Foundations and Cells of Neuroscience: History, Neurons, Glia, BBB, and Signaling

Founders and History of Neuroscience

  • Ramon y Cajal and Camillo Golgi are considered founding figures of neuroscience.

    • Cajal: helped identify the distinct shapes of neurons and how neurons are structured and connected.

    • Golgi: developed staining methods to view neurons under a microscope.

    • They worked independently in different labs; both were awarded the Nobel Prize in Physiology or Medicine in 1906 for their complementary discoveries.

    • Field is very young; just over a century old (roughly 119 years).

  • Nobel Prize detail and interpersonal drama:

    • Despite sharing the prize, their speeches were famously petty toward each other.

  • Historical misconception: Cajal believed neurons did not regenerate after death, due to the limitations of technology at the time.

  • Bonus historical note about Sigmund Freud:

    • Freud conducted research on Landry’s eels before psychoanalysis, identifying electrical components in cells.

  • Foundations and subfields introduced:

    • Neuroscience uses biology, chemistry, and psychology together to understand how the brain influences thoughts, feelings, and actions.

    • Subfields include:

    • Physiological psychology (synonymously linked with neuroscience; how physiology affects mind).

    • Psychopharmacology (drugs and brain-behavior relationships; planning for signaling and drug effects).

    • Neuropsychology (clinically oriented; illness/injury effects on behavior; dementia tracking and therapy planning).

    • Cognitive neuroscience (youngest subfield; studies how thought processes are affected by brain activity; uses imaging).

    • Imaging techniques: functional magnetic resonance imaging (fMRI) to visualize brain activity during cognitive tasks.

  • Mind-body problem and philosophical orientations:

    • Cartesian dualism (Descartes): mind and body are separate but interact; brain and mental experience interact somehow.

    • Monism: reality is either physical (brain-based) or spiritual; modern neuroscience tends toward physical explanations for conscious experiences.

    • Consensus today: the world is largely biological, but human experience is enriched by spiritual or psychological dimensions; brain function and cognitive enrichment are interdependent and plastic.

  • Scientific approach and humility:

    • Scientists do not claim to prove things; they propose best-supported explanations and remain open to alternative explanations.

    • The Socratic method influences scientific writing: acknowledge uncertainty and possible alternatives.

    • Occam’s Razor (simplest explanation is often best) and Morgan’s Canon (simplest behavioral explanation is likely correct) guide interpretation of data.

  • Categories for explaining behavior (useful heuristics):

    • Physiological explanations (e.g., caffeine’s effects on alertness).

    • Ontogenic/genetic explanations (genetic predispositions).

    • Structural explanations (neural architecture, connectivity).

    • Evolutionary explanations (developmental/adaptive history).

    • Functional explanations (what a behavior achieves or why it exists).

    • Real-world note: various factors interact; avoid over-interpretation of a single mechanism.

  • Quick recap prompt to students:

    • The brain is a complex, interactive system where biology, chemistry, and experience intertwine.

    • Expect lively debates at conferences; focus on evidence and reproducibility.

Foundations and Subfields of Neuroscience (core definitions and scope)

  • Neuroscience integrates biology, chemistry, and psychology to understand how brain activity relates to thoughts, feelings, and actions.

  • Subfields and brief descriptions:

    • Physiological psychology: how physiology affects the mind.

    • Psychopharmacology: how drugs affect brain and behavior; signaling dynamics.

    • Neuropsychology: clinical focus on illness/injury impact on cognition/behavior; dementia tracking and therapy design.

    • Cognitive neuroscience: how thought processes are implemented in neural circuits; often uses imaging to link brain activity to cognition.

  • Imaging and techniques:

    • Functional magnetic resonance imaging (fMRI): visualizes brain activity while performing cognitive tasks; highly complex data.

Mind-Body Problem: Dualism vs. Monism (and contemporary stance)

  • Cartesian dualism (Descartes): mind and body are distinct yet interact; the brain is the seat of mental life.

  • Monism (modern leaning): mental states are produced by brain activity; consciousness arises from neural processes.

  • Contemporary consensus:

    • The brain and mind are deeply interconnected.

    • Conscious experience benefits from cognitive enrichment and neural plasticity; environment shapes biology and vice versa.

  • Practical implications:

    • Brain health depends on cognitive stimulation; a bored brain may exhibit changes in biology (plasticity, neurotransmitter dynamics).

