Neurology, brainstem
Introduction "Brainstem" is the usual collective name for
the most basic elements of the brain, sometimes including the spinal cord
and everything in its immediate vicinity - but usually "brainstem" refers
only to the parts between the emotional brain in the center and the spinal
cord. Spinal cord and brainstem perform the most basic control functions of
the body, as evidenced by the fact that damage to it is usually severe, very
difficult or not repairable, and as far as the brainstem is concerned, mostly
fatal.
The standard descriptions of the brainstem begin with the parts most important
to the rest of the brain, which are on the top of it. In this description the
evolutionary view on things is used, together with the goal to show how
neurology works for the individual experiencing everyday life. Which
automatically means that you have to start from scratch, i.e. with zero
assumption about previous knowledge and at the bottom
of the structural organization. Just like it is done in nature.
An attempt has
been made to alternate between technical details versus why things are as
they are, based on evolutionary arguments and what seems plausible
practically.
Of course, a single web page can
only give the main aspects, but it is attempted to give such information that
further reading into the matter is much less of problem that it
presently is, e.g. in view of the several naming conventions that are used
in standard literature - an overview of the latter can be found here
.
The external sources are denoted by brown arrows, most of them
referring to Wikipedia, being widely available and a natural
point for most laymen to start a search in these matters.
Origins
The spinal cord itself is a product of previous evolution that started
with the creation of connecting cells between perceptual organs and limbs,
becoming specialized into cells called "neurons", looking like a wire-with-switch:
The green part is the active cell body, with its dendrites, and the wire attached
(with its electrically isolating cover) is called the axon
(portrayed on the right is a muscle thread). In the famous (simplified) example, it is at one
end connected to sense organs in the knee and at the other end to muscles in
the leg. Tap the knee, and the leg stretches automatically (from here
):
In present-day reality, another neuron is connected to the sense organs
in the knee and fires when these are "touched" - that "firing" is a
electro-chemical pulse due to some self-amplifying process in the cell body,
and this pulse travels down the axon ...:
... towards another neuron it can excite (usually more than one
active neuron connection is necessary to fire a neuron).
Another important point to note is that neurons are not simply mechanically
attached to each other like electrical wires have to be. In fact, they look
like they are actually isolated from each other, with a small gap between
the end of the axon and the body or dendrite of the neighbouring neuron:
The contact isn't made by electricity, but by the flow of chemicals.
This has a highly important purpose: the contact through this gap, called
"synaps", can be influenced by other chemicals floating in the
neighbourhood. That is: the contact can be "modulated". One (in)famous
substance that does this "modulating" is nicotine. The stuff that keeps you
smoking while you don't want to any more - more on that later.
The
outwardly most important kind of chemical working here are called "neurotransmitters".
There are the ones acting inside the neuron, of which there are two kinds:
the already mentioned exciting ones, of which the most abundant is
glutamate:
Not at all very complicated ... Note: most illustrations concentrate on the inner structure of these
molecules - for its functions only the external shape and the associated
charge distribution counts - the chemical reactions it takes part in are of
the kind of "lock and key". There is also a variant that does the
reverse: blocking or "inhibiting" the neighbouring neuron. This is
necessary for more precise control: if a limb moves to fast, it has to be
slowed down quickly - not only mechanically but firstly by the controls.
This cannot wait until the exciting impulse has worn off - it has to be done
actively. This is the role of the inhibiting neurons and their
neurotransmitters, of which the most abundant one is GABA (it looks like
glutamate). Please
note that nature has a strong preference for this kind of "force and
counterforce" combination - they result in more stable equilibriums.
So these basic beginnings evolved into a system of movement that became more complicated during evolution, with limbs with
multiple articulations, the spinal cord also becoming more complicated, with
concentrations of neurons for mutual coordination and, most importantly,
more precise control. This control is regulated through a feedback process,
that is to say: there are also neurons in the muscles of the limbs that
report back tensions and positions - in neurology this is called the
"proprioscopic" information. The signals to the muscles are adjusted on the
basis of the signals about how far the limb has progressed in its planned
direction. This feedback and coordination is executed in conglomerates of
cells called ganglia ("knots"), strewn throughout the spinal cord.
In animals with multiple limbs, the limbs themselves also have to be
coordinated. In animals of the type of centipedes (more correctly: similarly
build far ancestors), this automatically
led to series of knots, which, in view of its importance, got hidden into
the vertebral column as what is now known as the spinal chord.
In
more evolved creatures, the demand on control grew bigger and bigger, until
finally it wasn't sustainable within the vertebral column, and the newer
bigger neuronal structures grew out above the spinal chord, and became what
is now called the brainstem.
The boundary between the present day
human spinal cord and brainstem is somewhat vague - where the spinal cord
mainly consists of long strands neuron wires, the axons, with regions with a
number of neuron nodes between them, the brain stem is dominated by more
cohesive structures referred to as "nuclei", with much wiring in between. In
the pictures below the boundary is therefore kept somewhat vague:
|
|
|
Spinal chord |
Brainstem |
The spinal cord provides the basic wiring and control functions for
movement and the autonomic body functions. Having such importance that it is
safely hidden in the spine.
