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Brain Energy 2013 – Current Advances in Brain Maintenance Brain Energy 2013 – Current Advances in Brain Maintenance The Norwegian Academy of Science and Letters 29 August 2013 Cover: The lactate receptor GPR81 (green) is located in neurons, shown here in hippocampal cortex CA1. The receptor is concentrated in the pyramidal cell somatodendritic compartment including spines (white arrows in closeup), and to a lesser degree in vascular endothelium. Neurons were labelled for MAP2 (red), nuclei with DAPI (blue). Electronmicroscopy shows lactate receptor labelling (10 nm immunogold, red arrowheads) at the postsynaptic membrane (between black arrowheads), Illustrated by a synapse between a nerve terminal (t) and a dendritic spine (s) in stratum radiatum of CA1. See: Lauritzen KH, Morland C, Puchades M, Holm-Hansen S, Hagelin EM, Lauritzen F, Attramadal H, Storm-Mathisen J, Gjedde A, Bergersen LH (2013) Lactate Receptor Sites Link Neurotransmission, Neurovascular Coupling, and Brain Energy Metabolism. Cereb Cortex Epub 2013 May 21. Brain Energy 2013 – Current Advances in Brain Maintenance When: Thursday 29th August 2013 – Where: The Norwegian Academy of Science and Letters, Drammensveien 78, Oslo Why: Understanding mechanisms that keep the brain in good repair and able to adapt to new challenges, at all ages, will help address a major challenge facing mankind: how to retain an individual’s autonomy towards the end of life. Organized by: Linda H Bergersen, Vidar Gundersen, Jon Storm-Mathisen; Dep Oral Biology, and Inst Basic Medical Sciences, UiO Supported by: The Medical Faculty (equal opportunity grant to LHB), and Molecular Life Science - MLSUiO, University of Oslo Registration: Email to [email protected], [email protected] or [email protected] before 20th August 08:30 Registration, Coffee 08:50 The organizers Welcome - introduction 09:00 Linda H Bergersen Lactate is a volume transmitter – 'New Deal' for understanding brain function and brain disease 09:30 Pierre Magistretti Role of lactate as a signalling molecule for neuronal plasticity and memory 10:00 Albert Gjedde On the roles of energy metabolites as volume transmitters in brain: in vivo molecular imaging of neuroenergetics in aging and age-related neurodegenerative diseases 10:30 Ole Petter Ottersen Astrocytic endfeet: new insight in their function and organization 11:00 Coffee – Refreshments 11:30 Vidar Gundersen Glial signalling 12:00 Peter Somogyi Network state-dependent firing of distinct cell types in the awake and sleeping hippocampus 12:30 David Attwell Regulation of brain energy supply: where does the action start? 13:00 Martin Lauritzen Neurophysiology of local control of brain blood flow and energy metabolism 13:30 Lunch 14:15 Lene Juel Rasmussen Mitochondrial function regulates nucleotide metabolism and affects genomic stability: mechanisms and biomarkers for cognitive function 14:45 Vilhelm Bohr Base excision DNA repair and neurodegeneration 15:15 Tone Tønjum Glial responses to meningitis pathogens 15:45 Bjørnar Hassel When bacteria meet the brain: energy-metabolic interactions between abscess-forming bacteria and brain cells 16:15 Coffee – Refreshments 16:45 Magnar Bjørås Maintenance of brain function by Neil dependent repair of oxidative DNA damage 17:15 Arne Klungland Dynamics of methyl modifications in DNA and RNA: roles in brain and endurance running 17:45 Fred Gage (Rusty Gage) Functional consequences of exercise and enrichment on adult neurogenesis and cognition 18:15 Discussion Brain Maintenance (Moderator: Ole Petter Ottersen, Rector of the University of Oslo ) 19:00 End of Symposium – Sparkling wine Lactate is a volume transmitter – 'New Deal' for understanding brain function and brain disease Linda Hildegard Bergersen Department of Oral Biology and Department of Anatomy, University of Oslo, Oslo, Norway, and Department of Neuroscience and Pharmacology, University of Copenhagen, Copenhagen, Denmark [email protected] Lactate is a fuel for the brain, but of disputed importance. Some effects of lactate in the CNS, such as neuroprotection, are not easily explained in terms of energy metabolism, but suggest receptor mechanisms. In a recent study (Lauritzen KH et al 2013 Cereb Cortex), we show that the lactate receptor GPR81, also known as HCA1, downregulates cAMP production in the brain in response to physiologically increased lactate levels, such as occur in physical exercise, and by the specific GPR81 agonist 3,5-dihydroxybenzoate, which occurs in fruits and berries. The receptor is concentrated at the postsynaptic membranes of glutamatergic synapses on cortical pyramidal cells, including in the hippocampus. GPR81 is also enriched at the blood-brain barrier: the GPR81 density at endothelial cell membranes are about twice the GPR81 density at membranes