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Excitatory Local Interneurons Enhance Tuning of Sensory Information
Assisi, Collins
Stopfer, Mark
Bazhenov, Maxim
PLoS Computational Biology
Neurons in the insect antennal lobe represent odors as spatiotemporal patterns of activity that unfold over multiple time scales. As these patterns unspool they decrease the overlap between odor representations and thereby increase the ability of the olfactory system to discriminate odors. Using a realistic model of the insect antennal lobe we examined two competing components of this process –lateral excitation from local excitatory interneurons, and slow inhibition from local inhibitory interneurons. We found that lateral excitation amplified differences between representations of similar odors by recruiting projection neurons that did not receive direct input from olfactory receptors. However, this increased sensitivity also amplified noisy variations in input and compromised the ability of the system to respond reliably to multiple presentations of the same odor. Slow inhibition curtailed the spread of projection neuron activity and increased response reliability. These competing influences must be finely balanced in order to decorrelate odor representations.
Author Summary
The antennal lobe of insects and the olfactory bulb of vertebrates represent the first centers of the olfactory system where information about odor properties can be reorganized and optimized for further processing. Complex excitatory and inhibitory synaptic interactions within the antennal lobe and the olfactory bulb alter the responses of the principal neurons throughout the duration of the odor stimulation. These dynamic changes progressively increase the difference between firing patterns evoked by structurally similar odors, potentially helping the animal distinguish one odor from another. However, this process, called odor decorrelation, appears to oppose another important goal of olfactory processing, to minimize the inevitable noisy variations in representations of the same odor encountered under different environmental conditions; such variations could potentially lead to misclassification. It remains an interesting mystery how olfactory circuitry can solve these two seemingly contradictory goals as they process olfactory stimuli: first, separating different but chemically similar odors (sensitivity, capacity); and second, identifying representations of the same odor in a noisy environment (reliability). Our results suggest a balance between inhibitory and excitatory connections mediated by local antennal lobe interneurons enhances the decorrelation of similar odors while keeping the representation robust in the presence of noise.
Related Articles
Synaptic Learning Rules and Sparse Coding in a Model Sensory System
Finelli, Luca A.
Haney, Seth
Bazhenov, Maxim
Stopfer, Mark
Sejnowski, Terrence J.
PLoS Computational Biology
Neural circuits exploit numerous strategies for encoding information. Although the functional significance of individual coding mechanisms has been investigated, ways in which multiple mechanisms interact and integrate are not well understood. The locust olfactory system, in which dense, transiently synchronized spike trains across ensembles of antenna lobe (AL) neurons are transformed into a sparse representation in the mushroom body (MB; a region associated with memory), provides a well-studied preparation for investigating the interaction of multiple coding mechanisms. Recordings made in vivo from the insect MB demonstrated highly specific responses to odors in Kenyon cells (KCs). Typically, only a few KCs from the recorded population of neurons responded reliably when a specific odor was presented. Different odors induced responses in different KCs. Here, we explored with a biologically plausible model the possibility that a form of plasticity may control and tune synaptic weights of inputs to the mushroom body to ensure the specificity of KCs' responses to familiar or meaningful odors. We found that plasticity at the synapses between the AL and the MB efficiently regulated the delicate tuning necessary to selectively filter the intense AL oscillatory output and condense it to a sparse representation in the MB. Activity-dependent plasticity drove the observed specificity, reliability, and expected persistence of odor representations, suggesting a role for plasticity in information processing and making a testable prediction about synaptic plasticity at AL-MB synapses.
Author Summary
The way in which the brain encodes, processes, transforms, and stores sensory information is a fundamental question in systems neuroscience. One challenge is to understand how neural oscillations, synchrony, population coding, and sparseness interact in the process of transforming and transferring information. Another question is how synaptic plasticity, the ability of synapses to change their strength, interacts efficiently with these different coding strategies to support learning and information storage. We approached these questions, rarely accessible to direct experimental investigation, in the olfactory system of the locust, a well-studied example. Here, the neurons in the antennal lobe carry neural representations of odor identity using dense, spatially distributed, oscillatory synchronized patterns of neural activity. Odor information cannot be interpreted by considering their activity independently. On the contrary, in the mushroom body—the next processing region, involved in the storage and retrieval of olfactory memories and analogous to the olfactory cortex—odor representations are sparse and carried by more selective neurons. Sparse information coding by ensembles of neurons provides several important advantages including high memory capacity, low overlap between stored objects, and easy information retrieval. How is this sparseness achieved? Here, with a rigorous computational model of the olfactory system, we demonstrate that plasticity at the input afferents to the mushroom body can efficiently mediate the delicate tuning necessary to selectively filter intense sensory input, condensing it to the sparse responses observed in the mushroom body. Our results suggest a general mechanism for plasticity-enabled sparse representations in other sensory systems, such as the visual system. Overall, we illustrate a potential central role for plasticity in the transfer of information across different coding strategies within neural systems.
Related Articles
Temporal Features of Spike Trains in the Moth Antennal Lobe Revealed by a Comparative Time-Frequency Analysis
Capurro, Alberto
Baroni, Fabiano
Kuebler, Linda S.
Kárpáti, Zsolt
Dekker, Teun
Lei, Hong
Hansson, Bill S.
Pearce, Timothy C.
Olsson, Shannon B.
PLoS ONE
The discrimination of complex sensory stimuli in a noisy environment is an immense computational task. Sensory systems often encode stimulus features in a spatiotemporal fashion through the complex firing patterns of individual neurons. To identify these temporal features, we have developed an analysis that allows the comparison of statistically significant features of spike trains localized over multiple scales of time-frequency resolution. Our approach provides an original way to utilize the discrete wavelet transform to process instantaneous rate functions derived from spike trains, and select relevant wavelet coefficients through statistical analysis. Our method uncovered localized features within olfactory projection neuron (PN) responses in the moth antennal lobe coding for the presence of an odor mixture and the concentration of single component odorants, but not for compound identities. We found that odor mixtures evoked earlier responses in biphasic response type PNs compared to single components, which led to differences in the instantaneous firing rate functions with their signal power spread across multiple frequency bands (ranging from 0 to 45.71 Hz) during a time window immediately preceding behavioral response latencies observed in insects. Odor concentrations were coded in excited response type PNs both in low frequency band differences (2.86 to 5.71 Hz) during the stimulus and in the odor trace after stimulus offset in low (0 to 2.86 Hz) and high (22.86 to 45.71 Hz) frequency bands. These high frequency differences in both types of PNs could have particular relevance for recruiting cellular activity in higher brain centers such as mushroom body Kenyon cells. In contrast, neurons in the specialized pheromone-responsive area of the moth antennal lobe exhibited few stimulus-dependent differences in temporal response features. These results provide interesting insights on early insect olfactory processing and introduce a novel comparative approach for spike train analysis applicable to a variety of neuronal data sets.
Related Articles
Characterization and Coding of Behaviorally Significant Odor Mixtures
Riffell, Jeffrey A.
Lei, Hong
Christensen, Thomas A.
Hildebrand, John G.
Current biology : CB
For animals to execute odor-driven behaviors, the olfactory system must process complex odor signals and maintain stimulus identity in the face of constantly changing odor intensities [1–5]. Surprisingly, how the olfactory system maintains identity of complex odors is unclear [6–10]. We took advantage of the plant-pollinator relationship between the Sacred Datura (Datura wrightii) and the moth Manduca sexta [11, 12] to determine how olfactory networks in this insect's brain represent odor mixtures. We combined gas chromatography and neural-ensemble recording in the moth's antennal lobe to examine population codes for the floral mixture and its fractionated components. Although the floral scent of D. wrightii comprises at least 60 compounds, only nine of those elicited robust neural responses. Behavioral experiments confirmed that these nine odorants mediate flower-foraging behaviors, but only as a mixture. Moreover, the mixture evoked equivalent foraging behaviors over a 1000-fold range in dilution, suggesting a singular percept across this concentration range. Furthermore, neural-ensemble recordings in the moth's antennal lobe revealed that reliable encoding of the floral mixture is organized through synchronized activity distributed across a population of glomerular coding units, and this timing mechanism may bind the features of a complex stimulus into a coherent odor percept.
