LECT 6. RESPIRATION COMPETENCIES. Students, after mastering materials of the present lecture, should be able:

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1 LECT 6. RESPIRATION COMPETENCIES Students, after mastering materials of the present lecture, should be able: To explain the process of respiration (the oxidation of substrates particularly carbohydrates or the synthesis of metabolic energy used for plant growth and maintenance) To explain reactions, enzymes and products involved the respiration 2 1

2 Glucose is the most commonly cited substrate of respiration Glucose serves as the primary energy source for the brain and is also a source of energy for cells throughout the body Blood glucose is normally maintained between 70 mg/dl and 110 mg GUT4: glucose transporters (protrein) LECTURE FLOW QUESTIONS SUMMARY OF RESPIRATION 1. DEFINITION 2. THE SITE OF RESPIRATION 3. MAIN STEPS OF RESPIRATION 4. COMPARING ENERGY YIELD 5. FEEDBACK CONTROL OF RESPIRATION 6. RESPIRATION AND PLANT CARBON BALANCE 7. FACTORS AFFECTING RESPIRATION 2

3 QUESTIONS What is the importance of respiration How many steps does the respiration have What is the first and second step of respiration Where does the glycolysis take place Where does the TCA cycle take place Where does the Electron Transport Chain take place What is the compound exported from cytosol to mitochondria in the respiration What is the final acceptor of electrons in the respiration How many NADP does the glycolysis produce How many NADP does the TCA cycle produce How many ATP does the glycolysis produce How many ATP does the TCA cycle produce What is the meaning of ADP:O ratios QUESTIONS What is the ADP:O ratio of FADH2 How many ATP does the respiration produces totally How efficient is the respiration in the conversion of energy What does it mean by feedback control of respiration How is the effect of tissue stage of development on the respiration How is the effect of O2 on the respiration How is the effect of CO2 on the respiration How is the effect of plant injury on the respiration How is the foliar respiration of trees in term of height How is the response of foliar respiration to nitrogen How is the woody respiration in term of wood diameter 3

4 Respiration = Dark Respiration CH2O + O2 CO2 + H2O + energy Biochemistry of respiration Glycolysis ( glyco = sugar; lysis = untie ) Citric acid cycle ( Krebs cycle ) Electron transport and ATP synthesis Respiration and carbon economy of whole plants Hans Krebs, 1953 Nobel Prize Glycolysis break a 6-carbon sugar into two 3Carbon sugars (triose phosphate) takes some energy then strip electrons from these 3-C sugars releases a bit of energy in the form of ATP and NADH. Leftover products: 3C sugars Pyruvate and Malate (still embody substantial free energy) Citric acid/krebs cycle complete oxidation of pyruvate/malate to produce CO2, H2O, reducing power (NADH, FADH2) and ATP Electron Transport Chain launder NADH, FADH2 to ATP across inner mitochondrial membrane and to Membrane) 4

5 What is respiration? 1. DEFINITION Respiration is the process whereby the energy stored in carbohydrates, produced during photosynthesis, is released in a controlled manner. The energy (free energy) released during respiration is incorporated into a form (ATP) that can be readily utilized for the maintenance and development of the plant. Respiration is tightly coupled to other pathways Biosynthesis of Nucleotides Biosynthesis of Proteins Biosynthesis of Lipids Biosynthesis of Cell wall components Biosynthesis of Phytohormones Biosynthesis of Plant pigments 5

6 Where does the respiration take place? 2. THE SITE OF RESPIRATION 1. Mitochondria are the main site of ATP synthesis in eukaryote cells and as such are vital for the health and survival of the cell 2. From a chemical standpoint, respiration is most commonly expressed in terms of the oxidation of the six-carbon sugar glucose This equation represents a coupled redox reaction that oxidizes completely glucose to CO 2 with oxygen serving as the ultimate electron acceptor and reduced to water. The substrate for respiration most commonly cited, in a functioning plant cell, is glucose that is actually derived from such sources as the glucose polymer starch, the disaccharide sucrose, fructose-containing polymers (fructosans), and other sugars, as well as lipids (primarily triacylglycerol), organic acids, and on occasion, proteins The amount of energy release is roughly 2880 kj (686 kcal) per mole (180 g) of glucose oxidized that is coupled to the synthesis of ATP. 6

