Biological Rhythms. Today s lecture
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1 Biological Rhythms (a review of general endocrinology) 35 Neuroendocrine control: homeostatic responses and biological rhythms. A role for anticipation or feed-forward mechanisms or scheduled events. Biological rhythms and CN: what are they and how have been approached (what are oscillators, entrainment, light and melatonin, central and peripheral clocks). CN, running activity rhythm and ovulatory LH surge. CN signal transduction. Today s lecture control and story lines 1
2 Take Home Message (THM) It is thought that circadian regulation of physiology and behavior imparts survival advantages to organisms that use clocks. In mammals, a master clock resident in the CN synchronizes other central and peripheral oscillators to evoke this regulation. This master oscillator consists of interlocking transcriptional-translational feedback loops, and it regulates both core clock genes necessary for oscillator maintenance as well as specific output genes that directly or indirectly mediate physiology under circadian control. It is now clear that both neuroanatomic and molecular outputs of the clock are necessary for proper circadian clock function. It s all in the timing: many clocks, many outputs. Panda and Hogenesch JB (2004). Genomic Institute of Novartis Research Foundation and Dept of Neuropharmacology, cripps Research Institute. J of Biological Rhythms 19 (5), , Neuroendocrine control anticipation what if sensor e.g. t C/F afferent story line integration center (concept vs anatomy) (single cell vs multiple sites) (single vs multiple integrators) negative feedback story line efferent story line effector Throughout the course I have hinted that feedforward control helps to anticipate needed homeostatic responses (e.g. GLP-1). 2
3 Neuroendocrine control anticipation or what if sensor e.g. t C/F afferent story line integration center (concept vs anatomy) (single cell vs multiple sites) (single vs multiple integrators) negative feedback story line efferent story line effector A similar anticipation mechanism is based on the presence of biological clocks, central and peripherals, at cellular and sub-cellular levels. ingle system integrator e.g. gonadal hierarchy 2 P4 receptor C growth hierarchy IGF receptor C thyroid hierarchy T3 receptor C N TC GnRH /GRH TRH AP AP AP gonad liver thyroid 2 / P4 IGF T3 / T4 3
4 Multiple systems integrator Hypothalamic nuclei / areas PVN, paraventricular nucleus ARC, arcuate nucleus VMH, ventro-medial nucleus DMH, dorso-medial nucleus LH, lateral hypothalamus M, median eminence CF neuronal blood ( Neuronal = N Blood-born - BB ) BB xtra-hypothalamic nuclei / areas A, amigdala AP, area postrema DMN, dorso motor nucleus of vagus NT, nucleus tractus solitarious PBN, para braquial nucleus MFB, medial forebrain bundle PVN M/AP PP AN DMH VMH PVN LH PBN AP DMN NT from a central hypothalamic integrator to a diffused brain integrator BB N ARC M A e.g. food intake Neuroendocrine control the PVN or integrator concept and the CN or anticipation concept PVN hierarchy - Fb receptor C error signal M AP NT DMN midbrain gland controlled variable 4
5 Neuroendocrine control C NC h nt R-h R-nt R-h R-nt C NC effect if glucose is detected in the blood??? Glycemia up Insulin up Glycemia down example A B C D feedback homeostatic control e.g. glucose on ßcells Neuroendocrine control C NC h nt R-h R-nt R-h R-nt C NC effect if glucose is detected in the duodenum tract??? Duodenal glucose up GLP-1 up ßcell primed example feedforward A B C D anticipation e.g. GLP-1 on ßcells 5
6 Neuroendocrine control Theoretically, a feedforward mechanism anticipates the logistics needed to carry on a specific physiological effect A B C D feedback feedforward Keep this thought in mind for the rest of this lecture. o what??? and the rhythms??? Biological rhythms Chronobiology, or the study of biological rhythms, concerns itself with the timing of events within and external to animals. ingle cells and animals have evolved timing systems that are important for every type of behaviors and physiology. hours of anesthesia electrical current plasma prolactin plasma melatonin examples Migration and hibernation versus day and night??? memory score 6
