1 NIH Public Access Author Manuscript Published in final edited form as: Eur J Pharmacol September 24; 571(1): Time dependent decreases in central α 7 nicotinic acetylcholine receptors associated with haloperidol and risperidone treatment in rats Alvin V. Terry Jr. and Debra A. Gearhart Department of Pharmacology and Toxicology, Medical College of Georgia, Augusta, Georgia, 30912, United States Abstract α 7 nicotinic acetylcholine receptor deficits may contribute to cognitive dysfunction in schizophrenia; however, the contribution of antipsychotic drug exposure to these deficits is unknown. In this study, rats were treated orally with haloperidol (2.0 mg/kg/day) or risperidone (2.5 mg/kg/day) for 15 or 90 days. Subsequent immunoassays indicated that both antipsychotics were associated with α 7 nicotinic receptor decreases in the basal forebrain and prefrontal cortex when administered for 90 (but not 15) days, a result that was confirmed in autoradiographic experiments. These data suggest that haloperidol and risperidone may be associated with time dependent decreases in an important neurobiological substrate of memory. Keywords schizophrenia; memory; cognition; antipsychotic; acetylcholine; cholinergic 1. Introduction Low affinity α 7 nicotinic acetylcholine receptors comprise approximately 10% of the total number of nicotinic receptors in mammalian brain, yet they are believed to modulate a variety of calcium-dependent events in neurons including neurotransmitter release (McGehee et al., 1995; Gray et al., 1996), postsynaptic signaling (Chang & Berg, 1999; Hefft et al., 1999) and neuronal survival (Messi et al., 1997; Berger et al., 1998). There is also a significant body of evidence to support an important role of α 7 nicotinic receptors in the cognitive dysfunction associated with certain chronic psychiatric illnesses such as schizophrenia. This premise is supported by postmortem evidence of α 7 nicotinic receptor deficits in the hippocampus and frontal cortex of schizophrenic patients (Guan et al., 1999) and by linkage analysis implicating chromosome 15q14 (the region that includes the α 7 neuronal nicotinic acetylcholine receptor gene). Polymorphisms in the core promoter of the α 7 neuronal nicotinic acetylcholine receptor gene (CHRNA7; GeneBank accession no. Z23141) are associated with reduced inhibition of the P50 evoked response to repeated auditory stimuli (i.e., indicative of sensory gating abnormalities) in schizophrenia (reviewed, Freedman et al., 2003). α 7 nicotinic receptor deficits may also contribute to abnormalities of smooth pursuit eye movements, sustained attention, Corresponding Author: Alvin V. Terry Jr., Ph.D., Professor of Pharmacology and Toxicology, CJ-1020, Medical College of Georgia, 1120 Fifteenth Street, Augusta, Georgia , United States, Phone , Fax , Publisher's Disclaimer: This is a PDF file of an unedited manuscript that has been accepted for publication. As a service to our customers we are providing this early version of the manuscript. The manuscript will undergo copyediting, typesetting, and review of the resulting proof before it is published in its final citable form. Please note that during the production process errors may be discovered which could affect the content, and all legal disclaimersthat apply to the journal pertain.
