Olfactory learning in Drosophila

Animals are able to form associative memories and benefit from past experience. In classical conditioning an animal is trained to associate an initially neutral stimulus by pairing it with a stimulus that triggers an innate response. The neutral stimulus is commonly referred to as conditioned stimulus (CS) and the reinforcing stimulus as unconditioned stimulus (US). The underlying neuronal mechanisms and structures are an intensely investigated topic. The fruit fly Drosophila melanogaster is a prime model animal to investigate the mechanisms of learning. In this thesis we propose fundamental circuit motifs that explain aspects of aversive olfactory learning as it is observed in the fruit fly. Changing parameters of the learning paradigm affects the behavioral outcome in different ways. The relative timing between CS and US affects the hedonic value of the CS. Reversing the order changes the behavioral response from conditioned avoidance to conditioned approach. We propose a timing-dependent biochemical reaction cascade, which can account for this phenomenon. In addition to form odor-specific memories, flies are able to associate a specific odor intensity. In aversive olfactory conditioning they show less avoidance to lower and higher intensities of the same odor. However the layout of the first two olfactory processing layers does not support this kind of learning due to a nested representation of odor intensity. We propose a basic circuit motif that transforms the nested monotonic intensity representation to a non-monotonic representation that supports intensity specific learning. Flies are able to bridge a stimulus free interval between CS and US to form an association. It is unclear so far where the stimulus trace of the CS is represented in the fly's nervous system. We analyze recordings from the first three layers of olfactory processing with an advanced machine learning approach. We argue that third order neurons are likely to harbor the stimulus trace

Event Timing in Associative Learning

Associative learning relies on event timing. Fruit flies for example, once trained with an odour that precedes electric shock, subsequently avoid this odour (punishment learning); if, on the other hand the odour follows the shock during training, it is approached later on (relief learning). During training, an odour-induced Ca++ signal and a shock-induced dopaminergic signal converge in the Kenyon cells, synergistically activating a Ca++-calmodulin-sensitive adenylate cyclase, which likely leads to the synaptic plasticity underlying the conditioned avoidance of the odour. In Aplysia, the effect of serotonin on the corresponding adenylate cyclase is bi-directionally modulated by Ca++, depending on the relative timing of the two inputs. Using a computational approach, we quantitatively explore this biochemical property of the adenylate cyclase and show that it can generate the effect of event timing on associative learning. We overcome the shortage of behavioural data in Aplysia and biochemical data in Drosophila by combining findings from both systems

Olfactory learning in Drosophila

Animals are able to form associative memories and benefit from past experience. In classical conditioning an animal is trained to associate an initially neutral stimulus by pairing it with a stimulus that triggers an innate response. The neutral stimulus is commonly referred to as conditioned stimulus (CS) and the reinforcing stimulus as unconditioned stimulus (US). The underlying neuronal mechanisms and structures are an intensely investigated topic. The fruit fly Drosophila melanogaster is a prime model animal to investigate the mechanisms of learning. In this thesis we propose fundamental circuit motifs that explain aspects of aversive olfactory learning as it is observed in the fruit fly. Changing parameters of the learning paradigm affects the behavioral outcome in different ways. The relative timing between CS and US affects the hedonic value of the CS. Reversing the order changes the behavioral response from conditioned avoidance to conditioned approach. We propose a timing-dependent biochemical reaction cascade, which can account for this phenomenon. In addition to form odor-specific memories, flies are able to associate a specific odor intensity. In aversive olfactory conditioning they show less avoidance to lower and higher intensities of the same odor. However the layout of the first two olfactory processing layers does not support this kind of learning due to a nested representation of odor intensity. We propose a basic circuit motif that transforms the nested monotonic intensity representation to a non-monotonic representation that supports intensity specific learning. Flies are able to bridge a stimulus free interval between CS and US to form an association. It is unclear so far where the stimulus trace of the CS is represented in the fly's nervous system. We analyze recordings from the first three layers of olfactory processing with an advanced machine learning approach. We argue that third order neurons are likely to harbor the stimulus trace

