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Anatomical terminology |
An embryo is the initial stage of development for 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 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 three germ layers that will form all of the different parts of the body. Neurulation forms the nervous 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.
First attested in English in the mid-14c., the word embryon derives from Medieval Latin embryo, itself from Greek ἔμβρυον (embruon), lit. "young one", [1] which is the neuter of ἔμβρυος (embruos), lit. "growing in", [2] from ἐν (en), "in" [3] and βρύω (bruō), "swell, be full"; [4] the proper Latinized form of the Greek term would be embryum.
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]
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. [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]
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 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]
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 embryo with one small cell (the apical cell) and one large cell (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]
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]
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 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 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]
Developmental biology is the study of the process by which animals and plants grow and develop. Developmental biology also encompasses the biology of regeneration, asexual reproduction, metamorphosis, and the growth and differentiation of stem cells in the adult organism.
Ontogeny is the origination and development of an organism, usually from the time of fertilization of the egg to adult. The term can also be used to refer to the study of the entirety of an organism's lifespan.
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. The sexual fusion of haploid cells is called karyogamy, the result of which is the formation of a diploid cell called the zygote or zygospore.
Development of the human body is the process of growth to maturity. The process begins with fertilization, where an egg released from the ovary of a female is penetrated by a sperm cell from a male. The resulting zygote develops through mitosis and cell differentiation, and the resulting embryo then implants in the uterus, where the embryo continues development through a fetal stage until birth. Further growth and development continues after birth, and includes both physical and psychological development that is influenced by genetic, hormonal, environmental and other factors. This continues throughout life: through childhood and adolescence into adulthood.
A genetic chimerism or chimera is a single organism composed of cells with more than one distinct genotype. Animal chimeras can be produced by the merger of two embryos. In plants and some animal chimeras, mosaicism involves distinct types of tissue that originated from the same zygote but differ due to mutation during ordinary cell division.
Blastulation is the stage in early animal embryonic development that produces the blastula. In mammalian development, the blastula develops into the blastocyst with a differentiated inner cell mass and an outer trophectoderm. The blastula is a hollow sphere of cells known as blastomeres surrounding an inner fluid-filled cavity called the blastocoel. Embryonic development begins with a sperm fertilizing an egg cell to become a zygote, which undergoes many cleavages to develop into a ball of cells called a morula. Only when the blastocoel is formed does the early embryo become a blastula. The blastula precedes the formation of the gastrula in which the germ layers of the embryo form.
Gastrulation is the stage in the early embryonic development of most animals, during which the blastula, or in mammals the blastocyst, is reorganized into a two-layered or three-layered embryo known as the gastrula. Before gastrulation, the embryo is a continuous epithelial sheet of cells; by the end of gastrulation, the embryo has begun differentiation to establish distinct cell lineages, set up the basic axes of the body, and internalized one or more cell types including the prospective gut.
Endoderm is the innermost of the three primary germ layers in the very early embryo. The other two layers are the ectoderm and mesoderm. Cells migrating inward along the archenteron form the inner layer of the gastrula, which develops into the endoderm.
The blastocyst is a structure formed in the early embryonic development of mammals. It possesses an inner cell mass (ICM) also known as the embryoblast which subsequently forms the embryo, and an outer layer of trophoblast cells called the trophectoderm. This layer surrounds the inner cell mass and a fluid-filled cavity or lumen known as the blastocoel. In the late blastocyst, the trophectoderm is known as the trophoblast. The trophoblast gives rise to the chorion and amnion, the two fetal membranes that surround the embryo. The placenta derives from the embryonic chorion and the underlying uterine tissue of the mother. The corresponding structure in non-mammalian animals is an undifferentiated ball of cells called the blastula.
In zoology and developmental anatomy, the notochord is an elastic, rod-like anatomical structure found in animals of the phylum Chordata. A notochord is one of five synapomorphies, or shared derived characteristics, used to identify a species as a chordate.
