Anatomical identification of spectral receptor types in the retina and lamina of the Australian orchard butterfly,Papilio aegeus aegeus D.

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.     

Neural organisation in the first optic ganglion of the nocturnal bee <Emphasis Type="Italic">Megalopta genalis</Emphasis>

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     

The first optic ganglion of the bee

The nine receptor cells in each ommatidium of the worker bee end as six short visual fibres in the lamina and as three long visual fibres in the medulla. Behavioural and physiological evidence for regional variation in spectral sensitivity prompted observations on the morphology of the visual units. The distribution, branching pattern, diameter and the arrangement of axonal protusions of the characteristic receptor-cell axons were studied in various regions of the lamina. The six short visual fibres and two of the long visual fibres in each laminar cartridge are uniform over the total eye surface. Only the receptor axons of the ninth cell a UV and polarised light-sensitive cell, show obvious regional variation. In view of the regional constancy in morphology of eight of the nine receptor-cell axons, the regional variations in spectral sensitivity demand either functional subdivision of morphologically indistinguishable photoreceptors (e.g., content of different visual pigments) or a highly complex connectivity pattern of their axons in the first optic ganglion.     

The first optic ganglion of the bee

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.     

The first optic ganglion of the bee

Each visual unit (ommatidium) of the compound eye of the honey bee contains nine retinula cells, six of which end as axons in the first synaptic ganglion, the lamina, and three in the second optic ganglion, the medulla. A technique allowing light- and electron microscopy to be performed on the same silver-impregnated sections has made it possible to follow all types of retinula axons of one ommatidium to their terminals in order to study the shape of the terminal branches with their position in the cartridge. 1. The axons of retinula cells 1-6 (numbered according to Menzel and Snyder, 1974) end as three different types of short visual fibres (svf) in the lamina; the axons of retinula cells 7-9 run through the lamina to terminate in the medulla and are known as long visual fibres (lvf). Retinula cells of each type are identified by the location of their cell bodies and by the direction of their microvilli. The retinula cells 1 and 4 (group I according to Gribakin, 1967) end as svf type 1 with three tassel-like branches in stratum B of the first synaptic region. The pair of cells 3, 6 and the pair 2, 5 (group II) end in the first synaptic region in stratum A. Cells 3 and 6 have forked endings, svf type 2, whereas cells 2 and 5 have tapered endings, svf type 3. The remaining retinula cells 7, 8 and 9 have long fibres. Nos. 7 and 8 (group III) have tapered endings and are termed lvf types 1 and 2, respectively. The 9th cell is the lvf type 3 with a highly branched ending. 2. The nine axons in the bundle from one ommatidium have relative positions which do not change from the proximal retina to the monopolar cell body layer. 3. By following silver-stained retinula cells and their corresponding axons, it is possible to describe mirror-image arrangements of fibres in the axon bundles in different parts of the eye. This correlation of numbered retinula cells with specific axon types, together with the highly organized pattern in an axon bundle, allows the correlation between histological and physiological findings on polarization and colour perception.     

A neural network to improve dim-light vision? Dendritic fields of first-order interneurons in the nocturnal bee <Emphasis Type="Italic">Megalopta genalis</Emphasis>

Using the combined Golgi-electron microscopy technique, we have determined the three-dimensional dendritic fields of the short visual fibres (svf 1–3) and first-order interneurons or L-fibres (L1-4) within the first optic ganglion (lamina) of the nocturnal bee Megalopta genalis. Serial cross sections have revealed that the svf type 2 branches into one adjacent neural unit (cartridge) in layer A, the most distal of the three lamina layers A, B and C. All L-fibres, except L1-a, exhibit wide lateral branching into several neighbouring cartridges. L1-b shows a dendritic field of seven cartridges in layers A and C, dendrites of L2 target 13 cartridges in layer A, L3 branches over a total of 12 cartridges in layer A and three in layer C and L4 has the largest dendritic field size of 18 cartridges in layer C. The number of cartridges reached by the respective L-fibres is distinctly greater in the nocturnal bee than in the worker honeybee and is larger than could be estimated from our previous Golgi-light microscopy study. The extreme dorso-ventrally oriented dendritic field of L4 in M. genalis may, in addition to its potential role in spatial summation, be involved in edge detection. Thus, we have shown that the amount of lateral spreading present in the lamina provides the anatomical basis for the required spatial summation. Theoretical and future physiological work should further elucidate the roles that this lateral spreading plays to improve dim-light vision in nocturnal insects. B.G. is grateful for grants 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. would like to thank the Smithsonian Tropical Research Insitute, the Swedish Research Council, the Crafoord Foundation, the Wenner-Gren Foundation and the Royal Physiographic Society of Lund for their ongoing support.     

