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Neuronal Cell Types: Primer
Neuronal Cell Types: Primer
Neuronal Cell Types: Primer
R497
which send an axon out of the diameter), but send an axon that circuit function as well as
structure where their soma is runs for millimeters within the communicating with the brain’s
located, and intrinsic neurons, cerebellum. The granule cells are distant regions.
which make synapses only within tightly packed and the The major class of projection
the structure where their soma is cerebellum is large; in absolute neurons in the neocortex are the
located. (Alternative terms are terms, they are the most pyramidal cells. They have a wide
‘principal cells’ and numerous single type of neuron variety of shapes and
‘interneurons’, respectively.) Note in the nervous system. Two other projections. Their cell bodies can
that the distinction applies only interneurons are the basket and be located in any of the cortical
to the cell’s axonal projection, stellate cells. The Golgi cells — layers except layer 1. Many,
not to the source of its inputs: an named for their discoverer, though possibly not all, pyramidal
intrinsic neuron can receive Camillo Golgi, not for his staining cells project to distant regions.
synaptic inputs from cells located method — are much larger, with The term ‘pyramidal cell’ is used
within the same structure or from dendritic arbors that span all for neurons in several brain
distant ones. The following levels of the cerebellar cortex. structures (there are
paragraphs give examples of hippocampal pyramidal cells) and
neuronal types in the cerebellum Neurons of the neocortex generally denotes a large cell
and the retina, where the types This simple list of cerebellar with a roughly triangular soma
are pretty well understood, and neurons, and the images of from which arise distinct sets of
the neocortex, which has proved isolated cells in Figure 2, must apical and basal dendrites. Thus,
a much harder nut to crack. unfortunately ignore the the structure of the classical
gorgeous architecture with which pyramidal cell suggests that its
Neurons of the cerebellum the cells are interlaced — the job is to combine functionally
The cerebellum has long been a brain as Sherrington’s “great varied inputs from several cellular
popular model system, in large enchanted loom”. The neocortex layers, make a decision about
part because of its orderly and surely has a similar orderliness, those inputs, and send a
relatively simple architecture. It were we to understand it, but in message via a spike train to
contains a single type of the neocortex the overall plan is another region. The term has
projection neuron, the Purkinje less obvious and the taxonomy of drifted, however, and is now used
cell (Figure 2). Not only do these the neuronal types much less for cells that have neither
neurons have a highly clear. In contrast to the triangular somas nor the classic
stereotyped architecture, but cerebellum (or the retina, see branching pattern. It is believed
they are essentially two below), projection neurons in the that many cell types exist within
dimensional — they have neocortex outnumber intrinsic the large class of neocortical
somewhat the shape of a leaf. neurons, making up about 80% pyramidal cells, but there is at
This makes them easy to of all cortical neurons. This present no powerful and
conceptualize and easy of access difference may occur because universally agreed way to
for recording. projection neurons of the distinguish them.
The cerebellum contains five neocortex, in contrast to those of Even though intrinsic neurons
main types of intrinsic neuron. the cerebellum or retina, often comprise the minority of
Granule cells have tiny cell send axon collaterals that neocortical neurons, an
bodies (5–8 µm) and small terminate within the cortex. This extremely large number of types
dendritic arbors (~50 µm means that they have a local have been identified; the subset
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Current Biology
Figure 3. Patterns of connectivity in the retina, illustrated for three levels of the inner plexiform layer.
The level of stratification of the different cell types is an initial wiring diagram of the retina: it defines which cells can contact each other.
identified type have a regular If there are ~60 cell types in the marker that reveals the cell’s
mosaic. A mixture of cells from retina, how many are there in the arborizations, because selective
different types does not. cortex? From plausible expression of the protein in
assumptions of spacing, cell combination with neuronal
Future prospects number and dendritic field structure is unambiguous. This
For the first time in any diameter, one observer estimated combined approach is less
substantially complex CNS the number at ~1,000. At the cumbersome and more robust
structure, the inventory of cell least, this estimate serves the than strictly morphological
types in the retina appears to be rhetorical purpose of pointing out methods, and more reliable than
virtually complete. Evidence for how important is the number of depending solely on the type-
this comes partly from a new, types for any understanding of specificity of expression of a
unbiased, morphological this structure. If there are 1000 newly identified gene. It will
sampling technique; from the types of neuron in the cortex (or probably be the main way of
sheer volume of well-visualized even 100), there is a huge gulf identifying new cell types for the
cells that is easily generated by between that number and our foreseeable future.
modern cell-filling methods; from understanding of the cortex’s
the three-dimensional resolving physiological diversity. A Further reading
power of digital microscopy; and resolution, one way or the other, Blackshaw, S., and Livesey, R. (2002).
Applying genomics technologies
from the law of coverage, which would tell us which conceptual
to neural development. Curr. Opin.
dictates that a true cell type be models of the cortex are Neurobiol. 12, 110–114.
evenly spaced. Many types are plausible. Masland, R.H. (2001). The
now terminally identified: the New probes originating from fundamental plan of the retina.
combination of morphology, genome screening promise to Nat. Neurosci. 4, 877–886.
Rockhill, R.L., Euler, T., and Masland,
gene expression, and mosaic help sort the cells, although few R.H. (2000). Spatial order within
make further subdivision of the instances of truly cell-type- but not between types of retinal
type extremely unlikely. Fine- specific gene expression have neurons. Proc. Natl. Acad. Sci.
tuning of the classification will been discovered so far. The USA 97, 2303–2307.
Shepherd, G.M. (ed) (2004). The
still occur, and a few orphan common occurrence is partial Synaptic Organization of the Brain
types remain among the wide- specificity, in which several types (Oxford University Press).
field cells, which can cover the of neuron express the gene in Stevens, C.F. (1998). Neuronal
retina using small absolute question. Although it is diversity: Too many cell types for
comfort? Curr. Biol. 8, R708–R710.
numbers of cells. But the survey disappointing that more precise
probably encompasses 98% of specificity is not widespread,
Massachusetts General Hospital, 50
all of the retina’s neurons and it mixed expression is nonetheless Blossom St., Wellman 429, Boston,
is sure that no major players extremely valuable. The Massachusetts 02114, USA. Email:
have been missed. ambiguity adds only a need for a masland@helix.mgh.harvard.edu.