Sex Ratio
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The sex ratio (or gender ratio) is usually defined as the
ratio In mathematics, a ratio shows how many times one number contains another. For example, if there are eight oranges and six lemons in a bowl of fruit, then the ratio of oranges to lemons is eight to six (that is, 8:6, which is equivalent to the ...
of
male Male (symbol: ♂) is the sex of an organism that produces the gamete (sex cell) known as sperm, which fuses with the larger female gamete, or ovum, in the process of fertilization. A male organism cannot reproduce sexually without access to ...
s to
female Female (Venus symbol, symbol: ♀) is the sex of an organism that produces the large non-motile ovum, ova (egg cells), the type of gamete (sex cell) that fuses with the Sperm, male gamete during sexual reproduction. A female has larger gamet ...
s in a
population Population typically refers to the number of people in a single area, whether it be a city or town, region, country, continent, or the world. Governments typically quantify the size of the resident population within their jurisdiction using a ...
. As explained by
Fisher's principle Fisher's principle is an evolutionary model that explains why the sex ratio of most species that produce offspring through sexual reproduction is approximately 1:1 between males and females. A. W. F. Edwards has remarked that it is "probably the mo ...
, for evolutionary reasons this is typically about 1:1 in species which reproduce sexually. Many species deviate from an even sex ratio, either periodically or permanently. Examples include
parthenogenic Parthenogenesis (; from the Greek grc, παρθένος, translit=parthénos, lit=virgin, label=none + grc, γένεσις, translit=génesis, lit=creation, label=none) is a natural form of asexual reproduction in which growth and development ...
species, periodically mating organisms such as aphids, some
eusocial Eusociality (from Greek εὖ ''eu'' "good" and social), the highest level of organization of sociality, is defined by the following characteristics: cooperative brood care (including care of offspring from other individuals), overlapping generat ...
wasp A wasp is any insect of the narrow-waisted suborder Apocrita of the order Hymenoptera which is neither a bee nor an ant; this excludes the broad-waisted sawflies (Symphyta), which look somewhat like wasps, but are in a separate suborder. Th ...
s, bees,
ant Ants are eusocial insects of the family Formicidae and, along with the related wasps and bees, belong to the order Hymenoptera. Ants evolved from vespoid wasp ancestors in the Cretaceous period. More than 13,800 of an estimated total of ...
s, and
termite Termites are small insects that live in colonies and have distinct castes (eusocial) and feed on wood or other dead plant matter. Termites comprise the infraorder Isoptera, or alternatively the epifamily Termitoidae, within the order Blatto ...
s. The
human sex ratio In anthropology and demography, the human sex ratio is the ratio of males to females in a population. Like most sexual species, the sex ratio in humans is close to 1:1. In humans, the natural ratio at birth between males and females is sligh ...
is of particular interest to anthropologists and demographers. In human societies, sex ratios at birth may be considerably skewed by factors such as the age of mother at birth and by
sex-selective abortion Sex-selective abortion is the practice of terminating a pregnancy based upon the predicted sex of the infant. The selective abortion of female fetuses is most common where male children are valued over female children, especially in parts of Eas ...
and infanticide. Exposure to pesticides and other environmental contaminants may be a significant contributing factor as well. As of 2014, the global sex ratio at birth is estimated at 107 boys to 100 girls (1,000 boys per 934 girls)..


Types

In most species, the sex ratio varies according to the age profile of the population. It is generally divided into four subdivisions: * — ratio at fertilization * — ratio at birth * — ratio in sexually mature organisms :This is equivalent to the (ASR), which is defined as the ratio of adult males to females in a population. :*The
operational sex ratio In the evolutionary biology of sexual reproduction, operational sex ratio (OSR) is the ratio of sexually competing males that are ready to mate to sexually competing females that are ready to mate, or alternatively the local ratio of fertilizable fe ...
(OSR) is the ratio of ''sexually active'' males to females in a population, and is therefore derived from a subset of the individuals included when calculating the ASR. Although some research has equated these two values, they are conceptually different. * — ratio in post-reproductive organisms These definitions can be somewhat subjective since they lack clear boundaries.


