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1.
Summary The lamina ganglionaris of the superposition eye of Cloeon dipterum is composed of separate optic cartridges arranged in a hexagonal pattern. Each optic cartridge consists of one central, radially branched monopolar cell (Li) surrounded by a crown of seven retinula cell terminals and two more unilaterally branched monopolar cells (La1/La2) situated close together outside the cartridge. Projections to neighbouring cartridges have not been observed.In most cases, synaptic contacts could be seen between a presynaptic retinula cell and more than two other postsynaptic profiles, which belong to monopolar cells or sometimes to glial cells.Seven retinula cell fibers of one ommatidium pass in a bundle through the basement membrane, run into their respective cartridges without changing orientation and terminate at approximately equal levels in the lamina. Long visual fibers with endings in the medulla are not visible in the superposition eye lamina, but are present in the lateral apposition eye. The relationship between the behaviour of the animal, optic mechanisms of the superposition eye and the structure of the lamina is discussed.  相似文献   

2.
The Lamina ganglionaris (first optic neuropile) of the decapod crustacean Pandalus borealis has its optic cartridges (synaptic compartments) arranged in horizontal rows. Each optic cartridge contains seven receptor axon terminals and the branching axis fibres of five monopolar second order neurons. Four types of monopolar neurons are classified. Their cell bodies are arranged in two layers. The inner layer contains the cell bodies of exclusively one of these types, and each cartridge is invaded by two neurons of this neuron type (type M 1:a and M 1:b). The outer layer contains the cell bodies of the remaining three types (M 2, M3 and M4). One gives rise to a large radially branched axis fibre in the centre of the cartridge. The other two have wide branches which may make inter-cartridge contacts, one proximally and the other distally in the plexiform layer, which is clearly bistratified. The receptor axons terminate in two levels corresponding to these strata. Two sets of tangenital fibres form networks in the proximal and the mid-portion of the lamina. Both networks have fibres with primary branches in the vertical plane and secondary branches in the horizontal plane. The fibres of the networks are derived from axons that pass from the second optic neuropile, the medulla externa.  相似文献   

3.
At the anterior rim of the first optic neuropile, or lamina, of the housefly's (Musca domestica) compound eye, the terminals of photoreceptors (R) innervate postsynaptic neurons in variable numbers to provide a continuous range of natural hypo- and hyperinnervations. Frequencies of photoreceptor synapses have been measured from quantitative electron microscopy on single sections of the lamina's unit synaptic modules, called cartridges. These are normally innervated by six photoreceptor terminals (6R cartridges). At the lamina's edge hypoinnervated cartridges (2R-5R) are found, whereas hyperinnervated cartridges (7R, 8R) are located at the equator between dorsal and ventral eye halves. In 2R cartridges each presynaptic terminal forms up to 1.5 times the normal, 6R cartridge number of synapses, thereby offsetting the reduced number of terminals and partially conserving the input upon the postsynaptic neurons. Thus the terminals have a reserve synaptogenic capacity never normally revealed. By comparison, terminals in 8R cartridges form about the same numbers of synapses as in "normal" eye regions, so that their postsynaptic neurons have a synaptic input increased by the extra number of terminals. The number of synapses formed between input terminals and target neurons is therefore not fixed but changes as a function of the total receptor terminal complement. The size of a photoreceptor terminal covaries to a certain extent with the number of its presynaptic sites; the spacing density of presynaptic sites over the terminals' surface in a 2R cartridge compared with an 8R cartridge increases far less (only 17%) than the increase in the number of sites (43%). The pair of postsynaptic cell interneurons in each 2R cartridge also shows a decrease in axonal diameter compared with those in 8R cartridges. Thus both the pre- and postsynaptic cells show size changes correlated with changes in their synaptic engagement.  相似文献   

