Добавил:
Опубликованный материал нарушает ваши авторские права? Сообщите нам.
Вуз: Предмет: Файл:

Книги по МРТ КТ на английском языке / The Embryonic Human Brain An Atlas of Developmental Stages. Third Edition. 2006. By Ronan O'Rahilly

.pdf
Скачиваний:
2
Добавлен:
05.10.2023
Размер:
37.7 Mб
Скачать

308

C h a p t e r 2 6 : TRIMESTER 3 AND THE NEWBORN

Choroid

fissure

Commissure

of fornix

Indusium griseum & hippocampus

Corpus callosum

Fornix

Stria terminalis

Thalamus

Ant. commissure

Figure 26–3. A comparison of the medial surface of the forebrain at (A) 80, (B) 95, (C) 150, and (D) approximately 265 mm. The left thalamus has been sectioned sagittally (shaded by horizontal lines). In A, B, and C the entire brain stem has been removed, whereas in D it has been sectioned transversely at the level of the midbrain and its caudal portion has been eliminated. The considerable increase in length of the corpus callosum (shown in yellow) after 100 mm can readily be appreciated by comparing B and C. In D the corpus callosum shows clearly the rostrum, genu, central part, and splenium. The commissural plate beneath the corpus callosum becomes attenuated (C) to form the septum pellucidum. Modified from models in Keibel and Mall (1912).

The upper right-hand drawings show schematically the arrangement of the pia mater over the corpus callosum at 100 mm, 170 mm, and at birth. As the corpus callosum grows posteriorly, a pial layer (b) is reflected backwards over the original layer (a), so that a double fold is formed. This is the velum interpositum, between the two layers of which blood vessels can pass forward beneath the corpus callosum, through the transverse fissure (asterisk). These vessels contribute to the tela choroidea of the third ventricle.

 

 

 

 

TRIMESTER 3 AND THE NEWBORN

 

309

 

b

M

 

 

 

Figure 26–4. Left lateral views of the

 

 

 

 

brain during the first half of prenatal

 

 

 

 

 

 

 

g

 

b

 

life. The gradual concealment of the

a

I

 

 

midbrain is evident. The future insula

 

 

 

 

(I) is relatively stationary, whereas,

 

 

 

 

 

 

30 mm

 

 

 

accompanying the sweep of the

 

 

 

 

developing temporal pole (γ ), the

 

 

 

 

 

 

 

 

 

 

I

 

future occipital pole (β) is displaced

 

 

 

a

g

b

along a similarly curved path (inset). By

 

 

 

 

approximately 200 mm the central

 

 

 

 

 

 

 

 

53 mm

 

 

sulcus begins to appear and the frontal,

 

 

 

 

frontoparietal, and temporal opercula

 

 

 

 

 

 

 

 

 

I

are covering the insula.

 

 

 

a

 

 

 

 

 

 

 

b

 

 

 

g

 

 

 

68 mm

 

g

 

 

 

 

 

 

 

 

 

a

 

 

 

 

 

I

 

 

 

 

 

a

I

b

 

 

96 mm

g

Frontoparietal

 

Frontal

Temporal

 

ca 200 mm

 

 

Functional Considerations

Cerebral electrical activity has been recorded ex utero in the embryonic period, and potentials have been detected in utero in trimester 3 both abdominally and per vaginam.

Prenatal behavior includes movements, sleep (accompanied by rapid eye movements in trimester 3) and wakefulness. Cardiac motion ban be detected by ultrasound at 31/2 weeks at a rate of 100 per minute. Body movements can be recognized ultrasonically at 5–6 weeks (stage 16) and consist initially of extension and flexion of the neck and the thoracic region. Reflex and spontaneous movements probably begin simultaneously. More complex movements follow rapidly and form coordinated patterns dependent on the

structural maturation of the nervous system. Movements include startles, hiccups, isolated limb movements, and head rotation; later during trimester 1 they include breathing movements, yawning, sucking, and swallowing (of amniotic fluid), all of which are similar to those occurring postnatally. All motor patterns seem to be present by the beginning of trimester 2, when movement is almost constant.

Functional development of the cochlea takes place during trimesters 2 and 3. Hearing (by bone conduction in a fluid environment) and response to sounds occur by the middle of prenatal life, and a little later the fetus can discriminate between different frequencies and between speech stimuli; music is welcomed.

See Figure 23 – 29

Somatic afferent

Visceral afferent

Visceral efferent

Somatic efferent

Inferior cerebellar peduncle

Figure 26–5. Dorsal views of the rhombencephalon (A) at 8 postfertilizational weeks (stage 23: cf. Figs. 23–29 and 23–30) and (B, C) in two different reconstructions from the newborn. The arrangement of the nuclei and tracts, as well as the location of the rhombencephalic nuclei at 8 weeks, are very similar to those in the newborn. The striking resemblance between the rhombencephalon of the newborn and that of the embryo at 8 weeks is based on the circumstance that the fundamental organization of the rhombencephalon is attained much earlier than that of other parts of the brain. Figure 26–5B is based on a reconstruction by Sabin (1901). Figure 26–5C is after Hikji (1933). The neurons in the nucleus of the tractus solitarius are still immature at birth (Denawit-Saubie´ et al., 1994).

