Catabolite repression of the Bacillus subtilis gnt operon exerted by two catabolite-responsive elements

Catabolite repression of Bacillus subtilis catabolic operons is supposed to occur via a negative regulatory mechanism involving the recognition of a cis-acting catabolite-responsive element (cre) by a complex of CcpA, which is a member of the GalR-LacI family of bacterial regulatory proteins, and the seryl-phos-phorylated form of HPr (P-ser-HPr), as verified by recent studies on catabolite repression of the gnt operon. Analysis of the gnt promoter region by deletions and point mutations revealed that in addition to the ere in the first gene (gntR) of the gnt operon (cre_down), this operon contains another ere located in the promoter region (cre_up). A translational gntR-lacZ fusion expressed under the control of various combinations of wild-type and mutant cre_down and cre_up was integrated into the chromosomal amyE locus, and then catabolite repression of p-galac-tosidase synthesis in the resultant integrants was examined. The in vivo results implied that catabolite repression exerted by cre_up was probably independent of catabolite repression exerted by cre_down; both creup and cre_down catabolite repression involved CcpA. Catabolite repression exerted by cre_up was independent of P-ser-HPr, and catabolite repression exerted by cre_down was partially independent of P-ser-HPr. DNase I footprinting experiments indicated that a complex of CcpA and P-ser-HPr did not recognize cre_up, in contrast to its specific recognition of cre_down. However, CcpA complexed with glucose-6-phosphate specifically recognized cre_up as well as cre_down, but the physiological significance of this complexing is unknown.     

Neuregulin 3 promotes excitatory synapse formation on hippocampal interneurons

Hippocampal GABAergic interneurons are crucial for cortical network function and have been implicated in psychiatric disorders. We show here that Neuregulin 3 (Nrg3), a relatively little investigated low‐affinity ligand, is a functionally dominant interaction partner of ErbB4 in parvalbumin‐positive (PV) interneurons. Nrg3 and ErbB4 are located pre‐ and postsynaptically, respectively, in excitatory synapses on PV interneurons in vivo. Additionally, we show that ablation of Nrg3 results in a similar phenotype as the one described for ErbB4 ablation, including reduced excitatory synapse numbers on PV interneurons, altered short‐term plasticity, and disinhibition of the hippocampal network. In culture, presynaptic Nrg3 increases excitatory synapse numbers on ErbB4⁺ interneurons and affects short‐term plasticity. Nrg3 mutant neurons are poor donors of presynaptic terminals in the presence of competing neurons that produce recombinant Nrg3, and this bias requires postsynaptic ErbB4 but not ErbB4 kinase activity. Furthermore, when presented by non‐neuronal cells, Nrg3 induces postsynaptic membrane specialization. Our data indicate that Nrg3 provides adhesive cues that facilitate excitatory neurons to synapse onto ErbB4⁺ interneurons.     

Control of the modularity of {Mo2O2S2}-containing polyoxometalates. Synthesis, structure and characterization of tri- and tetra-modular assemblies based on the [β-B-AsW9O33] anion

Reaction at pH = 2.3 of the [Mo₂O₂S₂(OH₂)₆]²⁺ aqua cation with the tetravacant ion [β-B-HAs₂W₈O₃₁]⁷⁻ leads to the formation of a red solid from which three mixed salts have been obtained as single crystals and characterized by X-ray diffraction analysis. Three mixed salts K-5a, RbNa-5b, DMACs-5c exhibit a similar molecular arrangement consisting in three {β-HAs₂W₉O₃₄} subunits mutually linked by three {Mo₂O₂S₂} groups. The triangular arrangement delimits a large open-cavity, lined on the periphery by three outer {As-OH} groups and closed at the bottom by a small hexagonal pocket formed by six terminal oxygen atoms. The central hexagonal cavity is filled either by a potassium, a rubidium or a cesium cation. The outer {As-OH} groups are pointed towards two directions labelled up and down, respectively. In K-6a the three {As-OH} bonds are in up configuration leading to the {up, up, up} isomer. The structure of RbNa-5b is rather consistent with the superposition of the two {up, up, up} and {down, up, up} isomers disordered over the same crystallographic site, while only the {down, up, up} isomer is present in DMACs-5c. In solution, ¹⁸³W NMR characterization of 6a as sodium salt results in a complicated spectrum consistent with the simultaneous presence of the four isomers, {up, up, up}, {down, up, up}, {down, down, up} and {down, down, down}, respectively. 5a reacts with three equivalents of iodine to give CsNa-6 isolated as single crystals. In 6, four β-{AsW₉O₃₃} moieties are located at the corner of a super tetrahedron and are mutually connected by six {Mo₂O₂S₂} linkers. The three outer {As-OH} groups can be selectively removed by iodine, this oxidation reaction consisting in fact in a deprotecting process permitting the extension of the arrangement from triangular to tetrahedral.     