Also visible are the area above it full of
parts of the emotion organs, and the many curls of the cortex that
surround all. |
The brainstem is too large for the spine, and therefore
protrudes above it. Where damage to the spinal cord can limit itself to
limbs, and to the cortex to mood changes if you get a bullet through it,
damage to the brainstem is almost always deadly. |
In the field of neurology, the functioning of the spinal
chord is one of the best-known areas, with e.g. twelve numbered nerve
bundles each with precisely known functions. The
associated parts of the spinal chord take care of the basic, say
"mechanical", functions of the body - damage to it leads to paralyzes,
but (mostly) not to loss of any higher bodily function. Together with
the brainstem, the two also provide the control of the basic "house holding"
functions the body, breathing, heart rate etc., see below for an overview
(an illustration of the site of Ben Best
):
Clearly visible even in this scaled down form is that the spinal cord also
provides the basic regulation of all other important organs. The parts of
the spinal cord that do the work are the ganglia, the smaller
conglomerates of neurons. This part of the
nervous system is called the (ortho-) sympathetic part standing for the
activating functions, left in the image. In addition, there is the
para-sympathetic part, right, driven by nuclei in the brainstem, for
regulating the functions at rest, such as digestion.
Of the twelve cranial nerves running to and from the organs, present in this picture
are the optic nerve (oculomotor nerve, III), the facial nerve (facial nerve,
VII) and the vagrant nerve (vagus, X - to various parts of the breast).
This part of the nervous system performs its functions independent of
the higher, conscious parts of the brain, and is therefore called the
autonomic nervous system.
So assuming that this all works well, you
have a functioning musculoskeletal system. Of limited use, if you do not
know where you want to go. So another thing is needed: an observational
system. But even with an observational plus motional system, it doesn't
make sense to develop it you don't know what to do with it. So from the most
primitive beginnings, there must have been some gain in it, in terms of
survival.
Which, by using these terms, almost gives away the answer:
the gain from the vary first stages of development consisted of the
possibility to flee the danger.
Which
raises the question "What is danger" , and how primitive life discovered it.
The answer to these questions is known from a context earlier than that of
the nervous system: "danger" is the presence of dead fellow species, and the
method to discover this is the sensing of the waste products of that death. In this
primitive world, the main danger is that of "being eaten", and being eaten
in the most primitive form is "bite by bite". This necessarily brings
residues of the eaten creature into the environment. These waste products
can be detect fellow species - that is a biochemical process, which is now called
"smell"
. (Incidentally, this also explains why some predator fish swallow
their prey at once, which is not favorable in terms of digestion - this is
to avoid detection). Discovering waste products of your own kind is an
excellent signal to go elsewhere, if you have a motional system. That
is why a motional and eventually musculoskeletal system once arose. Already
in single-celled
live forms, see the image above right. The simple motto was: "Get out!".
Destination irrelevant.
By the way: being touched was an equally bad
sign, at least in potentiality, so it is no wonder that smell and touch are the most
primitive senses an any living and moving creature. And
together with touch comes another one, but one that isn't as immediate as
the other two: heat. The sensing of all this, touch
and heat, is called the somatosensory system
. Its signals are
to humans known as "pain". In the specialized field of neurology, it is
called "nociception"
- the skin is
abundant with nociceptors, that detect mechanical, chemical or thermal
damage, and report to the spinal chord. At which location actions are taken
and signals relayed, that when arriving at the conscious level are called
"pain".
In a next stage it is also clearly advantageous if you know
where to go, if only not to go in the direction of the predator.
For which an image of the environment is a desirable asset. With all kinds
of intermediate steps that start with the ability to detect light and dark,
detect some direction in it, and so on, to the various forms of "eye" that
exist in the animal kingdom
.
Eyes are by far the most demanding observation system in terms of the nervous
system. If only because there are two of them, and the signals of the two
can be combined with great advantage into an image having depth. For which
it is necessary to do some heavy calculating in order to construct a single
image of reality. And when this goal has been
achieved, the musculoskeletal system must be given commands so as to guide
the whole correctly through tis virtual reality. A term used also because it signifies its demands, because modern computers are just recently
getting to grasp with this task. A task mainly performed inside the
brainstem.
So over to the description of its organization.
Mechanics
The following illustration gives the first global view
(front / rear view) of the brainstem - this and the other anatomical
engravings come from the atlas of Gray
- this one is Gray 690:
In these engravings the dotted structures are the nuclei, conglomerates of
neurons, and the striped ones are the bundles of neuron outputs or axons
that carry the large scale connections. Elsewhere in the brain the nuclei
are usually more clearly defined, and reasonably spherical, in the brainstem
this is less pronounced, giving rise to a series of names such as
nucleus, corpus, locus, formation and so on.
