CHAPTER 7: CELL CELL INTERACTIONS
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1 CHAPTER 7: CELL CELL INTERACTIONS CHAPTER SYNOPSIS Cells of multicellular organisms must communicate with one another so that they behave as a coordinated group of cells rather than just a bunch of independent ones. Recall the cliche the left hand doesn t know what the right hand is doing. The signals to which a cell responds are dependent on the kinds of receptor proteins associated with that cell. If the signal is just the right molecular shape, it and the receptor bind eliciting a response somewhere in the cell. There are four main types of cell signaling. Direct contact and paracrine signaling are important in early organismal development. Endocrine signaling via hormones provides widespread response in both plants and animals. Synaptic signaling found in animal nervous systems produces more localized responses at the chemical synapse between the neuron and the receptor cell. Intracellular receptors are small molecules that are able to pass through the plasma membrane of the target cell. Nitrous oxide is one example. It activates the enzyme that catalyzes synthesis of cyclic GMP. A superfamily of steroid hormone receptors have specific DNA binding sites normally occupied by an inhibitory protein. When the signal molecule binds to another site on the receptor the inhibitor is released. The receptor then binds to DNA to activate or suppress a certain gene. There are three superfamilies of cell surface receptors. The signals associated with these receptors bind to receptor proteins on the cell surface. Thus an extracellular signal is converted to an intracellular signal. In chemically-gated ion channels the receptor is a transmembrane protein that winds back and forth through the membrane several times (called a multi-pass protein). The center of this protein forms a channel through which specific ions can pass. Enzymatic receptors are linked to or themselves act as enzymes. Most of these are protein kinases; they add phosphate groups to proteins. They are single pass transmembrane proteins where the region that binds with the signal molecule is located outside the cell and the portion that initiates enzymatic activity is found within it. G protein-linked receptors are seven-pass transmembrane proteins that become activated when they bind to GTP. These receptors are important because they provide the mechanism of action for over half of the therapeutic drugs currently in use. Second messengers are small molecules or ions that alter the shape and behavior of receptor proteins to relay the signal message to enzymes or genes within a cell. camp is the most widely used second messenger in animal cells. Ultimately the camp binds to A-kinase, activating it to phosphorylate certain cell proteins. Calcium is normally sequestered outside a cell or within its endoplasmic reticulum. With the proper G protein signal inositol triphosphate is eventually produced, which opens calcium channels in the ER membrane. This influx of calcium triggers many activities. In most cases, a cellular signal is too insignificant to result in an adequate cellular response. Protein kinase cascades amplify the signal. In vision for example, a single light-activated rhodopsin molecule activates many transducin molecules that further split 10 5 cyclic GMP molecules. The expression of cell identity results from the presence of unique cell surface markers, some of these, like those associated with ABO blood types, are glycolipids. Others are proteins anchored in the plasma membrane the major histocompatibility complex (MHC) proteins, for example. Cells in a multicellular organism also establish physical connections between themselves. Tight junctions are an example of these within animals. Anchoring junctions are common in tissues exposed to stress. In cadherin junctions, desmosomes for example, a cell s cytoskeleton is attached to that of other cells or to the extracellular matrix. Adherens junctions connect actin filaments of cells via linking proteins called integrins. Communicating junctions, gap junctions in animals, and plasmodesmata in plants, allow the passage of ions or molecules from one cell to another. 54
