Neuronal correlates of changes in the defensive behavior of Helix lucorum L. during ontogeny

In experiments on the isolated nervous system, responses of the same command neurones of avoidance behaviour to electrical stimulation of the intestinal nerve were compared in adult snails (Helix pomatia) and in snails 1-20 days old. Dynamics of spike reactions of the command neurones to rhythmic nerve stimulation differed in adult and newborn snails. In the neurones of young snails, no sensitization, common in the adult animals, was observed. Study of excitatory input in the command neurones during rhythmic nerve stimulation suggests independence of habituation of synaptic input amplitude and spike response sensitization, because dynamics of habituation coincided in the young and adult snails while spike response underwent sensitization only in adult animals.     

Interrelationship between grape snail alimentary and defense behavior command neurons]

Food stimulus (carrot juice), releasing feeding behaviour in intact snails, evoked spike discharges in giant meta-erebral cells (considered to be command neurones of feeding behaviour) and subthreshold EPSPs in giant parietal cells (command neurones of avoidance behaviour) of a half-intact preparation. Tactile stimulation, eliciting avoidance reactions in intact snails, evoked hyperpolarization in command neurones of feeding behaviour and a spike discharge in command neurones of avoidance behaviour. Spikes induced into either of command neurones produced no changes in activity of other command cells. Inhibition of command neurones of feeding behaviour is assumed to be the basis of behavioural choice when food and tactile stimulus are presented simultaneously.     

Dynamic defensive and feeding reactions to the acquisition of sensitization in edible snails]

In snails changes of defensive and alimentary behaviour and of reactions of command and modulator neurones of these kinds of behaviour were studied during elaboration of sensitization. After a single action of 50% chinine solution on mollusc's head, a short-term (during 50-70 min) and long-term (hours or days) facilitation was found of animals defensive reactions and responses of command neurones of defensive behaviour to tactile and chemical sensory stimuli. Alimentary behaviour of the snails and reactions of modulator neurones of alimentary behaviour to carrot juice presentation were inhibited in sensitized animals. Dynamics differences of defensive responses to tactile and chemical stimuli in the same sensitized animals were observed. Short-term responses facilitation under sensitization correlated chiefly with depolarization of the membrane potential of defensive behaviour neurones and with an increase of plasmatic membrane excitability. Prolonged facilitation of responses was due mainly to a change of effectiveness of synaptic transmission. The described model of sensitization elaboration can be a base for studying of molecular-cellular mechanisms, underlying the learning.     

Cellular reactions to elaboration of a backward conditioned connection in Helix lucorum

After 10-15 food stimuli paired with electrical shock in semi-intact snail preparation, responses to strong tactile stimuli identified feeding behaviour neurones were studied. Inhibition evoked by tactile stimulation in these cells before learning procedure disappeared and in some cases noxious stimulus evoked synaptic activation corresponding to feeding reactions in the intact animal. Changes in second-order sensory neurones pre-synaptic to the command neurones of avoidance behaviour are suggested to be the mechanism of forward conditioned connection as well as the mechanism of backward conditioned connection.     

The facilitation of defensive reactions during food consumption in the snail helix: the participation of glucose and gastrin/cholecystokinin-like peptide]

Molecular and cellular mechanisms of the interrelations between the feeding and defense behaviour were studied in a snail Helix lucorum. The dynamics of defense reactions was investigated in snails with different levels of feeding motivation. Defense reactions were suppressed in hungry snails, while 15-20 min after the beginning of food intake they were facilitated. The facilitation depended on a duration of starvation. Injection of 0.5 ml of 5 mM glucose solution (up to the glucose level in the haemolymph of a food satiated snail, 1.6-2.0 mM) or injections of 20-30 ng of synthetic analogues of the gastrointestinal peptides (pentagastrin of octapeptide cholecystokinin, CCK-8) facilitated the defense reaction in a hungry snail. Parameters of the facilitation were similar to those in the period of food intake. Activity of the command neurons of defense behaviour (L-PPL1) after the carrot juice application to the lip of a semi-intact preparation from a hungry snail was glucose-dependent. Similar glucose-dependent changes of L-PPL1 activity were found after CCK-8, but not FMRFamide application during the perfusion with 0.5 mM glucose. L-PPL1, but not L-PPa2-3 neurons were most sensitive to glucose and CCK-8 level changes in the Ringer solution. Adaptive significance of the behavioural phenomena as well as glucose and gastrin/CCK-like peptide participation in these processes are discussed.     