    • Scientific writing favors hedging language over absolute proofs; openness to alternative explanations is essential.

Scientific Principles and Attitudes in Neuroscience

  • Socratic method in science:

    • Acknowledging uncertainty and presenting evidence for support, while proposing alternatives.

  • Occam’s Razor and Morgan’s Canon in practice:

    • Keep explanations simple unless complexity is warranted by data; prefer simpler interpretations of behavior when possible.

  • Example of minimal explanation: caffeine’s behavioral effect can be traced to adenosine antagonism, but avoid over-personal linear causation without mechanistic evidence.

  • Practical tip for students:

    • If your writing uses words like ‘prove’ or ‘causes’ without experimental demonstration, anticipate instructor critique; use language reflecting evidence-based inferences.

Neurons vs Glia: Core Roles and Ratios (structure and function)

  • Neurons (nerve cells): electrical and chemical signaling to communicate messages.

    • Central nervous system (CNS) neurons are located inside the skeleton; peripheral nervous system (PNS) neurons are outside.

    • Neurons are the wired network of the brain; often used as the primary metaphor in cognitive neuroscience (e.g., computer-like signaling).

  • Glia (glial cells): support systems; more diverse roles than originally thought; earlier view as passive scaffolding was revised.

    • Glia are from the Latin for glue; not only glue but essential functional components.

    • Glia can outnumber neurons in some brain regions in certain contexts (e.g., cerebellum has relatively more glia per neuron).

  • Examples of glial types and roles (overview):

    • Radial glia: migratory scaffolds during early brain development; help guide neurons to their destinations; numbers decline as development completes.

    • Astrocytes: star-shaped cells; most versatile and abundant glia; roles include nutrient delivery, waste clearance, CNS maintenance, synaptic modulation, and blood-brain barrier (BBB) support; can form glial scars after injury.

    • Oligodendrocytes: form myelin in CNS; can myelinate multiple axons; myelin here is a CNS-specific myelin; CNS myelin is less permissive to regrowth once damaged.

    • Schwann cells: form myelin in the peripheral nervous system; promote nerve regeneration and growth after injury.

    • Microglia: immune cells of the CNS; microscopic and mobile; transform shape when activated; two major states in some models: M1 (pro-inflammatory) and M2 (anti-inflammatory); act as cleanup and immune defense; can switch phenotypes to address injury or inflammation; can contribute to disease states if dysregulated.

  • Glial scar and its implications:

    • Astrocyte-driven glial scar forms after CNS injury to isolate damaged tissue and protect surrounding areas.

    • In chronic injuries, glial scar can become a barrier to regrowth; in some cases, scar tissue can be surgically removed or chemically broken down to allow regrowth.

  • Microglia details:

    • “Tiny” cells that respond to injury; can revert to quiescent state when not needed; activation leads to pro- or anti-inflammatory states depending on signals.

    • Glial activity is a target for therapies to modulate inflammation and tissue repair.

  • Clinical and research relevance:

    • Glia contribute to pain signaling (astrocytes), BBB integrity, immune responses, and neuron support.

    • Glial pathology is implicated in many neurological disorders (e.g., glioblastoma, multiple sclerosis, neurodegenerative diseases).

Neuron Anatomy: Dendrites, Soma, Axon, and Terminals (structure and flow of signals)

  • General architecture (tree analogy):

    • Dendrites: branched extensions that receive signals from other neurons (like leaves absorbing sunlight).

    • Soma (cell body): contains nucleus and major organelles; site of basic cellular metabolism (mitochondria, nucleus, endoplasmic reticulum).

    • Axon: long, insulated conduit for electrical signals; often described as the “electrical highway” or an HDMI-like cable; white matter coloration is largely due to myelin.

    • Axon terminals: knob-like endings that release neurotransmitters to communicate with the next neuron (synaptic boutons).

  • Key terms and structures:

    • Dendrites receive input; soma integrates input; axon conducts action potentials to terminals.

    • Synapse and transmission: chemical signaling via neurotransmitters stored in vesicles; neurotransmitters released into the synaptic cleft and received by receptors on the postsynaptic neuron.

    • Neurotransmitter vesicles: membrane-bound packages that release neurotransmitters into the synapse when an action potential arrives.