7 Matrix contains the TCA cycle (and other) soluble enzymes Two membranes Inner membrane contains metabolite transporters and the electron transport chain Numbers of mitochondria per cell Inner membrane invaginated vary but usually 100s/cell Mitochondria have their own DNA and Ribosomes Mitochondrial DNA Mitochondria have some of their own DNA, ribosomes and trna; 22 trnas & rrnas (16S and 12S), so mitochondria can make many of their own proteins. The DNA is circular and lies in the matrix in structures called "nucleoids". Each nucleoid may contain 4-5 copies of the mitochondrial DNA (mtdna). 7

8 What are the stages of glucose oxidation? 3. MAIN STEPS OF RESPIRATION The reactions of glucose oxidation can be subdivided into three stages: 1. Glycolysis 2. The tricarboxylic acid (TCA) cycle, and 3. The electron transport chain (terminal oxidation) Anaerobic Respiration C6H12O6 + O2 2 CH2O5 + 2 H2O + 2 ATP or Glucose + Oxygen 2 Ethanol + 2 Water + 2 ATP 8

9 3 Main Respiration Steps 1. Glycolysis 2. Glucose (C6H12O6) Pyruvate (C3H4O3): Breakdown of Glucose to a 3-Carbon Compound Called called Pyruvate This occurs in Cytosol Some ATP and NADH Are also Formed Storage Energy Molecules NADH is Formed from NAD Similar Type of EnergyStoring Rx as NADP + H2 NADPH2 NAD + H NADH 2. Krebs Cycle/Citric Acid Cycle Tricarboxylic acid Cycle (TCA Cycle) occurs in Mitochondrial Matrix A Cyclic Series of Rxs that Completely Break down Pyruvate to CO2 and Various Carbon Skeletons Skeletons Are Used in other Metabolic Pathways to Make various Compounds Proteins Lipids Cell Wall Carbohydrates DNA This is the step where CO2 is Plant Hormones given off by the Plant Plant Pigments Many other Biochemical Compounds 9

10 3. Electron Transport Chain Oxidative Phosphorylation Series of proteins in the mitochondria helps transfer electrons (e-) from NADH to oxygen Releases a lot of energy This occurs on Mitochondrial Inner Membrane (Proteins Bound to Membrane) Released energy is used to drive the reaction ADP + P ATP Many ATP are made Oxygen is required for this step Water is produced 10

11 Overview of aerobic respiration GLYCOLISIS: SUCROSE or STARCH PYRUVATE Sucrose Glucose + Fructose Starch Glu 1-P Glu 1-P Glu 6-P Glucose Glu 6-P Fructose Fruc 6-P Glu 6-P Fruc 6-P Fru 6-P Fruc 1,6-P Fru 1,6-P Dihydroxyacetone P + Glyceraldehyde 3-P Glyceraldehyde 3-P 1,3 Bisphosphoglycerate 1,3 Bisphosphoglycerate 3 phosphoglycerate 3 phosphoglycerate 2 phosphoglycerate 2 phosphoglycerate Phosphoenol pyruvate (PEP) PEP Pyruvate Anaerobic Respiration (Fermentation) Pyruvate Lactate or Pyruvate Ethanol 11

12 Citric Acic Cycle (Krebs Cycle) 1. Pyurvate Acetyl-CoA 2. Acetyl-CoA Citrate 3. Citrate Isocitrate 4. Isocitrate α-ketoglutarate 5. α-ketoglutarate SuccinylCoA 6. Succinyl-CoA Succinate 7. Succinate Fumarate 8. Fumarate Malate 9. Malate Oxaloacetate 12

