Review

Biology and genetics of human head

and body lice
Aurélie Veracx and Didier Raoult
Aix Marseille Université, Unité de Recherche sur les Maladies Infectieuses et Tropicales Emergentes (URMITE), Unité Mixte de
Recherche (UMR) 63, Centre National de la Recherche Scientifique (CNRS) 7278, Institut de Recherche pour le Développement
(IRD) 198, Institut National de la Santé et de la Recherche Médicale (INSERM) 1095, 13005 Marseille, France

Head lice and body lice have distinct ecologies and differ are also currently re-emerging among homeless populations
slightly in morphology and biology, questioning their in industrialized countries [3].
taxonomic status. Over the past 10 years many genetic
studies have been undertaken. Controversial data sug- Bacteria found in lice and louse-transmitted diseases
gest that not only body lice but also head lice can serve as Body lice are responsible for the transmission of at least
vectors of Bartonella quintana, and a better understand- three bacterial diseases (Figure 1). Of these, two belong to
ing of louse epidemiology is crucial. Here, we review the a subgroup of Proteobacteria (Rickettsia prowazekii and
taxonomic studies based on biology and genetics, includ- Bartonella quintana) and one is a spirochete (Borrelia
ing genomic data on lice, lice endosymbionts, and louse- recurrentis). R. prowazekii is the etiologic agent of epidemic
transmitted bacteria. We recommend that studies of hu- typhus, B. recurrentis causes louse-borne relapsing fever,
man lice employ morphological and biological character- and B. quintana causes trench fever [4]. Two other bacteria
istics in conjunction with transcriptomic date because lice have been found in body lice, Acinetobacter spp. and Serratia
seem to differ mainly in gene expression (and not in gene marcescens [5], but it is not known if they can be transmitted
content), leading to different phenotypes. to humans by lice biting. Head lice have not been considered
vectors of human diseases. However, recently, they have
Human lice, an appropriate model of coevolution also been found to be infected by B. quintana [6–9]. Never-
Human lice theless, their role in trench fever transmission remains
The order of Phthiraptera (lice) is divided into two main undetermined. Head lice were also found to be infected with
groups: the sucking lice that comprise the Anoplura sub- Acinetobacter baumannii, but the clinical significance of this
order and the chewing lice that comprise three other finding is unknown [10]. Body lice and head lice harbor the
suborders: Amblycera, Ishnocera, and Rhynchophthirina same endosymbiotic microorganisms (Candidatus Riesia
(Figure 1) [1]. Lice are obligate ectoparasites, and each host pediculicola) that seem to be essential for the production
species has its own type of louse [2]. Indeed, parasite of nutritional components, such as B vitamins, that are
speciation often occurs at approximately the same time lacking in host blood [11,12]. The primary endosymbiont
as speciation of the host (cospeciation). The two genera of and the bacterial pathogens harbored by body lice all possess
sucking lice that parasitize humans are Pthirus and Ped- genomes that are reduced in size compared to their free-
iculus (Figure 1), which include two species of medical living close relatives [13]. Thus, lice offer an appropriate
importance, Pthirus pubis (pubic louse) and Pediculus model for understanding the coevolution of vectors, sym-
humanus. The latter is of great public health concern bionts, and pathogens in a specific niche in allopatry [13].
and consists of two ecotypes: head lice and body lice. Both
ecotypes have the same life cycle, beginning with an egg Overview
stage of approximately 7 days, followed by three instars of We provide here the first exhaustive review of data on human
approximately 3 days each before becoming adults that are head lice and body lice. First, we focus on relevant compara-
capable of reproducing. Both lice need to take regular blood tive studies on human head and body lice based on their
meals (approximately five times per day) on human skin to morphology and biology before the advent of molecular
survive. However, they live in different ecological niches. biology tools. Second, we present information on the body
Head lice live in human hair and are very commonly found louse genome, the genome of its symbiont, and some data on
among children. Due to bite reaction, they are responsible the genome of the pathogens transmitted by body lice. Final-
for a very intense pruritus that may lead to high irritation ly, the main genetic studies on human lice performed during
and even wound infection. Body lice live in clothes and are the past 10 years are reviewed and discussed, and some
associated with a lack of clothing hygiene and cold weather. inferences are made regarding the evolution of human lice.
They are often found in jails and unstable countries but
Human lice taxonomy before molecular biology
The morphology and biology of head and body lice, as
Corresponding author: Raoult, D. ([email protected]) reported over several decades, were used to assess their
Keywords: Pediculus humanus; taxonomy; biology; genetic; species; head lice;
body lice. taxonomic status (Table 1).

