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The Neuroanatomical Organization of Projection Neurons Associated with Different Olfactory Bulb Pathways in the Sea Lamprey, Petromyzon marinus
Green, Warren W.
Basilious, Alfred
Dubuc, Réjean
Zielinski, Barbara S.
PLoS ONE
Although there is abundant evidence for segregated processing in the olfactory system across vertebrate taxa, the spatial relationship between the second order projection neurons (PNs) of olfactory subsystems connecting sensory input to higher brain structures is less clear. In the sea lamprey, there is tight coupling between olfaction and locomotion via PNs extending to the posterior tuberculum from the medial region of the olfactory bulb. This medial region receives peripheral input predominantly from the accessory olfactory organ. However, the axons from olfactory sensory neurons residing in the main olfactory epithelium extend to non-medial regions of the olfactory bulb, and the non-medial bulbar PNs extend their axons to the lateral pallium. It is not known if the receptive fields of the PNs in the two output pathways overlap; nor has the morphology of these PNs been investigated. In this study, retrograde labelling was utilized to investigate the PNs belonging to medial and non-medial projections. The dendrites and somata of the medial PNs were confined to medial glomerular neuropil, and dendrites of non-medial PNs did not enter this territory. The cell bodies and dendrites of the non-medial PNs were predominantly located below the glomeruli (frequently deeper in the olfactory bulb). While PNs in both locations contained single or multiple primary dendrites, the somal size was greater for medial than for non-medial PNs. When considered with the evidence-to-date, this study shows different neuroanatomical organization for medial olfactory bulb PNs extending to locomotor control centers and non-medial PNs extending to the lateral pallium in this vertebrate.
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
The Spinobulbar System in Lamprey
Buchanan, James T.
Einum, James F.
Brain research reviews
Locomotor networks in the spinal cord are controlled by descending systems which in turn receive feedback signals from ascending systems about the state of the locomotor networks. In lamprey, the ascending system consists of spinobulbar neurons which convey spinal network signals to the two descending systems, the reticulospinal and vestibulospinal neurons. Previous studies showed that spinobulbar neurons consist of both ipsilaterally- and contralaterally-projecting cells distributed at all rostrocaudal levels of the spinal cord, though most numerous near the obex. The axons of spinobulbar neurons ascend in the ventrolateral spinal cord and brainstem to the caudal mesencephalon and within the dendritic arbors of reticulospinal and vestibulospinal neurons. Compared to mammals, the ascending system in lampreys is more direct, consisting of excitatory and inhibitory monosynaptic inputs from spinobulbar neurons to reticulospinal neurons. The spinobulbar neurons are rhythmically active during fictive locomotion, representing a wide range of timing relationships with nearby ventral root bursts including those in phase, out of phase, and active during burst transitions between opposite ventral roots. The spinobulbar neurons are not simply relay cells because they can have mutual synaptic interactions with their reticulospinal neuron targets and they can have synaptic outputs to other spinal neurons. Spinobulbar neurons not only receive locomotor inputs but also receive direct inputs from primary mechanosensory neurons. Due to the relative simplicity of the lamprey nervous system and motor control system, the spinobulbar neurons and their interactions with reticulospinal neurons may be advantageous for investigating the general organization of ascending systems in the vertebrate.
lamprey; spinal cord; locomotion; spinobulbar; reticulospinal
Related Articles
Presynaptic G protein-coupled receptors dynamically modify vesicle fusion, synaptic cleft glutamate concentrations and motor behavior
Gerachshenko, Tatyana
Schwartz, Eric
Bleckert, Adam
Photowala, Huzefa
Seymour, Andrew
Alford, Simon
The Journal of neuroscience : the official journal of the Society for Neuroscience
Understanding how neuromodulators regulate behavior requires investigating their effects on functional neural systems, but also their underlying cellular mechanisms. Utilizing extensively characterized lamprey motor circuits, and the unique access to reticulospinal presynaptic terminals in the intact spinal cord that initiate these behaviours, we have investigated effects of presynaptic G protein-coupled receptors on locomotion from the systems level, to the molecular control of vesicle fusion. 5-HT inhibits neurotransmitter release via a Gβγ interaction with the SNARE complex that promotes kiss-and-run vesicle fusion. In the lamprey spinal cord we demonstrate that while presynaptic 5-HT receptors inhibit evoked neurotransmitter release from reticulospinal command neurons, their activation does not abolish locomotion, but rather modulates locomotor rhythms. Liberation of presynaptic Gβγ causes substantial inhibition of AMPA receptor-mediated synaptic responses, but leaves NMDA receptor-mediated components of neurotransmission largely intact. Because Gβγ binding to the SNARE complex is displaced by Ca2+-synaptotagmin binding, 5-HT-mediated inhibition displays Ca2+ sensitivity. We show that as Ca2+ accumulates presynaptically during physiological bouts of activity, 5-HT/Gβγ-mediated presynaptic inhibition is relieved leading to a frequency-dependent increase in synaptic concentrations of glutamate. This frequency dependent phenomenon mirrors a shift in the vesicle fusion mode and a recovery of AMPA receptor-mediated EPSCs from inhibition without a modification of NMDA receptor EPSCs. We conclude that activation of presynaptic 5-HT GPCRs state-dependently alters vesicle fusion properties to shift the weight of NMDA vs AMPA receptor-mediated responses at excitatory synapses. We have therefore identified a novel mechanism in which modification of vesicle fusion modes may profoundly alter locomotor behaviour.
fictive locomotion; kiss-and-run; G protein-coupled receptors; serotonin; presynaptic; Gβγ
Related Articles
A parallel cholinergic brainstem pathway for enhancing locomotor drive
Smetana, Roy
Juvin, Laurent
Dubuc, Réjean
Alford, Simon
Nature neuroscience
The brainstem locomotor system is believed to be organized serially from the mesencephalic locomotor region (MLR) to reticulospinal neurons, which in turn, project to locomotor neurons in the spinal cord. In contrast, we now identify in lampreys, brainstem muscarinoceptive neurons receiving parallel inputs from the MLR and projecting back to reticulospinal cells to amplify and extend durations of locomotor output. These cells respond to muscarine with extended periods of excitation, receive direct muscarinic excitation from the MLR, and project glutamatergic excitation to reticulospinal neurons. Targeted block of muscarine receptors over these neurons profoundly reduces MLR-induced excitation of reticulospinal neurons and markedly slows MLR-evoked locomotion. Their presence forces us to rethink the organization of supraspinal locomotor control, to include a sustained feedforward loop that boosts locomotor output.
Related Articles
Olfactory consciousness and gamma oscillation couplings across the olfactory bulb, olfactory cortex, and orbitofrontal cortex
Mori, Kensaku
Manabe, Hiroyuki
Narikiyo, Kimiya
Onisawa, Naomi
Frontiers in Psychology
The orbitofrontal cortex receives multi-modality sensory inputs, including olfactory input, and is thought to be involved in conscious perception of the olfactory image of objects. Generation of olfactory consciousness may require neuronal circuit mechanisms for the "binding" of distributed neuronal activities, with each constituent neuron representing a specific component of an olfactory percept. The shortest neuronal pathway for odor signals to reach the orbitofrontal cortex is olfactory sensory neuron—olfactory bulb—olfactory cortex—orbitofrontal cortex, but other pathways exist, including transthalamic pathways. Here, we review studies on the structural organization and functional properties of the shortest pathway, and propose a model of neuronal circuit mechanisms underlying the temporal bindings of distributed neuronal activities in the olfactory cortex. We describe a hypothesis that suggests functional roles of gamma oscillations in the bindings. This hypothesis proposes that two types of projection neurons in the olfactory bulb, tufted cells and mitral cells, play distinct functional roles in bindings at neuronal circuits in the olfactory cortex: tufted cells provide specificity-projecting circuits which send odor information with early-onset fast gamma synchronization, while mitral cells give rise to dispersedly-projecting feed-forward binding circuits which transmit the response synchronization timing with later-onset slow gamma synchronization. This hypothesis also suggests a sequence of bindings in the olfactory cortex: a small-scale binding by the early-phase fast gamma synchrony of tufted cell inputs followed by a larger-scale binding due to the later-onset slow gamma synchrony of mitral cell inputs. We discuss that behavioral state, including wakefulness and sleep, regulates gamma oscillation couplings across the olfactory bulb, olfactory cortex, and orbitofrontal cortex.
olfactory cortex; orbitofrontal cortex; tufted and mitral cells; olfactory bulb; olfactory consciousness; gamma synchronization
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.
