Musculoskeletal System - Shoulder Development - Revision history

Z8600021 at 04:04, 10 April 2019

2019-04-10T04:04:35Z

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	{{Shoulder Timeline table}}		{{Shoulder Timeline table}}

			{{Shoulder Timeline table2}}


	Acromial secondary ossification centers began appear at age 10 and clavicular ones, while uncommon, began forming at age 11. Fusion of acromial primary and secondary ossification centers began at age 14 and was generally complete after age 16.{{#pmid:25319561\|PMID25319561}} A similar time course has been shown in Japanese subjects.{{#pmid:26321557\|PMID26321557}}		Acromial secondary ossification centers began appear at age 10 and clavicular ones, while uncommon, began forming at age 11. Fusion of acromial primary and secondary ossification centers began at age 14 and was generally complete after age 16.{{#pmid:25319561\|PMID25319561}} A similar time course has been shown in Japanese subjects.{{#pmid:26321557\|PMID26321557}}


			:'''Links:''' {{Upper limb ossification timeline}}
	==Embryonic Shoulder==		==Embryonic Shoulder==
	{\|		{\|

Z8600021 at 03:59, 10 April 2019

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	* Japanese subjects - acromion secondary ossification ~~centres~~ appear at ~~around~~ 10 ~~years~~ of age and ~~had fused at 15 to~~ 16 ~~years~~.{{#pmid:26321557\|PMID26321557}}		Acromial secondary ossification centers began appear at age 10 and clavicular ones, while uncommon, began forming at age 11. Fusion of acromial primary and secondary ossification centers began at age 14 and was generally complete after age 16.{{#pmid:25319561\|PMID25319561}} A similar time course has been shown in Japanese subjects.{{#pmid:26321557\|PMID26321557}}
	==Embryonic Shoulder==		==Embryonic Shoulder==
	{\|		{\|

Z8600021 at 03:10, 10 April 2019

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	{{Shoulder Timeline table}}		{{Shoulder Timeline table}}


			* Japanese subjects - acromion secondary ossification centres appear at around 10 years of age and had fused at 15 to 16 years.{{#pmid:26321557\|PMID26321557}}
	==Embryonic Shoulder==		==Embryonic Shoulder==
	{\|		{\|

Z8600021 at 03:03, 10 April 2019

2019-04-10T03:03:42Z

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	* '''~~Scapula development is governed by genetic interactions~~ of ~~Pbx1 with its family members~~ and ~~with Emx2 via their cooperative control of Alx1~~'''{{#pmid:~~20627960~~\|~~PMID20627960~~}} "~~The~~ genetic pathways ~~underlying shoulder blade~~ development ~~are largely unknown~~, ~~as gene~~ networks controlling ~~limb morphogenesis have limited influence on~~ scapula ~~formation~~. ~~Analysis~~ of ~~mouse mutants for Pbx~~ and ~~Emx2 genes has suggested their potential roles~~ in girdle development. In this study, ~~by generating compound mutant mice~~, ~~we examined~~ the ~~genetic control~~ of ~~scapula development by Pbx genes~~ and ~~their functional relationship with Emx2~~. ~~Analyses~~ of ~~Pbx~~ and ~~Pbx1;Emx2 compound mutants revealed that Pbx genes share overlapping functions in shoulder development~~ and that ~~Pbx1 genetically interacts with Emx2~~ in ~~this~~ process. ~~Here~~, ~~we provide a biochemical basis~~ for ~~Pbx1;Emx2 genetic interaction by showing that Pbx1~~ and ~~Emx2 can bind specific DNA sequences as heterodimers~~. ~~Moreover~~, the ~~expression of genes crucial for~~ scapula ~~development is altered in these mutants~~, ~~indicating~~ that ~~Pbx genes act upstream of essential pathways for scapula formation~~. In ~~particular~~, ~~expression~~ of ~~Alx1~~, ~~an effector~~ of ~~scapula blade patterning~~, ~~is absent in all compound mutants~~. ~~We demonstrate~~ that ~~Pbx1~~ and ~~Emx2 bind in vivo~~ to a conserved ~~sequence upstream~~ of ~~Alx1 and cooperatively activate~~ its ~~transcription via this potential regulatory element~~. ~~Our results establish an essential role~~ for ~~Pbx1 in genetic interactions with its family members~~ and ~~with Emx2~~ and ~~delineate novel regulatory networks~~ in shoulder girdle development."		* '''Review - Genetics of scapula and pelvis development: An evolutionary perspective'''{{#pmid:30797513\|PMID30797513}} "In tetrapods, the scapular and pelvic girdles perform the important function of anchoring the limbs to the trunk of the body and facilitating the movement of each appendage. This shared function, however, is one of relatively few similarities between the scapula and pelvis, which have significantly different morphologies, evolutionary histories, embryonic origins, and underlying genetic pathways. The scapula evolved in jawless fish prior to the pelvis, and its embryonic development is unique among bones in that it is derived from multiple progenitor cell populations, including the dermomyotome, somatopleure, and neural crest. Conversely, the pelvis evolved several million years later in jawed fish, and it develops from an embryonic somatopleuric cell population. The genetic networks controlling the formation of the pelvis and scapula also share similarities and differences, with a number of genes shaping only one or the other, while other gene products such as PBX transcription factors act as hierarchical developmental regulators of both girdle structures. Here, we provide a detailed review of the cellular processes and genetic networks underlying pelvis and scapula formation in tetrapods, while also highlighting unanswered questions about girdle evolution and development."

