The regionalisation of cell fate in the embryonic ectoderm was studied by analyzing the distribution of graft-derived cells in the chimaeric embryo following grafting of wheat germ agglutinin-gold-labelled cells and culturing primitive-streak-stage mouse embryos. Embryonic ectoderm in the anterior region of the egg cylinder contributes to the neuroectoderm of the prosencephalon and mesencephalon. Cells in the distal lateral region give rise to the neuroectoderm of the rhombencephalon and the spinal cord. Embryonic ectoderm at the archenteron and adjacent to the middle region of the primitive streak contributes to the neuroepithelium of the spinal cord. The proximal-lateral ectoderm and the ectodermal cells adjacent to the posterior region of the primitive streak produce the surface ectoderm, the epidermal placodes and the cranial neural crest cells. Some labelled cells grafted to the anterior midline are found in the oral ectodermal lining, whereas cells from the archenteron are found in the ...
We wish to understand how limbs are positioned with respect to the dorso-ventral axis of the body in vertebrate embryos, and how different regions of limb bud ectoderm, i.e. dorsal ectoderm, apical ridge and ventral ectoderm, originate. Signals from dorsal and ventral ectoderm control dorso-ventral patterning while the apical ectodermal ridge (AER) controls bud outgrowth and patterning along the proximo-distal axis. We show, using cell-fate tracers, the existence of two distinct ectodermal compartments, dorsal versus ventral, in both presumptive limb and flank of early chick embryos. This organisation of limb ectoderm is the first direct evidence, in vertebrates, of compartments in non-neural ectoderm. Since the apical ridge appears to be confined to this compartment boundary, this positions the limb. The mesoderm, unlike the ectoderm, does not contain two separate dorsal and ventral cell lineages, suggesting that dorsal and ventral ectoderm compartments may be important to ensure appropriate ...
Cranial placodes are evolutionary innovations of vertebrates. However, they most likely evolved by redeployment, rewiring and diversification of preexisting cell types and patterning mechanisms. In the second part of this review we compare vertebrates with other animal groups to elucidate the evolutionary history of ectodermal patterning. We show that several transcription factors have ancient bilaterian roles in dorsoventral and anteroposterior regionalisation of the ectoderm. Evidence from amphioxus suggests that ancestral chordates then concentrated neurosecretory cells in the anteriormost non-neural ectoderm. This anterior proto-placodal domain subsequently gave rise to the oral siphon primordia in tunicates (with neurosecretory cells being lost) and anterior (adenohypophyseal, olfactory, and lens) placodes of vertebrates. Likewise, tunicate atrial siphon primordia and posterior (otic, lateral line, and epibranchial) placodes of vertebrates probably evolved from a posterior proto-placodal region in
Header}} ==Introduction== [[File:Trilaminar_embryo.jpg,thumb,300px,The trilaminar embryo]] The top layer of the early [[T#trilaminar embryo,trilaminar embryo]] germ layers ({{ectoderm}}, {{mesoderm}} and {{endoderm}}) formed by {{gastrulation}}. The ectoderm can be though of as having 4 early regions: neural plate, neural crest, surface ectoderm and placodes. Note that there are other pages describing neural (central nervous system; brain and spinal cord) and neural crest (peripheral nervous system; sensory and sympathetic ganglia). Epidermis (integumentary, skin contribution) development will be briefly mentioned due to its ectoderm origin. The ectoderm contributes to the human embryo: # nervous system, both central (neural plate) and peripheral ({{neural crest}}). # epidermis of the skin (surface ectoderm) and pigmented cells ({{neural crest}}). # head regions that contribution sensory and endocrine structures ({{placodes}}). # adrenal gland medullary cells ({{neural crest}}). {, ...
The ectoderm is established during mammalian gastrulation from the anterior-distal portion of the pre-streak primitive ectoderm. Although fated spatially, transplantation experiments demonstrate that at this early stage, the cells are pluripotent and retain the ability to contribute to tissues derived from other germ layers, such as mesoderm (Beddington, 1982; Lawson et al., 1991). As gastrulation proceeds, the ectodermal precursors move in a proximal and posterior direction (Lawson et al., 1991), so that at the completion of gastrulation they comprise the entire inner layer of the embryo. At this stage, the cells have lost pluripotence, and have a differentiation potential limited to the ectodermal lineages (Carey et al., 1995). In lower vertebrates, the formation of ectoderm proceeds via a bipotent intermediate, definitive ectoderm, which differentiates to form the two major ectodermal lineages: surface ectoderm and neurectoderm (Hemmati-Brivanlou and Melton, 1997). The transient and dynamic ...