For axon bundles there is a similar "confusion" with terms such as
peduncle (Latin for "stem"), fasciculus ("bundle"),
lemniscus ("band") or just "fiber". Another term
that is dropped is that of 'decussation': this means that the associated
bundles cross over to the other side of the brainstem - the illustration
shows this happens a lot. This is for the overall coordination of the body.
This illustration
gives partly the view from outside, and partly some of the most prominent
internal structures. "Reading" from the bottom upwards, the first thing that
draws the attention from the outside, the brainstem being up to that point
just a boring pipe-like structure, is called the "olive", or when
focussing on function: the olivary nucleus.
This bottom part of the brainstem is called the "medulla oblongata"
of for short "medulla". It is where the first larger scale
coordinating structures are located. The next obvious
things are the two sketchy drawn protrusions. These are the connections to
the much larger cerebellum (not shown), and the point where the cerebellum
is attached to the brainstem - see further on. This
middle part of the brainstem is called the "pons" ("bridge"). One
also finds here the first "house holding" nuclei that regulate heart rate, breathing rate, etcetera, and the sources of the first two
modulating neurotransmitters. Above the pons but not really
visible comes the upper part of the brainstem, that traditionally is divided
in "tectum" and "tegmentum" ("house" and "roof"). The
tectum is where the secondary phase of the auditory and optic information
processing takes place. Especially the optic processing is important and
takes space, making the tectum quite prominent in primitive species.
The top is also the location where one finds the rest of the "house holding"
nuclei, and that of the sources of the second pair of modulating
neurotransmitters.
More insight into functional relations between all
these structures is given by more schematic drawings like the next one, a
view from behind (from here
):
Again: this has to be "read" from the bottom upwards.
So the
first larger structure is the olivary nucleus, (more correct: the "inferior",
i.e. the bottom and larger one of two) followed
by others like the cerebellum and the red nucleus, with
many smaller ones in between. The
red nucleus.is the last nucleus undisputedly within the brainstem.
The olivary nucleus is also the first one having a clearly laminated
structure, of the type generally called a "neural network", having
the folded look familiar from the cortex. The olivary nucleus probably does
the first larger-scale "mechanical" coordinating work. It is followed by the much larger
cerebellum, which is probably an averaging device: a mechanism that learns
from lots of previous behaviour the fine-control of present behaviour
.
The coordinating role of many of these structures can be surmised by the
presence of the vestibular nucleus, that is connected to the organ
of balance, in the ear. This information has to be combined with the optic
information coming from the eye, to correct the latter for the attitude of the head, in
order to get the correct horizon in the total picture of reality that is
going to be build. Which is taken one step further
with the red nucleus, that is in animals and generally associated
with "gait": the coordination of multiple limbs for the propagation of the
organism. And in human infants with early limb coordination.
Also shown on both illustrations is the
thalamus, the first structure that undisputedly lies outside of the
brainstem. Its present (main) role is that of a relay station between
spinal chord, brain stem and the rest of the brain further on: the emotion
organs and the cortex. This is discussed a shortly at the end of this article,
and in more detail in that on the emotion organs (the first ones of these
are the onion shaped things at the top of Gray 690, which are the
putamen and the caudate nucleus).
The next step here is
to take a more detailed look into the coordinating structures of the
brainstem itself. First an overview of the first large one, the olivary
nucleus and its direct surroundings, in a horizontal dissection (Gray 694):
The olivary nucleus is the first one with the structural form familiar
from the cortex, the cerebellum and many other ones, known for their look
with many folds and curls of what is essentially one sheet of tissue filled
with layers of neuron cell bodies, the "gray matter", with the rest of the
space filled by the axon outputs of them, having a slightly lighter outward
appearance and called "white matter".
This kind of structure falls
under a more general category called "neural network", which is almost
better known for its technical versions, as a type of software that learns
itself how to do things (more correctly: it taeches itself from a large
number of correct and incorrect examples it is fed). A first example being to recognize handwriting on
bank
statements. Of course this name was chosen at the time
because one knew that in neurology too, neural networks are the ones that do the learning. They have in common that they consist of layers of
"switches" or "logical gates" or biologically neurons, with layers of connections in
between these. With at least two layers: the input and the output.
So
let's have a look at a basic biological neural network - having the
following
general scheme:
Where the structure immediately tends towards a spherical shape because the layers
themselves tend to be more voluminous than the input and output bundles.
Which scheme immediately raises the question: why not do this work at
the location of where the input bundles came from?
That would almost certainly be indeed what would happen, so the application
of this kind of scheme lies in the cases where there are two or more input
bundles, with information that has to be coordinated.
