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While temperature fluctuations pose significant challenges to the nervous system, many vital neuronal systems in poikilothermic animals function over a broad temperature range. Using the gastric mill pattern generator in the Jonah crab, we previously demonstrated that temperature-induced increases in leak conductance disrupt neuronal function and that neuropeptide modulation provides thermal protection. Here, we show that neuropeptide modulation also increases temperature robustness in Dungeness and green crabs. As in Jonah crabs, higher temperatures increased leak conductance in both species' pattern-generating lateral gastric neuron and terminated rhythmic gastric mill activity. Likewise, increasing descending modulatory projection neuron activity or neuropeptide transmitter application rescued rhythms at elevated temperatures. However, decreasing input resistance using dynamic clamp only restored the rhythm in half of the experiments. Thus, neuropeptide modulation increased temperature robustness in both species, demonstrating that neuropeptide-mediated temperature compensation is not limited to one species, although the underlying cellular compensation mechanisms may be distinct., Competing Interests: Competing interests The authors declare no competing or financial interests., (© 2024. Published by The Company of Biologists Ltd.)

Large insects actively ventilate their tracheal system even at rest, using abdominal pumping movements, which are controlled by a central pattern generator (CPG) in the thoracic ganglia. We studied the effects of respiratory gases on the ventilatory rhythm by isolating the thoracic ganglia and perfusing its main tracheae with various respiratory gas mixtures. Fictive ventilation activity was recorded from motor nerves controlling spiracular and abdominal ventilatory muscles. Both hypoxia and hypercapnia increased the ventilation rate, with the latter being much more potent. Sub-threshold hypoxic and hypercapnic levels were still able to modulate the rhythm as a result of interactions between the effects of the two respiratory gases. Additionally, changing the oxygen levels in the bathing saline affected ventilation rate, suggesting a modulatory role for haemolymph oxygen. Central sensing of both respiratory gases as well as interactions of their effects on the motor output of the ventilatory CPG reported here indicate convergent evolution of respiratory control among terrestrial animals of distant taxa. [ABSTRACT FROM AUTHOR]

The heart and pyloric rhythms of crustaceans have been studied separately and extensively over many years. Local and hormonal neuromodulation and sensory inputs into these central pattern generator circuits play a significant role in an animal's response to perturbations, but are usually lost or removed during in vitro studies. To examine simultaneously the in vivo motor output of the crustacean heart and pyloric rhythms, we used photoplethysmography. In the population measured (n=49), the heart rhythm frequency ranged from 0.3 to 2.3 Hz. The pyloric rhythm varied from 0.2 to 1.6 Hz. We observed aweak correlation between the frequencies of the heart and pyloric rhythms. During multiple hour-long recordings, many animals held at a controlled temperature showed strong inhibitory bouts in which the heart decreased in frequency or become quiescent and the pyloric rhythm decreased in frequency. We measured the simultaneous responses of the rhythms to temperature ramps by heating or cooling the saline bath while recording both the heart and pyloric muscle movements. Q10, critical temperature (temperature at which muscle function is compromised) and changes in frequency were calculated for each of the rhythms tested. The heart rhythm was more robust to high temperature than the pyloric rhythm. [ABSTRACT FROM AUTHOR]

Blood feeding is an essential and signature activity of the medicinal leech species Hirudo verbana. Despite keen interest in understanding the neuronal substrates of this behavior, a major component of the nervous system associated with feeding has remained overlooked. In this study, for the first time, we report on the presence and characteristics of five stomatogastric ganglia (STGs) comprising the visceral stomatogastric nervous system (STN) of the leech. Although a brief report was published by Ruth Hanke in 1948 indicating that a ring of three ganglia (not five) was associated with the cephalic ganglia, this information was never integrated into subsequent neurobiological studies of feeding. Here, the anatomical features of the STGs are described, as are the morphological and electrophysiological characteristics of neurons originating in them. We also determined that two of the five STGs (STG-1 and STG-3) each contained two relatively large (ca. 40 μm diameter) serotonergic neurons. The STN was also enriched with dopaminergic and serotonergic arborizations; however, no intrinsic dopaminergic somata were observed. The trajectory of the serotonergic large lateral (LL) neuron, a command-like cell for feeding, was documented to project directly to the STN and not to the jaw and pharyngeal musculature as previously reported, thus reopening the important question of how the LL cell activates and coordinates biting activity with pharyngeal swallowing. Additional studies revealed that the LL cell is excited by blood serum applied to the lip and is strongly inhibited by dopamine. These findings provide a new foundation for understanding the regulation and modulation of neural networks involved in feeding. [ABSTRACT FROM AUTHOR]

Discontinuous gas exchange (DGE) is the best studied among insect gas exchange patterns. DGE cycles comprise three phases, which are defined by their spiracular state: closed, flutter and open. However, spiracle status has rarely been monitored directly; rather, it is often assumed based on CO2 emission traces. In this study, we directly recorded electromyogram (EMG) signals from the closer muscle of the second thoracic spiracle and from abdominal ventilation muscles in a fully intact locust during DGE. Muscular activity was monitored simultaneously with CO2 emission, under normoxia and under various experimental oxic conditions. Our findings indicate that locust DGE does not correspond well with the commonly described three-phase cycle.We describe unique DGE-related ventilation motor patterns, coupled to spiracular activity. During the open phase, when CO2 emission rate is highest, the thoracic spiracles do not remain open; rather, they open and close rapidly. This fast spiracle activity coincides with in-phase abdominal ventilation, while alternating with the abdominal spiracle and thus facilitating a unidirectional air flow along the main trachea. A change in the frequency of rhythmic ventilation during the open phase suggests modulation by intratracheal CO2 levels. A second, slow ventilatory movement pattern probably serves to facilitate gas diffusion during spiracle closure. Two flutter-like patterns are described in association with the different types of ventilatory activity.We offer a modified mechanistic model for DGE in actively ventilating insects, incorporating ventilatory behavior and changes in spiracle state. [ABSTRACT FROM AUTHOR]

