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Lecture 1 - Flash (Large) - 20120912 02.29.56PM
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  1. NEUR 3200 Principles of Neuroscience
  2. Announcements
  3. Lecture 1
  4. Reading
  5. Lecture 1 outline
  6. What is Neuroscience?
  7. Clinical neuroscience
  8. Neuroscience involves learning a language
  9. Lecture 1 outline
  10. Cells of the nervous system
  11. Internal structures of a cell
  12. Neurons: Various classifications possible
  13. Massive diversity of neuronal structure
  14. Neuron structure: synapses
  15. Synaptic signaling can be functionally fast or slow
  16. Synaptic signaling can be functionally fast or slow
  17. Neuronal function
  18. Molecular functions
  19. Cellular functions
  20. Cellular functions
  21. Systems-level functions
  22. Behavioural functions
  23. Lecture 1 outline
  24. Glia
  25. Astrocytes
  26. Glial stem cells classes as astrocytes
  27. Oligodendrocytes
  28. Oligodendroglial precursors
  29. Microglial cells
  30. Lecture 1 outline
  31. Comparative neuronal organization
  32. Sponges (Porifera phylum)
  33. Hydra (Cnidaria phylum)
  34. Hydra (Cnidaria phylum)
  35. Hydra (Cnidaria phylum)
  36. Hydra (Cnidaria phylum)
  37. Hydra (Cnidaria phylum)
  38. Flatworms (Phylum Platyhelminthes)
  39. Flatworms (Phylum Platyhelminthes)
  40. Flatworms (Phylum Platyhelminthes)
  41. Myotatic reflex in humans
  42. Neuronal activity in the myotatic reflex
  43. Phyla Annelida and arthropoda
  44. Phyla Annelida and arthropoda
  45. Phylum Chordata
  46. Phylum Chordata
  47. Phylum Chordata
  48. Lecture 1 outline
  49. Big chordate brains allow complex behaviour
  50. Human evolution: Primates
  51. Human evolution: Apes
  52. Human evolution: Homo sapiens
  53. Human culture is more than our biology
  54. Comparison of brain sizes
  55. Evolution of the human brain
  56. Why did big brains evolve? Possible answer…
  57. Big brains cause problems…
  58. Size is just part of the story…
  59. Lecture 1 outline
  60. Organization of the human nervous system
  61. Lecture 1 outline
  62. Organization of the human nervous system
  63. Organization of the human nervous system
  64. Structural organization of the brain
  65. Structural organization of the brain
  66. Organization of long-distance connections
  67. Lecture 1 outline
  68. Histology
  69. Fixing the brain
  70. Sectioning the brain
  71. Staining the brain
  72. Studying cell structure
  73. Studying cell structure
  74. Studying cell structure
  75. Alternatives to Golgi
  76. Staining for specific molecules
  77. Nissl stain
  78. Weil stain
  79. In situ hybridization
  80. Analysis of neural activity by ISH
  81. Immunohistochemistry (immunocytochemistry)
  82. Immuno vs. in situ
  83. Examining the slice
  84. Allen Brain Atlas
  85. Slide 84
  86. Slide 85
  87. Neuroscience Information Framework (aside)
  88. Neuroscience Information Framework (aside)
  89. Neuroscience Information Framework (aside)
  90. Neuroscience Information Framework (aside)
  91. Lecture 1 outline
  92. Determining neuronal circuits
  93. Neural tracing methods
  94. Anterograde labeling
  95. Neural tracing methods
  96. Neural tracing methods
  97. Functional analysis of neural systems
  98. Mechanical brain lesion/ablation
  99. Chemical brain lesions
  100. Brain lesions: Pros and cons
  101. Stereotaxic surgery
  102. Stereotaxic surgery
  103. Stereotaxic surgery
  104. Functional analysis of neural systems
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NEUR 3200 Principles of Neuroscience John Stead Announcements Office hours moved to after class on Friday (11.30-12.30), this week only Online quizzes have been fixed (hopefully) Note that they appear in numerical order, not chronological order The impact of copyright on this course http://thebuglepodcast.com/?p=355 Lecture 1 Introducing the nervous system Reading Purves et al. Chapter 1 Lecture 1 outline What is Neuroscience? Cells of the nervous system Neuron structure Neuron function Types of glia Evolution of nervous systems Brain size Organization of the human nervous system Studying the brain Histology Determining neuronal circuits Functional analysis of neural systems What is Neuroscience? Relatively new field Term first introduced in the 1960s, when many different sciences converged with the goal of understanding the biological basis of behaviour and cognition. Included: Neuroanatomists (gross and microscopic structure) Neurochemists Neurophysiologists (bioelectrical properties) Psychologists (organization and neural substrates of behaviour and cognition) Rapidly expanding field SfN’s first meeting in 1971: 1100 delegates SfN’s 41th meeting in 2011: 32357 delegates Clinical neuroscience Useful to understand differentiation between clinical areas Neurology Study of the diseases of the brain Neurosurgery Study of the surgical treatment of neurological disease Psychiatry Study of behavioural, emotional, and mental diseases Clinical psychologists share similar objectives, but are not licensed physicians (and generally cannot perscribe pharmaceuticals) Neuroscience involves learning a language Names of components of the nervous system typically reflect location, and appearance Location Dorsal, ventral Anterior (rostral), posterior (caudal) Medial, lateral E.g. ventromedial nucleus of the hypothalamus Appearance Hippocampus (like a sea horse; hippos is Greek for horse) Amygdala (like an almond; amygdalē is Greek for almond) Cerebrum (main brain; cerebrum is Latin for brain) Cerebellum (little brain; cerebellum is Latin for little brain) Lecture 1 outline What is Neuroscience? Cells of the nervous system Neuron structure Neuron function Types of glia Evolution of nervous systems Brain size Organization of the human nervous system Studying the brain Histology Determining neuronal circuits Functional analysis of neural systems Cells of the nervous system Main cell types that define the structure and function of the nervous system Neuron (Greek for ‘nerve’) Glia (Greek for ‘glue’) Human brain has ~100 billion neurons ~300 billion glia ~100,000 billion synapses Internal structures of a cell Neuroscience5e-Fig-01.03-1R.jpg Neurons: Various classifications possible Function Sensory, motor, interneuron Location Spinal, cortical, hippocampal, cerebellar…. Identity of neurotransmitter Glutamatergic, collinergic… Shape Pyramidal neuron (triangular cell body) Granule cell (extremely small neurons, look like granules) Mitral neuron (large and triangular, shaped like a mitre http://ars.sciencedirect.com/content/image/1-s2.0-S0194599800701616-gr5.jpg Massive diversity of neuronal structure Neuron structure: synapses Typically a single axon, usually branched (signal output) Multiple dendrites extend from cell body (signal input) Dendrites may have spines Neurons communicate through synapses Can be between 1 and 100,000 synapses per neuron Most synapses are found in a tangle of dendrites, axons, and glial cell processes together referred to as the neuropil Most synapses are axodendritic or axosomatic Other types exist, but are rare Somasomatic, dendrodendritic, axoaxonic If found in somatic, motor or autonomic nervous systems (PNS), synapses referred to as junctions Synaptic signaling can be functionally fast or slow Fast or slow refers to the speed of response, not neurotransmitter release/movement Fast neurotransmission Post-synaptic response has rapid onset and ending Generally mediate by animo acid neurotransmitters Excitatory (glutamate or aspartate) Inhibitory (GABA or glycine) Neurotransmitters signal change in postsynaptic membrane permeability Hyperpolarization or depolarization Synaptic signaling can be functionally fast or slow Slow neurotransmission Post-synaptic response is slower and lasts longer Generally mediate by monoamine and neuropeptide neurotransmitters Monoamines include dopamine, norepinephrine, serotonin Neuropeptides include beta-endorphin, somatostatin, vasopressin, corticotrophin-releasing hormone Neurotransmitters typically bind metabotropic receptors, signaling changes to proteins within post-synaptic cell Fast and slow transmission can occur at the same synapse, and neurons can produce and release more than one neurotransmitter Neuronal function The function of neurons can be conceptualized and studied at many different levels: Molecular Cellular Systems Behavioural Molecular functions Studies focus on the actions and interactions of molecules within neurons Regulation of gene transcription and translation Post-translational regulation of proteins Synthesis, storage, release and detection of neurotransmitters Intracellular signaling cascades initiated by neurotransmission Cellular functions Two main categories of intercellular interactions Synaptic communication between neurons Non-synaptic interactions between neurons and glia Interactions between neurons can be classified into one of three patterns Long hierarchical neuronal connections Highly sequential transmission of information, typical of primary sensory and motor systems Cellular functions Interactions between neurons can be classified into one of three patterns Local circuit neurons Connections mainly with neurons very nearby Neurons often have small cell bodies (no need to sustain long projections) Typically involve interneurons which amplify or limit transmission in nearby neurons, often without actually generating action potentials (due to small axon) Single-source divergent circuitry Less common, but characteristic of some neurons in hypothalamus, pons, and medulla One neuron innervates many target cells in distant brain regions modulating synaptic activity in several regions Systems-level functions Individual systems are composed of a generally sequential series of interneuronal connections If any link in the chain is missing, function of the entire system is compromised or eliminated Some systems detect and integrate sensory information Hearing, seeing, feeling tasting, balancing Some systems initiate and generate responses Motor responses (ie: skeletal movement) Visceral responses (regulate body temperature, appetite) Behavioural functions Behavioural research focuses on interactions between individuals, and between individuals and their environment Examination of interactions between systems i.e. integration of sensory input with specific motor outputs Examination of higher mental activity i.e. memory, speech, conciousness Lecture 1 outline What is Neuroscience? Cells of the nervous system Neuron structure Neuron function Types of glia Evolution of nervous systems Brain size Organization of the human nervous system Studying the brain Histology Determining neuronal circuits Functional analysis of neural systems Glia While neurons are all specialized for electrical signaling (sometimes over long distances), glia have many functions Do not participate directly in electrical signaling Three main categories of differentiated glial cell Astrocyte Oligodendrocyte Microglia Two additional categories of glial precursor Glial stem cell Oligodendrocyte precursor Astrocytes Literally ‘star-like