The horizontal cells in the lobula plate of the blowfly,Phaenicia sericata

SummarySummary Combining intracellular recording and dye injection techniques, the horizontal cells of the blowfly,Phaenicia (= Lucilia) sericata, were studied.Anatomy In each lobula plate, one finds a set of three cells, termed NH-, EH- and SH-cell. EH occurs in two distinct anatomical forms, EH1 and EH2, differing in their respective branching patterns of the axon at the frontal surface of the lobula plate. Each cell's dendrite covers approximately a third of the surface of the lobula plate corresponding to a third of the visual field of the ipsilateral eye. These dendrites possess postsynaptic spines. The axons of all three cells pass along the frontal surface of the lobula plate within the inner chiasma; they cross the optic peduncle and enter the central protocerebrum where they form a second arborization, the axonal arborization consisting of dorsally extending collaterals. The axons terminate in the posterior slope of the ventrolateral protocerebrum. The axonal arborization as well as the axonal terminals possess telodendritic knobs. Ultrastructural investigations show that the lobula plate-dendrite possesses exclusively postsynaptic chemical synapses, and that the axonal arborisation and the axonal terminals possess pre- as well as postsynaptic chemical synapses. The very endings of the axons are exclusively presynaptic.Physiology The horizontal cells respond to stimulation within the ipsi- and/or contralateral receptive field. Regressive motion within the contralateral receptive field induces EPSPs and action potentials of small amplitude (10–35 mV); progressive motion is ineffective. Within the ipsilateral receptive field, regressive motion hyperpolarizes the cell membrane whereas progressive motion induces a strong depolarizing membrane potential-shift with superimposed fast potential changes of noisy appearance. Thus, the horizontal cells respond to rotational movement of the surround around the high axis of the animal: clockwise rotation excites the horizontal cells of the right lobula plate and counterclockwise motion those of the left lobula plate, respectively. However, this compound potential behaviour can only be recorded in the lobula plate-axon and main dendrites, whereas the horizontal cells respond tocontralateral regressive motion with action potentialsonly in their axonal terminals in the posterior slope; no graded potentials can be recorded in this cell region if stimulation occurs within theipsilateral receptive field. It is discussed that the previously described graded potentials for the axonal terminals (Hausen 1976b) can only be measured if the cells are already damaged. The probable cause of this change in response behaviour from action potentials to a compound potential behaviour (consisting of graded potentials and action potentials though of small amplitude) is discussed.This research was supported by the Deutsche Forschungsgemeinschaft through grants Ec56/1a + b and a Heisenberg stipend EC 56/3, funds from the SFB 114, and a grant from the National Science Foundation (NSF BMS 74-21712) awarded to the author and L.G. Bishop. I am indebted to Dr. A. Whittle and particularly to Prof. K. Meller for their invaluable help in ultrathin sectioning and to Mrs. B. Decker who introduced me to the technique of cutting serial semithin sections. Prof. K. Hamdorf helped with many stimulating discussions. I am most grateful to Dr. W. Broughton for kindly correcting the English style.     

Orientation tuning of motion-sensitive neurons shaped by vertical-horizontal network interactions

We measured the orientation tuning of two neurons of the fly lobula plate (H1 and H2 cells) sensitive to horizontal image motion. Our results show that H1 and H2 cells are sensitive to vertical motion, too. Their response depended on the position of the vertically moving stimuli within their receptive field. Stimulation within the frontal receptive field produced an asymmetric response: upward motion left the H1/H2 spike frequency nearly unaltered while downward motion increased the spike frequency to about 40% of their maximum responses to horizontal motion. In the lateral parts of their receptive fields, no such asymmetry in the responses to vertical image motion was found. Since downward motion is known to be the preferred direction of neurons of the vertical system in the lobula plate, we analyzed possible interactions between vertical system cells and H1 and H2 cells. Depolarizing current injection into the most frontal vertical system cell (VS1) led to an increased spike frequency, hyperpolarizing current injection to a decreased spike frequency in both H1 and H2 cells. Apart from VS1, no other vertical system cell (VS2-8) had any detectable influence on either H1 or H2 cells. The connectivity of VS1 and H1/H2 is also shown to influence the response properties of both centrifugal horizontal cells in the contralateral lobula plate, which are known to be postsynaptic to the H1 and H2 cells. The vCH cell receives additional input from the contralateral VS2-3 cells via the spiking interneuron V1.     

