Embryology

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1 - morula, 2 - blastula
1 - blastula, 2 - gastrula with blastopore; orange - ectoderm, red - endoderm.
File:Human embryo 8 weeks.JPG
Dissection of human embryo, 38 mm - 8 weeks

Embryology (from Greek ἔμβρυον, embryon, "the unborn, embryo"; and -λογία, -logia) is the branch of biology that studies the development of gametes (sex cells), fertilization, and development of embryos and fetuses. Additionally, embryology is the study of congenital disorders that occur before birth.[1]

Embryonic development of animals

After cleavage, the dividing cells, or morula, becomes a hollow ball, or blastula, which develops a hole or pore at one end.

Bilaterals

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 include 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 a rise to the digestive organs, the gills, lungs or swim bladder if present, and kidneys or nephrites.
  • The middle layer, or mesoderm, gives rise to the muscles, skeleton if any, and blood system.
  • The outer layer of cells, or ectoderm, gives rise to the nervous system, including the brain, and skin or carapace and hair, bristles, or scales.

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.

Drosophila melanogaster (fruity fly)

File:Vinegar fly.jpg
Figure 1.1.1A Drosophila melanogaster
Figure 1.1.1B Drosophila melanogaster larvae contained in lab apparatus to be used for experiments in genetics and embryology.

Click here to read the main article on Drosophila embryogenesis

Drosophila melanogaster, a fruit fly, is a model organism in biology on which much research into embryology has been done (see figure 1.1.1A and figure 1.1.1B).[2] Before fertilization, the female gamete produces an abundance of mRNA - transcribed from the genes that encode bicoid protein and nanos protein.[3][4] These mRNA molecules are stored to be used later in what will become a developing embryo. The male and female Drosophila gametes exhibit anisogamy (differences in morphology and sub-cellular biochemistry). The female gamete is larger than the male gamete because it harbors more cytoplasm and, within the cytoplasm, the female gamete contains an abundance of the mRNA previously mentioned.[5][6] At fertilization, the male and female gametes fuse (plasmogamy) and then the nucleus of the male gamete fuses with the nucleus of the female gamete (karyogamy). Note that before the gametes' nuclei fuse, they are known as pronuclei.[7] A series of nuclear divisions will occur without cytokinesis (division of the cell) in the zygote to form a multi-nucleated cell (a cell containing multiple nuclei) known as a syncytium.[8][9] All the nuclei in the syncytium are identical, just as all the nuclei in every somatic cell of any multicellular organism are identical in terms of the DNA sequence of the genome.[10] Before the nuclei can differentiate in transcriptional activity, the embryo (syncytium) must be divided into segments. In each segment, a unique set of regulatory proteins will cause specific genes in the nuclei to be transcribed. The resulting combination of proteins will transform clusters of cells into early embryo tissues that will each develop into multiple fetal and adult tissues later in development (note: this happens after each nucleus becomes wrapped with its own cell membrane).

Outlined below is the process that leads to cell and tissue differentiation.

Maternal-effect genes - subject to Maternal (cytoplasmic) inheritance.

Zygotic-effect genes - subject to Mendelian (classical) inheritance.

  1. Gap genes establish 3 broad segments of the embryo.[14][15][16]
  2. Pair-rule genes define 7 segments of the embryo within the confines of the second broad segment that was defined by the gap genes.[17]
  3. Segment-polarity genes define another 7 segments by dividing each of the pre-existing 7 segments into anterior and posterior halves.[18][19]
  • Homeotic (homeobox) genes use the 14 segments as pinpoints for specific types of cell differentiation and the histological developments that correspond to each cell type.[20][21]

Humans

Humans are bilaterals and deuterostomes.

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 after conception (tenth week of pregnancy), the developing human is then called a fetus.

