Neurotransmitters in the Retina

Helga Kolb
Created: May 1, 2005; Updated: June 1, 2009.

General Characteristics
Today's research on the retina focuses a great deal of attention on neurotransmission
between the neurons of the retina. Various techniques using autoradiography,
immunocytochemistry and molecular biology are being used to mark neurons for
neurochemicals, their synthesizing enzymes, calcium binding proteins and receptors and
transporters of these neurochemicals. Cells immunostained with antibodies to the various
neurotransmitter candidates give particularly spectacular images by confocal microscopy
because they become stained to their finest dendrites and so can be readily classified
against their Golgi-stained equivalents. Furthermore, the whole population of
neurotransmitter specific neurons are stained so one can understand their topographical
organization into mosaics across the entire retina. Some immunocytochemistry for the
common neurotransmitter candidates has been performed on the human retina (1, 2) but
most have been done in the cat, rabbit and mouse retinas and so far the findings are the
same in general. The consistency of cell types staining across species boundaries, in
mammals at least, suggest that most, with a few exceptions, of the neurotransmitters,
neuromodulators and neuropeptides discovered in nonhuman retinas are present in
human retina too.

The Neurotransmitter of Neurons of the Vertical Pathways

through the Retina is Glutamate
Glutamate is the neurotransmitter of the neurons of the vertical pathways through the
retina. All photoreceptor types, rods and cones, use the excitatory amino acid glutamate
to transmit signals to the next order neuron in the chain (see chapter on glutamate and
Massey (3) for review). There was originally some evidence for the closely related amino
acid, aspartate, being present in rods but the later sophisticated techniques of
demonstrating amino acid signatures in retinal neurons cannot confirm aspartate as a
retinal neurotransmitter at all (4). Uptake, release and action of glutamate and agonists
upon second-order neurons in slice preparations or isolated cells in tissue culture have
also all confirmed glutamate to be the neurotransmitter acting at the first synapse in the
retina (see Lasater and Lam (5) for review). The action of the photoreceptor

NLM Citation: Kolb H. Neurotransmitters in the Retina. 2005 May 1 [Updated 2009 Jun 1]. In:
Kolb H, Fernandez E, Nelson R, editors. Webvision: The Organization of the Retina and Visual
System [Internet]. Salt Lake City (UT): University of Utah Health Sciences Center; 1995-.
2 Webvision

Figure 1. Distribution of Glutamate immunoreactivity in human retina.

neurotransmitter upon the second-order neurons is through two different types of

sensory channels though. The one type of postsynaptic receptor type is the metabotropic
glutamate channel (mGluR6) that involves a second messenger cascade and cyclic GMP
for activation of the channel (in the ON-center bipolar cell) whereas the other is an
ionotropic channel via at least two types of AMPA receptors and Na ions (OFF-center
bipolar and horizontal cells) (6-9). Photoreceptors in most vertebrates including human,
havea content of D2 dopamine receptors somewhere upon there surface (10).
Glutamate is also considered to be the neurotransmitter of all bipolar cells and most
ganglion cells in the vertebrate retina including the monkey retina (Kalloniatis and Marc,
personal communication and Marc et. al. (4)). In an immunocytochemical study of the
human retina a similar conclusion was drawn by us. All the ganglion cells appeared to
label strongly with antibodies to glutamate (2) (see Fig. 1).
Bipolar cells have receptor channels that are either of the metabotropic type, mGluR6,
(APB sensitive) or ionotropic type (AMPA) at their dendrites in the OPL, while their
axonal ending in the IPL have channels and receptors for GABA (A, B and C types), D1
dopamine and glycine because, of course, all kinds of amacrine cells are presynaptic at
these sites in the IPL neuropil. Ganglion cells are as diverse in receptor sensors as the
bipolar cells with the addition of receptors to acetylcholine (11), and the first appearance
in the retina of NMDA glutamate receptors that are typical in the brain (12, 13).
Neurotransmitters in the Retina 3

Figure 2. VGLUT3 immunostaining (A, red) shows a small-field amacrine cell. It also Immunostains for
glycine (B,C).