The synaptic organization of visual interneurons in the lobula complex of flies

Summary The synaptic organization of three classes of cobalt-filled and silver-intensified visual interneurons in the lobula complex of the blowfly Calliphora (Col A cells, horizontal cells and vertical cells) was studied electron microscopically. The Col A cells are regularly spaced, columnar, small field neurons of the lobula, which constitute a plexus of arborizations at the posterior surface of the neuropil and the axons of which terminate in the ventrolateral protocerebrum. They show postsynaptic specializations in the distal layer of their lobula-arborizations and additional presynaptic sites in a more proximal layer; their axon terminals are presynaptic to large descending neurons projecting into the thoracic ganglion. The horizontal and vertical cells are giant tangential neurons, the arborizations of which cover the anterior and posterior surface of the lobula plate, respectively, and which terminate in the perioesophageal region of the protocerebrum. Both classes of these giant neurons were found to be postsynaptic in the lobula plate and pre- and postsynaptic at their axon terminals and axon collaterals. The significance of these findings with respect to the functional properties of the neurons investigated is discussed.     

On the neuronal basis of figure-ground discrimination by relative motion in the visual system of the fly

A new class of large-field tangential neurones (Figure Detection (FD-) cells) has been found and analysed in the lobula plate, the posterior part of the third visual ganglion, of the fly by combined extra-and intracellular recording as well as Lucifer Yellow injection. The FD-cells are likely to play a prominent role in figure-ground discrimination. Together with the Horizontal Cells, the output elements of the neuronal network underlying the optomotor course control reaction, they seem to be appropriate to account for the characteristic yaw torque response to relative motion. The FD-cells might thus compensate for the deficits of the Horizontal Cells with respect to figureground discrimination (see Egelhaaf, 1985a).The FD-cells are directionally selective for either front-to-back (FD 1, FD 4) or back-to-front motion (FD 2, FD 3). Their excitatory receptive fields cover part of (FD 1, FD 2, FD 3) or the entire horizontal extent (FD 4) of the visual field of one eye. Their most important common property in the context of figureground discrimination is that they are more sensitive to relatively small objects than to spatially extended patterns. Their response to a small figure is much reduced by simultaneous large-field motion in front of the ipsi-as well as the contralateral eye. This large-field inhibition is either directionally selective or bidirectional, depending on the FD-cell under consideration. The main dendritic arborization of all FD-cells resides in the lobula plate. Their axonal projections lie in either the ipsi-or contralateral posterior optic foci and, thus, in the same area as the terminals of the Horizontal Cells. The FD-cells are, therefore, appropriate candidates for output elements of the optic lobes involved in figure-ground discrimination.     

Tangential medulla neurons in the mothManduca sexta. Structure and responses to optomotor stimuli

InManduca sexta, large tangential cells connect the medulla via the lobula valley (LoV) tract to the midbrain and the contralateral medulla. Tract neurons have been stained and recorded to determine their responses to optomotor stimulation. Neurons in the LoV-tract comprise a physiologically and anatomically heterogeneous population:

1. Motion insensitive medulla tangential (Mt) neurons arise from cell bodies in the ventral rind. Heterolateral cells arborize massively in both medullae and one or both halves of the midbrain. Mt-neurons respond to changes in light intensity. Physiological and anatomical evidence argues for their monocularity and transmission from the medulla on the side of the soma to the central brain and the contralateral medulla.
2. Motion sensitive neurons with cell bodies behind the protocerebral bridge connect the midbrain to the ipsior contralateral medulla. Direction-selective responses are characterized by excitation to motion in the preferred and inhibition in the opposite direction with maxima either in a horizontal or vertical direction. Peak values appear at contrast frequencies of appr. 3/s. The results suggest that these neurons are binocular and relay information from the midbrain to the medulla. They have been labelled as centrifugal medulla tangential (cMt) neurons.

