traumatic insemination

Traumatic insemination, also known as hypodermic insemination, is the mating practice in some species of invertebrates in which the male pierces the female's abdomen with his aedeagus and injects his sperm through the wound into her abdominal cavity (hemocoel). The sperm diffuse through the female's hemolymph, reaching the ovaries and resulting in fertilization.

The process is detrimental to the female's health. It creates an open wound which impairs the female until it heals, and is susceptible to infection. The injection of sperm and ejaculatory fluids into the hemocoel can also trigger an immune reaction in the female. Bed bugs, which reproduce solely by traumatic insemination, have evolved a pair of sperm-receptacles, known as the spermalege. It has been suggested that the spermalege reduces the direct damage to the female bed bug during traumatic insemination. However experiments found no conclusive evidence for that hypothesis; as of 2003, the preferred explanation for that organ is hygienic protection against bacteria.

The evolutionary origins of traumatic insemination are disputed. Although it evolved independently in many invertebrate species, traumatic insemination is most highly adapted and thoroughly studied in bed bugs, particularly ''Cimex lectularius''. Traumatic insemination is not limited to male-female couplings, or even couplings of the same species. Both homosexual and inter-species traumatic inseminations have been observed.

Mechanics

In humans and other vertebrates, blood and lymph circulate in two different systems, the circulatory system and lymphatic system, which are enclosed by systems of capillaries, veins, arteries, and nodes. This is known as a closed circulatory system. Insects, however, have an open circulatory system in which blood and lymph circulate unenclosed, and mix to form a substance called hemolymph. All organs of the insect are bathed in hemolymph, which provides oxygen and nutrients to all of the insect's organs.

Following traumatic insemination, sperm can migrate through the hemolymph to the female's ovaries, resulting in fertilization. The exact mechanics vary from taxon to taxon. In some orders of insects, the male genitalia ( paramere) enters the female's genital tract, and a spine at its tip pierces the wall of the female's bursa copulatrix. In others, the male penetrates the outer body wall. In either case, following penetration, the male ejaculates into the female. The sperm and ejaculatory fluids diffuse through the female's hemolymph. The insemination is successful if the sperm reach the ovaries and fertilize an ovum.

Female resistance to traumatic insemination varies from one species to another. Females from some genera, including ''Cimex'', are passive prior to and during traumatic insemination. Females in other genera resist mating and attempt to escape. This resistance may not be an aversion to pain caused by the insemination, as observational evidence suggests that insects do not feel pain.

Research into the paternity of offspring produced by traumatic insemination has found "significant" last-sperm precedence. That is, the last male to traumatically inseminate a female tends to sire most of the offspring from that female.

Evolutionary adaptation

Many reasons for the evolutionary adaptation of traumatic insemination as a mating strategy have been suggested. One is that traumatic insemination is an adaptation to the development of the mating plug, a reproductive mechanism used by many species. Once a male finishes copulating, he injects a glutinous secretion into the female's reproductive tract, thereby "literally glu ngher genital tract closed". Traumatic insemination allows subsequent males to bypass the female's plugged genital tract, and inject sperm directly into her circulatory system.

Others have argued that the practice of traumatic insemination may have been an adaptation for males to circumvent female resistance to mating to eliminate courtship time, allowing one male to inseminate many mates when contact between them is brief; or that it evolved as a new development in the sperm competition as a means to deposit sperm as close to the ovaries as possible.

This bizarre method of insemination probably evolved as male bed bugs competed with each other to place their sperm closer and closer to the mother lode of eggs, the ovaries. Some male insects evolved long penises with which they enter the vagina but bypass the female's storage pouch and deposit their sperm further upstream close to the ovaries. A few males, notably among bed bugs, evolved traumatic insemination instead, and eventually this strange procedure became the norm among these insects.

It has recently been discovered that members of the plant bug genus ''Coridromius'' (Miridae) also practice traumatic insemination. In these bugs, the male intromittent organ is formed by the coupling of the aedeagus with the left paramere, as in bed bugs. Females also exhibit paragenital modifications at the site of intromission, which include grooves and invaginated copulatory tubes to guide the male paramere. The evolution of traumatic insemination in ''Coridromius'' represents a third independent emergence of this form of mating within the true bugs.

Health repercussions

While advantageous to the reproductive success of the individual male, traumatic insemination imposes a cost on females: reduced lifespan and decreased reproductive output. "These ostsinclude (i) repair of the wound, (ii) leakage of blood, (iii) increased risk of infection through the puncture wound, and (iv) immune defence against sperm or accessory gland fluids that are introduced directly into the blood."

The male bed bug aedeagus has been shown to carry five (human) pathogenic microbes, and the exoskeleton of female bed bugs nine, including ''Penicillium chrysogenum'', '' Staphylococcus saprophyticus'', ''Stenotrophomonas maltophilia'', '' Bacillus licheniformis'', and '' Micrococcus luteus''. Tests with blood agar have shown some of these species can survive ''in vivo''. This suggests infections from these species may contribute to the increased mortality rate in bed bugs due to traumatic insemination.

The successive woundings each require energy to heal, leaving less energy available for other activities. Also, the wounds provide a possible point of infection which can reduce the female's lifespan. Once in the hemolymph, the sperm and ejaculatory fluids may act as antigens, triggering an immune reaction.

There is a tendency for dense colonies of bed bugs kept in laboratories to go extinct, starting with adult females. In such an environment, where mating occurs frequently, this high rate of adult female mortality suggests traumatic insemination is very detrimental to the female's health. The damage done, and the (unnecessarily) high mating rate of captive bed bugs, have been shown to cause a 25% higher-than-necessary mortality rate for females.

Bed bug adaptation

The effects of traumatic insemination are deleterious