cells’ Lots of local projections Located in brain and spinal cord Function to maintain appropriate chemical environment for neuronal signaling Glial stem cells classes as astrocytes A subset of astrocytes remain as stem cells, and are able to differentiate into any cell type within the CNS (neuron or glia) Mainly located near ventricles (subventricular zone; SVZ) Oligodendrocytes Generate myelin in CNS around some (not all) axons In PNS, myelin is generated by Schwann cells Oligodendroglial precursors These glial stem cells can replicate and generate multiple cell types (even neuron in some conditions) Unlike SVZ astrocytes, mainly produce oligodendrocytes cells Microglial cells Mainly derived from hematopoetic stem cells in bone marrow (which also give rise to all blood cells) from which they travel to brain Similar to macrophages in other tissues (can be classified as a form of macrophage) Remove cell debris following injury or apoptosis Release cytokines that modulate inflammation, immune activation Lecture 1 outline What is Neuroscience? Cells of the nervous system Neuron structure Neuron function Types of glia Evolution of nervous systems Brain size Organization of the human nervous system Studying the brain Histology Determining neuronal circuits Functional analysis of neural systems Comparative neuronal organization Neurons are basically the same in all animals Only the arrangement of those neurons changes dramatically between species Many insights can therefore be gained from simple organisms Sponges (Porifera phylum) Simplest multicellular animals Immobile, but smooth muscle sphincters control movement of water through body walls Smooth muscle cells contain sensors that detect stretch, chemicals, etc No separation between sensory cells and effector cells Animals lack any nervous system (no neurons) Hydra (Cnidaria phylum) Cnidaria phylum includes jellyfish, coral, anenomae Able to move and actively feed Body wall has 3 layers Ectoderm Mesoderm Endoderm First organisms to evolve neurons, most likely in the form of sensory neurons Some Cnidaria have only sensory neurons, other have both sensory and motor Sensory neurons derived from ectoderm Hydra (Cnidaria phylum) Sensory neurons are bipolar One end projects towards stimulus, other end to effector Information flows from input to output Information flow already unidirectional in the most primitive neurons http://dev.biologists.org/content/138/8/1447/F4.expansion.html Hydra (Cnidaria phylum) Why evolve sensory neurons? Increased sensitivity Due to cellular specialization for sensation Faster effector cell responses Due to cellular specialization for effectors Stronger behavioural responses One sensory neuron can signal to multiple effectors Enables input from different sensory modalities One effector can receive input from many sensory neurons, each detecting a different stimulus Hydra (Cnidaria phylum) Hydra also evolved motor neurons Receive inputs from sensory neurons Send outputs to effector cells Nervous system composed of 2 layers: sensory and motor Enables signal divergence and convergence Divergence Signal amplification One sensory neuron innervates many motor neurons, innervating many more effector cells Response could be mediated by many different parts of body Convergence Signals from many widespread sensory neurons can converge on a single motor neuron Hydra (Cnidaria phylum) Neurons have diffuse distribution called a nerve net More concentrated in regions with specialized functions, such as mouth and tentacles Aggregation of neurons is termed centralization http://www.tutorvista.com/content/biology/biology-ii/control-and-coordination/control-and-coordination-animals.php Flatworms (Phylum Platyhelminthes) Bilaterally symmetrical predators with anterior-posterior and dorsal-ventral axes http://bioweb.uwlax.edu/bio203/s2008/geske_rich/images/OrgWebTaeniaPile3.JPG Flatworms (Phylum Platyhelminthes) Bilaterally symmetrical predators with anterior-posterior and dorsal-ventral axes Many neurons cluster in ganglions, connected by nerve cords Centralization increases speed and efficiency of communication between neurons Cephalic ganglia is largest ganglion, with sensory input from front of flatworm http://media-3.web.britannica.com/eb-media/36/73336-004-54B86827.jpg Flatworms (Phylum Platyhelminthes) Simplest animals to have a third type of neuron: interneurons Intermediate between sensory and motor neurons Interneurons enable a range of new functions Allow greater convergence and divergence, therefore response complexity Act as switches in neural networks, turning sensory-motor connections on or off Act as pattern detectors or pattern generators i.e. transmit signal if sensory input from cells A and B but not C Act as pacemakers Some interneurons can generate intrinsic rhythms Myotatic reflex in humans While the human brain is vastly complex, some circuits are similar to those in simple organisms Myotatic reflex (knee-jerk) has similar complexity to flatworm (sensory, motor and interneuron components) Neuronal activity in the myotatic reflex Extracellular recordings optimally suited to record the frequency of action potentials Phyla Annelida and arthropoda Annelids segmented worms Arthropods spiders, centipedes, crustacians, insects Phyla Annelida and arthropoda Body segments are repeated along the anterior-posterior axis Enables greater complexity of body plans, without massive increases in genetic complexity Individual segments of body and associated nervous system van diverge in structure and function All complex invertebrates have ventral nerve cord, pair of ganglia in each segment, and longitudinal axon bundles between ganglia http://www.earthlife.net/insects/images/anatomy/nerves.gif Phylum Chordata Includes all vertebrates, Tunicates (e.g. sea squirts) and Cephalochordata (lancelets) http://science.kennesaw.edu/~jdirnber/Bio2108/Lecture/LecBiodiversity/34_05Tunicate-L.jpg http://comenius.susqu.edu/biol/202/animals/deuterostomes/cephalochordata/uwinnipeg-Lancelet.jpg IMG_5008.JPG Phylum Chordata Chordates share Notocord (dorsal cartilaginous rod) Replaced in vertebrates by spinal column, with cartilage remaining only as intervertebral discs Hollow dorsal nerve cord above notocord Evolves to spinal cord and brain in more advanced members of phylum http://www.biolsci.org/ms/getimage.php?name=ijbsv02p0149g01.jpg&type=thumb Phylum Chordata Characteristics of chordate nervous system Bilaterally symmetrical and segmented Crossed organization: Each hemisphere receives information from and controls the opposite side of the body Spinal cord is dorsal (at the back) to the heart and gut Lecture 1 outline What is Neuroscience? Cells of the nervous system Neuron structure Neuron function Types of glia Evolution of nervous systems Brain size Organization of the human nervous system Studying the brain Histology Determining neuronal circuits Functional analysis of neural systems Big chordate brains allow complex behaviour Behavioral complexity correlates with the evolution of cerebral hemispheres and cerebellum Increased size and folding (to fit more tissue into the skull) Mouse Human Human evolution: Primates Humans are members of the primate order, and ape suborder ~275 primates species, which share a number of features Excellent color vision Eyes in front of face: enhance depth perception Females: Usually only one infant per pregnancy; infants require more care Larger brains Human evolution: Apes Among the apes, we are most closely related to the chimpanzee Diverged ~5-10 million years ago We are the only surviving member of the hominid family. All hominids were thought to walk upright Family had 4 known members, only 1 remains H. habilis, H. erectus, H. neanderthalensis and H. sapiens Human evolution: Homo sapiens Humans (homo sapiens) probably originated in Africa ~200,000 years ago Co-existed with other hominids, including Neanderthals Archeological and genetic data point to an African origin for modern humans Human culture is more than our biology Culture: Learned behaviors that are passed on from one generation to the next through teaching and learning Brain is highly flexible: We perform many tasks today that our brains were not originally selected for early during evolution Example: programming computers Our biology thus provides the capacity for complex thought/tasks, while we learn these tasks through our cultural environment Environment thus plays a major role in brain development… Comparison of brain sizes Jerison (1973): Principle of Proper Mass Species exhibiting more complex behaviors will possess relatively larger brains Big animals have big brains Comparing species, doubling the body weight will by accompanied by a 2/3 increase in brain weight This allows brain sizes to be compared, while correcting for total body size Animals with unusually large brains display unusually complex behaviours Including humans and dolphins Evolution of the human brain Why did big brains evolve? Possible answer… Some foods are easy to find, so do not require complex behaviours and brains Apes that eat fruit (including humans) have unusually large brains, as finding fruit requires Knowledge of which trees contain fruit Ability to navigate trees to find fruit Understanding the seasonal nature of fruit availability, and when fruit is ripe to eat Success in competition for the limited amount of available fruit Extensive learning skills, so we can learn when/where to find fruit Once we adapted to eating this complex food source, it was probably easy to adapt to other food sources (including hunting and scavenging) Hunting monkeys in packs has been observed in some chimpanzee tribes Big brains cause problems… Size is just part of the story… Genetically, humans and chimpanzees are are 98.7% identical Gene function relates both to the sequence of the gene, and what it does (how much the gene is expressed) Microarrays are tools for simultaneously analyzing expression levels of thousands of genes Divergence in gene expression profiles between humans and chimps was far greater for brain than for other tissues See Enard et al. (2002) Intra- and interspecific variation in primate gene expression patterns. Science. 