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The Independent Florida alligator

Either your web browser doesn't support Javascript or it is currently turned off. In the latter case, please turn on Javascript support in your web browser and reload this page. Free to read. Author contributions: E. Developing cortical GABAergic interneurons rely on genetic programs, neuronal activity, and environmental cues to construct inhibitory circuits during early postnatal development. Disruption of these events can cause long-term changes in cortical inhibition and may be involved in neurological disorders associated with inhibitory circuit dysfunction.

We hypothesized that tonic glutamate signaling in the neonatal cortex contributes to, and is necessary for, the maturation of cortical interneurons. To test this hypothesis, we used mice of both sexes to quantify extracellular glutamate concentrations in the cortex during development, measure ambient glutamate-mediated activation of developing cortical interneurons, and manipulate tonic glutamate signaling using subtype-specific NMDA receptor antagonists in vitro and in vivo.

Consistent with elevated ambient glutamate, putative parvalbumin-positive interneurons in the cortex identified using GGAD1-eGFP reporter mice exhibit a transient, tonic NMDA current at the end of the first postnatal week.

Dysfunction of cortical inhibition is associated with multiple neurological disorders, including epilepsy.

Establishing inhibitory cortical networks requires in utero proliferation, differentiation, and migration of immature GABAergic interneurons, and subsequent postnatal morphological maturation and circuit integration. Here, we demonstrate that ambient glutamate provides tonic activation of immature, putative parvalbumin-positive GABAergic interneurons in the neonatal cortex via high-affinity NMDA receptors. When this activation is blocked, GABAergic interneuron maturation is disrupted, and cortical networks exhibit lasting abnormal hyperexcitability.

We conclude that temporally precise activation of developing cortical interneurons by ambient glutamate is critically important for establishing normal cortical inhibition. Inhibitory GABAefrgic interneurons INs control cortical function by constraining local neuronal activity and orchestrating circuit synchronization Hensch et al. During embryonic development, cells destined to become cortical INs are generated in the medial and caudal ganglionic eminences Xu et al.

These newborn INs migrate to the cortex, before radially migrating into the cortical lamina Angevine and Sidman, ; Berry and Rogers, Once in the cortex, a combination of genetic programs and environmental cues drive IN localization, morphological differentiation, and synaptic connectivity Huang et al.

GABAergic synaptic activity is established before glutamatergic, and GABA is excitatory in early development due to developmentally specific expression of chloride transporters Kaila et al.

The underlying developmental events that shape IN activity and maturation are critically important for long-term brain function but yet not fully understood.

Glutamate excitation arising from spontaneous network activity and sensory input is an important driver of cortical network maturation Chattopadhyaya et al. IN migration, morphology, and synaptic connectivity all rely on glutamate receptor activation during development. Indeed, dysfunctional cortical inhibition is thought to be involved in the pathophysiology of epilepsy, schizophrenia, and autism Powell et al.

Thus, glutamate-mediated excitation of INs during development has far reaching effects, beyond the cell-specific effects on INs themselves. This ensures that extracellular glutamate levels are low and that glutamatergic neurotransmission is spatially and temporally constrained. In the first week of rodent postnatal life, however, EAAT expression and functional glutamate uptake are minimal Ullensvang et al.

During the second and third weeks of life, cortical astrocytes rapidly increase EAAT expression and expand their processes, thereby constraining synaptic glutamate signaling Diamond and Jahr, ; Thomas et al. This creates a developmental window in which glutamate signaling is unconstrained by astrocytes Hanson et al. The obligate GluN1 subunit interacts with one or more of the four unique GluN2 subunits GluN2A-D , which are differentially expressed throughout the brain and through development Monyer et al.

GluN2D expression peaks in the cortex during early postnatal development Monyer et al. In addition to their effects on glutamate affinity, GluN2 subunit compositions affect magnesium affinities, channel kinetics, and calcium permeability of NMDARs Paoletti et al. They also a have prolonged deactivation time course following removal of glutamate, and do not show rapid desensitization Vicini et al. Cortical brain slices containing sensorimotor cortex were prepared from mice between the ages of P2 and P21 based on previous protocols Andresen et al.