2 Terry and Gearhart Page 2 and cognitive performance often observed in schizophrenia (reviewed Martin et al., 2004). Collectively, these findings have lead to the suggestion that a reduction in α 7 nicotinic receptor expression could serve as a biomarker for schizophrenia (Perl et al., 2003; Freedman et al., 2005). It is important to note, however, that the extent to which antipsychotic drugs contribute to α 7 nicotinic receptor alterations in schizophrenia is unknown given that the brains of antipsychotic-naïve patients have rarely been analyzed. This is an important consideration since many schizophrenia patients are treated with antipsychotic drugs for decades. In previous work in our laboratory, we have detected time dependent alterations in several cholinergic marker proteins including choline acetyltransferase, the vesicular acetylcholine transporter, and the high affinity choline transporter in rats treated with either representative first or second generation antipsychotics (see Terry et al., 2007 and review, Terry and Mahadik, 2007). Typically, we have observed elevations in cholinergic markers early in treatment (e.g., 15 days) followed by decreases below control levels at longer time periods (e.g., 90 days and beyond). We have also observed persistent decreases in α 7 nicotinic receptors in association with risperidone administered for 180 days (Terry et al., 2005), but have not determined if such a biphasic effect (as observed with other cholinergic proteins) occurs with α 7 nicotinic receptors. Therefore, in the present study in rats, the effects chronic oral treatment with a representative first generation antipsychotic, haloperidol, and a representative second generation antipsychotic, risperidone, on the levels of α 7 nicotinic receptors in several (memory-related) brain regions were investigated. The antipsychotics were delivered in drinking water and dosed on a mg/kg/24 hour basis for 15 or 90 days. This dosing approach was shown previously to result in plasma levels in rats that are considered within the therapeutic range for humans (Terry et al., 2005; Terry et al., 2007). α 7 nicotinic receptors were subsequently measured by ELISA and autoradiographic methods. 2. Materials and Methods 2.1 Subjects and Treatment Three month old Wistar rats were treated with haloperidol (2.0 mg/kg/day) or risperidone (2.5 mg/kg/day) orally in drinking water as described previously (see Terry et al., 2007) for periods of 15 or 90 days. All procedures employed during this study were reviewed and approved by the Medical College of Georgia Institutional Animal Care and Use Committee and are consistent with AAALAC guidelines. Measures were taken to minimize pain or discomfort in accordance with the National Institute of Health Guide for the Care and Use of Laboratory Animals (NIH Publications No ) revised Significant efforts were also made to minimize the total number of animals used while maintaining statistically valid group numbers. 2.2 ELISA for α 7 nicotinic acetylcholine receptors Euthanasia of rats, brain dissections, and preparation of brain lysates were done according to Gearhart et al. (2006). Briefly, basal forebrain, hippocampal formation, cortex, and prefrontal cortex were dissected and then homogenized in RIPA buffer containing protease inhibitors and glycerol. The Micro BCA Protein Assay Kit (Pierce Biotechnology; Rockford, IL) was used to determine the total protein concentration in each brain lysate. An indirect ELISA was devised to measure the relative levels α 7 nicotinic receptor protein in brain lysates. As an internal control for day-to-day variation in the ELISA methods, brain lysates from vehicle, haloperidol, and risperidone-treated rats were assayed on the same ELISA plate. Brain lysates (15-25 μg total protein/μl) were diluted in sodium carbonate buffer (ph 9.6), and then the diluted lysates (0.5 ug protein/50 μl/well) were adsorbed to Nunc Maxisorp TM ELISA plates (overnight at 5 C). Microwells were rinsed once with 300 μl of wash buffer (ph 7.4 phosphate-buffered saline containing 0.05% (v/v) Triton X-100), before blocking the wells for one hour with 300 μl/well
3 Terry and Gearhart Page 3 1% (w/v) nonfat dry milk in PBST. Wells were rinsed with wash buffer (300 μl/well), before adding 50 μl/well of α 7 nicotinic receptor antibody (Chemicon AB5637; rabbit polyclonal; diluted 1:100 in blocking buffer). Incubation with primary antibody was carried out at room temperature for two hours, and then the microwells were rinsed with wash buffer (five times, 300 μl/well). The primary antibody was detected using peroxidase (HRP)-conjugated goat antirabbit IgG (Jackson Immunoresearch # , diluted 1:10,000 in blocking buffer, 50 μl/well). After one hour at room temperature, the microplate was rinsed five times with wash buffer (300 μl/well). Tetramethylbenzidine (100 μl/well, KPL ) was the peroxidase substrate; after minutes at room temperature, 1M HCl (100 μl/well) was added to stop the peroxidase reaction (color changed from blue to yellow). The absorbance of the yellow reaction product was measured at 450 nm using a microplate spectrophotometer (BioTek Instruments, Inc., Winooski, VT). 