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A sex ratio is 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 males to females in a
population Population is a set of humans or other organisms in a given region or area. Governments conduct a census to quantify the resident population size within a given jurisdiction. The term is also applied to non-human animals, microorganisms, and pl ...
. 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. However, many species deviate from an even sex ratio, either periodically or permanently. These include
parthenogenic Parthenogenesis (; from the Greek + ) is a natural form of asexual reproduction in which the embryo develops directly from an egg without need for fertilization. In animals, parthenogenesis means the development of an embryo from an unfertiliz ...
and androgenetic species, periodically mating organisms such as aphids, some
eusocial Eusociality ( Greek 'good' and social) is the highest level of organization of sociality. It is defined by the following characteristics: cooperative brood care (including care of offspring from other individuals), overlapping generations wit ...
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 ...
s,
bee Bees are winged insects closely related to wasps and ants, known for their roles in pollination and, in the case of the best-known bee species, the western honey bee, for producing honey. Bees are a monophyletic lineage within the superfamil ...
s,
ant Ants are Eusociality, eusocial insects of the Family (biology), family Formicidae and, along with the related wasps and bees, belong to the Taxonomy (biology), order Hymenoptera. Ants evolved from Vespoidea, vespoid wasp ancestors in the Cre ...
s, and
termite Termites are a group of detritivore, detritophagous Eusociality, eusocial cockroaches which consume a variety of Detritus, decaying plant material, generally in the form of wood, Plant litter, leaf litter, and Humus, soil humus. They are dist ...
s.


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 ** The tertiary sex ratio 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 f ...
(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 conceptually distinct, researchers have sometimes equated the ASR with the OSR, particularly in experimental studies of animals where the difference between the two values may not always be readily apparent. * — 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 certain species that have distinct unisexual individuals, each producing either male or female gametes, either directly (in animals) or indirectly (in seed plants). Dioecious reproduction is ...
species, what is the equilibrium sex ratio maintained by natural selection? # For a sequential hermaphrodite, what is the equilibrium sex order and time of sex change? # 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. This means that 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 of ...
. 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 ''Sphaeralcea ...
''. A study performed by Danforth observed no significant difference in the number of males and females from the 1:1 sex ratio.


Human sex ratio

The
human sex ratio The human sex ratio is the ratio of males to females in a population in the context of anthropology and demography. In humans, the natural sex ratio at birth is slightly biased towards the male sex. It is estimated to be about 1.05 worldwide or ...
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. As the practice overwhelmingly targets female fetuses, sex-selective abortion often specifically refers to female-selective abortion. ...
and
infanticide Infanticide (or infant homicide) is the intentional killing of infants or offspring. Infanticide was a widespread practice throughout human history that was mainly used to dispose of unwanted children, its main purpose being the prevention of re ...
. Exposure to pesticides and other environmental contaminants may be a significant contributing factor as well. As of 2024, the global sex ratio at birth is estimated at 107 boys to 100 girls (1,000 boys per 934 girls). (2023 estimates) By old age, the sex ratio reverses, with 81 older men for every 100 older women; across all ages, the global population is nearly balanced, with 101 males for every 100 females.


