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We have recently shown that experience of flight remarkably enhanced subsequent terrestrial phonotaxis in females in response to the male calling song. Here, we elucidated the possible roles of octopamine and serotonin in the enhancing effect of flying on phonotactic behavior. Octopamine is known to be released into the hemolymph during flight in insects; however, the octopamine receptor antagonist epinastine did not abolish the effects of flight in our study. On the contrary, the drug significantly potentiated the influence of flying on phonotactic behavior. The octopamine receptor agonist chlordimeform, at a concentration of 2 mM, which was previously found to activate aggression in crickets, dramatically reduced the phonotactic response. However, at a 10-times-lower dose, chlordimeform produced a light but significant decrease in the time that females took to reach the source of the calling song. A similar effect was produced by octopamine itself, which hardly passes the blood-brain barrier in insects. The effect of flight was completely abolished in female crickets treated with alpha-methyl tryptophan (AMTP). AMPT suppresses the synthesis of serotonin, decreasing its content in the nervous systems of insects, including crickets. An activation of the serotonin synthesis with 5-hydroxytryptophan mimicked the effect of flight by increasing the number of visits to and the time spent in the zone near the source of the calling song. The 5-HT content in the third thoracic ganglion was significantly higher in flyers compared to the control group. In contrast, no changes in the octopamine level were observed in the third thoracic ganglion, which is known to play a crucial role in decision-making involved in intraspecific interactions. Therefore, the results suggest that although octopamine is known to be released into the hemolymph during flight, it is likely to inhibit rather than activate the central mechanisms related to phonotaxis. The weak facilitating effect of a low dose of chlordimeform can be attributed to the activation of peripheral octopaminergic receptors. Our results suggest that the serotoninergic system may contribute to the facilitation of female phonotactic behavior by flying. We suggest that both flying and serotonin enhance sexual motivation in females and, by these means, impact their behavioral response to the male calling song.

Intense species-specific locomotion changes the behavioural and cognitive states of various vertebrates and invertebrates. However, whether and how reproductive behaviour is affected by previous increased motor activity remains largely unknown. We addressed this question using a model organism, the pond snail Lymnaea stagnalis . Intense crawling in shallow water for two hours had previously been shown to affect orienting behaviour in a new environment as well as the state of the serotonergic system in L. stagnalis . We found that the same behaviour resulted in an increased number of egg clutches and the total number of eggs laid in the following 24 h. However, the number of eggs per clutch was not affected. This effect was significantly stronger from January to May, in contrast to the September-December period. Transcripts of the egg-laying prohormone gene and the tryptophan hydroxylase gene, which codes for the rate-limiting enzyme in serotonin synthesis, were significantly higher in the central nervous system of snails that rested in clean water for two hours after intense crawling. Additionally, the neurons of the left (but not the right) caudo-dorsal cluster (CDC), which produce the ovulation hormone and play a key role in oviposition, responded to stimulation with a higher number of spikes, although there were no differences in their resting membrane potentials. We speculate that the left-right asymmetry of the response was due to the asymmetric (right) location of the male reproductive neurons having an antagonistic influence on the female hormonal system in the hermaphrodite mollusc. Serotonin, which is known to enhance oviposition in L. stagnalis , had no direct effect on the membrane potential or electrical activity of CDC neurons. Our data suggest that (i) two-hour crawling in shallow water enhances oviposition in L. stagnalis , (ii) the effect depends on the season, and (iii) the underlying mechanisms may include increased excitability of the CDC neurons and increased expression of the egg-laying prohormone gene.

