The Development of the Notochord in Chick Embryos

Total Page:16

File Type:pdf, Size:1020Kb

The Development of the Notochord in Chick Embryos The Development of the Notochord in Chick Embryos by A. JURAND1 From the Institute of Animal Genetics, Edinburgh WITH EIGHT PLATES THE literature on the notochord, a structure characteristic of all vertebrates, is very extensive, due to the phylogenetic importance of this organ, its role in early embryonic development, and its central position in the developing vertebral column. As early as 1834 the notochord tissue was described by Miiller as being similar in appearance to the parenchyma of plants. Surprisingly, however, in the chick embryo, which is so widely used by embryologists, its development has not very often been the subject of descriptive or experimental investigations. From the early days most work on this fundamental organ was done on fish and amphibians, probably because the notochord in lower vertebrates is more suitable for investigations, as it persists longer, carrying out its function as an embryonic and larval skeleton. Papers have been published in the past describing the fate of the notochord in birds, but mostly in connexion with wider problems, such as the development of the vertebral column (Bruni, 1912; Harman, 1922; Piiper, 1928; Kuhlenbeck, 1930; Williams, 1942; and others) or dealing with some details connected with the notochord, as, for instance, its sheath (Klaatsch, 1895; Held, 1921; Baitsell, 1925; Duncan, 1957 a, b; and others). The life-span of the notochord in birds and mammals is comparatively short. Its transient role, which is to provide rigidity of the longitudinal axis of the embryo, and its inductive potencies in cartilage formation become obsolete as soon as the definite skeleton begins to be formed. In chick embryos the whole period of development and existence of the notochord is limited to about 10 days (i.e. from the 2nd until the 1 lth day of embryonic life). During this period the formation of the prechordal material, its moulding into a rod-like organ, and, later, profound intracellular changes take place. The rate of the morpho- genetic process, as well as the process of internal cytodifferentiation, is un- doubtedly much faster in the notochord than in embryonic structures which develop more slowly but persist longer. For the reasons given, investigations on the morphogenesis and cytodifferen- tiation of the chick notochord appear to be particularly interesting. In the 1 Author's address: The Institute of Animal Genetics, West Mains Road, Edinburgh 9, U.K. [J. Embryol. exp. Morpb. Vol. 10, Part 4, pp. 602-21, December 1962] A. JURAND—DEVELOPMENT OF THE NOTOCHORD 603 normal developmental table of the chick prepared by Hamburger & Hamilton (1951), which still remains the best guide for all embryological work on the development of chick embryos, the changes in the notochord are not actually taken into account, as this work was based exclusively on external characteristics. Some general information on the development of the chick notochord can be found in monographs or textbooks such as those of Waddington (1952, 1956), Hamilton (1952), and Romanoff (1960). More detailed data based on light microscopy have been published by Kuhlenbeck (1930) and Williams (1942). It seemed desirable to investigate more closely the process of normal develop- ment of the chick notochord by means of light- and electron-microscope tech- niques in order to try to co-ordinate all that is known so far. MATERIAL AND METHODS Chick embryos at all developmental stages from the 5th to the 31st were obtained from Brown Leghorn eggs incubated at 38-5° C. (The stages are num- bered throughout the paper according to Hamburger & Hamilton (1951).) For histological reasons, after being dissected from the eggs while the entire yolk was immersed in Pannet-Compton's saline solution, the embryos were fixed for 1 hour in Carnoy's fluid (with chloroform), rinsed thoroughly with abso- lute alcohol, and, after clearing in methyl benzoate, were embedded in wax (m.p. 54° C). Serial transverse or sagittal sections were stained with methyl green-pyronine. For electron-microscope examination the embryos were fixed in toto for 15-45 minutes, depending on age, after being dissected in Pannet-Compton's solution. As a fixative, a modified 1 or 2 per cent, osmium tetroxide solution buffered with sodium barbitone and sodium acetate was used (Palade, 1952; Caulfield, 1957). As the original Palade buffer proved unsatisfactory, a number of modifications were tried. As a result of these technical approaches it was established that the best preservation of tissues in general, and of the notochord in particular, can be achieved by using a 2 per cent, osmium tetroxide solution made up according to the following formula: Sodium barbitone B.D.H. 0-117 g. Sodium acetate cryst. 