Golgi studies of the first optic ganglion of the ant,Cataglyphis bicolor

The neurons of the first optic ganglion (the lamina) in the desert ant, Cataglyphis bicolor, have been studied with the light microscope after Golgi silver impregnation. The different types of retinal and laminal fibres and their configuration are compared with the results obtained in the bee. The first synaptic region in the visual system of the ant lies proximally to the fenestrated layer below the basement membrane and the layer containing the monopolar cell bodies. The synaptic region can be separated into three morphologically different zones: (1) The most distal layer where the short visual fibres end at two different levels. The short visual fibres and some laminal fibres (monopolar cell fibres) also show lateral elements in this region. (2) The second layer appears almost free of branches of retinal and laminal fibres. (3) The most proximal layer, which has a characteristically dense horizontal structure resulting from the lateral elements of long visual, centrifugal, monopolar and tangential fibres. Nine cell axons arising from each ommatidium leave the retina. Six of these are short visual fibres and end at two different levels in the lamina. Three different types of short visual fibres can be distinguished by their different terminal depths and lateral branching pattern. The remaining three fibres, the long visual fibres, terminate in the medulla. They can be distinguished from each other by their lateral elements in the lamina neuropile. The five morphologically different laminal fibre types (axons of the monopolar cells in the lamina) have different shapes and different arborizations at different levels. Tangential, centrifugal and incerta sedis-fibres, which originate either from cell bodies in the cell body layer at the periphery of the outer chiasma or more centrally, terminate in the synaptic region of the lamina. Consideration is given to the clearly demarkated arrangement and length of the branching pattern of retinal and laminal fibres at different levels of the synaptic region of the lamina. In addition, a hypothetical connectivity pattern is discussed.     

Fine structure of the first optic ganglion (lamina) of the cockroach, Periplaneta americana

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.     

The Organization of the lamina ganglionaris of the prawn,Pandalus borealis (Kröyer)

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.     

The organization of the lamina ganglionaris of the hemipteran insects,Notonecta glauca,Corixa punctata and Gerris lacustris

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.     

The organization of perpendicular fibre pathways in the insect optic lobe.

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.     

The retina-lamina projection in the visual system of the bee,Apis mellifera

Single Golgi impregnated visual cells and their axons were treated from the retina to the first synaptic layer (lamina) in serial electron microscopic sections. This analysis of the retina-lamina projection was undertaken in the upper dorso-median eye region which is known to be involved in the perception of polarized light. For identification of individual visual cells and their fibres a numbering system was used which relates the number of each of the nine visual cells within one retinula to the transverse axis of the rhabdom (TRA) (Fig. 1). Because of the twist of the retinula along its course to the basement membrane (Fig. 6), individual visual cells change their position relative to any eye-constant co-ordinate system. Each axon bundle originating from one 9-celled retinula performs a 180 degrees-rotation before entering the lamina (Fig. 2). The direction of rotation (clockwise or counter-clockwise), which may differ even between adjacent bundles, is related to the two mirror-image types of rhabdoms in the corresponding retinulae and is opposite to the direction of rhabdom twist. Thus, even in small groups of the in total 5500 ommatidia in the eye of the bee, two types of retinulae exist which can be characterized by the geometry of the rhabdoms as well as by the direction of rotation of the retinulae and the axon bundles (Fig. 1). Visual cell numbers 1, 2, and 9, the microvilli of which are oriented in the direction of TRA, form three long visual fibres terminating in the second synaptic layer (medulla). In cross sections of laminar pseudocartridges they appear as the smallest fibre profiles arranged in a symmetrical line of the pseudocartridge bundle (=the transverse axis of the pseudocartridge; TPA) (Fig. 4). The remaining six fibres (cell numbers 3-8) only project to the lamina (short visual fibres; svf's). Two of them (cell numbers 5 and 6), which are the largest cells in the proximal retinula and have their microvilli perpendicularly arranged to TRA (Fig. 1), give rise to the two thickest axons of the underlaying pseudocartridge. In cross sections, t he connecting line of these two axons is orthogonally oriented to TPA (Fig. 5). A model was developed, in which all long visual fibres originate from ultraviolet receptors and in which the polarization sensitivity of the basal ninth cell is enhanced by the twist of the rhabdom. Finally, this model is discussed in light of behavioral experiments revealing the ultraviolet receptors as the only cells involved in the detection of polarized light.     

