Freemartin
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A freemartin or free-martin (sometimes martin heifer) is an infertile
female Female ( symbol: ♀) is the sex of an organism that produces the large non-motile ova (egg cells), the type of gamete (sex cell) that fuses with the male gamete during sexual reproduction. A female has larger gametes than a male. Females ...
cattle Cattle (''Bos taurus'') are large, domesticated, cloven-hooved, herbivores. They are a prominent modern member of the subfamily Bovinae and the most widespread species of the genus ''Bos''. Adult females are referred to as cows and adult ma ...
with
masculinized Virilization or masculinization is the biological development of adult male characteristics in young males or females. Most of the changes of virilization are produced by androgens. Virilization is most commonly used in three medical and biology ...
behavior and non-functioning
ovaries The ovary is an organ in the female reproductive system that produces an ovum. When released, this travels down the fallopian tube into the uterus, where it may become fertilized by a sperm. There is an ovary () found on each side of the body. T ...
. Phenotypically, the animal appears female, but various aspects of female reproductive development are altered due to acquisition of
anti-Müllerian hormone Anti-Müllerian hormone (AMH), also known as Müllerian-inhibiting hormone (MIH), is a glycoprotein hormone structurally related to inhibin and activin from the transforming growth factor beta superfamily, whose key roles are in growth differen ...
from the male
twin Twins are two offspring produced by the same pregnancy.MedicineNet > Definition of TwinLast Editorial Review: 19 June 2000 Twins can be either ''monozygotic'' ('identical'), meaning that they develop from one zygote, which splits and forms two em ...
. Genetically, the animal is chimeric: karyotypy of a sample of cells shows XX/XY chromosomes. The animal originates as a female (XX), but acquires the male (XY) component in utero by exchange of some cellular material from a male twin, via vascular connections between placentas: an example of
microchimerism Microchimerism is the presence of a small number of cells in an individual that have originated from another individual and are therefore genetically distinct. This phenomenon may be related to certain types of autoimmune diseases although the res ...
.''In utero'' cell transfer between porcine littermates
in ''Reproduction, Fertility, and Development'' (2011; 23(2): 297–302. - doi: 10.1071/RD10165)
The chimerism is mainly present in the
hematopoietic stem cells Hematopoietic stem cells (HSCs) are the stem cells that give rise to other blood cells. This process is called haematopoiesis. In vertebrates, the very first definitive HSCs arise from the ventral endothelial wall of the embryonic aorta within ...
.


History

Freemartins are known to have been described by the Roman writer
Varro Marcus Terentius Varro (; 116–27 BC) was a Roman polymath and a prolific author. He is regarded as ancient Rome's greatest scholar, and was described by Petrarch as "the third great light of Rome" (after Vergil and Cicero). He is sometimes calle ...
, who called them .EARLY RECOGNITION OF THE FREEMARTIN CONDITION IN HEIFERS TWINBORN WITH BULLS
by W.W. Swett, C.A. Matthews, and R. R. Graves, in the Journal of Agricultural Research, vol. 61, no. 8 (October 15, 1940)
The 18th-century physician John Hunter discovered that a freemartin always has a male twin. It was hypothesized early in the 20th century that masculinizing factors travel from the male twin to the female twin through the vascular connections of the placenta because of the vascular fusion and affect the internal anatomy of the female. Several researchers made the discovery that a freemartin results when a female fetus has its
chorion The chorion is the outermost fetal membrane around the embryo in mammals, birds and reptiles (amniotes). It develops from an outer fold on the surface of the yolk sac, which lies outside the zona pellucida (in mammals), known as the vitell ...
fuse in the
uterus The uterus (from Latin ''uterus'', plural ''uteri'') or womb () is the organ in the reproductive system of most female mammals, including humans that accommodates the embryonic and fetal development of one or more embryos until birth. The ...
with that of a male twin. The result was published in 1916 by Tandler and Keller. The discovery was made independently by American biologist Frank R. Lillie, who published it in ''
Science Science is a systematic endeavor that Scientific method, builds and organizes knowledge in the form of Testability, testable explanations and predictions about the universe. Science may be as old as the human species, and some of the earli ...
'' in 1916. Both teams are now credited with the discovery. In rural areas
folklore Folklore is shared by a particular group of people; it encompasses the traditions common to that culture, subculture or group. This includes oral traditions such as tales, legends, proverbs and jokes. They include material culture, ranging ...
often claimed this condition was not just peculiar to cattle, but extended also to human twins; this belief perpetuated for generations, as was mentioned in the writings of
Cuthbert Bede Edward Bradley (25 March 1827 – 12 December 1889) was an English clergyman and novelist. He was born in Kidderminster and educated at Durham University from which he took his pen name Cuthbert Bede, B.A. His most popular book was '' The Adve ...
.