Sex ratio theory

Sex ratio theory is a field of academic study which seeks to understand the sex ratios observed in nature from an evolutionary perspective. It continues to be heavily influenced by the work of
Eric Charnov Eric Lee Charnov (born October 29, 1947) is an American evolutionary ecologist. He is best known for his work on foraging, especially the marginal value theorem, and life history theory, especially sex allocation and scaling/allometric rules. ...
. He defines five major questions, both for his book and the field in general (slightly abbreviated here): #For a
dioecious Dioecy (; ; adj. dioecious , ) is a characteristic of a species, meaning that it has distinct individual organisms (unisexual) that produce male or female gametes, either directly (in animals) or indirectly (in seed plants). Dioecious reproductio ...
species, what is the equilibrium sex ratio maintained by natural selection? #For a
sequential hermaphrodite Sequential hermaphroditism (called dichogamy in botany) is a type of hermaphroditism that occurs in many fish, gastropods, and plants. Sequential hermaphroditism occurs when the individual changes its sex at some point in its life. In particular, ...
, what is the equilibrium sex order and time of
sex change Sex change is a natural or artificial process in which an individual's sex is changed. Sex change may also refer to: Biology and medicine *Sex reassignment therapy *Sex reassignment surgery *Sequential hermaphroditism, a phenomenon whereby some ...
? #For a
simultaneous hermaphrodite Simultaneous hermaphroditism is one of the two types of hermaphroditism, the other type being sequential hermaphroditism. In this form of hermaphroditism an individual has sex organs of both sexes and can produce both gamete types even in the sa ...
, what is the equilibrium allocation of resources to male versus female function in each breeding season? #Under what conditions are the various states of hermaphroditism or dioecy evolutionarily stable? When is a mixture of sexual types stable? #When does selection favour the ability of an individual to alter its allocation to male versus female function, in response to particular environmental or life history situations? Biological research mostly concerns itself with sex ''allocation'' rather than sex ratio, sex allocation denoting the allocation of energy to either sex. Common research themes are the effects of local mate and resource competition (often abbreviated LMC and LRC, respectively).


Fisher's principle

Fisher's principle (1930) explains why in most species, the sex ratio is approximately 1:1. His argument was summarised by
W. D. Hamilton William Donald Hamilton (1 August 1936 – 7 March 2000) was a British evolutionary biologist, recognised as one of the most significant evolutionary theorists of the 20th century. Hamilton became known for his theoretical work expounding a ...
(1967) as follows, assuming that parents invest the same whether raising male or female offspring: :# Suppose male births are less common than female. :# A newborn male then has better mating prospects than a newborn female, and therefore can expect to have more offspring. :# Therefore parents genetically disposed to produce males tend to have more than average numbers of grandchildren born to them. :# Therefore the genes for male-producing tendencies spread, and male births become more common. :# As the 1:1 sex ratio is approached, the advantage associated with producing males dies away. :# The same reasoning holds if females are substituted for males throughout. Therefore 1:1 is the equilibrium ratio. In modern language, the 1:1 ratio is the
evolutionarily stable strategy An evolutionarily stable strategy (ESS) is a strategy (or set of strategies) that is ''impermeable'' when adopted by a population in adaptation to a specific environment, that is to say it cannot be displaced by an alternative strategy (or set o ...
(ESS). This ratio has been observed in many species, including the bee ''
Macrotera portalis ''Macrotera portalis'' is a species of communal, ground nesting, partially bivoltine bees found in arid grasslands and desert regions of North America. An oligolectic bee, ''M. portalis'' gathers pollen only from plants in the genus ''Sphaeralc ...
''. A study performed by Danforth observed no significant difference in the number of males and females from the 1:1 sex ratio.


