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• W2988649809 abstract "Article Figures and data Abstract Introduction Results Discussion Materials and methods References Decision letter Author response Article and author information Metrics Abstract Understanding memory formation, storage and retrieval requires knowledge of the underlying neuronal circuits. In Drosophila, the mushroom body (MB) is the major site of associative learning. We reconstructed the morphologies and synaptic connections of all 983 neurons within the three functional units, or compartments, that compose the adult MB’s α lobe, using a dataset of isotropic 8 nm voxels collected by focused ion-beam milling scanning electron microscopy. We found that Kenyon cells (KCs), whose sparse activity encodes sensory information, each make multiple en passant synapses to MB output neurons (MBONs) in each compartment. Some MBONs have inputs from all KCs, while others differentially sample sensory modalities. Only 6% of KC>MBON synapses receive a direct synapse from a dopaminergic neuron (DAN). We identified two unanticipated classes of synapses, KC>DAN and DAN>MBON. DAN activation produces a slow depolarization of the MBON in these DAN>MBON synapses and can weaken memory recall. https://doi.org/10.7554/eLife.26975.001 Introduction Associative memory helps animals adapt their behaviors to a dynamically changing world. The molecular mechanisms of memory formation are thought to involve persistent changes in the efficiency of synaptic transmission between neurons. In associative learning, persistent changes in synaptic efficacy correlated with memory formation have been found at points of convergence between two neuronal representations: one providing information from sensory inputs about the outside world and a second indicating whether the current environment is punitive or rewarding. Such sites of convergence have been identified for multiple forms of associative learning (Medina et al., 2002; Ardiel and Rankin, 2010; Tovote et al., 2015; Kandel and Schwartz, 1982). However, a comprehensive synaptic level description of connectivity at such a site of convergence is not available for an animal as complex as the fruit fly, Drosophila. The mushroom body (MB) is the center of associative learning in insects (Erber et al., 1980; Heisenberg et al., 1985; de Belle and Heisenberg, 1994; Dubnau et al., 2001; McGuire et al., 2001; Mizunami et al., 1998). Sensory information enters the MB via the calyx, where the dendritic claws of Kenyon cells (KCs) receive synaptic inputs from projection neurons of olfactory and other modalities including visual, gustatory and thermal (Vogt et al., 2016; Kirkhart and Scott, 2015; Yagi et al., 2016; Caron et al., 2013; Stocker et al., 1990; Wong et al., 2002; Strausfeld, 1976; Tanaka et al., 2004; Liu et al., 2015; Frank et al., 2015). The parallel axonal fibers of the KCs form the MB-lobes, the output region of the MB. A pattern of sparse activity in the KC population represents the identity of the stimulus. This sparseness is maintained through two mechanisms. First, individual KCs generally only spike when they receive simultaneous inputs from multiple projection neurons (Gruntman and Turner, 2013). Second, overall KC excitability is regulated by feedback inhibition from a GABAergic neuron, MB-APL, that arborizes throughout the MB (Papadopoulou et al., 2011; Lin et al., 2014a; Tanaka et al., 2008; Liu and Davis, 2009). Thus, only a small subset of KCs respond to a given sensory stimulus (Perez-Orive et al., 2002; Turner et al., 2008; Honegger et al., 2011; Murthy et al., 2008). Upon this representation of the sensory world, dopaminergic or octopaminergic neurons convey information of punishment or reward and induce memories that associate the sensory stimulus with its valence (Schroll et al., 2006; Schwaerzel et al., 2003; Liu et al., 2012; Burke et al., 2012; Riemensperger et al., 2005; Mao and Davis, 2009; Heisenberg, 2003; Claridge-Chang et al., 2009). The functional architecture of the MB circuit is best understood in adult Drosophila (Figure 1) (Ito et al., 1998; Lin et al., 2007; Tanaka et al., 2008; Strausfeld et al., 2003; Crittenden et al., 1998; Ito et al., 