    • Terminal buttons (boutons): presynaptic release sites that connect to the postsynaptic dendrites (or soma).

  • Axon-specific internal structures:

    • Microtubules: intracellular “fiber optic” cables that transport vesicles and organelles along the axon; tau protein can become tangled in disease (e.g., Alzheimer’s).

    • One axon per neuron (in most neurons) with an orthodromic direction (signal travels from cell body to terminal).

    • Interneurons may lack an axon (relay interneuron role in local circuits).

  • The standard neuron model and its origin:

    • The classic neuron model (pyramidal neuron as a common example) is heavily derived from the squid giant axon, which helped establish foundational principles of neuronal signaling.

    • Squid giant axon provided a long, accessible system to study action potentials in isolation.

  • Common neuron types mentioned:

    • Purkinje neurons (cerebellum): large, highly dendritic; critical for motor coordination.

    • Pyramidal neurons (cortex): cone-shaped cell bodies; central to higher cognitive processes.

    • Kenyon cells (honeybee): involved in memory and navigation in insects; illustrate diversity of neural processing across species.

    • Bipolar neurons (retina): neurons with two processes; important for vision processing.

  • Common note on neuron variability:

    • Not all neurons look the same; a standard model (often the pyramidal neuron) is used for teaching, but there is wide diversity across brain regions and species.

The Axon, Myelin, and Conduction: Speeding the Signal (why myelin matters)

  • Myelin and white matter:

    • Myelin is a fatty, lipid-rich insulation around axons; white color arises from lipid content.

    • It speeds signal transmission by insulating the axon and reducing membrane capacitance.

  • Nodes of Ranvier and saltatory conduction:

    • Myelinated segments alternate with exposed gaps (nodes of Ranvier).

    • Action potentials jump from node to node (saltatory conduction), increasing conduction velocity.

    • The term saltatory conduction involves sodium ions; precise ionic interactions occur at nodes during signaling.

  • Consequences of demyelination:

    • Diseases like multiple sclerosis (MS) and spinal cord injury (SCI) slow or block signal transmission due to exposed axon segments.

    • Resulting symptoms can include slowed reflexes, numbness, pain, and motor deficits.

  • Terminology and directionality:

    • Orthodromic conduction: signal travels in the natural direction (cell body to terminal).

    • Antidromic conduction: reverse direction; observed rarely in natural conditions but possible with artificial stimulation.

  • Myelin-producing cell types and locations:

    • Oligodendrocytes: myelinate CNS axons; can myelinate multiple axons; CNS myelin often resists regrowth when damaged.

    • Schwann cells: myelinate PNS axons; support regrowth after injury; peripheral nerves may recover better due to Schwann-mediated regrowth.

  • Important concept: absolute refractory period and relative refractory period:

    • After an action potential, neurons enter a brief period during which another action potential cannot be fired (absolute refractory).

    • This is followed by a relative refractory period where a stronger stimulus could trigger another action potential.

  • Electrical basis of signaling (membrane dynamics):

    • Neurons have a resting potential of approximately Vrest70mVV_{rest} \,\approx\,-70\,\text{mV} due to ionic gradients and selective permeability.

    • Action potentials require depolarization to a threshold around V<em>th55mVV<em>{th} \,\approx\,-55\,\text{mV}, after which a rapid rise to a peak occurs ( often reported as up to V</em>peak+55mVV</em>{peak} \approx +55\,\text{mV} in some classic; actual peak varies by neuron).

    • The membrane is polarized at rest; depolarization opens voltage-gated channels; after peak, repolarization restores the resting state with possible hyperpolarization.

  • Ion channels and pumps (basic mechanisms):

    • Ion channels: selective pores in the membrane that allow specific ions (Na+, K+, Ca2+) to pass when open.

    • Leaky channels and voltage-gated channels regulate ion flow; opening of channels changes the membrane potential.

    • Sodium-Potassium Pump (Na+/K+ ATPase): actively moves Na+ out and K+ in to restore resting gradient after activity; commonly represented as:
      3Na+  out,  2K+  inper ATP.3\,\text{Na}^+\;\text{out},\; 2\,\text{K}^+\;\text{in} \text{per ATP}.

    • Transporters: energy-dependent mechanisms that help with molecule movement and maintenance of gradients (akin to security checks like TSA).

