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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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g c

Raphe magnus nuclei

Somatic afferent

Visceral afferent

Visceral efferent

Somatic efferent

Inferior cerebellar peduncle

Figure 23–29. Dorsal view of a reconstruction of the rhombencephalon (silver-impregnated), showing nuclei and tracts. The caudal direction is towards the top of the page. The right embryonic half is on the left of the drawing. The cerebellum has not been included. The white line on the right-hand side of the drawing represents the cut edge of the roof of the fourth ventricle. Black triangles indicate the levels of the reconstructed cross sections in Figure 23–30. The nucleus funiculi teretis (Kappenkern of the genu of the facial nerve) is included. The two (ascending and descending) intramural parts of the facial nerve are indicated by the numeral 7.

The inset shows the four vestibular nuclei.

The rhombic lip, an important proliferative area in the alar plate, is the source of two superficially situated migratory areas: the so-called olivo-arcuate caudally, and the pontine rostrally. Two of the three migratory areas found at stage 23 have been reconstructed (Muller¨ and O’Rahilly, 1990c, Fig. 1). The term “olivo-arcuate migration” (Essick, 1912), however, is unsatisfactory, because it has been shown (in the monkey) that the olivary nuclei arise mainly from the ventricular layer.

The rhombencephalon at this stage is already very complicated and comparable in many respects to that of the newborn (Fig. 26–5). Its rapid development suggests an early onset of functional activity.

7

Vestibulo-

spinal tr.

and cuneo-

Figure 23–30. Reconstructed coronal sections of the rhombencephalon. The levels of the sections (A)–(F) in rostrocaudal succession are indicated in Figure 23–29 by black triangles. (A) Section showing the nuclei of the trigeminal nerve and the transition to the cerebellum via the inferior cerebellar peduncle. The position of the trigeminal nuclei here is conditioned by the slight obliquity of this slice. The inset in the lower right-hand corner of the page shows fiber bundles of the motor part, of the intermediate, and of the sensory portion of the trigeminal nerve. (B) Area of the abducent and facial nerves, showing the superior olivary nucleus, and the superior vestibular nucleus and its fibers to the medial longitudinal fasciculus. (C) Entry of the cochlear nerve. Ventral and dorsal cochlear nuclei are lateral to the inferior cerebellar peduncle. The fibers from the dorsal cochlear nucleus that run medially constitute the intermediate acoustic striae. A dagger indicates the tectobulbar and tectospinal tracts. (D) Entry of the sensory glossopharyngeal fibers that run directly to the tractus solitarius. Olivocerebellar fibers can be seen passing towards the inferior cerebellar peduncle. (E) and (F) Inferior olivary, vagal, accessory, and hypoglossal nuclei. A small bundle of corticospinal (pyramidal) fibers is present in all the sections.

250

C h a p t e r 2 3 : THE BRAIN AT THE END OF THE EMBRYONIC PERIOD

Figs. 23 – 29 &

23 – 31

a b

c

Three decussations are shown from dorsal to ventral:

(a)lemniscal produced by cuneate fibers in the medulla (c);

(b)lemniscal produced by gracile fibers in the medulla (g); and

(c)pyramidal in the cervical spinal cord and shown in greater detail in Figure 23-32.

Figure 23–31. Section showing in the median plane, from above downwards, spinal cord (cervical region with pyramidal decussation), nerve roots, septum medullae (with olivary nuclei immediately at each side), basilar artery, third ventricle, and diencephalon. Laterally, and again from above downwards, the jugular process (medial to which is the leptomeningeal or future subarachnoid space), otic capsule (including sections of three semicircular ducts), dura (a prominent line encircling both spinal cord and brain), and a small portion of the cerebral hemisphere.

THE BRAIN AT THE END OF THE EMBRYONIC PERIOD

251

Cuneatus

Gracilis

 

Lat. corticospinal

Motor

neurons

Ant. corticospinal

Figure 23–32. The pyramidal decussation in the transitional region of the medulla oblongata and spinal cord.

