Musculoskeletal System - Axial Skeleton Development: Difference between revisions
mNo edit summary |
|||
(113 intermediate revisions by 2 users not shown) | |||
Line 1: | Line 1: | ||
{{Header}} | |||
== Introduction == | == Introduction == | ||
[[File:Axial_skeleton.jpg|thumb|300px|Adult axial skeleton]] | [[File:Axial_skeleton.jpg|thumb|300px|Adult axial skeleton]] | ||
During the | During the third week the paraxial {{mesoderm}} forms into "balls" of mesoderm paired either side of the neural groove, called [[Somitogenesis|somites]], that are patterned by the [[Notochord|notochord]]. Similar regions of each somite differentiate initially into 2 parts: the dermomyotome (dermal and muscle component) and the '''sclerotome''' (forms vertebral column). Each somite undergoes a segmental shift to form a vertebral body and the intervertebral disc. | ||
The sclerotome mesenchyme first differentiates to form {{cartilage}}, that ossifies through [[Bone_Histology|endochondral ossification]] to form [[Bone Development|bone]]. The vertebral body begins as a bony collar that expands into regions of dying cartilage. The vertebral arch, enclosing the spinal cord, forms later and the arch remains open dorsally (linked by a ligament) to allow continued growth of the spinal cord. | |||
The axial skeleton consists of: | The axial skeleton consists of: {{skull}}, Auditory Ossicles, Hyoid bone, Vertebral column, Chest ({{sternum}}, {{ribs}}) | ||
The vertebral column is a series of bone segments (vertebra) separated by specialized joints (intervertebral disc). | The vertebral column is a series of bone segments (vertebra) separated by specialized joints (intervertebral disc). | ||
In the adult, the vertebra form rostro-caudally: 7 cervical, 12 thoracic, 5 lumbar, 1 sacrum, coccyx. There has been identified a population variability in the | In the adult, the vertebra form rostro-caudally: 7 cervical, 12 thoracic, 5 lumbar, 1 sacrum, coccyx. There has been identified a population variability in the distribution of the number of vertebra. | ||
Skeletal ossification continues postnatally, through puberty until mid-20s. Abnormalities of vertebral column development can lead to defects including scoliosis. | |||
See this series of articles by O'Rahilly, Müller and others on human vertebral column development using embryos from the [[Carnegie Collection]].{{#pmid:517765|PMID517765}}{{#pmid:7216919|PMID7216919}}{{#pmid:6833119|PMID6833119}}{{#pmid:2323997|PMID2323997}}{{#pmid:2182589|PMID2182589}}{{#pmid:7961131|PMID7961131}} | |||
<br> | |||
:[[Embryology_History|Historic Embryology]]: [[Paper - Vertebral Regional Determination in Young Human Embryos|1908 Vertebrae]] | [[Book_-_Manual_of_Human_Embryology_11C|1910 Vertebral Column]] | [[Paper - Development of the thoracic vertebrae in man|1914 Thoracic Vertebrae]] | |||
<br> | |||
{{Axial Skeleton table}} | |||
<br> | |||
{{Musculoskeletal Links}} | |||
==Some Recent Findings== | |||
{| | |||
|-bgcolor="F5FAFF" | |||
| | |||
{{ | * '''Rib Cage Morphogenesis in the Human Embryo: A Detailed Three-Dimensional Analysis'''{{#pmid:31344324|PMID31344324}} "Formation of the skeletal structure in the human embryo has important consequences in terms of support, protection, and function of organs and other systems. We aimed to describe the formation of the {{rib}} cage during the embryonic period, in order to detect prominent features and identify the possible factors affecting rib cage morphology. We employed high-resolution digitized imaging data (n = 34) obtained in human embryos with Carnegie stage (CS) between {{CS17}} and {{CS23}}. The rib cage became detectable as cartilage formation at CS17, expanding outward from the dorsal side of the chest-abdominal region. Ribs elongated progressively to surround the chest, differentiating into the upper and lower rib cage regions by CS{{CS20}}. The ends of corresponding ribs in the upper region elongated toward each other, leading to their joining and sternum formation between CS21 and CS23, while the lower region of the rib cage remained widely open. The rib cage area with the largest width shifted from the 5th rib pair at CS{{CS17}} to the 9th pair at CS23. The depth of the rib cage was similar across the upper region at CS{{CS17}}, with the major portion remaining in the middle part after CS{{CS20}}. The heart was located beneath the rib pairs providing the largest depth, while the liver was located beneath the rib pairs providing the largest width. Formation of the sternum, development of spinal kyphosis, and organization of larger internal organs within the thoracic and abdominal cavity are possible factors affecting rib cage morphology." | ||
{{ | * '''Evolutionary selection and morphological integration in the vertebral column of modern humans'''{{#pmid:31675109|PMID31675109}} "The main objective is to quantify integration, modularity, and response to selection in the presacral vertebral column of modern humans. Seventeen linear variables on each presacral vertebra were collected in 108 modern humans producing a total of ~39,000 measurements. Then, we studied patterns and magnitudes of integration at regional, vertebral, and intra-vertebral levels. Additionally, we calculated the ability of vertebrae to respond to selection by quantifying differences in evolvability, flexibility, and constraint throughout the spine. The results indicate that caudal vertebrae are more evolvable than those located more cranially in the presacral vertebral column, following an increasing pattern of evolvability from the cervical to the lumbar region. Additionally, the atlas and fifth lumbar vertebra show the lowest values of integration, while central thoracic vertebrae display the highest magnitudes of integration. These results could be related to three main factors: body plan organization expressed by the Hox genes, the strong developmental constraints that determine the number of mammalian vertebrae, and, finally, the functional requirements of an adaptation to bipedal locomotion in the human lineage." | ||
* '''Ossification of the vertebral column in human foetuses: histological and computed tomography studies'''{{#pmid:24068685|PMID24068685}} "There is no agreement in the literature as to the time of the onset and progress of the vertebral column ossification. The aim of the present study was to determine the precise sequence of ossification of the neural arches and vertebral centra.Histological and radiographic studies were performed on 27 human foetuses aged from 9 to 21 weeks. It was found that the ossification of vertebrae commences in foetuses aged 10 and 11 weeks. Ossification centres appear first for neuralarches in the cervical and upper thoracic vertebrae and by the end of 11th week they are present in all thoracic and lumbar neural arches. In the vertebral centrain foetus of 10 weeks ossification was found in the lower 7 thoracic and first lumbar vertebrae. By the end of 11th week ossification is present in the lower 4 cervical, all thoracic, all lumbar and 4 sacral vertebral centra." | |||
