Name two complementary coding schemes for external stimuli
Time Coding = detailed firing structure of response
Frequency Coding = number of action potential per time unit
Name four properties of grid cells
Within one module:
aligned orientation
similar grid scale
spatial phase shifts between grid cells
single grid cell module can not represent spatial location
-> without 1. or 2. one module could encode space
Neighboring grid cells:
similar spatial freq
similar orientation
different phase
Tick all correct answers (eight choices)
The cortex has 8 layers.
The “nucleus” has a laminar structure and contains small cell bodies.
The midbrain receives input from all spatially organized sensory organs and forms multisensory maps of the outer world.
The afferent fibers enter the spinal cord via the dorsal root.
Every organism with neurons has a brain.
The right and left hemisphere are connected via glial cells.
The reason for different neuronal morphologies lies in the computations they perform.
lost info :(
Correct: 3., 4., 7.
Rest is incorrect, bc
cortex has 6 layers
nuclei in the brain are clusters of neurons
some animals (e.g. jellyfish) have neurons but no centralized brain
right and left hemisphere are connected via corpus callosum
Given an ion X, calculate its Nernst potential given that (X_out/X_in) is 100.
E = 58 x log10(100) = 58 x 2 = 116 (mV)
Why do sodium-channels close right after they are opened?
Due to inactivation mechanisms:
sodium channels open -> Na+ intake -> depolarization -> inactivation by blocking further Na+ preventing further depolarization and allowing neuron to return to its resting state of negative voltage -> it is closed until next spike
Explain the signal transduction of an action potential in a myelinated axon
Action potential is propagated by saltatory conduction, where the electrical signal jumps between nodes of Ranvier (= gaps in the myelin sheath).
At each node, voltage-gated Na⁺ channels open, regenerating the action potential.
Conduction speed increases by reducing ion leakage and membrane capacitance along the myelin insulation.
Explain the relationship between information and uncertainty.
Information is defined as the reduction of uncertainty.
-> neuron's response conveys more information when it reduces uncertainty about the external world
-> greater neural selectivity or variability in response to different inputs increases the information content, because it helps distinguish between possible causes of sensory input.
What are Shannon’s axioms of uncertainty measures?
For equiprobable states uncertainty grows with the number that states
Uncertainty should be continuous in probability of states
“Branching” property (let’s discuss some other time)
How can you measure uncertainty?
Shannon’s entropy:
Entropy of a system = measure of uncertainty about it’s state
-> describes the capacity of a system to carry information
Given three spikes: sketch when the uncertainty is minimal and when it is maximal.
Explain important aspects in the emergence of functional circuits during development
early developement: activity-independent organization mechanisms where brain wires itself using chem. signals
afterwards, activity-dependent plasticity refines these connections based on experience and neural activity
Spontaneous activity in immature circuits for axonal refinement even before sensory input is fully available
Over time, synaptic pruning and experience-driven modifications to further sculpt circuits
Activity and Connectivity influence each other:
-> activity via synaptic plasticity forms conntectios & connection dynamics allow activity at different strengths
can happen at different scales: synaptic or circuit
When photoreceptors absorb photons, are they depolarized or hyperpolarized and is it graded potential or action potential?
Hyperpolarized and graded potential
Why is the dLGN of the thalamus not just a relay station?
because it actively & dynamically filters and modulates visual information before sending it to the cortex
For which feature are simple cells in the primary cortex selective and how do they work?
simple cells = elongated spatial RFs with neighboring on/off subfields
-> selective for orientation
How it works:
-> best response when a stimulus is at the right orientation and location within RF
-> RF have distinct ON and OFF subregions
-> respond to light or darkness
In which three “areas” do retinal ganglion cells perform redundancy reduction?
Spatial contrast, temporal contrast, spectral contrast
Name the two main processing streams in the extrastriate areas of the visual cortex.
Dorsal (parietal) stream (“where”): motion and spatial perception/depth → spatial location
Ventral (temporal) stream (“what”): color, form, fine detail → object recognition
Sketch the waveform strategy of continuous interleaved sampling.
How can you decode an animal’s location when measuring neural activity?
decode an animal’s location by recording from spatially tuned neurons like place cells or grid cells, which fire at specific positions
Decoding methods:
Bayesian decoding (uses prior knowledge of each cell's tuning curve to estimate the most likely position)
Population vector decoding (treats the firing rates as weighted vectors to compute a position estimate)
Explain the principle of experience-dependent correlated activity in the hippocampus with replay of neural activity in sleep.
Principle of experience-dependent correlated activity = Neurons that fire together get wired to gether (Hebbs Postulate)
Context of replay of neural activity in sleep:
in slow-wave sleep, the brain replays activity patterns in a compressed and coordinated way, reflects the same sequence of place cell activation seen during waking behavior.