2.3 Quantitative Receptor Autoradiography 2.4 Statistical Analyses 3. Results 3.1 ELISA Data In order to further assess the effects of chronic antipsychotic administration on α 7 nicotinic receptor densities at the 90 day treatment time point, autoradiographic analyses of brain tissues harvested from rats treated with haloperidol, risperidone, or vehicle were conducted with the subtype specific cholinergic radioligand [ 125 I]-α-bungarotoxin. Test subjects (N=7-8) were sacrificed by decapitation and the brains flash frozen. Using a Microm HM cryostat (-18 C), the left hemisphere of each brain was serially sectioned (16 μm) up to the midline, and sections were adhered onto chrome alum/gelatin-coated slides. The concentration of [ 125 I]-αbungarotoxin used was 1.0 nm, the incubation time was 120 minutes, and the duration of film exposure was 160 hours. Please refer to previous publications for more detailed descriptions of the autoradiographic procedures (Hernandez and Terry, 2005; Terry et al., 2005). Statistical analyses were made using a one or two-way analysis of variance (ANOVA) with repeated measures when indicated. Dunnett s procedure was used to examine post hoc differences (P value <0.05 considered significant). Figure 1 depicts the ELISA results from the four selected brain regions in rats treated with vehicle, haloperidol, or risperidone for 15 or 90 days. Results from drug-treated rats are presented as a percent of control (vehicle-treated rats). Statistically significant time x brain region interactions were observed for both haloperidol and risperidone, which had a P-value for the interaction of <0.05. The statistically significant interaction indicates that the effect of time on the normalized OD values was different in the different brain regions. As indicated in Figure 1, post hoc comparisons revealed that both haloperidol and risperidone were associated with significant decreases (P<0.05) in α 7 nicotinic receptor protein in the basal forebrain as well as prefrontal cortex. Significant decreases were also observed in the cortex at 90 days in animals administered risperidone. 3.2 Autoradiographic Data The results of autoradiographic experiments are depicted in Figure 2. Representative autoradiograms are illustrated in the top left of the figure. The pattern of [ 125 I]-α-bungarotoxin binding was similar to that observed in previous studies (Hernandez and Terry, 2005;Terry et al., 2005). Statistical analyses revealed that there were significant treatment-related differences in binding densities F 2,18 =4.2, P=0.03, a highly significant difference in binding in the different brain regions F 6,12 =253.7, P<0.001, without a significant treatment x brain region
4 Terry and Gearhart Page 4 4. Discussion Acknowledgements References interaction F 105,143 =1.3, P=0.31. Post hoc statistical comparisons revealed the following: 1) risperidone was associated with statistically significant decreases (P<0.05) in [ 125 I]-αbungarotoxin binding in the prefrontal cortex, and the CA1, CA2, and CA3 regions of the hippocampus, while haloperidol was associated with significant decreases (P<0.05) in binding in the CA1 and CA2 regions of the hippocampus. A perusal of the histograms in Figure 2 reveals however, that [ 125 I]-α-bungarotoxin binding was at least slightly decreased by both haloperidol and risperidone treatment in all of the brain regions that were analyzed (an effect that replicates an earlier study at 90 days, Terry et al., 2005). The results of this study can be summarized as follows: 1) neither haloperidol nor risperidone significantly affected the density of α 7 nicotinic receptors in memory-related brain regions in the rat when they were administered for 15 days; 2) both antipsychotics were, however, associated with significant α 7 nicotinic receptor decreases in the basal forebrain and prefrontal cortex when administered for 90 days (i.e., by as much as 22% in the case of haloperidol and 15% in the case of risperidone); 3) quantitative receptor autoradiography experiments with the radioligand [ 125 I]-α-bungarotoxin on tissues from animals that were treated with antipsychoitcs for 90 days revealed layer specific decreases in α 7 nicotinic receptors in the hippocampus and confirmed ELISA results in the prefrontal cortex. These data support an accruing body of evidence (see Terry and Mahadik, 2007) that representative first and second generation antipsychotics exert a time-dependent effect on cholinergic marker proteins in memory-related regions of mammalian brain. While we did detect a time dependent decrease in α 7 nicotinic receptors in the present study, we did not observe an increase above baseline at the early time point followed by a decease below baseline with longer treatment as we have found with other cholinergic proteins such as the vesicular acetylcholine transporter (Terry et al., 2007) in animals treated with haloperidol or risperidone. The mechanism for the time-dependent effect of these antipsychotics on cholinergic proteins is currently unknown, but it has also been observed in association with first generation