|} | |||
--[[ | {| class="wikitable mw-collapsible mw-collapsed" | ||
! More recent papers | |||
|- | |||
| [[File:Mark_Hill.jpg|90px|left]] {{Most_Recent_Refs}} | |||
== | Search term: [http://www.ncbi.nlm.nih.gov/pubmed/?term=Axial+Skeleton+Development ''Axial Skeleton Development''] | [http://www.ncbi.nlm.nih.gov/pubmed/?term=vertebral+column+Development ''Vertebral Column Development''] | [http://www.ncbi.nlm.nih.gov/pubmed/?term=Vertebra+Development ''Vertebra Development''] | [http://www.ncbi.nlm.nih.gov/pubmed/?term=Intervertebral+disc+Development ''Intervertebral Disc Development''] | [http://www.ncbi.nlm.nih.gov/pubmed/?term=rib+Development ''Rib Development''] | [http://www.ncbi.nlm.nih.gov/pubmed/?term=Scoliosis ''Scoliosis''] | ||
|} | |||
{| class="wikitable mw-collapsible mw-collapsed" | |||
! Older papers | |||
|- | |||
| {{Older papers}} | |||
* '''Foxa1 and foxa2 are required for formation of the intervertebral discs'''{{#pmid:23383217|PMID23383217}} "The intervertebral disc (IVD) is composed of 3 main structures, the collagenous annulus fibrosus (AF), which surrounds the gel-like nucleus pulposus (NP), and hyaline cartilage endplates, which are attached to the vertebral bodies. ... The NP forms from the embryonic notochord. Foxa1 and Foxa2, transcription factors in the forkhead box family, are expressed early during notochord development. ...Embryos lacking only Foxa1 or Foxa2 from the notochord were indistinguishable from control animals, demonstrating a functional redundancy for these genes in IVD formation. In addition, we provide in vivo genetic evidence that Foxa genes are required for activation of Shh in the notochord." | |||
* '''Role of environmental factors in axial skeletal dysmorphogenesis'''{{#pmid:20544699|PMID20544699}} "Approximately 1 in 1000 live births is afflicted with an axial skeletal defect. Although many of the known human teratogens can produce axial skeletal defects, the etiology of over half of the observed defects is unknown." | |||
* '''Embryology and bony malformations of the craniovertebral junction.'''{{#pmid:21193993|PMID21193993}} "The embryology of the bony craniovertebral junction (CVJ) is reviewed with the purpose of explaining the genesis and unusual configurations of the numerous congenital malformations in this region. ...A logical classification of this seemingly unwieldy group of malformations is thus possible based on their ontogenetic lineage, morbid anatomy, and clinical relevance. Representative examples of the main constituents of this classification scheme are given, and their surgical treatments are selectively discussed." | |||
|} | |||
== Textbooks == | == Textbooks == | ||
[[File:Developing_vertebra.jpg|thumb|Developing vertebra]] | |||
* '''The Developing Human: Clinically Oriented Embryology''' (8th Edition) by Keith L. Moore and T.V.N Persaud - Moore & Persaud Chapter 15 the skeletal system | * '''The Developing Human: Clinically Oriented Embryology''' (8th Edition) by Keith L. Moore and T.V.N Persaud - Moore & Persaud Chapter 15 the skeletal system | ||
* '''Larsen’s Human Embryology''' by GC. Schoenwolf, SB. Bleyl, PR. Brauer and PH. Francis-West - Chapter 11 Limb Dev (bone not well covered in this textbook) | * '''Larsen’s Human Embryology''' by GC. Schoenwolf, SB. Bleyl, PR. Brauer and PH. Francis-West - Chapter 11 Limb Dev (bone not well covered in this textbook) | ||
Line 37: | Line 69: | ||
* Briefly summarise the development of the limbs. | * Briefly summarise the development of the limbs. | ||
* Describe the developmental abnormalities responsible for the following malformations: selected growth plate disorders; congenital dislocation of the hip; scoliosis; arthrogryposis; and limb reduction deformities. | * Describe the developmental abnormalities responsible for the following malformations: selected growth plate disorders; congenital dislocation of the hip; scoliosis; arthrogryposis; and limb reduction deformities. | ||
== Development Overview == | == Development Overview == | ||
Below is a very brief overview using simple figures of 3 aspects of early musculoskeletal development. More detailed overviews are shown on other notes pages Mesoderm and Somite, Vertebral Column, Limb in combination with serial sections and Carnegie images. | Below is a very brief overview using simple figures of 3 aspects of early musculoskeletal development. More detailed overviews are shown on other notes pages Mesoderm and Somite, Vertebral Column, Limb in combination with serial sections and Carnegie images. | ||
=== Mesoderm | ===Mesoderm=== | ||
{| class="prettytable" width=100% | {| class="prettytable" width=100% | ||
Line 54: | Line 84: | ||
|- | |- | ||
| [[File:Mesoderm cartoon 03.jpg]] | | [[File:Mesoderm cartoon 03.jpg]] | ||
| Segmentation of the paraxial mesoderm into somites continues caudally at 1 somite/90minutes and a cavity ( | | Segmentation of the paraxial mesoderm into somites continues caudally at 1 somite/90minutes and a cavity (intra-embryonic coelom) forms in the lateral plate mesoderm separating somatic and splanchnic mesoderm. | ||
Note | Note intra-embryonic coelomic cavity communicates transiently with extra-embryonic coelom through portals (holes) initially on lateral margin of embryonic disc. | ||
|- | |- | ||
| [[File:Mesoderm cartoon 04.jpg]] | | [[File:Mesoderm cartoon 04.jpg]] | ||
Line 64: | Line 94: | ||
|} | |} | ||
=== | ===Sclerotome=== | ||
The {{Sclerotome}} forms from the ventromedial portion (region) of each somite with a mesenchymal cell organisation. The left and right sclerotome from each somite level contributes the vertebra and intervertebral disc of the entire axial skeleton. | |||
{| class="prettytable" width=100% | {| class="prettytable" width=100% | ||
Line 87: | Line 119: | ||
|} | |} | ||
=== | ===Axial Features=== | ||
* human occipitocervical segmentation<ref><pubmed>14529047</pubmed></ref> | |||
* head and vertebral column boundary - located between the 5th and 6th somites{{#pmid:1605355|PMID1605355}} | |||
* craniovertebral junction - caudal occiput, atlas, and axis{{#pmid:21822704|PMID21822704}} | |||
== Embryonic Development == | |||
===Week 8=== | |||
[[File:Stage22 vertebra and spinal cord 1.jpg|600px]] | |||
'''Human embryo''' (Stage 22) vertebra and spinal cord development. Note the structure of the vertebral arch, the dorsal ligament allowing expansion of the arch to accommodate spinal cord growth. | |||
==Skull== | |||
[[File:Skull_anterior.gif]] | |||
See [[Musculoskeletal_System_-_Skull_Development|Skull Development]] | |||
:'''Links:''' [[Musculoskeletal_System_-_Skull_Development|Skull Development]] | |||
==Vertebral Column== | |||
===Cervical Vertebra=== | |||
[[File:Vertebra ossification sequence.jpg|thumb|Vertebral element ossification between species.{{#pmid:20956304|PMID20956304}}]] | |||
* '''C1''' - (atlas) from fourth occipital and first cervical sclerotomes. Posterior arch ossifies from 2 centres in the lateral masses fusing postnatally between 3 - 5th years. | |||