1997; Aso et al., 2014a; Pech et al., 2013). In each MB, the parallel axonal fibers of ~2000 KCs can be divided into 16 compartmental units by the dendrites of 21 types of MB output neurons (MBONs) and the axon terminals of 20 types of dopaminergic neurons (DANs). A large body of behavioral and physiological studies suggests that these anatomical compartments are also parallel units of associative learning (see e.g. Hige et al., 2015a; Lin et al., 2014b). In each compartment, the dendrites of a few MBONs overlap with axon bundles of hundreds of KCs. Punishment and reward activate distinct sets of DANs. DAN input to a compartment has been shown to induce enduring changes in efficacy of KC>MBONs synapses in those specific KCs that were active in that compartment at the time of dopamine release (Hige et al., 2015a). The valence of the memory appears to be determined by which compartment receives dopamine during training, while the sensory specificity of the memory is determined by which KCs were active during training (Liu et al., 2012; Heisenberg, 2003; Burke et al., 2012). Figure 1 Download asset Open asset Diagram of the α lobe of the mushroom body. (A) An image of the adult brain showing the antennal lobes (AL), the mushroom bodies (MB) and an example of one of the ~50 types of projection neurons (PN) that carries olfactory information from the AL to the MB calyx and the lateral horn (LH). See Aso et al. (2014a) for more detail. The approximate position of the ~40 × 50 x 120 μm volume imaged by FIBSEM is indicated by the red dashed lines. (B) Magnified view of the α/β lobes showing the imaged volume. The α/β neurons bifurcate in the α1 compartment and project to the α and β lobes. The white box indicates the portion of the α3 compartment shown in Video 2. (C) Simplified diagram of the circuit organization in the α lobe. The projection patterns of the axons of dopaminergic neurons (DANs) and the dendrites of the MB output neurons (MBONs) onto the parallel axonal fibers of Kenyon cells define three compartmental units in the α lobe. The DANs (green) and MBONs with dendrites in the α1, α2 and α3 compartments (purple), known from previous light microscopic studies (see Aso et al., 2014a for more detail), are indicated. Arrows indicate the main presynaptic sites of each of the extrinsic neuron types. The names of neurons (shown in the rectangles with rounded corners) are color-coded to reflect their main neurotransmitter: black, dopamine; orange, acetylcholine; green, glutamate; blue, GABA. In addition to MBONs with dendrites in the α lobe, all three compartments receive projections from the GABAergic MBON-γ1pedc>α/β (dark blue) and the glutamatergic MBON-β1>α feedforward neurons (magenta), whose dendrites lie in other MB lobes. https://doi.org/10.7554/eLife.26975.002 Compartments can have distinct rates of memory acquisition and decay, and the 16 compartments together appear to form a set of parallel memory units whose activities are coordinated through both direct and indirect inter-compartmental connections (Aso and Rubin, 2016; Cohn et al., 2015; Perisse et al., 2016; Aso et al., 2014a). The DANs which project to the α1 compartment, the ventral-most compartment of the vertical lobe (Figure 1), play a key role in the formation of appetitive long-term memory of nutritional foods (Yamagata et al., 2015). DANs that project to the other α lobe compartments, α2 and α3, play roles in aversive long-term memory (Aso and Rubin, 2016; Séjourné et al., 2011; Pai et al., 2013). All three of these compartments receive feedforward inputs from GABAergic and glutamatergic MBONs whose dendrites lie in other MB compartments (Aso et al., 2014a) known to be involved in aversive or appetitive memory (Aso and Rubin, 2016; Aso et al., 2010; Burke et al., 2012; Perisse et al., 2016). In addition, two types of MB-intrinsic neurons send arbors throughout the MB-lobes: a large GABAergic neuron, MB-APL, which provides negative feedback important for sparse coding (Papadopoulou et al., 2011; Lin et al., 2014a), and the MB-DPM neuron, which is involved in memory consolidation and sleep regulation (Waddell et al., 2000; Keene et al., 2006; Haynes et al., 2015; Yu et al., 2005a; Cervantes-Sandoval