Adaptive changes in the output of neural circuits underlying rhythmic behaviors are relayed to muscles via motor neuron activity. Presynaptic and postsynaptic properties of neuromuscular junctions can impact the transformation from motor neuron activity to muscle response. Further, synaptic plasticity occurring on the time scale of inter-spike intervals can differ between multiple muscles innervated by the same motor neuron. In rhythmic behaviors, motor neuron bursts can elicit additional synaptic plasticity. However, it is unknown whether plasticity regulated by the longer time scale of inter-burst intervals also differs between synapses from the same neuron, and whether any such distinctions occur across a physiological activity range. To address these issues, we measured electrical responses in muscles innervated by a chewing circuit neuron, the lateral gastric (LG) motor neuron, in a well-characterized small motor system, the stomatogastric nervous system (STNS) of the Jonah crab, Cancer borealis. In vitro and in vivo, sensory, hormonal and modulatory inputs elicit LG bursting consisting of inter-spike intervals of 50–250 ms and inter-burst intervals of 2–24 s. Muscles expressed similar facilitation measured with paired stimuli except at the shortest inter-spike interval. However, distinct decay time constants resulted in differences in temporal summation. In response to bursting activity, augmentation occurred to different extents and saturated at different inter-burst intervals. Further, augmentation interacted with facilitation, resulting in distinct intra-burst facilitation between muscles. Thus, responses of multiple target muscles diverge across a physiological activity range as a result of distinct synaptic properties sensitive to multiple time scales. [ABSTRACT FROM AUTHOR]

The neural circuits underlying divergent courtship behaviors of closely related species provide a framework for insight into the evolution of motor patterns. In frogs, male advertisement calls serve as unique species identifiers and females prefer conspecific to heterospecific calls. Advertisement calls of three relatively recently (~8.5 Mya) diverged species - Xenopus laevis, X. petersii and X. victorianus - include rapid trains of sound pulses (fast trills).We show that while fast trills are similar in pulse rate (~60 pulses s-1) across the three species, they differ in call duration and period (time from the onset of one call to the onset of the following call). Previous studies of call production in X. laevis used an isolated brain preparation in which the laryngeal nerve produces compound action potentials that correspond to the advertisement call pattern (fictive calling). Here, we show that serotonin evokes fictive calling in X. petersii and X. victorianus as it does in X. laevis. As in X. laevis, fictive fast trill in X. petersii and X. victorianus is accompanied by an N-methyl-D-aspartate receptor-dependent local field potential wave in a rostral hindbrain nucleus, DTAM. Across the three species, wave duration and period are strongly correlated with species-specific fast trill duration and period, respectively. When DTAM is isolated from the more rostral forebrain and midbrain and/or more caudal laryngeal motor nucleus, the wave persists at species-typical durations and periods. Thus, intrinsic differences within DTAM could be responsible for the evolutionary divergence of call patterns across these related species. [ABSTRACT FROM AUTHOR]

During the post-embryonic developmental growth of animals, a number of physiological parameters such as locomotor performance, dynamics and behavioural repertoire are adjusted to match the requirements determined by changes in body size, proportions and shape. Moreover, changes inmovement parameters also cause changes in the dynamics of self-generated sensory stimuli, to which motion-detecting sensory systems have to adapt. Here, we examined head movements and swimming kinematics of Xenopus laevis tadpoles with a body length of 10-45 mm (developmental stage 46-54) and compared these parameters with fictive swimming, recorded as ventral root activity in semi-intact in vitro preparations. Head movement kinematics was extracted from high-speed video recordings of freely swimming tadpoles. Analysis of these locomotor episodes indicated that the swimming frequency decreased with development, along with the angular velocity and acceleration of the head, which represent selfgenerated vestibular stimuli. In contrast, neither head oscillation amplitude nor forward velocity changed with development despite the ~3-fold increase in body size. The comparison between free and fictive locomotor dynamics revealed very similar swimming frequencies for similarly sized animals, including a comparable developmental decrease of the swimming frequency. Body morphology and the motor output rhythm of the spinal central pattern generator therefore develop concurrently. This study thus describes development-specific naturalistic head motion profiles, which form the basis for more natural stimuli in future studies probing the vestibular system. [ABSTRACT FROM AUTHOR]

