BCH301

Course: BCH 301- Lipid Biochemistry (3 Credits - Compulsory)

Course Duration: Two hours per week for 15 weeks (30 hours)

Practical: Three hour per week for 15 wks (45 hours)

As taught in 2011/2012 session

Courseware Developed by :

1. DR. (MRS.) A.T. OLADIJI

B.Sc.(Hons.), M.Sc. Ph.D. (ILORIN) Biochemistry [email protected], [email protected] 2. DR. N.O. MUHAMMAD B.Sc.(Hons.), M.Sc. Ph.D. (ILORIN) Biochemistry [email protected], [email protected]

3. MRS. F. A. SULAIMAN B.Sc.(Hons.), M.Sc. (ILORIN) Biochemistry [email protected], [email protected]

2. LECTURER DETAILS:

1. DR. N.O. MUHAMMAD B.Sc.(Hons.), M.Sc. Ph.D. (ILORIN) Biochemistry [email protected], [email protected] Lecturer’s Office Address: Block 5, Room 5G 22, Department of Biochemistry Consultation hours: MONDAYS 10-12 noon. (subject to change) 2. MRS. F. A. SULAIMAN B.Sc.(Hons.), M.Sc. (ILORIN) Biochemistry [email protected], [email protected] Lecturer’s Office Address: Block 5, Room 5G12, Department of Biochemistry Consultation hours: MONDAYS 10-12 noon. (subject to change)

3.0 COURSE DETAILS: 3.1 Course Content

Classification of Lipids. Blood lipids and the lipoprotein systems. Lipid micelles monolayers and bilayers. Oxidation of fats. General biosynthesis of lipids, phospholipids and

sphingolipids. Unsaturated and essential fatty acids. Adipose tissue. Regulation of the metabolism of fats, ketosis, cholesterol metabolism. 30h (T); 45h (P); C PR: BCH 201

3.2 Course Description:

This course is designed to introduce 300 level students in the Department of Biochemistry to the macromolecule, Lipid. They will be taught the basic classification of lipids, be made to understand what blood lipids are and the lipoprotein systems. Fats are always synthesized and oxidized depending on the energy demand/status of the system. They will be taught how the body synthesizes such lipids as; phospholipids and sphingolipids.

Unsaturated fatty and essential fatty acids will be explained into details. The students will also be made to understand what adipose tissues are, how the metabolism of fats are regulated, ketosis and how cholesterol are metabolized.

3.3 Course Justification:

Lipids constitute a class of biomolecules important to life. Its importance to life will be emphasized in this course, as it will provide knowledge and all necessary information on how the body metabolizes the lipids obtained from the diet. How the body store excess fatty acid will also be discussed. Problems associated with excessive intake of some classes of lipids will be enumerated.

3.4 Course Objectives:

At the end of the course, the students will be able to:

§ Group various lipids into the acceptable classifications

§ explain the term ‘blood lipids’

§ discuss the various lipoprotein systems

§ define and explain lipid micelles, monolayers and bilayers

§ discuss the pathway of the oxidation of fats

§ explain the biosynthetic pathway of lipids, phospholipids and sphingolipids

§ list and explain the various types of unsaturated and essential fatty acids

§ discuss and explain the adipose tissue as well as the regulation of fats’ metabolism

§ explain by ketosis and cholesterol metabolism.

3.5 Course Requirements:

-This is a compulsory course for all 300 level students studying Biochemistry. Passing BCH 201 is a prerequisite for registering for this course, in view of this, students are expected to participate in all the course activities and have a minimum of 75% attendance to be able to write the final examination.

-Students are expected to treat the study questions and assignments, as they will be expected to supply answers to them in class when asked.

-They are also expected to have e-mail accounts.

3.6 Methods of Grading:

¡ Continous assessment - 20 %

¡ Practicals/Attendance - 10%

¡ Examinations - 70%

¡ Total - 100%

3.7 Course Delivery Strategies:

Course delivery will be by:

¡ Face-to-face method

¡ Lecture method

¡ Participatory method

¡ Assignments will be given out to students periodically, individually and in group.

4. LECTURE CONTENT

Week 1: Introduction to lipids

¡ Objectives: At the end of the week, students should:

- Be able to define lipids

- Be able to list the functions of lipids

¡ Description:

FIRST HOUR: The course outline will be introduced with emphasis on the objectives.

