Development and maintenance of tendons and ligaments.

Tendons and ligaments are fibrous connective tissues vital to the transmission of force and stabilization of the musculoskeletal system. Arising in precise regions of the embryo, tendons and ligaments share many properties and little is known about the molecular differences that differentiate them. Recent studies have revealed heterogeneity and plasticity within tendon and ligament cells, raising questions regarding the developmental mechanisms regulating tendon and ligament identity. Here, we discuss recent findings that contribute to our understanding of the mechanisms that establish and maintain tendon progenitors and their differentiated progeny in the head, trunk and limb. We also review the extent to which these findings are specific to certain anatomical regions and model organisms, and indicate which findings similarly apply to ligaments. Finally, we address current research regarding the cellular lineages that contribute to tendon and ligament repair, and to what extent their regulation is conserved within tendon and ligament development.

Bone morphology is regulated modularly by global and regional genetic programs.

Bone protrusions provide stable anchoring sites for ligaments and tendons and define the unique morphology of each long bone. Despite their importance, the mechanism by which superstructures are patterned is unknown. Here, we identify components of the genetic program that control the patterning of Sox9+/Scx+ superstructure progenitors in mouse and show that this program includes both global and regional regulatory modules. Using light-sheet fluorescence microscopy combined with genetic lineage labeling, we mapped the broad contribution of the Sox9+/Scx+ progenitors to the formation of bone superstructures. Then, by combining literature-based evidence, comparative transcriptomic analysis and genetic mouse models, we identified Gli3 as a global regulator of superstructure patterning, whereas Pbx1, Pbx2, Hoxa11 and Hoxd11 act as proximal and distal regulators, respectively. Moreover, by demonstrating a dose-dependent pattern regulation in Gli3 and Pbx1 compound mutations, we show that the global and regional regulatory modules work in a coordinated manner. Collectively, our results provide strong evidence for genetic regulation of superstructure patterning, which further supports the notion that long bone development is a modular process.This article has an associated 'The people behind the papers' interview.

Histological development and integration of the Zebrafish Weberian apparatus.

BACKGROUND: The Weberian apparatus enhances hearing in otophysan fishes, including Zebrafish (Danio rerio). Several studies have examined aspects of morphological development of the Weberian apparatus and hearing ability in Zebrafish. A comprehensive developmental description including both hard and soft tissues is lacking. This information is critical for both interpretation of genetic developmental analyses and to better understand the role of morphogenesis and integration on changes in hearing ability. RESULTS: Histological development of hard and soft tissues of the Weberian apparatus, including ossicles, ear, swim bladder, and ligaments are described from early larval stages (3.8 mm notochord length) through adult. Results show a strong relationship in developmental timing and maturation across all regions. All required auditory elements are present and morphologically integrated early, by 6.5 mm SL. Dynamic ossification patterns and changes in shape continue throughout the examined developmental period. CONCLUSIONS: This study provides the first comprehensive histological description of Weberian apparatus development in Zebrafish. Morphological integration was found early, before increases in hearing ability were detected in functional studies (>10 mm total length), suggesting morphological integration precedes functional integration. Further research is needed to examine the nature of the functional delay, and how maturation of the Weberian apparatus influences functionality.

Early development of the vertebral column.

The segmental organization of the vertebrate body is most obviously visible in the vertebral column, which consists of a series of vertebral bones and interconnecting joints and ligaments. During embryogenesis, the vertebral column derives from the somites, which are the primary segments of the embryonic paraxial mesoderm. Anatomical, cellular and molecular aspects of vertebral column development have been of interest to developmental biologists for more than 150 years. This review briefly summarizes the present knowledge on early steps of vertebral column development in amniotes, starting from sclerotome formation and leading to the establishment of the anatomical bauplan of the spine composed of vertebral bodies, vertebral arches, intervertebral discs and ribs, and their specific axial identities along the body axis.

The development of zebrafish tendon and ligament progenitors.