Figure 3. A-C, VGLUT3 cell has dendritic branching in S1 and S2. Dopamine cells branch above the main
plexus of the VGLUT3 cell. D-F, The MAP-1 ganglion cells (red) branch in the same layers as the dendrites
of the VGLUT3 cell.
4 Webvision

Figure 4. Electron microscopy of VGLUT3 immunostained processes in the IPL of the rat retina.

Amacrine cells
Recently an amacrine cell type has also been shown to contain the glutamate transporter,
vesicular glutamate transporter 3, shortened to VGLUT3 (14, 15). VGLUTs are used to
concentrate cystolic glutamate into the synaptic vesicles. Typically in the retina the
VGLUT1 isoform is expressed in photoreceptors and bipolar cells and the VGLUT2 in
ganglion cells. The amacrine cell type that can be immunostained to the antibody to
VGLUT3 is found to be a small-field amacrine cell with varicose processes that are
restricted in branching to the OFF laminae (S1 and S2) of the IPL (Fig. 2A). It is also
Neurotransmitters in the Retina 5

immunoreactive to the transmitter glycine as seen in Fig. 2, B and C, but not reactive for
GABA, dopamine (Fig. 3, A-C) or acetylcholine. Fig. 3, A-C, shows that the dopamine cell
and its dendrites run above the branches of the VGLUT3 cell and indeed appear to be able
to make synaptic contact with the cell bodies of the latter cell type, in the way we know
dopamine cell processes do (see below and amacrine cell chapter). The VGLUT3
amacrines appear to be in a good position to interact with OFF center ganglion cell,
stained for MAP-1, dendrites as shown in Fig. 3, D-F.
Electron microscopy of VGLUT3 dendrites in the IPL, indicate that they are postsynaptic
to OFF cone bipolar cell axons (Fig. 4B) and presynaptic to OFF ganglion cell dendrites
(Fig. 4A).
Recent evidence from the metabolic mapping technique of Marc and Jones (16) indicates
that a dopaminergic amacrine cell type also has a content of glutamate. However, from the
above evidence on VGLUT3 immunostaining it is clear that the dopaminergic type 1 cell
does not use that particular vesicular glutamate transporter.

Gamma Aminobutyric Acid

The classical inhibitory neurotransmitter gamma aminobutyric acid (GABA) occurs in
many different varieties of amacrine cells, and in one or more classes of horizontal cell in
most vertebrate retinas (4). There is still some controversy over whether GABA is
contained within horizontal cells in monkey and human retina.
In this figure (Fig. 5a) taken from human peripheral retina, it can be seen that there is
heavy staining with antibodies to GABA in the inner plexiform layer (three heavier bands
can be discerned) and in about half of the amacrine cell bodies in the lower row of
amacrine cells in the inner nuclear layer. Some displaced amacrines and interplexiform
cells are also revealed with GABA immunocytochemistry (2). However, the horizontal
cells are not stained at all in peripheral retina although they are in foveal retina apparently
(Cuenca, personal communication).
A valuable identification of individual cell types that contain GABA has come from
double staining techniques upon Golgi stained cell types (17). The amacrine cell that are
contain GABA are all of the large-field types. Thus we know now that A10, A13, A17,
A19, and the interplexiform cell accumulate GABA and probably use it as their primary
neurotransmitter (Fig. 5b). Many large field amacrine cells also colocalize GABA with
another neurotransmitter. Thus the GABAergic A17 colocalizes serotonin, the
acetylcholine (starburst amacrine) colocalizes GABA and so does the dopamine A18 cell
(18). Neuropetides (see later) are also commonly colocalized with GABA i.e. substance P
in A22 is almost certainly the secondary transmitter to GABA as the primary. GABAergic
amacrine cells and IPCs act upon bipolar, amacrine and ganglion cell processes or cell
bodies in the neuropils of the retina via all the three varieties of GABA receptors (a, b and
c types). It appears clear that A17 type or the type that makes reciprocal synapse with the
rod bipolar axon terminal uses a GABAc receptor (see chapter by Haohua Qian) (19).
6 Webvision

Figure 5a. Expression pattern of GABA immunoreactivity in human retina.

