Extracting synaptic conductances from single membrane potential traces

In awake animals, the activity of the cerebral cortex is highly complex, with neurons firing irregularly with apparent Poisson statistics. One way to characterize this complexity is to take advantage of the high interconnectivity of cerebral cortex and use intracellular recordings of cortical neurons, which contain information about the activity of thousands of other cortical neurons. Identifying the membrane potential (Vm) to a stochastic process enables the extraction of important statistical signatures of this complex synaptic activity. Typically, one estimates the total synaptic conductances (excitatory and inhibitory) but this type of estimation requires at least two Vm levels and therefore cannot be applied to single Vm traces. We propose here a method to extract excitatory and inhibitory conductances (mean and variance) from single Vm traces. This "VmT method" estimates conductance parameters using maximum likelihood criteria, under the assumption are that synaptic conductances are described by Gaussian stochastic processes and are integrated by a passive leaky membrane. The method is illustrated using models and is tested on guinea-pig visual cortex neurons in vitro using dynamic-clamp experiments. The VmT method holds promises for extracting conductances from single-trial measurements, which has a high potential for in vivo applications.Comment: Neuroscience (in press

Spectral functions of isoscalar scalar and isovector electromagnetic form factors of the nucleon at two-loop order

We calculate the imaginary parts of the isoscalar scalar and isovector electromagnetic form factors of the nucleon up to two-loop order in chiral perturbation theory. Particular attention is paid on the correct behavior of Im $\sigma_N(t)$ and Im $G_{E,M}^V(t)$ at the two-pion threshold $t_0=4 m_\pi^2$ in connection with the non-relativistic 1/M-expansion. We recover the well-known strong enhancement near threshold originating from the nearby anomalous singularity at $t_c = 4m_\pi^2-m_\pi^4/M^2 = 3.98 m_\pi^2$. In the case of the scalar spectral function Im $\sigma_N(t)$ one finds a significant improvement in comparison to the lowest order one-loop result. Higher order $\pi\pi$-rescattering effects are however still necessary to close a remaining 20%-gap to the empirical scalar spectral function. The isovector electric and magnetic spectral functions Im $G_{E,M}^V(t)$ get additionally enhanced near threshold by the two-pion-loop contributions. After supplementing their two-loop results by a phenomenological $\rho$-meson exchange term one can reproduce the empirical isovector electric and magnetic spectral functions fairly well.Comment: 10 pages, 6 figures, submitted to Physical Review

Generation of photovoltage in graphene on a femtosecond time scale through efficient carrier heating

Graphene is a promising material for ultrafast and broadband photodetection. Earlier studies addressed the general operation of graphene-based photo-thermoelectric devices, and the switching speed, which is limited by the charge carrier cooling time, on the order of picoseconds. However, the generation of the photovoltage could occur at a much faster time scale, as it is associated with the carrier heating time. Here, we measure the photovoltage generation time and find it to be faster than 50 femtoseconds. As a proof-of-principle application of this ultrafast photodetector, we use graphene to directly measure, electrically, the pulse duration of a sub-50 femtosecond laser pulse. The observation that carrier heating is ultrafast suggests that energy from absorbed photons can be efficiently transferred to carrier heat. To study this, we examine the spectral response and find a constant spectral responsivity between 500 and 1500 nm. This is consistent with efficient electron heating. These results are promising for ultrafast femtosecond and broadband photodetector applications.Comment: 6 pages, 4 figure

Lowest Q^2 Measurement of the gamma*p -> Delta Reaction: Probing the Pionic Contribution

To determine nonspherical angular momentum amplitudes in hadrons at long ranges (low Q^2), data were taken for the p(\vec{e},e'p)\pi^0 reaction in the Delta region at Q^2=0.060 (GeV/c)^2 utilizing the magnetic spectrometers of the A1 Collaboration at MAMI. The results for the dominant transition magnetic dipole amplitude and the quadrupole to dipole ratios at W=1232 MeV are: M_{1+}^{3/2} = (40.33 +/- 0.63_{stat+syst} +/- 0.61_{model}) (10^{-3}/m_{\pi^+}),Re(E_{1+}^{3/2}/M_{1+}^{3/2}) = (-2.28 +/- 0.29_{stat+syst} +/- 0.20_{model})%, and Re(S_{1+}^{3/2}/M_{1+}^{3/2}) = (-4.81 +/- 0.27_{stat+syst} +/- 0.26_{model})%. These disagree with predictions of constituent quark models but are in reasonable agreement with lattice calculations with non-linear (chiral) pion mass extrapolations, with chiral effective field theory, and with dynamical models with pion cloud effects. These results confirm the dominance, and general Q^2 variation, of the pionic contribution at large distances.Comment: 6 pages, 3 figures, 1 tabl

A measurement of the axial form factor of the nucleon by the p(e,e'pi+)n reaction at W=1125 MeV

