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Embryology (from
Greek Greek may refer to: Greece Anything of, from, or related to Greece, a country in Southern Europe: *Greeks, an ethnic group. *Greek language, a branch of the Indo-European language family. **Proto-Greek language, the assumed last common ancestor ...
ἔμβρυον, ''embryon'', "the unborn,
embryo An embryo is an initial stage of development of a multicellular organism. In organisms that reproduce sexually, embryonic development is the part of the life cycle that begins just after fertilization of the female egg cell by the male sperm ...
"; and -λογία, '' -logia'') is the branch of
animal Animals are multicellular, eukaryotic organisms in the Kingdom (biology), biological kingdom Animalia. With few exceptions, animals Heterotroph, consume organic material, Cellular respiration#Aerobic respiration, breathe oxygen, are Motilit ...
biology Biology is the scientific study of life. It is a natural science with a broad scope but has several unifying themes that tie it together as a single, coherent field. For instance, all organisms are made up of cells that process hereditar ...
that studies the
prenatal development Prenatal development () includes the development of the embryo and of the fetus during a viviparous animal's gestation. Prenatal development starts with fertilization, in the germinal stage of embryonic development, and continues in fetal devel ...
of gametes (sex cells),
fertilization Fertilisation or fertilization (see spelling differences), also known as generative fertilisation, syngamy and impregnation, is the fusion of gametes to give rise to a new individual organism or offspring and initiate its development. Pro ...
, and development of
embryo An embryo is an initial stage of development of a multicellular organism. In organisms that reproduce sexually, embryonic development is the part of the life cycle that begins just after fertilization of the female egg cell by the male sperm ...
s and
fetus A fetus or foetus (; plural fetuses, feti, foetuses, or foeti) is the unborn offspring that develops from an animal embryo. Following embryonic development the fetal stage of development takes place. In human prenatal development, fetal develo ...
es. Additionally, embryology encompasses the study of
congenital A birth defect, also known as a congenital disorder, is an abnormal condition that is present at birth regardless of its cause. Birth defects may result in disabilities that may be physical, intellectual, or developmental. The disabilities c ...
disorders that occur before birth, known as teratology. Early embryology was proposed by Marcello Malpighi, and known as preformationism, the theory that organisms develop from pre-existing miniature versions of themselves.
Aristotle Aristotle (; grc-gre, Ἀριστοτέλης ''Aristotélēs'', ; 384–322 BC) was a Greek philosopher and polymath during the Classical Greece, Classical period in Ancient Greece. Taught by Plato, he was the founder of the Peripatet ...
proposed the theory that is now accepted, epigenesis. Epigenesis is the idea that organisms develop from seed or egg in a sequence of steps. Modern embryology, developed from the work of Karl Ernst von Baer, though accurate observations had been made in Italy by anatomists such as Aldrovandi and
Leonardo da Vinci Leonardo di ser Piero da Vinci (15 April 14522 May 1519) was an Italian polymath of the High Renaissance who was active as a painter, Drawing, draughtsman, engineer, scientist, theorist, sculptor, and architect. While his fame initially re ...
in the Renaissance.


Comparative embryology


Preformationism and epigenesis

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 Aristotle (; grc-gre, Ἀριστοτέλης ''Aristotélēs'', ; 384–322 BC) was a Greek philosopher and polymath during the Classical Greece, Classical period in Ancient Greece. Taught by Plato, he was the founder of the Peripatet ...
. Much early embryology came from the work of the Italian anatomists Aldrovandi, Aranzio,
Leonardo da Vinci Leonardo di ser Piero da Vinci (15 April 14522 May 1519) was an Italian polymath of the High Renaissance who was active as a painter, Drawing, draughtsman, engineer, scientist, theorist, sculptor, and architect. While his fame initially re ...
, Marcello Malpighi, Gabriele Falloppio, Girolamo Cardano, Emilio Parisano, Fortunio Liceti, Stefano Lorenzini, Spallanzani, Enrico Sertoli, and
Mauro Ruscóni Mauro Ruscóni (18 November 1776 in Pavia - 27 March 1849 in Tremezzina) is an Italian physician and zoologist. Biography Coming from a respected merchant family, he finished his studies in his native town and in the 1790s, swept away by the po ...
. According to epigenesis, the form of an animal emerges gradually from a relatively formless egg. As
microscopy Microscopy is the technical field of using microscopes to view objects and areas of objects that cannot be seen with the naked eye (objects that are not within the resolution range of the normal eye). There are three well-known branches of mi ...
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. CLEVAGE Cleavage is the very beginning steps of a developing embryo. Cleavage refers to the many mitotic divisions that occur after the
egg An egg is an organic vessel grown by an animal to carry a possibly fertilized egg cell (a zygote) and to incubate from it an embryo within the egg until the embryo has become an animal fetus that can survive on its own, at which point the a ...
is fertilized by the sperm. The ways in which the cells divide is specific to certain types of animals and may have many forms.