2 CELL-CELL INTERACTIONS 55 CHAPTER OBJECTIVES Know what receptor proteins are, where they are located, and what they do. Explain the amplification process associated with protein kinase cascades. Describe the four types of cell signaling. Differentiate between intracellular receptors and cell surface receptors in terms of function. Know the three cell surface receptor superfamilies, the basic structures of each and how each functions to convert an extracellular signal to an internal one. Know what cell surface markers are, where they are located, and give examples. Describe the three primary kinds of junctions that exist between cells. Compare and contrast communicating junctions, gap junctions, and plasmodesmata. Understand how camp and calcium function as second messengers and why they are necessary. KEY TERMS adenyl cyclase adherens junction anchoring junction autocrine signal calcium camp cell surface receptor cell junction chemical synapse communicating junction desmosome endocrine signal G protein hormone intracellular receptor ion channel neurotransmitter nitric oxide paracrine signal protein kinase receptor protein second messenger tissue tight junction CHAPTER OUTLINE 7.0 Introduction I. CELLS OF MULTICELLULAR ORGANISMS TOUCH AND COMMUNICATE WITH EACH OTHER fig 7.1 A. Send and Receive Chemical Signals B. Coordinate Activities to Behave as Group, Not Individuals 7.1 Cells signal one another with chemicals I. RECEPTOR PROTEINS AND SIGNALING BETWEEN CELLS A. Cellular Communication Is Common in Nature 1. Occurs in all multicellular organisms 2. Uses a variety of molecules a. Includes peptides, large proteins, amino acids, nucleotides, steroids, lipids b. Even includes NO, a dissolved gas c. Some are attached to cell surface d. Some are released from cell
3 56 CHAPTER 7 B. Cell Surface Receptors 1. Cell responds to certain signals, ignores the rest fig Feat accomplished by receptor proteins a. Have three dimensional shape b. Signal molecule binds to receptor if correct shape c. Induces shape change in receptor protein d. Results in response by cell C. The Hunt for Receptor Proteins 1. Characterizing a small number of receptor proteins difficult 2. Recent techniques have fostered recent advances a. Monoclonal antibodies: Used to bind to particular receptors b. Gene analysis: Genetic engineering identifies and isolates receptor protein genes II. TYPES OF CELL SIGNALING A. Cells Communicate Through Four Basic Mechanisms fig Associated with distance between cells 2. Some cells send signals to selves a. Called autocrine signaling b. Plays important role in reinforcing developmental changes B. Direct Contact 1. Cells are very close 2. Molecules of plasma membrane bind in specific ways 3. Example: Cell interaction in early development fig 7.3a C. Paracrine Signaling 1. Molecules released by cells and taken up by neighboring cells 2. Paracrine signals are short-lived with local effects fig 7.3b 3. Plays important role in early development D. Endocrine Signaling 1. Released signal molecule collected and distributed via blood stream 2. Molecules called hormones, signaling is endocrine fig 7.3c 3. Used by animals and plants E. Synaptic Signaling 1. Nervous system s neurons produce neurotransmitters 2. Released from neurons close to the target cells, persist briefly fig 7.3d 3. Site of release called chemical synapse 7.2 Proteins in the cell and on its surface receive signals from other cells I. INTRACELLULAR RECEPTORS A. Common Elements of Cell Signaling Pathways 1. Intracellular receptors pass through target cell plasma membrane 2. Nature of receptors that receive the signal tbl Function of intracellular receptors fig 7.4 a. Lipid-soluble or small molecules pass directly across membrane b. Bind to receptors in cytoplasm or nucleus c. Trigger a variety of responses
4 CELL-CELL INTERACTIONS 57 B. Receptors that Act as Gene Regulators 1. Include similarly structured steroid hormone receptors 2. Genes may be evolved from single ancestral gene 3. Grouped in intracellular receptor superfamily 4. Each receptor has DNA binding site occupied by inhibitory protein fig 7.5 a. Signal molecule binding to another site on receptor releases inhibitor b. Receptor binds then to DNA to activate or suppress gene 5. Lipid-soluble signals last longer in blood than water-soluble ones 6. Target cell s response can vary enormously a. Binding site on target DNA differs from cell to cell affecting different genes b. Most eukaryotic genes have complex controls C. Receptors that Act as Enzymes 1. Example: Nitrous oxide (NO) gas 2. Binds to guanylyl cyclase in neighboring cells 3. Activated enzyme catalyzes synthesis of cyclic GMP 4. Recently recognized as signal molecule in vertebrates a. NO initiated response relaxes smooth muscle surrounding blood vessels b. Blood vessels expand, increasing blood flow II. CELL SURFACE RECEPTORS A. Most Signal Molecules Are Water-Soluble 1. Cell surface receptors cannot diffuse through cell membranes fig Signals bind to receptor proteins on cell surface 3. Convert extracellular signal to intracellular signal 4. Produces change in cell s cytoplasm 5. Include three superfamilies B. Chemically Gated Ion Channels 1. Receptor is multi-pass transmembrane protein fig 7.6a 2. Winds across plasma membrane several times 3. Center of protein forms a pore through which ions can pass 4. Ion channel opens or closes when neurotransmitter binds to protein a. Called chemical gating b. Type of ion determined by three-dimensional shape of ion channel C. Enzymatic Receptors 1. Acts as or are directly linked to enzymes fig 7.6b 2. Binding between signal molecule and receptor activates the enzyme 3. Most are protein kinases, add phosphate groups to proteins 4. Single pass transmembrane protein a. Signal molecule binds outside cell b. Portion initiating enzyme activity is in cell s cytoplasm D. G Protein-Linked Receptors 1. GTP binding G protein assists membrane-bound enzymes or ion channels fig How G protein-linked receptors work a. G proteins are mediators that initiate cytoplasmic signals b. Link cell surface receptor and cytoplasmic signal pathways 1) Signal has relatively short life span determined by GTP 2) Signal arrives, a G protein within the G protein-linked receptor site c. Signal molecule binds to receptor, receptor changes shape 1) G protein twists and binds to GTP
