embryology

1. The branch of biology that deals with the formation, early growth, and development of living organisms.
2. The embryonic structure or development of a particular organism.

The study of the development of an organism, commencing with the union of male and female gametes. Embryology literally means the study of embryos, but this definition is restrictive. An embryo is an immature organism contained within the coverings of an egg or within the body of the mother. Strictly speaking, the embryonic period ends at metamorphosis, hatching, or birth. Since developmental processes continue beyond these events, the scope of embryology is customarily broadened to encompass the entire life history of an organism. Embryology may, in this wider context, consider the mechanisms of both asexual reproduction and regeneration.

Animals

The production of male and female gametes is commonly considered to be the first phase in animal development. The differentiating gametes arise from diploid stem cells in the gonads. Cell division by meiosis reduces the number of chromosomes carried by a mature gamete to one-half that present in the stem cell. See also Gametogenesis.

The union of gametes (spermatozoon and ovum), representing the second phase of development, creates a diploid zygote with the potential to form an entire organism. Two events must occur for successful fertilization: the ovum must respond to contact with the spermatozoon by making preparations for further development, an event called activation, and the haploid nucleus of the spermatozoon must combine with the haploid nucleus of the ovum, an event called amphimixis.

Fertilization is the typical method to initiate development, but it is not the only method. In a few animals, the ovum develops independently by parthenogenesis, that is, without the participation of a spermatozoon.

A period of cell proliferation, converting the unicellular zygote into a multicellular embryo, represents the third phase of development. Cleavage is a modified form of cell division by mitosis, distinguished by little or no growth between the divisions. The cells of the embryo, or blastomeres, become progressively smaller at the end of each division, so the embryo maintains the relative size and shape of the zygote. Small, fluid-filled spaces form between the cleaving blastomeres, and these spaces eventually coalesce to create an internal cavity, or blastocoele. Upon the appearance of a blastocoele, the cells of an embryo are referred to collectively as the blastoderm. See also Blastulation.

The fourth phase of development is poorly delineated from cleavage, because the cells of the embryo continue to divide. Gastrulation is distinguished from cleavage by extensive cell rearrangements that lead, in most animals, to the establishment of three germ layers: an outer ectoderm, a middle mesoderm, and an inner endoderm. Endodermal and mesodermal cells of the blastoderm migrate to the inside of the embryo, while ectodermal cells remain on the surface, where they spread to completely cover the body.

Control of development passes from the cytoplasm to the nucleus immediately prior to gastrulation. Responding to cytoplasmic cues, the nuclei begin to specify the production of proteins that make the cells qualitatively different from one another. In a few invertebrates, the transfer of control from cytoplasm to nucleus actually fixes the developmental fate of a cell. In most other organisms, and particularly in vertebrates, the determination of cell fate is not finalized until the blastoderm has rearranged into the three germ layers. See also Cell lineage; Gastrulation; Germ layers.

The organization of cells into the tissues and organs of the body, constituting the fifth phase of development, is closely allied with gastrulation. Blastodermal rearrangements during creation of the germ layers shift cells into new positions and bring about new intercellular relationships. The developmental fate of a cell can, to a considerable degree, be the consequence of its new position. The influence exerted by one group of cells over the developmental fate of a neighboring group is called induction. Induction occurs by the transmission of chemical substances, called inducing agents.

Differentiation, or the process by which a cell becomes specialized, correlates to a reduction in the amount of genetic information that is expressed. Determination, or the fixation of a developmental fate, occurs when a cell has such a limited amount of usable genetic information that it must commit to a terminal pathway of differentiation. See also Cell differentiation.

Cellular differentiation is just one aspect of morphogenesis, or the development of form. Morphogenesis must be considered at all levels of organization, ranging from the individual cell to the whole organism. Such a broad perspective complicates the formulation of general theories of development. Presently, no comprehensive theory exists, but there are some embryologists who anticipate that a theory is possible once activities of the DNA molecule have been fully integrated into the topic of development. See also Animal morphogenesis; Reproduction (animal).

Plants

Reproductive development in multicellular plants is generally divided into three phases: gametogenesis, fertilization, and embryogenesis. The zygote produced by the fusion of male and female gametes divides to form a multicellular embryo with meristematic regions that ultimately produce the adult plant.