The possible roles for tract neurons in visually guided behaviour are discussed.     

Motion sensitive interneurons in the optomotor system of the fly

The three horizontal cells of the lobula plate of the blowflyCalliphora erythrocephala were studied anatomically and physiologically by means of cobalt impregnations and intracellular recordings combined with Procion and Lucifer Yellow injections. The cells are termed north, equatorial and south horizontal cell (HSN, HSE, HSS) and are major output neurons of the optic lobe. 1. The dendritic arborizations of the HSN, HSE, HSS reside in a thin anterior layer of the lobula plate and extend over the dorsal, equatorial and ventral parts of this neuropil, respectively. Due to the retinotopic organization of the optic lobe, these parts correspond anatomically to respective regions of the ipsilateral visual field. Homologue horizontal cells in both lobula plates of the same animal and in different animals are highly variable with respect to their individual dendritic branching patterns. They are extraordinarily constant, on the other hand, with regard to the position and size of their dendritic fields as well as their dendritic branching density distributions. Each cell covers about 40% of the total area of the lobula plate and shows the highest dendritic density near the lateral margin of the neuropil which subserves the frontal eye region. The axons of the horizontal cells are relatively short and large in diameter; they terminate in the posterior ventrolateral protocerebrum. 2. The horizontal cells are directionally selective motion sensitive visual interneurons responding preferentially to progressive (front to back) motion in the ipsilateral visual field with graded depolarization of their axons and superimposed action potentials. Stimulation with motion in the reverse direction leads to hyperpolarizing graded responses. The HSE and HSN are additionally activated by regressive motion in the contralateral visual field.     

Two classes of visual motion sensitive interneurons differ in direction and velocity dependency of in vivo calcium dynamics

Neurons exploit both membrane biophysics and biochemical pathways of the cytoplasm for dendritic integration of synaptic input. Here we quantify the tuning discrepancy of electrical and chemical response properties in two kinds of neurons using in vivo visual stimulation. Dendritic calcium concentration changes and membrane potential of visual interneurons of the fly were measured in response to visual motion stimuli. Two classes of tangential cells of the lobula plate were compared, HS-cells and CH-cells. Both neuronal classes are known to receive retinotopic input with similar properties, yet they differ in morphology, physiology, and computational context. Velocity tuning and directional selectivity of the electrical and calcium responses were investigated. In both cell classes, motion-induced calcium accumulation did not follow the early transient of the membrane potential. Rather, the amplitude of the calcium signal seemed to be related to the late component of the depolarization, where it was close to a steady state. Electrical and calcium responses differed with respect to their velocity tuning in CH-cells, but not in HS-cells. Furthermore, velocity tuning of the calcium response, but not of the electrical response differed between neuronal classes. While null-direction motion caused hyperpolarization in both classes, this led to a calcium decrement in CH-cells, but had no effect on the calcium signal in HS-cells, not even when calcium levels had been raised by a preceding excitatory motion stimulus. Finally, the voltage-[Ca2+]i-relationship for motion-induced, transient potential changes was steeper and less rectifying in CH-cells than in HS-cells. These results represent an example of dendritic information processing in vivo, where two neuronal classes respond to identical stimuli with a similar electrical response, but differing calcium response. This highlights the capacity of neurons to segregate two response components.     