antipsychotics in older studies which focused on elucidating the mechanism for their adverse motor effects (Miller and Chouinard, 2003). Given the important role of α 7 nicotinic receptors in neuronal function, cognition, and the pathophysiology of schizophrenia, it is important that the mechanism for this effect be elucidated. It is also important to determine if such receptor effects are associated with other antipsychotics given their widespread use in the schizophrenia population as well in a variety of other psychiatric and neurologic illnesses (e.g., autism, bipolar disease, Parkinson s disease, Alzheimer s disease). A better understanding of such effects might facilitate the development of new antipsychotics that do not affect α 7 nicotinic receptors (or other cholinergic proteins) and the development of adjunctive therapeutic agents designed to improve cognition. Specifically, a thorough knowledge of the chronic effects of different antipsychotic drugs on cholinergic marker proteins might facilitate more patient specific (i.e., tailored) therapeutic strategies that involve a cholinergic replacement strategy. The authors wish to acknowledge the technical support of Kristy Bouchard, Samantha Warner, Sameera Warsi, and Mary-Louise Middlemore. This work was supported by the National Institute of Mental Health (R01 MH to AVT). Berger F, Gage FH, Vijayaraghavan S. Nicotinic receptor-induced apoptotic cell death of hippocampal progenitor cells. J Neurosci 1998;18: [PubMed: ]
5 Terry and Gearhart Page 5 Chang KT, Berg DK. Nicotinic acetylcholine receptors containing alpha7 subunits are required for reliable synaptic transmission in situ. J Neurosci 1999;19: [PubMed: ] Freedman R, Olincy A, Ross RG, Waldo MC, Stevens KE, Adler LE, Leonard S. The genetics of sensory gating deficits in schizophrenia. Curr Psychiatry Rep 2003;5: [PubMed: ] Freedman R, Ross R, Leonard S, Myles-Worsley M, Adams CE, Waldo M, Tregellas J, Martin L, Olincy A, Tanabe J, Kisley MA, Hunter S, Stevens KE. Early biomarkers of psychosis. Dialogues Clin Neurosci 2005;7: [PubMed: ] Gearhart DA, Middlemore ML, Terry AV Jr. ELISA methods to measure cholinergic markers and nerve growth factor receptors in cortex, hippocampus, prefrontal cortex, and basal forebrain from rat brain. J Neurosci Meth 2006;150: Gray R, Rajan AS, Radcliffe KA, Yakehiro M, Dani JA. Hippocampal synaptic transmission enhanced by low concentrations of nicotine. Nature 1996;383: [PubMed: ] Guan ZZ, Zhang X, Blennow K, Nordberg A. Decreased protein level of nicotinic receptor alpha7 subunit in the frontal cortex from schizophrenic brain. Neuroreport 1999;10: [PubMed: ] Hefft S, Hulo S, Bertrand D, Muller D. Synaptic transmission at nicotinic acetylcholine receptors in rat hippocampal organotypic cultures and slices. J Physiol 1999;515: [PubMed: ] Hernandez CM, Terry AV Jr. Repeated nicotine exposure in rats: effects on memory function, cholinergic markers and nerve growth factor. Neuroscience 2005;130: [PubMed: ] Martin LF, Kem WR, Freedman R. Alpha-7 nicotinic receptor agonists: potential new candidates for the treatment of schizophrenia. Psychopharmacology (Berl) 2004;174: [PubMed: ] Messi ML, Renganathan M, Grigorenko E, Delbono O. Activation of alpha7 nicotinic acetylcholine receptor promotes survival of spinal cord motoneurons. FEBS Lett 1997;411: [PubMed: ] McGehee DS, Heath MJ, Gelber S, Devay P, Role LW. Nicotine enhancement of fast excitatory synaptic transmission in CNS by presynaptic receptors. Science 1995;269: [PubMed: ] Miller R, Chouinard G. Loss of striatal cholinergic neurons as a basis for tardive and L-dopa-induced dyskinesias, neuroleptic-induced supersensitivity psychosis and refractory schizophrenia. Biol Psychiatry 1993;34: [PubMed: ] Perl O, Ilani T, Strous RD, Lapidus R, Fuchs S. The alpha7 nicotinic acetylcholine receptor in schizophrenia: decreased mrna levels in peripheral blood lymphocytes. FASEB J 2003;17: [PubMed: ] Terry AV Jr, Gearhart DA, Mahadik SP, Warsi S, Davis LW, Waller JL. Chronic exposure to typical or atypical antipsychotics in rodents: temporal effects on central alpha 7 nicotinic acetylcholine receptors. Neuroscience 2005;136: [PubMed: ] Terry AV Jr, Mahadik SP. Time Dependent Cognitive Deficits Associated with First and Second Generation Antipsychotics: Cholinergic Dysregulation as a Potential Mechanism. J Pharmacol Exper Ther 2007;320: [PubMed: ] Terry AV Jr, Gearhart DA, Warner SE, Zhang G, Bartlett MG, Middlemore ML, Beck WD, Mahadik SP, Waller JL. Oral haloperidol or risperidone treatment in rats: temporal effects on nerve growth factor receptors, cholinergic neurons, and memory performance. Neuroscience 2007;146: [PubMed: ]
6 Terry and Gearhart Page 6 Fig 1. ELISA results for α 7 nicotinic receptor protein expressed as percent of vehicle-treated group (mean ± S.E.M.) in the brain regions indicated from rats treated with vehicle, haloperidol (2.0 mg/kg/24 h), or risperidone (2.5 mg/kg/24 h) for 15 or 90 days. Rats were euthanized at each of the time points indicated and then the dissected brain regions were processed for ELISA. *P<0.05 = significantly different from vehicle control. N=5-6.