0-078 g. Sucrose 0-18 g. Osmium tetroxide 0-2 g. 0-1 N hydrochloric acid 2 ml. Distilled water up to 10 ml. The pH of the above buffer solution without osmium tetroxide is 8-15. Its osmotic pressure is similar to that of the original Palade's buffer. The use of a higher pH than that recommended by Palade and other authors greatly im- proves the results, and so do the replacement of sodium chloride by sucrose 604 A. JURAND—DEVELOPMENT OF THE NOTOCHORD (Caulfield, 1957) and the increased percentage of osmium tetroxide (Pease, 1960). In preliminary investigations, a methyl-butyl methacrylate mixture proved to be unsatisfactory as an embedding medium, presumably due to its tendency to polymerization damage. All final embeddings were carried out with Araldite epoxy resin made up according to the original formula recommended by Glauert & Glauert (1958). After being fixed at about 4° C. the embryos were dehydrated with 35, 70, and 95 per cent, alcohol (5-10 minutes in each), and then with three changes of absolute alcohol. Embedding was carried out either with prepolymerized methacrylate mixture after two changes of liquid methacrylate or with the final Araldite mixture after replacing the absolute alcohol by two changes of 1:2- epoxy-propane. In the majority of cases ultra-thin sections were cut transversely to the longi- tudinal axis at the following levels: (1) In head-process and head-fold embryos, approximately in the middle of the length of the chorda-mesoblast concentration. (2) In embryos with 1-12 somites, at the level of the first somite or just anterior to it. (3) In older embryos in the cervical region of the notochord. The methacrylate sections were viewed without using electron 'stains', whereas the Araldite sections were contrasted either by floating, after being mounted on grids with collodion-carbon films, on potassium permanganate (1 per cent.) with uranyl acetate (2-5 per cent.) solution (modification of Lawn's method, 1960) for 20 minutes, or mounted straight on uncovered grids (e.g. Athene, No. 483) and subsequently stained by immersing in saturated lead acetate solution in ether-absolute alcohol (1:1) for 20-30 minutes. The latter method proved to be the more satisfactory. Some Araldite sections were treated by floating them on 10 per cent, phosphotungstic acid solution for 15 minutes and then washed for 5 minutes (Watson, 1958). The material used in the present investigations consisted of 185 chick embryos at various stages. RESULTS Light-microscope observations The aim of this part of the investigations was to have a preliminary and more general look at the developmental process taking place in the notochord, as, for example, the changes in the arrangement of the cells in the course of morpho- genesis of the organ. In particular, an attempt was made to re-examine the changes in the relationship of the chorda-mesoblast and, subsequently, of the notochord to the surrounding derivatives of the germ-layers, to determine the time of formation of the rod-like notochord, and to trace the vacuoliza- tion process and the subsequent appearance of the first involutive changes in the cells. A. JURAND—DEVELOPMENT OF THE NOTOCHORD 605 The histological material was also used in observations on the distribution of dividing cells by counting them at early stages (5-9) to get more information about the mechanism of elongation and growth of the notochord. The chorda-mesoblast condensation first appears at stage 5, at a point anterior to Hensen's node, as an extension of the primitive streak along the axis, in the form of an elongated, dorso-ventrally flattened strip of loosely arranged cells (Plate 1, fig. 1). It is distinctly separated from the medullary plate throughout its entire length by a definite boundary, but at the same time it remains very closely attached to it. The contact surface between the chorda-mesoblast and the medullary plate, which appears to be very intimate, is, however, not even; at regular intervals of about 30-50 ^ along the longitudinal axis of that surface distinct indentations can be found in the medullary plate, into which notochordal cells protrude. These indentations, which continue up to stage 7, can be seen particularly well in sagittal sections. It seems that during these stages the whole chorda-mesoblast tissue is being anchored by means of these indentations to the bottom of the medullary plate (Plate 2, fig. 11). Laterally, the central, comparatively loose mass of the presumptive notochord tissue is confluent with the paraxial mesoblast whose cells are, however, much looser than the axially situated prechordal cells. On the ventral side the chorda-mesoblast condensation at this stage, as well as in a few later stages, appears to be intimately fused with the unicellular layer of the endoderm. At the median line, therefore, the endodermal cells are difficult to distinguish from the prospective notochord cells as described by Hamilton (1952) and also by Fraser (1954) in somewhat older embryos in the pharyngeal region. The unification of both tissues at the head-process stage was described by Ussow (1906) as 'entochorda'.
