BIOLOGY. Cell Communication CAMPBELL. Reece Urry Cain Wasserman Minorsky Jackson. Lecture Presentation by Nicole Tunbridge and Kathleen Fitzpatrick

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1 CAMPBELL BIOLOGY TENTH EDITION Reece Urry Cain Wasserman Minorsky Jackson 11 Cell Communication Lecture Presentation by Nicole Tunbridge and Kathleen Fitzpatrick

2 Cellular Messaging Cells can signal to each other and interpret the signals they receive from other cells and the environment Signals are most often chemicals The same small set of cell signaling mechanisms shows up in diverse species and processes

3 Figure 11.1

4 Figure 11.1a Epinephrine

5 Concept 11.1: External signals are converted to responses within the cell Communication among microorganisms provides some insight into how cells send, receive, and respond to signals

6 Evolution of Cell Signaling The yeast, Saccharomyces cerevisiae, has two mating types, a and Cells of different mating types locate each other via secreted factors specific to each type Signal transduction pathways convert signals received at a cell s surface into cellular responses The molecular details of signal transduction in yeast and mammals are strikingly similar

7 Figure Exchange of mating factors Receptor a α factor α Yeast cell, mating type a a factor Yeast cell, mating type α

8 Figure Exchange of mating factors Receptor a α factor α Yeast cell, mating type a a factor Yeast cell, mating type α 2 Mating a α

9 Figure Exchange of mating factors Receptor a α factor α Yeast cell, mating type a a factor Yeast cell, mating type α 2 Mating a α 3 New a/ cell a/ α

10 Pathway similarities suggest that ancestral signaling molecules that evolved in prokaryotes and single-celled eukaryotes were adopted for use in their multicellular descendants Cell signaling is critical in the microbial world A concentration of signaling molecules allows bacteria to sense local population density in a process called quorum sensing

11 Figure Individual rod-shaped cells 0.5 mm 2 2 Aggregation in progress 3 Spore-forming structure (fruiting body) 2.5 mm Fruiting bodies

12 Figure 11.3a 0.5 mm 1 Individual rod-shaped cells

13 Figure 11.3b 0.5 mm 2 Aggregation in progress

14 Figure 11.3c 0.5 mm 3 Spore-forming structure (fruiting body)

15 Figure 11.3d 2.5 mm Fruiting bodies

16 Local and Long-Distance Signaling Cells in a multicellular organism communicate via signaling molecules In local signaling, animal cells may communicate by direct contact Animal and plant cells have cell junctions that directly connect the cytoplasm of adjacent cells Signaling substances in the cytosol can pass freely between adjacent cells

17 Figure 11.4 Plasma membranes Cell wall Gap junctions between animal cells Plasmodesmata between plant cells (a) Cell junctions (b) Cell-cell recognition

18 In many other cases, animal cells communicate using secreted messenger molecules that travel only short distances Growth factors, which stimulate nearby target cells to grow and divide, are one class of such local regulators in animals This type of local signaling in animals is called paracrine signaling

19 Synaptic signaling occurs in the animal nervous system when a neurotransmitter is released in response to an electric signal Local signaling in plants is not well understood beyond communication between plasmodesmata

20 In long-distance signaling, plants and animals use chemicals called hormones Hormonal signaling in animals is called endocrine signaling; specialized cells release hormones, which travel to target cells via the circulatory system The ability of a cell to respond to a signal depends on whether or not it has a receptor specific to that signal

21 Figure 11.5 Local signaling Secreting cell Target cells Electrical signal triggers release of neurotransmitter. Neurotransmitter diffuses across synapse. Local regulator (a) Paracrine signaling Long-distance signaling Secretory vesicles Endocrine cell (b) Synaptic signaling Target cell specifically binds hormone. Target cell Hormone travels in bloodstream. (c) Endocrine (hormonal) signaling Blood vessel