7 Terry and Gearhart Page 7 Fig 2. Top Left: Representative autoradiograms illustrating receptors labeled by [ 125 ]-α-bungarotoxin (ligand at the α 7 nicotinic receptor), in 16 μm sagittal sections of brains from Wistar rats treated with vehicle, haloperidol (2.0 mg/kg/24 h), or risperidone (2.5 mg/kg/24 h) for 90 days. PFC = prefrontal cortex; HIPP = hippocampus; CTX = cortex (layers I-VI); SI = substantia inominata. Histograms illustrate the effects of 90 days of antipsychotic treatment on the density of receptors labeled by [ 125 ]-α-bungarotoxin in different (memory-related) brain regions in the rat. *P<0.05 = significantly different from vehicle control. VEH = vehicle, HAL = haloperidol; RISP = risperidone. N=7-8 rats per group.
2009 H1N1 Influenza ( Swine Flu ) Hemagglutinin ELISA kit Catalog Number : SEK001 To achieve the best assay results, this manual must be read carefully before using this product and the assay is run as
EGFR (py1045)/ Pan EGFR (Human) ELISA Kit Catalog Number KA2156 96 assays Version: 01 Intended for research use only www.abnova.com I. INTRODUCTION EGFR (py1045)/pan EGFR (Human) ELISA (Enzyme-Linked Immunosorbent
Influenza A H1N1 (Swine Flu 2009) Hemagglutinin / HA ELISA Pair Set Catalog Number : SEK001 To achieve the best assay results, this manual must be read carefully before using this product and the assay
Toxoplasma gondii IgM ELISA Kit Catalog Number KA0226 96 assays Version: 01 Intended for research use only www.abnova.com Table of Contents Introduction... 3 Intended Use... 3 Background... 3 Principle
Phospho-Akt (pser 473 ) ELISA Kit for detection of human, mouse, or rat phospho-akt (pser 473 ) in cell and tissue lysates Catalog Number RAB0011 Storage Temperature 20 C TECHNICAL BULLETIN Product Description
STAT3 (py705)/ Pan STAT3 (Human/Mouse/Rat) ELISA Kit Catalog Number KA2176 96 assays Version: 02 Intended for research use only www.abnova.com Table of Contents Introduction... 3 Principle of the Assay...
STAT1 (ps727) (Human/Mouse) ELISA Kit Catalog Number KA2171 96 assays Version: 01 Intended for research use only www.abnova.com I. INTRODUCTION STAT1 (ps727) (Human/Mouse) ELISA (Enzyme-Linked Immunosorbent
colorimetric sandwich ELISA kit datasheet For the quantitative detection of rat TNF-alpha concentrations in cell culture supernatants. general information Catalogue Number Product Name Species cross-reactivity
A helping hand for your research Product Manual Human Apolipoprotein A1 EIA Kit Catalog Number: 83901 96 assays 1 Table of Content Product Description 3 Assay Principle 3 Kit Components 3 Storage 4 Reagent
Human Immunodeficiency Virus type 1 (HIV-1) p24 / Capsid Protein p24 ELISA Pair Set Catalog Number : SEK11695 To achieve the best assay results, this manual must be read carefully before using this product
Product Manual Human Leptin ELISA Kit Catalog Numbers MET-5057 MET-5057-5 96 assays 5 x 96 assays FOR RESEARCH USE ONLY Not for use in diagnostic procedures Introduction Leptin is a polypeptide hormone
H5N1 ( Avian Flu ) Hemagglutinin ELISA Pair Set Catalog Number : SEK002 To achieve the best assay results, this manual must be read carefully before using this product and the assay is run as summarized
RayBio Human Phospho-DDR2 (Tyr740) and Total DDR2 ELISA Kit Catalog #: PEL-DDR2-Y740-T User Manual Last revised March 22, 2018 Caution: Extraordinarily useful information enclosed ISO 13485 Certified 3607
Schizophrenia: Uncovering a molecular origin so humanity may better deal with insanity : What is Schizophrenia? Literally split mind A mental disorder comprising most major psychotic disorders; characterized
Global Histone H3 Acetylation Assay Kit Catalog Number KA0633 96 assays Version: 06 Intended for research use only www.abnova.com Table of Contents Introduction... 3 Intended Use... 3 Background... 3 Principle