1471-4922/$ – see front matter ß 2012 Elsevier Ltd. All rights reserved. http://dx.doi.org/10.1016/j.pt.2012.09.003 Trends in Parasitology, December 2012, Vol. 28, No. 12 563
 Review Trends in Parasitology December 2012, Vol. 28, No. 12

Order: Phthiraptera (lice)

Suborders: Hosts
- Amblycera
- Ischnocera Chewing lice Birds and mammals
- Rhynchophrina
- Anoplura
Hamophthiriidae
Neolinognathidae
Hoplopleuridae
Enderleinellidae
Polyplacidae
Linognathidae
Ratemiidae Only eutherian mammals
Microthoraciidae Sucking lice
Echinophthiriidae
Hybophthiridae
Haematopinidae
Pecaroecidae
Pedicinidae
Pthiridae
Pediculidae
Human lice Bacteria isolated Disease

- Pthiridae
Pthirus pubis

- Pediculidae
Pediculus humanus Rickesia prowazekii Epidemic typhus
humanus (Body lice) Bartonella quintana Trench fever
Borrelia recurrens Relapsing fever
Acinetobacter baumannii No reported case
Serraa marcescens No reported case

Pediculus humanus Bartonella quintana No reported case

capis (Head lice) Acinetobacter baumannii No reported case
Wolbachia pipiens No reported case

TRENDS in Parasitology

Figure 1. Classification of the Phthiraptera. List of the main suborders of chewing and sucking lice, the main families of sucking lice, and details on the two families of
human lice and the diseases that they can transmit.

