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An embryo is an initial stage of development of a
multicellular organism A multicellular organism is an organism that consists of more than one cell, in contrast to unicellular organism. All species of animals, land plants and most fungi are multicellular, as are many algae, whereas a few organisms are partially ...
. In
organism In biology, an organism () is any living system that functions as an individual entity. All organisms are composed of cells ( cell theory). Organisms are classified by taxonomy into groups such as multicellular animals, plants, and ...
s that reproduce sexually, embryonic development is the part of the life cycle that begins just after
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. Proc ...
of the female
egg cell The egg cell, or ovum (plural ova), is the female reproductive cell, or gamete, in most anisogamous organisms (organisms that reproduce sexually with a larger, female gamete and a smaller, male one). The term is used when the female gamete i ...
by the male sperm cell. The resulting fusion of these two cells produces a single-celled
zygote A zygote (, ) is a eukaryotic cell formed by a fertilization event between two gametes. The zygote's genome is a combination of the DNA in each gamete, and contains all of the genetic information of a new individual organism. In multicel ...
that undergoes many cell divisions that produce cells known as blastomeres. The blastomeres are arranged as a solid ball that when reaching a certain size, called a morula, takes in fluid to create a cavity called a blastocoel. The structure is then termed a blastula, or a blastocyst in
mammal Mammals () are a group of vertebrate animals constituting the class Mammalia (), characterized by the presence of mammary glands which in females produce milk for feeding (nursing) their young, a neocortex (a region of the brain), fu ...
s. 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 three germ layers that will form all of the different parts of the body. Neurulation forms the
nervous system In biology, the nervous system is the highly complex part of an animal that coordinates its actions and sensory information by transmitting signals to and from different parts of its body. The nervous system detects environmental changes ...
, 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 Birth is the act or process of bearing or bringing forth offspring, also referred to in technical contexts as parturition. In mammals, the process is initiated by hormones which cause the muscular walls of the uterus to contract, expelling the ...
or hatching.


Etymology

First attested in English in the mid-14c., the word ''embryon'' derives from
Medieval Latin Medieval Latin was the form of Literary Latin used in Roman Catholic Western Europe during the Middle Ages. In this region it served as the primary written language, though local languages were also written to varying degrees. Latin function ...
''embryo'', itself from Greek (''embruon''), lit. "young one", which is the neuter of (''embruos''), lit. "growing in", from ἐν (''en''), "in" and βρύω (''bruō''), "swell, be full"; 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). The development of a zygote into a multicellular embryo proceeds through a series of recognizable stages, often divided into cleavage, blastula, gastrulation, and organogenesis. Cleavage is the period of rapid mitotic cell divisions that occur after fertilization. During cleavage, the overall size of the embryo does not change, but the size of individual cells decrease rapidly as they divide to increase the total number of cells. Cleavage results in a blastula. 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. 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. Before implanting into the uterine wall the embryo is sometimes known as the ''pre-implantation embryo'' or ''pre-implantation conceptus''. Sometimes this is called the pre-embryo a term employed to differentiate from an embryo proper in relation to embryonic stem cell discourses. 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
flatworm The flatworms, flat worms, Platyhelminthes, or platyhelminths (from the Greek πλατύ, ''platy'', meaning "flat" and ἕλμινς (root: ἑλμινθ-), ''helminth-'', meaning "worm") are a phylum of relatively simple bilaterian, unsegmen ...
s to humans) are called triploblastic. During gastrulation of triploblastic animals, the three germinal layers that form are called the ectoderm, mesoderm, and
endoderm Endoderm is the innermost of the three primary germ layers in the very early embryo. The other two layers are the ectoderm (outside layer) and mesoderm (middle layer). Cells migrating inward along the archenteron form the inner layer of the g ...
. All tissues and organs of a mature animal can trace their origin back to one of these layers. For example, the ectoderm will give rise to the skin epidermis and the nervous system, the mesoderm will give rise to the vascular system, muscles, bone, and connective tissues, and the endoderm will give rise to organs of the digestive system and epithelium of the digestive system and respiratory system. 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. 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. 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. 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, whereas in
zebrafish The zebrafish (''Danio rerio'') is a freshwater fish belonging to the minnow family (Cyprinidae) of the order Cypriniformes. Native to South Asia, it is a popular aquarium fish, frequently sold under the trade name zebra danio (and thus often ...
, embryonic development is considered finished when a bone called the
cleithrum The cleithrum (plural cleithra) is a membrane bone which first appears as part of the skeleton in primitive bony fish, where it runs vertically along the scapula. Its name is derived from Greek κλειθρον = "key (lock)", by analogy with "clav ...
becomes visible. 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 interpret the scope of
embryology Embryology (from Greek ἔμβρυον, ''embryon'', "the unborn, embryo"; and -λογία, ''-logia'') is the branch of animal biology that studies the prenatal development of gametes (sex cells), fertilization, and development of embryos ...
broadly as the study of the development of animals.