and Davis, 2012; Keene et al., 2004). Previous EM studies in the MB lobes of cockroaches (Mancini and Frontali, 1970, Mancini and Frontali, 1967), locusts (Leitch and Laurent, 1996), crickets, ants, honey bees (Schürmann, 1974, 2016) and Drosophila (Technau, 1984) identified KCs by their abundance, fasciculating axons and small size. Additionally, Leitch and Laurent (1996) identified large GABA immunoreactive neurons that contact KC axons in the locust pedunculus. While these data provided early insights to guide modeling of the MB circuit, the volumes analyzed were limited and most neuronal processes could not be definitively assigned to specific cell types. In this paper, we report a dense reconstruction of the three compartments that make up the α lobe of an adult Drosophila male (Figure 1). Because we performed a dense reconstruction, with the goal of determining the morphology and connectivity of all cells in the volume, we have confidence that we have identified all cell types with processes in the α lobe. Comprehensive knowledge of the connectivity in the α lobe has allowed us to address several outstanding issues. The first concerns the nature of KC>MBON connectivity. Although each KC passes through all three compartments, it is not known if individual KCs have en passant synapses in each compartment. Thus, it remains an open question whether the sensory representation provided to each compartment and each MBON within a compartment is the same or whether different MBONs within a compartment might sample from non-overlapping sets of KCs, and thus use independent sensory representations for learning. It was also not known which, if any, other cell types are direct postsynaptic targets of KCs. The second concerns dopamine modulation. What are the locations of dopaminergic synapses and what does this distribution imply about the targets of dopaminergic modulation as well as volume versus local transmission? Cell-type-specific rescue of dopamine receptor mutants suggests that dopamine acts presynaptically in the KCs of KC>MBON synapses (Kim et al., 2007; Qin et al., 2012a; Liu et al., 2012; Ichinose et al., 2015). However, postsynaptic mechanisms have also been proposed (Cassenaer and Laurent, 2012; Pai et al., 2013) and a recent study detected expression of dopamine receptors in MBONs (Crocker et al., 2016), raising the possibility that MBONs might also be direct targets of DAN modulation. Behavioral, imaging and electrophysiological data (Aso et al., 2010; Hige et al., 2015a; Cohn et al., 2015) indicate that dopamine modulation respects the borders between compartments, but we do not know whether these borders have a distinct structure, such as a glial sheet. The third concerns the two MBON types that send feedforward projections into the α lobe. These MBONs have important roles in associative learning as revealed by behavioral assays and have been postulated to integrate memories of opposing valence and different time scales (Aso et al., 2014a; Aso and Rubin, 2016; Aso et al., 2014b; Perisse et al., 2016). However, we do not know which cell types these feedforward MBON projections targets within the MB. The fourth concerns the two neurons, MB-APL and MB-DPM, which arborize throughout the MB and are thought to regulate MB function globally (Liu and Davis, 2009; Lin et al., 2014a). What is their local synaptic connectivity within the α lobe and what can this tell us about how they perform their roles? Finally, the three compartments of the α lobe differ in important aspects, including valence of the memory formed, the time course of memory formation and retrieval, and the numerical complexity of their DAN inputs and MBON outputs. Are there obvious differences in the microcircuits of different compartments? In this paper, we report the answers to these questions. In addition, we demonstrate the utility of detailed anatomy at the electron microscopic level to provide novel insights: We show that nearly all cell types in the α lobe contain more than one morphological class of synaptic vesicle, raising the possibility that these cells utilize multiple neurotransmitters. In addition, we describe two prevalent sets of synaptic motifs—from DANs to MBONs and from