Biosensor was produced and loaded into brain slices as previously described Dulla et al. Each biosensor image had a ms exposure time using nm excitation. Images were acquired every 10 s. Ten frames of autofluorescence were recorded both before loading of the biosensor and in equivalent unloaded slices. Subsequently, 1 m m glutamate was washed in for 10—15 min to saturate the biosensor.

Autofluorescence was subtracted from the other conditions, and fluorescence resonance energy transfer FRET ratios were calculated. Calculating ambient glutamate levels was based on an established protocol Herman and Jahr, Nucleated patch formation was monitored visually and with continuously repeated 1 mV, ms voltage steps. Once nucleated, patches were separated from the cell body, they were brought to the surface of the slice, slowly raised to 40 mV, and placed within the flow of a rapid solution switcher AutoMate Scientific with a custom 6-barrel flow pipe.

NMDA and glutamate dose—response curves were generated by baseline adjusting all responses and then normalizing to the maximum NMDA or glutamate responses, respectively, on a cell-by-cell basis.

Conversion factors were calculated for each age as follows:. For quantifying ambient glutamate in cortical slices, L5Ps were patched in whole-cell configuration at P3 and P Normalized currents at each age were converted to equivalent concentrations of n m glutamate by multiplying them by the conversion factors for each age as follows:. SEM was calculated for ambient glutamate estimates by propagating proportional errors from the EC 50 values, maximum response ratios, and the NMDA-normalized APV-sensitive holding currents to the final estimated [ Glutamate ] Young, Slices were prepared as described above.

Cell type was confirmed based on electrophysiological properties, including membrane resistance and action potential AP firing frequency, as well as reporter expression. The change in holding current was recorded and access resistance was monitored throughout drug applications. Holding current values from voltage-clamp recordings were normalized to the capacitance of the recorded neuron to calculate current density. INs or pyramidal cells were identified and whole-cell recordings were established as described above.

Drug-induced hyperpolarizations were calculated by subtracting the baseline resting potential average of 90 s from the resting potential after drug wash-on average of 30 s. Additionally, current injections were performed before and after drug applications in current-clamped cells. The amplitude threshold was set to 15 pA, and all events were confirmed manually.

After centrifuging remaining tissue at RPM for 10 min, the cells were resuspended in L15 media and added to a Percoll gradient upper concentration Tissue was centrifuged for another 10 min at RPM, and the upper half of the interface containing myelin and debris was removed.

The lower gradient was diluted with L15 and centrifuged at RPM for 6 min, after which the supernatant was discarded. The cDNA was diluted GluN2D sense and antisense riboprobes were in vitro transcribed from linearized plasmids with T7 and T3 polymerases, respectively Roche Diagnostics. Riboprobes were precipitated, subjected to alkaline hydrolysis for 3.

The prehybridization treatments, riboprobe hybridization, and posthybridization washes were performed on brain sections as previously described Swanger et al. For riboprobe detection, the sections were incubated with peroxidase-conjugated anti-fluorescein antibodies , Roche Diagnostics at room temperature for 1 h, washed with TBS with 0.

The sections were then incubated with alkaline phosphatase-conjugated anti-digoxigenin antibodies , Roche Diagnostics followed by five 10 min washes with TBS with 0. Slides were stained with DAPI, rinsed briefly with H 2 O, and coverslipped with polyvinyl chloride mounting media containing propyl gallate Swanger et al.

Sense and antisense probes were hybridized on separate slides in parallel, detected with the same detection solutions, and imaged with the same acquisition settings. ImageJ was used to prepare the images for presentation, and the intensity settings for each channel are the same across all images at a given magnification. DQP was first dissolved to m m stock solution in DMSO and then diluted, before injections, to a final concentration of 5 m m in vehicle.