Figure 26–6. The main cerebral sulci and gyri mentioned in Table 26–1.

 

B

Cavum anterius

 

 

Cavum

A

Corpus

posterius

callosum

 

 

 

C

100%

50%

20%

6

Birth

months

Figure 26–7. The cavum septi pellucidi is a closed, median cleft between the two laminae of the septum pellucidum. It may perhaps develop by necrosis within the commissural plate. It is constant up to about

250 mm, its posterior portion (cavum Vergae) is usually closed by birth, and its anterior portion by about 3 postnatal months. It remains open subsequently in about 20%. (A) A coronal section at about the middle of prenatal life. (B) A median section early in the third trimester. Three more or less parallel features are (1) the pericallosal continuation of the anterior cerebral arteries (not shown here), (2) the corpus callosum, and (3) the cavum septi pellucidi. (C) A graph of the percentage of presence (ordinate) plotted against 0–7 postnatal months (abscissa) (Shaw and Alvord, 1969).

Figure 26–8. Superior view of the brain around the time of birth. Acknowledgement with Figure 26–9.

A

C

B

D

Figure 26–9. The brain around the time of birth. (A) and (B) Lateral and medial photographs. Courtesy of Marvin D. Nelson, M.D., and Floyd H. Gilles, M.D., Children’s Hospital, Los Angeles. (C) and (D) Lateral and medial views at 335 mm GL, 490 mm CH, biparietal diameter 94 mm, occipitofrontal diameter 108 mm, head circumference 355 mm, body weight 3,340 g, and fresh brain weight 403 g. The pattern of sulci and gyri is essentially similar to that of the adult. The names given here apply to the gyri. Arrows indicate the central and parieto-occipital sulci. Abbreviations: Fl, 2, 3, superior, middle, and inferior frontal gyri, Tl, 2, 3, superior, middle, and inferior temporal gyri; a., angular gyrus; s., supramarginal gyrus. Based on photographs in Feess-Higgins and Larroche (1987).

TRIMESTER 3 AND THE NEWBORN

313

Figure 26–10. (A) and (B) The arterial system of the newborn. The vessels

A marked are the three cerebral arteries (A.C., M.C., P.C.), the posterior communicating (P. co.), the anterior choroidal (A. chor.), the superior cerebellar (Sup. cbl), and the anterior and posterior inferior cerebellar arteries (AICA, PICA). The asterisk indicates the hypophysis. The posterior communicating is relatively large prenatally and in the newborn. In the adult, however, the blood flow in the posterior cerebral (initially merely a collateral branch of the posterior communicating) has been transferred from the carotid to the basilar system. These drawings are based on graphic reconstructions made by Padget (1948), whose work should be studied for further details. The venous system has been illustrated by Padget (1957, Fig. 17).

B

Anterior & middle cerebral

Internal carotid

Posterior communicating

Posterior cerebral

Basilar

Vertebral

314

C h a p t e r 2 6 : TRIMESTER 3 AND THE NEWBORN

A

B

C D

Figure 26–11. Examples of images during trimester 2. (A) and (B) Horizontal sections at 23 postmenstrual weeks. (C) and (D) Views of the maternal pelvis showing the fetal brain at 26 postmenstrual weeks. All four views present some pathological features. Courtesy of Robert Harris, M.D., Dartmouth Hitchcock Medical Center, Hanover, NH.

PRENATAL LIFE

315

 

 

TABLE 26–2. Examples of Events, Features, and Substances in the Prenatal Brain

 

Substances and/or Events

Reference

 

 

A. Embryonic Period

 

Nasal plate contains TH-ir neuronsa

Verney et al. (1996)

GnRH-ir cells migrating to forebrain

Verney et al. (1996),

 

Schwanzel-Fukuda et al. (1996)

Calbindin-ir postmitotic cell bodies in mesencephalic floor plate

Verney et al. (2001a,b)

Reelin-positive cells (subpial neurons) in neocortical anlage form a continuous horizontal

Meyer et al. (2000)b

row at telencephalic surface

 

GABA dispersed in different layers of the ventricular eminence(s)

Verney (2003)

Calretinin-ir neurons in mesencephalic floor plate

Verney et al. (2001a,b)

Calbindin-ir axons in thalamocortical bundle of internal capsule

 

Enzymes of the catecholamine pathwaya

Zecevic and Verney (1995)

Preplate (primordial plexiform layer) with reelin-ir cells

Meyer et al. (2000)

