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This article is about the animal. For the disease known as head lice, see Head-louse infestation.
The head louse (Pediculus humanus capitis) is an obligate ectoparasite of humans.[1] Head lice are wingless insects spending their entire life on human scalp and feeding exclusively on human blood.[1] Humans and chimpanzees are the only known host of this specific parasite, but many other species of lice are known which infest most orders of mammals and also birds.[1] The head louse differs from the related body louse in preferring to attach eggs to scalp hair rather than to clothing. Although the two species are visually identical, they do not interbreed in the wild, although they will interbreed in laboratory conditions. From genetic studies of them, they are thought to have diverged as species about 107,000 years ago, when many humans began to wear a significant amount of clothing.[2][3] A yet more distantly related species of hair-clinging louse, the pubic or crab louse (Pthirus pubis), also infests humans. It is visually different from the other two species, and is much closer in appearance to the lice which infest other primates.[4] Lice infestation of any part of the body is known as pediculosis.[5] Head lice differ from other hematophagic ectoparasites such as the flea in that lice spend their entire life cycle on a host.[6] Head lice cannot fly, and their short stumpy legs render them incapable of jumping, or even walking efficiently on flat surfaces.[6]
[edit] Adult morphologyLike other insects of the suborder Anoplura, adult head lice are small (1–3 mm long), dorso-ventrally flattened (see anatomical terms of location), and entirely wingless.[7] The thoracic segments are fused, but otherwise distinct from the head and abdomen, the latter being composed of seven visible segments.[8] Head lice are grey in general, but their precise color varies according to the environment in which they were raised.[8] After feeding, consumed blood causes the louse body to take on a reddish color.[8] [edit] Head One pair of antenna, each with five segments, protrude from the insect's head. Head lice also have one pair of eyes. Eyes are present in all species within Pediculidae (the family of which the head louse is a member), but are reduced or absent in most other members of the Anoplura suborder.[7] Like other members of Anoplura, head lice mouthparts are highly adapted for piercing skin and sucking blood.[7] These mouthparts are retracted into the insect's head except during feeding.[8][9] [edit] ThoraxSix legs project from the fused segments of the thorax.[8] As is typical in Anoplura, these legs are short and terminate with a single claw and opposing "thumb".[8] Between its claw and thumb, the louse grasps the hair of its host.[8] With their short legs and large claws, lice are well adapted to clinging to the hair of their host. However, these adaptations leave them quite incapable of jumping, or even walking efficiently on flat surfaces.[6] Like other members of Anoplura, wings are entirely absent.[7] [edit] AbdomenThere are seven visible segments of the louse abdomen.[8] The first six segments each have a pair of spiracles through which the insect breathes.[8] The last segment contains the anus and (separately) the genitalia.[8] [edit] Gender differencesIn male lice, the front two legs are slightly larger than the other four. This specialized pair of legs is used for holding the female during copulation. Males are slightly smaller than females and are characterized by a pointed end of the abdomen and a well-developed genital apparatus visible inside the abdomen. Females are characterized by two gonopods in the shape of a W at the end of their abdomen. [edit] Louse eggs Like most insects, head lice are oviparous. Louse eggs contain a single embryo, and are attached near the base of a host hair shaft.[10][11] Egg laying behavior is temperature dependent, and likely seeks to place the egg in a location that will be conducive to proper embyro development (which is, in turn, temperature dependent). In cool climates, eggs are generally laid within 1 cm of the scalp surface.[10][11] In warm climates, and especially the tropics, eggs may be laid 6 inches (15 cm) or more down the hair shaft.[12] To attach each egg, the adult female secretes a glue from her reproductive organ. This glue quickly hardens into a "nit sheath" that covers the hair shaft and the entire egg except for the operculum – a cap through which the embryo breathes.[11] The glue was previously thought to be chitin-based, but more recent studies have shown it to be made of proteins similar to hair keratin.[11] Each egg is oval-shaped and about 0.8 mm in length.[11] They are tan- to coffee-colored so long as they contain an embryo, but appear white after hatching.[11][12] Typically, a hatching time of six to nine days after oviposition is cited by authors.[10][13] However, these data are from work with body lice (not head lice) that show hatching time and hatching probability are extremely temperature dependent.[14][15] As of 2008[update], head louse hatching time and probability in situ (i.e., on a human head) have not been carefully examined. After hatching, the louse nymph leaves behind its egg shell, still attached to the hair shaft. The empty egg shell remains in place until physically removed by abrasion or the host, or until it slowly disintegrates, which may take months or years.[13] [edit] Nits
The term nit refers to either a louse egg or a louse nymph.[16] With respect to eggs, this rather broad definition includes the following:[17]
This has produced some confusion in, for example, school policy (see The "no-nit" policy) because, of the three items listed above, only eggs containing viable embryos have the potential to infest or re-infest a host.[18] Some authors have reacted to this confusion by restricting the definition of nit to describe only a hatched or nonviable egg: 
Others have retained the broad definition while simultaneously attempting to clarify its relevance to infestation:
