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6 Chapter 1 Introduction 1.1.2 Hippocampus and its role in AD Hippocampus has been the subject of particular interests to physiologists and neu- roscientists due to its role in the memory, as well as its involvement in the neu- rological disorders such as AD and epilepsy. Information concerning its function and anatomy maybe found in texts and reference works on neurology and neu- roanatomy of the subject (Duvernoy, 2005; Andersen et al., 2006; Taupin, 2008). Hippocampal formation, including hippocampus proper (also known as cornu Am- monis 1 , CA), subiculum, and dentate gyrus, is located in the limbic lobe on the medial aspect of the hemisected brain. It belongs to the most primitive archicor- tex of the cerebral cortex. In the description by Santiago Ramón y Cajal (1911; for English translations, see Ramón y Cajal, 1995, also Ramón y Cajal, 1968), the hippocampal proper is stratified into alveus, stratum oriens, stratum pyramidale, stratum radium, stratum lacunosum, and stratum modeluare, which are usually grouped into three archicortical layers, viz. molecular (stratum radium, stratum lacunosum and stratum modeluare), pyramidal (stratum pyramidale), and polymorphic layers (stratum oriens), from deep to superficial, with ventricular surface covered by the alveus. Based on the cell morphology, Lorente de Nó (1934) divided the hippocampus proper into 4 subfields, namely CA1–4 (see Figure 1.4). CA1, extending from subiculum, is the largest subfield, characterized by pyramidal cells with triangular cell body, in contrast to larger and ovoid cell bodies in CA2 and CA3. CA1 is also known as the ‘vulnerable sector’ or Sommer’s sector, because of its inordinate sen- sitivity to ischemia and hypoxia, while CA3 as the ‘resistant sector’ or Spielmeyer sector. CA2 is a narrow region between CA1 and CA3, of which the characteriza- tion and the distinct existence have been subject to discussion, while now can be distinguished by molecular and genetic criteria (Lein et al., 2005; Mercer et al., 2007). The CA4 subfield originally described by Lorente de Nó is the polymorphic layer of the dentate gyrus, consisting of interneurons. 1 Latin for ‘Ammon’s horn’, after the Egyptian deity imn ‘Ammon’.
Chapter 1 Introduction 7 Figure 1.3: Limbic system, location of hippocampus (red), medial view. Image source: Martini et al. (2002). The dentate gyrus consists of a superficial molecular, the granular in the middle, and the polymorphic being the deepest. The subiculum is connected to the three layers of the hippocampus proper, while develops into a six-layered structure to the end close to the parahippocampal neocortex. Hippocampus is accessed by the association areas of cortex through the entorhinal cortex, which provides the major input to the hippocampal formation. Other afferents to the hippocampus include indusium griseum, cingulate gyrus, spetal nuclei, and parahippocampal gyrus. The efferent fibers carrying the principal output from hippocampus follow the fiber through the alveus, fimbria and fornix. The two contralateral hippocampi on each hemisphere are interconnected by fornix commissure. A full description of the synaptic organization of the hippocampus is provided by Johnston and Amaral (1997).
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