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Bertram Gerber

Publications and source records attributed to Bertram Gerber.

13 recordsLinked to original sources

Outcome expectations drive learned behaviour in larval Drosophila.

Why does Pavlov's dog salivate? In response to the tone, or in expectation of food? While in vertebrates behaviour can be driven by expected outcomes, it is unknown whether this is true for non-vertebrates as well. We find that, in the Drosophila larva, odour memories are expressed behaviourally only if animals can expect a positive outcome from doing so. The expected outcome of tracking down an odour is determined by comparing the value of the current situation with the value of the memory for that odour. Memory is expressed behaviourally only if the expected outcome is positive. This uncovers a hitherto unrecognized evaluative processing step between an activated memory trace and behaviour control, and argues that learned behaviour reflects the pursuit of its expected outcome. Shown in a system with a simple brain, an apparently cognitive process like representing the expected outcome of behaviour seems to be a basic feature of behaviour control.

Animals↗

The Drosophila larva as a model for studying chemosensation and chemosensory learning: a review.

Understanding the relationship between brain and behavior is the fundamental challenge in neuroscience. We focus on chemosensation and chemosensory learning in larval Drosophila and review what is known about its molecular and cellular bases. Detailed analyses suggest that the larval olfactory system, albeit much reduced in cell number, shares the basic architecture, both in terms of receptor gene expression and neuronal circuitry, of its adult counterpart as well as of mammals. With respect to the gustatory system, less is known in particular with respect to processing of gustatory information in the central nervous system, leaving generalizations premature. On the behavioral level, a learning paradigm for the association of odors with food reinforcement has been introduced. Capitalizing on the knowledge of the chemosensory pathways, we review the first steps to reveal the genetic and cellular bases of olfactory learning in larval Drosophila. We argue that the simplicity of the larval chemosensory system, combined with the experimental accessibility of Drosophila on the genetic, electrophysiological, cellular, and behavioral level, makes this system suitable for an integrated understanding of chemosensation and chemosensory learning.

Animals↗

Light-induced activation of distinct modulatory neurons triggers appetitive or aversive learning in Drosophila larvae.

During classical conditioning, a positive or negative value is assigned to a previously neutral stimulus, thereby changing its significance for behavior. If an odor is associated with a negative stimulus, it can become repulsive. Conversely, an odor associated with a reward can become attractive. By using Drosophila larvae as a model system with minimal brain complexity, we address the question of which neurons attribute these values to odor stimuli. In insects, dopaminergic neurons are required for aversive learning, whereas octopaminergic neurons are necessary and sufficient for appetitive learning. However, it remains unclear whether two independent neuronal populations are sufficient to mediate such antagonistic values. We report the use of transgenically expressed channelrhodopsin-2, a light-activated cation channel, as a tool for optophysiological stimulation of genetically defined neuronal populations in Drosophila larvae. We demonstrate that distinct neuronal populations can be activated simply by illuminating the animals with blue light. Light-induced activation of dopaminergic neurons paired with an odor stimulus induces aversive memory formation, whereas activation of octopaminergic/tyraminergic neurons induces appetitive memory formation. These findings demonstrate that antagonistic modulatory subsystems are sufficient to substitute for aversive and appetitive reinforcement during classical conditioning.

Animals↗

Olfactory learning and behaviour are 'insulated' against visual processing in larval Drosophila.

We investigate the organization of behaviour across sensory modalities, using larval Drosophila melanogaster. We ask whether olfactory learning and behaviour are affected by visual processing. We find that: (1) Visual choice does not affect concomitant odour choice. (2) Visual context does not influence odour learning, nor do changes of visual context between training and test affect retrieval of odour memory. (3) Larvae cannot solve a biconditional discrimination task, despite generally permissive conditions. In this task, larvae are required to establish conditional associations: in light, one odour is rewarded and the other one is not, whereas in dark the opposite contingency is established. After such training, choice between the two odours is equal under light and dark testing conditions, suggesting that larvae do not establish odour memories specifically for one visual context only. Together, these data suggest that, in larval Drosophila, olfactory learning and behaviour are 'insulated' against visual processing.

Animals↗

The carrot, not the stick: appetitive rather than aversive gustatory stimuli support associative olfactory learning in individually assayed Drosophila larvae.

