Chemical Oscillations and Circadian Rhythms. Peter Ruoff University of Stavanger Stavanger, Norway
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1 Chemical Oscillations and Circadian Rhythms Peter Ruoff University of Stavanger Stavanger, Norway
2 Early Events Fechner publishes the observation of an oscillating electric current in an electrochemical cell Ostwald observes that dissolution of chromium in acid is periodic Morgan discovers that CO production from formaldehyde can occur periodically Bray discovers periodic H 2 O 2 decomposition in the presence of iodate Belousov tries to publish his observation of what we know today is the Belousov-Zhabotinsky reaction Zhabotinsky s starts to publish studies on Belousov s oscillatory reaction Prague conference on chemical and biological oscillators FKN mechanisms of the Belousov-Zhabotinsky reaction.
3
4 The Belousov-Zhabotinsky Reaction (stirred system) 1M H 2 SO 4 organic subsrate KBrO 3 or NaBrO 3 Ce 3+, Ce 4+ (Mn 2+, Fe(phen) 3 2+ or Ru(bpy) 3 2+ ) Fe-catalyzed system Oscillatory fluorescence in the Ru-catalyzed system (which is also light-sensitive)
5 The Belousov-Zhabotinsky Reaction (unstirred thin layer in a Petri-dish) Traveling oxidation waves
6 The FKN mechanism (1972) Dick Field Endre Körös Dick Noyes
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8 3 overall reactions in the BZ reaction Process B occurs when the concentration of Br-ion has moved below a critical concentration.
9 Relaxation type of oscillations (oxidation spikes) long bromide ion consuming period short autocatalytic period and bromide ion regeneration
10 The Oregonator (Field & Noyes 1974) Species: X = HBrO 2 Y = Br Z = Ce(IV) A, B = BrO 3 P = HOBr The Field-Noyes equations show close analogy to the Huxley-Hodgkin equations.
11 The Oregonator
12 Excitability in the BZ reaction (Oregonator) (experiment) Ruoff, CPL, 1982
13 Silver Ion Induced Oscillations [Ce(IV)] [Ce(IV)] [Br - ] [Br - ]
14 Oxidation pulse propagation
15 Facilitation phenomena
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17 Amplified Oregonator
18 Amplified Oregonator
19 Bromide-Ion Induced Excitability (Reduction Spikes) subcritical perturbation
20 Bromide-Ion Induced Excitability (Reduction Spikes) supercritical perturbation
21 Oxidation and Reduction Spikes in the Amplified Oregonator Large k O5, k O6 Low k O5, k O6
22 Regions of oscillations, bistability, and Ag + and Br - excitable steady states of amplified Oregonator in k 5 -k 6 space
23 Phase response curves in the BZ reaction (oxidation spikes) Phase shift = peak time (after perturbation) - peak time (no perturbation) Outline of experiment Perturbation by Br Perturbation by Ag+
24 Circadian rhythms are important in daily and seasonal adaptations of organisms to their environments
25 History of Circadian Rhythms. 325 BC: Androsthenes (from Thasus) observes that plants sleep Tamarindus indica day night
26 The French astronomist De Mairan finds in 1729 that leaf movements in Mimosa plants continue in darkness
27
28 Linné s flower clock CARL von LINNÉ ( ) Tiger Jaw (Faucaria tigrinis, South Africa, open flowers 5 pm, closes 8 pm)
29 Leaf movements under free-running conditions Phaseolus coccineus. Typical course of the circadian leaf movements under constant light (weak intensity). The phase shifts within six days by roughly 17 hours compared to the normal day. The length of one period is thus about 27 hours (circles in 24-hour-intervals; E. BÜNNING and M. TAZAWA, 1957). Time lapse movie of leaf movements in bean (Phaseolus) seedlings by Roger P. Hangarter, Indiana University.
30 Flashing and glow circadian rhythms in Gonyaulax polyedra (von der Heyde et al. (1992) JBR ) With increasing temperature flashing and glow rhythms change their amplitudes in opposite directions. Gonyaulax polyedra
31 Circadian variation in blood pressure and heart attacks daily variation of blood pressure Variation of heart attacks during the day
32 Properties of circadian rhythms circadian rhythms are generated already within a single cell and can occur under constant environmental conditions (free-running conditions). We call these rhythms for endogeneous rhythms. they can be phase shifted by sudden changes in light, temperature, drugs, etc. they can track external periodic variations such as lights on/off or temperature variations. This property is called entrainment. the circadian period is unaffected by different (but constant) temperatures. This property is called temperature compensation. Chronobiologists require this property for calling an oscillator a clock.
33 Examples of temperature compensated rhythms (from E. Bünning: The Physiologcal Clock, Berlin, 1964)
34 The 1957 suggestion by Hastings and Sweeny how temperature compensation may occur Practically all physiological reactions, enzyme catalyzed reactions, elementary reactions, etc. are quite dependent upon temperature. Their rates increase by a factor of 2-3 when temperature is increased by 10 C ( Van t Hoff s rule ). This makes it difficult pinpointing a certain process as the candidate how temperature compensation may arise. Hastings and Sweeney proposed in 1957 a general mechanism based on opposing reactions, suggesting that temperature compensation is a systemic property.
