Ovariole

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A generalized insect ovariole of the panoistic type, showing the germarium, developing oocytpes in follicles, and the oviduct. Insect ovariole diagram 2.svg
A generalized insect ovariole of the panoistic type, showing the germarium, developing oocytpes in follicles, and the oviduct.

An ovariole is a tubular component of the insect ovary, and the basic unit of egg production. [1] Each ovariole is composed of a germarium (the germline stem cell niche) at the anterior tip, a set of developing oocytes contained within follicles, and a posterior connection to a common oviduct. [2] While most insects have two ovaries, the number of ovarioles within each ovary varies across insect species. [3] This number may also be variable across individuals within a species, or between the left and right ovaries within an individual.

Contents

Types

Types of ovarioles: panoistic lack nurse cells, meroistic have nurse cells that are located either adjacent to the oocyte (polytrophic) or in the germarium (telotrophic). Ovariole type diagram.svg
Types of ovarioles: panoistic lack nurse cells, meroistic have nurse cells that are located either adjacent to the oocyte (polytrophic) or in the germarium (telotrophic).

Ovarioles are often classified into one of several types by the presence and position of nurse cells. [2] These specialized cells provide nutrition and molecules important for embryonic patterning to the developing oocyte. Ovarioles that lack nurse cells are referred to as panoistic and ovarioles with nurse cells are referred to as meroistic.

Meroistic ovarioles are further classified according to where nurse cells are located. [2] In polytrophic meroistic ovarioles, nurse cells are adjacent to the developing oocyte. In telotrophic meroistic ovarioles, nurse cells are located in the germarium and connect to developing ooctypes via nutritive cords.

In Drosophila melanogaster

A diagram of the ovary of the fruit fly Drosophila melanogaster, showing multiple ovarioles within the two ovaries, each containing multiple developing oocytes. DrosophilaOvary.png
A diagram of the ovary of the fruit fly Drosophila melanogaster, showing multiple ovarioles within the two ovaries, each containing multiple developing oocytes.

In the fruit fly Drosophila melanogaster , a common model organism for developmental research, each ovary typically contains between 16 and 20 polytrophic meroistic ovarioles. [4] These ovarioles continuously produce eggs through division and differentiation of the germline stem cells, located in the anterior tip of the germarium. There are also several populations of somatic support cells in the germarium, including terminal filament cells, cap cells, and anterior escort cells.

The process of oogenesis within the Drosophila ovariole has been divided into 14 identifiable stages. [5] Developing oocytes are arranged within the ovariole in an ontogenic series, with early stage oocytes toward the anterior and later stage oocytes posterior. At the end of stage 14, the egg passes through the lateral oviduct before entering the common oviduct and then exiting via the uterus.

Related Research Articles

<span class="mw-page-title-main">Egg cell</span> Female reproductive cell in most anisogamous organisms

The egg cell or ovum is the female reproductive cell, or gamete, in most anisogamous organisms. The term is used when the female gamete is not capable of movement (non-motile). If the male gamete (sperm) is capable of movement, the type of sexual reproduction is also classified as oogamous. A nonmotile female gamete formed in the oogonium of some algae, fungi, oomycetes, or bryophytes is an oosphere. When fertilized, the oosphere becomes the oospore.

A maternal effect is a situation where the phenotype of an organism is determined not only by the environment it experiences and its genotype, but also by the environment and genotype of its mother. In genetics, maternal effects occur when an organism shows the phenotype expected from the genotype of the mother, irrespective of its own genotype, often due to the mother supplying messenger RNA or proteins to the egg. Maternal effects can also be caused by the maternal environment independent of genotype, sometimes controlling the size, sex, or behaviour of the offspring. These adaptive maternal effects lead to phenotypes of offspring that increase their fitness. Further, it introduces the concept of phenotypic plasticity, an important evolutionary concept. It has been proposed that maternal effects are important for the evolution of adaptive responses to environmental heterogeneity.

<span class="mw-page-title-main">Germ cell</span> Gamete-producing cell

A germ cell is any cell that gives rise to the gametes of an organism that reproduces sexually. In many animals, the germ cells originate in the primitive streak and migrate via the gut of an embryo to the developing gonads. There, they undergo meiosis, followed by cellular differentiation into mature gametes, either eggs or sperm. Unlike animals, plants do not have germ cells designated in early development. Instead, germ cells can arise from somatic cells in the adult, such as the floral meristem of flowering plants.

