[Sequence] Neuroanatomy

Introductory Posts

Particular Anatomical Structures

Organizational Principles

Related Sequences

Brain Ontogeny: From The Tube

In 1969, H. Chandler Elliott said

Every brain system grows logically from the tube.

He was right. Today, we’ll learn why.

Primary Neurulation

A blastocyst has no brain. To correct this unfortunate situation, every vertebrate genome contains instruction for constructing a neural tube. This structure emerges via folding:

Brain Ontogeny- Primary Neurulation
This process is called primary neurulation. The resultant embryonic structures become the following systems in adults:

  • Neural crest cells evolves into the peripheral nervous system (i.e., nerves gathering information from your muscles, skin, and organs).
  • Neural tube cells evolves into the central nervous system (i.e., your brain and spinal nerves).
  • The neural canal evolves into the ventricular system (i.e., the hollow cavities in your brain filled with cerebrospinal fluid).

In the human embryo, primary neurulation is completed in the fourth week. It turns out this developmental milestone is relatively more prone to error than others – 1 in 500 embryos suffer from neural tube defects.

Vesicle Differentiation

The neural tube does not stay uniformly shaped for long: soon, three bumps (vesicles) emerge.  These vesicles appear at three weeks – even before primary neurulation is complete! Informally, these bumps comprise forebrain, midbrain, and hindbrain. Two weeks later, the human neural tube has differentiated into five vesicles.

Brain Ontogeny- Vesicle Differentiation

“Encephalon” means brain.

Vesicle-Structure Maps

Developmentally, the neural tube can be divided into five regions: tel-, di-, mes-, met-, and myel- encephalon.

Structurally, the central nervous system can be divided into three structures: cerebrum, cerebellum, and spinal cord.

We can usefully combine these perspectives:

Brain Ontogeny- Vesicle-Structure Overview

The brainstem and cerebellum are highly conserved across species. Most biological innovation is driven through the cerebrum (telencephalon).

Here is an anatomical view of the same relationships:

Brain Ontogeny- Vesicle-Structure Anatomy

Finally, here is a topological view of these same vesicle-structure maps:

Brain Ontogeny- Vesicle-Structure Topology (1)

 

You may notice how this graphic coheres well with my post on cranial nerves.

From this level of abstraction, we are better equipped to create structure-function maps, also known as neural architectures. To be continued.

[Graphic] Cranial Nerves

The brain receives information from three distinct sources:

  1. The endocrine axes.
  2. The spinal nerves
  3. The cranial nerves

This infographic explores the third source of information.

Loosely, cranial nerves exist because sensory information coming from the head doesn’t make much sense coming through the spinal cord.

Cranial Nerve Infographic (4)

Note that PSNS stands for Para-Sympathetic Nervous System (a branch of the autonomic nervous system).

 

Towards Body Architecture [Architecture v1.5]

Body Architecture v1.5

The autobiographical origin of this post is rather involved:

  • First, I wanted to understand the medial prefrontal cortex generated emotion.
  • Next, I wanted to understand how affective neuroscience interacted with the endocrine system.
  • Next, I wanted to understand the interactions between nervous & endocrine systems more generally.
  • Finally, I wanted to understand how nervous & endocrine systems together regulate the organism as a whole.

At the end of this adventure, the following concepts stand out to me as particularly significant:

  • Thirteen pairs of cranial nerves (CN) complement information derived from the spinal nerves.
  • The HP (hypothalamus-pituitary) axes represent neuroendocrine systems, such as the HPA axis (stress), HPT axis (metabolism) and HPG axis (sexuality). These are how the central nervous system drives the endocrine system, despite the blood-brain barrier.
  • The autonomic nervous system (i.e., the sympathetic & parasympathetic branches) intertwines with the endocrine system to maintain homeostasis.
  • The enteric nervous system (500 million neurons within the gut) is connected to the brain via the vagal nerve (CN 10), but can operate independently.

A few general musings to close:

  • Within anatomy, the various bodily systems feel much more studied than their interactions.
  • The endocrine and nervous systems seem to control the body at different time scales: nerve control is immediate, hormonal control is long-lasting.
  • The distinction between autonomic nervous system & autonomic endocrine system could be used to stiffen a conceptual divide between emotion and mood.
  • This process of consolidation of anatomical knowledge has really emphasized, for me, how the neocortex is not the only actor on the stage of my architectural research.