5 58 CHAPTER 7 2) G protein-gtp complex diffuses away from receptor d. Activated G protein complex can initiate other events 1) Activation is short-lived 2) GTP has life span of seconds to minutes e. G proteins can activate many pathways, but in transient manner 1) Must be continual source of signals for pathway to stay on 2) When external signal stops, pathway shuts down 3. Largest family of cell surface receptors a. More than 100 identified b. All have similar structure, may be derived from same ancestral sequence c. All composed of seven-pass transmembrane protein fig Evolutionary origin of G protein-linked receptors a. Same seven-pass structure in a variety of systems b. Light-activated bacteriorhodopsin proton pump of bacterial photosynthesis c. Yeast mating factor protein recognition receptor d. Various sensory receptors including vertebrate rhodopsin 5. Discovery of G proteins a. Theorized by Rodbell, isolated and purified by Gilman b. Involved in mechanism of half of medicines currently in use c. Further investigation should show how cells communicate in general 7.3 Follow the journey of information into the cell I. INITIATING THE INTRACELLULAR SIGNAL A. Second Messengers Relay Message 1. Also called intracellular mediators 2. Small molecules or ions that change shape and behavior of receptor proteins 3. Include cyclic adenosine monophosphate (camp) and calcium B. camp 1. Used as second messenger by all known animal cells fig Example: Epinephrine binding to G protein-linked β-adrenergic receptor a. Binding epinephrine activates G protein b. Enzyme adenylyl cyclase produces large amounts of camp in target cell fig 7.9a 1) camp binds to α-kinase 2) In muscle cells, activates it to phosphorylate cell proteins c. Action dependent on cell type, in muscle stimulates glycogen to glucose C. Calcium 1. Ca ++ levels in cytoplasm low, high outside cell and in ER 2. Chemically-gated calcium channels in ER membrane act as switches a. Influx of Ca ++ from inside ER to cytoplasm triggers many activities b. Skeletal muscles contract, some endocrine cells release hormones c. Receptor activates G protein, which activates phospholipase C enzyme d. Phospholipase C catalyzes production of inositol triphosphate (IP 3 ) e. IP 3 binds to Ca ++ channels opening them fig 7.9b 3. Also initiates response by binding to calmodulin fig 7.10 II. AMPLIFYING THE SIGNAL: PROTEIN KINASE CASCADES A. Receptors at Surface Receive Signal, But Response Is Elsewhere 1. Second messengers relay signal to enzymes or genes 2. Most receptors use other protein messengers to amplify signal to nucleus
6 CELL-CELL INTERACTIONS 59 B. Mechanism of the Amplification Process 1. Receptor phosphorylates stage-one protein in one of 2 ways a. Directly adds phosphate b. Activates G protein to activate a second phosphorylating protein 2. These in turn activate stage-two, then stage-three proteins fig Signal amplified since one signal at each step initiates multiple proteins C. The Vision Amplification Cascade 1. Single light-activated rhodopsin activates many transducin molecules a. Each transducin causes modification of cyclic GMP fig 7.12 b. One rhodopsin ultimately causes split of 10 5 cyclic GMPs fig Rod cells can detect brief flashes of only 5 photons D. The Cell Division Amplification Cascade 1. Cell division controlled by receptor that acts as a protein kinase 2. Receptor phosphorylates ras protein 3. Ras in turn activates multiple phosphorylation cascades 4. Hyperactive ras gene (as in cancer) results in uncontrolled cell division 7.4 Cell surface proteins mediate cell cell interactions I. THE EXPRESSION OF CELL IDENTITY A. Tissues Are a Fundamental Property of Multicellular Organisms 1. All cells within a tissue are identified as members of that tissue 2. Identification results from the presence of unique cell surface markers B. Tissue-Specific Identity Markers 1. Glycolipids a. Lipids with carbohydrate heads b. Markers on surface of red blood cells identify A, B, O blood types c. Cell populations of glycolipids change as cells differentiate 2. MHC Proteins a. Immune system cell surface markers distinguish between self and not self b. Immune system self marker proteins 1) Major histocompatibility complex proteins 2) Single pass proteins anchored in the plasma membrane 3) Many are members of immunoglobulin receptor superfamily fig 7.14 c. Immune system cells inspect other cells, destroy ones with not self markers II. INTERCELLULAR ADHESION A. Cell Junctions Are Long-Lasting Physical Connections Between Cells fig Nature of the connection determines what tissue is like 2. Tissue function dependent on how individual cells arranged within it III. TIGHT JUNCTIONS A. Cell Junctions Are Divided into Three Categories fig Connect adjacent cells to prevent small molecules from leaking fig Cells act as wall within an organ 3. Molecules sequestered within a region
