Renal System Development - Revision history

Z8600021 at 23:42, 10 August 2020

2020-08-10T23:42:29Z

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	== References ==		== References ==
	<references/>		<references/>

			===Reviews===

			{{#pmid:32765290}}

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 |}
 ==Embryonic Kidney==
 Embryonic stage descriptions based on [[Carnegie Collection]] embryos.<ref name=O'RahillyMüller1987>{{Ref-O'RahillyMüller1987}}</ref>

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	* '''Spatiotemporal heterogeneity and patterning of developing renal blood vessels'''{{#pmid:29627966\|PMID29627966}} "Here, we examine the developing kidney vasculature to assess its 3-dimensional structure and transcriptional heterogeneity. First, we observe that endothelial cells (ECs) grow coordinately with the kidney bud as early as {{ME10.5}}, and begin to show signs of specification by {{ME13.5}} when the first arteries can be identified. We then focus on how ECs pattern and remodel with respect to the developing nephron and collecting duct epithelia. ECs circumscribe nephron progenitor populations at the distal tips of the ureteric bud (UB) tree and form stereotyped cruciform structures around each tip. Beginning at the renal vesicle (RV) stage, ECs form a continuous plexus around developing nephrons. The endothelial plexus envelops and elaborates with the maturing nephron, becoming preferentially enriched along the early distal tubule."		* '''Spatiotemporal heterogeneity and patterning of developing renal blood vessels'''{{#pmid:29627966\|PMID29627966}} "Here, we examine the developing kidney vasculature to assess its 3-dimensional structure and transcriptional heterogeneity. First, we observe that endothelial cells (ECs) grow coordinately with the kidney bud as early as {{ME10.5}}, and begin to show signs of specification by {{ME13.5}} when the first arteries can be identified. We then focus on how ECs pattern and remodel with respect to the developing nephron and collecting duct epithelia. ECs circumscribe nephron progenitor populations at the distal tips of the ureteric bud (UB) tree and form stereotyped cruciform structures around each tip. Beginning at the renal vesicle (RV) stage, ECs form a continuous plexus around developing nephrons. The endothelial plexus envelops and elaborates with the maturing nephron, becoming preferentially enriched along the early distal tubule."

	* '''Conserved and Divergent Features of Human and Mouse Kidney Organogenesis'''{{#pmid:29449453\|PMID29449453}} "Human kidney function is underpinned by approximately 1,000,000 nephrons, although the number varies substantially, and low nephron number is linked to disease. Human kidney development initiates around 4 weeks of gestation and ends around 34-37 weeks of gestation. Over this period, a reiterative inductive process establishes the nephron complement. Studies have provided insightful anatomic descriptions of human kidney development, but the limited histologic views are not readily accessible to a broad audience. In this first paper in a series providing comprehensive insight into human kidney formation, we examined human kidney development in 135 anonymously donated human kidney specimens. We documented kidney development at a macroscopic and cellular level through histologic analysis, RNA in situ hybridization, immunofluorescence studies, and transcriptional profiling, contrasting human development (4-23 weeks) with {{mouse}} development at selected stages (embryonic day {{ME15.5}} and postnatal day 2). The high-resolution histologic interactive atlas of human kidney organogenesis generated can be viewed at the GUDMAP database (www.gudmap.org) together with three-dimensional reconstructions of key components of the data herein. At the anatomic level, human and mouse kidney development differ in timing, scale, and global features such as lobe formation and progenitor niche organization. The data also highlight differences in molecular and cellular features, including the expression and cellular distribution of anchor gene markers used to identify key cell types in mouse kidney studies." {{mouse}}
	\|}		\|}
	{\| class="wikitable mw-collapsible mw-collapsed"		{\| class="wikitable mw-collapsible mw-collapsed"
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	\|-		\|-
	\| {{Older papers}}		\| {{Older papers}}
	* '''Reciprocal Spatiotemporally Controlled Apoptosis Regulates Wolffian Duct Cloaca Fusion'''{{#pmid:29326158\|PMID29326158}} "The epithelial Wolffian duct (WD) inserts into the cloaca (primitive bladder) before metanephric kidney development, thereby establishing the initial plumbing for eventual joining of the ureters and bladder. Defects in this process cause common anomalies in the spectrum of congenital anomalies of the kidney and urinary tract (CAKUT). However, developmental, cellular, and molecular mechanisms of WD-cloaca fusion are poorly understood. Through systematic analysis of early WD tip development in mice, we discovered that a novel process of spatiotemporally regulated apoptosis in WD and cloaca was necessary for WD-cloaca fusion."		* '''Conserved and Divergent Features of Human and Mouse Kidney Organogenesis'''{{#pmid:29449453\|PMID29449453}} "Human kidney function is underpinned by approximately 1,000,000 nephrons, although the number varies substantially, and low nephron number is linked to disease. Human kidney development initiates around 4 weeks of gestation and ends around 34-37 weeks of gestation. Over this period, a reiterative inductive process establishes the nephron complement. Studies have provided insightful anatomic descriptions of human kidney development, but the limited histologic views are not readily accessible to a broad audience. In this first paper in a series providing comprehensive insight into human kidney formation, we examined human kidney development in 135 anonymously donated human kidney specimens. We documented kidney development at a macroscopic and cellular level through histologic analysis, RNA in situ hybridization, immunofluorescence studies, and transcriptional profiling, contrasting human development (4-23 weeks) with {{mouse}} development at selected stages (embryonic day {{ME15.5}} and postnatal day 2). The high-resolution histologic interactive atlas of human kidney organogenesis generated can be viewed at the GUDMAP database (www.gudmap.org) together with three-dimensional reconstructions of key components of the data herein. At the anatomic level, human and mouse kidney development differ in timing, scale, and global features such as lobe formation and progenitor niche organization. The data also highlight differences in molecular and cellular features, including the expression and cellular distribution of anchor gene markers used to identify key cell types in mouse kidney studies." {{mouse}}

