Книги по МРТ КТ на английском языке / The Embryonic Human Brain An Atlas of Developmental Stages. Third Edition. 2006. By Ronan O'Rahilly
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Amygdaloid complex
Figure 25–2. A comparison of coronal sections at (A) 20 mm (stage 20, 7 weeks) and (B) 240 mm (approximately 24–28 weeks), at the junction of trimesters 2 and 3. Section B1 is slightly anterior to section B2. A comparison between the embryonic and the fetal sections indicates the further development after 7 weeks. Some of the fetal features are already foreshadowed even at 8 weeks (stage 23). The most striking difference is in the corpus callosum, which, after 100 mm, extends rapidly in a caudal direction, thereby covering the roof of the third ventricle. Hence, during the embryonic period, the longitudinal fissure leads directly onto the roof of the third ventricle, whereas beginning in trimester 2, the fissure becomes blocked below by the corpus callosum.
Another very important feature is the descent of the fibers of the internal capsule. They traverse the region of the ventricular eminences at the end of the embryonic period, and during the fetal period (B, brown arrow) they pass first between two telencephalic structures, the putamen and the caudate nucleus, and then between two diencephalic components, the globus pallidus and the thalamus. Even in the adult brain, slight histological (vascular) differences can be seen between the two telencephalic elements and the two diencephalic constituents, emphasizing their different prosencephalic origins.
At least two subcortical afferent systems are said to “wait” in the subplate in the second half of trimester 2: thalamocortical fibers and basal forebrain fibers (Kostovic´ et al, 1992). Shortly thereafter, thalamocortical fibers penetrate the cortical plate, where intensive synaptic formation is occurring (Kostovic´ et al., 1992). The first sign of the nucleus basalis complex has been shown by cholinesterase reactivity at the beginning of the fetal period, and reactive bundles develop also (Kostovic,´ 1986).
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TRIMESTER 2 |
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TABLE 25–2. Summary of Development of Hippocampal Formation |
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A. Embryonic Period |
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Figure |
Hippocampus |
Dentate Area |
Fibers |
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15–4 |
Hippocampal thickening on both sides |
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of lamina terminalis |
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16–6 B,C |
Hippocampal thickening extends from |
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future frontal to future temporal pole. |
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16–11 |
Thickening is accompanied by a cell- |
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free marginal layer |
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18–2 |
Hippocampal primordium has grown |
Dentate area is a small rim intercalated |
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and reaches forebrain septum |
between hippocampal thickening |
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and area epithelialis. |
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18–5 |
Rostral and temporal parts are alike |
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20–13 |
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Hippocamposeptal and |
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precommissural fibers |
21–17 |
Thin sheath of marginal cells appears |
Migrating cells form primordium |
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opposite hippocampal thickening |
of dentate area |
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23–24 |
About 5 rows of scattered marginal cells |
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B. Fetal Period |
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GL (mm) |
Weeks |
Hippocampus |
Dentate Gyrus |
Fibers |
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32 |
8 |
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Migrating cells |
Fimbria |
44 |
9 |
Distinct pyramidal layer. |
Patches of cells (Fig. 25–3A′ ) |
Columns of fornix |
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CA 1–3 distinguishable |
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49 |
9 1 |
H1, H2, and H3 can be distinguished |
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2 |
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50–83 |
9 1 –12 |
Marginal cells in marginal layer |
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2 |
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56 |
10 |
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Spherical cell mass (Fig. 25–3B′ ). |
Hippocampal commissure |
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Fimbriodentate sulcus |
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61 |
10 |
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Polymorphic and granular layers |
Alveus forming |
65 |
10 1 |
Marginal layer with Cajal–Retzius cells |
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2 |
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79–85 |
12 |
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Granular layer more distinct |
Alveus complete |
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and clearly separated from H3 |
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120 |
14 |
Involution complete; hippocampal |
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matrix wider than subicular |
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150 |
17 |
Fig. 25–3C |
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Formation of synapses |
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mainly in marginal layer |
200 |
21 |
Fig. 25–3D; ventricular layer |
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Connections in subdivisions |
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seems to be exhausted |
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of hippocampus |
≥300 |
32–45 Hippocampus has grown in volume; |
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by 20 weeks |
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walls of hippocampal sulcus |
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are fused; molecular layer |
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still has some Cajal–Retzius cells |
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Newborn |
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Hippocampal surface only one-third |
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of adult area |
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Sources: A: based on authors’ research. B: based on data from various authors, including Humphrey (1965), Rakic and Yakovlev (1968), Kahle (1969), Bogolepova (1995), Hevner and Kinney (1996), and Bogolepova (1997).
