Effects of ageing on visual discrimination learning in Drosophila melanogaster

Two experiments examined conditioned suppression of the Proboscis Extension Response (PER), unconditionally released by sucrose stimulation of gustatory tarsal receptors, in young (7-day-old), middle-aged (30-day-old) and old (50-day-old) Drosophila melanogaster males, reared at 25°C. Individual flies were trained in a differential conditioning procedure in which a white stimulus signalled a quinine reinforcer, whereas a black stimulus was non-reinforced. When trained from the outset with the discriminative procedure, flies of all ages acquired the discrimination, although the acquisition of PER suppression to the white stimulus was retarded in middle-aged and old flies. The retardation of the acquisition of PER suppression in middle-aged and old flies was replicated in a second study in which the flies received simple conditioning to the white stimulus prior to discrimination training.     

The effects of yeast on sexual behavior in Drosophila melanogaster

The presence of yeast in the diet of virgin Drosophila melanogaster females increases the probability that the females will copulate, although it does not increase their attractiveness. Females raised on yeast move more slowly while being courted by males and elicit a greater number of attempted copulations per minute.     

Drosophila Conditioned Courtship: Two Ways of Testing Memory

In Drosophila, courtship reduction in male flies that have previous experience of courting a mated female is a result of the counterconditioning of an attractive unconditioned stimulus (US)—the aphrodisiac—which becomes an aversive conditioned stimulus (CS) after being paired with an aversive US—the antiaphrodisiac. In a retention test with a virgin female lacking the antiaphrodisiac, males retain a lower level of courtship for 3 hr after training. However, a measure of courtship suppression, the learning index (LI), decreases significantly after only 1 hr. In contrast, in the retraining test with a mated female, the LI shows no decrease for 8 hr but falls below significance 16 hr after training. These results are discussed in terms of the transfer of training. Nonspecific transfer and nonassociative behavioral modifications play little, if any, role in the transfer of training. The retraining test is recommended as a new protocol for studying conditioned courtship. According to the model proposed here, in tests with a virgin female, the duration of memory retention is limited by the retention of the direct association between the CS and the aversive motivational system or by the retention of an internal representation of the US. In retraining tests, the CS–US association seems to be the only factor involved in transfer 3 or more hours after training.     

Induced variation of serotonin in Drosophila melanogaster and its relation to learning performance

In insects the level of serotonin can be modified through the alimentary tract. Drosophila fed on chemically defined media to which has been added a synthesis inhibitor or an immediate precursor of this neurotransmitter have their serotonin level decreased or increased, respectively. (A 4- to 5-day treatment is optimal). Such treated flies, if tested in a procedure of Pavlovian conditioning, have different learning performances. With the same duration of starvation, nearly the same threshold for sucrose solutions, but different amounts of serotonin, the flies (blindly trained) showed conditioned inhibition of the tarsal (or proboscis-extension) reflex according to the amount of serotonin. The highest level of this biogenic amine gave the best performance.     

Ultrastructural plasticity associated with hippocampal-dependent learning: a meta-analysis

In order to develop a profile of how individual synapses in the hippocampal formation alter their structure following learning experience, a meta-analysis synthesized the available literature on morphological change following hippocampal-dependent learning. Analysis of the 132 calculated effect sizes suggest a consistent profile of morphological change in the hippocampus following learning experience. Across the hippocampal formation, dendritic complexity, spine density, and the size of perforated postsynaptic densities showed consistent increases following training. Both the density of synapses in general and perforated synapses in particular showed unique responses to training, depending on the duration of training and/or different cell layers of the hippocampal formation. Most importantly, it seems that this profile, while consistent, is small and specific--only a select few of the morphological parameters typically measured in anatomical studies of plasticity showed significant change following training. Collectively, these data suggest that the distinct electrophysiological properties of neocortical versus hippocampal synapses may be at least partially mediated by distinct morphological cascades. That is, on the basis of theory, and with the support of the current data, it seems that synaptogenesis correlates with enduring neocortical plasticity, while structural changes correlate with more transient hippocampal plasticity. To be able to state these conclusions with conviction, however, more data are needed in several key areas for continued pursuit of the morphological correlates of hippocampal-dependent learning.     