Despite the importance of tendons and ligaments for transmitting movement and providing stability to the musculoskeletal system, their development is considerably less well understood than that of the tissues they serve to connect. Zebrafish have been widely used to address questions in muscle and skeletal development, yet few studies describe their tendon and ligament tissues. We have analyzed in zebrafish the expression of several genes known to be enriched in mammalian tendons and ligaments, including scleraxis (scx), collagen 1a2 (col1a2) and tenomodulin (tnmd), or in the tendon-like myosepta of the zebrafish (xirp2a). Co-expression studies with muscle and cartilage markers demonstrate the presence of scxa, col1a2 and tnmd at sites between the developing muscle and cartilage, and xirp2a at the myotendinous junctions. We determined that the zebrafish craniofacial tendon and ligament progenitors are neural crest derived, as in mammals. Cranial and fin tendon progenitors can be induced in the absence of differentiated muscle or cartilage, although neighboring muscle and cartilage are required for tendon cell maintenance and organization, respectively. By contrast, myoseptal scxa expression requires muscle for its initiation. Together, these data suggest a conserved role for muscle in tendon development. Based on the similarities in gene expression, morphology, collagen ultrastructural arrangement and developmental regulation with that of mammalian tendons, we conclude that the zebrafish tendon populations are homologous to their force-transmitting counterparts in higher vertebrates. Within this context, the zebrafish model can be used to provide new avenues for studying tendon biology in a vertebrate genetic system.

Development of the Human Biceps Brachii Tendon and Coracoglenoid Ligament (7th-12th Week of Development).

The goal of this study is to clarify the development of the long head of the biceps brachii tendon (LHBT) and to verify the existence and development of the coracoglenoid ligament. Histological preparations of 22 human embryos (7-8 weeks of development) and 43 human fetuses (9-12 weeks of development) were studied bilaterally using a conventional optical microscope. The articular interzone gives rise to the LHBT, glenoid labrum, and articular capsule. During the fetal period, it was observed that in 50 cases (58%), the LHBT originated from both the glenoid labrum and the scapula, while in 36 cases (42%), it originated only from the glenoid labrum. The coracoglenoid ligament, first described by Sappey in 1867, is a constant structure that originates at the base of the coracoid process and projects toward the glenoid labrum zone, which is related to the origin of the LHBT. The coracoglenoid ligament was more easily identifiable in the 36 cases in which the LHBT originated only from the glenoid labrum. We suggest that the coracoglenoid ligament is a constant anatomical structure, is not derived from the articular interzone unlike the LHBT, and contributes to the fixation of the glenoid labrum in the scapula in cases in which the LHBT originated only from the glenoid labrum. We postulate that, when the LHBT is fixed only at the glenoid labrum, alterations in the coracoglenoid ligament could lead to a less sufficient attachment of the glenoid labrum to the scapula which could predispose to a superior labral lesion.

Role of notochord cells and sclerotome-derived cells in vertebral column development in fugu, Takifugu rubripes: histological and gene expression analyses.

Despite the common structure of vertebrates, the development of the vertebral column differs widely between teleosts and tetrapods in several respects, including the ossification of the centrum and the function of the notochord. In contrast to tetrapods, vertebral development in teleosts is not fully understood, particularly for large fish with highly ossified bones. We therefore examined the histology and gene expression profile of vertebral development in fugu, Takifugu rubripes, a model organism for genomic research. Ossification of the fugu centrum is carried out by outer osteoblasts expressing col1a1, col2a1, and sparc, and the growing centra completely divide the notochord into double cone-shaped segments that function as intercentral joints. In this process, the notochord basal cells produce a thick notochord sheath exhibiting Alcian-blue-reactive cartilaginous properties and composing the intercentral ligament in cooperation with the external ligament connective tissue. Synthesis of the matrix by the basal cells was ascertained by an in vitro test. Expression of twist2 indicates that this connective tissue is descended from the embryonic sclerotome. Notochord basal cells express sox9, ihhb, shh, and col2a1a, suggesting that the signaling system involved in chondrocyte proliferation and matrix production also functions in notochord cells for notochord sheath formation. We further found that the notochord expression of both ntla and shh is maintained in the fugu vertebral column, whereas it is turned off after embryogenesis in zebrafish. Thus, our results demonstrate that, in contrast to zebrafish, a dynamic morphogenesis and molecular network continues to function in fugu until the establishment of the adult vertebral column.

Roles for Nkx2-5 and Gata3 in the ontogeny of the murine smooth muscle gastric ligaments.