Recommended publications
  • Works Neuroembryology
    Swarthmore College Works Biology Faculty Works Biology 1-1-2017 Neuroembryology D. Darnell Scott F. Gilbert Swarthmore College, [email protected] Follow this and additional works at: https://works.swarthmore.edu/fac-biology Part of the Biology Commons Let us know how access to these works benefits ouy Recommended Citation D. Darnell and Scott F. Gilbert. (2017). "Neuroembryology". Wiley Interdisciplinary Reviews: Developmental Biology. Volume 6, Issue 1. DOI: 10.1002/wdev.215 https://works.swarthmore.edu/fac-biology/493 This work is brought to you for free by Swarthmore College Libraries' Works. It has been accepted for inclusion in Biology Faculty Works by an authorized administrator of Works. For more information, please contact [email protected]. HHS Public Access Author manuscript Author ManuscriptAuthor Manuscript Author Wiley Interdiscip Manuscript Author Rev Dev Manuscript Author Biol. Author manuscript; available in PMC 2018 January 01. Published in final edited form as: Wiley Interdiscip Rev Dev Biol. 2017 January ; 6(1): . doi:10.1002/wdev.215. Neuroembryology Diana Darnell1 and Scott F. Gilbert2 1University of Arizona College of Medicine 2Swarthmore College and University of Helsinki Abstract How is it that some cells become neurons? And how is it that neurons become organized in the spinal cord and brain to allow us to walk and talk, to see, recall events in our lives, feel pain, keep our balance, and think? The cells that are specified to form the brain and spinal cord are originally located on the outside surface of the embryo. They loop inward to form the neural tube in a process called neurulation.
    [Show full text]
  • The Genetic Basis of Mammalian Neurulation
    REVIEWS THE GENETIC BASIS OF MAMMALIAN NEURULATION Andrew J. Copp*, Nicholas D. E. Greene* and Jennifer N. Murdoch‡ More than 80 mutant mouse genes disrupt neurulation and allow an in-depth analysis of the underlying developmental mechanisms. Although many of the genetic mutants have been studied in only rudimentary detail, several molecular pathways can already be identified as crucial for normal neurulation. These include the planar cell-polarity pathway, which is required for the initiation of neural tube closure, and the sonic hedgehog signalling pathway that regulates neural plate bending. Mutant mice also offer an opportunity to unravel the mechanisms by which folic acid prevents neural tube defects, and to develop new therapies for folate-resistant defects. 6 ECTODERM Neurulation is a fundamental event of embryogenesis distinct locations in the brain and spinal cord .By The outer of the three that culminates in the formation of the neural tube, contrast, the mechanisms that underlie the forma- embryonic (germ) layers that which is the precursor of the brain and spinal cord. A tion, elevation and fusion of the neural folds have gives rise to the entire central region of specialized dorsal ECTODERM, the neural plate, remained elusive. nervous system, plus other organs and embryonic develops bilateral neural folds at its junction with sur- An opportunity has now arisen for an incisive analy- structures. face (non-neural) ectoderm. These folds elevate, come sis of neurulation mechanisms using the growing battery into contact (appose) in the midline and fuse to create of genetically targeted and other mutant mouse strains NEURAL CREST the neural tube, which, thereafter, becomes covered by in which NTDs form part of the mutant phenotype7.At A migratory cell population that future epidermal ectoderm.