Related Articles
Glomerular interactions in olfactory processing channels of the antennal lobes
Heinbockel, Thomas
Shields, Vonnie D. C.
Reisenman, Carolina E.
Journal of comparative physiology. A, Neuroethology, sensory, neural, and behavioral physiology
An open question in olfactory coding is the extent of interglomerular connectivity: do olfactory glomeruli and their neurons regulate the odorant responses of neurons innervating other glomeruli? In the olfactory system of the moth Manduca sexta, the response properties of different types of antennal olfactory receptor cells are known. Likewise, a subset of antennal lobe glomeruli has been functionally characterized and the olfactory tuning of their innervating neurons identified. This provides a unique opportunity to determine functional interactions between glomeruli of known input, specifically, (1) glomeruli processing plant odors and (2) glomeruli activated by antennal stimulation with pheromone components of conspecific females. Several studies describe reciprocal inhibitory effects between different types of pheromone-responsive projection neurons suggesting lateral inhibitory interactions between pheromone component-selective glomerular neural circuits. Furthermore, antennal lobe projection neurons that respond to host plant volatiles and innervate single, ordinary glomeruli are inhibited during antennal stimulation with the female's sex pheromone. The studies demonstrate the existence of lateral inhibitory effects in response to behaviorally significant odorant stimuli and irrespective of glomerular location in the antennal lobe. Inhibitory interactions are present within and between olfactory subsystems (pheromonal and non-pheromonal subsystems), potentially to enhance contrast and strengthen odorant discrimination.
Electrophysiology; Glomerulus; Neural coding; Olfaction; Pheromone
Related Articles
In the lamprey, spinal locomotor activity can be initiated by pharmacological microstimulation in several brain areas: rostrolateral rhombencephalon (RLR); dorsolateral mesencephalon (DLM); ventromedial diencephalon (VMD); and reticular nuclei. During DLM- or VMD-initiated locomotor activity in in vitro brain/spinal cord preparations, application of a solution that focally depressed neuronal activity in reticular nuclei often attenuated or abolished the locomotor rhythm. Electrical microstimulation in the DLM or VMD elicited synaptic responses in reticulospinal (RS) neurons, and close temporal stimulation in both areas evoked responses that summated and could elicit action potentials when neither input alone was sufficient. During RLR-initiated locomotor activity, focal application of a solution that depressed neuronal activity in the DLM or VMD abolished or attenuated the rhythm. These new results suggest that neurons in the RLR project rostrally to locomotor areas in the DLM and VMD. These latter areas then appear to project caudally to RS neurons, which probably integrate the synaptic inputs from both areas and activate the spinal locomotor networks. These pathways are likely to be important components of the brain neural networks for the initiation of locomotion and have parallels to locomotor command systems in higher vertebrates.
locomotion; command; descending; reticulospinal; central pattern generators
Related Articles
Muscarinic Receptor Activation Elicits Sustained, Recurring Depolarizations in Reticulospinal Neurons
Smetana, R. W.
Alford, S.
Dubuc, R.
Journal of neurophysiology
In lampreys, brain stem reticulospinal (RS) neurons constitute the main descending input to the spinal cord and activate the spinal locomotor central pattern generators. Cholinergic nicotinic inputs activate RS neurons, and consequently, induce locomotion. Cholinergic muscarinic agonists also induce locomotion when applied to the brain stem of birds. This study examined whether bath applications of muscarinic agonists could activate RS neurons and initiate motor output in lampreys. Bath applications of 25 μM muscarine elicited sustained, recurring depolarizations (mean duration of 5.0 ± 0.5 s recurring with a mean period of 55.5 ± 10.3 s) in intracellularly recorded rhombencephalic RS neurons. Calcium imaging experiments revealed that muscarine induced oscillations in calcium levels that occurred synchronously within the RS neuron population. Bath application of TTX abolished the muscarine effect, suggesting the sustained depolarizations in RS neurons are driven by other neurons. A series of lesion experiments suggested the caudal half of the rhombencephalon was necessary. Microinjections of muscarine (75 μM) or the muscarinic receptor (mAchR) antagonist atropine (1 mM) lateral to the rostral pole of the posterior rhombencephalic reticular nucleus induced or prevented, respectively, the muscarinic RS neuron response. Cells immunoreactive for muscarinic receptors were found in this region and could mediate this response. Bath application of glutamatergic antagonists (6-cyano-7-nitroquinoxaline-2,3-dione/D-2-amino-5-phosphonovaleric acid) abolished the muscarine effect, suggesting that glutamatergic transmission is needed for the effect. Ventral root recordings showed spinal motor output coincides with RS neuron sustained depolarizations. We propose that unilateral mAchR activation on specific cells in the caudal rhombencephalon activates a circuit that generates synchronous sustained, recurring depolarizations in bilateral populations of RS neurons.
Related Articles
Aversive cues fail to activate fos expression in the asymmetric olfactory-habenula pathway of zebrafish
deCarvalho, Tagide N.
Akitake, Courtney M.
Thisse, Christine
Thisse, Bernard
Halpern, Marnie E.
Frontiers in Neural Circuits
The dorsal habenular nuclei of the zebrafish epithalamus have become a valuable model for studying the development of left-right (L-R) asymmetry and its function in the vertebrate brain. The bilaterally paired dorsal habenulae exhibit striking differences in size, neuroanatomical organization, and molecular properties. They also display differences in their efferent connections with the interpeduncular nucleus (IPN) and in their afferent input, with a subset of mitral cells distributed on both sides of the olfactory bulb innervating only the right habenula. Previous studies have implicated the dorsal habenulae in modulating fear/anxiety responses in juvenile and adult zebrafish. It has been suggested that the asymmetric olfactory-habenula pathway (OB-Ha), revealed by selective labeling from an lhx2a:YFP transgene, mediates fear behaviors elicited by alarm pheromone. Here we show that expression of the fam84b gene demarcates a unique region of the right habenula that is the site of innervation by lhx2a:YFP-labeled olfactory axons. Upon ablation of the parapineal, which normally promotes left habenular identity; the fam84b domain is present in both dorsal habenulae and lhx2a:YFP-labeled olfactory bulb neurons form synapses on the left and the right side. To explore the relevance of the asymmetric olfactory projection and how it might influence habenular function, we tested activation of this pathway using odorants known to evoke behaviors. We find that alarm substance or other aversive odors, and attractive cues, activate fos expression in subsets of cells in the olfactory bulb but not in the lhx2a:YFP expressing population. Moreover, neither alarm pheromone nor chondroitin sulfate elicited fos activation in the dorsal habenulae. The results indicate that L-R asymmetry of the epithalamus sets the directionality of olfactory innervation, however, the lhx2a:YFP OB-Ha pathway does not appear to mediate fear responses to aversive odorants.
behavior; asymmetry; alarm pheromone; fos; fam84b
Related Articles
Perception of Odors Linked to Precise Timing in the Olfactory System
Rebello, Michelle R.
McTavish, Thomas S.
Willhite, David C.
Short, Shaina M.
Shepherd, Gordon M.
Verhagen, Justus V.
PLoS Biology
The temporal dynamics of glomeruli activity can be behaviorally discerned by mice down to 13 milliseconds.
While the timing of neuronal activity in the olfactory bulb (OB) relative to sniffing has been the object of many studies, the behavioral relevance of timing information generated by patterned activation within the bulbar response has not been explored. Here we show, using sniff-triggered, dynamic, 2-D, optogenetic stimulation of mitral/tufted cells, that virtual odors that differ by as little as 13 ms are distinguishable by mice. Further, mice are capable of discriminating a virtual odor movie based on an optically imaged OB odor response versus the same virtual odor devoid of temporal dynamics—independently of the sniff-phase. Together with studies showing the behavioral relevance of graded glomerular responses and the response timing relative to odor sampling, these results imply that the mammalian olfactory system is capable of very high transient information transmission rates.