The parasitic wasp Cotesia congregata suppresses feeding in its host, the caterpillar Manduca sexta, during specific periods of wasp development. We examined both feeding behaviour and the neurophysiology of the mandibular closer muscle in parasitized and unparasitized control M. sexta to determine how the wasp may accomplish this. To test whether the wasps activated a pre-existing host mechanism for feeding cessation, we examined the microstructure of feeding behaviour in caterpillars that stopped feeding due to illness-induced anorexia or an impending moult. These microstructures were compared with that shown by parasitized caterpillars. While there were overall differences between parasitized and unparasitized caterpillars, the groups showed similar progression in feeding microstructure as feeding ended, suggesting a common pattern for terminating a meal. Parasitized caterpillars also consumed less leaf area in 100 bites than control caterpillars at around the same time their feeding microstructure changed. The decline in food consumption was accompanied by fewer spikes per burst and shorter burst durations in chewing muscle electromyograms. Similar extracellular results were obtained from the motorneuron of the mandibular closer muscle. However, chewing was dramatically re-activated in non-feeding parasitized caterpillars if the connectives posterior to the suboesophageal ganglion were severed. The same result was observed in unparasitized caterpillars given the same treatment. Our results suggest that the reduced feeding in parasitized caterpillars is not due to damage to the central pattern generator (CPG) for chewing, motor nerves or chewing muscles, but is more likely to be due to a suppression of chewing CPG activity by ascending or descending inputs., Competing Interests: Competing interests The authors declare no competing or financial interests., (© 2023. Published by The Company of Biologists Ltd.)

During swimming in the amphibian Xenopus laevis, efference copies of rhythmic locomotor commands produced by the spinal central pattern generator (CPG) can drive extraocular motor output appropriate for producing image-stabilizing eye movements to offset the disruptive effects of self-motion. During metamorphosis, X. laevis remodels its locomotor strategy from larval tail-based undulatory movements to bilaterally synchronous hindlimb kicking in the adult. This change in propulsive mode results in head/body motion with entirely different dynamics, necessitating a concomitant switch in compensatory ocular movements from conjugate left-right rotations to non-conjugate convergence during the linear forward acceleration produced during each kick cycle. Here, using semi-intact or isolated brainstem/spinal cord preparations at intermediate metamorphic stages, we monitored bilateral eye motion along with extraocular, spinal axial and limb motor nerve activity during episodes of spontaneous fictive swimming. Our results show a progressive transition in spinal efference copy control of extraocular motor output that remains adapted to offsetting visual disturbances during the combinatorial expression of bimodal propulsion when functional larval and adult locomotor systems co-exist within the same animal. In stages at metamorphic climax, spino-extraocular motor coupling, which previously derived from axial locomotor circuitry alone, can originate from both axial and de novo hindlimb CPGs, although the latter's influence becomes progressively more dominant and eventually exclusive as metamorphosis terminates with tail resorption. Thus, adaptive interactions between locomotor and extraocular motor circuitry allows CPG-driven efference copy signaling to continuously match the changing spatio-temporal requirements for visual image stabilization throughout the transitional period when one propulsive mechanism emerges and replaces another. [ABSTRACT FROM AUTHOR]

The mechanisms of rhythmic motor pattern generation have been studied in detail in vitro, but the long-term stability and sources of variability in vivo are often not well described. The crab stomatogastric ganglion contains the well-characterized gastric mill (chewing) and pyloric (filtering of food) central pattern generators. In vitro, the pyloric rhythm is stereotyped with little variation, but intercircuit interactions and neuromodulation can alter both rhythm cycle frequency and structure. The range of variation of activity in vivo is, with few exceptions, unknown. Curiously, although the patterngenerating circuits in vivo are constantly exposed to hormonal and neural modulation, the majority of published data show only the unperturbed canonical motor patterns typically observed in vitro. Using long-term extracellular recordings (N=27 animals), we identified the range and sources of variability of the pyloric and gastric mill rhythms recorded continuously over 4 days in freely behaving Jonah crabs (Cancer borealis). Although there was no evidence of innate daily rhythmicity, a 12 h light-driven cycle did manifest. The frequency of both rhythms increased modestly, albeit consistently, during the 3 h before and 3 h after the lights changed. This cycle was occluded by sensory stimulation (feeding), which significantly influenced both pyloric cycle frequency and structure. This was the only instance where the structure of the rhythm changed. In unfed animals the structure remained stable, even when the frequency varied substantially. So, although central pattern generating circuits are capable of producing many patterns, in vivo outputs typically remain stable in the absence of sensory stimulation. [ABSTRACT FROM AUTHOR]

Many neuropeptides are members of peptide families, with multiple structurally similar isoforms frequently found even within a single species. This raises the question of whether the individual peptides serve common or distinct functions. In the accompanying paper, we found high isoform specificity in the responses of the lobster (Homarus americanus) cardiac neuromuscular system to members of the pyrokinin peptide family: only one of five crustacean isoforms showed any bioactivity in the cardiac system. Because previous studies in other species had found little isoform specificity in pyrokinin actions, we examined the effects of the same five crustacean pyrokinins on the lobster stomatogastric nervous system (STNS). In contrast to our findings in the cardiac system, the effects of the five pyrokinin isoforms on the STNS were indistinguishable: they all activated or enhanced the gastric mill motor pattern, but did not alter the pyloric pattern. These results, in combination with those from the cardiac ganglion, suggest that members of a peptide family in the same species can be both isoform specific and highly promiscuous in their modulatory capacity. The mechanisms that underlie these differences in specificity have not yet been elucidated; one possible explanation, which has yet to be tested, is the presence and differential distribution of multiple receptors for members of this peptide family. [ABSTRACT FROM AUTHOR]