    [Show full text]
  • Role of Notochord in Specification of Cardiac Left-Right Orientation In
    DEVELOPMENTAL BIOLOGY 177, 96±103 (1996) ARTICLE NO. 0148 Role of Notochord in Speci®cation of Cardiac Left± View metadata, citation and similar papers at core.ac.uk brought to you by CORE Right Orientation in Zebra®sh and Xenopus provided by Elsevier - Publisher Connector Maria C. Danos and H. Joseph Yost1 Department of Cell Biology and Neuroanatomy, University of Minnesota, 4-135 Jackson Hall, 321 Church Street S.E., Minneapolis, Minnesota 55455 The left±right body axis is coordinately aligned with the orthogonal dorsoventral and anterioposterior body axes. The developmental mechanisms that regulate axis coordination are unknown. Here it is shown that the cardiac left±right orientation in zebra®sh (Danio rerio) is randomized in notochord-defective no tail and ¯oating head mutants. no tail (Brachyury) and ¯oating head (Xnot) encode putative transcription factors that are expressed in the organizer and notochord, structures which regulate dorsoventral and anterioposterior development in vertebrate embryos. Results from dorsal tissue extirpation and cardiac primordia explantation indicate that cardiac left±right orientation is dependent on dorsoanterior structures including the notochord and is speci®ed during neural fold stages in Xenopus laevis. Thus, the notochord coordinates the development of all three body axes in the vertebrate body plan. q 1996 Academic Press, Inc. INTRODUCTION lations early in development or by genetic mutation, re- sulting in a population frequency of approximately 50% In all vertebrates examined, left±right asymmetries are reversal of the normal left±right orientations (for review, consistently aligned with respect to the anterioposterior see Yost, 1995b). This suggests that in the absence of normal and dorsoventral axes.
    [Show full text]
  • Update on the Notochord Including Its Embryology, Molecular Development, and Pathology: a Primer for the Clinician
    Open Access Review Article DOI: 10.7759/cureus.1137 Update on the Notochord Including its Embryology, Molecular Development, and Pathology: A Primer for the Clinician Tushar Ramesh 1 , Sai V. Nagula 1 , Gabrielle G. Tardieu 2 , Erfanul Saker 2 , Mohammadali Shoja 3 , Marios Loukas 2 , Rod J. Oskouian 4 , R. Shane Tubbs 5 1. Neurology, University of Alabama at Birmingham 2. Department of Anatomical Sciences, St. George's University School of Medicine, Grenada, West Indies 3. Neurological Surgery, University of Alabama at Birmingham 4. Swedish Neuroscience Institute 5. Neurosurgery, Seattle Science Foundation Corresponding author: Gabrielle G. Tardieu, [email protected] Abstract The notochord is a rod-like embryological structure, which plays a vital role in the development of the vertebrate. Though embryological, remnants of this structure have been observed in the nucleus pulposus of the intervertebral discs of normal adults. Pathologically, these remnants can give rise to slow-growing and recurrent notochord-derived tumors called chordomas. Using standard search engines, the literature was reviewed regarding the anatomy, embryology, molecular development, and pathology of the human notochord. Clinicians who interpret imaging or treat patients with pathologies linked to the notochord should have a good working knowledge of its development and pathology. Categories: Genetics, Neurology, Other Keywords: notochord, nucleus pulposus, chordoma, spine, embryology, development Introduction And Background The notochord, namesake of the phylum chordata (Figure 1), plays a central role in vertebrate development. It is most prominent in the first trimester, wherein it guides the folding of the embryo and regulates the differentiation and maturation of surrounding tissues. It is a transient embryologic entity in humans, thought to be completely absent or present in minute quantities, within the nucleus pulposus of intervertebral discs in adults.