Mapping of the Anatomical Circuit of CaM Kinase-Dependent Courtship Conditioning in Drosophila

Globally inhibiting CaM kinase activity in Drosophila, using a variety of genetic techniques, disrupts associative memory yet leaves visual and chemosensory perception intact. These studies implicate CaM kinase in the plastic processes underlying learning and memory but do not identify the neural circuitry that specifies the behavior. In this study, we use the GAL4/UAS binary expression system to define areas of the brain that require CaM kinase for modulation of courtship conditioning. The CaM kinase-dependent neurons that determine the response to the mated female during conditioning and those involved in formation and expression of memory were found to be located in distinct areas of the brain. This supports the idea that courtship conditioning results in two independent behavioral modifications: a decrement in courtship during the conditioning period and an associative memory of conditioning. This study has allowed us for the first time to genetically determine the circuit of information flow for a memory process in Drosophila. The map we have generated dissects the behavior into multiple components and will provide tools that allow both molecular and electrophysiological access to this circuit.     

Excitatory conditioning of individual Drosophila melanogaster

Testing individual animals from a heterogenic population of Drosophila melanogaster, we demonstrate conditioning of the proboscis extension reflex. The presentation of paired (conditioning) stimuli produced (a) an increase in the average number of conditioned responses over trials, (b) measured differences in performance levels among individual subjects, and (c) a sex difference, with more males conditioned than females, and those that did did so more quickly. The presentation of unpaired (control) stimuli produced significantly lower average levels of acquisition responding and a change in the distribution of individual response patterns. Neither central excitatory state nor sensitization induced by the conditioned or unconditioned stimuli directly affected the conditioned response, whereas unconditioned stimulus preexposure adversely affected performance levels. Presenting the unpaired (extinction) stimuli after conditioning produced less of a decline in responding than did an extinction procedure with removal of the unconditioned stimulus. With the ability to identify individual differences in acquisition and extinction patterns, and given the relatively large number of individuals that can be tested simultaneously on the automated stimulation apparatus, it is now possible to make precise behavioral measurements on samples large enough for the behavior-genetic analysis of D. melanogaster with conditioning as the phenotype.     

Selection for conditionability in Drosophila melanogaster.

Successful bidirectional selective breeding of Drosophila melanogaster for excitatory conditionability is reported, using the reliable measures of individual differences (first described by Holliday & Hirsch, 1984, 1986a, 1986b) to test 1,324 animals. Bidirectional selective breeding for good and poor conditioning has produced, respectively, one population in which the percentage of animals showing good conditioning has increased over 25 generations from 19% to 77% and another in which it has decreased over 23 generations to 0%-4%. No increase in a measure of sensitization induced by an unconditioned stimulus (the central excitatory state) accompanied the increase in the percentage of good conditioners in the population selected for good conditioning, whose level of conditioned responding exceeds that of the sensitization measure.     

Selection for central excitation in Drosophila melanogaster

In two populations of Drosophila melanogaster, bidirectional selection and single-pair matings for high and low expression of central excitatory state (CES) succeeded in producing from one a high, but not a low, CES line, and from the other a low, but not a high, CES line. Compared with results from the selection studies of the blow fly, Phormia regina, in which one major gene correlate of CES has been found, the results from this study suggest that in D. melanogaster there are several gene correlates of CES, opposite alleles of some of which have become fixed in the divergent lines.     

Learned suppression of photopositive tendencies in Drosophila melanogaster

A task was designed to teach individual flies to avoid a lighted area after they had displayed an initial preference for it. The flies walked in a T-maze and chose between a lighted and a darkened alley leading, respectively, to a lighted and a darkened vial. Flies that were photopositive on a first trial were subjected to an aversive stimulus (a filter paper inserted into the lighted vial and wetted with a quinine solution), and they performed 16 training trials; they learned to avoid the lighted vial. The flies trained with water instead of quinine in the lighted vial still display avoidance of the lighted vial, but to a lesser extent. The flies trained with a dry filter paper in the lighted vial did not show any increase in avoidance during training. Like the flies trained with no quinine at all, those trained to avoid the lighted vial under a partial reinforcement condition (one half of the trials with quinine, the other half with a dry vial) did not master the task. Finally, removal of the quinine after an avoidance acquisition criterion was reached resulted in an extinction process.     

