Ontogeny of the anuran urostyle and the developmental context of evolutionary novelty.

Developmental novelties often underlie the evolutionary origins of key metazoan features. The anuran urostyle, which evolved nearly 200 MYA, is one such structure. It forms as the tail regresses during metamorphosis, when locomotion changes from an axial-driven mode in larvae to a limb-driven one in adult frogs. The urostyle comprises of a coccyx and a hypochord. The coccyx forms by fusion of caudal vertebrae and has evolved repeatedly across vertebrates. However, the contribution of an ossifying hypochord to the coccyx in anurans is unique among vertebrates and remains a developmental enigma. Here, we focus on the developmental changes that lead to the anuran urostyle, with an emphasis on understanding the ossifying hypochord. We find that the coccyx and hypochord have two different developmental histories: First, the development of the coccyx initiates before metamorphic climax whereas the ossifying hypochord undergoes rapid ossification and hypertrophy; second, thyroid hormone directly affects hypochord formation and appears to have a secondary effect on the coccygeal portion of the urostyle. The embryonic hypochord is known to play a significant role in the positioning of the dorsal aorta (DA), but the reason for hypochordal ossification remains obscure. Our results suggest that the ossifying hypochord plays a role in remodeling the DA in the newly forming adult body by partially occluding the DA in the tail. We propose that the ossifying hypochord-induced loss of the tail during metamorphosis has enabled the evolution of the unique anuran bauplan.

Enigmatic human tails: A review of their history, embryology, classification, and clinical manifestations.

The presence of a human tail is a rare and intriguing phenomenon. While cases have been reported in the literature, confusion remains with respect to the proper classification, definition, and treatment methods. We review the literature concerning this anatomical derailment. We also consider the importance of excluding underlying congenital anomalies in these patients to prevent neurological deficits and other abnormal manifestations.

Human fetal anatomy of the coccygeal attachments of the levator ani muscle.

In contrast to the attachments to the pubis and rectum, there is little information on fetal development of the coccygeal attachment of the levator ani muscles. We find that at 9 weeks, the coccygeus muscle is a large muscle facing the piriformis or gluteus maximus and inserting onto the ischial spine, whereas the levator ani is restricted to the area near the pubis. By 12 weeks, the levator ani also obtains attachment to the ischial spine immediately ventral to the coccygeus muscle. The most superior part of the coccygeus muscle occupies a space at an angle between the pelvic splanchnic and pudendal nerves. Notably, medial to the coccygeus muscle, a third parasagittal muscle (previously termed the sacrococcygeus anterior) appears by 12 weeks, increases in mass by 18 weeks, and connects and mixes with the dorsal end of the levator ani by 18-20 weeks. Thus, the coccygeal attachment of the levator ani appears not to depend on the dorsal extension of the muscle itself but on fusion with the sacrococcygeus anterior. Therefore, the final levator sheet is formed medial (internal) to the coccygeus muscle and originates from two distinct anlage.

Caudal agenesis and associated caudal spinal cord malformations.

Caudal agenesis is a rare congenital anomaly resulting from an insult to the structures of the caudal eminence. It may be associated with anomalies of other structures derived from the caudal eminence: the hindgut and the urogenital system. Patients are more likely to present first to the pediatric surgeon (for evaluation of gastrointestinal anomalies), the urologist (for urogenital malformation or dysfunction), or the orthopedic surgeon (for lower extremity abnormalities), than to the neurosurgeon. Characteristic external features of the buttocks, hips, and lower extremities may suggest the diagnosis. MR imaging is the diagnostic modality of choice and should be used in all patients with suggestive external features or other caudal anomalies. The level of bone anomaly corresponds well to the level of weakness but not sensory loss. Sensation is usually relatively preserved. The caudal spinal cord is often truncated in cases of high bone lesions and tethered, with occasional association with a dysraphic lesion, in cases of low bone lesions. Early neurosurgical intervention is preferred in all cases of recognized occult spinal dysraphism. Progressive neurologic deficits may develop later in life in patients with unrecognized tethered cord or dural stenosis and require neurosurgical repair on diagnosis. A better understanding of the embryology of the caudal region and investigation of the teratogens that may interfere with this stage of development should lead to more effective treatment and prevention of caudal agenesis and the associated caudal anomalies.

Sacrococcygeal dysgenesis association.

In the malformation analysis of 445 patients ascertained only for a sacrococcygeal malformation, a new phenotype, the sacrococcygeal dysgenesis association (SDA), was delineated in 34%. In addition, sirenomelia patients were found in 12%, the VATER association in 27%, and 27% could not be classified. Heterogeneity in the patients with sacrococcygeal malformations was identified by the differences found in their associated malformations. SDA patients have a relatively small average number (3.3) of anomalies per patient as compared with 9.3 in sirenomelia and 6.2 in VATER patients. SDA abnormalities occurred to a significant degree only in 6 of 20 designated malformation categories (vertebral, rib, pelvic, lower limb, central nervous system [CNS], renal) in contrast to 17 in VATER and 18 in sirenomelia patients. The SDA vertebral malformation pattern also differed from that of VATER/sirenomelia patients as did the high sacrococcygeal agenesis:dysgenesis ratio and low thoracolumbar vertebrae and/or rib hypersegmentations. Most significantly, SDA patients had a large number of CNS anomalies and CNS-related dysfunctions of the urinary and distal intestinal tracts but no anatomic urinary or intestinal tract malformations. This contrasted sharply with the markedly increased occurrences of anatomic abnormalities in these body regions of the sirenomelia and VATER patients. Demographic data such as patient survival, twinning and, particularly, the high (28%) incidence of maternal diabetes in the SDA further support its differentiation from VATER/sirenomelia patients.