Figure 5b. Large-field amacrine cells in all retinas use gabaergic transmission. Illustration of large-field cell
types in cat retina.
Neurotransmitters in the Retina 7

Figure 6. Expression pattern of glycine immunoreactivity in human retina.

However, for most amacrine cells, what GABA receptors are associated with which
morphological or physiological subtype of amacrine cell, still need more elucidation.

Glycine
The other classic inhibitory neurotransmitter glycine, accounts for most of the small-field
types of amarine cell. All amacrine cells in the vertebrate retina can be accounted for by
the two inhibitory neurotransmitters GABA and glycine (4). In addition one or more
types of bipolar cell are also thought to contain glycine in mammalian retinas including
monkey and human.
In Fig. 6 it can be seen that the immunostaining for glycine is just as strong in the inner
plexiform layer as GABA staining. About the same number of amacrine cells are revealed.
However there is an addition of some small bipolar cell bodies in the inner nuclear layer
and the occasional large cell body of a ganglion cell type in the ganglion cell layer (2).
Pourcho and Goebel (20) showed quite clearly that tritiated glycine accumulated in Golgi
stained AII, A4, and A8 cells in cat retina. More recently as many as 10 different
morphological types of small field amacrine cells have been demonstrated by
immunocytochemistry and GFP (green fluorescent protein) staining in mouse retina (21,
22). The A8 cell was the most strongly glycinergic of the small field amacrine cells
according to Pourcho and Goebel, with the rod amacrine AII cell being also very clearly
glycinergic. Fig. 7 shows the commonest glycinergic amacrine cells present in the mouse
retina (22). These cells are presumably also so in the cat and primate retinas.
8 Webvision

Figure 7. Expression of glycine immunoreactivity in small-field amacrine cells of mouse retina. A-B, aII
amacrine cells; C, Type 2 cell (A2 cell); D, Type 3 cells (A4 cell); E, Type 4 cell (A6 cell); F, Type 7 cell (A9?
cell): G, A8 cell (Kolb et al. (24)). (Adapted from Waessle et al. (22)).

Glycine receptors are found on all the neurons that are postsynaptic to these glycinergic
(all small-field) amacrine cells. Thus receptors are found on certain bipolar cell axons, and
on many amacrine and ganglion cell dendrites. Again like for the GABA receptors, linking
receptor type to morphological type of postsynaptic cell is still a hot topic for research in
the retina (22, 23).

Dopamine Is Present in Amacrine Cells in the Mammalian

Retina
The neuromodulator dopamine is found in one or more types of amacrine cell in the
mammalian retina. The most robustly stained dopaminergic cell after
immunocytochemistry to the rate limiting enzyme of dopamine synthesis, tyrosine
hydroxylase (Toh), is recognized as an A18 cell of the Golgi descriptions (24, 25) (Fig. 8).
Neurotransmitters in the Retina 9

Figure 8. Immunostaining with antibodies to TOH. A18 amacrine cells stain and have overlapping
dendrites that form into rings.