A BMP regulatory network controls ectodermal cell fate decisions at the neural plate border.[2] During ectodermal patterning the neural crest and preplacodal ectoderm are specified in adjacent domains at the neural plate border. BMP signalling is required for specification of both tissues, but how it is spatially and temporally regulated to achieve this is not understood. Here, using a transgenic zebrafish BMP reporter line in conjunction with double-fluorescent in situ hybridisation, we show that, at the beginning of neurulation, the ventral-to-dorsal gradient of BMP activity evolves into two distinct domains at the neural plate border: one coinciding with the neural crest and the other abutting the epidermis. In between is a region devoid of BMP activity, which is specified as the preplacodal ectoderm. We identify the ligands required for these domains of BMP activity. We show that the BMP-interacting protein Crossveinless 2 is expressed in the BMP activity domains and is under the control of ...
Development of the human lens begins at the 4 mm embryonic stage. Unlike the rest of the eye, which is derived mostly from the neural ectoderm, the lens is derived from the surface ectoderm. The first stage of lens differentiation takes place when the optic vesicle, which is formed from outpocketings in the neural ectoderm, comes in proximity to the surface ectoderm. The optic vesicle induces nearby surface ectoderm to form the lens placode. At the 4 mm stage, the lens placode is a single monolayer of columnar cells. As development progresses, the lens placode begins to deepen and invaginate. As the placode continues to deepen, the opening to the surface ectoderm constricts and the lens cells forms a structure known as the lens vesicle. By the 10 mm stage, the lens vesicle has completely separated from the surface ectoderm. After the 10mm stage, signals from the developing neural retina induces the cells closest to the posterior end of the lens vesicle begin to elongate toward the anterior end ...
We have characterized the early stages of murine hindlimb morphogenesis in the legless (lgl)mutant and non-mutant littermates. Initially the entire ventral ectoderm expresses many genetic markers characteristic of the AER (en-1, fgf-8, msx-2, dlx-2, cd44, and cx-43). Subsequently, the expression dom …
In addition to nourishing the embryo, extra-embryonic tissues (EETs) contribute to early em-bryonic patterning, primitive hematopoiesis, and fetal health. These tissues are of major importance for human medicine, as well as for efforts to improve livestock efficiency, but they remain incompletely understood. In bovines, EETs are accessible easily, in large amounts, and prior to implantation. We took advantage of this system to describe, in vitro and in vivo, the cell types present in bovine EETs at Day 18 of development. Specifically, we characterized the gene expression patterns and phenotypes of bovine extra-embryonic ectoderm (or trophoblast; bTC), endoderm (bXEC), and mesoderm (bXMC) cells in culture and compared them to their respective in vivo micro-dissected cells. After a week of culture, certain characteristics (e.g., gene expression) of the in vitro cells were altered with respect to the in vivo cells, but we were able to identify cores of cell-type-specific (and substrate-independent) genes
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The role of β-amyloid (Aβ) in the pathogenesis of Alzheimers disease (AD) is still considered crucial. The state of Aβ aggregation is critical in promoting neuronal loss and neuronal function impairment. Recently, we demonstrated that Acetylcholine (ACh) is neuroprotective against the toxic effects of Aβ in the cholinergic LAN-2 cells. In biophysical experiments, ACh promotes the soluble Aβ peptide conformation rather than the aggregation-prone β-sheet conformation. In order to better understand the biological role of ACh in AD, we studied the effect of Aβ on the phosphorylation of the cytosolic phospholipase A2 (cPLA2) in the TB neuroectodermal cell line, which differentiates toward a neuronal phenotype when cultured in the presence of retinoic acid (RA). We chose the phosphorylated form of cPLA2 (Ser505, Phospho-cPLA2) as a biomarker to test the influence of ACh on the effects of Aβ in both undifferentiated and RA-differentiated TB cells. Our results show that TB cells are responsive ...
MITSUNAGA NAKATSUBO Keiko , KUSUNOKI Shinichiro , KAWAKAMI Hayato , AKASAKA Koji , AKIMOTO Yoshihiro Medical molecular morphology : official journal of the Japanese Society for Clinical Molecular Morphology 42(2), 63-69, 2009-06-01 医中誌Web 参考文献48件 ...
