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PubMed2003

Melanopsin and rod-cone photoreceptive systems account for all major accessory visual functions in mice.

In the mammalian retina, besides the conventional rod-cone system, a melanopsin-associated photoreceptive system exists that conveys photic information for accessory visual functions such as pupillary light reflex and circadian photo-entrainment. On ablation of the melanopsin gene, retinal ganglion cells that normally express melanopsin are no longer intrinsically photosensitive. Furthermore, pupil reflex, light-induced phase delays of the circadian clock and period lengthening of the circadian rhythm in constant light are all partially impaired. Here, we investigated whether additional photoreceptive systems participate in these responses. Using mice lacking rods and cones, we measured the action spectrum for phase-shifting the circadian rhythm of locomotor behaviour. This spectrum matches that for the pupillary light reflex in mice of the same genotype, and that for the intrinsic photosensitivity of the melanopsin-expressing retinal ganglion cells. We have also generated mice lacking melanopsin coupled with disabled rod and cone phototransduction mechanisms. These animals have an intact retina but fail to show any significant pupil reflex, to entrain to light/dark cycles, and to show any masking response to light. Thus, the rod-cone and melanopsin systems together seem to provide all of the photic input for these accessory visual functions.

Animals↗
PubMedDate not supplied

Horizontal cell sensitivity in the cat retina during prolonged dark adaptation.

The effects of dark adaptation on the response properties of ganglion cells have been documented extensively in the cat retina. To pinpoint the different retinal mechanisms that underlie these effects, we studied the response characteristics of cat horizontal (H) cells during prolonged dark adaptation. H-cell responses were recorded intracellularly in the optically intact, in vivo eye. To disentangle rod and cone contributions, sensitivity changes during dark adaptation were tracked with white light and with monochromatic lights that favored either rod or cone excitation. Stable, long-lasting recordings allowed us to measure changes of sensitivity for adaptation periods up to 45 min. Thresholds for white light and 503-nm monochromatic light decreased steadily and in parallel. The maximum increase of sensitivity, after extinguishing a photopic adaptation light, was 1.8 log units only, reached after about 35 min. Sensitivity for 581-nm lights also increased steadily, but at a shallower slope. The steady increase of sensitivity was concomitant with a linear shift in resting membrane potential and with an increase in relative rod contribution to the threshold responses. Even though small-amplitude responses were rod dominated after prolonged dark adaptation, sensitivity to rod signals remained relatively low, compared to sensitivity of cone responses or to the absolute sensitivity of ganglion cells. This suggests that the cone-H-cell pathway plays no role in the dark-adapted cat retina.

Adaptation, Ocular↗
PubMedDate not supplied

Cholecystokinin-like immunoreactive amacrine cells in the rat retina.

High levels of endogenous cholecystokinin (CCK) are present in the rat retina (Eskay & Beinfeld, 1982), but the cellular localization and physiological actions of CCK in the rat retina are uncertain. The goals of this study were to characterize the cells containing CCK, identify cell types that interact with CCK cells, and investigate the effects of CCK on rod bipolar cells. Rat retinas were labeled with antibody to gastrin-CCK (gCCK) using standard immunofluorescence techniques. Patch-clamp methods were used to record from dissociated rod bipolar cells from rats and mice. Gastrin-CCK immunoreactive (-IR) axons were evenly distributed throughout the retina in stratum 5 of the inner plexiform layer of the rat retina. However, the gCCK-IR somata were only detected in the ganglion cell layer in the peripheral retina. The gCCK-IR cells contained glutamate decarboxylase, and some of them also contained immunoreactive substance P. Labeled axons contacted PKC-IR rod bipolar cells, and recoverin-IR ON-cone bipolar cells. CCK-octapeptide inhibits GABA(C) but not GABA(A) mediated currents in dissociated rod bipolar cells.

Amacrine Cells↗
PubMed2006

P2X2 receptors on ganglion and amacrine cells in cone pathways of the rat retina.