7 Biological rhythms The golden hamster is the chronobiologists favorite animal subject because of their impressive daily rhythms in their use of a running-wheel kept in their cage. They sleep during the day and run all night. methods Biological rhythms - a clock methods time 7
8 Biological rhythms - a clock light dark But, what happens if you do not know if it is day or night??? constant darkness or constant light activity rhythms Biological rhythms - a clock light dark constant darkness or constant light activity rhythms ffect of constant darkness or constant light. Both conditions are free running conditions. 8
9 Biological rhythms - a clock splitting rhythm after prolonged exposure to constant light (each horizontal line represents two days of data) activity rhythms ffect of constant darkness or constant light. Both conditions are free running conditions. Biological rhythms - a clock vening () / morning (M) oscillators in mammalian photoperiodism. A, oscillators entrained to a short night rhythm B, as nights get longer (PRC) the oscillators drift apart since their free running rhythms differ C, in long nights oscillators adopt an entrainment pattern where they overlap minimally A B approach??? if light pulses here (phase response curve) PRC entrainment (zeitgebers) C one clock vs. two clocks entrainment vs. resetting (hold this model until the examples at the end of this lecture) 9
10 Biological rhythms - PRC approach??? effect of a light pulse (phase response curve) PRC entrainment (zeitgebers) all individuals are running in constant darkness, a free running condition. Biological rhythms - Melatonin tryptophane melatonin melatonin serotonin Melatonin is derived from the aminoacid tryptophane, the same as the NT serotonin 10
11 Biological rhythms - Melatonin CF plasma melatonin Melatonin is released from the pineal gland into the CF and blood during the night Biological rhythms - Melatonin L:D ummer condition melatonin L:D Winter condition PRC e.g. long day vs short night cfos e.g. short day vs long night melatonin light induced ummer vs Winter condition 11
12 Biological rhythms - Melatonin ummer condition induced by exogenous melatonin infusion Winter condition induced by exogenous melatonin infusion e.g. long day vs short night e.g. short day vs long night melatonin exogenous melatonin induced ummer vs Winter condition Biological rhythms - Melatonin Free running rhythm of drinking behavior in rats free running rhythm melatonin vehicle exogenous melatonin entrained rats to the period of the injection regime melatonin 12
13 Biological rhythms - CN Blinded rat with intact CN Blinded rat with a lesioned CN All individuals are blind, thus they are running in constant darkness, a free running condition melatonin o, light may not be necessary uprachiasmatic nucleus Blinded rat with intact CN Blinded rat with a lesioned CN CN All individuals are blind, thus they are running in constant darkness, a free running condition 13
14 uprachiasmatic nucleus o, what might be the relationship among biological rhythms, light, the CN and melatonin??? CN uprachiasmatic ß nucleus o, what might be the relationship among biological rhythms, light, the CN and melatonin??? CN Melatonin is released from the pineal gland into the CF and blood during the night 14
15 uprachiasmatic nucleus o, what might be the relationship among biological rhythms, light, the CN and melatonin??? CN Melatonin is released from the pineal gland into the CF and blood during the night uprachiasmatic nucleus Hypothalamic nuclei / areas PVN, paraventricular nucleus ARC, arcuate nucleus VMH, ventro-medial nucleus DMH, dorso-medial nucleus LH, lateral hypothalamus M, median eminence xtra-hypothalamic nuclei / areas A, amigdala AP, area postrema DMN, dorso motor nucleus of vagus NT, nucleus tractus solitarious PBN, para braquial nucleus MFB, medial forebrain bundle CN neuronal blood CF PVN M/AP PP AN and what might be their relationship with a PVN integration center or with diffused centers? N CN ( Neuronal = N Blood-born - BB ) DMH VMH PVN ARC M A LH PBN BB AP DMN NT 15
16 uprachiasmatic nucleus neuronal blood CF PVN PVN M/AP PP AN CN CN Theoretically, a feedforward mechanism anticipates the logistics needed to carry on a specific physiological effect uprachiasmatic nucleus the PVN or integrator concept and the CN or anticipation concept CN e.g. repro rhythm to clock 16