KCs to DANs—that were unanticipated despite the extensive anatomical, physiological, behavioral and theoretical studies that have been performed on the insect MB. We characterize these novel DAN to MBON connections using behavioral and physiological assays and find that DAN activation produces a slow depolarization of postsynaptic MBONs and can weaken memory recall. Results Data acquisition, segmentation and proofreading The brain of a 5-day-old adult male fly was fixed, embedded and trimmed as described in Materials and methods. A ~40 × 50 x 120 μm volume (Figure 1B) encompassing the vertical lobe of the MB was imaged by focused ion-beam milling scanning electron microscopy (FIBSEM) (Xu et al., 2017) over a 5-week imaging run (Videos 1 and 2). The assembled volume has isotropic voxels (8 × 8 × 8 nm) allowing image data to be viewed with the same resolution along any axis. Video 1 Download asset This video cannot be played in place because your browser does support HTML5 video. You may still download the video for offline viewing. Download as MPEG-4 Download as WebM Download as Ogg A portion of the dataset that was used for connectome reconstruction shown at down-sampled resolution. Approximately, 9600 sequential x-y imaging planes are shown covering a 35 × 35 × 77 µm region of the complete image volume (40 × 50 × 120 µm). The original voxel size was 8 × 8 × 8 nm; the video has been down sampled by a factor of eight, making the voxel size shown 64 × 64 × 64 nm. The video progresses from top of the vertical lobe, which is ensheathed in glia, through the α3 and α2 compartments as indicated by the black bracket in Figure 1B. https://doi.org/10.7554/eLife.26975.003 Video 2 Download asset This video cannot be played in place because your browser does support HTML5 video. You may still download the video for offline viewing. Download as MPEG-4 Download as WebM Download as Ogg A portion of the data set that was used for connectome reconstruction shown at the resolution at which the data was acquired, 8 × 8 × 8 nm voxels. The region shown corresponds to the portion of the α3 compartment indicated by the white box in Figure 1B. https://doi.org/10.7554/eLife.26975.004 The portion of the imaged volume that contained the α lobe was identified based on the morphologies of the KCs and ensheathing glia. We then reconstructed the shapes of the individual neurons in this selected volume, as well as mapped the locations of synapses. This process entailed the application of machine vision algorithms for synapse detection and image segmentation—that is, assigning each voxel to a particular neuron. These procedures have been previously published (Takemura et al., 2015; Plaza et al., 2014; Parag et al., 2014) and are briefly described in Materials and methods. The results of these automated processes were then reviewed and edited by trained human proofreaders; a total of ~8 person years was devoted to proofreading. The neuronal processes that entered or left the α lobe were traced until they exited the imaged volume; this information was helpful in distinguishing cell types, as described below. Cell type identification Because of the constrained size of the imaged volume, only the portions of the neurons that have processes in the α lobe were reconstructed. To identify the cell type of each partially reconstructed cell in the EM volume, we compared their morphologies to existing images of the relevant MB cell types from light microscopy (Figure 2) (Aso et al., 2014a). Our confidence in our ability to make correct correspondences by this approach was increased by the completeness of both EM and light microscopy datasets. We found only one cell type in our EM reconstructions that had not been described at the light level, a single MBON that we named (MBON-α2sp). All the other reconstructed arbors could be assigned to one of the neurons previously identified at the light level (Figures 2 and 3) except in the α1 compartment, where we reconstructed a few arbors that were not large enough to allow unambiguous assignment based on comparing light and EM morphologies and whose branches exited the imaged volume before connecting to an identified cell. Based on comparison with light microscopic