{| class=" | A comparison of vertebral element ossification between different species also in relation to Hox gene expression.{{#pmid:20956304|PMID20956304}} | ||
* For each taxon, circles indicate centra, and squares indicate left and right neural arches. | |||
* Colours represent the order of ossification. | |||
* Hox expression boundaries in the mouse (placentals) and vertebral segment identity are shown at right. | |||
* Conserved timing of V7 centrum ossification across mammals, including sloths (bradypus) | |||
* Overlap in Hox5-6 expression in the V6–V9 region of the sloth neck. | |||
bradypus - three-toed sloths are the only members of this genus | |||
dasypus - armadillo genus in the Dasypodidae family | |||
===Sacrum=== | |||
'''Evolution of the sacrum'''{{#pmid:3688211|PMID3688211}} "In order to study the formation of the sacrum during the primate evolution, a new way of numbering mammalian vertebrae is presented; this demonstrates that the thoracolumbosacral complex is fixed at 22 vertebrae in 80% and at 22 +/- 1 in 100% of the cases. The shift of a vertebra from one type to another occurs either at the thoracolumbar or at the lumbosacral junction and not at the cervicothoracic junction. Rarely does the shift take place at the sacrococcygeal junction. Data from 318 primates reveal that the seven original lumbar vertebrae of the Old World monkeys are reduced in the great apes by a caudad "thoracization" of one to two lumbar vertebrae and a cephalad sacralization of one to four lumbar vertebrae. In the apes, sacralization is not total and different stages that are intermediate between lumbar and sacral are described. In Homo sapiens there is a total sacralization of the last two original lumbar vertebrae. In addition, development of the sacral wings (alae) is minimal in apes and reaches its maximum in hominids. The tendency of the hominoid sacrum to incorporate the last lumbar vertebrae and to widen markedly provides for an enhanced articulation of the sacrum with the ilium and offers a firm base of support for the trunk during erect posture. This is necessary for the support of the weight of the trunk above the sacrum and for the stabilization of the body during bipedal posture and locomotion. Encephalization did not play any major role in the widening of the sacrum since the former by far preceded the latter." | |||
==Intervertebral Disc== | |||
[[File:Mouse-_intervertebral_disc_development_02.jpg|thumb|Mouse intervertebral disc (IVD) expression of transcription factor Nfatc1{{#pmid:20214815|PMID20214815}}]] | |||
The adult intervertebral disc (IVD) has to bear the same loads as the vertebra and also have flexibility to allow axial column movement. This is achieved by a complex structure (cartilaginous end-plate) that links the vertebra above and below the disc to a outer dense fibrous structure (annulus fibrosus) containing a gel-like core region (nucleus pulposus). Some research in this area focusses on the degeneration of the IVD with ageing. | |||
* cartilaginous end-plate - that anchor the discs to the adjacent vertebral body bones | |||
* annulus fibrosus - cells are derived from the sclerotome | |||
* nucleus pulposus - cells are derived from the notochord{{#pmid:20568241|PMID20568241}} | |||
===Annulus Fibrosus=== | |||
* cells are derived from the sclerotome | |||
===Nucleus Pulposus=== | |||
* cells are derived from the notochord{{#pmid:20568241|PMID20568241}} | |||
* notochordal marker brachyury | |||
* proteoglycan rich extracellular matrix | |||
==Rib Development== | |||
[[File:Keibel Mall 232.jpg|thumb|200px|Embryonic rib development([[Week 7]], Stage {{CS18}}, Human Embryo {{CE109}}, length 11 mm.]] | |||
[[File:Gray0965.jpg|thumb|200px|Adult Ribcage and Sternum]] | |||
Humans form 12 paired {{ribs}} from the cartilaginous costal processes of the developing thoracic vertebrae. Early rib development occurs at 7 weeks (((GA}} week 9) from lateral plate mesoderm and continues postnatal with secondary ossification centres appearing at 15 years of age. | |||
A recent paper has looked at rib cage development{{#pmid:31344324|PMID31344324}} between embryonic [[Week 7]] to [[Week 9]], {{CS17}} to {{CS23}}. First identifying cartilage formation at CS17{{CS17}} that expanding outward from the dorsal side of the chest-abdominal region. Ribs elongated progressively to surround the chest, differentiating into the upper and lower rib cage regions by CS{{CS20}}. | |||
* First 7 “true” ribs connect to the {{sternum}} through the costal cartilages about day 45. | |||
* Lower 5 “false” ribs do not connect to the {{sternum}}. | |||
<gallery caption="Human Embryonic Rib Development"> | |||
File:Keibel_Mall_260.jpg|Embryo 13 mm CRL | |||
File:Keibel_Mall_261.jpg|Embryo 17 mm CRL | |||
File:Keibel_Mall_262.jpg|Embryo 16 mm CRL | |||
File:Keibel_Mall_263.jpg|Embryo 32 mm CRL | |||
</gallery> | |||
[[File:Mouse_E12.5_Sox9_Expression.jpg|250px|alt=Mouse_E12.5_Sox9_Expression]] | |||
{{Sox}}9 expression in the Mouse ({{ME12.5}}) rib primordial.{{#pmid:23236180|PMID23236180}} | |||
{| class="wikitable mw-collapsible mw-collapsed" | |||
! Historic Rib papers | |||
|- | |- | ||
| [[ | | {{Ref-Bardeen1910}} | ||
| | [[Book_-_Manual_of_Human_Embryology_11C#Chondrogenous_Period|Chondrogenous Period]] and [[Book_-_Manual_of_Human_Embryology_11C#Ribs|Rib Development]] | ||
:"Ossification begins in the ribs before it does in the vertebrae. Centres appear in the bodies of the sixth and seventh ribs toward the end of the second month and then rapidly come to view in the other ribs. The centre in the first rib usually appears before that in the twelfth. All are usually present by the end of the second month, but that in the twelfth may not appear until later. In two specimens out of 29 fetuses with an estimated age of 55 to 110 days, Mall (1906) found a centre of ossification in the costal element of the seventh cervical vertebra." | |||
|} | |||
Developmental rib abnormalities may be isolated or can occur with other congenital anomalies. | |||
==Sternum Development== | |||
The sternum and sternal ribs derive from the somatic layer of the lateral plate mesoderm.{{#pmid:10803330|PMID10803330}}{{#pmid:26467243|PMID26467243}} | |||
{| class="wikitable mw-collapsible mw-collapsed" | |||
! Historic Sternum papers | |||
|- | |- | ||
| {{Ref-Paterson1900}} | |||
{{Ref-Keith1902}} | |||
[[Book_-_Human_Embryology_and_Morphology_19#Formation_of_Ribs|Body Wall, Ribs, and Sternum]] | |||
{{Ref-WhiteheadWaddell1911}} | |||
|} | |||
==Molecular== | |||
Like many other embryonic structures there are two separate considerations: | |||
# '''Pattern Formation''' - sclerotome differentiation and segmentation | |||
# '''Overt Differentiation''' - mesoderm differentiation to cartilage differentiation and ossification to bone. | |||
===Notch=== | |||
:'''Links:''' [[Developmental_Signals_-_Notch|Notch]] | |||
===Nuclear factor of activated T-cells, cytoplasmic 1=== | |||