			* '''Congenital Anatomical Variant of the Clavicle'''{{#pmid:28296289\|PMID28296289}} "The aim of this study is to present a rare abnormality of the clavicle (Code: SGS01) that was discovered in an ossuary in the Church of San Gaetano (Sulmona, central Italy; XVII-XIX centuries CE). In the middle third, the clavicle had three areas with losses of substance in the form of oval-shaped foramina with maximum diameters of 1-2 cm that were located in the anterior and superior surfaces of the diaphysis. The margins of these foramina were well defined and rounded, and the surfaces of the canal walls were smooth. Additionally, there were no zones of bony activity or reactive changes around the foramina. This new congenital anomaly of the clavicle and blood vessels is consistent with a variant that might have originated during fetal growth in which the subclavian vein or artery remained included during the process of ossification of the clavicle."

			* '''Developmental origin of the clavicle, and its implications for the evolution of the neck and the paired appendages in vertebrates'''{{#pmid:27279028\|PMID27279028}} "In fish, the pectoral appendage is adjacent to the head, but during vertebrate evolution a long neck region emerged via caudal relocation of the pectoral appendage. The pectoral appendage is comprised of endochondral portions, such as the humerus and the scapula, and a dermal portion, such as the clavicle, that contributes to the shoulder girdle. In the search for clues to the mechanism of the caudal relocation of the pectoral appendage, the cell lineage of the rostral lateral plate mesoderm was analyzed in chickens. It was found that, despite the long neck region in chickens, the origin of the clavicle attached to the head mesoderm ranged between 1 and 14 somite levels. Because the pectoral limb bud and the endochondral pectoral appendage developed on 15-20 and 15-24 somite levels, respectively, the clavicle-forming region corresponds to the embryonic neck, which suggests that the relocation would have been executed by the expansion of the source of the clavicle. The rostral portion of the clavicle-forming region overlaps the source of the cucullaris muscle, embraces the pharyngeal arches caudally, and can be experimentally replaced with the head mesoderm to form the cucullaris muscle, which implies that the mesodermal portion could have been the head mesoderm and that the clavicle would have developed at the head/trunk boundary. The link between the head mesoderm and the presumptive clavicle appears to have been the developmental constraint needed to create the evolutionarily conserved musculoskeletal connectivities characterizing the gnathostome neck. In this sense, the dermal girdle of the ganathostomes would represent the wall of the branchial chamber into which the endochondral pectoral appendage appears to have attached since its appearance in evolution."

