CELLULAR RESPIRATION and FERMENTATION

Energy flow and chemical recycling in ecosystems

Two major catabolic pathways
‡ Fermentation: partial degradation of organic molecules in the absence of oxygen ‡ Cellular respiration: complete breakdown of organic molecules, more efficient, uses oxygen ± Most processes occur in mitochondria ± General reaction: organic compound + O2 CO2 + H2O + energy

± Example glucose: C6H12O6 + 6O2 6CO2 + 6H2O + energy (ATP + heat)

How is energy harvested?
‡ Redox reactions yield energy in catabolic pathways ‡ General definition: Redox reactions = reactions that result in the transfer of one or more electrons from one reactant to another ‡ Oxidation: loss of electrons ‡ Reduction: addition of electrons ‡ General reaction: Xe- + Y X + Ye± X (electron donor) is the reducing agent and reduces Y ± Y (electron recipient) is the oxidizing agent and oxidizes X

How is energy harvested?
‡ Oxygen is one of the most potent oxidizing agents (most electronegative atom)  As electrons ³fall´ from a less electronegative atom to a more electronegative atom, they lose free potential energy ‡ Example: methane combustion ‡ Good fuels: carbs and fats

each catalyzed by a specific enzyme .‡ Fuels are broken down gradually in a series of steps.

but via a coenzyme (NAD+ = nicotinamide adenine dinucleotide)  NAD+: oxidizing agent in many redox steps .Principles of cellular respiration ‡ Transfer of electrons to oxygen is not direct.

‡ The stepwise ³fall´ of electrons during cellular respiration: ± Energy is tapped to synthesize ATP as electrons ³fall´ from NADH to oxygen ± Reaction steps are called electron transport chain .

Three stages during cellular respiration 1. 3. Glycolysis Krebs cycle (Citric acid cycle. tricarboxylic acid cycle) Electron transport chain and oxidative phosphorylation . 2.

Glucose: the example .

Glycolysis (³Splitting of sugar´) ‡ Glucose is split into two three-carbon sugars. which are oxidized and rearranged to pyruvate ‡ Ten steps in two phases ‡ Net yield: 2 ATP and 2 NADH per glucose ‡ In cytosol .

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Energy investment phase .

Energy payoff phase .

The Krebs cycle ‡ Completes the energy-yielding oxidation of organic molecules ‡ Pyruvate is channeled into mitochondrion and modified into Acetyl CoA .

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are produced in the Krebs cycle from each acetyl group.and 8H+ produced. .+ H+ 3 H2O are needed to provide 3O for 2CO2. one gets to CoA. 2 CO2 and 8e. So. There are 3H to start. These 3H2O give 6H+ and 6e-. totally 8 e.O CH3 ²C ²S ²CoA 2CO2 + HS²CoA + e.

Krebs cycle summary ‡ Each molecule pyrovate is broken down into 3 CO2 ‡ Each cycle yields ± 4 NADH ± 1 FADH2 ± 1 GTP ‡ Per glucose molecule two cycles! ‡ Within mitochondrial matrix .

The electron transport chain (ETC) ‡ Within inner membrane of mitochondrion (cristae) ‡ Electrons carried by NADH are transferred along the ETC to oxygen .

The electron transport chain (ETC) ‡ Produces a proton (H+) gradient between the intermembrane space and the matrix .

The electron transport chain (ETC) .

‡ H+ concentration is a proton-motive force: H+ diffuse back to the matrix through ³molecular mill´ generation of ATP ‡ Chemiosmosis: The coupling of the redox reactions of the electron transport chain to ATP synthesis .

Cellular respiration ± summary ‡ Most energy flows from glucose NADH ETC protonmotive force ATP ‡ Efficiency of respiration = 40% of initial energy of glucose .

Fermentation ‡ Anaerobic catabolism of sugars through a different ³finish´ of glycolysis ‡ NAD+ is recycled by transferring electrons from NADH to pyruvate or derivatives of pyruvate in various fermentation pathways .

In anaerobic condition: 2 .

such as yeasts: 2 .In organisms that can grow anaerobically.

Alcohol fermentation ‡ ‡ Pyruvate is converted to ethanol in two steps Many bacteria and some fungi can carry out alcohol fermentation .

applications .Alcohol fermentation .

When inadequate oxygen is present. for example. in a muscle cell undergoing vigorous contraction: .

Lactic acid fermentation ‡ Pyruvate is reduced directly by NADH to form lactate (ionized form of lactic acid). ‡ Muscle cells switch to lactic acid fermentation when O2 is scarce ‡ Some fungi and bacteria can carry out lactic acid fermentation .

Lactic acid fermentation .applications .

Fermentation and cellular respiration: common features ‡ Both use glycolysis to oxidize sugars to pyruvate with a net production of 2 ATP by substrate-level phosphorylation ‡ Both use NAD+ as an electron acceptor .