[[File:Mouse-_intervertebral_disc_development_02.jpg|thumb|Mouse intervertebral disc (IVD) expression of transcription factor Nfatc1{{#pmid:20214815|PMID20214815}}]] | |||
* Transcription factor (Nfatc1) has been identified in developing mouse intervertebral disc (IVD). | |||
* The NFAT family of transcription factors regulates cytokine gene expression by binding to the promoter/enhancer regions of antigen-responsive genes, usually in cooperation with heterologous DNA-binding partners. | |||
* The activation of NFAT proteins is controlled by calcineurin, the calmodulin-dependent phosphatase. | |||
:'''Links:''' [http://www.ncbi.nlm.nih.gov/omim/600489 OMIM - NFATC1] | |||
===Transforming growth factor beta=== | |||
Two functions in IVD development:{{#pmid:20214815|PMID20214815}} | |||
# prevent chondrocyte differentiation in the presumptive IVD | |||
# promote differentiation of annulus fibrosus from sclerotome | |||
==Abnormalities== | |||
===Spina Bifida=== | |||
[[File:USA spina bifida rates.jpg|thumb|USA spina bifida rates.]] | |||
[[Abnormal_Development_-_Folic_Acid_and_Neural_Tube_Defects|Folic Acid and Neural Tube Defects]] | |||
===Absent Cervical Spine Pedicle=== | |||
[[File:Absent cervical spine pedicle.jpg|400px]] | |||
'''Absent cervical spine pedicle'''{{#pmid:21062465|PMID21062465}} | |||
The vertebral pedicles (Latin, ''pediculus'' ="small foot") are paired processes that project dorsally and connect the body of the spinal vertebra to the arch. Absence is a rare abnormality characterized by the absence of a pedicle of the affected vertebral body, seen most frequently at the level C6 followed by the level C5 and C7. | |||
===Scoliosis=== | |||
[[File:Braune 1877 plate 30 fig1.jpg|thumb|Historic image scoliosis (Braune 1877)]] | |||
[[File:Scoliometry_01.jpg|thumb|Scoliometry{{#pmid:23226218|PMID23226218}}]] | |||
[[File:Cheneau brace.jpg|thumb|Chêneau Brace{{#pmid:21266084|PMID21266084}}]] | |||
[[File:Scoliosis.jpg]] [[File:Scoliosis xray.jpg]] | |||
* frequency of congenital scoliosis is approximately 0.5, or 1 in 1,000 births. | |||
* assymetric growth impairment of vertebral bodies | |||
* lateral deviation of spine (Lateral flexion, Forward flexion, Rotation of vertebral column on long axis) | |||
* compensated by movement of vertebral column above and below affected region (producing a primary and two secondary curves) | |||
* progresses rapidly in adolescence and becomes fixed once bone growth is completed. | |||
* both genetic and environmental terartogens implicated. | |||
** a large number of chromosomal deletions relating to regions {{Chr2}}p13 – 15, {{Chr6}}q13 and {{Chr15}}q12 have been identified. | |||
:'''Links:''' [http://www.ncbi.nlm.nih.gov/pubmed/?term=Scoliosis ''Scoliosis''] | |||
===VACTERL=== | |||
{| | |||
|-bgcolor="FEF9E7" | |||
| {{ICD-11}} - {{ICD11weblink}}1452617987 LD2F.11 VATER association] - ''VACTERL/VATER is an association of congenital malformations typically characterized by the presence of at least three of the following: vertebral defects, anal atresia, cardiac defects, tracheo-esophageal fistula, renal anomalies, and limb abnormalities.'' | |||
|} | |} | ||
{{VACTERL}}/VATER are the acronyms used to describe a multi-system congenital malformations including at least three of the following: | |||
# '''V'''ertebral defects | |||
# '''A'''nal atresia | |||
# '''C'''ardiac defects | |||
# '''T'''racheo-esophageal fistula | |||
# '''R'''enal anomalies | |||
# '''L'''imb abnormalities | |||
{{VACTERL Diagnosis collapse table}} | |||
:'''Links:''' {{VACTERL}} | |||
===Sagittal Cleft Vertebra=== | |||
Sagittal cleft vertebra, "butterfly vertebrae" or anterior rachischisis, is an abnormal vertebral body caused by failure of fusion of the 2 lateral chondrification centres during development. The term "butterfly vertebrae" is based upon the butterfly wings appearance of the 2 hemivertebrae on x-ray. See a recent review.{{#pmid"31448202|PMID31448202}} | |||
:'''Links:''' [http://www.ncbi.nlm.nih.gov/pubmed/?term=Sagittal+Cleft+Vertebra ''Sagittal Cleft Vertebra''] | |||
===Spondylocostal Dysostosis=== | |||
A rare heritable axial skeleton growth disorder characterized by abnormal development of vertebra, multiple segmentation defects, and ribs. | |||
:'''Links:''' [https://www.ncbi.nlm.nih.gov/books/NBK8828/ GeneReviews] | [http://www.ncbi.nlm.nih.gov/pubmed/?term=Spondylocostal+Dysostosis ''Spondylocostal Dysostosis''] | |||
== References == | == References == | ||
<references/> | <references/> | ||
Line 101: | Line 315: | ||
===Reviews=== | ===Reviews=== | ||
{{#pmid:21193993}} | |||
{{#pmid:19651306}} | |||
{{#pmid:19575673}} | |||
{{#pmid:19247958}} | |||
{{#pmid:18157900}} | |||
===Articles=== | |||
{{#pmid:19590010}} | |||
{{#pmid:19520072}} | |||
{{#pmid:19395637}} | |||
===Search PubMed=== | ===Search PubMed=== | ||
'''Search Pubmed:''' [http://www.ncbi.nlm.nih.gov/sites/entrez?db=pubmed&cmd=search&term=Axial%20Skeleton%20Development Axial Skeleton Development] | [http://www.ncbi.nlm.nih.gov/sites/entrez?db=pubmed&cmd=search&term=Vertebra%20Development Vertebra Development] | [http://www.ncbi.nlm.nih.gov/sites/entrez?db=pubmed&cmd=search&term=Intervertebral%20Disc%20Development Intervertebral Disc Development] | [http://www.ncbi.nlm.nih.gov/sites/entrez?db=pubmed&cmd=search&term=Axial%20Skeleton%20Abnormalities Axial Skeleton Abnormalities] | |||
'''Search Pubmed:''' [http://www.ncbi.nlm.nih.gov/sites/entrez?db=pubmed&cmd=search&term= | |||
==Additional Images== | ==Additional Images== | ||
Line 118: | Line 341: | ||
File:Bone-structure.jpg|Bone structure | File:Bone-structure.jpg|Bone structure | ||
File:Developing_vertebra.jpg|Developing vertebra | File:Developing_vertebra.jpg|Developing vertebra | ||
File:Fetal_head_lateral.jpg|Fetal head lateral (12 weeks) | File:Fetal_head_lateral.jpg|Fetal head lateral (12 weeks) | ||
File:Fetal_head_medial.jpg|Fetal head medial (12 weeks) | File:Fetal_head_medial.jpg|Fetal head medial (12 weeks) | ||
File:Fetal_head_section.jpg|Fetal head section (12 weeks) | File:Fetal_head_section.jpg|Fetal head section (12 weeks) | ||
File:Human axial skeleton- axis development 01.jpg|Historic plate - human axis development | |||
</gallery> | |||
===Historic=== | |||
{| class="wikitable mw-collapsible mw-collapsed" | |||
! Human Embryology And Morphology (1921) | |||
|- | |||
| Keith, A. [[Book - Human Embryology and Morphology (1921)|'''Human Embryology And Morphology''']] (1921) Longmans, Green & Co.:New York. | |||
[[Human Embryology and Morphology_5|5 Spinal Column and Back]] | |||
<gallery> | |||
File:Keith1921 fig047.jpg|Fig. 47. Pyramids of the Spine | |||
File:Keith1921 fig048.jpg|Fig. 48. Curves of the Spinal Column | |||
File:Keith1921 fig049.jpg|Fig. 49. Lumbo-sacral Region Spine Foetus end of the 2nd month | |||
File:Keith1921 fig050.jpg|Fig. 50. Variation in costal element of the seventh Cervical Vertebra | |||
File:Keith1921 fig051.jpg|Fig. 51. Sclerotome, muscle plate and skin plate | |||
File:Keith1921 fig052.jpg|Fig. 52. Where remnants of the Notochord may occur in the Adult | |||