Benefits of physical exercise for brain functions are well documented in mammals, including humans. In this review, we will summarize recent research on the effects of species-specific intense locomotion on behavior and brain functions of different invertebrates. Special emphasis is made on understanding the biological significance of these effects as well as underlying cellular and molecular mechanisms. The results obtained in three distantly related clades of protostomes, Nematodes, Molluscs and Artropods, suggest that influence of intense locomotion on the brain could have deep roots in evolution and wide adaptive significance. In C. elegans , improved learning, nerve regeneration, resistance to neurodegenerative processes were detected after physical activity; in L. stagnalis -facilitation of decision making in the novel environment, in Drosophila -increased endurance, improved sleep and feeding behavior, in G. bimaculatus -improved orientation in conspecific phonotaxis, enhanced aggressiveness, higher mating success, resistance to some disturbing stimuli. Many of these effects have previously been described in mammals as beneficial results of running, suggesting certain similarity between distantly-related species. Our hypothesis posits that the above modulation of cognitive functions results from changes in the organism's predictive model. Intense movement is interpreted by the organism as predictive of change, in anticipation of which adjustments need to be made. Identifying the physiological and molecular mechanisms behind these adjustments is easier in experiments in invertebrates and may lead to the discovery of novel neurobiological mechanisms for regulation and correction of cognitive and emotional status., Competing Interests: The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest., (Copyright © 2022 Dyakonova, Mezheritskiy, Boguslavsky, Dyakonova, Chistopolsky, Ito and Zakharov.)

In the cricket Gryllus bimaculatus, flying occurs soon after the last imaginal molt and precedes the mating behavior in natural conditions. Here, we tested the hypothesis that flying may improve subsequent behavioral performance in a novel environment in female crickets. We developed a behavioral set-up to test female cricket responsiveness to male calling song as well as their ability to locate and find the source of the song. The male song was produced by a loudspeaker hidden behind the fabric wall of a spacious square arena. Forced flight prior to the test promoted female sexual searching behavior in the novel environment. After the flight, more females reached the hidden source zone, spent more time near the source and finally more of them climbed over the wall section immediately in front of the hidden loudspeaker. At the same time, their behavior in the arena did not differ from the control group when the calling song was not delivered, suggesting that flight exerts its behavioral effects by influencing sexual motivation. Our results support the suggestion that preceding intense locomotion facilitates sexual searching behavior of females in a novel environment.

Simulations of neural activity are commonly based on differential equations. We address the question what can be achieved with a simplified discrete model. The proposed model resembles artificial neural networks enriched with additional biologically inspired features. A neuron has several states, and the state transitions follow endogenous patterns which roughly correspond to firing behavior observed in biological neurons: oscillatory, tonic, plateauing, etc. Neural interactions consist of two components: synaptic connections and extrasynaptic emission of neurotransmitters. The dynamics is asynchronous and event-based; the events correspond to the changes in neurons activity. This model is innovative in introducing discrete framework for modeling neurotransmitter interactions which play the important role in neuromodulation. We simulate rhythmic activity of small neural ensembles like central pattern generators (CPG). The modeled examples include: the biphasic rhythm generated by the half-center mechanism with the post-inhibitory rebound (like the leech heartbeat CPG), the triphasic rhythm (like in pond snail feeding CPG) and the pattern switch in the system of several neurons (like the switch between ingestion and egestion in Aplysia feeding CPG). The asynchronous dynamics allows to obtain multi-phasic rhythms with phase durations close to their biological prototypes. The perspectives of discrete modeling in biological research are discussed in the conclusion.

The role of serotonin in the immediate and delayed influence of physical exercise on brain functions has been intensively studied in mammals. Recently, immediate effects of intense locomotion on the decision-making under uncertainty were reported in the Great Pond snail, Lymnaea stagnalis (Korshunova et al., 2016). In this animal, serotonergic neurons control locomotion, and serotonin modulates many processes underlying behavior, including cognitive ones (memory and learning). Whether serotonin mediates the behavioral effects of intense locomotion in mollusks, as it does in vertebrates, remains unknown. Here, the delayed facilitating effects of intense locomotion on the decision-making in the novel environment are described in Lymnaea . Past exercise was found to alter the metabolism of serotonin, namely the content of serotonin precursor and its catabolites in the cerebral and pedal ganglia, as measured by high-performance liquid chromatography. The immediate and delayed effects of exercise on serotonin metabolism were different. Moreover, serotonin metabolism was regulated differently in different ganglia. Pharmacological manipulations of the serotonin content and receptor availability suggests that serotonin is likely to be responsible for the locomotor acceleration in the test of decision-making under uncertainty performed after exercise. However, the exercise-induced facilitation of decision-making (manifested in a reduced number of turns during the orienting behavior) cannot be attributed to the effects of serotonin., (Copyright © 2020 Aonuma, Mezheritskiy, Boldyshev, Totani, Vorontsov, Zakharov, Ito and Dyakonova.)