Gogo Receptor Contributes to Retinotopic Map Formation and Prevents R1-6 Photoreceptor Axon Bundling

Background

Topographic maps form the basis of neural processing in sensory systems of both vertebrate and invertebrate species. In the Drosophila visual system, neighboring R1–R6 photoreceptor axons innervate adjacent positions in the first optic ganglion, the lamina, and thereby represent visual space as a continuous map in the brain. The mechanisms responsible for the establishment of retinotopic maps remain incompletely understood.

Results

Here, we show that the receptor Golden goal (Gogo) is required for R axon lamina targeting and cartridge elongation in a partially redundant fashion with local guidance cues provided by neighboring axons. Loss of function of Gogo in large clones of R axons results in aberrant R1–R6 fascicle spacing. Gogo affects target cartridge selection only indirectly as a consequence of the disordered lamina map. Interestingly, small clones of gogo deficient R axons perfectly integrate into a proper retinotopic map suggesting that surrounding R axons of the same or neighboring fascicles provide complementary spatial guidance. Using single photoreceptor type rescue, we show that Gogo expression exclusively in R8 cells is sufficient to mediate targeting of all photoreceptor types in the lamina. Upon lamina targeting and cartridge selection, R axons elongate within their individual cartridges. Interestingly, here Gogo prevents bundling of extending R1-6 axons.

Conclusion

Taken together, we propose that Gogo contributes to retinotopic map formation in the Drosophila lamina by controlling the distribution of R1–R6 axon fascicles. In a later developmental step, the regular position of R1–R6 axons along the lamina plexus is crucial for target cartridge selection. During cartridge elongation, Gogo allows R1–R6 axons to extend centrally in the lamina cartridge.     

Photoreceptor projection and termination pattern in the lamina of gonodactyloid stomatopods (mantis shrimp)

The apposition compound eyes of gonodactyloid stomatopods are divided into a ventral and a dorsal hemisphere by six equatorial rows of enlarged ommatidia, the mid-band (MB). Whereas the hemispheres are specialized for spatial vision, the MB consists of four dorsal rows of ommatidia specialized for colour vision and two ventral rows specialized for polarization vision. The eight retinula cell axons (RCAs) from each ommatidium project retinotopically onto one corresponding lamina cartridge, so that the three retinal data streams (spatial, colour and polarization) remain anatomically separated. This study investigates whether the retinal specializations are reflected in differences in the RCA arrangement within the corresponding lamina cartridges. We have found that, in all three eye regions, the seven short visual fibres (svfs) formed by retinula cells 1–7 (R1–R7) terminate at two distinct lamina levels, geometrically separating the terminals of photoreceptors sensitive to either orthogonal e-vector directions or different wavelengths of light. This arrangement is required for the establishment of spectral and polarization opponency mechanisms. The long visual fibres (lvfs) of the eighth retinula cells (R8) pass through the lamina and project retinotopically to the distal medulla externa. Differences between the three eye regions exist in the packing of svf terminals and in the branching patterns of the lvfs within the lamina. We hypothesize that the R8 cells of MB rows 1–4 are incorporated into the colour vision system formed by R1–R7, whereas the R8 cells of MB rows 5 and 6 form a separate neural channel from R1 to R7 for polarization processing.This research was supported by the Swiss National Science Foundation (PBSKB-104268/1), the Australian Research Council (LP0214956) and the American Air Force (AOARD/AFOSR) (F62562-03-P-0227).     

brakeless is required for lamina targeting of R1-R6 axons in the Drosophila visual system

Photoreceptors in the Drosophila eye project their axons retinotopically to targets in the optic lobe of the brain. The axons of photoreceptor cells R1-R6 terminate in the first optic ganglion, the lamina, while R7 and R8 axons project through the lamina to terminate in distinct layers of the second ganglion, the medulla. Here we report the identification of the gene brakeless (bks) and show that its function is required in the developing eye specifically for the lamina targeting of R1-R6 axons. In mosaic animals lacking bks function in the eye, R1-R6 axons project through the lamina to terminate in the medulla. Other aspects of visual system development appear completely normal: photoreceptor and lamina cell fates are correctly specified, R7 axons correctly target the medulla, and both correctly targeted R7 axons and mistargeted R1-R6 axons maintain their retinotopic order with respect to both anteroposterior and dorsoventral axes. bks encodes two unusually hydrophilic nuclear protein isoforms, one of which contains a putative C(2)H(2) zinc finger domain. Transgenic expression of either Bks isoform is sufficient to restore the lamina targeting of R1-R6 axons in bks mosaics, but not to retarget R7 or R8 axons to the lamina. These data demonstrate the existence of a lamina-specific targeting mechanism for R1-R6 axons in the Drosophila visual system, and provide the first entry point in the molecular characterization of this process.     

