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PubMed1976

Microtubules and actin filaments in teleost visual cone elongation and contraction.

Teleost retinal cones contract in light and elongate in darkness. This paper describes the disposition of microtubules and cytoplasmic filaments in cone cells of 2 species of fish (Haemulon sciurus and Lutjanus griseus). In Haemulon, the neck-like "myoid" region of the cone changes in length from 5 mu to 75 mu. Maximal observed rates of elongation and contraction are comparable to that of chromosome movement in mitosis (2-3 mu/min). Microtubules presumably participate in cone elongation, since numerous longitudinal microtubules are present in the myoid region, and colchicine blocks dark-induced elongation. Myoid shortening, on the other hand, appears to be an active contractile process. Disruption of microtubules in dark-adapted cones does not produce myoid shortening in the absence of light, and light-induced myoid shortening is blocked by cytochalasin-B. Cone cells possess longitudinally-oriented thin filaments which bind myosin subfragment-1 to form arrowhead complexes typical of muscle actin. Myoid thin filaments are clearly observed in negatively stained preparations of isolated cones which have been disrupted with detergent after attachment to grids. These myoid filaments are not, however, generally preserved by conventional fixation, though bundles of thin filaments are preserved in other regions of the cell. Thus, actin filaments are poorly retained by fixation in precisely the region of the cone cell where contraction occurs. Cone cells also possess longitudinally-oriented thick filaments 130-160 A in diameter. That these thick filaments may be myosin is suggested by the presence of side-arms with approximately 150 A periodicity. The linear organization of the contractile apparatus of the retinal cone cell makes this cell a promising model for morphological characterization of the disposition of actin and myosin filaments during contraction in a nonmuscle cell.

Actins↗

PubMed1977

Analysis of electrical noise in turtle cones.

1. Properties of the light-sensitive voltage noise in cones in the retina of the turtle, Pseudemys scripta elegans, have been studied by intracellular recording.2. Suppression of the noise by light was a function of the hyperpolarizing response of a cone but not of the size or pattern of illumination.3. Power density spectra of the noise were fitted in many cones by the product of two Lorentzians with characteristic time constants tau(1) and tau(2) averaging 40 and 7 msec respectively. The spectra of some cells were peaked and could be fitted by a resonance curve.4. Spectra in dim light exhibited decreased low frequency power. They could often be fitted by a product of two Lorentzians using the same value of tau(2) as used in darkness but decreasing tau(1) and the zero frequency asymptote. An e-fold reduction in tau(1) occurred with lights which hyperpolarized by 4-7 mV.5. Injection of hyperpolarizing currents of about 0.1-0.2 nA into weakly coupled cones reduced the noise, and also reduced the sensitivity to dim flashes.6. The variance-voltage relation during steady illumination of different intensities differed from cone to cone. Dim lights increased the noise in some cells and decreased it in others, but moderately bright lights which gave steady responses of more than about one third maximal reduced the noise in all cells.7. When the cell was transiently depolarized during the differentiated component following steady illumination, the noise was less than it was after prolonged darkness.8. In the after-effect of bright light, the time course of recovery of noise was the same as that of flash sensitivity and voltage. The noise was reduced e-fold for hyperpolarizations averaging 3 mV while for sensitivity this reduction occurred for 1.3 mV. For a given hyperpolarization the noise was lower during the after-effect than during steady dim illumination.9. When a series of dim flashes was delivered to a cone, no significant increase in variance over the dark noise was detected during the photo-response. This implies that each photoisomerization evokes no more than about 1.5 muV at the peak of the response in a coupled cone, corresponding to about 50 muV in an isolated cone.10. The elementary shot events underlying the noise are about 100 muV in amplitude in an isolated cone, have a characteristic time constant of 16-60 msec and reflect unit conductance fluctuations of about 16 pS (S, Siemen identical with Omega(-1)).11. It is concluded that the noise source is internal to the cones. We postulate that the noise arises from opening and closing of the light-sensitive ionic channels in the outer segment, and that in darkness there is a residual concentration of the blocking substance which on average closes up to about one third of the channels. It seems likely that the unit event involves a considerable number of blocking molecules and ionic channels.