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Developmental expression of two closely related fibroblast growth factor receptors, bek and flg, is described from early postimplantation until advanced organogenesis. Transcripts of bek and flg were first seen in the primitive ectoderm of egg-cylinder- stage embryos. Later, starting with somitogenesis, and then throughout embryogenesis, they were actively transcribed both in the mesoderm and neuroectoderm. Bek was expressed also in the surface ectoderm and in various epithelia, whereas flg expression was restricted mainly to the mesenchyme. In the limb bud bek transcripts displayed a gradient-like distribution and appeared earlier than flg. The two receptors, in contrast to their almost identical ligand binding specificity, displayed distinct spatial specificities throughout development, suggesting that developmental localization may contribute to functional specificity. The role of bek and flg in gastrulation and in epithelial-mesenchymal interactions of organogenesis will be discussed. ...
Egfr signaling is required in a narrow medial domain of the head ectoderm (here called head midline ) that includes the anlagen of the medial brain (including the dorsomedial and ventral medial domain of the brain, termed DMD and VMD respectively), the visual system (optic lobe, larval eye) and the stomatogastric nervous system (SNS). These head midline cells differ profoundly from their lateral neighbors in the way they develop. Three differences are noteworthy: (1) Like their counterparts in the mesectoderm, the head midline cells do not give rise to typical neuroblasts by delamination, but stay integrated in the surface ectoderm for an extended period of time. (2) The proneural gene l sc, which transiently (for approximately 30 minutes) comes on in all parts of the procephalic neurectoderm while neuroblasts delaminate, is expressed continuously in the head midline cells for several hours. (3) Head midline cells, similar to ventral midline cells of the trunk, require the Egfr pathway. In ...
Images provided by Claire Anderson. The publication of Liem et al. 1995 (Liem Jr., K.F, Tremml G, Roelink H, Jessell T.M. Dorsal differentiation of neural plate cells induced by BMP-mediated signals from epidermal ectoderm. Cell. 1995; 82: 969-979. PMID: 7553857 ) was referred for the probe information. Authors indicated further probe information could be found in another publication (Francis PH, Richardson MK, Brickell PM, Tickle C. Bone morphogenetic proteins and a signalling pathway that controls patterning in the developing chick limb. Development. 1994 Jan;120(1):209-18. PMID: 8119128). The indicated probe sequence was extracted from NCBI (GenBank: X75914.1 ...
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Ras - dorsal-ventral anterior localization (Ras-dva) family. Ras-dva subfamily. Ras-dva (Ras - dorsal-ventral anterior localization) subfamily consists of a set of proteins characterized only in Xenopus leavis, to date. In Xenopus Ras-dva expression is activated by the transcription factor Otx2 and begins during gastrulation throughout the anterior ectoderm. Ras-dva expression is inhibited in the anterior neural plate by factor Xanf1. Downregulation of Ras-dva results in head development abnormalities through the inhibition of several regulators of the anterior neural plate and folds patterning, including Otx2, BF-1, Xag2, Pax6, Slug, and Sox9. Downregulation of Ras-dva also interferes with the FGF-8a signaling within the anterior ectoderm. Most Ras proteins contain a lipid modification site at the C-terminus, with a typical sequence motif CaaX, where a = an aliphatic amino acid and X = any amino acid. Lipid binding is essential for membrane attachment, a key feature of most Ras proteins. ...
Bone morphogenetic protein 4, or BMP4, is a transforming growth factor that causes the cells of the ectoderm to differentiate into skin cells. Without BMP4 the ectoderm cells would develop into neural cells. Axial mesoderm cells under the ectoderm secrete inhibitory signals called chordin, noggin and follistatin. These inhibitory signals prevent the action of BMP4, which would normally make the cells ectoderm; as a result, the overlying cells take their normal course and develop into neural cells. The cells in the ectoderm that circumscribe these neural cells do not receive the BMP4 inhibitor signals and as a result BMP4 induces these cells to develop into skin cells.[8] Neural plate border specifiers are induced as a set of transcription factors. Distalless-5, PAX3 and PAX7 prevent the border region from becoming either neural plate or epidermis.[1] These induce a second set of transcription factors called neural crest specifiers, which cause cells to become neural crest cells. In a newly ...
