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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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278

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

 

 

 

Figure 24–18. 42 mm. Median view.

 

 

(A) The prosencephalic septum is

 

 

relatively reduced in its rostrocaudal

 

 

extent. The paraphysis is

 

 

inconspicuous compared with that of

 

 

stage 23. The dorsal thalamus now

 

 

extends over half of the lateral

 

 

diencephalic wall. The marginal ridge

 

 

between the dorsal and ventral

 

 

thalami is no longer clear, although

 

 

the sulcus medius is visible (Fig.

 

 

24–28). The posterior commissure

 

 

and the commissure of the superior

 

 

colliculi are separated. The floor of

 

 

the mesencephalon has become

 

 

greatly thickened by an increase in

 

 

ascending and descending fibers. The

 

 

isthmic groove, formerly the isthmic

 

 

recess, is scarcely visible. The

 

 

rostrocaudal extent of the fourth

 

 

ventricle is relatively less. The ventral

 

 

thalamus becomes so reduced that

 

 

the former dorsal thalamus, now

 

 

termed merely the thalamus, borders

 

 

directly on the subthalamus, as

 

 

established by Richter (1965) using

 

Fig. 24 – 28

excellent fetal preparations. (B) The

 

 

overlapping of the left cerebral

 

 

hemisphere on the left half of the

 

 

diencephalon. The hippocampus, the

 

 

dentate area, the area epithelialis, and

 

 

the choroid fissure are shown as if the

 

 

prosencephalon were transparent.

 

 

The lamina affixa is cross-hatched.

 

 

The olfactory bulb is no longer

 

 

directed caudally, a change that is

 

 

perhaps related to a rostral extension

 

 

of the nasal cavities (Hochstetter,

 

 

1919). In slightly older fetuses the

 

 

olfactory bulb grows rostrally and

 

 

becomes thinner and longer,

 

 

concomitant with the lengthening of

 

 

the frontal part of the cerebral

 

 

hemisphere. The site of the entrance

 

 

of the lateral prosencephalic

 

 

fasciculus into the diencephalon is

 

 

stippled. Based partly on Hines

 

 

(1922). The levels of two sections,

 

 

Figures 24–27 and 24–28, are

 

 

indicated. Three-dimensional

 

 

reconstructions, prepared

 

Fig. 24 – 27

ultrasonically, of the ventricles in vivo

 

 

in fetuses of 38 or 40 mm are

 

 

illustrated by Blaas et al. (1995, 1998).

 

EARLY POSTEMBRYONIC PHASE

279

 

Vermis

 

 

Epiphysis

 

Nodule

 

Hemisphere

 

B

Flocculus

C

M

 

 

 

Internal

 

 

Vermis

A

cerebellar

 

 

 

 

 

swelling

Figure 24–19. Topographical changes in the cerebellum of the first trimester. The flocculonodular lobe (or vestibulocerebellum) is the archicerebellum, which is formed differently from the rest of the vermis and is considered by some not to belong to the cerebellum sensu stricto. Both the vermis and the hemispheres will contribute to the paleocerebellum (or spinocerebellum) and the neocerebellum (or corticopontocerebellum). (A) At stage 20 the floccular region begins to become tilted from a rostrolateral to an occipitolateral position. The internal cerebellar swellings are well-separated. Further growth leads to their fusion, which begins in stage 23. (B) At 9 weeks (42 mm) the flocculonodular lobe is pointed in an occipital direction. Growth of the extraventricular cerebellum leads to an eversion of the cerebellar hemispheres and a lengthening of the vermis, resulting in the shape of a dumbbell. (C) At 13 weeks (95 mm) several folia and fissures can be distinguished in the vermis.

B and C are based on reconstructions in Streeter (1912, where No. 86, 30 mm, is given incorrectly instead of No. 886, 42 mm).

PL

Nodule

2

Figure 24–20. Median section through embryonic and fetal vermian region showing chief fissures and folia. (See also Table 24-2). Modified from Rauber-Kopsch, Anatomie des Menschen.

Abbreviations: 1, fissura prima. 2, fissura secunda. ANT., anterior lobe. PL, posterolateral fissure. POST., posterior lobe. Prepyr., prepyramidal fissure. Pyr., pyramid. Uv., uvula.

