Thehead louse (Pediculus humanus capitis) is anobligateectoparasite ofhumans.[1] Head lice are winglessinsects that spend their entire lives on the humanscalp and feed exclusively on humanblood.[1] Humans are the only knownhosts of this specific parasite, whilechimpanzees andbonobos host a closely related species,Pediculus schaeffi. Other species oflice infest most orders of mammals and all orders of birds.
Lice differ from otherhematophagic ectoparasites such asfleas in spending their entire lifecycle on a host.[2] Head lice cannot fly, and their short, stumpy legs render them incapable of jumping, or even walking efficiently on flat surfaces.[2]
The non-disease-carrying head louse differs from the related disease-carryingbody louse (Pediculus humanus humanus) in preferring to attach eggs to scalp hair rather than to clothing. The twosubspecies are morphologically almost identical, but do not normally interbreed. From genetic studies, they are thought to have diverged as subspecies about 30,000–110,000 years ago, when many humans began to wear a significant amount of clothing.[3][4] However, the degree of separation is contentious as they can produce fertile offspring in a laboratory.[5]
A much more distantly related species of hair-clinging louse, the pubic orcrab louse (Pthirus pubis), also infests humans. It is morphologically different from the other two species and is much closer in appearance to the lice which infest other primates.[6] Louse infestation of the body is known aspediculosis,pediculosis capitis for head lice,pediculosis corporis for body lice, andphthiriasis for pubic lice.[7]
Like other insects of the suborderAnoplura, adult head lice are small (2.5–3 mm long), dorsoventrally flattened (seeanatomical terms of location), and wingless.[8] Thethoracic segments are fused, but otherwise distinct from thehead andabdomen, the latter being composed of seven visiblesegments.[9] Head lice are grey in general, but their precise color varies according to the environment in which they were raised.[9] After feeding, consumed blood causes the louse body to take on a reddish color.[9]
One pair ofantennae, each with five segments, protrudes from the insect's head. Head lice also have one pair ofeyes. Eyes are present in all species within the Pediculidae family, but are reduced or absent in most other members of the Anoplura suborder.[8] Like other members of the Anoplura, head louse mouthparts are highly adapted for piercing the skin and sucking blood.[8] These mouth parts are retracted into the insect's head except during feeding.[9][10]
Sixlegs project from the fused segments of thethorax.[9] As is typical in the Anoplura, these legs are short and terminate with a singleclaw and opposing "thumb".[9] Between its claw and thumb, the louse grasps the hair of its host.[9] With their short legs and large claws, lice are well adapted to clinging to the hair of their host. These adaptations leave them incapable of jumping, or even walking efficiently on flat surfaces. Lice can climb up strands of hair very quickly, allowing them to move quickly and reach another host.[2]
Seven segments of the louseabdomen are visible.[9] The first six segments each have a pair ofspiracles through which the insect breathes.[9] The last segment contains theanus and (separately) thegenitalia.[9]
In male lice, the front two legs are slightly larger than the other four. This specialized pair of legs are 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 abdomens.
Head louse egg (nit) attached to hair shaft of host
Like most insects, head lice areoviparous. Females lay about three or four eggs per day. Louseeggs (also known asnits), are attached near the base of a host hair shaft.[11][12] Eggs are usually laid on the base of the hair, 3–5 mm off the scalp surface.[11][12] In warm climates, and especially the tropics, eggs may be laid 6 inches (15 cm) or more down the hair shaft.[13]
To attach an 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 large parts of the egg except for the operculum, a cap through which the embryo breathes.[12] The glue was previously thought to bechitin-based, but more recent studies have shown it to be made ofproteins similar to hairkeratin.[12]
Each egg is oval-shaped and about 0.8 mm in length.[12] They are bright, transparent, and tan to coffee-colored so long as they contain an embryo, but appear white after hatching.[12][13] Head lice hatch typically six to nine days afteroviposition.[11][14]
After hatching, the lousenymph 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 six or more months.[14]
SEM images of a head louse egg
Louse egg attached to a hair shaft of its host
The female reproductive organ secretes a glue that quickly hardens into a "nit sheath" to cover the hair shaft and large parts of the egg, except for the operculum.
