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“Head direction cells are specialized neurons that fire only when an animal faces a certain range selleck products of directions in the horizontal plane, independent of the location and speed of the animal [2 and 3]. These neurons, which exist in a variety of brain regions [11], are already almost fully developed at the time when animals begin exploring the outside world, at the age of postnatal day 16–18 (P16–P18), a few days after the eyes open at P14–P15 [8 and 9]. The present study was designed to determine whether head direction tuning is present at even earlier ages, before the eyelids open and at a time

when rat pups still spend nearly all of their time in the nest [12]. We specifically asked whether directional tuning differences are maintained across experiences. If relative firing directions are maintained from one experimental trial to another, before the appearance of vision, it would point to strong innate components in the mechanism for directional tuning in the brain. A total of 163 cells were sampled from 14 rat pups while the pups moved around twice for 10 min in a circular or square recording box. Eighty-six of these cells were recorded

during the last 3–4 days before eye opening; 77 cells were recorded 1–2 days after eye opening. No cells were recorded for more than one block of trials. The total number of recording blocks (sessions) was 57. Pre-eye-opening data were obtained on P11 in one rat, P12 in three rats, P13 in six rats, P14 in eight rats, and P15 in one rat; post-eye-opening data were collected on P14 in one rat, P15 in eight rats, and P16 in eight Metformin price rats. Individual rats were recorded for 2–6 days. The tetrodes were placed in presubiculum in seven rats, in parasubiculum

in four rats, at the border between pre- and parasubiculum in two rats, and in medial entorhinal cortex (MEC) in one rat (Figure 1; Figure S1 available online). The tetrodes were distributed across deep and superficial layers of pre- and parasubiculum and deep layers of MEC. The pups moved freely across the recording arena and covered the entire range of head directions. Median running speeds increased from 7.6 ± 0.1 cm/s before eye opening to 9.4 ± 0.2 cm/s after eye opening (means across animals ± SEM; t(102) = 6.9, p < 0.001). Mean coverage of the recording box increased from 85.7% ± 0.8% to 91.5% ± 0.8% (t(102) = 5.0, p < 0.001). Head-direction-tuned cells were Protirelin present from the first day when cells could be identified in the target area (P11 and upward; Figures 1 and 2A). To compare directional tuning before and after eye opening, we computed, for each cell, the length of the mean vector for the distribution of firing rates across the 360° of possible head directions. Cells were classified as head direction cells if their mean vector was longer than the 95th percentile of a distribution of mean vector lengths for shuffled firing rates (Figure 2B). Before eye opening, 59 out of 86 cells (68.6%) passed this criterion.