Related Articles
Parallel Representation of Stimulus Identity and Intensity in a Dual Pathway Model Inspired by the Olfactory System of the Honeybee
Schmuker, Michael
Yamagata, Nobuhiro
Nawrot, Martin Paul
Menzel, Randolf
Frontiers in Neuroengineering
The honeybee Apis mellifera has a remarkable ability to detect and locate food sources during foraging, and to associate odor cues with food rewards. In the honeybee's olfactory system, sensory input is first processed in the antennal lobe (AL) network. Uniglomerular projection neurons (PNs) convey the sensory code from the AL to higher brain regions via two parallel but anatomically distinct pathways, the lateral and the medial antenno-cerebral tract (l- and m-ACT). Neurons innervating either tract show characteristic differences in odor selectivity, concentration dependence, and representation of mixtures. It is still unknown how this differential stimulus representation is achieved within the AL network. In this contribution, we use a computational network model to demonstrate that the experimentally observed features of odor coding in PNs can be reproduced by varying lateral inhibition and gain control in an otherwise unchanged AL network. We show that odor coding in the l-ACT supports detection and accurate identification of weak odor traces at the expense of concentration sensitivity, while odor coding in the m-ACT provides the basis for the computation and following of concentration gradients but provides weaker discrimination power. Both coding strategies are mutually exclusive, which creates a tradeoff between detection accuracy and sensitivity. The development of two parallel systems may thus reflect an evolutionary solution to this problem that enables honeybees to achieve both tasks during bee foraging in their natural environment, and which could inspire the development of artificial chemosensory devices for odor-guided navigation in robots.
dual pathway odor coding; mixture coding; antennal lobe; computational model; honeybee foraging
Related Articles
Role of histamine as a putative inhibitory transmitter in the honeybee antennal lobe
Sachse, Silke
Peele, Philipp
Silbering, Ana F
Gühmann, Martin
Galizia, C Giovanni
Frontiers in Zoology
Odors are represented by specific spatio-temporal activity patterns in the olfactory bulb of vertebrates and its insect analogue, the antennal lobe. In honeybees inhibitory circuits in the AL are involved in the processing of odors to shape afferent odor responses. GABA is known as an inhibitory transmitter in the antennal lobe, but not all interneurons are GABAergic. Therefore we sought to analyze the functional role of the inhibitory transmitter histamine for the processing of odors in the honeybee AL.
We optically recorded the representation of odors before, during and after histamine application at the input level (estimated from a compound signal), and at the output level (by selectively measuring the projection neurons). For both, histamine led to a strong and reversible reduction of odor-evoked responses.
We propose that histamine, in addition to GABA, acts as an inhibitory transmitter in the honeybee AL and is therefore likely to play a role in odor processing.
Related Articles
Neuronal Processing of Complex Mixtures Establishes a Unique Odor Representation in the Moth Antennal Lobe
Kuebler, Linda S.
Olsson, Shannon B.
Weniger, Richard
Hansson, Bill S.
Frontiers in Neural Circuits
Animals typically perceive natural odor cues in their olfactory environment as a complex mixture of chemically diverse components. In insects, the initial representation of an odor mixture occurs in the first olfactory center of the brain, the antennal lobe (AL). The contribution of single neurons to the processing of complex mixtures in insects, and in particular moths, is still largely unknown. Using a novel multicomponent stimulus system to equilibrate component and mixture concentrations according to vapor pressure, we performed intracellular recordings of projection and interneurons in an attempt to quantitatively characterize mixture representation and integration properties of single AL neurons in the moth. We found that the fine spatiotemporal representation of 2–7 component mixtures among single neurons in the AL revealed a highly combinatorial, non-linear process for coding host mixtures presumably shaped by the AL network: 82% of mixture responding projection neurons and local interneurons showed non-linear spike frequencies in response to a defined host odor mixture, exhibiting an array of interactions including suppression, hypoadditivity, and synergism. Our results indicate that odor mixtures are represented by each cell as a unique combinatorial representation, and there is no general rule by which the network computes the mixture in comparison to single components. On the single neuron level, we show that those differences manifest in a variety of parameters, including the spatial location, frequency, latency, and temporal pattern of the response kinetics.
odor processing; network; intracellular; electrophysiological recording; single cell
Related Articles
Topological Reorganization of Odor Representations in the Olfactory Bulb
Yaksi, Emre
Judkewitz, Benjamin
Friedrich, Rainer W
PLoS Biology
Odors are initially represented in the olfactory bulb (OB) by patterns of sensory input across the array of glomeruli. Although activated glomeruli are often widely distributed, glomeruli responding to stimuli sharing molecular features tend to be loosely clustered and thus establish a fractured chemotopic map. Neuronal circuits in the OB transform glomerular patterns of sensory input into spatiotemporal patterns of output activity and thereby extract information about a stimulus. It is, however, unknown whether the chemotopic spatial organization of glomerular inputs is maintained during these computations. To explore this issue, we measured spatiotemporal patterns of odor-evoked activity across thousands of individual neurons in the zebrafish OB by temporally deconvolved two-photon Ca2+ imaging. Mitral cells and interneurons were distinguished by transgenic markers and exhibited different response selectivities. Shortly after response onset, activity patterns exhibited foci of activity associated with certain chemical features throughout all layers. During the subsequent few hundred milliseconds, however, MC activity was locally sparsened within the initial foci in an odor-specific manner. As a consequence, chemotopic maps disappeared and activity patterns became more informative about precise odor identity. Hence, chemotopic maps of glomerular input activity are initially transmitted to OB outputs, but not maintained during pattern processing. Nevertheless, transient chemotopic maps may support neuronal computations by establishing important synaptic interactions within the circuit. These results provide insights into the functional topology of neural activity patterns and its potential role in circuit function.
Author Summary
Many sensory brain areas contain topographic maps where the physical location of neuronal activity contains information about a stimulus feature. In the first central processing center of the olfactory pathway, the olfactory bulb, chemically distinct odors often elicit spatially segregated input activity so that general chemical features are initially represented in a topographic fashion. It is, however, unclear whether this "chemotopic" organization of odor representations is maintained at subsequent stages of odor processing. To address this question, we visualized activity patterns across thousands of individual neurons in the intact olfactory bulb of zebrafish over time using two-photon calcium imaging. Our results demonstrate that odor-evoked activity across the output neurons of the olfactory bulb is chemotopically organized shortly after stimulus onset but becomes more widely distributed during the subsequent few hundred milliseconds of the response. This reorganization of olfactory bulb output activity is most likely mediated by inhibitory feedback and reduces the redundancy in activity patterns evoked by related stimuli. These results indicate that topographically organized activity maps in the olfactory bulb are not maintained during information processing, but contribute to the function of local circuits.
Two-photon calcium imaging in the zebrafish olfactory bulb reveals that mitral cells show more selective responses to odors than interneurons, and odor-evoked firing patterns of populations of mitral cells evolve over hundreds of milliseconds to become more distinct for different odors, thus providing more information about odor identity.
Related Articles
Spatio-temporal activity patterns of odor-induced synchronized potentials revealed by voltage-sensitive dye imaging and intracellular recording in the antennal lobe of the cockroach
Watanabe, Hidehiro
Ai, Hiroyuki
Yokohari, Fumio
Frontiers in Systems Neuroscience
In animals, odor qualities are represented as both spatial activity patterns of glomeruli and temporal patterns of synchronized oscillatory signals in the primary olfactory centers. By optical imaging of a voltage-sensitive dye (VSD) and intracellular recording from secondary olfactory interneurons, we examined possible neural correlates of the spatial and temporal odor representations in the primary olfactory center, the antennal lobe (AL), of the cockroach Periplaneta americana. Voltage-sensitive dye imaging revealed that all used odorants induced odor-specific temporal patterns of depolarizing potentials in specific combinations of anterior glomeruli of the AL. The depolarizing potentials evoked by different odorants were temporally synchronized across glomeruli and were termed "synchronized potentials." These observations suggest that odor qualities are represented by spatio-temporal activity patterns of the synchronized potentials across glomeruli. We also performed intracellular recordings and stainings from secondary olfactory interneurons, namely projection neurons and local interneurons. We analyzed the temporal structures of enanthic acid-induced action potentials of secondary olfactory interneurons using simultaneous paired intracellular recording from two given neurons. Our results indicated that the multiple local interneurons synchronously fired in response to the olfactory stimulus. In addition, all stained enanthic acid-responsive projection neurons exhibited dendritic arborizations within the glomeruli where the synchronized potentials were evoked. Since multiple local interneurons are known to synapse to a projection neuron in each glomerulus in the cockroach AL, converging inputs from local interneurons to the projection neurons appear to contribute the odorant specific spatio-temporal activity patterns of the synchronized potentials.