252

C h a p t e r 2 3 : THE BRAIN AT THE END OF THE EMBRYONIC PERIOD

Figure 23–33. The arteries in a dorsal view, with a key drawing on the left. This is the first published reconstruction of the arteries at stage 23. The right and left sides were reconstructed separately, and the two sides are slightly different. Most cranial nerves, the optic chiasma, and part of the optic tracts are shown. The main components of the circulus arteriosus have been present since stage 16 (Fig. 16–5) and the circle has been complete since stage 19 (Fig. 21–23). The arteries to the choroid plexus of the lateral ventricles come from one of the deep branches of the anterior cerebral artery (anterior choroid a.) and from the posterior cerebral artery (posterior choroid a.). Two anterior communicating arteries are present, a frequent finding later in life. Apart from differences in the proportions, the arterial pattern resembles closely that of the adult.

THE BRAIN AT THE END OF THE EMBRYONIC PERIOD

253

Figure 23–34. Reconstruction of the arteries in a right lateral view with a key drawing below. The overlapping that occurs in a dorsal representation is omitted. The posterior communicating artery is clearer than in Figure 23–33. Noteworthy are the serpentine course of the internal carotid and the many striatal branches that penetrate the anterior perforated substance. The ramifications of the middle cerebral are not shown. The anterior cerebral gives off a branch to the olfactory bulb and significant branches to the medial surface of the cerebral hemisphere. An example of the differing proportions in the embryo is seen in the posterior communicating artery, which joins the internal carotid long before the latter divides into the anterior and middle cerebral arteries.

The internal carotid develops early (stages 11–13) and is followed by the posterior communicating (stage 14), basilar and vertebral (stage 16), anterior, middle, and posterior cerebral (stage 17), and finally the anterior communicating (stage 21), thereby completing the circulus arteriosus (Figs. 21–22 and 21–23). Initially the posterior communicating is an important channel, and its distal end constitutes the stem of the posterior cerebral. It supplies the hindbrain until the vertebral system is completed and the basilar artery becomes dominant. The hypoglossal artery disappeared after stage 15, and the stapedial artery likewise at about stage 20, whereas the trigeminal artery is still present.

The main channels of the venous system correspond to the reconstructions of a stage 21 (24 mm) embryo shown by Padget (1957). The cavernous sinus is not yet present. The beginning development of the superior sagittal sinus is shown in Figure 23–9. Bilateral foramina for emissary veins are present in the parietal part of the chondrocranium. The capsuloparietal foramen constantly contains a vein that connects the transverse sinus to the external surface of the chondrocranium. A bilateral emissary foramen in the occipital contains a vein that connects the sigmoid sinus with the external surface (Muller¨ and O’Rahilly, 1980).

254

C h a p t e r 2 3 : THE BRAIN AT THE END OF THE EMBRYONIC PERIOD

A

B

C

Rh.

M

 

M

 

 

 

 

 

 

 

Rh.

 

Di.

Di.

 

Di.

 

 

 

Hemispheres

 

 

 

 

 

Hem.

Hem.

 

 

 

Figure 23–35. Examples of in vivo ultrasonic images at 6–7 12 postfertilizational weeks, i.e., during the embryonic period.

(A) 12.9 mm. Obliquely coronal section showing cerebral hemispheres. (B) 15 mm. Sagittal view showing mesencephalic flexure. (C) 15 mm. Another obliquely coronal section showing hemispheres.

 

THE BRAIN AT THE END OF THE EMBRYONIC PERIOD

255

D

E

 

Cerebellum

Lateral recess

M

M

Rh.

 

Di.

 

Cbl

Choroid

 

 

 

 

plexus

 

 

 

 

 

 

V4

 

 

 

Medulla

 

Spinal cord

 

oblong.

 

 

 

Figure 23–35. (Continued ) (D) 18 mm. Sagittal view showing the spinal cord. (E) 25 mm. Coronal section, the “hole in the head” view. Courtesy of Dr. Harm-Gerd Blaas, Trondheim, Norway.

256

C h a p t e r 2 3 : THE BRAIN AT THE END OF THE EMBRYONIC PERIOD

A B C D E

Figure 23–35. (Continued ) In vivo ultrasonic 3D reconstructions of the ventricular system at 5 12 –9 postfertilizational weeks. The isthmic region in these views appears narrower than that seen in graphic reconstructions.