K-ASSAY KAMIYA BIOMEDICAL COMPANY KAMIYA BIOMEDICAL COMPANY Estrone EIA kit For the quantitative determination of estrone in dried fecal extracts, urine and tissue culture media Cat. No. KT-720 For Research
Cogs 107b Systems Neuroscience www.dnitz.com lec9_02042010 neuromodulators and drugs of abuse principle of the week: functional anatomy Professor Nitz circa 1986 neurotransmitters: mediating information
DIAGNOSTIC AUTOMATION, INC. 21250 Califa Street, Suite 102 and 116, Woodland hills, CA 91367 Tel: (818) 591-3030 Fax: (818) 591-8383 firstname.lastname@example.org email@example.com www.rapidtest.com
Package Insert Apolipoprotein A1 (Apo A1) ELISA 1 x 96 Wells For Research Use Only (RUO). Not for use in clinical, diagnostic or therapeutic procedures. v. 1.0 Eagle Biosciences, Inc. 20A Northwest Blvd.,
EpiQuik Total Histone H3 Acetylation Detection Fast Kit (Colorimetric) Base Catalog # PLEASE READ THIS ENTIRE USER GUIDE BEFORE USE The EpiQuik Total Histone H3 Acetylation Detection Fast Kit (Colorimetric)
h AFLATOXIN M1 CAT. NO. 961AFLMO1M Competitive ELISA Immunoassay for the quantitative detection of Aflatoxin M1 in milk, milk powder and cheese. General Aflatoxins are toxic metabolites produced by a variety
Mycoplasma pneumoniae IgG ELISA Kit Catalog Number KA2260 96 assays Version: 01 Intended for research use only www.abnova.com Table of Contents Introduction... 3 Intended Use... 3 Background... 3 Principle
RayBio Human Phosphotyrosine BTK ELISA Kit Catalog #: PEL-BTK-Y User Manual Last revised August 10, 2016 Caution: Extraordinarily useful information enclosed ISO 13485 Certified 3607 Parkway Lane, Suite
Hepatitis A virus IgM ELISA Kit Catalog Number KA0285 96 assays Version: 04 Intended for research use only www.abnova.com Table of Contents Introduction... 3 Intended Use... 3 Background... 3 Principle
AlphaScreen : A Straightforward and Powerful Alternative to ELISA Martina Bielefeld-Sévigny Ph.D., R&D Director Overview AlphaScreen - an alternative to ELISA Why an alternative to ELISA? Assay principle
ab109908 Complex II Enzyme Activity Microplate Assay Kit Instructions for Use For the quantitative measurement of Complex II activity in samples from Human, Rat, Mouse and Cow This product is for research
Dopamine Discovery Day August 30, 2012 Rikshospitalet Store auditorium, Oslo, Norway Organized by Linda H. Bergersen & Vidar Gundersen Institute of Basic Medical Sciences & Centre for Molecular Biology
EpiQuik Global Histone H4 Acetylation Assay Kit Base Catalog # PLEASE READ THIS ENTIRE USER GUIDE BEFORE USE The EpiQuik Global Histone H4 Acetylation Assay Kit is suitable for specifically measuring global
Campylobacter Antigen ELISA Kit Catalog Number KA3204 96 assays Version: 01 Intended for research use only www.abnova.com Table of Contents Introduction... 3 Intended Use... 3 Background... 3 Principle
Chemical Control of Behavior and Brain 1 of 9 I) INTRO A) Nervous system discussed so far 1) Specific 2) Fast B) Other systems extended in space and time 1) Nonspecific 2) Slow C) Three components that
9/12/11 Chapter 12 Nervous Tissue Overview of the nervous system Cells of the nervous system Electrophysiology of neurons Synapses Neural integration Subdivisions of the Nervous System 1 Subdivisions of
WHO Prequalification of Diagnostics Programme PUBLIC REPORT Product: Bioelisa HCV 4.0 Number: PQDx 0165-060-00 Abstract Bioelisa HCV 4.0 with product codes 3000-1115 and 3000-1116, manufactured by Biokit
LH (Bovine) ELISA Kit Catalog Number KA2280 96 assays Version: 05 Intended for research use only www.abnova.com Table of Contents Introduction... 3 Intended Use... 3 Background... 3 Principle of the Assay...
