visual anatomy & physiology first edition martini & ober chapter 22 (sections 22.1-22.4)...
TRANSCRIPT
![Page 1: Visual Anatomy & Physiology First Edition Martini & Ober Chapter 22 (Sections 22.1-22.4) Metabolism & Cellular Respiration Lecture 8](https://reader030.vdocument.in/reader030/viewer/2022020922/56649f0e5503460f94c2271c/html5/thumbnails/1.jpg)
Visual Anatomy & PhysiologyFirst Edition
Martini & Ober
Chapter 22(Sections 22.1-22.4)
Metabolism & Cellular RespirationLecture 8
![Page 2: Visual Anatomy & Physiology First Edition Martini & Ober Chapter 22 (Sections 22.1-22.4) Metabolism & Cellular Respiration Lecture 8](https://reader030.vdocument.in/reader030/viewer/2022020922/56649f0e5503460f94c2271c/html5/thumbnails/2.jpg)
2
Lecture Overview
• Enzymes and control of metabolic reactions• Energy and metabolic reactions• Cellular respiration
– Overview– ATP as the biological energy carrier– Oxidation/Reduction– Steps in Cellular Respiration
• Glycolysis• The Citric Acid (Krebs) Cycle • The Electron Transport Chain
• Relationship of anabolism to catabolism
![Page 3: Visual Anatomy & Physiology First Edition Martini & Ober Chapter 22 (Sections 22.1-22.4) Metabolism & Cellular Respiration Lecture 8](https://reader030.vdocument.in/reader030/viewer/2022020922/56649f0e5503460f94c2271c/html5/thumbnails/3.jpg)
3
Enzymes and Metabolic Reactions
Enzymes – Biological catalysts
• control rates of metabolic reactions• lower activation energy needed to start reactions• globular proteins with specific shapes• not consumed in chemical reactions
• substrate specific
• shape of active site determines which substrate(s) the enzyme can act on
Figure from: Hole’s Human A&P, 12th edition, 2010
![Page 4: Visual Anatomy & Physiology First Edition Martini & Ober Chapter 22 (Sections 22.1-22.4) Metabolism & Cellular Respiration Lecture 8](https://reader030.vdocument.in/reader030/viewer/2022020922/56649f0e5503460f94c2271c/html5/thumbnails/4.jpg)
5
Enzymes Lower Activation Energy
Enzymes lower the barriers that block chemical reactions, i.e., they lower the activation energy needed to begin energetically favorable reactions
![Page 5: Visual Anatomy & Physiology First Edition Martini & Ober Chapter 22 (Sections 22.1-22.4) Metabolism & Cellular Respiration Lecture 8](https://reader030.vdocument.in/reader030/viewer/2022020922/56649f0e5503460f94c2271c/html5/thumbnails/5.jpg)
6
Control of Metabolic ReactionsMetabolic pathways
• series of enzyme-controlled reactions leading to formation of a product• each new substrate is the product of the previous reaction
Factors that alter activity of enzymes
• heat• radiation• substrate concentration• required cofactors• changes in pH
Enzyme names commonly• reflect the substrate• have the suffix – ase• sucrase, lactase, protease, lipase, hydrolase, oxidase
![Page 6: Visual Anatomy & Physiology First Edition Martini & Ober Chapter 22 (Sections 22.1-22.4) Metabolism & Cellular Respiration Lecture 8](https://reader030.vdocument.in/reader030/viewer/2022020922/56649f0e5503460f94c2271c/html5/thumbnails/6.jpg)
7
Cofactors and CoenzymesCofactors
• make some enzymes active• ions or coenzymes
Coenzymes• complex organic molecules that act as cofactors (so coenzymes ARE cofactors)• vitamins• NAD+
Vitamins are essential organic substances that human cells cannot synthesize, i.e., they must come from the diet
- required in very small amounts
- examples - B vitamins: Thiamine (B1), niacin
The protein parts of enzymes that need a nonprotein part (coenzymes, cofactors) to work are called apoenzymes
![Page 7: Visual Anatomy & Physiology First Edition Martini & Ober Chapter 22 (Sections 22.1-22.4) Metabolism & Cellular Respiration Lecture 8](https://reader030.vdocument.in/reader030/viewer/2022020922/56649f0e5503460f94c2271c/html5/thumbnails/7.jpg)
8
Overview of Cellular Metabolism
Figure from: Martini, Anatomy & Physiology, Prentice Hall, 2001
Metabolism – All the chemical reactions that occur in an organism
KEEP THIS OVERALL SCHEME IN MIND AS WE GO INTO DETAILS!!
