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ZENK-immunoreactivity in the caudomedial nidopallium of female European starlings relative to the type of song stimulus to which they were exposed. Both means and means adjusted to the response to other factors in the statistical model (photoexperience) are depicted, together with 95% confidence whiskers.

ZENK-immunoreactivity in the caudomedial nidopallium of female European starlings relative to photoexperience (naïve vs. experienced). Both means and means adjusted to the response to other factors in the statistical model (song stimulus) are depicted, together with 95% confidence whiskers.

Adult song recognition learning in songbirds is an ethologically relevant framework for mechanistic examination of nonassociative learning. For example, in colonial zebra finches (Taeniopygia guttata), each male sings one unique song his entire adult life. Recognition of specific songs can thus be used for individual identification, even when the bird is out of sight. Song carries social information that is used to establish and maintain relationships (Woolley and Doupe 2008; Riebel 2009; Lin et al. 2014). Song habituation would therefore support appropriate behavioral interactions with mates and neighbors versus intruders.

Here, we were motivated to expand our understanding of the molecular underpinnings involved in nonassociative song recognition learning. We focused on the mechanistic Target of Rapamycin (mTOR) signaling cascade. mTOR is appealing as a mediator of nonassociative learning because it integrates multiple upstream signals of the environment and is activated by triggers of experience-dependent plasticity (Hoeffer and Klann 2010; Costa-Mattioli and Monteggia 2013). mTOR, as a core member of mTOR complex 1 (mTORC1), can mediate synaptic plasticity by regulating mRNA-to-protein translation (Cammalleri et al. 2003; Kelleher et al. 2004; Richter and Klann 2009; Costa-Mattioli and Monteggia 2013). mTOR signaling makes demonstrated contributions to learning and memory but very little is known about its role in nonassociative learning (Hoeffer and Klann 2010; Costa-Mattioli and Monteggia 2013; Fifield et al. 2013; Graber et al. 2013; Fortress et al. 2015). We have, however, recently discovered that mTOR signaling in the auditory forebrain is required for tutor song copying in juvenile zebra finches (Ahmadiantehrani and London 2017). We therefore asked if mTORC1 signaling displayed signatures of nonassociative learning mechanisms in adults.

Schematic of 3-d-song recognition paradigm variations, including infusions, duration of training song playbacks, and testing song conditions, for the five experiments. Headers indicate song playback events on Days 2 and 3. Conditions for each experiment (Expt) are listed across the rows; a dashed line separates experiments. On Day 1, birds for all experimental groups were placed into acoustic isolation chambers. Training song playbacks on Day 2 varied in length depending on the experiment and group. Blocks illustrate duration of habituation training; the number within the block denotes the duration of training song playbacks in hours (h). Note Expt 3 has a group with a 1.5 h separation between two 3 h training song playbacks. Arrows indicate Rapamycin (Rapa), SC79, or DMSO (Veh) infusions 30 min prior to Day 2 training song playbacks. Testing conditions for pS6 activation on Day 3 included birds who experienced novel (Nov) or familiar (Fam) song playbacks for 30 min, or who were left in silence (Sil) for control; these groups are labeled Trained-Novel, Trained-Familiar, Trained-Silence, respectively.

Levels of mTOR signaling cascade activation in the adult auditory forebrain depend on prior training song experience (Experiment 1). (A) Representative images of pS6 and total S6 immunostaining in male and female auditory forebrain in the Trained-Silence, Trained-Familiar, and Trained-Novel groups. Boxed insets of each image are shown in a higher magnification directly below. Field L is indicated with a dashed oval. Bottom two rows are high magnification images to show cellular specificity of the pS6 and total S6 immunostaining. Scale bars = 500 µm (auditory forebrain), 250 µm (boxed insets), and 25 µm (high magnification images). (B) Sagittal schematic of the zebra finch brain, Dorsal (D) is up, Posterior (P) is to the right. Telencephalic regions are highlighted in gray. The area included in the auditory forebrain images in A (indicated by dashed square) is enlarged to show neuroanatomical detail. Scale bar = 1 mm. (CMM) caudomedial mesopallium; (NCM) caudomedial nidopallium; (HP) hippocampus. (C) pS6+/S6+ cell density ratios in the CMM (left) and NCM (right) in the three playback conditions. Bars represent the group mean SEM. Open circles denote the individual birds. CMM group means SEM: 0.11 0.015 (Trained-Silence), 0.4 0.039 (Trained-Familiar), and 0.73 0.042 (Trained-Novel). NCM group means SEM: 0.061 0.01 (Trained-Silence), 0.18 0.024 (Trained-Familiar), and 0.23 0.035 (Trained-Novel). ***P < 0.001. n = 3 males and 3 females per condition. Note y-axis scales in C are different.

Inhibition, but not constitutive activation, of mTOR signaling during training specifically alters the pS6 levels in response to familiar song playbacks (Experiment 2). (A) Representative high-magnification images of pS6+ cells in the CMM and NCM of adult birds from each experimental condition. Images shown are from a mix of males and females. Scale bar = 100 µm. (B) pS6+/S6+ cell density ratios in the CMM (left) and NCM (right). CMM group means SEM: 0.32 0.036 (DMSO-Sil), 0.86 0.46 (DMSO-Nov), 0.57 0.07 (DMSO-Fam), 0.16 0.043 (Rapa-Fam), and 0.5 0.062 (SC79-Fam). NCM group means SEM: 0.33 0.033 (DMSO-Sil), 0.65 0.038 (DMSO-Nov), 0.63 0.029 (DMSO-Fam), 0.26 0.087 (Rapa-Fam), and 0.59 0.075 (SC79-Fam). (C) Dysregulation of mTOR signaling during song playback training sessions does not alter ZENK induction patterns after familiar song exposure. Hippocampus-normalized ZENK-positive (ZENK+) cell density in the CMM (left) and NCM (right). CMM group means SEM: 0.28 0.03 (DMSO-Sil), 1.15 0.15 (DMSO-Nov), 0.57 0.11 (DMSO-Fam), 0.6 0.13 (Rapa-Fam), and 0.56 0.074 (SC79-Fam). NCM group means SEM: 0.37 0.054 (DMSO-Sil), 1.28 0.19 (DMSO-Nov), 0.73 0.12 (DMSO-Fam), 0.69 0.15 (Rapa-Fam), and 0.65 0.072 (SC79-Fam). Bars represent the experimental condition mean SEM. Open circles denote the individual birds in each experimental group. (Sil) Trained-Silence, (Fam) Trained-Familiar, (Nov) Trained-Novel. *P < 0.05, **P < 0.005, and ***P < 0.001, between indicated groups. n = 2 males and 2 females per group. 153554b96e

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