    [Show full text]
  • The Dorsal Neural Tube Organizes the Dermamyotome and Induces Axial Myocytes in the Avian Embryo
    Development 122, 231-241 (1996) 231 Printed in Great Britain © The Company of Biologists Limited 1996 DEV3253 The dorsal neural tube organizes the dermamyotome and induces axial myocytes in the avian embryo Martha S. Spence1, Joseph Yip2 and Carol A. Erickson1,* 1Section of Molecular and Cellular Biology, University of California, Davis, CA 95616, USA 2Department of Neurobiology, School of Medicine, University of Pittsburgh, Pittsburgh, PA 15261, USA *Author for correspondence (e-mail: [email protected]) SUMMARY Somites, like all axial structures, display dorsoventral reported previously to induce sclerotome. Thus, we have polarity. The dorsal portion of the somite forms the der- demonstrated that in the context of the embryonic envi- mamyotome, which gives rise to the dermis and axial mus- ronment, a dorsalizing signal from the dorsal neural tube culature, whereas the ventromedial somite disperses to can compete with the diffusible ventralizing signal from the generate the sclerotome, which later comprises the notochord. vertebrae and intervertebral discs. Although the neural In contrast to dorsal neural tube, pieces of ventral neural tube and notochord are known to regulate some aspects of tube, dorsal ectoderm or neural crest cells, all of which this dorsoventral pattern, the precise tissues that initially have been postulated to control dermamyotome formation specify the dermamyotome, and later the myotome from it, or to induce myogenesis, either fail to do so or provoke only have been controversial. Indeed, dorsal and ventral neural minimal inductive responses in any of our assays. However, tube, notochord, ectoderm and neural crest cells have all complicating the issue, we find consistent with previous been proposed to influence dermamyotome formation or to studies that following ablation of the entire neural tube, regulate myocyte differentiation.
    [Show full text]
  • An AOP-Based Ontology for Neural Tube Closure Caused by Disturbance in Retinoic Acid Signaling
    An AOP-based Ontology for Neural Tube Closure Caused by Disturbance in Retinoic Acid Signaling Cellular behavior Authors: Yvonne C.M. Staal1, Nancy Baker2, 3 4 Lyle D. Burgoon , George Daston , For each cell type information on its behavior was 5 1 Thomas B. Knudsen , Aldert H. Piersma collected from the available literature to map molecular interactions and genetic signals. 1. RIVM: National Institute of Public Health and the Environment, Bilthoven, The Netherlands; 2. Leidos, RTP, NC, United States; Table 1: example of information on cellular behavior for neuroectoderm fusion. 3. US Army Engineer Research and Development Center, RTP, NC, United States; Cell type Behavior Signal 4. Proctor & Gamble Company, Cincinnati OH, …. …. …. …. United States; neuroectoderm fusion inhibited by BMP 5. NCCT, US EPA/ORD, RTP, NC, United States neuroectoderm fusion requires Grhl2 Contact: [email protected] neuroectoderm fusion requires correct cell polarity Retinoic acid (RA) balance and leading neuroectoderm fusion inhibited by RhoA to neural tube defects neuroectoderm fusion requires Lrp6 Retinoid signaling plays an important role in embryo- neuroectoderm fusion involves Ephrin fetal development and its disruption is teratogenic. The biology of the RA pathway, leading to defects in neuroectoderm fusion requires Grhl2 neural tube closure was the basis for the construction of an ontology for developmental toxicity. neuroectoderm fusion requires Lrp6 We are constructing an ontology from an AOP network neuroectoderm fusion requires Traf4 that incorporates feedback-loops, which can be used for risk assessment. neuroectoderm fusion requires Cdx2 Fig 1: Schematic visualization (top to bottom) of neural tube closure. BMP inhibits dorso-lateral hinge point neuroectoderm fusion requires Pax3 (DLHP) formation, whereas this is stimulated by Noggin.