SECOND HOUR: the definition and importance of lipid will be explained in detail.

¡ Study questions:

1. What are lipids?

2. List five importance of lipids

3. Why are lipids insoluble or partially soluble in water?

4. State the characteristics of lipids.

5. Spingomyelin is an example of lipids. T/F

¡ Reading list:

1. 1,3,5 Biochemistry, Third edition (2005) by Voet and Voet, Wiley, ISBN: 978-0-471-19350-0.

2. 1,3,5Lehninger Principles of Biochemistry, Fourth Edition (2005) by David L. Nelson and Michael M. Cox (pages 343-368)

3. 1,3,5Harper’s Illustrated Biochemistry, (2003) twenty-sixth edition. McGraw-Hill companies limited. ISBN-0-07-121766-5 (pages 92-99, 111-121, 122- 129, 173- 179, 180-189, 197-204 and 205-218)

4. 3 Related texts on the internet.

Week 2: Classification of lipids

¡ Objective: At the end of the week, students should:

--Be able to classify the various lipids

--Be able to draw the structures of the various lipids

--Be able to differentiate between unsaturated and saturated fatty acids

¡ Description:

First hour: Classification of Lipids

Second hour: Structures of the various classes of lipids

¡ Study questions:

1. What are glycoproteins?

2. How will you classify sphingolipids structurally?

3. List the various classes of lipids.

4. Give one example for each class of lipid

5. Give the systematic name, trivial name and the structures of the following fatty acids. i. 18:1 ii. 22:1 iii. 20:4 iv. 22:6

¡ Reading list:





Week 3: Lipoprotein Systems


- be able to classify lipoprotein

- understand the role of lipoprotein in cholesterol transportation

- be able to describe why HDL-Cholesterol is termed good cholesterol while LDL-Cholesterol is termed bad cholesterol

¡ Description:

FIRST HOUR: The lipoprotein system is very important for cholesterol transportation. This process will be explained in detail to the students.

SECOND HOUR: Students will be exposed to the fate of cholesterol ingested or synthesized.

¡ Study questions:

1. List the various classes of lipoprotein.

2. In a tabular form, differentiate between the lipoproteins

3. What active role do lipoproteins play in the transportation of cholesterol?

4. What options are available for the treatment of hypercholesterolemia?

5. Discuss the process of cholesterol transport from the liver to the periphery tissues.

6. Discuss the regulation of cholesterol biosynthesis.

¡ Reading list:

1. 1,3,5 Biochemistry, Third edition (2005) by Voet and Voet, Wiley, ISBN: 978-0-471-19350-0




Week 4: Lipoproteins in relation to diseases

¡ Objective:At the end of the week, students should be able to:

- List some diseases associated with defective lipoprotein metabolism

- Discuss the significance of other lipids present in the blood

¡ Description:

FIRST HOUR: forms of lipids in the blood.

SECOND HOUR: Diseases associated with lipid transportation and their prevention.

¡ Study questions:

1. Discuss the role of lipid in the onset of atherosclerosis.

2. Lipids have been implicated in the development of CHD. Discuss.

3. Lipids are water insoluble, yet they need to be deposited in cells for metabolic purposes. Discuss how this problem is solved biochemically.

4. Write short notes on chylomicrons and high density lipoproteins.

5. Explain the contribution of low density lipoprotein and very low density lipoprotein

to the transport of lipids in the system.

¡ Reading list:

1. 1,3,5 Biochemistry, Third edition(2005) by Voet and Voet, Wiley, ISBN: 978-0-471-19350-0


3. 1,3,5Harper’s Illustrated Biochemistry, (2003) twenty-sixth edition. McGraw-Hill companies limited. ISBN-0-07-121766-5(pages 92-99, 111-121, 122- 129, 173- 179, 180-189, 197-204 and 205-218)


Week 5: Lipid micelles (monolayers), bilayers and liposomes

¡ Objective:At the end of the week, students should:

-Be able to describe what is meant by micelles (monolayers), bilayers and liposomes

-They should be able to give a diagrammatic representation of the three and

-They should be able to differentiate between the three.

¡ Description:

FIRST HOUR: The students will be exposed to the plasma membrane, its components and the proposed arrangements of these components in the membrane, using the fluid mosaic model.

SECOND HOUR: The structures of micelles, bilayers and liposomes will be drawn and short notes written on each.

