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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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268

C h a p t e r 2 4 : EARLY POSTEMBRYONIC PHASE

Figure 24–3. 33 mm. Projection of diencephalic tracts. The entrance of the Stammbundel¨ (Fig. 24–2) has not been included. The dorsal thalamus (stippled) takes up almost half of the medial surface. Some of the diencephalic nuclei are shown (hatched). The subthalamic and entopeduncular nuclei, and the globus pallidus externus, are clearly identifiable but have not been represented in the reconstruction. The preoptico-hypothalamotegmental tract is indicated by a dagger. The accessory optic tract ends in the region of the subthalamic nucleus, as described by Gilbert (1935) and Cooper (1946b), although the fibers in the adult are connected to the mesencephalic lateral and dorsal terminal nuclei (Fredericks et al., 1988).

Figure 24–4. 33 mm. Schematic presentation of the inferior olivary nucleus. In addition to the accessory olivary nucleus (medial to the hypoglossal nerve) and the principal olivary nucleus (lateral to the hypoglossal nerve) seen in stage 23, further subdivisions can now be distinguished: (1) the medial accessory olivary nucleus, a rostral part with tall cells; (2) a mediocaudal portion consisting of medium-sized cells related to internal arcuate fibers; (3) probably the caudal medial accessory nucleus, a group between the first two and containing small cells;

(4) probably the dorsal accessory nucleus, situated dorsolaterally and containing small cells; and (5) the main nucleus, the most voluminous part, which consists of tall cells. Projections to the cerebellum were present already in stage 23 (Fig. 23–30D). The olivocerebellar fibers join the inferior cerebellar peduncle. In cross sections the olivo-arcuate migrating cells are seen as superficial and intermediate strands, and the cellular axes are arranged tangentially.

EARLY POSTEMBRYONIC PHASE

269

C-Shaped Structures

A B

1

2

3

4

C

1

2

3

4

Figure 24–5. Associated with the growth of the relatively fixed corpus striatum and the expansion of the cerebral hemispheres in a curved direction to form the temporal lobes, a number of structures develop in a C-shaped manner: (A) the caudate nucleus (head, body, and tail); (B and C) the choroid plexus and the choroid fissure through which it passes; the fornix (columns, body, and crura) and its continuation, the fimbria; the hippocampus and dentate gyrus, the rostral portions of which are found later merely as a remnant termed the indusium griseum;

(C) the corpus callosum, which halts (as the splenium) without descending into the temporal lobe.

Central part

D

E

Parahippocampal gyrus

Figure 24–6. C-shaped structures (Continued ). (D) the lateral ventricle (anterior horn, central part, and inferior horn); (E) the limbic lobe (cingulate and parahippocampal gyri); and the frontal, parietal, and temporal lobes of the hemisphere.

270

C h a p t e r 2 4 : EARLY POSTEMBRYONIC PHASE

Figure 24–7. 33 mm. Lateral view showing the ventricles. Some recesses are well formed, e.g., the supramamillary and the inframamillary, whereas others, e.g., the suprapineal, are not clear. The long cavity of the isthmus is narrowing because of the growth of the cerebellum, but it still extends into a well-developed isthmic recess.

TABLE 24–1. Important C-Shaped Structures in the Brain

Structure

Stage of C-Shaped Appearance

Figure 24–5

Number in B & C

 

 

 

 

Caudate nucleus

23

A

1

Choroid plexus

22,23

B,C

2

Fornix and fibria

Fetal period

B,C

3

Hippocampus

22

B,C

4

Corpus callosum

Fetal period

C

 

Inferior horn of lateral ventricle

22,23

Figure 24–6D

 

 

 

 

 

 

 

EARLY POSTEMBRYONIC PHASE

271

Ventricular

 

Cortical

Figure 24–8. 27.5 mm. The cerebral cortex. The

 

 

cortical plate is present on the surface of the entire

 

layer

 

plate

 

 

 

 

 

 

neopallium and is thickest (approximately 12 rows of

 

 

 

 

 

 

 

 

 

 

cells) mostly where it began in stage 21: in the lateral

 

 

 

 

 

wall opposite the lateral ventricular eminence (Fig.

 

 

 

 

 

24–10). The subpial layer occupies approximately half

 

 

 

 

 

the thickness of the cortical plate, which is relatively

 

 

 

 

 

less than in stage 23. The subplate is at least twice the

 

 

 

 

 

thickness of the cortical plate, which is substantially

 

 

 

 

 

greater than in stage 23. This is based on an increase of

 

 

 

 

 

fibers, some of which lead to the lateral prosencephalic

 

 

 

 

 

fasciculus (Fig. 24–11). Serotonin and vesicular

 

 

 

 

 

 

monoamine transporter fibers are present in the

 

 

 

 

 

 

subplate and intermediate layer (Verney et al., 2002).