17 CN - repro rhythm to clock Blinded rat with intact CN (have repro rhythm) Blinded rat with a lesioned CN (do not have repro rhythm) e.g. repro rhythm to clock individuals are blind, thus they are running in constant darkness, a free running condition CN - repro rhythm to clock The CN is responsible for the precise timing of the LH surge: e.g. repro rhythm to clock CN lesions eliminates the ovarian cycle (behavior & LH surge). The LH surge maintains its exact relationship to the locomotor activity in a constant light environment, suggesting they are controlled by the same circadian mechanism. Lengthening / shortening of activity cycle by pharmacological treatment or light entrainment alters the estrous behavior and the LH surge. Pentobarbital in proestrous AM temporarily blocks the LH surge, which occurs at precisely the normal time the next day. and how would you use this knowledge to make a clock??? 17
18 R Na / K pump camp ----> PKA ----> channel / enzyme AC Protein synthesis teroid + R ----> R 5 3 mrna R XX1 HR DNA 1 additional TF Cellular response e.g. repro rhythm to clock a hint how would you use this knowledge to make a clock??? CN - repro rhythm to clock e.g. repro rhythm to clock and what the hell is this supposed to mean??? 18
19 CN - repro rhythm to clock (5) (2) (4) (1) (3) e.g. repro rhythm to clock a negative feedback CN - repro rhythm to clock e.g. repro rhythm to clock a negative feedback 19
20 CN - repro rhythm to clock photoreceptor CN clock neuron retinal ganglion cell Ca++ glutamate cytoplasmic responses e.g. repro rhythm to clock CCGs neurogenic or paracrine nuclear responses Target cell CN - repro rhythm to clock signal transduction 20
21 CN - repro rhythm to clock signal transduction Phosphorylation of CRB Oscillators (CN & others) CN times rhythmic events: 1 CN lesions eliminates rhythmic events. Fetal CN transplantation to CN lesioned rats restore rhythmic activities. 2 The restored rhythm depends on donor CN (exp: 22h vs 24h rhythm). 3 In some cases, this restoration does not depend on re-establishment of synaptic connections since encapsulated CN (allows diffusion of small chemicals but not neuronal outgrowth) is able to restore activity rhythms. CN oscillators 4 However, in some other cases, neuronal conections may be necessary to restore rhythmic events. 21
22 Oscillators (CN & others) Per luciferase transgene expression Circadian rhythms in isolated brain regions: in vitro studies extra CN oscillators Oscillators (CN & others) Circadian rhythms in isolated brain regions: in vitro studies Per luciferase transgene expression extra CN oscillators 22
23 Oscillators (CN & others) Circadian rhythms in isolated brain regions: in vitro studies Per luciferase transgene expression in pineal gland PRC extra CN oscillators Oscillators (CN & others) the PVN or integrator concept and the CN or anticipatory concept CN and how do oscillators get entrained or reseted (zeitgeber)? 23
24 Oscillators (CN & others) CN Oscillators (CN & others) CN CN 24
25 Its all in the timing xperimental endpoint: PRC studies using neuronal firing rate in CN slices xperimental results: ignaling in the CN is selectively responsive and integrative xample of entrainment or zeitgeber pathways Its all in the timing CN 25
26 Its all in the timing Take Home Message (THM) It is thought that circadian regulation of physiology and behavior imparts survival advantages to organisms that use clocks. In mammals, a master clock resident in the CN synchronizes other central and peripheral oscillators to evoke this regulation. This master oscillator consists of interlocking transcriptional-translational feedback loops, and it regulates both core clock genes necessary for oscillator maintenance as well as specific output genes that directly or indirectly mediate physiology under circadian control. It is now clear that both neuroanatomic and molecular outputs of the clock are necessary for proper circadian clock function. It s all in the timing: many clocks, many outputs. Panda and Hogenesch JB (2004). Genomic Institute of Novartis Research Foundation and Dept of Neuropharmacology, cripps Research Institute. J of Biological Rhythms 19 (5), ,
27 uggest Reading uggest Reading 27
28 uggest Reading uggest Reading 28
29 uggest Reading uggest Reading 29
30 uggest Reading Its timing and memories hippocampal cells help rats think ahead in a maze test 30
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