			* '''Reciprocal Spatiotemporally Controlled {{Apoptosis}} Regulates Wolffian Duct Cloaca Fusion'''{{#pmid:29326158\|PMID29326158}} "The epithelial Wolffian duct (WD) inserts into the cloaca (primitive bladder) before metanephric kidney development, thereby establishing the initial plumbing for eventual joining of the ureters and bladder. Defects in this process cause common anomalies in the spectrum of congenital anomalies of the kidney and urinary tract (CAKUT). However, developmental, cellular, and molecular mechanisms of WD-cloaca fusion are poorly understood. Through systematic analysis of early WD tip development in mice, we discovered that a novel process of spatiotemporally regulated apoptosis in WD and cloaca was necessary for WD-cloaca fusion."

	* '''{{Zebrafish}} Pronephros Development'''{{#pmid:28409341\|PMID28409341}} "The pronephros is the first kidney type to form in vertebrate embryos. The first step of pronephrogenesis in the zebrafish is the formation of the intermediate mesoderm during gastrulation, which occurs in response to secreted morphogens such as BMPs and Nodals. Patterning of the intermediate mesoderm into proximal and distal cell fates is induced by retinoic acid signaling with downstream transcription factors including wt1a, pax2a, pax8, hnf1b, sim1a, mecom, and irx3b. In the anterior intermediate mesoderm, progenitors of the glomerular blood filter migrate and fuse at the midline and recruit a blood supply. More posteriorly localized tubule progenitors undergo epithelialization and fuse with the cloaca. The Notch signaling pathway regulates the formation of multi-ciliated cells in the tubules and these cells help propel the filtrate to the cloaca. The lumenal sheer stress caused by flow down the tubule activates anterior collective migration of the proximal tubules and induces stretching and proliferation of the more distal segments. Ultimately these processes create a simple two-nephron kidney that is capable of reabsorbing and secreting solutes and expelling excess water-processes that are critical to the homeostasis of the body fluids. The zebrafish pronephric kidney provides a simple, yet powerful, model system to better understand the conserved molecular and cellular progresses that drive nephron formation, structure, and function." {{Zebrafish}}		* '''{{Zebrafish}} Pronephros Development'''{{#pmid:28409341\|PMID28409341}} "The pronephros is the first kidney type to form in vertebrate embryos. The first step of pronephrogenesis in the zebrafish is the formation of the intermediate mesoderm during gastrulation, which occurs in response to secreted morphogens such as BMPs and Nodals. Patterning of the intermediate mesoderm into proximal and distal cell fates is induced by retinoic acid signaling with downstream transcription factors including wt1a, pax2a, pax8, hnf1b, sim1a, mecom, and irx3b. In the anterior intermediate mesoderm, progenitors of the glomerular blood filter migrate and fuse at the midline and recruit a blood supply. More posteriorly localized tubule progenitors undergo epithelialization and fuse with the cloaca. The Notch signaling pathway regulates the formation of multi-ciliated cells in the tubules and these cells help propel the filtrate to the cloaca. The lumenal sheer stress caused by flow down the tubule activates anterior collective migration of the proximal tubules and induces stretching and proliferation of the more distal segments. Ultimately these processes create a simple two-nephron kidney that is capable of reabsorbing and secreting solutes and expelling excess water-processes that are critical to the homeostasis of the body fluids. The zebrafish pronephric kidney provides a simple, yet powerful, model system to better understand the conserved molecular and cellular progresses that drive nephron formation, structure, and function." {{Zebrafish}}
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	* '''Histone deacetylase 1 and 2 regulate Wnt and p53 pathways in the ureteric bud epithelium'''{{#pmid:25758227\|PMID25758227}} "Histone deacetylases (HDACs) regulate a broad range of biological processes through removal of acetyl groups from histones as well as non-histone proteins. Our previous studies showed that Hdac1 and Hdac2 are bound to promoters of key renal developmental regulators and that HDAC activity is required for embryonic kidney gene expression. However, the existence of many HDAC isoforms in embryonic kidneys raises questions concerning the possible specificity or redundancy of their functions. We report here that targeted deletion of both the Hdac1 and Hdac2 genes from the ureteric bud (UB) cell lineage of mice causes bilateral renal hypodysplasia. One copy of either Hdac1 or Hdac2 is sufficient to sustain normal renal development."		* '''Histone deacetylase 1 and 2 regulate Wnt and p53 pathways in the ureteric bud epithelium'''{{#pmid:25758227\|PMID25758227}} "Histone deacetylases (HDACs) regulate a broad range of biological processes through removal of acetyl groups from histones as well as non-histone proteins. Our previous studies showed that Hdac1 and Hdac2 are bound to promoters of key renal developmental regulators and that HDAC activity is required for embryonic kidney gene expression. However, the existence of many HDAC isoforms in embryonic kidneys raises questions concerning the possible specificity or redundancy of their functions. We report here that targeted deletion of both the Hdac1 and Hdac2 genes from the ureteric bud (UB) cell lineage of mice causes bilateral renal hypodysplasia. One copy of either Hdac1 or Hdac2 is sufficient to sustain normal renal development."