The vertical-horizontal neurone (VH) in the lobula plate of the blowfly,Phaenicia

Summary The anatomy and physiology of a motion-sensitive neurone, the vertical-horizontal (VH-) cell in the third visual neuropil (lobula plate) of the blowfly,Phaenicia was studied by intracellular recordings combined with dye injection. The cell possesses two dendritic fields in different layers of the lobula plate. The axon runs jointly with those of the vertical cells along the caudal surface of the lobula plate and terminates in the central protocerebrum lateral to the esophageal canal. The receptive field of the VH-cell is subdivided into two physiologically different parts which correspond to the two dendritic fields: if the input reaches the dendritic field residing in a more caudal layer (V-layer), the cell responds maximally to vertical pattern motion; whereas if the input reaches the dendritic field residing in a more rostral layer (H-layer), the cell responds maximally to horizontal pattern motion. The VH-neurone responds maximally to a contrast frequency of approximately / 1.8 Hz which coincides with the contrast frequency dependence of optomotor (following) responses. It is, therefore, considered to be a likely candidate mediating the pitch response (Blondeau and Heisenberg 1982) in flies.     

The Intrinsic Electrophysiological Characteristics of Fly Lobula Plate Tangential Cells: III. Visual Response Properties

In this last paper in a series (Borst and Haag, 1996; Haag et al., 1997) about the lobula plate tangential cells of the fly visual system (CH, HS, and VS cells), the visual response properties were examined using intracellular recordings and computer simulations. In response to visual motion stimuli, all cells responded mainly by a graded shift of their axonal membrane potential. While ipsilateral motion resulted in a graded membrane potential shift, contralateral motion led to distinct EPSPs. For HS cells, simultaneous extracellular recorded action potentials of a spiking interneuron, presumably the H2 cell, corresponded to the EPSPs in the HS cell in a one-to-one fashion. When HS cells were hyperpolarized during ipsilateral motion, they mainly produced action potentials, but when they were hyperpolarized during contralateral motion only a slight increase of EPSP amplitude, could be observed. Intracellular application of the sodium channel blocker QX 314 abolished action potentials of HS cells while having little effect on the graded membrane response to ipsilateral motion. HS and CH cells were also studied with respect to their spatial integration properties. For both cell types, their graded membrane response was found to increase less than linearly with the size of the ipsilateral motion pattern. However, while for HS cells various amounts of hyperpolarizing current injected during motion stimulation led to different saturation levels, this was not the case for CH cells. In response to a sinusoidal velocity modulation, CH cells followed pattern motion only up to 10 Hz modulation frequency, but HS cells still revealed significant membrane depolarizations up to about 40 Hz.In the computer simulations, the compartmental models of tangential cells, as derived in the previous papers, were linked to an array of local motion detectors. The model cells revealed the same basic response features as their natural counterparts. They showed a response saturation as a function of stimulus size. In CH-models, however, the saturation was less pronounced than in real CH-cells, indicating spatially nonuniform membrane resistances with higher values in the dendrite. As in the experiments, HS models responded to high-frequency velocity modulation with a higher amplitude than did CH models.     