BACKGROUND: The Brachyury (T) gene is required for the formation of posterior mesoderm and for axial development in both mouse and zebrafish embryos. In these species, and in Xenopus, the gene is expressed transiently throughout the presumptive mesoderm, and transcripts then persiste in notochord and posterior tissues. In Xenopus embryos, expression of the Xenopus homologue of Brachyury, Xbra, can be induced in presumptive ectoderm by basic fibroblast growth factor (FGF) and activin; in the absence of functional FGF or activin signalling pathways, expression of the gene is severely reduced. Ectopic expression of Xbra in presumptive ectoderm causes mesoderm to be formed. As Brachyury and its homologues encode sequence-specific DNA-binding proteins, it is likely that each functions by directly activating downstream mesoderm-specific genes. RESULTS: We show that expression in Xenopus embryos of RNA encoding a dominant-negative FGF receptor inhibits the mesoderm-inducing activity of Xbra. We ...
Eye formation in the human embryo begins at approximately three weeks into embryonic development and continues through the tenth week. Cells from both the mesodermal and the ectodermal tissues contribute to the formation of the eye. Specifically, the eye is derived from the neuroepithelium, surface ectoderm, and the extracellular mesenchyme which consists of both the neural crest and mesoderm. Neuroepithelium forms the retina, ciliary body, iris, and optic nerves. Surface ectoderm forms the lens, corneal epithelium and eyelid. The extracellular mesenchyme forms the sclera, the corneal endothelium and stroma, blood vessels, muscles, and vitreous. The eye begins to develop as a pair of optic vesicles on each side of the forebrain at the end of the 4th week of pregnancy. Optic vesicles are outgrowings of the brain which make contact with the surface ectoderm and this contact induces changes necessary for further development of the eye. Through a groove at the bottom of the optic vesicle known as ...
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The development of a vertebrate embryo is a complex process marked by several morphogenetic events, which create a highly reproducible pattern. The vertebrate limb has emerged as a model for studying pattern formation in the embryo mainly because limb manipulations do not affect embryo survival. Within the developing limb, experimental manipulation of the embryo resulted in the identification of the classical signaling centers known as the Zone of Polarizing Activity (ZPA) and the Apical Ectodermal Ridge (AER). The molecular signals required for function of the ZPA and AER have been identified. They are Sonic hedgehog (Shh) from the ZPA and Fibroblast Growth Factors (Fgfs) from the AER. The functions of each of these molecules are now beginning to be understood. Analysis of Shh and hedgehog (Hh) signaling target genes has shown that Hh activation in the limb bud mesoderm is required for normal limb development. It has been stated that Hh signaling in the limb bud ectoderm cannot occur because ...
Vertebrate sensory organs arise from epithelial thickenings called placodes. Along with neural crest cells, cranial placodes are considered ectodermal novelties that drove evolution of the vertebrate head. The anterior-most placode generates the endocrine lobe [adenohypophysis (ADH)] of the pituitary, a master gland controlling growth, metabolism, and reproduction. In addition to known ectodermal contributions, we use lineage tracing and time-lapse imaging in zebrafish to identify an endodermal contribution to the ADH. Single-cell RNA sequencing of the adult pituitary reveals similar competency of endodermal and ectodermal epithelia to generate all endocrine cell types. Further, endoderm can generate a rudimentary ADH-like structure in the near absence of ectodermal contributions. The fish condition supports the vertebrate pituitary arising through interactions of an ancestral endoderm-derived proto-pituitary with newly evolved placodal ectoderm. ...
A process that stimulates and directs a morphogenetic influence on a population of cells. These populations are often differentiating cells or adjacent neighboring cells that are not fixed on a developmental path. Embryonic induction plays a regulatory role in the development and construction of embryos.
TY - CHAP. T1 - The breast. AU - Heys, Steven D.. PY - 2006/1/1. Y1 - 2006/1/1. N2 - BASIC BIOLOGY; Anatomy of the breast; Embryology; The breast is a modified sweat gland which originates from the ectodermal layer of the embryo between the fifth and sixth week of gestation. It arises from the milk lines, which are two ridges of ectodermal thickening, running from the axilla to the groin. Although most of the milk line eventually disappears, a prominent ridge remains in the pectoral area to form the breast. The ectoderm in this area subsequently grows into the underlying mesoderm as a series (15-20) of buds. These ectodermal buds, which are initially solid, eventually form the lactiferous ducts and their associated alveoli. The adjacent mesenchyme develops into the surrounding adipose and connective tissues. During the final 2 months of gestation, the ducts become canalised and a mammary pit is formed in the ectoderm. The lactiferous ducts subsequently communicate with the mammary pit. ...