  • Membrane structure and selective permeability:

    • Lipid bilayer forms a hydrophobic interior, resisting water-soluble substances.

    • Channel proteins create selective pathways for ions; lipid-soluble substances diffuse more readily through the membrane.

    • Gradient-driven diffusion and active transport interact to maintain resting potential and enable signaling.

  • Practical visualization and student notes:

    • A common visualization uses an exaggerated cartoon with a polarized cell at rest, depolarization, and repolarization dynamics across the axon length.

    • The AP worksheet referenced by the instructor provides step-by-step guidance through membrane potential changes and channel dynamics.

Structure and Function of Glia: In-Depth Look at Supportive Cells

  • Highlights of glial diversity and their importance:

    • Glia are not just passive support; they are essential for signaling, protection, and repair.

    • The ratio of neurons to glia varies by region (e.g., cortex vs. cerebellum) and species; glia can outnumber neurons in some contexts.

  • Specific glial types and roles:

    • Radial glia: migratory scaffolds during early development; guide neuronal placement; numbers decrease after development.

    • Astrocytes: the most prominent glial cells; multiple roles including delivering nutrients, clearing debris, supporting synaptic function, modulating signals, and maintaining the blood-brain barrier (BBB); can also form glial scars after injury.

    • Oligodendrocytes: produce myelin in the CNS; can wrap multiple axons; CNS myelin limits regrowth if damaged.

    • Schwann cells: myelinate peripheral nerves; promote peripheral nerve growth and regeneration throughout life.

    • Microglia: immune cells of the CNS; highly dynamic; can adopt pro- or anti-inflammatory states; clear debris and support immune responses; can transform shapes when activated (resembling a Swiffer when cleaning up).

  • Microglia states and activation:

    • Activation status can shift between pro-inflammatory (M1) and anti-inflammatory (M2) states; this plasticity is a target for therapies to regulate neuroinflammation.

  • Astrocyte functions and BBB involvement:

    • Astrocytes maintain the blood-brain barrier (BBB) by interacting with endothelial cells; endfeet of astrocytes envelop blood vessels and regulate nutrient transport.

    • BBB consists of endothelial tight junctions that tightly regulate substances entering the brain; astrocyte endfeet actively transport nutrients and protect the brain from toxins.

    • Abnormal BBB permeability can contribute to stroke and drug delivery challenges.

  • Blood-Brain Barrier (BBB) and meninges overview:

    • BBB integrates with meninges to form a protective barrier around the brain.

    • Endothelial cells, tight junctions, basal lamina, pericytes, and astrocyte endfeet jointly contribute to BBB integrity.

    • Meninges provide protective layers around the brain and are discussed in more detail below (dura mater, arachnoid, pia mater).

  • Clinical terms and practical notes:

    • Glioblastoma: highly malignant glial tumor with poor prognosis; glial proliferation complicates treatment.

    • Friable tissue: characteristic of certain tumors; pathological term describing tissue that crumbles easily; glial scar tissue tends to be non-friable but can be physically removed if needed.

The Blood-Brain Barrier and Meniges: Protection, Pathways, and Pathologies

  • BBB key components and their roles:

    • Endothelial cells form tight junctions that limit paracellular diffusion.

    • Astrocyte endfeet regulate transport and support barrier integrity; BBB prevents many toxins from entering the brain (e.g., most large or lipophobic molecules).

    • Small, lipid-soluble molecules may diffuse through; larger or hydrophilic molecules require specific transporters.

  • Important brain barriers and protective layers:

    • BBB (blood-brain barrier): protects brain from toxins; regulates nutrient transport; can be compromised during strokes or disease.

    • Dura mater: tough outer membrane; provides mechanical protection; robust structure (etymology: “tough mother”).

    • Arachnoid mater: middle layer; spider-web-like appearance; contains CSF-containing subarachnoid space.

    • Pia mater: delicate inner membrane that adheres to the brain; closely follows brain contours.

  • Subarachnoid space and CSF:

    • Subarachnoid space contains cerebrospinal fluid (CSF); CSF helps cushion the brain and remove waste.

    • Hydrocephalus is an excess accumulation of CSF; aqueducts and CSF circulation pathways are important for nutrient delivery and toxin removal.

  • Cribriform plate and vulnerability:

    • A thin, porous area near the nasal cavity; particularly vulnerable to injury and infections; associated with a site of vulnerability in the context of Alzheimer's disease and cocaine use.

