Varvara E. Dyakonova

Complex avoidance behaviour and its neurochemical regulation in the land snail Cepaea nemoralis

1. In hot plate experiments, the pulmonate land snail Cepaea nemoralis displays a biphasic passive/active avoidance behaviour composed of retraction and subsequent searching mediated by antagonistic muscular systems. The switch, between the behaviours, is under neuronal control. 2. Leu- and met-enkephalin, as well as FMRFamide-antiserum, attenuated the retraction response and potentiated the searching behaviour. Opposite effects were achieved by injection of antisera to the enkephalins. 3. Both retraction and searching behaviours were potentiated by 5-HT. Methysergide antagonized the effects of the enkephalins on the searching behaviour. 4. We conclude that endogenous opioids act antagonistic to FMRFamide in the neuronally controlled switch between passive and active avoidance behaviour.

What are the elements of motivation for acquisition of conditioned taste aversion

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.

Serotonin precursor (5-hydroxytryptophan) causes substantial changes in the fighting behavior of male crickets, Gryllus bimaculatus

This study demonstrates that injection of the serotonin precursor 5-HTP causes substantial changes in the behavioral state, fighting behavior and ability to establish winner–loser relationships in male crickets (Gryllus bimaculatus). The characteristic features of 5-HTP-treated crickets include an elevated posture, enhanced general activity, longer duration of fighting, enhanced rival singing and a decreased ability to produce a clear fight loser. In addition, 5-HTP-treated males showed a slightly delayed latency to spread their mandibles, a decreased number of attacks and an equal potential to win in comparison to controls (physiological solution-treated males). The obtained results imply a significant role for serotonin in the regulation of social status-related behaviors in G. bimaculatus. Specifically, these data indicate that a decrease in serotonergic activity may be functionally important for the control of loser behavior and that some behavioral features of dominant male crickets are likely to be connected with the activation of the serotonergic system.

Memory block: A consequence of conflict resolution

ABSTRACT 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. Summary: Snails engage in the concept of ‘necessity knows no law’ as memories not to respond to a food substance are overpowered by hunger.

Coordination of rhythm-generating units via NO and extrasynaptic neurotransmitter release

The buccal ganglia of the mollusc, Lymnaea stagnalis, contain two distinct but interacting rhythm-generating units: the central pattern generator for the buccal rhythm and nitrergic B2 neurons controlling gut motility. Nitric oxide (NO) has previously been demonstrated to be involved in the activation of the buccal rhythm. Here, we found that NO-generating substances (SNP and SNAP) activated the buccal rhythm while slowing the endogenous rhythm of B2 bursters. The inhibitor of NO-synthase, L-NNA, the NO scavenger PTIO, or the inhibitor of soluble guanylyl cyclase, ODQ, each produced opposite, depolarising effects on the B2 neuron. In isolated B2 cells, only depolarising effects of substances interfering with NO production or function (PTIO, L-NNA and ODQ) were detected, whereas the NO donors had no hyperpolarising effects. However, when an isolated B2 cell was placed close to its initial position in the ganglion, hyperpolarising effects could be obtained with NO donors. This indicates that extrasynaptic release of some unidentified factor(s) mediates the hyperpolarising effects of NO donors on the B2 bursters. The results suggest that NO is involved in coordination between the radula and foregut movements and that the effects of NO are partially mediated by the volume chemical neurotransmission of as yet unknown origin.

Previous Motor Experience Enhances Courtship Behavior in Male Cricket Gryllus bimaculatus

In crickets Gryllus bimaculatus, flight has been shown to be able to promote aggressive encounters between males and to suppress escape behavior. The aim of this study was to examine the influence of flight on male behavior in male–female interactions. We found that flown males demonstrate enhanced courtship behavior. The latency of calling song was significantly shorter, while the relative total duration of singing as well as the duration of singing episodes longer in flown males than in the control. Mating rate was also significantly higher in the experimental group containing flown males. The results suggest that, in addition to previously reported effects on aggressiveness and escape, flying has a profound accelerating effect on male courtship behavior.