To be continued. This was fun!

Neural Roots Of Mind [Architecture v1.5]

Part Of: [Mental Architecture v1.5] sequence

Overview

This data collection effort will become larger over the coming months. Eventually, I hope to break this up into a more formal knowledge representation system. It would be nice to create (inter-referencing, version-controlled) tables & software models containing:

  • Behavior Explananda Database: Linking modules or modular circuits to different behaviors.
  • Lesion Explanada Database: Counting how many syndromes can be explained on the current architecture.
  • Confidence Distribution Model: Formalizing my Bayesian intuitions on the architecture, and linking to various streams of evidence & constraint.
  • Cortical Maps: Displaying localization information topographically, and noticing areas where localizations overlap (logically possible, but worthy of further research)

Implementation Notes

This document is not close to representative of my entire research project, but it is better to have at least something written down.

Finally, please note that Schema Memory, Semantic Memory, and Episodic Memory are not represented in what follows. These memory systems each comprise their own representational format, but following the principle that “modules tend to save their work near where they are computed”.

Without further ado, here is my December 2015 mapping between my mental architecture, and the brain.

Modular Localizations

The following are cognitive networks (or streams, or circuits) which have been fully or partially localized. Here lie 17 distinct networks.

  1. The Action Instigator stream is responsible for activating and deactivating motor activity. It corresponds with the task-positive network and task-negative network.
  2. The Face Computation network, which includes the Face Detector, Facial Identity Lookup, and Face Expression modules.
  3. The Body Detector network, which includes Body Part Detector and Whole Body Detector modules.
  4. The Attention (Top-Down) network, which includes the Preparatory Set circuit, and the Task Switching circuit.
    • Localization: The Preparatory Set circuit operates in cingulo opercular PFC.
    • Localization: The Task Switching circuit operates in fronto-parietal PFC. 
    • Theory Basis: I am leveraging A dual-networks architecture of top-down control (Dosenbach et al, 2008)
  5. The Attention (Bottom-Up) network comprises the Salience Computation circuit and the Salience Orientation circuit.
    • Localization: The Salience Computation circuit operates in the dorsal frontal lobe. This is the home of the salience map. 
    • Localization: The Salience Orientation circuit operates in ventral frontal lobe, including Frontal Eye Fields, and is strongly right-lateralized. 
    • Theory Basis: I am leveraging The Attention System of the Human Brain: 20 Years After (Petersen and Posner, 2012).
  6. The Attention (Arousal) network comprises the top-down attention network.
  7. The Audio (Spatial) stream, which includes a left-lateralized Language Production module. 
  8. The Audio (Perceptual) stream, which includes the Voice Recognition and left-lateralized Language Comprehension modules.
    1. Theory Basis: I am leveraging The Cortical Organization Of Speech Processing (Hickok and Peoppel, 2007)
  9. The Event Analysis complex, which comprises a Biological Motion and a Causal Inference module. 
    • Localization: The Biological Motion module operates in the Superior Temporal Sulcus.
    • Localization: The Causal Inference module, whose location is unknown…
  10. The Global Broadcasting network uses the Thalamus as a relay station to construct a global workspace.
  11. The Intimacy network, which comprises an Attachment and an Attraction module 
    • Localization: the Attachment module operates in the  location is the “senior executive of the emotional brain”; the Orbitofrontal Prefrontal Cortex.
    • Localization: The Attraction module, whose location is unknown…
  12. The Location Detector network, which comprises a Place Inference and a Landmark Inference module.
  13. The Memory Metacognition network, including the Recognition Heuristic and Confidence Heuristic modules. 
  14. The Smell stream begins with the Olfactory Bulb. It is the only sense organ that is not first routed into the Thalamus.
  15. The Taste stream begins with the Gustatory Cortex.
  16. The Vision (Action) stream (which travels dorsally towards the parietal lobe) also begins with the Primary Visual Cortex.
  17. The Vision (Perception) stream (which travels ventrally towards the temporal lobe) begins with the Primary Visual Cortex.