Early classification Nevertheless, data from double infestations in Ethiopia

The genus Pediculus was established by Linnaeus in 1758. support the distinction between the two species because,
This genus was applied to both head and body lice, which under conditions where interbreeding could theoretically
he termed P. humanus varieties 1 and 2 in 1767. In 1778, occur, head lice and body lice possessed distinct and non-
De Geer proposed naming these varieties P. humanus overlapping sizes [19].
capitis and P. humanus corporis, without determining
whether these varieties should be considered separate One or two species debate: physiological characteristics
species [14]. Head lice live in a different biotope (hair) than body lice
(inner part of clothes) and this may have an impact upon
One or two species debate: morphological physiological characteristics. In general, the two strains
characteristics show very similar developmental rates, but pre-imaginal
Generally, head lice are considered to be more heavily mortality of head lice is higher than for body lice [16]. Head
chitinized, smaller, darker, and with more pronounced lice are also more susceptible to starvation compared to
lateral indentations between segments of the abdomen body lice that are adapted to periodically being in clothes
than body lice [15]. Body lice were shown to have a set discarded by the host [16,17]. Head lice have a lower egg
of longitudinal muscles on the ventral body wall which is production than body lice, possibly due to their smaller
absent in head lice [16]. Another measure that distin- size, and their eggs are also slightly smaller [20]. In addi-
guishes between the two ecotypes is the length of anten- tion, the percentage of hatched eggs is higher in body lice
nae, which are longer in body lice, possibly due their [16]. Moreover, head lice are more active at lower tempe-
adaptation to darkness [17]. However, morphological rature, undoubtedly because they normally live on the
characteristics were proved to be inconsistent, with exposed scalp where the temperature is lower than within
the description of many intermediate forms [16,18]. clothes [17].
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Table 1. Chronology of the main biology-based studies on the taxonomic status of human head and body lice
Date Author Main observations Refs
1758 Linné The genus Pediculus was established
1767 Linné Description of Pediculus humanus varieties 1 and 2
1778 De Geer Description of Pediculus humanus capitis and Pediculus humanus corporis [14]
1861 Murray Lice imitate the color of the support upon which they live [26]
1915–1917 Fahrenholz Human lice description and classification based on various morphological [18]
characteristics, including size, shape and pigmentation
1917 Sikora; Bacot Evidence that P. capitis raised under P. corporis conditions become gradually [12,20]
indistinguishable from P. corporis
1917 Bacot In the laboratory, head lice sometimes lay eggs on clothes, but body lice rarely [20]
lay eggs on hairs and eggs are badly attached
Body lice have a homing instinct, but head lice do not
Head and body lice pair freely and their offspring are fertile
1917 Howlett When head lice are place on the body, they have a tendency to return to the head, [15]
but this tendency is less marked in the next generations
1918 Nuttall Feeding habits of P. capitis and P. corporis [17]
They represent extremes in the variation of the species P. humanus
1919 Nuttall Pigmentation is entirely dependent on the color of the background and is not a [26]
genetically transmitted characteristic
1919 Nuttall P. corporis is a descended from P. capitis in nature and some races of P. capitis are [17]
more labile than others
1919 Keilin and Nuttall Occurrence of an abnormal sex-ratio in the progeny of crosses and the appearance [23]
of hermaphrodites
Review of many cases supporting evidence of intermingling of the two forms of lice
1920 Nuttall Fahrenholz: description of human lice criticized [18]
Pigmentation is a poor criterion for differentiating lice
1924 Ewing Description of American lice and observation that human lice are hybrids [28–30]
1926 Description of mummy lice and comparison with contemporary lice conducted to
1929 develop an identification key for American lice
In some races of humans, a distinct variety of clothes louse developed from the
head louse, whereas this is not the case in other human races
1946 Busvine Confirmation that lice pigmentation depends on background color [27]
1948 Busvine Head lice reared in captivity without any signs of acquiring P. corporis characteristics [16]
1955 Alpatov Head lice may become body lice under body lice laboratory conditions [22]
1985 Busvine Description of head and body lice of distinct non-overlapping sizes in Ethiopia [19]