  • Meningitis: forms and red flags

    • Forms include viral, bacterial, parasitic, and fungal (fungal form is rarer and more dangerous).

    • Viral meningitis is most common; vaccines exist for some forms (e.g., certain viral meningitis agents).

    • Bacterial meningitis is more dangerous due to rapid progression; requires urgent care and isolation to prevent spread; red flag symptom includes a stiff neck (meningeal signs).

    • Parasitic meningitis is rare but recognized; fungal meningitis also exists; vaccines and treatment vary by organism.

    • Symptoms commonly include headache, fever, neck stiffness; stiff neck is a red flag prompting urgent medical evaluation.

  • Clinical management notes:

    • Vaccines exist for certain meningitis-causing agents; prompt treatment can be life-saving in bacterial meningitis.

    • Quarantine and isolation practices are used to prevent spread in suspected contagious cases.

  • CSF sampling and meninges details:

    • Spinal taps pierce the subarachnoid space to sample CSF for meningitis diagnosis.

    • Meningitis can be tied to the meninges being inflamed, leading to potential brain injury if swelling is not managed.

  • Neti pot caution:

    • Rare cases linked to amoebic meningitis from neti pot use with non-sterile water; use distilled or boiled water for nasal rinses.

  • The brain’s energy and metabolism in context of BBB:

    • Brain energy relies primarily on glucose; in starvation, ketone bodies can serve as an alternative fuel.

    • Thiamine (vitamin B1) is essential for carbohydrate metabolism; deficiency can impair cognitive function and memory.

    • Korsakoff syndrome (thiamine deficiency often linked to chronic alcoholism) causes severe memory impairment and confabulation (filling memory gaps with invented stories; patients may be highly suggestible).

    • Korsakoff has nicknames like “wet brain” or “pickle brain” due to memory impairment and confabulation; treatment depends on thiamine replacement and nutritional rehabilitation.

  • Nutritional and metabolic considerations:

    • Glucose is the brain’s dominant energy source under normal conditions.

    • In fasting or diabetes, ketones may become more prominent; improper balance can lead to metabolic complications (e.g., ketoacidosis).

    • Adequate vitamins (e.g., thiamine) are essential for cognitive functions; deficits can impair memory and executive function.

  • Memory and language symptoms as examples:

    • Anomia (difficulty naming objects) can result from temporal lobe or language network disruptions; a patient’s post-surgical outcome can include improved naming with time or therapy.

Neuronal Signaling: Action Potentials, Ion Flux, and the All-or-None Principle

  • Action potential concept:

    • Action potential is the electrical impulse that travels along axons; neurons with axons generate action potentials.

    • The phrase “action potential” reflects the all-or-none nature: once the threshold is reached, the signal propagates with consistent magnitude down the entire length of the axon.

    • All-or-none property ensures signal strength is preserved from origin to terminus.

  • Mechanistic steps of the action potential (simplified overview):

    • Resting state: membrane is polarized with a negative inside and positive outside; typical resting potential around Vrest70mVV_{rest} \approx -70\,\text{mV}.

    • Thresholding: depolarization to a critical level triggers voltage-gated Na+ channels to open; rapid Na+ influx drives the rising phase.

    • Peak: membrane potential reaches a positive value (e.g., up to around Vpeak+55mVV_{peak} \approx +55\,\text{mV} in many models).

    • Repolarization: Na+ channels inactivate, K+ channels open; K+ efflux returns the membrane toward resting potential.

    • Hyperpolarization: the membrane potential may dip below resting level (hyperpolarized) briefly.

    • Refractory periods: absolute refractory period prevents another action potential; relative refractory period requires stronger stimulation to trigger another spike.

  • Ion channels and transporters involved:

    • Ion channels: specific pores that allow Na+, K+, Ca2+ to pass when open.

    • Voltage-gated channels: open or close in response to changes in membrane potential; essential for the action potential.

    • Na+/K+ pump: restores ion gradients after an action potential by transporting Na+ out and K+ in against their concentration gradients (ATP-driven):
      3Na+ out,  2K+ inper ATP.3\,\text{Na}^+\text{ out},\;2\,\text{K}^+\text{ in} \quad \text{per ATP}.

    • Ion exchangers/transporters: actively regulate ion distributions and help maintain homeostasis.