of perivascular astrocytic processes, but about one seventh of that on synaptic membranes. There is only a slight signal in perisynaptic processes of astrocytes. In synaptic spines, as well as in adipocytes where the receptor was first identified, GPR81 immunoreactivity is located on subplasmalemmal vesicular organelles suggesting trafficking of the protein to and from the plasma membrane. In adipocytes, GPR81 is part of an autocrine / paracrine loop whereby increased production of lactate mediates the antilipolytic effect of insulin through downregulating cAMP. We suggest that, in the brain, the lactate receptor allows lactate released by activated neurons to act as a “volume transmitter” that tells brain cells to adjust their condition by curbing cAMP production. This would provide a means for linking neuronal activity, cerebral energy metabolism and energy substrate availability, including a neuronal glucose saving response to the supply of lactate. It is known that whereas brain cAMP levels increase acutely in arousal, augmenting cognition and memory, chronically increased cerebral cAMP such as in chronic stress and old age is associated with cognitive impairment. The new findings therefore point to the lactate receptor as a potential therapeutic target. Biography Linda Hildegard Bergersen obtained her PhD at the University of Oslo in 2001 under the supervision of Ole Petter Ottersen. After postdoc periods, in Oslo with Jon Storm-Mathisen and in Lausanne with Pierre Magistretti and Luc Pellerin in 2003, she established the “Brain and Muscle Energy Group” in Oslo in 2004, and in Copenhagen in 2011. Bergersen’s main research interest is brain energy supply, under physiological as well as pathological conditions. She has done pioneering work on the localization of lactate transporters and in a recent breakthrough discovered the lactate receptor GPR81 to be present and active in the brain, opening new opportunities for basic and translational research. Bergersen is currently a Professor of Physiology at the University of Oslo and part time Professor of Neurobiology of Aging at the University of Copenhagen, and elected member of the Norwegian Academy of Science and Letters. 1 Role of lactate as a signaling molecule for neuronal plasticity and memory Pierre J. Magistretti Brain Mind Intsitute, EPFL; Center for Psychiatric Neuroscience, CHUV/UNIL, Lausanne, Switzerland, and Biological and Environmental Sciences and Engineering Division, KAUST, Thuwal, Kingdom of Saudi Arabia [email protected] A tight metabolic coupling between astrocytes and neurons is emerging as a key feature of brain energy metabolism (Allaman et al 2011; Bélanger et al 2011). Over the years we have described two basic mechanisms of neurometabolic coupling. First the glycogenolytic effect of VIP - restricted to cortical columns - and of noradrenaline - spanning across functionally distinct cortical areas - indicating a regulation of brain homeostasis by neurotransmitters acting on astrocytes, as glycogen is exclusively localized in these cells (Magistretti, 2008). Second, the glutamate-stimulated aerobic glycolysis in astrocytes, mediated by the sodium-coupled reuptake of glutamate by astrocytes and the ensuing activation of the Na-K-ATPase, resulting in the release of lactate from astrocytes, which can then fuel the neuronal energy demands a mechanisms known as the ANLS (Pellerin and Magistretti 2011). We have recently shown that lactate derived from astrocytic glycogen is necessary for long-term memory consolidation and for induction of plasticity genes in neurons such as Arc and for maintenance of LTP (Suzuki et al, 2011). This key role of L-lactate in neuronal plasticity mechanisms was demonstrated in experiments in which specific pharmacological and gene expression downregulation interventions were implemented to prevent the production of L-lactate from glycogen - which is exclusively localized in astrocytes - and its release from these cells in the hippocampus during behavioral training (Suzuki et al, 2011). Such interventions completely prevented the establishment of in vivo LTP and long term memory and their effect was fully reversed by the intrahippocampal administration of L-lactate during the training session. The fact that glucose at equicaloric concentrations only marginally mimicked the rescuing effect of L-lactate, was taken as an unexpected indication that the primary mechanism of action of L-lactate on plasticity mechanisms was independent of its ability to act as an energy substrate. We therefore set out to investigate the molecular mechanisms at the basis of the function of L-lactate on neuronal plasticity. We have found that L-lactate stimulates the expression of synaptic plasticity-related genes such as Arc and Zif268 through
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