 Does not require oxygen and proceeds under aerobic or anaerobic conditions
 Some energy is captured, net yield of two ATP and two NADH molecules
 Take place in cytosol where ADP, NAD+, Pi float freely and used as necessary
 Endergonic reaction requiring ATP is phase 1
• Energy investment phase: broken down in 2 phosphorlyation reactions: PO4 is transferred from ATP to sugar
• Resulting sugar (fructose-1,6-biphosphate) is less stable and is broken enzymatically into two three carbon molecules:
dihydroxyacetone phosphate and glyceraldehyde-3-phosphate (G3P)
o Dihydroxyacetone phosphate is enzymatically converted to G2P so products are 2 G3P : 1 glucose
o Glucose + 2 ATP  2 G3P + 2 ADP
 Exergonic reaction yielding ATP and NADH is phase 2
• Energy capture phase: G3P converted to pyruvate
• G3P oxidized by removal of 2 electrons, immediately combining with NAD+ (hydrogen carrier molecule)
o NAD+ (oxidized) + 2 H (from G3P)  NADH (reduced) + H+

• Substrate-level phosphorylation: ATP forms when PO4 is transferred to ADP from phosphorylated intermediate
 In EIP, two molecules of ATP consumed but in ECP four molecules of ATP produced; glycolysis yields net energy profit of two
ATPs per glucose
• 2 G3P + 2 NAD+ + 4 ADP  2 pyruvate + 2 NADH + 4 ATP
o Acetyl coenzyme A (acetyl CoA): pyruvate molecules formed in glycolysis enter mitochondria, converting into this
 Occur in cyosol of aerobic prokaryotes
 Oxidative decarboxylation: pyruvate undergoes this process
• Carboxyl is removed as carbon dioxide, which diffuses out of cell
• Remaining two carbon fragment becomes oxidized and NAD+ accepts electrons removed during oxidation
• Oxidized two carbon fragment, an acetyl group, becomes attached to coenzyme A, yielding acetyl
• 2 pyruvate + 2 NAD+ + 2 CoA  2 acetyl CoA + 2 NADH + 2 CO2
o Four NADH molecules formed: 2 during glycolysis, 2 during formation of acetyl CoA from pyruvate
o Citric acid cycle (tricarboxylic acid (TCA) cycle or Krebs cycle): takes place in mitochondria
 Acetyl CoA transfers two carbon acetyl group to four carbon acceptor compound oxaloacetate forming citrate (six carbon
compound)
• Oxaloacetate + acetyl CoA ( citrate + CoA
 Citrate loses first then second carboxyl group as CO2
 One ATP formed per acetyl group by substrate-level phosphorylation; energy is transferred as NAD+ forming NADH
 For each acetyl group that enters the citric acid cycle, three molecules of NADH are produced
 Electrons are transferred to electron acceptor FAD, forming FADH2
 Two molecules of CO2 and about eight H are removed, forming three NADH and one FADH2
 Four carbon oxaloacetate has been regenerated and cycle continues
o Because two acetyl CoA are produced from each glucose, two cycles required per glucose; after two turns, original glucose has lost
all its carbons
o Citric acid cycle yields four CO2, six NADH, two FADH2, and two ATPs per glucose molecule
o Four molecules of ATP formed: two during glycolysis and two during citric acid cycle
o Oxidative phosphorylation: electrons passed along series of exergonic redox reactions, energy is used for ATP (endergonic),
because ATP synthesis is coupled to redox reactions in electron transport chain
o Electron transport chain: high energy electrons of hydrogen atoms are shuttled from one acceptor to another
 Series of electron carriers embedded in inner mitochondrial membrane of eukaryotes and in plasma membrane of aerobic
prokaryotes
 Each acceptor molecule becomes reduced as it accepts electrons and oxidized as it gives it up
 Electrons start with high energy content and lose some along the way
 Complex I: NADH-ubiquinone oxidoreductase; accepts electrons from NADH produced in glycolysis, formation of acetyl CoA,