Action Potentials↗

PubMed1975

Color-specific interconnections of cones and horizontal cells in the retina of the goldfish.

In Golgi preparations of goldfish retina we have observed three types of horizontal cell which receive exclusively from cones and one which receives exclusively from rods. The cone horizontal cells were designated H1, H2 and H3, in order of increasing dendritic spread, increasing separation from the outer synaptic layer, decreasing size of perikaryon, and decreasing density of cone contacts. Slender appendages with knobby terminal enlargements project horizontal cells by alalyzing serial 1 mum sections with the light microscope. The probable inputs, in terms of visual pigments in the cones which contact them, are: H1, red+green+blue; H2, green+blue; H3, blue. Analysis of previously published work suggests (1) that H1 cells generate monophasic or L-type responses, H2 cells generate biphasic or C1-type responses, and H3 cells generate triphasic or C2-type responses; (2) that H1 cells receive direct functional input at least from red-sensitive cones, H2 cells from green-sensitive cones, and H3 cells from blue-sensitive cones, and (3) that H1 constitute pathways from cones to H2 cells, and H2 cells, and H2 cells constitute pathways from cones and H1 cells to H3 cells. The precise location and route of the transfers, from H1 to H2 and from H2 to H3, are not yet known.

Animals↗

PubMed1977

Convergence of rod and cone signals in the cat's retina.

1. In an attempt to understand the convergence of rod and cone signals in the cat's retina, ganglion cells that received inputs from both rods and cones were stimulated using lights chosen to excite one or other receptor system or both together.2. If a mesopic background was chosen to allow the ganglion cell to be excited by a blue-green test flash primarily through rods and a deep red flash primarily through cones, one light could not be alternated with the other without eliciting a response from the cell.3. This appears to be a result of the different temporal properties of the scotopic and photopic systems. On the mesopic background responses to blue-green test flashes were transient. Responses to red test flashes arose with similar latency, but were more sustained.4. Rod and cone systems responded with similar latencies in the presence of the mesopic background that substantially light-adapted the rod system but left the full sensitivity of the cone system undiminished. When equivalently light-adapted, the cone system was faster.5. When brief flashes that acted through rods were presented with flashes that acted through cones the ganglion cell's response was the sum of the responses to the two flashes presented separately, as long as the flashes were weak. This linear relation ceased to hold when flashes were strong, but the breakdown appears not to be the result of mutual inhibition between rod and cone signals.6. When a background light excited both rod and cone systems it appeared to reduce sensitivity independently in each.7. The scotopic and photopic receptive fields of a given ganglion cell always were of the same type, on- or off-centre, and, within the limits of measurement, the central regions of the receptive fields were concentric and both the same size.

Action Potentials↗

PubMed1978

Suppression of cone signal in the dark-adapted frog retina as indicated by the electroretinogram.

Electroretinogram (ERG) cone acitvity is depressed in the dark-adapted frog retina. The strength of this effect is examined over a large range of flash energy, for 618 nm flashes extending up to about 4 log10 units above the "threshold" (10-25 micron V b-wave) of the cone ERG "released" in the early stage of rapid dark-adaptation (RDA). Cone signal depression is remarkably strong over this flash energy range. The cone ERG is practically absent for flashes up to about I log unit above cone RDA threshold. For stronger flashes, the suppression becomes time-dependent, that is, cone signal is very small for the first few hundred msec. after the flash, cone intrusion then becoming detectable. The results suggest that the cone suppression phenomena arises distally in the retina, probably near the receptor layer, and that cone signal intrusion a few hundred msec. after a strong flash may be due to light-adaptation of rods by the flash itself.

Animals↗

PubMed1976

Fine structure of growth cones in medullary raphe nuclei in the postnatal cat.