13 The electron transport chain (terminal oxidation) Complex I: NADH Dehydrogenase oxidizes NADH Complex II: Succinate Dehydroganase oxidizes reduced UQ (= ubiquinol) pumps 1H+ per e- Complex IV: Cytochrome c oxidase reduces O2 to H2O pumps 1H+ per e- uses electrochemical proton gradient to synthesize ATP oxidation of succinate (from citric acid cycle) e- are transferred via FADH2 does not pump protons Complex III: Cytochrome bc1 complex transfers e- to Ubiquinone (UQ) pumps 1H+ per e- Complex V: ATP synthase - complex I is a NADHubiquinone reductase - complex II is succinate dehydrogenase (part of the TCA cycle) - complex III is the ubiquinone -cytochrome c reductase - complex IV is cytochrome oxidase 13

14 14

15 Table. ADP/O ratios in isolated plant mitochondria Substrate ADP/O NADH (Malate) NADH (Succinate) NADH (External) NADH (Ascorbate)* *Artificial electron donor ADP/O = number of ATPs synthesized per two electrons transferred to oxygen 4. COMPARING ENERGY YIELD Glycolysis (per glucose): Krebs (per glucose): Net: 2ATP, 2NADH 1 mol glucose gives ΔGo = kj/mol 1 mol ATP takes ΔGo = 50.2 kj/mol 2ATP, 8NADH, 2FADH2 Total: 4ATP, 10NADH equivalents ATP:NADH ratio ~3 in Mitochondria Thus 4ATP + 10NADHx3 = 34 ATPs per glucose (more or less) (34 x 50.2 kj/mol)/(2880 kj/mol )= 59% conversion efficiency! (versus around 4% for glycolysis alone) 15

16 ENERGY PRODUCTION Site/Process Quantity ADP/O ATP Glycolysis 2 ATP TCA Cycle 2 ATP Cytosol 2 NADH 2.5 Mitochondrial Matrix 8 NADH 2.5 Mitochondrial Matrix 2 FADH2 1.5 TOTAL Conversion Efficiency (32 x 50.2 kj/mol)/(2880 kj/mol )= 55.8% Complex IV, the cytochrome c oxidase, is specifically inhibited by cyanide (CN-), azide (N3-), and carbon monoxide (CO). Cyanide and azide bind tightly to the ferric form of cytochrome a3, whereas carbon monoxide binds only to the ferrous form. The inhibitory actions of cyanide and azide at this site are very potent, whereas the principal toxicity of carbon monoxide arises from its affinity for the iron of hemoglobin 16

17 5. FEEDBACK CONTROL OF RESPIRATION Demand regulation Low amounts of ADP dramatically reduce the rate of mitochondrial respiration The rate of respiration increases when energy demand for growth, maintenance and transport processes is high that consume rapidly ATP leading to the production of ADP 17

18 RESPIRATION AND PLANT CARBON BALANCE On a wholeplant basis, respiration consumes from 30% to 70% of total fixed carbon Leaves account for about half of the total FACTORS AFFECTING RESPIRATION 1. Kind of Cell or Tissue. 2. Temperature 3. Low O2 can reduce aerobic respiration and increase anaerobic respiration Low O2 can reduce photorespiration CO2 5. Respiration generally has higher optimum and maximum temps than PS Rxs Oxygen 4. Young and developing cells (meristematic areas) usually have higher respiration rates. Developing and ripening fruit and seeds, too. Older cells and structural cells respire at lower rates Higher CO2 levels reduce rate of respiration (feedback inhibition). Seldom occurs except when O2 levels are limited (flooded, compacted soils) Plant Injury Injury will Increase Respiration 18

19 Kind of Cell or Tissue 19

20 Temperature Effect 20

21 21

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