And with these
thoughts in mind, lets look at the schematic representation of the
neighbourhood of the olivary nucleus, corresponding to the earlier
dissection (Gray 699, from here
):
With firstly a clear demonstration of what was earlier said about the
relation between left and right hand halves of the brainstem: the
information is frequently crossed over, in order to maintain overall
coordination
Also clearly visible are the multiple inputs and outputs
of the cerebellum, through what is called the inferior peduncles,
number 8. Also visible are the outputs of the olivary nucleus, number 4
(going to the cerebellum), but not its inputs, coming from the spinal chord,
below (due to clarity). Besides these connections,
there is also more detail on nuclei - represented are the gracilis,
cuneatus and cinerea nuclei as numbers 5, 6, en 7, that
relay the
proprioceptic (muscle feedback) and fine-touch information coming from the
spinal chord. They are also visible in the next overview
(a part of Gray 691),
which is a view from behind with some of the outer parts removed. This adds
the cochlear and vestubular nuclei: the information on
hearing and balance coming from the ear:
What is denoted by N.V. is the nervus vagus,
one of the twelf cranial nerves that connect brain to body - the vagus goes
towards the heart and surroundings (the torus).
So
everything points to the earlier suggestion that in this lower part of the
brainstem a lot of coordinating is going on, with the olivary nucleus
playing a major role, and all of most of it concerning the motor part in
connection with the short range senses, those sensing the body itself.
In the middle part of the brainstem, the pons, the long range senses,
sound and sight, come into play, among other things. Because sound and sight
turn out to be best discussed together with higher structures, they are
skipped for this moment.
So now over to the largest structure in the
neighbourhood: the cerebellum, see the next illustration (Gray 705) at more
or less the same location, but this time a sideward view:
Here visible is the global inner structure of gray and white matter (neuron
bodies and axons), and the location of the three peduncles connecting to the
cerebellum. Some extra things are shown: the trigeminal nerve, the
one that goes to the head (but not the eyes!). On
top of all, on the cut, it mentions "cerebral peduncles", the
connections towards the cortex (also: "cerebrum"). But there are a
lot of other connections too, over there ...
The cerebellum is described in a
separate article
,
not because it is too obscure, but because it is too clear. That is: the
cerebellum can be described almost completely, by invoking just one
assumption, and analyze it from there. The assumption is: the cerebellum is a
statistical averaging device. The cerebellum stores large amount of data on previous
behaviour, and averages that to use this (improved) average as a fine control on
present behaviour. That is: the cerebellum is the storage department of all
the endless exercises that athletes like tennis and golf players do. Or the
endless exercises that infants and older children do, then called "play".
With just this one assumption, it all makes sense. It is all different
aspects of the same process. And it fits perfectly well with the internal
structure of the cerebellum. And with the fact that it not yet replaced by
"the boss" of it all, the cerebrum or cortex. That looks like it outwardly, but has a
totally different function.
The next thing that comes into view on the outside, after the cerebellum
and still going upwards, are elements of the systems of hearing and sight, see the next
illustration, again of the same general area, but now seen from behind (Gray 709):
Here denoted as the
corpora quadrigemina ("foursome bodies") are four shapes bulging out of
the brainstem, giving rise to the assumption there are some spherical
structures hidden inside. These four are presently usually divided into to two's: the inferior colliculi
(below)
and
superior colliculi (above). The bottom pair are the source of
bundles of the auditory system, those on top of bundles of the optic system.
Here is a look inside (view from behind - part of Gray 691):
Also shown here is the cochlear nucleus, previously encountered because it
is situated lower in the brainstem. Like the inferior
colliculi associated with the auditory system. So what is their relation?
Here is the illustration that makes this relation clear with one stroke, and
which needed a stroke of luck to find it (from here
):
Again, read from the bottom upwards! Globally this is transparent: everything starts with the
information coming from the ear, that arrives through cochlear nerve. Which information is then processed through several stages, while going upwards through brainstem,
thalamus and finally the cortex. The process starts
with the cochlear nucleus that receives the
nerve. Then comes the associate of what until now was denoted as the
"olivary nucleus" but was in fact the "inferior olivary nucleus", which is
the (much) bigger one of two, and here is the second one, lying slightly
above it. This is probably the point where a sudden loud bang can lead to a
immediate shudder through the body. This is followed by the inferior colliculus,
just met. Its precise role in unknown (here), but probably similar to that
of the superior colliculus, of which more is known, so this is skipped
momentarily It is followed by the "medial geniculate body" which is a
nucleus that is a part of the thalamus. The thalamus relays its outcome to the
cortex through an axon bundle, that (for the sake of variation) this time is
called a "radiation" - the "auditory radiation" in Gray 691.
It is not difficult to guess what happens over here: here the sensory
information from the ear is combined, step by step, with all already
available other kinds of information, in order to construct an ever more
complete picture of reality. Which (probably) has its first final stage in
the thalamus. Which is then used "to ponder upon" by the cortex, our
consciousness.