The endosperm is a tissue produced inside the seeds of most of the flowering plants following double fertilization. It is triploid in most species, which may be auxin-driven. It surrounds the embryo and provides nutrition in the form of starch, though it can also contain oils and protein. This can make endosperm a source of nutrition in animal diet. For example, wheat endosperm is ground into flour for bread, while barley endosperm is the main source of sugars for beer production. Other examples of endosperm that forms the bulk of the edible portion are coconut "meat" and coconut "water", and corn. Some plants, such as certain orchids, lack endosperm in their seeds.
The blastocoel, also spelled blastocoele and blastocele, and also called cleavage cavity, or segmentation cavity is a fluid-filled or yolk-filled cavity that forms in the blastula during very early embryonic development. At this stage in mammals the blastula is called the blastocyst, which consists of an outer epithelium, the trophectoderm, enveloping the inner cell mass and the blastocoel.
A germ layer is a primary layer of cells that forms during embryonic development. The three germ layers in vertebrates are particularly pronounced; however, all eumetazoans produce two or three primary germ layers. Some animals, like cnidarians, produce two germ layers making them diploblastic. Other animals such as bilaterians produce a third layer between these two layers, making them triploblastic. Germ layers eventually give rise to all of an animal's tissues and organs through the process of organogenesis.
Organogenesis is the phase of embryonic development that starts at the end of gastrulation and continues until birth. During organogenesis, the three germ layers formed from gastrulation form the internal organs of the organism.
Plant embryonic development, also plant embryogenesis, is a process that occurs after the fertilization of an ovule to produce a fully developed plant embryo. This is a pertinent stage in the plant life cycle that is followed by dormancy and germination. The zygote produced after fertilization must undergo various cellular divisions and differentiations to become a mature embryo. An end stage embryo has five major components including the shoot apical meristem, hypocotyl, root meristem, root cap, and cotyledons. Unlike the embryonic development in animals, and specifically in humans, plant embryonic development results in an immature form of the plant, lacking most structures like leaves, stems, and reproductive structures. However, both plants and animals including humans, pass through a phylotypic stage that evolved independently and that causes a developmental constraint limiting morphological diversification.
In developmental biology, animal embryonic development, also known as animal embryogenesis, is the developmental stage of an animal embryo. Embryonic development starts with the fertilization of an egg cell (ovum) by a sperm cell (spermatozoon). Once fertilized, the ovum becomes a single diploid cell known as a zygote. The zygote undergoes mitotic divisions with no significant growth and cellular differentiation, leading to development of a multicellular embryo after passing through an organizational checkpoint during mid-embryogenesis. In mammals, the term refers chiefly to the early stages of prenatal development, whereas the terms fetus and fetal development describe later stages.
A neurula is a vertebrate embryo at the early stage of development in which neurulation occurs. The neurula stage is preceded by the gastrula stage; consequentially, neurulation is preceded by gastrulation. Neurulation marks the beginning of the process of organogenesis.
The development of fishes is unique in some specific aspects compared to the development of other animals.
Human embryonic development or human embryogenesis is the development and formation of the human embryo. It is characterised by the processes of cell division and cellular differentiation of the embryo that occurs during the early stages of development. In biological terms, the development of the human body entails growth from a one-celled zygote to an adult human being. Fertilization occurs when the sperm cell successfully enters and fuses with an egg cell (ovum). The genetic material of the sperm and egg then combine to form the single cell zygote and the germinal stage of development commences. Human embryonic development covers the first eight weeks of development, which have 23 stages, called Carnegie stages. At the beginning of the ninth week, the embryo is termed a fetus. In comparison to the embryo, the fetus has more recognizable external features and a more complete set of developing organs.
This glossary of developmental biology is a list of definitions of terms and concepts commonly used in the study of developmental biology and related disciplines in biology, including embryology and reproductive biology, primarily as they pertain to vertebrate animals and particularly to humans and other mammals. The developmental biology of invertebrates, plants, fungi, and other organisms is treated in other articles; e.g terms relating to the reproduction and development of insects are listed in Glossary of entomology, and those relating to plants are listed in Glossary of botany.