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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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188

C h a p t e r 2 1 : THE FIRST APPEARANCE OF THE CORTICAL PLATE IN THE CEREBRAL HEMISPHERES

Figure 21–1. An embryo attached to the placenta and showing the superficial vascular plexus of the head. In the cord an umbilical vessel is visible, as are also coils of intestine belonging to the normal, temporary umbilical hernia.

 

 

 

 

 

 

Figure 21–2. Right lateral view of the brain with

 

 

 

 

 

 

the membranous labyrinth added. A flat surface

 

 

 

 

 

 

marks the site of the future insula. Frontal,

 

 

 

 

 

 

occipital, and temporal poles are distinguishable

 

 

 

 

 

 

(α, β, γ ). The cerebral hemispheres overlap more

Di.

 

 

 

 

b

than half of the surface of the diencephalon. The

 

 

 

 

 

 

 

 

isthmic groove has deepened. Between 6 and 7 1

 

 

 

 

 

 

 

 

 

 

 

 

2

 

g

 

weeks the head rises in position. This is largely

 

 

 

 

 

 

because at stage 21 the growth and expansion of

 

 

 

 

 

 

the trunk are such that it could not remain in the

 

 

 

 

 

 

position it occupied at stage 18 (although in both

 

 

 

 

 

 

stages it rests on the ventral wall of the trunk). A

 

 

 

a

 

 

 

 

 

number of features typical of this stage were

 

 

 

 

 

 

 

 

 

 

 

 

described by Hochstetter (1919, his embryos Ha3

 

 

 

 

 

 

and Peh4).

THE FIRST APPEARANCE OF THE CORTICAL PLATE IN THE CEREBRAL HEMISPHERES

189

Isthmic

21 –

18

17

.

a

b

c

Figure 21–3. Graphic reconstruction from transverse sections to show a median view of the brain. The asterisk indicates the junction with the spinal cord. The pineal recess has formed. Ependymal cells over the posterior commissure form the subcommissural organ. The floor of the rhombencephalon is bent sharply, as described by Hochstetter (1919), and a transverse groove, the sulcus transversus rhombencephali, results. The isthmic recess (short arrow) is a guide to the interpeduncular nucleus. Interrupted lines delimit the rhombencephalon, mesencephalon, diencephalon, and telencephalon.

Striatal arteries in this embryo were shown by Padget (1948, Fig. 21).

As shown in the inset, the main internal grooves are (a) the sulcus limitans, (b) the hypothalamic sulcus, and (c) the marginal ridge.

190

C h a p t e r 2 1 : THE FIRST APPEARANCE OF THE CORTICAL PLATE IN THE CEREBRAL HEMISPHERES

Figure 21–4. (A) The overlap between the left cerebral hemisphere and the left half of the diencephalon. The positions of the subthalamic nuclei (medial and lateral mamillary nuclei, subthalamic nucleus proper, entopeduncular nucleus which is the future globus pallidus internus, and the globus pallidus externus) are shown schematically. The ventral part of the lateral geniculate body has developed (Table 20–1). The hippocampus, area dentata, area epithelialis, and the choroid fissure are indicated on the medial wall of the left hemisphere, which is shown as if transparent. The hippocampus reaches almost as far as the temporal pole.

The site of the future lamina affixa, which at this stage overlies the ventral thalamus, is marked by a dagger. The asterisk at the base of the telencephalon is at the site of a cistern. Parts not visible from the surface are shown by interrupted lines. The horizontal interrupted line delimits the diencephalon from the telencephalon. (B) Levels of Figures 21–11 to 21–17, which, because they are from a different embryo, do not necessarily show a

perfect fit.