This is an attempt to describe and explain so-called timbre-based music as a special system of musicking, communication, and psychological and social usage, which along with its corresponding beliefs constitutes a viable alternative to "frequency-based" music. Unfortunately, the current scientific research into music has been skewed almost entirely in favor of the frequency-based music prevalent in the West. Subsequently, whenever samples of timbre-based music attract the attention of Western researchers, these are usually interpreted as "defective" implementations of frequency-based music. The presence of discrete pitch is often regarded as the structural criterion that distinguishes music from non-music. We would like to present evidence to the contrary-in support of the existence of indigenous music systems based on the discretization and patterning of aspects of timbre, rather than pitch. This evidence comes mainly from extensive ethnographic research systematically conducted in eastern European and Asian parts of Russia from the 1890s. It involved the efforts of thousands of specialists and was coordinated by dozens of research institutions, and it has included not just ethnomusicology but linguistics, philology, organology, archaeology, anthropology, geography, and religious, and social studies. Much of the data has not been translated into Western languages. Although some Soviet-era publications were tainted by Marxist ideology, many researchers strove to provide accurate information (despite at times having been prosecuted for their work), and post-1990 research undertook a substantial revision of ideologically compromised concepts. Timbre-based tonal organization (TO) differs from that based on frequency in its personal orientation: musicking here occurs primarily for oneself and/or for close relatives/friends. Collective music-making is rare and exceptional. The foundation of timbre-based music seems to have vocal roots and rests on "personal song"-a system of personal identification through individualized patterns of rhythm, timbre, and pitch contour, utilized like a "human voice"-whose sound enables the recognition of a particular individual. The instrumental counterpart of the personalized singing tradition is the jaw harp tradition. The jaw harp is the principal musical instrument for at least 21 ethnicities in Russia, who occupy over half the territory of the country. The evolution of its TO forms the backbone for the development of timbre-based music art. Here, we provide the acoustic, socio-cultural, geographic, and chronological overview of timbre-based music., (Copyright © 2020 Nikolsky, Alekseyev, Alekseev and Dyakonova.)

Food deprivation for 1 day in the pond snail Lymnaea stagnalis before aversive classical conditioning results in optimal conditioned taste aversion (CTA) and long-term memory (LTM) formation, whereas 5-day food deprivation before training does not. We hypothesize that snails do in fact learn and form LTM when trained after prolonged food deprivation, but that severe food deprivation blocks their ability to express memory. We trained 5-day food-deprived snails under various conditions, and found that memory was indeed formed but is overpowered by severe food deprivation. Moreover, CTA-LTM was context dependent and was observed only when the snails were in a context similar to that in which the training occurred., (© 2015. Published by The Company of Biologists Ltd.)

The involvement of serotonin in mediating hunger-related changes in behavioral state has been described in many invertebrates. However, the mechanisms by which hunger signals to serotonergic cells remain unknown. We tested the hypothesis that serotonergic neurons can directly sense the concentration of glucose, a metabolic indicator of nutritional state. In the snail Lymnaea stagnalis, we demonstrate that completely isolated pedal serotonergic neurons that control locomotion changed their biophysical characteristics in response to glucose application by lowering membrane potential and decreasing the firing rate. Additionally, the excitatory response of the isolated serotonergic neurons to the neuroactive microenvironment of the pedal ganglia was significantly lowered by glucose application. Because hunger has been reported to increase the activity of select neurons and their responses to the pedal ganglia microenvironment, these responses to glucose are in accordance with the hypothesis that direct glucose signaling is involved in the mediation of the hunger-related behavioral state.