Motion sensitive interneurons in the optomotor system of the fly

The functional properties of the three horizontal cells (north horizontal cell, HSN; equatorial horizontal cell, HSE; south horizontal cell, HSS) in the lobula plate of the blowflyCalliphora erythrocephala were investigated electrophysiologically. 1. The receptive fields of the HSN, HSE, and HSS cover the dorsal, equatorial and ventral part of the ipsilateral visual field, respectively. In all three cells, the sensitivity to visual stimulation is highest in the frontal visual field and decreases laterally. The receptive fields and spatial sensitivity distributions of the horizontal cells are directly determined by the position and extension of their dendritic fields in the lobula plate and the dendritic density distributions within these fields. 2. The horizontal cells respond mainly to progressive (front to back) motion and are inhibited by motion in the reverse direction, the preferred and null direction being antiparallel. The amplitudes of motion induced excitatory and inhibitory responses decline like a cosine function with increasing deviation of the direction of motion from the preferred direction. Stimulation with motion in directions perpendicular to the preferred direction is ineffective. 3. The preferred directions of the horizontal cells show characteristic gradual orientation changes in different parts of the receptive fields: they are horizontally oriented only in the equatorial region and increasingly tilted vertically towards the dorsofrontal and ventrofrontal margins of the visual field. These orientation changes can be correlated with equivalent changes in the local orientation of the lattice of ommatidial axes in the pertinent compound eye. 4. The response amplitudes of the horizontal cells under stimulation with a moving periodic grating depend strongly on the contrast frequency of the stimulus. Maximal responses were found at contrast frequencies of 2–5 Hz. 5. The spatial integration properties of the horizontal cells (studied in the HSE) are highly nonlinear. Under stimulation with extended moving patterns, their response amplitudes are nearly independent of the size of the stimuli. It is demonstrated that this response behaviour does not result from postsynaptic saturation in the dendrites of the cells. The results indicate that the horizontal system is essentially involved in the neural control of optomotor torque responses performed by the fly in order to minimize unvoluntary deviations from a straight flight course.     

The organization of giant horizontal-motion-sensitive neurons and their synaptic relationships in the lateral deutocerebrum of Calliphora erythrocephala and Musca domestica

Summary Three giant horizontal-motion-sensitive (HS) neurons arise in the lobula plate. Their axons terminate ipsilaterally in the medial deutocerebrum and suboesophageal ganglion. Both Golgi impregnations and cobalt fills demonstrate that endings of the two HS cells, representing the upper and middle third of the retina, differ in shape and location from that of the HS cell subtending the lower third of the eye. This dichotomy is reflected by the terminals of a pair of centrifugal horizontal cells (CH), one of which invades lobula plate neuropil subtending the upper two-thirds of the retina. The other overlaps the dendrites of the HS cell subtending the lower one-third of the retina.The HS cells are cobalt-coupled to a variety of complexly arborizing descending neurons. In Musca domestica, gap-junction-like apposition areas have been observed between HS axon collaterals and descending neuron dendrites. The three HS cells also share conventional chemical synapses with postsynaptic elements, which include the dendritic spines of descending neurons. Unlike the giant vertical-motion-sensitive neurons of the lobula plate, whose relationships with descending neurons appear to be relatively simple, the horizontal cells end on a large number of descending neurons where they comprise one of several different populations of terminals. These descending neurons terminate within various centres of the thoracic ganglia, including neuropil supplying leg, neck, and flight muscle.     

Three descending interneurons reporting deviation from course in the locust

Three descending brain interneurons (DNI, DNM, DNC) are described from Locusta migratoria. All are paired, dorsally situated neurons, with soma in the protocerebrum, input dendrites in the proto- and deuterocerebrum, and a single axon running to the metathoracic ganglion and sometimes further. In DNI the soma and all cerebral arborizations lie ipsilateral to the axon. Discrete regions of arborization lie in the ipsilateral and medial ocellar tracts, the midprotocerebrum and the deuterocerebrum. In the other ganglia the axon branches only ipsilaterally, principally laterally in the flight motor neuropil but also towards the midline. DNC is similarly organized to DNI, but the cell crosses the midline in the brain. Soma, the single projection into a lateral ocellar tract, and the midprotocerebral arborization all lie contralateral to the axon. The deuterocerebral arborization is, however, ipsilateral to the axon. The pattern of projections in the remaining ganglia resembles that of DNI. The soma and all cerebral arborizations of DNM lie ipsilateral to the axon. The arborization is only weakly subdivided into protocerebral, deuterocerebral and medial ocellar tract regions. In the remaining ganglia the arborization extends bilaterally to similar areas of both left and right flight motor neuropil. A table of synonymy is given, equating the various names used for these neurons by previous authors. The morphology correlates well with the known input and output connections. They respond physiologically to deviations from the normal flight posture mediated by ocelli, eyes and wind hairs and connect to the thoracic flight apparatus.     