Development of the cell in flowering plants begins with a diploid megasporocyte located within the nucellar tissue of an immature ovule. This megasporocyte undergoes meiosis to form a tetrad of four haploid megaspores. In the most common pattern of development, three of these megaspores degenerate, leaving a single functional megaspore that undergoes several postmeiotic mitoses to form a mature megagametophyte (embryo sac) composed of seven cells and eight haploid nuclei. One of these haploid cells is the egg cell.

Development of the male gametes begins with numerous diploid cells (microsporocytes) located within the anthers of an immature flower. Each microsporocyte undergoes meiosis to form a tetrad of four haploid microspores, which then separate and enlarge to form mature pollen grains. Each microspore divides unequally to form a large vegetative cell, and a small generative cell located within the cytoplasm of the vegetative cell. The generative cell divides again, in either the maturing pollen grain or the elongating pollen tube, to form two genetically identical male gametes, the sperm cells.

The zygote is produced as part of a unique process known as double fertilization. One of the male gametes fuses with the egg cell to form the diploid zygote, while the other male gamete fuses with two polar nuclei, located near the center of the embryo sac, to form a triploid endosperm nucleus. Following double fertilization, the zygote develops into an embryo composed of two parts, the embryo proper and the suspensor. The embryo proper ultimately differentiates into the mature embryo, whereas the suspensor degenerates during later stages of development and is not usually present at maturity.

Flowering plants can be divided into two groups, monocots and dicots. In most dicots, the endosperm tissue is gradually absorbed by the developing embryo and is not present in the mature seed. Nutrients required for the germination of dicot seeds are generally stored in the embryonic leaves known as cotyledons. In contrast, most mature monocot seeds contain a significant amount of starchy endosperm tissue that serves as a source of nutrients for the germinating seedling.

Two important regions of the mature embryo are the root and the shoot apical meristems. The entire shoot system (stems, leaves, and flowers) of the adult plant forms from cells that are located in the shoot apical meristem of the mature embryo. The root apical meristem that is formed during embryogenesis becomes active during the early stages of germination and ultimately produces the entire root system of the adult plant. See also Apical meristem; Root (botany).

The final stages of embryogenesis in angiosperms include maturation, desiccation, and preparation for seed dormancy.

Different patterns of embryo development are found in gymnosperms and in the more primitive vascular and nonvascular plants. Double fertilization and the development of a nutritive endosperm tissue are features unique to the angiosperms. The haploid microgametophyte (germinating pollen grain) in most gymnosperms contains two male gametes, but only one of these participates in fertilization. The nutritive function served by the endosperm tissue in angiosperms is served in gymnosperms by the large haploid megagametophyte. Early divisions of the zygote are also different in gymnosperms; the zygote typically undergoes a series of free nuclear divisions during the earliest stages of embryogenesis, and multiple embryos often arise from a single zygote through a process known as polyembryony. Even more striking differences in embryogenesis are found in ferns and mosses, where the haploid or gametophytic phase of the life cycle is much more extensive.

Several major differences also exist between embryogenesis in plants and animals. Plant cells are surrounded by a cell wall that limits the contact and movement between adjacent cells. Embryogenesis in plants therefore proceeds without the morphogenetic movements that are characteristic of animal development. Morphogenesis in plants is also not limited to embryo development, but occurs throughout the life cycle. The mature plant embryo is therefore not simply a miniature version of the adult plant. See also Plant morphogenesis.

The study of the origin, growth, development, and function of an organism from fertilization to birth.

For more information on embryology, visit Britannica.com.

The study of the embryo; a major field of research in modern biology.

An expert in embryology.

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Morula, 8 cell stage

1 - morula, 2 - blastula

1 - blastula, 2 - gastrula with blastopore; orange - ectoderm, red - endoderm.

Embryology (from Greek ἔμβρυον, embryon, "unborn, embryo"; and -λογία, -logia) is a science which is about the development of an embryo from the fertilization of the ovum to the fetus stage. After cleavage, the dividing cells, or morula, becomes a hollow ball, or blastula, which develops a hole or pore at one end.