The dynamical stability of reverberatory neural circuits

The concept of reverberation proposed by Lorente de Nó and Hebb is key to understanding strongly recurrent cortical networks. In particular, synaptic reverberation is now viewed as a likely mechanism for the active maintenance of working memory in the prefrontal cortex. Theoretically, this has spurred a debate as to how such a potentially explosive mechanism can provide stable working-memory function given the synaptic and cellular mechanisms at play in the cerebral cortex. We present here new evidence for the participation of NMDA receptors in the stabilization of persistent delay activity in a biophysical network model of conductance-based neurons. We show that the stability of working-memory function, and the required NMDA/AMPA ratio at recurrent excitatory synapses, depend on physiological properties of neurons and synaptic interactions, such as the time constants of excitation and inhibition, mutual inhibition between interneurons, differential NMDA receptor participation at excitatory projections to pyramidal neurons and interneurons, or the presence of slow intrinsic ion currents in pyramidal neurons. We review other mechanisms proposed to enhance the dynamical stability of synaptically generated attractor states of a reverberatory circuit. This recent work represents a necessary and significant step towards testing attractor network models by cortical electrophysiology.     

Synaptic interactions between neurons involved in the production of abdominal posture in crayfish

Summary Pairs of neurons that produce or influence motor outputs in the abdominal positioning system of the crayfish (Procambarus clarkii) were impaled in isolated nerve cords with Lucifer Yellow-filled microelectrodes to determine their morphologies and the nature and extent of the synaptic interactions between them. Although the motor programs for positional adjustments can be produced by directly stimulating single interneurons, we found extensive interactions between these neurons, often involving the recruitment of one interneuron by another. The data indicate that the positioning interneurons do not operate as labelled lines, each independently producing a discrete position. Pairs of interneurons, each producing similar motor outputs when activated, were often found to be connected by unidirectional excitatory synapses. In contrast, central inhibition was commonly found between pairs of interneurons that produced antagonistic motor effects. Finally, the unidirectional interactions between positioning interneurons revealed a hierarchy of at least two tiers in this system. Based on these observations, we suggest that abdominal positioning in crustaceans is produced by constellations of interacting interneurons.Abbreviations FPI flexion-producing interneuron - EPI extension-producing interneuron - LI local interneuron - SFMN slow flexor motoneuron - SEMN slow extensor motoneuron     

Shunting inhibition improves robustness of gamma oscillations in hippocampal interneuron networks by homogenizing firing rates

Networks of GABAergic neurons are key elements in the generation of gamma oscillations in the brain. Computational studies suggested that the emergence of coherent oscillations requires hyperpolarizing inhibition. Here, we show that GABA(A) receptor-mediated inhibition in mature interneurons of the hippocampal dentate gyrus is shunting rather than hyperpolarizing. Unexpectedly, when shunting inhibition is incorporated into a structured interneuron network model with fast and strong synapses, coherent oscillations emerge. In comparison to hyperpolarizing inhibition, networks with shunting inhibition show several advantages. First, oscillations are generated with smaller tonic excitatory drive. Second, network frequencies are tuned to the gamma band. Finally, robustness against heterogeneity in the excitatory drive is markedly improved. In single interneurons, shunting inhibition shortens the interspike interval for low levels of drive but prolongs it for high levels, leading to homogenization of neuronal firing rates. Thus, shunting inhibition may confer increased robustness to gamma oscillations in the brain.     

The adhesion protein IgSF9b is coupled to neuroligin 2 via S-SCAM to promote inhibitory synapse development

Synaptic adhesion molecules regulate diverse aspects of synapse formation and maintenance. Many known synaptic adhesion molecules localize at excitatory synapses, whereas relatively little is known about inhibitory synaptic adhesion molecules. Here we report that IgSF9b is a novel, brain-specific, homophilic adhesion molecule that is strongly expressed in GABAergic interneurons. IgSF9b was preferentially localized at inhibitory synapses in cultured rat hippocampal and cortical interneurons and was required for the development of inhibitory synapses onto interneurons. IgSF9b formed a subsynaptic domain distinct from the GABA_A receptor– and gephyrin-containing domain, as indicated by super-resolution imaging. IgSF9b was linked to neuroligin 2, an inhibitory synaptic adhesion molecule coupled to gephyrin, via the multi-PDZ protein S-SCAM. IgSF9b and neuroligin 2 could reciprocally cluster each other. These results suggest a novel mode of inhibitory synaptic organization in which two subsynaptic domains, one containing IgSF9b for synaptic adhesion and the other containing gephyrin and GABA_A receptors for synaptic transmission, are interconnected through S-SCAM and neuroligin 2.     