anatomy (Aso et al., 2014a), we expect that these are segments of APL, DPM and MBON-γ1pedc>α/β (see below). One other difference between the observed light and EM morphologies was that MBON-α2p3p extends a few dendrites into α3 based on light microscopic analyses (Aso et al., 2014a), whereas in our EM reconstruction of this cell we found dendritic arborizations were confined to α2. Figure 2 Download asset Open asset Reconstructions of cells present in the α lobe. In panels (A-M) and (O), the upper image shows EM reconstructions generated as part of this study and the lower image shows the same cell type, segmented from previously acquired light microscopic images (Aso et al., 2014a). The EM reconstructions are limited to that portion of the neurons found in the α lobe, while the light images show the portion of each neuron found in the entire MB. (A) A total of 949 α/β Kenyon cells (KCs) were traced: 871 surface and core KCs (khaki); 78 posterior KCs (yellow). (B) The glutamatergic feedforward neuron, MBON-β1>α, arborizes in all three compartments of the α lobe. (C) The arborizations of the ipsi- and contralateral MBON-γ1pedc>α/β, GABAergic feedforward neurons, are shown separately in the upper panel. (D) The GABAergic APL neuron arborizes throughout the MB lobes and calyx. (E) The DPM neuron arborizes throughout the MB lobes. (F) The SIFamide neuron arborizes very widely, extending throughout the brain; only the αlobearborizations are shown. Panels (G-M) and (O) show compartment-specific MB output neurons (MBONs) and dopaminergic neurons (DANs). The α3 compartment has the axonal terminals of two DANs, PPL1-α3 (G), and the dendrites of two MBONs, MBON-α3 (H). The α2 compartment has two DANs, PPL1-α’2α2 (I), and four MBONs: a single MBON-a2sc (J); two MBON-α2p3p (K); and one newly found MBON, MBON-α2sp (L). The α1 compartment has 16 DANs, PAM-α1 (M in aggregate and N as individual cells), and two MBONs, MBON-α1 (O). https://doi.org/10.7554/eLife.26975.005 Figure 3 Download asset Open asset Profiles of reconstructed neurons in an EM cross-section at the depth of α3. (A) All the reconstructed neurons that have neurites at this depth are color-labeled using the same color scheme as in Figure 2. (B-E) Subsets of cell types are shown separately: (B) dendrites of the two MBON-α3 cells; (C) axonal projections of the two PPL1-α3 DANs; (D) axonal feedforward projects of MBON-β1>α and MBON-γ1pedc>α/β, of which only a few small profiles can be seen in a single section; and (E) APL, DPM and SIFamide neurons. Scale bars: 5 µm. https://doi.org/10.7554/eLife.26975.006 We reconstructed 949 KCs in the α lobe, a number that agrees well with the previous estimate of ~1000 α/β KCs obtained by counting genetically labeled cell nuclei with light microscopy (Aso et al., 2009). The α lobe KCs have been divided into three classes based on the location of their axons in the lobe: posterior (α/βp), surface (α/βs) or core (α/βc) (Tanaka et al., 2008; Strausfeld et al., 2003; Lin et al., 2007). The α/βp cells are clearly distinct in both morphology and synaptic connectivity (see below) and we assigned 78 neurons to this class, compared to ~90 estimated by Aso et al. (2014a). The remaining α/βs and α/βc KCs form a set of concentric layers in the α lobe arranged by birth order, with KCs that are born more recently occupying the more central, or core, layers. Following established nomenclature, we refer to KCs that occupy the outer most layer of the α lobe as surface, α/βs, and those occupying the inner layers as core, α/βc; the core KCs can be further divided into inner-core, α/βc(i) and outer-core, α/βc(o) (Tanaka et al., 2008). When the distinction is unimportant, we simply refer to the non-posterior KCs collectively as α/βsc. The relative spatial arrangements of these KC classes is illustrated in Figures 2A and 3A and Video 3. Video 3 Download asset This video cannot be played in place because your browser does support HTML5 video. You may still download the video for offline viewing. Download as MPEG-4 Download as WebM Download as Ogg All KCs in the α lobe, except outer-core (α/βc(o)). A total of 259 α/βc(i), 480 α/βs, and 78 α/βp KCs are shown colored in ivory, orange and