  • The role of myelin in conduction speed:

    • Myelin reduces membrane capacitance and allows current to travel more efficiently between nodes of Ranvier.

    • Saltatory conduction (jumping between nodes) makes signaling much faster in myelinated axons.

  • Directionality and realism in signaling:

    • In neurons with axons, signals generally move in the orthodromic direction (from soma toward terminals).

    • Antidromic conduction can be produced experimentally but is not a common natural direction for neural signaling.

  • The membrane’s resting and active states in the math/physics perspective:

    • Resting potential is a result of ionic gradients and selective permeability, with the inside typically around Vrest70mVV_{rest} \approx -70\,\text{mV} and the outside more positive.

    • The gradient and channel dynamics produce voltage changes that propagate as a wave of depolarization along the axon.

  • Practical notes and study aids:

    • The AP worksheet mentioned by the instructor provides a step-by-step breakdown of membrane potential changes and channel mechanics.

    • Visuals often include a schematic of channels opening, ion flow, and a propagating wave along the axon.

  • Clinical implications of signaling dynamics:

    • Demyelinating diseases slow conduction and alter the timing of neural communication, affecting motor control and sensation.

    • Abnormal ion channel function can underlie various neuropathies and epileptic activity.

Visualizing and Connecting: From Cells to Behavior

  • The brain’s connectivity and signaling scale from molecules to behavior:

    • Synapses: chemical communication occurs across the synaptic cleft via neurotransmitters released from presynaptic terminals into the cleft and binding to postsynaptic receptors.

    • Synaptic cleft: a tiny gap that ensures neurotransmitter release and receptor binding produce precise, timed signaling; opportunities for modulation and pathology.

  • The importance of context and history in interpretation:

    • Early models (e.g., squid giant axon) provided critical insights into action potentials; contemporary neuroscience integrates molecular, cellular, systems, and cognitive levels.

  • Everyday relevance and cautions:

    • When explaining brain function to non-experts, avoid overstatement; emphasize the best-supported mechanisms and remaining uncertainties.

  • Final reminder:

    • The brain’s complexity requires integrating multiple cell types (neurons and glia), barriers (BBB and meninges), and signaling systems (ionic flux, neurotransmitters, and neuromodulators) to understand behavior and disease.

Quick Reference: Key Terms and Concepts (glossary-style)

  • Neuron types and components:

    • Dendrites, Soma, Axon, Terminal buttons (boutons)

    • Pyramidal neuron, Purkinje neuron, Kenyon cells

  • Glia types and roles:

    • Astrocyte, Oligodendrocyte, Schwann cell, Microglia, Radial glia

  • Barriers and coverings:

    • Blood-brain barrier (BBB), Dura mater, Arachnoid, Pia mater, Cribriform plate

  • Membrane dynamics and signaling:

    • Resting potential Vrest70mVV_{rest} \approx -70\,\text{mV}

    • Threshold V<em>th55mVV<em>{th} \approx -55\,\text{mV}, Peak V</em>peak+55mVV</em>{peak} \approx +55\,\text{mV}

    • Saltatory conduction, Nodes of Ranvier, Orthodromic vs antidromic

    • Sodium-potassium pump: 3Na+ out,  2K+ in per ATP3\,\text{Na}^+\text{ out},\;2\,\text{K}^+\text{ in} \text{ per ATP}

    • Synapse, Neurotransmitter, Vesicles, Synaptic cleft

  • Health, disease, and nutrition:

    • Korsakoff syndrome, Confabulation, Thiamine (Vitamin B1)

    • Glial scar, Glioblastoma, Inflammation states (M1/M2)

    • Meningitis forms: viral, bacterial, parasitic, fungal

    • Hydrocephalus, Cribriform plate vulnerability

  • Philosophical and methodological notes:

    • Cartesian dualism, Monism, Occam’s Razor, Morgan’s Canon, Socratic method

  • Energy and metabolism:

    • Glucose as brain energy, Ketones in fasting, Oxygen necessity, Astrocyte-mediated nutrient transport

  • Neuroanatomical energy and support:

    • BBB transporters, Endothelial tight junctions, Astrocyte endfeet, Meningeal protection, CSF dynamics

  • Development and memory:

    • Radial glia as migratory scaffolds; astrocyte involvement in memory signaling; hippocampal and cortical memory circuits