Morphological aspects of the dynamic processes of growth cone formation and synaptogenesis have been studied in neonatal kitten (2-17 days) medullary raphe nuclei. The formation and elaboration of dendritic growth cones and primary dendritic trunks is actively taking place on the medium size class neurons (stellates) of these nuclei. The dendritic growth cones are morphologically distinctive due to their population of large dense-core vesicles and postsynaptic position. Another growth cone morphology, interpreted as axonal, is also described. This growth cone is typically found in close association or synaptic contact with the dendritic growth cones and contains, in addition to synaptic vesicles, a dense-core vesicle population distinguishable from that of the dendritic growth cone by the presence of a variety of vesicles containing an eccentrically positioned dense particle. No evidence of axo-axonic or dendrodendritic synapses has been found. Synaptogenesis was found to be occurring on somas, dendrites and dendritic growth cones throughout the medullary raphe nuclei, though this phenomenon was more apparent in indistinctly localized subnuclear spaces termed synaptogenic zones. Within these zones large class neurons are found to have greater densities of both axodendritic and axosomatic synapses than medium and small class neurons respectively. Axodendritic synaptic densities on primary and secondary dendrites of large and medium class neurons are greater than their respective axosomatic synapse densities, which may suggest that the latter forms at a later period of development.

Animals↗

PubMed1975

Receptive field organization of ganglion cells in the frog retina: contributions from cones, green rods and red rods.

1. The impulse discharge of ganglion cells was recorded with extracellular micro-electrodes in the excised and opened eye of the common frog, Rana temporaria. 2. When a single unit was isolated, the cell type was first determined according to the Maturana, Lettvin, McCulloch & Pitts (1960) classification with the aid of varying moving and stationary stimuli. 3. Class 4 cells respond only to a decrease of light when cones are stimulated but respond to an increase of light when green rods are stimulated. A distinct class of deviating class 4 cells was found that give a brief high frequency burst at 'off' from their small excitatory receptive fields (ERF); unlike typical class 4 cells they possess a purely inhibitory surrounding field (IRF).4. The contributions from the cones and the green and red rods were isolated by measuring the thresholds of the discharges with on-off stimuli of varying wave-lengths against strong yellow backgrounds, or against a very weak background or no background at all. The spatial distribution of the contributions to the ERF was determined by mapping threshold profiles, and additional information about ERF and IRF was obtained from area-threshold curves. 5. The cone-mediated ERFs were found to be 0-06-0-50 mm wide (1-5-12 degrees of visual field), which agrees well with the sizes of the dendritic trees of the ganglion cells. The green rod-mediated ERFs can be 0-5-1-5 mm wide and have less distinct boundaries than the cone-mediated. The green rod-mediated ERF of an individual ganglion cell is always larger than the cone-mediated ERF of the same cell. The red rod-mediated ERFs seem to be somewhat larger than the cone-mediated but smaller than the green rod-mediated. 6. The green rods contribute only to the on thresholds of class 1, 2 and 4 cells, but both to on and off in typical class 3 cells, while the cones contribute to on and off in classes 1-3 and only to off in class 4.7. When the red rods begin to contribute during dark adaptation they seem to enter the cone but not the green rod channels. 8. All three receptor types contribute to the IRF surrounding the ERF of classes 1, 2, 3 and deviating class 4 cells. Normal class 4 cells have no IRF. 9. The organization of the receptive fields is discussed in relation to the anatomy and electrophysiology of the cell types transmitting the signals from the receptors to the ganglion cells.

Action Potentials↗

PubMed1978

Influences of cones upon chromatic- and luminosity-type horizontal cells in pikeperch retinas.

1. The spectral sensitivity and spatial organization of cones and horizontal cells have been analysed by intracellular recording in pikeperch retinas. 2. The vast majority of cone recordings were obtained from orange-sensitive cones. They have an action spectrum which peaks at about 605 nm. Recordings from several green-sensitive cones have also been obtained. 3. The results of action spectrum measurements and spectral screening tests indicate that the vast majority of luminosity-type horizontal cells receive predominant input from the orange-sensitive cones. 4. Chromatic-type horizontal cells were recorded at more proximal levels of the retina than luminosity-type cells and were the classic red-depolarizing, green hyperpolarizing (R/G) type. 5. The action spectra of the depolarizing and hyperpolarizing responses of chromatic horizontal cells peak at about 650 and 530 nm, respectively. When the depolarizing mechanism is selectively depressed by a red background field, the action spectrum of the hyperpolarizing mechanism shows an enhanced sensitivity, peaks at 530--540 nm, and may approximate the action spectrum of the green-sensitive cones. 6. Small red fields evoke depolarizing responses from chromatic-type horizontal cells but do not seem to significantly activate the depolarizing surround mechanism of cones. 7. These and other results suggest that the colour-opponent properties of the chromatic-type horizontal cells are not fundamentally dependent upon feed-back to cones but rather originate from antagonistic interactions generated in post-receptor networks.