Plant embryos

Flowering plants ( angiosperms) create embryos after the fertilization of a haploid ovule by
pollen Pollen is a powdery substance produced by seed plants. It consists of pollen grains (highly reduced microgametophytes), which produce male gametes (sperm cells). Pollen grains have a hard coat made of sporopollenin that protects the gametoph ...
. The DNA from the ovule and pollen combine to form a diploid, single-cell zygote that will develop into an embryo. The zygote, which will divide multiple times as it progresses throughout embryonic development, is one part of a
seed A seed is an embryonic plant enclosed in a protective outer covering, along with a food reserve. The formation of the seed is a part of the process of reproduction in seed plants, the spermatophytes, including the gymnosperm and angiospe ...
. Other seed components include the
endosperm The endosperm is a tissue produced inside the seeds of most of the flowering plants following double fertilization. It is triploid (meaning three chromosome sets per nucleus) in most species, which may be auxin-driven. It surrounds the embryo ...
, 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 embryo with one small cell (the apical cell) and one large cell (the basal cell). The small, apical cell will eventually give rise to most of the structures of the mature plant, such as the stem, leaves, and roots. The larger basal cell will give rise to the suspensor, which connects the embryo to the endosperm so that nutrients can pass between them. 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. The dermal tissue will give rise to the
epidermis The epidermis is the outermost of the three layers that comprise the skin, the inner layers being the dermis and hypodermis. The epidermis layer provides a barrier to infection from environmental pathogens and regulates the amount of water ...
or outer covering of a plant, ground tissue will give rise to inner plant material that functions in
photosynthesis Photosynthesis is a process used by plants and other organisms to convert light energy into chemical energy that, through cellular respiration, can later be released to fuel the organism's activities. Some of this chemical energy is stored in ...
, resource storage, and physical support, and vascular tissue will give rise to connective tissue like the
xylem Xylem is one of the two types of transport tissue in vascular plants, the other being phloem. The basic function of xylem is to transport water from roots to stems and leaves, but it also transports nutrients. The word ''xylem'' is derived fr ...
and
phloem Phloem (, ) is the living tissue in vascular plants that transports the soluble organic compounds made during photosynthesis and known as ''photosynthates'', in particular the sugar sucrose, to the rest of the plant. This transport process i ...
that transport fluid, nutrients, and minerals throughout the plant. In heart stage, one or two
cotyledon A cotyledon (; ; ; , gen. (), ) is a significant part of the embryo within the seed of a plant, and is defined as "the embryonic leaf in seed-bearing plants, one or more of which are the first to appear from a germinating seed." The num ...
s (embryonic leaves) will form. Meristems (centers of
stem cell In multicellular organisms, stem cells are undifferentiated or partially differentiated cells that can differentiate into various types of cells and proliferate indefinitely to produce more of the same stem cell. They are the earliest type o ...
activity) develop during the torpedo stage, and will eventually produce many of the mature tissues of the adult plant throughout its life. At the end of embryonic growth, the seed will usually go dormant until germination. Once the embryo begins to germinate (grow out from the seed) and forms its first true leaf, it is called a
seedling A seedling is a young sporophyte developing out of a plant embryo from a seed. Seedling development starts with germination of the seed. A typical young seedling consists of three main parts: the radicle (embryonic root), the hypocotyl (embryo ...
or plantlet. Plants that produce
spore In biology, a spore is a unit of sexual or asexual reproduction that may be adapted for dispersal and for survival, often for extended periods of time, in unfavourable conditions. Spores form part of the life cycles of many plants, algae, ...
s instead of seeds, like
bryophyte The Bryophyta s.l. are a proposed taxonomic division containing three groups of non-vascular land plants ( embryophytes): the liverworts, hornworts and mosses. Bryophyta s.s. consists of the mosses only. They are characteristically limited in ...
s 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. 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. The structure and development of the rest of the embryo varies by group of plants. Since all land plants create embryos, they are collectively referred to as
embryophyte The Embryophyta (), or land plants, are the most familiar group of green plants that comprise vegetation on Earth. Embryophytes () have a common ancestor with green algae, having emerged within the Phragmoplastophyta clade of green algae as sis ...
s (or by their scientific name, Embryophyta). This, along with other characteristics, distinguishes land plants from other types of plants, such as
algae Algae (; singular alga ) is an informal term for a large and diverse group of photosynthetic eukaryotic organisms. It is a polyphyletic grouping that includes species from multiple distinct clades. Included organisms range from unicellular m ...
, which do not produce embryos.