and citric acid cycle
 Complex II: succinate-ubiqunone oxidoreuctase; accepts electrons from FADH2 produced in citric acid cycle
 Complex III: ubiquinone-cytochrome c oxidoreductase; product from I and II (reduced ubiquinone) is substrate; accepts
electrons from reduced ubiquinone and passes to cytochrome c
 Complex IV: cytochrome c oxidase; ccepts electrons from cytochrome c and uses electrons to reduce O and form water
• If no oxygen available to accept, then last cytochrome retains its electrons; each acceptor molecule in chain retains its
electrons (remains in reduced state), and entire chain is blocked all the way back to NADH
 No additional ATP produced by electron transport chain
o Chemiosmosis allows exergonic redox reactions to drive endergonic reaction in which ATP is produced by phosphorylating ADP
whereas in photosynthesis, ATP is produced by a comparable process
o ATPs: 2 from glycolysis, 2 from citric acid cycle, 32 – 34 from electron transport and chemiosmosis = 36 – 38 ATPs
mostly through oxidative phosphorylation (electron transport chain and chemiosmosis)
o Mitochondrial shuttle system
 In liver, kidney, heart: transfers electrons from NADH through inner mitochondrial membrane to NAD+ to ETC
 In skeletal muscle, brain: accepted by ubiquinone
o Rate of aerobic respiration regulated by how much ADP and phosphate are available
• Energy Yield of Nutrients Other Than Glucose
o Deamination: amino acids metabolized by amino group being first removed
o Β-oxidation: fatty acids are converted to acetyl CoA in the mitochondrial matrix then enters CAC
• Anaerobic Respiration and Fermentation
o Anaerobic respiration: does not use oxygen as final electron acceptor
 Electrons transferred from glucose to NADH then down ETC that is coupled to ATP synthesis by chemiosmosis but inorganic
substance replaces O as terminal electron acceptor; end product is CO2, Pi, ATP
 Nitrogen cycle: C6H12O6 + 12 KNO3  6 CO2 + 6 H2O + 12 KNO2 + energy (in chemical bonds of ATP)
 Fermentation: anaerobic pathway that does not involve ETC; only two ATPs are formed per glucose
o Facultative anaerobes: carry out aerobic respiration when oxygen is available
o Alcohol fermentation: carry out anaerobic respiration when oxygen is unavailable
 Decarboxylate pyruvate, release CO2 and form two carbon compound acetaldehyde
 C6H12O6  2 CO2 + 2 ethyl alcohol + energy (2 ATP)
o Lactate fermentation: NADH transfer H to pyruvate reducing it to lactate
 C6H12O6  2 lactate + energy (2 ATP)
Chapter 9: Photosynthesis: Capturing Energy
• Light
o Wavelength: distance from one wave peak to next
 Violet has shortest wavelength and red has longest
o Photons: small particles or packets of energy composing light
 Energy of photon is inversely proportional to wavelength
• Short wavelength has more energy per photon than long wavelength light
o When molecule absorbs photon of light energy, two things may happen
 Atom may return to ground state (condition in which all electrons are in normal lowest energy levels)
 Energy at ground state dissipates as heat or emission of light (fluorescence)
 Or energized electron may leave atom and be accepted by electron acceptor molecule which becomes reduced
• Chloroplasts
o Chlorophyll is confined to chloroplasts
o Chloroplasts are mainly in mesophyll (layer with many air spaces and high concentration of water vapor)
o Interior of leaf exchanges gases with outside through stomata (microscopic pores)
o Enclosed by outer and inner membranes
o Inner membrane encloses stroma (fluid-filled region containing most of enzymes required to produce carbohydrate mol.