22 Figure 11.5a Local signaling Target cells Secreting cell Local regulator (a) Paracrine signaling Secretory vesicles

23 Figure 11.5b Local signaling Electrical signal triggers release of neurotransmitter. Neurotransmitter diffuses across synapse. Target cell (b) Synaptic signaling

24 Figure 11.5c Long-distance signaling Endocrine cell Target cell specifically binds hormone. Hormone travels in bloodstream. Blood vessel (c) Endocrine (hormonal) signaling

25 The Three Stages of Cell Signaling: A Preview Earl W. Sutherland discovered how the hormone epinephrine acts on cells Sutherland suggested that cells receiving signals went through three processes Reception Transduction Response

26 In reception, the target cell detects a signaling molecule that binds to a receptor protein on the cell surface In transduction, the binding of the signaling molecule alters the receptor and initiates a signal transduction pathway; transduction often occurs in a series of steps In response, the transduced signal triggers a specific response in the target cell

27 Figure EXTRACELLULAR FLUID 1 Receptor Reception Plasma membrane CYTOPLASM Signaling molecule

28 Figure EXTRACELLULAR FLUID Plasma membrane CYTOPLASM Receptor 1 Reception 2 Transduction Relay molecules Signaling molecule

29 Figure EXTRACELLULAR FLUID Plasma membrane CYTOPLASM Receptor 1 Reception 2 Transduction Relay molecules 3 Response Activation of cellular response Signaling molecule

30 Animation: Overview of Cell Signaling

31 Concept 11.2: Reception: A signaling molecule binds to a receptor protein, causing it to change shape The binding between a signal molecule (ligand) and receptor is highly specific A shape change in a receptor is often the initial transduction of the signal Most signal receptors are plasma membrane proteins

32 Receptors in the Plasma Membrane G protein-coupled receptors (GPCRs) are the largest family of cell-surface receptors Most water-soluble signal molecules bind to specific sites on receptor proteins that span the plasma membrane

33 Figure 11.7 β 2 -adrenergic receptors Molecule mimicking ligand Plasma membrane Cholesterol

34 There are three main types of membrane receptors G protein-coupled receptors Receptor tyrosine kinases Ion channel receptors

35 G protein-coupled receptors (GPCRs) are cell surface transmembrane receptors that work with the help of a G protein G proteins bind the energy-rich GTP G proteins are all very similar in structure GPCR systems are extremely widespread and diverse in their functions

36 Receptor tyrosine kinases (RTKs) are membrane receptors that attach phosphates to tyrosines A receptor tyrosine kinase can trigger multiple signal transduction pathways at once Abnormal functioning of RTKs is associated with many types of cancers

37 A ligand-gated ion channel receptor acts as a gate when the receptor changes shape When a signal molecule binds as a ligand to the receptor, the gate allows specific ions, such as Na + or Ca 2+, through a channel in the receptor

38 Figure 11.8a Signaling molecule binding site Segment that interacts with G proteins G protein-coupled receptor

39 Figure 11.8b G protein-coupled receptor Plasma membrane Activated receptor Signaling molecule Inactive enzyme CYTOPLASM GDP G protein (inactive) Enzyme GDP GTP GDP GTP 1 2 Activated enzyme GTP GDP P i Cellular response 3 4

40 Figure 11.8ba G protein-coupled receptor Plasma membrane CYTOPLASM 1 Activated receptor GDP G protein (inactive) Signaling molecule Enzyme Inactive enzyme GDP GTP GDP GTP 2

41 Figure 11.8bb Activated enzyme GTP Cellular response 3 GDP P i 4

42 Figure 11.8c Signaling molecule (ligand) helix in the membrane Ligand-binding site Signaling molecule osines CYTOPLASM Receptor tyrosine kinase proteins (inactive monomers) Dimer 1 2 Activated relay proteins 6 ATP 6 ADP P P P P P P P P P P P P Cellular response 1 Cellular response 2 Activated tyrosine kinase regions (unphosphorylated dimer) Fully activated receptor tyrosine kinase (phosphorylated dimer) Inactive relay proteins 3 4