The correlated replay strengthens synaptic connections, supporting memory consolidation.
-> sleep helps stabilize and store experiences by reactivating and reinforcing learned patterns
Explain the difference between dedicated and intrinsic models.
Dedicated models:
assume that specific neurons or circuits are specialized to encode particular features or functions
-> fixed, labeled lines
Intrinsic models:
suggest that coding arises from the dynamics and interactions within a general-purpose network, without needing predefined feature-selective units.
-> emergent activity patterns
-> often more flexible and context-dependent
What is Weber’s law?
= discriminality of two nearby stimuli depends on ratio between intensities and not absolute magnitude
-> reflects how our perception scales with stimulus intensity.
Name one model to measure time.
Pacemaker–accumulator model =
Internal pacemaker generates pulses at a regular rate.
These pulses are sent to an accumulator when attention is directed to timing.
The total number of accumulated pulses is used to estimate elapsed time.
This model helps explain how we compare durations and why our sense of time can vary with attention or arousal.
Explain the circadian transcriptional circuit.
The circadian transcriptional circuit is a molecular feedback loop to generate ~24-hour rhythms using a gene expression cycle.
The circuit is within the suprachiasmatic nucleus in the hypothalamus and gets input form the retina
Which model describes the center-surround receptive field structure of retinal ganglion cells?
Difference-of-Gaussians (DoG) model
= represents how retinal ganglion cells respond to light in their receptive fields
models the center (excitatory) and surround (inhibitory) regions as two overlapping Gaussian functions
By subtracting the surround Gaussian from the center Gaussian, the model captures the antagonistic center-surround structure
How does processing in the retina reduce redundancy?
by emphasizing changes and differences in light rather than absolute intensity
Center-surround receptive fields filter out uniform areas and highlight contrast, removing spatial redundancy
Temporal adaptation allows cells to focus on changes over time, ignoring constant stimuli
Lateral inhibition sharpens edges and decorrelates neighboring signals
What are decorrelations methods and how do they differ?
= methods to reduce statistical dependencies between input signals to make information more efficient and less redundant
Methods:
Principal Component Analysis (PCA) – removes linear correlations by rotating the data to align with axes of maximum variance; outputs uncorrelated components but may still be statistically dependent.
Zero-phase Component Analysis (ZCA) – similar to PCA but preserves the original data orientation; results look more like the input, useful for early vision models.
Independent Component Analysis (ICA) – goes further by making components statistically independent, not just uncorrelated; good for separating mixed sources (e.g. natural scenes).
differ mainly in how far they go: PCA decorrelates linearly, ZCA also whitens but preserves structure, and ICA removes higher-order dependencies.
At what stage of the primate visual system can orientation-selective neurons be found?
in the primary visual cortex (V1)
What does “tuning” mean in the context of neurons in sensory systems?
= neuron's selective response to a specific range or type of stimulus (e.g. orientation, frequency, location, or direction)
-> A neuron is "tuned" to the stimulus that elicits its strongest response
What is population coding?
—
= way the brain represents information by combining the activity of many neurons, rather than relying on a single neuron
-> Each neuron contributes a part of the information based on its tuning curve, and the overall pattern across the population encodes the stimulus.
Which parameters determine coding precision in a population code?
Coding precision in a population code depends mainly on:
Number of neurons involved:
-> the more neurons, the higher the precision.
Tuning curve sharpness
-> narrower tuning means neurons respond more selectively, improving accuracy.
Neuronal variability (noise)
-> less noise leads to more reliable signals.
Correlation between neurons
-> lower correlations typically increase coding efficiency.
How can biases arise in a population code?
Non-uniform distribution of tuning preferences, where neurons are unevenly tuned across stimulus space.
Noise and variability in neuronal responses skewing the decoded estimate.
Correlations among neurons that distort the combined signal.
Inaccurate or incomplete decoding models that fail to capture true neural dynamics.
Give examples of population coding in
a) striate cortex (primary visual cortex)
and b) extrastriat
a) In the striate cortex (V1), population coding occurs in the combined activity of orientation-selective neurons, where the pattern of firing across many neurons represents the orientation of edges in the visual scene.
b) In extrastriate cortex (like area MT), populations of direction-selective neurons encode the overall motion direction of objects by integrating signals from V1 and other areas.
What percentage of cortex, roughly, is concerned with processing of visual information?
20-30%
What does receptive field remapping mean and where is it observed?
= rapid shift of a neuron's receptive field to a new location in anticipation of an eye movement (saccade).
-> It helps maintain stable visual perception despite frequent gaze shifts.
-> observed in areas like the parietal cortex and frontal eye fields.
What time scales are related to circadian timing?
