Phospholipids and Membrane Structure © Jones & Bartlett Learning, LLC © Jones & Bartlett Learning, LLC NOT for SALE OR DISTRIBUTION NOT for SALE OR DISTRIBUTION
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NOT FOR SALE OR DISTRIBUTION. © Jones & Bartlett Learning, LLC © Jones & Bartlett Learning, LLC NOT FOR SALE OR DISTRIBUTION NOT FOR SALE OR DISTRIBUTION © Jones & Bartlett Learning, LLC © Jones & Bartlett Learning, LLC NOT FOR SALE OR DISTRIBUTION NOT FOR SALE OR DISTRIBUTION © Jones & Bartlett Learning, LLC © Jones & Bartlett Learning, LLC NOT FOR SALE OR DISTRIBUTION NOT FOR SALE OR DISTRIBUTION © Jones & Bartlett Learning, LLC © Jones & Bartlett Learning, LLC NOT FOR SALE OR DISTRIBUTION NOT FOR SALE OR DISTRIBUTION © Jones & Bartlett Learning, LLC © Jones & Bartlett4 Learning, LLC NOT FOR SALE OR DISTRIBUTION NOT FOR SALE OR DISTRIBUTION Phospholipids and Membrane Structure © Jones & Bartlett Learning, LLC © Jones & Bartlett Learning, LLC NOT FOR SALE OR DISTRIBUTION NOT FOR SALE OR DISTRIBUTION 4.1 The Big Picture 4.2 Phospholipids Are the Basic Building Blocks of Cellular Membranes © Jones & Bartlett Learning, LLC © Jones & Bartlett Learning, LLC NOT FOR SALE4.3 TheOR Fluid–MosaicDISTRIBUTION Model Explains How PhospholipidsNOT FOR SALE OR DISTRIBUTION and Proteins Interact within a Cellular Membrane 4.4 The Smooth Endoplasmic Reticulum and Golgi Apparatus Build Most Eukaryotic Cellular Membrane Components© Jones & Bartlett Learning, LLC © Jones & Bartlett Learning, LLC 4.5 Chapter SummaryNOT FOR SALE OR DISTRIBUTION NOT FOR SALE OR DISTRIBUTION © Jones & Bartlett Learning, LLC © Jones & Bartlett Learning, LLC NOT FOR SALE OR DISTRIBUTION NOT FOR SALE OR DISTRIBUTION © Jones & Bartlett Learning, LLC © Jones & Bartlett Learning, LLC NOT FOR SALE OR DISTRIBUTION NOT FOR SALE OR DISTRIBUTION 117 3RD PAGES © Jones & Bartlett Learning, LLC. NOT FOR SALE OR DISTRIBUTION. © Jones & Bartlett Learning, LLC4.1 The Big Picture© Jones & Bartlett Learning, LLC NOT FOR SALE OR DISTRIBUTIONThe purpose of this chapterNOT is to FOR introduce SALE the OR general DISTRIBUTION structure and assembly of mem- branes. The presence of at least one membrane is required for any organism to be consid- ered alive, but this does not mean that all membranes are alike. Recall from Chapter 1 that, despite its relatively simple structure, the carbon atom is capable of forming a multitude © Jonesof different & Bartlett molecules; Learning, this is LLCespecially evident when one© considers Jones the& Bartlettmolecules Learning,that LLC NOT FORcompose SALE a membrane. OR DISTRIBUTION Unlike nucleic acids and proteins, NOTwhich FORare built SALE from ORsimple DISTRIBUTION nucleotide and amino acid subunits, membranes are much larger structures composed of a tremendous variety of molecules. Phospholipids are some of most abundant mol- ecules in membranes, so in this chapter we will focus on them; our goal is to discuss © Jones & Bartlett theLearning, common featuresLLC shared by all membranes,© Jones and &thus Bartlett complete Learning, our survey LLC of the NOT FOR SALE ORbuilding DISTRIBUTION blocks of cells. We will reserve theNOT details FOR concerning SALE membrane OR DISTRIBUTION specialization for later chapters. This chapter contains three major sections: Unlike nucleic acids and proteins, phospholipids are not composed of linear © Jones & Bartlett Learning, LLC polymers of a small© Jonesgroup of &subunits; Bartlett instead, Learning, they are branchedLLC molecules, and NOT FOR SALE OR DISTRIBUTION each branch is oftenNOT unique, FOR making SALE it OR nearly DISTRIBUTION impossible to memorize them all. Because of this, our strategy for learning phospholipid structure will rely heavily on generalizations, such that we can more easily understand the structural details of specific phospholipids in later chapters. © Jones & BartlettThe fluid–mosaic Learning, modelLLC explains how phospholipids© Jones and other& Bartlett molecules Learning, are LLC arranged to yield a biologically functional membrane. Our focus will be on the NOT FOR SALE OR DISTRIBUTION NOT FOR SALE OR DISTRIBUTION general concepts that apply to all membranes. Understanding the implications of these concepts will be crucial when we begin discussing specific membranes in later chapters, so this idea will likely require more attention than the first. Phospholipid synthesis and membrane formation occur in an assembly-line-like © Jones & Bartlett Learning,fashion. LLC This concerns the molecular© Jones mechanisms & Bartlett cells useLearning, to build phospho- LLC NOT FOR SALE OR DISTRIBUTIONlipids and organize them into functionalNOT FOR membranes. SALE NoticeOR DISTRIBUTION that the first two ideas are focused on what these structures are, while this idea addresses how they are made. It is important to understand the sequence of events, and the reasons