Author Summary
Olfactory receptor neurons respond to odors in the olfactory epithelium located in the nasal cavity in mammals. Each olfactory receptor neuron expresses only one olfactory receptor, out of several hundred encoded in the mammalian genome. Olfactory receptor neurons expressing the same olfactory receptor are scattered throughout the olfactory epithelium; however, their axons converge in one of thousands of glomeruli in the olfactory bulb. The glomeruli are the first neural relay station in the olfactory system, where olfactory receptor neurons transmit olfactory information to mitral cells. It is well established that different odors evoke different spatial patterns across the glomeruli. It is believed that the more similar the patterns, the more similar the evoked odor perceptions. Glomeruli also are activated in odor-specific sequences in time. These dynamics could increase the amount of information about odors by immense amounts. We used transgenic mice, whose mitral cells were made responsive to light, and asked how well they could discriminate the temporal dynamics of simple spatial patterns of light presented to the olfactory bulb after each sniff. Mice could detect the presence of temporal dynamics down to 13 ms, which provides ample resolution for them to be able to detect the dynamics in response to actual odors. Mice could also discern whether virtual odors, based on actual olfactory bulb activity, were dynamic or static and did so without reference to exact sniff-time. We conclude that both the spatial glomerular activity patterns and the temporal dynamics thereof are used in the mammalian olfactory system to encode odors.
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Sox10-dependent neural crest origin of olfactory microvillous neurons in zebrafish
Saxena, Ankur
Peng, Brian N
Bronner, Marianne E
The sense of smell in vertebrates is detected by specialized sensory neurons derived from the peripheral nervous system. Classically, it has been presumed that the olfactory placode forms all olfactory sensory neurons. In contrast, we show that the cranial neural crest is the primary source of microvillous sensory neurons within the olfactory epithelium of zebrafish embryos. Using photoconversion-based fate mapping and live cell tracking coupled with laser ablation, we followed neural crest precursors as they migrated from the neural tube to the nasal cavity. A subset that coexpressed Sox10 protein and a neurogenin1 reporter ingressed into the olfactory epithelium and differentiated into microvillous sensory neurons. Timed loss-of-function analysis revealed a critical role for Sox10 in microvillous neurogenesis. Taken together, these findings directly demonstrate a heretofore unknown contribution of the cranial neural crest to olfactory sensory neurons in zebrafish and provide important insights into the assembly of the nascent olfactory system.
DOI: http://dx.doi.org/10.7554/eLife.00336.001
eLife digest
Neurons have crucial roles in both the peripheral and central nervous systems. The role of the neurons in the sensory organs (the eyes, ears, and nose) is to sense stimuli—including light, sound, and odor—and transmit this sensory information to the neurons of the central nervous system for processing. The first step in sensing an odor relies on the peripheral nerves of the olfactory epithelium. This tissue, which lines the inside of the nasal cavity, includes two main types of olfactory sensory neurons: ciliated sensory neurons that detect volatile or easily evaporated substances and microvillous sensory neurons that detect pheromones, nucleotides, and/or amino acids.
During vertebrate embryogenesis, an embryo develops three distinct germ layers, the ectoderm, mesoderm, and endoderm, each of which gives rise to the different tissues of the body. The ectoderm has three parts—the external ectoderm, the neural crest, and the neural tube—and together they give rise to the neurons of the peripheral and central nervous systems. The neurons within the eye and ear are known to originate from a thickened portion of the ectoderm, and it has been proposed that olfactory neurons develop in a similar manner.
Now, Saxena et al. show that, unlike what happens in the eye and ear, some olfactory sensory neurons originate from the neural crest. By studying the development of the olfactory system in zebrafish, Saxena et al. discovered that microvillous neurons, but not ciliated neurons, develop from neural crest cells, and that the transcription factor Sox10 is critical for the development of microvillous neurons. By establishing that neural crest cells are involved in the development of a substantial proportion of olfactory sensory neurons, this work sets the stage for future studies of olfactory nerve growth and regeneration. It may also assist researchers working on anosmia (the inability to smell).
DOI: http://dx.doi.org/10.7554/eLife.00336.002
neural crest migration; olfactory development; microvillous sensory neurons; neurogenesis; Zebrafish
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Sex-specific processing of social cues in the medial amygdala
Bergan, Joseph F
Ben-Shaul, Yoram
Dulac, Catherine
Animal–animal recognition within, and across species, is essential for predator avoidance and social interactions. Despite its essential role in orchestrating responses to animal cues, basic principles of information processing by the vomeronasal system are still unknown. The medial amygdala (MeA) occupies a central position in the vomeronasal pathway, upstream of hypothalamic centers dedicated to defensive and social responses. We have characterized sensory responses in the mouse MeA and uncovered emergent properties that shed new light onto the transformation of vomeronasal information into sex- and species-specific responses. In particular, we show that the MeA displays a degree of stimulus selectivity and a striking sexually dimorphic sensory representation that are not observed in the upstream relay of the accessory olfactory bulb (AOB). Furthermore, our results demonstrate that the development of sexually dimorphic circuits in the MeA requires steroid signaling near the time of puberty to organize the functional representation of sensory stimuli.
DOI: http://dx.doi.org/10.7554/eLife.02743.001
eLife digest
Many animals emit and detect chemicals known as pheromones to communicate with other members of their own species. Animals also rely on chemical signals from other species to warn them, for example, that a predator is nearby. Many of these chemical signals—which are present in sweat, tears, urine, and saliva—are detected by a structure called the vomeronasal organ, which is located at the base of the nasal cavity.
When this organ detects a particular chemical signal, it broadcasts this information to a network of brain regions that generates an appropriate behavioral response. Two structures within this network, the accessory olfactory bulb and the medial amygdala, play an important role in modifying this signal before it reaches its final destination—a region of the brain called the hypothalamus. Activation of the hypothalamus by the signal triggers changes in the animal's behavior. Although the anatomical details of this pathway have been widely studied, it is not clear how information is actually transmitted along it.
Now, Bergan et al. have provided insights into this process by recording signals in the brains of anesthetized mice exposed to specific stimuli. Whereas neurons in the accessory olfactory bulb responded similarly in male and female mice, those in the medial amygdala showed a preference for female urine in male mice, and a preference for male urine in the case of females. This is the first direct demonstration of differences in sensory processing in the brains of male and female mammals.
These differences are thought to result from the actions of sex hormones, particularly estrogen, on brain circuits during development. Consistent with this, neurons in the medial amygdala of male mice with reduced levels of estrogen showed a reduced preference for female urine compared to control males. Similarly, female mice that had been previously exposed to high levels of estrogen as pups showed a reduced preference for male urine compared to controls.
In addition to increasing understanding of how chemical signals—including pheromones—influence the responses of rodents to other animals, the work of Bergan et al. has provided clues to the neural mechanisms that underlie sex-specific differences in behaviors.
DOI: http://dx.doi.org/10.7554/eLife.02743.002
medial amygdala; sensory representation; sexual dimorphism; vomeronasal system; pheromones; behavior; mouse
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
Effect of sniffing on the temporal structure of mitral/tufted cell output from the olfactory bulb
Carey, Ryan M.
Wachowiak, Matt
The Journal of neuroscience : the official journal of the Society for Neuroscience
Neural activity underlying odor representations in the mammalian olfactory system is strongly patterned by respiratory behavior; these dynamics are central to many models of olfactory information processing. We have previously found that sensory inputs to the olfactory bulb change both their magnitude and temporal structure as a function of sniff frequency. Here, we asked how sniff frequency affects responses of mitral/tufted (MT) cells – the principal olfactory bulb output neuron. We recorded from MT cells in anesthetized rats while reproducing sniffs recorded previously from awake animals and varying sniff frequency. The dynamics of a sniff-evoked response were consistent from sniff to sniff but varied across cells. Compared to the dynamics of receptor neuron activation by the same sniffs, the MT response was shorter and faster, reflecting a temporal sharpening of sensory inputs. Increasing sniff frequency led to moderate attenuation of MT response magnitude and significant changes in the temporal structure of the sniff-evoked MT cell response. Most MT cells responded with a shorter duration and shorter rise-time spike burst as sniff frequency increased, reflecting increased temporal sharpening of inputs by the olfactory bulb. These temporal changes were necessary and sufficient to maintain respiratory modulation in the MT cell population across the range of sniff frequencies expressed during behavior. These results suggest that the input-output relationship in the olfactory bulb varies dynamically as a function of sniff frequency, and that one function of the postsynaptic network is to maintain robust temporal encoding of odor information across different odor sampling strategies.