2002 Apr 12;296(5566):340-3. Lecture 1 outline What is Neuroscience? Cells of the nervous system Neuron structure Neuron function Types of glia Evolution of nervous systems Brain size Organization of the human nervous system Studying the brain Histology Determining neuronal circuits Functional analysis of neural systems Organization of the human nervous system Broadly divided into CNS and PNS Not a perfect subdivision as nerves carry axons from cell bodies in both CNS and peripheral ganglia Lecture 1 outline What is Neuroscience? Cells of the nervous system Neuron structure Neuron function Types of glia Evolution of nervous systems Brain size Organization of the human nervous system Studying the brain Histology Determining neuronal circuits Functional analysis of neural systems Organization of the human nervous system Broadly divided into CNS and PNS Not a perfect subdivision as nerves carry axons from cell bodies in both CNS and peripheral ganglia Organization of the human nervous system Neuroscience5e-Fig-01.10-2R.jpg Structural organization of the brain Neurons in the CNS are organized in one of two general manners layers or nuclei Layers of neurons Cerebral cortex Cerebellum Hippocampal formation Olfactory bulb Clusters of relatively compact neurons within ‘nuclei’, sharing similar connections and functions (called ganglia if in PNS) Nucleus accumbens Caudate nucleus Substantia nigra Raphe nuclei Structural organization of the brain Neurons in the CNS are organized in one of two general manners layers or nuclei http://ars.els-cdn.com/content/image/1-s2.0-S0306452204003185-gr1.jpg Organization of long-distance connections Tract Bundle of axons running between structures in the CNS Nerve Bundle of axons located in the PNS Analogous to tract Commissures Tract that crosses the midline of the brain Axon tracts and commissures are typically rich in myelin Higher lipid content looks whiter, hence name ‘white matter’ Gray matter is cell bodies and neuropil Lecture 1 outline What is Neuroscience? Cells of the nervous system Neuron structure Neuron function Types of glia Evolution of nervous systems Brain size Organization of the human nervous system Studying the brain Histology Determining neuronal circuits Functional analysis of neural systems Histology Histology is the study of tissue, sectioned as a thin slice and stained to reveal specific structures Histological procedures are broadly divided into four stages Fix the brain Slice the brain Stain the slice Examine the slice Fixing the brain First, the whole animal is typically perfused Entire blood supply replaced with saline or a fixative while the animal is terminally anaesthetized Removal of blood leads to better histological results Then, the brain is removed and fixed The brain is already a very delicate gelatinous structure On removal, it rapidly turns to mush due to autolysis (self-digestion) Fixation prevents autolysis, hardens the brain, and protects the brain from mould and bacteria Paraformaldehyde is a common fixative Sectioning the brain A brain section is a thin slice of the brain (typically 5-200 m thick) Sectioning typically done with a microtome, vibratome, or cryostat (microtome adapted for frozen tissues) Composed of a blade, a platform to hold the brain, and a mechanism to move the knife forward to take the next in a series of sections Section is mounted onto glass slide for staining Staining the brain Lots of different stains are available Depends on what you want to visualize Just a few examples… Studying cell structure Morphology of individual neurons first discerned by Camillo Golgi 1873 developed silver stain method http://jp.physoc.org/content/588/19/3639/F1.large.jpg Studying cell structure Golgi staining stains a small number of neurons completely Visualize cell body, axon, dendrites, dendritic spines Allowed identification of points at which neurons contacted each other Named synapses by Sherrington in 1897 Studying cell structure Golgi method involves soaking brain alternately in potassium dichromate and silver nitrate solutions Staining takes weeks ~1% of neurons become filled with dense precipitate Nobody knows how or why it works Alternatives to Golgi Markers can today be directly injected into single neurons via micropipettes Can be combined with electrophysiology to simultaneously examine structure and function Can also target specific neurons Staining for specific molecules Nissl stain Detects nucleic acids in ribosomes - stains cell bodies by targeting rough endoplasmic reticulum A range of different dyes perform similar functions, and are all considered to be Nissl stains, including Methylene blue Cresyl violet Thionine Toluidin blue O Allow visualization of clusters of cell bodies within nuclei Nissl stain http://brainmaps.org/ Weil stain Stains myelinated axons (others are available) In situ hybridization In situ hybridization DNA and RNA molecules bind to other DNA or RNA molecules with great specificity If you label a DNA molecule (eg: with radioactive isotope) it will bind only to DNA with a complementary sequence Allows you to detect mRNA produced from a specific gene RNA probe hybridizes to mRNA from the gene in situ (exactly in place where it is located) Analysis of neural activity by ISH c-fos is a transcription factor that is rapidly expressed after neuronal activation c-fos ISH identifies neurons that were active immediately prior to euthanasia Immunohistochemistry (immunocytochemistry) Use labeled antibodies to detect specific proteins Glial fibrillary acid protein (GFAP) in blue Microtubule-associated protein-2 (MAP2) in green Immuno vs. in situ In situ hybridization tells you where a gene is being expressed Which cells are producing the protein? Immunohistochemistry tells you where the mature protein is located Probably indicates where the protein acts Can be a long way from the site of synthesis Some signaling molecules active in the brain are mainly synthesized in the periphery (ie: insulin) These two techniques can therefore tell you very different things, as you are identifying either proteins or mRNA molecules Examining the slice Various different types of microscope, depending on what you want to visualize Light microscope Good for <x1500 magnification Fluorescence microscope Similar to light microscope, but uses different wavelengths of light allowing visualization of different colours of label Electron microscope Allows magnification of <x2,000,000 Confocal laser scanning microscope Scans tissue with a laser at various depths, allowing 3D reconstructions from within thick slices of tissue Allen Brain Atlas http://www.brain-map.org The Allen Brain Atlas (ABA) is an interactive, genome-wide image database of gene expression in the mouse brain. A combination of RNA in situ hybridization data, detailed Reference Atlases and informatics analysis tools are integrated to provide a searchable digital atlas of gene expression Shows Drd2 expression primarily in basal ganglia Neuroscience Information Framework (aside) The Neuroscience Information Framework is a central repository of online neuroscience resources including data, materials and tools http://neuinfo.org/ New resource, under development Explore the navigator Not that user-friendly, but a site to watch Affiliated with the Society for Neuroscience www.sfn.org Lots of information and tools also available on this site Neuroscience Information Framework (aside) Screen Shot 2012-07-24 at 12.41.15 PM.png Neuroscience Information Framework (aside) Screen Shot 2012-07-24 at 12.43.13 PM.png Neuroscience Information Framework (aside) Screen Shot 2012-07-24 at 12.44.11 PM.png Lecture 1 outline What is Neuroscience? Cells of the nervous system Neuron structure Neuron function Types of glia Evolution of nervous systems Brain size Organization of the human nervous system Studying the brain Histology Determining neuronal circuits Functional analysis of neural systems Determining neuronal circuits Early studies involved lesion of specific brain regions, nerve, or tracts in animals and determining where function was lost Limited tracing of individual axon paths first achieved in 1885 (Marchi & Algeri) Severing individual nerves causes distal portions of axons to degenerate Termed Wallerian degradation Myelin sheaths also degenerate Visible as black particles on light background http://neuropathology-web.org/chapter12/images12/12-3d.jpg Neural tracing methods Neurons can have very long projections, both to and from the cell body Generally cannot simply stain a cell and see where the projection goes, as there are millions of similar projections within the brain slice Anterograde labeling traces the path of projections leaving a cell (the axon) PHA-L is a protein (from kidney beans) which is taken up by neurons and transported down axons A specific brain region in injected with PHA-L, it is taken up into neurons are transported along axons Brain is sliced and stained with antibodies against PHA-L Anterograde labeling Injection of dye into dorsal root ganglion reveals axons from dorsal root ganglion into spinal cord Neuroscience5e-Fig-A04-0.jpg Neuroscience5e-Fig-09.01-2R.jpg Neural tracing methods Retrograde labeling method Used to identify which regions send axons to a specific region of interest Fluorogold injected into the region of interest Taken up by axon terminals and transported back to the cell bodies and dendrites When visualized under fluorescent conditions, fluorogold releases light so can be easily visualized Neural tracing methods Transneuronal tracing method Inject pseudorabies virus Infects neurons, and spreads to other neurons in contact with them The longer the experiment runs, the more neurons are infected Identify infected sites by immunohistochemistry for viral proteins Functional analysis of neural systems Neural systems can be understood, in part, through studies that focus on function Lesion studies To understand normal function of a brain region, we can destroy it, and see the effects Ablation = removal of part of the brain Lesion = damage, destroy, cut part of the brain Example Some regions of the human parietal lobe cortex are involved in spatial perception and memory Lesions impair the ability to draw and follow maps, and remember the location of objects that have just seen Also problems with mental arithmetic Mechanical brain lesion/ablation Aspirate surface brain tissue Radio frequency lesion Wire inserted into the brain, insulated except for the tip Ensures only a small localized region at the tip is damaged AC current in the wire produces high radio-frequency currents that heat the brain and cause damage Chemical brain lesions More selective than mechanical lesions Only kills neurons that have specific receptors Does not destroy other neurons/glia or neuronal projections in the affected region Common approaches: Excitotoxic lesions with kainic acid This amino acid is injected into the brain, and stimulates neurons through glutamate receptors Stimulation is so extreme that the excessively activated neurons die 6-hydroxydopamine (6-HD) lesions 6-HD is similar to dopamine and norepinephrine Specifically taken up by neurons that detect these neurotransmitters, and destroys them Brain lesions: Pros and cons Size and location of lesions cannot be perfectly controlled Impossible to perfectly replicate a lesion Interpretation is complex Each brain region contributes to a range of functions Each functional output of the brain is the product of many different brain regions Nevertheless, it remains one of the simplest, oldest, most useful, and most powerful tools available Question: How to you consistently target the correct brain region for ablation, or any other physical manipulation? Stereotaxic surgery Allows electrodes or cannulae (tubes for injection) to be precisely located within the brain Stereotaxic apparatus has a head holder, a holder for the electrode/cannula, and a device that positions the electrode holder precisely in three dimensions Stereotaxic surgery Stereotaxic atlas Describes location of brain structures in three dimensions Species-specific Co-ordinates are relative to a location termed Bregma Soft spot towards the front of the head of a newborn Anterior fontanelle Point at which the two frontal and two parietal bones meet Stereotaxic surgery Stereotaxic methods need to be verified after the experiment is completed Typically, you remove, slice and stain the brain to identify the precise location for your injection or lesion http___www Functional analysis of neural systems Electrophysiology Electrical recordings from live cells E.g. single-cell recordings with microelectrodes Neuroscience5e-Fig-02.02-1R.jpg
