The visual Wulst in birds shows a four-layered structure: apical part of the hyperpallium (HA), interstitial part of HA (IHA), intercalated part of hyperpallium (HI), and densocellular part of hyperpallium (HD). HD also connects with the hippocampus and olfactory system. Because HD is subjacent to HI, the two have been treated as one structure in many studies, and the fiber connections of HD have been examined by afferents and efferents originating outside HD. However, to clarify the difference between these two layers, they need to be treated separately. In the present study, the fiber connections of HD and HI were analyzed with tract-tracing techniques using a combination of injections of cholera toxin subunit B (CTB) for retrograde tracing and biotinylated dextran amine (BDA) for anterograde tracing. When the two tracers were bilaterally injected in HD, a major reciprocal connection was seen with the dorsolateral subdivision (DL) of the hippocampal formation. When CTB and BDA were bilaterally injected in HI, strong reciprocal connections were found between HI and HA. Next, projection neurons in HD and HI were examined by double staining for CTB combined with vesicular glutamate transporter 2 (vGluT2) mRNA in situ hybridization. After CTB was injected in DL or HA, many neurons revealed CTB1/vGluT21 in HD or HI, respectively. Furthermore, in situ hybridization showed that DL and HA contained neurons expressing various subunits of ionotropic glutamate receptors: AMPA, kainate, and NMDA types. These results suggest that glutamatergic neurons in HD and HI project primarily to DL and HA, respectively.
K E Y W O R D Sbird, fiber connection, glutamate receptor, hippocampal formation, vesicular glutamate transporter 2, Wulst, RRID: AB_10013220, RRID: AB_2313640
| I N TR ODU C TI ONThe avian Wulst (hyperpallium or dorsal pallium) is composed, ventrally from the pial surface, of four layers or 'pseudolayers ' (Medina, 2007): the apical part of the hyperpallium (HA), the interstitial part of the apical hyperpallium (IHA), the intercalated part of the hyperpallium (HI), and the densocellular part of the hyperpallium (HD) (Reiner et al., 2004). The Wulst extends caudally from the level of the olfactory bulbs to roughly half to three quarters of the way through the hemisphere, where it is bounded by medial pallial structures (Karten & Hodos, 1967;Puelles, Martinez-de-la-Torre, Paxinos, Watson, & Martinez, 2007). It is composed of two, relatively distinct, functional regions. A small 'rostral Wulst' is somatosensory in nature, whereas a much larger part, with which the present paper is particularly concerned, extends Abbreviations: AI, intermediate arcopallium; BDA, biotinylated dextran amine; CDL, dorsolateral corticoid area; CTB, cholera toxin B; DIP, dorsointermediate posterior thalamic nucleus; DL, dorsolateral subdivision of hippocampal formation; DLL, lateral part of dorsolateral anterior thalamic nucleus; DLM, medial part of dorsolateral anterior thalamic nucleus; DLP, dorsolateral posterior thalamic nucleus; ...