The gastric ligaments are superficial cord-like structures, located on the lesser curvature of the stomach, that extend from the pylorus to the esophagus. These ligaments have been documented in a wide variety of mammalian species, including humans, but their composition and ontogeny is unexplored. Here, we demonstrate that, during ontogeny, the gastric ligaments are first visible as extensions from a C-shaped domain of Gata3-expressing cells that surround the future pylorus; this domain will later give rise to the pyloric outer longitudinal muscle (OLM). The open ends of the C are located ventrally, and, beginning at embryonic day (E) 13.5, the ligaments grow anteriorly from these points. Whereas most other ligaments of the stomach are neurovascular in nature, the gastric ligaments are composed of smooth muscle cells that mature between E14.5 and E16.5. The gastric ligaments coexpress the transcription factors Gata3, Nkx2-5, and Sox9, and germline loss of Gata3 or conditional deletion of Nkx2-5 abrogates Sox9 expression and impairs gastric ligament smooth muscle development; similar phenotypes were previously seen in the OLM. In accord with this molecular contiguity between the OLM and gastric ligaments, three-dimensional image reconstruction highlights physical contiguity between these smooth muscle structures, suggesting that they may work together as a unit to control flexure of the pyloric region, a function similar to the ligament of Treitz at the duodenojejunal junction. These findings may have implications for our understanding of normal pyloric sphincter function, as well as the common human congenital pathology idiopathic hypertrophic pyloric stenosis.

Fetal development of ligaments around the tarsal bones with special reference to contribution of muscles.

Through a histological examination of eight mid-term human fetuses (10-15 weeks) and seven late-stage fetuses (30-34 weeks), we attempted to determine how and when fetal ligaments around the tarsal bones form the regular arrangement seen in adults. Ligaments along the dorsal aspect of the tarsal bones developed early as an elongation of the perichondrium, in contrast to the late development of the plantar-sided ligaments. In contrast, a distal elongation of the tibialis posterior tendon was a limited plantar ligament in the early stage; finally, it extended from the navicular, ran obliquely to cross the dorsal side of the fibularis longus tendon, and inserted to the lateral cuneiform and fourth metatarsal. In the late stage, the adductor hallucis muscle origin provided multiple ligamentous structures along the cuneiforms and metatarsals. The tarsal sinus contained multiple fibrous bundles (possibly, the putative interosseous talocalcanean ligaments) that were derived from (1) insertion tendons of the extensor digitorus brevis muscle and (2) the fibrous sheath of the extensor digitorus longus tendon. The aponeurotic origin of the quadratus plantae muscle seemed to contribute to formation of the long plantar ligament. Therefore, tarsal ligaments appeared likely to develop from the long tendons, their fibrous sheaths and aponeuroses and intramuscular tendons of the proper foot muscles. Under in utero conditions with little or no stress from the plantar side of the foot, the muscle-associated connective tissue seems to play a crucial role in providing a regular arrangement of the ligaments in accordance with tensile stress from muscle contraction.

Morphology of the ligament of Treitz likely depends on its fetal topographical relationship with the left adrenal gland and liver caudate lobe as well as the developing lymphatic tissues: a histological study using human fetuses.

To investigate the factors affecting the development of the ligament of Treitz, we examined sagittal and frontal histological sections of 35 human fetuses with a crown-rump length of 100-300 mm (approximately 16-38 weeks of gestation). The retropancreatic fascia consistently extended in a layer behind the pancreatic body and the splenic artery and vein, and also in front of the left renal vein and left adrenal. In 18 specimens, a connective tissue band was seen originating from the diaphragmatic crus around the esophageal opening and ending at the retropancreatic fascia to the left of the origin of the celiac artery. In 10 of these 18 specimens, these putative upper parts of the ligament contained striated muscles, or so-called Hilfsmuskel. Although most of other 17 specimens were larger fetuses, the left adrenal, the liver caudate lobe and the celiac ganglion made space for the ligament very limited. In 22 specimens including the above 18, the retropancreatic fascia extended inferiorly to approach the fourth portion of the duodenum (D4) or the duodenojejunal junction (DJJ). However, in 11 of the 22 examples of the putative lower part of the ligament, the connection between the duodenal muscle coat and the fascia was interrupted by developing lymphatic tissues. Consequently, the ligament of Treitz seemed to develop from both pleuroperitoneal membrane-derived cells and the retropancreatic fusion fascia, although the morphology was markedly modified by adjacent structures such as the adrenal gland. The ligament may "recover" after the adrenal becomes reduced in size after birth.