¡ Study questions:

1. What are micelles?

2. Explain what is meant by lipid monolayers and bilayers.

3. Give diagrammatic representations of micelles, bilayers and liposomes.

4. In your own words, briefly explain the fluid mosaic model for membrane structure.

5. List, draw and explain the three major amphipathic lipid aggregates that form in water.

6. Briefly explain the fluid mosaic model for membrane structure.

¡ Reading list:





Week 6: Oxidation of fats


- Be able to explain the various types of oxidations (β, ω and α) of fats that exist.

- Be able to draw the pathway for complete oxidation of saturated fatty acids

- Be able to oxidize unsaturated fatty acids by identifying the two major reactions and enzymes required for this.

- Be able to draw the pathway for the odd chain fatty acids

¡ Description:

FIRST HOUR: β, ω and α oxidation will be explained to the students

SECOND HOUR: They will be taught the necessary processes involved in the oxidation of saturated, unsaturated and odd chain fatty acids.

¡ Study questions:

1. With the aid of a simple diagram only, show the three stages involved in fatty acid oxidation.

2. Beta oxidation of unsaturated fatty acids has 4 basic steps catalyzed by 4 basic enzymes. Draw a pathway to illustrate this.

3. Acyl CoA synthethase is the enzyme catalyzing the reaction below. Write the product of the reaction.

Fatty acid + CoA + ATP -----à___________________+ AMP +_______

4. Draw the pathway for the breakdown of GM1 to ceramide showing the disease caused by enzyme defect or deficiency

5. Differentiate between hormones and eicosanoids

¡ Reading list:





Week 7: General biosynthesis of phospholipids and sphingolipids


-Be able to highlight the 7 major types of glycerophospholipids

-Be able to list the 5 major types of spingophospholipids.

-Be able to describe the pathway s leading to the biosynthesis of these lipids

¡ Description:

FIRST HOUR: The students will be taught the biosynthesis of each of the 12 lipids and other simpler lipids.

SECOND HOUR: They will be taught to differentiate them using a tabular format. Short notes will also be written on each.

¡ Study questions:

Fill this table: Name of Sphingolipid Name of X Formular of X

Ceramide

Phosphocholine

Cerebroside (N. glycolipid)

Di,tri,tetra (saccharide)

Ganglioside (GM2)

Fill this table: Membrane lipids Hydrophilic unit Hydrophobic unit

Cholesterol

Sphingomyelin

Glycolipid

Phosphoglyceride

¡ Reading list:





Week 8: Unsaturated and essential fatty acids

¡ Objective: At the end of the week, students should be able to:

- Define saturated and unsaturated fatty acids.

- Account for the differences in the properties of saturated and unsaturated fatty acids.

- Define and list the essential fatty acids.

- Draw the structures of the essential fatty acids

- Give some physiological importance of essential fatty acids

¡ Description:

FIRST HOUR: The inability of animals to incorporate double bonds at certain positions in lipids will be discussed. The need for the diet to therefore contain such lipids will be emphasized.

SECOND HOUR: Definition and lists of the essential fatty acids as well as their structures will be dealt with in details. Their physiological functions will also be discussed.

¡ Study questions:

1. Essential fatty acids include oleic and gamma linoleic acids. T/F

2. Draw the structures of the essential fatty acids.

3. Why are these lipids classified as essential?

4. Differentiate using the structures (of two fatty acid each), between saturated and unsaturated fatty acids

5. What are the essential fatty acids.

¡ Reading list:





Week 9: Adipose Tissue


-Be able to define the adipose tissue.

-Be able to explain the mobilization of fatty acids out of the adipose tissue.

-Be able to explain lipolysis.

¡ Description:

FIRST HOUR: The adipose tissue will be defined; the mobilization of fatty acids out

of the adipose tissue will be discussed. The term ‘Lipolysis’ will also be explained.

SECOND HOUR: The students will be taught the effect of a cascade reaction on fatty acid degradation and synthesis/degradation of glycogen. They will also be taught the hormonal regulation of fuel metabolism.

¡ Study questions:

1. Write a short note on the adipose tissue.

2. With the aid of a diagram, illustrate the cascade reaction involved in the mobilization of fatty acid from the adipose tissue.