			* '''Development of the shoulder girdle musculature'''{{#pmid:26676088\|PMID26676088}} "The muscles of the shoulder region are important for movements of the upper limbs and for stabilizing the girdle elements by connecting them to the trunk. They have a triple embryonic origin. First, the branchiomeric shoulder girdle muscles (sternocleidomastoideus and trapezius muscles) develop from the occipital lateral plate mesoderm using Tbx1 over the course of this development. The second population of cells constitutes the superficial shoulder girdle muscles (pectoral and latissimus dorsi muscles), which are derived from the wing premuscle mass. This muscle group undergoes a two-step development, referred to as the "in-out" mechanism. Myogenic precursor cells first migrate anterogradely into the wing bud. Subsequently, they migrate in a retrograde manner from the wing premuscle mass to the trunk. SDF-1/CXCR4 signaling is involved in this outward migration. A third group of shoulder muscles are the rhomboidei and serratus anterior muscles, which are referred to as deep shoulder girdle muscles; they are thought to be derived from the myotomes. It is, however, not clear how myotome cells make contact to the scapula to form these two muscles. In this review, we discuss the development of the shoulder girdle muscle in relation to the different muscle groups."
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	\| {{Older papers}}		\| {{Older papers}}
			* '''Scapula development is governed by genetic interactions of Pbx1 with its family members and with Emx2 via their cooperative control of Alx1'''{{#pmid:20627960\|PMID20627960}} "The genetic pathways underlying shoulder blade development are largely unknown, as gene networks controlling limb morphogenesis have limited influence on scapula formation. Analysis of mouse mutants for Pbx and Emx2 genes has suggested their potential roles in girdle development. In this study, by generating compound mutant mice, we examined the genetic control of scapula development by Pbx genes and their functional relationship with Emx2. Analyses of Pbx and Pbx1;Emx2 compound mutants revealed that Pbx genes share overlapping functions in shoulder development and that Pbx1 genetically interacts with Emx2 in this process. Here, we provide a biochemical basis for Pbx1;Emx2 genetic interaction by showing that Pbx1 and Emx2 can bind specific DNA sequences as heterodimers. Moreover, the expression of genes crucial for scapula development is altered in these mutants, indicating that Pbx genes act upstream of essential pathways for scapula formation. In particular, expression of Alx1, an effector of scapula blade patterning, is absent in all compound mutants. We demonstrate that Pbx1 and Emx2 bind in vivo to a conserved sequence upstream of Alx1 and cooperatively activate its transcription via this potential regulatory element. Our results establish an essential role for Pbx1 in genetic interactions with its family members and with Emx2 and delineate novel regulatory networks in shoulder girdle development."
	\|}		\|}
	== Textbooks ==		== Textbooks ==

Z8600021 at 02:27, 10 April 2019

2019-04-10T02:27:32Z

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	Search term: [http://www.ncbi.nlm.nih.gov/pubmed/?term=Shoulder+Development ''Shoulder Development''] \| [http://www.ncbi.nlm.nih.gov/pubmed/?term=Scapula+Development ''Scapula Development''] \| [http://www.ncbi.nlm.nih.gov/pubmed/?term=Clavicle+Development ''Clavicle Development'']		Search term: [http://www.ncbi.nlm.nih.gov/pubmed/?term=Shoulder+Development ''Shoulder Development''] \| [http://www.ncbi.nlm.nih.gov/pubmed/?term=Scapula+Development ''Scapula Development''] \| [http://www.ncbi.nlm.nih.gov/pubmed/?term=Clavicle+Development ''Clavicle Development'']

			\|}
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			! Older papers
			\|-
			\| {{Older papers}}
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	== Textbooks ==		== Textbooks ==

Z8600021 at 02:26, 10 April 2019

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	\| [[File:Mark_Hill.jpg\|90px\|left]] {{Most_Recent_Refs}}		\| [[File:Mark_Hill.jpg\|90px\|left]] {{Most_Recent_Refs}}

	Search term: [http://www.ncbi.nlm.nih.gov/pubmed/?term=Shoulder+Development ''Shoulder Development''] \| [http://www.ncbi.nlm.nih.gov/pubmed/?term=Scapula+Development ''Scapula Development'']		Search term: [http://www.ncbi.nlm.nih.gov/pubmed/?term=Shoulder+Development ''Shoulder Development''] \| [http://www.ncbi.nlm.nih.gov/pubmed/?term=Scapula+Development ''Scapula Development''] \| [http://www.ncbi.nlm.nih.gov/pubmed/?term=Clavicle+Development ''Clavicle Development'']

	\|}		\|}

Z8600021 at 02:25, 10 April 2019

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	\| [[File:Mark_Hill.jpg\|90px\|left]] {{Most_Recent_Refs}}		\| [[File:Mark_Hill.jpg\|90px\|left]] {{Most_Recent_Refs}}