4.
Panorpa larvae possess stemmata (lateral ocelli), which have the structure of compound eyes, and stemma lamina and stemma medulla neuropils. A distinct lobula neuropil is lacking. The stemma neuropils have a columnar organization. They contain lamina monopolar cells, and both short and long visual fibers. All the identified larval monopolar neurons have radially arranged dendrites along the entire depth of the lamina neuropil and a single terminal arborization within the medulla (L1/L2-type). The terminals of visual fibers have short spiny lateral projections. Long fibers possess en passant synapses within the lamina. The same principles of organization of first and second order visual neuropils are found in Panorpa imagines. In contrast to the larvae, a lobula neuropil is present. Adults have monopolar cells of the L1-type that are similar to the L1-neurons found in Diptera. The columnar organization, the presence of short and long visual fibers, and lamina monopolar neurons are thus features common to both visual systems, viz., the larval (stemmata) and the imaginal (compound eyes).  相似文献   

5.
In the housefly's first optic neuropile, or lamina, the axons of two classes of monopolar cell interneurons, L1 and L2, exhibit a daily rhythm of size changes: swelling during the day, and shrinking by night. At least for the L2 cells this rhythm is circadian. Moreover, epithelial glial cells that enwrap each lamina cartridge, its monopolar cell axons, and their surrounding crown of input photoreceptor terminals also change size, but in the opposite direction to the changes in L1 and L2-swelling by night and shrinking by day. The rhythmic changes in glia indicate the possible involvement of these cells in the lamina's circadian system. To examine their role in regulating the rhythmic changes of L1 and L2's axon sizes we have injected three chemicals into the haemolymph of the fly's head: fluorocitrate (FL) and iodoacetate (IAA), which affect the metabolism of glial cells, and octanol (OC), which closes gap junction channels. All chemicals exerted an effect on L1 and L2, which depended on the time of injection, the drug concentration, and the postinjection times at which we examined the fly's brains. Moreover, day/night changes in the axon sizes of L1 and L2 were increased in FL- and IAA-treated flies, indicating that glial cells may normally inhibit these changes by regulating the sizes of L1 and L2's axons during the day and night. In turn, lack of a day/night rhythm in L1 and L2 after OC injections shows that the rhythm's persistence depends on communication between the lamina cells through gap junction channels.  相似文献   

6.
Summary The gross structure as well as the neuronal and non-neuronal components of the lamina ganglionaris of the locust Schistocerca gregaria are described on the basis of light- and electron-microscopical preparations of Golgj (selective silver) and ordinary histological preparations. The array of optic cartridges within the lamina neuropile — their order and arrangement — and the composition of the cartridges are described. There are six types of monopolar neurons: three whose branches reach to other cartridges and three whose branches are confined to their own cartridges. Retinula axons terminate either in the lamina or the medulla neuropiles. There are three types of centrifugal neurons, two types of horizontal neuron, as well as glia and trachea in the lamina neuropile. The development of the lamina neuropile is described in terms of developing monopolar and centrifugal axons, growing retinula fibres, and composition of the developing optic cartridges.MSN was supported in part by a Fulbrights-Hays Scholarsship. We are grateful to the Science Research Council for its grant to PMJS.  相似文献   

7.
Summary The arrangement of first and second order neurons in an optic cartridge and the topographical relationships of the second order neurons within a cartridge and to groups of surrounding cartridges have been analyzed in the visual system of the bee, Apis mellifera, from light and electron microscope studies on Golgi preparations. At the level of the monopolar cell body layer, the nine retinula cell fibres of each ommatidium, the six short visual fibres arranged in a circle surrounding the three long visual fibres, become cartridges as a consequence of the appearance of the second order neurons (L-fibres) which join the R-fibre bundles. Two of the four different L-fibre types, L-1 and L-2, remain together in the centre of the cartridge throughout the lamina. The axons of the L-3 and L-4 fibres, however, have their position integrated into the circle formed by the endings of the short visual fibres. On the basis of further examination of light and especially electron microscopical Golgi material, the different L-fibres can be classified into four types which appear in each cartridge. The clear stratification in the first synaptic region (A, B and C) seems to be the best criterion for a morphological classification since such a classification necessarily also includes a functional basis. According to a naming system based on the position of the lateral processes, L-fibres with side branches in strata A, B and C are called L-1 fibres. Fibres with lateral processes in strata A and B are L-2 fibres; monopolar cell fibres with branches only in the second stratum B are L-fibres of type 3; and all monopolar cells with branches only in stratum C are called L-4 fibres. In addition to the branching pattern covering only the parent cartridge, two of the four fibre types (L-2 and L-4) have long collaterals reaching neighbouring cartridges: L-2 in stratum A and L-4 in stratum C. These collaterals presumably form a substrate for lateral interactions.  相似文献   