FSH (Rodent) ELISA Kit Catalog Number KA2330 96 assays Version: 06 Intended for research use only www.abnova.com Table of Contents Introduction... 3 Intended Use... 3 Background... 3 Principle of the Assay...
Chapter 10 Nervous System I Divisions of the Nervous System Cell Types of Neural Tissue neurons neuroglial cells Central Nervous System brain spinal cord Peripheral Nervous System nerves cranial nerves
Synaptic Communication Steven McLoon Department of Neuroscience University of Minnesota 1 Course News The first exam is next week on Friday! Be sure to checkout the sample exam on the course website. 2
Influenza A H1N1 HA ELISA Pair Set for H1N1 ( A/Puerto Rico/8/1934 ) HA Catalog Number : SEK11684 To achieve the best assay results, this manual must be read carefully before using this product and the
BIO 211: ANATOMY & PHYSIOLOGY I 1 Ch 10 A Ch 10 B This set CHAPTER 10 NERVOUS SYSTEM 1 BASIC STRUCTURE and FUNCTION Dr. Lawrence G. Altman www.lawrencegaltman.com Some illustrations are courtesy of McGraw-Hill.
2-Deoxyglucose Assay Kit (Colorimetric) Catalog Number KA3753 100 assays Version: 01 Intended for research use only www.abnova.com Table of Contents Introduction... 3 Background... 3 General Information...
The Nervous System Functional Neuroanatomy and How Neurons Communicate Mark Stanford, Ph.D. Santa Clara Valley Health & Hospital System Addiction Medicine and Therapy Services The Nervous System In response
OxisResearch A Division of OXIS Health Products, Inc. BIOXYTECH pl GPx Enzyme Immunoassay Assay for Human Plasma Glutathione Peroxidase For Research Use Only. Not For Use In Diagnostic Procedures. Catalog
Autonomic Nervous System Lanny Shulman, O.D., Ph.D. University of Houston College of Optometry Peripheral Nervous System A. Sensory Somatic Nervous System B. Autonomic Nervous System 1. Sympathetic Nervous
Egg Protein Residue ELISA Kit Catalog Number KA3208 96 assays Version: 01 Intended for research use only www.abnova.com Table of Contents Introduction... 3 Intended Use... 3 Background... 3 Principle of
Biomarkers Workshop In Clinical Trials Imaging for Schizophrenia Trials Research focused on the following areas Brain pathology in schizophrenia and its modification Effect of drug treatment on brain structure
CHAPTER 7 The Nervous System: Neurons and Synapses Chapter 7 Outline Neurons and Supporting Cells Activity in Axons The Synapse Acetylcholine as a Neurotransmitter Monoamines as Neurotransmitters Other
ab109714 ATP Synthase Enzyme Activity Microplate Assay Kit Instructions for Use For the quantitative measurement of ATP Synthase activity in samples from Human, Rat and Cow This product is for research
Section: Chapter 5: Multiple Choice 1. The structure of synapses is best viewed with a(n): p.155 electron microscope. light microscope. confocal microscope. nissle-stained microscopic procedure. 2. Electron
LIVE INTERACTIVE LEARNING @ YOUR DESKTOP Understanding the Brain: What Drugs Can Tell Us Presented by: Dr. Rochelle D. Schwartz-Bloom March 24, 2011 Understanding the Brain: What Drugs Can Tell Us Rochelle
The Protein Experts Manual Cytoskeleton, Inc. V. 6.0 Precision Red Advanced Protein Assay Reagent Cat. # ADV02 cytoskeleton.com Phone: (303) 322.2254 Fax: (303) 322.2257 Customer Service: firstname.lastname@example.org
Schizophrenia, Sensory Gating, and Nicotinic Receptors by Lawrence E. Adler, Ann Olincy, Merilyne Waldo, Josette Q. Harris, Jay Qriffith, Karen Stevens, Karen Flach, Herbert Nagamoto, Paula Bickford, Sherry
Synaptic Transmission Graphics are used with permission of: Pearson Education Inc., publishing as Benjamin Cummings (http://www.aw-bc.com) Page 1. Introduction Synaptic transmission involves the release
Mercodia Porcine C-peptide ELISA Directions for Use 10-1256-01 REAGENTS FOR 96 DETERMINATIONS Manufactured by Mercodia AB Sylveniusgatan 8A SE-754 50 Uppsala Sweden INTENDED USE Mercodia Porcine C-peptide
Estriol ELISA kit Catalog Number KA0316 96 assays Version: 05 Intended for research use only www.abnova.com Table of Contents Introduction... 3 Background... 3 Principle of the Assay... 3 General Information...
