Parallel mechanisms encode direction in the retina

In the retina, presynaptic inhibitory mechanisms that shape directionally selective (DS) responses in output ganglion cells are well established. However, the nature of inhibition-independent forms of directional selectivity remains poorly defined. Here, we describe a genetically specified set of ON-OFF DS ganglion cells (DSGCs) that code anterior motion. This entire population of DSGCs exhibits asymmetric dendritic arborizations that orientate toward the preferred direction. We demonstrate that morphological asymmetries along with nonlinear dendritic conductances generate a centrifugal (soma-to-dendrite) preference that does not critically depend upon, but works in parallel with the GABAergic circuitry. We also show that in symmetrical DSGCs, such dendritic DS mechanisms are aligned with, or are in opposition to, the inhibitory DS circuitry in distinct dendritic subfields where they differentially interact to promote or weaken directional preferences. Thus, pre- and postsynaptic DS mechanisms interact uniquely in distinct ganglion cell populations, enabling efficient DS coding under diverse conditions.     

Cellular mechanisms for direction selectivity in the retina

Direction selectivity represents a fundamental computation found across multiple sensory systems. In the mammalian visual system, direction selectivity appears first in the retina, where excitatory and inhibitory interneurons release neurotransmitter most rapidly during movement in a preferred direction. Two parallel sets of interneuron signals are integrated by a direction-selective ganglion cell, which creates a direction preference for both bright and dark moving objects. Direction selectivity of synaptic input becomes amplified by action potentials in the ganglion cell dendrites. Recent work has elucidated direction-selective mechanisms in inhibitory circuitry, but mechanisms in excitatory circuitry remain unexplained.     

Cholinergic mechanisms of interneuronal transmission in the retina

Acetylcholine, applied to the isolated perfused frog and human retina, induces a corneopositive potential. This electrogenic action of acetylcholine, in conjunction with existing data, confirms the view that transmission in the retina is cholinergic. The magnitude and the temporal course of the potential evoked by acetylcholine depends both on its concentration and on the state of adaptation of the retina. Photic stimulation reduces the response to acetylcholine; under these circumstances flashes are more effective than a steady illumination. On the other hand the response of the retina to light decreases during perfusion with acetylcholine. The positive component of the ERG is particularly strongly inhibited, leaving only the negative P_III. The results indicate that acetylcholine acts on synapses between the first and second retinal neurons. They can be explained in terms of the hypthesis of the desensitization of cholinergic receptors in the retina.     

Multiple mechanisms for contrast adaptation in the retina

The retina adapts to average light intensity but also to the range of light intensities (contrast). A study by Baccus and Meister, in this issue of Neuron, identifies three ways that ganglion cells and interneurons adapt to high contrast: shorten integration time, reduce gain, and depolarize. Only the depolarization decays, over tens of seconds.     

Cone photoreceptor reflectance variation in the northern tree shrew and thirteen-lined ground squirrel

In vivo images of human cone photoreceptors have been shown to vary in their reflectance both spatially and temporally. While it is generally accepted that the unique anatomy and physiology of the photoreceptors themselves drives this behavior, the exact mechanisms have not been fully elucidated as most studies on these phenomena have been limited to the human retina. Unlike humans, animal models offer the ability to experimentally manipulate the retina and perform direct in vivo and ex vivo comparisons. The thirteen-lined ground squirrel and northern tree shrew are two emerging animal models being used in vision research. Both models feature cone-dominant retinas, overcoming a key limitation of traditional rodent models. Additionally, each possesses unique but well-documented anatomical differences in cone structure compared to human cones, which can be leveraged to further constrain theoretical models of light propagation within photoreceptors. Here we sought to characterize the spatial and temporal reflectance behavior of cones in these species. Adaptive optics scanning light ophthalmoscopy (AOSLO) was used to non-invasively image the photoreceptors of both species at 5 to 10 min intervals over the span of 18 to 25 min. The reflectance of individual cone photoreceptors was measured over time, and images at individual time points were used to assess the variability of cone reflectance across the cone mosaic. Variability in spatial and temporal photoreceptor reflectance was observed in both species, with similar behavior to that seen in human AOSLO images. Despite the unique cone structure in these animals, these data suggest a common origin of photoreceptor reflectance behavior across species. Such data may help constrain models of the cellular origins of photoreceptor reflectance signals. These animal models provide an experimental platform to further explore the morphological origins of light capture and propagation.     

