The staging of human embryos, as distinct from seriation, depends on a morphological scheme devised by Streeter and completed by O’Rahilly, who proposed the term Carnegie stages. To avoid misconceptions and errors, and to place new findings in perspective, it is necessary to summarize the essentials of the Carnegie system: (1) Twenty-three stages cover the embryonic period, i. e. the first 8 postfertilizational weeks of development. (2) The system is based on internal as well as external features, and the use of only external criteria is subject to serious limitations. For example, precise delineation of stages 19–23 and of the embryonic-fetal transition depends on histological examination. (3) Prenatal measurements are not an integral component of the staging system, and hence a stage should never be assigned merely on the basis of embryonic length. A 20-mm embryo, for example, could belong to any of three stages. Measurements, however, are important for the assessment of age, and very few measurements are available for staged embryos. Presented here and based on accurate staging are the maximum diameter of the chorionic sac, the crown-heel length, the greatest length exclusive of the lower limbs, the biparietal diameter, the head circumference, the length of the hindbrain, the total length of the brain, and the lengths of the limbs as well as of their segments, including the foot length. (4) Prenatal ages are also not an integral part of the staging system and hence a stage should never be assigned merely on the basis of prenatal age. Ages, however, are of clinical importance and their estimate has been rendered more precise by accurate timing of fertilization followed by ultrasonography. Prenatal age is postfertilizational and hence some 2 weeks less than the postmenstrual interval. The term gestational age is ambiguous and should be discarded. Presented here is a new graph showing proposed estimates of age in relation to stages and based on current information.
The first systematic account of the neural crest in the human has been prepared after an investigation of 185 serially sectioned staged embryos, aided by graphic reconstructions. As many as fourteen named topographical subdivisions of the crest were identified and eight of them give origin to ganglia (Table 2). Significant findings in the human include the following. (1) An indication of mesencephalic neural crest is discernible already at stage 9, and trigeminal, facial, and postotic components can be detected at stage 10. (2) Crest was not observed at the level of diencephalon 2. Although pre-otic crest from the neural folds is at first continuous (stage 10), crest-free zones are soon observable (stage 11) in Rh.1, 3, and 5. (3) Emigration of cranial neural crest from the neural folds at the neurosomatic junction begins before closure of the rostral neuropore, and later crest cells do not accumulate above the neural tube. (4) The trigeminal, facial, glossopharyngeal and vagal ganglia, which develop from crest that emigrates before the neural folds have fused, continue to receive contributions from the roof plate of the neural tube after fusion of the folds. (5) The nasal crest and the terminalis-vomeronasal complex are the last components of the cranial crest to appear (at stage 13) and they persist longer. (6) The optic, mesencephalic, isthmic, accessory, and hypoglossal crest do not form ganglia. Cervical ganglion 1 is separated early from the neural crest and is not a Froriep ganglion. (7) The cranial ganglia derived from neural crest show a specific relationship to individual neuromeres, and rhombomeres are better landmarks than the otic primordium, which descends during stages 9-14. (8) Epipharyngeal placodes of the pharyngeal arches contribute to cranial ganglia, although that of arch 1 is not typical. (9) The neural crest from rhombomeres 6 and 7 that migrates to pharyngeal arch 3 and from there rostrad to the truncus arteriosus at stage 12 is identified here, for the first time in the human, as the cardiac crest. (10) The hypoglossal crest provides cells that accompany those of myotomes 1-4 and form the hypoglossal cell cord at stages 13 and 14. (11) The occipital crest, which is related to somites 1-4 in the human, differs from the spinal mainly in that it does not develop ganglia. (12) The occipital and spinal portions of the crest migrate dorsoventrad and appear to traverse the sclerotomes before the differentiation into loose and dense zones in the latter. (13) Embryonic examples of synophthalmia and anencephaly are cited to emphasize the role of the neural crest in the development of cranial ganglia and the skull.
Serial sections of 105 human embryos (including 20 silver preparations) from stage 11 (24 days) to stage 22 (54 days) were studied, and 23 graphic reconstructions were prepared. The hypoglossal nucleus is evident at stage 12 and becomes isolated from other efferent nuclei at stage 14. The first hypoglossal nerve fibers appear at stage 12. The roots unite at stage 14 and the main trunk arrives in the tongue at stage 15. Four occipital somites can be identified during stage 13, and the sclerotomic material forms two bilateral masses. The fourth sclerotome separates in stage 14 and develops like a vertebra. This and the remaining sclerotomic material form the basioccipital and exoccipital parts of the chondrocranium , which are the first to appear. Four occipital myotomes develop and grow towards the tongue as the "hypoglossal cord", which arrives prior to the hypoglossal nerve. The developmental similarity in the hypoglossal region between birds and mammals, combined with experimental studies in birds, renders it extremely likely that the hypoglossal musculature in mammals also is derived from occipital somites. The present study is the first in which this conclusion is adequately supported in the human. This investigation aids in the interpretation and timing of origin of variations (e.g., bipartite hypoglossal canal) and anomalies (e.g., persistent hypoglossal artery).
Two sites of fusion (a term preferred to closure) of the neural folds and two neuropores are found in the human embryo. No convincing embryological evidence of a pattern of multiple sites of fusion, such as has been described in the mouse, is available for the human. The construction of embryological details from information derived from other species or from the examination of later anomalies is liable to error. Neural tube defects are reviewed and although they have been considered on the basis of five, four, or three sites of fusion, interpretations based on two sites can as readily be envisaged.
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