Although the crustacean heart is modulated by a large number of peptides and amines, few of these molecules have been localized to the cardiac ganglion itself; most appear to reach the cardiac ganglion only by hormonal routes. Immunohistochemistry in the American lobster Homarus americanus indicates that pyrokinins are present not only in neuroendocrine organs ( pericardial organ and sinus gland), but also in the cardiac ganglion itself, where pyrokinin-positive terminals were found in the pacemaker cell region, as well as surrounding the motor neurons. Surprisingly, the single pyrokinin peptide identified from H. americanus, FSPRLamide, which consists solely of the conserved FXPRLamide residues that characterize pyrokinins, did not alter the activity of the cardiac neuromuscular system. However, a pyrokinin from the shrimp Litopenaeus vannamei [ADFAFNPRLamide, also known as Penaeus vannamei pyrokinin 2 (PevPK2)] increased both the frequency and amplitude of heart contractions when perfused through the isolated whole heart. None of the other crustacean pyrokinins tested (another from L. vannamei and two from the crab Cancer borealis) had any effect on the lobster heart. Similarly, altering the PevPK2 sequence either by truncation or by the substitution of single amino acids resulted in much lower or no activity in all cases; only the conservative substitution of serine for alanine at position 1 resulted in any activity on the heart. Thus, in contrast to other systems (cockroach and crab) in which all tested pyrokinins elicit similar bioactivities, activation of the pyrokinin receptor in the lobster heart appears to be highly isoform specific. [ABSTRACT FROM AUTHOR]

Cockroaches are remarkably stable runners, exhibiting rapid recovery from external perturbations. To uncover the mechanisms behind this important behavioral trait, we recorded leg kinematics of freely running animals in both undisturbed and perturbed trials. Functional coupling underlying inter-leg coordination was monitored before and during localized perturbations, which were applied to single legs via magnetic impulses. The resulting transient effects on all legs and the recovery times to normal pre-perturbation kinematics were studied. We estimated coupling architecture and strength by fitting experimental data to a six-leg-unit phase oscillator model. Using maximum-likelihood techniques, we found that a network with nearest-neighbor inter-leg coupling best fitted the data and that, although coupling strengths vary among preparations, the overall inputs entering each leg are approximately balanced and consistent. Simulations of models with different coupling strengths encountering perturbations suggest that the coupling schemes estimated from our experiments allow animals relatively fast and uniform recoveries from perturbations. [ABSTRACT FROM AUTHOR]

The modulatory effects of nitric oxide (NO) and cAMP on the rhythmic beating activity of the swimmeret motor neurones in the crayfish were examined. Swimmerets are paired appendages located on the ventral side of each abdominal segment that show rhythmic beating activity during forward swimming, postural righting behaviour and egg ventilation in gravid females. In isolated abdominal nerve cord preparations, swimmeret motor neurones are usually silent or show a continuous low-frequency spiking activity. Application of carbachol, a cholinergic agonist, elicited rhythmic bursts of motor neurone spikes. The co-application of L-arginine, the substrate for NO synthesis with carbachol increased the burst frequency of the motor neurones. The co-application of the NO donor SNAP with carbachol also increased the burst frequency of the motor neurones. By contrast, co-application of a NOS inhibitor, L-NAME, with carbachol decreased beating frequency of the motor neurones. These results indicate that NO may act as a neuromodulator to facilitate swimmeret beating activity. The facilitatory effect of L-arginine was cancelled by co-application of the soluble guanylate cyclase (sGC) inhibitor ODQ suggesting that NO acts by activating sGC to promote the production of cGMP Application of L-arginine alone or membrane-permeable cGMP analogue 8-Br-cGMP alone did not elicit rhythmic activity of motor neurones, but co-application of 8-Br-cGMP with carbachol increased bursting frequency of the motor neurones. Furthermore, application of the membrane-permeable cAMP analogue CPT-cAMP alone produced rhythmic bursting of swimmeret motor neurones, and the bursting frequency elicited by CPT-cAMP was increased by coapplication with L-arginine. Co-application of the adenylate cyclase inhibitor SQ22536 ceased rhythmic bursts of motor neurone spikes elicited by carbachol. These results suggest that a cAMP system enables the rhythmic bursts of motor neurone spikes and that a NO-cGMP signaling pathway increases cAMP activity to facilitate swimmeret beating. [ABSTRACT FROM AUTHOR]

The regulation of insect gut physiology is complex and involves the interactions of a number of mechanisms, including the neural regulation of gut contraction by altering neural input and the modulation of gut contractions by neuropeptides directly affecting the muscle. The FGLa-type allatostatins (FGLa/ASTs) are known brain/gut peptides with numerous physiological roles, including modulation of gut contraction and neural input. To further investigate the pleiotropic roles of FGLa/AST peptides in Locusta migratoria, we have examined the role of a locust FGLa/AST (Scg-AST-6) In the gut. Proctolin and Scg-AST-6 have opposing effects on gut contraction, where proctolin dose-dependently increases gut muscle tension, while Scg-AST-6 inhibits both muscle tension and spontaneous and neurogenic contractions in a dose-dependent manner. Results from neurophysiological recordings indicate that there may be a central pattern generator (CPG) within the ventricular ganglia regulated by descending inhibition, and the addition of Scg-AST-6 dose-dependently modulates this ventricular ganglion CPG. This work provides a comprehensive picture of how FGLa/ASTs may modulate and coordinate each region of the locust gut, and shows that FGLa/ASTs have both central effects, on the ventricular ganglion CPG, and peripheral effects on the gut muscle. Overall, this study shows how FGLa/ASTs contribute to the complex regulation and fine tuning of gut contraction. [ABSTRACT FROM AUTHOR]