	* '''~~Bmp7~~ functions via a polarity mechanism to promote cloacal septation'''{{#pmid:22253716\|PMID22253716}} "During normal development in human and other placental mammals, the embryonic cloacal cavity separates along the axial longitudinal plane to give rise to the urethral system, ventrally, and the rectum, dorsally. Defects in cloacal development are very common and present clinically as a rectourethral fistula in about 1 in 5,000 live human births. Yet, the cellular mechanisms of cloacal septation remain poorly understood. ...Our results strongly indicate that Bmp7/JNK signaling regulates remodeling of the cloacal endoderm resulting in a topological separation of the urinary and digestive systems. Our study points to the importance of Bmp and JNK signaling in cloacal development and rectourethral malformations."		* '''{{Bmp}}7 functions via a polarity mechanism to promote cloacal septation'''{{#pmid:22253716\|PMID22253716}} "During normal development in human and other placental mammals, the embryonic cloacal cavity separates along the axial longitudinal plane to give rise to the urethral system, ventrally, and the rectum, dorsally. Defects in cloacal development are very common and present clinically as a rectourethral fistula in about 1 in 5,000 live human births. Yet, the cellular mechanisms of cloacal septation remain poorly understood. ...Our results strongly indicate that Bmp7/JNK signaling regulates remodeling of the cloacal endoderm resulting in a topological separation of the urinary and digestive systems. Our study points to the importance of Bmp and JNK signaling in cloacal development and rectourethral malformations."

	* '''Size and location of the kidneys during the fetal period'''{{#pmid:21110022\|PMID21110022}} "The level of the left kidney was higher than the level of the right kidney in the fetal period. The posterior surface relations to the ribs showed certain ascendance during gestation, corresponding to vertebral levels. However, fetal kidneys do not reach the same level as adults at full term. The kidneys move farther apart from the midline of the body during the fetal period. The dimensions, weight, and volume of the kidneys increased with gestational age during the fetal period. The ratio between kidney weights and fetal body weights were determined, and we observed that the ratio decreased during the fetal period. There were no sex or laterality differences in any parameter." (See also [[Fetal Development]])		* '''Size and location of the kidneys during the {{fetal}} period'''{{#pmid:21110022\|PMID21110022}} "The level of the left kidney was higher than the level of the right kidney in the {{fetal}} period. The posterior surface relations to the ribs showed certain ascendance during gestation, corresponding to vertebral levels. However, fetal kidneys do not reach the same level as adults at full term. The kidneys move farther apart from the midline of the body during the fetal period. The dimensions, weight, and volume of the kidneys increased with gestational age during the fetal period. The ratio between kidney weights and fetal body weights were determined, and we observed that the ratio decreased during the fetal period. There were no sex or laterality differences in any parameter." (See also [[Fetal Development]])

	* '''Characterization of Mesonephric Development and Regeneration Using Transgenic Zebrafish.'''{{#pmid:20810610\|PMID20810610}} "The majority of previous studies have focused on the pronephros of zebrafish, which consists of only two nephrons and is structurally simpler than the mesonephros of adult fish and the metanephros of mammals. To evaluate the zebrafish system for more complex studies of kidney development and regeneration, we investigated the development and post-injury regeneration of the mesonephros in adult zebrafish." (See also [[Zebrafish Development]])		* '''Characterization of Mesonephric Development and Regeneration Using Transgenic Zebrafish.'''{{#pmid:20810610\|PMID20810610}} "The majority of previous studies have focused on the pronephros of zebrafish, which consists of only two nephrons and is structurally simpler than the mesonephros of adult fish and the metanephros of mammals. To evaluate the zebrafish system for more complex studies of kidney development and regeneration, we investigated the development and post-injury regeneration of the mesonephros in adult zebrafish." (See also [[Zebrafish Development]])

Z8600021 at 22:22, 23 Ocak 2020

2020-01-23T22:22:46Z

← Older revision		Revision as of 09:22, 24 Ocak 2020
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			* '''A practical guide to the stereological assessment of glomerular number, size, and cellular composition'''{{#pmid:31960613\|PMID31960613}} "The evaluation of a range of measures in the kidneys, such as developmental stage, rate and success, injury, and disease processes, relies on obtaining information on the three-dimensional structure of the renal corpuscles, and in particular the glomerular capillary tufts. To do this in the most accurate, comprehensive, and unbiased manner depends on a knowledge of stereological methods. In this article, we provide a practical guide for researchers on how to quantitate a number of structures in the kidneys, including the estimation of total glomerular number, glomerular capillary length and filtration surface area, and the cellular composition of individual glomeruli. Guidance is also provided on how to apply these methods to kidneys at different sizes and levels of maturity."