Etymology

The etymology of the term "freemartin" is uncertain: speculations include that "free" may indicate "willing" (referring to the freemartin's willingness to work) or "exempt from reproduction" (referring to its sterility, or to a farmer's decision to not bother trying to breed a freemartin, or both), or that it may be derived from a
Flemish Flemish (''Vlaams'') is a Low Franconian dialect cluster of the Dutch language. It is sometimes referred to as Flemish Dutch (), Belgian Dutch ( ), or Southern Dutch (). Flemish is native to Flanders, a historical region in northern Belgium; ...
word for a cow which gives no milk and/or has ceased to be capable of bearing offspring; "martin" is generally held to derive from an Irish or Gaelic word for "cow" or "heifer", although connections to
Martinmas Saint Martin's Day or Martinmas, sometimes historically called Old Halloween or Old Hallowmas Eve, is the feast day of Saint Martin of Tours and is celebrated in the liturgical year on 11 November. In the Middle Ages and early modern period, ...
have also been posited.


Mechanism

In most cattle twins, the blood vessels in the chorions become interconnected, creating a shared circulation for both twins. If both fetuses are the same sex this is of no significance, but if they are different, male
hormone A hormone (from the Greek participle , "setting in motion") is a class of signaling molecules in multicellular organisms that are sent to distant organs by complex biological processes to regulate physiology and behavior. Hormones are require ...
s pass from the male twin to the female twin. The male hormones (testosterone and anti-Müllerian hormone) then masculinize the female twin, and the result is a freemartin. The degree of masculinization is greater if the fusion occurs earlier in the pregnancy – in about ten percent of cases no fusion takes place and the female remains fertile. The male twin is largely unaffected by the fusion, although the size of the testicles may be slightly reduced. Testicle size is associated with fertility, so there may be some reduction in bull fertility. Freemartins behave and grow in a similar way to
castrated Castration is any action, surgical, chemical, or otherwise, by which an individual loses use of the testicles: the male gonad. Surgical castration is bilateral orchiectomy (excision of both testicles), while chemical castration uses pharmac ...
male cattle (steers).


Diagnosis

If suspected, a test can be done to detect the presence of the male Y-chromosomes in some circulating white blood cells of the subject. Genetic testing for the Y-chromosome can be performed within days of birth and can aid in the early identification of a sterile female bovine. Physical examination of the calf may also reveal differences: a subjective assessment is that frequently there is a lengthened tuft of hair at the ventral tip of the vulva in a freemartin heifer atypical in fertile heifer calves. Also, often many (but not all) freemartins will have a shortened length of
vagina In mammals, the vagina is the elastic, muscular part of the female genital tract. In humans, it extends from the vestibule to the cervix. The outer vaginal opening is normally partly covered by a thin layer of mucosal tissue called the hymen ...
compared with that of a fertile heifer. Commercial probes are available to check heifers for obvious freemartinism in lieu of doing a blood test


Other animals

A freemartin is the normal outcome of mixed twins in all cattle species which have been studied. It does not normally occur in most other mammals, though it has been recorded in
sheep Sheep or domestic sheep (''Ovis aries'') are domesticated, ruminant mammals typically kept as livestock. Although the term ''sheep'' can apply to other species in the genus '' Ovis'', in everyday usage it almost always refers to domesticated ...
,
goat The goat or domestic goat (''Capra hircus'') is a domesticated species of goat-antelope typically kept as livestock. It was domesticated from the wild goat (''C. aegagrus'') of Southwest Asia and Eastern Europe. The goat is a member of the a ...
s, and
pig The pig (''Sus domesticus''), often called swine, hog, or domestic pig when distinguishing from other members of the genus '' Sus'', is an omnivorous, domesticated, even-toed, hoofed mammal. It is variously considered a subspecies of ''Sus ...
s.


Uses

Freemartins are occasionally used in stem cell and
immunology Immunology is a branch of medicineImmunology for Medical Students, Roderick Nairn, Matthew Helbert, Mosby, 2007 and biology that covers the medical study of immune systems in humans, animals, plants and sapient species. In such we can see the ...
research. During fetal development cells are exchanged between the fused circulations of the bovine twins. Up to 95% of the freemartin's blood cells can be derived from those of her twin brother. Male-derived cells and their progeny can be easily visualised in the freemartin tissues, as only they contain the male
Y chromosome The Y chromosome is one of two sex chromosomes (allosomes) in therian mammals, including humans, and many other animals. The other is the X chromosome. Y is normally the sex-determining chromosome in many species, since it is the presence or abse ...
. Thus, by analysing these tissues, one can investigate the capacity of hematopoietic stem cells or other circulating cells to produce other tissues in addition to blood. The freemartin model allows one to analyse perfectly healthy and unmanipulated animals, without resorting to transplantation often used in stem cell research.