Extracellular ATP is known to mediate fast, excitatory neurotransmission through activation of ionotropic P2X receptors. In this study, the localization of the P2X(2) receptor (P2X(2)R) subunit was studied in rat retina by using immunofluorescence immunohistochemistry and preembedding immunoelectron microscopy. The P2X(2)R was observed in large ganglion cells as well as in a subset of amacrine cells. Double labeling revealed that 96% of all P2X(2)R-immunoreactive amacrine cells showed gamma-aminobutyric acid (GABA) immunoreactivity. Subsets of P2X(2)R-immunoreactive amacrine cells expressed nitric oxide synthase and substance P; however, no colocalization was observed with choline acetyltransferase, vasoactive intestinal peptide, or tyrosine hydroxylase. Nearest-neighbor analysis confirmed that P2X(2)Rs were expressed by a heterogeneous population of amacrine cells. The synaptic connectivity of P2X(2)R amacrine cells was also investigated. It was interesting that P2X(2)R-immunoreactive amacrine cell dendrites stratified in the sublaminae of the inner plexiform layer occupied by cone, but not rod bipolar cell axon terminals. Immunoelectron microscopy revealed that P2X(2)-immunoreactive amacrine cell processes were associated with cone bipolar cell axon terminals as well as other conventional synapses in the inner plexiform layer. Taken together, these data provide further evidence for the involvement of extracellular ATP in neuronal signaling in the retina, particularly within cone pathways.

Amacrine Cells↗
PubMed2004

Predominant loss of the photopic negative response in central retinal artery occlusion.

PURPOSE: To determine how the photopic negative response (PhNR) is affected in central retinal artery occlusion (CRAO). DESIGN: Observational case series. METHODS: Seven patients with unilateral CRAO were included. Full-field scotopic and photopic electroretinograms (ERGs) including the PhNR were recorded. Each ERG amplitude in the affected eye was expressed as a percentage of amplitude of the corresponding wave in the unaffected eye. RESULTS: Mean of the PhNR amplitude was reduced to 12.3 +/- 11.7% of that of unaffected eyes whereas the cone b-wave amplitude was attenuated to only 73.4 +/- 30.4%. This reduction of the PhNR amplitude was more significant than that of other waves including the rod b-wave, maximum a-wave and b-wave, cone a-wave and b-wave, and 30 Hz flicker ERG (P <.005). CONCLUSIONS: The PhNR was severely affected in CRAO despite relative preservation of the cone b-wave, implicating massive loss of ganglion cells and their axons.

Aged↗
PubMed2001

How patterns of bleached rods and cones become visual perceptual experiences: a proposal.

In an attempt to increase information about how mammalian visual systems create a perceptual experience out of a retinal photochemical bleach pattern, this article brings together recent rat physiological data acquired with large electrodes, an old cat behavioral experiment, and two complex human behaviors: reading and the reversible blindness people experience when the scene being viewed is stabilized on the retinal surface. The outcome suggests this juxtaposition of disparate data sets has been logical, reasonable, and informative. The link between rats and reading is the fact that both rat and human retinas convert bleach patterns into ganglion cell volleys 3 times a second. The probable trigger for these episodic retinal volleys is a more or less abrupt change in the pattern of bleached rods and cones, and we claim the absence of this trigger when the image is stabilized is responsible for the blindness. The cat behavioral experiment correlates performance on visual discrimination tasks with the number of nerve fibers remaining after lesions of the optic tract. The analysis of the result, which shows that as few as 2% of the normal number of nerve fibers supports perfect performance of such tasks, prompts the concept of a second dynamic visual system, operating in parallel with the anatomical nervous system pictured in the textbooks. The dynamic visual system model, which brings into the foreground important old facts that have been neglected and integrates them with new data, offers a synthesis that may be useful in interpreting classical visual behavioral phenomena.

Animals↗
PubMed2005

The organization of the inner retina in a pure-rod deep-sea fish.

The pure rod retina of a deep-sea eel species was used as a model system for the study of the differentiation of horizontal, bipolar, amacrine and ganglion cells. We wanted to test the hypothesis that the functional organization of the inner retina is less complex than in species with duplex, rod- and cone-containing retinae. We used immunocytochemistry, backfilling ganglion cells with fluorescent dextranes and microinjection of Lucifer Yellow, to visualize the micromorphology of the various cell types in a confocal microscope. The pure rod retina contains a single type of horizontal cell. The inner plexiform layer is 10-15 microm thick and shows three main sublayers. Bipolar terminals are found in all sublayers, but the majority are found in the inner sublamina b (PKC-immunoreactive cells, that in fish with duplex retinae receive a mixed rod-cone input). The neurochemical diversity of amacrine cells in terms of immunoreactivity does not differ from other teleosts; this similarity includes the pattern of dendritic stratification and ramification as revealed by microinjection. Ten different types of ganglion cells are distinguished based on the sizes of their perikaryon and dendritic field, and the stratification pattern in the inner plexiform layer. This is similar to the situation in catfish with retinae containing a single type of cone in addition to a majority of rods. In this respect, the differences between pure rod retinae and duplex retinae containing a single cone type were less obvious than hypothesized. In the deep-sea eel, the density of dendritic ramification in amacrine and ganglion cells was strongly reduced. This may be functionally related to the fact that vision in the deep sea environment relies exclusively on bioluminescence and is represented by burst-like emissions of point sources. This requires a mode of retinal signal processing that is less complex than in duplex retinae and involves a lower density of dendritic branching and synapses.