Helicobacter pylori IgA ELISA Kit Catalog Number KA0964 96 assays Version: 03 Intended for research use only www.abnova.com Table of Contents Introduction... 3 Intended Use... 3 Background... 3 Principle
The Nervous System Dr. ZHANG Xiong Dept. of Physiology ZJU School of Medicine Http://10.10.10.151 Part 1. Summary of the nervous system The Nervous System Central Nervous System Brain + Spinal Cord Peripheral
List of contents Ralstonia solanacearum PathoScreen Kit Lot number Item 96 wells 288 wells 480 wells Antibody coated 96-well microtiter plates 1 3 5 Peroxidase enzyme conjugate, ready to use 11 ml 33 ml
BL-1020: First-in-Class GABA-Enhanced Antipsychotic For Schizophrenia December 2012 Forward Looking Statements This presentation contains "forward-looking statements." These statements include words like
Overview Schizophrenia Psychology 372 Physiological Psychology Steven E. Meier, Ph.D. Listen to the audio lecture while viewing these slides Probably consists of more than one disorder Is the most devastating
Potato leafroll virus (PLRV) Reagent Set List of contents Lot number Item 96 wells 500 wells 1000 wells 5000 wells Capture antibody 0.150 ml 0.275 ml 0.525 ml 2.525 ml Alkaline phosphatase enzyme conjugate
RayBio Human ENA-78 ELISA Kit Catalog #: ELH-ENA78 User Manual Last revised April 15, 2016 Caution: Extraordinarily useful information enclosed ISO 13485 Certified 3607 Parkway Lane, Suite 100 Norcross,
Neurotransmitter Systems II Receptors Reading: BCP Chapter 6 Neurotransmitter Systems Normal function of the human brain requires an orderly set of chemical reactions. Some of the most important chemical
CSE511 Brain & Memory Modeling Lect 22,24,25: Memory Systems Compare Chap 31 of Purves et al., 5e Chap 24 of Bear et al., 3e Larry Wittie Computer Science, StonyBrook University http://www.cs.sunysb.edu/~cse511
- Neurotransmitters Of The Brain - INTRODUCTION Synapsis: a specialized connection between two neurons that permits the transmission of signals in a one-way fashion (presynaptic postsynaptic). Types of
CASE 4 A 32-year-old woman presents to her primary care physician s office with difficulty chewing food. She states that when she eats certain foods that require a significant amount of chewing (meat),
Neural Basis of Motor Control Central Nervous System Skeletal muscles are controlled by the CNS which consists of the brain and spinal cord. Determines which muscles will contract When How fast To what
Mercodia Porcine C-peptide ELISA Directions for Use 10-1256-01 REAGENTS FOR 96 DETERMINATIONS Manufactured by Mercodia AB, Sylveniusgatan 8A, SE-754 50 Uppsala, Sweden EXPLANATION OF SYMBOLS USED ON LABELS
Biology 105 Midterm Exam 3 Review Sheet The third midterm exam will cover the following lecture material (lectures 8, 9, 10, 11, 12, and 13): Skeletal System (from chapter 5 in the textbook), Nervous System
INTENDED USE The ELEGANCE Chlamydia pneumoniae IgG ELISA has been designed for the in vitro diagnostic measurement of anti- Chlamydia pneumoniae IgG in the screening of human serum. PRINCIPLES OF THE ELEGANCE
A. Introduction CNS Tour (Lecture 12) There are to a chemical pathways in the nervous system. These pathways also form different neurological structures B. Spinal Cord Receives sensory neurons from skin
ELEC ENG 3BB3: Cellular Bioelectricity Notes for Lecture 22 Friday, February 28, 2014 10. THE NEUROMUSCULAR JUNCTION We will look at: Structure of the neuromuscular junction Evidence for the quantal nature
THE NERVOUS SYSTEM Homeostasis Strand Introduction In general, a nervous system has three overlapping functions : 1. Sensory input conduction of signals from sensory receptors to integration centres 2.