Action Potentials↗

PubMed1979

The effects of background illumination on the photoresponses of red and green cones.

1. The photoresponses of light- and dark-adapted red and green cone photoreceptors were recorded intracellularly in the retina of the turtle, Pseduemys scripta elegans. Background illumination produced similar effects on both types of cones. 2. In response to the onset of a prolonged, steady background illumination the cone initially hyperpolarized to a peak which then sagged back to a steady-state polarization that was typically about one half the initial peak amplitude. This sag was observed for all backgrounds studied (dim as well as bright). 3. A resensitization was observed concomitantly with this sag; both the maximum increment and decrement responses grew in amplitude as light-adaptation proceeded. After about 2--3 min of background illumination, the amplitudes of these responses stabilized. 4. The dark-adapted cone produced graded responses to test pulses over a range of intensities spanning about 3.5 log units. The amplitudes of these responses were well fit by the relationship V = I.Vm/(I + sigma). 5. After 2--3 min of background illumination, 500 msec test pulses either brighter or dimmer than the background intensity were substituted for the background. The light-adapted intensity-response curves constructed from this data were similar to the dark-adapted curve but were shifted horizontally and slightly vertically, so that they still spanned about 3.5 log units of intensity. Thus, in the light-adapted cone, graded responses were elicited by a range of bright test pulses which would have produced saturated responses when delivered to the dark-adapted cone. 6. The 'off response' observed at the offset of the background became faster as the background intensity was increased. It also became faster with time following the onset of any particular background intensity. 7. It was concluded that cone sensitivity during any state of light-adaptation is determined by two mechanisms; response compression resulting from the instantaneous non-linearity between 'internal transmitter' concentration and membrane potential and a more active 'cellular adaptation' mechanism which is manifest as a shift in the intensity-response curve. In the steady-state condition of light-adaptation, most of the sensitivity changes are a result of the cellular adaptation mechanism. 8. Photopigment bleaching caused by the backgrounds, negative feed-back from horizontal cells and voltage dependent mechanisms in the cones could not account for this cellular adaptation. These effects of background illumination were interpreted in terms of the 'internal transmitter' hypothesis of phototransduction.

Adaptation, Ocular↗

PubMed1978

Mammalian cones: disc shedding, phagocytosis, and renewal.

During the past several years we have examined a variety of different mammalian retinas for ultrastructural evidence of cone disc shedding and RPE phagocytosis. In this paper we review our previously published evidence from squirrel and human retinas as well as present new evidence of cone disc shedding in rhesus monkey and cat. All these species show definitive evidence for the shedding of discs from cone outer segments and the phagocytosis of shed discs by apical processes of the RPE; both of these events closely resemble those described for mammalian rods. The occurrence of cone disc shedding leads directly to the conclusion that new membrane must be added to the cone outer segment in order to maintain its length. The successive evaginations, which are observed at the bases of cone outer segments, we consider to be indirect evidence for the addition of new discs. Finally, we propose a model for the structural organization of mammalian cone outer segments.

Animals↗

PubMed1977

Blue and green cone mechanisms in retinitis pigmentosa.

Pronounced abnormalities in the increment threshold curves for the blue and green cone mechanisms only 10 degrees above the fovea, and in some cases in the fovea as well, have been found with a 2.5 degrees stimulus in different genetic types of retinitis pigmentosa. Whenever cone thresholds were abnormal, thresholds determined by the blue cone mechanism (pi1 and pi3) were more elevated than those determined by the green cone mechanism (pi4). Thresholds determined by the green cone mechanism were consistently more elevated at low background intensities than at intermediate and high background intensities. Threshold elevations for the blue and green cone mechanisms relative to each other in the perifovea in a given patient tested with the 2.5 degrees stimulus could be simulated in normal observers tested with a single small stimulus. These findings support the idea that the summation pools for blue and green cone mechanisms are proportionally reduced below normal in retinitis pigmentosa, at least in the perifovea, and are compatible with the idea that both mechanisms are comparably involved at the photoreceptor level.