The reaction p(e,e'pi+)n was measured at the Mainz Microtron MAMI at an invariant mass of W=1125 MeV and four-momentum transfers of Q^2=0.117, 0.195 and 0.273 (GeV/c)^2. For each value of Q^2, a Rosenbluth separation of the transverse and longitudinal cross sections was performed. An effective Lagrangian model was used to extract the `axial mass' from experimental data. We find a value of M_A=(1.077+-0.039) GeV which is (0.051+-0.044) GeV larger than the axial mass known from neutrino scattering experiments. This is consistent with recent calculations in chiral perturbation theory.Comment: 14 pages, 5 figures, uses elsart.cl

A Comprehensive Workflow for General-Purpose Neural Modeling with Highly Configurable Neuromorphic Hardware Systems

In this paper we present a methodological framework that meets novel requirements emerging from upcoming types of accelerated and highly configurable neuromorphic hardware systems. We describe in detail a device with 45 million programmable and dynamic synapses that is currently under development, and we sketch the conceptual challenges that arise from taking this platform into operation. More specifically, we aim at the establishment of this neuromorphic system as a flexible and neuroscientifically valuable modeling tool that can be used by non-hardware-experts. We consider various functional aspects to be crucial for this purpose, and we introduce a consistent workflow with detailed descriptions of all involved modules that implement the suggested steps: The integration of the hardware interface into the simulator-independent model description language PyNN; a fully automated translation between the PyNN domain and appropriate hardware configurations; an executable specification of the future neuromorphic system that can be seamlessly integrated into this biology-to-hardware mapping process as a test bench for all software layers and possible hardware design modifications; an evaluation scheme that deploys models from a dedicated benchmark library, compares the results generated by virtual or prototype hardware devices with reference software simulations and analyzes the differences. The integration of these components into one hardware-software workflow provides an ecosystem for ongoing preparative studies that support the hardware design process and represents the basis for the maturity of the model-to-hardware mapping software. The functionality and flexibility of the latter is proven with a variety of experimental results

The first determination of Generalized Polarizabilities of the proton by a Virtual Compton Scattering experiment

Absolute differential cross sections for the reaction (e+p -> e+p+gamma) have been measured at a four-momentum transfer with virtuality Q^2=0.33 GeV^2 and polarization \epsilon = 0.62 in the range 33.6 to 111.5 MeV/c for the momentum of the outgoing photon in the photon-proton center of mass frame. The experiment has been performed with the high resolution spectrometers at the Mainz Microtron MAMI. From the photon angular distributions, two structure functions which are a linear combination of the generalized polarizabilities have been determined for the first time.Comment: 4 pages, 3 figure

Effective electro-optical modulation with high extinction ratio by a graphene-silicon microring resonator

Graphene opens up for novel optoelectronic applications thanks to its high carrier mobility, ultra-large absorption bandwidth, and extremely fast material response. In particular, the opportunity to control optoelectronic properties through tuning of Fermi level enables electro-optical modulation, optical-optical switching, and other optoelectronics applications. However, achieving a high modulation depth remains a challenge because of the modest graphene-light interaction in the graphene-silicon devices, typically, utilizing only a monolayer or few layers of graphene. Here, we comprehensively study the interaction between graphene and a microring resonator, and its influence on the optical modulation depth. We demonstrate graphene-silicon microring devices showing a high modulation depth of 12.5 dB with a relatively low bias voltage of 8.8 V. On-off electro-optical switching with an extinction ratio of 3.8 dB is successfully demonstrated by applying a square-waveform with a 4 V peak-to-peak voltage.Comment: 12 pages, including 7 figure

Measurement of the Transverse-Longitudinal Cross Sections in the p (e,e'p)pi0 Reaction in the Delta Region

Accurate measurements of the p(e,e?p)pi0 reaction were performed at Q^2=0.127(GeV/c)^2 in the Delta resonance energy region. The experiments at the MIT-Bates Linear Accelerator used an 820 MeV polarized electron beam with the out of plane magnetic spectrometer system (OOPS). In this paper we report the first simultaneous determination of both the TL and TL? (``fifth" or polarized) cross sections at low Q^{2} where the pion cloud contribution dominates the quadrupole amplitudes (E2 and C2). The real and imaginary parts of the transverse-longitudinal cross section provide both a sensitive determination of the Coulomb quadrupole amplitude and a test of reaction calculations. Comparisons with model calculations are presented. The empirical MAID calculation gives the best overall agreement with this accurate data. The parameters of this model for the values of the resonant multipoles are |M_{1+}(I=3/2)|= (40.9 \pm 0.3)10^{-3}/m_pi, CMR= C2/M1= -6.5 \pm 0.3%, EMR=E2/M1=-2.2 \pm 0.9%, where the errors are due to the experimental uncertainties.Comment: 10 pages, 3 figures, minor corrections and addition