In bilateral animals, the blastula develops in one of two ways that divides the whole animal kingdom into two halves (see: Embryological origins of the mouth and anus). If in the blastula the first pore (blastopore) becomes the mouth of the animal, it is a protostome; if the first pore becomes the anus then it is a deuterostome. The protostomes include most invertebrate animals, such as insects, worms and molluscs, while the deuterostomes includes more advanced animals including the vertebrates. In due course, the blastula changes into a more differentiated structure called the gastrula.

The gastrula with its blastopore soon develops three distinct layers of cells (the germ layers) from which all the bodily organs and tissues then develop:

• The innermost layer, or endoderm, gives rise to the digestive organs, lungs and bladder.
• The middle layer, or mesoderm, gives rise to the muscles, skeleton and blood system.
• The outer layer of cells, or ectoderm, gives rise to the nervous system and skin.

In humans, the term embryo refers to the ball of dividing cells from the moment the zygote implants itself in the uterus wall until the end of the eighth week after conception. Beyond the eighth week, the developing human is then called a fetus. Embryos in many species often appear similar to one another in early developmental stages. The reason for this similarity is because species have a shared evolutionary history. These similarities among species are called homologous structures, which are structures that have the same or similar function and mechanism having evolved from a common ancestor.

Contents 1 History 2 Vertebrate and invertebrate embryology 3 Modern embryology research 4 See also 5 References 6 Further reading 7 External links

History

Human embryo at six weeks gestational age

Histological film 10 day mouse embryo

Beetle larvae

As recently as the 18th century, the prevailing notion in human embryology was preformation: the idea that semen contains an embryo — a preformed, miniature infant, or "homunculus" — that simply becomes larger during development. The competing explanation of embryonic development was epigenesis, originally proposed 2,000 years earlier by Aristotle. According to epigenesis, the form of an animal emerges gradually from a relatively formless egg. As microscopy improved during the 19th century, biologists could see that embryos took shape in a series of progressive steps, and epigenesis displaced preformation as the favored explanation among embryologists.^[1]

Modern embryological pioneers include Gavin de Beer, Charles Darwin, Ernst Haeckel, J.B.S. Haldane, and Joseph Needham, while much early embryology came from the work of Aristotle and the great Italian anatomists: Aldrovandi, Aranzio, Leonardo da Vinci, Marcello Malpighi, Gabriele Falloppia, Girolamo Cardano, Emilio Parisano, Fortunio Liceti, Stefano Lorenzini, Spallanzani, Enrico Sertoli, Mauro Rusconi, etc.^[2] Other important contributors include William Harvey, Kaspar Friedrich Wolff, Heinz Christian Pander, Karl Ernst von Baer, and August Weismann.

After the 1950s, with the DNA helical structure being unravelled and the increasing knowledge in the field of molecular biology, developmental biology emerged as a field of study which attempts to correlate the genes with morphological change, and so tries to determine which genes are responsible for each morphological change that takes place in an embryo, and how these genes are regulated.

Vertebrate and invertebrate embryology

Many principles of embryology apply to both invertebrate animals as well as to vertebrates.^[3] Therefore, the study of invertebrate embryology has advanced the study of vertebrate embryology. However, there are many differences as well. For example, numerous invertebrate species release a larva before development is complete; at the end of the larval period, an animal for the first time comes to resemble an adult similar to its parent or parents. Although invertebrate embryology is similar in some ways for different invertebrate animals, there are also countless variations. For instance, while spiders proceed directly from egg to adult form many insects develop through at least one larval stage

Modern embryology research

Currently, embryology has become an important research area for studying the genetic control of the development process (e.g. morphogens), its link to cell signalling, its importance for the study of certain diseases and mutations and in links to stem cell research.

See also

• Ontogeny
• Embryogenesis
• Recapitulation theory
• Prenatal development
• Protostomes
• Deuterostomes
• Germ layers
• Epigenesis (biology)
• Developmental biology
• Cell signalling
• Hedgehog signaling pathway
• Morphogens
• French flag model
• Embryo drawing

References

1. ^ Campbell et al. (p. 987)
2. ^ Massimo De Felici, Gregorio Siracus, The rise of embryology in Italy: from the Renaissance to the early 20th Century, Int. J. Dev. Biol. 44: 515-521 (2000).
3. ^ Parker, Sybil. "Invertebrate Embryology," McGraw-Hill Encyclopedia of Science & Technology (McGraw-Hill 1997).

