Embryo

An embryo (/ˈɛmbrioʊ/ EM-bree-oh) is the initial stage

of development for a multicellular organism. In Embryo
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 cell. The resulting fusion of these two cells
produces a single-celled zygote that undergoes many
cell divisions that produce cells known as blastomeres.
The blastomeres (4-cell stage) are arranged as a solid
ball that when reaching a certain size, called a morula,
(16-cell stage) takes in fluid to create a cavity called a
blastocoel. The structure is then termed a blastula, or a
blastocyst in mammals.

The mammalian blastocyst hatches before implantating

into the endometrial lining of the womb. Once
implanted the embryo will continue its development
through the next stages of gastrulation, neurulation,
and organogenesis. Gastrulation is the formation of the A human embryo seven weeks after
three germ layers that will form all of the different conception (nine weeks gestational age)
parts of the body. Neurulation forms the nervous
Anatomical terminology
system, and organogenesis is the development of all
the various tissues and organs of the body.

A newly developing human is typically referred to as an embryo until the ninth week after conception,
when it is then referred to as a fetus. In other multicellular organisms, the word "embryo" can be used
more broadly to any early developmental or life cycle stage prior to birth or hatching.

Etymology
First attested in English in the mid-14th century, the word embryon derives from Medieval Latin embryo,
itself from Greek ἔμβρυον (émbryon) 'young one',[1] which is the neuter form of ἔμβρυος (émbryos)
'growing in'.[2] Morphologically it is derived from ἐν (en) 'in'[3] and βρύω (brýō) 'to swell, to be full'.[4]
The proper Latinized form of the Greek term would be embryum.
Development

Animal embryos
In animals, fertilization begins the process of embryonic
development with the creation of a zygote, a single cell
resulting from the fusion of gametes (e.g. egg and sperm).[5]
The development of a zygote into a multicellular embryo
proceeds through a series of recognizable stages, often
divided into cleavage, blastula, gastrulation, and
organogenesis.[6] 0:00

Cleavage is the period of rapid mitotic cell divisions that

occur after fertilization. During cleavage, the overall size of Embryonic development of salamander,
the embryo does not change, but the size of individual cells circa the 1920s
decrease rapidly as they divide to increase the total number of
cells.[7] Cleavage results in a blastula.[6]

Depending on the species, a blastula or blastocyst stage

embryo can appear as a ball of cells on top of yolk, or as a
hollow sphere of cells surrounding a middle cavity.[8] The
embryo's cells continue to divide and increase in number,
while molecules within the cells such as RNAs and proteins
actively promote key developmental processes such as gene
expression, cell fate specification, and polarity.[9] Before
implanting into the uterine wall the embryo is sometimes
known as the pre-implantation embryo or pre-implantation
conceptus.[10] Sometimes this is called the pre-embryo a
term employed to differentiate from an embryo proper in
relation to embryonic stem cell discourses.[11] Embryos (and one tadpole) of the
wrinkled frog (Rana rugosa)
Gastrulation is the next phase of embryonic development, and
involves the development of two or more layers of cells
(germinal layers). Animals that form two layers (such as
Cnidaria) are called diploblastic, and those that form three
(most other animals, from flatworms to humans) are called
triploblastic. During gastrulation of triploblastic animals, the
three germinal layers that form are called the ectoderm,
mesoderm, and endoderm.[8] All tissues and organs of a
mature animal can trace their origin back to one of these
layers.[12] For example, the ectoderm will give rise to the skin
epidermis and the nervous system,[13] the mesoderm will give
rise to the vascular system, muscles, bone, and connective Mouse and snake embryos
tissues,[14] and the endoderm will give rise to organs of the
digestive system and epithelium of the digestive system and
respiratory system.[15][16] Many visible changes in embryonic structure happen throughout gastrulation as
the cells that make up the different germ layers migrate and cause the previously round embryo to fold or
invaginate into a cup-like appearance.[8]

Past gastrulation, an embryo continues to develop into a mature multicellular organism by forming
structures necessary for life outside of the womb or egg. As the name suggests, organogenesis is the stage
of embryonic development when organs form. During organogenesis, molecular and cellular interactions
prompt certain populations of cells from the different germ layers to differentiate into organ-specific cell
types.[17] For example, in neurogenesis, a subpopulation of cells from the ectoderm segregate from other
cells and further specialize to become the brain, spinal cord, or peripheral nerves.[18]

