Rhombencephalon

By the time the midbrain flexure appears, the length of the rhombencephalon is greater than
that of the combined extent of the mesencephalon and prosencephalon. Rostrally, it exhibits
a constriction, the isthmus rhombencephali (see Fig. 17.3B), best viewed from the dorsal
aspect. Ventrally, the hindbrain is separated from the dorsal wall of the primitive pharynx
only by the notochord, the two dorsal aortae and a small amount of mesenchyme; on each
side, it is closely related to the dorsal ends of the pharyngeal arches.
The pontine flexure appears to ‘stretch’ the thin, epithelial roof plate, which becomes
widened. The greatest increase in width corresponds to the region of maximum convexity, so
that the outline of the roof plate becomes rhomboidal. By the same change, the lateral walls
become separated, particularly dorsally, and the cavity of the hindbrain, subsequently the
fourth ventricle, becomes flattened and somewhat triangular in cross-section. The pontine
flexure becomes increasingly acute until, at the end of the second month, the laminae of its
cranial (metencephalic) and caudal (myelencephalic) slopes are opposed to each other (see
Fig. 17.21); at the same time, the lateral angles of the cavity extend to form the lateral
recesses of the fourth ventricle.
At about 4 ½ weeks of development, when the pontine flexure is first discernible, the
association between the rhombomeres and the underlying motor nuclei of certain cranial
nerves can be seen. The general pattern of distribution of motor nuclei is as follows:
rhombomere 1 contains the trochlear nucleus, rhombomeres 2 and 3 contain the trigeminal
nucleus, rhombomeres 4 and 5 contain the facial nucleus, rhombomere 5 contains the
abducens nucleus, rhombomeres 6 and 7 contain the glossopharyngeal nucleus, and
rhombomeres 7 and 8 contain the vagal, accessory and hypoglossal nuclei. Rhombomeric
segmentation represents the ground plan of development in this region of the brainstem and
is pivotal for the development of regional identity (see Fig. 12.4). However, with further
morphogenesis, the obvious constrictions of the rhombomere boundaries disappear, and the
medulla once again assumes a smooth contour. The differentiation of the lateral walls of the
hindbrain into basal (ventrolateral) and alar (dorsolateral) plates has a similar significance to
the corresponding differentiation in the lateral wall of the spinal cord, and ventricular,
intermediate and marginal zones are formed in the same way.

Cells of the basal plate (ventrolateral lamina)

Cells of the basal plate form three elongated, discontinuous, columns that are positioned
ventrally and dorsally with an intermediate column between (Fig. 17.20).
The most ventral column is continuous with the anterior grey column of the spinal
cord and will supply muscles considered ‘myotomic’ in origin. It is represented in the caudal
part of the hindbrain by the hypoglossal nucleus, and it reappears at a higher level as the
nuclei of the abducens, trochlear and oculomotor nerves (somatic efferent nuclei). The
intermediate column is represented in the upper part of the spinal cord and caudal brainstem
(medulla oblongata and pons), and its neurones supply branchial (pharyngeal) and
postbranchial musculature. It is discontinuous, forming the elongated nucleus ambiguous in
the caudal brainstem, which gives fibres to the ninth, tenth and eleventh cranial nerves, and
continues into the cervical spinal cord as the origin of the accessory nerve. At higher levels,
parts of this column give origin to the motor nuclei of the facial and trigeminal nerves. The
nucleus ambiguus and the facial and trigeminal motor nuclei are termed branchial (special
visceral) efferent nuclei. Neurones in the most dorsal column of the basal plate (represented
in the spinal cord by the lateral grey column) innervate viscera. The column is discontinuous;
its large caudal part forms some of the dorsal nucleus of the vagus and its cranial part forms
the salivatory nucleus. These nuclei are termed general visceral (general splanchnic) efferent
nuclei, and their neurons give rise to preganglionic, parasympathetic nerve fibres.
It is important to note that the neurones of the basal plate and their three columnar
derivatives are only motor in the sense that some of their number form either motor
neurones or preganglionic parasympathetic neurones. The remainder, which greatly
outnumber the former, differentiate into functionally related interneurones and, in some loci,
into neuroendocrine cells.

Cell columns of the alar plate (dorsolateral lamina)

Cell columns of the alar plate are discontinuous and give rise to general visceral (general
splanchnic) afferent, special visceral (special splanchnic) afferent, general somatic afferent,
and special somatic afferent nuclei (their relative positions, in simplified transverse section,
are shown in Fig. 17.20). The general visceral afferent column is represented by a part of the
dorsal nucleus of the vagus, the special visceral afferent column by the nucleus of the tractus
solitarius, the general somatic afferent column by the afferent nuclei of the trigeminal nerve,
and the special somatic afferent column by the nuclei of the vestibulocochlear nerve. (The
relatively simple functional independence of these afferent columns implied by the foregoing
classification is, in the main, an aid to elementary learning. The emergent neurobiological
mechanisms are, in fact, much more complex and less well understood.) Although they tend
to retain their primitive positions, some of these nuclei are later displaced by differential
growth patterns, by the appearance and growth of neighbouring fibre tracts, and possibly by
active migration.
It has been suggested that a neurone tends to remain as near as possible to its
predominant source of stimulation, and that, to achieve this aim, it will migrate around
intervening structures, towards the greatest density of stimuli. The curious paths of the
axons arising from the facial nucleus and the nucleus ambiguus have been regarded as
exemplars of this phenomenon of neurobiotaxis. In a 10 mm embryo, the facial nucleus lies
in the floor of the fourth ventricle, occupying the position of the special visceral efferent
column, and it is placed at a higher level than the abducens nucleus. As growth proceeds,
the facial nucleus migrates at first caudally and dorsally, relative to the abducens nucleus,
and then ventrally to reach its adult position. As it migrates, the axons to which its somata
give rise elongate and their subsequent course is assumed to map out the pathway along
which the facial nucleus has travelled. Similarly, the nucleus ambiguus initially arises
immediately deep to the ventricular floor but, in the adult, it is more deeply placed and its
efferent fibres pass first dorsally and medially before curving laterally to emerge at the
surface of the medulla oblongata