	* '''Branching morphogenesis of the urinary collecting system in the human embryonic metanephros'''{{#pmid:30192900\|PMID30192900}} "An elaborate system of ducts collects urine from all nephrons, and this structure is known as the urinary collecting system (UCS). This study focused on how the UCS is formed during human embryogenesis. Fifty human embryos between the Carnegie stage {{CS14}} and {{CS23}} were selected from the [[Kyoto Collection]] at the Congenital Anomaly Research Center of Kyoto University, Japan. Metanephroses, including the UCS, were segmented on serial digital virtual histological sections. Three-dimensional images were computationally reconstructed for morphological and quantitative analyses. A CS timeline was plotted. It consisted of the 3-D structural morphogenesis of UCS and quantification of the total amount of end-branching, average and maximum numbers of generations, deviation in the metanephros, differentiation of the urothelial epithelium in the renal pelvis, and timing of the rapid expansion of the renal pelvis. The first UCS branching generation occurred by {{CS16}}. The average branching generation reached a maximum of 8.74 ± 1.60 and was already the twelfth in {{CS23}}. ...Differentiation may have continued up until the tenth generation to allow for renal pelvis expansion. The branching speed was not uniform. There were significantly more branching generations in the polar- than in the interpolar regions (P < 0.05). Branching speed reflects the growth orientation required to form the metanephros. Further study will be necessary to understand the renal pelvis expansion mechanism in {{CS23}}."		* '''Branching morphogenesis of the urinary collecting system in the human embryonic metanephros'''{{#pmid:30192900\|PMID30192900}} "An elaborate system of ducts collects urine from all nephrons, and this structure is known as the urinary collecting system (UCS). This study focused on how the UCS is formed during human embryogenesis. Fifty human embryos between the Carnegie stage {{CS14}} and {{CS23}} were selected from the [[Kyoto Collection]] at the Congenital Anomaly Research Center of Kyoto University, Japan. Metanephroses, including the UCS, were segmented on serial digital virtual histological sections. Three-dimensional images were computationally reconstructed for morphological and quantitative analyses. A CS timeline was plotted. It consisted of the 3-D structural morphogenesis of UCS and quantification of the total amount of end-branching, average and maximum numbers of generations, deviation in the metanephros, differentiation of the urothelial epithelium in the renal pelvis, and timing of the rapid expansion of the renal pelvis. The first UCS branching generation occurred by {{CS16}}. The average branching generation reached a maximum of 8.74 ± 1.60 and was already the twelfth in {{CS23}}. ...Differentiation may have continued up until the tenth generation to allow for renal pelvis expansion. The branching speed was not uniform. There were significantly more branching generations in the polar- than in the interpolar regions (P < 0.05). Branching speed reflects the growth orientation required to form the metanephros. Further study will be necessary to understand the renal pelvis expansion mechanism in {{CS23}}."

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	===Articles===		===Articles===
			{{#pmid:31960613}}

	{{#pmid:29158444}}		{{#pmid:29158444}}

Z8600021 at 23:57, 18 Haziran 2019

2019-06-18T23:57:29Z

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	Week 8 - {{CS22}} mesonephros<br>(now degenerating)		Week 8 - {{CS22}} mesonephros<br>(now degenerating)
	\|}		\|}

			====Wolffian body====
			Historic name for the developing combined renal (mesonephros) and genital (paramesonephrotic blastema) structures. This term is no longer used in describing development.

			{{Male Vignette}}

	===Metanephros===		===Metanephros===

Z8600021 at 23:42, 18 Haziran 2019

2019-06-18T23:42:50Z

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	===Mesonephros===		===Mesonephros===
			The mesonephros, historically called the Wolffian body, is the embryonic stage of renal development that is entirely lost during development of the later fetal /adult metanepros. The only developmental structure not lost is the {{mesonephric duct}}.

	{\|		{\|
	\| Human E24, Mouse E9.5 caudal to pronephros		\| Human E24, Mouse {{ME9.5}} caudal to pronephros
	* forms by induction from pronephros		* forms by induction from pronephros
	* pronephric duct now becomes mesonephric duct (also called Wolffian Duct)		* pronephric duct now becomes mesonephric duct (also called Wolffian Duct)
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	\| [[File:Stage 13 kidney sections 2.jpg\|200px]]		\| [[File:Stage 13 kidney sections 2.jpg\|200px]]

	Week 5 - ~~[[Carnegie stage 13\|~~Stage ~~13]]~~ mesonephros<br>(extending the length of the body)		Week 5 - Stage {{CS13}} mesonephros<br>(extending the length of the body)
	\| [[File:Stage22 mesonephros.jpg\|200px]]		\| [[File:Stage22 mesonephros.jpg\|200px]]

	Week 8 - ~~[[Carnegie stage 22\|Stage 22]]~~ mesonephros<br>(now degenerating)		Week 8 - {{CS22}} mesonephros<br>(now degenerating)
	\|}		\|}

	===Metanephros===		===Metanephros===
	[[File:Fetal_10wk_urogenital_1.jpg\|thumb\|200px\|Early fetal kidney (week 10)]]		[[File:Fetal_10wk_urogenital_1.jpg\|thumb\|200px\|Early fetal kidney (week 10)]]
	* Human E35-37, Mouse ~~E11~~ epithelia bud at end of mesonephric duct ureteric bud and associated metanephric mesenchyme		* Human E35-37, Mouse {{ME11}} epithelia bud at end of mesonephric duct ureteric bud and associated metanephric mesenchyme