why each step is necessary. Fully understanding this information will require © Jones & Bartlett Learning, LLC solid comprehension© Jones of the basic& Bartlett chemistry Learning, we discussed LLC in Chapter 1, because NOT FOR SALE OR DISTRIBUTION we will be discussingNOT enzymatic FOR SALE reactions OR that DISTRIBUTION give rise to the structurally complex molecules, including phospholipids, that compose membranes. © Jones4.2 & Bartlett Phospholipids Learning, LLC Are the Basic© Jones Building & Bartlett Learning, LLC NOT FOR SALEBlocks OR DISTRIBUTION of Cellular MembranesNOT FOR SALE OR DISTRIBUTION Key ConCeptS © Jones & Bartlett Learning, of the four majorLLC classes of molecules that serve© Jones as the building & Bartlett blocks for Learning, cells (sugars, LLCnucleic acids, proteins, phospholipids), phospholipids are the most structurally complex. NOT FOR SALE OR DISTRIBUTION phosphoglycerides are the most common phospholipids;NOT FOR they SALE are composed OR DISTRIBUTION of two fatty acyl groups and a polar head group attached to a glycerol backbone. the length and degree of saturation of the fatty acyl tails can vary between different phospholipids. the primary function of phospholipids is to form the lipid bilayer that is characteristic of all © Jones & Bartlett Learning, LLC cellular membranes. © Jones & Bartlett Learning, LLC phospholipid bilayers are selectively permeable to solutes. NOT FOR SALE OR DISTRIBUTION NOT FOR SALE OR DISTRIBUTION 118 Chapter 4 phospholipids and Membrane Structure 3RD PAGES 3RD PAGES © Jones & Bartlett Learning, LLC. NOT FOR SALE OR DISTRIBUTION. © Jones Phospholipids& Bartlett Learning, are structurally LLC and functionally different© from Jones the &other Bartlett cellular Learning,building LLC blocks we have discussed in Chapters 1, 2, and 3. Before we begin our detailed examina- NOT FOR SALE OR DISTRIBUTION NOT FOR SALE OR DISTRIBUTION tion of phospholipids, let’s compare their structural properties to these other basic building blocks. In Chapter 2, we learned that nucleic acids are, structurally, some of the simplest biomolecules in cells; they are composed of only four different nucleotide subunits, joined into linear polymers by one type of chemical bond (the phosphoester bond), and stabi- lized as linear pairs by© hydrogen Jones bonds.& Bartlett The complexityLearning, of LLCnucleic acids arises not from© Jones & Bartlett Learning, LLC their basic structure, butNOT from FOR their SALE tremendous OR sizeDISTRIBUTION—the average lengths of prokaryotic NOT FOR SALE OR DISTRIBUTION and eukaryotic genes are approximately 900 and 1,300 base pairs, respectively. Structures of this size can adopt a wide variety of shapes, giving rise to the different folded forms of nucleoid and chromatin (see Chapter 2, DNA Packaging Is Hierarchical). The© Jones same pattern & Bartlett is evident Learning, in proteins. LLC Polypeptides are composed© ofJones 20 different & Bartlett Learning, LLC aminoNOT acids, FOR joined SALE into linear OR polymers DISTRIBUTION by one type of chemical bond (theNOT peptide FOR bond), SALE OR DISTRIBUTION and stabilized by five different types of chemical bonds (see Chapter 3, Five Classes of Chemical Bonds Stabilize Protein Structure). The average length of a protein ranges from ~250 to ~300 to ~470 amino acids in archaea, prokaryotes, and eukaryotes, respectively. © Jones &Notice Bartlett that over Learning, the course LLCof evolution from archaea to ©eukaryotes, Jones &the Bartlett length of Learning,an aver- LLC NOT FORage SALE protein OR nearly DISTRIBUTION doubled. NOT FOR SALE OR DISTRIBUTION The pattern continues for sugars. Oligosaccharides and polysaccharides are some of the most diverse types of biomolecules, yet most of them are composed of fewer than 20 monosaccharide subunits joined by a small number of bonds (principally the glycosidic bond, a form of ether bond). But size alone does not account for the tremendous diversity © Jones & Bartlett Learning, LLC © Jones & Bartlett Learning, LLC of oligosaccharides and polysaccharides. Cells employ two additional strategies for increas- NOT FOR SALE OR DISTRIBUTION NOT FOR SALE OR DISTRIBUTION ing variation in these structures: branching and chemical modification. In branching, sugar subunits in these polymers form polygonal, ring-like structures, and the carbon atoms that form most of the vertices in these polygons contain functional groups capable of forming covalent bonds with similar groups on other sugars. This means that a single sugar sub- unit (a© monosaccharide)Jones & Bartlett can form Learning, bonds with LLC two or more neighboring sugars,© Jones forming & Bartlett a Learning, LLC branchedNOT network FOR SALE(see Figure OR 1-14). DISTRIBUTION This way, the same number of monosaccharidesNOT FOR SALEcan OR DISTRIBUTION be combined into multiple combinations, increasing