The embryonic period varies from species to species. In human development, the term fetus is used
instead of embryo after the ninth week after conception,[19] whereas in zebrafish, embryonic development
is considered finished when a bone called the cleithrum becomes visible.[20] In animals that hatch from
an egg, such as birds, a young animal is typically no longer referred to as an embryo once it has hatched.
In viviparous animals (animals whose offspring spend at least some time developing within a parent's
body), the offspring is typically referred to as an embryo while inside of the parent, and is no longer
considered an embryo after birth or exit from the parent. However, the extent of development and growth
accomplished while inside of an egg or parent varies significantly from species to species, so much so
that the processes that take place after hatching or birth in one species may take place well before those
events in another. Therefore, according to one textbook, it is common for scientists to interpret the scope
of embryology broadly as the study of the development of animals.[8]

Plant embryos
Flowering plants (angiosperms) create embryos after the
fertilization of a haploid ovule by pollen. The DNA from the
ovule and pollen combine to form a diploid, single-cell
zygote that will develop into an embryo.[21] The zygote,
which will divide multiple times as it progresses throughout
embryonic development, is one part of a seed. Other seed
components include the endosperm, which is tissue rich in
nutrients that will help support the growing plant embryo, and
the seed coat, which is a protective outer covering. The first
cell division of a zygote is asymmetric, resulting in an
The inside of a Ginkgo seed, showing the
embryo with one small cell (the apical cell) and one large cell
embryo
(the basal cell).[22] The small, apical cell will eventually give
rise to most of the structures of the mature plant, such as the
stem, leaves, and roots.[23] The larger basal cell will give rise to the suspensor, which connects the
embryo to the endosperm so that nutrients can pass between them.[22] The plant embryo cells continue to
divide and progress through developmental stages named for their general appearance: globular, heart,
and torpedo. In the globular stage, three basic tissue types (dermal, ground, and vascular) can be
recognized.[22] The dermal tissue will give rise to the epidermis or outer covering of a plant,[24] ground
tissue will give rise to inner plant material that functions in photosynthesis, resource storage, and physical
support,[25] and vascular tissue will give rise to connective tissue like the xylem and phloem that
transport fluid, nutrients, and minerals throughout the plant.[26] In heart stage, one or two cotyledons
(embryonic leaves) will form. Meristems (centers of stem cell activity) develop during the torpedo stage,
and will eventually produce many of the mature tissues of the adult plant throughout its life.[22] At the
end of embryonic growth, the seed will usually go dormant until germination.[27] Once the embryo begins
to germinate (grow out from the seed) and forms its first true leaf, it is called a seedling or plantlet.[28]

Plants that produce spores instead of seeds, like bryophytes and ferns, also produce embryos. In these
plants, the embryo begins its existence attached to the inside of the archegonium on a parental
gametophyte from which the egg cell was generated.[29] The inner wall of the archegonium lies in close
contact with the "foot" of the developing embryo; this "foot" consists of a bulbous mass of cells at the
base of the embryo which may receive nutrition from its parent gametophyte.[30] The structure and
development of the rest of the embryo varies by group of plants.[31]

Since all land plants create embryos, they are collectively referred to as embryophytes (or by their
scientific name, Embryophyta). This, along with other characteristics, distinguishes land plants from
other types of plants, such as algae, which do not produce embryos.[32]

Research and technology

Biological processes
Embryos from numerous plant and animal species are studied in biological research laboratories across
the world to learn about topics such as stem cells,[33] evolution and development,[34] cell division,[35] and
gene expression.[36] Examples of scientific discoveries made while studying embryos that were awarded
the Nobel Prize in Physiology or Medicine include the Spemann-Mangold organizer, a group of cells
originally discovered in amphibian embryos that give rise to neural tissues,[37] and genes that give rise to
body segments discovered in Drosophila fly embryos by Christiane Nüsslein-Volhard and Eric
Wieschaus.[38]

Assisted reproductive technology

Creating and/or manipulating embryos via assisted reproductive technology (ART) is used for addressing
fertility concerns in humans and other animals, and for selective breeding in agricultural species. Between
the years 1987 and 2015, ART techniques including in vitro fertilization (IVF) were responsible for an
estimated one million human births in the United States alone.[39] Other clinical technologies include
preimplantation genetic diagnosis (PGD), which can identify certain serious genetic abnormalities, such
as aneuploidy, prior to selecting embryos for use in IVF.[40] Some have proposed (or even attempted - see
He Jiankui affair) genetic editing of human embryos via CRISPR-Cas9 as a potential avenue for
preventing disease;[41] however, this has been met with widespread condemnation from the scientific
community.[42][43]