The operculum allows the embryo to breathe.
A first-stage nymph hatching from an egg
A first-stage nymph hatching from an egg (detail)
Empty shells are matte, collapsed and white in color. The termnit may include any of the following:[15]
Of these three, only eggs containing viable embryos have the potential to infest or reinfest a host.[16] However, ano nit policy is a common public health measure to prevent transmission of lice. Some authors have therefore restricted the definition of nit to describe only a hatched or nonviable egg:
Louse hatching
In many languages, the terms used for the hatched eggs, which were obvious for all to see, have subsequently become applied to the embryonated eggs that are difficult to detect. Thus, the term "nit" in English is often used for both. However, in recent years, my colleagues and I have felt the need for some simple means of distinguishing between the two without laborious qualification. We have, therefore, come to reserve the term "nit" for the hatched and empty egg shell and refer to the developing embryonated egg as an "egg".
Development ofPediculus humanus humanus (body lice), which is similar to that of head lice (Pediculus humanus capitis)
Head lice, like other insects of the orderPhthiraptera, arehemimetabolous.[1][10] Newly hatchednymphs willmoult three times before reaching the sexually mature adult stage.[1] Thus, mobile head lice populations may contain eggs, nits, three nymphalinstars, and the adults (male and female) (imago).[1]Metamorphosis during head louse 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] as well as the existence of reproductive organs in the adults. Aside from reproduction, nymph behavior is similar to the adult. Like adults, nymphs feed also only on human blood (hematophagia), and cannot survive long away from a host.[1] Outside their hosts lice cannot survive more than 24 hrs.[23] The time required for head lice to complete their nymph development to the imago lasts for 12–15 days.[1]
Nymph mortality in captivity is 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.[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 body. This results in death within a day or two.[1] Whether the high mortality recorded under experimental conditions is representative of conditions in the wild is unclear.[1]
Copulation inPediculus humanus humanus (Pediculus humanus capitis is similar), female is on top, with the male below. Dilation of the female's vagina has already occurred, and the male's dilator rests against his back (dorsal surface), out of the way. The male vesica, which contains the penis proper (not seen), is fully inserted into the vagina. Note the male's attachment with his specialized claws on the first leg pair to the specialized notch on the female's third leg pair.
Head licereproduce sexually, andcopulation is necessary for the female to produce fertile eggs.Parthenogenesis, the production of viable offspring by virgin females, does not occur inPediculus humanus.[1] Pairing can begin within the first 10 hours of adult life.[1] After 24 hours, adult lice copulate frequently, withmating occurring during any period of the night or day.[1][24] Mating attachment frequently lasts more than an hour.[24] Young males can successfully pair with older females, and vice versa.[1]
Experiments withP. h. 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.[24] The female laid only one egg after mating, and her entire body was tinged with red—a condition attributed to rupture of thealimentary canal during the sexual act.[24] Old females frequently die following, if not during, copulation.[24] During its lifespan of 4 weeks a female louse lays 50-150 eggs. Eggs hatch within 6–9 days, each nymphal stage last for 4–5 days and accordingly the period from egg to adults lasts for 18–24 days. Adult lice live for an additional 3–4 weeks.[25]
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 socioeconomic 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 14 years of age are the most frequently infested group.[26]
All stages except eggs are blood-feeders and bite the skin four to five times daily to feed. They inject saliva which contains ananticoagulant and suck blood. The digested blood is excreted as dark redfrass.[27]
Although 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 move towards shadows or dark-coloured objects in their vicinity.[24][28]
Lice have no wings or powerful legs for jumping, so they use the claws on their legs to move from hair to hair.[27] 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, hats, brushes, towels, clothing, beds, or closets.Head-to-head contact is by far the most common route of lice transmission.[29]
About 6–12 million people, mainly children, are treated annually for head lice in the United States alone. In the UK, it is estimated that two thirds of children will experience at least one case of head lice before leaving primary school.[30] High levels of louse infestations have also been reported from all over the world, including Australia, Denmark, France, Ireland, Israel, and Sweden.[16][31]
Analysis of the DNA of lice found onPeruvianmummies may indicate that some diseases (such astyphus) may have passed from theNew World to theOld World, instead of the other way around.[32][33]
The sequencing of the genome of the body louse was first proposed in the mid-2000s[34] and the annotated genome was published in 2010.[35] An analysis of the body and head louse transcriptomes revealed these two organisms are extremely similar genetically.[36]
Unlike otherbilateral animals, the 37mitochondrial genes of human lice are not on a singlecircular chromosome but extensively fragmented. For the head louse, and the body louse, they are on 20 minichromosomes, for the pubic louse 14 minichromosomes and the chimpanzee louse,18 minichromosomes.[37]
Human lice are divided into three deeply divergent mitochondrial clades known as A, B, and C.[38][39] Three subclades have been identified, D (a sister clade of A), E (a sister clade of C), and F (a sister clade of B).[40][41][42]
^abcdefghijklmnopqrsBuxton, Patrick A. (1947). "The biology ofPediculus humanus".The Louse; an account of the lice which infest man, their medical importance and control (2nd ed.).London: Edward Arnold. pp. 24–72.