(A)10.1 mm. The rhombencephalic cavity is at the top of the head.

(B)17 mm. The hand is distinct.

(C)21.5 mm. The mesencephalic cavity is relatively large.

(D)29 mm. The “praying feet” position is characteristic.

(E)38 mm. The volume of the cerebral hemispheres keeps increasing.

Color scheme for this figure: Yellow, lateral ventricle. Green, third ventricle. Red, mesencephalic cavity (future aqueduct). Blue, fourth ventricle. Courtesy of Dr. Harm-Gerd Blaas, Trondheim, Norway.

Figure 23–35. (Continued ) These two views are of the embryo marked B on page 254. With the addition of contours they were used in the calculation of volumes of the embryonic body and of the ventricular cavities (Blaas et al., 1998).

THE BRAIN AT THE END OF THE EMBRYONIC PERIOD

257

TABLE 23–1. Some Examples of Genes Already Known to Be Implicated in the Development of the Human Nervous System

Genes

Neural Region

Malformations

References

 

 

 

 

En-2

Mes-rhombencephalon

 

Song et al., 1996

HOXA3

Neural crest of pharyngeal arches

 

Goodman, 2003

HOXD13 (hand)

 

With HOXA13 the only two

Goodman, 2003

 

 

HOX genes mutated in

 

 

 

human

 

HOXA13 (genital)

 

 

Goodman, 2003

PAX 1

 

Spina bifida

Hol et al., 1996 (cited in Dahl

 

 

 

et al., 1997)

PAX3

Neural plate and early fusion

Dysraphia

Gerard´ et al., 1995

PAX3

Neural crest

Waardenburg syndrome;

Baldwin et al., 1992 (cited in

 

 

mutation in 50%

Dahl et al., 1997)

PAX5

Mes-rhombencephalic

 

Gerard´ et al., 1995

 

boundary and spinal cord

 

 

PAX 6

“Master control gene”

Aniridia;

Gerard´ et al., 1995

 

 

congenital cataract (Peter

MacDonald and Wilson, 1996

 

 

anomaly)

(cited in Dahl et al., 1997)

PAX 6

Early morphogenesis of eyes

 

Goodman, 2003

ROBO3

Crossing of corticospinal and

Gaze palsy with progressive

Jen et al., 2004

 

somatosensory axons in

scoliosis

 

 

medulla

 

 

SHH

Brain

Holoprosencephaly

Belloni et al., 1996

SHH

Organogenesis of brain, eye,

Holoprosencephaly

Muenke and Cohen, 2000

 

somites, spinal cord,

 

 

 

craniofacial structures

 

 

SHH

Signal of prechordal plate

Holoprosencephaly

Goodman, 2003

 

inducing forebrain

 

 

SHH

Differentiation of floor plate,

 

Roessler et al., 1996

 

rostrocaudal axis

 

 

SIX 3

 

Deletion on chromosome

Oliver et al., 1995

 

 

2p21–p22: holoprosencephaly

 

 

 

type 2

 

SIX 3

 

At 2p21 4 different mutations

Schell et al., 1996

 

 

in HPE patients

Wallis et al., 1999 (cited in

 

 

 

Muenke and Cohen, 2000

SIX 3

 

Gene mutated in some

Pasquier et al., 2000

 

 

patients with HPE

 

SIX 3

SIX 3 at 2p21 present from

 

Grenadino et al., 1999

 

5–7 weeks

 

 

ZIG2

Formation and differentiation

Holoprosencephaly with mild

Goodman, 2003

 

of neural tube

facial dysmorphism

 

 

 

Examples in Mouse and Rat Based on Current Literature

 

SIX

 

Expressed in neural plate rostrally

Otx2

 

Mesencephalon

 

En-1, En-2

 

Mes-rhombencephalon

Gbx2

 

Rhmbomere 1, cerebellum

Dlx-2

 

Medial ventricular eminence, ventral thalamus

Emx1, 2

 

Dentate gyrus