Dopamine (DA) activates fictive crawling behavior in the medicinal leech. To identify the cellular mechanisms underlying this activation at the level of crawl-specific motoneuronal bursting, we targeted potential cAMP-dependent events that are often activated through DAHike receptor signaling pathways. We found that isolated ganglia produced crawl-like motoneuron bursting after bath application of phosphodiesterase inhibitors (PDIs) that upregulated cAMP. This bursting persisted in salines in which calcium ions were replaced with equimolar cobalt or nickel, but was blocked by riluzole, an inhibitor of a persistent sodium current. PDI-induced bursting contained a number of patterned elements that were statistically similar to those observed during DA-induced fictive crawling, except that one motoneuron (CV) exhibited bursting during the contraction rather than the elongation phase of crawling. Although DA and the PDIs produced similar bursting profiles, intracellular recordings from motoneurons revealed differences in altered membrane properties. For example, DA lowered motoneuron excitability whereas the PDIs increased resting discharge rates. We suggest that PDIs (and DA) activate a sodium-influx-dependent timing mechanism capable of setting the crawl rhythm and that multiple DA receptor subtypes are involved in shaping and modulating the phase relationships and membrane properties of cell-specific members of the crawl network to generate crawling. [ABSTRACT FROM AUTHOR]

The relative importance of sensory Input for the production of centrally generated motor patterns is crucial to our understanding of how animals coordinate their body segments to locomote in legged locomotion, where terrain heterogeneity may require stride-by-strlde changes In leg placement, evidence suggests that sensory information Is essential for the timing of leg movement. In a previous study we showed that In cockroaches, renowned for rapid and stable running, a coordinated pattern can be elicited from the motor centres driving the different legs in the absence of sensory feedback. In the present paper, we assess the role of movement-related sensory Inputs in modifying this central pattern. We studied the effect of spontaneous steps as well as imposed transient and periodic movements of a single Intact leg, and demonstrate that, depending on the movement properties, the resulting proprioceptive feedback can significantly modify phase relationships among segmental oscillators of other legs. Our analysis suggests that feedback from front legs Is weaker but more phaslcally precise than from hind legs, selectively transferring movement-related Information In a manner that strengthens the Inherent rhythmic pattern and modulates local perturbations. [ABSTRACT FROM AUTHOR]

Autotomy (voluntary loss of an appendage) is common among diverse groups of vertebrates and invertebrates, and much attention has been given to ecological and developmental aspects of tail autotomy in lizards. Although most studies have focused on the ramifications for the lizard (behavior, biomechanics, energetics, etc.), the tail itself can exhibit interesting behaviors once segregated from the body. For example, recent work highlighted the ability of leopard gecko tails to jump and flip, in addition to being able to swing back and forth. Little is known, however, about the control mechanisms underlying these movements. Using electromyography, we examined the time-varying in vivo motor patterns at four sites (two proximal and two distal) In the tall of the leopard gecko, Eublepharis macularius, following autotomy. Using these data we tested the hypothesis that the disparity In movements results simply from overlapping pattern generators within the tail. We found that burst duration, but not cycle duration, of the rhythmic swings reached a plateau at approximately 150 s following autotomy. This is likely because of physiological changes related to muscle fatigue and ischemia. For flips and jumps, burst and cycle duration exhibited no regular pattern. The coefficient of variation in motor patterns was significantly greater for jumps and flips than for rhythmic swings. This supports the conclusion that the different tail behaviors do not stem from overlapping pattern generators, but that they rely upon Independent neural circuits. The signal controlling jumps and flips may be modified by sensory information from the environment. Finally, we found that jumps and flips are initiated using relatively synchronous activity between the two sides of the tall. In contrast, alternating activation of the right and left sides of the tail result in rhythmic swings. The mechanism underlying this change in tail behavior is comparable to locomotor gait changes in vertebrates. [ABSTRACT FROM AUTHOR]

The mammalian masticatory rhythm is produced by a brainstem timing network. The rhythm is relatively fixed within individual animals but scales allometrically with body mass (Mb) across species. It has been hypothesized that sensory feedback and feed- forward adjust the rhythm to match the jaw's natural resonance frequency, with allometric scaling being an observable consequence. However, studies performed with adult animals show that the rhythm is not affected by jaw mass manipulations, indicating that either developmental or evolutionary mechanisms are required for allometry to become manifest. The present study was performed to tease out the relative effects of development versus natural selection on chewing rate allometry. Thirty- one dog breeds and 31 mass-matched non-domestic mammalian species with a range in Mb from ∼2kg to 50kg were studied. Results demonstrated that the chewing rhythm did not scale with Mb among dog breeds (R=0.299, P>0.10) or with jaw length (Lj) (R=0.328, P>0.05). However, there was a significant relationship between the chewing rhythm and Mb among the non-domestic mammals (R=0.634, P<0.001). These results indicate that scaling is not necessary in the adult animal. We conclude that the central timing network and related sensorimotor systems may be necessary for rhythm generation but they do not explain the 1/3rd to 1/4th allometric scaling observed among adult mammals. The rhythm of the timing network is either adjusted to the physical parameters of the jaw system during early development only, is genetically determined independently of the jaw system or is uniquely hard-wired among dogs and laboratory rodents. [ABSTRACT FROM AUTHOR]