	Ureteric Bud		Ureteric Bud

Z8600021 at 14:08, 3 Haziran 2019

2019-06-03T14:08:30Z

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	===Urethra Development===		===Urethra Development===
	The entire human male and female urethra is {{endoderm}}al in origin based on the presence of FOXA1, KRT 7, uroplakin, and the absence of KRT10 staining.{{#pmid:30287094\|PMID30287094}} A recent study of male penile urethra describes a theory of a "two-step process of urethral plate canalization and urethral fold fusion to form the human penile urethra. Canalization ("opening zipper") opens the solid urethral plate into a groove, and fusion ("closing zipper")."{{#pmid:27397682\|PMID27397682}}		The entire human male and female urethra is {{endoderm}}al in origin based on the presence of FOXA1, KRT 7, uroplakin, and the absence of KRT10 staining.{{#pmid:30287094\|PMID30287094}} A recent study of male penile urethra describes a theory of a "two-step process of urethral plate canalization and urethral fold fusion to form the human penile urethra. Canalization ("opening zipper") opens the solid urethral plate into a groove, and fusion ("closing zipper")."{{#pmid:27397682\|PMID27397682}}{{#pmid:29155192\|PMID29155192}}

	==Kidney Ascent==		==Kidney Ascent==

Z8600021 at 14:03, 3 Haziran 2019

2019-06-03T14:03:06Z

← Older revision		Revision as of 01:03, 4 Haziran 2019
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	===Urethra Development===		===Urethra Development===
	The entire human male and female urethra is {{endoderm}}al in origin based on the presence of FOXA1, KRT 7, uroplakin, and the absence of KRT10 staining.{{#pmid:30287094\|PMID30287094}}		The entire human male and female urethra is {{endoderm}}al in origin based on the presence of FOXA1, KRT 7, uroplakin, and the absence of KRT10 staining.{{#pmid:30287094\|PMID30287094}} A recent study of male penile urethra describes a theory of a "two-step process of urethral plate canalization and urethral fold fusion to form the human penile urethra. Canalization ("opening zipper") opens the solid urethral plate into a groove, and fusion ("closing zipper")."{{#pmid:27397682\|PMID27397682}}

	==Kidney Ascent==		==Kidney Ascent==

Z8600021 at 13:59, 3 Haziran 2019

2019-06-03T13:59:06Z

← Older revision		Revision as of 00:59, 4 Haziran 2019
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	{{Trigone movie}}		{{Trigone movie}}

			===Urethra Development===
			The entire human male and female urethra is {{endoderm}}al in origin based on the presence of FOXA1, KRT 7, uroplakin, and the absence of KRT10 staining.{{#pmid:30287094\|PMID30287094}}

	==Kidney Ascent==		==Kidney Ascent==

Z8600021 at 04:31, 19 February 2019

2019-02-19T04:31:49Z

← Older revision		Revision as of 15:31, 19 February 2019
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	===Week 7===		===Week 7===

	* [[Carnegie stage ~~18]]~~ - Collecting tubules develop from the calices at stages [[Carnegie stage 17\|17]] and [[Carnegie stage 18\|18]]. They are surrounded by sharply outlined condensed primordia in the process of forming secretory tubules. Renal corpuscules are not yet present. By stage 18 the mesonephric duct and the ureter open almost independently into the vesico-urethral canal<ref name=Shikinami1926>{{Ref-Shikinami1926}}</ref>: i.e., the common excretory duct is disappearing. The cloacal membrane is ready to rupture.		* Carnegie stage {{CS18}} - Collecting tubules develop from the calices at stages [[Carnegie stage 17\|17]] and [[Carnegie stage 18\|18]]. They are surrounded by sharply outlined condensed primordia in the process of forming secretory tubules. Renal corpuscules are not yet present. By stage 18 the mesonephric duct and the ureter open almost independently into the vesico-urethral canal<ref name=Shikinami1926>{{Ref-Shikinami1926}}</ref>: i.e., the common excretory duct is disappearing. The cloacal membrane is ready to rupture.

			* Carnegie stage {{CS19}} - Lack of orientation in metanephrogenic tissue. Beginning formation of renal vesicles.<ref name=Streeter1957>{{Ref-Streeter1957}}</ref>