The following are cognitive networks which I have not yet localized. Here lie 4 distinct networks.

  1. The Folk Mechanics network, which computes expectancies based on, e.g., the continuity of motion and other physics principles
  2. The Folk Math network, which includes the Subitizing module.
  3. The Identity Maintenance network, which includes the Identity Constructor, Identity Evaluator, and Identity Stabilizer modules.
  4. The Reputation complex, including the Caregiving module (partially mediated by oxytocin) and the Appreciation module (its functional sibling).

The following are cognitive modules which have been fully or partially localized. Here lie 14 distinct modules.

  1. The Action Engine module operates in the Primary Motor Cortex.
  2. The Action Evaluator operates in the Premotor Cortex.
  3. The Action Planner module operates in the Supplementary Motor Area.
  4. The Agent Classifier module, which links to the Amygdala…
  5. The Embodied Self module operates in the Primary Somatosensory Cortex.
  6. The Memory Heuristic (Effort) module, which constructs the Feeling Of Effort, lives somewhere in the Prefrontal Cortex (PFC),
  7. The Homeostasis Regulator operates in the Hypothalamus.
  8. The Pavlovian Conditioning module operates in the Posterior Parietal Cortex.
  9. The Memory Constructor (Frame) module, which lives somewhere in the Prefrontal Cortex (PFC) …
  10. The Memory Constructor (Episodic) module operates in the Hippocampus.
  11. The Memory Constructor (Semantic) , which may be near the Lateral Occipital (LO) Complex…
  12. The Normative Centroid module, which lives somewhere in the Ventro-Medial Prefrontal Cortex, and activates the Amygdala.
  13. The Conflict Monitor module, which lives in the dorsal Anterior Cingulate Cortex (dACC).
  14. The Intuition Override module, which operates in the right Ventrolateral Prefrontal Cortex (rVPC).

The following are cognitive modules which I have not localized. Here lie 9 distinct modules.

  1. The Counterfactual Simulator module
    1. Theory Basis: this module is adapted from Alan Leslie’s work, and Mindreading (Nichols and Stich, 2003).
  2. The Lie Detection module
    1. Theory Basis: Default Credulity theory, as defined by Gilbert (1991) How Mental Systems Believe.
  3. The Metaphor Linker module
  4. The Mind Reading module
  5. The Moral Rigidity module
    1. Theory Basis: Sacred Value Protection Model. Thinking the unthinkable: sacred values and taboo cognitions (Tetlock, 2003)
  6. The Social Processing (Fairness) module.
  7. The Social Processing (Exclusion) module
  8. The Social Processing (Authority) module
  9. The Symbolic Marker module

 

Cerebral Cortex: A Cartographic Approach

Part Of: Neuroanatomy sequence
Followup To: The Thalamocortical Plasma Globe
Content Summary: 1100 words, 11 min read

Cortical Area & The Obstetric Dilemma

Last time, we learned that the brain is organized like a plasma globe: a sphere within a sphere. Today, we’ll be exploring a technique for reasoning about the cerebral cortex, or “outer sphere”. A few things you should know about this organ:

  • It weighs about a pound.
  • Stretched flat, it would cover an area of about 2.5 square feet.
  • It is about six millimeters thick.
  • It houses 20 billion neurons.

Does your neocortex have the most neurons? No, that title goes to the cerebellum, whose 100 billion neurons coordinates complex movements. But your brain must do a lot more than motor fine-tuning. Your brain perceives its environment, identifies objects, sets goals, makes decisions, feels emotions, and experiences consciousness. Where does your brain perform these tasks?  Primarily in the neocortex! Loosely speaking, the outer sphere does much of the “heavy lifting” for your brain.

Why is the human brain so wrinkled? After all, not all species have brains with this shape:

Cytoarchitecture- Obstetric Dilemma
Consider the following evolutionary pressures, together known as the obstetric dilemma.

  • A bigger neocortex does more work, and homo sapiens make its living (ecologically speaking) by intelligence. Thus, natural selection will select towards increased cortical area.
  • Compared to other animals, human childbirth is a uniquely dangerous affair. Why? Brain size. Thus, natural selection will select away from increased brain volume.

