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Behavioral analysis Phenotype = Genotype X Environment PHENOTYPE – the observable or measurable properties and characteristics of an individual as produced by its genotype and the environment GENOTYPE – the genetic make-up of an individual (the one that you’re born with and the one that is operating at a particular moment) ENVIRONMENT – broadly defined: ambient conditions, developmental conditions, social conditions, internal milieu GENE-ENVIRONMENT INTERPLAY: Gene-environment correlation; Gene-environment interaction DEFINITION OF BEHAVIOR Behavior seen as a function of an organism's environment BEHAVIORISM Behavior seen as a function of an organism's mental processes – COGNITIVE SCIENCE Behavior seen as a function of an organism's biology – NEUROSCIENCE, BEHAVIORAL GENETICS BEHAVIORAL GENETICS GENETIC AND EPIGENETIC THEORIES OF INFORMATION PROCESSING DNA DNA RNA RNA Proteins Proteins Epigenetic networks (open) FUNCTION FUNCTION MOLECULAR NEUROSCIENCE BEHAVIORAL GENETICS – WHAT IS IT ALL ABOUT? Research in behavioral genetics examines the effects of genotype and environment on a range of phenotypic traits such as anxiety, intelligence, sexual orientation, antisocial behavior etc. •Gene for aggression •Gene for homosexuality •Gene for memory •Gene for fear… WHO ARE THE SUBJECTS OF STUDY? HUMANS – families; twins: mono-/dizygous, reared together or apart; isolated populations ANIMALS – rodents, flies, fish, nematodes - normal, genetically segregating populations - selected lines - inbred strains (incl. recombinant and congenic strains) - genetically altered animals (gene amplification, knock-out, mutagenesis) How to correlate behaviors with gene functions? Behavioral analysis of mutants? GENETIC APPROACHES TO BRAIN-BEHAVIOR RELATIONSHIPS REVERSE GENETICS – genotype driven FORWARD GENETICS – phenotype driven The ethics of animal use • Utilitarianism: the view that an action is right if and only if it produces a better balance of benefits and harms than available alternative actions • Consider the “U.S. Government Principles for the Utilization and Care of Vertebrate Animals Used in Testing, Research, and Training” U.S Government Principles (summary) • Procedures involving animals should be designed and performed with due consideration of their relevance to human or animal health, the advancement of knowledge, or the good of society. • The animals selected for a procedure should be of an appropriate species and quality and the minimum number required to obtain valid results. [Alternative] methods…should be considered. • Proper use of animals, including the avoidance or minimization of discomfort, distress, and pain when consistent with sound scientific practices, is imperative. • Procedures with animals that may cause more than momentary or slight pain or distress should be performed with appropriate sedation, analgesia, or anesthesia. • Animals that would otherwise suffer severe or chronic pain or distress that cannot be relieved should be painless killed at the end of the procedure or, if appropriate, during the procedure. U.S Government Principles (continued) • The living conditions of animals should be appropriate for their species and contribute to their health and comfort….veterinary care shall be provided as indicated. • Investigators and other personnel shall be appropriately qualified and experienced for conducting procedures on living animals. • …exceptions…should be made by…an appropriate review group such as an IACUC The three “R’s”: a consensus in the research community •Reduce •Refine •Replace www.jax.org/phenome Paigen K, Eppig JT. A mouse phenome project. Mamm Genome. 2000 Sep;11(9):715-7. Moldin SO et al. Trans-NIH neuroscience initiatives on mouse phenotyping and mutagenesis. Mamm Genome. 2001 Aug;12(8):575-81. Grubb SC et al. A collaborative database of inbred mouse strain characteristics. Bioinformatics. 2004 Nov 1;20(16):2857-9. IACUC • Must have at least a laboratory animal veterinarian, a scientist, and a community member not affiliated with the institution and who does not use animals in research • Reviews all facilities and programs twice a year; identifies and ensures correction of any deficiencies • Reviews “Protocols”, the researcher’s or teacher’s proposal for animal use • Monitors animal use; can suspend any animal activity that does not meet standards The protocol review process (simplified) IACUC Protocol Review • Does the research address an important question? • Does the research require the use of animals? • Is the research necessary, in that it does not needlessly repeat previous work? • Is the experience of each animal in the study adequately described and justified? Factors control behaviors The three “E’s”: • Epigenetic factors • Environmental factors • Experimenter Epigenetic sources of behavioral differences in