<span class="mw-page-title-main">Oogenesis</span> Egg cell production process

Oogenesis, ovogenesis, or oögenesis is the differentiation of the ovum into a cell competent to further develop when fertilized. It is developed from the primary oocyte by maturation. Oogenesis is initiated in the embryonic stage.

<i>Drosophila</i> embryogenesis Embryogenesis of the fruit fly Drosophila, a popular model system

Drosophila embryogenesis, the process by which Drosophila embryos form, is a favorite model system for genetics and developmental biology. The study of its embryogenesis unlocked the century-long puzzle of how development was controlled, creating the field of evolutionary developmental biology. The small size, short generation time, and large brood size make it ideal for genetic studies. Transparent embryos facilitate developmental studies. Drosophila melanogaster was introduced into the field of genetic experiments by Thomas Hunt Morgan in 1909.

The border cells are a cluster of 6–8 cells that migrate in the ovariole of the fruit-fly Drosophila melanogaster, during the process of oogenesis. A fly ovary consists of a string of ovarioles or egg chambers arranged in an increasing order of maturity. Each egg chamber contains 16 central germline, nurse cells surrounded by a monolayer epithelium of nearly 1000 follicle cells. At stage 8 of oogenesis, these cells initiate invading the neighbouring nurse cells, and reach the oocyte boundary by Stage 10.

<span class="mw-page-title-main">Gurken localisation signal</span>

mRNA localization is a common mode of posttranscriptional regulation of gene expression that targets a protein to its site of function. Proteins are highly dependent on cellular environments for stability and function, therefore, mRNA localization signals are crucial for maintaining protein function. The Gurken localisation signal is an RNA regulatory element conserved across many species of Drosophila. The element consists of an RNA stem loop within the coding region of the messenger RNA that forms a signal for dynein-mediated Gurken mRNA transport to the dorsoanterior cap near the nucleus of the oocyte.

Orthodenticle (otd) is a homeobox gene found in Drosophila that regulates the development of anterior patterning, with particular involvement in the central nervous system function and eye development. It is located on the X chromosome. The gene is an ortholog of the human OTX1/OTX2 gene.

<span class="mw-page-title-main">Vitelline envelope</span> Outer proteinaceous layer outside the oocyte and egg

The insect vitelline envelope is the outer proteinaceous layer outside the oocyte and egg. The vitelline envelope, not being a cellular structure, is commonly referred to as a membrane. However, this is a technical misnomer as the structure is composed of protein and is not a cellular component. It varies in thickness between different insects and even varies at different parts of the egg. It lies inside the outer shell of the egg, which is commonly referred to as the chorion.

Staufen is a protein product of a maternally expressed gene first identified in Drosophila melanogaster. The protein has been implicated in helping regulate genes important in determination of gradients that set up the anterior posterior axis such as bicoid and oskar. Staufen proteins, abbreviated Stau, are necessary for cell localization during the oogenesis and zygotic development. It is involved in targeting of the messenger RNA encoding these genes to the correct pole of the egg cell.

Vasa is an RNA binding protein with an ATP-dependent RNA helicase that is a member of the DEAD box family of proteins. The vasa gene is essential for germ cell development and was first identified in Drosophila melanogaster, but has since been found to be conserved in a variety of vertebrates and invertebrates including humans. The Vasa protein is found primarily in germ cells in embryos and adults, where it is involved in germ cell determination and function, as well as in multipotent stem cells, where its exact function is unknown.

mir-279 is a short RNA molecule found in Drosophila melanogaster that belongs to a class of molecules known as microRNAs. microRNAs are ~22nt-long non-coding RNAs that post-transcriptionally regulate the expression of genes, often by binding to the 3' untranslated region of mRNA, targeting the transcript for degradation. miR-279 has diverse tissue-specific functions in the fly, influencing developmental processes related to neurogenesis and oogenesis, as well as behavioral processes related to circadian rhythms. The varied roles of mir-279, both in the developing and adult fly, highlight the utility of microRNAs in regulating unique biological processes.

The gene Maelstrom, Mael, creates a protein, which was first located in Drosophila melanogaster in the nuage perinuclear structure and has functionality analogous to the spindle, spn, gene class. Its mammalian homolog is MAEL.

Most insects reproduce oviparously, i.e. by laying eggs. The eggs are produced by the female in a pair of ovaries. Sperm, produced by the male in one testicle or more commonly two, is transmitted to the female during mating by means of external genitalia. The sperm is stored within the female in one or more spermathecae. At the time of fertilization, the eggs travel along oviducts to be fertilized by the sperm and are then expelled from the body ("laid"), in most cases via an ovipositor.