Animals↗
PubMed1993

Comparison of immunolocalization patterns for the synaptic vesicle proteins p65 and synapsin I in macaque monkey retina.

The distributions of the two synaptic vesicle proteins p65 [Matthew et al. (1981) J. Cell Biol., 91:257-269] and synapsin I [De Camilli et al. (1983) J. Cell Biol., 96:1337-1354] were compared in macaque monkey retina using pre-embedding immunocytochemistry for both light and electron microscopy. The monoclonal antibody AB-48 against p65 labeled ribbon-containing synaptic terminals of cone, rod, and bipolar cells as well as many conventional synapses of amacrine cells. In contrast, a polyclonal antiserum against synapsin I (SYN I) labeled many amacrine conventional synapses but no photoreceptor or bipolar ribbon synaptic terminals. Horizontal cell pre- and post-synaptic profiles in the outer plexiform layer were not labeled by either antibody. At the light microscopic level, the banding patterns in the inner plexiform layer also differed for the two antibodies, with four bands of AB-48 immunoreactivity in sublayers S1, S2, S4, and S5 but only three bands of SYN I immunoreactivity in S1, S3, and S5. SYN I also labeled varicose fibers in both the inner nuclear layer and the outer plexiform layer that are probably processes of dopaminergic and GABAergic interplexiform cells. Varicose fibers in the ganglion cell layer were labeled by both antibodies. These results provide the first electron microscopic immunocytochemical labeling for AB-48 and SYN I in intact retina and confirm that AB-48 labels both ribbon and conventional synaptic terminals, whereas SYN I labels only conventional synapses.

Animals↗
PubMed2004

Synaptic circuitry mediating light-evoked signals in dark-adapted mouse retina.

Light-evoked excitatory cation current (DeltaIC) and inhibitory chloride current (DeltaICl) of rod and cone bipolar cells and AII amacrine cells (AIIACs) were recorded from slices of dark-adapted mouse retinas, and alpha ganglion cells were recorded from flatmounts of dark-adapted mouse retinas. The cell morphology was revealed by Lucifer yellow fluorescence with a confocal microscope. DeltaIC of all rod depolarizing bipolar cells (DBCRs) exhibited similar high sensitivity to 500 nm light, but two patterns of DeltaICl were observed with slightly different axon morphologies. At least two types of cone depolarizing bipolar cells (DBCCs) were identified: one with axon terminals ramified in 70-85% of IPL depth and DBCR-like DeltaIC sensitivity, and the other with axon terminals ramified in 55-75% of IPL depth and much lower DeltaIC sensitivity. The relative rod/cone inputs to DBCs and AIIACs were analyzed by comparing the DeltaIC and DeltaICl thresholds and dynamic ranges with the corresponding values of rods and cones. On average, the sensitivity of a DBCR to the 500 nm light is about 20 times higher than that of a rod. The sensitivity of an AIIAC is more than 1000 times higher than that of a rod, suggesting that AIIAC responses are pooled through a coupled network of about 40 AIIACs. Interactions of rod and cone signals in dark-adapted mouse retinas appear asymmetrical: rod signals spread into the cone system more efficiently than cone signals into the rod system. The mouse synaptic circuitry allows small rod signals to be highly amplified and effectively transmitted to the cone system via rod/cone and AIIAC/DBCC coupling. Three types of alpha ganglion cells (alphaGCs) were identified. (1) ONGCs exhibits no spike activity in darkness, increased spikes in light, sustained inward DeltaIC, sustained outward DeltaICl of varying amplitude, and large soma (20-25 microm in diameter) with an alpha-cell-like dendritic field about 180-350 microm stratifying near 70% of the IPL depth. (2) Transient OFFalphaGCs (tOFFalphaGCs) exhibit no spike activity in darkness, transient increased spikes at light offset, small sustained outward DeltaIC in light, a large transient inward DeltaIC at light offset, a sustained outward DeltaICl, and a morphology similar to the ONalphaGCs except for that their dendrites stratified near 30% of the IPL depth. (3) Sustained OFFalpha GCs (sOFFalphaGCs) exhibit maintained spike activity of 5-10 Hz in darkness, sustained decrease of spikes in light, sustained outward DeltaIC, sustained outward DeltaICl, and a morphology similar to the tOFFalphaGCs. By comparing the response thresholds and dynamic ranges of alphaGCs with those of the pre-ganglion cells, our data suggest that the light responses of each type of alphaGCs are mediated by different sets of bipolar cells and amacrine cells.