Adolescent↗

PubMed1979

Rods and cones in the mouse retina. II. Autoradiographic analysis of cell generation using tritiated thymidine.

The period of cell genesis of rod and cone photoreceptor cells has been determined in the retinas of C57BL/6J mice. Embryonic mice were exposed to a single dose of 3H-thymidine at embryonic day (E) 10--18 by injecting pregnant mice intraperitoneally. Animals at postnatal ages were injected subcutaneously once between postnatal day (P) 0--10. The eyes were removed at one to three months of age. After fixation, they were embedded in glycol methacrylate, sectioned at 1.5 micrometers and prepared for autoradiographic analysis. All of the cone cells are generated over a relatively short time interval during the fetal period. In the posterior retina, the peak of cone cell genesis occurs at E13-E14, and no cones are generated after E16. The rods, by contrast, are generated later and over a longer time period. They first begin to be generated in the posterior retina on E13, but the peak of cell genesis is not reached until the day of birth, and some rods are generated as late as P5. For both rods and cones the peaks of cell genesis in the peripheral retina occur two to three days later than in the posterior retina. The findings demonstrate that rods and cones are developmentally distinct cell types in the mouse retina.

Animals↗

PubMed1977

The red and green cone visual pigments of deuternomalous trichromacy.

1. Three "simple" deuteranomalous trichromats match with abnormally low "red" tristimulus values throughout the spectrum and abnormally high "green" tristimulus values in the long wave end of the spectrum which become normal (and then low) in the yellow-green. The spectrum locus of this transition differs from one anomalous to the other. Differences in the matches of two of these cannot be due to differences in eye media transmissivities alone. Therefore these two deuteranomalous have different cone visual pigments. 2. The analytical anomaloscope was used in the confrontation of one deuteranomalous with six deuteranopes in turn. In each confrontation the deuteranope set the anomaloscope in his mode and adjusted the intensity of the monochromatic light for a match. Deuteranomalous matches were rejected by four of these six deuteranopes. 3. They were accepted by two of the six. These two rejected each other's matches in a way not attributable to differences in eye media transmissivity. 4. Three different psychophysical techniques were used to measure the action spectra of the long wave cones of these two deuternopes. All three methods reveal small but systematic differences in lambdamax and shape of the curve for the one deuteranope compared with that of the other. 5. In red-green spectral range, these spectra are accurately described by different linear combinations of the color matching functions of the same deuteranomalous whose matches the two deuteranopes accept. Linear combinations of those of a second deuteranomalous, with at least one different kind of cone, fit less well. 6. The wave length discrimination curve of the former deuteranomalous was measured with a new method. The curves of two normals were also obtained for comparison. Wave-length discrimination predictions from the Stiles (1946) line element theory were compared to the anomalous curve. The deuteranopic action spectra were used in the line element to compute this deuteranomalous' discrimination. There is reasonable first order correspondence between prediction and observation, but the prediction is sensitive to small changes in the derivatives of the logarithms of the action spectra. 7. Line element prediction of the deuteranomalous step-by-step luminous efficiency curve is insensitive to such uncertainties. The agreement with expectation from the above assumptions and the measured step-by-step deuteranomalous luminous efficiency curve in the red-green part of the spectrum is therefore good. 8. It is concluded that the erythrolabe in one deuternope's long wave cones has the action spectrum of this deuteranomalous' long and the erythrolabe in the other deuternope's long wave sensitive cones has that of this deuternomalous' medium wave cones. This leads to a general hypothesis about the nature of all forms of red-green colour vision defects transmitted recessively on the X chromosome.

Color Vision Defects↗

PubMed1978

The visual pigments of rods and cones in the rhesus monkey, Macaca mulatta.