Conditioned suppression of proboscis extension in Drosophila melanogaster

Food-deprived Drosophila melanogaster extend their proboscises following sucrose stimulation of the front tarsi (the proboscis extension reflex). Médioni and Vaysse (1975) reported that the inhibition of this response can be conditioned over trials if such proboscis extensions are punished by applying an aversive stimulus to the foreleg tarsi. In this study, Médioni and Vaysse's basic observations of conditioning were replicated, with a different strain of flies and a modified conditioning apparatus.     

Behavioral manipulation of retrieval in a spatial memory task for Drosophila melanogaster

A paradigm for operant conditioning of freely walking single Drosophila flies has been described previously. A fly can be conditioned to avoid one side of a small test chamber if the chamber is heated whenever the fly enters this side. In a subsequent memory test without heat the fly continues to avoid the previously heat-associated side. In this experimental design one cannot exclude that flies mark the heated side by an odor that they subsequently avoid during the test. As a final proof for associative learning in the present experiment, flies are trained in one chamber and tested for learning in another, similar one. Handling in the transfer experiment interferes with memory display, even if the fly is returned to the old chamber instead of a new one. Memory can be reactivated, however, by subjecting the fly to an additional brief training (priming), which is too short to establish significant learning in naive flies. For efficient priming, heat has to be applied to the same side as during training in the old chamber. Only then the fly subsequently shows a side preference and avoids the side of the new chamber, which in the old one had been associated with heat. The two chambers are similar but not identical The transfer experiment therefore raises the question as to what the flies use as spatial reference during training and test. In the light, they can be shown to orientate according to visual landmarks associated with the chamber. In complete darkness, where training and memory scores do not differ from those in the light, they are assumed to use a combination of tactile and idiothetic information for orienting.     

Stress affects dopaminergic signaling pathways in Drosophila melanogaster

Behaviors modulated by dopamine appear to be conserved across species. In the model system Drosophila melanogaster, as in mammals, dopamine modulates female sexual receptivity, a simple form of learning and responses to drugs of abuse. Synthesis, reuptake and binding of dopamine are also evolutionarily conserved. Since stress has been shown to affect dopaminergic signaling pathways in mammals, we investigated the consequences of exposure to diverse stressors on dopaminergic physiology in the fruit fly, D. melanogaster. Animals were exposed to a metabolic stress (starvation), an oxidative stress (via the superoxide anion generator paraquat) or a mechanical stress (gentle vortexing). Sexual maturity, reproductive status, gender and type of stress differentially affected survival. The stress paradigms also resulted in alterations in the activity of tyrosine hydroxylase, the rate-limiting enzyme in dopamine biosynthesis. Exposure to these stressors perturbed female sexual receptivity and ovarian development, which are modulated by dopamine, suggesting that dopaminergic physiology is affected as a consequence of stress. Transgenic Drosophila with reduced levels of neuronal dopamine displayed an altered response to these stressors, suggesting that, as in mammals, dopamine is a key element in the stress response.     

Mushroom Bodies Suppress Locomotor Activity in Drosophila melanogaster

Locomotor activity of single, freely walking flies in small tubes is analyzed in the time domain of several hours. To assess the influence of the mushroom bodies on walking activity, three independent noninvasive methods interfering with mushroom body function are applied: chemical ablation of the mushroom body precursor cells; a mutant affecting Kenyon cell differentiation (mushroom body miniature¹); and the targeted expression of the catalytic subunit of tetanus toxin in subsets of Kenyon cells. All groups of flies with mushroom body defects show an elevated level of total walking activity. This increase is attributable to the slower and less complete attenuation of activity during the experiment. Walking activity in normal and mushroom body-deficient flies is clustered in active phases (bouts) and rest periods (pauses). Neither the initiation nor the internal structure, but solely the termination of bouts seems to be affected by the mushroom body defects. How this finding relates to the well-documented role of the mushroom bodies in olfactory learning and memory remains to be understood.