Investigating the effects of fluidic connection between microbial fuel cells

Microbial fuel cells (MFCs) can ‘treat’ wastewater but individually are thermodynamically restricted. Scale-up might, therefore, require a plurality of units operating in a stack which could introduce losses simply through fluidic connections. Experiments were performed on two hydraulically joined MFCs (20 cm apart) where feedstock flowed first through the upstream unit (MFC_up) and into the downstream unit (MFC_down) to explore the interactive effect of electrical load connection, influent make-up and flow-rate on electrical outputs. This set-up was also used to investigate how calculating total internal resistance based on a dynamic open circuit voltage (OCV) might differ from using the starting OCV. When fed a highly conductive feedstock (~4,800 μS) MFC_down dropped approximately 180 mV as progressively heavier loads were applied to MFC_up (independent of flow-rate) due to electron leakages through the medium. The conductivities of plain acetate solutions (5 and 20 mM) were insufficient to induce losses in MFC_down even when MFC_up was operating at high current densities. However, at the highest flow-rate (240 mL/h) MFC_down dropped by approximately 100 mV when using 5 and 220 mV using 20 mM acetate. When the distance between MFCs was reduced by 5 cm, voltage drops were apparent even at lower flow-rates, (30 mL/h decreased the voltage by 115 mV when using 20 mM acetate). Shear flow-rates can introduce dissolved oxygen and turbulence all capable of affecting the anodic biofilm and redox conditions. Calculating total internal resistance using a dynamic OCV produced a more stable curve over time compared to that based on the starting constant OCV.     

Target and temporal pattern selection at neocortical synapses

We attempt to summarize the properties of cortical synaptic connections and the precision with which they select their targets in the context of information processing in cortical circuits. High-frequency presynaptic bursts result in rapidly depressing responses at most inputs onto spiny cells and onto some interneurons. These 'phasic' connections detect novelty and changes in the firing rate, but report frequency of maintained activity poorly. By contrast, facilitating inputs to interneurons that target dendrites produce little or no response at low frequencies, but a facilitating-augmenting response to maintained firing. The neurons activated, the cells they in turn target and the properties of those synapses determine which parts of the circuit are recruited and in what temporal pattern. Inhibitory interneurons provide both temporal and spatial tuning. The 'forward' flow from layer-4 excitatory neurons to layer 3 and from 3 to 5 activates predominantly pyramids. 'Back' projections, from 3 to 4 and 5 to 3, do not activate excitatory cells, but target interneurons. Despite, therefore, an increasing complexity in the information integrated as it is processed through these layers, there is little 'contamination' by 'back' projections. That layer 6 acts both as a primary input layer feeding excitation 'forward' to excitatory cells in other layers and as a higher-order layer with more integrated response properties feeding inhibition to layer 4 is discussed.     

Modelling the spatial and temporal constrains of the GABAergic influence on neuronal excitability

GABA (γ-amino butyric acid) is an inhibitory neurotransmitter in the adult brain that can mediate depolarizing responses during development or after neuropathological insults. Under which conditions GABAergic membrane depolarizations are sufficient to impose excitatory effects is hard to predict, as shunting inhibition and GABAergic effects on spatiotemporal filtering of excitatory inputs must be considered. To evaluate at which reversal potential a net excitatory effect was imposed by GABA (E_GABA^Thr), we performed a detailed in-silico study using simple neuronal topologies and distinct spatiotemporal relations between GABAergic and glutamatergic inputs.These simulations revealed for GABAergic synapses located at the soma an E_GABA^Thr close to action potential threshold (E_AP^Thr), while with increasing dendritic distance E_GABA^Thr shifted to positive values. The impact of GABA on AMPA-mediated inputs revealed a complex temporal and spatial dependency. E_GABA^Thr depends on the temporal relation between GABA and AMPA inputs, with a striking negative shift in E_GABA^Thr for AMPA inputs appearing after the GABA input. The spatial dependency between GABA and AMPA inputs revealed a complex profile, with E_GABA^Thr being shifted to values negative to E_AP^Thr for AMPA synapses located proximally to the GABA input, while for distally located AMPA synapses the dendritic distance had only a minor effect on E_GABA^Thr. For tonic GABAergic conductances E_GABA^Thr was negative to E_AP^Thr over a wide range of g_GABA^tonic values. In summary, these results demonstrate that for several physiologically relevant situations E_GABA^Thr is negative to E_AP^Thr, suggesting that depolarizing GABAergic responses can mediate excitatory effects even if E_GABA did not reach E_AP^Thr.     