 

 

 

 

 

Synapses in early fetuses are present above and below

 

 

 

 

 

the cortical plate, but never within it (Molliver et al.,

 

 

 

 

 

1973). The key drawing is a graphic reconstruction.

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

Intermediate

 

 

 

Subpial

 

Subplate

layer

 

 

layer

Figs 24 – 10

11

8

 

 

 

 

Figures 24–8 to 24–15 are from a fetus of 27.5 mm (silver-impregnated). The brain is slightly more developed than that of the 33 mm fetus; e.g., the cortical plate now extends to the occipital pole, and the brain is no longer as compact. Additional features include the external capsule, the hippocampal sulcus, and choroid villi in the roof of the third ventricle. The photomicrographs from this fetus are oriented so that the rostral end is uppermost. Figures 24–8 to 24–11 show a lateral view, with bars representing 0.15 mm. Figures 24–12 to 24–15 show a medial view, with bars representing 0.2 mm.

By the end of the trimester thalamocortical fibers penetrate the cortical plate and intensive synaptogenesis begins to occur (Kostovic and Rakic, 1990), followed later by a considerable decrease in synaptic number and density (Huttenlocher and Dabholkar, 1997). Synapses become mature during the first postnatal year and new synapses can form throughout life, particularly in the parahippocampal region.

272

C h a p t e r 2 4 : EARLY POSTEMBRYONIC PHASE

Lateral ventricle

V3

1

2

Tel.

Di.

Hemispheric stalk

Inferior horn

Figure 24–9. Unexpected migration from the telencephalon to the diencephalon.

1.Migration from a thin cellular layer near the di-telencephalic junction (cf Fig. 21–6) to the pulvinar and other thalamic components (Letinicˇ and Kostovic,´ 1997).

2.Migration from cells coating the corpus striatum adjacent to the sulcus terminalis to the pulvinar. These cells proliferate until the end of the fetal period (Rakic and Sidman, 1969).

TABLE 24–2. Simplified Classification of the Main

Morphological Divisions and Subdivisions of the

Cerebellum

 

Stage of

 

Appearance

 

(F, fetal period)

 

 

Cerebellar hemispheres & vermisa

 

Corpus cerebelli

 

Anterior lobe

F

Fissura prima

F

Posterior lobe

F

Horizontal fissure

F

Fissura secunda

F

Posterolateral fissure

17

Flocculonodular lobeb

23

Flocculus

21

Nodule

23

Cerebellar peduncles

 

Superior

18

Middle

F

Inferior

18

 

 

a Most of the hemispheres and vermis consist of the paleocerebellum (spinocerebellum) and the neocerebellum (corticopontocerebellum). b The flocculonodular lobe is the archicerebellum (vestibulocerebellum).

EARLY POSTEMBRYONIC PHASE

273

Figure 24–10. 27.5 mm. The corpus striatum, internal capsule, and adjacent structures. The claustrum is well developed, and a clear connection with the ventricular layer of the olfactory area still exists (Fig. 24–15). The external capsule has appeared. Fibers of the internal capsule are seen to reach the diencephalon through the hemispheric stalk, which is delimited by the sulcus terminalis.

The key drawing shows the area included in the photomicrograph. The arrival and origin of the fibers in the subplate are depicted. The ventricular eminences are said to expand to about 10 times the maximum thickness observed in other regions of the CNS during the second month, but reach their peak only at the middle of prenatal life (Rakic and Sidman, 1982). Their growth is supported by the striatal arteries (Nelson et al., 1991), which mature earlier than the arteries of the cerebral wall, that is, they develop a muscularis between 20 and 22 weeks. The ventricular layer contains a network of vessels that possess no muscularis at any age. The regression of the ventricular layer is accompanied by a regression of the capillaries (Nelson et al., 1991).

274

C h a p t e r 2 4 : EARLY POSTEMBRYONIC PHASE

Figure 24–11. 27.5 mm. An enlargement of the internal capsule at a more dorsal level. The key drawing is a graphic reconstruction.