olfaction; synchronized potentials; optical imaging; voltage-sensitive dye; intracellular recording; local interneurons; projection neurons; insects
Related Articles
Rapid Encoding and Perception of Novel Odors in the Rat
Wesson, Daniel W
Carey, Ryan M
Verhagen, Justus V
Wachowiak, Matt
PLoS Biology
To gain insight into which parameters of neural activity are important in shaping the perception of odors, we combined a behavioral measure of odor perception with optical imaging of odor representations at the level of receptor neuron input to the rat olfactory bulb. Instead of the typical test of an animal's ability to discriminate two familiar odorants by exhibiting an operant response, we used a spontaneously expressed response to a novel odorant—exploratory sniffing—as a measure of odor perception. This assay allowed us to measure the speed with which rats perform spontaneous odor discriminations. With this paradigm, rats discriminated and began responding to a novel odorant in as little as 140 ms. This time is comparable to that measured in earlier studies using operant behavioral readouts after extensive training. In a subset of these trials, we simultaneously imaged receptor neuron input to the dorsal olfactory bulb with near-millisecond temporal resolution as the animal sampled and then responded to the novel odorant. The imaging data revealed that the bulk of the discrimination time can be attributed to the peripheral events underlying odorant detection: receptor input arrives at the olfactory bulb 100–150 ms after inhalation begins, leaving only 50–100 ms for central processing and response initiation. In most trials, odor discrimination had occurred even before the initial barrage of receptor neuron firing had ceased and before spatial maps of activity across glomeruli had fully developed. These results suggest a coding strategy in which the earliest-activated glomeruli play a major role in the initial perception of odor quality, and place constraints on coding and processing schemes based on simple changes in spike rate.
Author Summary
Olfactory stimuli elicit temporally complex patterns of activity across groups of receptor neurons as well as across central neurons. It remains unclear which parameters among these complex activity patterns are important in shaping odor perception. To address this issue, we imaged from the olfactory bulb of awake rats as they detected and responded to odorants. We used a spontaneously expressed response to novel odorants—exploratory sniffing—as a behavioral measure of odor perception. This assay allowed us to measure the speed with which rats perform simple odor discriminations by monitoring changes in respiration. Rats discriminated a novel odorant from a learned one in as little as 140 ms. Simultaneously imaging the sensory input to the olfactory bulb carried by receptor neurons revealed that the bulk of the response time is due to the peripheral events underlying odorant detection (inhalation and receptor neuron activation), leaving only 50–100 ms for central processing and response initiation. In most trials, responses to a novel odorant began before the initial barrage of input had ceased and before spatial patterns of input to the bulb had fully developed. These results suggest a coding strategy in which the earliest inputs play a major role in the initial perception of odor quality and place constraints on coding schemes based on simple changes in firing rate.
Imaging the olfactory bulb of awake rats reveals that odor discrimination occurs about 100 ms after sensory input reaches the brain, sharply limiting the role that spike rate and temporal integration can play in coding odor identity.
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Decorrelation of Odor Representations via Spike Timing-Dependent Plasticity
Linster, Christiane
Cleland, Thomas A.
Frontiers in Computational Neuroscience
The non-topographical representation of odor quality space differentiates early olfactory representations from those in other sensory systems. Decorrelation among olfactory representations with respect to physical odorant similarities has been proposed to rely upon local feed-forward inhibitory circuits in the glomerular layer that decorrelate odor representations with respect to the intrinsically high-dimensional space of ligand–receptor potency relationships. A second stage of decorrelation is likely to be mediated by the circuitry of the olfactory bulb external plexiform layer. Computations in this layer, or in the analogous interneuronal network of the insect antennal lobe, are dependent on fast network oscillations that regulate the timing of mitral cell and projection neuron (MC/PN) action potentials; this suggests a largely spike timing-dependent metric for representing odor information, here proposed to be a precedence code. We first illustrate how the rate coding metric of the glomerular layer can be transformed into a spike precedence code in MC/PNs. We then show how this mechanism of representation, combined with spike timing-dependent plasticity at MC/PN output synapses, can progressively decorrelate high-dimensional, non-topographical odor representations in third-layer olfactory neurons. Reducing MC/PN oscillations abolishes the spike precedence code and blocks this progressive decorrelation, demonstrating the learning network's selectivity for these sparsely synchronized MC/PN spikes even in the presence of temporally disorganized background activity. Finally, we apply this model to odor representations derived from calcium imaging in the honeybee antennal lobe, and show how odor learning progressively decorrelates odor representations, and how the abolition of PN oscillations impairs odor discrimination.
olfaction; gamma oscillations; sparse synchronization; STDP; olfactory bulb; antennal lobe; odor learning; conditioning
Related Articles
Sparse Distributed Representation of Odors in a Large-scale Olfactory Bulb Circuit
Yu, Yuguo
McTavish, Thomas S.
Hines, Michael L.
Shepherd, Gordon M.
Valenti, Cesare
Migliore, Michele
PLoS Computational Biology
In the olfactory bulb, lateral inhibition mediated by granule cells has been suggested to modulate the timing of mitral cell firing, thereby shaping the representation of input odorants. Current experimental techniques, however, do not enable a clear study of how the mitral-granule cell network sculpts odor inputs to represent odor information spatially and temporally. To address this critical step in the neural basis of odor recognition, we built a biophysical network model of mitral and granule cells, corresponding to 1/100th of the real system in the rat, and used direct experimental imaging data of glomeruli activated by various odors. The model allows the systematic investigation and generation of testable hypotheses of the functional mechanisms underlying odor representation in the olfactory bulb circuit. Specifically, we demonstrate that lateral inhibition emerges within the olfactory bulb network through recurrent dendrodendritic synapses when constrained by a range of balanced excitatory and inhibitory conductances. We find that the spatio-temporal dynamics of lateral inhibition plays a critical role in building the glomerular-related cell clusters observed in experiments, through the modulation of synaptic weights during odor training. Lateral inhibition also mediates the development of sparse and synchronized spiking patterns of mitral cells related to odor inputs within the network, with the frequency of these synchronized spiking patterns also modulated by the sniff cycle.
Author Summary
In the paper we address the role of lateral inhibition in a neuronal network. It is an essential and widespread mechanism of neural processing that has been demonstrated in many brain systems. A key finding that would reveal how and to what extent it can modulate input signals and give rise to some form of perception would involve network-wide recording of individual cells during in vivo behavioral experiments. While this problem has been intensely investigated, it is beyond current methods to record from a reasonable set of cells experimentally to decipher the emergent properties and behavior of the network, leaving the underlying computational and functional roles of lateral inhibition still poorly understood. We addressed this problem using a large-scale model of the olfactory bulb. The model demonstrates how lateral inhibition modulates the evolving dynamics of the olfactory bulb network, generating mitral and granule cell responses that account for critical experimental findings. It also suggests how odor identity can be represented by a combination of temporal and spatial patterns of mitral cell activity, with both feedforward excitation and lateral inhibition via dendrodendritic synapses as the underlying mechanisms facilitating network self-organization and the emergence of synchronized oscillations.
Related Articles
Associative Conditioning Tunes Transient Dynamics of Early Olfactory Processing
Fernandez, Patricia C.
Locatelli, Fernando F.
Person-Rennell, Nicole
Deleo, Gregory
Smith, Brian H.