	Search term: [http://www.ncbi.nlm.nih.gov/pubmed/?term=Shoulder+Development ''Shoulder Development'']		Search term: [http://www.ncbi.nlm.nih.gov/pubmed/?term=Shoulder+Development ''Shoulder Development''] \| [http://www.ncbi.nlm.nih.gov/pubmed/?term=Scapula+Development ''Scapula Development'']

	~~<pubmed limit=5>Shoulder Development</pubmed>~~
	\|}		\|}
	== Textbooks ==		== Textbooks ==

Z8600021 at 02:13, 11 March 2018

2018-03-11T02:13:23Z

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	[[File:Stage20-23_limbs_b.jpg\|500px\|right]]		[[File:Stage20-23_limbs_b.jpg\|500px\|right]]
	== Introduction ==		== Introduction ==
			[[File:Keith1902 fig245.jpg\|thumb\|Developing shoulder skeleton]]
	[[File:Shoulder cartoon.jpg\|thumb\|right\|Adult shoulder skeleton]]		[[File:Shoulder cartoon.jpg\|thumb\|right\|Adult shoulder skeleton]]
	[[File:Appendicular_skeleton.jpg\|thumb\|Adult appendicular skeleton]]		[[File:Appendicular_skeleton.jpg\|thumb\|Adult appendicular skeleton]]

Z8600021 at 02:11, 11 March 2018

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	[[File:Shoulder cartoon.jpg\|thumb\|right\|Adult shoulder skeleton]]		[[File:Shoulder cartoon.jpg\|thumb\|right\|Adult shoulder skeleton]]
	[[File:Appendicular_skeleton.jpg\|thumb\|Adult appendicular skeleton]]		[[File:Appendicular_skeleton.jpg\|thumb\|Adult appendicular skeleton]]
	[[File:Limb bud geometry and patterning.jpg\|thumb\|Limb bud geometry and patterning~~<ref name="PMID20644713"><pubmed>~~20644713~~</pubmed>~~\| ~~[http://www.ncbi.nlm.nih.gov/pmc/articles/PMC2903596/ PMC2903596] \| [http://www.plosbiology.org/article/info%3Adoi%2F10.1371%2Fjournal.pbio.1000421 PLoS]</ref>~~]]		[[File:Limb bud geometry and patterning.jpg\|thumb\|Limb bud geometry and patterning{{#pmid:20644713\|PMID20644713}}]]

	The skeletal shoulder consists of: the clavicle (collarbone), the scapula (shoulder blade), and the humerus. Development of his region occurs through both forms of ossification processes.		The skeletal shoulder consists of: the clavicle (collarbone), the scapula (shoulder blade), and the humerus. Development of his region occurs through both forms of ossification processes.
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	* '''Scapula development is governed by genetic interactions of Pbx1 with its family members and with Emx2 via their cooperative control of Alx1'''~~<ref><pubmed>~~20627960~~</pubmed></ref>~~ "The genetic pathways underlying shoulder blade development are largely unknown, as gene networks controlling limb morphogenesis have limited influence on scapula formation. Analysis of mouse mutants for Pbx and Emx2 genes has suggested their potential roles in girdle development. In this study, by generating compound mutant mice, we examined the genetic control of scapula development by Pbx genes and their functional relationship with Emx2. Analyses of Pbx and Pbx1;Emx2 compound mutants revealed that Pbx genes share overlapping functions in shoulder development and that Pbx1 genetically interacts with Emx2 in this process. Here, we provide a biochemical basis for Pbx1;Emx2 genetic interaction by showing that Pbx1 and Emx2 can bind specific DNA sequences as heterodimers. Moreover, the expression of genes crucial for scapula development is altered in these mutants, indicating that Pbx genes act upstream of essential pathways for scapula formation. In particular, expression of Alx1, an effector of scapula blade patterning, is absent in all compound mutants. We demonstrate that Pbx1 and Emx2 bind in vivo to a conserved sequence upstream of Alx1 and cooperatively activate its transcription via this potential regulatory element. Our results establish an essential role for Pbx1 in genetic interactions with its family members and with Emx2 and delineate novel regulatory networks in shoulder girdle development."		* '''Scapula development is governed by genetic interactions of Pbx1 with its family members and with Emx2 via their cooperative control of Alx1'''{{#pmid:20627960\|PMID20627960}} "The genetic pathways underlying shoulder blade development are largely unknown, as gene networks controlling limb morphogenesis have limited influence on scapula formation. Analysis of mouse mutants for Pbx and Emx2 genes has suggested their potential roles in girdle development. In this study, by generating compound mutant mice, we examined the genetic control of scapula development by Pbx genes and their functional relationship with Emx2. Analyses of Pbx and Pbx1;Emx2 compound mutants revealed that Pbx genes share overlapping functions in shoulder development and that Pbx1 genetically interacts with Emx2 in this process. Here, we provide a biochemical basis for Pbx1;Emx2 genetic interaction by showing that Pbx1 and Emx2 can bind specific DNA sequences as heterodimers. Moreover, the expression of genes crucial for scapula development is altered in these mutants, indicating that Pbx genes act upstream of essential pathways for scapula formation. In particular, expression of Alx1, an effector of scapula blade patterning, is absent in all compound mutants. We demonstrate that Pbx1 and Emx2 bind in vivo to a conserved sequence upstream of Alx1 and cooperatively activate its transcription via this potential regulatory element. Our results establish an essential role for Pbx1 in genetic interactions with its family members and with Emx2 and delineate novel regulatory networks in shoulder girdle development."
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	* '''Before we Are Born''' (5th ed.) Moore and Persaud Chapter 16,17: p379-397, 399-405		* '''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		* '''Essentials of Human Embryology''' Larson Chapter 11 p207-228