For immunofluorescence-based quantification of GABAergic synapse density, DQP- and vehicle-treated littermates were processed, stained, and imaged in parallel. Secondary antibodies goat anti-rabbit and goat anti-mouse Cy3.

Sections were mounted using Fluoromount-G Southern Biotechnology. Mounted sections were then used for confocal imaging. Images were taken from L4—5 of somatosensory cortex.

Laser and imaging settings were identical from section to section and across imaging days. Synapses were counted in the projected images using ImageJ. Briefly, a mask was generated for both VGAT and Gephyrin images using a background subtraction and rolling ball radius of To quantify colocalization of eGFP with other molecular markers, immunohistochemical approaches were used to label neuronal tissue from G42 animals.

Secondary antibodies against mouse, rat, chicken, or rabbit, tagged with Alexa, -Cy3, or , were applied at a concentration of All INs showed characteristic fast-spiking properties. Cells were then held for 5—15 min at resting potential, and current steps were repeated.

The patch pipette was then withdrawn slowly to form an outside-out patch and allow the cell to reseal. Cells sat for at least 10 min after removal of the patch pipette. Slices were whole-mounted in Fluoromount-G Southern Biotechnology.

Two-dimensional reconstructions were made in ImageJ by SUM-projecting Z stacks of filled neurons and then tracing with the simple neurite tracer ImageJ plugin. Sholl analysis was done on the two-dimensional reconstructions in ImageJ with the Sholl analysis plugin. Additionally, three-dimensional reconstructions were made with Imaris Bitplane and branch points were quantified in Imaris, using the three-dimensional reconstructions.

All drugs were obtained from commercial sources. Power analyses were performed to determine the sample size for each experiment, and the sample populations used for each experiment are detailed below. Mice of either sex were used for all experiments, with an approximate ratio of female to male mice.

Data were not analyzed by gender, but no abnormal distributions of responses were seen. For all electrophysiology analyses, data analysis was automated with MATLAB scripts and applied in the same fashion to datasets across all experimental groups. For all experiments, analysis was done using between-subject comparisons; no within-subject comparisons were made.

Glutamate imaging : Data collected from 2 or 3 animals per group, 3 slices per animal. For comparisons of one or two experimental groups, a Student's t test was used. The Kolmogorov—Smirnov test was used to compare distributions of synaptic current properties. The Wilcoxon rank-sum test was used to compare two groups when data were not normally distributed.

In LMM analysis, data from the same animal are fitted with random intercepts to account for the correlation between repeated measurements on the same mouse. To monitor extracellular glutamate levels, we first used fluorescent glutamate biosensor imaging in acute cortical slices Dulla et al.

This approach allows semiquantitative evaluation of glutamate levels across broad areas of the cortex in acute cortical slices.

Subpicosecond Electro-optic Imaging Using Interferometric And Polarimetric Apparatus

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The Role of Rydberg States in Spectroscopy and Photochemistry

Print Send Add Share. Additional Physical Form: Also available online. Dates or Sequential Designation: Vol. Permission granted to University of Florida to digitize and display this item for non - profit research and educational purposes. Any reuse of this item in excess of fair use or other copyright exemptions requires permission of the copyright holder. Related Items Preceded by: Florida alligator. Xjj j.

Imzwu, Jmuxt/uj Makes

Metascore: User Score: 7. This is the first release for the new project from blink's Travis Barker and Mark Hoppus. User Score: 8.

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News Archive for January 2014

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The aim of this volume is to offer a balanced overview of molecular Rydberg spectroscopy as it has developed over recent decades. The former is aimed at spectral levels where Rydberg, valence-shell, and intermediate-type states interact, with a variety of photochemical consequences. The latter considers states extremely close to the ionization limit, from whereionization is possible with a very slight amount of additional energy. Recently developed techniques make it possible to produce ions in well-defined electronic, vibrational and rotational states, including states resulting from spin-orbit or Jahn-Teller splitting. It is then possible to study the structure and reactions of such state-selected ions as well as those of the corresponding neutral molecules. These techniques amount to badly needed high resolution photoelectron spectroscopy.

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