43 Figure 11.8ca Signaling molecule (ligand) helix in the membrane Ligand-binding site osines CYTOPLASM Receptor tyrosine kinase proteins (inactive monomers) 1

44 Figure 11.8cb Signaling molecule Dimer 2

45 Figure 11.8cc 6 ATP 6 ADP P P P P P P Activated tyrosine kinase regions (unphosphorylated dimer) Fully activated receptor tyrosine kinase (phosphorylated dimer) 3

46 Figure 11.8cd Activated relay proteins P P P P P P Cellular response 1 Cellular response 2 Inactive relay proteins 4

47 Figure 11.8d-1 1 Signaling molecule (ligand) Gate closed Ions Ligand-gated ion channel receptor Plasma membrane

48 Figure 11.8d-2 1 Signaling molecule (ligand) Gate closed Ions 2 Gate open Ligand-gated ion channel receptor Plasma membrane Cellular response

49 Figure 11.8d-3 1 Signaling molecule (ligand) Gate closed Ions 2 Gate open Ligand-gated ion channel receptor Plasma membrane Cellular response 3 Gate closed

50 Intracellular Receptors Intracellular receptor proteins are found in the cytoplasm or nucleus of target cells Small or hydrophobic chemical messengers can readily cross the membrane and activate receptors Examples of hydrophobic messengers are the steroid and thyroid hormones of animals An activated hormone-receptor complex can act as a transcription factor, turning on specific genes

51 Figure 11.9 Hormone (aldosterone) EXTRA- CELLULAR FLUID Receptor protein Plasma membrane Hormonereceptor complex mrna NUCLEUS DNA New protein CYTOPLASM

52 Figure 11.9a Hormone (aldosterone) EXTRA- CELLULAR FLUID Receptor protein NUCLEUS Plasma membrane Hormonereceptor complex CYTOPLASM

53 Figure 11.9b Hormonereceptor complex DNA mrna NUCLEUS New protein CYTOPLASM

54 Concept 11.3: Transduction: Cascades of molecular interactions relay signals from receptors to target molecules in the cell Signal transduction usually involves multiple steps Multistep pathways can greatly amplify a signal Multistep pathways provide more opportunities for coordination and regulation of the cellular response

55 Signal Transduction Pathways The binding of a signaling molecule to a receptor triggers the first step in a chain of molecular interactions Like falling dominoes, the receptor activates another protein, which activates another, and so on, until the protein producing the response is activated At each step, the signal is transduced into a different form, usually a shape change in a protein

56 Protein Phosphorylation and Dephosphorylation Phosphorylation and dephosphorylation of proteins is a widespread cellular mechanism for regulating protein activity Protein kinases transfer phosphates from ATP to protein, a process called phosphorylation Many relay molecules in signal transduction pathways are protein kinases, creating a phosphorylation cascade

57 Figure Signaling molecule Receptor Inactive protein kinase 1 Activated relay molecule Active protein kinase 1 Inactive protein kinase 2 P i ATP PP ADP Active protein kinase 2 P Inactive protein kinase 3 P i ATP PP ADP Active protein kinase 3 P Inactive protein P i ATP PP ADP Active protein P Cellular response

58 Figure 11.10a Signaling molecule Receptor Activated relay molecule Inactive protein kinase 1 Active protein kinase 1

59 Figure 11.10b Inactive protein kinase 1 Active protein kinase 1 Phosphorylation cascade Inactive protein kinase 2 P i ATP PP ADP Active protein kinase 2 P Inactive protein kinase 3 P i ATP PP ADP Active protein kinase 3 P

60 Figure 11.10c Inactive protein kinase ATP 3 ADP Active protein PP kinase 3 P i P Inactive protein P i ATP PP ADP Active protein P Cellular response

61 Protein phosphatases rapidly remove the phosphates from proteins, a process called dephosphorylation This phosphorylation and dephosphorylation system acts as a molecular switch, turning activities on and off or up or down, as required