	===Week 8===		===Week 8===

	* [[Carnegie stage ~~20]]~~ - The external surface of the metanephros is said to be slightly lobulated. A reconstruction of the urinary system has been published by Shikinami<ref name=Shikinami1926>{{Ref-Shikinami1926}}</ref> (fig. 5).		* Carnegie stage {{CS20}} - The external surface of the metanephros is said to be slightly lobulated. A reconstruction of the urinary system has been published by Shikinami<ref name=Shikinami1926>{{Ref-Shikinami1926}}</ref> (fig. 5). S-shaped lumen in renal vesicles. Spoon-shaped capsules (Bowman’s).<ref name=Streeter1957>{{Ref-Streeter1957}}</ref>
	* [[Carnegie stage ~~21]]~~ - Metanephros is spoon-shaped glomerular capsules are developing, ~~but~~ no large glomeruli ~~are present yet~~.		* Carnegie stage {{CS21}} - Metanephros is spoon-shaped glomerular capsules are developing, no large glomeruli.<ref name=Streeter1957>{{Ref-Streeter1957}}</ref>
	* [[Carnegie stage ~~22]]~~ - Metanephros ~~has a~~ few large glomeruli ~~present~~.		* Carnegie stage {{CS22}} - Metanephros few large glomeruli.<ref name=Streeter1957>{{Ref-Streeter1957}}</ref>
	* [[Carnegie stage ~~23]]~~ - Comparison between the mesonephros and the metanephros in staged embryos is lacking. In metanephros, the kidneys have ascended from a sacral level at stages 13–15 to a lumbar level at stages 17–23. At stage 23 they are generally at the level of lumbar vertebrae 1–3..		* Carnegie stage {{CS23}} - Short secretory tubules. Numerous large glomeruli. Long secretory tubules. High epithelium in some tubules. Increased number and convolutions of tubules.<ref name=Streeter1957>{{Ref-Streeter1957}}</ref> Comparison between the mesonephros and the metanephros in staged embryos is lacking. In metanephros, the kidneys have ascended from a sacral level at stages 13–15 to a lumbar level at stages 17–23. At stage 23 they are generally at the level of lumbar vertebrae 1–3..

	==Fetal Kidney==		==Fetal Kidney==

← Older revision		Revision as of 09:24, 24 Ocak 2020
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	* '''Spatiotemporal heterogeneity and patterning of developing renal blood vessels'''{{#pmid:29627966\|PMID29627966}} "Here, we examine the developing kidney vasculature to assess its 3-dimensional structure and transcriptional heterogeneity. First, we observe that endothelial cells (ECs) grow coordinately with the kidney bud as early as {{ME10.5}}, and begin to show signs of specification by {{ME13.5}} when the first arteries can be identified. We then focus on how ECs pattern and remodel with respect to the developing nephron and collecting duct epithelia. ECs circumscribe nephron progenitor populations at the distal tips of the ureteric bud (UB) tree and form stereotyped cruciform structures around each tip. Beginning at the renal vesicle (RV) stage, ECs form a continuous plexus around developing nephrons. The endothelial plexus envelops and elaborates with the maturing nephron, becoming preferentially enriched along the early distal tubule."		* '''Spatiotemporal heterogeneity and patterning of developing renal blood vessels'''{{#pmid:29627966\|PMID29627966}} "Here, we examine the developing kidney vasculature to assess its 3-dimensional structure and transcriptional heterogeneity. First, we observe that endothelial cells (ECs) grow coordinately with the kidney bud as early as {{ME10.5}}, and begin to show signs of specification by {{ME13.5}} when the first arteries can be identified. We then focus on how ECs pattern and remodel with respect to the developing nephron and collecting duct epithelia. ECs circumscribe nephron progenitor populations at the distal tips of the ureteric bud (UB) tree and form stereotyped cruciform structures around each tip. Beginning at the renal vesicle (RV) stage, ECs form a continuous plexus around developing nephrons. The endothelial plexus envelops and elaborates with the maturing nephron, becoming preferentially enriched along the early distal tubule."

	* '''Conserved and Divergent Features of Human and Mouse Kidney Organogenesis'''{{#pmid:29449453\|PMID29449453}} "Human kidney function is underpinned by approximately 1,000,000 nephrons, although the number varies substantially, and low nephron number is linked to disease. Human kidney development initiates around 4 weeks of gestation and ends around 34-37 weeks of gestation. Over this period, a reiterative inductive process establishes the nephron complement. Studies have provided insightful anatomic descriptions of human kidney development, but the limited histologic views are not readily accessible to a broad audience. In this first paper in a series providing comprehensive insight into human kidney formation, we examined human kidney development in 135 anonymously donated human kidney specimens. We documented kidney development at a macroscopic and cellular level through histologic analysis, RNA in situ hybridization, immunofluorescence studies, and transcriptional profiling, contrasting human development (4-23 weeks) with {{mouse}} development at selected stages (embryonic day {{ME15.5}} and postnatal day 2). The high-resolution histologic interactive atlas of human kidney organogenesis generated can be viewed at the GUDMAP database (www.gudmap.org) together with three-dimensional reconstructions of key components of the data herein. At the anatomic level, human and mouse kidney development differ in timing, scale, and global features such as lobe formation and progenitor niche organization. The data also highlight differences in molecular and cellular features, including the expression and cellular distribution of anchor gene markers used to identify key cell types in mouse kidney studies." {{mouse}}
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	{\| class="wikitable mw-collapsible mw-collapsed"		{\| class="wikitable mw-collapsible mw-collapsed"
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	\| {{Older papers}}		\| {{Older papers}}
	* '''Reciprocal Spatiotemporally Controlled Apoptosis Regulates Wolffian Duct Cloaca Fusion'''{{#pmid:29326158\|PMID29326158}} "The epithelial Wolffian duct (WD) inserts into the cloaca (primitive bladder) before metanephric kidney development, thereby establishing the initial plumbing for eventual joining of the ureters and bladder. Defects in this process cause common anomalies in the spectrum of congenital anomalies of the kidney and urinary tract (CAKUT). However, developmental, cellular, and molecular mechanisms of WD-cloaca fusion are poorly understood. Through systematic analysis of early WD tip development in mice, we discovered that a novel process of spatiotemporally regulated apoptosis in WD and cloaca was necessary for WD-cloaca fusion."		* '''Conserved and Divergent Features of Human and Mouse Kidney Organogenesis'''{{#pmid:29449453\|PMID29449453}} "Human kidney function is underpinned by approximately 1,000,000 nephrons, although the number varies substantially, and low nephron number is linked to disease. Human kidney development initiates around 4 weeks of gestation and ends around 34-37 weeks of gestation. Over this period, a reiterative inductive process establishes the nephron complement. Studies have provided insightful anatomic descriptions of human kidney development, but the limited histologic views are not readily accessible to a broad audience. In this first paper in a series providing comprehensive insight into human kidney formation, we examined human kidney development in 135 anonymously donated human kidney specimens. We documented kidney development at a macroscopic and cellular level through histologic analysis, RNA in situ hybridization, immunofluorescence studies, and transcriptional profiling, contrasting human development (4-23 weeks) with {{mouse}} development at selected stages (embryonic day {{ME15.5}} and postnatal day 2). The high-resolution histologic interactive atlas of human kidney organogenesis generated can be viewed at the GUDMAP database (www.gudmap.org) together with three-dimensional reconstructions of key components of the data herein. At the anatomic level, human and mouse kidney development differ in timing, scale, and global features such as lobe formation and progenitor niche organization. The data also highlight differences in molecular and cellular features, including the expression and cellular distribution of anchor gene markers used to identify key cell types in mouse kidney studies." {{mouse}}