Fig. 21 –

11

12

13

14 6, 15 17

16 9, 10

THE FIRST APPEARANCE OF THE CORTICAL PLATE IN THE CEREBRAL HEMISPHERES

191

Figure 21–5. An almost median section. The olfactory and lateral ventricles are continuous. Choroid folds can be seen in the lateral and fourth ventricles. In the diencephalon, the zona limitans intrathalamica is distinguishable between the dark, dorsal area (dorsal thalamus) and the less dark, ventral area (ventral thalamus). The posterior commissure is partly diencephalic and partly mesencephalic (commissure of the superior colliculi). The interpeduncular nucleus is immediately rostral to the tuberculum interpedunculare of the isthmus, which segment includes also the trochlear commissure and the isthmic recess. In the cerebellum, the internal fiber layer is detectable. The sulcus transversus rhombencephali is produced by the sharp bend of the floor of the rhombencephalon. The inferior olivary nucleus is ventral in the rhombencephalon. The base of the chondrocranium shows the dorsum sellae and the hypophysial fossa (containing the hypophysis cerebri), and continues rostrally into the nasal septum. The dens of cervical vertebra 2 is evident.

192

Hemispheric stalk

C h a p t e r 2 1 : THE FIRST APPEARANCE OF THE CORTICAL PLATE IN THE CEREBRAL HEMISPHERES

Lamina affixa

Sulcus terminalis Medial

ventricular eminence

Cortical plate

Lateral

prosencephalic fasciculus Globus pallidus externus

Figure 21–6. A portion of the right cerebral hemisphere adjacent to the diencephalon. The groove between the two (di-telencephalic sulcus) is filled with loose mesenchyme. The choroid plexus now presents stumpy villi limited by three to four rows of epithelial cells. The layer adjacent and ventral to the plexus is the future lamina affixa, which at this stage lies opposite the ventral thalamus. In the adjacent massa cellularis reuniens

(Richter, 1965) cellular strands (asterisk below) possibly indicate migrating cells. A deep groove, the sulcus terminalis, is situated medial to the medial ventricular eminence. The germinal zone (matrix and subventricular layer) of the ventricular eminences is bordered immediately by the postmitotic layer, without an intervening intermediate zone. The cortical plate, which is typical of stage 21, has begun to form opposite the ventricular eminences. It was seen by Molliver et al. (1973). The transition from the hemisphere to the diencephalon is the hemispheric stalk, which contains a thick bundle of fibers, the lateral prosencephalic fasciculus (Stammbundel¨ of His). The telencephalic area adjacent to it is the amygdaloid region. The diencephalic nucleus ventral to the lateral prosencephalic fasciculus is the globus pallidus externus.

The hemispheric stalk seen in this figure is the original connection between the diencephalon and the telencephalon. It will become greatly enlarged by fibers and tracts, especially by the continuation of the internal capsule.

The levels of Figures 21–6 to 21–10 are shown in Figure 21–4B. The bars represent 0.1 mm.

Optic tract

THE FIRST APPEARANCE OF THE CORTICAL PLATE IN THE CEREBRAL HEMISPHERES

193

Figure 21–7. Developmental comparison between the spinal cord and the brain wall as derivatives of the neural tube. The cortical plate develops within the primordial plexiform layer (preplate), so that it comes to lie between superficial and deep laminae, namely layer 1 and the subplate of the neocortex. The stages illustrated are 11 for the neural tube, and 17 for the spinal cord. The neocortical blocks are (left) up to stage 20, and (right) from stage 21.

194

C h a p t e r 2 1 : THE FIRST APPEARANCE OF THE CORTICAL PLATE IN THE CEREBRAL HEMISPHERES

A B

Sulcus

limitans

BASAL ALAR

Floor

plate

 

.

 

Hypothal

 

Ma.

 

sulcus

 

 

Ivf

C

Ch.

D

NH

AH

Ch.

 

 

Ad.