The activity of isolated neurons and the modulatory state of an isolated nervous system represent a recent behavioural state

ABSTRACT Behavioural/motivational state is known to influence nearly all aspects of physiology and behaviour. The cellular basis of behavioural state control is only partially understood. Our investigation, performed on the pond snail Lymnaea stagnalis whose nervous system is useful for work on completely isolated neurons, provided several results related to this problem. First, we demonstrated that the behavioural state can produce long-term changes in individual neurons that persist even after neuron isolation from the nervous system. Specifically, we found that pedal serotonergic neurons that control locomotion show higher activity and lower membrane potential after being isolated from the nervous systems of hungry animals. Second, we showed that the modulatory state (the chemical neuroactive microenvironment of the central ganglia) changes in accordance with the nutritional state of an animal and produces predicted changes in single isolated locomotor neurons. Third, we report that observed hunger-induced effects can be explained by the increased synthesis of serotonin in pedal serotonergic neurons, which has an impact on the electrical activity of isolated serotonergic neurons and the intensity of extrasynaptic serotonin release from the pedal ganglia. Summary: In the pond snail Lymnaea stagnalis, the behavioural state can produce long-term changes in individual neurons that persist even after neuron isolation from the nervous system as well as alterations in the neurochemical microenvironment of the central ganglia.

The activity of isolated snail neurons controlling locomotion is affected by glucose

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.

Effects of 5-HT and insulin on learning and memory formation in food-deprived snails

&NA; The pond snail Lymnaea stagnalis learns conditioned taste aversion (CTA) and consolidates it into long‐term memory (LTM). How well they learn and form memory depends on the degree of food deprivation. Serotonin (5‐HT) plays an important role in mediating feeding, and insulin enhances the memory consolidation process following CTA training. However, the relationship between these two signaling pathways has not been addressed. We measured the 5‐HT content in the central nervous system (CNS) of snails subjected to different durations of food deprivation. One‐day food‐deprived snails, which exhibit the best learning and memory, had the lowest 5‐HT content in the CNS, whereas 5‐day food‐deprived snails, which do not learn, had a high 5‐HT content. Immersing 1‐day food‐deprived snails in 5‐HT impaired learning and memory by causing an increase in 5‐HT content, and that the injection of insulin into these snails reversed this impairment. We conclude that insulin rescues the CTA deficit and this may be due to a decrease in the 5‐HT content in the CNS of Lymnaea. HighlightsOne‐day food‐deprived snails that exhibit the best CTA learning had a low 5‐HT content in their CNS.Five‐day food‐deprived snails, which did not learn, had a high level of 5‐HT content in their CNS.Insulin rescues the CTA deficit and this may be due to a decrease in the 5‐HT content in the CNS.

Discrete Modeling of Multi-transmitter Neural Networks with Neuronal Competition

We propose a novel discrete model of central pattern generators (CPG), neuronal ensembles generating rhythmic activity. The model emphasizes the role of nonsynaptic interactions and the diversity of electrical properties in nervous systems. Neurons in the model release different neurotransmitters into the shared extracellular space (ECS) so each neuron with the appropriate set of receptors can receive signals from other neurons. We consider neurons, differing in their electrical activity, represented as finite-state machines functioning in discrete time steps. Discrete modeling is aimed to provide a computationally tractable and compact explanation of rhythmic pattern generation in nervous systems. The important feature of the model is the introduced mechanism of neuronal competition which is shown to be responsible for the generation of proper rhythms. The model is illustrated with two examples: a half-center oscillator considered to be a basic mechanism of emerging rhythmic activity and the well-studied feeding network of a pond snail. Future research will focus on the neuromodulatory effects ubiquitous in CPG networks and the whole nervous systems.