As any microchip designer can tell you, wrinkles are a way to increase surface area, while holding volume constant!

Flattening The Lobes

And now, a short story. Cognitive neuroscience is rife with Latin terms for cortical areas. There are hundreds of them: “anterior cingulate cortex”, “fusiform gyrus”, “temporo-parietal junction”, etc. Over the past few years, as I consumed more of the field, I had slowly acclimated to hypotheses regarding the functions and interrelations for such areas. But given my lack of robust anatomical intuitions, these Latin names were just linguistic markers; I lacked an appreciation for geography.

While 3D models of different locations were mildly helpful, the pieces really fell into place when I discovered cortical flat maps.

While the neocortex is like a sheet, in some ways it is more accurate to imagine two sheets. Most people know that the brain has two hemispheres, but fewer know that these hemisphere’s are not (directly) connected? The two halves of your brain do talk to one another, but via a subterranean tunnel known as the corpus callosum.

Okay, here it is: a flat map of the human brain.

Cytoarchitecture- Lobes (2)

The brain is often divided into four to six different sections, or lobes. My map colors each lobe, illustrating the relationships between 3D brain and 2D map.

Here are my five lobes:

  1. Occipital (red, back of the brain). This lobe is primarily responsible for vision.
  2. Parietal (yellow, top of the brain). This lobe is primarily responsible for touch.
  3. Temporal (green, sides of the brain). This lobe is primarily responsible for concepts.
  4. Frontal Lobe (blue, front of the brain). This lobe is primarily responsible for personality.
  5. Limbic Lobe (purple, underneath the brain). This lobe is primarily responsible for emotion.

Flat maps are an underappreciated resource. But I’ll return to this point some other day. In the meantime, I should mention that these flat maps are imprecise elaborations of the flat maps created by the Gallant Lab. That is, my creations strike me a bit like this:

Cytoarchitecture- Metaphor To Ancient Cartography

I am planning to ultimately construct more precise flat maps using tools like Freesurfer.  But in the meantime, I’ll stick with my quick-and-easy 16th century cartographic approach. 🙂 

Primary vs Association Cortex

Let’s turn now to perception. In Tunneling Into The Soup, we discussed how sense organs (e.g., the rods and cones in your eye) translate the physical reality outside your body into neuron-compatible signals that your brain consumes. To make this intuitively compelling, I employed the following metaphor:

Perceptual Tunneling

Your eye transmits data to your brain via information highways known as the optic nerves. The traffic of these highways – sense data – captures only a subset of physical reality; call this subset an umwelt. Such highways drain their contents onto landing sites on the cortex known as primary areas.

Aristotle is wrong: you have more than five senses. Balance and body alignment are senses in just the same way as sight, hearing, touch, taste, and smell. I am currently aware of nine different types of sense, or sensory modalities.  But of these, three sense modalities are particularly important (and consume more space!) in the human brain. They are:

  1. Body sense-data. This area includes touch, and other senses associated with body sensation and position.
  2. Visual sense-data. Nerves that leave your eyes deliver their data here.
  3. Audio sense-data. Nerves that leave your ears deliver their data here.

These three primary areas of your cortex are marked in dark green:

Cytoarchitecture- Lobes + Primary Cortex (1)

Non-primary areas of the cortex (areas that are not dumping grounds) are called association areas. Association cortex is associated with two distinct functions:

  1. Non-modal computations like goal generation.
  2. Multi-modal computations such as “hand-eye coordination”.

Evolutionary Considerations

How does the human brain compare to that of other species? Consider again the mouse brain. It is obviously smaller than the human variety, but is the ratio of primary-vs-association cortex preserved?

It turns out that the answer is no. Because primary sensory processing is more immediately useful to survival, the neocortex of the mouse is actually dominated (>50%) by primary areas.

This biological fact reminds me of the (amusing) Expanding Earth conspiracy theory. In contrast to plate tectonics, the Expanding Earth claim is that continent size has remained constant, whereas the oceans have been expanding. Despite being complete bunk, it does provide a colorful metaphor to our genetic distinctions from the mouse. Our species has invested more heavily in association cortex (oceans) than primary cortex (continents).