Figure 21–8. A median view of the brain at stage 22 to show the termination of the alar and basal laminae, and that of the sulcus limitans in the brain. The chief interpretations of the ending of the sulcus limitans, as summarized by Richter (1965, Fig. 35), are as follows:

(A) The supramamillary recess (Kingsbury; Schulte and Tilney; Kuhlenbeck). (B) The prechiasmatic recess (His; Johnston; Streeter). (C) The interventricular foramen (Spatz; Kahle; Richter). Interpretation A is that supported by the present authors for the human embryo. (D) An enlargement of the commissural plate to show three subregions described in the rat: (a) the future anterior commissure; (b) the probable site of closure of the rostral neuropore; and (c) the future hippocampal commissure and the corpus callosum. With further development of these commissures, the embryonic lamina terminalis becomes reduced to the adult type (Ad.), which extends from the caudal limit of the commissure to the optic chiasma.

THE FIRST APPEARANCE OF THE CORTICAL PLATE IN THE CEREBRAL HEMISPHERES

195

Subpial layer

Cortical plate

Subplate

Intermediate

layer

Ventricular & subventricular layers

Figure 21–9. The cortical plate is visible within the cell-poor primordial plexiform layer at the periphery of the ventricular eminences. The plate separates the former primordial plexiform layer into the subpial layer, which is future layer 1, and the subplate (Fig. 21–7). The subplate contains thalamocortical fibers that arrived in the primordial plexiform layer already at the previous stage. Penetration of the cortical plate by monoamine fibers occurs at the end of trimester 1 (Verney et al., 1993). It is to be noted that the subpial layer and the subplate appear during the embryonic period and not, as has been claimed (Kostovic´ and Rakic, 1990), in the middle third of prenatal life. Transient fibers in the subplate are illustrated in Figure 23–22. Glial production begins from stem cells in the subventricular layer. From Muller¨ and O’Rahilly (1990b).

196

C h a p t e r 2 1 : THE FIRST APPEARANCE OF THE CORTICAL PLATE IN THE CEREBRAL HEMISPHERES

Nucleus of diagonal band

Lat. em.

Cortical plate

Med. em

Lat. olf. fibers

Piriform cortex

Figure 21–10. An overall view of the septal area showing the nucleus of the diagonal band. The prosencephalic septum is closely related to the hippocampal region, which lies dorsomedially. A faint groove separates the lateral and medial ventricular eminences. The cortical plate of the neocortex is contiguous to the piriform cortex. Lateral olfactory fibers are visible. The loose tissue surrounding the base of the telencephalon is a basal cistern (asterisk in key on next page.)

THE FIRST APPEARANCE OF THE CORTICAL PLATE IN THE CEREBRAL HEMISPHERES

197

21 – 11

12

V3

13

Mam. recess

 

 

Habenulo-

Subthalamic

Basilar a.

V3

 

inter-

 

nucleus

 

 

 

peduncular

 

 

 

 

 

 

 

tract

Figure 21–12. Section through the mamillary recess showing the

 

 

 

 

 

 

migratory stream from the mamillary area to the subthalamic nu-

 

 

 

cleus. The fibers, which are from the mamillotegmental tract, arise

 

 

 

in the lateral, as well as the medial, mamillary nucleus.

Mamillothalamic tract

Mamillary area

Figure 21–11. The di-mesencephalic transition showing the mamillary area with its budding surface (“leicht hockerige¨ Oberflache¨ ,” Hochstetter, 1919; “mamelones de crecimiento,” Orts Llorca, 1977). The buds are indicated by arrows. The habenulointerpeduncular tract runs between the ventricular and intermediate layers, and this part of the section is the caudal area of the diencephalon. Ventrally, its median cellular material seems to be a continuation of the interpeduncular nucleus of the midbrain. Large meshes occur in the subdural space and are generally near the cerebral wall, whereas smaller meshes are found near the skeletogenous layer.

V3

Figure 21–13. The mamillothalamic tract runs from the lateral mamillary nucleus to the thalamus. The nucleus appears on each side as a dark, basal area.

The levels of Figures 21–11 to 21–13 are shown also in Figure 21–4B. The bars represent 0.2 mm.