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 cell In multicellular organisms, stem cells are undifferentiated or partially differentiated cells that can differentiate into various types of cells and proliferate indefinitely to produce more of the same stem cell. They are the earliest type o ...
s, evolution and development,
cell division Cell division is the process by which a parent cell divides into two daughter cells. Cell division usually occurs as part of a larger cell cycle in which the cell grows and replicates its chromosome(s) before dividing. In eukaryotes, there a ...
, and
gene expression Gene expression is the process by which information from a gene is used in the synthesis of a functional gene product that enables it to produce end products, protein or non-coding RNA, and ultimately affect a phenotype, as the final effect. T ...
. Examples of scientific discoveries made while studying embryos that were awarded the
Nobel Prize in Physiology or Medicine The Nobel Prize in Physiology or Medicine is awarded yearly by the Nobel Assembly at the Karolinska Institute, Nobel Assembly at the Karolinska Institutet, Karolinska Institute for outstanding discoveries in physiology or medicine. The Nobel Pr ...
include the
Spemann-Mangold organizer The Spemann-Mangold organizer is a group of cells that are responsible for the induction of the neural tissues during development in amphibian embryos. First described in 1924 by Hans Spemann and Hilde Mangold, the introduction of the organizer pro ...
, a group of cells originally discovered in amphibian embryos that give rise to neural tissues, and genes that give rise to body segments discovered in '' Drosophila'' fly embryos by Christiane Nüsslein-Volhard and
Eric Wieschaus Eric Francis Wieschaus (born June 8, 1947 in South Bend, Indiana) is an American evolutionary developmental biologist and 1995 Nobel Prize-winner. Early life Born in South Bend, Indiana, he attended John Carroll Catholic High School in Birmi ...
.


Assisted reproductive technology

Creating and/or manipulating embryos via
assisted reproductive technology Assisted reproductive technology (ART) includes medical procedures used primarily to address infertility. This subject involves procedures such as in vitro fertilization (IVF), intracytoplasmic sperm injection (ICSI), cryopreservation of gamete ...
(ART) is used for addressing fertility concerns in humans and other animals, and for
selective breeding Selective breeding (also called artificial selection) is the process by which humans use animal breeding and plant breeding to selectively develop particular phenotypic traits (characteristics) by choosing which typically animal or plant ma ...
in agricultural species. Between the years 1987 and 2015, ART techniques including in vitro fertilization (IVF) were responsible for an estimated 1 million human births in the United States alone. Other clinical technologies include
preimplantation genetic diagnosis Preimplantation genetic diagnosis (PGD or PIGD) is the genetic profiling of embryos prior to implantation (as a form of embryo profiling), and sometimes even of oocytes prior to fertilization. PGD is considered in a similar fashion to prenatal ...
(PGD), which can identify certain serious genetic abnormalities, such as
aneuploidy Aneuploidy is the presence of an abnormal number of chromosomes in a cell, for example a human cell having 45 or 47 chromosomes instead of the usual 46. It does not include a difference of one or more complete sets of chromosomes. A cell with an ...
, prior to selecting embryos for use in IVF. 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; however, this has been met with widespread condemnation from the scientific community. 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. 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. IVF and other ART techniques, including cloning via interspecies somatic cell nuclear transfer (iSCNT), are also used in attempts to increase the numbers of endangered or vulnerable species, such as Northern white rhinos,
cheetah The cheetah (''Acinonyx jubatus'') is a large cat native to Africa and central Iran. It is the fastest land animal, estimated to be capable of running at with the fastest reliably recorded speeds being , and as such has evolved specialize ...
s, and sturgeons.


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. Some large-scale animal species cryoconservation efforts include " frozen zoos" in various places around the world, including in the UK's Frozen Ark, the Breeding Centre for Endangered Arabian Wildlife (BCEAW) in the United Arab Emirates, and the San Diego Zoo Institute for Conservation in the United States. 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. The
Svalbard Global Seed Vault The Svalbard Global Seed Vault ( no, Svalbard globale frøhvelv) is a secure backup facility for the world's crop diversity on the Norwegian island of Spitsbergen in the remote Arctic Svalbard archipelago. The Seed Vault provides long-term stor ...
in Norway maintains the largest collection of plant reproductive tissue, with more than a million samples stored at .


Fossilized embryos

Fossilized animal embryos are known from the Precambrian, and are found in great numbers during the Cambrian period. Even fossilized
dinosaur Dinosaurs are a diverse group of reptiles of the clade Dinosauria. They first appeared during the Triassic period, between 243 and 233.23  million years ago (mya), although the exact origin and timing of the evolution of dinosaurs is ...
embryos have been discovered.


See also

* Embryo loss * Pregnancy *
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 dev ...
* In vitro fertilisation *
Pre-embryo In human embryonic development a pre-embryo is a conceptus before implantation in the uterus. Pre-embryo in human embryonic development The word pre-embryo sometimes is used in ethical contexts to refer to a human conceptus at least between fer ...
*
Miscarriage Miscarriage, also known in medical terms as a spontaneous abortion and pregnancy loss, is the death of an embryo or fetus before it is able to survive independently. Miscarriage before 6 weeks of gestation is defined by ESHRE as biochemical ...
*
Abortion Abortion is the termination of a pregnancy by removal or expulsion of an embryo or fetus. An abortion that occurs without intervention is known as a miscarriage or "spontaneous abortion"; these occur in approximately 30% to 40% of preg ...


Notes


External links


UNSW Embryology - Educational website

A Comparative Embryology Gallery
{{Authority control Developmental biology Bioethics Fertility medicine Articles containing video clips