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I'm using total in bank . this bucket . so . and when they met you I was saying so for my and also for the movie we saw Dallas right side for what he can expect to cover up in the course of the job and we Two and his allies slide . Identity supports June lecture so that you can see are the top local how long ago properly before you get adapt to life . the brief discussion What is Neuroscience been told about cells in those system but single stall and mules little course of the cells onto the cells of New scientist and works on most . I miss him so I took him Evolution on including those system . we can be . the best majority of this course would in part to humans . I made a Nobody runs dolphins well guide . Seth was dismissive neuronal select . we got to humans . Rick actually learned some very moist basic principles of neural function or so ago with the simplest . they go through repeated our realization though system which is basically digital cinema system performance system . I'm very subdivision to all the beach . I'm and . that's why and I woke or will the about techniques . now that we Should Brendan be teaching this the concerns of bread in this course on the just going with this blog to that . the techniques the house in climate next semester on flight day sessions wihtin grad students to come in until about techniques however it's probably useful to does have some idea . so when the Clinton techniques were look before we get to have reinvested in the course material flow on it once we got into some point but it might be to die in my help . aggression those new songs true Immobile Neuroscience Israel . I am get some things . going use on touch on the slide . guess . cells . and it would resort Golgi . start . . . so clinical . 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I'm conditions and a commission on the wind but nothing that is good . mostly this artist's that's what I've mammalian deduced is up move on local and state is no from the Alps and the dime the brain is the physical blamed on every poem about consciousness that level of behavior can be supplied and dry the physical of the close to some metaphysical so when that doesn't exclusive perspective of of news long-distance only one Lecture gut . evolution doesn't foods we have a new an awesome is that Neurochemists Neurophysiologists situation of the movie . I'm borrowing magical Paul to all the move system . I am for cohesive dough to focus on localization and you'll Soft filter through of all ago on the code mission without too much focus on the one magical . sensation the Lord in all the fruit of new songs on the themselves perfused relatively recently in Sindh in terms of his ships arrived in the conflict . it's very Reference binding the first arm Society for Neuroscience general meeting Post sought to Neuroscience what in the consent to warm the house lobe fun Friday about eleven hundred delegates . the most recent sort neuroscience meeting in quotes in twenty eleven . you know Washington . I'm bad with those two thousand people attend the conference . it's nothing image last one is to say in the folds all and of the census is just too much going on of the school bus . we wanted it . I am . this course will still return to mend though the underlying Belgium close to one of what how things work . I've that close to the Belgian how things go work region more Clinical sought it . sadly it's food for the chic welcomed to me . if all you will be Difference most was when Various different compartments there is different . did some disciplines within the Clinical news on TV out the new or just to study diseases all the blame . not a neurosurgeon system to study how you treat your local disease for surgical techniques . yes the entrance . to study be able emotional I'm to mention diseases and the room anyway it's good fun going to sue him in a ways to Clinical for Golgi human form to Michael replied not the best in situ in many ways I robot and blueberries difference this is the fuel chemicals like or just in Human position where you can squad medication to types . the cake but cautious can subscribe occasion . but what causes to me speaking . all demanded to speak input called a small licensed physicians . 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Kevin can cause a lot next to . no you situ will become more more menu it says the help you on the sounds of the field on found reading a film simple . in the fields . from those system post office with a big layers of make sure we can meet the goal mould this next week to go a whole week the Bush to imagine . I'm depressed movement system of Allah and name to compile body from weather . submissions ventral and should response to medulla buses route . the assignment should place on the appearance will be up howling . squinty appearance of the oven only knew not to miss two new code of poll blade duplex and the Franco done time and said I've read countless in the campus was named all for the sequel because the bill because of that aisle . it bothered become the tell all AC or spite . 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I'm than partially to him also to axis allies to take . so if you like them is a classic encounter will know when fruit tree should it be did come to classes can be much better . so in the mid to late awaken all squares chins and and when somebody some interaction . but if you do led to the top and complex AP wrapping to come up . you didn't miss a lecture . hopefully next to a winnable use when they classes with Andrew in see the King . I'm from the DRG Shi dae disorder . it says the system . and we made them so cannot neurons and glia in this close block was unable cells you go for double cells involved in the blade basket . the blood system . we can't afford Why merrily on mulesing glia knew him being Greek who grieve with the move . I'm glia being Greek community was believed that this cells was issued to those holding together . the boy . I'm big on all these things progress the bow hundred in the new rules in the best way to around three times as many glia . that's with those better you'll see some books inside the studio hundred billion neurons of a thousand billion glia bulb outs . Boca where we all . um it was about a hundred thousand billion . find out since a mission of writing which is it does politics Playboy . we don't know very much about how the combined works . the so called Moon elements to the in flight ever cell . just a couple of house more relevant to us in new loans when of them what to call under which in their hearts going over them with how Alfred cell . it's is what generates ATP which most of which is basically the of is accessible chemical energy to Not AGP then he continued . Pudge was a CPI with an adjacent to the Impala . a source of of course this is inside cells . us which is located in the Sudbury right . Lucas is well contained you did nine economists in the story gene transcription takes place . the onto generate a message on my humans don't eyes and transported to the new to us into thought about them web the master are my strong right through into proteins using why the science center as well sense of proportion fleet in the Saw deposit . I was those blokes limbs all set on through these B school these men but if synapses in front is so good to of the end of the reticulum that you did read some stuff them in part to secure and than those offices put to death . and it was were taken . I'm a good rapport this was a really important for new loans because the good job for us . Israel could use medical into the stickles to than shit . I makes cords of areas blood cells . so in some evolution of the newest murals . it's a good rapport on its butt . great food the school's new posts place sand them down the axle to them who sold breakthrough in lease out the sauna . okay . I was of course not done joints . I'm single I told become cell phone . I'm from New England boss George O'Neil . . this was in pasta for neurons . because Mills mission exclusive doubt those in collections will be systems . as coaches by so function you go you must neural into good sense to me root you go you into new looks . where are those neural than the well look at ofNeural stabbed cell but is instead I in the choral section of campus and set about our fruit how I'm Spinal motion your result . lol umm cabling tulips . when you turn nobody really good reason to seek new onto his good play would go um Colin the gene Neurochemists asshole calling . the cost What placed on Shape when out the gravel neurons which are all basically the sentiments from shapes what coach Hugo granule cells . cells luscious more listed on cells and falling into buying the team can continue on . I'm doing the cool granule says because on the next Typically grains of sand the release will . when from the twenty oh of which are lives trying to neural to look at the end of time to brood The Boldin on a sense of fun of metal . I miss the example above of of March will sell much of neural . trying to shut it . I'm become stronger side and many more tools to pick of the shape of a blade so of of bishops that is trying to understand speed Dome but us away from can . a source and I took it . the amount that Douglass been structure . mammalian will the structure is blue will tend to do this in the same clues us was the function likely cause two hundred and on which when the boat . a few minutes I decided the pecans cell in June the allow function . I looked and listened directs with the function is was thinking my guest . getting function into it . . the . said oh . so should move to the bathroom shut does great . . potentially . Ross . . Google are words that goes well in the one being the rule will be included into being . do note though that the overwhelmed with Malcolm says . Moshe from the group was Mr Fulton includes reading together . I didn't one of my compared that with for job all day but no bribe the cell where it is close to the human and a output new addition century a relational which you could truly isn't going be doing very much integration tool . whether Types and the highlight will go this lineup is the retinal on the corn cell . I'm going to lose . unusual embedded in the in room that in the summer we met up for a while of that does very few emigrants cells in humans . I'm going cells of those where you situ how bright the actual slide not to connections . nobody move will know this or that single be precisely membrane of posts about to membrane . you get when it's neurotransmitter from the productive members Example post I want to be assigned a sick member . 