Internal

capsule Subplate

Cortical plate

Figs 24 – 15 13 11

EARLY POSTEMBRYONIC PHASE

275

Figure 24–12. 27.5 mm. The hippocampus is clearly recognizable because of its thinner ventricular layer, which is contiguous to the lamina epithelialis. The third ventricle is visible below, and its folded roof seems to show the beginning development of choroid villi. The thick fiber bundle on each side of the roof area is the stria medullaris thalami. The choroid plexuses of the lateral ventricles are well shown.

Presubiculum

Hippocampus

Stria

 

Lamina

medullaris

 

 

epithelialis

thalami

 

 

 

 

 

 

Figure 24–13. 27.5 mm. The hippocampus more ventrally has joined the prosencephalic septum. The hippocampal sulcus has appeared and is contiguous to the rhinal sulcus, as shown in the key drawing with Figure 24–11. The falx cerebri is evident. The lateral ventricles are seen to be continuous with the cavity of the telencephalon medium, which in turn blends with the third ventricle between the thalami. See also Figure 24–10. The sulcus

Hippocampus terminalis lies between the ventricular eminence and the dorsal

thalamus.

Lamina terminalis

l.-v.f.

Sulcus terminalis

Thalamus

 

dorsalis

V3

 

276

C h a p t e r 2 4 : EARLY POSTEMBRYONIC PHASE

 

 

A

B

L

Olfactory

tract

M

Olfactory bulb

Anterior olfactory nucleus

Olfactory tubercle

M L

Amygdaloid nuclei

Figure 24–14. Development of the olfactory tract. (A) Schematic ventral view of the telencephalon at stage 23 showing medial (M) and lateral (L) olfactory fibers. (B) The postnatal olfactory features.

Modified from Muller¨ and O’Rahilly (2004c, Cells Tissues Organs) by permission of S. Karger AG, Basel.

Olfactory ventricle

Circular sulcus

Septum

Telencephalic part of third ventricle

Commissural

plate

Figure 24–15. 27.5 mm. The dorsal portions of the olfactory bulbs are separated from the prosencephalic septum by the circular sulcus. The olfactory ventricles are evident. The falx cerebri has developed in the loose tissue of the subarachnoid space. Dark-appearing tissue emanating from the ventricular and subventricular layers of the right-hand olfactory bulb leads to the claustrum.

The transient subpial granular layer, which appears before the end of trimester 1, seems to arise from the same area as the claustrum (Fig. 22–12). Two streams of cells have been observed at 14 weeks: one from the medial and the other from the lateral angle of the subventricular layer of the olfactory bulb (Gadisseux

et al., 1992). The cells form a continuous lamina. The subpial granular layer represents a precursor pool for reelin-producing neurons that complement earlier-generated cells [Cajal–Retzius cells in the primordial plexiform layer] during the critical period of cortical migration (Meyer et al., 1999a). Reelin is essential for neuronal migration and cortical lamination. The nucleus basalis (of Meynert) develops during the fetal period. Cholinesterase-reactive fibers arising from it are distributed to the neocortex and the limbic cortex by the end of trimester 2 (Kostovic,´ 1986).

EARLY POSTEMBRYONIC PHASE

277

Figure 24–16. 42 mm. Lateral view, based on a solid (Born) reconstruction. The membranous labyrinth has been added. The insula is becoming deeper. The interval between the diencephalon and the rhombencephalon is wider than in stage 23, and the pontine flexure is less marked. Parts of the diencephalon, as well as the mesencephalon, still remain uncovered by the cerebral hemispheres. The (caudal) mesencephalic Blindsack is visible. The intraventricular part of the cerebellum lies within the fourth ventricle and the extraventricular portion is subdivided by the posterolateral fissure.

Figures 24–16 to 24–18 are from a fetus of 42 mm that was studied by Streeter (1918), Hines (1922), and Padget (1948, 1957). In Figure 24–16 and in subsequent illustrations, an orientation more similar to that of the adult is shown; i.e., the cerebral hemispheres have been “lifted” into a more horizontal position and the brain stem is almost vertical.

A

coronal

 

plane

 

tal

 

tomea

rbi

O

plane

 

A

transverse

plane

 

 

A coronal plane

Figure 24–17. 42 mm. The brain in situ. The face is positioned more or less vertically to allow easier comparison with the adult. The head is less spherical than in stage 23, and its rostrocaudal axis is lengthening. This seems to allow more space for the brain, which is now less compact. Padget (1948, Fig. 21) reconstructed the arteries of this fetus.

Planes have been added to show that prenatally a transverse plane through the trunk does not correspond to the orbitomeatal plane in the adult, which is accepted as the horizontal standard. Similarly, coronal planes are not comparable.