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
Carbon Dioxide Sensing Modulates Lifespan and Physiology in Drosophila
Poon, Peter C.
Kuo, Tsung-Han
Linford, Nancy J.
Roman, Gregg
Pletcher, Scott D.
PLoS Biology
Loss of an olfactory receptor that senses carbon dioxide increases longevity and fecundity and alters overall physiology in fruit flies.
For nearly all life forms, perceptual systems provide access to a host of environmental cues, including the availability of food and mates as well as the presence of disease and predators. Presumably, individuals use this information to assess the current and future states of the environment and to enact appropriate developmental, behavioral, and regulatory decisions. Recent work using the nematode worm, Caenorhabditis elegans, and the fruit fly, Drosophila melanogaster, has established that aging is subject to modulation through neurosensory systems and that this regulation is evolutionarily conserved. To date, sensory manipulations shown to impact Drosophila aging have involved general loss of function or manipulation of complex stimuli. We therefore know little about the specific inputs, sensors, or associated neural circuits that affect these life and death decisions. We find that a specialized population of olfactory neurons that express receptor Gr63a (a component of the olfactory receptor for gaseous phase CO2) affects fly lifespan and physiology. Gr63a loss of function leads to extended lifespan, increased fat deposition, and enhanced resistance to some (but not all) environmental stresses. Furthermore, we find that the reduced lifespan that accompanies exposure to odors from live yeast is dependent on Gr63a. Together these data implicate a specific sensory cue (CO2) and its associated receptor as having the ability to modulate fly lifespan and alter organism stress response and physiology. Because Gr63a is expressed in a well-defined population of neurons, future work may now be directed at dissecting more complex neurosensory and neuroendocrine circuits that modulate aging in Drosophila.
Author Summary
Sensory inputs, including taste and smell, can modulate lifespan in organisms such as fruit flies and nematodes. For example, the smell of live yeast is sufficient to accelerate aging in fruit flies that are nutrient restricted. However, the sensory pathways and specific olfactory cues that modulate aging are unknown. Here, we show that the olfactory receptor for carbon dioxide, Gustatory Receptor 63a (Gr63a), plays a role in determining longevity in Drosophila melanogaster. Flies lacking Gr63a function live longer, have increased fat storage, and exhibit greater reproductive output than control flies. Ablation of the neurons that express Gr63a also results in long-lived flies. Notably, the smell of live yeast does not affect the lifespan of flies that lack a functional Gr63a receptor, which suggests that carbon dioxide is a key regulatory molecule of this complex odor. Because Gr63a expression is restricted to a very select population of neurons, these results implicate a specific neural circuit in the modulation of fly lifespan.
Related Articles
Spinal and supraspinal postural networks
Deliagina, T.G.
Beloozerova, I.N.
Zelenin, P.V.
Orlovsky, G.N.
Brain research reviews
Different species maintain a particular body orientation in space (upright in humans, dorsal-side-up in quadrupeds, fish and lamprey) due to the activity of a closed-loop postural control system. We will discuss operation of spinal and supraspinal postural networks studied in a lower vertebrate (lamprey) and in two mammals (rabbit and cat).
In the lamprey, the postural control system is driven by vestibular input. The key role in the postural network belongs to the reticulospinal (RS) neurons. Due to vestibular input, deviation from the stabilized body orientation in any (roll, pitch, yaw) plane leads to generation of RS commands, which are sent to the spinal cord and cause postural correction. For each of the planes, there are two groups of RS neurons responding to rotation in the opposite directions; they cause a turn opposite to the initial one. The command transmitted by an individual RS neuron causes the motor response, which contributes to the correction of posture. In each plane, the postural system stabilizes the orientation at which the antagonistic vestibular reflexes compensate for each other. Thus, in lamprey the supraspinal networks play a crucial role in stabilization of body orientation, and the function of the spinal networks is transformation of supraspinal commands into the motor pattern of postural corrections.
In terrestrial quadrupeds, the postural system stabilizing the trunk orientation in the transversal plane was analyzed. It consists of two relatively independent sub-systems stabilizing orientation of the anterior and posterior parts of the trunk. They are driven by somatosensory input from limb mechanoreceptors. Each sub-system consists of two closed-loop mechanisms – spinal and spino-supraspinal. Operation of the supraspinal networks was studied by recording the posture-related activity of corticospinal neurons. The postural capacity of spinal networks was evaluated in animals with lesions to the spinal cord. Relative contribution of spinal and supraspinal mechanisms to the stabilization of trunk orientation is discussed.
Related Articles
Functional organization of a neural network for aversive olfactory learning in Caenorhabditis elegans
Ha, Heon-ick
Hendricks, Michael
Shen, Yu
Gabel, Christopher V.
Fang-Yen, Christopher
Qin, Yuqi
Colón-Ramos, Daniel
Shen, Kang
Samuel, Aravinthan D.T.
Zhang, Yun
Many animals use their olfactory systems to learn to avoid dangers, but how neural circuits encode naïve and learned olfactory preferences, and switch between those preferences, is poorly understood. Here, we map an olfactory network, from sensory input to motor output, which regulates the learned olfactory aversion of Caenorhabditis elegans for the smell of pathogenic bacteria. Naïve animals prefer smells of pathogens but animals trained with pathogens lose this attraction. We find that two different neural circuits subserve these preferences, with one required for the naïve preference and the other specifically for the learned preference. Calcium imaging and behavioral analysis reveal that the naïve preference reflects the direct transduction of the activity of olfactory sensory neurons into motor response, whereas the learned preference involves modulations to signal transduction to downstream neurons to alter motor response. Thus, two different neural circuits regulate a behavioral switch between naïve and learned olfactory preferences.
Related Articles
Contribution of supraspinal systems to generation of automatic postural responses
Deliagina, Tatiana G.
Beloozerova, Irina N.
Orlovsky, Grigori N.
Zelenin, Pavel V.
Frontiers in Integrative Neuroscience
Different species maintain a particular body orientation in space due to activity of the closed-loop postural control system. In this review we discuss the role of neurons of descending pathways in operation of this system as revealed in animal models of differing complexity: lower vertebrate (lamprey) and higher vertebrates (rabbit and cat). In the lamprey and quadruped mammals, the role of spinal and supraspinal mechanisms in the control of posture is different. In the lamprey, the system contains one closed-loop mechanism consisting of supraspino-spinal networks. Reticulospinal (RS) neurons play a key role in generation of postural corrections. Due to vestibular input, any deviation from the stabilized body orientation leads to activation of a specific population of RS neurons. Each of the neurons activates a specific motor synergy. Collectively, these neurons evoke the motor output necessary for the postural correction. In contrast to lampreys, postural corrections in quadrupeds are primarily based not on the vestibular input but on the somatosensory input from limb mechanoreceptors. The system contains two closed-loop mechanisms – spinal and spino-supraspinal networks, which supplement each other. Spinal networks receive somatosensory input from the limb signaling postural perturbations, and generate spinal postural limb reflexes. These reflexes are relatively weak, but in intact animals they are enhanced due to both tonic supraspinal drive and phasic supraspinal commands. Recent studies of these supraspinal influences are considered in this review. A hypothesis suggesting common principles of operation of the postural systems stabilizing body orientation in a particular plane in the lamprey and quadrupeds, that is interaction of antagonistic postural reflexes, is discussed.