Octopamine-like immunoreactivity was localized to a limited number (<40) of neurons in the Aplysia central nervous system, including three neurons in the paired buccal ganglia (BG) that control feeding movements. Application of octopamine (OA) to the BG circuit produced concentration-dependent (10-8-10-4 mol I-1) modulatory actions on the spontaneous burst activity of the bilaterally paired B67 pharyngeal motor neurons (MNs). OA increased B67's burst duration and the number of impulses per burst. These effects reflected actions of OA on the intrinsic tetrodotoxin-resistant driver potential (DP) that underlies B67 bursting. In addition to its effects on B67's burst parameters, OA also increased the rate and regularity of burst timing. Although the bilaterally paired B67 MNs both exhibited rhythmic bursting in the presence of OA, they did not become synchronized. In this respect, the response to OA differed from that of dopamine, another modulator of the feeding motor network, which produces both rhythmicity and synchrony of bursting in the paired B67 neurons. It is proposed that modulators can regulate burst synchrony of MNs by exerting a dual control over their intrinsic rhythmicity and their reciprocal capacity to generate membrane potential perturbations. In this simple system, dopaminergic and octopaminergic modulation could influence whether pharyngeal contractions occur in a bilaterally synchronous or asynchronous fashion. [ABSTRACT FROM AUTHOR]

The aerial respiratory behaviour of the mollusc Lymnaea stagnalis is an important homeostatic behaviour that can be operantly conditioned. The central pattern generator underlying this behaviour, as well as motorneurons innervating the respiratory orifice, the pneumostome, have been identified and their activity can be monitored in the semi-intact preparation using electrophysiological recordings. In this study, we used both intact animals and semi-intact preparations to identify novel changes in the respiratory central pattern generator following operant conditioning. In addition, we reared animals in the absence of this respiratory behaviour throughout development, to investigate whether previous experience and activity-dependent plasticity during development are essential to allow neural plasticity in the adult. We found that animals raised normally (allowed to perform aerial respiratory behaviour) exhibited the expected reduction in aerial respiratory behaviour following operant conditioning. Then, using the semi-intact preparation, we identified novel neural changes within the network as a result of the conditioning. These included specific changes at the level of the central pattern generator interneurons, as well as the motor output. In the differentially reared intact animals, there was no behavioural reduction as a result of operant conditioning, although their baseline respiratory behaviour was already significantly reduced as a result of their differential rearing. There were, however, significant differences found in the network parameters in the semi-intact preparation, similar to those observed in normally reared animals. We thus provide evidence for neural plasticity within the network in the absence of significant behavioural changes in differentially reared animals, and show that plasticity was not dependent on previous activity of the network during development. [ABSTRACT FROM AUTHOR]

Neuronal network flexibility enables animals to respond appropriately to changes in their internal and external states. We are using the isolated crab stomatogastric nervous system to determine how extrinsic inputs contribute to network flexibility. The stomatogastric system includes the well-characterized gastric mill (chewing) and pyloric (filtering of chewed food) motor circuits in the stomatogastric ganglion. Projection neurons with somata in the commissural ganglia (CoGs) regulate these rhythms. Previous work characterized a unique gastric mill rhythm that occurred spontaneously in some preparations, but whose origin remained undetermined. This rhythm includes a distinct protractor phase activity pattern, during which a key gastric mill circuit neuron (LG neuron) and the projection neurons MCN1 and CPN2 fire in a pyloric rhythm-timed activity pattern instead of the tonic firing pattern exhibited by these neurons during previously studied gastric mill rhythms. Here we identify a new extrinsic input, the post-oesophageal commissure (POC) neurons, relatively brief stimulation (30 s) of which triggers a long-lasting (tens of minutes) activation of this novel gastric mill rhythm at least in part via its lasting activation of MCN1 and CPN2. Immunocytochemical and electrophysiological data suggest that the POC neurons excite MCN1 and CPN2 by release of the neuropeptide Cancer borealis tachykinin-related peptide Ia (CabTRP Ia). These data further suggest that the CoG arborization of the POC neurons comprises the previously identified anterior commissural organ (ACO), a CabTRP la-containing neurohemal organ. This endocrine organ thus appears to also have paracrine actions, including activation of a novel and lasting gastric mill rhythm. [ABSTRACT FROM AUTHOR]

In locusts, a central pattern generator underlies the rhythmic movements of the ovipositor valves that serve to drive the abdomen into damp soil in order to lay eggs. We have investigated the role of nitric oxide (NO) in the control of this oviposition digging rhythm. NO increases the frequency of the rhythm by acting via sGC to elevate cGMP, which in turn acts via PKG. Increasing exogenous NO levels using the NO donors SNAP and PAPANONOate increased the cycle frequency of the fictive digging rhythm, as did increasing endogenous NO by bath application of the substrate for NOS, L-arginine. On the other hand, application of the NO scavenger PTIO decreased the cycle frequency, indicating that NO must normally exert a continuous and dynamic role on the central pattern generator underlying the oviposition rhythm. Inhibiting the main molecular target of NO, soluble guanylate cyclase, with ODQ reduced the cycle frequency of the rhythm, suggesting that NO mediated its effects via sGC and cyclic GMP. Further evidence for this was produced by bath application of 8-Br-cGMP, which increased the frequency of the rhythm. Bath application of the generic protein kinase inhibitor and a selective PKG inhibitor, H-7 and KT-5823, respectively, reduced the frequency of the rhythm, suggesting that PKG acted as a target for cGMP. Thus, we conclude that NO plays a key role in regulating the frequency of the central pattern generator controlling rhythmic egg-laying movements in locusts by acting via sGC/cGMP-PKG. [ABSTRACT FROM AUTHOR]

