Original article

Oxford, UK
International
IJD
Blackwell
0011-9059
Suppl.
44 Publishing,
Science,
Journal
2005
of
Ltd.
Dermatology

The biology of hair shape

Biology of
Thibaut andhair
Bernard
shape

Sebastien Thibaut, DES, and Bruno A. Bernard, Dr es Sci

LOreal Recherche, Clichy, France

Correspondence
B. A. Bernard, Dr es Sci
Head of Hair Biology Group
Center de Recherche C. Zviak
LOral
90 rue du Gnral Roguet
92110 Clichy
France
E-mail: bbernard@rd.loreal.com

Few studies have been devoted to the biology of African hair,
although its curly appearance is a characteristic trait. Pub-
lished reports have mainly focused on hair density and hair
cycle,1,2 growth rate,2 chemical analysis35 and scanning
electron microscopy.6 Many different definitions have been
used in the literature to describe the curly nature of hair.
Hrdy5,7,8 developed a nomenclature which has been widely
accepted. The terminology defines a kink as a sudden con-
striction and twisting of the hair shaft, producing an obvious
discontinuity in the curvature.7 Many theories were put for-
ward to explain why hair crimped and curled. One such the-
ory was based on the geometric shape of the hair shaft. It was
indeed generally considered that hair shape was related to the
cross-section of the hair shaft. A circular cross-section was
assumed to give rise to straight hair, while a pronounced ellip-
tical cross-section was supposed to produce curly hair. It was
well known that African hair had a large maximum diameter
and a small minimum diameter, resulting in the most elliptical
shaft found in any racial group. However, Hrdy had observed
that the curliest hair often was not the most elliptical, the rela-
tionship between hair shaft cross-sectional shape and hair
form being thus open to debate.
Other theories were based on the morphology of the hair
bulb, and an explanation of crimping as being a result of
keratinization within a retroverted hair bulb was proposed.9
It had been observed that the retroverted hair bulb had a golf
club-like shape which could determine the slope and direction
of the emerging hair shaft. Asymmetry of keratinization was
another morphological factor associated with crimping. This
asymmetry was believed to result in a bilaterality of fiber for-
mation,9 but this phenomenon remained unexplored. Other
researchers suggested that the inner root sheath (IRS) played
a major role in hair fiber molding. In all follicles, the IRS starts
2 keratinizing at a lower level than does the hair, thus building
a tube through which the hair emerges. It was postulated that
the IRS was thicker on the concave side of wool fibers,9 and
that this could play a part in molding the less keratinized hair
shaft into its crimped shape. Finally, it was suggested that hair
slope and whorl patterns were controlled by the dermis.10 Only
one study suggested that the shape and morphology of the hair
follicle itself may play an important role in the macroscopic
appearance of hair.11 Using 3D computer-aided reconstruc-
tion, it was indeed concluded that hair form was governed by
the shape of the hair follicle and that a straight follicle even with
an oval cross-section might produce a straight hair shaft.11
Recently, we provided evidence that human curly hair
follicles exhibited a retrocurvature similar to that of wool
hair follicles.12 The asymmetry of the curly hair bulb and the
alteration of the various programs of differentiation were
found to be very close to the morphological features of wool
follicles. In addition, it was demonstrated that such character-
istics were not specific to African hair type, as both curly
African and Caucasian hairs revealed the same pattern.
The direct comparison of straight hair and curly hair high-
lighted the altered differentiation programs involved in curly
hair generation. The proliferative compartment of the follicle,
identified as Ki-67 positive matrix cells, clearly extended
above the Auber line on the convex side. In contrast, hair cuti-
cle, IRS and outer root sheath (ORS) differentiation programs
started earlier on the concave side of the follicle, as shown by
hHb2, transglutaminase-1 and cytokeratin-14 expression,
respectively. Serial cross-sections of curly hair bulb labeled
with hHb2 antibody showed the progressive closure of the
shaft cuticle ring and confirmed the lack of axial symmetry at
this level, where the hair shaft keratin network was still loose
and not yet organized.13
The behavior of curly hair in in vitro culture provided fur-
ther useful observations.12,14 When the 2-mm-long proximal

International Journal of Dermatology 2005, 44 (Suppl. 1), 2 3 2005 The International Society of Dermatology
Thibaut and Bernard
part of the follicle was put into culture, thus avoiding any
influence of the appending sebaceous gland, arrector pili
muscle or dermal environment, curly hair maintained its curly
growth throughout the duration of the experiment. In other
words, the shape of the hair shaft appeared to be intrinsically
programmed by the lower half of the follicle, independent of
the dermal environment.
In addition, immunohistologic study of this anagen hair
follicle curvature revealed that some ORS cells expressed -
smooth muscle actin (-SMA) on the concave side. Immuno-
histochemical and immunoelectron microscopic studies of
human anagen follicles revealed the presence of -SMA in
fibroblasts located in the innermost layer of the transverse
collagenous fibers of the connective tissue sheath. These cells
are thought to play an important role in the hair growth
cycle15. Changes in -SMA expression in the dermal sheath
during the hair cycle indeed suggested that -SMA, usually
characteristic of dermal sheath cells (or myofibroblasts), could
be related to contractile processes that might control hair
follicle shortening. In nonmuscle cells, bundles of -SMA
filaments associated with other proteins perform specific
functions such as contraction, mechanical support, and cell
adhesion.16,17 -SMA expression could hold another key to
curly hair follicle morphology; its control might provide a
way of modifying hair shape, for example by altering the
ellipticity of the growing fiber. Indeed, both connective tissue
sheath thickening and curvature-restricted -SMA expression
undersigned a mechanical stress occurring at the site of hair
fiber structure building-up.13
In conclusion, in vitro growth strongly suggests that curva-
ture of curly hair is programmed from the basal area of the
follicle. This bending process is apparently linked to a lack of
axial symmetry in the lower part of the bulb, affecting con-
nective tissue sheath, ORS, IRS and the hair shaft cuticle. In
addition, curly shape seems to be independent of dermal envi-
ronment, as retrocurvature is maintained in in vitro culture.
The pronounced ellipsoidal cross-section of the hair shaft
could be caused by a heterogeneous and dissymmetric hair
fiber framework, in addition to an internal mechanical stress,
as suggested by -SMA expression in a restricted area of the
ORS. The demonstration of the bulbar origin of hair curvature
opens up a new field in the biological control of hair shape.
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