The human tail and spinal dysraphism.

Recent publications have endeavoured to differentiate between the true, or vestigial tail, and the pseudotail by clinical and pathological examination, and have indicated the benign nature of the true tail. The true tail arises from the most distal remnant of the embryonic tail, contains adipose, connective, muscle, and nerve tissue, and is covered by skin. Pseudotails represent a variety of lesions having in common a lumbosacral protrusion and a superficial resemblance to vestigial tails. A review of the case reports indicates spina bifida to be the most frequent coexisting anomaly with both. A review of occult spinal dysraphism shows it to be associated with cutaneous signs in more than 50% of instances. Three cases of spinal dysraphism with tail-like cutaneous structures are described and their radiological, operative, and pathological findings presented. The classification of each of the appendages into true tail or pseudotail remains obscure. Although the finding of these three tails was the subject of much curiosity, surgical treatment was clearly designed to adequately deal with the associated dysraphic state. The presence of a tail-like appendage in the lumbosacral region should alert the clinician to the possibility of underlying spinal dysraphism. Preoperative assessment must include a complete neurological history and examination as well as computed tomographic or magnetic resonance imaging.

Lower axial skeleton of the musk shrew (Suncus murinus).

Macroscopic structure as well as pre- and postnatal development of the lumbar, sacral, and caudal vertebrae of the musk shrew (Suncus murinus, Insectivora) were observed. The lumbar vertebrae possess two pairs of unusual processes, hyperapophyses and hypapophyses. The hyperapophyses are located on the dorsal surface of the caudal articular processes of all the lumbar vertebrae, whereas the hypapophyses are found on the caudal part of the ventral surface of the bodies in the first few lumbar vertebrae. The former gives attachment to the Mm. rotatores lumborum and the latter to the Mm. psoas major and minor. The articular processes of the lumbar vertebrae are oriented more horizontally compared with those in other mammals. The sacrum is very narrow transversely due to poor development of the ventrolateral wing. The auricular surface includes cranial parts of the wing and of the fused vertebral arches as well as the cranial articular process of the first sacral vertebra. In the caudal vertebrae, chevron bones are H-shaped when viewed ventrally, and give attachment to tendons of the caudal muscles. This report describes the relationships between the structural peculiarities of the lower axial skeleton and the locomotive habits of the musk shrew.

The human vertebral column at the end of the embryonic period proper. 4. The sacrococcygeal region.

The sacral and coccygeal vertebrae at 8 postovulatory weeks (the end of the embryonic period proper) have been studied by means of graphic reconstructions. The cartilaginous sacrum is now a definitive unit composed of five separable vertebrae, each of which consists of a future centrum and bilateral neural processes. The base of each neural process consists of an anterolateral or alar element, not present in the lumbar region, and a posterolateral part, which includes costal and transverse elements. The usual illustrations, in which the costal component is placed in the alar element, are incorrect. The future dorsal foramina (containing dorsal rami) face laterally in the embryo and are in line with the thoracicolumbar intervertebral foramina. Considerable differential growth is required to change the dorsal openings from a lateral to a dorsal positions. The intervertebral foramina transmit ventral rami, but pelvic foramina are not yet present. The lumbosacral plexus is completed by S.N.1-3; S.N.4, 5 and Co.N.1 form the pelvic plexus. The inferior hypogastric plexus and the hypogastric nerves are present. The sacrum takes part in the spina bifida occulta that characterises the entire length of the embryonic vertebral column. The coccygeal vertebrae, which are variable, were 4-6 in number in the present series. The first is the best developed. The ventriculus terminalis ends usually at the level of Co.V.1 and the spinal cord generally at Co.V.5. The coccygeal notochord ends commonly in bifurcation or trifurcation. 'Haemal arches' were not observed.

Human tails.

Two children and one infant with a "human tail" are presented. The patho-embryology of this medical curiosity is briefly discussed. Treatment is usually unnecessary but resection of part of the coccyx together with the "tail" may become indicated by coccygodynia or for aesthetic reasons.

A study of fetal growth retardation in teratological tests: an examination of the relationship between body weight and ossification of coccygeal vertebrae in mouse and rat fetuses.

We examined the relationship between body weight and ossification of coccygeal vertebrae in normal full-term fetuses of mice and rats to reliably evaluate fetal growth retardation in teratological tests. Correlation coefficients between body weight and number of ossified coccygeal vertebrae were positive in mice and rats. However, the coefficients were variable between groups and were not statistically significant in some groups. Averages of body weight and ossified coccygeal vertebrae were variable in the normal fetuses. However, the coefficient of variation in each group was nearly constant, and the component of variance between groups (sigma 1(2)) was small as compared with that within groups (sigma 0(2)). Therefore, the relationship between low body weight and retardation of ossification observed in teratological experiments may be evaluated by measuring relative differences of body weight and ossification between the treated group and the control group with the variance within groups (sigma 0) as a scale.