7 60 CHAPTER 7 B. Creating Sheets of Cells 1. Cells lining digestive tract only one cell layer thick 2. One surface faces inside, other surface faces extracellular space with blood vessels 3. Tight junctions encircle each cell in sheet-like belt 4. Ensure that materials pass through cells, not between cells C. Partitioning the Sheet 1. Partition plasma membranes of lining cells into separate compartments 2. Nutrient transport proteins must stay in proper orientation to function 3. Segregate different proteins on opposite sides of sheet, can t drift between IV. ANCHORING JUNCTIONS A. Mechanical Attachment 1. Attach cell s cytoskeleton to that of other cells or extracellular matrix 2. Common in sheets of tissues exposed to stress-like muscle, skin epithelium B. Cadherin and Intermediate Filaments: Desmosomes 1. Desmosomes connect cytoskeletons of adjacent cells fig Hemidesmosomes anchor epithelial cells to basement membrane 3. Cadherin single-pass transmembrane glycoproteins create critical link a. Cytoplasmic end linked to intermediate filaments b. Other end projects through membrane links to cadherin of next cell 4. More secure than connection to free-floating membrane proteins C. Cadherin and Actin Filaments 1. Cadherins also connect to cell s actin framework, less stable connection fig Migration of neurons in development 3. May provide roadmap for cells to find their destination D. Integrin-Mediated Links 1. Adherens junctions fig 7.20 a. One side connects to actin filaments of a cell b. Other side linked to neighbor cell or extracellular matrix 2. Linking proteins belong to superfamily of receptors called integrins 3. Integrin is transmembrane protein made of two glycoprotein subunits 4. Component of matrix that cell binds to depends on integrin combination V. COMMUNICATING JUNCTIONS A. Direct Cellular Connections 1. Some cells communicate through direct connections between cells 2. Pass chemical signals from one cell to another 3. Permit small molecules or ions to pass from cell to cell B. Gap Junctions In Animals fig Composed of connexons a. Six identical transmembrane proteins arranged in a circle b. Connexons of two adjacent cells must be perfectly aligned c. Small molecules like sugars and amino acids can pass 2. Are dynamic structures that can open and close a. Respond to factors like Ca ++ and H + ions b. If cell-damaged ions flow in, close gap junctions, seal off cell
8 CELL-CELL INTERACTIONS 61 C. Plasmodesmata in Plants fig Occur only at gaps in cell walls, provide cytoplasmic connections between cells 2. Function like animal cell gap junctions 3. Are lined with plasma membrane 4. Contain central tubule connecting ER of both cells 5. Play role in integrating activities of plant body INSTRUCTIONAL STRATEGY PRESENTATION ASSISTANCE: Students need to be secure in the material from the previous chapter to progress in this one. There is a lot of new terminology. Flash cards will help the students learn this information making them as much as using them. Table 7.1 is an invaluable summary of the chapter, but too many students will use it as a crutch and just memorize it. It s much better if they make their own table(s). The best part of the learning process is in the organization and gathering of information. Re-emphasize the importance of molecular binding to induce three-dimensional shape changes to effect a variety of changes within the cell. Stress the importance of protein kinases. The topic will come up again in gene expression and regulation. It s amazing how a simple addition of a PO 4 group can do so much in a cell! Don t let your students confuse desmosomes and plasmodesmata. They sound similar, but are very different structures. VISUAL RESOURCES: The amplification analogy in the book is very straight forward, but students will quickly identify with the shampoo commercial where you tell two friends and they tell two friends... while they see on TV one person, then two, then four, and so on. You could also do a class demo on this, distributing note cards, rubber bands, or other small, inexpensive objects. Make models of various cell cell interactions using plastic, paper, string, straws, etc. Better yet, have your students make them, collect them, choose the best, and you will have models to show in class next semester!
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