Amacrine Cells↗
PubMedDate not supplied

Cone and rod inputs to murine retinal ganglion cells: evidence of cone opsin specific channels.

To identify ultraviolet (UV) and middle- (M) wavelength-sensitive cone and rod signals in murine retinal ganglion cells, single ganglion cell responses were studied in anesthetized, light-adapted C57/BL6 mice with tungsten microelectrodes driven through the sclera and vitreous to the neural retina. One hundred fifty-four ganglion cells were examined in 43 retinas of 34 mice. The retina was stimulated with diffuse flashes and/or pulses of ultraviolet (360 nm) or green (520 nm) light in the presence and absence of a strong steady orange adapting light. Twelve ganglion cells were studied in the dark-adapted retina in order to identify the signals of rods. Three functionally different types of ganglion cells were found: (1) phasic responding cells (31%) with no spontaneous activity and large impulse amplitudes; (2) tonic responding cells (60%) with irregular, low frequency (5-10 Hz) spontaneous activity and smaller impulse amplitudes; and (3) metronome-like cells (9%) with regular, relatively high-frequency (20-40 Hz) spontaneous activity. A few cells (1%) had habituating responses. Every cell encountered was affected by diffuse stimulation. The more common two types were excited at either the ON or OFF or at both the ON and OFF phases of stimulation. Type III cells had weaker responses, sometimes only inhibited by turning off a light. In the light-adapted state, most cells received signals of the same polarity from UV- and M-cones but UV-cone inputs were usually more dominant, especially in ventral retina. A fraction of cells received signals from only UV- (18%) or only M- (3%) cones. In rare cases (2%) these cone inputs had an opposite polarity on the same cell. In the dark-adapted state, all cells were at least four or five logarithmic units more sensitive and more to green than ultraviolet light. The results indicate that co-expression of both UV-and M-cone opsins cannot be ubiquitous in murine retina. Some cones, especially UV cones, exist without the presence of any functional M-cone opsin. This must be the case to explain the presence of ganglion cells that receive inputs only from UV-cones and others that receive inputs of opposite polarity from UV- and M-cones. The results support the hypothesis that murine retina has the physiological capacity to relay signals to the brain that allow the sensing of chromatic contrast and color vision.

Adaptation, Physiological↗
PubMed2000

Neuronal nitric oxide synthase immunoreactive neurons in the mammalian retina.

The development of immunocytochemistry has led to a better understanding of synaptic transmission carried out by neuroactive substances in the mammalian brain, including the retina. In the mammalian retina, nitric oxide (NO) is widely accepted as a neuromodulator. Histochemistry based on NADPH-d and immunocytochemistry based on nitric oxide synthase (NOS) have been used to identify the presence of nitric oxide in the mammalian retina. Certain types of amacrine cells and a class of displaced amacrine cells have been labeled consistently in all mammalian retinae studied to date. Other cell types showing NADPH-d reactivity or NOS immunoreactivity varied between species. NADPH-d reactive or NOS immunoreactive amacrine cells may serve as a source of NO for amacrine, bipolar, and ganglion cells in the inner retina, whereas interplexiform cells, bipolar cells, and horizontal cells may serve as a source of NO for the outer retina of mammals.

Animals↗
PubMed1999

Quantitative estimations of foveal and extra-foveal retinal circuitry in humans.