Ectoderm, Endoderm, and Mesoderm: What do the 3 Germ Cell Layers Give Rise To? Definitions, Table, and Easy Mnemonics for the 3 Germ Cell Layers
Ectoderm, Endoderm, and Mesoderm: What do the 3 Germ Cell Layers Give Rise To? Definitions, Table, and Easy Mnemonics for the 3 Germ Cell Layers
Neural brain development as well stem cell proliferation with circuits plus crest and mind ectoderm drosophila cells in tumor formation progenitor | mathappsllc.com
EV:0100003. EFO:0000414. MESH:A16.254.425.273. MIAA:0000173. ZFA:0000016. CALOHA:TS-0216. MAT:0000155. EMAPA:16069. GAID:1304. BILA:0000036. EHDAA2:0000428. XAO:0000001. BTO:0000315. TAO:0000016. MAT:0000173. VHOG:0000153. NCIT:C12703. UMLS:C0013574. FBbt:00000111. FMA:69070. AAO:0000137. http://linkedlifedata.com/resource/umls/id/C0013574. http://www.snomedbrowser.com/Codes/Details/362851007. http://en.wikipedia.org/wiki/Ectoderm. ...
Zebrafish msxB, msxC and msxE function together to refine the neural-nonneural border and regulate cranial placodes and neural crest development. part 2
Previously, the researchers were not able to address the genes possible role in mouse eye formation because inactivation of Six3 significantly disrupted development of the area of the brain where the eye normally forms. The St. Jude team overcame this problem by taking advantage of Cre/loxP-technology, which allowed them to choose the time and place in which to remove Six3 function from specific cells. This permitted the investigators to remove Six3 activity from the presumptive lens ectoderm (PLE)--the area of the developing head where the lens will ultimately form in response to a series of biochemical signals. Following this systematic approach, the St. Jude team demonstrated that Six3 plays its important role in the PLE. The investigators also showed that a key consequence of removing Six3 during early development is that the PLE fails to undergo its normal thickening, an initial critical step in lens formation.. Our discovery helps to better unravel the regulatory pathway that controls ...
The main points in the foregoing paper maybe summarised as follows :. (1) A yolk nucleus of the type described by Bambeke, as occurring in the egg of Pholcus, is present in the developing egg of Flustrella hispida.. (2) Segmentation and cell-lineage have been followed out in detail up to the 32-cell stage.. (3) The formation of the endoderm has been traced.. (4) The oral and aboral ectoderm are differentiated as early as the 16-cell stage, and remain quite distinct from that time onwards.. (5) The ciliated ring of the larva is formed by the coalescence of several originally distinct rows of cells, and not by the hypertrophy of a single row.. (6) A stomach, comparable to that of Alcyonidium, is present also in Flustrella.. ...
The neural plate is a key developmental structure that serves as the basis for the nervous system. Opposite the primitive streak in the embryo, ectodermal tissue thickens and flattens to become the neural plate. The ends of the neural plate, known as the neural folds, push the ends of the plate up ...
The brain and spinal cord develop from the ectoderm. Following formation of the neural ectoderm, the neural preplate is formed and splits to form the neural plate. Closure of the neural plate forms the neural tube in a process called neurulation (see description in Neural Tube overview). The central hollow space of the neural tube later forms the fluid-filled brain ventricles. Neuroepithelium is generated in the neural tube walls and gives rise to immature nerve precursors called neuroblasts, the majority of which migrate and grow leading axonal appendages, and then aggregate in specific, genetically determined locations that will become the brainstem and spinal cord.The neuroectoderm then subdivides into ventricular and subventricular zones, which produce separate waves of migrating neuroblasts.. Following expansion of the anterior part of the neural tube and of the neural crest region, the tube twists and indentations appear and form the primary brain vesicles - the prosencephalon ...