24h time frame (night-day)
Name the brain structure that is crucially involved in circadian timing in mammals.
suprachiasmatic nucleus in the hypothalamus
Which duration ranges are relevant to interval timing?
hundreds of milliseconds to several minutes
Name psychophysical tasks for interval timing and explain briefly
Examples:
Temporal Production: The subject generates a specific time interval (e.g., pressing a button for 10 seconds).
Temporal Reproduction: The subject observes a time interval and then tries to reproduce it.
Temporal Discrimination: The subject judges which of two intervals is longer.
Temporal Estimation: The subject estimates the duration of a presented interval verbally or by other means.
What is the regression effect in magnitude estimation?
= tendency of people to overestimate small stimuli and underestimate large stimuli, causing their responses to regress toward the mean of the presented range.
This bias reflects imperfect scaling of perceived magnitude.
The likelihood of measurement M given a stimulus S is described by the conditional probability P(M | S).
How could you determine P(S | M)?
Name the method you are using and give its mathematical formulation.
using Bayes’ theorem:
where:
P(M|S) = likelihood of measurement given stimulus,
P(S) = prior probability of the stimulus,
P(M) = marginal probability of the measurement.
This method is called Bayesian inference.
What are population clocks?
= neural mechanisms where the time is represented by the evolving activity pattern across a population of neurons.
(Instead of relying on a single pacemaker, timing emerges from the dynamic, time-dependent changes in population activity, allowing the brain to track intervals and temporal sequences.)
Describe the components of a state-dependent network
A state-dependent network consists of:
Neurons with intrinsic dynamics, whose current activity depends on their previous state.
Recurrent connections, allowing feedback and interaction among neurons.
External inputs that drive the network and interact with its internal state.
Together, these components create time-dependent patterns of activity that encode information based on the network’s evolving state.
How can a state-dependent network be used to explain time processing in the brain?
by encoding temporal information in its changing internal activity patterns.
As the network evolves after a stimulus, different network states correspond to different elapsed times, allowing the brain to read out time intervals from these dynamic activity trajectories without needing a dedicated clock.
Name one advantage and one disadvantage of working with human subjects in neuroscience
Possible advantages:
the ability to test for unique cognitive effects – certain things cannot be tested well in animals, example Numerosity, or speech.
training and communication – training is faster because one can communicate with the human subject and describe the desired task, this cannot be done in animals. In humans it avoids overtraining effects.
The whole brain – with the techniques that we have, we can measure at a mid-level of spatial and temporal scale brain activity patterns across the entire brain.
Possible disadvantages:
The measured signal is not a direct reflection of neural activity, but an indirect reflection of neural activity. This is particularly problematic for comparisons across different populations where a change in the vascular response would be expected (e.g. age).
The typical methods for measuring neural activity in animals are unethical in humans,
The methods are costly and time intensive.
The temporal and spatial resolution is to coarse to see individual neurons or circuits.
Name two methods for measuring human brain activity other than MRI, and their temporal and spatial resolution
PET: spatial resolution on the order of 5-10 mm, and temporal resolution of minutes to hours
EEG: spatial resolution on the order of centimeters and temporal resolution of muss less than a ms
Single cell recordings: high spatial (μm) and temporal resolution (<< ms)
Here are two neuroscience research questions. One can be addressed with human MRI, one cannot. Please label which question CANNOT be tested with human MRI and say in your own words why not.
Does area X of the brain contain enough information to decode task 1 from task 2?
Are the connections between two neurons excitatory or inhibitory in nature?
The second question cannot be addressed with human MRI because the spatial scale of the measurements (mm) is too coarse to be able to resolve a single neuron and the fMRI signal cannot resolve inhibitory and excitatory activity
Write out the Larmor equation and describe it in your own words what the equation equates
State what it is used for in MRI?
The Larmor equation:
-> ω0 = larmor frequency of a particle (resonant frequency)
-> ɣ = particle’s gyromagnetic ratio (ratio between the particle’s magnetic moment to its angular momentum)
-> B0 = strength of the magnetic field,
Relevance for MRI:
-> allows precise tuning of radiofrequency pulses to match the resonance frequency of hydrogen nuclei in tissue
-> This is how the MRI system selectively excites and detects signals from specific regions, enabling spatial encoding and image formation.
What are the three electromagnetic components required to produce an image in MRI?
If that was too easy, can you state what each of the three components is used for?
A strong static magnetic field: required to set up the atomic particles to receive energy
A radio-frequency pulse (from a radio transmission coil): sending energy into the system at the particle’s resonant frequency to be measured.
Magnetic gradients in the 3 spatial planes: turned on and off at specific times within the sequence to either select the exicted space, or encode the spatial dimension through the frequency or the signal phase.
Describe the signal that is measured with MRI and how the differences between brain tissue are measured?
= damped sinusoidal signal, also called a free induction decay (FID) signal, with a frequency that is centered on the Larmor frequency for hydrogen protons.