The pond snail Lymnaea stagnalis is capable of being classically conditioned to avoid food and to consolidate this aversion into a long-term memory (LTM). Previous studies have shown that the length of food deprivation is important for both the acquisition of taste aversion and its consolidation into LTM, which is referred to as conditioned taste aversion (CTA). Here we tested the hypothesis that the hemolymph glucose concentration is an important factor in the learning and memory of CTA. One-day food deprivation resulted in the best learning and memory, whereas more prolonged food deprivation had diminishing effects. Five-day food deprivation resulted in snails incapable of learning or remembering. During this food deprivation period, the hemolymph glucose concentration decreased. If snails were fed for 2days following the 5-day food deprivation, their glucose levels increased significantly and they exhibited both learning and memory, but neither learning nor memory was as good as with the 1-day food-deprived snails. Injection of the snails with insulin to reduce glucose levels resulted in better learning and memory. Insulin is also known to cause a long-term enhancement of synaptic transmission between the feeding-related neurons. On the other hand, injection of glucose into 5-day food-deprived snails did not alter their inability to learn and remember. However, if these snails were fed on sucrose for 3min, they then exhibited learning and memory formation. Our data suggest that hemolymph glucose concentration is an important factor in motivating acquisition of CTA in Lymnaea and that the action of insulin in the brain and the feeding behavior are also important factors., (Copyright © 2013 Elsevier Inc. All rights reserved.)

Intraspecific aggression is influenced in numerous animal groups by the previous behavioral experiences of the competitors. The underlying mechanisms are, however, mostly obscure. We present evidence that a form of experience-dependent plasticity of aggression in crickets is mediated by octopamine, the invertebrate counterpart of noradrenaline. In a forced-fight paradigm, the experience of flying maximized the aggressiveness of crickets at their first encounter and accelerated the subsequent recovery of aggressiveness of the normally submissive losers, without enhancing general excitability as evaluated from the animals' startle responses to wind stimulation. This effect is transitory and concurrent with the activation of the octopaminergic system that accompanies flight. Hemocoel injections of the octopamine agonist chlordimeform (CDM) had similar effects on aggression but also enhanced startle responses. Serotonin depletion, achieved using alpha-methyl-tryptophan, enhanced startle responses without influencing aggression, indicating that the effect of CDM on aggression is not attributable to increased general excitation. Contrasting this, aggressiveness was depressed, and the effect of flying was essentially abolished, in crickets depleted of octopamine and dopamine using alpha-methyl-p-tyrosine (AMT). CDM restored aggressiveness in AMT-treated crickets, indicating that their depressed aggressiveness is attributable to octopamine depletion rather than to dopamine depletion or nonspecific defects. Finally, the flight effect was blocked in crickets treated with the octopamine receptor antagonist epinastine, or with the alpha-adrenoceptor and octopamine receptor antagonist phentolamine, but not with the beta-adrenoceptor antagonist propranolol. The idea that activity-specific induction of the octopaminergic system underlies other forms of experience-dependent plasticity of aggressive motivation in insects is discussed.

In the pond snail Lymnaea stagnalis, D-glucose action was investigated on electrical activity of identified central neurons. In the CNS preparations isolated from specimens that starved for 24-96 h, D-glucose added to a standard or HiDi saline at 500-700 microg/ml effectively hyperpolarized ca. 90% of feeding related neurons B1, SO and CGC. However, not all feeding-related neurons examined were responsive to glucose. Experiments on cells of the serotonergic Pedal A cluster have shown that hyperpolarizing action of D-glucose is retained following complete isolation of "hunger" neurons. Threshold concentration producing 1-3 mV hyperpolarization was ca. 50 microg/ml. The results suggest a direct glucose involvement in the mechanisms that control feeding behavior in Lymnaea.

We investigated the effect of food intake on the serotonin and dopamine levels of the CNS as well as on the spontaneous firing activity of the CGC in isolated preparations from starved, feeding and satiated animals. Furthermore we investigated the effects of 1 microM serotonin and/or dopamine and their mixture on the firing activity of the CGC. The HPLC assay of serotonin and dopamine showed that during food intake both the serotonin and dopamine levels of the CNS increased whereas in satiated animals their levels were not significantly more than the control levels. Recording from the CGC in isolated CNS preparation from starved, feeding or satiated animals showed that feeding increased the firing frequency of the CGC compared to the starved control. The application of 1 microM dopamine decreased the firing frequency whereas the application of 1 microM serotonin increased the firing frequency of the CGC. We conclude that during food intake the external and internal food stimuli increase the activity of the central monoaminergic system and also increase the levels of monoamines in the CNS. Furthermore, we also suggest that the increased dopamine and serotonin levels both affect the activity of the serotonergic neurons during the different phases of feeding.