			==Timeline==

			{{Shoulder Timeline table}}

	==Embryonic Shoulder==		==Embryonic Shoulder==
	{\|		{\|
	\| [[File:Human embryonic shoulder girdle 01.jpg\|600px]]		\| [[File:Human embryonic shoulder girdle 01.jpg\|600px]]
	\| Drawing of a model of the right shoulder girdle of the 17mm. (Robinson) embryo, viewed from behind.Human embryonic shoulder girdle (17mm CRL) ~~<ref name="PMID17232952"><pubmed>~~17232952~~</pubmed>~~\| ~~[http://www.ncbi.nlm.nih.gov/pmc/articles/PMC1289013 PMC1289013]</ref>~~		\| Drawing of a model of the right shoulder girdle of the 17mm. (Robinson) embryo, viewed from behind.Human embryonic shoulder girdle (17mm CRL){{#pmid:17232952\|PMID17232952}}

	The sternal segment (S.S.) has been cut coronally to expose the interior, and the acromial segment has been cut horizontally for the same purpose.		The sternal segment (S.S.) has been cut coronally to expose the interior, and the acromial segment has been cut horizontally for the same purpose.
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	* S. S. - sternal segment of clavicle		* S. S. - sternal segment of clavicle
	\|-		\|-
	\| Human embryonic shoulder girdle (17mm CRL) ~~<ref name="PMID17232952"><pubmed>~~17232952~~</pubmed>~~\| ~~[http://www.ncbi.nlm.nih.gov/pmc/articles/PMC1289013 PMC1289013]</ref>~~		\| Human embryonic shoulder girdle (17mm CRL){{#pmid:17232952\|PMID17232952}}
	\| '''17mm CRL (Robinson) embryo'''		\| '''17mm CRL (Robinson) embryo'''
	* Week 7 - difficult to estimate the amount of shrinkage in this historic material.		* Week 7 - difficult to estimate the amount of shrinkage in this historic material.
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	* Final differentiation of these cells into post-mitotic muscle fibers in the limb bud is regulated by another myogenic determination factor, Myogenin.		* Final differentiation of these cells into post-mitotic muscle fibers in the limb bud is regulated by another myogenic determination factor, Myogenin.

	Some text modified from~~<ref><pubmed>~~17592144~~</pubmed></ref>~~		Some text modified from{{#pmid:17592144\|PMID17592144}}

	'''Links:''' [[Musculoskeletal_System_-_Muscle_Development\|Muscle Development]]		'''Links:''' [[Musculoskeletal_System_-_Muscle_Development\|Muscle Development]]