Differences in brain tissue are measured by tissue specific differences in the speed of decay of the signal (or recovery of the longitudinal magnetization) that are reflected in the brightness of the individual voxels of the image.
How is a specific plane of the brain (or body) selected in MRI?
A magnetic gradient, that is perpendicular to the 2D slice to be measured, is turned on at the same time as the radio frequency pulse is given. This changes the strength of the main magnetic field along the direction of the 2D slice. Then only those protons in the 2D plane where the Larmor frequency (based on the static magnetic field strength) matches the given radio frequency will be excited.
What is the signal called that is measured with functional MRI?
BLOOD OXYGEN LEVEL DEPENDENT (BOLD) signal.
Describe how neuronal activity leads to a change in the fMRI signal that is then measured in functional MRI (fMRI)
Neuron activation
-> requiring energy from oxygen in blood
-> dilation of local blood vessels
-> more blood is send to that area than needed
-> increased amount of oygenated hemoglobin with less magnetic property than deoxygenated hemoglobin
-> MRI signals gets stronger in that region
-> change measured in fMRI
in longer:
(When neurons become active, they need energy, which requires oxygen. Oxygen is delivered by the blood. In response to this activity, local blood vessels dilate, sending more oxygen-rich blood to the area than is actually needed. This increases the amount of oxygenated hemoglobin, which is less magnetic than deoxygenated hemoglobin. As a result, the MRI signal becomes stronger (brighter) in that region. This change is what fMRI measures, and the system gradually returns to baseline after the activity ends.)
What evidence is there that neuronal activity is related to the fMRI signal?
Study showed: both single-neuron activity and fMRI signals in the visual cortex of monkeys.
-> When the animals saw a checkerboard pattern, the same brain area showed both neuron firing and increased fMRI (BOLD) signal.
Further analysis revealed that the fMRI signal matched best with local field potentials (LFPs), which reflect local synaptic input rather than long-distance action potentials. This suggests fMRI primarily reflects input and processing in a brain region, not just output.
Describe the experiment that was used to determine that the BOLD signal measured with fMRI is a linear signal.
What benefit does the linear system have for fMRI research?
In the experiment, researchers presented a short stimulus once, twice, and three times, with short pauses in between. They subtracted the BOLD signal for one stimulus from that of two, and then that from three, and aligned the resulting curves. The curves looked nearly identical, showing that BOLD responses add up linearly, even if they overlap in time.
This linearity means we can use a single, standard BOLD response (called the impulse response function) to predict the brain's response to any stimulus pattern.
-> simplifies fMRI data analysis and modeling.
Describe the shape and timing of the hemodynamic response function (HRF) or the fMRI response to a short stimulus.
How is it typically modeled?
How is it implemented in fMRI analyses?
The hemodynamic response function (HRF) rises and peaks around 6 seconds, then falls and dips slightly below baseline around 12 seconds before returning to normal.
It’s typically modeled as the difference of two gamma functions—one for the rise, one for the undershoot.
In fMRI analysis, this HRF is used as an impulse response function, and researchers convolve it with the stimulus timing to predict the expected BOLD signal.
Can we measure regional brain activity when we are at rest?
How is this activity measured?
What is the assumption made here?
What is the name of the most common network found at rest?
Yes, we can measure regional brain activity when we are at rest.
This activity is measured by examining how the time courses of brain activity correlate between different voxels.
The assumption made here is that the correlated brain activity between regions means the regions are connected, and not for instance, that some external stimulus caused this activity.
The most common set of brain regions (or network) found at rest is the Default Mode Network.
What is the basis of spike sorting of extracellular waveshapes in multichannel extracellular recording?
= each neuron produces a distinctive extracellular waveform (shape and timing) across multiple recording channels, from which features spikes from different neurons can be classified and separated, allowing identification of individual neurons' activity from the mixed signals.
What is the role of inhibitory neurons in generation of the oscillations?
They synchronize the activity of excitatory neurons creating rhythmic patterns
(-> regulates the timing and pacing of neural firing, creating rhythmic patterns that underlie oscillations like gamma and theta rhythms essential for information processing.)
Describe the concept of remapping of hippocampal place cells
= Hippocampal place cells change their firing in new or altered environments, reorganizing the spatial map to flexibly represent different contexts.
What type of dipole does excitatory synaptic input to the distal dendrite of pyramidal cell give rise to?
Excitatory synaptic input to the distal dendrite of a pyramidal cell generates a current dipole with a sink at the dendrite (where positive ions enter) and a source near the soma (return current).
-> creates a current dipole polarity oriented from the dendrites toward the soma.
What is the role of hippocampal sharpwave/ripples in memory consolidation?
= reactivation and replay of neural activity patterns from prior experiences during rest or sleep.
-> helps transfer memories from the hippocampus to the cortex, strengthening and stabilizing long-term memory storage.
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