Embryology - History of embryology as a science." Science Encyclopedia. Web. 06 Nov. 2009. <http://science.jrank.org/pages/2452/Embryology.html>.
"Germ layer." Encyclopædia Britannica. 2009. Encyclopædia Britannica Online. 06 Nov. 2009 <http://www.britannica.com/EBchecked/topic/230597/germ-layer>.

Further reading

• Scott F. Gilbert. Developmental Biology. Sinauer, 2003. ISBN 0-87893-258-5.
• Lewis Wolpert. Principles of Development. Oxford University Press, 2006. ISBN 0-19-927536-X.

External links

Wikimedia Commons has media related to: Embryology

• Embryo Research UK philosophy and ethics website discussing the ethics of embryology
• Human embryo research Canadian website covering the ethics of human embryo research
• Indiana University's Human Embryology Animations
• What is a human admixed embryo?
• UNSW Embryology Large resource of information and media
• Definition of embryo according to Webster

v • d • e Developmental biology > Human embryogenesis (development of embryo) and development of fetus First three weeks Week 1 Fertilization · Egg activation · Zygote · Cleavage · Morula · Blastula (Blastomere) · Blastocyst · Inner cell mass Week 2 (Bilaminar) Hypoblast · Epiblast Week 3 (Trilaminar) Germ layers Archenteron/Primitive streak (Primitive pit, Primitive knot/Blastopore, Primitive groove) · Gastrula/Gastrulation · Regional specification Ectoderm Surface ectoderm · Neuroectoderm · Somatopleure · Neurulation · Neural crest Endoderm Splanchnopleure Mesoderm Chorda- · Paraxial (Somite/Somitomere/Sclerotome/Myotome/Dermatome) · Intermediate · Lateral plate (Intraembryonic coelom, Splanchnopleure/Somatopleure) Extraembryonic/ uterus Trophoblast (Cytotrophoblast, Syncytiotrophoblast) Blastocoele · Yolk sack/exocoelomic cavity · Heuser's membrane · Extraembryonic coelom · Vitelline duct Umbilical cord (Umbilical artery, Umbilical vein, Wharton's jelly) · Allantois Placenta · Decidua (Decidual cells) · Chorionic villi/Intervillous space · Gestational sac (Amnion/Amniotic sac/Amniotic cavity, Chorion) Histogenesis Programmed cell death (Apoptosis) · Stem cells · Germ line development Organogenesis Limb development: Limb bud · Apical ectodermal ridge/AER other structures: Eye development · Cutaneous structure development · Heart development · Development of the urinary and reproductive organs (Some dates are approximate—see Carnegie stages and a timeline.)

v • d • e Human development of head and neck / branchial apparatus (GA 1.65) Nose Nasal placode · Olfactory pit · nasal prominences (Lateral, Medial) · Intermaxillary segment Mouth/neck Palate Primitive palate · Secondary palate 1st arch Frontonasal prominence · Maxillary prominence · Mandibular prominence (Meckel's cartilage) Anterior tongue: Lateral lingual swelling · Tuberculum impar Thyroid (pouches 3-5) Thyroid diverticulum · Thyroglossal duct · Ultimobranchial body General Pharyngeal groove (Cervical sinus) · Pharyngeal arch (1st, 2nd) · Pharyngeal pouch (Ultimobranchial body) respiratory system navs: anat nose,larynx/lower+thoracic cavity/physio/dev, noncongen/congen/tumors, symptoms+signs/eponymous, proc mouth navs: anat/dev, noncongen/congen face/congen GI/neoplasia, symptoms, proc endocrine navs: anat/physio/dev/hormones, noncongen/congen/neoplasia, symptoms+signs/eponymous, proc

v • d • e Ossification (GA 2.80) Upper limb Ossification of humerus · Ossification of ulna · Ossification of radius Lower limb Ossification of tibia Head cranium: Ossification of occipital bone · Ossification of frontal bone · Ossification of temporal bone · Ossification of the sphenoid · Ossification of ethmoid facial bones: Ossification of maxilla · Ossification of the mandible Other Ossification of scapula bone and cartilage navs: anat/physio/dev, noncongen/congen/neoplasia, symptoms+signs/eponymous signs, proc