Body lice were proved to be vector of several bacterial Intermingling of capitis and corporis in nature
diseases. R. prowazekii and B. quintana were shown to be Many cases support the intermingling of the two forms of
transmitted through the voluminous (blood-contaminated) lice when they invade each other’s feeding grounds
feces that enter through bite wounds, conjunctiva, and [15,20,23]. Capitis and corporis were shown to pair freely,
respiratory membranes [21]. The vectorial capacity of head and their offspring are fertile [20]. However, there was an
lice is debated, but they also produce voluminous blood- abnormal sex-ratio in the progeny of crosses, with a
contaminated feces [21]. The vectorial capacity of body lice marked decrease in the proportion of females to males
may reflect their greater blood intake during feeding epi- and the appearance of hermaphrodites [23]. Interestingly,
sodes resulting from the more difficult access to blood for in our laboratory, we found lice eggs on a cap from a
lice in clothes because they must deal with host body homeless person, confirming that head lice may lay eggs
movements [17]. This may lead to an increased internal on clothes [24]. Finally, a study undertaken in 2003 further
pressure in the corporis form that could explain its larger confirmed that head lice may be established on the body
average size, loss of angularity in the abdominal segments, [21].
and the more widely separated hairs upon the abdominal
surface, compared to the capitis form [17]. However, the Several subspecies or varieties debate
capitis and the corporis forms feed in the same way if they Fahrenholz classified lice into six subspecies on the basis of
are reared under the same conditions [17]. lice morphology and pigmentation: three subspecies of
capitis (P. capitis angustus, P. capitis maculatus, and P.
Rearing observations capitis capitis) and three subspecies of corporis (P. nigri-
The typical capitis, which are raised on humans under tarum, P. chinensis, and P. humanus humanus). Each of
conditions that are favorable for corporis, gradually be- these species occurs on what he referred to as different
come morphologically indistinguishable from corporis after ‘human races’ [18]. However, pigmentation as a criterion to
four to five generations [22,23]. The typical capitis and describe and differentiate between lice may lead to errors
corporis forms may represent the extremes in the variation in differentiation because unpigmented structures are
of the species P. humanus [17]. However, similar work difficult to observe and may be reported as being absent
could not confirm these observations [16]. even though they are effectively present [18]. Furthermore,
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it was reported that lice imitate the color of the skin upon insect repertoire. However, despite its small size, the body
which they live [25,26]. A series of color gradations accord- louse genome is functionally complete [13]; 90% of the
ing to louse origin, ranging from the black louse to the predicted body louse genes share homology and 80% of
light-gray louse, were described. However, the accuracy of the genes show orthology to other sequenced insect genes
these results has been challenged by several authors who [33]. The genome contains 10 773 protein-coding genes and
stated that the color difference is inconsistent because a 57 microRNAs. Lice belong to the hemimetabolic insects.
large variety of louse colors can be found on a single host The louse genome composition is interesting because lice
[26]. Moreover, additional experiments showed that the could constitute an outgroup of holometabolic insects and
pigmentation was entirely dependent on the color of the because they share more orthologous genes with this group
background and was not a genetically transmitted charac- than with the well-studied Drosophila melanogaster model
teristic [26]. The variability in louse colors on a single host [13]. The genome contains significantly fewer genes asso-
may be affected not only by the color of the skin but also by ciated with environmental sensing and response. First,
the color of the hair and clothing [26,27]. odorant and gustatory receptors, as well as odorant-
Ewing also used morphological characteristics to pro- binding proteins and chemosensory proteins, do not seem
pose an identification key for American lice that included to be necessary for host location and selection because their
five varieties of human lice: P. humanus nigritarum Fab- respective genes are dramatically fewer in number than in
ricius (also known as P. humanus corporis De Geer), P. other insects [13]. Second, the genome encodes the smallest
humanus marginatus Fahrenholz, P. humanus ameri- number of detoxifying enzymes compared with other insect
canus, a new variety, and P. humanus humanus Linnaeus genomes [33]. Its obligate parasitism of a single host
[28–30]. He worked on both contemporary and mummy lice species and its simple life history may be indicative of
because he was aware that America, a melting pot of an evolutionary process that resulted in a smaller number
human races, had also become a melting pot for hybrid of specific gene families. Moreover, the louse has a single
lice from different origins. insulin-like peptide (ilp) gene, which may reflect its
restricted and homogeneous diet [13].
First assumptions about the evolution of head and body
lice The mitochondrial (mt) genome
At the time, the predominant opinion was that corporis In eukaryotes, mt chromosomes are typically circular,
descended from capitis in nature [17,28]. Indeed, it was approximately 16 kb in length, and contain 37 genes
thought that when primitive humans lost the hair that [34]. However, in lice the 37 mt genes are located on 18
covered their bodies, and began to wear clothes, lice living minicircular chromosomes instead of one single chromo-
in hair evolved to adapt to this new ecologic niche. The some. Each of the minicircular chromosomes is 3–4 kb in
variation in the time required for the adaptation of the length and contains one to three genes [35]. The circular
typical capitis form to evolve into the typical corporis form chromosomes also contain three blocks of highly conserved
