A reading order for Kandel
Principles of Neural Science is 1,600 pages. Here is which chapters to read in which order so that each one lands on something you have already built, one a week for about a year.
On this pageOne chapter a week
KandelPrinciples of Neural ScienceThe reference text of neuroscience, read a chapter a week in the site order for about a year. is the reference text of neuroscience and reading it cover to cover is a mistake: the early chapters on molecular biology are the least relevant to an engineer, and the chapters that matter most are scattered. This order follows the site. Each block attaches to a phase so that the biology explains something you have already measured. Chapter numbers refer to the sixth edition; the titles are stable across editions and are what to search for.
Block one, with Phase 1: the cell and the spike
- With Phase 1Block one: the cell and the spikeMembrane, channels, the action potential, propagation. You will have heard one on a speaker before you read about it.
- With Phase 2Block two: from cell to signalSynapses, populations, and where the field on the scalp comes from. Read while your amplifier is on the desk.
- With Phase 3Block three: the systems BCIs readSensory pathways, motor cortex, attention. Read while your classifier is failing on day two.
- With Phase 4Block four: rhythms, sleep, plasticityThe chapters that explain the oscillations you have been recording since Phase 1.
- With Phases 5 and 6Block five: the clinic and the frontierDisorders, degeneration, and repair. Read alongside the regulatory and implant material.
Overview chapters on the brain and behaviour and on nerve cells and circuits (the first two or three chapters). Then the membrane potentialMembrane potential The voltage across a cell's membrane, about −70 millivolts at rest, set by ion concentrations and which channels are open. Related: Resting potentialResting potential The steady membrane voltage of a neuron that is not firing, typically −60 to −70 millivolts. , Action potentialAction potential The brief, all-or-nothing voltage spike a neuron fires, about a millisecond long and about 100 millivolts tall across its membrane. , Ion channelIon channel A protein pore in the membrane that lets specific ions through, often only when the voltage or a chemical tells it to open. . Glossary entry and passive properties, ion channelsIon channel A protein pore in the membrane that lets specific ions through, often only when the voltage or a chemical tells it to open. Related: Action potentialAction potential The brief, all-or-nothing voltage spike a neuron fires, about a millisecond long and about 100 millivolts tall across its membrane. , Hodgkin–Huxley modelHodgkin–Huxley model The 1952 set of four differential equations for the squid axon that reproduces the action potential from voltage-dependent sodium and potassium channels. . Glossary entry, and propagated signalling: the action potentialAction potential The brief, all-or-nothing voltage spike a neuron fires, about a millisecond long and about 100 millivolts tall across its membrane. Related: Membrane potentialMembrane potential The voltage across a cell's membrane, about −70 millivolts at rest, set by ion concentrations and which channels are open. , Ion channelIon channel A protein pore in the membrane that lets specific ions through, often only when the voltage or a chemical tells it to open. , SpikeSpike Shorthand for an action potential as seen by an electrode, a brief blip about a millisecond wide. , Extracellular recordingExtracellular recording Recording with an electrode near neurons rather than inside one; you see small, fast blips from nearby cells rather than the full membrane swing. . Glossary entry. Read these while the SpikerBoxSpikerBox Backyard Brains' small battery amplifier that lets you hear action potentials from a cockroach leg on a speaker; easy to build yourself. Related: Action potentialAction potential The brief, all-or-nothing voltage spike a neuron fires, about a millisecond long and about 100 millivolts tall across its membrane. , Instrumentation amplifierInstrumentation amplifier A precision differential amplifier with very high input impedance and high CMRR, built from three op-amps and sold as a single chip such as the INA128 or AD8232. . Glossary entry is on the bench. Four to six weeks.