Cytoarchitecture- Expanding Association Cortex (1)

Have humans invested equally in every part of the association cortex? No: the hominid line shows pronounced (6x) gains in the prefrontal cortex, and comparatively restrained growth elsewhere. We thus have reason to believe that most uniquely human behaviors must be supported by the prefrontal cortex. To be continued.

Takeaways

  • The outer sphere of your neural plasma globe – your neocortex – is responsible for nearly all of your higher thought.
  • Your neocortex covers an area of about 2.5 feet. Flat maps of the neocortex allow us to intuitively visualize the brain.
  • Your neocortex contains primary (direct sensory input) and association (abstract computations) areas.
  • Humans have more association areas, especially in the prefrontal cortex. This hints at a basis for our unique abilities.

v1.5: The Tripartite Mind

Part Of: [Cognitive Architecture] sequence
Followup To: [Mental Architecture v1.4]

Let’s review our theoretical trajectory.

The Autonomic Mind: Belief, Motivation, and Decision Making

Our first architectures (1.0 → 1.2) explored the autonomic mind, which comprises our most fundamental mental capacities.

The Algorithmic Mind: Attention, Consciousness, and Intelligence

Subsequent architectures (1.3 → 1.4) complemented this understanding with the algorithmic mind, by weaving together three theories:

  • The Global Workspace theory of consciousness (inspired by Baars).
  • The Interpretive Sensory Access theory of introspection (inspired by Carruthers).
  • The Emergent Working Memory theory of attention (inspired by Postle).

There is a close link between intelligence and consciousness, as evidenced by working memory’s strong correlation with both. Generally fluid intelligence (IQ) is essentially a measure of the precision of your attentional streams.

The Reflective Mind: Metacognition, Control, and Culture

The novel innovation of this architecture (1.5) could be entitled Prefrontal Cortex: The Final Frontier. It integrates two theories into the base corpus:

  • The Reflective Modulation theory of cognitive override (inspired by Stanovitch).
  • The Somatic Frame theory of culture (inspired by Damasio).

These five theories together constitute the foundation stones of my mental architecture. Let me call this synthesis the Attention-Modulated Tripartite Mind theory.

Putting Clothes On My Theory

A theory is a house, and the above merely represents its foundation. On this base, I will add details: the following posts are planned:

  • Towards Architectural Phylogeny. Explaining mental evolution across species is an important requirement for any mental architecture.
  • Towards Brain Architecture. If the brain implements the mind, then we should be able to localize mental software packages to their respective neural locations.
  • Towards Body Architecture. The brain’s computational powers serve the needs of an organism. Mental and brain architectures must be rooted in the concerns of anatomy and ecology.

The Thalamocortical Plasma Globe

Part Of: Neuroanatomy sequence
Content Summary: 400 words, 4 min read

Today, we embark on a (very) brief tour inside your head!

How Neurons Work

What is a brain? A brain is a collection of 120 billion neurons.

What is a neuron? A neuron is simply a cell. Since you are a eukaryote, all cells in your body – neuron cells, skin cells, etc – have a lot in common, including:

  • cell body (environmental barrier)
  • nucleus (genome storage)
  • mitochondrion (energy production)
  • endoplasmic reticulum (protein production)

In addition to these shared features, specialized tendrils (dendrites and axons) set neurons apart, giving them their web-like shape.

Brain- Neuron (1)

Neurons are useless individually. But when chained together, they may transmit pulses of electrochemical energy known as spike trains. input dendrites receive such signals and – sometimes! – fire, pushing the outgoing signal through the output axon. The output tendrils of a neuron connect with the input tendrils of other neurons; these connections are called synapses. 

In this way, neurons mediate a relationship between input and output signals. What other kinds of things do this? Mathematical devices known as functions, and electrical devices known as transistors.

Gray Matter vs White Matter

Let’s zoom out to consider the entire brain. A brain has two symmetric halves (the hemispheres) connected by a bridge (the corpus callosum). Here’s what it looks like from the inside:

Brain- Gray vs White Matter

Certain parts of the brain are gray, and others are white. Why? What happens when you put these under a microscope?

  • Gray matter turn out to be dense clumps of neuronal cells.
  • White matter aren’t made of whole neurons at all, but axons coated in a fatty substance (myelin). These tissues accelerates neural signals.