35 Importance of genetics to study the mechanisms of the circadian clock In the beginning of the 1970 s reseachers showed using the fly Drosophila and the fungus Neurospora crassa that circadian properties such as period length, temperature compensation, and entrainment behavior are inherited and determined by genes. Genes were identified, where alleles of these genes showed different properties of the circadian clock. Examples of these genes are period and timeless in Drosophila and frequency in Neurospora. This was the start of using model organisms in circadian rhythm research
36 Circadian clock properties are inherited
37 Model organisms in circadian rhythm research
38 Neurospora crassa: A model organism to study genetics and circadian rhythms eukaryotic filamentous fungus growths rapidly nonpathological easy to handle many mutants available circadian (conidiation) rhythm is easy assayable web resource: Fungal Genetics Stock Center
39 Neurospora in nature grows in the tropics/subtropics has also been found in New Mexico, Alaska, Spain, Portugal, Switzerland. Genome sequenced in ,225,835 bp 9,826 genes 7 chromosomes
40 Neurospora s three main cell types photo: I. W. Jolma, UiS DAPI stain of nuclei (from Fungal Genetics Stock Center) for more pictures or videos, click here
41 photo: Bodil Aase, UiS photo: Bodil Aaase
42 Nuclei stained with DAPI
43 Conidia photo: Bodil Aaase
44 Conidia photo: Bodil Aaase
45 Neurospora circadian rhythm in a Petri dish Period length 22 h Rhythm shows temperature and ph compensation
46 In Neurospora, the circadian rhythm is easily seen in so-called race tubes
47 Race tube assay of Neurospora's circadian rhythm Jason C. Thoen and Van Gooch: Time Lapse Video Showing an Internal Circadian Clock in Mold (Neurospora) Growth
48 Temperature Compensation in different frq mutants
49 ph Compensation compared to temperature compensation in different frq mutants ph compensation temperature compensation
50 FREQUENCY (FRQ) protein >sp P19970 FRQ_NEUCR FREQUENCY CLOCK PROTEIN - Neurospora crassa. MADSGDKSQGMRPPPFDSRGHPLPRRASPDKSITLENHRLARDTSSRVTS start of long FRQ SSALGVTESQPQLKSSPTRRNSSGESEPTNWFNQSNRNPAAAFHDESHIM start of short FRQ EVDPPFYQKETDSSNEESRYPPGRNPVHPPGGVQLPGFRPVAAHSTAADD YRSVIDDLTVENKRLKEELKRYKQFGSDVMRKEKLFEIKVHGLPRRKKRE coiled-coil LEATLRHFAASLGDSSESTSQRRKTGRHGTAVYSSGVSLSKHDSSSSSRS RPVDSAYNSMSTGRSSHAPHSSGPSLGRPSLTRAKSVGTQKVENYLRDTP DGLLPHHIVMTDKEKKKLVVRRLEQLFTGKISGRNMQRNQSMPSMDAPLA PEGTNMAPPRPPPEGLREACIQLQDGDNPRKNRSSKDNGSASNSGGDQTE LGGTGTGSGDGSGSGGRTGNNTSPPGAIAPDQRPTRPRDLDPDRVQIPSE potential GSK-3 P-site NMDYIRHLGLVSPEFLQGSRTSYQDVAPDAEGWVYLNLLCNLAQLHMVNV frq[7]:g D frq[1]:g S TPSFIRQAVSEKSTKFQLSADGRKIRWRGGTDGTKFSSDSSEDKSQQSPM S513 PEST-1 TEDTEDGSDKNGRRKKRKTQQASSEIGRFGPSRSPSDTFHYKPMFVHRNS PEST-1 NLS SSIETSLEESMSQGSEDAVDESNMGNSKWDFSGSGTTQQRRKRRYDGAIV YYTGAPFCTDLSGDPGDMSPTAQMTAGREVEGSGSGDEVEHVLQRTLSGS SLPIRPLSDDRARVAEVLDFDPGNPPELVADDGSSPNDEDFVFPWCEDPA KVRIQPIAKEVMEPSGLGGVLPDDHFVMLVTTRRVVRPILQRQLSRSTTS EDTAEFIAERLAAIRTSSPLPPRSHRLTVAPLQVEYVSGQFRRLNPAPLP PEST-2 PPAIFYPPFSTDSSWDDGDDLASDDEEVEEVEEDSYSEGQISRRANPHFS PEST-2 DNNTYMRKDDLAFDTETDVRMDSDDNRLSDSGHNMRAMMPRAEAVDGDDS PLAAVTGKEVDIVHTGSSVATAGGAESGYSSSMEDVSSS
51 frq: frequency Minimal Model of the Circadian Rhythms in Neurospora crassa
52 A theory for temperature compensation: The antagonistic balance equation This approach suggests that temperature compensation is a systemic property as suggested by Hastings and Sweeney in 1957.
53
54
55 Estimation of activation energy for k5 and k6
56
57 Model shows relaxation type of oscillations in frq-mrna: Short transcription phase long FRQ-protein degradation phase time, h
58 Temperature Compensation in different frq alleles
59 Applying the antagonistic balance equation: The Oregonator
60 The (closed) Belousov-Zhabotinsky reaction is not temperature compensated The Arrhenius plot to the left shows that the BZ reaction is not compensated. The overall E a for the period is ca. 73 kj/mol.
61
62
63 Thank you for your attention!
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