Synaptic NMDA Receptor-Dependent Ca2+ Entry Drives Membrane Potential and Ca2+ Oscillations in Spinal Ventral Horn Neurons

During vertebrate locomotion, spinal neurons act as oscillators when initiated by glutamate release from descending systems. Activation of NMDA receptors initiates Ca²⁺-mediated intrinsic membrane potential oscillations in central pattern generator (CPG) neurons. NMDA receptor-dependent intrinsic oscillations require Ca²⁺-dependent K⁺ (K_Ca2) channels for burst termination. However, the location of Ca²⁺ entry mediating K_Ca2 channel activation, and type of Ca²⁺ channel – which includes NMDA receptors and voltage-gated Ca²⁺ channels (VGCCs) – remains elusive. NMDA receptor-dependent Ca²⁺ entry necessitates presynaptic release of glutamate, implying a location at active synapses within dendrites, whereas VGCC-dependent Ca²⁺ entry is not similarly constrained. Where Ca²⁺ enters relative to K_Ca2 channels is crucial to information processing of synaptic inputs necessary to coordinate locomotion. We demonstrate that Ca²⁺ permeating NMDA receptors is the dominant source of Ca²⁺ during NMDA-dependent oscillations in lamprey spinal neurons. This Ca²⁺ entry is synaptically located, NMDA receptor-dependent, and sufficient to activate K_Ca2 channels at excitatory interneuron synapses onto other CPG neurons. Selective blockade of VGCCs reduces whole-cell Ca²⁺ entry but leaves membrane potential and Ca²⁺ oscillations unaffected. Furthermore, repetitive oscillations are prevented by fast, but not slow, Ca²⁺ chelation. Taken together, these results demonstrate that K_Ca2 channels are closely located to NMDA receptor-dependent Ca²⁺ entry. The close spatial relationship between NMDA receptors and K_Ca2 channels provides an intrinsic mechanism whereby synaptic excitation both excites and subsequently inhibits ventral horn neurons of the spinal motor system. This places the components necessary for oscillation generation, and hence locomotion, at glutamatergic synapses.     

On the role of ageing and musculoskeletal pain on dynamic balance in manual workers

The purpose of this study was to explore the interacting effects of age and musculoskeletal pain on balance in manual workers. Ninety male manual workers aged 51–72 yr were recruited and stratified according to lower extremity musculoskeletal pain intensity (pain/no pain) and work status (working/retired). The five-repetition sit-to-stand (STS) test was used to assess lower extremity function including completion time, stand time, sit time and dynamic rate of force development both in the upwards (RFD_up) and downwards moving phase (RFD_down). Dynamic balance was expressed as the range, velocity, standard deviation (SD), maximum Lyapunov Exponent and sample entropy of centre of pressure displacement in the anterior-posterior and medial-lateral direction, as well as free moment during the STS test. Except for higher age, no marked differences were seen between working and retired participants. Both age and musculoskeletal pain were negatively associated with motor function. Age × pain interactions showed that completion time, stand time, RFD_up and RFD_down were negatively associated with age for participants without pain, but positively for those with pain. Similar findings were seen for dynamic balance. These findings indicate that the effects of lower extremity musculoskeletal pain on lower extremity function and dynamic balance are age dependent.     

Synaptic information transfer in computer models of neocortical columns

Understanding the direction and quantity of information flowing in neuronal networks is a fundamental problem in neuroscience. Brains and neuronal networks must at the same time store information about the world and react to information in the world. We sought to measure how the activity of the network alters information flow from inputs to output patterns. Using neocortical column neuronal network simulations, we demonstrated that networks with greater internal connectivity reduced input/output correlations from excitatory synapses and decreased negative correlations from inhibitory synapses, measured by Kendall’s τ correlation. Both of these changes were associated with reduction in information flow, measured by normalized transfer entropy (nTE). Information handling by the network reflected the degree of internal connectivity. With no internal connectivity, the feedforward network transformed inputs through nonlinear summation and thresholding. With greater connectivity strength, the recurrent network translated activity and information due to contribution of activity from intrinsic network dynamics. This dynamic contribution amounts to added information drawn from that stored in the network. At still higher internal synaptic strength, the network corrupted the external information, producing a state where little external information came through. The association of increased information retrieved from the network with increased gamma power supports the notion of gamma oscillations playing a role in information processing.     