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Figure 25–3. Continued (C) and (D)
Development of the hippocampal formation during trimester 2. Cellular production and migration continue. As the corpus callosum develops, the more rostral parts of the hippocampal primordium regress. The formation, which became curved and multilayered in trimester 1, becomes S-shaped and then rolled during trimester 2. (C) A coronal section at 150 mm (17 weeks). The width of the pyramidal layer in CA1 narrows progressively from the subiculum towards CA2 to CA4. Its cells are small and immature. Three layers can be identified already in the dentate gyrus. The dentate area presents a wide marginal layer, the subiculum is double-layered, and the entorhinal cortex is bipartite. (D) At 203 mm (21 weeks). The pyramidal layer is still populated with mostly immature neurons, as is also the dentate gyrus. Fibers connecting the entorhinal cortex, hippocampus, and subiculum are present by about 19 weeks, and connections between other subdivisions of the hippocampal formation may be established about one week later. Connections with the neocortex, however, are still only beginning at 22 weeks (Hevner and Kinney, 1996). (E) Summary of the change in shape from 9 weeks to curved and multilayered at 10 weeks, S-shaped at 17 weeks, and infolded at 21 weeks.
(F) The hippocampal formation of the adult. C and D are drawn from photomicrographs in Arnold and Trojanowski (1996).
With the completion of the hippocampal arch, it reaches from the olfactory bulb to the region of the uncus. Its transformation into induseum griseum begins rostrally during trimester 2 (Kier et al., 1995, 1997).
Stippling, fimbria. Black, dentate gyrus. Horizontal hatching, hippocampus. Orange-brown, pia mater. The subiculum and the parahippocampal gyrus are left white.
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C h a p t e r 2 6 : TRIMESTER 3 AND THE NEWBORN |
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TABLE 26–1. The Usual Sequence of Appearance of Cerebral Sulci and Gyri Arranged in Groups |
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Sulci |
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Gyri |
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Group |
Weeks |
Lateral |
Medial |
Lateral |
Medial |
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A |
10–15 |
Lateral |
Callosal |
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B |
16–19 |
Parieto-occipital |
Cingulate |
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Gyrus rectus |
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calcarine |
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cingulate |
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Cuneus |
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C |
20–23 |
Central |
Collateral |
Superior temporal |
Parahippocampal |
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Superior temporal (T1) |
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D |
24–27 |
Precentral |
Occipitotemporal |
Precentral |
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Inferior temporal (T2) |
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Middle temporal |
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Postcentral |
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Inferior temporal |
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Superior frontal (F1) |
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Medial & lateral |
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Intraparietal |
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occipitotemporal |
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Postcentral |
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Superior frontal |
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Superior & inferior |
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parietal lobules |
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E |
28–31 |
Inferior frontal (F3) |
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Middle frontal |
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Middle frontal (F2) |
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Inferior frontal |
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Pars triangularis |
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F |
32–35 |
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Paracentral lobule |
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Source: Based largely on data supplied by Marvin D. Nelson, M.D., Los Angeles.
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Tectum |
A |
Tegmentum |
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Cerebral |
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Substantia nigra |
peduncle |
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Crus cerebri |
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B |
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Figure 26–1. A likely migration (arrow) of alar (A) into basal (B) territory has long been suspected in the midbrain. Moreover, the origin of the different cells of the substantia nigra is highly complex (Verney et al., 2001a,b). This section at the junction of trimesters 2 and 3 shows that the adult form and features have clearly been attained.