8.
Summary Neuronal elements, i.e. first and second order neurons, of the first optic ganglion of three waterbugs, N. glauca, C. punctata and G. lacustris, are analyzed on the basis of light and electron microscopy.Eight retinula cell axons, leaving each ommatidium, disperse to different cartridges as they enter the laminar outer plexiform layer. Such a pattern of divergence is one of the conditions for neuronal superposition; it is observed for all three species of waterbugs. The manner in which the receptors of a single bundle of ommatidia split of within the lamina, whereby information from receptors up to three or five horizontal rows away can converge upon the same cartridge, differs among the species. Six of the eight axons of retinula cells R1-6, the short visual fibers end at different levels within the bilayered lamina, whereas the central pair of retinula cells R7/8, the long visual fibers, run directly through the lamina to a corresponding unit of the medulla. Four types of monopolar cells L1–L4 are classified; their branching patterns seem to be correlated to the splitting and termination of retinula cell axons. The topographical relationship and synaptic organization between retinula cell terminals and monopolar cells in the two laminar layers are identified by examination of serial ultrathin sections of single Golgi-stained neurons.An attempt is made to correlate some anatomical findings, especially the neuronal superposition, to results from physiological investigations on the hemipteran retina.  相似文献   

9.
We describe visual responses of seventeen physiological classes of columnar neuron from the retina, lamina and medulla of the locust (Locusta migratoria) optic lobe. Many of these neurons were anatomically identified by neurobiotin injection. Characterisation of neuronal responses was made by moving and flash stimuli, and by two system identification techniques: 1. The first-order spatiotemporal kernel was estimated from response to a spatiotemporal white-noise stimulus; 2. A set of kernels to second order was derived by the maximal-length shift register (M-sequence) technique, describing the system response to a two-channel centre-surround stimulus. Most cells have small receptive fields, usually with a centre diameter of about 1.5°, which is similar to that of a single receptor in the compound eye. Linear response components show varying spatial and temporal tuning, although lateral inhibition is generally fairly weak. Second-order nonlinearities often have a simple form consistent with a static nonlinear transformation of the input from the large monopolar cells of the lamina followed by further linear filtering.Abbreviations LMC large monopolar cell - LVF long visual fibre - RF receptive field - SMC small monopolar cell - SVF short visual fibre  相似文献   

10.
High resolution serial photomicrography has been used to plot the axonal projection patterns between retina, lamina and medulla in the optic lobes of various insects with differing ommatidial receptor arrangements. Observations are reported on the cabbage white and skipper butterflies, the bee, locust, fly, backswimmer and waterbug. The patterns of these fibre pathways have previously eluded non-rigorous analyses primarily because of their physical dimensions but are revealed in this study to have striking precision and uniformity between species when examined at the level of individually identifiable cells. Axon bundles of the tracts between retina and lamina or lamina and medulla project between a single ommatidium and its corresponding lamina cartridge or between corresponding lamina and medulla cartridges. Lateral interweaving of axons between adjacent bundles is absent. The bundles preserve the retinotopic order within their total array, so transferring the pattern of retinulae directly upon the lamina and thence after horizontal inversion in the chiasma upon the medulla. Within the lamina neuropile on the other hand the trajectories of the individual terminals from a bundle have patterns which are species-specific, sometimes involving lateral divergences. In species with open-rhabdomere ommatidia the terminals distribute to a group of lamina cartidges with a pattern which resembles the receptor pattern in the overlying ommatidium. In species with fused-rhabdome ommatidia the terminals of a single retinula behave less interestingly and all enter the same cartridge, within which, again, each occupies a position related to its cell body position within the retinula. Long visual fibres in both eye types penetrate the lamina and terminate in the particular medulla cartridge that connects with the lamina cartridge underlying their ommatidium. The perpendicular fibre pathways therefore project the visual field exactly upon the medulla in all species while the lack of interweaving between adjacent fibre bundles precludes their involvement in lateral interactions between pathways with differing visual axes. Uniformity of these projection patterns between cell layers and species differences in retinular terminal locations in the lamina can be correlated with different modes of axon growth between and within neuropile layers during optic lobe neurogenesis. Further discussion surrounds the question of which particular receptors give rise to which type of axon, for which no clear generalization has yet emerged.  相似文献   