Golgi analysis of tangential neurons in the lobula plate of<Emphasis Type="Italic">Drosophila melanogaster</Emphasis>

The lobula plate (LP), which is the third order optic neuropil of flies, houses wide-field neurons which are exquisitely sensitive to motion. Among Diptera, motion-sensitive neurons of larger flies have been studied at the anatomical and physiological levels. However, the neurons ofDrosophila lobula plate are relatively less explored. AsDrosophila permits a genetic analysis of neural functions, we have analysed the organization of lobula plate ofDrosophila melanogaster. Neurons belonging to eight anatomical classes have been observed in the present study. Three neurons of the horizontal system (HS) have been visualized. The HS north (HSN) neuron, occupying the dorsal lobula plate is stunted in its geometry compared to that of larger flies. Associated with the HS neurons, thinner horizontal elements known as h-cells have also been visualized in the present study. Five of the six known neurons of the vertical system (VS) have been visualized. Three additional neurons in the proximal LP comparable in anatomy to VS system have been stained. We have termed them as additional VS AVS)-like neurons. Three thinner tangential cells that are comparable to VS neurons, which are elements of twin vertical system (tvs); and two cells with wide dendritic fields comparable to CH neurons of Diptera have been also observed. Neurons comparable to VS cells but with ‘tufted’ dendrites have been stained. The HSN and VS1-VS2 neurons are dorsally stunted. This is possibly due to the shape of the compound eye ofDrosophila which is reduced in the fronto-dorsal region as compared to larger flies     

Lobula plate and ocellar interneurons converge onto a cluster of descending neurons leading to neck and leg motor neuropil in Calliphora erythrocephala

Summary In the fly, Calliphora erythrocephala, a cluster of three Y-shaped descending neurons (DNOVS 1–3) receives ocellar interneuron and vertical cell (VS4–9) terminals. Synaptic connections to one of them (DNOVS 1) are described. In addition, three types of small lobula plate vertical cell (sVS) and one type of contralateral horizontal neuron (Hc) terminate at DNOVS 1, as do two forms of ascending neurons derived from thoracic ganglia. A contralateral neuron, with terminals in the opposite lobula plate, arises at the DNOVS cluster and is thought to provide heterolateral interaction between the VS4–9 output of one side to the VS4–9 dendrites of the other. DNOVS 2 and 3 extend through pro-, meso-, and metathoracic ganglia, branching ipsilaterally within their tract and into the inner margin of leg motor neuropil of each ganglion. DNOVS 1 terminates as a stubby ending in the dorsal prothoracic ganglion onto the main dendritic trunks of neck muscle motor neurons. Convergence of VS and ocellar interneurons to DNOVS 1 comprises a second pathway from the visual system to the neck motor, the other being carried by motor neurons arising in the brain. Their significance for saccadic head movement and the stabilization of the retinal image is discussed.     

Vertical motion detectors and their synaptic relations in the third optic lobe of the fly

The synaptic relations of the giant vertical cells in the lobula plate of the fly were investigated using electron microscopical procedures and Lucifer yellow dye backfill and injection techniques. Histological features of the giant vertical cells are described. The giant vertical cells are exclusively postsynaptic in the lobula plate. They function to integrate input from dense arrays of chemical synapses and have a wide spatial input from the lobula plate. The giant vertical cells are postsynaptic to perpendicularly occurring cells. There are two classes of cells presynaptic to the vertical cells, one of which contains large dense-core vesicles. The giant vertical cells are not the only cells postsynaptic to these two classes of perpendicualr cells. A second group of smaller tangential cells, the twin vertical cells, were also found postsynaptic to many of the same cells that synapsed with the giant vertical cells. The twin vertical cells and the giant vertical cells are therefore integrating some of the same information in the lobula plate.     