All this applies to the simpler of the two long-distance
sense organs, but it is no surprise the other one, sight, follows a
similar pattern. When one knows what to look for, it is not difficult to
find it - here is the Gray version (Gray 722):
In fact, this is a heavily edited version of Gray 722, to make it
correspond to the correct bodily (relative) locations, i.e. to make it
readable from the bottom upwards! So as to correspond to the sequence of
operations upon the optic information!
After which corrections, it corresponds closely with
what happens to the auditory information.
But it also shows the
extraordinary importance given to the eye: three nuclei and three separate
nerves are devoted to the control of the attitude of the eye - the oculomotor, trochlear and
abducent nucleus and nerve. The latter is devoted just to moving the eye so
as to look downwards. The reason for this is also
visible in the way the field of view of the two eyes is divided: not in
left-right, but in inside-outside. The reason for this is that it divides
the field of view in "close by and far away". Things that are close by are
projected by the lens on the red-colored part of the retina. And no doubt
get a more urgent treatment: what is close by, is by definition more urgent
and potentially more dangerous. And for a standing human,
most danger comes from below, on the ground - so a special nerve for that,
too. Also note the paths that are colored black: they
correlate the auditory information with the optic control.
Some more detail, from bottom to top: those interested in the eye itself
may look here
where it is explained how such a structure may evolve - nature has done it
lots of times in different ways in different species.
The crossing of the paths is part of the hardware used to construct a picture
having depth from the two flat ones. Then come nuclei
in the brainstem that control the movements of the eye - these are extra.
Followed by the optic analogue of the inferior colliculus: the superior
colliculus. Followed by the optic analogue of the
medial geniculate body: the lateral geniculate body ("lateral" is
"sideways") - what is denoted by "pulvinar" is the backside part of the thalamus. To finally arrive
at the cortex. Probably with almost or all of the work done. The cortex
probably just builds yet another and more extensive picture of reality, this
times including all kinds of dreams and wishes ...
One structure gets
extra attention because more and illuminating information on it is
available: the superior colliculus. Here a schematic representation
of its internal structure (from here
):
This cross-section shows why it has it spherical shape: there are three
bundles of axons coming in and leaving, starting or ending in their more
voluminous cell bodies. All three ending in their
own layer, denoted by visual input layer, multimodal input
layer, and motor output layer. With, not visible, lots of
connections between the three layers that do the coordinating. An archetypal
example of a neural network. Also note: again a
decussation. "Aqueduct" is "just" the tract filled
with brain fluid that runs up and down the spine. The name "tectospinal
tract" is a fine example of the naming mess of neurology: this denotes the
other name for the superior colliculi (and surroundings): the tectum.
This is an archetypal example of this kind of neural network, doing
associative work.
This ends part two of the description of the mechanical aspect of the
functions of the brainstem. Now over to part two of what some may find more
appealing: the neuronal chemistry.
Neuronal chemistry
As already has been shown there is a neurotransmitter inside a neuron that
stimulates the neuron it is connected to firing, glutamate, and another one
(in another kind of neuron) GABA, that inhibits the connected
neuron from firing. Each neuron has multiple connections coming into it, and
the fact if it actually fires, is a balance between the activating and the
inhibiting ones. That is: the output of neurons is digital: to fire or not
to fire, but
its inputs are not - if there are very many of them, it is almost like
analog (if this is not clear enough at this point consult this
).
These
connections take care of the normal operation of the system.
Now step
all the way back to the simple mono-cellular creature with its tail. This
tail clearly is "to get away", or at least this is one of its purposes. And
this "getting away" is usually due to some perceived danger, so this
situation certainly isn't a normal situation, but a situation where you want
things to go in a hurry. Humans would call it an "emergency".
In an emergency you want things to go extra fast and if needed extra
powerful. But you don't want this situation to last any longer than
necessary, because this takes extra energy and resources, and both energy
and resources are scarce goods, due to the competition, that have to be
gathered with some effort and difficulty.
So what you want is a way
to get the system quickly into a state of emergency, and also quickly out of
it.
This is provided by neurons in the middle of the brainstem. These
neurons produce neurotransmitters that get released in the neighbourhood of
the other neurons, get into the synapses, their connecting gap, and speed
things up or slows them down again. The "speeding up"
neurotransmitter has a familiar sounding European name: noredrenalin -
Americans call it norepinephrine. It is the neurological equivalent of what
adrenalin does, quite familiarly, within the rest of the body.
And its counterpart is serotonin, which is also getting quite well known, if
only because of its increasing use in psycho-pharmaca (though as yet
indirectly: these anti-depressives (SSRI's) block the serotonin re-intake in
the synaps
). Here is a schematic overview
(side-view) of the area where the modulating neurotransmitters are produced (from here
):
|
The here relevant structures are the locus
coeruleus (or "blue area") where lie the noradrenelin-producing
neurons, and the multiple raphe nuclei
(or "border nuclei") that produce serotonin.
In this schematic representation there may seem to be a lot more serotonin
producers, but in fact these nuclei are called "raphe" nuclei because they
lie at the border between left and right halve of the brainstem, and are
indeed quite thin, see the following horizontal cross section (Gray 711):
By the way: this being so thinly spread out might point to them not being
friendly to their direct environment.