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I knew how to be a lesson and enjoys then form cell border which is your temple . not communication to new successes and have small stuff in there long enough hundred of the fun acids in your eyes to a previous life . miss him so that I found a cut tangle had enjoyed tonight's Aldrin on the of those poses from cells . the collection is referred to as the new appeal . it's only really playful . that move Lots head of connections the dead between cells of references neuropil Most setups to come across awesome injury or axosomatic into motel when Tommy's just name it it's we an axle and enjoy . it's when I'm outsold I'm the slow mo the cell blog . true for fall with the most comatose on ABC's unity of the tribes lived so bored so bored Somasomatic dendrodendritic Max likes to lick his Among from of what you get with those all produced Iraq . I'm nucleus of does it in distinction to mention between saw enough to be dead in the session nervous system . 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I'm talking about bread without bending . limited effect . so I could sell oil move demanded packs within the posts on optics cell . so Fast neurotransmission isn't so new covers from brain to release him neurotransmitter which have been dying to meet up . I'd to underline shuttle . well I tell to open . closing a depolarization boy Hyperpolarization . the of posts are not sick cell member gave one to him that were read to subside after communication gospel when Knowledge told my in the cold takes all generating action potentials . fruit . do I will speak . watch I would because it is rapid or slow sometimes of eleven of the chattel Iraq was pulled against imposing a jury blue so seemingly due to the right amino on acid transmit if that adds to its mitral sparkly double will likely to be the menu includes letting go Most Coleman . you know most all went into the tree neurotransmitter . I'm so so that blue so the momentum is singing is me with his Jenny through Hyperpolarization all the politicians premature media . I told people elation . the posts I've two member . it's a good slow neurotransmission Sanchez missions continue to miss at lunch on receptors were brought metabotropic receptors became a culture percent to million in public metabotropic receptors . we're neurotransmitter the return of sensory friendly Chinese . maybe some tips to look after a teammate with the you it's me get a change of gene transcription . in the post and saw topicsCells Marcos . I'm so change the can was open changing transcription is gonna take on . the receipts . we neurotransmitter the brain how does closely with signaling cascade for anything to do just get it all in the giving the Oklahoma gene transcription . Babel a clothesline . so when he walked and top with a slowing rate of one of us would think about . is the define your terms mission to go changes though . the reactive if you loose balls to the sea . all the system . all the new . so the bottom of the moon . men like is the poll is for sample . it was also false notion of motion or less what you call it . it is issued and talking back to baking dish Cuomo and regions all the plane all of these laws had actually been making them making melodies in the Blaine more likely to be calm . in response to a false neurotransmission . so you know that getting to go ladies . they are up . I was like set points in the blanks of the human brain What is cool . and if we How playbook release in March and bundled in this way . wherefore girls certainly in much is going over this book . so the brain the Lion region with my which formed differently . Sioux Falls to Shows mission when she comes in . ok so these This lunar transmission gyrus begins mediate to blight Medal a means in your passwords . no I mean think of the brain norepinephrine serotonin . Functional had soared from the match but although peer endorphin from us that it was impression of sewage Caudate turf and release include month . could . . she's What newest record signalling um . I was sitting in his mediate . very little swelling saw action potentials in a couple weeks . in . so when you think about . blue dogs didn't come to this is function . how does one think about function to in for the complacent levels Unlike molecular function and family functions Systems function of the table function . so evolution to level when Coke Bell will change is essential when within a single Neural . I'm Regulation to transcription translation Able greetings all pull all Molecular functions idiotic of the season is slower neurotransmitter that situation metabotropic said to which initiates a single Cascade which was also the poem attendance at the example that would be a molecular function Molecular spawns within the cell . 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I was sent of those mules putting extreme you will accept if I'm reading what times that is because ago I told Doug all boys down in the lot . lay down one leg form with a must watch . so can I found were really long you'll connections recorded on long no connection . then you can sometimes tell . how will always the connections of in your just looking Until blog . in a few how the issues at school . but a new in house support . Bendigo the fellows go how we generated will for proteins to sustain them on actual statements but to those who Knowledge cell bodie facilities . I'm central article to read when you shoot on some bonus for the in the local . Best connections primarily the Dubai and the sensory on the motor system . . so I'm rest numerical instructions . I wound some and made some pretty good and I was causes my own little been told for the base connections in the eleven signaling . between you . she mediate question . and I'm sitting in bed tonight changes and depolarization Hyperpolarization and I settle Jews who grew to changes in Walsall to get integrated turned in on them but until then we'll cell which results in Burmese unusual smell to it because integrated strange impression call from Embley in second cell . rather I were the how not to show . while . and then the baby growing in the global function is all Russian control . I joked ensure the functions propagate it signal the time the signal for long-distance son to do transmit signal for already . of course you need action central . but if you only need to transmit signal for sure the students . the new doneness so the how to how bout signal when you yourself will call him . in just how to say she pops it . Chas machine on inflation with them spend a quiet little current . 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I was CNS are made in the complex along with the slow neurotransmission most part the a couple minutes ago . cells to improve local borough of the rock a new cats in the in the blanks that the Rapidly juices on goods collection of neurons . but then most defensible generation seven hundred items releasing so tired and probably right . so how does one when the band spent the rapid use percent projections to let cool for the two places to L the bright . in his right boot and palm like to see . K I'm after we had a single source for evidence Sectioning Doolittle in collection and sanitation projects in guts and Information fell infinitely large region so super well and they hypothalamus palms on in the medulla eyes in the medulla whether Iraq when you can simply take it . so every rock in that were on wheels out the leftists who has switches that the Center for the um the more genetic transmission ghosts of the sensible . in more detail . mummy focus on . neurotransmitters . poker . systems with function . most of the routes . can you lend other level of the molecular changes which would render continuum of the level the Celts our winter auction which took neural function in terms of how in your an ox within a much most system . so system is composed all the classic connections often in Haiti sequential series is more newborn infants who systems . the story of point in the motor systems way get blown connection to another connection to another connection . like ascending or descending from the blame . the the blame on receives What transmits information to the board . the thing about the sequential connections that you did get warm it all all bought me one more link in China which is missing . then the whole system . cells to work . I'm in the swap and um birth I'm afraid of with workers to remove course . otters are going to this place to grab hold . book is but is powerless to to the of spinal injury . like a pig walking to bus point use the seventeenth one so yeah the complete voltage albeit climate system . because your house . I've been all over trying those know what for . look the mood rules conceived left with much about it it's optional for lower Homo one of the bucket . in guts in systems where you have a image sequential processing . it is true . I'm turns into how we be sense inflation is integrate it . I'm Paul says up to phylum of the boys structures So Philippa systems this is seeing feeling tasting Phylum thing you can focus on Systems level functions in that way . and so systems initiate Joseph S this is the most fences and the vessel was pulled to the social being . the blue book . sober mood to is false is good . ot station below tree with a bow those close to the result and I Regulation beating in the hall so busy construction in your by its war do with our jaws what to do it herself with squeezing her not longer . I went to invest in your function comes all the federal Behavioral commander research groups is all I'm entitled to an end and you will between interactions between an individual . I'm to the environs . I went to the approaches tend to look at how good can systems interact . so systems that whomever will tell the sense he says . we systems that will inevitably lead to system . in the benighted rope . but that perspective . you may look at how will a change through the eyes fitness which soon Broadly said to system is going to result in a stupid with dense with the best American bodies as well put in that fruit among system so Ruby who are reading and integrate seeing the movie from systems where the Java to move out how feminist leads to which local sometimes in without stimulus for me it's still well to just do what you're thinking about men were systems will lose consciousness soul old did it to speech or the dog out . um dude you got integration between system . look at . commencing you somewhere in the course as mobile end of the . Wilson combat tequila or so . Sam . . we sat Types Lysol cause him to lead us . I'm . Seoul going to let . 