Examples in non-human species


Environmental and individual control

Spending equal amounts of resources to produce offspring of either sex is an
evolutionarily stable strategy An evolutionarily stable strategy (ESS) is a strategy (or set of strategies) that is ''impermeable'' when adopted by a population in adaptation to a specific environment, that is to say it cannot be displaced by an alternative strategy (or set o ...
: if the general population deviates from this equilibrium by favoring one sex, one can obtain higher reproductive success with less effort by producing more of the other. For species where the cost of successfully raising one offspring is roughly the same regardless of its sex, this translates to an approximately equal sex ratio.
Bacteria Bacteria (; singular: bacterium) are ubiquitous, mostly free-living organisms often consisting of one biological cell. They constitute a large domain of prokaryotic microorganisms. Typically a few micrometres in length, bacteria were among ...
of the genus ''
Wolbachia ''Wolbachia'' is a genus of intracellular bacteria that infects mainly arthropod species, including a high proportion of insects, and also some nematodes. It is one of the most common parasitic microbes, and is possibly the most common reproducti ...
'' cause skewed sex ratios in some
arthropod Arthropods (, (gen. ποδός)) are invertebrate animals with an exoskeleton, a Segmentation (biology), segmented body, and paired jointed appendages. Arthropods form the phylum Arthropoda. They are distinguished by their jointed limbs and Arth ...
species as they kill males. Sex-ratio of adult populations of pelagic copepods is usually skewed towards dominance of females. However, there are differences in adult sex ratios between families: in families in which females require multiple matings to keep producing eggs, sex ratios are less biased (close to 1); in families in which females can produce eggs continuously after only one mating, sex ratios are strongly skewed towards females. Several species of reptiles have
temperature-dependent sex determination Temperature-dependent sex determination (TSD) is a type of environmental sex determination in which the temperatures experienced during embryonic/larval development determine the sex of the offspring. It is only observed in reptiles and teleost fish ...
, where incubation temperature of eggs determines the sex of the individual. In the
American alligator The American alligator (''Alligator mississippiensis''), sometimes referred to colloquially as a gator or common alligator, is a large crocodilian reptile native to the Southeastern United States. It is one of the two extant species in the gen ...
, for example, females are hatched from eggs incubated between , whereas males are hatched from eggs . In this method, however, all eggs in a clutch (20–50) will be of the same sex. In fact, the natural sex ratio of this species is five females to one male. In birds, mothers can influence the sex of their chicks. In
peafowl Peafowl is a common name for three bird species in the genera '' Pavo'' and '' Afropavo'' within the tribe Pavonini of the family Phasianidae, the pheasants and their allies. Male peafowl are referred to as peacocks, and female peafowl are ref ...
, maternal body condition can influence the proportion of daughters in the range from 25% to 87%. In several groups of fish, such as
wrasses The wrasses are a family, Labridae, of marine fish, many of which are brightly colored. The family is large and diverse, with over 600 species in 81 genera, which are divided into 9 subgroups or tribes. They are typically small, most of them le ...
,
parrotfish Parrotfishes are a group of about 90 fish species regarded as a family (Scaridae), or a subfamily (Scarinae) of the wrasses. With about 95 species, this group's largest species richness is in the Indo-Pacific. They are found in coral reefs, ro ...
and
clownfish Clownfish or anemonefish are fishes from the subfamily Amphiprioninae in the family Pomacentridae. Thirty species of clownfish are recognized: one in the genus '' Premnas'', while the remaining are in the genus '' Amphiprion''. In the wild, t ...
,
dichogamy Sequential hermaphroditism (called dichogamy in botany) is a type of hermaphroditism that occurs in many fish, gastropods, and plants. Sequential hermaphroditism occurs when the individual changes its sex at some point in its life. In particular, ...
— or sequential hermaphroditism — is normal. This can cause a discrepancy in the sex ratios as well. In the
bluestreak cleaner wrasse The bluestreak cleaner wrasse, ''Labroides dimidiatus'', is one of several species of cleaner wrasses found on coral reefs from Eastern Africa and the Red Sea to French Polynesia. Like other cleaner wrasses, it eats parasites and dead tissue o ...
, there is only one male for every group of 6-8 females. If the male fish dies, the strongest female changes its sex to become the male for the group. All of these wrasses are born female, and only become male in this situation. Other species, like clownfish, do this in reverse, where all start out as non-reproductive males, and the largest male becomes a female, with the second-largest male maturing to become reproductive.


Domesticated animals

Traditionally, farmers have discovered that the most economically efficient community of animals will have a large number of females and a very small number of males. A herd of cows with a few bulls or a flock of hens with one rooster are the most economical sex ratios for domesticated livestock.