Acid-Base Balance Modulates Respiratory and Alimentary Behavior of the Mollusc Lymnaea stagnalis

An active reaction (change of pH) in hemolymph of the freshwater pulmonate mollusc Lymnaea stagnalis is studied in various physiological states: hunger, satiety, and alimentary excitation. In satiated animals the hemolymph pH is shifted to the acid area (7.6 ± 0.003) as compared with hungry animals (7.9 ± 0.05). The satiated individuals are characterized by an increased respiratory activity and decreased food consumption as compared with hungry animals. Acidification of the medium leads to an increase of excitability of the respiratory network interneurons (RPeD1, VD4, VD1/RPD2) and to inhibition of activity of the alimentary network interneurons (R/L CGC), whereas alkalization, to opposite effects. It is suggested that pH is one of factors coordinating activity of functional systems of Lymnaea stagnalis due to effect on neuronal correlates of the respiratory and alimentary behavior.     

Sensitization and habituation of command neurons during a defensive reflex in grape snails

All investigated neurones were classified into functional groups depending on their participation in unconditioned avoidance reflex of pneumostome closure. Habituation was produced by tactile stimuli applied with a frequency of 0.1 c/s. Spike responses of neurons participating in processing sensory information habituated gradually, while reactions of command neurones of avoidance behaviour became sensitized to the second or third stimulus of the series. Behavioral reaction was sensitized in parallel with command neurones in spite of habituation in other types of nerve cells. A conclusion is drawn that behavioral habituation is due to waning of responses in all the participating neurones, but the independnet process of sensitization is due to sensitization of command neurones. Probable mechanisms of neuronal sensitization are discussed.     

Overestimating resource value and its effects on fighting decisions

Much work in behavioral ecology has shown that animals fight over resources such as food, and that they make strategic decisions about when to engage in such fights. Here, we examine the evolution of one, heretofore unexamined, component of that strategic decision about whether to fight for a resource. We present the results of a computer simulation that examined the evolution of over- or underestimating the value of a resource (food) as a function of an individual's current hunger level. In our model, animals fought for food when they perceived their current food level to be below the mean for the environment. We considered seven strategies for estimating food value: 1) always underestimate food value, 2) always overestimate food value, 3) never over- or underestimate food value, 4) overestimate food value when hungry, 5) underestimate food value when hungry, 6) overestimate food value when relatively satiated, and 7) underestimate food value when relatively satiated. We first competed all seven strategies against each other when they began at approximately equal frequencies. In such a competition, two strategies--"always overestimate food value," and "overestimate food value when hungry"--were very successful. We next competed each of these strategies against the default strategy of "never over- or underestimate," when the default strategy was set at 99% of the population. Again, the strategies of "always overestimate food value" and "overestimate food value when hungry" fared well. Our results suggest that overestimating food value when deciding whether to fight should be favored by natural selection.     

A shift in prey color preference in the green toad Bufo viridis Laur. after food satiation]

In the behavioural experiments when showing simultaneously coloured stimuli (food targets) and also in choosing between two stimuli, red and blue, it had been shown with a high level of reliability that the hungry Bufo viridis which had not been fed for at least a week, in the overwhelming majority of cases chose the red-coloured food targets. Within 1-3 days after an abundant feeding, in the same animals the changes occurred in their preferences of the prey colour. The percentage of choosing the blue targets by the satiated toads significantly increases. In a separate series of experiments had been demonstrated that the changing motivation is conditioned by colour, since hungry and satiated Bufo viridis under conditions of simultaneous demonstration of four stimuli, black and three different grey stimuli, chose exclusively the black stimulus.     

