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Sex pheromones are
pheromone A pheromone () is a secreted or excreted chemical factor that triggers a social response in members of the same species. Pheromones are chemicals capable of acting like hormones outside the body of the secreting individual, to affect the behavi ...
s released by an organism to attract an individual of the same species, encourage them to mate with them, or perform some other function closely related with
sexual reproduction Sexual reproduction is a type of reproduction that involves a complex life cycle in which a gamete ( haploid reproductive cells, such as a sperm or egg cell) with a single set of chromosomes combines with another gamete to produce a zygote tha ...
. Sex pheromones specifically focus on indicating females for breeding, attracting the opposite sex, and conveying information on species, age, sex and genotype. Non-volatile pheromones, or cuticular contact pheromones, are more closely related to social insects as they are usually detected by direct contact with chemoreceptors on the antennae or feet of insects. Insect sex pheromones have found uses in monitoring and trapping of pest insects.


Evolution

Sex pheromones have evolved in many species. The many types of pheromones (i.e. alarm, aggregation, defense, sexual attraction) all have a common cause acting as chemical cues to trigger a response. However, sex pheromones are particularly associated with signaling mating behaviors or dominance. The odors released can be seen as a favorable trait selected by either the male or female leading to attraction and copulation. Chemical signaling is also used to find genetically different mates and thus avoid inbreeding. Females are often selective when deciding to mate, and chemical communication ensures that they find a high-quality mate that satisfies their reproductive needs.


Sexual selection

Odours may be a kind of male "ornament" selected for by female choice. They meet the criteria for such ornaments that
Charles Darwin Charles Robert Darwin ( ; 12 February 1809 – 19 April 1882) was an English naturalist, geologist, and biologist, widely known for his contributions to evolutionary biology. His proposition that all species of life have descended f ...
set out in ''
The Descent of Man, and Selection in Relation to Sex ''The Descent of Man, and Selection in Relation to Sex'' is a book by English naturalist Charles Darwin, first published in 1871, which applies evolutionary theory to human evolution, and details his theory of sexual selection, a form of biolo ...
''. After many years of study the importance of such chemical communication is becoming clear.
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 t ...
s usually compete for scarce
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 ...
s, which make
adaptive Adaptation, in biology, is the process or trait by which organisms or population better match their environment Adaptation may also refer to: Arts * Adaptation (arts), a transfer of a work of art from one medium to another ** Film adaptation, a ...
choices based on male traits. The choice can benefit the female directly and/or genetically. In tiger moths (''
Utetheisa ornatrix ''Utetheisa ornatrix'', also called the bella moth, ornate moth or rattlebox moth is a moth of the subfamily Arctiinae. It is aposematically colored ranging from pink, red, orange and yellow to white coloration with black markings arranged in v ...
''), females choose the males that produce the most pheromone; an honest signal of the amount of protective
alkaloid Alkaloids are a class of basic, naturally occurring organic compounds that contain at least one nitrogen atom. This group also includes some related compounds with neutral and even weakly acidic properties. Some synthetic compounds of similar st ...
s the male has, as well as an indicator of the size of female offspring (females fertilised by such males lay more eggs). Male
cockroach Cockroaches (or roaches) are a paraphyletic group of insects belonging to Blattodea, containing all members of the group except termites. About 30 cockroach species out of 4,600 are associated with human habitats. Some species are well-known ...
es form dominance hierarchies based on pheromone "badges", while females use the same pheromone for male choice. In most species, pheromones are released by the non-limiting sex. Some female
moth Moths are a paraphyletic group of insects that includes all members of the order Lepidoptera that are not butterflies, with moths making up the vast majority of the order. There are thought to be approximately 160,000 species of moth, many o ...
s signal, but this is cheap and low risk; it means the male has to fly to her, taking a high risk. This mirrors communication with other sensory modalities, e.g. male
frog A frog is any member of a diverse and largely carnivorous group of short-bodied, tailless amphibians composing the order Anura (ανοὐρά, literally ''without tail'' in Ancient Greek). The oldest fossil "proto-frog" ''Triadobatrachus'' is ...
s croak; male
bird Birds are a group of warm-blooded vertebrates constituting the class Aves (), characterised by feathers, toothless beaked jaws, the laying of hard-shelled eggs, a high metabolic rate, a four-chambered heart, and a strong yet lightweig ...
s are usually colourful. Male long-range pheromone signals may be associated with patchy resources for the female. In some species, both sexes signal. Males can sometimes attract other males instead, the sex pheromone acting as an aggregation pheromone.


External fertilization and chemical duets

It is likely that most externally fertilizing species (e.g. marine worms,
sea urchin Sea urchins () are spiny, globular echinoderms in the class Echinoidea. About 950 species of sea urchin live on the seabed of every ocean and inhabit every depth zone from the intertidal seashore down to . The spherical, hard shells ( tests) ...
s) coordinate their sexual behaviour (release of
sperm Sperm is the male reproductive cell, or gamete, in anisogamous forms of sexual reproduction (forms in which there is a larger, female reproductive cell and a smaller, male one). Animals produce motile sperm with a tail known as a flagellum ...
and eggs) using pheromones. This coordination is very important because
sperm Sperm is the male reproductive cell, or gamete, in anisogamous forms of sexual reproduction (forms in which there is a larger, female reproductive cell and a smaller, male one). Animals produce motile sperm with a tail known as a flagellum ...
are diluted easily, and are short-lived. Coordination therefore provides a selective advantage to both males and females: individuals that do not coordinate are unlikely to achieve fertilisation and hence to leave offspring. The main selective advantage of outcrossing is that it promotes the masking of deleterious recessive alleles, while inbreeding promotes their harmful expression.