	==Limb Bone==		==Limb Bone==
	[[File:Chicken-limb_sox9_wnt6.jpg\|thumb\|Limb sox9 and Wnt6 expression~~<ref name="~~PMID20334703~~"><pubmed>20334703</pubmed></ref>~~]]		[[File:Chicken-limb_sox9_wnt6.jpg\|thumb\|Limb sox9 and Wnt6 expression{{#pmid:20334703\|PMID20334703}}]]
	[[File:Chicken- limb bud chondrogenesis.jpg\|thumb\|Chicken- limb bud chondrogenesis]]		[[File:Chicken- limb bud chondrogenesis.jpg\|thumb\|Chicken- limb bud chondrogenesis]]
	Bone formation within the limb occurs by endochondral ossification of a pre-existing cartilage template. Ossification then replaces the existing cartilage except in the regions of articulation, where cartilage remains on the surface of the bone within the joint. Therefore bone development in the limb is initially about cartilage development or chondrogenesis.		Bone formation within the limb occurs by endochondral ossification of a pre-existing cartilage template. Ossification then replaces the existing cartilage except in the regions of articulation, where cartilage remains on the surface of the bone within the joint. Therefore bone development in the limb is initially about cartilage development or chondrogenesis.
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	# '''Chondrogenesis''' - the differentiation of mesoderm to form cartilage, which will be essentially the same program for all cartilage templates.		# '''Chondrogenesis''' - the differentiation of mesoderm to form cartilage, which will be essentially the same program for all cartilage templates.

	A recent study has identified that the overlying limb surface ectoderm potentially inhibits limb early chondrogenesis through Wnt6 signaling.~~<ref name="~~PMID20334703~~" />~~		A recent study has identified that the overlying limb surface ectoderm potentially inhibits limb early chondrogenesis through Wnt6 signaling.{{#pmid:20334703\|PMID20334703}}


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	===Postnatal Growth===		===Postnatal Growth===

	A ~~recent~~ paper has characterised the postnatal growth of male and female clavicles (data shown below).~~<ref><pubmed>~~19934713~~</pubmed></ref>~~		A paper has characterised the postnatal growth of male and female clavicles (data shown below).{{#pmid:19934713\|PMID19934713}}

	* 18 years of age the mean clavicle length +/-SD for females was 149+/-12 mm and for males it was 161+/-11 mm.		* 18 years of age the mean clavicle length +/-SD for females was 149+/-12 mm and for males it was 161+/-11 mm.
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	[[File:Gray0321.jpg\|thumb\|Hip bone]]		[[File:Gray0321.jpg\|thumb\|Hip bone]]

	The skeletal pelvis consists of: the sacrum and coccyx (axial skeleton), and pelvic girdle formed by a pair of hip bones (appendicular skeleton). Before puberty, he pelvic girdle also consists of three unfused bones: the ilium, ischium, and pubis. In chicken, the entire pelvic girdle originates from the somatopleure mesoderm (somite levels 26 to 35) and the ilium, but not of the pubis and ischium, depends on somitic and ectodermal signals.~~<ref><pubmed>~~18087724~~</pubmed></ref>~~		The skeletal pelvis consists of: the sacrum and coccyx (axial skeleton), and pelvic girdle formed by a pair of hip bones (appendicular skeleton). Before puberty, he pelvic girdle also consists of three unfused bones: the ilium, ischium, and pubis. In chicken, the entire pelvic girdle originates from the somatopleure mesoderm (somite levels 26 to 35) and the ilium, but not of the pubis and ischium, depends on somitic and ectodermal signals.{{#pmid:18087724\|PMID18087724}}


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	===Fibroblast Growth Factors===		===Fibroblast Growth Factors===

	* Fgf8 - morphogen gradient forms by a source-sink mechanism with freely diffusing molecules.~~<ref><pubmed>~~19741606~~</pubmed></ref>~~		* Fgf8 - morphogen gradient forms by a source-sink mechanism with freely diffusing molecules.{{#pmid:19741606\|PMID19741606}}

	===T-box Transcription Factors===		===T-box Transcription Factors===
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	===Reviews===		===Reviews===
			{{#pmid:17006658}}

			{{#pmid:11170011}}

	~~<pubmed>17006658</pubmed>~~		{{#pmid:3324657}}
	~~<pubmed>11170011</pubmed>~~
	~~<pubmed>~~3324657~~</pubmed>~~

	===Articles===		===Articles===
			{{#pmid:19763557}}

			{{#pmid:9272998}}

	~~<pubmed>19763557</pubmed>~~		{{#pmid:531584}}
	~~<pubmed>9272998</pubmed>~~
	~~<pubmed>~~531584~~</pubmed>~~