Coordination of locomotor activity of mollusc Lymnaea stagnalis at nutrition: Role of the internal medium acid-base balance (pH)

The correlation was studied between the hemolymph pH value and the locomotion rate in the mollusc Lymnaea stagnalis at different degree of activation of the food-procuring behavior. It is established that the hemolymph pH value remains unchanged in the satiated (hungry), freely moving Lymnaea individuals, and, hence, it does not determine the locomotor activity of molluscs in these functional states. In the course of food consumption, locomotion decreases significantly, which is due to an observed decrease of the hemolymph pH. This effect is realized through the corresponding changes of electrical activity of motoneurons of the locomotor network of the Lymnaea nervous system (PeA-cluster). It is suggested that the pH value plays the modulating role in coordination of activities of the nervous centers of the mollusc nutrition and locomotion.     

Species richness along multiple gradients: testing a general multivariate model in oak savannas

Behavioral responses of prey to their predators can critically alter community dynamics. Whether or not a prey responds, clearly depends on the effectiveness of that response. The effectiveness on the other hand is predicted to depend on predator behavior. Actively searching predators can render the behavioral responses in their prey ineffective. Nevertheless, most studies investigating the optimal reaction of prey treated predators as immobile elements of the environment. I experimentally manipulated activity of poolfrog ( Rana lessonae ) tadpoles by keeping them at low and high food levels, and exposed them to three species of invertebrate predators ( Aeshna cyanea, Anax imperator , and Dytiscus marginalis ), whose activity also was manipulated through different food levels. Satiated, less active predators were more likely to kill hungry, more active tadpoles, but hungry predators killed hungry and satiated tadpoles about equally often. This result suggests that reducing their activity is a more effective strategy for tadpoles if the predators themselves are less active. On the other hand, against hungry, highly motivated predators, the behavioral avoidance strategies were essentially ineffective. Antipredator behavior is generally thought to stabilize the dynamics of predator–prey systems. The results presented here, however, suggest that the community dynamical consequences of antipredator behavior also critically depend on decisions made by predators.     

Chemoreception by the asteroid Cross aster papposus (L.)

Chemoreception abilities were demonstrated in the solasterid asteroid Crossaster papposus (L.) from a level substratum in the current-swept Crouch Estuary, England. C. papposus showed wide variability of positive and negative responses to crude extracts of prey species and (unattractive) conspecifics. C. papposus has an olfactory (distance chemoreception) ability to whole, live potential prey and conspecifics. This was confirmed by contact chemoreception responses, which are stronger, to the same stimuli. The olfaction tests in the Y-maze yielded variable results in which the initial (rheopositive or rheonegative) responses demonstrated that conspecifics and prey species were avoided yet subsequent arm choices of rheopositive individuals showed significant selection for prey over controls and no avoidance of conspecifics. The arm choices for prey relate to a priority of orientation to food and feeding over intraspecific avoidance in hungry asteroids. Intraspecific avoidance functions more in dispersion than prevention of cannibalism and is less important than feeding to hungry asteroids. Interpretation of chemically-mediated responses of asteroids requires appreciation of their innate behavioural variability which has led to confusion in the literature. This is the first experimental study of contact and distance chemoreception among solasterids. Their chemosensory abilities have previously been contradictory which may be due to intraspecific avoidance in satiated individuals of some species.     

The effect of predation risk on the predatory behaviour of a sit-and-wait predator,Ranatra dispar (Heteroptera: Nepidae), the water stick insect

Two primary defence behaviours, fore-leg extension to enhance crypsis and swimming to bottom and remaining motionless, of a predatory water bug,Ranatra dispar, are described along with their subsequent effect on foraging behaviour. It was hypothesised that hungry predators would respond less and for a shorter duration compared with recently fed individuals when exposed to a model threat stimulus, thereby tending to take a higher risk of predation during feeding than satiated animals. A greater proportion of animals responded overall with the leg extension response compared with the swimming response, although the mean duration of the former was significantly shorter than the latter response. A significantly higher proportion of nonfasted predators responded, and for a longer duration than fasted individuals. The type of model used significantly effected the proportion of animals that responded with leg extensions but not on its duration. In contrast, both number of animals and the duration of the swim/motionless behaviour were significantly effected by stimulus type. The subsequent effect of these 2 behavioural responses on feeding behaviour was examined and showed that although about the same number of predators removed prey from their mouthparts during both responses, significantly more prey were dropped, and therefore lost, during swimming. The results clearly indicate the significant effects that defensive behaviours have on time budgets in foraging behaviour.     