Block two, with Phase 2: from cell to signal
Synaptic transmission chapters: the overview, transmitter release, and the postsynaptic receptors. Then the chapter on the EEGElectroencephalography (EEG) Recording the brain's electrical activity from electrodes on the scalp, a few tens of microvolts of blurred, summed cortical activity. Related: Electrocorticography (ECoG)Electrocorticography (ECoG) Recording from electrodes placed on the surface of the brain under the skull, usually during epilepsy surgery evaluation; cleaner and sharper than EEG. , Magnetoencephalography (MEG)Magnetoencephalography (MEG) Recording the tiny magnetic fields produced by cortical currents with superconducting sensors in a shielded room; less blurred by the skull than EEG, and far more expensive. , Volume conductionVolume conduction The spreading of currents from a source through the conductive tissues of the head, which is why one source shows up on many electrodes and why scalp EEG is blurry. , Microvolt (µV)Microvolt (µV) One millionth of a volt. Scalp EEG is tens of microvolts; a AA battery is 1.5 million of them. . Glossary entry and brain rhythms if your edition has one (the “sleep and wakefulness” and “seizures” chapters carry the EEGElectroencephalography (EEG) Recording the brain's electrical activity from electrodes on the scalp, a few tens of microvolts of blurred, summed cortical activity. Related: Electrocorticography (ECoG)Electrocorticography (ECoG) Recording from electrodes placed on the surface of the brain under the skull, usually during epilepsy surgery evaluation; cleaner and sharper than EEG. , Magnetoencephalography (MEG)Magnetoencephalography (MEG) Recording the tiny magnetic fields produced by cortical currents with superconducting sensors in a shielded room; less blurred by the skull than EEG, and far more expensive. , Volume conductionVolume conduction The spreading of currents from a source through the conductive tissues of the head, which is why one source shows up on many electrodes and why scalp EEG is blurry. , Microvolt (µV)Microvolt (µV) One millionth of a volt. Scalp EEG is tens of microvolts; a AA battery is 1.5 million of them. . Glossary entry material in older editions). This is where the electrodeElectrode The conductor that turns ion currents in tissue into electron currents in a wire; its chemistry and contact quality set the noise you live with. Related: Silver/silver-chloride (Ag/AgCl)Silver/silver-chloride (Ag/AgCl) The standard electrode material for biopotentials, because its chemistry produces a stable, low-noise, low-drift interface with salty gel. , Electrode impedanceElectrode impedance How hard it is to push current through the skin-electrode contact, typically 5 to 50 kilohms with gel and megohms dry, higher at low frequencies. , Half-cell potentialHalf-cell potential The DC voltage that appears wherever a metal meets an electrolyte; two electrodes with different half-cell potentials produce an offset that can swamp the signal. . Glossary entry-electrolyte interface meets the synaptic currents it records. Four weeks.
Block three, with Phase 3: the systems that BCIs read
The motor system chapters: the organization of movement, voluntary movement and the primary motor cortex, the premotor cortex and planning. Then the sensory coding chapter and the somatosensory system chapters. Then the chapters on the visual system, at least the retina and the primary visual cortex, for the SSVEPSteady-state visual evoked potential (SSVEP) When you stare at something flickering at a fixed rate, visual cortex oscillates at that rate, and the frequency can be read straight out of the EEG spectrum. Related: Canonical correlation analysis (CCA)Canonical correlation analysis (CCA) Finding the combination of channels that correlates best with reference sinusoids at each candidate frequency; the standard SSVEP decoder. , Brain-computer interface (BCI)Brain-computer interface (BCI) A system that turns measured brain activity into control of something outside the body, from a cursor to a wheelchair to a speech synthesizer. . Glossary entry and prosthesis work. Eight weeks.
Block four, with Phase 4: the rhythms, sleep, and plasticity
Sleep and wakefulness. The chapters on learning and memory: the cellular mechanisms of implicit memory (this is where LTP lives), and the hippocampus and explicit memory. The chapter on brain development and critical periods for plasticity. Six weeks.
Block five, with Phase 5 and 6: the clinic and the frontier
The chapters on diseases of the motor unit and on movement disorders (Parkinson’s, the basal ganglia, and thus DBSDeep brain stimulation (DBS) Electrodes implanted deep in the brain and pulsed continuously, the standard treatment for advanced Parkinson's disease and essential tremor. Related: Charge densityCharge density The charge injected per pulse divided by electrode area; keeping it below safe limits prevents electrode damage and tissue injury. . Glossary entry). The chapter on seizures and epilepsyEpilepsy A tendency to recurrent seizures; its monitoring units are where seizure-detection algorithms get validated and where ECoG data comes from. Related: Epilepsy monitoring unit (EMU)Epilepsy monitoring unit (EMU) The hospital ward where patients are recorded on video-EEG for days to capture seizures before surgery. , Seizure detectionSeizure detection Automatically flagging seizures in EEG, a real clinical need and a good first machine-learning project on the TUH corpus. . Glossary entry. The chapters on the auditory system (the cochlear implantCochlear implant The most successful neural prosthesis: an electrode array in the inner ear that stimulates the auditory nerve in up to 22 bands. Related: Vocoder (noise vocoder)Vocoder (noise vocoder) A simulation of what a cochlear implant delivers: the loudness envelope of each frequency band, applied to noise in that band. . Glossary entry’s biology) and the vestibular system. The chapter on spinal cord injury and repair if present. Six weeks.
What to skip
The detailed molecular chapters on gene expression, the chapters on development beyond the plasticity material, most of the chapters on emotion, motivation, and language unless a project takes you there. They are excellent; they are not on the path.
How to read each chapter
Fifteen minutes with the summary and the figures first, as the paper-reading page says. Then the text. Then five lines in the notebook: what the chapter explained that you had seen, one fact you did not expect, one thing you still do not understand. A year of five-line summaries is a personal textbook.