History

Human embryo at six weeks gestational age
File:10dayMouseEmb.jpg
Histological film 10-day mouse embryo

As recently as the 18th century, the prevailing notion in western 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. Much early embryology came from the work of the Italian anatomists Aldrovandi, Aranzio, Leonardo da Vinci, Marcello Malpighi, Gabriele Falloppio, Girolamo Cardano, Emilio Parisano, Fortunio Liceti, Stefano Lorenzini, Spallanzani, Enrico Sertoli, and Mauro Rusconi.[22] 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 favoured explanation among embryologists.[23]

After 1827

8–9-week human embryo

Karl Ernst von Baer and Heinz Christian Pander proposed the germ layer theory of development; von Baer discovered the mammalian ovum in 1827.[24][25][26] Modern embryological pioneers include Charles Darwin, Ernst Haeckel, J.B.S. Haldane, and Joseph Needham. Other important contributors include William Harvey, Kaspar Friedrich Wolff, Heinz Christian Pander, August Weismann, Gavin de Beer, Ernest Everett Just, and Edward B. Lewis.

After 1950

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 invertebrates as well as to vertebrates.[27] 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

References

Citations

  1. "Embryology Definition".<templatestyles src="Module:Citation/CS1/styles.css"></templatestyles>
  2. Carlson, Bruce M.; Kantaputra, Piranit N. (2014). "4 Molecular Basis for Embryonic Development". Human embryology and developmental biology (5th ed.). Philadelphia, PA: Elsevier/Saunders. p. 59. ISBN 978-1-4557-2794-0. the basic framework for understanding the molecular basis of embryonic development still rests largely on studies of developmental genetics in Drosophila<templatestyles src="Module:Citation/CS1/styles.css"></templatestyles>
  3. Alberts, Bruce; Johnson, Alexander; Lewis, Julian; Raff, Martin; Roberts, Keith; Walter, Peter (2008). "22 Development of Multicellular Organisms". Molecular biology of the cell (5th ed.). New York: Garland Science. p. 1334. ISBN 0-8153-4106-7. All the egg-polarity genes in these four classes are maternal-effect genes: it is the mother’s genome, not the zygotic genome, that is critical. Thus, a fly whose chromosomes are mutant in both copies of the Bicoid gene but who is born from a mother carrying one normal copy of Bicoid develops perfectly normally, with- out any defects in the head pattern. However, if that daughter fly is a female no functional Bicoid mRNA can be deposited into the anterior part of her own eggs, and all of these will develop into headless embryos regardless of the father’s genotype.<templatestyles src="Module:Citation/CS1/styles.css"></templatestyles>
  4. Alberts, Bruce; Johnson, Alexander; Lewis, Julian; Raff, Martin; Roberts, Keith; Walter, Peter (2008). "22 Development of Multicellular Organisms". Molecular biology of the cell (5th ed.). New York: Garland Science. p. 1334. ISBN 0-8153-4106-7. The future posterior end of the embryo con- tains a high concentration of mRNA for a regulator of translation called Nanos, which sets up a posterior gradient in the same way. The third signal is generated symmetrically at both ends of the egg<templatestyles src="Module:Citation/CS1/styles.css"></templatestyles>
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  7. Carlson, Bruce M.; Kantaputra, Piranit N. (2014). "2 Transport of Gametes and Fertilization". Human embryology and developmental biology (5th ed.). Philadelphia, PA: Elsevier/Saunders. p. 59. ISBN 978-1-4557-2794-0. after the head of the sperm enters the cytoplasm of the egg... the chromatin begins to spread out within the nucleus (now called a pronucleus) as it moves closer to the nuclear material of the egg.<templatestyles src="Module:Citation/CS1/styles.css"></templatestyles>
  8. Warn, RM (1986). "The cytoskeleton of the early Drosophila embryo". Journal of cell science. Supplement. 5: 311–28. PMID 3308915. This type of embryo shows a separation of mitosis from cytokinesis during the early stages of development. Most cells are only formed when a syncytium of approximately 6000 nuclei are present.<templatestyles src="Module:Citation/CS1/styles.css"></templatestyles>