Structural and functional significance of scrotal ligament: a comparative histological study.

Gubernaculum testes is the most important parameter in testicular migration. At the end of migration, it is described as scrotal ligament, which has implications in testicular torsion. The present study aims to examine the structure of scrotal ligament and compare it with gubernaculum. Sixteen adult cadaveric testicular specimens and fourteen fetal testicular specimens of different age groups were examined after getting ethical clearance from the institute ethics committee and consent from the parents. Meticulous dissection was done. The length, site of proximal, and distal attachment of scrotal ligament and gubernaculum were noted and histologically evaluated. A separate scrotal ligament could not be delineated in any adult specimens. It merged with testicular coverings. Histological examination showed the presence of patchy areas of dense collagen fibres of variable density amidst loose areolar connective tissue. In contrast, fetal specimens showed the presence of a definitive gubernaculum testes and revealed the presence of mesenchymal tissue, collagen, elastic fibres, and myocytes which varied according to gestational age of fetuses. Structure of scrotal ligament and gubernaculum testes is highly variable. Description of scrotal ligament as a firm attachment from lower pole of testes to scrotum is controversial, questioning its role as protective factor in testicular torsion.

[Anatomy of the cervical spine]. / Cervikalcolumnas anatomi.

BACKGROUND: Knowledge of biomechanics and the cervical spine's anatomy has become more topical as the incidence of whiplash neck disorders has increased. Unfortunately, injuries after traffic accidents are often brought to court, where the medical expert's knowledge is of utmost importance to ensure a correct medical evaluation. MATERIAL AND METHODS: The article is based on information identified through non-systematic searches of PubMed and on the author's experience as a professor of anatomy. RESULTS: The cervical spine is particularly vulnerable to forces perpendicular to the length axis. Stability depends largely on the soft tissue. Injuries of soft tissue (especially in ligaments and intervertebral discs) may lead to instability and periosteal reaction with subsequent new formation of bone. INTERPRETATION: The cervical spine is a relatively weak and vulnerable part of the body. One should consider locally restricted new formations of tissue with corresponding height reduction of the intervertebral disc as a sign of genuine injury.

Anatomy of the ligamentum venosum arantii and its contribution to the left hepatic vein and common trunk control. A study on cadaveric livers.

BACKGROUND: The control of the left hepatic vein (LHV) and the common trunk of the middle hepatic vein (MHV) and LHV (CT) is considered difficult during liver resection and could be improved by detailed knowledge on the ligamentum venosum Arantii (LV). AIM: The aim of this study was to describe the LV and its connections to the LHV and the CT and to present surgical relevance of the obtained data. MATERIAL AND METHODS: During autopsy of 50 cadavers of both sexes, the LV was exposed, measured and then dissected, simulating a surgical maneuver to facilitate the approach to the LHV and CT. The extrahepatic parts of the LHV, MHV and CT were measured. RESULTS: The LV was 52-70 mm long and 5-8 mm thick. It had a fibrotic structure and was not patent in 96% of the cases. The extrahepatic part of the LHV measured 3-19 mm, that of the MHV 3-18 mm and that of the CT 4-15 mm. CONCLUSION: LV dissection facilitated extraparenchymatous clamping of the hepatic veins: the extrahepatic parts of the LHV and CT measured > 3 mm in 86 and 84% of the cases, respectively.

Immunoexpression of constitutive and inducible cyclooxygenase isoforms in distinguishing and accessory structures of synovial joints in rat foetuses.

Joint formation is a developmental process regulated by various factors including bone morphogenetic proteins, transforming and growth factors, etc. Recently, a high expression of cyclooxygenase (COX) isoforms in the foetal cartilaginous elements was also revealed. On the other hand, various joint and skeletal abnormalities were seen in laboratory animal and human offspring, exposed in utero to several COX inhibitors. Immunoexpression of constitutive (COX-1) and inducible (COX-2) cyclooxygenase isoforms was evaluated in various articular structures of untreated and unfamiliar 21-day-old male rat foetuses. Both COX isoforms were detected in the articular cartilage and joint capsule, as well as in the intra-articular disc of the temporomandibular joint and meniscus of the knee joint. COX-1 immunostaining was revealed in the anterior and posterior cruciate ligament of the knee joint and the labrum of the hip and shoulder, whereas COX-2 immunoreactivity in those structures was not found. It could be concluded that both constitutive and inducible COX isoforms are physiologically expressed in various structures of synovial joints in rat foetuses at the end of prenatal development.