o Thylakoids: suspended in stroma, forms interconnected set of flat, disclike sacs
 Thylakoid lumen: fluid-filled interior space enclosed by thylakoid membrane
 Grana: stacks of thylakoid sacs
o Chlorophyll: main pigment of photosynthesis, absorbs light primarily in blue and red regions of visible spectrum
 Contains complex ring structure porphyrin ring (joined smaller rings composed of C and N; absorbs light energy)
 Or contains long hydrocarbon side chain that makes molecule nonpolar and anchors chlorophyll in membrane
 All chlorophyll molecules in thylakoid membrane are associated with specific chlorophyll-binding proteins
 Chlorophyll a: pigment initiates light-dependent reactions of photosynthesis; contains methyl group (CH3)
 Chlorophyll b: accessory pigment that also participates in photosynthesis; contains carbonyl group (CHO)
 Carotenoids: yellow and orange accessory photosynthetic pigment absorbing different wavelengths of light
o Absorption spectrum: plot of absorption of light of different wavelengths
o Action spectrum: graph of relative effectives of different wavelengths of light; rate v. wavelength
o Chlorophyll in chloroplasts is responsible for photosynthesis
• Overview of Photsynthesis
o Cell uses light energy captured by chlorophyll to power synthesis of carbohydrates
 6 CO2 + 12 H2O  C6H12O6 + 6 O2 + 6 H2O
o Reactions of photosynthesis divided into two phases:
 Light-dependent reaction: photo part of photosynthesis in thylakoids
• Begin as chlorophyll captures light causing electrons to move to higher energy state then transferred to acceptor molecule
and replace by electron from water then water splits and O is released
• Some energy used to phosphorylate ADP forming ATP
• Coenzyme NADP+ is reduced forming NADPH
 Carbon fixation reaction: synthesis part of photosynthesis in stroma
• ATP and NADPH from light-dependent reaction are used to transfer chemical energy
• Carbon fixation: “fix” C from CO2 to existing skeletons of organic molecules; no direct requirement for light
• The Light-Dependent Reactions
o 12 H2O + 12 NADP+ + 18 ADP + 18 Pi  6 O2 + 12 NADPH + 18 ATP
o Antenna complexes: arrangement of chlorophyll, accessory pigments, and pigment binding proteins into light gathering units in
thylakoid
 Absorbs light and transfers to reaction center (consists of chlorophyll molecules and proteins)
o Light then converted to chemical energy in reaction centers by a series of electron transfer reactions
o Two types of photosynthetic units
 Photosystem I: consists of pair of chlorophyll a molecules with absorption peak at 700 nm and referred to as P700
 Photosystem II: made up of pair of chlorophyll a molecules with absorption peak of 680 nm and referred to as P680
o When pigment molecule absorbs light energy, energy Is passed from one pigment to another with antenna complex until it reaches
reaction center. When reaches P700 or P680, electron is raised to higher energy level
o Noncyclic electron transport: in photosynthesis, linear flow of electrons, produced by photolysis of water, through photosystems I
and II; results in formation of ATP (by chemiosmosis) NADPH, and O2
 Pigment in antenna complex associated with photosystem I absorbs photon of light; energy is transferred from one pigment
molecule to another until reaches reaction center, where it excites an electron in a molecule of P700. Energized electron is
transferred to primary electron acceptor then electron goes through electron transport chain until passed to ferredoxin (iron-
containing protein). Ferredoxin transfers electron to NADP+ in presence of enzyme ferredoxin-NADP+ reductase. When NADP+
accepts two electrons, they unite with H+, thus reduced form is NADPH which is released into stroma. P700 becomes positively
charged when giving up electron to primary electron acceptor; missing electron is replaced by one donated by photosystem II
 Photosystem II is activated when pigment molecule in antenna complex absorbs photon of light energy. Energy is transferred
to reaction center, where it causes electron in P680 to move to higher energy level. Energized electron is accepted by primary
electron acceptor and then passes along electron transport chain until donated to P700
 Electron donated in ETC replaced by photolysis (light splitting) of water
• P680 molecule giving up energized electron to primary electron acceptor is positively charged (P680+). P680+ is oxidizing
agent pulling away O from water. Water broken into 2 electrons, 2 protons, 2 O. Each electron donated to P680+ which
then loses positive charge; proteins are released into thylakoid lymen
• Light splits water indirectly by causing P680 to become oxidized.