File:Keith1921 fig053.jpg|Fig. 53. Notochord to the basilar or parachordal cartilage | |||
File:Keith1921 fig054.jpg|Fig. 54. Parts of the first Cervical Vertebra | |||
File:Keith1921 fig055.jpg|Fig. 55. Membranous Basis of a Vertebra | |||
File:Keith1921 fig056.jpg|Fig. 56. Stages in the Development of a Vertebra | |||
File:Keith1921 fig057.jpg|Fig. 57. Order Centres of Ossification appear in the Bodies | |||
File:Keith1921 fig058.jpg|Fig. 58. Foetal Axis, Atlas, and Basi-occipital | |||
File:Keith1921 fig059.jpg|Fig. 59. Atlanto-axio-occipital Articulations | |||
File:Keith1921 fig060.jpg|Fig. 60. Bicipital Rib of a Lower Vertebrate (crocodile) | |||
File:Keith1921 fig061.jpg|Fig. 61. Half of a first Lumbar Vertebra | |||
File:Keith1921 fig062.jpg|Fig. 62. Elements composing the Sacrum | |||
File:Keith1921 fig063.jpg|Fig. 63. Human caudal or coccygeal region | |||
</gallery> | </gallery> | ||
|} | |||
==Terms== | |||
* '''centrum''' - (vertebral body) anatomical term referring to the main bony part of the vertebra that forms the majority of the axial skeleton. | |||
* '''Cobb angle''' - clinical method of measuring the degree of scoliosis and post-traumatic kyphosis. Named after the American orthopedic surgeon John Robert Cobb (1903 - 1967) an American orthopedic surgeon. | |||
* '''haemal arch''' - referring to the bony arch region within animal tail vertebra that contains blood vessels. | |||
* '''idiopathic scoliosis''' - clinical term for a spinal column deformity appearing usually postnatally after the age of 10 years old. | |||
* '''lordosis''' - clinical and anatomy term describing the curvature of the spine with the convexity toward the front. Normal to have lordosis in the cervical and lumbar regions of the spinal column. | |||
* '''neural arch''' - referring to the bony arch region within vertebra that contains the spinal cord. | |||
* '''notochord sheath''' - region surrounding the notochord. in teleost fish direct mineralization of this region, by intramembranous ossification, forms the initial vertebral centrum. (More? {{Zebrafish}}) | |||
* '''rib hump''' - clinical term for the prominence formed by ribs on the convexity of a curve, caused by rotation of the spine and attached ribs. | |||
* '''spinal column''' - term referring to the musculoskeletal elements (vertebrae, ligaments, and intervertebral discs), that surround the spinal cord and form the axial skeleton. | |||
* '''synsacrum''' - in birds sacral and lumbar vertebrae fused region forming elongated sacral region. | |||
* '''thoracic hypokyphosis''' - term referring to the backward curve in the upper spine is to great, round back, Scheuermann's disease, or simply kyphosis. | |||
==Terms== | ==Terms== | ||
{{ | {{Bone terms}} | ||
==External Links== | |||
{{External Links}} | |||
* [http://www.scoliosis-australia.org/scoliosis/about_scoliosis.html '''Scoliosis Australia'''] | |||
* [http://www.scoliosisjournal.com/ '''Scoliosis'''] - the official journal of the Society on Scoliosis Orthopaedic and Rehabilitation Treatment (SOSORT) and affiliated with the International Research Society of Spinal Deformities (IRSSD). | |||
* [http://www.youngwomenshealth.org/index.html '''Young Women's Health'''] - [http://www.youngwomenshealth.org/scoliosis-article.html scoliosis] | |||
{{Glossary}} | |||
{{ | {{Footer}} | ||
[[Category: | [[Category:Musculoskeletal]] [[Category:Axial Skeleton]] [[Category:Notochord]] [[Category:Bone]] [[Category:Cartilage]] [[Category:Vertebra]] |
Latest revision as of 23:43, 7 January 2020
Embryology - 14 Jun 2024 Expand to Translate |
---|
Google Translate - select your language from the list shown below (this will open a new external page) |
العربية | català | 中文 | 中國傳統的 | français | Deutsche | עִברִית | हिंदी | bahasa Indonesia | italiano | 日本語 | 한국어 | မြန်မာ | Pilipino | Polskie | português | ਪੰਜਾਬੀ ਦੇ | Română | русский | Español | Swahili | Svensk | ไทย | Türkçe | اردو | ייִדיש | Tiếng Việt These external translations are automated and may not be accurate. (More? About Translations) |
Introduction
During the third week the paraxial mesoderm forms into "balls" of mesoderm paired either side of the neural groove, called somites, that are patterned by the notochord. Similar regions of each somite differentiate initially into 2 parts: the dermomyotome (dermal and muscle component) and the sclerotome (forms vertebral column). Each somite undergoes a segmental shift to form a vertebral body and the intervertebral disc.
The sclerotome mesenchyme first differentiates to form cartilage, that ossifies through endochondral ossification to form bone. The vertebral body begins as a bony collar that expands into regions of dying cartilage. The vertebral arch, enclosing the spinal cord, forms later and the arch remains open dorsally (linked by a ligament) to allow continued growth of the spinal cord.
The axial skeleton consists of: skull, Auditory Ossicles, Hyoid bone, Vertebral column, Chest (sternum, ribs)
The vertebral column is a series of bone segments (vertebra) separated by specialized joints (intervertebral disc).
In the adult, the vertebra form rostro-caudally: 7 cervical, 12 thoracic, 5 lumbar, 1 sacrum, coccyx. There has been identified a population variability in the distribution of the number of vertebra.
Skeletal ossification continues postnatally, through puberty until mid-20s. Abnormalities of vertebral column development can lead to defects including scoliosis.
See this series of articles by O'Rahilly, Müller and others on human vertebral column development using embryos from the Carnegie Collection.[1][2][3][4][5][6]
skull (22) | Auditory Ossicles (6) | Hyoid bone (1) | Vertebral Column (26) | Thoracic cage (27) |
---|---|---|---|---|
|
|
|||
Links: axial skeleton |
Some Recent Findings
|
More recent papers |
---|
This table allows an automated computer search of the external PubMed database using the listed "Search term" text link.
More? References | Discussion Page | Journal Searches | 2019 References | 2020 References Search term: Axial Skeleton Development | Vertebral Column Development | Vertebra Development | Intervertebral Disc Development | Rib Development | Scoliosis |
Older papers |
---|
These papers originally appeared in the Some Recent Findings table, but as that list grew in length have now been shuffled down to this collapsible table.
See also the Discussion Page for other references listed by year and References on this current page.
|
Textbooks
- The Developing Human: Clinically Oriented Embryology (8th Edition) by Keith L. Moore and T.V.N Persaud - Moore & Persaud Chapter 15 the skeletal system
- Larsen’s Human Embryology by GC. Schoenwolf, SB. Bleyl, PR. Brauer and PH. Francis-West - Chapter 11 Limb Dev (bone not well covered in this textbook)
- Before we Are Born (5th ed.) Moore and Persaud Chapter 16,17: p379-397, 399-405
- Essentials of Human Embryology Larson Chapter 11 p207-228
Objectives
- Identify the components of a somite and the adult derivatives of each component.