    [Show full text]
  • Embryology and Teratology in the Curricula of Healthcare Courses
    ANATOMICAL EDUCATION Eur. J. Anat. 21 (1): 77-91 (2017) Embryology and Teratology in the Curricula of Healthcare Courses Bernard J. Moxham 1, Hana Brichova 2, Elpida Emmanouil-Nikoloussi 3, Andy R.M. Chirculescu 4 1Cardiff School of Biosciences, Cardiff University, Museum Avenue, Cardiff CF10 3AX, Wales, United Kingdom and Department of Anatomy, St. George’s University, St George, Grenada, 2First Faculty of Medicine, Institute of Histology and Embryology, Charles University Prague, Albertov 4, 128 01 Prague 2, Czech Republic and Second Medical Facul- ty, Institute of Histology and Embryology, Charles University Prague, V Úvalu 84, 150 00 Prague 5 , Czech Republic, 3The School of Medicine, European University Cyprus, 6 Diogenous str, 2404 Engomi, P.O.Box 22006, 1516 Nicosia, Cyprus , 4Department of Morphological Sciences, Division of Anatomy, Faculty of Medicine, C. Davila University, Bucharest, Romania SUMMARY Key words: Anatomy – Embryology – Education – Syllabus – Medical – Dental – Healthcare Significant changes are occurring worldwide in courses for healthcare studies, including medicine INTRODUCTION and dentistry. Critical evaluation of the place, tim- ing, and content of components that can be collec- Embryology is a sub-discipline of developmental tively grouped as the anatomical sciences has biology that relates to life before birth. Teratology however yet to be adequately undertaken. Surveys (τέρατος (teratos) meaning ‘monster’ or ‘marvel’) of teaching hours for embryology in US and UK relates to abnormal development and congenital medical courses clearly demonstrate that a dra- abnormalities (i.e. morphofunctional impairments). matic decline in the importance of the subject is in Embryological studies are concerned essentially progress, in terms of both a decrease in the num- with the laws and mechanisms associated with ber of hours allocated within the medical course normal development (ontogenesis) from the stage and in relation to changes in pedagogic methodol- of the ovum until parturition and the end of intra- ogies.
    [Show full text]
  • Ectoderm: Neurulation, Neural Tube, Neural Crest
    4. ECTODERM: NEURULATION, NEURAL TUBE, NEURAL CREST Dr. Taube P. Rothman P&S 12-520 [email protected] 212-305-7930 Recommended Reading: Larsen Human Embryology, 3rd Edition, pp. 85-102, 126-130 Summary: In this lecture, we will first consider the induction of the neural plate and the formation of the neural tube, the rudiment of the central nervous system (CNS). The anterior portion of the neural tube gives rise to the brain, the more caudal portion gives rise to the spinal cord. We will see how the requisite numbers of neural progenitors are generated in the CNS and when these cells become post mitotic. The molecular signals required for their survival and further development will also be discussed. We will then turn our attention to the neural crest, a transient structure that develops at the site where the neural tube and future epidermis meet. After delaminating from the neuraxis, the crest cells migrate via specific pathways to distant targets in an embryo where they express appropriate target-related phenotypes. The progressive restriction of the developmental potential of crest-derived cells will then be considered. Additional topics include formation of the fundamental subdivisions of the CNS and PNS, as well as molecular factors that regulate neural induction and regional distinctions in the nervous system. Learning Objectives: At the conclusion of the lecture you should be able to: 1. Discuss the tissue, cellular, and molecular basis for neural induction and neural tube formation. Be able to provide some examples of neural tube defects caused by perturbation of neural tube closure.