ETS
![Page 8: Visual Anatomy & Physiology First Edition Martini & Ober Chapter 22 (Sections 22.1-22.4) Metabolism & Cellular Respiration Lecture 8](https://reader030.vdocument.in/reader030/viewer/2022020922/56649f0e5503460f94c2271c/html5/thumbnails/8.jpg)
9
Overview of Glucose Breakdown
Figure from: Hole’s Human A&P, 12th edition, 2010
![Page 9: Visual Anatomy & Physiology First Edition Martini & Ober Chapter 22 (Sections 22.1-22.4) Metabolism & Cellular Respiration Lecture 8](https://reader030.vdocument.in/reader030/viewer/2022020922/56649f0e5503460f94c2271c/html5/thumbnails/9.jpg)
11
Overview of Catabolism
Figure from: Martini, Visual Anatomy & Physiology, Pearson, 2011
![Page 10: Visual Anatomy & Physiology First Edition Martini & Ober Chapter 22 (Sections 22.1-22.4) Metabolism & Cellular Respiration Lecture 8](https://reader030.vdocument.in/reader030/viewer/2022020922/56649f0e5503460f94c2271c/html5/thumbnails/10.jpg)
12
A Closer Look at Mitochondria
Strategically placed in cell where ATP demand is high
Concentration of enzymes in the matrix is so high that there is virtually no hydrating water. Enzyme-linked reactions and pathways are so crowded that normal rules of diffusion do not apply!
Figure from: Martini, Anatomy & Physiology, Prentice Hall, 2001
(Impermeable to charged or polar molecules)
![Page 11: Visual Anatomy & Physiology First Edition Martini & Ober Chapter 22 (Sections 22.1-22.4) Metabolism & Cellular Respiration Lecture 8](https://reader030.vdocument.in/reader030/viewer/2022020922/56649f0e5503460f94c2271c/html5/thumbnails/11.jpg)
13
Carbohydrate Metabolism
• Most cells generate ATP and other energy-yielding compounds via the catabolism of carbohydrate (and fats)
General Reaction sequence in carbohydrate catabolism
C6H12O6 + 6 O2 6 CO2 + 6 H2O + ENERGY
If the above reaction happened all at once, all the chemical energy contained in the carbohydrate would be DISSIPATED AS HEAT.
How does the body harness the energy from carbohydrates?
![Page 12: Visual Anatomy & Physiology First Edition Martini & Ober Chapter 22 (Sections 22.1-22.4) Metabolism & Cellular Respiration Lecture 8](https://reader030.vdocument.in/reader030/viewer/2022020922/56649f0e5503460f94c2271c/html5/thumbnails/12.jpg)
14
Harnessing Energy - Stepwise Breakdown of Carbohydrates
Figure from: Alberts et al., Essential Cell Biology, Garland Press, 1998
![Page 13: Visual Anatomy & Physiology First Edition Martini & Ober Chapter 22 (Sections 22.1-22.4) Metabolism & Cellular Respiration Lecture 8](https://reader030.vdocument.in/reader030/viewer/2022020922/56649f0e5503460f94c2271c/html5/thumbnails/13.jpg)
15
Energy for Metabolic Reactions
Energy• ability to do work or change something (potential, kinetic)• heat, light, sound, electricity, mechanical energy, chemical energy• changed from one form to another, but NEVER destroyed (law of conservation of energy)• involved in all metabolic reactions
Release of chemical energy• most metabolic processes depend on chemical energy• oxidation of glucose generates chemical energy• cellular respiration releases chemical energy slowly from molecules and makes it available for cellular use
![Page 14: Visual Anatomy & Physiology First Edition Martini & Ober Chapter 22 (Sections 22.1-22.4) Metabolism & Cellular Respiration Lecture 8](https://reader030.vdocument.in/reader030/viewer/2022020922/56649f0e5503460f94c2271c/html5/thumbnails/14.jpg)
16
Oxidation and Reduction Revisited
Oxidation• gain of O2
• loss of e-
• loss of H (since a H carries an electron with it) • increase in oxidation number, e.g., Fe2+ -> Fe3+
Oxidation Is Loss of electrons; Reduction Is Gain of electrons
“OIL RIG”
Reduction • loss of O2
• gain of e-
• gain of H • decrease in oxidation number, e.g., Fe3+ -> Fe2+
![Page 15: Visual Anatomy & Physiology First Edition Martini & Ober Chapter 22 (Sections 22.1-22.4) Metabolism & Cellular Respiration Lecture 8](https://reader030.vdocument.in/reader030/viewer/2022020922/56649f0e5503460f94c2271c/html5/thumbnails/15.jpg)
17
Energy of Organic Molecules• Carbohydrates like glucose store a great deal
of chemical energy (as H·)• As carbohydrates (C6H12O6) are oxidized to
CO2 they liberate their energy and lose electrons and H (H·)
• But there must be molecules to accept these electrons, i.e., some molecules must be reduced.