The mechanisms by which different life histories affect neural circuits are largely unknown. We show that the thermosensitivity and thermotolerance of neural circuit operation are affected in a complex dynamic fashion by photoperiod, prior heat experience and the sex of the animal. We compared thermosensitivity and thermotolerance of ventilatory motor pattern generation in locusts reared under two photoperiods (12:12 and 16:8; i.e. 12 h:12 h and 16 h:8 h L:D, respectively) before and after heat shock pre-treatment (HS: 3 h, 45°C) in order to determine the effect of daylength on properties of neural function. We monitored central pattern generator (CPG) output electromyographically from muscle 161 in the second abdominal segment during ramped increases in temperature and also measured the time taken for the circuit to fail at high temperatures and the time taken to recover on return to room temperature. There were effects of photoperiod, heat pre-treatment and: the sex of the animal on ventilatory rate, time-to-failure and time-to-recovery. The ventilatory motor pattern of 16:8 and 12:12 locusts responded differently to increasing and maintained high temperature stress in both control and heat shocked locusts. We found that 12:12 locusts were generally more robust than 16:8 locusts: they lived longer, they showed greater tolerance to high temperatures, and they recovered more quickly from temperature-induced circuit failure. A faster ventilatory rate in 12:12 animals at high temperatures may have accelerated evaporative cooling to mediate improved temperature tolerance. [ABSTRACT FROM AUTHOR]

To fully understand neuronal network operation, the influence of all inputs onto that network must be characterized. As in most systems, many neuronal and hormonal pathways influence the multifunctional motor circuits of the crustacean stomatogastric ganglion (STG), but the actions of only some of them are known. Therefore, we characterized the influence of the kinin peptide family on the gastric mill (chewing) and pyloric (filtering of chewed food) motor circuits in the STG of the crab Cancer borealis. The kinins are myoactive in arthropods and they occur within the arthropod central nervous system (CNS), but their CNS actions are not well characterized in any species. The pevkinins were first identified in the shrimp Penaeus vannamei, but they have yet to be studied in the STG of any species. We identified kinin-like immunolabeling (KLI) in the pericardial organs (POs) in C. borealis, but there was no KLI within the STG. The POs are a major source of hormonal influence on the STG. Pevkinin peptides activated the pyloric circuit and they caused a modest increase in the speed of ongoing pyloric rhythms. This modest influence on cycle speed resulted in part from pevkinin excitation of the lateral pyloric neuron, whose strengthened inhibitory synapse onto the pyloric pacemaker neurons limited the pevkinin-mediated increase in cycle speed. The pevkinin excitation of the pyloric rhythm was not strong enough to interfere with the previously documented, gastric mill rhythm-mediated weakening of the pyloric rhythm. Pevkinin also had little influence on the gastric mill rhythm. These results indicate that the kinin peptides have distinct and selective modulatory actions on the pyloric rhythm. [ABSTRACT FROM AUTHOR]

The aim of this study was to investigate the neural basis of operant conditioning in a semi-intact preparation of the pond snail, Lymnaea stagnalis. Lymnaea learns, via operant conditioning, to reduce its aerial respiratory behaviour in response to an aversive tactile stimulus to its open pneumostome. Here we report the successful conditioning of naive semi-intact preparations to show ‘learning in the dish’ and long-term memory that persists for at least 18 h. The neurons that generate this behaviour are readily identifiable and, for the first time, we have recorded from a neuron during a training paradigm that leads to long-term memory formation in the same preparation. Specifically, we recorded from the respiratory neuron Right Pedal Dorsal 1 (RPeD1), which is part of the respiratory central pattern generator and initiates the aerial respiratory behaviour. Previous studies have shown that long-term memory of this behaviour results in reduced RPeD1 activity. In the present study, we demonstrate that preventing RPeD1 impulse activity between training sessions reduces the number of sessions needed to produce long-term memory in our semi-intact preparation. [ABSTRACT FROM AUTHOR]

It is widely appreciated that the selection and modulation of locomotor circuits are dependent on the actions of higher-order projection neurons. In the leech, Hirudo medicinalis, locomotion is modulated by a number of cephalic projection neurons that descend from the subesophageal ganglion in the head. Specifically, descending brain interneuron Tr2 functions as a command-like neuron that can terminate or sometimes trigger fictive swimming. In this study, we demonstrate that Tr2 is dye coupled to the dopaminergic neural network distributed in the head brain. These findings represent the first anatomical evidence in support of dopamine (DA) playing a role in the modulation of locomotion in the leech. In addition, we have determined that bath application of DA to the brain and entire nerve cord reliably and rapidly terminates swimming in all preparations exhibiting fictive swimming. By contrast, DA application to nerve cords expressing ongoing fictive crawling does not inhibit this motor rhythm. Furthermore, we show that Tr2 receives rhythmic feedback from the crawl central pattern generator. For example, Tr2 receives inhibitory post-synaptic potentials during the elongation phase of each crawl cycle. When crawling is not expressed, spontaneous inhibitory post- synaptic potentials in Tr2 correlate in time with spontaneous excitatory post-synaptic potentials in the CV motor neuron, a circular muscle excitor that bursts during the elongation phase of crawling. Our data are consistent with the idea that DA biases the nervous system to produce locomotion in the form of crawling. [ABSTRACT FROM AUTHOR]