For an understanding of the basis for psychophysical measurement of visual resolution, quantitative morphological studies of retinal neuronal architecture are needed. Here we report on cell densities and retinal ganglion cell:cone ratio (RGC:C) from the foveal border to the peripheral retina (34 degrees eccentricity). Quantitative estimates of RGC and C densities were made using a modified disector method in three vertically sectioned human retinae and were adjusted for RGC displacement. In agreement with our previous data on humans, we found an RGC:C ratio close to 3 at 2-3 degrees eccentricity. Outside the foveal border, the ratio declined to 1.0 at 7.5 degrees eccentricity and to 0.5 at eccentricities larger than 19 degrees. Center-to-center separation of C and RGC in addition to center-to-center separation of estimated 'receptive fields' was calculated at corresponding locations along the superior and inferior hemimeridians. The center-to-center separation of estimated 'receptive fields' was found to be more closely related to resolution thresholds from the fovea to 19 degrees eccentricity than was the separation of RGC and C. On the basis of these quantitative estimates, models for neural circuitry involved in central and peripheral spatial vision can be discussed.

Cell Count↗
PubMedDate not supplied

Distribution of photoreceptor types in the retina of a marsupial, the tammar wallaby (Macropus eugenii).

Mammalian retinae generally contain low numbers of short-wavelength-sensitive cones (S-cones) and higher numbers of middle- to long-wavelength-sensitive cones (M-cones). Some recent studies found topographic differences between the different photoreceptor types and in some instances between photoreceptors and ganglion cells. To investigate this question further, we constructed topographical maps of the different photoreceptors found in an Australian marsupial, the tammar wallaby. We used two polyclonal antibodies that have been shown to label S-cones (JH455) or M-cones (JH492) in a range of mammals. In the tammar wallaby, the antisera clearly distinguish two cone types. JH455 recognizes a small subset of cones (S-cones) with a density of less than 500 cells/mm2 in the ventral retina. Their density increases towards the dorsal retina to about 1600-2000 cells/mm2. JH492 recognizes all remaining cones (M-cones), but also faintly labels most cone cells recognized by JH455. The distribution of M-cones, unlike that of the S-cones, shows a clear horizontal streak of high cell density through the central retina, just like the ganglion cells. Unlike the ganglion cells, however, the M-cones do not peak in the temporal retina but show a very broad peak (12,000-18,000 cells/mm2) in the central or even slightly nasal retina. Based on our findings, the retina of the tammar can be divided into three distinct regions: firstly, the dorsal retina, which has a low ganglion and low cone cell density but a high percentage of S-cones (30%), is thought to provide good spectral sensitivity; secondly, the central horizontal band of retina, which has a high ganglion and high cone cell density and therefore provides good spatial resolution; and thirdly, the ventral retina, which has a low ganglion cell but high cone cell density with few S-cones (5%) and is therefore thought to have a high contrast sensitivity but low acuity.

Animals↗
PubMed2002

Hammerhead ribozymes designed to cleave all human rod opsin mRNAs which cause autosomal dominant retinitis pigmentosa.

PURPOSE: Knockdown hammerhead ribozymes were designed to cleave at a sterically accessible site in long expression-competent mutant and normal human rod opsin mRNAs. Ribozyme suppression of mutant mRNA is expected to rescue autosomal dominant retinitis pigmentosa (adRP) caused by rod opsin mutations. METHODS: Energy minimization algorithms predicted regions in human rod opsin mRNA accessible to ribozyme cleavage. Opsin and ribozyme RNAs were generated by in vitro transcription. Ribozyme cleavage reactions were performed in vitro at various enzyme:substrate ratios with appropriate controls and analyzed on denaturing polyacrylamide gels. RESULTS: A GUC triplet in a predicted unhybridized loop was selected as the target cleavage site. Ribozymes were designed to stabilize catalytic core folding. Ribozyme reactions with normal and representative mutant (C187Y) opsin mRNAs demonstrated a decrease of long and short RNA targets and proportional appearance of cleavage products. Site-specific targeting was proved by lack of cleavage of a normal mRNA engineered with a silently altered cleavage motif. CONCLUSIONS: Knockdown ribozymes, targeting all known adRP mutants, cleaved human rod opsin mRNAs at the intended target site in vitro. These ribozymes may reduce total opsin mRNA and protein in rod photoreceptors as a gene therapy strategy. A beneficial outcome on rod survival in adRP is expected although normal rhodopsin levels, already in excess, would also decrease. Knockdown ribozymes attack an accessible site common to all mutant mRNAs to avoid redesign and optimization for each new dominant mutation. This strategy can be extended to any dominant disease affecting genes normally expressed in excess.

Base Sequence↗
PubMed2003

Light-evoked excitatory and inhibitory synaptic inputs to ON and OFF alpha ganglion cells in the mouse retina.