Responses of blowfly motion-sensitive neurons to reconstructed optic flow along outdoor flight paths

The retinal image flow a blowfly experiences in its daily life on the wing is determined by both the structure of the environment and the animal’s own movements. To understand the design of visual processing mechanisms, there is thus a need to analyse the performance of neurons under natural operating conditions. To this end, we recorded flight paths of flies outdoors and reconstructed what they had seen, by moving a panoramic camera along exactly the same paths. The reconstructed image sequences were later replayed on a fast, panoramic flight simulator to identified, motion sensitive neurons of the so-called horizontal system (HS) in the lobula plate of the blowfly, which are assumed to extract self-motion parameters from optic flow. We show that under real life conditions HS-cells not only encode information about self-rotation, but are also sensitive to translational optic flow and, thus, indirectly signal information about the depth structure of the environment. These properties do not require an elaboration of the known model of these neurons, because the natural optic flow sequences generate—at least qualitatively—the same depth-related response properties when used as input to a computational HS-cell model and to real neurons.     

Localized direction selective responses in the dendrites of visual interneurons of the fly

Background  

The various tasks of visual systems, including course control, collision avoidance and the detection of small objects, require at the neuronal level the dendritic integration and subsequent processing of many spatially distributed visual motion inputs. While much is known about the pooled output in these systems, as in the medial superior temporal cortex of monkeys or in the lobula plate of the insect visual system, the motion tuning of the elements that provide the input has yet received little attention. In order to visualize the motion tuning of these inputs we examined the dendritic activation patterns of neurons that are selective for the characteristic patterns of wide-field motion, the lobula-plate tangential cells (LPTCs) of the blowfly. These neurons are known to sample direction-selective motion information from large parts of the visual field and combine these signals into axonal and dendro-dendritic outputs.     

Synaptic effects of contralateral somatic afferents on spinal motoneurons

Electrical activity of flexor and extensor alpha-motoneurons of the lumbar segments of cat's spinal cord as recorded intracellularly during electric stimulation of afferents of the contralateral posterior limb. Contralateral postsynaptic potentials (PSP) were shown to be evoked by activation of cutaneous and high-threshold muscle afferents. The high-threshold afferents of various muscle nerves participate to varying degrees in the generation of contralateral PSP. Contralateral inhibitory postsynaptic potentials (IPSP) were recorded in both flexor and extensor motoneurons along with contralateral excitatory postsynaptic potentials (EPSP). There are no fundamental differences in their distribution between flexor and extensor neurons. Inhibitory influences as a rule are predominant in both during the first 20 msec, and EPSP are predominant in the interval between 20 and 100 msec. The balance of excitatory and inhibitory pathway activity was found to be not as stable as that of the homolateral pathways.I. P. Pavlov Institute of Physiology, Academy of Sciences of the USSR, Leningrad. Translated from Neirofiziologiya, Vol. 3, No. 4, pp. 418–425, July–August, 1971.     

A synergetic theory of environmentally-specified and learned patterns of movement coordination

By combining neuropharmacology and electrophysiology, we tried to determine whether the main neuronal mechanism responsible for direction-selective motion detection in the fly is based on an excitatory or an inhibitory synaptic interaction. By blocking inhibitory interactions with picrotoxinin, an antagonist of the inhibitory neurotransmitter GABA, we could abolish most of the directional selectivity of a large-field movement-sensitive neuron (H1-cell) in the lobula plate of the blowfly Calliphora erythrocephala. These modifications are similar to changes observed in the optomotor response of the fruitfly Drosophila melanogaster after application of picrotoxinin (Bülthoff and Bülthoff 1987a, b). Assuming a simplified logical model, these results are compatible with inhibitory synaptic interactions at the level of the elementary movement detectors. The picrotoxinin-induced changes in direction selectivity are not due to modifications of the peripheral visual processing in the retina and lamina. This was shown by simultaneous recordings of the electroretinogram and the H1-cell. The latencies between drug injections into various parts of the brain and their first effects on the H1-cell suggest that the inhibitory mechanism for motion detection is located in the medulla rather than in the lobula plate.