Two mechanisms for spreading depression in the chicken retina

Two mechanisms have been proposed to explain spreading depression (SD): one based on a release of glutamate (Van Harreveld, 1959), and the other on a release of potassium (Grafstein, 1956) from neuronal elements. Both glutamate and KCl cause transparency changes in the retina, comparable to those occurring in this tissue during SD. The glutamate effect is inhibited by MgCl₂ (10 mM), in contrast to the transparency change due to KCl which is not affected by Mg⁺⁺. Also SD is usually inhibited by MgCl₂ which suggests that such SDs are based on a glutamate release. Impairment of the tissue metabolism promotes SDs which are insensitive to MgCl₂. The resulting failure of the mechanisms that transport K⁺ and glutamate which leak out of the intracellular compartment back into the cells and fibers, seems to be involved in the generation of Mg⁺⁺ insensitive SDs. This may facilitate either K-based SDs or glutamate-based SDs since the inhibitory effect of Mg⁺⁺ is counteracted by an enhanced glutamate concentration. Both proposed mechanisms for SD seem to be possible under special circumstances.     

Water involvement in the mechanisms of retina electrical activity

Retina is an excitable system containing approximately 90% water. As we found that deuteration selectively changes amplitudes and latencies of retina biopotentials, specifically the ON and OFF responses, we used it to probe the role of water in those processes. A study of the retina deuteration kinetics was simultaneously performed. This revealed the existence of at least two retinal water compartments. The data suggested a third compartment also, with a lower motional "degree of freedom," existing where H2O-D2O exchange becomes important only after saturation by D2O of the first two compartments. Correlation of the electrophysiological effects of D2O with the kinetic data suggests that the ON response is related to the first water compartment and the OFF response to the third. The results point to independence on the ON and OFF response mechanisms and, very probably, to their different morphological origins.     

Regulatory mechanisms in melatonin biosynthesis in retina.

The vertebrate retina produces melatonin in a light-dependent rhythmic fashion, synchronized with, but independent from the rhythm of the hormone formation in the pineal gland. This review summarizes the current status of our knowledge on regulatory mechanisms involved in controlling the retinal melatonin biosynthesis. Special emphasis is given to the role and mode of action of dopamine and GABA, two established retinal neurotransmitters, as well as that of second messengers (cyclic AMP, calcium ions). Comparisons are made between lower vertebrates and mammals.     

Adrenoceptor mechanisms in the cardiovascular effects of cocaine in conscious squirrel monkeys.

The purpose of the current experiment was to study the role of various adrenoceptor subtypes in the cardiovascular response to cocaine in conscious squirrel monkeys. A variety of adrenoceptor antagonists were administered i.v. prior to the administration of 0.3 mg/kg cocaine (i.v.). Cocaine alone produced an increase in both blood pressure and heart rate. The non-selective alpha adrenoceptor antagonist phentolamine produced a dose-dependent antagonism of the pressor effect of cocaine, as did the alpha-1 selective antagonist prazosin. The alpha-2 selective antagonist yohimbine had no effect on the pressor effect of cocaine. The non-selective beta antagonist propranolol enhanced the pressor effect of cocaine as did the beta-1 selective antagonist atenolol. However, the effect of atenolol was not dose-dependent. The beta-2 selective antagonist ICI 118,551 and labetalol, which blocks both alpha and beta adrenoceptors, did not alter the pressor effect of cocaine. Propranolol, atenolol, and labetalol all antagonized the tachycardiac effect of cocaine in a dose-dependent manner, while the beta-2 antagonist ICI 118,551 did not. Phentolamine, prazosin and yohimbine also reduced the tachycardiac effect of cocaine, although these effects were dose-dependent only for yohimbine, which also significantly elevated baseline heart rate. These results indicate that alpha-1 adrenoceptor mechanisms mediate the pressor effect of cocaine, while beta-1 adrenoceptor mechanisms are involved in the tachycardiac effect of cocaine in squirrel monkeys. Propranolol potentiated cocaine's pressor effect through beta-2 independent mechanisms. Thus, neither alpha-2 nor beta-2 adrenoceptor mechanisms appear to be involved in cocaine's cardiovascular effects.     

Energetic costs of the winter arboreal microclimate: The gray squirrel in a tree

Heated taxidermic mounts of the gray squirrel were used to analyze the thermal environment of a small arboreal endotherm. Changes in the standard operative temperature (T _es) calculated from the temperatures of heated and unheated mounts agreed well with the power consumption (M–E) of mounts on the ground and on the wind-ward side of a 48-cm diameter tree trunk. As wind speed (u) rose and sky solar radiation (Q _r) decreased, the windward side of the tree trunk became an increasingly more stressful thermal environment than the leeward side of the trunk or the ground, producingM–E differences of more than 30%. Although theM–E of a ground mount and a limb mount 4 m in the air are dependent onQ _ras well asu, the ratio of the two value ofM–E is independent ofQ _r, poorly predicted byu and well predicted byu ^1/2.     