Fictional use

* In the
Aldous Huxley Aldous Leonard Huxley (26 July 1894 – 22 November 1963) was an English writer and philosopher. He wrote nearly 50 books, both novels and non-fiction works, as well as wide-ranging essays, narratives, and poems. Born into the prominent Huxle ...
novel ''
Brave New World ''Brave New World'' is a dystopian novel by English author Aldous Huxley, written in 1931 and published in 1932. Largely set in a futuristic World State, whose citizens are environmentally engineered into an intelligence-based social hiera ...
'', a "freemartin" (mentioned in chapters 1, 3, 11 and 17) is a woman who has been deliberately made sterile by exposure to hormones during fetal development; in the book, government policy requires freemartins to constitute 70% of the female population. A side effect of this is some freemartin women having developed beards. * The
Robert A. Heinlein Robert Anson Heinlein (; July 7, 1907 – May 8, 1988) was an American science fiction author, aeronautical engineer, and naval officer. Sometimes called the "dean of science fiction writers", he was among the first to emphasize scientific accu ...
novel ''
Beyond This Horizon ''Beyond This Horizon'' is a science fiction novel by American writer Robert A. Heinlein. It was originally published as a two-part serial in ''Astounding Science Fiction'' (April, May 1942, under the pseudonym Anson MacDonald) and then as a sin ...
'' lists "the clever and repulsively beautiful pseudo-feminine freemartins" as one of the genetically-engineered specialist types of humans that were created in the "Empire of the Great Khans" (chapter 2). *In the Robert Heinlein novel '' Farnham's Freehold'', the protagonist, Hugh Farnham, is given a companion ("bedwarmer") that is described as a "natural freemartin". *In the
Avram Davidson Abraham, ; ar, , , name=, group= (originally Abram) is the common Hebrew patriarch of the Abrahamic religions, including Judaism, Christianity, and Islam. In Judaism, he is the founding father of the special relationship between the Jews ...
story "The House the Blakeneys Built", the cattle are freemartins. *In the fantasy book series ''
Bazil Broketail ''Bazil Broketail'' is a 1992 fantasy novel by author Christopher Rowley, the name of its main character, and the name of the further series of books featuring him. Bazil is a leatherback dragon A dragon is a reptilian legendary creature tha ...
'' by
Christopher Rowley Christopher Rowley (born 1948) is an American writer specializing in the science fiction and fantasy genres. He is also a former journalist and television screenswriter. Christopher Rowley was born in 1948 in Lynn, Massachusetts, to an Ameri ...
, "freemartin" is the name for a breed of sterile female dragons. *In ''
Footfall ''Footfall'' is a 1985 science fiction novel by American writers Larry Niven and Jerry Pournelle. The book depicts the arrival of members of an alien species called the Fithp that have traveled to the Solar System from Alpha Centauri in a large ...
'' by
Larry Niven Laurence van Cott Niven (; born April 30, 1938) is an American science fiction writer. His best-known works are '' Ringworld'' (1970), which received Hugo, Locus, Ditmar, and Nebula awards, and, with Jerry Pournelle, '' The Mote in God's E ...
and
Jerry Pournelle Jerry Eugene Pournelle (; August 7, 1933 – September 8, 2017) was an American scientist in the area of operations research and human factors research, a science fiction writer, essayist, journalist, and one of the first bloggers. In the 1960s ...
, a lithely-built human woman uses the term "freemartin" to describe herself, while doubting her sexuality. *In
Nicola Griffith Nicola Griffith (; born 30 September 1960) is a British-American novelist, essayist, and teacher. She has won the Washington State Book Award, Nebula Award, James Tiptree, Jr. Award, World Fantasy Award and six Lambda Literary Awards. Person ...
's novel ''
Hild Hild or Hildr may refer to: * Hildr or Hild is one of the Valkyries in Norse mythology, a personification of battle * Hild or Hilda of Whitby is a Christian saint who was a British abbess and nun in the Middle Ages * Hild (Oh My Goddess!), the ult ...
'', the title character is sometimes referred to by others as a freemartin, in reference to her non-feminine character and social role. * In Lauren Groff's novel ''Matrix'' (New York: Riverhead Books, 2021), 12th-century English peasants use freemartins to pull wagons.


References


External links


Mosaicism and Chimerism at colostate.edu
{{Webarchive, url=https://web.archive.org/web/20060330180240/http://arbl.cvmbs.colostate.edu/hbooks/genetics/medgen/chromo/mosaics.html , date=2006-03-30
Picture at agric.gov.ab.ca
Bovine health Twin Female mammals