Furthermore, 0

Furthermore, 0.1% biocytin was included in the internal remedy for the purpose of labeling. between the peak and subsequent descending zero crossing of the sinusoid. The phases of inhibitory inputs that induced spikes shifted earlier as the baseline sinusoid rate of recurrence increased, while spiking output shifted to later on phases. Raises in magnitude of the inhibitory inputs shifted the spiking output to earlier phases. Pharmacological blockade of h-current abolished the phase selectivity of hyperpolarizing inputs eliciting spikes. A network computational model using cells possessing related rebound properties as found produces spatially periodic firing properties resembling grid cell firing when a simulated animal moves along a linear track. These results suggest that the ability of mEC stellate cells to open fire rebound spikes in response to a specific range of phases of inhibition could support complex attractor dynamics that provide completion and separation to keep up spiking activity of specific grid cell populations. (Domnisoru, Kinkhabwala, & Tank, 2013; Schmidt-Hieber & H?usser, 2013) that could contribute BRD9757 to the firing of entorhinal neurons inside a grid cell pattern (Hafting, Fyhn, Molden, Moser, & Moser, 2005). The intrinsic properties of stellate cells display raises in membrane potential oscillation period along the dorsal-to-ventral (D/V) axis of the mEC (Boehlen, Heinemann, & Erchova, 2010; Giocomo & Hasselmo, 2008a,b; Giocomo et al., 2007) that resemble the increasing gradient of grid cell firing field size and spacing (Hafting et al., 2005; Sargolini et al., 2006). Similarly, the intrinsic spiking rate of recurrence of grid cells measured by autocorrelograms differs along the D/V axis and shows changes with operating rate (Jeewajee, Barry, O’Keefe, & Burgess, 2008). These experimental data have encouraged the use of oscillatory dynamics to model grid cell properties inside a class of models termed oscillatory interference models (Blair, Welday, & Zhang, 2007; Burgess, Barry, Jeffery, & O’Keefe, 2005; Burgess, Barry, & O’Keefe, 2007; Hasselmo, Giocomo, & Zilli, 2007; Blair, Gupta, & Zhang, 2008; Burgess, 2008) that could link intrinsic cellular properties to grid cell properties. Another class of grid cell models, termed continuous attractor dynamic models, focuses on synaptic relationships between neurons that could interact with intrinsic properties. Attractor dynamic models use symmetric recurrent network contacts to generate grid cell firing patterns, and use variations in asymmetric synaptic relationships regulated by operating velocity to produce variations in grid cell firing field size and spacing (Burak & Fiete, 2009; Couey BRD9757 et al., 2013; Fuhs & Touretzky, 2006; Guanella, Kiper, & Verschure, 2007; Pastoll, Solanka, Vehicle Rossum, & Nolan, 2013). Recent data support elements of the network activity shown by attractor models including shared features of spacing and orientation within individual modules (Stensola et al., 2012) that shift collectively during environmental manipulations (Barry, Hayman, Burgess, & Jeffery, 2007; Yoon et TSPAN9 al., 2013). Attractor dynamic models possess properties of both pattern separation, to avoid having neural activity spread throughout the network, and pattern completion BRD9757 to keep up firing in the set of neurons within a human population coding a specific representation. Stellate cells of the mEC are inlayed in an inhibitory network. They share little to no direct synaptic contacts with one another, but instead interact indirectly through inhibitory interneurons (Couey et al., 2013; Pastoll et al., 2013). The strong inhibitory innervation of stellate cells coupled with their intrinsic properties suggests a functional part for rebound spiking. Rebound spikes happen in response to release from hyperpolarizing current pulses and are dependent on the presence of the h-current (= 17), 2 mM kynurenic acid and 100 M picrotoxin were added to the recording means to fix block glutamatergic and GABAergic synaptic transmission, respectively. All recordings were made between 35 and 37 C. Whole-cell pipettes were fabricated with borosilicate glass capillaries by means of a P-90 horizontal puller (Sutter Tools)..