The locus coeruleus and raphe
nuclei do not produce these neurotransmitters just for the brainstem, but for the
entirety of the brain. Inclusive of the cortex, see this illustration (from
here
):
So these neurons have axons that are quite long. Though for serotonin
this task is divided: the bottom ones produce for the spinal chord, middle
ones for the brainstem, and top ones for everything above that.
Another thing: according to
standard description of e.g. the nucleus raphe magnus
, the bottom ones also seem to play a role in the processing of the pain
signals. This is not a straightforward task: pain is of such an importance
that it should be relayed immediately. But in situations of emergency, the
opposite is the case: the pain should not detract the system from getting
its escape. Just try to write a computer program that
deals with this problem. With so many possible situations and combinations
thereof ... A task a modern programmer would delegate to a neural network
...
Which is also the task that a system that includes an emergency
mode faces: when to apply it, and when to get to rest again. A task that
nature too, of course, has delegated to a neural network. So where to find a
neural network in the brainstem, other than the "simple" coordinating ones
like the olivary nucleus?
What has been learned about the global
structure of a neural network is that it consists of layers of neurons
intertwined with layers of connections. Now well-known in the meantime is
that neuron-layers are darker in color and areas with connections, i.e.
filled with axons, are lighter. So one might try to look for a striped
structure in the brainstem. Now this isn't easy to be
found, but what one does find is a structure that has a "net"-like
appearance, which in the Latin of anatomy is called "reticular": "having a
net-like appearance". A net-like appearance can be created by two
layered structures, if the layers are more or less perpendicular to each
other. So our net-like structure might be a neural network, with something
extra.
So where is that net-like structure to be found in the
brainstem? Here are a few representations - first is Gray 694 again, a
cross-section:
An overtly net-like structure in the middle coming in two variants "formatio
reticularis grisea" and "formatio reticularis alba" or "darker
reticular formation" (more neuron bodies) and "lighter reticular formation"
(more connections).
And another look at Gray 690 (bottom halve) gives its sideways view:
So the reticular formation has the shape of a column running down (or
up) the center of the brainstem. In which one finds the more distinct
nuclei, for example at the point where the two halves meet the raphe nuclei,
see the schematic view below (from here
):
This scheme uses the ancient nomenclature, so here some translations: the
bottom third is the medulla, middle pons, and upper is tectum plus
tegmentum. The central
reticular nucleus is the (inferior) olivary nucleus, and the gigantocellular
formation ("area with very large neurons"), is the pedunculopontine
nucleus
and surroundings which will come by below.
Now take that the
reticular formation is the one that, amongst other things, decides when to
start the state of emergency, and when to stop it. But how does the reticular
formation recognize situations that should be called an emergency?
Now
it would be very handy if there were rules for this. Then nature could
select those individuals that apply the rules, just by having them survive
more than the individuals that don't know the rules.
Alas, such rules
are not present, or more correctly: they vary rather frequently. So,
formulated like this, every one knows the solution to the situation: the
individual has to find out the rules as they present themselves in the
environment. That is: effective behaviour has to be stimulated and
ineffective behaviour has to be discouraged - a process abbreviated as
"learning".
And the means for this stimulating and discouraging
should not be too different from what has gone before, so how about using
neurotransmitters for it? Coming from, as for noradrenalin etc., neurons that
produce them.
Which, of course, is just a constructed introduction
towards a known conclusion: in its top section, the brainstem produces a
stimulating and a discouraging neurotransmitter. The first one is called
dopamine, which by now is overly familiar. Because it not only stimulates the
individual to doing things, it does this with great emphasis. So much so,
that every situation that releases dopamine, is also a situation that has
the potentiality of leading to addiction. At least: so it seems. Drugs like
heroin relaease (extra) dopamine, and the earlier mentioned nicotine blocks
the reuptake of dopamine in the synaps, i.e. effectively increases its
level.
The other one is much less known, and called acetylcholine. It is highly
likely that this has no less an importance than dopamine - as seen before:
nature prefers systems of "force and counter force", that together determine
the end result.
Here again an overview of the brainstem, with the general area where they
are produced, the tegmentum, denoted in green:
This shows two dopamine producing areas: substantia nigra
("black substance" - more precise: substantia nigra (pars) compacta
- in fact, it is quite flat), and its direct surroundings that are
yellow-striped, and called the ventral tegmental area ("forwardly
tegmentum") or VTA. Here a hard to find illustration from a scientific
publication
that states that there is a distribution of tasks between them:
This makes much sense. And might apply to the other neurotransmitters too.
The "one size fits all" approach is rather more surprising.
The areas
that produce acetylcholine are slight more to the bottom, and are the already mentioned
pedunculopontine nucleus
of gigantocellalur nucleus.