 NET provides continuous supply of replacements for energized electrons that were given up by P700
o Cyclic electron transport: only photosystem I is involved; simple light-dependent reaction
 Energized electrons that originate from P700 at reaction center eventually return to P700
o Photophosphorylation: synthesis of ATP is coupled to transport of electrons that have been energized by protons of light
o ATP synthase: thylakoid membrane is impermeable to H+ except through certain channels formed by enzyme
• The Carbon Fixation Reactions
o Energy of ATP and NADPH is used in formation of organic molecules from CO2
o 12 NADPH + 18 ATP + 6 CO2  C6H12O6 + 12 NADP+ + 18 ADP + 18 Pi + 6 H2O
o Occurs in stroma through Calvin Cycle in three phases:
 CO2 uptake: single reaction in which molecule of CO2 reacts with ribulose biphosphate (RuBP; phosphorylated 5 C)
• Catalyzed by ribulose biphosphate carboxylase / oxygenase or rubisco
• Product is unstable, 6 C intermediate which immediately breaks down into 2 phosphoglycerate (PGA) with 3 C each
• C that was originally part of CO2 is now part of carbon skeleton, carbon is “fixed”
• Calvin cycle also known as C3 pathway because product of intial carbon fixation reaction is 3 C; plans is C3 plants
 Carbon reduction: energy and reducing power from ATP and NADPH are used to convert PGA to G3P
• Reaction of two G3P molecules is exergonic and leads to formation of glucose or fructose
 RuBP regeneration: series of 10 reactions
• 30 carbons are rearranged into six molecules of ribulose phosphate each of which becomes phosphorylated by ATP to
produce RuBP, five carbon compound with which cycle started
o Inputs required are six molecules of CO2, phosphates transferred from ATP, and electrons from NADPH
o Outputs are six carbons from CO2 are accounted for by harvest of hexose molecule; remaining G3P are used to synthesize RuBP
molecules with which more CO2 molecules may combine
o Photorespiration: occurs in presence of light, requires oxygen, produces CO2 and H2O but does not produce ATP and reduces
photosynthetic efficiency because it removes some of intermediates used in Calvin Cycle
o C4 plants first fix CO2 into four carbon compound, oxaloacetate; step precedes Calvin Cycle
o CAM plants initially fix carbon at night through formation of oxaloacetate; step precedes Calvin Cycle
o C4 pathway: CO2 is fixed through formation of oxaloacetate occurs before C3 pathway in different cells
 Bundle sheath cells: encircle veins of leaf
 Pathway occurs in mesophyll cells whereas Calvin Cycle occurs in bundle sheath cells
 Has phosphoenolpyruvate (PEP) carboxylase enzyme that has an extremely high affinity thus binding effectively
 Previous step formed oxaloacetate which is converted to malate which then passes onto chloroplasts within bundle sheath cells
where different enzyme catalyzes decarboxylation of malate to yield pyruvate and CO2
 CO2 then combines with RuBP to go through Calvin Cycle. Pyruvate formed returns to mesophyll cell
o Crassulacean acid metabolism (CAM) pathway: plants living in dry conditions have a special carbon fixation pathway
 Open stomata at night, admitting CO2, while minimizing water loss
 Use enzyme PEP carboxylase to fix CO2 forming oxaloacetate which is converted to malate and stored in cell vacuoles
 During day, stomata are closed and gas exchange cannot occur between plant and atmosphere, CO2 is removed from malate
by decarboxylation reaction
 Now CO2 is available within leaf tissue to be fixed into sugar by Calvin Cycle
o Important differences between CAM and C4 are:
 C4 and C3 occur in different location within leaf of C4 plant
 CAM and C3 pathways occur at different times within same cell of CAM plant
 C4 promotes rapid growth, CAM is successful adaptation to dry conditions (can survive in desert)
• Metabolic Diversity
o Photoautotrophs: land plants, algae, and certain prokaryotes
 Phototrophs: use light energy to make ATP and NADPH which temporarily hold chemical energy but are unstable and cannot be
stockpiled in cell
 Autotrophs: chemical energy of ATP and NADPH then drives carbon fixation, anabolic pathway in which stable organic
molecules are synthesized from CO2 and water
o Chemoheterotrophs: animals, fungi, and most bacteria
 Chemotrophs: obtain energy from chemicals, typically redox reactions
 Heterotrophs: cannot fix carbon, use organic molecules produced by other organisms as building blocks from which they
synthesize the carbon compounds they need
o Photoheterotrophs: able to use light energy but unable to carry out C fixation and must obtain C from organic compounds
o Chemoautotrophs: obtain energy from oxidation of reduced inorganic molecules
• Photosynthesis in Plants and in the Environment
o By fixing carbon, photoautotrophs are ultimate source of all organic molecules used as energy and carbon sources
o Photolysis of water by photosystem II releases O2 that all aerobic organisms require for aerobic respiration
Chapter 10: Chromosomes, Mitosis, and Meiosis
• Eukaryotic Chromosomes
o Chromosomes: major carriers of genetic information in eukaryotes which lie within nucleus
 Chromatin: material consisting of DNA and associated proteins; makes up chromosomes
o Genome: numerous amount of genes in an organism
o Histones: facilitates chromosome packaging; positive charge because high proportion of amino acids with basic side chains
 Associates with DNA which has negative charge because of phosphate groups to form nucleosomes
• Nucleosome: beadlike structure with 146 base pairs of DNA wrapped around a disc-shaped core of 8 histone molecules
• Scaffolding proteins: nonhistone proteins that help maintain chromosome structure
o Condensin: group of proteins required for chromosome compaction; binds to DNA and wraps it into coiled loops that are compacted
into a mitotic or meiotic chromosome
• The Cell Cycle and Mitosis
o Cell cycle: the stages through which a cell passes from one cell division to the next
o The cell cycle consists of two main phases, interphase and M phase.