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  11. Alberts, Bruce; Johnson, Alexander; Lewis, Julian; Raff, Martin; Roberts, Keith; Walter, Peter (2008). "22 Development of Multicellular Organisms". Molecular biology of the cell (5th ed.). New York: Garland Science. pp. 1333–1334. ISBN 0-8153-4106-7. In the stages before fertilization, the anteroposterior axis of the future embryo becomes defined by three systems of molecules that create landmarks in the oocyte (Figure 22–32)... The three sets of genes responsible for these localized deter- minants are referred to as the anterior, posterior, and terminal sets of egg- polarity genes.<templatestyles src="Module:Citation/CS1/styles.css"></templatestyles>
  12. Lua error in Module:Citation/CS1/Identifiers at line 47: attempt to index field 'wikibase' (a nil value).
  13. Alberts, Bruce; Johnson, Alexander; Lewis, Julian; Raff, Martin; Roberts, Keith; Walter, Peter (2008). "22 Development of Multicellular Organisms". Molecular biology of the cell (5th ed.). New York: Garland Science. p. 1336. ISBN 0-8153-4106-7. After the initial gradients of Bicoid and Nanos are created to define the antero- posterior axis, the segmentation genes refine the pattern.<templatestyles src="Module:Citation/CS1/styles.css"></templatestyles>
  14. Jäckle, Herbert; Hoch, Michael; Pankratz, Michael J.; Gerwin, Nicole; Sauer, Frank; Brönner, Günter (January 1992). "Transcriptional control by Drosophila gap genes". Journal Of Cell Science: 39–51. The segmented body pattern along the longitudinal axis of the Drosophila embryo is established by a cascade of specific transcription factor activities. This cascade is initiated by maternal gene products that are localized at the polar regions of the egg. The initial long-range positional information of the maternal factors, which are transcription factors (or are factors which activate or localize transcription factors), is transferred through the activity of the zygotic segmentation genes. The gap genes act at the top of this regulatory hierarchy. Expression of the gap genes occurs in discrete domains along the longitudinal axis of the preblastoderm and defines specific, overlapping sets of segment primordia.<templatestyles src="Module:Citation/CS1/styles.css"></templatestyles>
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  18. Martizez Arias, A; Baker, NE; Ingham, PW (May 1988). "Role of segment polarity genes in the definition and maintenance of cell states in the Drosophila embryo". Development (Cambridge, England). 103 (1): 157–70. PMID 3197626. Segment polarity genes are expressed and required in restricted domains within each metameric unit of the Drosophila embryo.<templatestyles src="Module:Citation/CS1/styles.css"></templatestyles>
  19. Carlson, Bruce M.; Kantaputra, Piranit N. (2014). "4 Molecular Basis for Embryonic Development". Human embryology and developmental biology (5th ed.). Philadelphia, PA: Elsevier/Saunders. p. 59. ISBN 978-1-4557-2794-0. In Drosophila, each stripe (segment) is subdivided into anterior and posterior halves. The posterior half of one segment and the anterior half of the next are collectively known as a parasegment.<templatestyles src="Module:Citation/CS1/styles.css"></templatestyles>
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  22. De Felici Massimo, Siracus Gregorio (2000). "The rise of embryology in Italy: from the Renaissance to the early 20th Century" (PDF). Int. J. Dev. Biol. 44: 515–521.<templatestyles src="Module:Citation/CS1/styles.css"></templatestyles>
  23. Campbell et al. (p. 987)
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  25. Lois N. Magner (2005). History of the Life Sciences. New York. Basel: Marcel Dekker. p. 166. ISBN 9780824743604.<templatestyles src="Module:Citation/CS1/styles.css"></templatestyles>
  26. Lua error in Module:Citation/CS1/Identifiers at line 47: attempt to index field 'wikibase' (a nil value).
  27. Parker, Sybil. "Invertebrate Embryology," McGraw-Hill Encyclopedia of Science & Technology (McGraw-Hill 1997).

Sources

Further reading

  • Lua error in Module:Citation/CS1/Identifiers at line 47: attempt to index field 'wikibase' (a nil value).
  • 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.
  • Carlson, Bruce M.; Kantaputra, Piranit N. (2014). Human embryology and developmental biology. Philadelphia, PA: Elsevier/Saunders. ISBN 978-1-4557-2794-0.<templatestyles src="Module:Citation/CS1/styles.css"></templatestyles> (click here for more information)

External links