yellow, respectively. https://doi.org/10.7554/eLife.26975.007 The α lobe, a linear structure formed by the continuous axons of the KCs, can be divided into three non-overlapping compartments, α1, α2 and α3 (Figures 1 and 2). Each compartment has a unique set of DANs and MBONs whose complex dendritic arbors demarcate the extent of the compartment (Figure 1; Video 4). The α3 compartment at the tip of the α lobe contains two PPL1-α3 DANs and two MBON-α3 cells (Figure 2G and H). The α2 compartment has two PPL1-α′2α2 DANs (neurons that innervate both the α2 compartment and the α′2 compartment of the α′ lobe) and four MBONs of three distinct types that differ based on the KCs they receive input from: one MBON-α2sc, two MBON-α2p3p and one MBON-α2sp (Figure 2I–L). The α1 compartment has 16 PAM-α1 DANs and 2 MBON-α1 cells (Figure 2M–O; Video 5). These cell numbers are per hemisphere for MBONs but per brain for DANs because each DAN innervates the MB in both hemispheres. In some cases, the distinction of arbors of the MBONs and DANs were not very clear in each compartment. In these cases, we used the characteristic axonal positions at which the neurites of these cells enter the MB lobes (Aso et al., 2014a) in making cell-type assignments. For example: PPL1-α3’s main axons all enter the lobe from the posterior side, whereas MBON-α3’s axons enter from the medial side. DPM has a thick main axon entering into the α lobe from the posterior medial side (Waddell et al., 2000) (see Figure 3—figure supplement 1 in Aso et al., 2014a), whereas APL has a very thin axon entering the vertical lobe from the posterior side. Further confirmation for the identities of the cell types assigned to these reconstructed arbors was provided by their distinct synaptic connectivity. For example, early in the process we found MBON dendritic arbors had no pre-synaptic sites, APL was pre-synaptic to KCs but not DANS, and DPM was pre-synaptic to both. These patterns enabled us to double-check the assignments that we had made based on morphology. Video 4 Download asset This video cannot be played in place because your browser does support HTML5 video. You may still download the video for offline viewing. Download as MPEG-4 Download as WebM Download as Ogg Tiling of the MBONs and DANs in the α lobe. Neurites of MBONs and DANs are confined to a single compartment where they are intermingled. Two MBON-α3, two PPL1-α3 and the one MBON-α2sc are shown in sequence. https://doi.org/10.7554/eLife.26975.008 Video 5 Download asset This video cannot be played in place because your browser does support HTML5 video. You may still download the video for offline viewing. Download as MPEG-4 Download as WebM Download as Ogg PAM-α1 DANs. Individual morphologies of 16 PAM-α1 neurons are displayed in sequence showing how the terminals of these cells collectively fill the compartment. https://doi.org/10.7554/eLife.26975.009 There are six additional cells with arbors in the α lobe, and each innervates all three compartments (Figure 2B–F): the ipsilateral APL and DPM, MB-intrinsic neurons that arborize widely throughout the MB lobes; a neuron expressing the neuropeptide SIFamide that arborizes broadly throughout the brain (Park et al., 2014; Verleyen et al., 2004); and the axons of the ipsilateral MBON-β1>α and the ipsi- and contralateral MBON-γ1pedc>α/β, which project from other MB lobes. In total, we reconstructed and identified portions of 983 neurons in the α lobe. Since we accounted for all major neurites and neuronal profiles and had only small fragmented bodies left unassigned to a specific neuronal type in the reconstructed volume (see below for the quantitative estimate), we are confident that there are no other cell types with significant arborization in the α lobe. Synapse number and morphology The resolution provided by EM allowed us to determine the number and location of chemical synapses between the cells we identified, information that was not available from previous light level analyses. We identified 89,406 presynaptic densities (Figures 4 and 5), using a combination of machine learning algorithms and human annotation. We then manually annotated a total of 224,697 postsynaptic sites in the α lobe, based primarily on