Dioecious plants secondary sex ratio and amount of pollen

It was found that the amount of fertilizing pollen can influence secondary sex ratio in dioecious plants. Increase in pollen amount leads to decrease in number of male plants in the progeny. This relationship was confirmed on four plant species from three families – '' Rumex acetosa'' (
Polygonaceae The Polygonaceae are a family of flowering plants known informally as the knotweed family or smartweed—buckwheat family in the United States. The name is based on the genus '' Polygonum'', and was first used by Antoine Laurent de Jussieu in 1 ...
), ''Melandrium album'' (Caryophyllaceae), ''
Cannabis sativa ''Cannabis sativa'' is an annual herbaceous flowering plant indigenous to Eastern Asia, but now of cosmopolitan distribution due to widespread cultivation. It has been cultivated throughout recorded history, used as a source of industrial fibe ...
'' and ''
Humulus japonicus ''Humulus japonicus'', known as Japanese hop, is an ornamental plant in the family Cannabaceae. Some authorities have it as a synonym of ''Humulus scandens''. Originally native to East Asian countries such as China, Japan, Korea, and extending i ...
'' ( Cannabinaceae).


Polyandrous and cooperatively breeding homeotherms

In
charadriiform Charadriiformes (, from ''Charadrius'', the type genus of family Charadriidae) is a diverse order of small to medium-large birds. It includes about 390 species and has members in all parts of the world. Most charadriiform birds live near water an ...
birds, recent research has shown clearly that
polyandry Polyandry (; ) is a form of polygamy in which a woman takes two or more husbands at the same time. Polyandry is contrasted with polygyny, involving one male and two or more females. If a marriage involves a plural number of "husbands and wives ...
and sex-role reversal (where males care and females compete for mates) as found in
phalarope __NOTOC__ A phalarope is any of three living species of slender-necked shorebirds in the genus ''Phalaropus'' of the bird family Scolopacidae. Phalaropes are close relatives of the shanks and tattlers, the ''Actitis'' and Terek sandpipers, a ...
s, jacanas,
painted snipe The Rostratulidae, commonly known as the painted-snipes, are a family of wading birds that consists of two genera: '' Rostratula'' and '' Nycticryphes''. Description The painted-snipes are short-legged, long-billed birds similar in shape to the ...
and a few
plover Plovers ( , ) are a widely distributed group of wading birds belonging to the subfamily Charadriinae. Description There are about 66 species in the subfamily, most of them called "plover" or "dotterel". The closely related lapwing subf ...
species is clearly related to a strongly male-biased adult sex ratio. Those species with male care and polyandry invariably have adult sex ratios with a large surplus of males, which in some cases can reach as high as six males per female. Male-biased adult sex ratios have also been shown to correlate with
cooperative breeding Cooperative breeding is a social system characterized by alloparental care: offspring receive care not only from their parents, but also from additional group members, often called helpers. Cooperative breeding encompasses a wide variety of group s ...
in mammals such as alpine marmots and wild
canids Canidae (; from Latin, ''canis'', " dog") is a biological family of dog-like carnivorans, colloquially referred to as dogs, and constitutes a clade. A member of this family is also called a canid (). There are three subfamilies found within ...
. This correlation may also apply to cooperatively breeding birds, though the evidence is less clear. It is known, however, that both male-biased adult sex ratios and cooperative breeding tend to evolve where caring for offspring is extremely difficult due to low secondary productivity, as in
Australia Australia, officially the Commonwealth of Australia, is a Sovereign state, sovereign country comprising the mainland of the Australia (continent), Australian continent, the island of Tasmania, and numerous List of islands of Australia, sma ...
and
Southern Africa Southern Africa is the southernmost subregion of the African continent, south of the Congo and Tanzania. The physical location is the large part of Africa to the south of the extensive Congo River basin. Southern Africa is home to a number of ...
. It is also known that in cooperative breeders where both sexes are philopatric like the
varied sittella The varied sittella (''Daphoenositta chrysoptera'') is a small, around 10–11 cm long, songbird native to Australia. It is also known as the Australian nuthatch, orange-winged sittella and the barkpecker. Taxonomy The varied sittella was fi ...
, adult sex ratios are equally or more male-biased than in those cooperative species, such as fairy-wrens,
treecreepers The treecreepers are a family (biology), family, Certhiidae, of small passerine Aves, birds, widespread in wooded regions of the Northern Hemisphere and sub-Saharan Africa. The family contains eleven species in two genus, genera, ''Certhia'' and ...
and the
noisy miner The noisy miner (''Manorina melanocephala'') is a bird in the honeyeater family, Meliphagidae, and is endemic to eastern and southeastern Australia. This miner is a grey bird, with a black head, orange-yellow beak and feet, a distinctive yellow ...
where females always disperse.