illustrates that some varieties of capitis are more labile than regions that may form a stable stem–loop to initiate repli-
others [17]. This finding was also stated later by Ewing: ‘in cation and transcription. The coding regions show single-
certain races of humans a distinct variety of clothes louse nucleotide polymorphisms. There is evidence of recombi-
developed from the head-louse type for that race, while in nation between minichromosomes that is probably facili-
other races, no clothes-louse type distinctive from head louse tated by the identical sequences present on different
developed’ [28]. We will discuss this topic later in this review. minichromosomes, thus explaining the extreme sequence
variation in the noncoding regions [35]. The recombination
The louse genome of these minichromosomes may be either homologous or
Chromosome structure nonhomologous. There are also different types of chimeric
Genome sequencing of the human body louse [13] con- mt minichromosomes, in addition to the 18 mt minichro-
firmed that body lice and head lice have the smallest mosomes [36]. This novel type of circular mt chromosome is
genomes of any insect reported to date (108 Mb for females also present in the other sucking lice, but not in chewing
and 109 Mb for males), as previously estimated by flow lice or the Psocoptera. Blood-feeding appears to have co-
cytometry in 2007 [31]. Lice are diploid organisms that evolved with minicircular mt chromosomes in sucking lice
have six chromosomes (five metacentric chromosomes and [35]. Moreover, the gene content of various eukaryotic
one telocentric chromosome) [32]. The average guanine- mitochondrial genomes (including P. humanus) was inves-
cytosine (GC) content of the P. humanus genome is 28%, tigated to determine the origin of each mt gene and recon-
making this genome unusually AT rich. Transposable stitute the origin of mitochondria. This work showed that
elements represent only 1% of the genome, which is mitochondria do not have a stable or unique form, and that
markedly less than for any sequenced insect genome. Both mitochondria of different organisms do not have the same
class I and class II mobile elements are present [13]. No evolutionary history or the same number of genes [37].
genes of prokaryotic origin have been found in the louse
genome, suggesting the absence of DNA transfer from The louse endosymbiont and its genome
Candidatus Riesia pediculicola to its host [13]. Generalities about the endosymbiont
The human louse endosymbiont is a bacterium belonging
Gene content and function to the family Enterobacteriaceae in the g-Proteobacteria
The expectation for the reproductive evolution of obligate class. Its closest relatives are species in the genus Arseno-
parasites would be a reduced genome with a reduced basal phonus, and it was termed Candidatus Riesia pediculicola
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[12]. Many studies were undertaken during the past 5 Moreover, this bacterium was described as the fastest-
years on louse endosymbionts [38–40]. The microorganism evolving insect P-endosymbiont, leading to the conclusion
is primarily located in a disc-shaped organ located on the that nucleotide substitution rates decrease as the age of
ventral side of the midgut (the mycetome) and is transmit- the endosymbiosis increases to slow the overall rate of
ted from the female louse to its progeny after its migration endosymbiont extinction [44].
to the ovaries [39,41,42].
Genomic data on louse-infesting bacteria
The endosymbiont genome As mentioned above, three main intracellular bacteria are
The genome of the obligatory louse endosymbiont contains transmitted by lice: R. prowazekii, B. quintana, and B.
less than 600 genes on a short, linear chromosome and a recurrentis [4]. Interestingly, in addition to all being highly
circular plasmid. When compared with the genome of other pathogenic, these bacteria share another common charac-
endosymbionts, only 24 genes are unique to Candidatus teristic: an unusually reduced genome size compared to
Riesia pediculicola, including genes coding for transport close relatives. Hence, B. recurrentis appears to be a
and binding proteins, as well as enzymes involved in degraded subset of the tick-borne relapsing fever-causing
lipopolysaccharide biosynthesis that may be essential for agent Borrelia duttonii [45]. In addition, B. quintana is
cell-wall stability during extracellular migration [13]. described as a genomic derivative of the zoonotic agent
There are 30 genes in all bacteria studied that are absent Bartonella henselae, which is transmitted among cats by
from Candidatus Riesia pediculicola. These genes are the cat flea and to humans by cat scratches or cat bites [46].
mainly exonucleases that are required for conjugation, Finally, R. prowazekii is also known to have a reduced
and enzymes that are involved in energy metabolism, thus genome and to contain hundreds of degraded genes [47]. In
reflecting the dependence of the symbiont on its louse host fact, as bacteria interact with their environment their
for nutrients. In return, the bacterium is thought to be genetic content varies through gene gain and loss. When
required by the louse for the production of pantothenic acid a bacterium becomes intracellular the possibility of gene
(vitamin B5) [43]. The genes encoding this function are exchange is reduced, leading to gene loss and a reduction in
situated together on the plasmid, and not on the linear genome size. However, intracellular bacteria of amoebae
chromosome of the bacteria. The reduction in genome size are in sympatry with many other bacteria and viruses,
and the high AT-bias suggest an ancient association leading to a very large genome [48]. In cases of intracellular
between the louse and its primary endosymbiont [13]. bacteria living in allopatry, new characteristics may not be
However, Candidatus Riesia pediculicola is an insect acquired, and the bacteria can become specialized to their
primary endosymbiont (P-endosymbiont) that has been environment and lose the capacity to adapt to a changing
associated with the louse for only 13–25 million years. environment. A greater reduction in genome size will lead