[Anatomical and histological study of the uterosacral ligament: practical surgical consequences]. / Evaluation anatomique et histologique du ligament utérosacré : conséquences pratiques en chirurgie.

OBJECTIVE: To define the vascular and nervous relationships of the uterosacral ligament and to analyze histologically its content for a better description of this structure. MATERIALS AND METHODS: Three fresh fetal cadavers, three embalmed and one fresh adult cadavers were used. The anatomical relationships of the uterosacral ligament were studied by dissecting one fresh fetal pelvis and two embalmed adult pelves. By histological and immunohistological examinations, eight biopsies of the cervical origin of the complexe ligamentaire utérosacral (USLC) were analyzed: four from fresh fetuses, two from a fresh adult cadaver and two from an embalmed adult cadaver. The specimens were stained with haematoxylin eosin safran (HES) coloration, with antinervous cell specific antibodies (PS100) and with antismooth muscle actine antibodies (to visualize vessel walls) before examination under optical microscope. RESULTS: On anatomic examination, the uterosacral ligament was covered by the visceral pelvic fascia. By removing this fascia, the uterosacral ligament appeared to be a condensation of nervous fibers made up of hypogastric and pelvic nerves forming the hypogastric plexus. Histologically, the uterosacral ligament contained connective tissue, nervous fibers, sympathetic nodes, vessels and fatty tissue. No structured ligamentous organization was identified. CONCLUSION: The uterosacral "ligament" is in fact a "ligament complex" integrating connective tissue as well as nervous and vascular elements. Radical wide excisions of the USLC during cancer or endometriosis surgery and uterosacral suspension during pelvic floor reconstructive surgery should be performed with caution in order to preserve pelvic innervation.

Examination of the Topographical Anatomy and Fetal Development of the Tendinous Annulus of Zinn for a Common Origin of the Extraocular Recti.

Purpose: The aim was to clarify the topographical anatomy of the common tendinous ring for the four rectus muscles in both adults and fetuses. Methods: We histologically examined the annular ligament for a common origin of the extraocular rectus muscles using 10 specimens from elderly individuals and 31 embryonic and fetal specimens. Results: At 6 to 8 weeks, each rectus carried an independent long tendon, individually originating from the sphenoid. Notably, we found additional origins from the optic or oculomotor nerve sheath. At 12 to 15 weeks, the lateral, inferior, and medial recti muscles were united to provide a C-shaped musculofibrous mass that was separated from the superior rectus originating from the edge of the optic canal opening. Morphologic features at 31 to 38 weeks were almost the same as those at 12 to 15 weeks, but the long and thick common tendon of the three recti reached the sphenoid body in the parasellar area. In adults, a ring-like arrangement of the rectus muscles ended at a site 8.1 to 12.0 mm anterior to the optic canal opening and independent of the superior rectus origin, the lateral, inferior, and medial recti formed a C-shaped muscle mass. The united origins of the three recti changed to a fibrous band extending along the superomedial wall of the orbital fissure. Conclusions: Consequently, none of the specimens we examined exhibited an annular tendon representing a common origin of the four recti, suggesting that the common tendinous ring includes only medial, lateral, and inferior rectus muscles with the superior rectus taking its origin independently.

Understanding Tendons: Lessons from Transgenic Mouse Models.

Tendons and ligaments are connective tissues that have been comparatively less studied than muscle and cartilage/bone, even though they are crucial for proper function of the musculoskeletal system. In tendon biology, considerable progress has been made in identifying tendon-specific genes (Scleraxis, Mohawk, and Tenomodulin) in the past decade. However, besides tendon function and the knowledge of a small number of important players in tendon biology, neither the ontogeny of the tenogenic lineage nor signaling cascades have been fully understood. This results in major drawbacks in treatment and repair options following tendon degeneration. In this review, we have systematically evaluated publications describing tendon-related genes, which were studied in depth and characterized by using knockout technologies and the subsequently generated transgenic mouse models (Tg) (knockout mice, KO). We report in a tabular manner, that from a total of 24 tendon-related genes, in 22 of the respective knockout mouse models, phenotypic changes were detected. Additionally, in some of the models it was described at which developmental stages these changes appeared and progressed. To summarize, only loss of Scleraxis and TGFß signaling led to severe tendon developmental phenotypes, while mice deficient for various proteoglycans, Mohawk, EGR1 and 2, and Tenomodulin presented mild phenotypes. These data suggest that the tendon developmental system is well organized, orchestrated, and backed up; this is even more evident among the members of the proteoglycan family, where the compensatory effects are much clearer. In future, it will be of great importance to discover additional master tendon transcription factors and the genes that play crucial roles in tendon development. This would improve our understanding of the genetic makeup of tendons, and will increase the chances of generating tendon-specific drugs to advance overall treatment strategies.