    [Show full text]
  • Embryology of the Spine and Associated Congenital Abnormalities Kevin M
    The Spine Journal 5 (2005) 564–576 Review Article Embryology of the spine and associated congenital abnormalities Kevin M. Kaplan, MDa,*, Jeffrey M. Spivak, MDa,b, John A. Bendo, MDa,b aNew York University-Hospital for Joint Diseases, Department of Orthopaedic Surgery, 14th Floor, 301 East 17th Street, New York, NY 10003, USA bHospital for Joint Diseases Spine Center, Department of Orthopaedic Surgery, 14th Floor, 301 East 17th Street, New York, NY 10003, USA Received 11 June 2004; accepted 18 October 2004 Abstract BACKGROUND CONTEXT: The spine is a complex and vital structure. Its function includes not only structural support of the body as a whole, but it also serves as a conduit for safe passage of the neural elements while allowing proper interaction with the brain. Anatomically, a variety of tissue types are represented in the spine. Embryologically, a detailed cascade of events must occur to result in the proper formation of both the musculoskeletal and neural elements of the spine. Alterations in these embryologic steps can result in one or more congenital abnormalities of the spine. Other body systems forming at the same time embryologically can be affected as well, resulting in associated defects in the cardiopulmonary system and the gastrointestinal and genito- urinary tracts. PURPOSE: This article is to serve as a review of the basic embryonic development of the spine. We will discuss the common congenital anomalies of the spine, including their clinical presentation, as examples of errors of this basic embryologic process. STUDY DESIGN/SETTING: Review of the current literature on the embryology of the spine and associated congenital abnormalities.
    [Show full text]
  • 26 April 2010 TE Prepublication Page 1 Nomina Generalia General Terms
    26 April 2010 TE PrePublication Page 1 Nomina generalia General terms E1.0.0.0.0.0.1 Modus reproductionis Reproductive mode E1.0.0.0.0.0.2 Reproductio sexualis Sexual reproduction E1.0.0.0.0.0.3 Viviparitas Viviparity E1.0.0.0.0.0.4 Heterogamia Heterogamy E1.0.0.0.0.0.5 Endogamia Endogamy E1.0.0.0.0.0.6 Sequentia reproductionis Reproductive sequence E1.0.0.0.0.0.7 Ovulatio Ovulation E1.0.0.0.0.0.8 Erectio Erection E1.0.0.0.0.0.9 Coitus Coitus; Sexual intercourse E1.0.0.0.0.0.10 Ejaculatio1 Ejaculation E1.0.0.0.0.0.11 Emissio Emission E1.0.0.0.0.0.12 Ejaculatio vera Ejaculation proper E1.0.0.0.0.0.13 Semen Semen; Ejaculate E1.0.0.0.0.0.14 Inseminatio Insemination E1.0.0.0.0.0.15 Fertilisatio Fertilization E1.0.0.0.0.0.16 Fecundatio Fecundation; Impregnation E1.0.0.0.0.0.17 Superfecundatio Superfecundation E1.0.0.0.0.0.18 Superimpregnatio Superimpregnation E1.0.0.0.0.0.19 Superfetatio Superfetation E1.0.0.0.0.0.20 Ontogenesis Ontogeny E1.0.0.0.0.0.21 Ontogenesis praenatalis Prenatal ontogeny E1.0.0.0.0.0.22 Tempus praenatale; Tempus gestationis Prenatal period; Gestation period E1.0.0.0.0.0.23 Vita praenatalis Prenatal life E1.0.0.0.0.0.24 Vita intrauterina Intra-uterine life E1.0.0.0.0.0.25 Embryogenesis2 Embryogenesis; Embryogeny E1.0.0.0.0.0.26 Fetogenesis3 Fetogenesis E1.0.0.0.0.0.27 Tempus natale Birth period E1.0.0.0.0.0.28 Ontogenesis postnatalis Postnatal ontogeny E1.0.0.0.0.0.29 Vita postnatalis Postnatal life E1.0.1.0.0.0.1 Mensurae embryonicae et fetales4 Embryonic and fetal measurements E1.0.1.0.0.0.2 Aetas a fecundatione5 Fertilization
    [Show full text]
  • Development of the Notochord in Normal and Malformed Human Embryos and Fetuses