The ability to learn is universal among animals; we investigate associative learning between odors and "tastants" in larval Drosophila melanogaster. As biologically important gustatory stimuli, like sugars, salts, or bitter substances have many behavioral functions, we investigate not only their reinforcing function, but also their response-modulating and response-releasing function. Concerning the response-releasing function, larvae are attracted by fructose and repelled by sodium chloride and quinine; also, fructose increases, but salt and quinine suppress feeding. However, none of these stimuli has a nonassociative, modulatory effect on olfactory choice behavior. Finally, only fructose but neither salt nor quinine has a reinforcing effect in associative olfactory learning. This implies that the response-releasing, response-modulating and reinforcing functions of these tastants are dissociated on the behavioral level. These results open the door to analyze how this dissociation is brought about on the cellular and molecular level; this should be facilitated by the cellular simplicity and genetic accessibility of the Drosophila larva.

Animals↗

Experimental psychology: event timing turns punishment to reward.

Can relief from pain be a pleasure? If so, noxious events should--despite their typically aversive effects--also have a 'rewarding' after-effect. Through training fruitflies by using an electric shock paired with an odour, we show here that the shock can condition either avoidance of this odour or approach to it. These opposing behaviours depend on the relative timing of the shock and odour presentations during training, and indicate that a shock can act as either an aversive reinforcer or an appetitive one.

Animals↗

An engram found? Evaluating the evidence from fruit flies.

Is it possible to localize a memory trace to a subset of cells in the brain? If so, it should be possible to show: first, that neuronal plasticity occurs in these cells. Second, that neuronal plasticity in these cells is sufficient for memory. Third, that neuronal plasticity in these cells is necessary for memory. Fourth, that memory is abolished if these cells cannot provide output during testing. And fifth, that memory is abolished if these cells cannot receive input during training. With regard to olfactory learning in flies, we argue that the notion of the olfactory memory trace being localized to the Kenyon cells of the mushroom bodies is a reasonable working hypothesis.

Animals↗

The effect of cumulative experience on the use of elemental and configural visual discrimination strategies in honeybees.

We addressed the question of whether the amount of individual experience determines the use of elemental or configural visual discrimination strategies in free-flying honeybees Apis mellifera. We trained bees to fly into a Y-maze to collect sucrose solution on a rewarded stimulus presented in one of the arms of the maze. Stimuli were colour disks, violet (V), green (G) or yellow (Y), which were of equal psychophysical salience for honeybees. Training followed an A+, BC+ design, followed by an AC versus BC test. Training consisted of 6 (3 A+ and 3 BC+), 20 (10 A+ and 10 BC+) or 40 (20 A+ and 20 BC+) acquisition trials. Elemental models of compound processing predict a preference for the non-trained stimulus AC while configural models predict a preference for the trained stimulus BC. Our results show that increasing the number of acquisition trials results in a change of the internal representation of stimuli. After six training trials, bees favoured an elemental strategy and preferred AC to BC during the tests. Generally, increasing the number of training trials resulted in an increase of the choice of BC. Thus, short training favoured processing of the compound as the sum of its elements (elemental account) while long training favoured its processing as being different from the sum of its elements (configural account). Additionally, we observed that the change in stimulus processing was also influenced by stimulus similarity. Colour perceptual similarity favoured configural processing with increasing experience.

Animals↗

Hydroxyurea-induced partial mushroom body ablation does not affect acquisition and retention of olfactory differential conditioning in honeybees.

The mushroom bodies (MBs), a paired structure in the insect brain, play a major role in storing and retrieving olfactory memories. We tested whether olfactory learning and odor processing is impaired in honeybees in which MB subunits were partially ablated. Using hydroxyurea (HU) to selectively kill proliferating cells, we created honeybees with varying degrees of MB lesions. Three-dimensional reconstructions of brains were generated to analyze the drug-induced morphological changes. These reconstructions show that, with few exceptions, only the MBs were affected by the drug, while other brain areas remained morphometrically intact. Typically, lesions affected only the MB in one hemisphere of the brain. To preclude HU-induced physiologic deficits in the antennal lobe (AL) affecting olfactory learning, we measured the responses to odors in the AL using an in vivo calcium imaging approach. The response patterns did not differ between the AL of intact versus ablated brain sides within respective specimens. We, therefore, carried out side-specific classical discriminative olfactory conditioning of the proboscis extension reflex (PER) with control bees and with HU-treated bees with or without MB ablations. All experimental groups learned equally to discriminate and respond to a rewarded (CS+) but not to an unrewarded (CS-) conditioned stimulus during acquisition and retention tests. Thus, our results indicate that partial MB lesions do not affect this form of elemental olfactory learning.

