Medcelična signalizacija. Lodish 4:
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1 Medcelična signalizacija Lodish 4:
2 Šest stopenj zunajceličnega signaliziranja Sinteza signalne molekule v signalizacijski celici Sproščanje signalne molekule iz signalizacijske celice Transport signalne molekule do tarčnih celic Zaznavanje signala z receptorji na tarčnih celicah Odziv tarčne celice (sprememba metabolizma, funkcije, proženje razvoja) Prekinitev signala celičnega odziva
3 Tudi evkariontski mikroorganizmi uporabljajo signaliziranje med celicami, običajno v povezavi s parjenjem ali diferenciacijo v spremenjenih pogojih v okolju. FEROMONI: snovi, ki vplivajo na vedenje ali izražanje proteinov drugih osebkov iste vrste. Višji organizmi uporabljajo zunajcelično signaliziranje znotraj posameznega organizma (hormoni, nevrotransmiterji, faktorji rasti). Pri živalih razlikujemo glede na doseg delovanja signalih molekul 4 tipe zunajceličnega signaliziranja: endokrino, parakrino, avtokrino in signaliziranje z neposrednim delovanjem v PM-zasidranih proteinov na sosednje celice.
4 Endokrino signaliziranje Hormoni: epinefrin, norepinefrin, inzulin, glukagon, trijodotironin, tetrajodotironin, steroidi
5 Parakrino signaliziranje Nevrotransmiterji (acetilholin, glutamat, glicin, serotonin, epinefrin, norepinefrin ), faktorji rasti (EGF, PDGF ), citokini (interlevkini, interferoni), prostaglandini
6 Avtokrino signaliziranje Faktorji rasti (EGF, PDGF ), citokini (interlevkini, interferoni), prostaglandini
7 Signaliziranje z neposrednim delovanjem PMzasidranih proteinov na sosednje celice. Membranski faktorji rasti (EGF), MHC-antigen kompleksi
8 T-cell antigen receptor (TCR)-mediated activation of T lymphocite (macrophage, dendritic cell, B-cell) Cytokine secretion T-cell maturation into helper and cytotoxic c.
9 Receptorji ligandna specifičnost efektorska specifičnost Različni receptorji lahko specifično vežejo isti ligand (npr. ACh), vezava pa sproži različne odzive v tarčnih celicah (npr. vezava ACh na AChR v skeletni mišici izzove skrčenje mišice, na AChR v srčni mišici pa upočasni frekvenco krčenja). Isti receptor na različnih tarčnih celicah lahko ob vezavi istega liganda sproži različen odziv v tarčni celici (npr. vezava ACh na AChR v srčni mišici upočasni frekvenco krčenja srca, na AChR acinarne celice v pankreasu pa izločanje mešičkov s prebavnimi encimi). Različni receptorji lahko specifično vežejo različne ligande, vezava pa sproži enak odziv v tarčni celici (npr. tako vezava epinefrina kot glukagona, vsakega na svoj receptor na jetrni celici, povzroči izločanje glukoze v kri) (sekundarni prenašalci: camp)
10 Signalizacijske molekule Male lipofilne snovi, ki pasivno prehajajo membrano in delujejo na znotrajcelične receptorje. Hidrofilne molekule, ki se vežejo na površinske receptorje. Lipofilne molekule, ki se vežejo na površinske receptorje. Plini: NO.
11 Mali lipofilni hormoni, pasivno prehajajo membrano in se vežejo na znotrajcelične receptorje. - (- or +) - Affected stability of mrna.
12 Nekaj primerov malih lipofilnih hormonov steroidni hormoni (kortizol, progesteron, estradiol, testosteron) tiroksin retinoidi
13 Steroidi nastajajo iz holesterola in delujejo nekaj ur ali dni. Pogosto vplivajo na rast in diferenciacijo tkiv. Tiroksin (dve obliki: tetra- in trijodotironin) nastaja iz tiroglobulina s proteolitično cepitvijo in se sprošča v kri. Pospešuje izražanje številnih genov za katabolične encime. Retinoidi nastajajo iz retinola (vitamin A) in imajo zelo raznolike učinke zaradi različnih lastnosti ligandov in receptorjev; sodelujejo pri delitvi, diferenciaciji in odmiranju celic.