The Journal of neuroscience : the official journal of the Society for Neuroscience
Odors evoke complex spatiotemporal responses in the insect antennal lobe (AL) and mammalian olfactory bulb. However, the behavioral relevance of spatiotemporal coding remains unclear. In the present work we combined behavioral analyses with calcium imaging of odor induced activity in the honey bee AL to evaluate the relevance of this temporal dimension in the olfactory code. We used a new way for evaluation of odor similarity of binary mixtures in behavioral studies, which involved testing if a match of odor sampling time is necessary between training and testing conditions for odor recognition during associative learning. Using graded changes in the similarity of the mixture ratios, we found high correlations between the behavioral generalization across those mixtures and a gradient of activation in AL output. Furthermore, short odor stimuli of 500 ms or less affected how well odors were matched with a memory template, and this time corresponded to a shift from a sampling-time-dependent to a sampling-time-independent memory. Accordingly, 375 ms corresponded to the time required for spatiotemporal AL activity patterns to reach maximal separation according to imaging studies. Finally, we compared spatiotemporal representations of binary mixtures in trained and untrained animals. AL activity was modified by conditioning to improve separation of odor representations. These data suggest that one role of reinforcement is to "tune" the AL such that relevant odors become more discriminable.
olfaction; synchrony; transients; spatiotemporal coding; plasticity; calcium imaging; discrimination
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Learned odor discrimination in Drosophila without distinct combinatorial odor maps in the antennal lobe
DasGupta, Shamik
Waddell, Scott
Current biology : CB
A unifying feature of mammalian and insect olfactory systems is that olfactory sensory neurons (OSNs) expressing the same unique odorant receptor gene converge onto the same glomeruli in the brain (1–7). Most odorants activate a combination of receptors and thus distinct patterns of glomeruli, forming a proposed combinatorial spatial code that could support discrimination between a large number of odorants (8–11). OSNs also exhibit odor-evoked responses with complex temporal dynamics (11), but the contribution of this activity to behavioral odor discrimination has received little attention (12). Here we investigated the importance of spatial encoding in the relatively simple Drosophila antennal lobe. We show that Drosophila can learn to discriminate between two odorants with one functional class of Or83b-expressing OSNs. Furthermore, these flies encode one odorant from a mixture, and cross-adapt to odorants that activate the relevant OSN class, demonstrating that they discriminate odorants using the same OSNs. Lastly, flies with a single class of Or83b-expressing OSNs recognize a specific odorant across a range of concentration indicating that they encode odorant identity. Therefore flies can distinguish odorants without discrete spatial codes in the antennal lobe, implying an important role for odorant-evoked temporal dynamics in behavioral odorant discrimination.
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Ensemble Response in Mushroom Body Output Neurons of the Honey Bee Outpaces Spatiotemporal Odor Processing Two Synapses Earlier in the Antennal Lobe
Strube-Bloss, Martin F.
Herrera-Valdez, Marco A.
Smith, Brian H.
PLoS ONE
Neural representations of odors are subject to computations that involve sequentially convergent and divergent anatomical connections across different areas of the brains in both mammals and insects. Furthermore, in both mammals and insects higher order brain areas are connected via feedback connections. In order to understand the transformations and interactions that this connectivity make possible, an ideal experiment would compare neural responses across different, sequential processing levels. Here we present results of recordings from a first order olfactory neuropile – the antennal lobe (AL) – and a higher order multimodal integration and learning center – the mushroom body (MB) – in the honey bee brain. We recorded projection neurons (PN) of the AL and extrinsic neurons (EN) of the MB, which provide the outputs from the two neuropils. Recordings at each level were made in different animals in some experiments and simultaneously in the same animal in others. We presented two odors and their mixture to compare odor response dynamics as well as classification speed and accuracy at each neural processing level. Surprisingly, the EN ensemble significantly starts separating odor stimuli rapidly and before the PN ensemble has reached significant separation. Furthermore the EN ensemble at the MB output reaches a maximum separation of odors between 84–120 ms after odor onset, which is 26 to 133 ms faster than the maximum separation at the AL output ensemble two synapses earlier in processing. It is likely that a subset of very fast PNs, which respond before the ENs, may initiate the rapid EN ensemble response. We suggest therefore that the timing of the EN ensemble activity would allow retroactive integration of its signal into the ongoing computation of the AL via centrifugal feedback.
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Friends and Foes from an Ant Brain's Point of View – Neuronal Correlates of Colony Odors in a Social Insect
Brandstaetter, Andreas Simon
Rössler, Wolfgang
Kleineidam, Christoph Johannes
PLoS ONE
Successful cooperation depends on reliable identification of friends and foes. Social insects discriminate colony members (nestmates/friends) from foreign workers (non-nestmates/foes) by colony-specific, multi-component colony odors. Traditionally, complex processing in the brain has been regarded as crucial for colony recognition. Odor information is represented as spatial patterns of activity and processed in the primary olfactory neuropile, the antennal lobe (AL) of insects, which is analogous to the vertebrate olfactory bulb. Correlative evidence indicates that the spatial activity patterns reflect odor-quality, i.e., how an odor is perceived. For colony odors, alternatively, a sensory filter in the peripheral nervous system was suggested, causing specific anosmia to nestmate colony odors. Here, we investigate neuronal correlates of colony odors in the brain of a social insect to directly test whether they are anosmic to nestmate colony odors and whether spatial activity patterns in the AL can predict how odor qualities like "friend" and "foe" are attributed to colony odors.
Methodology/Principal Findings
Using ant dummies that mimic natural conditions, we presented colony odors and investigated their neuronal representation in the ant Camponotus floridanus. Nestmate and non-nestmate colony odors elicited neuronal activity: In the periphery, we recorded sensory responses of olfactory receptor neurons (electroantennography), and in the brain, we measured colony odor specific spatial activity patterns in the AL (calcium imaging). Surprisingly, upon repeated stimulation with the same colony odor, spatial activity patterns were variable, and as variable as activity patterns elicited by different colony odors.
Ants are not anosmic to nestmate colony odors. However, spatial activity patterns in the AL alone do not provide sufficient information for colony odor discrimination and this finding challenges the current notion of how odor quality is coded. Our result illustrates the enormous challenge for the nervous system to classify multi-component odors and indicates that other neuronal parameters, e.g., precise timing of neuronal activity, are likely necessary for attribution of odor quality to multi-component odors.
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Nonassociative Plasticity Alters Competitive Interactions Among Mixture Components In Early Olfactory Processing
Locatelli, Fernando F
Fernandez, Patricia C
Villareal, Francis
Muezzinoglu, Kerem
Huerta, Ramon
Galizia, C. Giovanni
Smith, Brian H.
The European journal of neuroscience
Experience related plasticity is an essential component of networks involved in early olfactory processing. However, the mechanisms and functions of plasticity in these neural networks are not well understood. We studied nonassociative plasticity by evaluating responses to two pure odors (A and X) and their binary mixture using calcium imaging of odor elicited activity in output neurons of the honey bee antennal lobe. Unreinforced exposure to A or X produced no change in the neural response elicited by the pure odors. However, exposure to one odor (e.g. A) caused the response to the mixture to become more similar to the other component (X). We also show in behavioral analyses that unreinforced exposure to A caused the mixture to become perceptually more similar to X. These results suggest that nonassociative plasticity modifies neural networks in such a way that it affects local competitive interactions among mixture components. We used a computational model to evaluate the most likely targets for modification. Hebbian modification of synapses from inhibitory local interneurons to projection neurons most reliably produces the observed shift in response to the mixture. These results are consistent with a model in which the antennal lobe acts to filter olfactory information according to its relevance for performing a particular task.
Olfaction; learning; antennal lobe; plasticity; Apis mellifera
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The neuronal architecture of the mushroom body provides a logic for associative learning
Aso, Yoshinori
Hattori, Daisuke
Yu, Yang
Johnston, Rebecca M
Iyer, Nirmala A
Ngo, Teri-TB
Dionne, Heather
Abbott, LF
Axel, Richard
Tanimoto, Hiromu
Rubin, Gerald M
We identified the neurons comprising the Drosophila mushroom body (MB), an associative center in invertebrate brains, and provide a comprehensive map describing their potential connections. Each of the 21 MB output neuron (MBON) types elaborates segregated dendritic arbors along the parallel axons of ∼2000 Kenyon cells, forming 15 compartments that collectively tile the MB lobes. MBON axons project to five discrete neuropils outside of the MB and three MBON types form a feedforward network in the lobes. Each of the 20 dopaminergic neuron (DAN) types projects axons to one, or at most two, of the MBON compartments. Convergence of DAN axons on compartmentalized Kenyon cell–MBON synapses creates a highly ordered unit that can support learning to impose valence on sensory representations. The elucidation of the complement of neurons of the MB provides a comprehensive anatomical substrate from which one can infer a functional logic of associative olfactory learning and memory.
DOI: http://dx.doi.org/10.7554/eLife.04577.001
eLife digest
One of the key goals of neuroscience is to understand how specific circuits of brain cells enable animals to respond optimally to the constantly changing world around them. Such processes are more easily studied in simpler brains, and the fruit fly—with its small size, short life cycle, and well-developed genetic toolkit—is widely used to study the genes and circuits that underlie learning and behavior.