balance control; postural reflexes; reticulospinal neurons; pyramidal tract neurons; rubrospinal neurons; unilateral labyrinthectomy; galvanic vestibular stimulation
Related Articles
Reorganization of neuronal circuits of the central olfactory system during postprandial sleep
Yamaguchi, Masahiro
Manabe, Hiroyuki
Murata, Koshi
Mori, Kensaku
Frontiers in Neural Circuits
Plastic changes in neuronal circuits often occur in association with specific behavioral states. In this review, we focus on an emerging view that neuronal circuits in the olfactory system are reorganized along the wake-sleep cycle. Olfaction is crucial to sustaining the animals' life, and odor-guided behaviors have to be newly acquired or updated to successfully cope with a changing odor world. It is therefore likely that neuronal circuits in the olfactory system are highly plastic and undergo repeated reorganization in daily life. A remarkably plastic feature of the olfactory system is that newly generated neurons are continually integrated into neuronal circuits of the olfactory bulb (OB) throughout life. New neurons in the OB undergo an extensive selection process, during which many are eliminated by apoptosis for the fine tuning of neuronal circuits. The life and death decision of new neurons occurs extensively during a short time window of sleep after food consumption (postprandial sleep), a typical daily olfactory behavior. We review recent studies that explain how olfactory information is transferred between the OB and the olfactory cortex (OC) along the course of the wake-sleep cycle. Olfactory sensory input is effectively transferred from the OB to the OC during waking, while synchronized top-down inputs from the OC to the OB are promoted during the slow-wave sleep. We discuss possible neuronal circuit mechanisms for the selection of new neurons in the OB, which involves the encoding of olfactory sensory inputs and memory trace formation during waking and internally generated activities in the OC and OB during subsequent sleep. The plastic changes in the OB and OC are well coordinated along the course of olfactory behavior during wakefulness and postbehavioral rest and sleep. We therefore propose that the olfactory system provides an excellent model in which to understand behavioral state-dependent plastic mechanisms of the neuronal circuits in the brain.
olfactory bulb; olfactory cortex; adult neurogenesis; cell elimination; behavioral state; slow-wave sleep; sharp waves; sensory experience
Related Articles
Movement- and behavioral state–dependent activity of pontine reticulospinal neurons
Thankachan, Stephen
Fuller, Patrick M.
Lu, Jun
Forty-five years ago Shik and colleagues were the first to demonstrate that electrical stimulation of the dorsal pontine reticular formation induced fictive locomotion in decerebrate cats. This supraspinal motor site was subsequently termed the "mesencephalic locomotor region (MLR)". Cholinergic neurons of the pedunculopontine tegmental nucleus (PPT) have been suggested to form, or at least comprise in part, the neuroanatomical basis for the MLR, but direct evidence is lacking. In an effort to clarify the location and activity profiles of pontine reticulospinal neurons supporting locomotor behaviors, we employed in the present study a retrograde tracing method in combination with single unit recordings and antidromic spinal cord stimulation as well as characterized the locomotor- and behavioral state-dependent activities of both reticulospinal and non-reticulospinal neurons. The retrograde labeling and antidromic stimulation responses suggested a candidate group of reticulospinal neurons that were non-cholinergic and located just medial to the PPT cholinergic neurons and ventral to the cuneiform nucleus (CnF). Unit recordings from these reticulopsinal neurons in freely behaving animals revealed that the preponderance of neurons fired in relation to motor behaviors and that some of these neurons were also active during REM sleep. By contrast, non-reticulospinal neurons, which likely included cholinergic neurons, did not exhibit firing activity in relation to motor behaviors. In summary, the present study provides neuroanatomical and electrophysiological evidence that non-cholinergic, pontine reticulospinal neurons may constitute the major component of the long-sought neuroanatomic MLR in mammals.
mesencephalic locomotor region; rat; single unit activity; sleep-wake REM sleep
Related Articles
Changes in Maternal Gene Expression in Olfactory Circuits in the Immediate Postpartum Period
Canavan, Sofija V.
Mayes, Linda C.
Treloar, Helen B.
Frontiers in Psychiatry
Regulation of maternal behavior in the immediate postpartum period involves neural circuits in reward and homeostasis systems responding to cues from the newborn. Our aim was to assess one specific regulatory mechanism: the role that olfaction plays in the onset and modulation of parenting behavior. We focused on changes in gene expression in olfactory brain regions, examining nine genes found in previous knockout studies to be necessary for maternal behavior. Using a quantitative PCR (qPCR)-based approach, we assessed changes in gene expression in response to exposure to pups in 11 microdissected olfactory brain regions. Over the first postpartum days, all nine genes were detected in all 11 regions (at differing levels) and their expression changed in response to pup exposure. As a general trend, five genes (Dbh, Esr1, FosB, Foxb1, and Oxtr) were found to decrease their expression in most of the olfactory regions examined, while two genes (Mest and Prlr) were found to increase expression. Nos1 and Peg3 levels remained relatively stable except in the accessory olfactory bulb (AOB), where greater than fourfold increases in expression were observed. The largest magnitude expression changes in this study were found in the AOB, which mediates a variety of olfactory cues that elicit stereotypic behaviors such as mating and aggression as well as some non-pheromone odors. Previous analyses of null mice for the nine genes assessed here have rarely examined olfactory function. Our data suggest that there may be olfactory effects in these null mice which contribute to the observed maternal behavioral phenotypes. Collectively, these data support the hypothesis that olfactory processing is an important sensory regulator of maternal behavior.
olfactory bulb; accessory olfactory bulb; olfactory tubercle; piriform cortex; entorhinal cortex; amygdala; hippocampus
Related Articles
Projections from the posterolateral olfactory amygdala to the ventral striatum: neural basis for reinforcing properties of chemical stimuli
Ubeda-Bañon, Isabel
Novejarque, Amparo
Mohedano-Moriano, Alicia
Pro-Sistiaga, Palma
de la Rosa-Prieto, Carlos
Insausti, Ricardo
Martinez-Garcia, Fernando
Lanuza, Enrique
Martinez-Marcos, Alino
BMC Neuroscience
Vertebrates sense chemical stimuli through the olfactory receptor neurons whose axons project to the main olfactory bulb. The main projections of the olfactory bulb are directed to the olfactory cortex and olfactory amygdala (the anterior and posterolateral cortical amygdalae). The posterolateral cortical amygdaloid nucleus mainly projects to other amygdaloid nuclei; other seemingly minor outputs are directed to the ventral striatum, in particular to the olfactory tubercle and the islands of Calleja.
Although the olfactory projections have been previously described in the literature, injection of dextran-amines into the rat main olfactory bulb was performed with the aim of delimiting the olfactory tubercle and posterolateral cortical amygdaloid nucleus in our own material. Injection of dextran-amines into the posterolateral cortical amygdaloid nucleus of rats resulted in anterograde labeling in the ventral striatum, in particular in the core of the nucleus accumbens, and in the medial olfactory tubercle including some islands of Calleja and the cell bridges across the ventral pallidum. Injections of Fluoro-Gold into the ventral striatum were performed to allow retrograde confirmation of these projections.
The present results extend previous descriptions of the posterolateral cortical amygdaloid nucleus efferent projections, which are mainly directed to the core of the nucleus accumbens and the medial olfactory tubercle. Our data indicate that the projection to the core of the nucleus accumbens arises from layer III; the projection to the olfactory tubercle arises from layer II and is much more robust than previously thought. This latter projection is directed to the medial olfactory tubercle including the corresponding islands of Calleja, an area recently described as critical node for the neural circuit of addiction to some stimulant drugs of abuse.
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My DS is 8 weeks tomorrow. He has issues with gas and i think he might have silent reflux (going to talk with dr at 2 month appt next week). I was trying to transition him from sleeping/napping in rock & play or swing into his crib when the reflux seemed to flare up from him trying to sleep flat. And he sometimes sleeps good in rock & play but other times his gas wakes him up. My question is what the heck am i supposed to do to get him to sleep/nap when his gas gives him so much trouble? And what about the reflux? I assume i shouldn't try the crib until i find out about the reflux and get him on meds?