Collagen fibril morphology and organization: implications for force transmission in ligament and tendon.

Connective tissue mechanical behavior is primarily determined by the composition and organization of collagen. In ligaments and tendons, type I collagen is the principal structural element of the extracellular matrix, which acts to transmit force between bones or bone and muscle, respectively. Therefore, characterization of collagen fibril morphology and organization in fetal and skeletally mature animals is essential to understanding how tissues develop and obtain their mechanical attributes. In this study, tendons and ligaments from fetal rat, bovine, and feline, and mature rat were examined with scanning electron microscopy. At early fetal developmental stages, collagen fibrils show fibril overlap and interweaving, apparent fibril ends, and numerous bifurcating/fusing fibrils. Late in fetal development, collagen fibril ends are still present and fibril bundles (fibers) are clearly visible. Examination of collagen fibrils from skeletally mature tissues, reveals highly organized regions but still include fibril interweaving, and regions that are more randomly organized. Fibril bifurcations/fusions are still present in mature tissues but are less numerous than in fetal tissue. To address the continuity of fibrils in mature tissues, fibrils were examined in individual micrographs and consecutive overlaid micrographs. Extensive microscopic analysis of mature tendons and ligaments detected no fibril ends. These data strongly suggest that fibrils in mature ligament and tendon are either continuous or functionally continuous. Based upon this information and published data, we conclude that force within these tissues is directly transferred through collagen fibrils and not through an interfibrillar coupling, such as a proteoglycan bridge.

CRP2 transcript expression pattern in embryonic chick limb.

Members of the cysteine-rich protein (CRP) family are evolutionary conserved proteins that have been implicated in the processes of cell proliferation and differentiation via the cytoskeletal proteins. In this paper, we present the dynamic expression pattern of CPR2 transcripts during chick limb bud development. CRP2 transcripts are located in various tissues, including muscle, arteries, cartilage, ligaments and digit tendons and also in the apical ectodermal ridge and feather buds.

The structure and development of avian lumbosacral specializations of the vertebral canal and the spinal cord with special reference to a possible function as a sense organ of equilibrium.

The avian lumbosacral vertebral column and spinal cord show a number of specializations which have recently been interpreted as a sense organ of equilibrium. This sense organ is thought to support balanced walking on the ground. Although most of the peculiar structures have been described previously, there was a need to reevaluate the specializations with regard to the possible function as a sense organ. Specializations were studied in detail in the adult pigeon. The development of the system was studied both in the pigeon (semiprecocial at hatching) and in the chicken (precocial). Specializations in the vertebral canal consist of a considerable enlargement, which is not due to an increase in the size of the spinal nervous tissue, but to a large glycogen body embedded in a dorsal rhomboid sinus. The dorsal wall of the vertebral canal shows segmented bilateral dorsal grooves, which are covered by the meninges towards the lumen of the vertebral canal leaving openings in the midline and laterally. This results in a system of lumbosacral canals which look and may function similar to the semicircular canals in the inner ear. Laterally these canals open above ventrolateral protrusions or accessory lobes of the spinal cord which contain neurons. There are large subarachnoidal cerebrospinal fluid spaces, lateral and ventral to the accessory lobes. Movement of this fluid is thought to stimulate the lobes mechanically. As to the development of avian lumbosacral specializations, main attention was given to the organization of the lobes and the adjacent fluid spaces including the dorsal canals. In the pigeon the system is far from being adult-like at hatching but maturates rapidly after hatching. In the chicken the system looks already adult-like at hatching. The implications derived from the structural findings are discussed with regard to a possible function of the lumbosacral specializations as a sense organ of equilibrium. The adult-like organization in the newly hatched chickens, which walk around immediately after hatching, supports the assumed function as a sense organ involved in the control of locomotion on the ground.