Fruit flies can learn to approach odors that have previously been paired with food, and also to avoid any odors that have been paired with an electric shock, and a part of the brain called the mushroom body has a central role in this process. When odorant molecules bind to receptors on the fly's antennae, they activate neurons in the antennal lobe of the brain, which in turn activate cells called Kenyon cells within the mushroom body. The Kenyon cells then activate output neurons that convey signals to other parts of the brain.
It is known that relatively few Kenyon cells are activated by any given odor. Moreover, it seems that a given odor activates different sets of Kenyon cells in different flies. Because the association between an odor and the Kenyon cells it activates is unique to each fly, each fly needs to learn through its own experiences what a particular pattern of Kenyon cell activation means.
Aso et al. have now applied sophisticated molecular genetic and anatomical techniques to thousands of different transgenic flies to identify the neurons of the mushroom body. The resulting map reveals that the mushroom body contains roughly 2200 neurons, including seven types of Kenyon cells and 21 types of output cells, as well as 20 types of neurons that use the neurotransmitter dopamine. Moreover, this map provides insights into the circuits that support odor-based learning. It reveals, for example, that the mushroom body can be divided into 15 anatomical compartments that are each defined by the presence of a specific set of output and dopaminergic neuron cell types. Since the dopaminergic neurons help to shape a fly's response to odors on the basis of previous experience, this organization suggests that these compartments may be semi-autonomous information processing units.
In contrast to the rest of the insect brain, the mushroom body has a flexible organization that is similar to that of the mammalian brain. Elucidating the circuits that support associative learning in fruit flies should therefore make it easier to identify the equivalent mechanisms in vertebrate animals.
DOI: http://dx.doi.org/10.7554/eLife.04577.002
mushroom body; olfactory learning; associative memory; neuronal circuits; dopamine; plasticity; D. melanogaster
Related Articles
The banana code—natural blend processing in the olfactory circuitry of Drosophila melanogaster
Schubert, Marco
Hansson, Bill S.
Sachse, Silke
Frontiers in Physiology
Odor information is predominantly perceived as complex odor blends. For Drosophila melanogaster one of the most attractive blends is emitted by an over-ripe banana. To analyze how the fly's olfactory system processes natural blends we combined the experimental advantages of gas chromatography and functional imaging (GC-I). In this way, natural banana compounds were presented successively to the fly antenna in close to natural occurring concentrations. This technique allowed us to identify the active odor components, use these compounds as stimuli and measure odor-induced Ca2+ signals in input and output neurons of the Drosophila antennal lobe (AL), the first olfactory neuropil. We demonstrate that mixture interactions of a natural blend are very rare and occur only at the AL output level resulting in a surprisingly linear blend representation. However, the information regarding single components is strongly modulated by the olfactory circuitry within the AL leading to a higher similarity between the representation of individual components and the banana blend. This observed modulation might tune the olfactory system in a way to distinctively categorize odor components and improve the detection of suitable food sources. Functional GC-I thus enables analysis of virtually any unknown natural odorant blend and its components in their relative occurring concentrations and allows characterization of neuronal responses of complete neural assemblies. This technique can be seen as a valuable complementary method to classical GC/electrophysiology techniques, and will be a highly useful tool in future investigations of insect-insect and insect-plant chemical interactions.
Drosophila; gas chromatography; in vivo calcium imaging; olfaction; blend coding; insect antennal lobe
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Honeybees Learn Odour Mixtures via a Selection of Key Odorants
Reinhard, Judith
Sinclair, Michael
Srinivasan, Mandyam V.
Claudianos, Charles
PLoS ONE
The honeybee has to detect, process and learn numerous complex odours from her natural environment on a daily basis. Most of these odours are floral scents, which are mixtures of dozens of different odorants. To date, it is still unclear how the bee brain unravels the complex information contained in scent mixtures.
Methodology/Principal Findings
This study investigates learning of complex odour mixtures in honeybees using a simple olfactory conditioning procedure, the Proboscis-Extension-Reflex (PER) paradigm. Restrained honeybees were trained to three scent mixtures composed of 14 floral odorants each, and then tested with the individual odorants of each mixture. Bees did not respond to all odorants of a mixture equally: They responded well to a selection of key odorants, which were unique for each of the three scent mixtures. Bees showed less or very little response to the other odorants of the mixtures. The bees' response to mixtures composed of only the key odorants was as good as to the original mixtures of 14 odorants. A mixture composed of the other, non-key-odorants elicited a significantly lower response. Neither an odorant's volatility or molecular structure, nor learning efficiencies for individual odorants affected whether an odorant became a key odorant for a particular mixture. Odorant concentration had a positive effect, with odorants at high concentration likely to become key odorants.
Our study suggests that the brain processes complex scent mixtures by predominantly learning information from selected key odorants. Our observations on key odorant learning lend significant support to previous work on olfactory learning and mixture processing in honeybees.
Related Articles
Plant odorants interfere with detection of sex pheromone signals by male Heliothis virescens
Pregitzer, Pablo
Schubert, Marco
Breer, Heinz
Hansson, Bill S.
Sachse, Silke
Krieger, Jürgen
Frontiers in Cellular Neuroscience
In many insects, mate finding relies on female-released sex pheromones, which have to be deciphered by the male olfactory system within an odorous background of plant volatiles present in the environment of a calling female. With respect to pheromone-mediated mate localization, plant odorants may be neutral, favorable, or disturbing. Here we examined the impact of plant odorants on detection and coding of the major sex pheromone component, (Z)-11-hexadecenal (Z11-16:Ald) in the noctuid moth Heliothis virescens. By in vivo imaging the activity in the male antennal lobe (AL), we monitored the interference at the level of olfactory sensory neurons (OSN) to illuminate mixture interactions. The results show that stimulating the male antenna with Z11-16:Ald and distinct plant-related odorants simultaneously suppressed pheromone-evoked activity in the region of the macroglomerular complex (MGC), where Z11-16:Ald-specific OSNs terminate. Based on our previous findings that antennal detection of Z11-16:Ald involves an interplay of the pheromone binding protein (PBP) HvirPBP2 and the pheromone receptor (PR) HR13, we asked if the plant odorants may interfere with any of the elements involved in pheromone detection. Using a competitive fluorescence binding assay, we found that the plant odorants neither bind to HvirPBP2 nor affect the binding of Z11-16:Ald to the protein. However, imaging experiments analyzing a cell line that expressed the receptor HR13 revealed that plant odorants significantly inhibited the Z11-16:Ald-evoked calcium responses. Together the results indicate that plant odorants can interfere with the signaling process of the major sex pheromone component at the receptor level. Consequently, it can be assumed that plant odorants in the environment may reduce the firing activity of pheromone-specific OSNs in H. virescens and thus affect mate localization.
pheromone detection; antennal lobe; pheromone receptor; pheromone binding protein; olfaction
Related Articles
Keeping their distance? Odor response patterns along the concentration range
Strauch, Martin
Ditzen, Mathias
Galizia, C. Giovanni
Frontiers in Systems Neuroscience
We investigate the interplay of odor identity and concentration coding in the antennal lobe (AL) of the honeybee Apis mellifera. In this primary olfactory center of the honeybee brain, odors are encoded by the spatio-temporal response patterns of olfactory glomeruli. With rising odor concentration, further glomerular responses are recruited into the patterns, which affects distances between the patterns. Based on calcium-imaging recordings, we found that such pattern broadening renders distances between glomerular response patterns closer to chemical distances between the corresponding odor molecules. Our results offer an explanation for the honeybee's improved odor discrimination performance at higher odor concentrations.
calcium-imaging; honeybees; glomerular pattern; odor concentration coding; odor identity coding; chemical dissimilarity
Related Articles
Learning Modifies Odor Mixture Processing to Improve Detection of Relevant Components
Chen, Jen-Yung
Marachlian, Emiliano
Assisi, Collins
Huerta, Ramon
Smith, Brian H.