Bipolar cell and amacrine cell synaptic inputs to alpha ganglion cells (alphaGCs) in dark-adapted mouse retinas were studied by recording the light-evoked excitatory cation current (DeltaIC) and inhibitory chloride current (DeltaICl) under voltage-clamp conditions, and the cell morphology was revealed by Lucifer yellow fluorescence with a confocal microscope. Three types of alphaGCs were identified. (1) ONalphaGCs exhibits no spike activity in darkness, increased spikes in light, sustained inward DeltaIC, sustained outward DeltaICl of varying amplitude, and large soma (20-25 microm in diameter) with alpha-cell-like dendritic field approximately 180-350 microm stratifying near 70% of the inner plexiform layer (IPL) depth. (2) Transient OFFalphaGCs (tOFFalphaGCs) exhibit no spike activity in darkness, transient increased spikes at light offset, small sustained outward DeltaIC in light, a large transient inward DeltaIC at light offset, a sustained outward DeltaICl, and a morphology similar to the ONalphaGCs except for that their dendrites stratified near 30% of the IPL depth. (3) Sustained OFFalphaGCs exhibit maintained spike activity of 5-10 Hz in darkness, sustained decrease of spikes in light, sustained outward DeltaIC, sustained outward DeltaICl, and a morphology similar to the tOFFalphaGCs. By comparing the response thresholds and dynamic ranges of alphaGCs with those of the preganglion cells, our data suggest that the light responses of each type of alphaGCs are mediated by different sets of bipolar cells and amacrine cells. This detailed physiological analysis complements the existing anatomical results and provides new insights on the functional roles of individual synapses in the inner mammalian retina.

Action Potentials↗
PubMed1999

Long-term full-thickness embryonic rabbit retinal transplants.

PURPOSE: To establish the light and electron microscopic morphology of long-term full-thickness embryonic rabbit retinal transplants, with special attention paid to graft- host integration. METHODS: Eighteen rabbits received a complete embryonic neuroretina 19 days after conception. The transplants were positioned under the host retina, flat against the host retinal pigment epithelium with proper polarity, using a vitrectomy technique. After surviving 3 to 10 months, the transplants were examined by light and electron microscopy. RESULTS: The outer retina of the host had degenerated in all specimens. In 16 of the 18 eyes, well-laminated transplants with correct polarity, measuring up to 3.2 mm in length, were found. The transplants displayed long outer segments facing the host retinal pigment epithelium, and they were laminated to the level of the inner plexiform layer in which fusion with the host was often evident. Fusion was more prominent in the oldest transplants. Electron microscopy revealed bundles of neurites at different levels of maturation in close contact with Müller cell fimbriae at regular intervals along the graft-host border. CONCLUSIONS: Full-thickness embryonic rabbit retinal transplants positioned with correct polarity develop into large laminated retinas and survive without immunosuppression for at least 10 months. Host and graft adapt and almost reconstruct the normal retinal appearance. Ultrastructurally, well-developed photoreceptors and many normal synapse types are seen, and neuron sprouting is evident at the graft-host border.

Animals↗
PubMed1999

Early retinal development in the zebrafish, Danio rerio: light and electron microscopic analyses.

The morphological differentiation of the zebrafish retina was analyzed by using light (LM) and transmission electron (TEM) microscopy between the time of initial ganglion cell differentiation (approximately 32 hours postfertilization; hpf) and shortly after the point when the retina appears functional (approximately 74 hpf), i.e., when all major cell types and basic synaptic connections are in place. The results show that the inner retinal neurons, like the photoreceptor and ganglion cells, differentiate first within the ventronasal region, and differentiation subsequently spreads asymmetrically into the nasal and dorsal regions before reaching the ventrotemporal retina. In addition, we show that the attenuation of the optic stalk occurs in parallel with ganglion cell differentiation between 32 and 40 hpf. The first conventional synapses appear within the inner plexiform layer simultaneously with the first photoreceptor outer segment discs at 60 hpf; functional ribbon triads arise within photoreceptor synaptic terminals at 65 hpf; and synaptic ribbons occur within bipolar cell axon terminals at the time larvae exhibit their first visual responses (approximately 70 hpf). Although development is initially more advanced within the ventronasal region between 50 and 60 hpf, development across the retina rapidly equilibrates such that it is relatively comparable within all quadrants of the central retina by 70 hpf. An area within the temporal retina characterized by tightly packed and highly tiered cones emerges with subsequent development. Retinal differentiation in the zebrafish corresponds with that generally described in other vertebrates and can be correlated with the development of visual and electroretinographic responses in the animal.

Animals↗