It is very characteristic in wholemount appearance, with a large cell body and a dense
plexus of dendrites in stratum 1 of the inner plexiform layer. Holes or "rings" in the plexus
of criss-crossing stained dendrites are sites of amacrine cell bodies or large amacrine
dendrites (see chapter on amacrine cells) on which the processes of the Toh stained plexus
synapse. The Type 1 dopamine cell (A18) is known to synapse upon the AII rod amacrine
cell and possibly also upon A8 and A17 cells (26-29).
A second type of dopamine amacrine cell has also been described in the monkey and
human retinas (2, 30). The Type 2 CA cell has dendrites stratifying in stratum 3 of the IPL
and in vertical views of Toh processed human retinas these dendrites can be seen quite
clearly below those of the Type 1 (A18) dendritic plexus in stratum 1 (orange arrows, Fig.
9).
Type 1 dopamine cells provide ascending processes to the outer plexiform layer (see
above) which are known to synapse on the GABAergic interplexiform cell (see previous
chapter on feed-back loops).GABA and dopamine colocalize to the A18 cell type (Type 1
CA cell) and serotonin has also been found to co-exist in in the same amacrine cells in cat
retina (31).
Both D1 and D2 receptor types have been found on neurons of the inner and outer retina
in many vertebrates. It is thought that the D1 receptor is particularly associated with cells
10 Webvision

Figure 9. Vertical section of a type 1 TOH+ amacrine cell.

that are coupled by gap junctions, because of dopamine's known action in cyclic AMP
regulation of gap junction channels (32). Thus D1 receptors have been demonstrated in
association with horizontal cells of the OPL and some amacrine cells of the IPL. D1
receptors are robust on ganglion cell bodies as well (personal observations), despite the
fact that the dopamine amacrine cell is not known to make direct synapses upon ganglion
cells. D2 receptors are also found in the retina associated with photoreceptors in the outer
nuclear layer, outer limiting membrane and retinal pigment epithelium even. Also they
are present in the IPL but the target cells in this neuropil are not well understood yet.

Acetylcholine
The classic fast excitatory neurotransmitter of the peripheral nervous system,
acetylcholine (ACh), is found in a mirror symmetric pair of amacrine cells in the
vertebrate retina. In the rabbit such cells have been named starburst cell (33, 34). One of
the mirror pair occurs in the amacrine cell layer with dendrites in sublamina a (OFF
sublamina of the IPL). The other of the pair has its cell body displaced to the ganglion cell
layer and its dendrites stratify in sublamina b (ON sublamina of the IPL).
These ACh containing amacrine cells are common to almost all vertebrate retinas and
have been described morphologically in human retina too (25, 35) (see previous chapter
on amacrine cells). Both muscarinic and nicotinic receptors have been demonstrated in
the mammalian retina, particularly associated with transient phasic ganglion cells (Y
Neurotransmitters in the Retina 11

Figure 10. ACh containing amacrine cell stained with Lucifer yellow.

cells) (36, 37) and effects of ACh and antagonists on ganglion cell responses are
documented but not well understood (Fig. 10).

Serotonin
There are two types of serotonin-accumulating amacrine cell in the rabbit retina (38). One
of these is almost certainly the A17 cell or the reciprocal amacrine cell of the rod system
in the rabbit (see previous chapters) (Fig. 11). However, in cat retina, a completely
different amacrine cell type stains with antibodies against serotonin. One cell type in cat is
similar to the wide-field cell A20, while the other may be the A18 or dopamine cell (31).
Even where serotonin is strongly demonstrated in these amacrine cells in rabbit retina, it
is not thought to be the neurotransmitter.
12 Webvision

Figure 11. Serotonin accumulating amacrine cells in rabbit retina known as A17 GABAergic cells in cat and
primate retinas.

Serotonin co-exists with GABA, in the A17 cell and the latter is thought to be the
releasable transmitter (18, 39). A few cold-blooded vertebrates have a bistratified
amacrine cell and a bipolar cell type that immunostain for serotonin (40, 41).