See also

*
Evolution of sex Sexual reproduction is an adaptive feature which is common to almost all multicellular organisms and various unicellular organisms, with some organisms being incapable of asexual reproduction. Currently the adaptive advantage of sexual repro ...
*
Operational sex ratio In the evolutionary biology of sexual reproduction, operational sex ratio (OSR) is the ratio of sexually competing males that are ready to mate to sexually competing females that are ready to mate, or alternatively the local ratio of fertilizable fe ...
*
Sex allocation Sex is the trait that determines whether a sexually reproducing animal or plant produces male or female gametes. Male plants and animals produce smaller mobile gametes (spermatozoa, sperm, pollen), while females produce larger ones ( ova, of ...
*
Trivers–Willard hypothesis In evolutionary biology and evolutionary psychology, the Trivers–Willard hypothesis, formally proposed by Robert Trivers and Dan Willard in 1973, suggests that female mammals adjust the sex ratio of offspring in response to maternal condition, ...
*
XY sex-determination system The XY sex-determination system is a sex-determination system used to classify many mammals, including humans, some insects (''Drosophila''), some snakes, some fish ( guppies), and some plants ('' Ginkgo'' tree). In this system, the sex of an i ...
Humans: *
Human sex ratio In anthropology and demography, the human sex ratio is the ratio of males to females in a population. Like most sexual species, the sex ratio in humans is close to 1:1. In humans, the natural ratio at birth between males and females is sligh ...
*
List of countries by sex ratio The human sex ratio is the comparative number of males with respect to each female in a population. This is a list of sex ratios by country or region. Methodology The table's data is from ''The World Factbook'' unless noted otherwise. It show ...
*
Bride kidnapping Bride kidnapping, also known as marriage by abduction or marriage by capture, is a practice in which a man abducts the woman he wishes to marry. Bride kidnapping (hence the portmanteau bridenapping) has been practiced around the world and ...
*
Groom kidnapping Groom kidnapping, colloquially known as Pakaruah shaadi or Jabaria shaadi, is a phenomenon in the western parts of Bihar and eastern Uttar Pradesh states, more prominent in Munger and Dumka (now in Jharkhand) wherein eligible bachelors are abducte ...
*
Demographic transition In demography, demographic transition is a phenomenon and theory which refers to the historical shift from high birth rates and high death rates in societies with minimal technology, education (especially of women) and economic development, to l ...
*
Sex selection Sex selection is the attempt to control the sex of the offspring to achieve a desired sex. It can be accomplished in several ways, both pre- and post-implantation of an embryo, as well as at childbirth. It has been marketed under the title family ...
*
Sex-selective abortion and infanticide Sex-selective abortion is the practice of terminating a pregnancy based upon the predicted sex of the infant. The selective abortion of female fetuses is most common where male children are valued over female children, especially in parts of Eas ...
*
Youth bulge A population pyramid (age structure diagram) or "age-sex pyramid" is a graphical illustration of the distribution of a population (typically that of a country or region of the world) by age groups and sex; it typically takes the shape of a pyramid ...
Institutions: * Gender Balance Council


Notes


References

* Also printed as * *{{cite journal , author1=Trivers R.L. , author-link1=Robert Trivers , author2=Willard D.E., author-link2=Dan Willard , title=Natural selection of parental ability to vary the sex ratio of offspring , journal=Science , volume=179 , pages=90–2 , year=1973 , doi = 10.1126/science.179.4068.90 , pmid=4682135 , issue=4068 , bibcode = 1973Sci...179...90T , s2cid=29326420 *Rath, R.M., and Mishra A.K. (2005). ''Techniques for Sex Ratio Analysis.'' Association of Professional Geographers.


External links


CIA listing of sex ratios for individual countries (including age divisions)


Population Selection