• In cellular respiration, O2 becomes the final electron (H·) acceptor and is reduced to H2O
![Page 16: Visual Anatomy & Physiology First Edition Martini & Ober Chapter 22 (Sections 22.1-22.4) Metabolism & Cellular Respiration Lecture 8](https://reader030.vdocument.in/reader030/viewer/2022020922/56649f0e5503460f94c2271c/html5/thumbnails/16.jpg)
18
Harnessing Energy from Carbohydrates
General Reaction sequence in carbohydrate catabolism
C6H12O6 + 6 O2 6 CO2 + 6 H2O + ENERGY
OXIDATION
REDUCTION
Electrons (H·) “fall” in energy from organic molecules to oxygen during cellular respiration.
That is, e- LOSE potential energy during this process and this energy is captured to make ATP
However, electrons CANNOT be transferred directly from glucose to the electron transport chain. There are intermediates – activated carrier molecules
![Page 17: Visual Anatomy & Physiology First Edition Martini & Ober Chapter 22 (Sections 22.1-22.4) Metabolism & Cellular Respiration Lecture 8](https://reader030.vdocument.in/reader030/viewer/2022020922/56649f0e5503460f94c2271c/html5/thumbnails/17.jpg)
19
Activated Carrier Molecules
Figure from: Alberts et al., Essential Cell Biology, Garland Press, 1998
Some activated carriers: ATP, NADH, FADH2, GTP, NADPH
![Page 18: Visual Anatomy & Physiology First Edition Martini & Ober Chapter 22 (Sections 22.1-22.4) Metabolism & Cellular Respiration Lecture 8](https://reader030.vdocument.in/reader030/viewer/2022020922/56649f0e5503460f94c2271c/html5/thumbnails/18.jpg)
20
ATP – An Activated Carrier Molecule
• each ATP molecule has three parts:• an adenine molecule• a ribose molecule• three phosphate molecules in a chain
• ATP carries its energy in the form or P (phosphate)
• ATP is a readily interchangeable form of energy for cellular reactions (“common currency”)
These two components together are called a ?
High-energy bonds
Figure from: Hole’s Human A&P, 12th edition, 2010
![Page 19: Visual Anatomy & Physiology First Edition Martini & Ober Chapter 22 (Sections 22.1-22.4) Metabolism & Cellular Respiration Lecture 8](https://reader030.vdocument.in/reader030/viewer/2022020922/56649f0e5503460f94c2271c/html5/thumbnails/19.jpg)
21
NAD(H) – An Activated Carrier Molecule
NAD (and NADP) are specialized to carry high-energy e- and H atoms
Figure from: Alberts et al., Essential Cell Biology, Garland Press, 1998
A “packet” of energy = H·
These packets of energy will be passed to oxygen in the electron transport chain, and their energy used to drive the synthesis of ATP
Important carriers of e- in catabolism: NADH, FADH2
NAD+
NADH + H+
NAD+ NADH
![Page 20: Visual Anatomy & Physiology First Edition Martini & Ober Chapter 22 (Sections 22.1-22.4) Metabolism & Cellular Respiration Lecture 8](https://reader030.vdocument.in/reader030/viewer/2022020922/56649f0e5503460f94c2271c/html5/thumbnails/20.jpg)
22
Overview of Cellular Respiration
Figure from: Martini, Anatomy & Physiology, Prentice Hall, 2001
Cellular respiration
(aerobic)
Anaerobic
![Page 21: Visual Anatomy & Physiology First Edition Martini & Ober Chapter 22 (Sections 22.1-22.4) Metabolism & Cellular Respiration Lecture 8](https://reader030.vdocument.in/reader030/viewer/2022020922/56649f0e5503460f94c2271c/html5/thumbnails/21.jpg)
23
Overview of Glucose BreakdownFigure from: Hole’s Human A&P, 12th edition, 2010