Adenosine May Be a Retinal Neurotransmitter

The purine nucleotide, adenosine may be a neurotransmitter or neuromodulator in the
mammalian retina. Autoradiography and immunocytochemistry for adenosine has
revealed cell bodies in the amacrine and ganglion cell layers (42). Probably most of these
cells are amacrine cells but some in the ganglion cell layer may be true ganglion cells. In
human retina additional cells that could be bipolar or horizontal cell label too. K+ and
Neurotransmitters in the Retina 13

light evoked release of adenosine can be measured in rabbit and chick retinas. And the
vertical pathway neurotransmitter glutamate can induce adenosine release from [3H]-
adenosine preloaded rabbit retina. Additionally some effects of adenosine on the ERG
generated in the retina and on the activity of ganglion cell terminals in the superior
colliculus have been recorded and the information points to the strong likelihood that
adenosine does play a neurotransmitter role in the vertebrate retina (See review by
Blazynski and Perez (42)). Adenosine colocalizes with GABA, acetylcholine and serotonin
in various retinas. Much more research is needed in this area.

Substance P Occurs in an Amacrine Type and a Ganglion Cell

Type
Substance P (SP) is a neuropeptide belonging to the tachykinin family that include
neurokinin A, neuropeptide K and neurokinin B, as well. Substance P is thought to be a
neurotransmitter or neuromodulator in the vertebrate retina (See Kolb et al. (43) for a
review).
SP-IR amacrine cells appear to be of a single type in the human retina (Fig. 12). They are
large-field cells with large cell bodies (16 um diameter) lying in normal or displaced
positions on either side of the inner plexiform layer (IPL).
Their sturdy, spiny and appendage-bearing dendrites stratify in stratum 3 (S3) of the IPL,
where many overlapping fine dendrites intermingle to form a plexus of stained processes.
Either cell bodies or primary dendrites emit "axon-like" process which divides typically
into two long, fine processes that run in opposite directions for hundreds of microns in S5
and S3 before disappearing as distinct entities in the stained plexus in S3. Long fine
dendrites also pass from the dendritic plexus to run in S5 and down to the nerve fibre
layer to end as large varicosities at blood vessel walls. In addition fine processes are
emitted from the dendritic plexus that run in S1, and some pass up to the outer plexiform
layer (OPL) to run therein for short distances. The SP-IR amacrine cell has many
similarities to thorny type 2 amacrine cells described in Golgi studies. SP amacrine cells
co-localize GABA as a neurotransmitter (44).
In addition to the SP-IR amacrine cells, a ganglion cell type is immunostained with SP
(Fig. 13). Its 20- 22 um cell body gives rise to a radiate, sparsely-branched, wide-spreading
dendritic tree running in S3. Its dendrites and cell body become enveloped by the more
intensely SP-IR processes and boutons from the SP-IR amacrine cell type (Fig. 13, fine
arrows). The SP-IR ganglion cell type most resembles G21 of a Golgi study. A ganglion cell
type in rabbit retina has definitely been proved to contain substance P. Such ganglion cells
almost certainly colocalize a more standard neurotransmitter like glutamate too (45).

Other Neuropeptides
Immunostaining with antibodies against somatostatin has revealed a small population of
neurons in the ganglion cell layer in the rabbit retina (46). They are distributed only in the
14 Webvision

Figure 12. Substance P containing amacrine cell in human retina.

Figure 13. Substance P containing ganglion cell in human retina.

Neurotransmitters in the Retina 15

Figure 14. Somatostatin-IR amacrine cells of the monkey retina (47).

Figure 15. Vertical section of CRF-IR amacrine cells are probably the same type as those labelling with
somatostatin (47).

inferior retina and in the far peripheral circumference of the retina. However, in the
monkey and human retinas these amacrines are more uniformly distributed. The
somatostatin-IR amacrines have long fibers that distribute across the entire retina running
in three plexuses in the middle and outer and inner strata of the inner plexiform layer
(Fig. 14, from Marshak (47)). In the human and monkey most of the somatostatin-IR cells
have their cell bodies in the ganglion cell layer (Fig. 14) and they emit fibers or axon-like
processes that can be measured running 20 mm across the entire retina (48). Because
these axon like processes stay within the retina, not passing to the optic nerve, the
somatostatin-immunoreactive cells are likened to the associational neurons of Cajal (49).
Corticotropin releasing factor (CRF) is contained within a population of wide-field
tristratified amacrines with long axon like processes, very similar in morphology to those
16 Webvision

Figure 16. Vasoactive Intestinal peptide-IR amacrine cells of the monkey retina (47).