mice (Francis DD et al. Nat Neurosci 6:445-6, 2003) Crabbe JC and Phillips TJ (Nat Neurosci 6:440-2, 2003) Mother nature meets mother nurture Intrauterine position in mice according to number of flanking males (0M to 2M). In mice, 2M fetuses are exposed to a higher level of androgens than 1M or 0M. Females adjacent to males become partially masculinized in their behavior, and 2M fetuses, both male and female, show increased body weight, greater aggression and territoriality, and enhanced male traits including reproductive behavior (Ryan & Vandenbergh 2002). In rat, overall litter male/female composition may be more important than precise intrauterine position (reviewed by Ryan & Vandenbergh 2002). Conversely, proximity to female fetuses influences male physiological development: prostate glands of males flanked by two females were enlarged compared to controls (Timms et al. 1999); estradiol is implicated (vom Saal et al. 1997). Lathe, R. (2004) The individuality of mice. Genes, Brain & Behavior 3 (6), 317-327. Environmental factors • Housing environment • Enriched environment • Experimental environment Experimenter 이건 아니 자나 왜 이리 실험이 안 되는 거야? Experimental procedures Experiments without habituation ---- Experiments demanding habituation This order was chosen in the attempt to go from the least stressful to the most stressful test. Animals were received at least 2 weeks prior to the beginning of testing and given at least these 2 weeks for acclimation. They were group housed, 3–5/cage, and were at least 8 weeks of age at the start of the experimental series in order to minimize variability due to developmental factors. In addition, two partners (EMBL and Harwell) tested the influence of cage enrichment on behaviour in this test battery. Results are currently being analysed and will be compared and evaluated in the next workpackage meeting at the end of April/04. The “working” SOPs will be audited accordingly. Hierarchical Strategy for Phenotypic (Behavioral) Analysis in Transgenic Mice Rogers et al., 1997 – SHIRPA Crawley and Paylor, 1997 Gold LH (1999) Psychopharmacol. 147:2-4 Two words of behavior • Habituation • Sensitization Repeated stimulations make one boring or sensitive? Stephen Jay Gould on the evolution of Mickey Mouse Supernormal stimuli? vs. Repeated stimulations Repeated stimulation Elicited behavior is not plastic, but can change with repeated stimulation. Habituation effect Decreased response with repeated stimulation. Sensitization effect Increased response with repeated stimulation. Habituation effect Response Trials Dishabituation Dishabituation effect Response Trials Spontaneous recovery Retention Interval Response Trials Location of habituation processes Sensory adaptation Habituation Fatigue How is habituation different from sensory adaptation and fatigue? • Sometimes the habituated response will not recover after for a long time. • (Long Term Habituation) Day Day 11 Responses to Stimulus Day Day 21 21 Repeated Trials of stim Repeated Trials of stim A Habituation will be slower for stimuli which have a higher salience (i.e., intensity) Resp Resp to to weak weak coffee coffee flavor flavor Resp Resp to to Strong Strong coffee coffee flavor flavor Responses to Stimulus Repeated Trials Repeated Trials Dishabituation following the presentation of a novel stimulus. Present Response Response to to Present Stimulus B B stimulus stimulus A A Stimulus (light) (light) (tone) (tone) Responses to Stimulus Response Response to to stimulus stimulus A A (tone) (tone) Repeated Trials Generalization gradient of a habituated response. Responses to Stimulus 1700 1800 1900 2000 2100 2200 2300 Tone frequency of test stimulus following habituation at 2000 Hz Fashion and quality of experiments Sensitization effect Response Trials Sensitization (Arousal) In general Sensitization effects can be influenced by similar parameters as habituation. However, generalization is greater in Sensitization. - Following exposure to cutaneous pain, rats’ reactivity to a wide range of auditory stimuli is increased. Sensitization can be thought of as “arousal”. Habituation and Sensitization in Aplysia Sensitization Pathway Increased serotonin release Dual-Process theory of Habituation and Sensitization 2 underlying processes exist – 1) A Habituation process – 2) A Sensitization process The observable behavior is the sum of these two processes. The habituation effect is observed when the habituation process is greater than the sensitization process. Dual Process examples Sensitization effect Habituation effect S S + + NET - - H NET H Trials Trials Groves and Thompson (1970) S-R System: Habituation is thought to occur in the reflex arc. State System: Sensitization is thought to occur in the part of the nervous system that determines general responsiveness. Infant attention Looking Time Trials Opponent Process Theory of Motivation