- Give examples of sites of (a) endochondral and (b) intramembranous ossification and to compare these two processes.
- Identify the general times (a) of formation of primary and (b) of formation of secondary ossification centres, and (c) of fusion of such centres with each other.
- Briefly summarise the development of the limbs.
- Describe the developmental abnormalities responsible for the following malformations: selected growth plate disorders; congenital dislocation of the hip; scoliosis; arthrogryposis; and limb reduction deformities.
Development Overview
Below is a very brief overview using simple figures of 3 aspects of early musculoskeletal development. More detailed overviews are shown on other notes pages Mesoderm and Somite, Vertebral Column, Limb in combination with serial sections and Carnegie images.
Mesoderm
Cells migrate through the primitive streak to form mesodermal layer. Extraembryonic mesoderm lies adjacent to the trilaminar embryo totally enclosing the amnion, yolk sac and forming the connecting stalk. | |
Paraxial mesoderm accumulates under the neural plate with thinner mesoderm laterally. This forms 2 thickened streaks running the length of the embryonic disc along the rostrocaudal axis. In humans, during the 3rd week, this mesoderm begins to segment. The neural plate folds to form a neural groove and folds. | |
Segmentation of the paraxial mesoderm into somites continues caudally at 1 somite/90minutes and a cavity (intra-embryonic coelom) forms in the lateral plate mesoderm separating somatic and splanchnic mesoderm.
Note intra-embryonic coelomic cavity communicates transiently with extra-embryonic coelom through portals (holes) initially on lateral margin of embryonic disc. | |
Somites continue to form. The neural groove fuses dorsally to form a tube at the level of the 4th somite and "zips up cranially and caudally and the neural crest migrates into the mesoderm. |
Sclerotome
The sclerotome forms from the ventromedial portion (region) of each somite with a mesenchymal cell organisation. The left and right sclerotome from each somite level contributes the vertebra and intervertebral disc of the entire axial skeleton.
Mesoderm beside the notochord (axial mesoderm, blue) thickens, forming the paraxial mesoderm as a pair of strips along the rostro-caudal axis. | |
Paraxial mesoderm towards the rostral end, begins to segment forming the first somite. Somites are then sequentially added caudally. The somitocoel, is a cavity forming in early somites, which is lost as the somite matures. | |
Cells in the somite differentiate medially to form the sclerotome (forms vertebral column) and dorsolaterally to form the dermomyotome. | |
The dermomyotome then forms the dermotome (forms dermis) and myotome (forms muscle).
Neural crest cells migrate beside and through somite. | |
The myotome differentiates to form 2 components dorsally the epimere and ventrally the hypomere, which in turn form epaxial and hypaxial muscles respectively. The bulk of the trunk and limb muscle coming from the Hypaxial mesoderm. Different structures will be contributed depending upon the somite level. |
Axial Features
- human occipitocervical segmentation[13]
- head and vertebral column boundary - located between the 5th and 6th somites[14]
- craniovertebral junction - caudal occiput, atlas, and axis[15]
Embryonic Development
Week 8
Human embryo (Stage 22) vertebra and spinal cord development. Note the structure of the vertebral arch, the dorsal ligament allowing expansion of the arch to accommodate spinal cord growth.
Skull
- Links: Skull Development
Vertebral Column
Cervical Vertebra
- C1 - (atlas) from fourth occipital and first cervical sclerotomes. Posterior arch ossifies from 2 centres in the lateral masses fusing postnatally between 3 - 5th years.
A comparison of vertebral element ossification between different species also in relation to Hox gene expression.[16]
- For each taxon, circles indicate centra, and squares indicate left and right neural arches.
- Colours represent the order of ossification.
- Hox expression boundaries in the mouse (placentals) and vertebral segment identity are shown at right.
- Conserved timing of V7 centrum ossification across mammals, including sloths (bradypus)
- Overlap in Hox5-6 expression in the V6–V9 region of the sloth neck.
bradypus - three-toed sloths are the only members of this genus
dasypus - armadillo genus in the Dasypodidae family
Sacrum
Evolution of the sacrum[17] "In order to study the formation of the sacrum during the primate evolution, a new way of numbering mammalian vertebrae is presented; this demonstrates that the thoracolumbosacral complex is fixed at 22 vertebrae in 80% and at 22 +/- 1 in 100% of the cases. The shift of a vertebra from one type to another occurs either at the thoracolumbar or at the lumbosacral junction and not at the cervicothoracic junction. Rarely does the shift take place at the sacrococcygeal junction. Data from 318 primates reveal that the seven original lumbar vertebrae of the Old World monkeys are reduced in the great apes by a caudad "thoracization" of one to two lumbar vertebrae and a cephalad sacralization of one to four lumbar vertebrae. In the apes, sacralization is not total and different stages that are intermediate between lumbar and sacral are described. In Homo sapiens there is a total sacralization of the last two original lumbar vertebrae. In addition, development of the sacral wings (alae) is minimal in apes and reaches its maximum in hominids. The tendency of the hominoid sacrum to incorporate the last lumbar vertebrae and to widen markedly provides for an enhanced articulation of the sacrum with the ilium and offers a firm base of support for the trunk during erect posture. This is necessary for the support of the weight of the trunk above the sacrum and for the stabilization of the body during bipedal posture and locomotion. Encephalization did not play any major role in the widening of the sacrum since the former by far preceded the latter."
Intervertebral Disc
The adult intervertebral disc (IVD) has to bear the same loads as the vertebra and also have flexibility to allow axial column movement. This is achieved by a complex structure (cartilaginous end-plate) that links the vertebra above and below the disc to a outer dense fibrous structure (annulus fibrosus) containing a gel-like core region (nucleus pulposus). Some research in this area focusses on the degeneration of the IVD with ageing.
- cartilaginous end-plate - that anchor the discs to the adjacent vertebral body bones
- annulus fibrosus - cells are derived from the sclerotome
- nucleus pulposus - cells are derived from the notochord[19]
Annulus Fibrosus
- cells are derived from the sclerotome
Nucleus Pulposus
- cells are derived from the notochord[19]
- notochordal marker brachyury
- proteoglycan rich extracellular matrix
Rib Development
Humans form 12 paired ribs from the cartilaginous costal processes of the developing thoracic vertebrae. Early rib development occurs at 7 weeks (((GA}} week 9) from lateral plate mesoderm and continues postnatal with secondary ossification centres appearing at 15 years of age.
A recent paper has looked at rib cage development[7] between embryonic Week 7 to Week 9, 17 to 23. First identifying cartilage formation at CS1717 that expanding outward from the dorsal side of the chest-abdominal region. Ribs elongated progressively to surround the chest, differentiating into the upper and lower rib cage regions by CS20.
- First 7 “true” ribs connect to the sternum through the costal cartilages about day 45.
- Lower 5 “false” ribs do not connect to the sternum.
Sox9 expression in the Mouse (E12.5) rib primordial.[20]
Historic Rib papers |
---|
Bardeen CR. XI. Development of the Skeleton and of the Connective Tissues in Keibel F. and Mall FP. Manual of Human Embryology I. (1910) J. B. Lippincott Company, Philadelphia.