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I'm supposed to Immunohistochemistry which is looking out Species blog site is so unfair is tissue . and then decide . is this the only venue several different approaches of Oregon to live because it's a cell and into glial so much . I'm what you'd into which our differentiated good sense . I have include precursors to say she stem cells grown between generating of the glial Types Island from some cases involved in generating neural its individual nerves . we of neural stem cells are we not only going to start pre Just glucosamine is looking a glial cell phone or in the second of can . so as to sites with photos outside because they look like Starbucks the garland projections all and place . the Scarlets our families destroy glial Chawla g milk like soft . that Reference objections to making room I would live cells and connecting with humans as well located in the brain and spent occur . I'm good many functions Montag an appropriate chemical environment for neuronal signaling . 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I and said this could rapid rewards you places you go the subventricular the and as he's dead on Greece's wage its new the new Genesis killing supply Union you'll still find two groups . I was of the the subgranular zone of dentate gyrus in the hippocampal us which is generation you know and um . we wish to become integrated into the mature in campus . I sold the deal stem cells . though often preferably the protest . Show is truly are the solution of the subgranular zone . as Jews and . Jared's . we'll come back to Plato called for direct it . dentate gyrus neighbors beloved campus . could . so when you do . the amendment . in the CNS around so much when the likes of the summer cycling but and relax on our is my life is in session of system is to do it . actually those are abandoned and then go on the transmission Information wrestlers routine funny . neurons than that to the altar and the monument . fishing out crosses locker whales out songs insulated them a lecture . shouldn't occur in the new opera nervous system and of course a shrill cells to evolve not course . there stem cells solar could see our zeal to the new glial precursor . generate and ofNeural only good thoughts . if approved debate and generate a big tools of heart as well . Russo Generate Neural scored the . that's something that you can cut fluency in the allowed under the conditions and get them to . sometimes Generate continuous whether that she will generate new new loans can be little in the living writing is so mum out of contention . learn to read American girl sense . the riots . and when the light that comes much of the matter . so generated . in the body stem cells located in the bottom Allen on the metal and soul into Clinical smoke Labour that I am blessed in that amount which is the basic all conveniences . ventral is to to Nobody did through to cope with any affection and two I if you today and the juice or fruit on the cell injury or apoptosis them to feel stem cells will come in and get We Oval but debris and reticulum Ch about awful close and Brian of Sharia Note two when the bottoms . indeed our little one will cells between the two and end of the move cells and use which is used . 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I need to do things right . right to sell between of it . during the month I will try though cell the word in the current NSL known before . I've said this space is low that you can really have to MIT's album you'll connection . because everyone really into its can happen the sign no connections one . so that been in the human world and decided to really why are some pretty course this the cake that questions about no interaction on your function . in these new better boy simple . it's . Coast but was too late for this and in the we used our more medals through Sam's . so made it aspect of neuroscience . most those the room and the young doctor Neuroscience . do we still it all on our movement below of what would can cost them out of mouse workers well Psychiatry main demolished . pretty cells made a Rattler . but with those who work Human implying situ I'm making friends is about how the human brain . but I sure could . so it was so crude Simplest audible . the emotion that I will a that those who Simplest mid December . so says it all . I'm . I motioned to move in the old one . I mean itself right around then this is the US including debt . his preferred to is peripheral . I will Looking out Sponges head . loosing them . Gladwell book mentions about dieting . I had a limited the adults of very high fences . Smooth muscle . the closer our movements and blue so through the border were also demand filter Alps . food stocks . 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I'm that affects the well . well . Laguna Phylum concious August need him identify them . which is a hunter . where animal the news in oceans . we can always tentacles into school to have whistles into hands . I'm . she spends most destroy him . I intend in the the sounds of the bomb see I could Visceral call to action movie realms deluge of ghosts and assaults a warm bed and out of hand . you did not do Bendis be down . Included sound of them . I will take about a summer garden gumball around to it . it is a will to some good mood complex . the units . data from can rest functions can just wipe nervous to go . so today was to move a little cloud to be freed . the school as the what you of someone than say ten years now probe led to this what we do need elements notices them . a recent weeks and then the Liverpool as and for me to them and them just like our this day . the arm . the first organisms through the meal . good eye and said oh hello there were spaces . haha sensory Neuroscience Muslims same image as a sense in you . bill but all specialized cells specialized to detect sense information . I'm been charmed knits that sensory inflation to a secondary cell . the singer cell being muscles . the good sense to move to either form be Ectoderm destroy the um of the nervous system is delightful detect them in in humans . . when the child to the ascension you . I will . this included cortical . Best structure house and inputs . I doubt I'll put them in the first moved to in battle . well got that men men men of teachers . this is the most primitive room between Oak . it of course when you mix diced pieces apart when all species of Post bedroom but when it's printed the missing so we Note howl in the world to how you infants but I'm side of is open to make think that was new and the of of all of that . though . the setting of the best high but sheltered I haven't input . Gabby input from dendrites and at the outputs down to the axle . can . . so when the sensor new . strange venue . I'll sensation . I was out of sight so and so . she really seems like a blue ribbon is better to separate out with you functions . this letter is aggression . think about this image and spreads it because . to me to ask the question but how sensory most nukes . I'm rest were slowly open the situation Bundaberg and is why we didn't how sensory on most tools to discuss our from sense and at that . so will many would We Hobson studios in the first but when region . Increased sensitivity when cell and said she did that is in CA and function . to melting which is sensation . and mental tension our mission rules sensory elements inside the cell because the cell them how to make space for me at the top of them nestled in Saw design cell . so good tissue the are sensitivity . . I said she felt the effects cell responses . Cindy because we of the potential name any specialization for sensation cell . really I'm not ability to live the special would like to sell . cells . the offence he said . I'm just focus on God in a messy so . the Sox to Edinburgh . I was closest . when the winner situation where they measure when cell . the measurement receptors like when a in response within the single cell . I did sensory neural and signaling to lancelets molds of different types cells . so that symbol . decent edge information . sometimes when I'm rushed larger cells . I'm going human doesn't agree with me and mental effects of wilful to to muscle to back together . so However potential for extreme between sensory modalities . I'm hiding the truth and sensory modalities converge on the same effect the cell . the two nd awful cell . maybe you will commit to . to spend two potential and two . the good them and adding to the right . I mean really look for themselves . through good with sensory ability to detect sense information to lose between system about six . you type . they descend to new and special areas into contempt of the recession of tension is detecting um it chemical imbalance understand specialists function is taking light and the tool converge on the same effect if so then we how convergence of different sentiment dolphins . since this the job . so Purves medicine humans . same symptoms YouTube of them are dead . Phylum also evolved since you don't . so about snot . good grief specialists our is fruit signaling to with iTunes . the reasons why I've never to read the brain knows system the system in several phases just said to neurons in our of pieces from us sensory I'm two months . so let me take it was a two level after level . there of . blue fish . referred to divergence on the inclusions criminals being consumed signal of patients with another man ago when one died . I'm sense to post a little sensory neural was living in responses to lancelets local different effect . cells . the original can in the right many men cells or in this case many motor new role . Ben in the bind many more effective self time these two levels sensory tomorrow so I was from the great city location where would I been sent to new and . the bomb signaling to miss most of the Commission on the Africa mostly on to . and soon to local food that can affect cells' Moussa . um so expression receive this in CNS could call all business Phyla . I'm sure the writing will pull the somewhat outlets where we can address to an animal I'd touch it moved from input to the intense cold . on the road isn't good . I'm trying to improve said . Mendes how exactly does a good to see out . the single coming to when tentacles of the building . resulting in either spells which is completely What question . or two . convergence will have Signals from many widespread sensory neurons . she converging on the ice single most normal course There Wilks going different sensory modalities out well . . nervous system . the sentences that good but sensory on but soon you'll spend on a black day brain that it's centralization and new into system by circulation . neurons in been place . I'm We Carl say where hundred euro with whom I'd better than that how to do all that meat which ago and the new aunts . the movement in ten schools for sensation of movement as well the cake . completed useful completed our of London but will complete the human . I'm separation low distribution love on that . each of the moon those around him al . mom those are and tentacles . when points which is an accumulation of new loans which would even approach a dog basic put it right . well this little as two of the mortgage