Caecal parameters of rats fed diets supplemented with inulin in exchange for sucrose

Abstract

The effects of different modes of inulin supplementation on caecal fermentation were evaluated in rats. Groups S and IN were fed diets containing 5% of sucrose or inulin, respectively, for the whole experimental period of 40 days. Group IN/S was fed IN and S diets, whereas group S/IN was fed S and IN diets, in the first and the second 20-day period, respectively. Groups IN_up and IN_down were fed diets in which the content of inulin increased from 1 – 5% and decreased from 5 – 1%, every 8 days, respectively. The common effects of inulin on caecal fermentation, i.e. enlargement of tissue, acidification of digesta, a decrease in activities of potentially harmful bacterial enzymes (β-glucuronidase and β-glucosidase), and an increase in the total volatile fatty acids concentration and pool, were especially observed in the IN, S/IN and IN_up groups. The results suggested that the intensity of caecal fermentation is increased when inulin is present at a relatively high dietary level and that these changes are easily reversible after inulin withdrawal from feed.     

Glutamate as the transmitter at fast and slow neuromuscular junctions of larval diptera

Summary Excitatory postsynaptic currents (EPSCs) produced by stimulating either the fast or slow motor axon to the bodywall muscles of larval sheepfly, are reduced by 60% in the presence of 2 × 10^–4 M L-glutamate. L-aspartate at 2 × 10^–4 M is without effect on either the fast or slow EPSC. Extracellularly recorded slow EPSCs are also reduced by iontophoresis of L-glutamate onto the active site. Iontophoresis of L-aspartate has no effect on the slow EPSC. These results are discussed in relation to a possible transmitter role for glutamate and aspartate at fast and slow excitatory neuromuscular synapses of the bodywall muscles of larval sheepfly.Abbreviation EPSC excitatory postsynaptic current The author is indebted to Professor P.N.R. Usherwood for use of facilities and for helpful discussion during the course of this work. The author is supported by the S.R.C.     

Inhibitory Contribution to Suprathreshold Corticostriatal Responses: An Experimental and Modeling Study

Neostriatal neurons may undergo events of spontaneous synchronization as those observed in recurrent networks of excitatory neurons, even when cortical afferents are transected. It is necessary to explain these events because the neostriatum is a recurrent network of inhibitory neurons. Synchronization of neuronal activity may be caused by plateau-like depolarizations. Plateau-like orthodromic depolarizations that resemble up-states in medium spiny neostriatal neurons (MSNs) may be induced by a single corticostriatal suprathreshold stimulus. Slow synaptic depolarizations may last hundreds of milliseconds, decay slower than the monosynaptic glutamatergic synaptic potentials that induce them, and sustain repetitive firing. Because inhibitory inputs impinging onto MSNs have a reversal potential above the resting membrane potential but below the threshold for firing, they conform a type of “shunting inhibition”. This work asks if shunting GABAergic inputs onto MSNs arrive asynchronously enough as to help in sustaining the plateau-like corticostriatal response after a single cortical stimulus. This may help to begin explaining autonomous processing in the striatal micro-circuitry in the presence of a tonic excitatory drive and independently of spatio-temporally organized inputs. It is shown here that besides synaptic currents from AMPA/KA- and NMDA-receptors, as well as L-type intrinsic Ca²⁺- currents, inhibitory synapses help in maintaining the slow depolarization, although they accomplish the role of depressing firing at the beginning of the response. We then used a NEURON model of spiny cells to show that inhibitory synapses arriving asynchronously on the dendrites can help to simulate a plateau potential similar to that observed experimentally. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Quantitative Organization of GABAergic Synapses in the Molecular Layer of the Mouse Cerebellar Cortex