Effect of serotonin on the activity of the neurons involved in the realization of the avoidance reflex of the snail

Bath application of 10(-5) mol/l of serotonin (5-HT) elicited a 50% increase of summary EPSPs recorded in command neurones for avoidance behaviour. No significant changes of rest potential and input resistance were seen in these cells. 5-HT evoked an increase of spontaneous level of firing in motoneurones involved in the same reflex, as well as an increase in the number of spikes which paralleled increase of EPSPs to the same stimulus in command neurones. In sensory cells, presynaptic to the command neurones, application of 5-HT evoked a significant increase of excitability and of input resistance. Monosynaptic EPSPs recorded in the command neurones showed a 40% increase after serotonin application. It is concluded that the major locus of plastic changes evoked by 5-HT application in the neuronal chain underlying avoidance reflex is the synaptic contact between sensory and command neurones.     

Interconnection between parameters of motor activity and blood glucose concentration in newborn rats at starvation and under glucose load conditions

The study is carried out on the one-, 3-, 5-, 7, and l0-day old free moving or loosely fixed rat pups (P1–10). In satiated and submitted to the 24-h starvation rat pups, parameters of spontaneous periodical motor activity (SPMA) and the blood glucose content were determined. The total glucose level in hungry rat pups was, on average, 1.5–2.5 times lower than in satiated animals. Administration of glucose to hungry rat pups increased 6–11 times its concentration in blood as compared with the animals not obtaining glucose. The glucose administration to the satiated rat pups led to a rise of its level in blood from 2 to 5 times as compared with intact animals, which was 2–3 times less than in the case of hungry animals. Analysis of pattern of motor activity recorded under conditions of the glucose deficit caused by the 24-h starvation of rat pups did not reveal significant changes of ratio of rhythmical components. Introduction of glucose to hungry and to the lesser degree to satiated rat pups led to potentiation of the minute rhythm of activity in all age groups. The exception was the first day after birth when the glucose administration to satiated rat pups promoted an enhancement of the decasecond rhythm and a decrease of the total level of motor activity. Comparison of ontogenetic dynamics of the SPMA parameters and the glucose content in blood of hungry rat pups revealed the clearly expressed regularity absent in the satiated animals: the glucose level in blood was higher during activity than in the state of rest. The performed study has shown that intensity, duration, and, to a degree, pattern of SPMA in the newborn rat pups depend on the level of satiety, and can be significantly changed in the artificially produced hypo- or hyperglycemia. The existing ontogenetic fluctuations in the character of reaction at performance of the glucose tolerance test can be connected both with morphofunctional maturation of the motor system and with immaturity of various chains of carbohydrate metabolism.     

Pathophysiologic alterations in Biomphalaria glabrata infected with Angiostrongylus costaricensis

Hemolymph glucose, alkaline phosphatase, lactic dehydrogenase, and creatine phosphokinase in Biomphalaria glabrata infected with Angiostrongylus costaricensis were significantly higher on day 27 postinfection (PI) than in uninfected snails. Hemolymph total calcium from infected snails was less on days 6, 12, and 27 PI than that from controls. Total hemolymph protein was similar for controls and infected animals during the entire study. Throughout the study the mean number of amoebocytes/mm³ hemolymph from infected snails was significantly less than that for controls. Mean total wet weights of digestive gland and foot muscle from infected and uninfected snails was similar throughout the study. Mean μg glycogen/mg wet weight of digestive gland from infected snails was significantly greater on days 24, 27, and 28 PI than that from controls. Mean μg glycogen/mg wet weight of foot muscle from infected snails was significantly reduced between days 12 and 28 PI from that of uninfected snails. It is suggested that hemolymph glucose and digestive gland glycogen in infected snails are augmented by glycogen breakdown in the foot muscle of parasitized animals. Elevations in hemolymph enzymes are due to tissue destruction by larvae emerging from the foot muscle of infected snails. Parasite-induced derangements in shell metabolism underlie observed changes in hemolymph calcium in infected snails.     