			* '''Reciprocal Spatiotemporally Controlled {{Apoptosis}} Regulates Wolffian Duct Cloaca Fusion'''{{#pmid:29326158\|PMID29326158}} "The epithelial Wolffian duct (WD) inserts into the cloaca (primitive bladder) before metanephric kidney development, thereby establishing the initial plumbing for eventual joining of the ureters and bladder. Defects in this process cause common anomalies in the spectrum of congenital anomalies of the kidney and urinary tract (CAKUT). However, developmental, cellular, and molecular mechanisms of WD-cloaca fusion are poorly understood. Through systematic analysis of early WD tip development in mice, we discovered that a novel process of spatiotemporally regulated apoptosis in WD and cloaca was necessary for WD-cloaca fusion."

	* '''{{Zebrafish}} Pronephros Development'''{{#pmid:28409341\|PMID28409341}} "The pronephros is the first kidney type to form in vertebrate embryos. The first step of pronephrogenesis in the zebrafish is the formation of the intermediate mesoderm during gastrulation, which occurs in response to secreted morphogens such as BMPs and Nodals. Patterning of the intermediate mesoderm into proximal and distal cell fates is induced by retinoic acid signaling with downstream transcription factors including wt1a, pax2a, pax8, hnf1b, sim1a, mecom, and irx3b. In the anterior intermediate mesoderm, progenitors of the glomerular blood filter migrate and fuse at the midline and recruit a blood supply. More posteriorly localized tubule progenitors undergo epithelialization and fuse with the cloaca. The Notch signaling pathway regulates the formation of multi-ciliated cells in the tubules and these cells help propel the filtrate to the cloaca. The lumenal sheer stress caused by flow down the tubule activates anterior collective migration of the proximal tubules and induces stretching and proliferation of the more distal segments. Ultimately these processes create a simple two-nephron kidney that is capable of reabsorbing and secreting solutes and expelling excess water-processes that are critical to the homeostasis of the body fluids. The zebrafish pronephric kidney provides a simple, yet powerful, model system to better understand the conserved molecular and cellular progresses that drive nephron formation, structure, and function." {{Zebrafish}}		* '''{{Zebrafish}} Pronephros Development'''{{#pmid:28409341\|PMID28409341}} "The pronephros is the first kidney type to form in vertebrate embryos. The first step of pronephrogenesis in the zebrafish is the formation of the intermediate mesoderm during gastrulation, which occurs in response to secreted morphogens such as BMPs and Nodals. Patterning of the intermediate mesoderm into proximal and distal cell fates is induced by retinoic acid signaling with downstream transcription factors including wt1a, pax2a, pax8, hnf1b, sim1a, mecom, and irx3b. In the anterior intermediate mesoderm, progenitors of the glomerular blood filter migrate and fuse at the midline and recruit a blood supply. More posteriorly localized tubule progenitors undergo epithelialization and fuse with the cloaca. The Notch signaling pathway regulates the formation of multi-ciliated cells in the tubules and these cells help propel the filtrate to the cloaca. The lumenal sheer stress caused by flow down the tubule activates anterior collective migration of the proximal tubules and induces stretching and proliferation of the more distal segments. Ultimately these processes create a simple two-nephron kidney that is capable of reabsorbing and secreting solutes and expelling excess water-processes that are critical to the homeostasis of the body fluids. The zebrafish pronephric kidney provides a simple, yet powerful, model system to better understand the conserved molecular and cellular progresses that drive nephron formation, structure, and function." {{Zebrafish}}
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	* '''Histone deacetylase 1 and 2 regulate Wnt and p53 pathways in the ureteric bud epithelium'''{{#pmid:25758227\|PMID25758227}} "Histone deacetylases (HDACs) regulate a broad range of biological processes through removal of acetyl groups from histones as well as non-histone proteins. Our previous studies showed that Hdac1 and Hdac2 are bound to promoters of key renal developmental regulators and that HDAC activity is required for embryonic kidney gene expression. However, the existence of many HDAC isoforms in embryonic kidneys raises questions concerning the possible specificity or redundancy of their functions. We report here that targeted deletion of both the Hdac1 and Hdac2 genes from the ureteric bud (UB) cell lineage of mice causes bilateral renal hypodysplasia. One copy of either Hdac1 or Hdac2 is sufficient to sustain normal renal development."		* '''Histone deacetylase 1 and 2 regulate Wnt and p53 pathways in the ureteric bud epithelium'''{{#pmid:25758227\|PMID25758227}} "Histone deacetylases (HDACs) regulate a broad range of biological processes through removal of acetyl groups from histones as well as non-histone proteins. Our previous studies showed that Hdac1 and Hdac2 are bound to promoters of key renal developmental regulators and that HDAC activity is required for embryonic kidney gene expression. However, the existence of many HDAC isoforms in embryonic kidneys raises questions concerning the possible specificity or redundancy of their functions. We report here that targeted deletion of both the Hdac1 and Hdac2 genes from the ureteric bud (UB) cell lineage of mice causes bilateral renal hypodysplasia. One copy of either Hdac1 or Hdac2 is sufficient to sustain normal renal development."