    Int. J. De... BioI. 35: 345.352 (1991) 345 Development of the notochord in normal and malformed human embryos and fetuses MIRNA SA RAGA BABIC Department of Histology and Embryology, Faculty of Medicine, Study at Split, University of Zagreb, Republic of Croatia, Yugoslavia ABSTRACT In view of its possible involvement in early embryogenesis and teratogenesis, the developmental characteristics of the human notochord were studied by light and electron microscopy and immunohistochemistry on 20 human conceptuses (5th-22nd week). At the earliest embryonic stages examined, the notochord is closely related to both the pharyngeal endoderm and the neuroectoderm of the posterior (tail) end of the neural tube. In both regions the interspace is bridged by slender cytoplasmic processes, lined with basal lamina and filled with amorphous extracellular material containing collagen types III and IV and laminin, The notochordal cells express cytokeratin brightly and vimentin weakly. As embryonic age progresses, the notochord gradually separates from the epithelia, becomes the axis of developing spinal column and undergoes progressive cell degenera- tion and rearrangement within the vertebral bodies, This is associated with extensive production of extracellular material and the first appearance of fibronectin. Intracellularly, the expression of vimentin gradually increases, while that of cytokeratin slightly weakens. Changes in the notochord parallel other developmental events in axial organs. In anencephalic fetuses the course of the notochord is irregular and partly interrupted with segments outside the basichondrocranium in specimens with craniorachischisis. KEY WORDS: notochord, human emblJos, skill! base, axial slnutllres Introduction few light and electron microscopic studies (Kunimoto, 1918; Peacock, 1951, 1952; Trout et al., 1982a,b; Shinohara and Tanaka, Although a transitory embryonic organ, the notochord plays an 1988) were performed on human samples.
    [Show full text]
  • Switching on the Notochord
    Downloaded from genesdev.cshlp.org on September 28, 2021 - Published by Cold Spring Harbor Laboratory Press PERSPECTIVE Switching on the notochord Vincent T. Cunliffe and Philip W. Ingham1 Developmental Genetics Programme, University of Sheffield, Sheffield S10 2TN, UK The notochord is a defining characteristic of the chor- ductive interactions are nevertheless critical for the es- date embryo. It is a dorsally located rod of tensile meso- tablishment of some cell lineages (Nishida 1997). In dermal tissue that lies immediately beneath the neural these organisms, notochordal fate is induced by vegetal tube. In vertebrates the notochord functions as a skeletal blastomeres in just 10 cells of the 110-cell embryo (Na- element during early embryogenesis and as a source of katani and Nishida 1994). These 10 cells then divide signals that pattern the neural tube and paraxial meso- twice to produce the entire set of 40 cells that comprises derm. Despite the recent identification of mutations af- the larval notochord. Initial expression of the Ciona in- fecting notochord development in zebrafish, the rather testinalis homolog of the vertebrate Brachyury gene, Ci- modest progress in isolating genes expressed in the no- Bra, coincides precisely with the restriction of early blas- tochord during its differentiation has limited our under- tomeres to a notochordal fate (Yasuoh and Satoh 1993). standing of the molecular mechanisms underpinning its Expression of Brachyury is then maintained exclusively structure and function. In a recent study, Takahashi et in the notochord, in contrast to the situation in verte- al. (1999), using embryos of the primitive chordate, the brate embryos, where Brachyury is expressed in both no- ascidian, have redressed the balance, identifying a large tochord and ventral–posterior mesoderm (Fig.
    [Show full text]