<i>Homeotic protein bicoid</i> Protein-coding gene in the species Drosophila melanogaster

Homeotic protein bicoid is encoded by the bcd maternal effect gene in Drosophilia. Homeotic protein bicoid concentration gradient patterns the anterior-posterior (A-P) axis during Drosophila embryogenesis. Bicoid was the first protein demonstrated to act as a morphogen. Although bicoid is important for the development of Drosophila and other higher dipterans, it is absent from most other insects, where its role is accomplished by other genes.

Oogonial stem cells (OSCs), also known as egg precursor cells or female germline cells, are diploid germline cells with stem cell characteristics: the ability to renew and differentiate into other cell types, different from their tissue of origin. Present in invertebrates and some lower vertebrate species, they have been extensively studied in Caenorhabditis elegans, Drosophila melanogaster. OSCs allow the production of new female reproductive cells (oocytes) by the process of oogenesis during an organism's reproductive life.

<span class="mw-page-title-main">Fusome</span>

The fusome is a membranous structure found in the developing germ cell cysts of many insect orders. Initial description of the fusome occurred in the 19th century and since then the fusome has been extensively studied in Drosophila melanogaster male and female germline development. This structure has roles in maintaining germline cysts, coordinating the number of mitotic divisions prior to meiosis, and oocyte determination by serving as a structure for intercellular communication.

<i>Sarcophaga barbata</i> Fly species

Sarcophaga barbata is a species from the genus Sarcophaga and the family of flesh fly, Sarcophagidae. It is most closely related to S. plinthopyga, S. securifera, and S. bullata of the same genus. The species was first discovered by Eugene Thomson in 1868. S. barbata has also been found in the Middle East near carcasses, where the larvae can thrive. S. barbata is also a prominent organism in scientific research and has been used to study L-3-glycerophosphate oxidation and location within the mitochondria.

<span class="mw-page-title-main">Ovarian stem cell</span>

Ovarian stem cells are oocytes formed in ovarian follicle before birth in female mammals. They do not form post-natally, and are depleted throughout reproductive life. In humans it is estimated that 500,000–1,000,000 primordial follicles are present at birth, decreasing rapidly with age until roughly age 51 when ovulation stops, resulting in menopause. The origin of these oocytes remains under discussion. The publication of a study in 2004 proposing germ cell renewal in adult mice sparked a debate on the possibility of stem cells in the postnatal ovary. An increasing number of studies suggest that stem cells exist within the mammalian ovary and can be manipulated in vitro to produce oocytes, but whether such ovarian stem cells have the potential to differentiate into oocytes remains uncertain.

The germ cell nest forms in the ovaries during their development. The nest consists of multiple interconnected oogonia formed by incomplete cell division. The interconnected oogonia are surrounded by somatic cells called granulosa cells. Later on in development, the germ cell nests break down through invasion of granulosa cells. The result is individual oogonia surrounded by a single layer of granulosa cells. There is also a comparative germ cell nest structure in the developing spermatogonia, with interconnected intracellular cytoplasmic bridges.

References

  1. King, Robert C.; Mulligan, Pamela K.; Stansfield, William D. (2013). A dictionary of genetics (8th ed.). Oxford University Press. doi:10.1093/acref/9780199766444.001.0001. ISBN   9780199376865.
  2. 1 2 3 Büning, Jürgen (1994-07-31). The Insect Ovary: Ultrastructure, previtellogenic growth and evolution. Springer Science & Business Media. ISBN   978-0-412-36080-0.
  3. Robertson, J. G. (2011-02-14). "Ovariole numbers in Coleoptera". Canadian Journal of Zoology. 39 (3): 245–263. doi:10.1139/z61-028.
  4. Gilboa, Lilach (2015-06-01). "Organizing stem cell units in the Drosophila ovary". Current Opinion in Genetics & Development. Developmental mechanisms, patterning and organogenesis. 32: 31–36. doi:10.1016/j.gde.2015.01.005. ISSN   0959-437X. PMID   25703842.
  5. McLaughlin, John M.; Bratu, Diana P. (2015), Bratu, Diana P.; McNeil, Gerard P. (eds.), "Drosophila melanogaster Oogenesis: An Overview", Drosophila Oogenesis: Methods and Protocols, vol. 1328, New York, NY: Springer, pp. 1–20, doi:10.1007/978-1-4939-2851-4_1, ISBN   978-1-4939-2851-4, PMID   26324426 , retrieved 2022-06-21