11.
The forewing stretch receptor (SR) neuron makes monosynaptic connections with wing depressor motoneruons; in this article the pharmacology of its output onto the first baslar motoneuron (BA1) has been investigated. The SR, like other insect afferents that have been studied so far, appears to be cholinergic; transmission was suppressed reversibly by the nicotinic antagonist gallamine (10?4M) and irreversibly by α-bungarotoxin (10?6 M). The choline reuptake blocker hemicholinium-3 (10?4 M) also caused a reversible reduction in the amplitude of SR excitatory postsynaptic potentials (EPSPs) recorded in BA1. The receptor subtype nonselective muscarinic antagonists atropine (10?4 M), scopolamine (10?4 M), and quinuclidinyl benzilate (10?5 M), unlike nicotinic antagonists, caused an augmentation in EPSP amplitude. This effect does not appear to be caused by an increase in sensitivity of the motoneuron to acetylcholine (ACh), since atropine produced a marked reduction rather than an increase in the amplitude of responses to ACh pressure applied to the soma of BA1. Scopolamine only caused a modest reduction in the amplitude of ACh somatic responses. The simplest explanation for these observations is that muscarinic antagonists bring about an increase in EPSP amplitude by blockade of presynaptic autoreceptors that normally down-regulate the release of ACh from SR terminals. The effects of muscarinic receptor subtype-selective antagonists indicate that presynaptic receptors in this preparation may have a pharmacological profile more similar to that of vertebrate M2 receptors than to that of M1 or M2 subtypes. The functional significance of autoreceptors in this preparation are discussed. © 1995 John Wiley & Sons, Inc.  相似文献   

12.
The structural organization of the first optic ganglion (lamina) of the cockroach (Periplaneta americana) was investigated by the use of light and electron microscopy. Each compound eye of the cockroach is composed of up to 2000 visual units (ommatidia) of the fused rhabdom type. The ommatidia themselves consist of eight receptor cells which terminate as axons in either the first or second optic ganglion. Three different short visual fibre types end in two separate strata in the lamina, and one long fibre type ends in the second optic ganglion. Monopolar second-order neurons with wide field branching patterns in the middle stratum of the first synaptic region have postsynaptic contacts with short visual fibres. Horizontal fibre elements with branching patterns at different levels of the lamina apparently form three horizontal plexuses with presynaptic and/or postsynaptic connections to first- and secondorder neurons. The lack of well-organized fibre cartridges containing a constant number of first and second order neurons in each fascicle and the presence of only unistratified wide field monopolar cells could represent, as compared to other insect orders, a primitive stage in the development of the first optic ganglion.  相似文献   

13.
We have examined the generation and development of glial cells in the first optic ganglion, the lamina, of Drosophila melanogaster. Previous work has shown that the growth of retinal axons into the developing optic lobes induces the terminal cell divisions that generate the lamina monopolar neurons. We investigated whether photoreceptor ingrowth also influences the development of lamina glial cells, using P element enhancer trap lines, genetic mosaics and birthdating analysis. Enhancer trap lines that mark the differentiating lamina glial cells were found to require retinal innervation for expression. In mutants with only a few photoreceptors, only the few glial cells near ingrowing axons expressed the marker. Genetic mosaic analysis indicates that the lamina neurons and glial cells are readily separable, suggesting that these are derived from distinct lineages. Additionally, BrdU pulse-chase experiments showed that the cell divisions that produce lamina glia, unlike those producing lamina neurons, are not spatially or temporally correlated with the retinal axon ingrowth. Finally, in mutants lacking photoreceptors, cell divisions in the glial lineage appeared normal. We conclude that the lamina glial cells derive from a lineage that is distinct from that of the L-neurons, that glia are generated independently of photoreceptor input, and that completion of the terminal glial differentiation program depends, directly or indirectly, on an inductive signal from photoreceptor axons.  相似文献   