62 Small Molecules and Ions as Second Messengers Many signaling pathways involve second messengers Second messengers are small, nonprotein, water-soluble molecules or ions that spread throughout a cell by diffusion Second messengers participate in pathways initiated by GPCRs and RTKs Cyclic AMP and calcium ions are common second messengers

63 Cyclic AMP Cyclic AMP (camp) is one of the most widely used second messengers Adenylyl cyclase, an enzyme in the plasma membrane, converts ATP to camp in response to an extracellular signal

64 Figure Adenylyl cyclase Phosphodiesterase Pyrophosphate H 2 O ATP camp AMP

65 Figure 11.11a Adenylyl cyclase Pyrophosphate ATP camp

66 Figure 11.11b Phosphodiesterase H 2 O camp AMP

67 Many signal molecules trigger formation of camp Other components of camp pathways are G proteins, G protein-coupled receptors, and protein kinases camp usually activates protein kinase A, which phosphorylates various other proteins Further regulation of cell metabolism is provided by G-protein systems that inhibit adenylyl cyclase

68 Figure First messenger (signaling molecule such as epinephrine) G protein Adenylyl cyclase GTP G protein-coupled receptor ATP camp Second messenger Protein kinase A Cellular responses

69 Understanding of the role of camp in G protein signaling pathways helps explain how certain microbes cause disease The cholera bacterium, Vibrio cholerae, produces a toxin that modifies a G protein so that it is stuck in its active form This modified G protein continually makes camp, causing intestinal cells to secrete large amounts of salt into the intestines Water follows by osmosis and an untreated person can soon die from loss of water and salt

70 Calcium Ions and Inositol Triphosphate (IP 3 ) Calcium ions (Ca 2+ ) act as a second messenger in many pathways Ca 2 + can function as a second messenger because its concentration in the cytosol is normally much lower than the concentration outside the cell A small change in number of calcium ions thus represents a relatively large percentage change in calcium concentration

71 Figure Endoplasmic reticulum (ER) ATP Plasma membrane Mitochondrion Nucleus Ca 2+ pump ATP CYTOSOL ATP EXTRACELLULAR FLUID Key High [Ca 2+ ] Low [Ca 2+ ]

72 A signal relayed by a signal transduction pathway may trigger an increase in calcium in the cytosol Pathways leading to the release of calcium involve inositol triphosphate (IP 3 ) and diacylglycerol (DAG) as additional second messengers These two are produced by cleavage of a certain phospholipid in the plasma membrane

73 Figure EXTRA- CELLULAR FLUID Signaling molecule (first messenger) G protein CYTOSOL G protein-coupled receptor GTP Phospholipase C PIP 2 DAG Endoplasmic reticulum (ER) lumen IP 3 -gated calcium channel Ca 2+ IP 3 (second messenger) Nucleus

74 Figure EXTRA- CELLULAR FLUID Signaling molecule (first messenger) G protein CYTOSOL G protein-coupled receptor GTP Phospholipase C PIP 2 DAG Endoplasmic reticulum (ER) lumen Nucleus IP 3 -gated calcium channel Ca 2+ Ca 2+ (second messenger) IP 3 (second messenger)

75 Figure EXTRA- CELLULAR FLUID Signaling molecule (first messenger) G protein CYTOSOL G protein-coupled receptor GTP Phospholipase C PIP 2 DAG Endoplasmic reticulum (ER) lumen Nucleus IP 3 -gated calcium channel Ca 2+ Ca 2+ (second messenger) Various proteins activated IP 3 (second messenger) Cellular responses

76 Animation: Signal Transduction Pathways

77 Concept 11.4: Response: Cell signaling leads to regulation of transcription or cytoplasmic activities The cell s response to an extracellular signal is called the output response