Alteration of thermoregulation behavior in juvenile fish in relation to satiation level

Thermoregulation behavior of satiated and hungry juvenile sterlet Acipenser ruthenus, the Siberian sturgeon Acipenser baeri, and the goldfish Carassius auratus was studied. Significant alterations of thermoregulation behavior appear in juvenile fish one day after food deprivation. Hungry fish preferred, on average, lower temperatures, their temperature range was broader, and the locomotor activity was higher than in satiated fish.     

Assessing Projection Bias in Consumers’ Food Preferences

The aim of this study is to test whether projection bias exists in consumers’ purchasing decisions for food products. To achieve our aim, we used a non-hypothetical experiment (i.e., experimental auction), where hungry and non-hungry participants were incentivized to reveal their willingness to pay (WTP). The results confirm the existence of projection bias when consumers made their decisions on food products. In particular, projection bias existed because currently hungry participants were willing to pay a higher price premium for cheeses than satiated ones, both in hungry and satiated future states. Moreover, participants overvalued the food product more when they were delivered in the future hungry condition than in the satiated one. Our study provides clear, quantitative and meaningful evidence of projection bias because our findings are based on economic valuation of food preferences. Indeed, the strength of this study is that findings are expressed in terms of willingness to pay which is an interpretable amount of money.     

Humoral response of the snail Biomphalaria glabrata to trematode infection: observations on a circulating hemagglutinin

The hemolymph of invertebrates often contains molecules that agglutinate vertebrate erythrocytes and that may function as humoral mediators of "non-self" recognition. The objectives of this study were to 1) determine if exposure of M line or 10-R2 strain Biomphalaria glabrata snails to infection with the trematodes Echinostoma paraensei and Schistosoma mansoni could increase agglutinating activity in snail hemolymph, and 2) identify particular hemolymph molecules with such activity. In some host-parasite combinations, such as juvenile M line snails and E. paraensei, infection provoked significant elevations in titer from as early as 2 days postinfection (dpi) through 15 dpi. In other combinations, as with 10-R2 snails and E. paraensei or S. mansoni, host responses were comparatively modest, yet still measurable. In general, E. paraensei and S. mansoni elicited different responses from the same host strain, and M line and 10-R2 snails responded differently to the same parasite. Further study of the response of juvenile M line snails to E. paraensei indicated that hemolymph agglutinating activity could be inhibited by several monosaccharides (including L-fucose) and by EDTA and EGTA. An affinity column containing L-fucose agarose beads was used to purify molecules with agglutinating activity from the hemolymph of such snails. The fraction eluted from the column by 0.2 M L-fucose was shown by SDS-PAGE to contain a broad band of 80-120 kD and, less consistently, a 200 kD band. Following extensive dialysis to remove L-fucose, this fraction had agglutinating activity. As a previous study has shown that the hemolymph of E. paraensei-infected snails contains significantly increased quantities of 80-120 kD polypeptides, it is concluded that polypeptides in this size range are responsible, at least in part, for the increased hemolymph agglutination activity in such snails.     

Effect of serotonin and noradrenaline on the magnitude of the response of the command neurons for defensive behavior in Helix lucorum L

Changes in synaptic responses of identified command neurones of avoidance behaviour to the electric nerve stimulation were investigated in the isolated nervous system of the snail during bath application of serotonin or noradrenaline. Serotonin (10(-5) M) elicited an increase of summary EPSP amplitude in the cells without changes of input resistance and resting potential. Noradrenaline (10(-5) M) application evoked an increase of EPSP amplitude, accompanied by an increase of the input resistance. Mechanisms of serotonin and noradrenaline influence on synaptic responses are discussed.