Together with noradrenalin and
serotonin, these four modulating
neurotransmitters perform quite basic functions in the entire brain and its
most fundamental functioning, as illustrated by the following experiments
(from the site of Ben Best
):
|
Cutting the fibers from the substantia nigra makes cats comatose.
Destruction of the locus ceruleus eliminates rapid eye movement (REM)
sleep in cats. Destruction of the raphe nuclei results in cats that
cannot sleep. |
For its role in these kind of functions, the reticular formation has
been given a prominent role in what is called the "reticular activating
system"
- the structures that determine the transitions between the states of awakening
and sleep, and alertness in general.
That is all about the chemistry
of the nervous system for now. But there is also a body that must follow it. This
is also done by means of chemistry, but this time in the bloodstream.
Neurotransmitters that are released into the bloodstream are usually not
called neurotransmitters, but "hormones". So here is where for example
adrenalin comes into play. That was much more easily detectable as something
having to do with emergencies, then frequently called "stress".
Since there is a very strict separation between nervous system and body,
called the "blood-brain barrier" (in view of the danger of infection), there
is a special organ (or two) that serves the purpose of getting
neurotransmitters or hormones into the bloodstream. Now usually these organs
are dealt with together with the emotion organs, further upwards, but in the
"bottom up" approach used here this seems rather silly: getting the
neurotransmitters also into the bloodstream is an integral part of their
purpose. So here they are, at least the two most important ones:
The structure colored red is the hypothalamus, and the grey-colored globe
attached to it is the pituitary gland. The hypothalamus controls things and
does some of the producing, the pituitary gland does the rest of the
producing and the actual transfer into the bloodstream:
Here list of the tasks that the hypothalamus performs
(Wikipedia, 02-04-2012):
|
The hypothalamus coordinates many hormonal and behavioural
circadian rhythms, complex patterns of neuroendocrine outputs,
complex homeostatic mechanisms, and important behaviours. The
hypothalamus must therefore respond to many different signals,
some of which are generated externally and some internally. The
hypothalamus is thus richly connected with many parts of the
central nervous system, including the brainstem reticular
formation and autonomic zones, the limbic forebrain
(particularly the amygdala, septum, diagonal band of Broca, and
the olfactory bulbs, and the cerebral cortex). |
This as far as the chemistry of the brainstem is involved.
Miscellaneous
There are a lot of other structures in the brainstem, and some of them
deserve some more attention.
First of all there is the peri-aqueductal grey or
PAG
,
located here:
The name is Latin that translates into "grey area surrounding the waterway".
Where the "waterway" is the channel filled with brain fluid that runs down
and up the brainstem and spinal chord - the "IV ventricle" is a
larger part of it.
The fact that the PAG is "grey" means that it is
largely filled with neuron bodies and has only short range connections between
them, i.e. it is a kind of calculating structure. Known about it is that it
plays a role in the relaying of the pain signals. And from human experience
it is known that pain signals can be partly or totally blocked. And from
other cultures it is known that individuals are able to be trained in this. And
the PAG does indeed receive input from the cortex. So it
might well be that the PAG is the place where all this is controlled.
In this context a note about the system of pain signalling. About that
much is known, so here no more details are given. But one thing is striking:
there is a separate system for slow pain and for acute pain. Now the silly
question is: why a separate system for slow pain, if you can use the also
present and certainly not disposable quicker one? One
part of the answer might be that quicker signalling requires more energy
and/or resources. So much so, that it is more efficient to have two systems.
The
brainstem, middle and top, is also the place where one finds the higher
structures involved in the "house holding" functions such as control of
blood pressure, heart rate, bladder control, etcetera, each one having its
own nucleus or area, as illustrated below (from the site of Ben Best
,
with its orientation adapted to the standard of the present site):
Also denoted here is the divide in (ortho-)sympathic and parasympathic
parts of the nervous system, the first one standing for the activating
functions, and the latter for those while at rest, like the digestive
system.
An example that is not in the illustration is that of the
nucleus of
mastication, the processing of food in the mouth. That requires at
least the coordination of the movements of jaw and tongue. This nucleus
relieves the conscious mind of continuously having to check these movements,
while still retaining overall control.
So this form of coordination
and integration already take place inside the brainstem. It is no more than
natural to assume that many structures higher in the brain, perform similar
and higher kinds of coordination and integration. As will be met in the next
layer beyond the brainstem.
Global integration
This as far as the functioning of the brainstem as individual structure is
concerned. Here some information on how it is build into the entire nervous
system, though somewhat limited because the structures that follow
have not yet been described.
These other parts are emotion organs in the
middle (yellow below), and the cortex on top (green), see this illustration:
Funnily, the connections to the more nearby emotion organs get less
attention than those to the cortex, the latter constituted by the longer and
more visible axon bundles. Here is one the more close quarter connections that
is usually overlooked:
The green area is where the bundles running from brainstem to hypothalamus
are running.