Mulhost pathogens (generalists) with an extended vectros capacity

Associated with vectors that feed on a broad range of hosts
Borrelia duonii Bartonella henselae Rickesia conorii

Host range size of the vector

Survival potenal in novel hosts
Opportunies for gene acquision
Genome size and gene regulaon
Capability to cope with environmental changes

Pathogenicity

Single host pathogens (specialists) restricted to Pediculus humanus

Associated with a vector that has a narrow host range (only humans)

Borrelia recurrens Bartonella quintana Rickesia prowazekii

TRENDS in Parasitology

Figure 2. Reductive evolution leading to a higher pathogenicity. Reductive evolution of the highly pathogenic bacteria associated with Pediculus humanus, a narrow host-
range vector compared to less-virulent closely related bacteria associated with broad host-range vectors.

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to deregulation and a higher level of pathogenicity [49–51]. Genetic studies of human head and body lice
This explains why the bacteria of the genera Borrelia, Genetic tools questioned the division of human lice into
Bartonella and Rickettsia comprise both highly pathogenic head lice and body lice (Table 2). The first study was based
bacteria with small genomes, that are transmitted by a on the 18S rRNA gene [52], and subsequent studies focused
very specific vector (the louse), and less pathogenic bacte- on mt genes [53–56] and intergenic spacers [24,57]. These
ria, with larger genomes, that are transmitted by ticks or studies revealed that there are three clades of head lice,
fleas that feed on a larger variety of hosts (Figure 2) [45]. one of which may also be body lice (Clade A) [53,54].

Table 2. Summary of the main genetic studies on human head and body lice
DNA Type Gene Fragment length Date First author Title Ref
Mitochondrial DNA Cytochrome oxidase 524 bp 2002 Leo Evidence from mitochondrial DNA that [55]
subunit 1 (COI) head and body lice of humans are
conspecific
610 bp 2003 Kittler Molecular evolution of Pediculus [56]
humanus and the origin of clothing
524 bp 2003 Yong The geographic segregation of human [52]
lice preceded that of Pediculus
humanus capitis and Pediculus
humanus humanus
854 bp 2004 Reed Genetic analysis of lice supports direct [54]
contact between modern and archaic
humans
383 bp 2008 Raoult Molecular identification of lice from [53]
pre-Columbian mummies
827 bp 2008 Light Geographic distributions and origins of [65]
human head lice based on
mitochondrial data
Cytochrome b (Cyt b) 440 bp 2003 Kittler Molecular evolution of Pediculus [56]
humanus and the origin of clothing
671 bp 2004 Reed Genetic analysis of lice supports direct [54]
contact between modern and archaic
humans
356 bp 2008 Raoult Molecular identification of lice from [53]
pre-Colombian mummies
316 bp 2010 Li Genotyping of human lice suggest [24]
multiple emergences of body lice from
local head louse populations
NADH dehydrogenase 4 (ND4) 579 bp 2003 Kittler Molecular evolution of Pediculus [56]
humanus and the origin of clothing
Nuclear DNA Elongation factor 1a (EF-1a) 485 bp 2003 Kittler Molecular evolution of Pediculus [56]
humanus and the origin of clothing
348 bp 2003 Yong a The geographic segregation of human [52]
lice preceded that of Pediculus
humanus capitis and Pediculus
humanus humanus
RNA polymerase II (RPII) 601 bp 2003 Kittler Molecular evolution of Pediculus [56]
humanus and the origin of clothing
18S rRNA gene, small ribosomal 1474–1493 bp 2003 Yong The geographic segregation of human [52]
subunit rRNA lice preceded that of Pediculus
humanus capitis and Pediculus
humanus humanus
1195 bp 2005 Leo Unraveling the evolution of the head [66]
and Barker and body lice of humans
Microsatellites 130–180 bp 2005 Leo The head and body lice of humans are [67]
genetically distinct; evidence from
double infestations
Intergenic spacers 133–155 bp 2010 Li Genotyping of human lice suggests [24]
323–328 bp multiple emergences of body lice from
165–185 bp local head louse populations
156–189 bp
cDNA Transcript prediction 2012 Olds Comparison of the transcriptional [58]
profiles of head and body lice
Analysis based on data Comparison of phylogenetic and population 2008 Light What’s in a name: the taxonomic status [68]
available in GenBank genetic approaches of human head and body lice
Bayesian coalescent modeling approach for 2011 Toups Origin of clothing lice indicates early [69]
estimation of effective migration rates clothing use by anatomically modern
humans in Africa
a
The EF-1a sequences of this study are contaminated by fungi [67].