Figure 17. Vertical section of neuropeptide Y-IR amacrine cells are wide-field and unistratified in stratum 1
of the IPL (47).

containing somatostatin (Fig. 15) (47). Colocalization of the two peptides has not been
attempted yet but it seems probably that they are one and the same cell type.
Vasoactive Intestinal peptide (VIP) immunostains a population of amacrine cells that can
be normally place in the INL or displaced to the ganglion cell layer (Fig. 16). They appear
to have a medium size dendritic field and dendrites that branch diffusely through the
middle strata of the IPL. They may be equivalent to the A12 type of Mariani (50).
Amacrine cells that immunostain for neuropeptide Y are quite regularly stained in the
primate and rodent retina. These amacrine cells have a characteristic sparsely branched
wide-field dendritic tree with dendrites running in stratum 1 of the IPL (Fig. 17) (47).
Recent research wherby NPY amacrine cells can be selectively ablated indicates that these
Neurotransmitters in the Retina 17

Figure 18. Two type of amacrine (1 and 2, ) and a type of bipolar cell known to contact blue cones is
immunostained with the peptide cholecystokinin (47).

cells have a role in low spatial frequency tuning of certain large ganglion cell types (51).
Both ON and OFF center ganglion cell types are said to be equally affected by loss of NPY
amacrine cells, which is a strange finding considering the OFF stratum branching of these
particuar amacrine cells. Presumably another chain of amacrine cells is involved in spatial
tuning of ganglion in addition to the NPY type.
Using antibodies to the glycine extended form of cholecystokinin, Marshak and colleagues
were able to demonstrate the presence of two different morphological types of amacrine
cell and a single morphological type of bipolar cell in the monkey retina (47, 52). The
amacrines come as pairs that are either bistratified in the strata 2 and 4 and a larger
bodied monstratified pair, again branching in either stratum 2 or 4 (Fig. 18). The blue
cone specific bipolar cell is well characterized by CCK immunostaining and its
distribution, contact with blue cones specfically and its probably ON center physiology is
now well known (52, 53) (see chapter on S-cone pathways).
18 Webvision

Figure 19. NADPH-diaphorase histochemical staining of an amacrine cell in monkey retina.

NADPH-Diaphorase Staining and the Possibility That There are

Nitric Oxide Containing Neurons in the Retina
Nitric oxide is formed in many nerve cells of the peripheral and central nervous system. It
is known to play a role in second messenger cascades by activating guanylyl cyclase and
elevating cyclic GMP levels. Since many neurotransmitters and transduction events utilize
cyclic GMP in the retina (notable phototransduction in the photoreceptors and activation
of metabotropic glutamate receptors) the idea that nitric oxide might play a key role in
retinal neurotransmission has been forwarded. Nitric oxide synthase the enzyme required
for synthesis of nitric oxide requires NADPH as a cofactor in the cell. Thus the simple
histochemical technique to reveal NADPH in a cell by the reduction of tetrazolium salts
has proved to be a good marker of neurons that contain NADPH-diaphorase and hence
possibly also nitric oxide synthase. An antibody has been made against the synthase but
Neurotransmitters in the Retina 19

Figure 20. Drawing from wholemount retina stained with NADPH-diaphorase of the commonly stained
amacrine cell.