	* '''~~Bmp7~~ functions via a polarity mechanism to promote cloacal septation'''{{#pmid:22253716\|PMID22253716}} "During normal development in human and other placental mammals, the embryonic cloacal cavity separates along the axial longitudinal plane to give rise to the urethral system, ventrally, and the rectum, dorsally. Defects in cloacal development are very common and present clinically as a rectourethral fistula in about 1 in 5,000 live human births. Yet, the cellular mechanisms of cloacal septation remain poorly understood. ...Our results strongly indicate that Bmp7/JNK signaling regulates remodeling of the cloacal endoderm resulting in a topological separation of the urinary and digestive systems. Our study points to the importance of Bmp and JNK signaling in cloacal development and rectourethral malformations."		* '''{{Bmp}}7 functions via a polarity mechanism to promote cloacal septation'''{{#pmid:22253716\|PMID22253716}} "During normal development in human and other placental mammals, the embryonic cloacal cavity separates along the axial longitudinal plane to give rise to the urethral system, ventrally, and the rectum, dorsally. Defects in cloacal development are very common and present clinically as a rectourethral fistula in about 1 in 5,000 live human births. Yet, the cellular mechanisms of cloacal septation remain poorly understood. ...Our results strongly indicate that Bmp7/JNK signaling regulates remodeling of the cloacal endoderm resulting in a topological separation of the urinary and digestive systems. Our study points to the importance of Bmp and JNK signaling in cloacal development and rectourethral malformations."

	* '''Size and location of the kidneys during the fetal period'''{{#pmid:21110022\|PMID21110022}} "The level of the left kidney was higher than the level of the right kidney in the fetal period. The posterior surface relations to the ribs showed certain ascendance during gestation, corresponding to vertebral levels. However, fetal kidneys do not reach the same level as adults at full term. The kidneys move farther apart from the midline of the body during the fetal period. The dimensions, weight, and volume of the kidneys increased with gestational age during the fetal period. The ratio between kidney weights and fetal body weights were determined, and we observed that the ratio decreased during the fetal period. There were no sex or laterality differences in any parameter." (See also [[Fetal Development]])		* '''Size and location of the kidneys during the {{fetal}} period'''{{#pmid:21110022\|PMID21110022}} "The level of the left kidney was higher than the level of the right kidney in the {{fetal}} period. The posterior surface relations to the ribs showed certain ascendance during gestation, corresponding to vertebral levels. However, fetal kidneys do not reach the same level as adults at full term. The kidneys move farther apart from the midline of the body during the fetal period. The dimensions, weight, and volume of the kidneys increased with gestational age during the fetal period. The ratio between kidney weights and fetal body weights were determined, and we observed that the ratio decreased during the fetal period. There were no sex or laterality differences in any parameter." (See also [[Fetal Development]])

	* '''Characterization of Mesonephric Development and Regeneration Using Transgenic Zebrafish.'''{{#pmid:20810610\|PMID20810610}} "The majority of previous studies have focused on the pronephros of zebrafish, which consists of only two nephrons and is structurally simpler than the mesonephros of adult fish and the metanephros of mammals. To evaluate the zebrafish system for more complex studies of kidney development and regeneration, we investigated the development and post-injury regeneration of the mesonephros in adult zebrafish." (See also [[Zebrafish Development]])		* '''Characterization of Mesonephric Development and Regeneration Using Transgenic Zebrafish.'''{{#pmid:20810610\|PMID20810610}} "The majority of previous studies have focused on the pronephros of zebrafish, which consists of only two nephrons and is structurally simpler than the mesonephros of adult fish and the metanephros of mammals. To evaluate the zebrafish system for more complex studies of kidney development and regeneration, we investigated the development and post-injury regeneration of the mesonephros in adult zebrafish." (See also [[Zebrafish Development]])