Biology 104: Spring 2004 : Nervous system: (material from chapters 12 and 13)

The anterior part of the neural tube expands (driven by inside fluid pressure!)
and subdivides into five parts

Telencephalon-----Cerebrum (very large in mammals, esp. humans)
                 Hippocampus ("memory storage")

Diencephalon----- Optic vesicle neural retina and pigmented retina
                 Thalamus and epithalamus, and hypothalamus

Mesencephalon-----Tectum, including the optic tectum

Metencephalon-----Cerebellum

Myelencephalon-----Medulla

An interesting fact about the neural tube: cell bodies move to the surface of the neural tube before and during each mitotic division (see fig 12, page 403)

For many years, scientists saw that all the mitotic cells were next to the neural tube, and drew the logical conclusion (that only those cells were going through the cell cycle).
But it turned out that all the cells were making DNA, but that they shift back toward the lumen to divide! (This is also true of other pseudostratified epithelia, like somites)

When neural tube cells undergo transverse cytokinesis, then one daughter cell remains connected at the lumen and continues cell-cycling. but the other daughter cell differentiates as a nerve cell.

Nerve cells never again enter the S period, once they have formed an axon and dendrites.
Differentiated nerve cells never divide, & never even make DNA

Nerve axons (and also dendrites) are strands of cytoplasm laid down behind an actively crawling amoeboid tip, called the "growth cone" .

The "wiring pattern" of the nervous system is produced by several different mechanisms that guide crawling of growth cones

Sometimes, there are gradients of chemotactic attractants. (chemotaxis)
Other times, there are adhesion gradients. (haptotaxis)
Other times, they are guided along glial cells
This is part of a major field of research: sometimes called "neural connectivity"

A lot of such research is done on grasshoppers.
Sea slugs are studied because they have giant nerve cells.

In vertebrates, two of the most intensively-studied regions are the cerebellum
and the optic nerve (retino-tectal connection)

The cerebellum is necessary for balance, and has relatively few differentiated cell types (~7) each one of which forms very stereotyped shapes.

One kind of cerebral cell are the Purkinje cells each Purkinje cell has a "dendritic arbor" which is flat but otherwise looks like, and is as complicated as, a tree. (and forms around a hundred thousand synapses) One of the other cell types extends long axons that are exactly perpendicular to the plane of flattening of the Purkinje cells.

Mutations in certain genes cause incorrect wiring of the cerebellum and the phenotype of these mutations is that the mice can't keep their balance, and the names of each of 30+ genes are "staggerer", "reeler", "waltzer", "weaver" etc.

(as usual, genes get named after what is abnormal in mutants)

Each of these genes codes for some protein needed to cause the correct wiring pattern of the cerebellum.

In the neural retina, the rod and cone cells detect light; but are located directly next to the pigmented retina.

A completely different kind of cells, called "ganglion cells", are the ones that send axons back to the brain.

Please note that the word ganglion means a mass of nerve cells that are clumped together somewhere outside the nervous system. For example, the pairs of sensory ganglia along the sides of the spinal cord, and also the many autonomic ganglia.

The ganglion cells of the retina really are not in ganglia; so it was misleading to give them the name "ganglion cells". (& try not to let it confuse you)

In each eye, you have about a million ganglion cells.
One axon extends from each of these ganglion cells, and "grows" back from the eyeball to the bottom of the brain, where it crosses the optic nerve from the other eye, in a structure called the optic chiasm) and then extends all the way to the optic tectum. (except in mammals, where it goes somewhere else; never mind where)

"Growth" of nerve fiber tips is really active crawling (amoeboid)
(Simple growth is rarely if ever used to create embryological patterns, but people and textbooks habitually say that cells "grow" to a certain shape or location. They don't mean it literally.)

At the tectum, each ganglion cell axon forms a synapse; The spatial pattern of these ganglion cells connections is map-like. This is the best-studied example of a "neural projection"

The nervous system contains many (hundreds?) of examples of neural projections, in which nerves connect in a spatial pattern that is a map of the relative locations of the other ends of these same nerves. There are sensory projections and motor projections. It has long been "A Holy Grail" for neurobiology to discover the control mechanisms that guide axons to form projections.
Thousands of research papers have been published on the subject

In order to get to the optic tectum, the ganglion cell axon growth cones have to be guided along the following routes: (& the guidance mechanism for each step needs to be explained)

1) Converge toward the "blind spot" inside the retina itself;

2) Extend from the eyeball back to the brain, and downward to the floor of the diencephalon;

3) Pass the optic nerve fibers coming from the other eye (at the Optic Chiasm)

4) Extend up the opposite side of the brain

5) Spread out across the roof of the mesencephalon to form a map-like arrangement of synapses
(which is for some reason upside-down and backwards! for example, ganglion cells from the upper left corner of the retina connect their axons to the lower right corner of the optic tectum

Stereo vision depends on the brain being able to compare what is seen by one eye, in comparison to the other.

In animals with stereo vision, about half the axons turn back at the optic chiasm, and then extend to the same side of the brain. For example the right retina extends half its fibers to the right brain.

Surprisingly, in Siamese Cats and White Rats, and White Mice something goes wrong with this "turning back" mechanism, and all the fibers from the right eye go to the left brain,
and all the fibers from the left eye go to the right brain. As a result, these animals not only lack stereo vision, that also seem to "see double" in half of each eye.

 


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