Fusion of the cerebellar hemispheres, although not accepted by Sidman and Rakic (1982), had been supported by Streeter (1912), Jakob (1929), Hochstetter (1939), and Larsell (1947), and is confirmed here. The paired cerebellar

hemispheres began to unite at stage 23, thereby initiating the formation of the vermis (Fig. 23–26). The fusion is probably not completed, however, until the end of the first trimester (Feess-Higgins and Larroche, 1987).

280

 

 

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

 

 

 

 

 

FISSURES

FOLIA

 

 

 

 

Horizontal

 

 

 

 

 

 

Tuber

A

 

 

1

Prepyramidal

 

 

 

1

 

 

Pyramid

 

Hemisphere

 

 

 

 

2

2

Postpyr. = secunda

 

 

 

 

 

 

 

 

 

 

 

 

3

 

Uvula

 

 

 

3

Posterolateral

 

 

 

 

 

 

Nodule

1

Horizontal

 

 

2

B

Tonsil

3

Paraflocculus

Flocculus

Tonsil

Paraflocculus

Flocculus

C

Pyramid 1

2

Uvula

3

Nodule

Figure 24–21. Simplified dorsal view of the fetal cerebellum with only the chief features indicated. The fissura prima is not visible in these views. (A) at 13 weeks (95 mm) from the same embryo as in Figure 24–19C. (B) and (C) later in the first half of prenatal life. Based on Streeter in Keibel and Mall (1912).

Histogenesis of the Cerebellum. Homeobox genes are believed to be involved in the initial development of the external germinal layer, and interaction with glial fibers is necessary for the internal cellular migration. Cell proliferation in the external germinal layer is the main cause of the increase in the cerebellar surface.

Piriform (Purkinje) cells are generated at the beginning of the fetal period (Rakic and Sidman, 1970) and their characteristic shape is attained by the middle of prenatal

life. Their maturation continues after birth (Zecevic and Rakic, 1976). Modified radial glial (Bergmann) cells are necessary for the migration of the Purkinje cells (Choi and Lapham, 1980).

Photomicrographs of the fetal cerebellum have been published by Rakic and Sidman (1970), Zecevic and Rakic (1976), and Choi and Lapham (1980).

The development of the human cerebellum is different from that in the rat (Muller¨ and O’Rahilly, 1990a,b).

EARLY POSTEMBRYONIC PHASE

A

Marginal

Ventricular

Stage 13

14

15

18

23

LAYERS:

Intermediate

Molecular Æ fiber

Ext. germinal

Piriform

B

Deep nucleus

Ventricle

Ventricular

281

Molecular

Piriform

Granular

White matter

Ependyma

Figure 24–22. (A) The embryonic period as interpreted by the authors. A marginal layer becomes added (stage 14) to the ventricular layer. An intermediate layer is seen at stages 15–18. A sheath of fibers is developing superficially at stage 18. The external germinal layer begins to appear in stage 23. The dentate nucleus will form from a cellular lamina between two layers of fibers. (See Figure 21–19.) (B) The fetal period. The external germinal layer, which began to form at the end of the embryonic period, continues to receive cells from the rhombic lip. Near the end of the fetal period its cells migrate internally (purple arrow) and gives rise to the internal granular layer and basket cells. The external germinal layer disappears after birth. In the adult (last column) two Purkinje cells are shown, one in a sagittal, the other in a transverse plane. Their axons reach the deep nuclei. A mossy fiber is shown leading to a glomerulus which contains a mossy fiber rosette. The glomerulus is contacted by granule and Golgi cells. The dendrites of Purkinje, granular, and Golgi cells reach the molecular layer.