If white matter doesn’t process information, why is there so much of it?

The Neural Plasma Globe

To answer this question, we turn to the gray matter. Where does it live? In only two places: the wrinkly surface of the brain (the cerebral cortex), and in an evolutionarily ancient part of the brain (the thalamus, and neighboring midbrain structures). Think: spheres within spheres. 

We can see now why there is so much white matter. White matter comprises information highways which transport information to and from different areas of the cerebral cortex. Some of these highways directly connect cortical regions, but much travels through the thalamic “central hub”. We will call such highways thalamocortical radiations.

With tractography (a sister technology to the MRI scan), we can now directly visualize such radiations. There is something almost poetic about these highways… you could say that the brain is like a plasma globe.

Brain- Plasma Globe analogy (1)

 

 

Next time, we’ll explore some implications of this metaphor.

Mental Architecture v1.4

Part Of: [Mental Architecture] project
Followup To: [Mental Architecture v1.3]

 

Less of a jump than v1.3, but still a worthwhile place to “save my work”.  Significant new features include:

  • Consolidation Of Working Memory
  • Improved Sensory Specificity
  • A Home For Emotional Processing
  • Explicit Brain Architecture

Consolidation Of Working Memory

Adopts the Alternative View, contra the “Standard Model”, as put forward by Postle (2006) Working Memory As An Emergent Property of the Mind and Brain. Abolishes the Visuo-spatial Sketchpad and the Phonological Loop and replaces it w/ the contents of perceptual memory (and semantic memory, etc).

Improved Sensory Specificity

Reintroduces some of the sensory details already present in Mental Architecture v1.0, and adds a few organizing distinctions (e.g., exteroceptive vs interoceptive)

A Home For Emotional Processing

For months, my Emotion Generator module has lacked a comfortable home. Now, following Damasio’s gesture at homeostasis, I finally have a home for the “big six” emotions (Anger, Disgust, Fear, Joy, Sadness, Surprise). Specifically, I conjecture that these emotions are produced in topographic map located in the middle of a processing stream, whose data flows from interoceptive (e.g., hormonal) sense organs.

Explicit Brain Architecture

  • The Vision processing stream starts with Area V1 at the back of the head.
  • The Audio processing stream includes six tonotopic maps within the primary auditory cortex.
  • The Temporal Indexing module, which serves episodic memory creation, is performed by the hippocampus.
  • The Salience Attention modules comprises the bottom-up attention network.
  • The Directed Attention module comprises the top-down attention network.
  • The Central Executive module is housed in the pre-frontal cortex (PFC).
  • The Embodied Self is the Sensory Neuromatrix which is the cortical homunculus.
  • The Action Instigator module is modulated by the direct & indirect pathways of the basal ganglia.
  • The Action Engine is the Motor Neuromatrix which is the primary motor cortex.
  • The Action Evaluator is the premotor cortex.
  • The Action Planner module is the Supplementary Motor Area (SMA).
  • The Pavlovian Conditioning module is the posterior parietal cortex.
  • The Agent Detector module includes the Face Detection submodule which is operated by part of the fusiform gyrus.
  • The Favoritism Engine module is partially mediated by oxytocin.

Administrivia

Module Changelog

  • Removal: the Sensory Slave Modules modules b/c these are none other than perceptual systems.
  • Removal: the Modality Integration module from episodic memory, as this ability is provided by the two attentional networks.
  • Edit: relocating the Emotion Generators module from episodic memory & into interoceptive sensory processing.
  • Addition: Reflex Conditioning modules which bypassing knowledge queries en route to making hurried decisions.
  • Addition: Homeostasis Regulator modules which drive the brain’s endocrine system output.
  • Addition: a Body Ownership module within the sensory neuromatrix. The thing responsible for e.g., the rubber hand illusion.

Open Tasks

  • Incorporate Feature Integration Theory.
  • Incorporate the Somatic Marker Hypothesis.

Open Questions

  • What are the implementation details of semantic memory “files”, and frame memory “folders”?
  • Is lexical memory distinct from semantic memory?

Frustratingly Unaddressed Human Explananda

  • Music
  • Humor