![Page 22: Visual Anatomy & Physiology First Edition Martini & Ober Chapter 22 (Sections 22.1-22.4) Metabolism & Cellular Respiration Lecture 8](https://reader030.vdocument.in/reader030/viewer/2022020922/56649f0e5503460f94c2271c/html5/thumbnails/22.jpg)
24
Overview of Glucose BreakdownOccurs in three major of reaction series…
1. Glycolysis (glucose to pyruvate; in cytoplasm) 2. Citric acid cycle (finishes oxidation begun in glycolysis;
in the matrix of mitochondria)3. Electron transport chain (uses e- transfer to make
ATP; on inner membranes of mitochondria)
Produces• carbon dioxide• water• ATP (chemical energy)• heat (energy has changed form from chemical)
Includes• anaerobic reactions (without O2) - produce little ATP• aerobic reactions (requires O2) - produce most ATP
![Page 23: Visual Anatomy & Physiology First Edition Martini & Ober Chapter 22 (Sections 22.1-22.4) Metabolism & Cellular Respiration Lecture 8](https://reader030.vdocument.in/reader030/viewer/2022020922/56649f0e5503460f94c2271c/html5/thumbnails/23.jpg)
25
Glycolysis• series of ten reactions
• breaks down glucose (6C) into 2 molecules of pyruvic acid (pyruvate) (3C)
• occurs in cytosol
• anaerobic phase of cellular respiration (that is, it can continue to work with OR without O2)
• yields 2 ATP and 2 NADH molecules per glucose
Glucose (6C) 2 Pyruvate (3C)NAD+ NADH
ADP + Pi ATP
![Page 24: Visual Anatomy & Physiology First Edition Martini & Ober Chapter 22 (Sections 22.1-22.4) Metabolism & Cellular Respiration Lecture 8](https://reader030.vdocument.in/reader030/viewer/2022020922/56649f0e5503460f94c2271c/html5/thumbnails/24.jpg)
26
Overview of Glycolysis
glucose (6C) → 2 pyruvate (3C)
Products of glycolysis
- ATP
- NADH
- Pyruvate
NOTE what happens with and without O2 being available…
Figure from: Hole’s Human A&P, 12th edition, 2010
![Page 25: Visual Anatomy & Physiology First Edition Martini & Ober Chapter 22 (Sections 22.1-22.4) Metabolism & Cellular Respiration Lecture 8](https://reader030.vdocument.in/reader030/viewer/2022020922/56649f0e5503460f94c2271c/html5/thumbnails/25.jpg)
27
Metabolism of Pyruvate Without O2
• process of forming lactate from glucose is anerobic glycolysis
• important for regenerating NAD+ so glycolysis can continue to generate ATP for the cell
Glucose (6C) 2 Pyruvate (3C)
NAD+ NADH + H+
ADP + Pi ATP
Pyruvic acid (pyruvate)NADH + H+ NAD+
Lactic acid (lactate)
O2
![Page 26: Visual Anatomy & Physiology First Edition Martini & Ober Chapter 22 (Sections 22.1-22.4) Metabolism & Cellular Respiration Lecture 8](https://reader030.vdocument.in/reader030/viewer/2022020922/56649f0e5503460f94c2271c/html5/thumbnails/26.jpg)
28
Overview of Aerobic Reactions
If oxygen is available – • pyruvic acid is used to produce acetyl CoA• citric acid (Krebs) cycle begins• electron transport chain functions• carbon dioxide and water are formed• maximum of 36 molecules of ATP produced per glucose molecule
Figure from: Hole’s Human A&P, 12th edition, 2010
![Page 27: Visual Anatomy & Physiology First Edition Martini & Ober Chapter 22 (Sections 22.1-22.4) Metabolism & Cellular Respiration Lecture 8](https://reader030.vdocument.in/reader030/viewer/2022020922/56649f0e5503460f94c2271c/html5/thumbnails/27.jpg)
29
Citric Acid Cycle
What happens…