3. Why are adipose cells unable to phosphorylate endogenous glycerol?

4. Use a simple diagram to illustrate synthesis and degradation of triacylglycerols by adipose tissue.

5. In mammals, the major site of accumulation of triacylglycerol is---------------

¡ Reading list:





Week 10: Regulation of the metabolism of fats


-Be able to explain how fatty acid degradation is regulated.

-Be able to explain the effect of insulin on fuel metabolism

-Be able to explain the effect of epinephrine on fuel metabolism

-Be able to explain the effect of glucagon fuel metabolism

¡ Description:

FIRST HOUR: The effect of such hormones as norepinephrine, epinephrine, Insulin and glucagon will be explained.

SECOND HOUR: The students will be taught the various ways by which fatty acid metabolism can be regulated.

¡ Study questions:

1. Hormones regulate fatty acid metabolism. Discuss.

2. Write short notes on the following hormones: Insulin, Glucagon, epinephrine and Norepinephrin.

3. Discuss the regulation of lipolysis.

4. Insulin stimulates the storage of fuels and the synthesis of proteins in a variety of ways. Discuss.

5. Catecholamine increase the amount of glucose released into the blood by the liver and decrease the utilization of glucose by muscle. T/F-----------

¡ Reading list:





WEEK 11- CONTINUOUS ASSESSMENT TEST

¡ Week 11: Continous Assessment Test

Students will be assessed based on the topics treated in the previous 10 weeks The test shall be essay/paper based.

Week 12: Ketosis


- Be familiar with terms such as ketone bodies, ketosis and ketonuria.

- Be able to list and draw the structure of the ketone bodies

- Understand when and where ketone bodies are produced

- Draw the pathway for synthesis of ketone bodies

¡ Description:

FIRST HOUR: The students will be familiarized with the list and structures of existing ketone bodies. They will be taught when and where ketone bodies are produced.

SECOND HOUR: In times of fasting and starvation, the body finds alternative route for its energy need. At such times, ketone bodies have been found to be very useful. The students will therefore be taught the pathway by which ketone bodies are produced and the effect of excess ketone body in blood and urine.

¡ Study questions:

1. Define ketosis and ketonuria.

2. How is the production of ketone bodies regulated?

3. Draw the structures of the ketone bodies.

4. What enzymes are involved in the production of ketone bodies?

5. In which compartment of the cell are ketone bodies produced?

¡ Reading list:





Week 13: Cholesterol metabolism


- Be familiar with the structure of cholesterol

- Be able to list the importance of cholesterol in the body

- Understand the four stages in the pathway of cholesterol biosynthesis.

- Be able to discuss the treatment options available for treatment of hypercholesterolemia.

- Understand the regulatory mechanism of cholesterol biosynthesis

¡ Description:

FIRST HOUR: the structure of cholesterol will be drawn, the importance of cholesterol in the body will be listed, and the four stages in the pathway of cholesterol biosynthesis will be taught.

SECOND HOUR: Cholesterol to the non scientist is generally thought to be a non useful component of the diet. The topic will discuss the good in cholesterol as well as the bad that can arise due to unregulated synthesis. Lecture will include the location for synthesis, enzymes involved and other metabolites of cholesterol.

¡ Study questions:

1. List the enzymes involved in cholesterol biosynthesis.

2. List the stages of cholesterol biosynthesis.

3. How is cholesterol biosynthesis regulated?

4. Give five importance of cholesterol.

5. What options are available for the treatment of hypercholesterolemia?

¡ Reading list:





Week 14 : Revision

¡ Class Interactive sessions for discussion of problems.

¡ Discussion of the CA test questions and problem solving.

¡ Treatment of problems arising from the practical periods.

Week 15 : Revision continued

¡ Activities of the previous week will be continued.

¡ Students must be well prepared for the examinations by the end of this week.

Study Questions:

1. The release of arachidonate into the cells is catalyzed by --------------------------------- 2. Arachidonate serve as the precursors for a large number of products called------------ 3. --------------------is the final product generated in a reaction catalyzed by lipoxygenase 4. -------------------, ------------------- and ------------------- are the products of cyclization

of arachidonate catalyzed by cyclooxygenase 5. The enzyme that catalyzes formation of double bonds from C9 and above is called---- 6. ------------------- is a precursor of arachodonoyl COA

7. Membrane lipids are in constant motion and three major kinds of lipid motions exist in the membrane. Highlight them.

8. Use a simple pathway to illustrate ω-oxidation of fatty acids.

9. There are two general strategies for attaching head groups to glycerol phospholipids. Explain.

10 The enzymes unique to ω-oxidation of fatty acids are located in which organelle of the liver and kidney?

11. The preferred substrates for omega oxidation are fatty acids of ---------- or -------------Carbon atoms.

12. Operational Definition of Lipids How is the definition of “lipid” different from the types of definitions used for other biomolecules as amino acids, nucleic acids, and proteins?