Holoblastic

Holoblastic cleavage is the complete division of cells. Holoblastic cleavage can be radial (see: Radial cleavage), spiral (see: Spiral cleavage), bilateral (see:
Bilateral cleavage In embryology, cleavage is the division of cells in the early development of the embryo, following fertilization. The zygotes of many species undergo rapid cell cycles with no significant overall growth, producing a cluster of cells the same size ...
), or rotational (see:
Rotational cleavage In embryology, cleavage is the division of cells in the early development of the embryo, following fertilization. The zygotes of many species undergo rapid cell cycles with no significant overall growth, producing a cluster of cells the same size ...
). In holoblastic cleavage the entire egg will divide and become the embryo, whereas in meroblastic cleavage some cells will become the embryo and others will be the yolk sac.


Meroblastic

Meroblastic cleavage is the incomplete division of cells. The division furrow does not protrude into the yolky region as those cells impede membrane formation and this causes the incomplete separation of cells. Meroblastic cleavage can be bilateral (see:
Bilateral cleavage In embryology, cleavage is the division of cells in the early development of the embryo, following fertilization. The zygotes of many species undergo rapid cell cycles with no significant overall growth, producing a cluster of cells the same size ...
), discoidal (see:
Discoidal cleavage In embryology, cleavage is the division of cells in the early development of the embryo, following fertilization. The zygotes of many species undergo rapid cell cycles with no significant overall growth, producing a cluster of cells the same size ...
), or centrolecithal (see: Centrolecithal).


Basal phyla

Animals that belong to the basal phyla have holoblastic radial cleavage which results in radial symmetry (see: Symmetry in biology). During cleavage there is a central axis that all divisions rotate about. The basal phyla also has only one to two embryonic cell layers, compared to the three in bilateral animals ( endoderm, mesoderm, and ectoderm).


Bilaterians

In
bilateral animals The Bilateria or bilaterians are animals with bilateral symmetry as an embryo, i.e. having a left and a right side that are mirror images of each other. This also means they have a head and a tail (anterior-posterior axis) as well as a belly and ...
cleavage can be either holoblastic or meroblastic depending on the species. During gastrulation the blastula develops in one of two ways that divide the whole animal kingdom into two-halves (see: '' Embryological origins of the mouth and anus''). If in the blastula the first pore, or blastopore, becomes the mouth of the animal, it is a
protostome Protostomia () is the clade of animals once thought to be characterized by the formation of the organism's mouth before its anus during embryonic development. This nature has since been discovered to be extremely variable among Protostomia's mem ...
; if the blastopore becomes the anus then it is a
deuterostome Deuterostomia (; in Greek) are animals typically characterized by their anus forming before their mouth during embryonic development. The group's sister clade is Protostomia, animals whose digestive tract development is more varied. Some exa ...
. The protostomes include most
invertebrate Invertebrates are a paraphyletic group of animals that neither possess nor develop a vertebral column (commonly known as a ''backbone'' or ''spine''), derived from the notochord. This is a grouping including all animals apart from the chordate ...
animals, such as insects, worms and molluscs, while the
deuterostome Deuterostomia (; in Greek) are animals typically characterized by their anus forming before their mouth during embryonic development. The group's sister clade is Protostomia, animals whose digestive tract development is more varied. Some exa ...
s include the vertebrates. In due course, the blastula changes into a more differentiated structure called the gastrula. Soon after the gastrula is formed, three distinct layers of cells (the germ layers) from which all the bodily organs and tissues then develop.


Germ layers

* The innermost layer, or endoderm, give 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.


''Drosophila melanogaster'' (fruit fly)

Drosophila have been used as a developmental model for many years. The studies that have been conducted have discovered many useful aspects of development that not only apply to fruit flies but other species as well. Outlined below is the process that leads to cell and tissue differentiation. #