Analysis and Synthesis of the Dynamic Response of Retinal Neurons

theory of linear systems analysis is developed in a form directly applicable to the treatment of the Limulus retina. The dynamics of the retina may conveniently be characterized by means of a spatiotemporal transfer function, which summarizes the response of the system to moving sinusoidal gratings ( analysis ). The response of the retina to an arbitrary stimulus may then be calculated by addition of the response to suitably weighted sinusoidal stimuli ( synthesis ). Responses were obtained from the in-situ retina by means of extracellular recording of impulse activity in single optic nerve fibers. Test ommatidia were chosen in the interior of the retina, to avoid asymmetries introduced by the edge of the retina. Stimuli which varied in both space and time were produced under computer control on the screen of a display oscilloscope, and were conveyed to the Limulus eye by means of a fiber-optic taper. Transfer functions were measured using counterphase modulation of cosine gratings according to a sum-of-sinusoids temporal signal, a procedure equivalent to the use of moving gratings, for ommatidia with symmetrical receptive fields. By means of these transfer functions, the responses of the Limulus eye to visual stimuli moving at various velocities were predicted in a parameter-free Fourier synthesis calculation. There was good agreement between these predictions and the measured responses to these stimuli. A quantitative model for the dynamic, integrative action of the Limulus retina is developed, based on the original formulation for the steady state given by the Hartline- Ratliff equations. The model comprises an excitatory generator potential, and dynamic processes of self and lateral inhibition. An explicit expression for the spatiotemporal transfer function is obtained in terms of transfer functions for the generator potential, self-inhibitory, and lateral-inhibitory transductions, and spatial transforms of the lateral inhibitory kernel and the point-spread characteristic of the experimental and physiological optics. Explicit functional forms for these component transductions are adopted. The parameters which occur in these expressions serve to incorporate information about the subcellular physiology of retinal neurons into the quantitative description of the function of the retina as a whole. Procedures are described for the estimation of these parameters from empirical transfer function data. Transfer functions calculated from the model on the basis of parameters obtained with these procedures show good agreement with the corresponding empirical transfer functions. The parameter values obtained in this way are, in general, quite consistent with the results of many more direct (and frequently more invasive) measurements reported in the literature. In particular, the inhibitory kernel, as determined from our transfer function measurements, shows a small crater in the vicinity of the test-ommatidium. The dynamical model can be used to describe the response of the retina in the vicinity of its boundary, as well as in the interior. An analysis, based on the Wiener-Hopf technique, is given for the response of peripheral retinal neurons. The predictions derived from this theory were compared with experiment through the use of illumination patterns in which one half of the retina was kept in darkness, while the remaining half was presented with a moving stimulus. This procedure permitted the calibration of model transfer functions by means of methods appropriate only for interior ommatidia, while simulating the neural environment at the edge of a homogeneous retina. Significant differences between the responses to stimuli which moved toward and away from the simulated edge were observed experimentally, in good agreement with the predictions of the theory. Similar behavior was also observed at the actual anatomical boundary of the eye

Temporal Changes in the Ommatidial Structure of the Cockroach, Leucophaea Maderae

A circadian rhythm in eye sensitivity to light has been previously reported for Leucophaea maderae. Temporal changes in eye cell morphology that could be correlated with those changes in eye sensitivity to light were examined. Rhabdom area, screening pigment organization and palisade layer area about the rhabdom were the parameters measured to detect structural change through time. Measurements of those parameters from tissue samples obtained from the anterior one-third of compound eyes surgically removed at midday, light offset, midnight and light onset from roaches entrained to a 12-h light / 12-h dark photoperiodic cycle were used to assess the daily pattern of morphological changes. Eyes were removed at subjective midday and subjective midnight from roaches free-running under constant conditions of temperature and darkness to detect circadian changes. All roaches received food and water ad libitum. Tissue samples were fixed, embedded, sectioned and the sections were examined and photographed using a Zeiss transmission electron microscope to test for time-related morphological differences. The extent of pigment organization was determined by counting the number of pigment granules found within a 10µm diameter circle centered about the rhabdom. The rhabdom area and the palisade layer area were determined by the Jandel PC3-D computer program. The rhabdom area did not vary throughout the day. The organization of screening pigment granules and the palisade layer area did vary on a daily basis. In animals maintained under constant environmental conditions the rhythm of the organization of the pigment granules did not persist. In contrast, temporal changes in the palisade layer area did persist for three cycles with a pattern similar to that in roaches held under LD12:12 and thus could be considered a circadian rhythm controlled by a pacemaker

Coexpression of Spectrally Distinct Rhodopsins in Aedes aegypti R7 Photoreceptors

The retina of the mosquito Aedes aegypti can be divided into four regions based on the non-overlapping expression of a UV sensitive Aaop8 rhodopsin and a long wavelength sensitive Aaop2 type rhodopsin in the R7 photoreceptors. We show here that another rhodopsin, Aaop9, is expressed in all R7 photoreceptors and a subset of R8 photoreceptors. In the dorsal region, Aaop9 is expressed in both the cell body and rhabdomere of R7 and R8 cells. In other retinal regions Aaop9 is expressed only in R7 cells, being localized to the R7 rhabdomere in the central and ventral regions and in both the cell body and rhabdomere within the ventral stripe. Within the dorsal-central transition area ommatidia do not show a strict pairing of R7–R8 cell types. Thus, Aaop9 is coexpressed in the two classes of R7 photoreceptors previously distinguished by the non-overlapping expression of Aaop8 and Aaop2 rhodopsins. Electroretinogram analysis of transgenic Drosophila shows that Aaop9 is a short wavelength rhodopsin with an optimal response to 400–450 nm light. The coexpressed Aaop2 rhodopsin has dual wavelength sensitivity of 500–550 nm and near 350 nm in the UV region. As predicted by the spectral properties of each rhodopsin, Drosophila photoreceptors expressing both Aaop9 and Aaop2 rhodopsins exhibit a uniform sensitivity across the broad 350–550 nm light range. We propose that rhodopsin coexpression is an adaptation within the R7 cells to improve visual function in the low-light environments in which Ae. aegypti is active