13 Melting Points of Lipids The melting points of a series of 18-carbon fatty acids are: stearic acid, 69.6 _C; oleic acid, 13.4 _C; linoleic acid, _5 _C; and linolenic acid, _11 _C. (a) What structural aspect of these 18-carbon fatty acids can be correlated with the melting point? Provide a molecular explanation for the trend in melting points. (b) Draw all the possible triacylglycerols that can be constructed from glycerol, palmitic acid, and oleic acid. Rank them in order of increasing melting point. (c) Branched-chain fatty acids are found in some bacterial membrane lipids. Would their presence increase or decrease the fluidity of the membranes (that is, give them a lower or higher melting point)? Why?

14. Preparation of Béarnaise Sauce During the preparation of béarnaise sauce, egg yolks are incorporated into melted butter to stabilize the sauce and avoid separation. The stabilizing agent in the egg yolks is lecithin (phosphatidylcholine). Suggest why this works.

15. Hydrophobic and Hydrophilic Components of Membrane Lipids A common structural feature of membrane lipids is their amphipathic nature. For example, in phosphatidylcholine, the two fatty acid chains are hydrophobic and the

phosphocholine head group is hydrophilic. For each of the following membrane lipids, name the components that serve as the hydrophobic and hydrophilic units: (a) phosphatidylethanolamine; (b) sphingomyelin; (c) galactosylcerebroside; (d) ganglioside; (e) cholesterol.

16. Properties of Lipids and Lipid Bilayers Lipid bilayers formed between two aqueous phases have this important property: they form two-dimensional sheets, the edges of which close upon each other and undergo self-sealing to form liposomes. (a) What properties of lipids are responsible for this property of bilayers? Explain. (b) What are the consequences of this property for the structure of biological membranes?

17. Length of a Fatty Acid Molecule The carbon–carbon bond distance for single- bonded carbons such as those in a saturated fatty acyl chain is about 1.5 Å. Estimate the length of a single molecule of palmitate in its fully extended form. If two molecules of palmitate were placed end to end, how would their total length compare with the thickness of the lipid bilayer in a biological membrane?

18. Temperature Dependence of Lateral Diffusion The experiment described in Figure 11–17 was performed at 37°C. If the experiment were carried out at 10°C, what effect would you expect on the rate of diffusion? Why?

5. GENERAL READING LIST: Suggested Further Reading List:

General Gurr, M.I. & Harwood, J.L. (1991) Lipid Biochemistry: An Introduction, 4th edn, Chapman & Hall, London.A good general resource on lipid structure and metabolism, at the intermediate level. Vance, D.E. & Vance, J.E. (eds) (2002) Biochemistry of Lipids, Lipoproteins, and Membranes, New Comprehensive Biochemistry,Vol. 36, Elsevier Science Publishing Co., Inc., New York. An excellent collection of reviews on various aspects of lipid structure, biosynthesis, and function. Structural Lipids in Membranes Bogdanov, M. & Dowhan, W. (1999) Lipid-assisted protein folding. J. Biol. Chem. 274, 36,827–36,830. A minireview of the role of membrane lipids in the folding of membrane proteins. De Rosa, M. & Gambacorta, A. (1988) The lipids of archaebacteria.Prog. Lipid Res. 27, 153–175. Dowhan, W. (1997) Molecular basis for membrane phospholipid diversity: why are there so many lipids? Annu. Rev. Biochem. 66, 199–232. Gravel, R.A., Kaback, M.M., Proia, R., Sandhoff, K., Suzuki, K., & Suzuki, K. (2001) The GM2 gangliosidoses. In The Metabolic and Molecular Bases of Inherited Disease, 8th edn (Scriver, C.R., Sly, W.S., Childs, B., Beaudet, A.L., Valle, D., Kinzler, K.W., & Vogelstein, B., eds), pp. 3827–3876, McGraw-Hill, Inc., New York. This article is one of many in a four-volume set that contains definitive descriptions of the clinical, biochemical, and genetic aspects of hundreds of human metabolic diseases—an authoritative source and fascinating reading. Hoekstra, D. (ed.) (1994) Cell Lipids, Current Topics in Membranes, Vol. 4, Academic Press, Inc., San Diego. Lipids as Signals, Cofactors, and Pigments