 M phase involves two main processes, mitosis and cytokinesis. Mitosis, a process involving the nucleus, ensures that each new
receives the same number and types of chromosomes as were present in the original nucleus. Cytokinesis, which generally
begins before mitosis is complete, is the division of the cell cytoplasm to form two cells.
 Interphase: the time when no cell division is occurring.
• G1 phase – growth and normal metabolism take place. Typically the longest phase.
• S phase – DNA replicates and histone proteins are synthesized so that the cell can make duplicate copies of its
chromosomes.
• G2 phase – increased protein synthesis
o Mitosis, the nuclear division that produces two nuclei containing chromosomes identical to the parental nucleus.
 Prophase: Chromosome compaction, when the long chromatin fibers begin a coiling process that makes them shorter and
thicker. The chromatin can then be distributed to the daughter cells with less likelihood of tangling. After compactin, the
chromatin is referred to as chromosomes. It is now apparent that each chromosome was duplicated and consists of a pair of
sister chromatids, which contain identical, double-stranded DNA sequences. Each chromatid includes a constricted region
called the centromere. Sister chromatids are physically linked by a ring-shaped protein complex called cohesin. Cohesions
extend along the length of the sister chromatid arms and are particularly concentrated at the centromere. Kinetochore, a
multiprotein complex to which microtubules can bind. Animal cells have a pair of centrioles in the middle of each microtubule-
organizing center. microtubules radiate from each pole forming the mitotic spindle, a structure that separates the duplicated
chromosomes during anaphase.
 Metaphase: all the cell’s chromosomes align at the cell’s metaphase plane. Each chromatid is completely condensed and
appears thick and distinct.
 Anaphase: sister chromatids separate. Once chromatids are no longer attached to their duplicates, each chromatid is called a
chromosome. The now-separated chromosomes move to opposite poles, using the spindle mircrotubules as tracks.
 Telophase: chromosomes decondense by partially uncoiling. A new nuclear envelope forms around each set of chromosomes.
Spindle microtubules disappear and the nucleoli reorganize.
 Cytokinesis, the division of the cytoplasm to yield two daughter cells. In animal cells, an actomyosin contractile ring attached to
the PM encircles the cell and contracts, producing a cleavage furrow that gradually deepens and eventually separates the
cytoplasm into two daughter cells, each with a complete nucleus. In plant cells, cytokinesis occurs by forming a cell plate as a
line of vesicles originating in the Golgi complex. The vesicles contain materials to construct both a primary cell wall for each
daughter cell and a middle lamella that cements the primary cell walls together.
o Prokaryotes reproduce asexually, generally by binary fission, a process in which one cell divides into two offspring cells.
• Sexual Reproduction and Meiosis
o Asexual reproduction – a single parent splits to produce two or more individuals.
o In Sexual reproduction – two haploid sex cells or gametes fuse to form a single diploid zygote.
o A diploid cell has a characteristic number of chromosome pairs per cell. The members of each pair, called homologous
chromosomes, are similar in length, shape and other features and carry genes affecting the same kinds of attributes of the
organism. A haploid cell contains only one member of each homologous pair.
o A diploid cell undergoing meiosis completes two successive cell divisions, yielding four haploid cells. Sexual life cycles in eukaryotes
require meiosis, which makes it possible for each gamete to contain only half the number of chromosomes in the parent cell.

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