Adult actinopterygian fishes typically perform steady forward swimming using either their pectoral fins or their body axis as the primary propulsor. In most species, when axial undulation is employed for swimming, the pectoral fins are tucked (i.e. adducted) against the body; conversely, when pectoral fins are beating, the body axis is held straight In contrast to adults, larval fishes can combine their pectoral fin and body-axis movements during locomotion; however, little is known about how these locomotor modes are coordinated. With this study we provide a detailed analysis of the coordinated fin and axial movements during slow and fast swimming by examining forward locomotion in larval zebrafish (Danio rerio L.). In addition, we describe the musculature that powers pectoral fin movement in larval zebrafish and discuss its functional implications. As larvae, zebrafish alternate their pectoral fins during slow swimming (0.011±0.001 mm ms-1) in conjunction with axial undulations of the same frequency (18-28 Hz). During fast swimming (0.109± 0.030 mm ms-1 36-67 Hz), the fins are tucked against the body and propulsion occurs by axial undulation alone. We show that during swimming, larval fishes can use a similar limb-axis coordination pattern to that of walking and running salamanders. We suggest that the fin-axis coordination observed in larval zebrafish may be attributed to a primitive neural circuit and that early terrestrial vertebrates may have gained the ability to coordinate limbs and lateral bending by retaining a larval central pattern generator for limb-axis coordination in the adult life history stage. [ABSTRACT FROM AUTHOR]

The article reports on a study which shows that the large-amplitude depolarization that is the cause of bursts of action potentials in the motor neurons in crab ventilatory system, is produced by a plateau differential. These motor neurons were found to satisfy several experimental tests which have been suggested for plateau potentials.

The article studies the maintenance of the functional motor pattern produced by the central pattern generator responsible for the gill ventilation rhythm in the shore crab Carcinus maenas. This motor pattern was found to maintain relatively constant phase relationships between the various motor bursts in the rhythm. The two purposes of the phase relationships in the ventilatory motor pattern are also discussed.

Neuronal inputs to microcircuits are often present as multiple copies of apparently equivalent neurons. Thus far, however, little is known regarding the relative influence on microcircuit output of activating all or only some copies of such an input. We examine this issue in the crab ( Cancer borealis ) stomatogastric ganglion, where the gastric mill (chewing) microcircuit is activated by modulatory commissural neuron 1 (MCN1), a bilaterally paired modulatory projection neuron. Both MCN1s contain the same co-transmitters, influence the same gastric mill microcircuit neurons, can drive the biphasic gastric mill rhythm, and are co-activated by all identified MCN1-activating pathways. Here, we determine whether the gastric mill microcircuit response is equivalent when stimulating one or both MCN1s under conditions where the pair are matched to collectively fire at the same overall rate and pattern as single MCN1 stimulation. The dual MCN1 stimulations elicited more consistently coordinated rhythms, and these rhythms exhibited longer phases and cycle periods. These different outcomes from single and dual MCN1 stimulation may have resulted from the relatively modest, and equivalent, firing rate of the gastric mill neuron LG (lateral gastric) during each matched set of stimulations. The LG neuron-mediated, ionotropic inhibition of the MCN1 axon terminals is the trigger for the transition from the retraction to protraction phase. This LG neuron influence on MCN1 was more effective during the dual stimulations, where each MCN1 firing rate was half that occurring during the matched single stimulations. Thus, equivalent individual- and co-activation of a class of modulatory projection neurons does not necessarily drive equivalent microcircuit output., Competing Interests: Competing interestsThe authors declare no competing or financial interests., (© 2020. Published by The Company of Biologists Ltd.)

The heart and pyloric rhythms of crustaceans have been studied separately and extensively over many years. Local and hormonal neuromodulation and sensory inputs into these central pattern generator circuits play a significant role in an animal's response to perturbations, but are usually lost or removed during in vitro studies. To examine simultaneously the in vivo motor output of the crustacean heart and pyloric rhythms, we used photoplethysmography. In the population measured ( n =49), the heart rhythm frequency ranged from 0.3 to 2.3 Hz. The pyloric rhythm varied from 0.2 to 1.6 Hz. We observed a weak correlation between the frequencies of the heart and pyloric rhythms. During multiple hour-long recordings, many animals held at a controlled temperature showed strong inhibitory bouts in which the heart decreased in frequency or become quiescent and the pyloric rhythm decreased in frequency. We measured the simultaneous responses of the rhythms to temperature ramps by heating or cooling the saline bath while recording both the heart and pyloric muscle movements. Q 10 , critical temperature (temperature at which muscle function is compromised) and changes in frequency were calculated for each of the rhythms tested. The heart rhythm was more robust to high temperature than the pyloric rhythm., Competing Interests: Competing interestsThe authors declare no competing or financial interests., (© 2019. Published by The Company of Biologists Ltd.)