holder can import a reopening on guard . . . women whose mind again . with the altitude problems . the same guy I'm part of the time . how many keys . it was and attitudes of honey almonds . these are the choice to match her predecessor and I'm to do and dorsal ventral axes in those ago and bought Baltimore why I'm bad at all . very very switch . the reason that those course but for us . I went was Flatworms is because they don't hire any sort of from menstruation system and how many you who don't only go that route between the bed using stuff . then the outside so they the devil ocean . what if using it probably will . on the coast to move goods and to bed diffusion . but is limited to the cell is Columbia of class . organs of the two national . the biggest loser think I will go bust if our was good to me to walk . I used a type Will by the sun example of how one is tapeworm yet in the Gulf of . yes humans . Moyes system arrival were screwed over nuts . I'm the biggest long before in the wall was in it was type wind . I had this piece of white . to be a skill based type of place for me to go again . so from the US Smooth visit and brought suits most of which she won a total that I promptly and exalted the total . because most small really going to tunnel dimensions because I dunno how to nerve . very much a complexity that boat . blue next level of complexity all know system . the good Neurons clustering together in groups . those groups and tools all called ganglia . the news is that you can measure said that is in one place . the sentiment place . but outgunned . how do Neuroscience of three general . communications between the between humans and event cost better you see because of communications which are independent of watching section . if you're going to the weather what he asked me to bottom section nervous tension sitting nicely to in Phyla gun you up . but can be communicated occupation premature to be communicating through the auction said to me . McGrady potentials good . centralization . so that's another thing could course Cerebral gondola which all good the nougat . I know the folks with open because is going axes and all forms of about a news in the direction that is when the direction is useful to Madame de move since Information drilling into the pollen is going into it . so that I was told that this onto the post in axis . I've led to an accumulation of fish of sensory information . in the throes of the organism . I was starting to get the post punk song but isn't to blame but it's on to that little bit like a blank . I've done duo were laid down its bowling games collections all . I'm ofNeural the of . they should be killed on the um cent inflation from a local on the barley . I will soon send out most to mention to dot on the barley . this guy mulesing connected to each other up by the sound of cool . so you know you like the nervous system and of me knew from the local disco . these move cause brain damage mainly so that was going to go equipment just a single one down about . they will start again first thing to look a bit like Brian . it lists all of that move the system complexity . if the Simplest adult how can tools . so there would not grow a good closer neurons and system when they longer just going to call me sensory the motel . when L potentially going form sensory seeking to neural the mud so all from center to motor . um where it's There improves boyhood neural glad I didn't you . range of new functions possible . I'm . could . so us and then the dry winter sensory to today with sentiment so when Increased Shiite how steep the but what goes into conclusions . going out with us . as increase the capacity that would have been great to see dot of condition where not even going to single convergence from different sources into men effects . we can dive and it's Neural so I switch switches in neural networks turning sensory motor connections . I'm all good for example very soon . I would get of whom switch . they had to stop and take tools were popping degenerate soon . you made so they may be beneficial to organism to do we have was spelt . AP Defense information from system your eye and a banal from see . the Greek as well . that's the you um me . you hurled about releasing the credit . I need to detect movements . I'm to detect job that I've been chemical signals that would put it up . Brett its goal . then maybe the Bedok label because the doctor putting the food Bregma new presidency you and that was that ultimately could do so be ready to be complex most well you didn't win when he could of been so insensitive functions . like if he goes slightly different set of circumstances . responses were with and we can be complex behaviors . because the presence that all these in tulips . I can pacemakers Jews generation all in total with them . so compelling to humans live a better job or misconduct rather rude about . I mentioned humans just because when adult setting see . Neural neural connections which mimic or going humans have a few signs will go in humans . I'm for that longer wear them a token outline toxic reflex in humans . should familiar with the modern reflex in the can off many of the word . you basically get the the main point out that Lecture saw the cake simple week since . . . we are . I really don't know why are you address to tune them out and that is possible to move out of Minnesota history . I don't know . well them . when is it . then he made it to the . two men setting reflex . basically I'm sensory pleasure . inside bad a mental day so basically it's the connection that a Masters in the muscle . I'm advanced about stretching the muscle is and will bind to it . so with this system which composed and Ben . today was of course incursion but when sense to neural items to most Neural squealed them into your . so since the November muscle which it stretch . into the local . the continuum . the Nazis with the emotion you want . not to the same muscle . I don't us out with on a new home . due to them sign ups to a second most in your which goes to the antagonist all the muscle . I'm by see what happens is so . signal . then be structure set to . the direct connection Types of the most Neural . so just to Neural Canadian curling construction to stretch causes contraction . I wound up being denied knowing of three mom that I should know . I'm so very simple to new and function . dot is rushing to him how do spells unite us . blunt I do miss the muscle and make us a test . so the same toy and I was contract now mostly gotta make sure all the Muslim League . we're axes . I'm that it was into his knuckle . true destination . um since Jr and stimulates most new . since the nation to stand through new under the lights . this continuum which inhibits the medicine you on preventing contraction all this muscle guy from a tissue over the identity muscles quote in conjunction with each other . many Lecture Corcoran Smooth self I will try and if the same view . the IMO . we started at auction pensions forming in the sense in your . then Dennis and profiles like citation in the big spend so most neural the mosaic was naked al . I'm the Indians mule . because of the underlying between Dr essential sparring for you'll sensory I'm fruit motor and interneuron does because it slice of the top the signal to cost thousands in you and I'm a cross sign ups into much doing on the team's new . so this notion of including a contraction nestled cell . into Images sailing from its new install . Sen the actions and tools like his sons . so the birth of Nestle doesn't contract . I decided so I'm eyes the most good . the symbols could get composed of three helps balloons sentiment is no end . I'm not pursuing the idea is that we can expect to see him problems . level of system complex . in the White think . the move . Identity went on a limestone glia mode which will the snow the institution a takeover know I'm come back . well I found it . but that's does . and . . . . . . . you have . . . . . . the dog . the other brother . the women . thinking . the singer did so . how onto the . even on to it genetic . so and that would shrink this time . yes yes . . . . just happened . but uses well sorry baby . you won't use it yet and you have changes in . Slade . she rest in him . as adults of the mitral sentenced to six . rates . the best public . we are protein generation feeling . so what was being given you be eager to connect and then with the speaker showed its hunting evolution . region four weeks . the text to speech and words to be to make it work in the lake . some are we to recap the resurrection . three when to . you can pick a genetic connection between us . etc put it interesting . so when led . colossal thinking about all joked . great art and good . he is reported to the boil from the norm etc The . you either . it's lovely and out complex problems . um . it's Apes . I tell you what they fruit yet . will the world . I'm quitting . yes . I'm so . she was on . it did . the bill . the site it all . it's a good one . can we . so yes votes though . yet I didn't get on . that and the overlap between that you can do that . who . and my sis . oh . when fine . although we weren't we . when . she which . 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I . the . . . . he . . you will deal with . I . . . . the guy . thank you much good . instead let's continue on . Malcolm Webb opened fire with them too . I'm problems . the internet and segmented Wilson has audio to M'sia in the backyard . I'm close to the tapeworm today in the drops . I'm understood in good are all components of the same time . just because the place and try to make good writing . how did change in the design goes for and problems though both of these Phyla is vice the sign . so often posited in which why this centipedes crustaceans insects . the produce . though I hate Saw admitted to report and Canada West . so I could smooth out . in the group will a White House . houses centipedes oil chemical and other the good for you in the Playhouse in the alley and some on here . so will pinch and also some things well . squatters Alex point . hundred . so if I'll let are in . so instead of going will securely yum . the group drinking but how in the press them into Egypt . and all I was a member all be of the British thousand Society . well slice in my usual thousand is a stick insects . I had understood its the complex sentiment two miles in a binding mental chains . well he's of grumbling be greetings to techniques and so DOI of the places first . I went with our email about from my be going now . moose . the difference . and so ends the Billboard need is going . it's worrying the problems in the Adelaide to multiple . the would not go segmented bonus . so if the stories glia notes will mean previously been reading the lesson Kaylee those who've been of the rings around the blog . I'm in the fragmentation of AC repeat instructions all down to Costa on the actions . this is going to improve the evolution of baby complexity . because you can basically I've . Merle . independent units abroad . in