14.
Summary Neurons within the compound eye of the flyCalliphora erythrocephala, suspected of containing gamma-aminobutyric acid were revealed immunocytochemically, using antibodies directed against gamma-aminobutyric acid (GABA) and glutamic acid decarboxylase (GAD). The GABA content within putative GABAergic neurons was increased by high affinity uptake of GABA and selective blocking of GABA metabolism with Gabaculine. Only neuronal populations which were labelled with the GABA as well as the GAD antibodies were presumed to be GABAergic. The first optic neuropil (lamina) exhibited two distinct GA-BAergic fibre populations amongst a larger population comprised of fourteen cell classes. One fibre population was formed by the axons of the photopic photoreceptors R7 which pass through the lamina and terminate in the second optic neuropil (the medulla). The identity of R7 was established from longitudinal and transverse sections of the retina where R7 can be unequivocally distinguished from the six scotopic photoreceptors R1-6 and the other photopic receptor, R8.The other fibre population matched the profiles in the lamina of terminals of efferent C2 neurons. These neurons project distally from beneath the medulla out to the lamina ganglionaris where each retinotopic unit (cartridge) contains a characteristic hook-like terminal arbor distally. We propose from these data that the photoreceptors R7 and the efferent C2 neurons use GABA as a neurotransmitter.  相似文献   

15.
Summary The crustacean optic neuropiles, the lamina ganglionaris and especially the medulla externa, show a specific pattern of green fluorescence with the fluorescence histochemical method of Falck-Hillarp. Normally, only the terminals and the cell bodies fluoresce, but in reserpine-treated animals exogenous catecholamines are taken up by the whole adrenergic neuron and are thus visualized as a whole. Incubating crayfish optic neuropiles in dopamine or -methylnoradrenaline after reserpine treatment demonstrated a tangential neuron connecting the lamina and the medulla externa. The morphology of this tangential neuron differs from the two types of tangential neurons, Tan1 and Tan2, previously characterized with Golgi techniques. The catecholaminergic neuron thus constitutes a third tangential neuron type. Acknowledgement. The present study has been supported by the Swedish Natural Science Research Council, grant B 2760-009, the Magnus Bergvall foundation, and the Swedish Medical Research Council, grant 04X-712, the latter to Prof. Bengt Falck to whom we extend our gratitude. We are also indebted to Mrs. Rita Wallén and Miss Maria Walles for their skilled technical assistance. Reserpine (Serpasil®) was generously given to us by Hässle-Ciba-Geigy AB  相似文献   

16.
Summary Five monopolar cells and two long visual fibres are a consistent component of the lamina cartridge of the ventral half of the eye of the dragonfly Sympetrum rubicundulum. They communicate with the chiasm via a cartridge axon bundle comprising a minimum of ten fibres. The arrangement of these elements is documented with respect to the ommatidial photoreceptor axon bundle innervating them. These relationships are described both within the lamina cortex and in the cross-section of the underlying cartridge.  相似文献   

17.
Each neural unit (cartridge) in the first optic ganglion (lamina) of the nocturnal bee Megalopta genalis contains nine receptor cell axons (6 short and 3 long visual fibres), and four different types of first-order interneurons, also known as L-fibres (L1 to L4) or lamina monopolar cells. The short visual fibres terminate within the lamina as three different types (svf 1, 2, 3). The three long visual fibres pass through the lamina without forming characteristic branching patterns and terminate in the second optic ganglion, the medulla. The lateral branching pattern of svf 2 into adjacent cartridges is unique for hymenopterans. In addition, all four types of L-fibres show dorso-ventrally arranged, wide, lateral branching in this nocturnal bee. This is in contrast to the diurnal bees Apis mellifera and Lasioglossum leucozonium, where only two out of four L-fibre types (L2 and L4) reach neighbouring cartridges. In M. genalis, L1 forms two sub-types, viz. L1-a and L1-b; L1-b in particular has the potential to contact several neighbouring cartridges. L2 and L4 in the nocturnal bee are similar to L2 and L4 in the diurnal bees but have dorso-ventral arborisations that are twice as wide. A new type of laterally spreading L3 has been discovered in the nocturnal bee. The extensive neural branching pattern of L-fibres in M. genalis indicates a potential role for these neurons in the spatial summation of photons from large groups of ommatidia. This specific adaptation in the nocturnal bee could significantly improve reliability of vision in dim light. B.G. is grateful for travel awards from the Royal Physiographic Society, the Per Westlings Fond, the Foundation of Dagny and Eilert Ekvall and the Royal Swedish Academy of Sciences. E.J.W. acknowledges the receipt of a Smithsonian Short-Term Research Fellowship and thanks the Swedish Research Council, the Crafoord Foundation, the Wenner–Gren Foundation and the Royal Physiographic Society of Lund for their ongoing support. W.T.W. was supported by general research funds from the Smithonian Tropical Research Institute  相似文献   