^abcdMaunder, J. W. (1983). "The Appreciation of Lice".Proceedings of the Royal Institution of Great Britain.55:1–31.
^Buxton, Patrick A. (1947). "The crab lousePhthirus pubis".The Louse; an account of the lice which infest man, their medical importance and control (2nd ed.). London: Edward Arnold. pp. 136–141.
^abcBuxton, Patrick A. (1947). "The Anoplura or Sucking Lice".The Louse; an account of the lice which infest man, their medical importance and control (2nd ed.). London: Edward Arnold. pp. 1–4.
^abcdefghijBuxton, Patrick A. (1947). "The Anatomy ofPediculus humanus".The Louse; an account of the lice which infest man, their medical importance and control (2nd ed.). London: Edward Arnold. pp. 5–23.
^abcdefBurkhart CN, Burkhart CG (July 2005). "Head lice: scientific assessment of the nit sheath with clinical ramifications and therapeutic options".Journal of the American Academy of Dermatology.53 (1):129–33.doi:10.1016/j.jaad.2005.01.134.PMID15965432.
^Pollack RJ, Kiszewski AE, Spielman A (August 2000). "Overdiagnosis and consequent mismanagement of head louse infestations in North America".The Pediatric Infectious Disease Journal.19 (8):689–93, discussion 694.doi:10.1097/00006454-200008000-00003.PMID10959734.S2CID2557006.
^Mumcuoglu KY, Meinking TA, Burkhart CN, Burkhart CG (August 2006). "Head louse infestations: the 'no nit' policy and its consequences".International Journal of Dermatology.45 (8):891–6.doi:10.1111/j.1365-4632.2006.02827.x.PMID16911370.S2CID38910169.
^Mumcuoglu KY, Miller J, Gofin R, et al. (September 1990). "Epidemiological studies on head lice infestation in Israel. I. Parasitological examination of children".International Journal of Dermatology.29 (7):502–6.doi:10.1111/j.1365-4362.1990.tb04845.x.PMID2228380.S2CID39798857.
^Mumcuoglu KY (May 2006). "Effective treatment of head louse with pediculicides".Journal of Drugs in Dermatology.5 (5):451–2.PMID16703782.
^Mumcuoglu KY, Barker SC, Burgess IE, et al. (April 2007). "International guidelines for effective control of head louse infestations".Journal of Drugs in Dermatology.6 (4):409–14.PMID17668538.
^Pittendrigh BR, Clark JM, Johnston JS, Lee SH, Romero-Severson J, Dasch GA (November 2006). "Sequencing of a new target genome: the Pediculus humanus humanus (Phthiraptera: Pediculidae) genome project".Journal of Medical Entomology.43 (6):1103–11.doi:10.1603/0022-2585(2006)43[1103:SOANTG]2.0.CO;2 (inactive 2024-11-22).PMID17162941.S2CID25046936.{{cite journal}}: CS1 maint: DOI inactive as of November 2024 (link)