Help, DS doesn't sleep/nap well
Posted 01/04/2013
Comments (2)
Posted 01/04/2013
hey, not sure i can offer much in terms of answers to your questions but just wanted to let you know that my DS had the same issues in his first few months. he's almost 5 months now and gas and reflux are no longer causing any sleep problems whatsoever. just wanted to let you know that they will resolve themselves -- and hopefully sooner rather than later.
re your questions, i'd hold off on trying the crib. for reflux, there really isn't much you can do other than having baby sleep on an incline, giving meds and trying to not lie him down too soon after feeding. for the gas, i never found any tricks -- tried probiotics for baby, changing my diet, nothing seemed to help.
netsirk203 · Original Poster
Posted 01/04/2013
Thanks! At least i know it will be better soon. One way or another!

subset: commoncrawl


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Key things you need to know aboutMarilyn Mullane:
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9 Facts About Hospital Quality
Patients' risk of dying during a hospital stay for heart bypass surgery could be 87.4% lower, on average, at a hospital rated 5 stars compared to a hospital rated 1 star for that procedure. Choosing the right hospital is critical to your health. View Slideshow
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Board members generously give of their time, talent and leadership, as they serve in an unpaid capacity.
The Board comprises nine members appointed by Harris County Commissioners' Court. Each member serves a term of two years.
Elvin Franklin Jr., chairDaisy Stiner, vice chair
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Dr. Donald R. Butts
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Lawrence D. Finder
Dr. Kimberly Monday
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E. Dale Wortham
Mailing Address:P.O. Box 66769Houston, Texas 77266-6769
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Premium Bare Steel Front Winch Bumper (NT04-H1751-B) by Fab Fours®. Building on our reputation for the highest quality and design in the truck market, the Fab Fours winch bumper focuses on strength and durability. This bumper is designed to be the ultimate in functionally-stylish front end protection while also providing more tire clearance to eliminate rubbing. Key features include the integrated fog and turn lights, D-Ring mounts and ability to accommodate up to 12,000 lb. winches. Available in either a Full Guard, Pre-runner Guard or No Guard Bumper.
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Unilever is to reposition its Peperami brand later this year in a bid to portray it as a healthy snack choice.
The move will coincide with the launch of two additional lines of the product, with a barbecue variant of the original Peperami set to launch in April, and a beef jerky product to be released in September. A new strapline for the brand is also set to be rolled out, marking the end of the current "Peperami: it's a bit of an animal" tagline, which was first introduced in 1993.
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Welcome to the UFO!
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The Untenured Faculty Organization was formed in 2009 to provide support and opportunities for professional development to pre-tenure faculty. We meet regularly and offer frequent workshops on matters of interest to junior faculty such as the RTP process, grant funding, teaching, technology and other topics.
Click on the links to see UFO officers, UFO events, plus the agendas and minutes of UFO meetings.
Upcoming Events
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Literature & the arts
Literature and the arts play a vital role in the culture and traditions of New England. Western Massachusetts in particular has had a rich history of fostering writers and poets, musicians, dancers, and actors. The Department of Special Collections and University Archives seeks to document not only the lives and work of writers and performers in our region, but the creative and artistic process; showing not just the inspiration, but the perspiration as well.
View all collections in Literature and the arts
Significant collections
- Arts and arts management
- Significant collections under the National Arts Policy Archive and Library include materials from the National Endowment for the Arts, the National Assembly of State Arts Agencies, and the Arts Extension Service.
- Poetry
- SCUA houses significant collections for the poets Robert Francis, Madeleine de Frees, and Anne Halley, as well as small collections for William Carlos Williams and Wallace Stevens. The records of the Massachusetts Review are an important literary resource.
- Prose writing
- Collections of note include the papers of writers William J. Lederer (author of The Ugly American, Nation of Sheep, and Their Own Worst Enemy), William Manchester (The Death of a President and American Caesar), Jodi Picoult (many novels from Songs of the Humpback Whale in 1992 to The Storyteller in 2013), Mary Doyle Curran (The Parish and the Hill).
- Journalism
- Journalists and photojournalists associated with traditional print and new media, including an important collection for the Liberation News Service, a media service for the alternative press, and the Social Change Periodicals Collection, which includes alternative and radical small press publications. The papers of Sidney Topol provide insight into the technical development of cable television.
- Literary criticism and linguistics
- The papers of literary scholars associated with the University; records of the Massachusetts Review.
- Performing arts
- The vibrant performing arts community in western Massachusetts is well represented in SCUA through groups ranging from the Arcadia Players Baroque music ensemble to theater troupes such as Double Edge Theater, the Valley Light Opera, and the New World Theater. Among the most significant national collections are the Roberta Uno Asian Women Playwrights Collection and the papers of African American expatriate actor and director Gordon Heath, while the James Ellis Theatre Collection includes nearly 8,000 printed volumes on the English and American stage, 1750-1915, along with numerous broadsides, graphics, and some manuscript materials. Musical collections include the papers of Philip Bezanson and Charles Bestor, the score collection of Julian Olevsky, and the Katanka Fraser Political Music Collection.
Within its holdings, SCUA houses collections of the published works of W.E.B. Du Bois, Robert Francis, Anne Halley, William J. Lederer, William Manchester, Thomas Mann, William Morris, Wallace Stevens, and William Butler Yeats, as well as the personal poetry libraries of Halley, Francis, and Stevens. The department also has an extensive collection of Science Fiction magazine fiction and Scottish literature.
Massachusetts AFL-CIO
Massachusetts AFL-CIO Records, 1902-1995.
Call no.: MS 369
Formed in 1887 as the Massachusetts branch of the American Federation of Labor, the Massachusetts AFL-CIO currently represents the interests of over 400,000 working people in the Commonwealth. Like its parent organization, the national AFL-CIO, the Mass. AFL-CIO is an umbrella organization, a union of unions, and engages in political education, legislative action, organizing, and education and training.
The official records of the Massachusetts AFL-CIO provide insight into the aims and administrative workings of the organization. These includes a nearly complete run of proceedings and reports from its conventions since 1902, except for a five year gap 1919-1923, minutes and agendas for the meetings of the Executive Council, and the President's files (1982- ). The collection is particularly strong in the period since about 1980.
- Labor unions--Massachusetts
- Massachusetts AFL-CIO
Patterson, Charles H.
Charles H. Patterson Papers, 1930-1958.
Call no.: FS 089
For many years, Charles H. Patterson served as head of the Department of Language and Literature at Massachusetts Agricultural College. Born in Smithsonville, Ont., in 1863, Patterson received both a BA (1887) and MA (1893) from Tufts University before launching his teaching career. He joined the faculty at MAC as an assistant professor of English, in 1916, after 13 years at West Virginia University. A former professional actor, he taught courses in modern literature, with a particular interest in drama, and served as department chair for nearly a decade before his sudden death in 1933.
The Patterson Papers contain a small selection of correspondence and notes on English composition and literature as taught at Massachusetts Agricultural College. Most noteworthy, perhaps, is a draft of Patterson's unpublished book, The Amazing Boucicault.
- Boucicault, Dion, 1820-1890
- Drama--Study and teaching
- Massachusetts Agricultural College--Faculty
- Massachusetts Agricultural College. Department of English
- Patterson, Charles H
Perreault, Alida
Alida Perreault Papers, 1906-1957 (Bulk: 1928-1933).
Call no.: MS 808
Alida Capistrant was the eleventh of twelve children in a large French-Canadian family in South Hadley, Massachusetts born on July 24,1914. Her parents both immigrated from Quebec in 1885. On September 30, 1895 they were married in South Hadley. The Capistrant family rented their home until 1912 when they purchased their first house in South Hadley. Alida had an active social life as a teenager and considered attending college or university, but did not pursue any further education until about 1943, when she studied at the Providence Hospital School of Nursing in Holyoke, Massachusetts. Two years later she married James Perreault and the couple had two children, a daughter, Marcia (Perreault) Matthieu and a son, David James Perreault. They lived in South Hadley until 2003, when they moved to Chandler, Arizona to be near their daughter. Alida Perreault died of complications from Alzheimer's disease on April 7, 2006, and James died in 2008. Both are buried in Saint Rose Cemetery in South Hadley next to Alida's family.
Alida's correspondence during her high school years (1928-1932) reveal a young woman with a substantial network of friends and family. The bulk of the collection consists of letters from several friends, including two potential romantic interests. Letters document daily activities, family happenings, and later Alida's interest in a career as a nurse and her leadership role in the South Hadley Women's Club.