18.
Whole-cell patch-clamp recordings were obtained from 116 freshly dissociated neuronal somata from the optic lobe of adult locusts (Schistocerca gregaria). Prerequisites were a papain treatment and the directed transfer of somata to the recording chamber by dabbing. Of the recorded somata, 65 were from lamina and 51 from other optic lobe neurons. All somata supported voltage-activated outward currents and some (24% of optic lobe, 3% of lamina neurons) also fast inward currents. Most lamina neurons supported an outward current that activated (V 1/2=−8.5 mV) and inactivated rapidly and a sustained outward current. Some lamina and most optic lobe neurons expressed only a sustained outward current (V 1/2=−9.4 mV). GABA and histamine elicited inward currents at negative holding potentials. Most optic lobe (95%) but only 18% of lamina neurons showed a γ-aminobutyric acid (GABA) current, whereas a similar percentage of optic lobe (50%) and lamina neurons (67%) expressed a histamine current. Both currents reversed near the chloride equilibrium potential, were reversibly reduced by picrotoxin, and did not show rundown. Thus, they likely represent chloride currents mediated by ionotropic receptors. Our data indicate that the lamina neurons recorded mainly represent monopolar cells postsynaptic to histaminergic photoreceptors. The optic lobe neurons, on which GABA and histamine apparently act as inhibitory neurotransmitters, are more heterogeneous. Accepted: 30 November 1997  相似文献   

19.
Summary The nine receptor cells examined in each ommatidium of the butterfly Papilio aegeus aegeus can be named according to their positional orientation across the fused rhabdom. Six of them end as short visual fibres (svf) in the second stratum of the lamina, whereas the remaining three retinula cells (lvf) pass together with the lamina fibres (L-fibres) the first optic ganglion and the outer chiasma to end in the three most distal layers of the second optic ganglion, the medulla. The organization of the retinula-cell axons within the pseudocartridge and the cartridge remains almost uniform throughout the first optic ganglion. Five L-fibres, which have their origin in the fenestrated layer (FL), join each laminar cartridge before entering the neuropil of the first optic region. Four of these L-fibres (L-1, L-2, L-3 and L-4) could be definitely located and characterized using Golgi-stained light- and electron-microscopic techniques. Whereas L-1 and L-3 show a lateral branching pattern reaching only fibres of the same cartridge, L-2 and L-4 have long collaterals interconnecting several neighbouring cartridges in a characteristic pattern. Serial sections of silver-impregnated retinula-cell axons as well as L-fibres were investigated for their synaptic connectivity patterns between and within these fibres. These cellular interactions and possible information processing are discussed.  相似文献   

20.
The optic nerve of Necturus maculosus consists of a homogeneous population of astroglia and bundles of unmyelinated axons. The glial cell processes ramify within the nerve roughly delineating fascicles of axons and come together at the periphery to form a complete external limiting membrane interrupted only by narrow clefts between adjacent processes. They are frequently “attached” to one another, forming specialized junctions. Blood vessels are entirely outside the nerve which is surrounded by a basal lamina. The temperature dependence of the glial membrane potential is accurately predicted by the Nernst relation. The membrane potential is unaffected by changes in Cl, Na, Li, and guanidinium which are apparently impermeant. The permeability of the glial membrane to other cations is in the sequence Tl> K> Rb> Cs> NH4. This suggests that the chemical nature of the site of potassium permeability in glial cells is similar to that in the neuron.  相似文献   

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