Bell, R.M., Exton, J.H., & Prescott, S.M. (eds) (1996) Lipid Second Messengers, Handbook of Lipid Research, Vol. 8, Plenum Press, New York. Binkley, N.C. & Suttie, J.W. (1995) Vitamin K nutrition and osteoporosis. J. Nutr. 125, 1812–1821. Brigelius-Flohé, R. & Traber, M.G. (1999) Vitamin E: function and metabolism. FASEB J. 13, 1145–1155. Chojnacki, T. & Dallner, G. (1988) The biological role of dolichol. Biochem. J. 251, 1–9. Clouse, S.D. (2002) Brassinosteroid signal transduction: clarifying the pathway from ligand perception to gene expression. Mol. Cell 10, 973–982. Lemmon, M.A. & Ferguson, K.M. (2000) Signal-dependent membrane targeting by pleckstrin homology (PH) domains. Biochem. J. 350, 1–18. Prescott, S.M., Zimmerman, G.A., Stafforini, D.M., & McIntyre, T.M. (2000) Platelet-activating factor and related lipid mediators. Annu. Rev. Biochem. 69, 419–445. Schneiter, R. (1999) Brave little yeast, please guide us to Thebes: sphingolipid function in S. cerevisiae. BioEssays 21, 1004–1010. Suttie, J.W. (1993) Synthesis of vitamin K-dependent proteins. FASEB J. 7, 445–452. Vermeer, C. (1990) _-Carboxyglutamate-containing proteins and the vitamin K-dependent carboxylase. Biochem. J. 266, 625–636. Describes the biochemical basis for the requirement of vitamin K in blood clotting and the importance of carboxylation in the synthesis of the blood-clotting protein thrombin. Viitala, J. & Järnefelt, J. (1985) The red cell surface revisited. Trends Biochem. Sci. 10, 392–395. Includes discussion of the human A, B, and O blood type determinants. Weber, H. (2002) Fatty acid-derived signals in plants. Trends Plant Sci. 7, 217–224. Zittermann, A. (2001) Effects of vitamin K on calcium and bone metabolism. Curr. Opin. Clin. Nutr. Metab. Care 4, 483–487. Composition and Architecture of Membranes Boon, J.M. & Smith, B.D. (2002) Chemical control of phospholipid distribution across bilayer membranes. Med. Res. Rev. 22, 251–281. Intermediate-level review of phospholipid asymmetry and factors that influence it. Dowhan, W. (1997) Molecular basis for membrane phospholipids diversity: why are there so many lipids? Annu. Rev. Biochem. 66, 199–232. Ediden, M. (2002) Lipids on the frontier: a century of cell membrane bilayers. Nat. Rev. Mol. Cell Biol. 4, 414–418. A short review of how the notion of a lipid bilayer membrane was developed and confirmed. Haltia, T. & Freire, E. (1995) Forces and factors that contribute to the structural stability of membrane proteins. Biochim. Biophys. Acta 1241, 295–322. Good discussion of the secondary and tertiary structures of membrane proteins and the factors that stabilize them. von Heijne, G. (1994) Membrane proteins: from sequence to structure. Annu. Rev. Biophys. Biomol. Struct. 23, 167–192. A review of the steps required to predict the structure of an integral protein from its sequence. White, S.H., Ladokhin, A.S., Jayasinghe, S., & Hristova, K. (2001) How membranes shape protein structure. J. Biol. Chem.276, 32,395–32,398. Brief, intermediate-level review of the forces that shape transmembrane helices. Wimley, W.C. (2003) The versatile _ barrel membrane protein. Curr. Opin. Struct. Biol. 13, 1–8. Intermediate-level review.

6. LEGEND: Key for the Reading List:

1. 1 - Available in the University Library

2. 2 - Available in local bookshops

3. 3 - Available on the Web

4. 4 - Personal collection

5. 5 - Departmental library

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