ART techniques are also used to improve the profitability of agricultural animal species such as cows and
pigs by enabling selective breeding for desired traits and/or to increase numbers of offspring.[44] For
example, when allowed to breed naturally, cows typically produce one calf per year, whereas IVF
increases offspring yield to 9–12 calves per year.[45] IVF and other ART techniques, including cloning via
interspecies somatic cell nuclear transfer (iSCNT),[46] are also used in attempts to increase the numbers
of endangered or vulnerable species, such as Northern white rhinos,[47] cheetahs,[48] and sturgeons.[49]
Cryoconservation of plant and animal biodiversity
Cryoconservation of genetic resources involves collecting and storing the reproductive materials, such as
embryos, seeds, or gametes, from animal or plant species at low temperatures in order to preserve them
for future use.[50] Some large-scale animal species cryoconservation efforts include "frozen zoos" in
various places around the world, including in the UK's Frozen Ark,[51] the Breeding Centre for
Endangered Arabian Wildlife (BCEAW) in the United Arab Emirates,[52] and the San Diego Zoo Institute
for Conservation in the United States.[53][54] As of 2018, there were approximately 1,700 seed banks used
to store and protect plant biodiversity, particularly in the event of mass extinction or other global
emergencies.[55] The Svalbard Global Seed Vault in Norway maintains the largest collection of plant
reproductive tissue, with more than a million samples stored at −18 °C (0 °F).[56]

Fossilized embryos
Fossilized animal embryos are known from the Precambrian, and are found in great numbers during the
Cambrian period. Even fossilized dinosaur embryos have been discovered.[57]

See also
Embryo loss
Pregnancy
Prenatal development
In vitro fertilisation
Pre-embryo
Miscarriage
Abortion

Notes
1. ἔμβρυον (https://www.perseus.tufts.edu/hopper/text?doc=Perseus%3Atext%3A1999.04.005
7%3Aentry%3De%29%2Fmbruon) Archived (https://web.archive.org/web/20130531161449/
http://www.perseus.tufts.edu/hopper/text?doc=Perseus%3Atext%3A1999.04.0057%3Aentr
y%3De%29%2Fmbruon) 2013-05-31 at the Wayback Machine, Henry George Liddell,
Robert Scott, A Greek-English Lexicon, on Perseus
2. ἔμβρυος (https://www.perseus.tufts.edu/hopper/text?doc=Perseus%3Atext%3A1999.04.005
7%3Aentry%3De%29%2Fmbruos) Archived (https://web.archive.org/web/20130531041203/
http://www.perseus.tufts.edu/hopper/text?doc=Perseus%3Atext%3A1999.04.0057%3Aentr
y%3De%29%2Fmbruos) 2013-05-31 at the Wayback Machine, Henry George Liddell,
Robert Scott, A Greek-English Lexicon, on Perseus
3. ἐν (https://www.perseus.tufts.edu/hopper/text?doc=Perseus%3Atext%3A1999.04.0057%3A
entry%3De%29n) Archived (https://web.archive.org/web/20130531101258/http://www.perse
us.tufts.edu/hopper/text?doc=Perseus%3Atext%3A1999.04.0057%3Aentry%3De%29n)
2013-05-31 at the Wayback Machine, Henry George Liddell, Robert Scott, A Greek-English
Lexicon, on Perseus
 4. βρύω (https://www.perseus.tufts.edu/hopper/text?doc=Perseus%3Atext%3A1999.04.0057%
3Aentry%3Dbru%2Fw) Archived (https://web.archive.org/web/20130531101318/http://www.
perseus.tufts.edu/hopper/text?doc=Perseus%3Atext%3A1999.04.0057%3Aentry%3Dbru%2
Fw) 2013-05-31 at the Wayback Machine, Henry George Liddell, Robert Scott, A Greek-
English Lexicon, on Perseus
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original on 24 September 2015. Retrieved 8 August 2015.
External links
UNSW Embryology - Educational website (http://php.med.unsw.edu.au/embryology/index.ph
p?title=Main_Page)
A Comparative Embryology Gallery (https://web.archive.org/web/20060206140431/http://rav
en.zoology.washington.edu/embryos/)

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