Calretinin-positive cells continuous with CR-ir cells in the ventricular zone

 

GABA-positive cells and fibers in primordial plexiform layer

Zecevic and Milosevic (1997)

GABA neurons may establish contacts with other terminals or cells

 

Compartmentalization of the preplate: subpial monolayer of horizontal CR-ir cells

Meyer et al. (2000)

Reelin-positive subpial cells

 

Calretinin pioneering cells with first corticofugal fibers

Meyer et al. (2000)

Cortical plate

Muller¨ and O’Rahilly (1990b)

Retrobulbar reelin-ir cells migrating to “marginal zone” of cortex

Meyer et al. (2000)

GABA-ir cells in zona limitans intrathalamica and ventral thalamus

Kultas-Ilinsky et al. (2004)

B. Trimester 1, Postembryonic Phase

 

VMAT2 (vesicular monoamine transporter)

Verney et al. (2002)

SERT serotonin transporter

 

GABA (γ -amino butyric acid)-immunoreactivity in the telencephalic wall

Zecevic and Milosevic (1997)

Initial organization of cortical development

 

Neurofilament protein-labelled fibers, calretinin-ir, calbindin-ir neurons (in ascending

Zecevic et al. (1999)

caudorostral gradient)

 

Cells of motor thalamic nuclei express calcium-binding protein, fibers coexpress GABA

Kultas-Ilinsky et al. (2004)

TH-ir axons in nucleus accumbens and amygdaloid complex coexpress the calbindin

Verney et al. (2001b)

D28K phenotype

 

NPH (carrier molecule of vasopressin and oxytocin) in cells arising from hypothalamic

Mai et al. (1997)

sulcus (potentially functional hypothalamohypophysial system)

 

D1R (dopamine receptor 1)-ir in striosomal bodies and neuropil

Brana et al. (1996)

GAD67-positive local circuit neurons in the thalamic nuclei (outside afferent receiving

Kultas-Ilinsky et al. (2004)

territory of basal nuclei)

 

Apoptosis-regulatory Bcl-2 oncoprotein in hippocampus and brainstem

Chan and Yew (1998)

Synaptophysin indicates synaptogenesis in corpus striatum

Ulfig et al. (2000)

 

 

 

(Continued )

316

C h a p t e r 2 6 : PRENATAL LIFE

 

 

 

 

TABLE 26–2. (Continued )

 

 

Substances and/or Events

 

Reference

 

 

 

C. Trimester 2

 

 

SERT-ir fibers in the internal capsule

 

Verney et al. (2002)

4 neuropeptides produced by several neuron-populations are present in the corpus

Brana et al. (1995)

striatum: SRIF (somatostatin), ENK (enkephalin), SP (substance P), DYN (dynorphin)

 

Monocarboxylate transporters MCT1 and MCT2 in the visual cortex

Fayol et al. (2004)

Immunoreactivity of neural cell adhesion molecule L1 in parallel fibers of the molecular

Tsuru et al. (1996)

layer and in the Purkinje cell layer of cerebellum

 

Slit-2 required for guiding both preand post-crossing callosal axons (repels the growth

Shu and Richards (2001)

cones away from the median plane)

 

 

Subpial granular layer expresses calretinin and reelin

Meyer and Wahle (1999b)

NPY (neuropeptide Y)-ir neurons in subplate

 

Delalle et al. (1997)

Diffuse and cellular AKAP (a kinase anchoring protein) 79-immunity in striatum

Ulfig et al. (2001a,b)

Migration of ventricular cells into cortex coincides with maximum density of

Meyer et al. (1999)

reelin-producing cells of subpial granular layer

 

 

Synaptotagmin-ir fibers in subplate and cortical plate

Ulfig et al. (2002)

Vimentin-positive radial glia in thalamus and corpus striatum

Ulfig et al. (1999)

Calretinin and SMI in ventral parts of dorsal thalamus

Kultas-Ilinsky et al. (2004)

Edg-2 involved in myelination

 

Briese and Ulfig (2003)

Synaptoporin-ir fibers in subplate and cortical plate

Ulfig et al. (2002)

Synaptogenesis in auditory and prefrontal cortices

Huttenlocher and Dabholkar (1997)

 

 

 

a See also Table 17–3 for development of dopaminergic neurons.

b Neocortical formation using immunohistochemistry for reelin, calretinin and glutamic acid decarboxylase.

PRENATAL LIFE

317

Figure 26–12. Graph showing fresh brain weight plotted against estimated weeks after fertilization. This is a Gompertz prediction with a 95% confidence band. After McLennan, Gilles, and Neff in Gilles, Leviton, and Dooling (1983).

Figure 26–13. The diameter of the fixed fetal brain (n = 156) modified from Dunn (1921). The measurements are from the frontal to the occipital pole, and from the right to the left temporal pole.