Note that all these quotations appear to reject the notion that louse nymphs are nits, and may indicate that the nit definition is currently in flux. [edit] Development and nymphs Head lice, like other insects of the order Phthiraptera, are hemimetabolous.[1][9] Newly hatched nymphs will molt three times before reaching the sexually-mature adult stage.[1] Thus, mobile head lice populations contain members of up to four developmental stages: three nymphal instars, and the adult (imago).[1] Metamorphosis during head lice development is subtle. The only visible differences between different instars and the adult, other than size, is the relative length of the abdomen, which increases with each molt.[1] Aside from reproduction, nymph behavior is similar to the adult. Nymphs feed only on human blood (hematophagia), and cannot survive long away from a host.[1] Like adult head lice, the nymph cannot fly or jump. However, there are reports of the light-weighted nymphs being blown by wind.[1] Similarly, after a molt, the discarded exoskeleton can be later shed by the host, and may be mistakenly interpreted as a viable louse.[6] The time required for head lice to complete their nymph development to the imago depends on feeding conditions. At minimum, eight to nine days are required for lice having continuous access to a human host.[1] This experimental condition is most representative of head lice conditions in the wild. Experimental conditions where the nymph has more limited access to blood produces more prolonged development, ranging from 12 to 24 days.[1] Nymph mortality in captivity is high – about 38% – especially within the first two days of life.[1] In the wild, mortality may instead be highest in the third instar.[1] Nymph hazards are numerous. Failure to completely hatch from the egg is invariably fatal, and may be dependent on the humidity of the egg's environment.[1] Death during molting can also occur, although it is reportedly uncommon.[1] During feeding, the nymph gut can rupture, dispersing the host's blood throughout the insect. This results in death within a day or two.[1] It is unclear if the high mortality recorded under experimental conditions is representative of conditions in the wild.[1] [edit] Reproduction Adult head lice reproduce sexually, and copulation is necessary for the female to produce fertile eggs. Parthenogenesis, the production of viable offspring by virgin females, does not occur in Pediculus humanus.[1] Pairing can begin within the first 10 hours of adult life.[1] After 24 hours, adult lice copulate frequently, with mating occurring during any period of the night or day.[1][20] Mating attachment frequently lasts more than an hour.[20] Young males can successfully pair with older females, and vice versa.[1] Experiments with Pediculus humanus humanus (body lice) emphasize the attendant hazards of lice copulation. A single young female, confined with six or more males, will die in a few days, having laid very few eggs.[1] Similarly, death of a virgin female was reported after admitting a male to her confinement.[20] The female laid only one egg after mating, and her entire body was tinged with red – a condition attributed to rupture of the alimentary canal during the sexual act.[20] Old females frequently die following, if not during, intercourse.[20] Females lay about three to four eggs daily.[21] During its lifespan of four weeks a female louse lays 50–150 eggs (nits).[citation needed] [edit] Lifespan and colony persistenceA generation lasts for about one month.[citation needed] The number of children per family, the sharing of beds and closets, hair washing habits, local customs and social contacts, healthcare in a particular area (e.g. school) and socio-economic status were found to be significant factors in head louse infestation. Girls are two to four times more frequently infested than boys. Children between 4 and 13 years of age are the most frequently infested group.[22] [edit] Behavior[edit] FeedingAll stages are blood-feeders and they bite the skin four to five times daily to feed. "To feed, the louse bites through the skin and injects saliva which prevents blood from clotting; it then sucks blood into its digestive tract. Bloodsucking may continue for a long period if the louse is not disturbed. While feeding, lice may excrete dark red feces onto the skin."[23] [edit] Position on hostAlthough any part of the scalp may be colonized, lice favor the nape of the neck and the area behind the ears, where the eggs are usually laid. Head lice are repelled by light, and will move towards shadows or dark-colored objects in their vicinity.[20][24] [edit] MigrationLice have no wings and no powerful legs for jumping, so they move by using their claw-like legs to transfer from hair to hair.[23] Normally head lice infest a new host only by close contact between individuals, making social contacts among children and parent-child interactions more likely routes of infestation than shared combs, brushes, towels, clothing, beds or closets. Head-to-head contact is by far the most common route of lice transmission. [edit] DistributionAbout 6–12 million people, mainly children, are treated annually for head lice in the United States alone. High levels of louse infestations have also been reported from all over the world including Israel, Denmark, Sweden, UK, France and Australia.[18][25] [edit] As a disease vectorHead lice (Pediculus humanus capitis) are not known to be vectors of diseases, unlike body lice (Pediculus humanus humanus), which are known vectors of epidemic or louse-borne typhus (Rickettsia prowazeki), trench fever (Rochalimaea quintana) and louse-borne relapsing fever (Borrellia recurrentis). [edit] Ancient lice – use in archaeogenetics
Lice are also important in the field of Archaeogenetics. Because most "modern" human diseases have in fact recently jumped from animals into humans through close agricultural contact, and also given fact that Neolithic human populations were too scattered to support contagious "crowd" diseases, lice (along with such parasites as intestinal tapeworms) are considered to be one of the few ancestral disease infestations of humans and other hominids. As such, analysis of mitochondrial lice DNA has been used to map early human and archaic human migrations and living conditions. Because lice can only survive for a few hours or days without a human host, and because lice species are so specific to certain species or areas of the body, the evolutionary history of lice reveals much about human history. It has been demonstrated, for example, that some varieties of human lice went through a population bottleneck about 100,000 years ago (supporting the Single origin hypothesis), and also that hominid lice lineages diverged around 1.18 million years ago (probably infesting Homo erectus) before re-uniting around 100,000 years ago. This recent merging seems to argue against the Multi-regional origin of modern human evolution and argues instead for a close proximity replacement of archaic humans by a migration of anatomically modern humans, either through inter-breeding, fighting, or being more fit to use available resources. Analysis of the DNA of lice found on Peruvian mummies has led to some surprising findings. The research indicates that some diseases (like typhus) may have passed from the New World to the Old World, instead of the other way around.[26] [edit] See also
[edit] References
[edit] External links
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