282

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

Figure 24–23. 46 mm. A nearly median section. The dorsal thalamus, which appears dark because it is cut at the level of the ventricular layer, is well shown. It is separated from the ventral thalamus by a band of gray material, which is probably the tract of the zona intrathalamica. When the individual thalamic nuclei are said to appear depends on the criteria selected, and these are mainly the establishment of cell-poor boundaries and the presence of nerve fibers, followed by differentiation into nuclear groups. Further information: Dekaban (1954), Fabiani and Barontini (1956), Yamadori (1965). The lateral geniculate body shows lamination during trimester 2. The pulvinar, which begins to develop at the commencement of trimester 2, is derived mainly from the medial ventricular eminence and grows greatly during the second half of prenatal life. Migrating cells from the medial ventricular eminence cover the fetal corpus striatum adjacent to the sulcus terminalis (constituting the “corpus gangliothalamicum” of Rakic and Sidman, 1969) and are said to participate in the formation of other thalamic nuclei (Letinic,´ and Kostovic,´ 1997). See Figure 24–9.

The epiphysis cerebri, the posterior commissure, and the commissure of the superior colliculi are all recognizable. The three subdivisions of the axis (termed X, Y, and Z by O’Rahilly, Muller,¨ and Meyer, 1983, in their account of the occipitocervical region) are evident. A key drawing is provided with Figure 23-6.

Inf. cerebellar peduncle

EARLY POSTEMBRYONIC PHASE

283

Figure 24–24. An ultrasonic image at 35 mm. The mesencephalon and the diencephalon can be identified, and the choroid plexus of the lateral ventricle (cf. Fig. 24–26) is evident. Courtesy of Dr. Harm-Gerd Blaas, Trondheim, Norway.

284

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

Figure 24–25. (A) dorsal and (B) lateral views of the brain at 42 mm, with parts of the hemisphere removed. In A the external capsule, not clearly distinguishable in stage 23, is now visible. In B the basal nuclei, choroid fissure, and internal and external capsules are exposed. The part of the cerebral hemisphere removed here is indicated in Figure 24–28 by a dotted line. These instructive views are (with a few corrections) based on Kollmann’s Handatlas der Entwicklungsgeschichte des Menschen (1907).

C H A P T E R

CH

24

TRIMESTER 1: LATER POSTEMBRYONIC PHASE

GL

100 mm

13W

Approximately 50–100 mm in Greatest Length; Approximately 9–13 Postfertilizational Weeks

The cerebral hemispheres continue their rapid enlargement and reach a size approximately three times that at the end of the embryonic period. The cor-

pus callosum and the fornix are noteworthy among the expanding C-shaped structures. The increase of the forebrain angle during trimester 1 brings about a “rotation” of the olfactory bulbs such that they now point caudorostrally rather than rostrocaudally. The anterior commissure

is more evident and is developing connections with olfactory centers and with the amygdaloid nuclei. Folia and fissures are appearing in the vermis cerebelli. The posterior horn of the lateral ventricle is beginning to evaginate.

Mathematical and hormonal criteria have been used (by Guyot) to indicate an advance in fetal development when the GL is approximately 90 mm at some 90 postfertilizational days.

The Embryonic Human Brain: An Atlas of Developmental Stages, Third Edition. By O’Rahilly and Muller¨ Copyright C 2006 John Wiley & Sons, Inc.

285

286

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

Fig. 24 – 26

Figure 24–26. A sagittal section at 40 mm. The relatively great size of the choroid plexus is noticeable. The key shows the hippocampal thickening. (The cortical plate is present in most of the hemispheres.) The dorsal thalamus is closely related to the ventricular eminences. Thalamocortical fibers are already numerous, although the thalamic nuclei have yet to form. The habenulo-interpeduncular and mamillotegmental tracts can be followed to the mesencephalic tegmentum. The decussation of the trochlear nerves is identifiable in the isthmus. The internal cerebellar swelling is large. The rhombic lip can be seen here and again further caudally at the end of the medulla oblongata.

LATER POSTEMBRYONIC PHASE

287

Figure 24–27. 50 mm. A coronal section showing the anterior commissure, the caudate nucleus, the internal capsule, and the putamen. Based on a photomicrograph in Feess-Higgins and Larroche (1987).

Figure 24–28. 37 mm. A coronal section showing the interventricular foramina and a number of important relationships. Based on an excellent photo-micrograph in Richter (1965). The dotted line indicates the part of the cerebral hemisphere removed in Figure 24–25B.

Cholinesterase-reactive fibers from the nucleus basalis complex are distributed widely to the neocortex and limbic cortex by the end of trimester 2 (Kostovic,´ 1986).

Fig.

24 – 25