Chondrogenous Period and Rib Development
|
Developmental rib abnormalities may be isolated or can occur with other congenital anomalies.
Sternum Development
The sternum and sternal ribs derive from the somatic layer of the lateral plate mesoderm.[21][22]
Historic Sternum papers |
---|
Paterson The sternum - its early development and ossification in man and mammals. (1900) J Anat Physiol. 35(1): 21-32 PMID 17232454
Keith A. Human Embryology and Morphology. (1902) London: Edward Arnold. Body Wall, Ribs, and Sternum Whitehead RH. and Waddell JA. The early development of the mammalian sternum (1911) Amer. J Anat. 12: 89-106. |
Molecular
Like many other embryonic structures there are two separate considerations:
- Pattern Formation - sclerotome differentiation and segmentation
- Overt Differentiation - mesoderm differentiation to cartilage differentiation and ossification to bone.
Notch
- Links: Notch
Nuclear factor of activated T-cells, cytoplasmic 1
- Transcription factor (Nfatc1) has been identified in developing mouse intervertebral disc (IVD).
- The NFAT family of transcription factors regulates cytokine gene expression by binding to the promoter/enhancer regions of antigen-responsive genes, usually in cooperation with heterologous DNA-binding partners.
- The activation of NFAT proteins is controlled by calcineurin, the calmodulin-dependent phosphatase.
- Links: OMIM - NFATC1
Transforming growth factor beta
Two functions in IVD development:[18]
- prevent chondrocyte differentiation in the presumptive IVD
- promote differentiation of annulus fibrosus from sclerotome
Abnormalities
Spina Bifida
Folic Acid and Neural Tube Defects
Absent Cervical Spine Pedicle
Absent cervical spine pedicle[23]
The vertebral pedicles (Latin, pediculus ="small foot") are paired processes that project dorsally and connect the body of the spinal vertebra to the arch. Absence is a rare abnormality characterized by the absence of a pedicle of the affected vertebral body, seen most frequently at the level C6 followed by the level C5 and C7.
Scoliosis
- frequency of congenital scoliosis is approximately 0.5, or 1 in 1,000 births.
- assymetric growth impairment of vertebral bodies
- lateral deviation of spine (Lateral flexion, Forward flexion, Rotation of vertebral column on long axis)
- compensated by movement of vertebral column above and below affected region (producing a primary and two secondary curves)
- progresses rapidly in adolescence and becomes fixed once bone growth is completed.
- both genetic and environmental terartogens implicated.
- Links: Scoliosis
VACTERL
ICD-11 - LD2F.11 VATER association - VACTERL/VATER is an association of congenital malformations typically characterized by the presence of at least three of the following: vertebral defects, anal atresia, cardiac defects, tracheo-esophageal fistula, renal anomalies, and limb abnormalities. |
VACTERL/VATER are the acronyms used to describe a multi-system congenital malformations including at least three of the following:
- Vertebral defects
- Anal atresia
- Cardiac defects
- Tracheo-esophageal fistula
- Renal anomalies
- Limb abnormalities
VACTERL Clinical Diagnosis | |||||||||||||||||||||||||||
---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|
|
- Links: VACTERL
Sagittal Cleft Vertebra
Sagittal cleft vertebra, "butterfly vertebrae" or anterior rachischisis, is an abnormal vertebral body caused by failure of fusion of the 2 lateral chondrification centres during development. The term "butterfly vertebrae" is based upon the butterfly wings appearance of the 2 hemivertebrae on x-ray. See a recent review.{{#pmid"31448202|PMID31448202}}
- Links: Sagittal Cleft Vertebra
Spondylocostal Dysostosis
A rare heritable axial skeleton growth disorder characterized by abnormal development of vertebra, multiple segmentation defects, and ribs.
- Links: GeneReviews | Spondylocostal Dysostosis
References
- ↑ O'Rahilly R & Meyer DB. (1979). The timing and sequence of events in the development of the human vertebral column during the embryonic period proper. Anat. Embryol. , 157, 167-76. PMID: 517765
- ↑ O'Rahilly R, Muller F & Meyer DB. (1980). The human vertebral column at the end of the embryonic period proper. 1. The column as a whole. J. Anat. , 131, 565-75. PMID: 7216919
- ↑ O'Rahilly R, Müller F & Meyer DB. (1983). The human vertebral column at the end of the embryonic period proper. 2. The occipitocervical region. J. Anat. , 136, 181-95. PMID: 6833119
- ↑ O'Rahilly R, Müller F & Meyer DB. (1990). The human vertebral column at the end of the embryonic period proper. 3. The thoracicolumbar region. J. Anat. , 168, 81-93. PMID: 2323997
- ↑ O'Rahilly R, Müller F & Meyer DB. (1990). The human vertebral column at the end of the embryonic period proper. 4. The sacrococcygeal region. J. Anat. , 168, 95-111. PMID: 2182589
- ↑ Müller F & O'Rahilly R. (1994). Occipitocervical segmentation in staged human embryos. J. Anat. , 185 ( Pt 2), 251-8. PMID: 7961131
- ↑ 7.0 7.1 Okuno K, Ishizu K, Matsubayashi J, Fujii S, Sakamoto R, Ishikawa A, Yamada S, Yoneyama A & Takakuwa T. (2019). Rib Cage Morphogenesis in the Human Embryo: A Detailed Three-Dimensional Analysis. Anat Rec (Hoboken) , 302, 2211-2223. PMID: 31344324 DOI.
- ↑ Arlegi M, Veschambre-Couture C & Gómez-Olivencia A. (2019). Evolutionary selection and morphological integration in the vertebral column of modern humans. Am. J. Phys. Anthropol. , , . PMID: 31675109 DOI.
- ↑ Skórzewska A, Grzymisławska M, Bruska M, Lupicka J & Woźniak W. (2013). Ossification of the vertebral column in human foetuses: histological and computed tomography studies. Folia Morphol. (Warsz) , 72, 230-8. PMID: 24068685
- ↑ Maier JA, Lo Y & Harfe BD. (2013). Foxa1 and Foxa2 are required for formation of the intervertebral discs. PLoS ONE , 8, e55528. PMID: 23383217 DOI.
- ↑ Alexander PG & Tuan RS. (2010). Role of environmental factors in axial skeletal dysmorphogenesis. Birth Defects Res. C Embryo Today , 90, 118-32. PMID: 20544699 DOI.
- ↑ Pang D & Thompson DN. (2011). Embryology and bony malformations of the craniovertebral junction. Childs Nerv Syst , 27, 523-64. PMID: 21193993 DOI.
- ↑ <pubmed>14529047</pubmed>
- ↑ Christ B & Wilting J. (1992). From somites to vertebral column. Ann. Anat. , 174, 23-32. PMID: 1605355
- ↑ Raybaud C. (2011). Anatomy and development of the craniovertebral junction. Neurol. Sci. , 32 Suppl 3, S267-70. PMID: 21822704 DOI.
- ↑ 16.0 16.1 Hautier L, Weisbecker V, Sánchez-Villagra MR, Goswami A & Asher RJ. (2010). Skeletal development in sloths and the evolution of mammalian vertebral patterning. Proc. Natl. Acad. Sci. U.S.A. , 107, 18903-8. PMID: 20956304 DOI.
- ↑ Abitbol MM. (1987). Evolution of the sacrum in hominoids. Am. J. Phys. Anthropol. , 74, 65-81. PMID: 3688211 DOI.