78 Nuclear and Cytoplasmic Responses Ultimately, a signal transduction pathway leads to regulation of one or more cellular activities The response may occur in the cytoplasm or in the nucleus Many signaling pathways regulate the synthesis of enzymes or other proteins, usually by turning genes on or off in the nucleus The final activated molecule in the signaling pathway may function as a transcription factor

79 Figure Growth factor Receptor Reception Phosphorylation cascade Transduction CYTOPLASM Inactive transcription factor DNA Active transcription factor P Response NUCLEUS mrna Gene

80 Figure 11.15a Growth factor Receptor Reception Phosphorylation cascade Transduction CYTOPLASM Inactive transcription factor NUCLEUS

81 Figure 11.15b Phosphorylation cascade Transduction CYTOPLASM Inactive transcription factor DNA Active transcription factor P Response Gene NUCLEUS mrna

82 Other pathways regulate the activity of enzymes rather than their synthesis For example, a signal could cause opening or closing of an ion channel in the plasma membrane, or a change in cell metabolism

83 Figure Reception Binding of epinephrine to G protein-coupled receptor (1 molecule) Transduction Inactive G protein Active G protein (10 2 molecules) Inactive adenylyl cyclase Active adenylyl cyclase (10 2 ) Response Glycogen Glucose 1-phosphate (10 8 molecules) ATP Inactive protein kinase A Cyclic AMP (10 4 ) Active protein kinase A (10 4 ) Inactive phosphorylase kinase Active phosphorylase kinase (10 5 ) Inactive glycogen phosphorylase Active glycogen phosphorylase (10 6 )

84 Figure 11.16a Reception Binding of epinephrine to G protein-coupled receptor (1 molecule) Response Glycogen Glucose 1-phosphate (10 8 molecules)

85 Figure 11.16b Transduction Inactive G protein Active G protein (10 2 molecules) Inactive adenylyl cyclase Active adenylyl cyclase (10 2 ) ATP Inactive protein kinase A Cyclic AMP (10 4 ) Active protein kinase A (10 4 )

86 Figure 11.16c Transduction Inactive protein kinase A Cyclic AMP (10 4 ) Active protein kinase A (10 4 ) Inactive phosphorylase kinase Active phosphorylase kinase (10 5 ) Inactive glycogen phosphorylase Active glycogen phosphorylase (10 6 )

87 Signaling pathways can also affect the overall behavior of a cell, for example, a signal could lead to cell division

88 Regulation of the Response A response to a signal may not be simply on or off There are four aspects of signal regulation to consider Amplification of the signal (and thus the response) Specificity of the response Overall efficiency of response, enhanced by scaffolding proteins Termination of the signal

89 Signal Amplification Enzyme cascades amplify the cell s response to the signal At each step, the number of activated products is much greater than in the preceding step

90 The Specificity of Cell Signaling and Coordination of the Response Different kinds of cells have different collections of proteins These different proteins allow cells to detect and respond to different signals The same signal can have different effects in cells with different proteins and pathways Pathway branching and cross-talk further help the cell coordinate incoming signals

91 Figure Signaling molecule Receptor Relay molecules Activation or inhibition Response 1 Response 2 Response 3 Response 4 Response 5 Cell A: Pathway leads to a single response. Cell B: Pathway branches, leading to two responses. Cell C: Cross-talk occurs between two pathways. Cell D: Different receptor leads to a different response.

92 Figure 11.17a Signaling molecule Receptor Relay molecules Response 1 Response 2 Response 3 Cell A: Pathway leads to a single response. Cell B: Pathway branches, leading to two responses.

93 Figure 11.17b Activation or inhibition Response 4 Response 5 Cell C: Cross-talk occurs between two pathways. Cell D: Different receptor leads to a different response.