In humans

No study has led to the isolation of true
human sex pheromones No study has led to the isolation of true human sex pheromones, though various researchers have investigated the possibility of their existence. Sex pheromones are chemical (olfactory) signals, pheromones, released by an organism to attract an ind ...
. While humans are highly dependent upon visual cues, when in close proximity, smells also play a role in sociosexual behaviors. An inherent difficulty in studying human pheromones is the need for cleanliness and odorlessness in human participants.


Signalling

Different species use a wide variety of chemical substances to send sexual signals. The first to be described chemically was
bombykol Bombykol is a pheromone released by the female silkworm moth to attract mates. It is also the sex pheromone in the wild silk moth (''Bombyx mandarina''). Discovered by Adolf Butenandt in 1959, it was the first pheromone to be characterized chemica ...
, the
silkworm moth The domestic silk moth (''Bombyx mori''), is an insect from the moth family Bombycidae. It is the closest relative of ''Bombyx mandarina'', the wild silk moth. The silkworm is the larva or caterpillar of a silk moth. It is an economically im ...
's sex pheromone, which is a complex alcohol, (E,Z)-10,12-hexadecadienol, discovered in 1959. It is detected in the antennae of the male moth by a pheromone-binding protein which carries the bombykol to a receptor bound to the membrane of a nerve cell. The chemicals used by other moths are species-specific. For example, the Eastern spruce budworm ''
Choristoneura fumiferana ''Choristoneura fumiferana'', the eastern spruce budworm, is a species of moth of the family Tortricidae native to the eastern United States and Canada. The caterpillars feed on the needles of spruce and fir trees. Eastern spruce budworm po ...
'' female pheromones contain a 95:5 mix of ''E-'' and ''Z'' 11-tetradecenal
aldehyde In organic chemistry, an aldehyde () is an organic compound containing a functional group with the structure . The functional group itself (without the "R" side chain) can be referred to as an aldehyde but can also be classified as a formyl grou ...
s, while the sex pheromones of other species of
spruce budworm ''Choristoneura'' is a genus of moths in the family Tortricidae The Tortricidae are a family of moths, commonly known as tortrix moths or leafroller moths, in the order Lepidoptera. This large family has over 11,000 species described, and ...
contain
acetate An acetate is a salt formed by the combination of acetic acid with a base (e.g. alkaline, earthy, metallic, nonmetallic or radical base). "Acetate" also describes the conjugate base or ion (specifically, the negatively charged ion called a ...
s and
alcohol Alcohol most commonly refers to: * Alcohol (chemistry), an organic compound in which a hydroxyl group is bound to a carbon atom * Alcohol (drug), an intoxicant found in alcoholic drinks Alcohol may also refer to: Chemicals * Ethanol, one of se ...
s. Sexual development in the freshwater
green alga The green algae (singular: green alga) are a group consisting of the Prasinodermophyta and its unnamed sister which contains the Chlorophyta and Charophyta/Streptophyta. The land plants ( Embryophytes) have emerged deep in the Charophyte alga a ...
''
Volvox ''Volvox'' is a polyphyletic genus of chlorophyte green algae in the family Volvocaceae. It forms spherical colonies of up to 50,000 cells. They live in a variety of freshwater habitats, and were first reported by Antonie van Leeuwenhoek in 17 ...
'' is initiated by a
glycoprotein Glycoproteins are proteins which contain oligosaccharide chains covalently attached to amino acid side-chains. The carbohydrate is attached to the protein in a cotranslational or posttranslational modification. This process is known as glycos ...
pheromone. It is one of the most potent known biological effector molecules, as it can trigger sexual development at a
concentration In chemistry, concentration is the abundance of a constituent divided by the total volume of a mixture. Several types of mathematical description can be distinguished: '' mass concentration'', '' molar concentration'', '' number concentration'', ...
as low as 10−16 moles per litre. Kirk and Kirk showed that sex-inducing pheromone production can be triggered experimentally in somatic cells by heat shock.


Uses

Sex pheromones have found applications in pest monitoring and pest control. For monitoring,
pheromone trap A pheromone trap is a type of insect trap that uses pheromones to lure insects. Sex pheromones and aggregating pheromones are the most common types used. A pheromone-impregnated lure, as the red rubber septa in the picture, is encased in a conven ...
s are used to attract and catch a sample of pest insects to determine whether control measures are needed. For control, much larger quantities of a sex pheromone are released to disrupt the mating of a pest species. This can be either by releasing enough pheromone to prevent males from finding females, effectively drowning out their signals, or by mass trapping, attracting and removing pests directly. For example, research on the control of the spruce bud moth ('' Zeiraphera canadensis)'' has focused on the use of the pheromone E-9-tetradecenyl-acetate, a chemical the spruce bud moth releases during mating.


References

{{reflist , 30em Sexual reproduction Pheromones Chemical ecology