- Capistrant family--Correspondence
- Family--Massachusetts--History
- High school students--Massachusetts
- South Hadley (Mass.)--History
- Williston Northampton School (Easthampton, Mass.)
- Fogg, Esther
- McEwan, William
- Mitkiewitcz, Freddie A.
Types of material
- Correspondence
- Greeting cards
- Invitations
Rare book collections
The Department of Special Collections and University Archives houses over 35,000 volumes reflecting an evolving history of collecting at UMass Amherst. Beginning in the late 1860s with a focus on agriculture and the natural sciences, SCUA has developed into a resource for the study of regional and local history in New England, emphasizing our varied cultural, social, religious, and political histories.
Beyond New England, SCUA has built strength in several distinct areas, ranging from the history of social change to the extraordinary collection of Japanese rarities collected by the geologist Benjamin Smith Lyman during the Meiji era. Other noteworthy collections include those pertaining to the culture of the Cold War: a growing collection of books printed in East Germany and one of the largest collections of materials in the United States from the Solidarity movement in Poland.
All books and periodicals held by SCUA are cataloged in the Library's online catalog and summary descriptions of most major book collections, but not individual titles, are included in SCUA's own online catalog, UMarmot.
Selected areas of collecting interest:
Agriculture, horticulture, natural history
The library holds key works in apiculture, entomology, gardening, landscape design, organic agriculture, pomology, sustainability, and viticulture, with numerous works in animal husbandry. Materials date back to the 16th century, however the strength of the collections lies in the late 18th and 19th centuries.
Cookery in New England
The collections of Beatrice A. McIntosh, Athena Savas, and Lynette Foucher, among others, include books, pamphlets, and ephemera relating to the culinary history of New England, including many thousands of cookbooks published by church and community organizations.
European history and culture
Diverse collections ranging from materials on Revolutionary-era Europe, 1789-1848 (the Binet and Brabançonne Collections); Anglo-American Political Economy; twentieth century German history (the Harold Gordon Collection on the Interwar period and the Hans Joachim Ring Collection on East German cinema); and Communist-era Poland (Basia Jakubowska Schlatner Solidarity Collection).
Gay and Lesbian Literature
The centerpiece extensive of our holdings is the collection of gay rights pioneer Barbara Gittings and her partner, Kay Tobin Lahusen, which includes books on the history of homosexuality in America, works by and about gay writers, gay activism, and related topics.
Literary collections
Books by and about Robert Francis, Archibald MacLeish, William Manchester, William Lederer, and the Broadside Press, among others, as well as the poetry libraries of Francis, Wallace Stevens, and Anne Halley. Although the literary collections focus largely on New England writers, SCUA houses fine collections of the works of William Morris and William Butler Yeats, signed first editions of works by Thomas Mann, and collections of French and Scottish writers.
New England history and culture
Local and regional histories, novels, and other writing about Massachusetts from the eighteenth century to the present. These include an array of election, ordination, installation, dedication, fast-day, mission, farewell, and funeral sermons; Fourth of July orations; and addresses to or by benevolent, cultural, and civic organizations in the Commonwealth. SCUA also collects works printed in small towns and rural districts of Massachusetts from the eighteenth and nineteenth centuries.
Social change
Valuable collections for the history of antislavery in New England and politics of the left. The John P. Roche and Steven Siteman Collections focus on the American left from the late 19th century through the 1950s, with some European materials and materials from the political right.
Santerre Franco-American Collection
Richard Santerre Franco-American Collection, 1872-1978.
Call no.: RB 009
An historian from Lowell, Mass., Richard Santerre received his doctorate from Boston College in 1974 for his dissertation Le Roman Franco-Americain en Nouvelle Angleterre, 1878-1943. For more than twenty years he published regularly on the history of French and French-Canadian immigrants in New England, particularly Massachusetts, while doing so, assembling a significant collection of books on the subject.
With titles in both French and English, the Santerre Collection deals with the wide range of Franco-American experience in New England, touching on topics from literature and the arts to religion, benevolent societies, language, the process of assimilation, biography, and history. The collection includes several uncommon imprints regarding French American communities in Lowell, Lawrence, New Bedford, and Worcester, Mass., as well as in Maine, New Hampshire, and Rhode Island, and it includes publications of associations such as the Ralliement Français en Amérique, the Association Canado-Americain, and the Alliance Française de Lowell.
- Franco-Americans--Connecticut
- Franco-Americans--Massachusetts
- Franco-Americans--New Hampshire
- Franco-Americans--Rhode Island
- French Canadians
- Santerre, Richard
Solander, Arvo A.
Arvo A. Solander Papers, 1930-1958.
Call no.: MS 587
Graduating from Harvard in the thick of the Great Depression, Arvo A. Solander worked as a civil and sanitary engineer for a variety of state and federal agencies, including the Civil Works Administration and the Civilian Conservation Corps. During the 1930s, as opportunity arose, he filled positions as a road engineer, in the design and construction of water and sewage plants, in pollution control, as a safety engineer in the shellfish industry, and in mosquito control, taking jobs throughout Massachusetts and as far away as Tennessee. After using his talents as an officer in the Sanitary Corps during the Second World War, based primarily in Arkansas, Solander returned home to Massachusetts and opened a private engineering office in South Hadley. He worked as a civil engineer and surveyor until his death in January 1976.
The Arvo Solander Papers consists of twenty-four bound volumes documenting thirty years of varied work as an engineer, including his contributions to the construction of the Quabbin Reservoir. Within the bound volumes are a wide range of reports, typescripts, sketches and diagrams, graphs, contracts and design specifications, photographs, and postcards.
- Civil engineers
- Civilian Conservation Corps (U.S.)
- Depressions--1929
- Fisheries--Massachusetts
- Mosquitoes--Control
- Quabbin Reservoir (Mass.)
- Roads--Design and construction
- Sanitary engineers
- Sewage disposal plants--Design and construction
- United States. Federal Civil Works Administration
- Water--Pollution--Tennessee
- Water-supply--Massachusetts
- Westfield State Sanatorium
- World War, 1939-1945
- Wrentham State School
- Solander, Arvo A
Types of material
- Photographs
- Scrapbooks
Stockbridge, Levi, 1820-1904
Levi Stockbridge Papers, 1841-1878.
Call no.: RG 003/1 S76
Born in Hadley, Mass., in 1820, Levi Stockbridge was one of the first instructors at Massachusetts Agricultural College and President from 1879-1882. Known for his work on improving crop production and for developing fertilizers, Stockbridge was an important figure in the establishment of the college's Experiment Station. After filling in as interim President of MAC in 1879, he was appointed president for two years, serving during a period of intense financial stress. After his retirement in 1882, he was named an honorary professor of agriculture.
The Stockbridge Papers include correspondence, personal notebooks, travel diary, journal as a farmer (1842-1845), writings, lectures, notes on experiments, clippings, photocopies of personal and legal records, and biographical material, including reminiscences by Stockbridge's daughter. Also contains auction records, notebook of Amherst, Massachusetts town records (1876-1890), and printed matter about Amherst and national elections, including some about his candidacy for Congress on Labor-Greenback party ticket 1880. Also contains papers (13 items) of Stockbridge's son, Horace Edward Stockbridge (1857-1930), agricultural chemist and educator, including a letter (1885) from him to the elder Stockbridge, written from Japan while he was professor at Hokkaido University.
- Agriculture--Experimentation--History
- Agriculturists--Massachusetts--History
- Amherst (Mass.)--Politics and government--19th century
- Greenback Labor Party (U.S.)--History
- Japan--Description and travel--19th century
- Legislators--Massachusetts--History--19th century
- Massachusetts Agricultural College
- Massachusetts Agricultural College--Students
- Massachusetts Agricultural College. President
- Massachusetts Cattle Commission
- Massachusetts--Politics and government--1865-1950
- Stockbridge family
- Stockbridge, Horace E. (Horace Edward),1857-1930
- Stockbridge, Levi, 1820-1904
Types of material
- Diaries
Stone, George E. (George Edward), 1860-1941
George Edward Stone Papers, 1890-1957.