	===Search PubMed===		===Search PubMed===
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	{{Footer}}		{{Footer}}
	[[Category:Limb]][[Category:Bone]]		[[Category:Limb]][[Category:Bone]][[Category:Joint]]

Z8600021 at 04:52, 22 Mayıs 2017

2017-05-22T04:52:07Z

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	* '''Scapula development is governed by genetic interactions of Pbx1 with its family members and with Emx2 via their cooperative control of Alx1'''<ref><pubmed>20627960</pubmed></ref> "The genetic pathways underlying shoulder blade development are largely unknown, as gene networks controlling limb morphogenesis have limited influence on scapula formation. Analysis of mouse mutants for Pbx and Emx2 genes has suggested their potential roles in girdle development. In this study, by generating compound mutant mice, we examined the genetic control of scapula development by Pbx genes and their functional relationship with Emx2. Analyses of Pbx and Pbx1;Emx2 compound mutants revealed that Pbx genes share overlapping functions in shoulder development and that Pbx1 genetically interacts with Emx2 in this process. Here, we provide a biochemical basis for Pbx1;Emx2 genetic interaction by showing that Pbx1 and Emx2 can bind specific DNA sequences as heterodimers. Moreover, the expression of genes crucial for scapula development is altered in these mutants, indicating that Pbx genes act upstream of essential pathways for scapula formation. In particular, expression of Alx1, an effector of scapula blade patterning, is absent in all compound mutants. We demonstrate that Pbx1 and Emx2 bind in vivo to a conserved sequence upstream of Alx1 and cooperatively activate its transcription via this potential regulatory element. Our results establish an essential role for Pbx1 in genetic interactions with its family members and with Emx2 and delineate novel regulatory networks in shoulder girdle development."		* '''Scapula development is governed by genetic interactions of Pbx1 with its family members and with Emx2 via their cooperative control of Alx1'''<ref><pubmed>20627960</pubmed></ref> "The genetic pathways underlying shoulder blade development are largely unknown, as gene networks controlling limb morphogenesis have limited influence on scapula formation. Analysis of mouse mutants for Pbx and Emx2 genes has suggested their potential roles in girdle development. In this study, by generating compound mutant mice, we examined the genetic control of scapula development by Pbx genes and their functional relationship with Emx2. Analyses of Pbx and Pbx1;Emx2 compound mutants revealed that Pbx genes share overlapping functions in shoulder development and that Pbx1 genetically interacts with Emx2 in this process. Here, we provide a biochemical basis for Pbx1;Emx2 genetic interaction by showing that Pbx1 and Emx2 can bind specific DNA sequences as heterodimers. Moreover, the expression of genes crucial for scapula development is altered in these mutants, indicating that Pbx genes act upstream of essential pathways for scapula formation. In particular, expression of Alx1, an effector of scapula blade patterning, is absent in all compound mutants. We demonstrate that Pbx1 and Emx2 bind in vivo to a conserved sequence upstream of Alx1 and cooperatively activate its transcription via this potential regulatory element. Our results establish an essential role for Pbx1 in genetic interactions with its family members and with Emx2 and delineate novel regulatory networks in shoulder girdle development."
	\|}		\|}
			{\| 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=Shoulder+Development ''Shoulder Development'']

			<pubmed limit=5>Shoulder Development</pubmed>
			\|}
	== Textbooks ==		== 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		* '''The Developing Human: Clinically Oriented Embryology''' (8th Edition) by Keith L. Moore and T.V.N Persaud - Moore & Persaud Chapter 15 the skeletal system

← Older revision		Revision as of 12:13, 11 March 2018
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	[[File:Stage20-23_limbs_b.jpg\|500px\|right]]		[[File:Stage20-23_limbs_b.jpg\|500px\|right]]
	== Introduction ==		== Introduction ==
			[[File:Keith1902 fig245.jpg\|thumb\|Developing shoulder skeleton]]
	[[File:Shoulder cartoon.jpg\|thumb\|right\|Adult shoulder skeleton]]		[[File:Shoulder cartoon.jpg\|thumb\|right\|Adult shoulder skeleton]]
	[[File:Appendicular_skeleton.jpg\|thumb\|Adult appendicular skeleton]]		[[File:Appendicular_skeleton.jpg\|thumb\|Adult appendicular skeleton]]