1. New microspectrophotometric measurements have been made of the photo-pigments of individual rods and cones from the retina of the rhesus monkey (Macaca mulatta). The measuring beam was passed transversely through isolated outer segments. 2. The transverse absorbance for rods ranged from 0.02 to 0.04 and that for cones from 0.01 to 0.03. 3. The mean absorbance spectrum for rods (n = 25) had a peak of 502 +/- 2.7 nm. A digitonin extract from the same group of eyes gave a lambda-max. of 499 +/- 1 nm. 4. Of a sample of 82 cones, 40 were 'red' (P565 nm) and 42 were 'green' (P536 nm). The mean absorbance spectrum for the green cones is very similar to the Dartnall nomogram, but that for the red cones is narrower. 5. No bleachable, blue-sensitive outer segments were recorded, although structures were found that absorbed at short wave-lengths and were neither photosensitive nor dichroic. 6. If the long wave-length and middle wave-length cone pigments of the rhesus monkey are assumed to be identical to those of man and if additional assumptions are made about the lengths of human outer segments and about prereceptoral absorption, it is possible to derive psychophysical sensitivities that closely resemble the pi5 and pi4 mechanisms of W. S. Stiles.

Animals↗

PubMed1975

Goldfish retina: functional polarization of cone horizontal cell dendrites and synapses.

In serial electron micrographs we observed that dendrites of goldfish cone horizontal cells are either central or lateral in ribbon synaptic triads, depending on cone and horizontal cell type. The chromatic properties of cone horizontal cell responses may be explained if the cone horizontal cells act as interneurons, receiving from cones through their central processes but acting on cones through their lateral processes.

Animals↗

PubMed1977

Responses and receptive-field organization of cones in perch retinas.

1. Cones in the retinas of two closely related species of perch, the walleye and sauger (S, vitreum vitreum and S. canadense), are remarkably large. This paper reports a first series of intracellular recordings obtained from 77 of these cones. 2. A small spot of light evokes a sustained hyperpolarizing response from perch cones which may exceed 10 mV in amplitude, is graded with stimulus intensity, and is markedly reduced when the spot is decentered. Most cones seem to be orange sensitive with peak sensitivity at about 600 nm. 3. Enlarging the stimulus diameter from 0.04 to 0.25 mm produces a modest increase in the hyperpolarizing response. However, larger stimuli which illuminate surrounding regions of the retina often evoke a delayed depolarizing potential which antagonizes the sustained phase of the cone's hyperpolarizing response to central illumination. 4. The outer diameter of the region of the antagonistic surround is at least 2.2 mm in extent. An annulus evokes a depolarizing response only if the central region of the receptive field is simultaneously activated. 5. The present results provide the first direct evidence that the receptive fields of cones in fish retinas have an antagonistic center-surround organization. Luminosity-type horizontal cells probably serve as the interneurons which mediate the depolarizing influence of the surround.

Animals↗

PubMed1979

Rod sensitivity relative to cone sensitivity in retinitis pigmentosa.

The present study evaluates rod sensitivity relative to cone sensitivity in retinitis pigmentosa patients. From perimetric measures of absolute thresholds to long- and short-wavelength stimuli, we find the results can be characterized in three different ways. For one group of patients it appears that only cone function is present in all parts of the visual field. For another group of patients, despite large losses in absolute sensitivity, rod sensitivity still exceeds cone sensitivity by at least the normal factor, suggesting concomitant rod and cone sensitivity losses. For a third group of patients it appears that central retinal thresholds are determined primarily by cones, midperipheral retinal thresholds determined by a composite of rod and cone function, and far- peripheral thresholds determined predominately by rods. The argument is put forward that the different groups of threshold results represent different disease mechanisms rather than different stages in the progression of a single disease process.

Adult↗

PubMed1979

Blue-sensitive cones of the cat produce a rodlike electroretinogram.

Two cone mechanisms are identifiable in the strongly yellow light-adapted electroretinogram (ERG) of the arterially perfused cat eye. One has its maximum spectral sensitivity near 555 nm; the other has its maximum near 450 nm. The former cone system produces a much larger signal with characteristics of a typical cone or inhibitory ERG. The latter cone system produces a small, saturable signal (less than 5 microV) which resembles a rodlike or excitatory ERC. The results imply that the latter ERG is generated by blue-sensitive cones, which form a small fraction of the total cone population and share some physiological and perhaps anatomical properties of rods.

Animals↗