14
15
16 Hidrofilne signalizacijske molekule, ki se vežejo na površinske receptorje
17 Nekaj primerov hidrofilnih signalizacijskih molekul (poli)peptidni hormoni (inzulin, glukagon) male nabite molekule (Tyr epinefrin, His histamin)
18 Kateholamini dopamin epinefrin (adrenalin) norepinefrin (noradrenalin)
19 Lipofilni hormoni, ki delujejo na membranske receptorje Eikozanoidni hormoni (20 C) PG delujejo kot lokalni mediatorji z avto- ali parakrinim signaliziranjem
20
21 Vrste receptorjev za signalizacijske molekule na površini celic Receptorji, povezani s Tyr-kinazo Z G-proteini povezani receptorji (metabotropni receptorji) Receptorji ionski kanalčki (ionotropni receptorji) Receptorji z lastno encimsko aktivnostjo (guanilil ciklazno, fosfatazno, kinazno)
22 Receptorji povezani s Tyr-kinazo Receptorji za citokine, interferone, HGF
23 Primer: receptor za IgE (Fc RI) Signaliziranje pri alergičnem imunskem odzivu Histamine release alergic response mastocit, bazofilec Simons & Toomre (2000) Nat. Rev. Molec. Cell Biol. 1,
24 Medcelična signalizacija (prenos živčnega signala) Lodish, H. et al.: Nerve cells. In: Molecular cell biology, W.H. Freeman & Co., New York. Alberts, B. et al.: Membrane transport of small molecules and the electrical properties of membranes. In: Molecular Biology of the Cell, Garland Science, New York.
25 Structure of typical mammalian neurons
26 Neurons communicate with many other cells green - MAP2 Hippocampal interneurons red - synaptotagmin
27 A synapse
28 Reception, conduction and transmission of electric signals Ion channels in neuronal plasma membrane:
29 Origin of the resting PM potential in a typical vertebrate neuron
30 Electric potential (E) across the cell PM Nernst equation:
31 Effect of changes in ion permeability on PM potential
32 Passive spread of depolarization Membrane depolarization spreads passively only over a short distance Only resting K + channels Lenght constants
33 Voltage-gated ion channels generate action potentials (cycles of membrane depolarization, hyperpolarization and return to the resting potential)
34 Ion permeabilities during an action potential
35 Structure and function of the voltage-gated Na + channel
36 Unidirectional conduction of action potential
37 Myelination of axons membrane layers
38 Formation and structure of a myelin sheath
39 Myelination increases the velocity of signal conduction from ~1 m/s to m/s. 12 µm myelinated vertebrate axon 600 µm unmyelinated squid axon 12 m/s Radical reduction of neuronal volume & ATP consumption Evolution of vertebrate brain
40 Signal transmission at a chemical synapse (frog neuro-muscular junction)
41 Chemical synapse - schematically ~20 nm
42 Neurotransmitters
43 Synaptic vesicle cycle
44 Excitatory and inhibitory responses in postsynaptic cells Frog skeletal muscle - nicotinic AChR (ligand-gated ion channel) Frog heart muscle - muscarinic AChR (G protein-coupled R)
45 Activation of ligand-gated ion channels at a NM junction
46 Nicotinic ACh receptor
47 Fast synapses
48 Muscarinic ACh receptor (M2) in the heart muscle PM
49 Slow synapses
50 Electric synapse - schematically
51
52 Ukrivljanje membran
53 McMahon & Gallop (2005) Nature 438,
54 Mechanisms of membrane deformation McMahon & Gallop (2005) Nature 438,
55 Phospholipids have different shapes and lipid shape affects membrane curvature Gangliozidi SM Inverted-cone PS Cone Sprong et al. (2001) Nat. Rev. Molec. Cell. Biol. 2,
56 Lipid shaping - Enzymes that change acyl chain composition of lipids Palmitoyl CoA CtBP/BARS Scales & Scheller (1999) Nature 401, Enzymes that change headgroup size of lipids
57 Role of LPAAT-mediated conversion of LPA to PA in one of the steps of synaptic vesicle formation (fission) LPAAT LPAAT Schmidt et al. (1999) Nature 401,
58 Indirect scaffolding Epsin (N-BAR) PH/PX domain clathrin Direct scaffolding
59 Amphipatic helix and membrane curvature BAR domains McMahon & Gallop (2005) Nature 438,
60 Further reading: McMahon, H.T. and Gallop, J.L. (2005): Membrane curvature and mechanisms of dynamic cell membrane remodelling. Nature 438, McMahon, H.T. et al. (2010): Membrane curvature in synaptic vesicle fusion and beyond. Cell 140,
Medcelična signalizacija -
Medcelična signalizacija - - prenos živčnega signala Lodish, H. et al.: Nerve cells. In: Molecular cell biology, W.H. Freeman & Co., New York. Alberts, B. et al.: Membrane transport of small molecules
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