As our bodies form, cells within the embryo divide and separate. Certain cells come together to form the outer layer, or ectoderm, of the early embryo, and give rise to tissue such as the skin and nervous system (spine, peripheral nerves and brain). Other cells come together to form the mesoderm or middle layer of the embryo, and eventually give rise to tissue like muscle, heart or bone. Once cells have been assigned to the different regions - mesoderm or ectoderm - a mysterious mechanism draws boundaries between them that mark their permanent separation. Any defect in these boundaries leads to disorganized mixing of cell populations, severe embryo abnormalities and eventually lethality.. Until now, adherence was thought to be the principle force responsible for the separation of the ectoderm from the mesoderm in embryonic cells. The Differential Adhesion Hypothesis used to explain the process of embryonic tissue separation - which has been accepted until now - postulates that cells from each ...
Now so far the little hollow bead of cells is basically two layers of tissue thick, an outer layer, called the ectoderm, and an inner layer, called the endoderm, and these are called your germ layers. For those organisms that stop developing at this point with that classy mouth-anus combo, they only get two germ layers. Theyre called diploblastic and they were born that way, its totally okay. But for more complex organisms whose mouths are separate from our anuses (Yes!), we develop a third layer of tissue, making us triploblasts. Here, the ectoderm is going to end up being the animals skin and nerves and spinal cord and most of its brain while the endoderm ends up becoming the digestive tract: the esophagus and colon and liver and stuff. And in addition some of the cells being breaking off between the endoderm and the ectoderm and form another layer called the mesoderm. These cells will eventually end up as the muscles and the circulatory system and the reproductive system, and, in the case ...
In the early 20th century, a set of famous experiments by Hans Spemann and Hilde Mangold showed that the formation of nervous tissue is induced by the underlying mesoderm. For decades, though, the nature of the induction process defeated every attempt to figure it out, until finally it was resolved by genetic approaches in the 1990s. Induction of neural tissue requires inhibition of the gene for a so-called bone morphogenetic protein, or BMP. Specifically the protein BMP4 appears to be involved. Two proteins called Noggin and Chordin, both secreted by the mesoderm, are capable of inhibiting BMP4 and thereby inducing ectoderm to turn into neural tissue. It appears that a similar molecular mechanism is involved for widely disparate types of animals, including arthropods as well as vertebrates. In some animals, however, another type of molecule called Fibroblast Growth Factor or FGF may also play an important role in induction ...
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22 days: … --, Ectoderm --, Superficial ectoderm and neuroectoderm --, Neuro plate --, Neural groove and Neural fold folds (Neural crest begin to form) --, Neural tube ...
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Ectoderm is one of three germ layers--groups of cells that coalesce early during the embryonic life of all animals except maybe sponges, and from which organs and tissues form. As an embryo develops, a single fertilized cell progresses through multiple rounds of cell division. Eventually, the clump of cells goes through a stage called gastrulation, during which the embryo reorganizes itself into the three germ layers: endoderm, ectoderm, and mesoderm. After gastrulation, the embryo goes through a process called neurulation, which starts the development of nervous system.. Format: Articles Subject: Processes ...
Figure 3. Clk2 promotes neural induction and suppresses epidermis differentiation via inhibition of BMP signal. (a) Clk2 induced neural tissues in X. laevis ectodermal explants. clk2‐GR and clk2 SYA‐GR mRNAs (500 pg, 1,000 pg and 1,500 pg) were injected into the animal hemisphere of 4‐cell stage embryos. At the blastula stage, embryos were transferred to a medium containing DEX and the ectodermal explants were isolated. At the neurula stage, the explants were subjected to an RT‐PCR analysis to examine the expression of marker genes. Expression of early neural markers (sip1, sox2 and sox3), an anterior neural marker (otx2) and a posterior neural marker (hoxb9) was upregulated by Clk2 overexpression, whereas the differentiated neural marker (ncam) was only slightly increased. The expression of epidermal keratin was reduced in the presence of Clk2. Embryo and −RT indicate whole embryos examined in the presence or absence of reverse transcriptase, respectively. histone h4 was used as ...
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Arl13b is a gene known to regulate ciliogenesis. Functional alterations in this genes activity have been associated with Joubert syndrome. We found that in Arl13 null mouse embryos the orientation of the optic cup is inverted, such that the lens is abnormally surrounded by an inverted optic cup whose retina pigmented epithelium is oddly facing the surface ectoderm. Loss of Arl13b leads to the dis ...
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