A model for non-monotonic intensity coding

Peripheral neurons of most sensory systems increase their response with increasing stimulus intensity. Behavioural responses, however, can be specific to some intermediate intensity level whose particular value might be innate or associatively learned. Learning such a preference requires an adjustable transformation from a monotonic stimulus representation at the sensory periphery to a non-monotonic representation for the motor command. How do neural systems accomplish this task? We tackle this general question focusing on odour intensity learning in the fruit fly; whose first- and second-order olfactory neurons show monotonic stimulus response curves. Nevertheless, flies form associative memories specific to particular trained odour intensities. Thus, downstream of the first two olfactory processing layers, odour intensity must be re-coded to enable intensity-specific associative learning. We present a minimal, feed-forward, three-layer circuit, which implements the required transformation by combining excitation, inhibition, and, as a decisive third element, homeostatic plasticity. Key features of this circuit motif are consistent with the known architecture and physiology of the fly olfactory system, whereas alternative mechanisms are either not composed of simple, scalable building blocks or not compatible with physiological observations. The simplicity of the circuit and the robustness of its function under parameter changes make this computational motif an attractive candidate for tuneable non-monotonic intensity coding

Calcium in Kenyon Cell Somata as a Substrate for an Olfactory Sensory Memory in Drosophila

Animals can form associations between temporally separated stimuli. To do so, the nervous system has to retain a neural representation of the first stimulus until the second stimulus appears. The neural substrate of such sensory stimulus memories is unknown. Here, we search for a sensory odor memory in the insect olfactory system and characterize odorant-evoked Ca2+ activity at three consecutive layers of the olfactory system in Drosophila: in olfactory receptor neurons (ORNs) and projection neurons (PNs) in the antennal lobe, and in Kenyon cells (KCs) in the mushroom body. We show that the post-stimulus responses in ORN axons, PN dendrites, PN somata, and KC dendrites are odor-specific, but they are not predictive of the chemical identity of past olfactory stimuli. However, the post-stimulus responses in KC somata carry information about the identity of previous olfactory stimuli. These findings show that the Ca2+ dynamics in KC somata could encode a sensory memory of odorant identity and thus might serve as a basis for associations between temporally separated stimuli

Event Timing in Associative Learning: From Biochemical Reaction Dynamics to Behavioural Observations

Associative learning relies on event timing. Fruit flies for example, once trained with an odour that precedes electric shock, subsequently avoid this odour (punishment learning); if, on the other hand the odour follows the shock during training, it is approached later on (relief learning). During training, an odour-induced Ca++ signal and a shock-induced dopaminergic signal converge in the Kenyon cells, synergistically activating a Ca++-calmodulin-sensitive adenylate cyclase, which likely leads to the synaptic plasticity underlying the conditioned avoidance of the odour. In Aplysia, the effect of serotonin on the corresponding adenylate cyclase is bi-directionally modulated by Ca++, depending on the relative timing of the two inputs. Using a computational approach, we quantitatively explore this biochemical property of the adenylate cyclase and show that it can generate the effect of event timing on associative learning. We overcome the shortage of behavioural data in Aplysia and biochemical data in Drosophila by combining findings from both systems

Search for heavy resonances decaying to a top quark and a bottom quark in the lepton+jets final state in proton–proton collisions at 13 TeV

info:eu-repo/semantics/publishe

Optimasi Portofolio Resiko Menggunakan Model Markowitz MVO Dikaitkan dengan Keterbatasan Manusia dalam Memprediksi Masa Depan dalam Perspektif Al-Qur`an

Risk portfolio on modern finance has become increasingly technical, requiring the use of sophisticated mathematical tools in both research and practice. Since companies cannot insure themselves completely against risk, as human incompetence in predicting the future precisely that written in Al-Quran surah Luqman verse 34, they have to manage it to yield an optimal portfolio. The objective here is to minimize the variance among all portfolios, or alternatively, to maximize expected return among all portfolios that has at least a certain expected return. Furthermore, this study focuses on optimizing risk portfolio so called Markowitz MVO (Mean-Variance Optimization). Some theoretical frameworks for analysis are arithmetic mean, geometric mean, variance, covariance, linear programming, and quadratic programming. Moreover, finding a minimum variance portfolio produces a convex quadratic programming, that is minimizing the objective function ðð¥with constraintsð ð 𥠥 ðandð´ð¥ = ð. The outcome of this research is the solution of optimal risk portofolio in some investments that could be finished smoothly using MATLAB R2007b software together with its graphic analysis