their adjacency to a presynaptic density (see Materials and methods). Of these postsynaptic sites, 93% could be traced back to the main arbors of an identified cell. The remaining 7% of postsynaptic sites were typically in small branches that could not be reliably traced to a particular cell; the small size and discontinuous nature of these neurites indicates that they are fragments of identified cells rather than collectively constituting an additional cell type. For 86% of synapses, we were able to identify the cell types of both the pre- and postsynaptic cells. The fact that we did a dense reconstruction, mapping the vast majority of synapses, allowed us to determine quantitative properties of the network of neuronal connections that would not be revealed by a sparser sampling approach. Figure 4 shows examples of the synaptic morphologies we observed in the MB α lobe, and Figure 5 shows examples from a higher quality dataset (with 4 × 4 × 4 nm voxels and imaged with higher signal to noise) collected from selected regions of a second brain. While the ~100 x slower imaging required to collect data at this higher resolution precluded imaging the entire volume, these selected areas allowed us to catalog the various synaptic motifs present in the MB with greater confidence. Figure 4 with 2 supplements see all Download asset Open asset Examples of synaptic motifs in the α lobe. (A) Five KCs are shown, converging once in the α lobe to form a rosette synapse (arrow). (B,C) EM cross-section of the rosette synapse formed by these five KCs. Each KC is colored in (B) with the same color as the corresponding reconstructed cell in (A). (C) The same EM image as (B) with a dendrite of an MBON (asterisk). Presynaptic specializations of the KCs are indicated by red arrowheads at which KCs contact with both the MBON and neighboring KC. (D) A PAM-α1 dopaminergic neuron synapses onto MBON-α1 and KCs in the α1 compartment; the red arrowhead marks the presynaptic specialization in PAM-α1. (E) Two KCs synapse onto a PPL1-α3 dopaminergic neuron. An adjacent MBON (asterisk) also appears to receive input from one of these KCs. (F) The MBON-β1>α feedforward neuron makes an axon-axonal synapse onto the MBON-γ1pedc>α/β feedforward neuron, as well as a synapse onto MBON-α3 dendrites, in the α3 compartment; the presynaptic specialization in MBON-β1>α is marked by a red arrowhead. (G) The MBON-γ1pedc>α/β feedforward neuron synapses onto MBON-α3 dendrites; the presynaptic specialization in MBON-γ1pedc>α/β is marked by a red arrowhead. Scale bar: 500 nm, applies to panels (B)-(G). https://doi.org/10.7554/eLife.26975.010 Figure 5 Download asset Open asset Images from the higher resolution dataset and examples of the distribution of synapses on a KC. (A) A triangular motif of KC<>KC>MBON synapses. Presynaptic densities in two adjacent KCs (arrowheads) contact to an MBON (asterisk); the KCs also appear to make reciprocal contacts. (B) A rosette synapse formed by a postsynaptic MBON (asterisk) surrounded by five KCs. (C) The α3 and α2 portion of a core KC that has a total of 63 presynaptic sites in the α lobe (red puncta) is shown. This KC makes 49 synapses onto MBONs; the remaining 14 synapses are onto other cell types such as APL and DPM. (D) Sites where the same KC as in (C) is postsynaptic (black puncta) are also shown: Of the 114 inputs this KC receives in the α lobe, 94 come from 65 other core KCs; 13 from 11 different surface KCs; four from DANs (three times in α3 and once in α1); and three from APL. Note that because multiple synapses can occur in close proximity, the number of distinct puncta visible is smaller than the number of synapses and that red and black puncta are often co-localized, indicating the KC is pre- and postsynaptic at the same site on its axon. (E) We found three kinds of synaptic vesicles in neurons in the α lobe: rounded clear vesicles (white arrowheads), small-r" @default.
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• W2988649809 title "Author response: A connectome of a learning and memory center in the adult Drosophila brain" @default.
• W2988649809 doi "https://doi.org/10.7554/elife.26975.037" @default.
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