Locatelli, Fernando
Bazhenov, Maxim
The Journal of Neuroscience
Honey bees have a rich repertoire of olfactory learning behaviors, and they therefore are an excellent model to study plasticity in olfactory circuits. Recent behavioral, physiological, and molecular evidence suggested that the antennal lobe, the first relay of the olfactory system in insects and analog to the olfactory bulb in vertebrates, is involved in associative and nonassociative olfactory learning. Here we use calcium imaging to reveal how responses across antennal lobe projection neurons change after association of an input odor with appetitive reinforcement. After appetitive conditioning to 1-hexanol, the representation of an odor mixture containing 1-hexanol becomes more similar to this odor and less similar to the background odor acetophenone. We then apply computational modeling to investigate how changes in synaptic connectivity can account for the observed plasticity. Our study suggests that experience-dependent modulation of inhibitory interactions in the antennal lobe aids perception of salient odor components mixed with behaviorally irrelevant background odors.
antennal lobe; honey bees; olfaction; olfactory learning
Related Articles
Serotonin Modulates Olfactory Processing in the Antennal Lobe of Drosophila
Dacks, Andrew M.
Green, David S.
Root, Cory M.
Nighorn, Alan J.
Wang, Jing W.
Journal of neurogenetics
Sensory systems must be able to extract features of environmental cues within the context of the different physiological states of the organism and often temper their activity in a state-dependent manner via the process of neuromodulation. We examined the effects of the neuromodulator serotonin on a well-characterized sensory circuit, the antennal lobe of Drosophila melanogaster, using two-photon microscopy and the genetically expressed calcium indicator, G-CaMP. Serotonin enhances sensitivity of the antennal lobe output projection neurons in an odor-specific manner. For odorants that sparsely activate the antennal lobe, serotonin enhances projection neuron responses and causes an offset of the projection neuron tuning curve, most likely by increasing projection neuron sensitivity. However, for an odorant that evokes a broad activation pattern, serotonin enhances projection neuron responses in some, but not all, glomeruli. Further, serotonin enhances the responses of inhibitory local interneurons, resulting in a reduction of neurotransmitter release from the olfactory sensory neurons via GABAB receptor-dependent presynaptic inhibition, which may be a mechanism underlying the odorant-specific modulation of projection neuron responses. Our data suggest that the complexity of serotonin modulation in the antennal lobe accommodates coding stability in a glomerular pattern and flexible projection neuron sensitivity under different physiological conditions.
olfaction; neuromodulation; serotonin; antennal lobes
Related Articles
Complex and non-redundant signals from individual odor receptors that underlie chemotaxis behavior in Drosophila melanogaster larvae
Grewal, Jeewanjot S.
Nguyen, Christine
Robles, Raquel
Cho, Christina
Kir, Karolina
Fledderman, Nicole
Gacharna, George
Wesolowski, Michael
Klinger, Christie
Vallejo, Pedro
Menhennett, Lorien
Nagaraj, Abhiram
Ebo, Chineze
Peacy, Garrett
Davelis, Eftihia
Kucher, David
Giers, Sarah
Kreher, Scott A.
Biology Open
The rules by which odor receptors encode odors and allow behavior are still largely unexplored. Although large data sets of electrophysiological responses of receptors to odors have been generated, few hypotheses have been tested with behavioral assays. We use a data set on odor responses of Drosophila larval odor receptors coupled with chemotaxis behavioral assays to examine rules of odor coding. Using mutants of odor receptors, we have found that odor receptors with similar electrophysiological responses to odors across concentrations play non-redundant roles in odor coding at specific odor concentrations. We have also found that high affinity receptors for odors determine behavioral response thresholds, but the rules for determining peak behavioral responses are more complex. While receptor mutants typically show loss of attraction to odors, some receptor mutants result in increased attraction at specific odor concentrations. The odor receptor mutants were rescued using transgenic expression of odor receptors, validating assignment of phenotypes to the alleles. Vapor pressures alone cannot fully explain behavior in our assay. Finally, some odors that did not elicit strong electrophysiological responses are associated with behavioral phenotypes upon examination of odor receptor mutants. This result is consistent with the role of sensory neurons in lateral inhibition via local interneurons in the antennal lobe. Taken together, our results suggest a complexity of odor coding rules even in a simple olfactory sensory system.
Odor receptors; Olfaction; Drosophila; Or42a; Or42b
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Zam: DAP irate as its logo stands out in Tanda Putera
Among the film Tanda Putera 's most outspoken critics is the DAP, which, among others, has held several press conferences to speak out against it. But why?
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I was working on updating a classic American recipe for my Paleo Magazine column this weekend and realized I had no idea how to properly prepare...Read More
How To Peel A Grapefruit
I la-la-love pink grapefruit, and the Thai Pink Grapefruit Salad from Well Fed 2 is a both unexpected (dried shrimp! coconut! lime!) and delicious. The combo of sweet, salty, cool, and creamy is irresistible — as is pink grapefruit all on its own.
We're coming into grapefruit season now, so put away those shameful serrated grapefruit spoons and take a look at this technique I learned from my dad for peeling a grapefruit. Be sure to stick around to the end of the video because my husband/videographer Dave always includes a tasty little blooper or something silly to close out the show.
More Like This...
In each issue of Paleo Magazine, I share the story of a traditional recipe and adapt it to fit into a healthier Paleo lifestyle. In a recent...Read More
Aweeeee! I love your video. Very cool presentation. Awesome shirt I might add. You look very pretty. 🙂
"Be sure to stick around to the end of the video because my husband/videographer Dave always includes a tasty little extra to close out the show." Where's the tasty extra?
Pingback: How To Peel A Grapefruit | Paleo Digest
Great technique! I have been cooking for 50+ years and didn't know this technique – shows you can always learn something new. Thanks!
So handy! And you are so adorable 🙂
My favorite snack is to break the segments into small bite sized pieces and put in freezer!
Great tutorial, I always just slice mine down the middle and scoop the fruit out, but I'll be testing this out.
I finally tracked down dried shrimp. I'm making the grapefruit salad tonight and I'm totally excited about it. 🙂
This post changed my life. I love grapefruit and just bought some in hopes to fight off a terrible no good awful cold, but was not sure what I was going to do with them. I love juicing them with ginger, but I'm currently too lazy to pull out the juicer and clean it. I peeled them using this technique, grated some fresh ginger over it, and topped with a sprinkle of coconut sugar. Spicy and delightful! Thank you for sharing this 🙂
Where did you find dried shrimp? i've looked all over and all i can find is fish food not human food 🙁
Ok, here is a crazy question. I know it is way out there but here it goes. WHERE IS THE RECIPE?!
I forgot to say thank you.:)


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Luca Tedesco (an Italian Developer and researcher) has just released a video ,which can be found on Twitter , that demonstrates the technique of iOS Jailbreaking specifically for 64-bit smartphones. This video comes just hours prior to the long awaited 7 September event ,in which Apple is to announce it's latest flagship phone ,the iPhone […] Read More
iPhone Cydia Applications
People who are familiar with Android OS might feel some of the software functionalities provided in Android OS should be made part of iOS but there are no easy ways of getting them then Jailbreak tweaks. One such kind of functionality is Haptic Feedback on Keyboard which gives you real vibration feel of some activity going on […] Read More
Finally the wait is over, WeatherIcon has been updated to work with iOS 6 on iPhone 5, iPod Touch and iPad. It is one of the most anticipated Jailbreak tweak which allows you to show real time weather updates right on the icon of weather application. Thereby you can get glimpse of current weather right on the […] Read More
How to Install Siri On iPhone 4, iPad 2, iPod touch With Twitter & Facebook Integration Using Free WordJelly Siri Server [VIDEO]
Since the launch of iPhone 4S siri assistant feature, fans have been waiting for the solution which allows it port on older devices. Although many solutions were surfaced on cydia app store but none of them has achieved the same perfection which came with iPhone 4S Siri Assistant feature. One of the famous solution came […] Read More
Recently we reported you about a native iOS 5 hidden feature which allow users to make FaceTime calls over 3G Network through simple plist modification rather than using any third party app. Now jailbreak community has released tweak which allows you to modify the required Plist through Graphical User Interface (GUI) rather than doing it manually using iFile. Developer 3Gvenice has named […] Read More
Release of iPhone 4S and iPad 2 Jailbreak has opened the gates for jailbreak to come up with special apps and tweak which works the support of Siri and make things done easier then before. We have seen similar successful attempts in recent days such as SiriToggles, SiriUtils which allows you to change your settings simply through your voice command. […] Read More
We have reviewed various cydia tweaks such as SiriToggles, SiriUtils which really enhances iPhone 4S capabilities by allowing you to launch Apps, Turn ON/OFF WiFi/Bluetooth, Change Brightness And More using Siri Assistant feature . Now another jailbreak app called Sireet has just hit Cydia store which enables you to Tweet using native Siri Assistant feature. All you have to do is […] Read More
SiriUtils Lets You Launch Apps, Turn ON/OFF WiFi/Bluetooth, Change Brightness And More [Jailbreak Tweak]
It was only couple of days back when iPhone 4S untethered jailbreak was released and now iOS developers have already started releasing new jailbreak tweaks and apps specially to enhance the capability of Siri Assistant feature. Just now we reviewed SiriToggles which allows you to Enable/Disable certain iPhone settings. Now similar kind of jailbreak tweak called SiriUtils […] Read More
Have you ever wished of opening apps or change settings through Voice Commands specially after the release of iPhone 4S Siri Assistant feature? After the release of iPhone 4S and iPad 2 Untethered Jailbreak, Thanks to iOS jailbreak community who have already started using iPhone 4S Siri Assistant feature to bring this functionality on iOS […] Read More
We have already reviewed WeeSpaces, WeeCompose, WeeKillBackground, WeeSearch jailbreak tweaks brings various new features to iOS 5 Notification Center which are not available in native Notification Center app. Another amazing jailbreak tweak has been released for Notification Center called WeeSlide which not only give fast access to Volume and Brightness Controls but also adds a WiFi […] Read More
Previously we have reviewed many jailbreak tweaks which allows you to change your iPhone Slide to Unlock Text such as Slider Enhancer. Now we another similar kind of jailbreak tweak has hit the cydia app store called SlideMyName which allows you to replace your Slide to unlock text with your iDevice. This jailbreak tweak has come from […] Read More
I am sure you have already jailbroken your iPhone 4S after the release of Greenpois0n Absinthe tool for both Windows and Mac operating system. Jailbreaking tool which provides Untethered jailbreak for iPhone 4S running iOS 5.0 or 5.0.1 firmware and iPad 2 (Wi-Fi only, GSM & CDMA) devices running iOS 5.0.1 firmware. Therefore, in this tutorial we have prepared a […] Read More


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The Effingham Chamber of Commerce had a ribbon cutting July 20 for Energy One Credit Union, which has moved its office from inside the Effingham ...
