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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.


100 cm

50

C H A P T E R 26

SUPPLEMENT: EARLY

POSTNATAL LIFE

Years

5

10

A

B

Figure 26–1. (A) A boy at 14 postnatal days, showing the myelin in the thalamic region, brain stem, and cerebellum. (B) A girl at 20 postnatal days. The hypophysis, optic chiasma and tracts, and mamillary bodies (superimposed) are clearly visible, as are the corpus callosum, epiphysis, tectum, tentorium cerebelli, and fourth ventricle.

The Embryonic Human Brain: An Atlas of Developmental Stages, Third Edition. By O’Rahilly and Muller¨

Copyright C 2006 John Wiley & Sons, Inc.

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320

C

C h a p t e r 2 6 : EARLY POSTNATAL LIFE

Figure 26–14. (Continued ) (C) At 3 years, most of the adult features are already clearly visible. Structures seen more readily in this image include the septum pellucidum, the aqueduct, and the spheno-occipital joint. In the cerebellum the fissura prima, the nodule, and the tonsil are distinguishable. Many of the white areas represent either fat or bone marrow, whereas solid bone does not produce signals.

Postnatal Prolongation of Fetal growth

Human development shows a general retardation relative to other primates, resulting in retention of juvenile features (neoteny). The human infant is relatively undeveloped and helpless at birth, and the completion of growth and maturation is postponed. It has been proposed (particularly by Adolf Portmann) that because human birth is

accelerated, growth rates during the first postnatal year follow the fetal trends shown by other primates. Indeed it has been estimated that were birth to be delayed in proportion to retarded development in general, then existence in utero would be increased to 21 months. The maintenance of fetal growth rates postnatally allows the brain to increase considerably in mass. By plotting brain weight versus body weight it has been shown that the high fetal slope of the graph is, in the human, continued well into postnatal life.


EARLY POSTNATAL LIFE

321

D E

Figure 26–14. (Continued ) (D) A coronal view at 2 postnatal months, showing myelination bilaterally in the white matter of the cerebellum.

(E) At 4 years, showing the considerable advance in myelination, e.g., in the corona radiata, corpus callosum, fornices, thalami, colliculi, arbor vitae of the cerebellum, and cerebellar peduncles. The lateral, third, and fourth ventricles are distinguishable, as are also the tentorium cerebelli and the cerebellar folia.

A–E, courtesy of Marvin D. Nelson, M.D., Children’s Hospital, Los Angeles.

Myelin(iz)ation

Myelination in the central nervous system begins after its onset in the peripheral system in trimester 2. It continues for a number of years after birth at a gradually decreasing rate. In the CNS, myelination is undertaken by oligodendrocytes and is very slow, and one axon can be enveloped in several sheaths. The determination of the onset and end of myelination depends on whether light or electron microscopy is used, and the appearances on magnetic resonance images lag about a month behind histological data.

Myelination expresses the functional maturity of the brain and is correlated with psychomotor development, although transmission of impulses and functional activity begin before myelin sheaths develop. Large afferent tracts become myelinated early, and tracts that appear early in development generally undergo early myelination, e.g., the medial longitudinal fasciculus. Although myelination is found in the pyramidal decussation by the middle of

prenatal life (Wozniak´ and O’Rahilly, 1982), in the pyramidal tracts it begins late in trimester 3 and is not completed until about two years. Cortical association fibers are among the last to become myelinated.

At birth the human brain is only moderately myelinated (Fig. 26–14A,B) and the cerebral hemispheres contain little myelin. Myelin can be shown histologically in the brain stem, the cerebellar white matter, and the posterior limb of the internal capsule, with extensions to the thalamus and the basal nuclei. Myelination is greatest during the first two postnatal years and is practically complete in early adulthood, although it continues throughout life. Tables showing the progress of myelination are available (e.g., Larroche, 1966; Gilles et al., 1983).

A few examples of magnetic resonance imaging serve to illustrate the increasing degree of myelination. Postnatal myelination in the central nervous system has been studied extensively by Brody and Kinney and their colleagues (Brody et al., 1987; Kinney et al., 1988).


BIBLIOGRAPHY

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