Animals↗

Olfactory learning in individually assayed Drosophila larvae.

Insect and mammalian olfactory systems are strikingly similar. Therefore, Drosophila can be used as a simple model for olfaction and olfactory learning. The brain of adult Drosophila, however, is still complex. We therefore chose to work on the larva with its yet simpler but adult-like olfactory system and provide evidence for olfactory learning in individually assayed Drosophila larvae. We developed a differential conditioning paradigm in which odorants are paired with positive ("+" fructose) or negative ("-" quinine or sodium chloride) gustatory reinforcers. Test performance of individuals from two treatment conditions is compared-one received odorant A with the positive reinforcer and odorant B with a negative reinforcer (A+/B-); animals from the other treatment condition were trained reciprocally (A-/B+). During test, differences in choice between A and B of individuals having undergone either A+/B- or A-/B+ training therefore indicate associative learning. We provide such evidence for both combinations of reinforcers; this was replicable across repetitions, laboratories, and experimenters. We further show that breaks improve performance, in accord with basic principles of associative learning. The present individual assay will facilitate electrophysiological studies, which necessarily use individuals. As such approaches are established for the larval neuromuscular synapse, but not in adults, an individual larval learning paradigm will serve to link behavioral levels of analysis to synaptic physiology.

Animals↗

Olfactory blocking and odorant similarity in the honeybee.

Blocking occurs when previous training with a stimulus A reduces (blocks) subsequent learning about a stimulus B, when A and B are trained in compound. The question of whether blocking exists in olfactory conditioning of proboscis extension reflex (PER) in honeybees is under debate. The last published accounts on blocking in honeybees state that blocking occurs when odors A and B are similar (the "similarity hypothesis"). We have tested this hypothesis using four odors (1-octanol, 1-nonanol, eugenol, and limonene) chosen on the basis of their chemical and physiological similarity (experiment 1). We established a generalization matrix that measured perceptual similarity. Bees in the "block group" were first trained with an odor A and, in the second phase, with the mixture AB. Bees in the "novel group" (control group) were first trained with an odor N and, in the second phase, with the mixture AB. After conditioning, bees in both groups were tested for their response to B. We assayed all 24 possible combinations for the four odors standing for A, B, and N. We found blocking in four cases, augmentation in two cases, and no difference in 18 cases; odor similarity could not account for these results. We also repeated the experiments with those six odor combinations that gave rise to the similarity hypothesis (experiment 2: 1-hexanol, 1-octanol, geraniol) and found augmentation in one and no effect in five cases. Thus, blocking is not a consistent phenomenon, nor does it depend on odor similarity.

Analysis of Variance↗

A role for Synapsin in associative learning: the Drosophila larva as a study case.

Synapsins are evolutionarily conserved, highly abundant vesicular phosphoproteins in presynaptic terminals. They are thought to regulate the recruitment of synaptic vesicles from the reserve pool to the readily-releasable pool, in particular when vesicle release is to be maintained at high spiking rates. As regulation of transmitter release is a prerequisite for synaptic plasticity, we use the fruit fly Drosophila to ask whether Synapsin has a role in behavioral plasticity as well; in fruit flies, Synapsin is encoded by a single gene (syn). We tackled this question for associative olfactory learning in larval Drosophila by using the deletion mutant syn(97CS), which had been backcrossed to the Canton-S wild-type strain (CS) for 13 generations. We provide a molecular account of the genomic status of syn(97CS) by PCR and show the absence of gene product on Western blots and nerve-muscle preparations. We found that olfactory associative learning in syn(97CS) larvae is reduced to approximately 50% of wild-type CS levels; however, responsiveness to the to-be-associated stimuli and motor performance in untrained animals are normal. In addition, we introduce two novel behavioral control procedures to test stimulus responsiveness and motor performance after "sham training." Wild-type CS and syn(97CS) perform indistinguishably also in these tests. Thus, larval Drosophila can be used as a case study for a role of Synapsin in associative learning.

Animals↗