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Connecting the CMA and MOCA
The Cleveland Museum of Art and MOCA Cleveland have joined forces to present two contemporary installations by celebrated multi-media artist Janet Cardiff—Forty-Part Motet at the CMA and The Paradise Institute at MOCA. In celebration of these two innovative works of art the CMA's first Friday happy hour event, MIX, will be a special collaboration with MOCA's WtF (Welcome to Fridays) series. Join us Friday, June 7 from 5-9 p.m. at MIX: Connections for cocktails, live music and immersive art at both the CMA and MOCA. A free trolley will run all evening between the two museums. In anticipation of this exciting evening, we discuss Janet Cardiff and her groundbreaking installations.


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This is a well organized and orthodox summary of the recognized methods of treatment of tuberculosis, offered as a professorship thesis to the University of Buenos Aires. The author has classified the subtitles under the three general heads (1) treatment of the soil, (2) treatment of the bacillus, (3) treatment of the focus. Under each of the general headings there are chapters carefully divided into paragraphs, each contributing its quota necessary to complete the entire field indicated in the title. The presentation constitutes an excellent compendium relating to all the methods of treatment, non-surgical and surgical, recognized as appropriate according to present day standards.
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Bayern Munich visits Viktoria Plzen on Tuesday knowing a win could see it qualify for the knockout stages of the Champions League.
The defending champions sit three points clear of nearest rival Manchester City at the top of Group D after three consecutive wins, scoring 11 goals and conceding just one.
Franck Ribery's form has been key to Bayern's success so far, as the France international winger is the German side's top scorer in the tournament having found the net three times. Two of those goals came in Bayern's 5-0 destruction of Plzen at the Allianz Arena last month with Mario Gotze, Bastian Schweinsteiger and David Alaba also scoring in a comprehensive win.
Coach Pep Guardiola will look for his Bayern side to replicate that performance, well aware that the Bundesliga champions will qualify for the round of 16 if they take all three points and Manchester City beats CSKA Moscow.
Bayern should be boosted by the prospective returns of winger Arjen Robben and midfielder Toni Kroos, who both missed Saturday's 2-1 win over Hoffenheim with groin injuries. Defender Dante is still a doubt due to a lack of fitness, while Xherdan Shaqiri and Claudio Pizarro (thigh), Thiago Alcantara (ankle) and Holger Badstuber (knee) are all unavailable.
Plzen - competing in the competition for only the second time - is on the brink of elimination and needs a shock win to have a realistic chance of reaching the last 16.
The Czech outfit has shipped 11 goals in three straight defeats and has scored in just one game, a 3-2 defeat to CSKA.
Attacker Tomas Wagner could be crucial if Plzen is to get a result after finding the net eight times in all competitions this season, while Daniel Kolar and Stanislav Tecl - six goals each - may also play a pivotal role.
Pavel Vrba's men have no major injury worries, with defender David Limbersky poised to make his Champions League comeback after recovering from a thigh injury.
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- 3 loaves
- 55 Min
This proves that pumpkin isn't only for Halloween. It's a rich, homemade taste for anytime. Plus, the addition of nuts and dates gives this fall bread an extra-special crunch, making it even better than more traditional pumpkin breads.
What You'll Need:
- 4 eggs
- 3 cups sugar
- 1 cup peanut oil
- 1/3 cup water
- 1 (30-ounce) can pumpkin pie mix
- 3 1/2 cups all-purpose flour
- 2 teaspoons baking soda
- 1 1/2 teaspoons salt
- 1 teaspoon ground cinnamon
- 1 1/2 cups chopped dates
- 1 cup chopped roasted peanuts
What To Do:
- Preheat oven to 350 degrees F. Grease and flour three 9 x 5-inch loaf pans.
- In a large bowl, lightly beat the eggs. Gradually beat in the sugar, oil, water, and pumpkin. Add the flour, baking soda, salt, and cinnamon; mix well. Stir in the dates and peanuts. Divide the batter evenly among the loaf pans.
- Bake 55 to 60 minutes or until the bread begins to pull away from the sides of the pans. Cool in pans on wire racks for 30 minutes, then remove from pans and cool completely on racks.
Sometimes I use raisins instead of dates, or even some of both.
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text:
Eyes on the Dollar: New York Islanders
Cam Lewis
December 14 2015 08:00AM
Remember a few years ago when the New York Islanders dealt Matt Moulson and a first and second round pick in either 2014 or 2015 to the Buffalo Sabres for soon-to-be UFA Thomas Vanek? Then a few months later they traded him to Montreal for Sebastien Collberg and second round pick? That was awful. It was like, Alexei Yashin trade/contract/buyout bad. It looked like they gave away a lottery pick and Moulson for 47 games of Vanek, and a couple of throw-ins. Garth Snow opted to give Buffalo the first round pick in 2015 rather than 2014, giving him one summer to turn the franchise around, otherwise his team would be on the losing end of a horrible trade.
Jaroslav Halak, Mikhail Grabovski, and Nikolai Kulemin were signed as free agents, Johnny Boychuk and Nick Leddy were acquired through trade, and the team did a complete 180, going from eighth in the Metropolitan Division to third in just one year. This season, the Islanders are one of the best teams in the Eastern Conference and appear to be one their way to establishing themselves as legit Cup contender. That said, a trade request from travis Hamonic and two key impending free agents will make 2016 another interesting summer for Snow.
Future Free Agent Profiles: Brent Burns
Cam Lewis
December 11 2015 07:30AM
By now, we all know that this summer's free agent crop is absolutely loaded with talent. In fact, it might actually be the best group of players to hit the open market at the same time since the NHL implemented the salary cap back in 2005. But what about the 2017 UFA class? While next, next summer's group doesn't boast names like Steven Stamkos and Anze Kopitar, there are still a handful of franchise changing players who could possibly become free agents.
Like Brent Burns, for example, who, aside from owning arguably the best beard in hockey, has cemented himself as one of the league's elite offensive defencemen over the past few seasons. Since the beginning of the 2013 lockout season, Erik Karlsson and P.K. Subban are the only defencemen in the NHL who have scored more points than Burns has, and this season, nobody in the league has produced more individual shot attempts.