- Acetyl CoA enters cycle - Citric Acid is converted to various intermediates - Several important products are produced in these interconversions of citric acid…
• ATP is produced•NAD+ is reduced to NADH and FAD is reduced to FADH2
• CO2 produced
In mitochondria…Figure from: Hole’s Human A&P, 12th edition, 2010
![Page 28: Visual Anatomy & Physiology First Edition Martini & Ober Chapter 22 (Sections 22.1-22.4) Metabolism & Cellular Respiration Lecture 8](https://reader030.vdocument.in/reader030/viewer/2022020922/56649f0e5503460f94c2271c/html5/thumbnails/28.jpg)
30
Source of e- for the Electron Transport Chain
Notice the flow of electrons to the Electron Transport Chain
Figure from: Hole’s Human A&P, 12th edition, 2010
![Page 29: Visual Anatomy & Physiology First Edition Martini & Ober Chapter 22 (Sections 22.1-22.4) Metabolism & Cellular Respiration Lecture 8](https://reader030.vdocument.in/reader030/viewer/2022020922/56649f0e5503460f94c2271c/html5/thumbnails/29.jpg)
31
Electron Transport Chain• NADH and FADH2 carry electrons to the ETC• ETC series of electron carriers located in cristae of mitochondria• energy from electrons transferred to ATP synthase• ATP synthase catalyzes the oxidative phosphorylation of ADP to ATP• water is formed
*Chemiosmosis
Figure from: Hole’s Human A&P, 12th edition, 2010
![Page 30: Visual Anatomy & Physiology First Edition Martini & Ober Chapter 22 (Sections 22.1-22.4) Metabolism & Cellular Respiration Lecture 8](https://reader030.vdocument.in/reader030/viewer/2022020922/56649f0e5503460f94c2271c/html5/thumbnails/30.jpg)
32
Oxidative Phosphorylation
Figure from: Martini, Anatomy & Physiology, Prentice Hall, 2001
Chemiosmosis, Chemiosmotic coupling, or Chemiosmotic phosphorylation
![Page 31: Visual Anatomy & Physiology First Edition Martini & Ober Chapter 22 (Sections 22.1-22.4) Metabolism & Cellular Respiration Lecture 8](https://reader030.vdocument.in/reader030/viewer/2022020922/56649f0e5503460f94c2271c/html5/thumbnails/31.jpg)
33
Summary of Cellular Respiration
Figure from: Martini, Anatomy & Physiology, Prentice Hall, 2001
![Page 32: Visual Anatomy & Physiology First Edition Martini & Ober Chapter 22 (Sections 22.1-22.4) Metabolism & Cellular Respiration Lecture 8](https://reader030.vdocument.in/reader030/viewer/2022020922/56649f0e5503460f94c2271c/html5/thumbnails/32.jpg)
34
Is This All the Metabolism There Is?
Definitely NOT!
Notice how many lines connect pyruvate and Acetyl CoA to the rest of metabolism
![Page 33: Visual Anatomy & Physiology First Edition Martini & Ober Chapter 22 (Sections 22.1-22.4) Metabolism & Cellular Respiration Lecture 8](https://reader030.vdocument.in/reader030/viewer/2022020922/56649f0e5503460f94c2271c/html5/thumbnails/33.jpg)
35
Intermediates of Metabolism
Acetyl CoA (2C) and Pyruvate (3C) are important:
Allow interconversion of different types of molecules so cell’s needs can be met
Figure from: Martini, Visual Anatomy & Physiology, Pearson, 2011
![Page 34: Visual Anatomy & Physiology First Edition Martini & Ober Chapter 22 (Sections 22.1-22.4) Metabolism & Cellular Respiration Lecture 8](https://reader030.vdocument.in/reader030/viewer/2022020922/56649f0e5503460f94c2271c/html5/thumbnails/34.jpg)
36
Summary of Catabolism of Proteins, Carbohydrates, and Fats
Acetyl CoA is a common intermediate in the breakdown of most fuels.