Organization of the lamina ganglionaris of the optic lobe of the butterfly Pararge aegeria (Linné) (Lepidoptera : Satyridae)

The present work reports on a neuroanatomical study of the butterfly Pararge aegeria (Lepidoptera : Satyridae) focusing on the lamina ganglionaris underlying two different regions of the retina of the compound eye: the dorsal rim area and the large dorsal region. No differences between both lamina regions, concerning the structure of the cartridges and the morphology of the identified neurons, could be detected. After passing the basement membrane, the visual cell axons are organized in retinotopic bundles (pseudocartridges), in which the axons of the 9 visual cells (V1 and 5, D2, 4, 6, 8, H3 and 7, B9) are arranged in the same way as in the retina. In the pseudocartridge there are no synaptic contacts. Before entering the lamina cartridge, the bundles rotate 90 °. The cartridges are joined by the fibres of 4 monopolar cells (L1, L2, L3 and L4), which could be identified and located inside the lamina cartridges in serial EM-sections. Golgi impregnations revealed the morphology of these fibres. Thus, the regional specialization of the retina (dorsal rim area and large dorsal region) does not seem to be reflected at the level of the first visual neuropil. Additionally, the cartridges of both lamina regions were investigated qualitatively for synaptic contacts among fibres. In addition to monadic chemical synapses and multiple contact synapses with presynaptic ribbons, cell contacts are also facilitated by invaginations and bridges. These cellular interactions and their functional implications are discussed.     

The organisation of the lamina ganglionaris of the crabs Scylla serrata and Leptograpsus variegatus

Summary The gross structure and neuronal elements of the first optic ganglion of two crabs, Scylla serrata and Leptograpsus variegatus, are described on the basis of Golgi (selective silver) and reduced silver preparations. Of the eight retinula cells of each ommatidium, seven end within the lamina, while the eighth cell sends a long fibre to the external medulla. Five types of monopolar neurons are described, three types of large tangential fibres, and one fibre which may be centrifugal. The marked stratification of the lamina is produced by several features. The main synaptic region, the plexiform layer, is divided by a band of tangential fibres; the short retinula fibres end at two levels in the plexiform layer; and two types of monopolar cells have arborisations confined to the distal or proximal parts of the plexiform layer. The information presently available concerning the retina-lamina projection in Crustacea is examined. Some of the implications of retina and lamina structure are discussed in conjunction with what is known about their electrophysiology.     

Neurosecretory cells in the optic lobes of the brain and activity rhythms in Lycosa tarentula (Araneae: Lycosidae)

Several paired groups of neurosecretory cells (NS) were identified in the dorsal cortical neurons of the optic lobes of the brain of Lycosa tarentula (Araneae). Two large bottle-shaped cells (NS A1, A2) and a cluster of ca. 20 smaller cells (NS B) were found between the lamina and medulla of the anterior median eyes (AM). The forward oriented bundles of NS B axons run alongside large fibres linked to the synaptic zones of the indirect eyes. In front of the arcuate body, an islet of about 10 fusiform cells (NS C1) sends short axons close to the internal cortical border. Other large cells (NS C2, C3) are found from the medulla of the AM to the anterior border of the central body. Their long axons end deeply in the brain neuropil. NS B and C1 function synchronously. The secretory cycles of NS A1 and A2 seem to be in opposition. The activity of these three types of NS depends on the phase of the day. Anatomical relationships of NS A, B and C1 with visual afferent/efferent fibres via synaptic buttons indicate a role of these cells in the modulation of circadian rhythms of visual and locomotor activity. On the other hand, NS C2 and C3, the functioning of which is not synchronous, might be involved in the modulation or control of the elementary movements of L. tarentula when active or at rest.     

Ocellar projections within the central nervous system of the worker honey bee,Apis mellifera

Summary The projections of ocellar fibres within the brain and thorax of the honey bee, Apis mellifera, were established using a modified cobalt sulphide technique, supplemented by serial sectioning of the brain for the light microscope.The results are: 5 large fibres in each lateral nerve and 12 in the median nerve have wide-field terminal arborisations in ocellar association areas on either side of the posterior slope area. 9 medium-sized fibres in each lateral nerve and 12 in the median nerve form a second ocellar association area on each side of the perioesophageal foramen. A group of fine fibres, stained via the ocellar nerves, arborise just below and anterior to the protocerebral bridge. 10 medium-sized fibres run from the level of the ocellar nerve tracts to the first and second thoracic ganglia, branching in a number of discrete areas within each ganglion. These fibres also form a restricted ocellar association area within the suboesophageal ganglion. A few fibres run between the higher-order optic centres and the ocellar tract. The large- and mediumsized fibres give off short, stout spines from their axons within the ocellar tracts.