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The Roundup
Nation World HQ
December 11 2015 06:30AM
Leafs fans still think Stamkos is coming, Oilers and Flames both heating up, a look back at the Evander Kane trade, Prust happy with fine for spear on Marchand, NHL expansion, 2016 draft lottery and salary cap, a big left hand, fantasy hockey and more in this week's Nation Roundup brought to you by DraftKings
The NHL's 2016-17 cap ceiling and this summer's UFA class
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December 09 2015 04:55PM
A couple of days ago at the league's Board of Governors Meeting, Commissioner Gary Bettman said that the NHL's salary cap could go up by roughly $3 million next season. He also went on to say that this was a "very, very, very rough projection" and that next year's cap will "depend on a variety of different factors" like the strength of the Canadian dollar and the revenue the league can generate. It's important to remember that this time last year, Bettman suggested that the cap for the 2015-16 season would be around $73 million, but as we know, it ended up being less than that.
If this is the case, and everything goes according to plan, we'll be looking at somewhere around a $74.5 million salary cap ceiling for the 2016-17 season, which could be interesting as we head into a summer that boasts one of the best free agent classes of recent memory.
Top to Bottom: Checking in with each team two months into the season
Cam Lewis
December 07 2015 02:58PM
We're two months into the NHL season! The Jets and Lightning are underachieving because of issues on special teams, it seems like an AHL team could compete for a playoff spot in the Pacific Division, and the Dallas Stars are scoring like a team straight out of the 1980s. Let's look up and down the league at our legitimate contenders, bubble teams, and write-offs.


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Web posted Saturday, July 11, 1998 4:28 a.m. CT
New ENMU cage coach Tyler eyes positive attitude
By TRAVIS HARSCH
Globe-News Sports Writer
There are two kinds of basketball coaches in the world - those who see a .500 record as half wins and those who see the same record as half losses.
Terry Tyler, named the new head coach at Eastern New Mexico on June 23, sees the wins.
"My attitude when I came down here to interview was that I didn't come down here to interview for a job, I came down to be a coach," Tyler said. "I want to come down here and turn this program around. I don't want to just have a title, I want to have an impact on the program and be a part of this university."
Tyler, an assistant coach at Notre Dame last season, will try to turn the program around from an 8-18 record last season. He compared his new school's home, the Lone Star Conference, to one which has won wide acclaim in recent years as a top basketball league.
"In the Big East (of which Notre Dame is a member), it was very competitive, and there were no easy games. It's probably one of the top conferences in the country," Tyler said.
"I think this (the LSC) is a competitive conference, with the South Division and the North Division, and it's competitive. You've got good basketball teams, so there are not going to be easy games."
Tyler replaces Earl Diddle, who led the Greyhounds to a 156-124 mark in his 10 seasons at the helm. Diddle resigned following the 1997-98 season, during which ENMU was 3-13 in the LSC. Tyler said the Greyhounds may return a few players from that team, but his won't be the only new face in Greyhound Arena next season.
"We have three or four returning, but the majority is going to be new players," Tyler said. "I feel the outlook is interesting, looking at last year's record, there is a challenge, and we've got a lot of work in front of us."
Tyler, who played 11 seasons in the NBA, said he hopes to use that professional knowledge to help his players.
"I learned a lot. I found out that it is a business," Tyler said. "I think that is what I will pass on to my players, I'm sure they dream about going to the next level, and I will share my experiences with them. But at the same time, I will make sure they understand that life goes on after basketball."
A lot of that knowledge from Notre Dame and the NBA Tyler has put to use; he wants his teams to play a quick-paced game.
"I like the up-tempo game, and I think that's something that the kids like to play," Tyler said. "But there are going to be times when we are going to run set plays too, and we're going to play a man-to-man defense."
In contrast to the fast-break style his team will employ, the relaxed atmosphere of this region helped draw Tyler to Portales.
"I lived in Dallas for a few years and I always felt I liked this part of the country and I liked the pace of everything," Tyler said. "I knew this was a good fit for me; it was the opportunity of a lifetime. I felt it had always been my calling to be a head coach and I decided I wanted to be in this area. I was familiar with it.
Although Tyler is familiar with the area, he wasn't very well-acquainted with ENMU when he first looked at the head coaching position, he said.
"When I saw the opening, I went ahead and applied for it, not knowing much about the school, although I did look it up on the Internet to find out a little bit about the program," Tyler said. "It was always my goal to be a head coach. It just worked out that they liked my resume and interviewed me, along with two other candidates for the job, and I was selected."
Tyler also hopes his optimism will spread to his team as the Greyhounds strive for a season anyone can see optimistically.
"I want our players to feel good about themselves, academically and athletically," Tyler said. "I want them to feel like they can go out on the court and play with anybody."
We'd love to hear your feedback. © 1998 Amarillo Globe-NewsWebmaster
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»0 Comments Soccer on TV Posted by Soccer America on Jun. 25, 10:15 a.m. Gol TV, TeleFutura and Univision provide live game coverage of the Copa America. The USA takes part as a guest team in the South American championship for the first time since finishing fourth in 1995. The Americans' first game is Thursday -- four days after lifting the Gold Cup title -- against Argentina. The U-20 World Cup, which will air ... »0 Comments Soccer on TV Posted by Soccer America on Jun. 22, 2:00 p.m. The U.S. women's national team continues preparations for the 2007 World Cup when it takes on Brazil in a friendly (a rematch of the 2004 Olympic gold medal game won by the USA) on Saturday in New Jersey (ESPN2 & Univision). The U.S. men meet archrival Mexico in the Gold Cup final on Sunday (Fox Soccer Channel and Univision). In ... »0 Comments Soccer on TV Posted by Soccer America on Jun. 18, 10:00 a.m. Boca Juniors takes a 3-0 lead into the second leg of the Libertadores Cup semifinal on Wednesday (Fox Sports en Espanol). On Thursday, the USA faces Canada in the Gold Cup semifinals (Fox Soccer Channel & Telefutura) while Mexico clashes with Guadeloupe (Telefutura). ESPN2's Thursday night MLS clash is Houston-Chivas USA.
»0 Comments Soccer on TV Posted by Soccer America on Jun. 11, 11:45 a.m. The weekend is highlighted by four Gold Cup quarterfinals, including the USA-Panama rematch of the 2005 final on Saturday on Fox Soccer Channel and Univision. Fans can watch David Beckham'sfinal game for Real Madrid, which hosts Real Madrid with La Liga title on the line on Sunday. In addition, there is a full slate of MLS games. »0 Comments Soccer on TV Posted by Soccer America on Jun. 8, 10:15 a.m. Montreal hosts Portland in USL action on Friday, when Mexico opens its Gold Cup campaign against Cuba. Then comes Big Saturday: The USA's Gold Cup clash with Trinidad & Tobago (Fox Soccer Channel, TeleFutura) and three other Gold Cup clashes; two MLS games; and Gol TV tracks the Spanish La Liga race between Barcelona and Real Madrid, airing both ... »0 Comments Games on TV Posted by Soccer America on Jun. 4, 9:00 a.m. This week's live televised soccer action kicks off on Tuesday with Brazil's friendly against Turkey (Fox) and Mexico's friendly against Paraguay (Telemundo). Then there's European Championship qualifying, Libertadores Cup semifinal action, the Gold Cup kickoff (the USA opens on Thursday) and MLS's Wednesday & Thursday night games. Montreal takes on Portland in USL's Friday night game. The weekend brings ... »0 Comments Soccer On TV Posted by Soccer America on Jun. 1, 9:00 a.m. Get an early start to your weekend soccer viewing with England's friendly against Brazil (3 pm ET on Fox). Friday night's USL's game (Fox) is an all-Canadian affair. MLS action on Saturday overlaps with the USA's friendly against China. Setanta USA and Fox Soccer Channel provide broadcasts of European Championship qualifying on Saturday.
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will this affect the CVC price? funny GS d/g CVC and now its up. funny how when it was 18.85 no D/G, after it goes down 5$ they d/g it, what ever. thanks! jmo
Sentiment: Hold
Think Cablevision as the eyes and ears for the big market makers and if you will "big brother" They compute storage data and these headends get the big picture before their subscribers. selling co. assets is usually a sign of foreboding times..probably find out six months out if this company is going to survive.I see too much at&t plus Verizon fios and all this smartphone technology stealing market share. BUT THE REAL CATALYST was those stupid reative annoying optimium commercials with that goofy clown idiot.
Sentiment: Sell