A second and third bundle is visible in the next one:
The area on the left is partly filled with the mammilotegmental tract
, a number of connections between the brainstem mammilary bodies,
belonging to the emotion organs and probably involved in the evaluation of
behaviour. The area on the right connect to what is
called the habenula
or habenular nucleus , the
interpeduncular nucleus
. The role of the habenula is less well-known but is considered among the
more ancient structures in this neighbourhood (just like the hypothalamus)
and closely related to the brainstem.
The third batch of connections
are those between the nuclei of the four the modulating neurotransmitters
and the rest of the brain. All four of them follow the same general pattern
as shown before but here with more detail (from
, but again with its orientation changed to the standard used here):
|
RN: raphe nuclei; BS: brain stem; Th: thalamus; S: septal nuclei; HC:
hippocampus; Amy: amygdala; H: hypothalamus; LC: locus
ceruleus; RF: reticular formation.
The hypothalamus is located directly at the H - the
protrusion below it is the pituitary gland . The hippocampus is denoted
as a destination (HC) but not visible itself. |
|
This is a schematic representation In reality the main branch that
encircles the brain lies directly on top of the corpus callosum, flush with
it, in a layer called the cingulum
. Here is a fMRI -recording of the connections running from the cingulum into
the cortex:
This is what happens in your brain when you feel it suddenly awash with some
emotion: these connections flush it with one or a combination of
the four modulating neurotransmitters.
For the more visible even longer range connections it was the easiest to
find a suitable illustration - the following one is Gray 764:
The gray globes near the top are the thalamus's and the gray triangles
sideways of them are parts of the putamen and environment.
The heart-shaped thing in the middle is the cerebellum that facilitates fine control. Both
are also connected to this
scheme, which is not shown (for clarity, probably). Drawn in the spinal cords are some nerves going
to muscles etcetera.
Conclusion The opinion that the brainstem just does the
housekeeping and has no further influence on our daily lives is still
widespread. So here some of the things it does do while we are awake.
First of all: the brainstem takes over in situations of high
emergency. When ultimate speed and/or power are required. There are extreme
outcomes on both sides: on the one hand there are instances where the
mother defends her child against the bear, and sailors carrying 200 pound bombs
to throw them overboard (the disaster with the USS Forrestal
)
And on the other hand there are the cases where people fall into complete
apathy.
There are the situations of lesser but still quite high
urgency but that endure a bit longer, where people start to behave
illogically and ineffectively - this is called "stress". The overloaded
brainstem overloads the entire system.
There are those situations in
normal everyday life where people fall into behaviour that is quite ineffective
and in the extreme self-destructive, that is called "addiction". The real
thing one is addicted to are the things in the brainstem - either
noradrenalin (excitement) or dopamine (almost everything else). This
has a weird extension in that it can also apply to processes higher in the
brain. Probably all extremes that go with abstract thoughts and ideas are
caused by addiction to dopamine
.
A common trait
among all these effects caused by the functioning of the brainstem, and
there are of course many more, is that they are very digital in nature: it is
"on" or "off". Now that is never a good idea to begin
with, and that is what nature improved upon with the next layers of
neurological development and evolution.
To prepare for these next layers, in separate
articles, a final section on the anatomical transition from brainstem to
the higher structures.
Follow on
For the step upwards the next layers it suffices to repeat some earlier used
illustrations. Because the brainstem is quite small relative to the other
parts of the brain, its illustration usually includes parts of these other
structures. The first one is Gray 690:
The reticular formation is undisputedly part of the brainstem, but the red
nucleus already a matter of some dispute: functionally it is part of the
brainstem being associated with "gait", i.e. a mechanical coordinating
function. Anatomically it is beyond the point where the brainstem splits
visibly into two halves. But perhaps the latter has less importance.
Also a case for discussion is the corpus subthalamicus or more
usually the subthalamic nucleus (or in very ancient nomenclature:
Nucleus of Luys). It has its main connections upwards, towards the basal
ganglia, where it seems to have the role of a "clock circuit", getting the
flow of information through the basal ganglia that themselves have only
passive interconnections. But that is the subject of the next article.
Then comes the
thalamus, the first structure that undisputedly lies outside of the
brainstem. Its present (main) role is that of a relay station between
spinal chord, brain stem and the rest of the brain further on: the emotion
organs and the cortex. This is discussed in more detail in the article on the emotion organs.
Gray 690 also shows the first ones of the latter, in the form of the onion shaped things at the top, which are the
putamen and the caudate nucleus.
Two more overviews
focus on the functional relations. The first one (from behind) already
encountered, includes the end point, the cortex:
And the second one, in side view, slightly more anatomical:
Clearly shown is that the subthalamic nucleus still has the flat shape
by getting squeezed into the tubular surroundings of the brainstem. With the
thalamus being the first to be able to really "blossom out" into a more or
less spherical structure, that is the mark of most structures above it.
The story continues with the description of the emotion organs, here
-
for a direct route to the connection of neurology with language, go to Abstraction ladder
.
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