In the cerebellar cortex, interneurons of the molecular layer (stellate and basket cells) provide GABAergic input to Purkinje cells, as well as to each other and possibly to other interneurons. GABAergic inhibition in the molecular layer has mainly been investigated at the interneuron to Purkinje cell synapse. In this study, we used complementary subtractive strategies to quantitatively assess the ratio of GABAergic synapses on Purkinje cell dendrites versus those on interneurons. We generated a mouse model in which the GABA_A receptor α1 subunit (GABA_ARα1) was selectively removed from Purkinje cells using the Cre/loxP system. Deletion of the α1 subunit resulted in a complete loss of GABA_AR aggregates from Purkinje cells, allowing us to determine the density of GABA_AR clusters in interneurons. In a complementary approach, we determined the density of GABA synapses impinging on Purkinje cells using α-dystroglycan as a specific marker of inhibitory postsynaptic sites. Combining these inverse approaches, we found that synapses received by interneurons represent approximately 40% of all GABAergic synapses in the molecular layer. Notably, this proportion was stable during postnatal development, indicating synchronized synaptogenesis. Based on the pure quantity of GABAergic synapses onto interneurons, we propose that mutual inhibition must play an important, yet largely neglected, computational role in the cerebellar cortex.     

Effects of stimulus timing on transmitter release and postsynaptic membrane potential at crayfish neuromuscular junctions

Summary Different synaptic terminals of the single excitor axon to the opener muscle of crayfish (Procambarus clarkii) often release transmitter in a very different manner when stimulated with the same equal-interval, doublet, or triplet patterns. Compared to synapses that show little facilitation (low F_e synapses), highly facilitating (high F_e) synapses show greater percentage increases in several measures of synaptic efficacy when stimulated with any of these patterns. Low F_e synapses usually show the greater absolute changes in these measures of synaptic efficacy. Changes in the span and pattern of doublets and triplets can independently affect both pre- and postsynaptic measures of synaptic efficacy at either low F_e or high F_e synapses.Abbreviations EJP excitatory junctional potential - MJP spontaneous miniature EJP - F _e ratio of EJP at 1 Hz to EJP amplitude at 10 Hz - F ₁ zero-time facilitation - A ₂,B ₂,C ₂ doubly corrected EJP amplitude of a particular pulse - average amplitude of doubly corrected EJPs in a train of equal-interval, doublets, and triplets, respectively - A_m, B_m, C_m maximum depolarization reached by a particular EJP - time constant of decay     

Calcium requirement of long-term depression and rebound potentiation in cerebellar Purkinje neurons

Cerebellar Purkinje neurons (PNs) receive two main excitatory inputs, from climbing fibers and parallel fibers, and inhibitory inputs, from GABAergic interneurons. The synapses formed by parallel fibers and by inhibitory interneurons on PNs are able to undergo long-lasting in efficacy. Thus, the excitatory parallel fiber-PN synapse undergoes long-term fibers. Synaptic inhibition can be potentiated by climbing fiber activity by a mechanism named rebound potentiation, resulting in a more powerful inhibitory effect of GABAergic interneurons. The induction of both long-term depression and rebound potentiation requires a transient elevation of the cytoplasmic calcium concentration ([Ca²⁺]_i). The [Ca²⁺]_i-transient is caused by Ca²⁺ entry through voltage-gated Ca²⁺ channels and, possibly, by release of Ca²⁺ from IP_3- and ryanodine-sensitive stores. Direct Ca²⁺ entry through synaptic AMPA receptor channels seems not to contribute significantly to the Ca²⁺ signal mediating the induction of both long-term depression and rebound potentiation.     

Information processing in the femur-tibia control loop of stick insects

The complicated response characteristics of the identified nonspiking interneuron type E4 upon elongation stimuli to the femoral chordotonal organ (fCO) can be obtained by a computer simulation using the neuronal network simulator BioSim, if the following assumptions were introduced: (1) The interneurons receive direct excitatory input from position- and velocity-sensitive fCO afferents but also, in parallel delayed inhibition from the same velocity-sensitive afferents. (2) Position-sensitive afferents in part show adaptation with a rather long time-constant. A subsequent experimental analysis demonstrated that all these assumptions fit the reality: (1) Interneurons of type E4 receive direct excitatory input from fCO afferents. (2) Interneurons of type E4 are affected by velocity dependent delayed inhibitory inputs from the fCO. (3) The fCO does contain adapting position-sensitive sensory neurons, which have not been described before. The described principle of the information processing is also able to generate the response in interneurons of type E6 with less steep amplitude-velocity characteristic due to a different weighting of the direct excitation and delayed inhibition.Abbreviations EPSP excitatory postsynaptic potential - FETi fast extensor tibiae motor neuron - fCO femoral chordotonal organ - FT-control loop femur-tibia control loop - IPSP inhibitory postsynaptic potential - SETi slow extensor tibiae motor neuron