staining and identification of morphological cell types that contain this enzyme have been
unsatisfactory to date for the retina. Thus the NADPH-diaphorase histochemical
technique is still more reliable.
In monkey retina, three types of amacrine cell and one type of ganglion cell appear clearly
stained with NADPH-diaphorase. The commonest type is the one shown above (Fig. 19
and Fig. 20) and occurs at a maximum density of 280 cell/mm2 at 1 mm from the fovea
(Cuenca, personal communication). It has a large cell body that lies either in the amacrine
cell layer or can be displaced to the ganglion cell layer. The dendrites radiate out from the
cell body like the spokes of a wheel, contain, many fine spines, and lie on one plane in
stratum 3 of the IPL. Fine axon like processes arise from the main dendritic tree that is
about 300 um in diameter to run for mm in all directions. The cell is probably the
20 Webvision

Figure 21. Summary diagram of the organization of neurotransmitters in the retina.

equivalent of a spiny amacrine described by Mariani (50) and a cell type stained by Dacey
as axon bearing (54).

Amacrine Cell Populations and Mosaics Arrangements Are

Revealed by Neurotransmitter Immunocytochemistry
Table 1 lists the amacrine cell types that have now become associated with particular
neurotransmitter substances in the mammalian retina. The list is compiled primarily from
data of cat and rabbit retinas. However, some clearly recognizable amacrine cell types of
the monkey and human retina have been included. The list is as yet incomplete
concerning an exact classification of each cell type by neurotransmitter signature because
of the difficulty of agreeing on correspondence between different authors classification
schemes and making cross species comparisons. The review article by Vaney (18) is the
most comprehensive treatment on the subject of the differences between cat and rabbit
retina in this regard.
Immunocytochemical staining of amacrine cells, when done on wholemounts of retina,
can reveal every cell of the population that are immunoreactive to the antibody used.
Thus we are increasingly acquiring population maps and distribution maps of all the
different types of amacrine cells according to neurotransmitter content. Most amacrine
types are arranged in regular mosaics and the individual cells have certain overlap
Neurotransmitters in the Retina 21

characteristics that can be calculated from nearest neighbor statistics. The cat and monkey
amacrine mosaics peak in cell density with closest packing of their smallest dendritic trees
in the fovea or area centralis. Then from center to periphery the neurons distribute evenly
in concentric rings of decreasing density and increasing dendritic field sizes. The rabbit
amacrine mosaics peak along the horizontally-organized visual streak and fall off
therefrom linearly into superior and inferior retina (18). Some of the more sparsely
distributed neurotransmitter types, have unique distributions. For example, the
somatostatin-immunoreactive associational neurons in rabbit retina are located almost
exclusively in inferior retina.
The commonest amacrine cell type of the cat and rabbit retina is the glycinergic AII
amacrine (estimated 512,000 total in cat retina) followed by the serotonin-accumulating
amacrines (between 170,000 and 230,000 cells in rabbit), cholinergic cells (approximately
130,000 in rabbit retina) and substance P-containing cell types (39,000 cells in cat retina).
Dopaminergic amacrine cells are some of the lowest density populations of cells (3-5,000
in cat; 6-8,000 in rabbit). Somatostatin occurs in a small population of only 1000 cells in
rabbit retina (Fig. 21).

Table 1. Common neurotransmitter substances found in mammalian amacrine cells

Neurotransmitter Amacrine cell type
rabbit cat primate
GABA A2, A10, A13, A17, A19, A22, A20 same as cat
glycine All A3, A4 same as cat
DAPI-3 All
A8
acetylcholine “starburst” Ca, Cb “starburst” “starburst”
a-type, b-type a-type, b-type
dopamine Toh-IR A18 CA-type1
CA1, CA2, CA3 CA-type2
serotonin S1, S2 A20 and A22 A17
substance P tri-stratified A22 thomy 2
VIP Tri-stratified
somatostatin association neuron long axon-like
processes
nitric oxide NADPH-diaphorase cell multiaxonal spiny
Compiled from Vaney (18)

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