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TRENDS in Parasitology

Figure 3. Head lice nits. Picture taken from a highly infested homeless person.

Recently, a transcriptome study of human head and concern gene expression levels [58]. Indeed, 14 putative
body lice revealed that there is only one gene that is differentially expressed genes were identified by compar-
present in body lice but not in head lice. Otherwise, the ing head louse and body louse data. Nine head louse genes
main differences identified between head lice and body lice were more highly expressed: genes encoding a putative

(a) (b)

(c)
TRENDS in Parasitology

Figure 4. Body lice nits. Pictures taken from clothes of a highly infested homeless person: (a) a piece of pants, (b) the armpit, and (c) the collar.

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enzymatic polyprotein, a putative cuticle protein, a cyto- 4 Badiaga, S. and Brouqui, P. (2012) Human louse-transmitted
infectious diseases. Clin. Microbiol. Infect. 18, 332–337
chrome P450, a putative triadia, a putative glucose dehy-
5 La Scola, B. et al. (2001) Detection and culture of Bartonella quintana,
drogenase precursor, a putative trypsin-4 precursor, a Serratia marcescens, and Acinetobacter spp. from decontaminated
putative parathyroid precursor, and two hypothetical pro- human body lice. J. Clin. Microbiol. 39, 1707–1709
teins. Five other genes were expressed at lower levels and 6 Angelakis, E. et al. (2011) Altitude-dependent Bartonella quintana
encode an agglutinin isolectin 2 precursor, a putative genotype C in head lice, Ethiopia. Emerg. Infect. Dis. 17, 2357–2359
7 Angelakis, E. et al. (2011) Bartonella quintana in head louse nits.
Bardet–Biedl syndrome 4 (bbs4), a histone H2B.3, as well
FEMS Immunol. Med. Microbiol. 62, 244–246
as a predicted protein and a hypothetical protein of unknown 8 Bonilla, D.L. et al. (2009) Bartonella quintana in body lice and head lice
function. Thus, head lice and body lice have almost the same from homeless persons, San Francisco, California, USA. Emerg. Infect.
genomic content but are phenotypically different (different Dis. 15, 912–915
ecotypes) as a result of differential gene expression. 9 Sasaki, T. et al. (2006) First molecular evidence of Bartonella quintana
in Pediculus humanus capitis (Phthiraptera: Pediculidae), collected
from Nepalese children. J. Med. Entomol. 43, 110–112
Concluding remarks 10 Bouvresse, S. et al. (2011) No evidence of Bartonella quintana but
Body lice are only found in one lineage (Clade A). The detection of Acinetobacter baumannii in head lice from elementary
theory that body lice evolved from head lice when humans schoolchildren in Paris. Comp. Immunol. Microbiol. Infect. Dis. 34,
began to wear clothes [56] is incompatible with genetic 475–477
11 Buchner, P. (1920) Zur Kenntnis der Symbiose niederer pflanzlicher
studies. The data suggest that evolution of body lice from Organismen mit Pedikuliden. Biol. Zentbl. 39, 535–540 (in German)
head lice, and vice versa, takes place constantly among 12 Sikora, H. (1919) Vorläufige Mitteilung über Myzetome bei
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