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 of ...
: 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 (; : bacterium) are ubiquitous, mostly free-living organisms often consisting of one Cell (biology), biological cell. They constitute a large domain (biology), domain of Prokaryote, prokaryotic microorganisms. Typically a few micr ...
of the genus ''
Wolbachia ''Wolbachia'' is a genus of gram-negative bacteria infecting many species of arthropods and filarial nematodes. The symbiotic relationship ranges from parasitism to obligate mutualism. It is one of the most common parasitic microbes of arthrop ...
'' cause skewed sex ratios in some
arthropod Arthropods ( ) are invertebrates in the phylum Arthropoda. They possess an arthropod exoskeleton, exoskeleton with a cuticle made of chitin, often Mineralization (biology), mineralised with calcium carbonate, a body with differentiated (Metam ...
species as they kill males. Sex-ratio of adult populations of pelagic
copepods Copepods (; meaning 'oar-feet') are a group of small crustaceans found in nearly every freshwater and saltwater habitat. Some species are planktonic (living in the water column), some are benthic (living on the sediments), several species have ...
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 observed in reptiles and teleost fish, ...
, where incubation temperature of eggs determines the sex of the individual. In the American alligator, 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 two bird species of the genus '' Pavo'' and one species of the closely related genus '' Afropavo'' within the tribe Pavonini of the family Phasianidae (the pheasants and their allies). Male peafowl are referred t ...
, maternal body condition can influence the proportion of daughters in the range from 25% to 87%.
Dichogamy Sequential hermaphroditism (called dichogamy in botany) is one of the two types of hermaphroditism, the other type being simultaneous hermaphroditism. It occurs when the organism's sex changes at some point in its life. A sequential hermaphrodit ...
(sequential hermaphroditism) is normal in several groups of fish, such as
wrasses The wrasses are a family, Labridae, of marine ray-finned fish, many of which are brightly colored. The family is large and diverse, with over 600 species in 81 genera, which are divided into nine subgroups or tribes. They are typically small, ...
,
parrotfish Parrotfish (named for their mouths, which resemble a parrot's beak) are a clade of fish placed in the tribe Scarini of the wrasse family (Labridae). Traditionally treated as their own family (Scaridae), genetic studies have found them to be dee ...
and
clownfish Clownfishes or anemonefishes (genus ''Amphiprion'') are saltwater fishes found in the warm and tropical waters of the Indo-Pacific. They inhabit mainly coral reefs and have a distinctive colouration typically consisting of white vertical bars on ...
. 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 off ...
, 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 Sorrel (''Rumex acetosa''), also called common sorrel or garden sorrel, is a perennial herbaceous plant in the family Polygonaceae. Other names for sorrel include spinach dock and narrow-leaved dock ("dock" being a common name for the genus ''Ru ...
'' (
Polygonaceae The Polygonaceae are a Family (biology), family of flowering plants known Common name, informally as the knotweed family or smartweed—buckwheat family in the United States. The Botanical name, name is Basionym, based on the genus ''Polygonum'', ...
), ''
Silene alba ''Silene latifolia'', commonly known as white campion, is a dioecious flowering plant in the family Caryophyllaceae, native to most of Europe, Western Asia and northern Africa. It is a herbaceous annual, occasionally biennial or a short-lived p ...
'' (Caryophyllaceae), ''
Cannabis sativa ''Cannabis sativa'' is an annual Herbaceous plant, herbaceous flowering plant. The species was first classified by Carl Linnaeus in 1753. The specific epithet ''Sativum, sativa'' means 'cultivated'. Indigenous to East Asia, Eastern Asia, the pla ...
'' and ''
Humulus japonicus ''Humulus japonicus'', known as Japanese hops, 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 ...
'' ( Cannabinaceae).


Polyandrous and cooperatively breeding homeotherms

In charadriiform 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 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, and also ...
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 members of a widely distributed group of wader, wading birds of subfamily Charadriinae. The term "plover" applies to all the members of the subfamily, though only about half of them include it in their name. Species lis ...
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 marmot The alpine marmot (''Marmota marmota'') is a large ground-dwelling squirrel, from the genus of marmots. It is found in high numbers in mountainous areas of central and southern Europe, at heights between in the Alps, Carpathians, Tatras and N ...
s and wild
canids Canidae (; from Latin, ''canis'', "dog") is a biological family of caniform carnivorans, constituting a clade. A member of this family is also called a canid (). The family includes three subfamilies: the Caninae, and the extinct Borophaginae ...
. 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 country comprising mainland Australia, the mainland of the Australia (continent), Australian continent, the island of Tasmania and list of islands of Australia, numerous smaller isl ...
and
Southern Africa Southern Africa is the southernmost region of Africa. No definition is agreed upon, but some groupings include the United Nations geoscheme for Africa, United Nations geoscheme, the intergovernmental Southern African Development Community, and ...
. 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 songbird native to Australia. About 10–11 cm long, it is also known as the Australian nuthatch, orange-winged sittella and the barkpecker. Taxonomy The varied sittella was fir ...
, 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. The family contains nine species in one genus, ''Certhia''. Their plumage is dull-coloured. As t ...
and the noisy miner where females always disperse.


See also

*
Evolution of sex Sexually reproducing animals, plants, fungi and protists are thought to have evolved from a common ancestor that was a single-celled eukaryotic species. Sexual reproduction is widespread in eukaryotes, though a few eukaryotic species have sec ...
*
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 f ...
*
Sex allocation Sex allocation is the allocation of resources to male versus female reproduction in sexual species.Charnov EL. (1982). ''The Theory of Sex Allocation''. Princeton University Press, Princeton, NJ. Sex allocation theory tries to explain why many spec ...
*
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 present in many mammals (including humans), some insects (''Drosophila''), some snakes, some fish (guppy, guppies), and some plants (''Ginkgo'' tree). In this system, the sex of an ...
Humans: *
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 shows ...
*
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 t ...
* Groom kidnapping *
Demographic transition In demography, demographic transition is a phenomenon and theory in the Social science, social sciences referring to the historical shift from high birth rates and high Mortality rate, death rates to low birth rates and low death rates as societi ...
*
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. As the practice overwhelmingly targets female fetuses, sex-selective abortion often specifically refers to female-selective abortion. ...
*
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 ...


References


Further reading

* Also printed as * * *


External links


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


{{Authority control Population Selection Biostatistics