Call no.: FS 085
Professor of Botany, Massachusetts Agricultural College.
Correspondence, lecture notes, reports, notes on experiments, drawings depicting original apparatus, scrapbooks of printed botanical illustrations, student papers, genealogies, memorabilia, and photographs; together with papers reflecting administrative and official duties; correspondence, notes, and news clippings on psychic phenomena; and autobiographical notes, including reflections on Massachusetts Agricultural College and on Emily Dickinson.
- Botany--Massachusetts
- Dickinson, Emily, 1830-1886
- Horticulture--Massachusetts
- Massachusetts Agricultural College--Faculty
- Massachusetts Agricultural College. Botany Department
- Plant physiology--Massachusetts
- Barlow, Waldo
- Stone, George E. (George Edward), 1860-1941
Types of material
- Herbaria
- Photographs
Storrsville (Mass.) Lyceum Debating Society
Storrsville Lyceum Debating Society Minutebook, 1842-1846.
Call no.: MS 016 bd
Club that met weekly or bi-weekly in Storrsville, Massachusetts, to debate questions of local, national, and international interest including religion, abolition and slavery, human nature, penal reform, the lure of the West, intemperance, and war and peace. Single minutebook includes two versions of the constitution, proposed and debated questions, the teams, the outcome, and notations of any additional activities that took place during the formal meetings.
- Ciceronean Debating Club (Dana, Mass.)
- Dana (Mass. : Town)--Intellectual life--19th century
- Debates and debating--Massachusetts--Dana (Town)--History
- Storrsville (Dana, Mass. : Town)--Intellectual life--19th century
- Storrsville Lyceum Debating Society (Dana, Mass.)--Archives
Types of material
- Minute books


text:
Madera musician produces new CDs for kids
Children's recording artist Oklin Bloodworth of Madera has recorded and produced two more children's albums in time for Valentines Day.
Children's recording artist Oklin Bloodworth of Madera has recorded and produced two more children's albums in time for Valentines Day.
Recent stories about a 12-year-old boy being found responsible for raping a 7-year-old girl have stunned Madera County.
A fundraising dinner was held Saturday at Fourth Street Church of God to benefit Doors of Hope Pregnancy Center.
Local teacher and historian Bill Coate received a standing ovation during a school board meeting Tuesday after Tenisha Armstrong, 29, Coate's former student, discussed the impact he has made on her life.
Mary Jean Wilson, executive director of the Shunammite Place, knows how it feels to struggle with a troubled past. She battled and overcame traumatic childhood experiences which haunted her earlier in life.
The rate of development of the mountain areas of Madera County is up to 45 years ahead of what was projected in 1975, mainly in the Yosemite Lakes park area.
As the kids locked their arms around the bows, the wagons twisted, bounced, and turned through the tortuous Cerro Alto Mountain Pass about 30 miles east of El Paso.
The 24-year-old woman involved in an eight-car crash Jan. 27 died Monday in a Fresno hospital.
The injured woman whose identity was in question after an eight-car pileup on State Route 99 last week is 24-year-old Tondi Mason of Modesto, a friend says.


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Are risk takers more important than others?
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iHerb Customer Reviews
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Items 1-8 of 8 total


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Damascus Nightsby Philip Boehm Rafik Schami
In the classical Arab tradition of tale-telling, here is a magical book that celebrates the power of storytelling, delightfully transformed for modern sensibilities by an award-winning author.The time is present-day Damascus, and Salim the coachman, the city's most famous storyteller, is mysteriously struck dumb. To break the spell, seven friends gather for seven nights to present Salim with seven wondrous "gifts"-seven stories of their own design.Upon this enchanting frame of tales told in the fragrant Arabian night, the words of the past grow fainter, as ancient customs are yielding to modern turmoil. While the hairdresser, the teacher, the wife of the locksmith sip their tea and pass the water pipe, they swap stories about the magical and the mundane: about djinnis and princesses, about contemporary politics and the difficulties of bargaining in a New York department store. And as one tale leads to another... and another... all of Damascus appears before your eyes, along with a vision of storytelling-and talk-as the essence of friendship, of community, of life.A sly and graceful work, a delight to readers young and old, Damascus Nights is, according to Publishers Weekly, "a highly atmospheric, pungent narrative."
An Ermine in Czernopolby Philip Boehm Gregor Von Rezzori
An NYRB Classics Original. Set just after World War I, An Ermine in Czernopol centers on the tragicomic fate of Tildy, an erstwhile officer in the army of the now-defunct Austro-Hungarian Empire, determined to defend the virtue of his cheating sister-in-law at any cost. Rezzori surrounds Tildy with a host of fantastic characters, engaging us in a kaleidoscopic experience of a city where nothing is as it appears--a city of discordant voices, of wild ugliness and heartbreaking disappointment, in which, however, "laughter was everywhere, part of the air we breathed, a crackling tension in the atmosphere, always ready to erupt in showers of sparks or discharge itself in thunderous peals." This first complete English translation of The Ermine of Czernopol makes a masterpiece of postwar literature available to American readers.
Letters to Milenaby Franz Kafka Philip Boehm
Kafka's terrible illness was progressing. He was a man who needed so much time, and who had so painfully little. Kafka lived his whole life as few do; these letters are a testimony to his intense vitality and to his genius as a writer.
A Woman in Berlin: Eight Weeks in the Conquered Cityby Anonymous Philip Boehm
A New York Times Book Review Editors' Choice. For eight weeks in 1945, as Berlin fell to the Russian army, a young woman kept a daily record of life in her apartment building and among its residents. "With bald honesty and brutal lyricism" ( Elle), the anonymous author depicts her fellow Berliners in all their humanity, as well as their cravenness, corrupted first by hunger and then by the Russians. "Spare and unpredictable, minutely observed and utterly free of self-pity" (The Plain Dealer,Cleveland), A Woman in Berlin tells of the complex relationship between civilians and an occupying army and the shameful indignities to which women in a conquered city are always subject - the mass rape suffered by all, regardless of age or infirmity. A Woman in Berlin stands as "one of the essential books for understanding war and life" (A. S. Byatt, author of Possession).
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text: Valero, Oscar On Banach fixed point theorems for partial metric spaces. (English) Zbl 1087.54020 Appl. Gen. Topol. 6, No. 2, 229-240 (2005). Summary: We prove several generalizations of the Banach fixed point theorem for partial metric spaces (in the sense of O'Neill) given in [{\it S. Oltra} and {\it O. Valero}, Rend. Ist. Math. Univ. Triest. 36, 17--26 (2004; Zbl 1080.54030)], obtaining as a particular case of our results the Banach fixed-point theorem of {\it S. G. Matthews} [Ann. N. Y. Acad. Sci. 728, 183--197 (1994; Zbl 0911.54025)], and some well-known classical fixed-point theorems when the partial metric is, in fact, a metric. Cited in 37 Documents MSC:54H25Fixed-point and coincidence theorems in topological spaces54E50Complete metric spaces54E99Topological spaces with richer structures68Q55SemanticsKeywords:dualistic partial metric; complete; quasi-metric BibTeX


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A wizard (Nicolas Cage) trains a reluctant protege (Jay Baruchel) in the art of magic to help him protect Manhattan from a powerful adversary (Alfred Molina).
The trailer for The Sorcerer's Apprentice is so frenetic we feared the movie might be an equally chaotic mess. For some critics it is, for others it's not so bad, and a few even really like it.
"...manages to hit every mark and gives us a rare blockbuster that should please viewers of all types."
"...isn't bad as these things go. It's more diverting than the National Treasure movies, which, like this one, were produced by Jerry Bruckheimer, were directed with aggressive impersonality by Jon Turteltaub and starred Nicolas Cage."
"...It's all sugar and caffeine, no nutrition. In place of a plot, there's a premise; in place of carefully crafted action, there are stupefying exercises in computer-generated imagery, and in place of an ending, there's a hook for the sequel..."
"A noisy, f/x-spewing cauldron of a movie..."
"A tired relic of summer-movie cliches ... Apprentice lurches from one been-there-done-that sequence to another."