Receptive field mechanisms of ganglion cells in the cat retina

On the basis of anatomical and physiological results of the vertebrate retina, a method is proposed for analysing the respective fields of ganglion cells in the cat retina. In the model, we assume the following: (a) Ganglion cells receive their input from bipolar and/or amacrine cells. (b) The nonlinearity of ganglion cell responses is due to the activities of transient type amacrine cells. The method has been proved to be effective. According to the results of this investigation, the receptive field properties of X type and Y type ganglion cells are heterogeneous. Thus, it may be considered that their receptive fields consist of center and surround mechanisms. The receptive field properties of X-cells are almost linear and the X-cells seem to receive most of their input from bipolar cells. On the other hand, the ones of Y-cells are highly nonlinear. Consequently, it is conceivable that the Y-cells receive their input mainly from transient type amacrine cells.     

Structural organization of the retina in the tree shrew (Tupaia glis)

This investigation was undertaken to establish the gross and ultrastructural organization of the photoreceptors and retina in the Malayan tree shrew (Tupaia glis). Photographs of the fundus revealed no specialization or differentiation of a central foveal region. Histologic sections revealed a single row of relatively short and thick cones distributed uniformly throughout the retina. Electron micrographs of the retina indicated that the receptor outer segments are closely invested by pigment-filled epithelial processes and an amorphous interstitial material. The internal fine structure of the receptor outer segments revealed the characteristic stacks or arrays of bimembranous discs. The ellipsoid portions of the cone inner segments include tightly packed and extraordinarily large mitochondria. These mitochondria consist of unique patterns of concentric cristae arranged in highly ordered whorls of lamellar configurations. The cone synaptic pedicles contain a unique system of tubules not previously described in synaptic endings. Histologic sections indicated that only cone populations are located in the central region of the retina, whereas histologic, histochemical, and ultrastructural comparisons suggested that photoreceptors with some "rodtype" features are located more peripherally. The relatively small proportion of these rodtype receptors among the great preponderance of cone populations is in general accord with the tree shrew's diurnal habits as well as its great reliance on photopic vision and its visually guided behavior.     

Spectral sensitivity of ground squirrel cones measured with ERG flicker photometry

Summary Ground squirrels have dichromatic color vision. The spectral sensitivities of the two classes of cones found in the retinas of two species of ground squirrel were measured using ERG flicker photometry. The spectral sensitivity curves for these cone classes were closely fit by curves from wavelength-dependent visual pigment nomograms. One cone type had an average peak sensitivity of 518.9 nm (California ground squirrels,Spermophilus beecheyi) or 517.0 nm (thirteen-lined ground squirrels,Spermophilus tridecemlineatus). The second type of cone found in these ground squirrels had an average peak sensitivity of 436.7 nm. An examination of the variation in spectral sensitivity among individual animals suggests that the sensitivity peaks for the middle-wavelength cone cover a range of not greater than 4 nm.     

Spectral inference reveals principal cone-integration rules of the zebrafish inner retina

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Candidate molecular mechanisms for establishing cell identity in the developing retina

In the developing nervous system, individual neurons must occupy appropriate positions within circuits. This requires that these neurons recognize and form connections with specific pre- and postsynaptic partners. Cellular recognition is also required for the spacing of cell bodies and the arborization of dendrites, factors that determine the inputs onto a given neuron. These issues are particularly evident in the retina, where different types of neurons are evenly spaced relative to other cells of the same type. This establishes a reiterated columnar circuitry resembling the insect retina. Establishing these mosaic patterns requires that cells of a given type (homotypic cells) be able to sense their neighbors. Therefore, both synaptic specificity and mosaic spacing require cellular identifiers. In synaptic specificity, recognition often occurs between different types of cells in a pre- and postsynaptic pairing. In mosaic spacing, recognition is often occurring between different cells of the same type, orhomotypic self-recognition. Dendritic arborization can require recognition of different neurites of the same cell, or isoneuronal self-recognition. The retina is an extremely amenable system for studying the molecular identifiers that drive these various forms of recognition. The different neuronal types in the retina are well defined, and the genetic tools for marking cell types are increasingly available. In this review we will summarize retinal anatomy and describe cell types in the retina and how they are defined. We will then describe the requirements of a recognition code and discuss newly emerging candidate molecular mechanisms for recognition that may meet these requirements.     

Studies on the mechanisms underlying horizontal-bipolar interaction in the carp retina.

Responses of on-center bipolar cells and horizontal cells were recorded simultaneously in the carp retina, and the effect of polarization of horizontal cells on the bipolar cells was studied. Hyperpolarization by extrinsic current of horizontal cells elicited in the bipolar cells a hyperpolarizing response which, unlike the electrical coupling betweeen adjacent horizontal cells, was accompanied by a change in membrane conductance. The bipolar cell responses elicited by polarization of external horizontal cells showed a negative reversal potential, while those elicited by polarization of intermediate horizontal cells showed a positive reversal potential. It was suggested that the external horizontal cells modify the cone-bipolar transmission which involves the conductance change of subsynaptic potassium and/or chloride channels, while the intermediate horizontal cells modify the rod-bipolar transmission which involves the conductance change of sodium channels.