Summary
Intracellular recording and dye injection (Procion Yellow) techniques were applied to a set of nine giant, homolateral cells of the lobula plate of dipterans (Phaenicia, Sarcophaga —the “big vertical cells” (or V-cells) of Pierantoni (1974).
Anatomical Findings
The V-cells can be divided into 4 distinctly different groups which, in principle, agree with the classification (though into 3 groups) given by Strausfeld (1976 a). In each case, the cell body lies in the caudal cell body layer of the optic pedunculus with the cell axon extending at the caudal surface of the lobula plate. The axons run through the optic pedunculus and terminate in the ventrolateral part of the protocerebrum. The large bifurcated dendrites and their small dendritic branches form a fan which lies in a frontal plane perpendicular to the columnar arrangement of this ganglion and penetrate the whole dorso-ventral region of the lobula plate. The highest density of dendritic branches is found in the anterior part of the retino-topic projection. It decreases gradually towards the projection of the posterior retina in the medial lobula plate. The dendritic field widths (Fig. 9) extend approximately 60 degrees in the horizontal and 200 degrees in the vertical direction; thus, there is a considerable degree of overlap in the horizontal direction. The posterior V-cells V6 to V8 possess a more extended horizontal field width in the dorsal part of the eye. These anatomical findings were verified by physiological measurements of the receptive field widths.
Physiological Findings
The anterior V-cells (V1 to V3) are directionally sensitive to vertical movement: upward and downward pattern motion causes hyper-polarizing and depolarizing DC-membrane potential shifts, respectively. The lateral and posterior V-cells (V4 to V8) exhibit an additional sensitivity to horizontal movement giving rise to depolarizing and hyperpo-larizing DC-membrane potential shifts with progressive and regressive pattern motion, respectively (Fig. 14). The directional sensitivity of V9 is not known. All V-cells respond to monocular, ipsilateral stimulation only. In addition to their directional sensitivity to moving patterns they respond to light intensity changes with a strong increase in the fluctuations of the transmembrane potential (which is proportional to the logarithm of the intensity). The average DC-membrane potential changes only very slightly (+3 mV) to a hundredfold increase in light intensity. These graduated potentials are sometimes accompanied by small, rapid spike-like potentials. Regular action potentials have never been observed under natural conditions, but can be elicited by hyperpolarizing current injection. The ionic basis of the observed potential behaviour is discussed.
On physiological and anatomical grounds, the V-cells are believed to be output elements of the lobula plate with connections to descending neurones of the ventral nerve cord (Strausfeld and Obermayer, 1976) and to heterolateral elements. The anatomical and physiological properties of a heterolateral element (VS1) are discussed and evidence is presented to show that this cell is postsynaptic to the V-cells.
In general, the results of this study accord with results obtained onCalliphora (Hausen, 1976c; Hengstenberg, 1977).
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We wish to thank the “Deutsche Forschungsgemeinschaft” for support of these investigations through grant ec 56/1b. This research was furthermore supported by grants NSF BMS 74-21712 and NIH 1 ROL EY 01513-01. We are most grateful to Dr. N.J. Strausfeld for his help in identifying some of these cells. We wish to thank Prof. Hamdorf, Drs. Buchner, Franceschini, Hausen, Mr. D. Krieghoff, S. Razmjoo, D. Whittle for fruitful discussions and critical reading of the manuscript. We thank Mr. S. Soohoo for his aid in the computer analysis. We are indebted to Mr. D. Aranovich for designing some of the electronic control circuits, Mr. J. Wilson for building stimulus equipment, Mrs. I. Paas and Mr. J. Eppinger for help with the figures, and Mrs. Hundt and Mrs. B. Hadamczyk for the typing of the manuscript.
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Eckert, H., Bishop, L.G. Anatomical and physiological properties of the vertical cells in the third optic ganglion ofPhaenicia sericata (Diptera, Calliphoridae). J. Comp. Physiol. 126, 57–86 (1978). https://doi.org/10.1007/BF01342651
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DOI: https://doi.org/10.1007/BF01342651