Acetyl CoA can be generated by carbohydrates, fats, or amino acids
Acetyl CoA can be converted into fatty acids
Figure from: Hole’s Human A&P, 12th edition, 2010
![Page 35: Visual Anatomy & Physiology First Edition Martini & Ober Chapter 22 (Sections 22.1-22.4) Metabolism & Cellular Respiration Lecture 8](https://reader030.vdocument.in/reader030/viewer/2022020922/56649f0e5503460f94c2271c/html5/thumbnails/35.jpg)
37
Pyruvate is a Key Junction in Metabolism
Figure from: Martini, Anatomy & Physiology, Prentice Hall, 2001
Pyruvate is used to synthesize amino acids and Acetyl CoA
Pyruvate can also be used to synthesize glucose via gluconeogenesis.
*
Lipolysis
Lipo-
genesis
GlycogenesisGlycogenolysis
![Page 36: Visual Anatomy & Physiology First Edition Martini & Ober Chapter 22 (Sections 22.1-22.4) Metabolism & Cellular Respiration Lecture 8](https://reader030.vdocument.in/reader030/viewer/2022020922/56649f0e5503460f94c2271c/html5/thumbnails/36.jpg)
38
Carbohydrate StorageExcess glucose can be
• stored as glycogen by glycogenesis (liver and muscle cells) • stored as fat by lipogenesis• converted to amino acids
Figure from: Hole’s Human A&P, 12th edition, 2010
![Page 37: Visual Anatomy & Physiology First Edition Martini & Ober Chapter 22 (Sections 22.1-22.4) Metabolism & Cellular Respiration Lecture 8](https://reader030.vdocument.in/reader030/viewer/2022020922/56649f0e5503460f94c2271c/html5/thumbnails/37.jpg)
39
Terms to Know…
• Glycolysis – metabolism of glucose to pyruvate• Gluconeogenesis – metabolism of pyruvate to
glucose (making CHO from non-CHO source)• Glycogenesis – metabolism of glucose to
glycogen• Glycogenolysis – metabolism of glycogen to
glucose• Lipolysis – breakdown of triglyceride into
glycerol and fatty acids• Lipogenesis – creation of new triglyceride (fat)
-olysis breakdown of -neo new-genesis creation of
![Page 38: Visual Anatomy & Physiology First Edition Martini & Ober Chapter 22 (Sections 22.1-22.4) Metabolism & Cellular Respiration Lecture 8](https://reader030.vdocument.in/reader030/viewer/2022020922/56649f0e5503460f94c2271c/html5/thumbnails/38.jpg)
40
Review
• Enzymes are biological catalysts– Highly specific for their substrate– Lower activation energy needed to start a reaction– Are not consumed during reaction– May require cofactors/coenzymes– Effectiveness is greatly affected by temperature,
pH, and the presence of required cofactors
• The goal of metabolism is to provide the cell with energy (catabolism) and materials for the manufacture of cellular components (anabolism)
![Page 39: Visual Anatomy & Physiology First Edition Martini & Ober Chapter 22 (Sections 22.1-22.4) Metabolism & Cellular Respiration Lecture 8](https://reader030.vdocument.in/reader030/viewer/2022020922/56649f0e5503460f94c2271c/html5/thumbnails/39.jpg)
41
Review
• Cells derive energy mainly from carbon compounds like carbohydrates and fats– These substances contain a great deal of energy
stored in their chemical bonds– This energy must be liberated in stepwise
fashion– Activated carriers serve as intermediates to
capture the energy liberated at each step
• Energy is the ability to do work– May be potential or kinetic– Changes form– Is never destroyed (only converted to another
form)
![Page 40: Visual Anatomy & Physiology First Edition Martini & Ober Chapter 22 (Sections 22.1-22.4) Metabolism & Cellular Respiration Lecture 8](https://reader030.vdocument.in/reader030/viewer/2022020922/56649f0e5503460f94c2271c/html5/thumbnails/40.jpg)
44
Review
• Anabolism is intimately tied to catabolism– Energy derived from catabolism is used to drive anabolic
reactions– Some molecules are important junctions between
catabolism and anabolism – Acetyl CoA
• Generated from pyruvate, fatty acids, and amino acids• Can be used to synthesize fatty acids and other molecules• CANNOT be used to generate pyruvate
– Pyruvate • Can be synthesized from glucose and amino acids• can be used to synthesize amino acids, glucose and acetyl CoA
– NOTE, however, that in humans fatty acids cannot be converted to glucose