v • d • e Prenatal development/Mammalian development of circulatory system (GA 5) Vascular Blood island Development of arteries Dorsal aorta · Aortic arches Development of veins Cardinal veins · Ducts of Cuvier Fetal circulation umbilical cord: Umbilical vein → Ductus venosus → Inferior vena cava → Heart → Pulmonary artery → Ductus arteriosus → Umbilical artery yolk sac: Vitelline veins · Vitelline arteries Heart development Primitive heart tube Truncus arteriosus · Bulbus cordis · Primitive ventricle · Primitive atrium · Sinus venosus Septa/ostia Endocardial cushions/Septum intermedium · Ostium primum · Septum primum (Ostium secundum) · Septum secundum (Foramen ovale) · Aorticopulmonary septum Other Atrial canal heart navs: anat/physio/dev, noncongen/congen/neoplasia, symptoms+signs/eponymous, proc

v • d • e Prenatal development/Mammalian development of nervous system (GA 9.733) General neural development/ neurulation/neurula Notochord · Neuroectoderm · Neural plate · Neural folds · Neural groove Neural crest (Cranial, Trunk, Cardiac) · Neural tube (Neuromere/Rhombomere, Cephalic flexure) Alar plate (sensory) · Basal plate (motor) Glioblast · Neuroblast note: mostly ecoderm, but mesoderm is precursor for epineurium, perineurium, and endoneurium central nervous system navs: anat/physio/dev, noncongen/congen/neoplasia, symptoms+signs/eponymous, proc

v • d • e Prenatal development/Mammalian development of eye and ear (GA 10.1002) Eye development Optic vesicles · Optic stalk · Optic cup · Lens placode Auditory development Auditory vesicle · Auditory pit eye navs: anat/adnexa anat/pathways/physio/dev, noncongen/congen/neoplasia, eponymous signs, proc ear navs: anat/physio/dev, noncongen/congen, eponymous signs, proc

v • d • e Prenatal development/Mammalian development of respiratory system (overview) (GA 11.1071) Upper Nasal placode Lower Laryngotracheal groove · Lung buds respiratory system navs: anat nose,larynx/lower+thoracic cavity/physio/dev, noncongen/congen/tumors, symptoms+signs/eponymous, proc

v • d • e Prenatal development/Mammalian development of digestive system (GA 11.1101) Gut Upper GI tract and accessory Stomodeum Foregut: upper GI (Buccopharyngeal membrane, Rathke's pouch, Tracheoesophageal septum) · accessory (Pancreatic bud, Hepatic diverticulum) Lower GI tract Midgut Hindgut: Urorectal septum Proctodeum Abdominopelvic Mesentery Dorsal mesentery · Ventral mesentery Thoracic diaphragm Septum transversum digestive system navs: anat of tract,glands,perit,diaphragm/physio/dev, noncongen/congen/congen of d+w/neoplasia, symptoms+signs/eponymous, proc

v • d • e Prenatal development/mammalian embryogenesis - Development of the urinary and reproductive organs (GA 11.1204) Common Urinary/Reproductive system Cloacal membrane - Cloaca - Urethral groove - Urogenital sinus - Urachus - Urogenital folds Kidney development Nephrogenic cord - Nephrotome - Pronephros - Mesonephros (Mesonephric tubules) Ureteric bud - Metanephric blastema Fetal genital development Primarily internal Gonadal ridge - Sex cord (Cortical cords, Testis cords) Pronephric duct/Wolffian duct/mesonephric duct - Müllerian duct/paramesonephric ducts (Vaginal plate) Primarily external Genital tubercle - Phallus - Labioscrotal folds Gubernaculum - Processus vaginalis Homologues List of homologues of the human reproductive system urinary system navs: anat/physio/dev, noncongen/acid+base/congen/neoplasia, symptoms+signs/eponymous, proc reproductive system navs: anat female,male/physio/dev, noncongen/congen/neoplasia, symptoms+signs/eponymous, proc

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