In the cricket Gryllus bimaculatus, the opiate antagonist naloxone, 30 microg in 50 microl into hemolymph per animal, caused a release of intraspecific aggression in male-losers and in females. Naloxone had no significant effect on the aggression of winners and isolated males. The agonist of mu-opiate receptors DAGO, 45 microg, suppressed aggression in male winners and decreased the percentage and duration of contact fights between previously isolated males. The results suggest that, during social contacts, the activity of the endogenous opioid system may contribute to suppressing aggression in subordinate males, as well as in females.

1. In the cricket Gryllus bimaculatus, effects of opiate receptor antagonist naloxone, 9 or 30 microg per animal, on aggressive behavior were investigated. 2. Naloxone had no significant impact on aggression of isolated and dominant males. In contrast, the drug caused a dramatic release of social aggression in female and subordinate male crickets. 3. The results suggest that activity of the opioid system contributes to suppress aggression in subordinate males, as well as in females, during social contacts.

1. Opioid- and FMRFamide (FMRFa)-ergic systems are believed to play antagonistic behavioral roles in both higher and lower animals. In our previous experiments on a snail, behavioral choice has been demonstrated to be dependent on a balance between FMRFa and enkephalins [7]. Here, we examined if the disturbance of the balance causes changes in the activity of both systems. Opiate receptor blocker naloxone was applied and its effect on c-jun expression of met-enkephalin (MEnk)- and FMRFa-ergic neurons was examined immunocytochemically in terrestrial gastropod snail Cepaea nemoralis. 2. In control, untreated snails, central neurons with c-jun/AP-1-like-immunoreactivity were found to occur. These included MEnk-, FMRFa- and 5HT-immunoreactive (-ir) neurons, as was revealed by double-labelling. 3. After treatment with naloxone for 4 h, the following changes were observed: (i) increase in the number of MEnk-ir neurons; increase in the number of neurons showing c-jun/AP-1 and MEnk double-labeling; (ii) disappearance of c-jun/AP-1-immunoreactivity from some FMRFa-ir neurons. 4. It is suggested that immediate early genes are involved in the mechanisms responsible for the reciprocal regulation of the opioid and antiopioid neuropeptide systems.

Immunocytochemical techniques for double staining were employed to investigate the morphological basis for interactions between enkephalins and other neuroactive compounds in the behavior of the gastropod mollusc Cepaea nemoralis. Coexistence of each of the two enkephalins with FMRFamide, serotonin or GABA-like immunoreactivity was found in certain neurons in cerebral, parietal, and pedal ganglia. Tyrosine hydroxylase-immunoreactive neurons were occasionally seen in close apposition to, but never colocalized with, the enkephalins. A comparison between these anatomical observations and previous behavioral studies suggests that in gastropod molluscs cotransmission of enkephalins with classical transmitters may, at least partly, reflect synergism of these substances in the control of definite behavioral programs.

The monoaminergic system of the pond snail Lymnaea stagnalis changed markedly following prolonged exposure to chlorpromazine (CPZ). HPLC-ED indicated that levels of serotonin (5-HT), but not those of dopamine, were significantly decreased (e.g., to less than 40% after 30 days of treatment with 1 microM CPZ). Glyoxylate-induced fluorescence was depressed to undetectable levels in selected subpopulations of 5-HT neurons. Performance of 5-HT-dependent motor behaviors was impaired, and a considerably decreased firing activity was observed in affected 5-HT neurons. The present results, in accord with past ones with haloperidol, suggest that a previously overlooked mechanism of monoamine down-regulation may contribute to affects of antipsychotic drugs.