Fermentation and cellular respiration: differences ‡ Fermentation: ± NAD+ is recycled by passing the electrons of NADH to an organic molecule ± One molecule glucose yields 2 ATP (energy stored in pyruvate is unavailable to the cell) ‡ Respiration: ± Electrons of NADH are ultimately passed to O2. generating ATP by oxidative phosphorylation ± Additional ATP is generated from the oxidation of pyruvate in the Krebs cycle ± One molecule glucose yields ~38 ATP .

Other carbohydrates as well as fats and proteins can also be broken down ‡ Intermediate product enter at different steps of glycolysis and Krebs cycle .

 Metabolic pathways of respiration also important for the synthesis of molecules .

Feedback mechanisms control cellular respiration ‡ E.g. phosphofructokinase (catalyzes third step of glycolysis): ± inhibited by ATP and citrate (first product of the Krebs cycle) ± stimulated by AMP ± allosteric regulation  Rate of glycolysis and Krebs cycle are synchronized .

Facultative anaerobes ‡ Some yeast and many bacteria that can survive using either fermentation or respiration ‡ At a cellular level. human muscle cells can behave as facultative anaerobes Saccharomyces Yeast Cells .

5 billion years old ± appeared long before appreciable quantities of O2 accumulated in the atmosphere (green algae and plants hadn¶t evolved yet)  First prokaryotes may have generated ATP exclusively from glycolysis .Evolutionary perspective ‡ Glycolysis ± the most widespread metabolic pathway and occurs in the cytosol without membrane-enclosed organelles ± probably evolved early in the history of life ‡ Oldest bacterial fossils ± are over 3.

g.Other carbohydrates as well as fats and proteins can also be broken down Carbohydrates: ‡ Polysaccharides (e.g. starch) can be hydrolyzed to glucose monomers that enter glycolysis ‡ Hexose sugars other than glucose (e. fructose) are modified to enter glycolysis .

etc.Other carbohydrates as well as fats and proteins can also be broken down Proteins: ‡ Must first be digested to individual amino acids ‡ Amino acids must have their amino groups removed (excreted as ammonia.) ‡ Carbon skeletons are then modified by enzymes and enter as intermediaries into glycolysis or the Krebs cycle depending on their structure . urea.

then enter the Krebs cycle as acetyl CoA ‡ A gram of fat will generate twice as much ATP as a gram of carbohydrate via aerobic respiration .Other carbohydrates as well as fats and proteins can also be broken down Fats ‡ Must be digested to glycerol and fatty acids ‡ Glycerol can be converted to an intermediate of glycolysis ‡ Fatty acids are split into twocarbon fragments via beta oxidation.

Metabolic pathways of respiration also important for the synthesis of molecules ‡ Not all organic molecules of food are completely oxidized to make ATP ‡ Intermediaries in glycolysis and the Krebs cycle can be diverted to anabolic pathways: ± E. a human cell can synthesize about half the 20 different amino acids by modifying compounds from the Krebs cycle ± Glucose can be synthesized from pyruvate and fatty acids from acetyl CoA ‡ Cells can convert one kind of molecule to another using glycolysis and Krebs cycle: ± E. excess carbohydrates and proteins can be converted to fats .g.g.

Feedback mechanisms control cellular respiration ‡ As ATP levels drop. catabolism speeds up to produce more ATP ‡ Excess ATP leads to inhibition of catabolism ‡ Control is based mainly on regulating the activity of enzymes at strategic points in the catabolic pathway ‡ E. phosphofructokinase (catalyzes third step of glycolysis): ± inhibited by ATP and citrate (first product of the Krebs cycle) ± stimulated by AMP  Rate of glycolysis and Krebs cycle are synchronized .g.

Metabolic pathways are complex« .

Review of key processes ‡ During glycolysis and Krebs cycle ATP is produced directly through substrate-based phosphorylation ‡ In the electron transport chain ATP is produced indirectly through oxidative phosphorylation much greater yield .

Substrate-based phosphorylation ‡ Directly catalyzed by an enzyme ‡ Remember: enzymes ± lower the activation energy needed for a reaction to proceed ± Speed up reactions ± Are substrate-specific ± Make reactions controllable through feedback mechanisms .

Oxidative phosphorylation ‡ Ultimately driven by the loss of electrons from food molecules ‡ Electrons are shuttled via NADH (and FADH2) .

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2.1. Acetyl CoA adds its two-carbon fragment Citrate is converted to isocitrate by removal and subsequent addition of a water molecule .

resulting compound is oxidized. reducing NAD+ to NADH.3. reducing NAD+ to NADH CO2 is lost. remaining molecule is attached to CoA 4. resulting compound is oxidized. . CO2 is lost.

which is eventually transferred to ADP ATP Two hydrogens are transferred to FAD FADH2 6. CoA is displaced by a phosphate group.5. .

7. 8. Addition of water rearranges bonds Compound is oxidized. reducing NAD+ NADH Cycle starts again  .

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