Bipolar Cells of the Mouse Retina: A
Gene Gun, Morphological Study
VINCENZO PIGNATELLI AND ENRICA STRETTOI
*
Istituto di Neuroscienze del Consiglio Nazionale delle Ricerche, sede di Pisa, Area della
Ricerca Consiglio Nazionale delle Ricerche, Via Giuseppe Moruzzi 1, 56100 Pisa, Italy
ABSTRACT
One of the key elements concerning our understanding of the organization of the mouse
retina is the complete classification of the various types of bipolar cells. With the present
study, we tried to contribute to this important issue. Unfortunately, most of the antibodies
that stain specifically bipolar cells in the retina of other mammals hardly work for the retina
of the mouse. We succeeded in overcoming this limitation by using a relatively novel tech-
nique based on the gene gun transfer of fluorescent dyes to cells. Hence, we were able to stain
a considerable number of bipolar cells that could be characterized according to morphological
and comparative criteria. We also performed a complete morphometric analysis of a subset of
bipolar cells stained by anti–neurokinin-3 receptor antibodies. We found nine types of cone
bipolar cells and one type of rod bipolar cell; these data are consistent with the findings of
previous studies on the retinas of other mammals, such as rabbits, rats, and monkeys and
with a recent study based on the mouse retina (Ghosh et al. [2004] J Comp Neurol 469:70 –
82). Our results also confirm the existence of a common structural similarity among mam-
malian retinas. It remains to be elucidated what is exactly the functional role of the various
types of cone bipolar cells and what is the specific contribution they provide to the perception
of a given visual stimulus. Most probably, each bipolar cell type constitutes a specialized
channel for the computation of a selected component of the visual stimulus. More complex
signal coding, involving the coordinated activity of various types of bipolar cells, could also be
postulated, as it has been shown for ganglion cells (Meister [1996] Proc Natl Acad Sci U S A
93:609 – 614). J. Comp. Neurol. 476:254 –266, 2004. © 2004 Wiley-Liss, Inc.
Indexing terms: bipolar cells; gene gun; morphology
The central nervous system operates through both se-
rial and parallel processing; the latter implicates that
multiple, anatomically distinct areas or neural circuits
execute various tasks at the same time, upon receiving a
similar input signal (Grunert et al., 1994; Prut et al., 2001;
Zaborszky, 2002; Chiry et al., 2003; Scott and Johnsrude,
2003). The retina makes no exception to this working
strategy: many types of anatomically and functionally dis-
tinct ganglion cells, the last neurons in the retinal visual
pathway, operate in parallel to provide the brain, through
multiple sets of channels, information about the position,
chromatic features, contrast, duration, and so on, of a
given visual stimulus (Rodieck, 1998). The complexity of
ganglion cell physiology originates partly from complex
synaptic interactions taking place in the outer and inner
plexiform layers of the retina; in turn, the complexity of
retinal networks is reflected in the high variety of cellular
types that compose each of the various classes of retinal
neurons (Masland and Raviola, 2000).
In the past few years, the full catalog of neurons has
been provided for the retina of various mammals and the
puzzle composition is quickly progressing toward the iden-
tification of all the retinal cells for an increasing number
of species. We know that the rabbit retina (and thus pre-
sumably, the retina of most mammals) contains 2 types of
photoreceptors, 2 types of horizontal cells, approximately
a dozen types of bipolar cells (McGillem and Dacheux,
2001), 23 types of amacrine cells, and, finally, 12 varieties
of ganglion cells (Masland, 2001b). Shape, size, pattern of
dendritic and axonal stratification, connectivity, staining
properties, and so on, contribute to the definition of a cell
Grant sponsor: TeleThon Project E833; Grant sponsor: National Insti-
tutes of Health; Grant number: R01 EY 12654.
*Correspondence to: Enrica Strettoi, Istituto di Neuroscienze del CNR,
sede di Pisa, Area della Ricerca, Via G. Moruzzi 1, 56100 Pisa, Italy.
E-mail: strettoi@in.cnr.it
Received 18 November 2003; Revised 8 March 2004; Accepted 19 April
2004
DOI 10.1002/cne.20207
Published online in Wiley InterScience (www.interscience.wiley.com).
THE JOURNAL OF COMPARATIVE NEUROLOGY 476:254 –266 (2004)
© 2004 WILEY-LISS, INC.