94 Signaling Efficiency: Scaffolding Proteins and Signaling Complexes Scaffolding proteins are large relay proteins to which other relay proteins are attached Scaffolding proteins can increase the signal transduction efficiency by grouping together different proteins involved in the same pathway In some cases, scaffolding proteins may also help activate some of the relay proteins

95 Figure Signaling molecule Plasma membrane Receptor Scaffolding protein Three different protein kinases

96 Termination of the Signal Inactivation mechanisms are an essential aspect of cell signaling If ligand concentration falls, fewer receptors will be bound Unbound receptors revert to an inactive state

97 Concept 11.5: Apoptosis integrates multiple cell-signaling pathways Cells that are infected or damaged or have reached the end of their functional lives often undergo programmed cell death Apoptosis is the best understood type Components of the cell are chopped up and packaged into vesicles that are digested by scavenger cells Apoptosis prevents enzymes from leaking out of a dying cell and damaging neighboring cells

98 Figure µm

99 Apoptosis in the Soil Worm Caenorhabditis elegans In worms and other organisms, apoptosis is triggered by signals that activate a cascade of suicide proteins in the cells programmed to die When the death signal is received, an apoptosis inhibiting protein (Ced-9) is inactivated, triggering a cascade of caspase proteins that promote apoptosis The chief caspase in the nematode is called Ced-3

100 Figure Ced-9 protein (active) inhibits Ced-4 activity Mitochondrion Deathsignaling molecule Ced-9 (inactive) Cell forms blebs Receptor for deathsignaling molecule Ced-4 Ced-3 Inactive proteins Active Ced-4 Active Ced-3 Activation cascade Other proteases Nucleases (a) No death signal (b) Death signal

101 Figure 11.20a Ced-9 protein (active) inhibits Ced-4 activity Mitochondrion Receptor for deathsignaling molecule Ced-4 Ced-3 Inactive proteins (a) No death signal

102 Figure 11.20b Deathsignaling molecule Ced-9 (inactive) Cell forms blebs Active Ced-4 Active Ced-3 Activation cascade Other proteases Nucleases (b) Death signal

103 Apoptotic Pathways and the Signals That Trigger Them In humans and other mammals, several different pathways, including about 15 caspases, can carry out apoptosis Apoptosis can be triggered by signals from outside the cell or inside it Internal signals can result from irreparable DNA damage or excessive protein misfolding

104 Apoptosis evolved early in animal evolution and is essential for the development and maintenance of all animals For example, apoptosis is a normal part of development of hands and feet in humans (and paws in other mammals) Apoptosis may be involved in some diseases (for example, Parkinson s and Alzheimer s); interference with apoptosis may contribute to some cancers

105 Figure Interdigital tissue Cells undergoing apoptosis 1 mm Space between digits

106 Figure 11.21a 1 mm Interdigital tissue

107 Figure 11.21b 1 mm Cells undergoing apoptosis

108 Figure 11.21c 1 mm Space between digits

109 Figure 11.UN01a 1 Mating factor activates receptor. Mating factor G protein-coupled receptor Shmoo projection forming P Formin GDP 2 GTP G protein binds GTP and becomes activated. 3 Fus3 Phosphorylation cascade Fus3 P Phosphorylation cascade activates Fus3, which moves to plasma membrane. 4 P Formin Fus3 P Fus3 phosphorylates formin, activating it. 5 Formin Actin subunit Microfilament Formin initiates growth of microfilaments that form the shmoo projections.

110 Figure 11.UN01aa 1 Mating factor activates receptor. Mating factor G protein-coupled receptor GDP 2 GTP G protein binds GTP and becomes activated. Phosphorylation cascade 3 Fus3 Fus3 P Phosphorylation cascade activates Fus3, which moves to plasma membrane.

111 Figure 11.UN01ab Shmoo projection forming P Formin P Fus3 Actin subunit Formin Formin P 4 Fus3 phosphorylates formin, Microfilament activating it. 5 Formin initiates growth of microfilaments that form the shmoo projections.

112 Figure 11.UN01b Wild type

113 Figure 11.UN01c Fus3

114 Figure 11.UN01d formin

115 Figure 11.UN02 Receptor 1 Reception 2 Transduction 3 Response Relay molecules Activation of cellular response Signaling molecule

116 Figure 11.UN03

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