- ↑ 18.0 18.1 18.2 Sohn P, Cox M, Chen D & Serra R. (2010). Molecular profiling of the developing mouse axial skeleton: a role for Tgfbr2 in the development of the intervertebral disc. BMC Dev. Biol. , 10, 29. PMID: 20214815 DOI.
- ↑ 19.0 19.1 Risbud MV, Schaer TP & Shapiro IM. (2010). Toward an understanding of the role of notochordal cells in the adult intervertebral disc: from discord to accord. Dev. Dyn. , 239, 2141-8. PMID: 20568241 DOI.
- ↑ Plummer NW, Spicher K, Malphurs J, Akiyama H, Abramowitz J, Nürnberg B & Birnbaumer L. (2012). Development of the mammalian axial skeleton requires signaling through the Gα(i) subfamily of heterotrimeric G proteins. Proc. Natl. Acad. Sci. U.S.A. , 109, 21366-71. PMID: 23236180 DOI.
- ↑ Sadler TW. (2000). Embryology of the sternum. Chest Surg. Clin. N. Am. , 10, 237-44, v. PMID: 10803330
- ↑ Mekonen HK, Hikspoors JP, Mommen G, Köhler SE & Lamers WH. (2015). Development of the ventral body wall in the human embryo. J. Anat. , 227, 673-85. PMID: 26467243 DOI.
- ↑ Guggenberger R, Andreisek G, Scheffel H, Wildermuth S, Leschka S & Stolzmann P. (2010). Absent cervical spine pedicle and associated congenital spinal abnormalities - a diagnostic trap in a setting of acute trauma: case report. BMC Med Imaging , 10, 25. PMID: 21062465 DOI.
- ↑ Postema SG, van der Sluis CK, Waldenlöv K & Norling Hermansson LM. (2012). Body structures and physical complaints in upper limb reduction deficiency: a 24-year follow-up study. PLoS ONE , 7, e49727. PMID: 23226218 DOI.
- ↑ Zaborowska-Sapeta K, Kowalski IM, Kotwicki T, Protasiewicz-Fałdowska H & Kiebzak W. (2011). Effectiveness of Chêneau brace treatment for idiopathic scoliosis: prospective study in 79 patients followed to skeletal maturity. Scoliosis , 6, 2. PMID: 21266084 DOI.
- ↑ Solomon BD. (2011). VACTERL/VATER Association. Orphanet J Rare Dis , 6, 56. PMID: 21846383 DOI.
Reviews
Pang D & Thompson DN. (2011). Embryology and bony malformations of the craniovertebral junction. Childs Nerv Syst , 27, 523-64. PMID: 21193993 DOI.
Iimura T, Denans N & Pourquié O. (2009). Establishment of Hox vertebral identities in the embryonic spine precursors. Curr. Top. Dev. Biol. , 88, 201-34. PMID: 19651306 DOI.
Alexander T, Nolte C & Krumlauf R. (2009). Hox genes and segmentation of the hindbrain and axial skeleton. Annu. Rev. Cell Dev. Biol. , 25, 431-56. PMID: 19575673 DOI.
Mallo M, Vinagre T & Carapuço M. (2009). The road to the vertebral formula. Int. J. Dev. Biol. , 53, 1469-81. PMID: 19247958 DOI.
Tyl RW, Chernoff N & Rogers JM. (2007). Altered axial skeletal development. Birth Defects Res. B Dev. Reprod. Toxicol. , 80, 451-72. PMID: 18157900 DOI.
Articles
Mead TJ & Yutzey KE. (2009). Notch pathway regulation of chondrocyte differentiation and proliferation during appendicular and axial skeleton development. Proc. Natl. Acad. Sci. U.S.A. , 106, 14420-5. PMID: 19590010 DOI.
Skuntz S, Mankoo B, Nguyen MT, Hustert E, Nakayama A, Tournier-Lasserve E, Wright CV, Pachnis V, Bharti K & Arnheiter H. (2009). Lack of the mesodermal homeodomain protein MEOX1 disrupts sclerotome polarity and leads to a remodeling of the cranio-cervical joints of the axial skeleton. Dev. Biol. , 332, 383-95. PMID: 19520072 DOI.
Wilson V, Olivera-Martinez I & Storey KG. (2009). Stem cells, signals and vertebrate body axis extension. Development , 136, 1591-604. PMID: 19395637 DOI.
Search PubMed
Search Pubmed: Axial Skeleton Development | Vertebra Development | Intervertebral Disc Development | Axial Skeleton Abnormalities
Additional Images
Historic
Human Embryology And Morphology (1921) |
---|
Keith, A. Human Embryology And Morphology (1921) Longmans, Green & Co.:New York.
|
Terms
- centrum - (vertebral body) anatomical term referring to the main bony part of the vertebra that forms the majority of the axial skeleton.
- Cobb angle - clinical method of measuring the degree of scoliosis and post-traumatic kyphosis. Named after the American orthopedic surgeon John Robert Cobb (1903 - 1967) an American orthopedic surgeon.
- haemal arch - referring to the bony arch region within animal tail vertebra that contains blood vessels.
- idiopathic scoliosis - clinical term for a spinal column deformity appearing usually postnatally after the age of 10 years old.
- lordosis - clinical and anatomy term describing the curvature of the spine with the convexity toward the front. Normal to have lordosis in the cervical and lumbar regions of the spinal column.
- neural arch - referring to the bony arch region within vertebra that contains the spinal cord.
- notochord sheath - region surrounding the notochord. in teleost fish direct mineralization of this region, by intramembranous ossification, forms the initial vertebral centrum. (More? zebrafish)
- rib hump - clinical term for the prominence formed by ribs on the convexity of a curve, caused by rotation of the spine and attached ribs.
- spinal column - term referring to the musculoskeletal elements (vertebrae, ligaments, and intervertebral discs), that surround the spinal cord and form the axial skeleton.
- synsacrum - in birds sacral and lumbar vertebrae fused region forming elongated sacral region.
- thoracic hypokyphosis - term referring to the backward curve in the upper spine is to great, round back, Scheuermann's disease, or simply kyphosis.
Terms
Bone Terms | ||
---|---|---|
Bone Development
| ||
|
External Links
External Links Notice - The dynamic nature of the internet may mean that some of these listed links may no longer function. If the link no longer works search the web with the link text or name. Links to any external commercial sites are provided for information purposes only and should never be considered an endorsement. UNSW Embryology is provided as an educational resource with no clinical information or commercial affiliation.
- Scoliosis Australia
- Scoliosis - the official journal of the Society on Scoliosis Orthopaedic and Rehabilitation Treatment (SOSORT) and affiliated with the International Research Society of Spinal Deformities (IRSSD).
- Young Women's Health - scoliosis
Glossary Links
- Glossary: A | B | C | D | E | F | G | H | I | J | K | L | M | N | O | P | Q | R | S | T | U | V | W | X | Y | Z | Numbers | Symbols | Term Link
Cite this page: Hill, M.A. (2024, June 14) Embryology Musculoskeletal System - Axial Skeleton Development. Retrieved from https://embryology.med.unsw.edu.au/embryology/index.php/Musculoskeletal_System_-_Axial_Skeleton_Development
- © Dr Mark Hill 2024, UNSW Embryology ISBN: 978 0 7334 2609 4 - UNSW CRICOS Provider Code No. 00098G