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)
Diencephalon----- Optic vesicle neural retina and pigmented retina 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). 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. 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)
A lot of such research is done on grasshoppers.
In vertebrates, two of the most intensively-studied regions are the cerebellum 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.
"Growth" of nerve fiber tips is really active crawling (amoeboid) 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. 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 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,
|