the group . today Joe and generate a well known . segments . he the sunset a genes my own in the not imply sort . don't need anymore genes to generate less local will fall sleep . so again how differently I e long bodie with most of . units to read with very little expansion of of of genetic complexity compare with of the day with a single segment . Cuba old . the compared that's um segmentation boarding the Doomben how . two more specialized functions like how light and bodie Paul which Evans of agro boy called attention Evolves clad little boy calls evolve into a head . so that everyone could divergence . news complex in guts books these are live from is all that eventual know could so that's good for them was committed from us we ars go full that is ventral . it's also said was nerve cord eyes and saw that you would is going to have us all with no coat . that and have done us any segments of the blog . what a lot of thought . and I'm so of launching lots of balloons between those kind it . so Phylum Chordata writes that's drink cases to the Sunday that includes us . so the armed the IPO . I was napping . lunch will lancelets lentil most locals Lancelet system can offer is the right to left . I was a hive time . merchant it's done of them it is Example cuticles sea squirts what they did that come with . I'm soo flow of life is to make that step would be to Motherwell friends and I evolve into the rebate it will then develop into be is Immobile strain structure when it sought to more fruit . the bed the other examples all correlates I was the ha of prime . I'd take his models . you also the result was in court . nobody in there sometimes . I'm certain to correlates . Shoaib and However admitted correlates I need explained . correlates to a house and called a note to cord which is a good full costs less in this mode . so it suits him . today is basically writing region team . along the entire length of the bedding . I'm not true and important in terms of some from from primitive locomotion in cannot Nissl squinted eye on the Bruce . region central Laing . I have opposing muscle groups we will know Amanda crew going to we go in a very good cry and not a true I'm good to swim in a very on the patient's what . ok so damn good wishes found within . and correlates of course in humans without a cause largest one will go on . once one can of Cola bottling . bless me to hide the remnants of this move could Joondalup and the Genesis we haven't that scored just cartilage . the doctor thing gets mad replaced with bottom . completed a place to store my best in the conflict which may we see in the discs between them but it today that is moved remember of the women's all medical . so . when notocord Joe and Andrew enjoyed that . and to choose . the nervous system . I prefer was born but emotion . they said to do this ganglion plague of the skin . maybe I don't let which than I would larger in about of the world salt pinch off . I'm that ultimately pinch off into achieved . on that too is molding go on to form the nervous system or down the aisle to post a axis . I'm . most of her in her all crutches segment us a magical segmented of course we know there will vessels which are all down with an awful . and correlates not cross it was a sian rather warm palm . the website of the left hemisphere the blame will regulate movement all the beyond whites hemisphere . on the way so I bought a of the bawdy no windows . what we had a cough but it is crossed . um . I'm rats was cold and about dorsal to the hall on the Gulf . two nd rhinestones . more later . so I did over the top . corn evolution . we sought to give brains localized we have a blanket you cannot last less than a does have a bit of an accumulation been swelling of the funds and that more funds in the Dubai can but is now in the from the swing going off to mush and almost tissue . and to really referred to as Brian . but of course most of the correlates to we think about . I have cause to brains . um I've read roof to more more more complex pieces this is the cells in golf and then . this group could tax enable cortex than graphs playfully August of from going . I'm not always basal folded up in Canada mouse brain . the curve in the mouse playing other outline of history . they don't get these which danc relations and folding level in the mouse plane because the master who needs as much closer to the snow fall . um but move the moon complex and tenuous the greater cortex in the . I'm the only way to get about what's been in sort of go is quite less local folder . when Chris and Amanda the primates this center is somewhat different climates pieces or tactile a toy actually TV watching from the place up the mouse to dab on infant pregnancy . I've been for free lot more cash . I moved all knowledge brains because when how much more complex behaviors all that may be over them . um . I'm into the apes was close to wait to to chimps of those two boys and ten million years ago . the Welsh coast of the members of the imminent coming of the bowl . no sign how does erectus them a lesson on the site and off of but we only select . so humans as basis . without hominid that it all really . as might be around three thousand units . Odom suggested I saw Saw emotion for humans of spaces . although of the climb would Humans abode of which causes with of all planets . the site I mentioned right side up . point to an African or Jim among humans . beyond this is right on gentle but no doubt many points in schools lack even sure someone died or services and the guy that which is part of the genetic data pointing to . um I've can origin from on humans when they raised circuits India Knight . Lisa was cold when white next week Human into aging quite so um . so gene right with most of all the gene you go is region of the past into a cold be seeking to in many soft light . the sequence all the many soft like a in safe a published new kite . and so decided that I can look of the Structural essentially than most of the . um uh meals . plus the forms all this mini stop light in the UK . all small the the fourth repeat all day to go but today repeat and I'm in the name and most alot about for sure beats . you get from the grounds twenty per cent where it not much longer I will be the lounge and fifty rupees long to go to different in size . I miss the differences structure . but think it's incumbent different from structure . the essential you can write again about killed together with a little boy to the grind . that's it . election coverage found Study rind of one month . you can do now . and today got to find three minutes only for use on the site . you can publish them to come . I need a guy can find fruit in his on of those winters the differences between these populations in though . he wore were put to the whole swarm of small lot else . most heart frequency in Japan the melon UK . so the quilt is a different mode called to detect the same types exist . when fruit to put the Russians form . I'm Africa . I with our niece of over twenty different lineages which all souls a different aisle slices in the fall collection . K so say on this issue into so many of us the this but found the genetic diversity . in the bathroom publishing to bed . it's cold call for a great villain the population can reach anywhere else in the will . and here a of the very soul . oh basal a simplistic explanation . is that . humans are rejected is so hard ask and then left Africa uncle played to the West the world . back in the left More with a subset of complex and with the subset of tombs . that idea a subset denied by the sea . part expression we have next to a diversity out after them we have inside out . so as we left Africa . we have small populations . small populations tend to lose tonight but those the cost of the most published is from Nova on the expression the Dickens when those two . this is that of his many many piece of inflation . from genetics that indicate outlaw region in Some Tanaka . . I live . I'm . understanding with the If behavior essentially . and . I'm good right . just look of the blood in the live every year . infants by one . of course we left . I'm as that for example we consider culture . in the difficult go the minimum behaviors . I was being called one generation Mexican drug teaching . I'm programming . Joanna intentions because our brains also flexible . many thing about Evolution of the human blamed the and the evolution process there is good too . I was too . new be directed towards appear whether compartment computers known can address . I benefits in some good Evolution read them . I'm selective first since those ocean the smell be outspoken computers but instead when outgrown clothes . I will to live and able to rapidly assimilate . Edna the news and humans have human use and is in society . what we on out . this culture of the new outfits ago in Toronto the Jonesboro Africa budget thousand years . but with the echo of the rest couple thousand words . . simply simplicity so mostly it's that good men good sign . so can root out this label culture news of their book and there . transmit to mention generation generation generation where we generation builds on the provision which is in the Evolution stall . necessary but you evolution is unclear because adult to our brains were points when hundred houses see . understanding these . and good but complex cultures generally take complex culture true for the job is like complex language though I'll be traveling to Memphis in the present and Excellent across many many generations were . said the edges then she set up . I'm a winner and pushes permissive will . the descendant of the main dishes which included the balance of culture . I'm identical to bend . you're not going complex in the blame capacity to put in a situation where we ot mum . the move . I am Study that was published a book was published most simply because spoken out in articles local principal problem maps . I'm between Sydney . though the bodies showed dogs . you conclude its power bill the brain . Visible the spaces is going to be basal was bloody what . so I've seen an increase in the right including an increase in price on it whether through relationship between those two increases well if you double the lead . man the brain flow increased by about two . so the baby can grow with blog reading . Essendon the blame for that but can the growing at the same thought . you will go to Venice . you will lead the U brains because the boy in the so do . um . it needs systems allow front of the proceeds as to ablation also put out much information . I'm . that trend of doubling to two threads to I would flee if any spaces is guided into Brian Quinn expects course starts . I'm the most surprising when you Google the biggest brains the you expect crossroads . which I'm writing which essentially describes the leisure between doubling DOI price increase and lancelets what to post . if you all . Kewell . the lead and inspection . Amy's you don't . when that indiscretion dimensions well . Lisa means the goal I'd go to Brighton . the move would